You are on page 1of 670

CAB International 2009.

The Mango, 2nd Edition: Botany, Production and Uses


(ed. R.E. Litz) 1
1 Introduction: Botany and Importance
S.K. Mukherjee
1
and R.E. Litz
2
1
Calcutta University, Calcutta, India
2
University of Florida, Florida, USA
1.1 Introduction 1
1.2 Description of Mango 2
The tree 2
Flowers 2
The fruit 3
The seeds and polyembryony 4
1.3 History of Cultivation 5
Origin of Mangifera indica 5
Domestication of mango 9
Distribution 10
1.4 Germplasm Conservation 11
Genetic erosion 11
Collection and documentation of Mangifera germplasm 12
Relevance of germplasm resources to mango improvement 12
1.5 Importance of Mango 12
Cultivars 12
1.6 Production and Uses 14
1.1 Introduction
Mango has become a major fruit crop of the tropics and subtropics, particu-
larly in Asia, where the mango has always been the most important fruit crop
and where it has been considered the king of fruits (Purseglove, 1972). A
generation ago, the Green Revolution culminated, creating surpluses of sta-
ple and horticultural crops in many developing countries. The Green Revo-
lution was the result of nearly a century of effort of applying Mendelian
genetics to crop improvement (i.e. conventional breeding) together with the
optimization of agronomic and horticultural practices and the successful
management of insect pests and diseases. However, improvement of tree
S.K. Mukherjee and R.E. Litz 2
crops has lagged far behind eld crops for several reasons: their heterogene-
ity, polyploidy, lengthy juvenile period, time required for evaluation of trees
in the eld, and the relatively high cost of maintaining tree plantings. For the
most part, fruit cultivars continue to be ancient selections, many of which
have serious problems, including alternate bearing, lack of disease resistance,
low yields, etc. The rapid growth of mango production in recent years has
been due to its expansion into new growing regions of the New World, China
and parts of Africa; the planting of regular bearing selections; and the adop-
tion of modern eld practices, which include irrigation management, control
of owering, etc. Agricultural practices are currently undergoing another
revolution, as integrated pest and disease management replaces the earlier
reliance on agrichemicals, and emerging elds within biotechnology begin to
impact cultivar development.
1.2 Description of Mango
The tree
The mango tree is believed to have evolved as a canopy layer or emergent
species of the tropical rainforest of South and South-east Asia (Kaur et al.,
1980; Bompard, Chapter 2, this volume). Mature trees can attain a height of
40 m or more, and can survive for several hundred years. Mango trees that
have been domesticated by selection from openly pollinated seedling popu-
lations show variation in tree architecture (i.e. shape and size). The tree is an
arborescent evergreen. Leaves are simple and alternate, with petioles that
range in length from 1 to 12.5 cm. Leaf morphology is highly variable, de-
pending on the cultivar: leaves can be lanceolate, oblong, ovate and interme-
diate types involving these forms. Leaf length ranges from 12 to 38 cm and
width can be between 213 cm. Young leaves are copper-coloured, changing
gradually to light and then dark green with age. The leaves are spirally
arranged in whorls and are produced in ushes. The canopy is normally
oval, elongated or dome shaped. The juvenile period of seedling trees can
range from 3 to 7 years. The root system consists of a long, vigorous taproot
and abundant surface feeder roots.
Flowers
Mango owers are borne on terminal pyramidal panicles, and are glabrous
or pubescent; the inorescence is rigid and erect, up to 30 cm long, and is
widely branched, usually tertiary, although the nal branch is always cymose.
The inorescence is usually densely owered with hundreds of small ow-
ers, which are 510 mm in diameter. The owers are either monoecious or
polygamous, and both monoecious and polygamous owers are borne
within a single inorescence (Plate 1). The pistil aborts in male owers. The
ratio of monoecious to polygamous owers is strongly inuenced by
Introduction: Botany and Importance 3
environmental and cultural factors. The owers have four or ve sepals and
petals that are ovate to ovoid to lanceolate and also thinly pubescent. The
oral disc also is four- or ve-lobed, eshy and large and located above the
base of the petals. There are ve large, eshy stamens, only one or two of
them being fertile; the remaining stamens are sterile staminodes that are sur-
mounted by a small gland. In addition, two or three smaller laments arise
from the lobes of the nectaries. The stamens are central. The ovule is anatro-
pous and pendulous. It is believed that the owers are cross-pollinated by
ies (see Davenport, Chapter 5, this volume).
Mukherjee (1951a, 1953) investigated the pollen morphology of mango
and 12 other Mangifera species. Their pollen grains were tricolpate of
almost the same size. Mondal et al. (1982, cited in Kostermans and Bom-
pard, 1993) attempted to correlate pollen morphology with taxonomic
relationships of 17 Mangifera species based upon different characteristics
of the exine and sporoderm. They demonstrated that all of the species of
section II (subgenus Limus) possess coarse exine; whereas there was no
clear correlation with pollen type in species within section I (subgenus
Mangifera).
The fruit
Description
The mango fruit is a large, eshy drupe, containing an edible mesocarp of
varying thickness. The mesocarp is resinous and highly variable with respect
to shape, size, colour, presence of bre and avour. The avour ranges from
turpentine to sweet. The exocarp is thick and glandular. There is a character-
istic beak that develops laterally on the proximal end of the fruit. A sinus is
always present above the beak. Fruit shape varies, including elongate,
oblong and ovate or intermediate forms involving two of these shapes. Fruit
length can range from 2.5 to > 30 cm, depending on the cultivar. The endo-
carp is woody, thick and brous; the bres in the mesocarp arise from the
endocarp.
The mango fruit is climacteric (see Brecht and Yahia, Chapter 14, this
volume), and increased ethylene production occurs during ripening. Chloro-
phyll, carotenes, anthocyanins and xanthophylls are all present in the fruit.
The skin is generally a mixture of green, red and yellow pigments, although
fruit colour at maturity is genotype dependent. During ripening the chloro-
plasts in the peel become chromoplasts, which contain yellow and red pig-
ments (Krishnamurthy and Subramanyam, 1970; Akamine and Goo, 1973;
Salunkhe and Desai, 1984; Mitra and Baldwin, 1997). Peel colour obviously is
cultivar dependent (see Knight et al., Chapter 3, this volume). Fruit of Bom-
bay Green is green; Carabao, Manila, Mulgoa and Arumanis are greenish-
yellow; Dashehari and Alphonso are yellow; and Haden, Keitt and
Tommy Atkins have a red blush. The red blush is due to the presence of
anthocyanins (Lizada, 1991). The pulp carotenoids in ripe fruit also vary with
respect to cultivar (Mitra and Baldwin, 1997).
S.K. Mukherjee and R.E. Litz 4
Flavour
Flavour of the mango mesocarp is a function of carbohydrates, organic acids,
lactones, monoterpene hydrocarbons and fatty acids (Mitra and Baldwin,
1997). During fruit maturation, starch that accumulates in the chloroplasts is
hydrolysed to sucrose, glucose and fructose (Medlicott et al., 1986; Selvaraj
et al., 1989; S. Kumar et al., 1994); sucrose is present in slightly higher concen-
trations than either fructose or glucose. Organic acid content decreases dur-
ing ripening (Krishnamurthy and Subramanyam, 1970). The dominant
organic acid is citric acid, but glycolic acid, malic acid, tartaric acid and oxalic
acids are also present (Sarker and Muhsi, 1981; Medlicott and Thompson,
1985). The peach-like avour of mangoes is attributed to the presence of lac-
tones (Lakshminarayana, 1980; Wilson et al., 1990).
Nutrition
Mango fruit contain amino acids, carbohydrates, fatty acids, minerals, organic
acids, proteins and vitamins. During the ripening process, the fruit are ini-
tially acidic, astringent and rich in ascorbic acid (vitamin C). Ripe mangoes
contain moderate levels of vitamin C, but are fairly rich in provitamin A and
vitamins B
1
and B
2
. Perry and Zilva (1932) determined the vitamin A, C and
D content of the fruit of three Indian mango cultivars, and found that the
pulp of mangoes is a concentrated source of vitamin C. The pulp of mango
fruit contains as much vitamin A as butter, although vitamin D is not present
in a signicant quantity. Fruit acidity is primarily due to the presence of malic
and citric acids. In addition, oxalic, malonic, succinic, pyruvic, adipic, galac-
turonic, glucuronic, tartaric, glycolic and mucic acids are also present (Jain et al.,
1959; Fang, 1965). Acidity is cultivar related; for example, immature Florida
cultivars have low acidity (0.51.0%) in comparison with Alphonso (3%).
During ripening, acidity decreases to 0.10.2%. Following fruit set, starch
accumulates in the mesocarp. Free sugars, including glucose, fructose and
sucrose, generally increase during ripening; however, the sucrose content
increases three- to fourfold due to the hydrolysis of starch. Sucrose is the
principal sugar of ripe mangoes. The sucrose content of ripe fruit of three
Indian cultivars, Alphonso, Pairie and Totapuri, ranges from 11 to 20%
representing 15 to 20% of the total soluble solids (Popenoe, 1932).
The seeds and polyembryony
Mango seeds are solitary, large and at, ovoid oblong and surrounded by the
brous endocarp at maturity. The testa and tegumen are thin and papery.
Embryos are dicotyledonous. Seeds of monoembryonic mango types contain
a single zygotic embryo, whose cotyledons can be unequal in size or lobed in
shape. The seeds of polyembryonic mango types contain one or more embryos
(Plate 2); usually one embryo is zygotic, whereas the remaining embryos are
derived directly from the nucellus, a maternal tissue. Nucellar embryos
apparently lack a suspensor. Polyembryony has also been reported in
Mangifera casturi, M. laurina and M. odorata (Bompard, 1993). Certain
Introduction: Botany and Importance 5
polyembryonic cultivars reportedly can produce seeds with adventitious
nucellar embryos only, for example Strawberry (Juliano, 1934), Carabao
and Pico (Juliano and Cuevas, 1932) and Olour and Cambodiana
(Maheshwari et al., 1955). Early studies suggested that polyembryony
appeared to be a polygenic trait (Juliano, 1934; Sturrock, 1968), segregating as
a recessive character in the progeny of controlled crosses. Recent studies,
however, have demonstrated that the polyembryony trait is inherited as a
dominant character (Aron et al., 1998). Several studies have shown that nucel-
lar seedlings can be distinguished from the single zygotic seedling of poly-
embryonic seeds by isozymes (Schnell and Knight, 1992; Degani et al., 1993)
and DNA markers, for example single sequence repeats (SSRs) (Eiadthong et al.,
1999a), amplied fragment length polymorphisms (AFLPs) (Kashkush et al.,
2001) and inter-simple-sequence-repeats (ISSRs) (Gonzalez et al., 2002). Mango
seeds are considered to be recalcitrant, and cannot survive for more than a
few days or weeks at ambient temperatures (Parisot, 1988). This important
characteristic of mango seeds would have inhibited the long distance dis-
persal of mango by seed until recent times.
1.3 History of Cultivation
Origin of Mangifera indica
The largest number of Mangifera species occurs in the Malay Peninsula, the
Indonesian archipelago, Thailand, Indochina and the Philippines (Mukher-
jee, 1985; Bompard, 1989; see Bompard, Chapter 2, this volume). The most
recent classication of Mangifera species was based upon oral morphology
(Kostermans and Bompard, 1993) and included 69 species, most of which are
included in two subgenera Mangifera and Limus with another 11 species
occupying an uncertain position (Table 1.1). Eiadthong et al. (1999b) described
the phylogenetic relationships among Mangifera species using genomic
restriction fragment length polymorphisms (RFLPs) and amplication of
chloroplast DNA (cpDNA), and suggested that the Mangifera species should
be classied using molecular data. In the next few years, it is likely that
molecular biology will have a major impact on phylogenetic studies involving
mango and its relatives.
Mangifera species with a single fertile stamen are distributed in north-
eastern India, Myanmar, Thailand and the Malay Peninsula. Many of the
mango relatives have small fruits with thin, acidic esh, large seeds, abun-
dant bre and astringent resinous substances that are localized near the skin.
In addition to M. indica, edible fruit is produced by at least 26 other species
in the genus, primarily species found in South-east Asia (Gruezo, 1992).
Mangifera caesia, known as binjai or kemang in South-east Asia, is culti-
vated in Java, where it bears fruit in the mango off-season (Bompard, 1992a).
Mangifera foetida is less commonly cultivated due to its highly astringent
fruit; however, the fruit is widely used for pickling and as a substitute for
tamarind (Bompard, 1992b). Mangifera kemang and M. altissima are consumed
S
.
K
.

M
u
k
h
e
r
j
e
e

a
n
d

R
.
E
.

L
i
t
z
6
Table 1.1. Classication of Mangifera species according to Kostermans and Bompard (1993).
Genus Subgenus Section Species
Mangifera Mangifera Marchandora Pierre M. gebede Miq
Euantherae Pierre M. caloneura Kurz
M. cochinchinensis Engler
M. pentandra Hooker f.
Rawa Kosterm. M. andamanica King M. minutifolia Evard.
M. gracilepes M. nicobarica Kosterm.
M. grifthii Hooker f. M. paludosa Kosterm.
M. merrillii Mukherji M. parvifolia Boerl. & Koorders
M. microphylla Griff. ex Hooker f.
Mangifera Ding Hou M. altissima Blanco. M. mucronulata Bl.
M. applanata Kosterm. M. oblongifolia Hooker f.
M. austro-indica Kosterm. M. orophila Kosterm.
M. austro-yunnanensis Hu M. pedicellata Kosterm.
M. casturi Kosterm. M. pseudo-indica Kosterm.
M. collina Kosterm. M. quadrida Jack
M. dewildei Kosterm. M. rigida Bl.
M. dongnaiensis Pierre M. rubropetala Kosterm.
M. ava Evard. M. rufocostata Kosterm.
M. indica L. M. similis Bl.
M. lalijiwa Kosterm. M. sulauesiana Kosterm.
M. laurina Bl. M. sumbawaensis Kosterm.
M. linearifolia (Mukherji) Kosterm. M. sylvatica Roxb.
M. longipetiolata King M. swintonioides Kosterm.
M. magnica Kochummen M. timorensis Bl.
M. minor Bl. M. torquenda Kosterm.
M. monandra Merr. M. zeylanica (Bl.) Hooker f.
I
n
t
r
o
d
u
c
t
i
o
n
:

B
o
t
a
n
y

a
n
d

I
m
p
o
r
t
a
n
c
e
7
Limus (Marchand)
Kosterm.
M. blommesteinii Kosterm. M. leschenaultii Marchand
M. caesia Jack M. macrocarpa Bl.
M. decandra Ding Hou M. odorata Griff.
M. foetida Lour. M. pajang Kosterm.
M. kemanga Bl. M. superba Hooker f.
M. lagenifera Griff.
Species of uncertain
position
M. acutigemma Kosterm. M. persiciformis Wu & Ming
M. bompardii Kosterm. M. subsessifolia Kosterm.
M. bullata Kosterm. M. taipa Buch.-Hamilton
M. campospermoides
Kosterm.
M. transversalis Kosterm.
M. hiemalis Liang Jian Ying M. utana Utana
M. maingayii Hooker f.
S.K. Mukherjee and R.E. Litz 8
as fresh fruit or used green as a salad (Angeles, 1992; Bompard, 1992a).
Mangifera pajang has the largest fruit in the genus, and is an attractive fruit.
Mangifera odorata is grown in the Philippines and Indonesia, and has occa-
sionally been used as a rootstock for mango (Ochse, 1931; Bompard, 1992c).
Mangifera odorata is widely grown in the humid lowlands of South-east Asia
in areas that are unsuitable for mango as a mango substitute. Mangifera lau-
rina and M. pentandra are appreciated as salad ingredients (Bompard, 1992d).
In addition, M. grifthii, M. minor, M. monandra, M. quadrida and M. similis
have palatable fruit that are considered to have great potential (Gruezo,
1992). All mango cultivars belong to the species M. indica.
According to De Candolle (1884), It is impossible to doubt that it (the
mango) is a native of south Asia or of the Malay Archipelago, when we see
the multitude of varieties cultivated in those countries, the number of ancient
names, in particular a Sanskrit name, its abundance in the gardens of Bengal,
of Deccan peninsula, and of Ceylon even in Rheedes time (i.e. 1683).
Although the centre of origin and diversity of the genus Mangifera is now
rmly established as being in South-east Asia, the origin of M. indica has been
a matter of speculation for many years. The fossil record provides few clues,
as only a single fossil bearing the imprint of a leaf of M. pentandra has ever
been found (Seward, 1912). Mangifera indica is believed to have rst appeared
during the Quatenary period (Mukherjee, 1951b). Blume (1850) considered
that mango might have originated from several related species, primarily
located in the Malay archipelago.
On the basis of ancient accounts of travellers and the written historical
record, it was believed for many years that mango must have originated in
India and spread outwards from there to South-east Asia and thence to the
New World and Africa. Because north-eastern India is at the northernmost
edge of the distribution of the Mangifera species, Hooker (1876) suggested
that mango might have been naturalized in India. The historical record pro-
vides a sometimes conicting account of the distribution of mango. Miquel
(1859) did not record it as being wild in the Indonesian archipelago. Accord-
ing to Rumphius (1741), the mango was introduced into certain islands of the
Indonesian archipelago within recent times; however, the mango was in cul-
tivation in Java at least as early as ad 9001100, when the temple at Borobo-
dur was built and faced with carvings of the Buddha in contemplation under
a mango tree (Plate 3). Based upon taxonomic and recent molecular evidence,
it is now apparent that the mango probably evolved within a large area
including north-western Myanmar, Bangladesh and north-eastern India (see
Bompard, Chapter 2, this volume).
Polyembryonic and monoembryonic M. indica
Within M. indica, there are two distinct types that can be distinguished on the
basis of their mode of reproduction and their respective centres of diversity:
a subtropical group with monoembryonic seed (Indian type) and a tropical
group with polyembryonic seed (South-east Asian). A few polyembryonic
cultivars occur along the west coast of India; however, they may have been
introduced into Goa from South-east Asia, perhaps by the Portuguese from
Introduction: Botany and Importance 9
their colonies of Malacca in the Malay Peninsula or Timor in the Indonesian
archipelago. Kumar et al. (2001) estimated the genetic relatedness among ten
polyembryonic and monoembryonic cultivars from the west coast of south-
ern India using genomic and chloroplast DNA RFLP analysis. The cultivars
could be grouped on the basis of embryo type (i.e. monoembryonic and poly-
embryonic) and had distinctly different genetic backgrounds. They con-
cluded that polyembryonic mangoes could not have originated in India, and
must have been introduced, probably from South-east Asia.
Domestication of mango
Historical record
It is probable that mango cultivation originated in India, where De Candolle
(1884) estimated that mango cultivation appeared to have begun at least 4000
years ago. In the early period of domestication, mango trees probably yielded
small fruit with thin esh. Such fruit can be found today in north-eastern
India and in the Andaman Islands (Anonymous, 1992). Folk selections of
superior seedlings over many hundreds of years would have resulted in
larger fruit with thicker esh. Mukherjee (1950a, b) described many of these
primitive selections from Orissa in north-eastern India; they demonstrated
great variation in fruit shape and size.
The mango is a very important cultural and religious symbol of India.
Buddhist pilgrims Fa-Hien and Sung-Yun mentioned in their travel notes
that the Gautama Buddha was presented with a mango grove by Amradarika
(c.500 bc) as a place for meditation (Popenoe, 1932). According to Burns and
Prayag (1921), a mango tree is depicted in friezes on the stupa of Bharut,
which was constructed c.100 bc. Other travellers to India, including the Chi-
nese Hwen Tsung (ad 632645), the Arabs Ibn Hankal (ad 902968) and Ibn
Batuta (ad 13251349) and the Portuguese Lurdovei de Varthema (ad 1503
1508), all described the mango. The Indian subcontinent was the birthplace
of some of the earliest highly developed civilizations, and over the centuries,
India exerted strong cultural, religious and commercial inuence over South
and South-east Asia. In successive waves, Hinduism, Buddhism and Islam
were introduced into South-east Asia from India. To this day, many com-
monly used words in Indonesia are derived from both Sanskrit and Tamil.
One of the most widely used words for mango in Malaysia and Java (Indone-
sia) is mangga, which is derived from the Tamil manga. Traders and monks
from India possibly introduced superior selections of mango into South-east
Asia; however, vegetative propagation was unknown in India until after the
arrival of the Portuguese in Goa in the 15th century. Moreover, the most im-
portant mango selections of Thailand, Cambodia, Vietnam, Malaysia, Indo-
nesia and the Philippines historically have all been of the polyembryonic
type, and have traditionally been seed propagated. Until the establishment
of Portuguese enclaves on the coast of India beginning in the late 15th century,
mango cultivars did not exist in India, as there was no known method
for vegetatively propagating superior selections (see Iyer and Schnell,
S.K. Mukherjee and R.E. Litz 10
Chapter 4, this volume). However, under the Moghul emperor Akbar (1556
1605), the best selections of seedling mangoes were propagated by approach
grafting and were planted in large orchards. The Lakh Bagh, a mango
orchard of 100,000 trees, was planted near Darbhanga in Bihar. Perhaps noth-
ing more eloquently attests to the importance of this fruit and the esteem in
which it was held than this vast mango orchard. The Ain-i-Akbari, an ency-
clopedic work that was written during the reign of Akbar, contains a lengthy
account of the mango, and includes information about the quality of the fruit
and varietal characteristics. There was evidently a strong body of informa-
tion about mango cultivation that had accumulated up to that time. Most of
the mango cultivars of India had their origin in those years, and have been
maintained under cultivation for over 400 years by vegetative propagation.
Alphonso, Dashehari, Langra, Rani Pasand, Safdar Pasand and other
mango cultivars were selected during that time. Relics of orchards from the
time of Akbar are found in different parts of India, and it has been suggested
that they could still provide valuable material for selection of superior mango
cultivars.
Distribution
Spreading from the centres of domestication
The global spread of mangoes and their cultivation outside their original
centres of domestication probably did not occur until the beginning of the
European voyages of discovery and colonialization in the 15th and 16th
centuries. Because mango seeds are recalcitrant, and cannot survive for more
than a few days or weeks, mango germplasm in the early days must have
been transported as ripe fruit, seedlings or, later on, as grafted plants. It is
believed that the Portuguese transported the mango from their colonies in
India to their African colonies, although Purseglove (1972) suggested that it
might also have been introduced to Africa via Persia and Arabia in the 10th
century by Arab traders. The Portuguese later introduced the mango into
Brazil from their African colonies of Mozambique and Angola. Spaniards,
who encountered a mango-growing civilization in the Philippines after
Magellans passage across the Pacic Ocean, introduced polyembryonic
mango types to their New World colonies through the Pacic trading ports
of Mexico and Panama. The most important, traditional mango cultivar in
Mexico remains the Manila, reecting its Philippine origin. Carabao and
Manila are probably identical. The mango was introduced to the West Indies
in the mid- to late 18th century, probably from Brazil. The rst introductions
of mango into Florida (USA) occurred in 1861, and involved the No. 11
polyembryonic seedling from Cuba. Seven years later, another polyembry-
onic selection, Peach was introduced into the state (Knight and Schnell,
1993). Many of the early introductions into Florida proved to be unproduc-
tive, although Mulgoba was planted on a small commercial scale (this culti-
var is referred to as Mulgoa in India, Mulgoba in the USA and Malgoa in
Malaysia).
Introduction: Botany and Importance 11
Secondary centres of diversity
In 1910, a seedling of Mulgoba came into production in Florida. Its fruit had
a highly attractive red blush, and appeared to bear more heavily than its
parent(s) (Wolfe, 1962). This selection was named Haden. Although Haden
was not superior with respect to fruit quality in comparison to the imported
germplasm from India, its genetic base was much wider. During the 20th
century, more introductions of mango germplasm into Florida occurred from
South-east Asia (the Philippines, Cambodia), India and elsewhere. It was at
one time believed that these introductions of mango germplasm created a
secondary centre of diversity of the species (Knight and Schnell, 1993).
Eldon, Glenn, Lippens, Osteen, Parvin, Smith, Springfels, Tommy
Atkins and Zill are progeny of Haden. Saigon seedlings were selections
made from Cambodiana, a polyembryonic introduction from Indochina.
From Saigon seedlings, Alice, Herman and Florigon were selected.
Based upon more recent genetic analysis involving microsatellite markers, it
is now estimated that the majority of Florida cultivars are descended from
only four monoembryonic Indian mango cultivar accessions, i.e., Mulgoba,
Sandersha, Amini and Bombay, together with the polyembryonic Tur-
pentine from the West Indies (Schnell et al., 2006). The Florida mango culti-
vars have been found to be highly adaptable to many agroecological areas
and bear regularly, whereas many of the outstanding Indian cultivars have
been unproductive outside their centre of domestication, and are alternate
bearing. These selections also have a highly attractive red blush at maturity,
rm esh, a high esh to seed ratio and a regular bearing habit. Some of the
Florida cultivars, for example Tommy Atkins, Keitt, etc. are also moder-
ately resistant to anthracnose, the most important production and posthar-
vest problem of mango in many areas. In the latter half of the 20th century,
plantings of Florida cultivars have been established in many countries and
now form the basis of international trade of mangoes.
Current distribution
The mango is cultivated commercially throughout the tropics and in many
subtropical areas. It is grown at the equator and at a latitude of 3537 in
southern Spain. According to Knight and Schnell (1993), The process that
began in Florida introduction of superior germplasm from abroad followed
by selection of improved cultivars adapted to local conditions is now
underway in many areas.
1.4 Germplasm Conservation
Genetic erosion
The Mangifera species have their centre of diversity and origin in South-east
Asia, a region that has experienced great economic development in recent
years. Vast wooded areas have been completely or partially deforested either
for expanding agriculture or for removal of tropical hardwoods for export.
S.K. Mukherjee and R.E. Litz 12
This has caused great genetic erosion within many species and genera. The
Mangifera species, like many other tropical fruit trees, are canopy and emer-
gent trees of the tropical rainforest (Kaur et al., 1980). These trees are widely
scattered in the tropical rainforest, ower erratically and reproduce from
large seeds that deteriorate rapidly. As such, they are particularly vulnerable
and in danger of extinction.
Collection and documentation of Mangifera germplasm
The International Plant Genetic Resources Institute (IPGRI), formerly known
as the International Board for Plant Genetic Resources (IBPGR), commis-
sioned an ecogeographical study of known Mangifera genetic resources (Muk-
herjee, 1985). Based upon this documentation, a joint IBPGR-International
Union for the Conservation of Nature (IUCN)-World Wildlife Fund (WWF)
project was initiated to collect wild mangoes on the island of Borneo and in
the Malay Peninsula (Bompard, 1989), the regions that held the highest con-
centrations of Mangifera species. Kostermans and Bompard (1993), in the lat-
est revision of the taxonomy of Mangifera, recognized 69 species, many of
which were collected during the course of this project (Table 1.1). Because of
the loss of natural habitat, the establishment of in situ and ex situ germplasm
collections of Mangifera species was considered to be imperative.
Relevance of germplasm resources to mango improvement
The genetic improvement of mango hitherto has depended on the utilization
of the genetic variability found within a single species, M. indica. According
to Mukherjee (1985), A concerted sampling strategy should be devised for ex
situ samples to meet urgent needs for use in research for improvement of the
crop through breeding or as rootstocks. Sources of resistance to mango mal-
formation, anthracnose, powdery mildew, gall midge are urgently needed.
1.5 Importance of Mango
Cultivars
A partial list of the principal mango cultivars has been provided in Table 1.2.
This list includes many cultivars that were identied in a survey of world
mango production compiled by Watson and Winston (1984). The distribution
of mango cultivars outside their centres of domestication can be attributed
primarily to three historical events: (i) the movement of Indian varieties
(monoembryonic) along the trade routes of the Portuguese to Africa and
South America; (ii) the spread of South-east Asian varieties (polyembryonic)
across the Pacic Ocean to Central and South America by the Spaniards; and
(iii) the identication of improved mango cultivars initially in Florida and
Introduction: Botany and Importance 13
Table 1.2. Most important mango cultivars in major producing countries.
Continent Country Cultivars
Africa Cote dIvoire Amelie, Kent
Egypt Alphonso, Bullocks Heart, Hindi be Sennara,
Langra, Mabrouka, Pairie, Taimour, Zebda
Kenya Boubo, Ngowe, Batawi
Mali Amelie, Kent
South Africa Fascell, Haden, Keitt, Kent, Sensation, Tommy
Atkins, Zill
Asia Bangladesh Aswina, Fazli, Gopal Bhog, Himsagar, Khirsapati,
Langra
China Gui Fei, Tainong No. 1, Keitt, Sensation, Zill, Zihua,
Jin Huang
India Alphonso, Banganapalli, Bombay, Bombay Green,
Chausa, Dashehari, Fazli, Fernandian,
Himsagar, Kesar, Kishen Bhog, Langra, Mallika,
Mankurad, Mulgoa, Neelum, Pairi, Samar
Behisht, Suvarnarekha, Totapuri, Vanraj, Zardalu
Indonesia Arumanis, Dodol, Gedong, Golek,
'
Madu,
'
Manalagi
Israel Haden, Tommy Atkins, Keitt, Maya, Nimrod, Kent,
Palmer
Malaysia Apple Rumani, Arumanis, Golek, Kuala Selangor 2,
Malgoa
Myanmar Aug Din, Ma Chit Su, Sein Ta Lone,
'
Shwe Hin Tha
Pakistan Anwar Ratol, Began Pali, Chausa, Dashehari,
Gulab Khas, Langra, Siroli, Sindhri,
Suvarnarekha, Zafran
The Philippines Carabao, Manila Super, Pico
Taiwan Irwin, Jin-hwung, Keitt, Tommy Atkins, Tainong
No. 1, Tsar-swain
Thailand Nam Doc Mai, Ngar Charn, Ok Rong, Keow Savoey,
Pimsen Mum
Australia Calypso, Kensington Pride
North and
Central
America
Costa Rica Haden, Irwin, Keitt, Mora, Tommy Atkins
Dominican
Republic
Haden, Keitt, Kent, Tommy Atkins
Guatemala Haden, Keitt, Kent, Tommy Atkins
Haiti Francine, Madame Francis
Mexico Ataulfo, Haden, Keitt, Kent, Manila, Palmer,
Sensation, Tommy Atkins, Van Dyke
USA Keitt, Kent, Tommy Atkins
South
America
Brazil Bourbon, Coite, Coquinho, Coracao, Espada,
Haden, Itamaraca, Keitt, Mamao, Palmer, Rosa,
Tommy Atkins, Uba, Van Dyke
Colombia Vallenato
Ecuador Haden, Keitt, Kent, Tommy Atkins
Peru Haden, Keitt, Kent, Tommy Atkins
Venezuela Haden, Keitt, Kent, Tommy Atkins
S.K. Mukherjee and R.E. Litz 14
later in other new mango-producing areas, as a result of open and controlled
pollination among local and introduced mango germplasm from India and
South-east Asia.
Further information about many of the mango cultivars, including their
fruit characters, is available in Knight et al. (Chapter 3, this volume), and in
publications by Burns and Prayag (1921) for mangoes of Maharashtra, Naik
and Gangolly (1950) for south Indian mangoes, Singh and Singh (1956) for
Uttar Pradesh mangoes, Mukherjee (1948) for Bengal mangoes and Camp-
bell (1992) for Florida mangoes.
Because many clonally propagated mango cultivars have unique local
and/or regional names, there is considerable confusion in nomenclature. The
Indian Agricultural Research Institute (IARI), New Delhi, has been recog-
nized by the International Society for Horticultural Science (ISHS) as the
International Registration Authority for Mango, whose mission is to consoli-
date superuous names of mango cultivars. The potential for molecular, for
example randomly amplied polymorphic DNA (RAPD), markers, to resolve
much of this confusion has been demonstrated by Schnell and Knight (1992),
Degani et al. (1993), Schnell et al. (1995), Eiadthong et al. (1999a), Kashkush et al.
(2001) and Gonzalez et al. (2002) (see Bompard, Chapter 2 and Iyer and
Schnell, Chapter 4, this volume).
There is little variation among seedlings derived from polyembryonic
mangoes. None the less, a certain amount of variability does occur, probably
as a result of somatic mutation. Thus, in Indonesia there are several Aru-
manis selections that are denoted numerically, for example Arumanis 1,
Arumanis 2, etc. In addition, although Philippine mango cultivars are dis-
tinguished by different names, for example Carabao, Manila, Philippine,
etc., the differences among them are quite subtle.
1.6 Production and Uses
The mango is the most important fruit of Asia, and currently ranks fth in
total production (in metric tonnes) among major fruit crops worldwide, after
Musa (bananas and plantains) (105,815,354 t), Citrus (all types) (105,440,168
t), grapes (65,584,233 t) and apples (59,444,377 t) (FAOSTAT, 2006). According
to the Food and Agriculture Organization of the United Nations (FAO)
database (FAOSTAT, 2006), world mango production has increased from
16,903,407 t in 1990 to 28,221,510 t in 2005. Much of this new production has
occurred outside the traditional centres of mango culture of South and
South-east Asia. In 1990, India produced approximately 51% of the worlds
mangoes, but by 2005, Indias share had declined to approximately 38%,
despite the substantial increase in mango production since 1990 (from 8,645,405
to 10,800,000 t between 1990 and 2005). The current leading producing nations
after India include (in metric tonnes) China (3,450,000), Thailand (1,800,000),
Pakistan (1,673,900), Mexico (1,600,000), Indonesia (1,478,204), Brazil (1,000,000)
and the Philippines (950,000). Although world production has increased by
67% between 1990 and 2005, mango exports have increased almost sixfold
Introduction: Botany and Importance 15
from 158,030 to 907,782 t, with total export value estimated to be
US$583,763,000 (FAOSTAT, 2006). The major exporting countries are (in met-
ric tonnes) Mexico (212,505), India (156,222) and Brazil (111,181). As a result,
mangoes are widely available as fresh fruit and as processed products (i.e.
dried fruit, dairy products, juice, pickles, etc.).
Mangoes are an important component of the diet in many less developed
countries in the subtropics and tropics. In regions of the world that have
experienced low living standards and serious nutritional deciencies, their
attractiveness and avour have also enhanced the quality of life. Surplus
production has increasingly been processed and fruit of certain cultivars is
destined for export as fresh fruit. Approximately 1% of mango production is
utilized for processing for juice, nectars, preserves (including chutney), fruit
leather, dried fruit slices, frozen pulp and as a avouring for baked goods,
ice cream, yoghurt, etc. (see Raymundo et al., Chapter 17, this volume). No
part of the fruit is wasted. In India and the subcontinent, the seed is used for
extraction of starch amchur, and the peels (skin) have been used as a source
of anacardic acid. Mango wood is a low quality timber, and the bark of the
tree is an important source of tannins for curing leather.
References
Akamine, E.K. and Goo, T. (1973) Respiration and ethylene production during ontogeny
of fruit. Journal of the American Society for Horticultural Science 98, 381383.
Angeles, D.E. (1992) Mangifera altissima Blanco. In: Verheij, E.W.M. and Coronel, R.E.
(eds) Plant Resources of South-east Asia No.2: Edible Fruits and Nuts. Pudoc-DLO,
Wageningen, the Netherlands, pp. 206207.
Anonymous (1992) Annual Report. Central Institute of Horticulture for Northern Plains,
Lucknow, India.
Aron, Y., Czosnek, H., Gazit, S. and Degani, C. (1998) Polyembryony in mango
(Mangifera indica L.) is controlled by a single dominant gene. HortScience 33,
12411242.
Blume, C.L. (1850) Museum Botanicum Lugduno Batavum 1, 190193.
Bompard, J.M. (1989) Wild Mangifera species in Kalimantan (Indonesia) and in Malay-
sia. Final Report. International Board for Plant Genetic Resources, Rome.
Bompard, J.M. (1992a) Mangifera caesia Jack. and Mangifera kemanga Blume. In: Ver-
heij, E.W.M. and Coronel, R.E. (eds) Plant Resources of South-east Asia No.2: Edible
Fruits and Nuts. Pudoc-DLO, Wageningen, the Netherlands, pp. 207209.
Bompard, J.M. (1992b) Mangifera foetida Lour. and Mangifera pajang Kostermans. In:
Verheij, E.W.M. and Coronel, R.E. (eds) Plant Resources of South-east Asia No.2:
Edible Fruits and Nuts. Pudoc-DLO, Wageningen, the Netherlands, pp. 209211.
Bompard, J.M. (1992c) Mangifera laurina Blume and Mangifera pentandra Hooker f. In:
Verheij, E.W.M. and Coronel, R.E. (eds) Plant Resources of South-east Asia No.2:
Edible Fruits and Nuts. Pudoc-DLO, Wageningen, the Netherlands, pp. 216218.
Bompard, J.M. (1992d) Mangifera odorata Grifth. In: Verheij, E.W.M. and Coronel, R.E.
(eds) Plant Resources of South-east Asia No.2: Edible Fruits and Nuts. Pudoc-DLO,
Wageningen, the Netherlands, pp. 218221.
Bompard, J.M. (1993) The genus Mangifera re-discovered: the potential of wild species
to mango cultivation. Acta Horticulturae 341, 6977.
S.K. Mukherjee and R.E. Litz 16
Burns, W. and Prayag, S.M. (1921) The Book of Mango. Bulletin 103, Department of
Agriculture, Bombay, India.
Campbell, R.J. (1992) A Guide to Mangoes in Florida. Fairchild Tropical Garden, Miami,
Florida.
De Candolle, A.P. (1884) Origin of Cultivated Plants. Hafner, London.
Degani, C., Cohen, M., Reuveni, O., El-Batsri, R. and Gazit, S. (1993) Frequency and
characteristics of zygotic seedlings from polyembryonic mango cultivars, deter-
mined using isozymes as genetic markers. Acta Horticulturae 341, 7885.
Eiadthong, W., Yonemoni, K., Kanzaki, S., Sugiura, A., Utsunomiya, N. and Subhadrab-
andhu, S. (1999a) Amplied fragment length polymorphism (AFLP) analysis for
studying the genetic relationships among Mangifera species in Thailand. Journal of
the American Society for Horticultural Science 125, 160164.
Eiadthong, W., Yonemoni, K., Sugiura, A., Utsunomiya, N. and Subhadrabandhu, S.
(1999b) Analysis of phylogenetic relationship in Mangifera by restriction site analy-
sis of an amplied region of cpDNA. Scientia Horticulturae 80, 145155.
Fang, T.T. (1965) Chromatographic fractionation of nonnitrogenous organic acids of
mango and guava fruits by silica gel column. Memoirs of the College of Agriculture,
National Taiwan Museum 8, 236.
FAOSTAT (2006) Available at: http://faostat.fao.org/site/340/default.aspx (accessed 22
October 2006).
Gonzalez, A., Coulson, M. and Brettell, R. (2002) Development of DNA markers (ISSRs)
in mango. Acta Horticulturae 575, 139143.
Gruezo, W.S. (1992) Mangifera L. In: Verheij, E.W.M. and Coronel, R.E. (eds) Plant Re-
sources of South-east Asia No.2: Edible Fruits and Nuts. Pudoc-DLO, Wageningen,
the Netherlands, pp. 203206.
Hooker, J.D. (1876) The Flora of British India 2. Reeve, London.
Jain, N.L., Krishnamurthy, G.V. and Lal, G. (1959) Nonvolatile organic acids in unripe
pickling mangoes and salted mango slices by paper chromatography. Food Science
8, 115117.
Juliano, J.B. (1934) Origin of embryos in the strawberry mango. Philippine Journal of
Science 54, 553563.
Juliano, J.B. and Cuevas, N.L. (1932) Floral morphology of the mango (Mangifera indica
L.) with special reference to the Pico variety from the Philippines. Philippine Agri-
culturist 21, 449472.
Kashkush, K., Jinggui, F., Tomer, E., Hillel, J. and Lavi, U. (2001) Cultivar identication
and genetic map of mango (Mangifera indica). Euphytica 122, 129136.
Kaur, A., Ha, C.O., Jong, K., Sands, V.E., Chan, H.T., Soepadmo, E. and Ashton, P.S.
(1980) Apomixis may be widespread among trees of the climax rain forest. Nature
271, 440442.
Knight, R.J., Jr and Schnell, R.A. (1993) Mango (Mangifera indica L.) introduction and
evaluation in Florida and its impact on the world industry. Acta Horticulturae 341,
125135.
Kostermans, A.J.G.H. and Bompard, J.M. (1993) The Mangoes: Botany, Nomenclature,
Horticulture, Cultivation and Utilization. Academic Press, London.
Krishnamurthy, S. and Subramanyam, H. (1970) Respiratory climacteric and chemical
changes in the mango fruit Mangifera indica L. Journal of the American Society for
Horticultural Science 95, 333337.
Kumar, N.V.H., Narayanaswamy, P., Prasod, D.T., Mukunda, G.K. and Sondhu, S.N.
(2001) Estimation of genetic diversity of commercial mango (Mangifera indica L.)
cultivars using RAPD markers. Journal of Horticultural Science and Biotechnology
76, 529533.
Introduction: Botany and Importance 17
Kumar, S., Das, D.K., Singh, A.K. and Prasad, U.S. (1994) Sucrose metabolism during
maturation and ripening of mango cultivars. Plant Physiology and Biochemistry 21,
2732.
Lakshminarayana, S. (1980) Mango. In: Nagy, S. and Shaw, P.E. (eds) Tropical and
Subtropical Fruits. AVI Publishing Co., Westport, Connecticut, USA, pp. 184257.
Lizada, M.C. (1991) Post harvest physiology of mango a review. Acta Horticulturae
291, 437449.
Maheshwari, P., Sachar, R.C. and Chopra, R.N. (1955) Embryological studies in mango
(Mangifera indica L.). In: Proceedings of the 42nd Indian Scientic Congress, Baro-
da, India, p. 233.
Medlicott, A.P. and Thompson, A.K. (1985) Analysis of sugars and organic acids
in ripening mango fruit (Mangifera indica L. var. Keitt) by high performance
liquid chromatography. Journal of the Science of Food and Agriculture 36,
56566.
Medlicott, A.P., Bhogol, M. and Reynolds, S.B. (1986) Changes in peel pigmentation
during ripening of mango fruit (Mangifera indica var. Tommy Atkins). Annals of
Applied Biology 109, 651656.
Miquel, F.A.G. (1859) Flora van Nederlandsch Indie 1, 627634.
Mitra, S.K. and Baldwin, E.A. (1997) Mango. In: Mitra, S.K. (ed.) Postharvest Physiology
and Storage of Tropical and Subtropical Fruits. CAB International, Wallingford, UK,
pp. 85122.
Mukherjee, S.K. (1948) The varieties of mango (M. indica L.) and their classication.
Bulletin of the Botanical Society of Bengal 2, 101133.
Mukherjee, S.K. (1950a) Wild mangoes of India. Science and Culture 15, 469471.
Mukherjee, S.K. (1950b) Mango. Its allopolyploid nature. Nature 150, 196197.
Mukherjee, S.K. (1951a) Pollen analysis in Mangifera in relation to fruit set and taxonomy.
Journal of the Indian Botanical Society 30, 4955.
Mukherjee, S.K. (1951b) Origin of mango. Indian Journal of Genetics and Plant Breeding
11, 4956.
Mukherjee, S.K. (1953) Origin, distribution and phylogenetic afnities of the species of
Mangifera L. Journal of the Linnean Society, Botany 55, 6583.
Naik, K.C. and Gangolly, S.R. (1950) A Monograph on Classication and Nomenclature
of South Indian Mangoes. Government Press, Madras, India.
Ochse, J.J. (1931) Fruits and Fruiticulture in the Dutch East Indies. G. Kolff, Batavia,
(Jakarta), Indonesia.
Parisot, E. (1988) Etude de la croissance rhythmique chez de jeunes manguiers (Mangifera
indica L.). Description, germination et conservation de graines polyembryonnees
de manguier. Fruits 43, 97105.
Perry, E.O.V. and Zilva, S.S. (1932) Preliminary Report on Vitamin Content of the Mango.
Empire Marketing Board, London.
Popenoe, W. (1932) Manual of Tropical and Subtropical Fruits. Macmillan Co., New
York.
Purseglove, J.W. (1972) Mangoes west of India. Acta Horticulturae 24, 107174.
Rumphius, G.E. (17411750) Herbarium Amboinense. Vol. 16. Den Haag, Amsterdam.
Salunkhe, D.K. and Desai, B.B. (1984) Mango. In: Postharvest Biotechnology of Fruits,
Vol. 1. CRC Press, Boca Raton, Florida, pp. 7794.
Sarker, S. and Muhsi, A.A. (1981) A study on the content and interconversions of or-
ganic acids of mango (Mangifera indica L.) at various stages of fruit development.
Bangladesh Journal of Agricultural Science 8, 6975.
Schnell, R.J. and Knight, R.J., Jr (1992) Frequency of zygotic seedlings from ve polyem-
bryonic mango rootstocks. HortScience 27, 174176.
S.K. Mukherjee and R.E. Litz 18
Schnell, R.J., Ronning, C.M. and Knight, R.J., Jr (1995) Identication of cultivars and
validation of genetic relationships in Mangifera indica L. using RAPD markers.
Theoretical and Applied Genetics 90, 269274.
Schnell, R.J., Brown, J.S., Olano, C.T., Meerow, A.W., Campbell, R.J. and Kuhn, D.N.
(2006) Mango genetic diversity analysis and pedigree inferences for Florida culti-
vars using microsatellite markers. Journal of the American Society for Horticultural
Science 13, 214224.
Selvaraj, Y., Kumar, R. and Pal, D.K. (1989) Changes in sugars, organics, amino acids,
lipids, lipid constituents and aroma characteristics or ripening mango (Mangifera
indica L.) fruit. Journal of Food Science and Technology 26, 306311.
Seward, A.C. (1912) Dictyledonous leaves from Assam. Records of the Geological Sur-
vey of India 42, 100.
Singh, L.B. and Singh, R.N. (1956) A Monograph on the Mangoes of UP. Superintendent
of Printing, Uttar Pradesh Government, Lucknow, India.
Sturrock, T.T. (1968) Genetics of mango polyembryony. Proceedings of the Florida State
Horticultural Society 81, 311314.
Watson, B.J. and Winston, E.C. (1984) Plant genetic improvement. In: Proceedings of the
First Australian Mango Research Workshop. Commonwealth Scientic and Indus-
trial Research Organization (CSIRO), Canberra, pp. 104138.
Wilson, C.W., Shaw, P.E. and Knight, R.J., Jr (1990) Importance of some lactones and
2,5-dimethyl-4-hydroxy-3-(2H)-furanone to mango (Mangifera indica L.) aroma.
Journal of Agricultural Food Chemistry 38, 15561559.
Wolfe, H.S. (1962) The mango in Florida 1887 to 1962. Proceedings of the Florida
State Horticultural Society 75, 387391.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 19
2 Taxonomy and Systematics
J.M. Bompard
Les Mazes, Montaud, France
2.1 Introduction 19
2.2 The Genus Mangifera L. 20
Distribution 20
Ecology and habitat 20
2.3 Taxonomy and Systematics 22
Taxonomic history 22
2.4 Phytogeography 28
Species distribution 28
Subgenera and section distribution 29
2.5 Interspecic Molecular Characterization 30
2.6 Region of Origin of the Genus 31
2.7 Origin of the Common Mango 32
The common mango in South-east Asia 32
2.8 Conclusion 35
Potential contribution of wild species to mango cultivation 35
Source of rootstock 35
Hybridization 36
Potential of wild species 36
2.1 Introduction
The genus Mangifera is one of the 73 genera (c.850 species) belonging to the
family of Anacardiaceae, in the order of Sapindales. Anacardiaceae is a fam-
ily of mainly tropical species, with a few representatives in temperate regions.
Malesia, which is the phytogeographic region extending from the Malay
Peninsula south of the Kangar-Pattani line to the Bismarck Archipelago east
of New Guinea (Whitmore, 1975) contains more species in the Anacardiaceae
than any other area. Within Malesia occurrence is mainly in Western Malesia
(Ding Hou, 1978b).
J.M. Bompard 20
Apart from mango, several other cultivated fruit trees belong to the fam-
ily, for example the ambarella or Otaheite apple (Spondias dulcis Forst.) prob-
ably from Melanesia, and the yellow and purple mombins (Spondias mombin
L. and S. purpurea L., respectively) from tropical America, the Bouea species
from IndoMalesia, dragon plums (Dracontomelum spp.) from IndoMalesia
and the Pacic region, kafr plum (Harpephyllum caffrum Bernh. ex K. Krause)
and the marula plum (Sclerocarya caffra Sond.) of southern Africa. The cashew
(Anacardium occidentale L.) is from tropical America and the pistachio (Pistacia
vera L.) from Iran and Central Asia. Anacardiaceous species also yield other
valuable products: wood (several genera), gums and resins (Pistacia spp.),
varnishes (Rhus spp. and Melanorrhoea spp., lacquer trees) and tanning
materials (Rhus spp. and Schinopsis spp.). It is also a family well known for
the dermal irritation produced by some of its members, such as the poison
ivies and oaks (Rhus spp.) in North America, rengas (Gluta spp.) in South-
east Asia and other species including some Mangifera species whose resinous
sap may induce a mild to strong allergic reaction.
2.2 The Genus Mangifera L.
Distribution
The range of natural distribution of the 69 Mangifera species is mainly
restricted to tropical Asia, and extends as far north as 27 latitude and as far
east as the Carolines Islands. Wild mangoes occur in India, Sri Lanka, Ban-
gladesh, Myanmar, Sikkim, Thailand, Kampuchea, Vietnam, Laos, southern
China, Malaysia, Singapore, Indonesia, Brunei, the Philippines, Papua New
Guinea and the Solomon and Carolines Islands. The highest species diver-
sity, c.29 species, occurs in western Malesia, especially in peninsular Malay-
sia and in Borneo and Sumatra, which represent the heart of the distribution
range of the genus (Fig. 2.1).
Ecology and habitat
The majority of Mangifera species occur as a rule as scattered individuals in
tropical lowland rainforests on well-drained soils. The species are distrib-
uted mostly below 300 m, but can occur up to c.1000 m above sea level, on
well-drained soils (44 species), in periodically inundated areas (ten species)
and in certain types of swamp forest (i.e. M. gedebe, M. grifthii and M. parvi-
folia). Three species are mainly found in sub-montane forests above 1000 m
and occasionally up to 1700 m above sea level (M. bompardii, M. dongnaiensis
and M. orophila). There are also species that are adapted to seasonally dry
climates in deciduous or semi-deciduous forests (e.g. M. caloneura, M. collina,
M. timorensis and M. zeylanica). A few species occur north of the Tropic of
Cancer, for example M. austro-yunnanensis and M. persiciformis in China, M.
Taxonomy and Systematics 21
sylvatica Roxb. in Sikkim and southern China, at altitudes of 6001900 m above
sea level; apparently wild M. indica can also be found outside the tropics.
Wild mangoes are large trees, 3040 m (occasionally 54 m) in height, with
tall columnar boles. Several species are exploited for their timber. The major-
ity of wild mangoes occur as scattered individuals at very low densities in
lowland forests on well-drained soils. Some of these are very rare; there are
normally one to three trees above 40 cm in diameter/10 ha. Only a few spe-
cies (M. gedebe, M. grifthii and M. parvifolia) are gregarious in certain types of
swamp forest. Most species are evergreen although a few are deciduous in
the rainforests following a dry period, and stand bare for a short time before
ushes of new leaves appear. A deciduous habit that is not linked to a sea-
sonal climate also occurs in other genera of Anacardiaceae (Ding Hou, 1978b).
In the rainforest of western Malesia, Mangifera species ower and fruit
very irregularly. As with many other genera in the region, mast or general
fruiting at intervals of 38 years is the dominant pattern. In mast years, the
ground beneath the trees can be covered with mangoes, whose strong smell
attracts many animals. Isolated owering may occur after a dry period and
is generally followed by a poor fruit crop. The occurrence of owering of a
few species, for example the lanjut (M. lagenifera) is only once every 510
years. There seem to be clear reproductive barriers between species in the
wild, although limited hybridization among cultivated species has been
reported (see section 2.8, Conclusion, this chapter).
30 N
4
2
5
4
3
6
13
27
27
28
9
5
7
4
3
5
2
20 N
10 N
0 N
10 N
30 N
20 N
10 N
0 N
10 N
80 100 120 160
80 100 120 160 140
Fig. 2.1. Distribution of Mangifera species in the range of the genus. Numbers shown indicate
the number of wild species in each area: Sri Lanka, India and Sikkim, Andaman and Nicobar
Islands, Myanmar, Thailand, Indochina, China, peninsular Malaysia, Sumatra, Borneo, Java,
Lesser Sunda Islands, Sulawesi, Moluccas, the Philippines, New Guinea, and Solomon Islands
(the Caroline Islands not represented).
J.M. Bompard 22
These widely scattered towering tree species, often with an inaccessible
crown, are undercollected and poorly represented in herbarium collections
(Bompard, 1995). Because of their irregular owering, the owers and fruits
of a few species are still unknown. Collecting plant material is consequently
very difcult, and plant explorations are still yielding new records or new
species. Many species have been recently recorded for the rst time, even
from peninsular Malaysia, a country that has already been rather well combed
by botanists, having one of the highest collecting indices in the Malesian
region. Other species still await to be discovered. Sadly some species of very
limited range may already have been lost to posterity by deforestation.
Our very meagre knowledge of the wild mangoes is due to the fact that
identication at the species level from leaves only is often difcult because of
intraspecic variation in vegetative characters. Moreover, many of the origi-
nal species were based on very poor specimens. Consequently, frequent mis-
identication of herbarium material has resulted in much confusion, requiring
a critical revision of all the specimens in these collections. It is not uncommon
that the same species has been described from different places under differ-
ent names. For instance, M. inocarpoides described by Merrill and Perry from
New Guinea in 1941, M. camptosperma and M. reba (recorded by Pierre in
South Vietnam in 1897) are now recognized to be a single species M. gedebe
Miquel, a species initially named in 1861 from a specimen collected in Suma-
tra. Mangifera longipes Grifth is now treated as M. laurina Blume, because
this name takes precedence as it was validly published 4 years earlier.
After thorough study of herbarium collections and eld collections, a
number of species have been newly described. Sixty-nine species are now
recorded, including 13 species of uncertain afnities, in contrast with the
49 species recognized by Mukherjee (1949). As more collections are made,
there will doubtless be further taxonomic adjustments made to the genus
Mangifera.
2.3 Taxonomy and Systematics
Taxonomic history
Subdivision of the genus
An historical review of the subdivisions of the genus Mangifera shows that
two major groups have been rather consistently recognized in taxonomic
treatments. Hooker (1862) was the rst to recognize two sections based on
the characters of the ower disc: section I with a disc broader than the ovary,
and section II with a disc stalk-like or wanting. These sections were later
named by Marchand (1869) Amba, an Indian name for the common mango,
and Limus, a Sundanese name for M. foetida in West Java, respectively. He
also added a section Manga for M. leschenaultii, which in fact belongs to the
section Limus.
In his monograph of the Anacardiaceae, Engler (1883) maintained Hook-
ers sections, and subdivided group A (Hookers section I) into two groups,
Taxonomy and Systematics 23
one group with four or ve petals and the other group with four petals. He
considered the following sequence of morphological characters to be impor-
tant for classication: (i) texture of the leaves; (ii) number of fertile stamens;
(iii) prominence of veins; (iv) pilosity of inorescences; and (v) leaf shape.
Pierre (1897) further divided the genus Mangifera into ve sections based
on ower characters, i.e. number of stamens, the attachment of stamens to
the disk, and the style. Two of these ve sections namely section I Euan-
therae, with a short thick ower disc and 412 fertile stamens, and section V
Marchandora then consisting of M. camptosperma (currently considered a syn-
onym of M. gedebe) are still maintained as they form clear-cut sections.
In his monograph, Mukherjee (1949) recognized two unnamed sections,
conserving Hookers subdivision. Ding Hou (1978a) adopted the same
method in his revision of the Malesian Anacardiaceae recognizing only
Hookers two original sections and providing them with proper names and
synonyms: section Mangifera (section I Hooker, section Amba Marchand,
group A Engler, sections Euantherae and Marchandora Pierre) and section
Limus (section II Hooker, sections Limus and Manga Marchand, group B
Engler, and sections Eudiscus and Microdiscus Pierre).
Most recent classication of the genus
The taxonomic classication referred to herein follows that proposed by Kos-
termans and Bompard (1993). This treatise includes the results of collections
and surveys carried out between 1986 to 1998 in Borneo and peninsular
Malaysia, which were initiated and sponsored by the International Institute
for Genetic Resources (now Biodiversity International) and the World Wide
Fund for Nature.
1
It was published under the auspices of the International
Board for Plant Genetic Resources (now International Plant Genetics Resources
Institute) and the Linnean Society of London.
The most recent treatment of Mangifera reects the current status of what
is still fragmentary knowledge. It can provide a basis for further studies
involving all aspects of the wild relatives of mango, but particularly their
potential in mango breeding. Determining phylogenetic afnities based upon
molecular markers could change our thinking about relationships among
Mangifera species and among the cultivated forms of M. indica (see Interspe-
cic Molecular Characterization section, this chapter).
The morphological characters used for identication have been placed in
the following sequence of importance:
Shape of the oral disc (see section Subdivision of the genus). 1.
Number of fertile stamens. 2.
Seed labyrinthine or not. 3.
Shape of secondary branches of the inorescences: open or lax panicle, 4.
owers glomerulate or sub-glomerulate, the ramications racemoid or
spike like.
Pubescence of the inorescence. 5.
Shape, number and attachment of the nerves (ridges or ngers) at the 6.
inner surface of the petals.
J.M. Bompard 24
Shape and size of the petals. 7.
Flowers tetra- or pentamerous (not a very constant character and often 8.
overlapping).
Reticulation of the leaves, especially of the lower surface. 9.
Shape of the leaf (only fully grown leaves of sterile branches can be used). 10.
Texture of the leaves. 11.
Deciduous or non-deciduous trees. 12.
Colour of the owers. 13.
Shape, colour and smoothness of the fruit. 14.
Number and size of the stone bres. 15.
Kostermans (Kostermans and Bompard, 1993) raised the sections to the
rank of subgenus, i.e. subgenus Limus (Marchand) Kosterm., having a disc
narrower than the base of the ovary, stalk-like or even lacking and subgenus
Mangifera (Ding Hou) Kosterm., having a disc broader than the base of the
ovary, cushion-like, often divided in four or ve lobes.
SUBGENUS LIMUS (MARCHAND) KOSTERM. Mangifera species of the subgenus Limus
are quite distinctive and show only remote afnity with the common mango.
This taxon is more primitive than the subgenus Mangifera and may be ances-
tral to it, although the two subgenera may have originated from two different
ancestors. The subgenus Limus consists of 11 species, which are native to the
rainforests of western Malesia (peninsular Thailand, Malay Peninsula, Suma-
tra, West Java and Borneo), with the exception of M. foetida, which extends to
the east, possibly as far as New Guinea, and M. odorata which is only known
in cultivation.
Kostermans divided the subgenus Limus into two sections: (i) section
Deciduae for deciduous trees (i.e. M. caesia, M. kemanga, M. pajang, M. superba
and possibly M. blommesteinii, M. decandra and M. lagenifera); and (ii) section
Perennes for non-deciduous species (i.e. M. foetida, M. leschenaultii, M. macro-
carpa and M. odorata) (Kostermans and Bompard, 1993). In deciduous trees,
the bracts enclosing the buds leave a characteristic collar of dense, narrow
scars, which persist on old twigs and are especially prominent in M. caesia
and M. kemanga.
Mangifera lagenifera and M. decandra have ten stamens, ve of which are
fertile. The other nine species have only one (and rarely two) fertile stamen(s)
and two to four staminodes. The two species with ve fertile stamens (M.
decandra and M. lagenifera) and M. superba, M. caesia, M. kemanga and M. blom-
mesteinii, whose leaves are apically aggregated into rosettes at the end of mas-
sive twigs are particularly distinctive. The fruits of these species are broadly
ellipsoid or pear shaped, not compressed, and have dirty whitish or pinkish
mesocarp and lanceolate, and brous, non-ligneous leathery endocarp.
Mangifera subsessilifolia shows some afnity with M. lagenifera and M.
blommesteinii; however, it has been placed among the species of uncertain
taxonomic position due to a lack of complete study material. This is not a
very rare species, but owering and fruiting seem to occur at intervals of, or
> 5 years, similar in this respect to M. lagenifera, which can be found growing
Taxonomy and Systematics 25
in old orchards in peninsular Malaysia. The owers and fruits of M. sub-
sessilifolia are still unknown.
Mangifera foetida, M. odorata, M. caesia and M. kemanga are widely cultivated
in the humid lowlands of the Malay Peninsula, Sumatra, Borneo, Java and Bali.
They have also been introduced elsewhere in South-east Asia; M. caesia, M. foe-
tida and M. odorata are grown in the southern part of the Philippines, M. foetida
is grown in Myanmar, and M. odorata is found in Indochina. They have been
described in general reviews of tropical fruit (Ochse and Bakhuizen, 1931; Ochse
et al., 1961; Molesworth, 1967; Verheij and Coronel, 1991).
Mangifera pajang, an endemic and commonly cultivated species in Bor-
neo, is hardly known outside its native island. This deciduous tree has very
stout twigs, with leaves more or less aggregate at the apices. The globose
fruits, up to 20 cm in diameter, are the largest known fruits in the genus. The
rough, potato-brown rind (0.51 cm thick) can be peeled off like that of a
banana. Its bright, deep yellow, thick and brous esh is sweet with a dis-
tinctive taste (Kostermans, 1965; Bompard, 1991a). In orchards in Borneo
where M. foetida and M. pajang are both cultivated, forms with leaves and
fruits having intermediate characters are occasionally found.
Mangifera caesia, M. foetida, M. pajang and especially M. odorata are impor-
tant in tropical humid regions where the common mango cannot be grown
satisfactorily. Mangifera pajang has potential as an ornamental tree, having
brilliant rose-red blossoms (Philipps et al., 1982).
SUBGENUS MANGIFERA. The subgenus Mangifera contains most of the species (47),
and is divided into four sections: (i) section Marchandora Pierre; (ii) section Euan-
therae Pierre; (iii) section Rawa Kosterm.; and (iv) section Mangifera Ding Hou.
Section Marchandora Pierre. This section has only one species, M. gedebe Miquel
(syn. M. camptosperma Pierre, M. inocarpoides Merr. and Perry, M. reba Pierre).
The labyrinthine seed is unique to this species, wherein the inner integu-
ments penetrate the cotyledons and form numerous irregular folds. The at,
discus-like fruit has only a very thin mesocarp. Mangifera gedebe grows in
inundated places along rivers or lakes. The seed oats in water and is dis-
persed during periods of high water, and this may explain its wide distribu-
tion, from Myanmar through Malesia to New Guinea and the Bougainville
Island.
Section Euantherae Pierre. The three species in this section (M. caloneura Kurz
(syn. M. duperreana Pierre), M. cochinchinensis Engler and M. pentandra Hook.
f.) appear to be the most primitive among the species of the subgenus
Mangifera. The owers are characterized by the presence of ve fertile sta-
mens. The three species are mainly conned to Myanmar, Thailand, Indo-
china and the north of the Malay Peninsula. The region is in the transition
zone from the humid tropical rainforest to monsoon forest, and these species
show an adaptation to low rainfall. Mangifera cochinchinensis, which occurs in
south-eastern Thailand and in Vietnam, has small oblong fruits with a thin
seed; the fruits are much relished by local people in southern Vietnam,
although they are very acidic. Mangifera caloneura and M. pentandra are closely
J.M. Bompard 26
related, and can be mistaken for M. indica. However, their leaves are more
leathery, have a more conspicuously dense reticulation, and the panicles are
much more hirsute than the common mango. Mangifera caloneura occurs from
Myanmar through Thailand to Indochina, in lowland evergreen forests, as
well as in semi-deciduous forests. It is cultivated for its acidic-sweet fruit,
and has been planted along the streets of Vientiane and Ho Chi Minh City
(Saigon). Mangifera pentandra, apparently native to the northern Malay Pen-
insula close to the Kra isthmus transition zone, is found in old orchards, in
scattered locations, especially in Kedah and possibly also in peninsular Thai-
land. It is also grown in the Anambas Islands and in Sabah, where it might
have been introduced in early times. It is a prolic bearer, with small man-
goes, c.8 cm length, and ripening green or yellow. The pale orange, watery
pulp has a sweet taste and few bres.
Section Rawa Kosterm. This group, consisting of nine species, is not well
delimitated. Most species have thick twigs and rather coriaceous leaves
seated on protruding pedestals. The small, hardly attened ovoid or ellip-
soid fruits that are black or partly red at maturity in several species are also
characteristic. Rawa is the Malay word for marsh, indicating that these spe-
cies usually are found in periodically or permanently inundated areas. The
ve species that occur in west Malesia (M. gracilipes, M. grifthii, M. micro-
phylla, M. paludosa and M. parvifolia) grow primarily in the swamps of south
peninsular Malaysia, in central coastal areas of east Sumatra and western
Borneo, and occasionally in peripheral uplands. It has also been reported
from the Andaman Islands and from Thailand (Sreekumar et al., 1996; Eiad-
thong et al., 2000a).
Mangifera andamanica and M. nicobarica are endemics from the Andaman
and Nicobar Islands, respectively. Mangifera merrillii is a rare species endemic
to the Philippines and M. minutifolia is known solely from a single collection
from southern Vietnam. Mangifera grifthii and M. microphylla are the only
cultivated species within section Rawa. The former species is considered to
be representative of the section, and is cultivated along the eastern coast of
peninsular Malaysia and in western Borneo, and rarely in Sumatra. The fruits
are small (35 cm long) and oblong or ovoid; the skin is rose-red, turning
purplish black at maturity. The rind is thin and easily removed from the
orange-yellow pulp, which is juicy and pleasantly sweet. Different forms are
recognized by local people, according to the size and taste of fruits. Mangifera
microphylla is a related, but less well-known species, having thinner leaves
and a rather similar fruit.
Section Mangifera Ding Hou. With more than 30 species, section Mangifera is
by far the largest. The common mango and the related M. laurina belong
here. Species within the section have the same distribution range as the
genus. The section may be divided into three groups based on oral structure
and organ number variation: (i) those having pentamerous owers; (ii) those
having tetramerous owers; and (iii) an intermediate group of species hav-
ing both pentamerous and tetramerous owers. Within these three groups, it
is possible to distinguish species with either puberulous or glabrous panicles.
Taxonomy and Systematics 27
Only characteristics of representative species within each group, especially
those found in cultivation, are described below.
Pentamerous owers (14 species): Three species, M. laurina, M. minor and
M. sylvatica, show afnity with the common mango. Mangifera laurina is a
species of the lowland forests of Malesia, where it is also under cultivation in
old orchards. It can be distinguished from the common mango by having lax
and widely pyramidal, glabrous or sparingly puberulous panicles. The ow-
ers are smaller and are not glomerulate; the petals have a different shape,
texture and colour. The fruit resemble those of a small common mango, with
orange-yellow pulp, which is almost liquid at maturity. It is generally con-
sumed when unripe. Several forms are in cultivation; however these are now
becoming rare. Mangifera laurina is well suited to the humid tropical lowlands,
fruiting well in areas where the common mango cannot be grown satisfacto-
rily; moreover, it appears to be highly resistant to anthracnose (Bompard,
1991b).
Mangifera minor occurs east of Wallaces line, from Sulawesi to New
Guinea (east Malesia) and to the Carolines Islands in the east. It is adapted to
a wide range of ecological conditions, growing equally well in dry savannahs
and in tropical rainforests up to 1300 m. The fruit is obliquely oblong, 510 cm
long, much narrowed, the tip obtuse, with a distinct beak and sinus. It is
found in cultivation, although the yellowish fruit pulp is acidic and scant.
Mangifera sylvatica is found from Sikkim (up to 1200 m) to northern Myan-
mar and Thailand, and apparently also in Yunnan up to 1900 m. The fruit is
obliquely ovate, 810 cm long, much compressed distally forming a hook,
has scanty whitish-yellow pulp which is almost breless. Other species are
occasionally found in cultivation, for example M. rufocostata, which is esteemed
by the Banjarese people of South Kalimantan for its very sour fruits that are
used to prepare a spicy condiment with chilli.
Tetramerous owers (15 species): Mangifera altissima is apparently endemic
to the Philippines, where it occurs mainly at low elevations in the forests
from northern Luzon to Mindoro (Brown, 1950). The fruit is mango shaped,
ovoid or ellipsoid, slightly compressed, up to 8 cm length, green or some-
what yellow when ripe, with whitish, sweetish-acidic esh. It is commonly
found in dooryards, and thrives in regions with distinct wet and dry seasons
(Angeles, 1991).
Mangifera torquenda occurs wild in west Malesia, and is cultivated in
south Sumatra and in Borneo, where it is common in the forests and orchards
of eastern Kalimantan. The sub-globose fruit, c.7.5 cm long and 6.5 cm in
diameter, is yellow-green with darker spots at maturity, and has a thin rind.
The pale yellow pulp has a rather pleasant sweet-acid, slightly resinous taste
and a light turpentine smell. Short bres are attached to the seed. It is closely
related to M. longipetiolata.
Mangifera magnica is a common species in the rainforests of western
Malesia, occasionally cultivated in central Sumatra and in West Kalimantan,
where it has a special importance in the myths of Land Dayak peoples. The
fruit is ovoid oblong, up to 12 cm long, 10 cm in diameter, only slightly
compressed, greyish green with brown spots. The pulp is whitish, soft at
J.M. Bompard 28
maturity, sweetish acid. Sweeter forms are reported in central Kalimantan
(J.J. Afriastini, personal communication). The stone is unique in the genus in
that it lacks bres adhering to it.
Mangifera quadrida is found from peninsular Malaysia to the Moluccas.
The fruit is ellipsoid-globose, 68 cm long, green covered with black dots turn-
ing completely black at maturity, and has a pale yellow, sweet-acid pulp.
Another form is recognized by its more coriaceous leaves, smaller fruits, c.4 cm
long, having dark yellow pulp, purplish around the stone, and a sweet, palat-
able taste, somewhat like prunes. Both forms are cultivated in old orchards.
Tetra- and pentamerous owers (four species, and also M. indica): Mangifera
casturi is related to M. quadrida, from which it can be distinguished by leaf
and fruit characters. It has never been collected in the wild, and is a favourite
among the Banjarese people in south Kalimantan. The fruits are small, a little
compressed and up to 6 cm in length, becoming completely black at matu-
rity. The orange pulp is very sweet and palatable, and resembles honey
mango or mangga madu grown in East Java. Although M. casturi bears
heavily, it has a strong- to alternate-bearing habit. It is an excellent fruit for
the humid tropical lowlands, and appears to be resistant to anthracnose. Sev-
eral differently named forms exist; these have polyembryonic seeds. Mangifera
rubropetala is also only known in cultivation, and may be a primitive race of
M. indica.
SPECIES OF UNCERTAIN TAXONOMIC POSITION. There is a group of 11 disparate spe-
cies of uncertain taxonomic position that cannot be placed with certainty due
to the absence of adequate material. There are three species only known in
China.
2.4 Phytogeography
Species distribution
An examination of the present distribution of the genus shows that the larg-
est number of Mangifera species in either subgenera is found in western
Malesia on the Sunda shelf. A decreasing number of species occurs towards
the genus boundary east of Wallaces line in east Malesia, and in its northern
and western range of distribution. While peninsular Malaysia and the islands
of Sumatra and Borneo have the highest diversity of species, the number of
species becomes gradually lower in east Malesia, especially in the Lesser
Sunda Islands, Moluccas and New Guinea. This is explained by the geologic
and paleogeographic features of the Malesian region which spans two large
partly submerged continental shelves, the Asiatic shelf (Sunda Shelf linking
the Malay Peninsula with the islands of Sumatra, Java, Borneo and Palawan)
and the Australasian shelf (Sahul Shelf linking the Aru islands and New
Guinea with Australia). During the last glaciation period (c.22,50011,000 bp)
the shelves were regions of land uncovered by the lowering of sea level, and
present day peninsular Malaysia, Sumatra and Borneo were connected by
Taxonomy and Systematics 29
land bridges during the late period of maximal sea lowering. During the cool
periods of glacial maxima, the Malesian forest was reduced in extent, but
there is no evidence that it was reduced to isolated island forests. The Sunda-
land and Papuasian rainforest blocks are therefore comparable to refugia in
terms of species richness and the high degree of endemism (Whitmore, 1981).
Mangifera has undergone major species development in west Malesia, which
has remained relatively stable over a long period of time. The current vegeta-
tion of west Malesia probably differs very little from that at the end of the
Tertiary (van Steenis, 1950). A lower number of Mangifera species is found in
Java and the Philippines, regions less often connected with Asia during the
Pleistocene.
Only three species occur in New Guinea, which is largely covered with
rainforest. These include M. minor, M. mucronulata and the widely distributed
M. gedebe. Mangifera foetida also occurs, but may have been introduced. Mangifera
minor occurs from Celebes and the Philippines to the Solomon Islands; M.
mucronulata is found in the Moluccas, New Guinea and the Solomon Islands.
The distribution of these species suggests a late immigration of a Laurasian
genus from Sundaland via the Philippines, Sulawesi and the Moluccas into
New Guinea, which is supported by the geological history of the region. No
Mangifera species have ever been recorded from northern Australia.
Very few species are found in peninsular India and Sikkim. From present-
day distribution, there is little evidence of migration of species into the sub-
continent of India after its collision with Eurasia in the middle Eocene
(Audley-Charles et al., 1981). According to Mehrotra et al. (1998), fossil leaves
described as Eomangiferophyllum damalgiriensis Mehr. from the Upper Palaeo-
cene in north-eastern India are an analogue of the modern genus Mangifera.
Mangifera sylvatica occurs along the northern limit of the range of
Mangifera, with more or less discontinuity, from Sikkim to northern Thailand
and to the southern part of Yunnan, where it is reported in mountains up to
1900 m above sea level (Anonymous, 1980). The few species that grow in
southern China are very poorly known: M. austro-yunnanensis from western
Yunnan, M. persiciformis from south-eastern Yunnan and southern Guizhou
at latitudes up to 26N and M. hiemalis, the winter mango from Guangxi
near the northern border Vietnam. In the revised Flora of China (Min and Bar-
fod, 2008), M. austro-yunnanensis is considered to be conspecic with M.
indica, M. hiemalis is treated as a synonym of M. persiciformis, and M. laurina
is recorded from the lowland forests of south Yunnan.
Subgenera and section distribution
The species distribution is especially meaningful when the ranges of the spe-
cies of each subgenus and section are considered separately.
Subgenus Limus
All species of the subgenus Limus are restricted to the Malesian area (M. foetida
and M. macrocarpa occurring in peninsular Thailand), whereas all the species
J.M. Bompard 30
with ve fertile stamens, considered the most primitive condition, are con-
ned to west Malesia (M. decandra in Sumatra and Borneo; M. lagenifera in the
two latter areas and in peninsular Malaysia). Only M. caesia, M. foetida and
closely related M. leschenaultii occur in east Malesia.
Subgenus Mangifera
In the subgenus Mangifera, M. gedebe is the only species belonging to the sec-
tion Marchandora, and has the widest range within the genus, extending from
Myanmar through Malesia to New Guinea and Bougainville Island. Section
Euantherae is centred in the region from Myanmar to Vietnam. Mangifera
pentandra is only known from peninsular Malaysia, the Anambas Islands and
Borneo. Section Rawa is mainly in western Malaysia and shows notable
diversication in the swamps and peripheral uplands in the south of penin-
sular Malaysia, east central Sumatra (notably the Riau province) and west
Borneo. During the glacial period this area, termed the Riouw pocket (Cor-
ner, 1978), formed a vast plain connecting the Malay Peninsula, Sumatra and
Borneo, and is believed to have been lled with swamps. Mangifera merrillii
is an endemic of the Philippines, M. minutifolia is an endemic of Vietnam, M.
andamanica and M. nicobarica are endemics of the Andamans and Nicobar
Islands. None of the species of section Mangifera occurring in mainland
South-east Asia, north of the isthmus of Kra, are found in eastern Malesia;
however, it would be interesting to assess the genetic relatedness of M. syl-
vatica and M. minor, and also M. laurina, which may prove to be phylogeneti-
cally very closely related.
2.5 Interspecic Molecular Characterization
Molecular biology techniques now make it possible to assess genetic related-
ness in a more precise way. Published data support some of the groupings based
on anatomical characters (Kostermans and Bompard, 1993) but not entirely.
RAPD (random amplication of polymorphic DNA) markers were rst
used in mango by Schnell and Knight (1993) and Schnell et al. (1995). Nine
Mangifera species were analysed and compared to the traditional taxonomic
groupings. The unweighted pair group method of arithmetic averages
(UPGMA) cluster analysis for the subgenus Limus was not supportive of the
separation between sections Perennes and Deciduae, which, admittedly, has a
weak taxonomic basis. It conrmed the relatedness between M. foetida and
M. pajang. The UPGMA cluster analysis of the subgenus Mangifera supported
the current taxonomy based on ower morphology. It showed the related-
ness between M. quadrida and M. torquenda (both placed in the group of
species with tetramerous owers), but also with M. casturi, although the lat-
ter species has tetra- and pentamerous owers. One of the most signicant
results was the evidence for the existence of interspecic hybridization within
the studied species of the section Mangifera (see also Yonemori et al., 2002).
Phylogenetic relationships among 14 Mangifera species of Thailand were
analysed by comparing amplied fragment length polymorphism (AFLP)
Taxonomy and Systematics 31
markers (Eiadthong et al., 2000b), and by comparing sequences of the inter-
nal transcribed spacer (ITS) region of nuclear ribosomal DNA (nrDNA)
(Yonemori et al., 2002). They demonstrated that the common mango was
closely related to M. laurina, M. sylvatica and M. oblongifolia of subgenus
Mangifera to which M. indica belongs. A close relationship between M. indica
and M. sylvatica has been corroborated by Nishiyama et al. (2006), who com-
pared signal intensity of genomic in situ hybridization (GISH) on somatic
metaphase chromosomes of M. indica, using labelled DNA of eight wild
Mangifera species.
Furthermore, Eiadthong et al. (2000b) and Yonemori et al. (2002) have
demonstrated that M. odorata, M. foetida and M. macrocarpa (of subgenus
Limus) were related to M. indica. It is not surprising in the case of M. odorata
whose hybrid origin (M. foetida M. indica) has now been established, but
this calls into question the position of the section Perennes.
Results of molecular studies do not permit a comprehensive view of the
phylogenetic relationships among the genera. So far, they are rather support-
ive of the groupings based on phenotype within the subgenus Mangifera
(notably sections Rawa and Euantherae), but not for the subgenus Limus which
will need to be redescribed, and likely restricted to the group of species
related to M. caesia (M. kemanga, M. lagenifera, M. superba and possibly M.
decandra). More studies will be needed to infer phylogenetic relationships
within the section Mangifera. Keeping in mind the frequent misidentications
in collections and botanic gardens, herbarium specimens of studied material
must be deposited in the national herbaria so that its taxonomic position can
be ascertained in case of doubt.
2.6 Region of Origin of the Genus
Based on morphological, phytogeographical and fossil evidence, Mukherjee
(1953) argued that:
although the highest number of species of both sections is concentrated in the
Malay Peninsula [19 were then recorded], the centre of origin of the genus
cannot be restricted to that area alone, as both the phylogenetically older
species, i.e. with pentacyclic owers (M. duperreana, now reduced to M.
caloneura, and M. lagenifera), occur in Siam and Indochina, and the former is
absent from Malay Peninsula.
He concluded that the genus had its origin somewhere in the Myanmar
ThailandIndochina area or in the Malayan area. Careful identication of the
greatest part of herbarium materials available has allowed a more accurate
delimitation of the distribution ranges of the Mangifera species, notably of the
subgenus Limus, and has revealed, among other things, that M. lagenifera
does not occur north of Kra isthmus contrary to Mukherjees assertions. Fur-
thermore, the ten-stamen species, M. decandra, which was described by Ding
Hou in 1972 and hence was unknown to Mukherjee, is conned to Borneo
and Sumatra, and to date has not been recorded from peninsular Malaysia.
J.M. Bompard 32
Without overemphasizing the present great species diversity of subgen-
era in the Malay Peninsula, Borneo and Sumatra, the available evidence
points to a Sundaic origin for the genus. This, however, must not minimize
the particular importance of the region stretching from Myanmar to Indo-
china as another centre of diversication, as attested by the range of species
belonging to the section Euantherae. Unfortunately, many of the species of
this region remain poorly known, and it can be expected that plant collecting
in the region will yield interesting new ndings. The speciation that occurred
in this region with a likely radiation centre today traced by the range of the
section Euantherae, is of special signicance as it has given rise to the com-
mon mango.
2.7 Origin of the Common Mango
The common mango apparently originated in regions on the western border
of the secondary centre of diversication mentioned above. Truly wild mango
trees have been recorded in Bangladesh (Chittagong Hill tract, c.23N), north-
eastern India (undoubtedly indigenous in the evergreen tracts of valley of
Assam according to Kanjilal et al., 1937), and in Myanmar where it was
reported as not unfrequent in the tropical and lower mixed forests all over
Burma from Arracan and Pegu down to Tenasserim (Kurz, 1877). It would
be desirable to assess its afnity with the species of the section Euantherae, as
well as with species of other sections of the subgenus Mangifera that occur in
the same area and region. It is also believed to be wild in the sub-Himalayan
tract, in deep gorges of the Baraitch and Gonda hills in Oudh, and the outer
hills in Kamaon and Garhwal (Brandis, 1874). The common mango has been
grown and disseminated for such a long time in India that semi-wild trees
can be found in the forests throughout the subcontinent. The fruits of wild
trees are said to be small and of poor quality. Watt (1891) mentioned two so-
called almost unaltered wild varieties existed under cultivation in Tirhoot,
one originating from Kangra, a very variable one, and the other from Sikkim
which was evidently the progenitor of the varieties cultivated in Malda.
The common mango in South-east Asia
The Linnean binomial (Mangifera indica) indicates in this instance the place
where the common mango was selected and improved, and not necessarily its
place of its origin. It has been traditionally accepted that mango was domesti-
cated several millennia ago in India (see Mukherjee and Litz, Chapter 1, this
volume); however, it cannot be excluded that domestication occurred inde-
pendently in several areas, possibly in the south-western and south-eastern
regions of its centre of origin, or later differentiated in those two regions. This
hypothesis would account for the differences that exist between the local
polyembryonic cultivars of Myanmar, Thailand, Indochina and Indonesia,
and the monoembryonic Indian cultivars. Note that polyembryony occurs
Taxonomy and Systematics 33
also in the cultivated M. casturi, M. laurina and M. odorata. Aron et al. (1998)
have demonstrated that polyembryony in mango is under the control of a
single dominant gene.
According to Juliano (1937), Bijhouwer suggested that there were two
main centres of domestication of mango, one in India with monoembryonic
mangoes, the other in the Saigon area, Indonesia and the Philippines with
polyembryonic mangoes. The Saigon area must in fact be extended to
southern Vietnam, other parts of Indochina, Thailand and Myanmar, which
were recognized by Valmayor (1962) as homes of polyembryonic mangoes.
Notwithstanding, the origin of polyembryonic mangoes is probably better
placed in Myanmar, and possibly the eastern part of Assam. According to
Brandis (1874), in Burma, the mango is not generally grafted, and seeds of a
good kind, as a rule, produce fruit of a similar description. There are only a
few polyembryonic mango cultivars in India. They are restricted to the south-
western coastal region, and geographically isolated from the polyembryonic
mangoes of Myanmar and South-east Asia. Analysis of genetic relatedness
using RAPD markers among polyembryonic and monoembryonic cultivars
grown in the west coast of southern India suggest that the polyembryonic
types are unlikely to have originated from India and might have been intro-
duced from South-east Asia (Ravishankar et al., 2004).
Indian Buddhist monks might have introduced the common polyembry-
onic mango to South-east Asia, rst along land trade routes through Myan-
mar, where they might have found better races, and from there into insular
South-east Asia. It is well established that some local names of the common
mango currently used in parts of Indonesia are of Sanskrit origin (ampelam
and its cognates), and are sometimes used to designate M. laurina, which is a
truly native species. Vernacular names do not always travel with a plant, and
even if they did so in the case of the common mango, it is very unlikely that
these introductions were the rst ones and that they came obligatorily from
India. In the absence of a comprehensive classication of the innumerable
South-east Asian cultivated forms of the common and wild mangoes, includ-
ing the countless primitive races, we have to rely on linguistics and the rich
history and prehistory of this region.
Vernacular names
The different local names of the common mango in Indonesia (pauh, ampe-
lam and its variants, and mangga) bear evidence of a long history of con-
tacts with mainland Asia and India, and point to possible introduction at
different times from different places. In some parts of Indonesia, the vernacu-
lar names paoh or pauh refer either to primitive races of the common
mango, or to native species, as a rule the ones most closely resembling the
common mango, for example: pauh asal (= native mango) for M. pentandra
in peninsular Malaysia; pahohutan or pahutan (= forest mango) for M.
altissima in the Philippines; and pao pong (= forest mango) for M. minor in
Flores, Lesser Sunda Islands. Pau is a word belonging to Austronesian lan-
guages, nowadays spoken over a very wide area from Madagascar to the
Easter Islands by people who originate from mainland Asia. These languages
J.M. Bompard 34
are still spoken by certain minority populations in Vietnam, Cambodia and
the Mergui Archipelago off the coast of Myanmar (Bellwood et al., 1995). In
Cambodia, which was occupied by the Chams from about the 3rd to the 15th
century ad, pa:uh is a Chamic word. Sva:y, used by the Khmers (as in
sva:y srok meaning mango of the village (M. indica), and sva:y prey, wild
mango (i.e. M. caloneura) as attested in pre-Angkorian Khmer inscriptions
dating from the 6th to the 8th century ad (Pou and Martin, 1981)) is of Austro-
Asiatic origin. Sva:y has cognates in south Vietnam (xoay) and in Asian
languages spoken by aboriginal people in peninsular Malaysia. Wai, another
cognate, is a vernacular name of M. minor in several parts of New Guinea.
Pawley and Ross (1995) proposed wai and pau(q) as the reconstructed
Proto-Oceanic terms referring, respectively, to a generic name for mango,
and a species that is probably M. indica.
Nowadays, these two words are generic terms for mango fruits that
rather closely resemble M. indica. In the same way, thayet which is the com-
mon vernacular name referring to M. indica in Myanmar (sinnin thayet and
taw-thayet for M. caloneura and M. sylvatica, respectively), or mamuang in
Thai languages are probably generic names.
Obviously, linguistic evidence alone provided by these vernacular names
is not sufcient to prove the time and place of an introduction. None the less,
it is signicant that in mainland South-east Asia none of the vernacular names
of the common mango exhibits signs of an Indian inuence, moreover, cog-
nates of these names are also applied to primitive races in some parts of
insular South-east Asia.
Evidence of early trade in South-east Asia
The history of plant domestication in mainland South-east Asia has undoubt-
edly involved introduction of plants by people migrating from the mainland
into insular South-east Asia. In more recent times, there is evidence of con-
tacts and sea trade since at least the rst centuries ad between mainland and
insular South-east Asia to indicate that there have been numerous opportuni-
ties for introduction of the common mango from different places at different
times prior to the 4th century (before the Indianization of early South-east
Asian states) into present-day Malaysia and Indonesia.
Recent studies based on archaeological evidence stress the long unrecog-
nized importance of South-east Asian trade (emanating from South-east
Asia) between ports established along the Java Sea, those of mainland Asia,
and India, back to the 1st century ad, and possibly earlier (Walker and San-
toso, 1984). Trade routes connected the developing population centres of the
mainland, such as the earliest known South-east Asian political entity, Funan,
an advanced agrarian society located on the southern Vietnam coast, which
became inuenced by the Indians and reached the zenith of its commercial
prosperity in the middle of the 3rd century (Hall, 1985). Increasingly, king-
doms organized according to the Indian concept of royalty were established
in the Indonesian archipelago, for example Kutai in East Kalimantan (4th
century) and Central Java (8th to 9th century), the latter being famous for the
Buddhist temple at Borobudur, where sculptures depict the mango tree.
Taxonomy and Systematics 35
It is highly probable that the eventual introductions of superior cultivars
of polyembryonic mangoes from the south-west coast of India, between the
6th and 14th century, the height of classical South-east Asian civilization and
also the golden age of early south Indian civilization (Hall, 1985), were not
the rst ones.
During the 16th and 17th centuries, the Portuguese and Spaniards con-
tributed to the widest distribution of superior varieties in the archipelago, espe-
cially to the east. The name mango itself derives from the Tamil man-kay or
man-ga (see Mukherjee and Litz, Chapter 1, this volume), which the Portu-
guese adapted as manga and mangueira when they colonized west India.
Superior Philippine cultivars originated through introduction of culti-
vars from Indonesia, for example Dodol into Mindanao, and from Indo-
china, for example Carabao and Pico in Luzon, the Visayas and northern
Mindanao (Wester, 1920; Bondad et al., 1984). However, these introductions
dating from the rst half of the 17th century were also preceded by the intro-
duction of primitive races of the common mango as well as other species into
the Sulu Archipelago and Mindanao through contacts with north Borneo, as
attested by their local names quoted by Wester (1920), that is mampalam
(M. indica, and possibly also M. laurina), baonoh (M. caesia) and wannih (M.
odorata).
The South-east Asian M. indica germplasm includes many races that defy
classication. Natural cross-pollination has undoubtedly occurred with
native species, such as M. laurina, which was also brought into cultivation in
several areas before the introduction of M. indica.
2.8 Conclusion
Potential contribution of wild species to mango cultivation
To date, the improvement and breeding of common mango has depended on
the use of genetic variability within a single species, M. indica. Mukherjee
(1957) observed that similarity in chromosome number and pollen morphol-
ogy in different species suggests close compatibility during hybridization
and stock-scion relationship if other species are used as stock for the com-
mon mango. Biotechnology opens new perspectives for mango improve-
ment (Litz, 2004). As noted by Litz et al. (Chapter 18, this volume), the
transformation of mango with genes from other species could address a
number of plant breeding objectives.
Source of rootstock
Grafting experiments between M. indica and other species are reported in the
literature, for example budding of M. indica on M. foetida and M. odorata in
Java (Ochse and Bakhuizen, 1931), M. odorata on M. indica in the Philippines
(Wester, 1920), and M. indica on M. zeylanica in Sri Lanka (Gunaratman, 1946).
J.M. Bompard 36
Mangifera indica Madu in Java, and M. laurina in Sabah have been used as
rootstocks for M. casturi. Trials of grafted M. caesia on M. indica (Wester, 1920)
and M. indica on M. kemanga or M. caesia (Ochse and Bakhuizen, 1931) were
unsuccessful, as these two species have distinct bark features and only remote
afnity with the common mango. Better compatibility can be expected using
species more closely related to the common mango within the subgenus
Mangifera. In West Kalimantan, M. laurina is occasionally used as a rootstock
for the common mango on periodically inundated riverbanks. It has been
tried as a rootstock by the Department of Agriculture in Sabah (Lamb, 1987).
Campbell (2004) reported that M. casturi, M. grifthii, M. laurina, M. odorata,
M. pentandra and M. zeylanica grafted on M. indica had a high percentage of
success.
Several species that can grow in permanently inundated areas (i.e. M.
gedebe, M. quadrida, M. grifthii and other species of the section Rawa) repre-
sent a potential source of rootstock for the development of mango cultivation
on poorly drained soils or in areas liable to prolonged ood. Other species
may be a source of dwarng rootstocks.
Hybridization
From our observations in Borneo, natural interspecic hybridization involv-
ing various cultivated Mangifera species can occasionally occur. Suspected
hybrids were observed between wild M. gedebe and cultivated M. laurina in
the lakes area along the Mahakam River in East Kalimantan, where impor-
tant populations of M. gedebe occur; between cultivated M. foetida and M. pajang,
two species showing close afnity, in different areas of Kalimantan where
both species are grown together; and between closely related M. caesia and
M. kemanga in cultivation. A hybrid origin has been suggested for M. odorata
(M. indica M. foetida), which is unknown in the wild (Ding Hou, 1978a).
Based on AFLP analysis, Teo et al. (2002) and Kiew et al. (2003) have con-
rmed that M. odorata is a hybrid between M. foetida and M. indica. The index
of similarity showed that M. odorata is closer to M. foetida (76% similarity)
than it is to M. indica (66%). Yamanaka et al. (2006) showed a high genetic
similarity among 11 landraces of M. odorata from the Malaysian Agricultural
Research and Development Institute (MARDI) gene bank. Higher variability
can be expected from Sumatra and Java samples.
Existing information about experimental interspecic hybridization is
scarce. According to Mukherjee et al. (1968), successful crosses between
M. odorata and M. zeylanica were made in India.
Potential of wild species
There is little doubt that wild mangoes are potentially valuable in breeding
programmes. Some species have important horticultural implications as they
demonstrate many desirable characteristics (Bompard, 1993). Fairchild (1948)
Taxonomy and Systematics 37
noted that crosses between the common mango and related ve-stamen spe-
cies of the section Euantherae might produce hybrids with better pollinating
quality. Mangifera pentandra, which is grown in peninsular Malaysia and
Sabah, is a prolic bearer, due to its high proportion of hermaphrodite to
male owers.
Stress resistance
In the Malesian rainforests, wild mangoes thrive well under an ever-humid
climate, without a prolonged dry season, i.e. is in areas with an annual rain-
fall > 4000 mm and no monthly mean < 100 mm and where the common
mango cannot be grown satisfactorily. Species, occurring in subtropical areas,
including primitive races of the common mango, or in high altitude tropical
forests, should be evaluated for cold tolerance, opening up the possibilities
for mango production in subtropical and Mediterranean areas. Mangifera lau-
rina and other species related to the common mango that grow in the rainfor-
est (e.g. M. minor in New Guinea) are apparently immune to anthracnose.
Sharma and Choudhury (1976) also observed that trees of an unknown wild
race found in the Tripura State (north-eastern India) were free from mango
malformation.
Potential new fruits
Extensive, yet largely unrecorded variability also exists among the non-indica
species under cultivation. Sadly, this gene pool is barely represented in exist-
ing collections, and is rapidly vanishing. An increasing number of horticul-
turists are demonstrating a keen interest in the wild relatives of the mango.
It is hoped that local peoples who have contributed to the recognition and
maintenance of these species can benet from future innovative mango
breeding.
Since early times, local peoples have planted seeds collected from trees
that were observed to produce better quality fruits in the forests around their
settlements. In areas now completely devoid of lowland primary forest, espe-
cially in Sumatra and Borneo, the only wild relatives still found are those
which have been integrated into indigenous agroforests which represent
gene banks for an amazing diversity of fruit trees. A tenuous but constant
selection pressure over many centuries has resulted in improved selections
of several species. Today, some of these selections hold economic importance
for their intrinsic characteristics. In Malesia, forms of M. odorata and M. foetida
with sweeter and less brous esh have been identied. The wani, a form of
M. caesia from Bali and Borneo, has green-skinned fruit with milky white soft
esh and a sweet taste quite different from the fruit of common forms of M.
caesia. In addition, there are many interesting selections of M. casturi, M. grif-
thii and M. torquenda.
Further improvement of these wild mangoes is especially desirable
owing to their local economic importance in the wet tropical regions. Use of
vegetative propagation methods must be encouraged. With proper selection,
there is every reason to believe that other Mangifera species can become valu-
able commercial fruits.
J.M. Bompard 38
Acknowledgement
Thanks are due to Dr Dawn Frame who assisted in correcting the text.
Note
1
Surveys were carried out in Kalimantan in cooperation with the Indonesian Institute of
Science (LIPI) and the Indonesian Commission on Germplasm, and in Malaysia with the
Forest Research Institute of Malaysia (FRIM).
References
Angeles, D.E. (1991) Mangifera altissima Blanco. In: Verheij, E.W.M. and Coronel, R.E.
(eds) Plant Resources of South-east Asia No.2: Edible Fruits and Nuts. Pudoc-DLO,
Wageningen, the Netherlands, pp. 206207.
Anonymous (1980) Mangifera L. In: Cheng, M. and Ming, T.L. (eds) Flora Reipublicae
Popularis Sinicae. Vol. 45(1). Science Press, Beijing, Peoples Republic of China,
pp. 7378.
Aron, Y., Czosnek, H., Gazit, S. and Degani, C. (1998) Polyembryony in mango
(Mangifera indica L.) is controlled by a single dominant gene. HortScience 33,
12411242.
Audley-Charles, M.G., Hurley, A.M. and Smith, A.G. (1981) Continental movements in
the mesozoic and cenozoic. In: Whitmore, T.C. (ed.) Wallaces Line and Plate Tec-
tonics. Clarendon Press, Oxford, UK, pp. 923.
Bellwood, P., Fox, J.J. and Tryon, D. (eds) (1995) The Austronesians, Historical and Com-
parative Perspectives. The Australian National University, Canberra.
Bompard, J.M. (1991a) Mangifera caesia Jack and Mangifera kemanga Blume, Mangifera
foetida Lour. and Mangifera pajang Kostermans. In: Verheij, E.W.M. and Coronel,
R.E. (eds) Plant Resources of South-east Asia No.2: Edible Fruits and Nuts. Pudoc-
DLO, Wageningen, the Netherlands, pp. 207211.
Bompard, J.M. (1991b) Mangifera laurina Blume and Mangifera pentandra Hooker f.;
Mangifera odorata. In: Verheij, E.W.M. and Coronel, R.E. (eds) Plant Resources of
South-east Asia No.2: Edible Fruits and Nuts. Pudoc-DLO, Wageningen, the Neth-
erlands, pp. 216220.
Bompard, J.M. (1993) The genus Mangifera re-discovered: the contribution of wild spe-
cies to mango cultivation. Acta Horticulturae 341, 6977.
Bompard, J.M. (1995) Surveying Mangifera in the tropical rain forests of South-east Asia.
In: Guarino, L., Ramanatha Rao, V. and Reid, R. (eds) Collecting Plant Genetic Diver-
sity. Technical Guidelines. CAB International, Wallingford, UK, pp. 627637.
Bondad, N.D., Rivera, F.N., Agcopra, D.B. and Minh Tam Aurin (1984) Philippine man-
goes and their relationship to South-east Asian cultivars. Philippine Geographical
Journal 28, 5971.
Brandis, D.D. (1874) Forest Flora of North West and Central India. Allen, London,
pp. 125127.
Brown, W.H. (1950) Useful Plants of the Philippines. Vol. 2. Bureau of Science, Manila,
the Philippines, pp. 336340.
Campbell, R.J. (2004) Graft compatibility between Mangifera species and Mangifera
indica L. turpentine rootstocks and their subsequent horticultural traits. Acta Horti-
culturae 645, 311313.
Taxonomy and Systematics 39
Corner, E.J.H. (1978) The Freshwater Swamp-forest of South Johore and Singapore. Bo-
tanic Gardens, Parks and Recreation Department, Singapore.
Ding Hou (1972) A new species of Mangifera. Reinwardtia 8, 323327.
Ding Hou (1978a) Flora Malesiana Praecursores 56 (Anacardiaceae). Blumea 24,
141.
Ding Hou (1978b) Anacardiaceae. 4. Mangifera. In: van Steenis, C.G.G.J. (ed.) Flora
Malesiana I. Vol. 8. Rijksherbarium, Leiden, the Netherlands, pp. 423440.
Eiadthong, W., Yonemori, K., Sugiura, A., Utsunomiya, N. and Subhadrabandhu, S. (2000a)
Records of Mangifera species in Thailand. Acta Horticulturae 509, 213224.
Eiadthong, W., Yonemori, K., Sugiura, A., Utsunomiya, N. and Subhadrabandhu, S.
(2000b) Amplied fragments length polymorphism analysis for studying genetic
relationships among Mangifera species in Thailand. Journal of the American Society
for Horticultural Science 125, 160164.
Engler, A. (1883) Anacardiaceae. In: de Candolle, A.P. (ed.) Monographiae Phanerog-
amarum. Vol. 4. Masson, Paris, pp. 195215.
Fairchild, D. (1948) The mango relatives of Cochin China; those with ve-stamen ow-
ers. Proceedings of the Florida State Horticultural Society 61, 250255.
Gunaratman, S.C. (1946) The cultivation of mango in the dry zone of Ceylon (part II).
Tropical Agriculturist 102, 2330.
Hall, K.R. (1985) Maritime Trade and State Development in Early South-east Asia. Allen
and Unwin, Sydney.
Hooker, J.D. (1862) Mangifera. In: Bentham, G. and Hooker, J.D. (eds) Genera Plantarum.
Vol. I. Reeve and Co., London, p. 420.
Juliano, J.B. (1937) Embryos of Carabao mango (Mangifera indica Linn.). Philippines
Agriculturist 25, 749760.
Kanjilal, U.N., Kanjilal, P.C. and Das, A. (1937) Mangifera. Flora of Assam 1(2), 335336.
Kiew, R., Teo, L.L. and Gan, Y.Y. (2003) Assessment of the hybrid status of some Malesian
plants using Amplied Fragment Length Polymorphism. Telopea 10, 225233.
Kostermans, A.J.G.H. (1965) New and critical Malesian plants VII. Reinwardtia 7,
1946.
Kostermans, A.J.G.H. and Bompard, J.M. (1993) The Mangoes. Botany, Nomenclature,
Horticulture, and Utilization. Academic Press, London.
Kurz, S. (1877) Forest Flora of British Burma. Vol. 1. Burma Government Printers, Cal-
cutta, pp. 304305.
Lamb, A. (1987) The potential of some wild and semi-wild fruit trees in Sabah and the
progress made by the Department of Agriculture, Sabah in establishing a germplasm
pool. Paper presented in Forest Research Institute Malaysia, Kepong, Malaysia.
Litz, R.E. (2004) Biotechnology and mango improvement. Acta Horticulturae 645, 8592.
Marchand, L. (1869) Rvision du Groupe des Anacardiaces. Baillire, Paris.
Mehrotra, R.C., Dilcher, D.L. and Awasthi, N. (1998) A palaeocene Mangifera-like leaf
fossil from India. Phytomorphology 48, 91100.
Min, T. and Barfod, A. (2008) Anacardiaceae, Mangifera. In: Wu, Z.Y., Raven, P.H. and
Hong, D.Y. (eds) Flora of China. Vol. 11. Missouri Botanical Garden Press, St Louis,
Missouri and Science Press, Beijing, pp. 338339. Available at: http://ora.huh.
harvard.edu/china/mss/volume11/index.htm#alphabetical_list (accessed 5 August
2008).
Molesworth, A.B. (1967) Malayan Fruits. An Introduction to the Cultivated Species.
Donald Moore Press, Singapore.
Mukherjee, S.K. (1949) A monograph of the genus Mangifera L. Lloydia 12, 73136.
Mukherjee, S.K. (1953) Origin, distribution and phylogenetics afnities of the species of
Mangifera L. Journal of the Linnean Society, Botany 55, 6583.
J.M. Bompard 40
Mukherjee, S.K. (1957) Cytology of some Malayan species of Mangifera. Cytologia 22,
239241.
Mukherjee, S.K., Singh, R.N., Majumder, P.K. and Sharma, P.K. (1968) Present position
regarding breeding of mango (Mangifera indica L.) in India. Euphytica 17, 462467.
Nishiyama, K., Choia, Y.A., Honshoa, C., Eiadthong, W. and Yonemori, K. (2006) Ap-
plication of genomic in situ hybridization for phylogenetic study between
Mangifera indica L. and eight wild species of Mangifera. Scientia Horticulturae
110, 114117.
Ochse, J.J. and Bakhuizen, R.C. (1931) Fruits and Fruitculture in the Dutch East Indies.
G. Kolff, Batavia (Jakarta), Indonesia.
Ochse, J.J., Soul, M.J, Dijkman, M.J. and Wehlburg, C. (1961) Tropical and Subtropical
Agriculture. Vol. 2. MacMillan, New York.
Pawley, A. and Ross, M. (1995) The prehistory of Oceanic languages: a current view. In:
Bellwood, P., Fox, J.J. and Tyron, D. (eds) The Austronesians. The Australian Na-
tional University, Canberra, pp. 3974.
Philipps, A., Philipps, S.M. and Philipps, C. (1982) Some ornamental plants of Sabah.
Nature Malaysiana 7(4), 2027.
Pierre, L. (1897) Flore Forestire de la Cochinchine. Vol.1, fasc. 23. Doin, Paris.
Pou, S. and Martin, M. (1981) Les noms des plantes dans lpigraphie ancienne khmre.
Asie du Sud Est et Monde Insulindien 12, 370.
Ravishankar, K.V., Chandrashekara, P., Sreedhara, S.A., Dinesh, M.R., Lalitha, A. and
Saiprasad, G.V.S. (2004) Diverse genetic bases of Indian polyembryonic and mo-
noembryonic mango (Mangifera indica L) cultivars. Current Science 87, 870871.
Schnell, R.J. and Knight, R.J., Jr (1993) Genetic relationships among Mangifera spp. based
on RAPD markers. Acta Horticulturae 341, 8692.
Schnell, R.J., Ronning, C.M. and Knight, R.J. (1995) Identication of cultivars and valida-
tion of genetic relationships in Mangifera indica L. using RAPD markers. Theoreti-
cal and Applied Genetics 90, 269274.
Sharma, D.K. and Choudhury, S.S. (1976) Occurrence of an unknown wild race of
Mangifera in Tripura. Current Science 45, 305306.
Sreekumar, P.V., Veenakumari, K. and Padhye, P.M. (1996) Mangifera grifthii (Anacar-
diaceae): an addition to the Indian mangoes, from Andaman Islands, India. Ma-
layan Nature Journal 50, 8587.
Teo, L.L., Kiew, R., Set, O., Lee, S.K. and Gan, Y.Y. (2002) Hybrid status of kuwini,
Mangifera odorata (Anacardiaceae) veried by amplied fragment polymorphism.
Molecular Ecology 11, 14651469.
Valmayor, R.V. (1962) The Mango, its Botany and Production. University of the Philip-
pines, Laguna, the Philippines.
van Steenis, C.G.G.J. (1950) The delimitation of Malaysia and its main plant geographi-
cal divisions. In: van Steenis, C.G.G.J. (ed.) Flora Malesiana. Series I. Sijthoff and
Noordhoff Publishers, Groningen, the Netherlands, pp. 7075.
Verheij, E.W.M. and Coronel, R.E. (eds) (1991) Plant Resources of South-east Asia No.2:
Edible Fruits and Nuts. Pudoc-DLO, Wageningen, the Netherlands.
Walker, M.J. and Santoso, S. (1984) Romano-Indian pottery in Indonesia. In: van de
Velde, P. (ed.) Prehistoric Indonesia, a Reader. Foris, Dordrecht, the Netherlands,
pp. 376383.
Watt, G. (1891) A Dictionary of the Economic Products of India. Vol. 5. Ofce of the
Superintendent, Government Printing, Calcutta; W.H. Allen, London, pp. 146157.
Wester, P.J. (1920) The Mango. Bureau of Agriculture Bulletin No. 18. Bureau of Agricul-
ture, Manila, the Philippines.
Whitmore, T.C. (1975) Tropical Rain Forests of the Far East. Clarendon, Oxford, UK.
Taxonomy and Systematics 41
Whitmore, T.C. (1981) Paleoclimate and vegetation history. In: Whitmore, T.C. (ed.) Wal-
laces Line and Plate Tectonics. Clarendon, Oxford, UK, pp. 3642.
Yamanaka, N., Hasran, M., Xu, D.H., Tsunematsu, H., Idris, S. and Ban, T. (2006) Ge-
netic relationship and diversity of four Mangifera species revealed through AFLP
analysis. Genetic Resources and Crop Evolution 53, 949954.
Yonemori, K., Honsho, C., Kanzaki, S., Eiadthong, W. and Sugiura, A. (2002) Phyloge-
netic relationships of Mangifera species revealed by ITS sequences of nuclear ribo-
somal DNA and a possibility of their hybrid origin. Plant Systematics and Evolution
231, 5975.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
42 (ed. R.E. Litz)
3 Important Mango Cultivars and
their Descriptors
R.J. Knight, Jr, R.J. Campbell and I. Maguire
1
University of Florida, Florida, USA
Fairchild Tropical Botanic Garden, Florida, USA
3.1 Introduction 43
3.2 Criteria for Cultivar Description 44
3.3 Mango Cultivars 45
Alfa (Brazil) 45
Alphonso (India) 45
Amelie (West Africa) 46
Arumanis (Indonesia) 46
Ataulfo (Mexico) 46
B74 (Calypso) (Australia) 47
Banganpalli (India) 47
Beta (Brazil) 47
Bombay Green (India) 47
Cambodiana (Vietnam) 48
Carabao (Philippines) 48
Chausa (India) 48
Cogshall (Florida, USA) 49
Corao de Boi (Brazil) 49
Dasheheri (India) 49
Espada (Brazil) 49
Ewais (Egypt) 50
Excellent Succari (Egypt) 50
Extrema (Brazil) 50
Fajri (India) 50
Fernandin (India) 51
Genovea (Egypt) 51
Glenn (Florida, USA) 51
Golek (Indonesia) 51
Haden (Florida, USA) 52
Himsagar (India) 52
Hindi Besennara (Egypt) 52
Hindi Khassa (Egypt) 52
Mango Cultivars and Descriptors 43
Irwin (Florida, USA) 53
Julie (West Indies) 53
Keitt (Florida, USA) 53
Kensington (Australia) 54
Kent (Florida, USA) 54
Khanefy (Egypt) 54
Kyo Savoy (Thailand) 55
Langra (India) 55
Mabrouka (Egypt) 55
Madame Francis (Haiti) 55
Mallika (India) 56
Manila (Mexico) 56
Manzanillo (Mexico) 56
Mesk (Egypt) 56
Mulgoa (India to Florida, USA) 57
Nabeel (Egypt) 57
Nam Doc Mai (Thailand) 57
Neelum (India) 58
Nuwun Chan (Thailand) 58
Okrung (Thailand) 58
Osteen (Florida, USA) 59
Pairi (India) 59
Palmer (Florida, USA) 59
Rosa (Brazil) 59
Sensation (Florida, USA) 60
Suvarnarekha (India) 60
Tahar (Israel) 60
Taimour (Egypt) 61
Tommy Atkins (Florida, USA) 61
Totapuri (India) 61
Turpentine (West Indies) 62
Vallenato (Colombia) 62
Van Dyke (Florida, USA) 62
White Succari (Egypt) 62
Zebda (Egypt) 63
3.4 Conclusion 63
3.1 Introduction
The mango (Mangifera indica L.) has traditionally been grown in an area that
extends southwards and eastwards from India through Myanmar and Vietnam
to Indonesia. It probably is not indigenous to the Philippines, where it has long
been cultivated (Valmayor, 1962), but Mangifera species are endemic there (Bon-
dad, 1982). This crop is best adapted to a warm tropical monsoon climate, with a
pronounced dry season followed by rains. Fruit of the best quality is usually
produced in such areas, but specic races are known to fruit in humid regions.
For example, some Mangifera species bear dependably on the island of Borneo,
where most standard cultivars do not mature normal crops of fruit. Numerous
Mangifera species closely related to the common mango are indigenous to
R.J. Knight et al. 44
Borneo and nearby parts of Malaysia and Indonesia, and this region is the prob-
able centre of diversity for this genus (see Bompard, Chapter 2 this volume).
Most crops long cultivated over an extended time and area show consider-
able diversity, reecting different selection criteria in different regions of cul-
ture as well as genetic responses to varied environmental inuences. Certainly
this is true of the mango. Indian cultivars differ markedly from those grown
in South-east Asian countries and in Egypt. An additional factor that has
promoted genetic diversity in the group recently has been the widespread
introduction of this crop into new areas of cultivation, many in the western
hemisphere, over the last 500 years. In this manner, genetically diverse germ-
plasm has been brought from widely dispersed areas of the original range of
the species and grown in mixed plantings where, through the cross-pollination
natural to the species, new genetic combinations have been made and selected
under many varying conditions of microclimate. In Florida, since the late 18th
century, enough such importations and genetic recombinations have occurred
to qualify the southern part of this state as a secondary centre of diversity for
the crop. A new group of mango clones designated the Florida cultivars has
been exported to Brazil, Israel, Australia and other places where the process of
increasing diversity under new and varying cultural and environmental con-
ditions continues (Knight and Schnell, 1994; Schnell et al., 2006).
Some Florida cultivars, most notably Haden, have been important in aid-
ing the establishment of a modern mango industry in other parts of the world
(Knight and Schnell, 1994), and the phenomenon rst observed in Florida is now
occurring elsewhere; we are presented with the prospect of the importation of
cultivars of outstanding merit from their countries of origin to be grown, rst
experimentally and then commercially, in new regions. For this reason it is
important to become familiar with the characteristics of a group of cultivars that
currently are known in the commerce and/or horticulture of one or more coun-
tries, and that may have potential for expanded culture or use in breeding.
3.2 Criteria for Cultivar Description
In the recent past, efforts to assemble lists of mango descriptors produced
two publications that cover the subject (Mukherjee, 1985; IBPGR (now Inter-
national Plant Genetic Resources Institute, IPGRI), 1989) and provide people
who manage collections with morphological criteria to identify cultivars. The
Descriptor List used by IPGRI documents passport data (identifying the acces-
sion and information recorded by collectors), characterization (recording char-
acters considered to be highly heritable which can easily be seen in the eld
and are expressed in all environments) and preliminary evaluation, which records
a limited number of additional traits thought desirable by a consensus of users
of the crop. Plant data are important in preliminary evaluation, and include for
the tree, habit and height of the mature tree; for the leaf, shape, length and
width, and colour of the young leaf; and for the inorescence, position, shape,
density of owers, length, colour, hairiness, presence or absence of leafy bracts,
and percentage of owers in an average inorescence. (Some research indicates
Mango Cultivars and Descriptors 45
that both leafy bracts and number of perfect owers are inuenced by local
conditions and vary in their expression with differing environments.)
Additional plant data used in initial evaluation include those for the ower,
diameter in millimetres, type (pentamerous, tetramerous or both), nature of
disc (swollen, broader or narrower than ovary, reduced or absent) and number
of stamens; fruit, length, width and thickness, weight, shape, skin colour (which
may be compared with reference cultivars), skin thickness, skin texture, ratio
of pulp to skin and stone, texture of pulp, adherence of skin to pulp, bre in
pulp and its quantity and length, and stem insertion; and stone, length, weight,
veins and pattern of venation, presence or absence of bres and their length.
Additional plant data for leaves, inorescence and fruit have been col-
lected and some of these, notably season (maturity period), productivity, eat-
ing quality and attractiveness are quite important. Unfortunately, from the
viewpoint of those who expect to apply these criteria outside the Indian sub-
continent, reference cultivars are for the most part Indian and many are not
readily available outside India. Other important characters that have been
evaluated or proposed for evaluation include susceptibility to stress (drought,
wind, ooding), susceptibility to specic diseases and pests, molecular
markers, cytological characters and identied genes. Because of the extreme
comprehensiveness of this list and the limited availability of many of the
proposed descriptor evaluations at this time, we have tried to utilize such
information as is available to make the comparison, identication and evalu-
ation of specic well-known cultivars a practical possibility.
3.3 Mango Cultivars
A list of mango cultivars that are of interest in areas other than their places of
origin, with descriptions intended to help differentiate them, follows (see
Plates 440). Spelling and name variants in some cases represent efforts to
transliterate from other orthographies to the Roman alphabet, and in others
reect regional differences in usage.
Alfa (Brazil)
A monoembryonic cultivar developed by EMBRAPA Cerrado, Brazil, from
crossing Mallika Van Dyke. The tree is semi-dwarf in habit and high-
yielding, resistant to Oidium mangiferae and malformation, and moderately
resistant to anthracnose (Colletotrichum gloeosporioides); the fruit is large (435 g),
pink-red, rm, medium brous and of good quality (16% total soluble solids
(TSS), 0.23% acidity) (Pinto et al., 2004).
Alphonso (India)
Also known as Appus, Badami, Gundu, Haphus, Kagdi, Khader and
Khader Pasand. The tree is moderately large, with broadly rounded, dense
R.J. Knight et al. 46
canopy; the fruit (Plate 4) is yellow, ovate-oblique, averaging 6 cm long by
5 cm broad, weighing 225325 g (mean 226 g); the skin is thin; the esh is
rm to soft, low in bre, yellow, sweet with characteristic aroma and with a
very pleasant taste preferred by many who know this cultivar, bringing pre-
mium prices on Indian and international markets. Seed is monoembryonic in
a large, woody stone; the quality is excellent; ripening fruit in late to midsea-
son. Bearing is irregular, medium to heavy in India, but light and irregular in
Florida (Prasad, 1977; R.J. Knight, Jr, personal communication, 1995).
Amelie (West Africa)
Also known as Gouverneur in the Caribbean. The tree is tall with a rounded,
dense canopy; the fruit is green to orange-yellow with the advance of the season,
rounded, 1015 cm long by approximately 10 cm broad by approximately 7.8 cm
thick and weighing 300600 g (average 360 g). The skin is thick and separated
with difculty; the esh is soft, juicy, melting, without bre, a deep orange
colour, sweet and perfumed, free from turpentine, and provides the best of
mango tastes. Seed is monoembryonic in a medium-sized, elongate, narrow
stone that adheres to the esh, having a few short, pliable bres that are not
objectionable; the quality is excellent; the season early. The fruit closely resem-
bles that of Julie. Amelie is exported to France, along with Kent, from
Burkina Faso, Ivory Coast and Mali. Amelie is increasing in popularity on the
French market, chiey in Paris and the surrounding area. It brings lower prices
than cultivars with blushed fruit because the consumer is not always aware
when it is ripe (Naville, 1985, 1986; R. LePrette, personal communication, 1996).
Arumanis (Indonesia)
Also referred to as Harumanis. The tree is vigorous and tall with a slightly
open canopy. The fruit (Plate 5) is greenish yellow with large, light-yellow
dots, elongate oblong with rounded base, 1114 cm long by 6.67.5 cm broad
by 4.756.5 cm thick, weighing 200350 g. The skin is thick, tough and easily
separated, the esh rm and juicy with little bre, lemon yellow, sweet,
slightly insipid with a strong aroma, of poor to fair quality. Seed is polyem-
bryonic in a thick, woody stone; this cultivar ripens midseason and bears
regularly. Relatively easy to propagate by graftage, scionwood survives well;
widely planted in humid parts of the world where many better-quality culti-
vars fail to fruit (R.J. Knight, Jr, personal communication, 1995).
Ataulfo (Mexico)
A polyembryonic cultivar sold in North American markets under the name
Ataulfo and as Champagne. Originated in Tapachula, Chiapas, Mexico
reportedly from seed brought from Costa Rica in about 1930. The tree is
Mango Cultivars and Descriptors 47
vigorous and upright, a mid-range producer with production averages of
1020 t/ha possible. The tree is not highly adaptive to different climatic/
edaphic conditions. It is moderately resistant to anthracnose disease. The
fruit (Plate 6) is small (200300 g), elongate, of good quality, sweet with slight
acidity, yellow, rm, standing shipping stress well, and ripens from early to
midseason (Campbell et al., 2002; Magallanes-Cedeo, 2004).
B74 (Calypso) (Australia)
A monoembryonic cultivar that originated from the controlled cross of Sensa-
tion Kensington. The tree is upright, with low to moderate vigour and is
highly productive, with good tolerance of ower and fruit diseases; the fruit
(Plate 7) is moderately large (457.4 38.1 g), ovate (10.12 0.27 cm long by
9.13 0.28 cm wide), bre-free and rm, bright yellow overlaid with red blush,
with extended shelf life and potential for shipment to overseas markets; ripens
late in the season; patented (Whiley, 2001; Whiley and Hofman, 2006).
Banganpalli (India)
Also called Beneshan and Chappatai. The tree is medium sized with a
rounded canopy; the fruit is primrose-yellow, ovate-oblique, large and the
skin smooth, thin and shiny, esh rm to meaty with juice moderately abun-
dant, without bre, maize-yellow, with pleasant aroma and sweet taste. Seed
is monoembryonic, in an oblong stone covered with sparse bres; quality
good; ripens midseason and bears heavily (Singh, 1960).
Beta (Brazil)
A cultivar developed by EMBRAPA Cerrado, Brazil, from crossing Amra-
pali Winters (M20222 United States Department of Agriculture (USDA)).
The tree is moderately vigorous and free of malformation, high-yielding but
irregular, moderately resistant to anthracnose and Oidium; the fruit is small
(310 g), yellow, rm with low bre, of excellent quality (24.8% TSS, 0.16%
acidity) (Pinto et al., 2004).
Bombay Green (India)
Also called Bhojpuri, Bombai, Hiralal Bombai, Kali Bombai, Laile Alipur,
Malda, Sarauli and Sheeri-Dhan. The tree is tall and erect; the fruit (Plate
8) is apple green with ochre blush at the base and on some exposed parts,
dots abundant, with brown specks in the middle, ovate with beak almost
missing, medium sized, with tough, thick, non-adhering smooth skin; the
esh is cadmium-orange, rm and juicy with scanty bre just under the skin,
R.J. Knight et al. 48
very sweet with pleasant aroma, of very good quality; seed is monoembryonic
in a full, thick, medium-sized stone. This cultivar ripens early in the season
and is a medium bearer. Bombay Yellow is said to be practically identical to
this cultivar but for a slight difference in fruit colour. The present Bombay
Green is said to be a degenerate form of the original one (Singh, 1960). In
Jamaica it is sometimes called Peter, which suggests a confusion with Pairi,
but the Jamaican Peter is without the bright red blush normal to Pairi.
Cambodiana (Vietnam)
Also known as Xoai Voi. The tree is moderately vigorous, with a dense,
rounded canopy; the fruit (Plate 9) is greenish yellow, unblushed with a few
small white dots, oblong to ovate, 911.5 cm long by 6.57.5 cm broad by
56 cm thick, weighing 220340 g; the skin is thin, tender and adherent; the
esh contains little bre, is tender and melting, lemon yellow, sweet and
mildly subacid with a pleasant aroma; the seed is polyembryonic in a thick,
woody stone. Ripens early in the season. Brought to Florida in 1902, where it
gave rise to the Saigon landrace (Campbell, 1992).
Carabao (Philippines)
The tree is vigorous, forming a large and dense canopy; the fruit (Plate 10) is
greenish to bright yellow, brushed with a few small green dots, long and
slender, with base rounded to slightly attened, 1113 cm long by 6.57 cm
broad by 66.5 cm thick, weighing 270440 g; the skin is thick, medium tough
and easily separated; the esh is without bre, tender and melting, lemon
yellow, spicy and sweet with a mild aroma, of good to excellent quality; seed
is polyembryonic in a thin, papery stone. Ripens early in the season (Camp-
bell, 1992). This is a heavy bearer that may alternate; however, it can be
induced to fruit by potassium nitrate treatment in the tropics (Bondad and
Linsangan, 1979). It is highly resistant to bacterial black spot (Xanthomonas
campestris pv. mangiferaeindicae) in Queensland (Mayers et al., 1988). It was
introduced to Florida in 1909. Carabao is important in commerce between
the Philippines and Japan and is increasingly imported into the USA.
Chausa (India)
Also called Samar Bahisht Chausa and Khajari. The tree is tall and spread-
ing; the fruit is canary yellow to raw sienna when fully ripe, with numerous
obscure medium-sized dots with minute specks inside them, oblong with
prominent beak, obtuse to rounded, medium sized; the skin is thin and some-
what adhering, pulp raw sienna, soft and juicy with scanty ne, long bres
near the skin; the fruit is very sweet with a luscious, delightful aroma, of
excellent quality; seed is monoembryonic in a thick, medium-sized oblong
Mango Cultivars and Descriptors 49
stone with ne, short bres all over the surface and a tuft of long bres on
the ventral edge. Ripens late in the season and is a light bearer (Singh, 1960).
Cogshall (Florida, USA)
A monoembryonic cultivar that originated on Pine Island in Lee County. The
tree is relatively small, forming a rounded canopy, moderately susceptible to
anthracnose and consistently productive; the fruit is medium to large, aver-
aging about 350 g, yellow with a bright crimson blush, oblong (1114 cm long
by 7.58.5 cm broad by 6.28 cm thick) of excellent quality, rich and sweet in
taste, with tender skin and soft esh. Ripens early to midseason over about 4
weeks, a season longer than some cultivars. It is recommended for the home
garden, not commercial planting, in Florida but is now grown commercially
on Mauritius and marketed in France. Seedling of Haden (Campbell and
Campbell, 1995; Schnell et al., 2006).
Corao de Boi (Brazil)
The tree is vigorous, precocious and productive; the fruit is greenish yellow and
intense red on the side exposed to the sun, cordiform, medium sized, pulp yel-
low and brous. The seed is polyembryonic. There are two seasons in So Paulo,
JanuaryFebruary and SeptemberDecember. This is one of the best-known
commercial cultivars in So Paulo state (Sampaio, 1980; A.C. Pinto, personal
communication, 1996; L.C. Donadio, personal communication, 1996).
Dasheheri (India)
Also known as Dasheri and Aman Dusehri. The tree is of medium height
and moderate vigour, spreading, with a rounded, medium-dense canopy; the
fruit is primrose to canary yellow with abundant light-yellow dots, oblong to
oblong-oblique with base rounded to obliquely rounded, medium sized, skin
smooth, medium thick, tough and non-adhering; the esh is yellow, rm,
with almost no bre, scanty juice and a delightful aroma, very sweet taste, of
excellent quality; seed is monoembryonic in a thick, medium-sized stone. Rip-
ens midseason and is heavy bearing; fruit keeps well (Singh, 1960).
Espada (Brazil)
The tree is tall and develops rapidly, with a dense canopy, very productive;
the fruit is intense green or greenish yellow, oblong-elongate with a concave
base, medium sized, with smooth, thick skin; the esh has much bre, is egg-
yellow, with a strong aroma of turpentine. The quality is considered good for
fresh consumption. The polyembryonic seed is in an oblong stone, covered
R.J. Knight et al. 50
with soft bres and many nerves. There are two seasons per year in So
Paulo, JanuaryFebruary and NovemberDecember (Sampaio, 1980; A.C.
Pinto, personal communication, 1996).
Ewais (Egypt)
A polyembryonic cultivar of major commercial importance. The tree is vigor-
ous, the fruit small (275 g), yellow with no blush, with small, light-brown
slightly corky dots, ovate-oblong in shape (11.7 cm long by 7.2 cm wide by
6.3 cm thick), with adherent skin of intermediate thickness, relatively free of
disease; esh orange, juicy but susceptible to jelly seed, with no objectionable
bre, sweet and agreeable in taste, of very good quality. The stone is large
(38.5 g). Fruit ripens midseason (Knight and Sanford, 1998). In warm subtrop-
ics this cultivar has shown a tendency for owering in the warm season, with
fruit ripening during the cool winter. It has good anthracnose tolerance.
Excellent Succari (Egypt)
A polyembryonic mango of minor commercial importance. The tree is vigor-
ous, ripening fruit in late midseason. The fruit is small (280 g), green with a
yellow overlay and small, yellow smooth dots, ovate-oblong in shape (11 cm
long by 7 cm wide by 6.4 cm thick), with non-adherent skin of intermediate
thickness quite free of surface disease; the esh is orange, melting (without
jelly seed) and juicy with no objectionable bres, a delightfully sweet taste
and excellent quality; stone large (36.6 g) (Knight and Sanford, 1998). It has
moderate to good anthracnose tolerance in the warm subtropics.
Extrema (Brazil)
The tree is upright, vigorous and productive. The fruit is yellow with green-
ish areas, ovate-reniform, weighing 350400 g, with smooth and thin skin,
and yellow, watery esh with almost no bres with a moderately resinous,
agreeable taste. The quality is considered good for fresh consumption and
processing. The polyembryonic seed is in a large, brous stone. Ripens early
in the season (Sampaio, 1980; A.C. Pinto, personal communication, 1996).
Fajri (India)
Also spelled Fazli. The tree is of medium size and moderately vigorous,
with rounded, open canopy. The fruit is light chrome yellow with small,
dark-coloured fairly sparse dots, obliquely oval with base slightly rounded
and beak distinct to slightly prominent, large (averaging 14.3 cm long by
9.8 cm broad, weighing 500 g on average) with a medium-thick skin that is
Mango Cultivars and Descriptors 51
smooth with some inclination to be warty, and rm to soft, breless esh of
a light cadmium yellow with a pleasant aroma and a sweet taste, having juice
that may be scanty to moderately abundant, of good to very good quality.
The seed is monoembryonic in a large, oblong stone that is covered with a
sparse, short and soft bre. Ripens midseason to late (Gangolly et al., 1957; N.
Balasundaram, India, personal communication, 1990).
Fernandin (India)
The tree is moderately vigorous with a dense, rounded canopy; the fruit is
bright yellow with an attractive bright-red blush, ovate-oblique, averaging
12.2 cm long by 8.5 cm broad, weighing 450 g; the skin is rough and warty,
thick and adherent, esh bright yellow, moderately to abundantly juicy,
thick, with no objectionable bre, with delightful to piquant aroma and sweet
to very sweet, delicious taste, of superior quality; seed is monoembryonic;
season late (Gangolly et al., 1957; Singh, 1960).
Genovea (Egypt)
A polyembryonic cultivar of minor commercial importance. The fruit is small
(234.5 g), green with a yellow overlay and medium-sized smooth yellow
dots, ovate-oblong in shape (11 cm long by 6 cm wide by 5.6 cm thick), a thin
adherent skin relatively free of surface disease; esh orange, rm (no jelly
seed) and juicy with no objectionable bres, a sweet agreeable taste of ac-
ceptable quality; stone large (53 g) (Knight and Sanford, 1998).
Glenn (Florida, USA)
The tree is moderately vigorous, small to medium with dense, rounded can-
opy of compact growth; the fruit (Plate 11) is bright yellow with orange-red
blush, with numerous small yellow and white dots, oval to oblong with a
rounded base, 9.512.5 cm long by 7.58.5 cm broad by 78 cm thick, weigh-
ing 400620 g; the skin is thin, tough and easily separated, esh soft and juicy,
with little bre, deep yellow, rich and spicy with a strong, pleasant aroma, of
excellent quality; seed is monoembryonic in a thick, woody stone. Ripens
early in the season and is a regular bearer. This is a seedling of Haden
(Campbell, 1992; Schnell et al., 2006).
Golek (Indonesia)
The tree is moderately vigorous with an upright, open canopy; the fruit
(Plate 12) is greenish yellow with an orange overlay and prominent white
dots, oblong with rounded base, 9.512.5 cm long by 68 cm broad by 5.5
6.5 cm thick, weighing 200365 g; the skin is thin, tough and easily separated;
R.J. Knight et al. 52
the esh is soft and juicy with abundant bre (not objectionable), deep yel-
low, sweet, insipid with a mild aroma, of poor to fair quality; the seed is
polyembryonic in large, woody stone with abundant ne bre. Ripens mid-
season (R.J. Knight, Jr, personal communication, 1995).
Haden (Florida, USA)
The tree is vigorous, with a large, spreading canopy; the fruit (Plate 13) is
bright yellow with a deep crimson or red blush and numerous large yellow
dots, oval with a rounded base, 10.514 cm long by 910.5 cm broad by 8.5
9.5 cm thick, weighing 510680 g; the skin is thick, tough and adherent; the
esh is rm and juicy with abundant bre, deep yellow, rich and sweet with
a pleasant aroma, of good to excellent quality; the seed is monoembyonic in
a medium-thick woody stone. Ripens early to midseason and bearing is
sometimes irregular. This is a seedling of Mulgoba Turpentine and is the
rst of the Florida mango cultivars, introduced in 1910 and since grown in
many other countries. It is the seed parent of numerous other cultivars
(Campbell, 1992; Knight and Schnell, 1994; Schnell et al., 2006).
Himsagar (India)
The tree is vigorous, tall, with a dense, spreading canopy; the fruit (Plate 14)
is greenish yellow to bright yellow with no blush, with light-yellow dots,
ovate with a attened base, 1215 cm long by 8.59.5 cm broad by 7.58.5 cm
thick, weighing 465585 g; the skin is thin, tough and easily separated; the
esh is rm and juicy with no bre, orange, rich and sweet with a mild aroma,
of good to excellent quality; the seed is monoembryonic in a thick, woody
stone. This is a late midseason cultivar that bears well (R.J. Knight, Jr, per-
sonal communication, 1995).
Hindi Besennara (Egypt)
A polyembronic cultivar of major commercial importance. The tree is of me-
dium vigour, ripening fruit early to midseason. The fruit (Plate 15) is of small
to medium size (319 g), green with orange overlay, with small white corky dots,
oblong-cylindrical in shape (15.4 cm long by 6.7 cm wide by 6.5 cm thick) with
thick, non-adherent skin relatively free of surface disease; the esh is orange,
yielding and juicy with no objectionable bres, pleasantly sweet in taste, of
very good quality; the stone is large (47.2 g) (Knight and Sanford, 1998).
Hindi Khassa (Egypt)
A polyembryonic cultivar of major commercial importance. The tree is vig-
orous, ripening fruit in late midseason. The fruit is of medium size (461 g),
Mango Cultivars and Descriptors 53
yellow with no blush, with intermediate-sized smooth, light-yellow dots,
oblong-cylindrical in shape (16 cm long by 6.6 cm wide by 6.9 cm thick), with
thick, adherent skin relatively free of surface disease; esh is orange, rm
and juicy with no objectionable bres, of mediocre taste and a quality not
suitable for export; stone is large (55 g) (Knight and Sanford, 1998).
Irwin (Florida, USA)
The tree is small to medium, moderately vigorous, with open canopy. The fruit
(Plate 16) is bright yellow with a crimson or bright red blush, numerous large
white dots, ovate with rounded base, 11.513 cm long by 89 cm broad by 6.5
7.5 cm thick, weighing 340450 g; the skin is medium-thick, tender and adherent;
the esh is soft, tender, melting and juicy without bre, lemon yellow, sweet and
mild with a pleasant aroma, of good quality; the seed is monoembryonic in a
thin, papery stone. The stone may be seedless following cool weather at ower-
ing time. This is an early, regular and heavy bearer. The fruit is usually soft with
a short postharvest life, but it is often exported from tropical America to Europe.
It is a seedling of Lippens Haden (Campbell, 1992; Schnell et al., 2006).
Julie (West Indies)
Also called St Julienne. The tree is compact (dwarf), with a dense canopy;
the fruit (Plate 17) is greenish yellow with a light pink to maroon blush and
numerous small white dots, rounded with attened apex, pronouncedly
compressed laterally, 79.5 cm long by 47.5 cm broad by 25.5 cm thick,
weighing 200325 g with a thin, tender skin and soft, melting, juicy, orange
esh with scanty bre, of a rich, spicy avour with a strong, pleasant aroma,
of good quality; seed is monoembryonic in a thin, papery stone. This cultivar
ripens midseason and is a regular producer of small crops. The fruit is often
severely infected with anthracnose disease, but its unique taste is preferred
by many West Indians, and it is exported to the London market (C.W. Campbell,
personal communication, 1996).
Keitt (Florida, USA)
The tree is medium sized, moderately vigorous, upright with open canopy;
the fruit (Plate 18) is greenish yellow, with a pink or red blush, numerous
small white or yellow dots, oval, with rounded base, 1315 cm long by
911 cm broad by 8.510 cm thick, weighing 5102000 g; the skin is thick,
tough and adherent; the esh is rm and juicy, with little bre, lemon yellow,
sweet and mild with a pleasant aroma, of good to excellent quality; the seed is
monoembryonic in a thick and woody stone. This cultivar ripens late in the
season. It is a seedling of Brooks. After Tommy Atkins it is the most com-
mercially important cultivar in the export mango industry of the western
R.J. Knight et al. 54
hemisphere. It is resistant to anthracnose disease, packing and shipping stress
and is heavily productive (Campbell, 1992; Schnell et al., 2006). It is highly sus-
ceptible to bacterial black spot in Queensland (Mayers et al., 1988).
Kensington (Australia)
Also known as Kensington Pride and Bowen. Kensington has a large, vig-
orous tree with spreading canopy; the fruit (Plate 19) is yellow with an orange-
red blush on the shoulder, round ovate with a attened base and a slight beak,
10.513 cm long by 8.59.6 cm broad by 7.58.5 cm thick, weighing 350750 g;
the skin is thick, tender and adherent; the esh is soft and juicy, with moderate
to little bre, sweet with a characteristic avour that makes it the most popu-
lar cultivar in Australian markets, of excellent quality; seed is polyembryonic
in a moderately thick, woody stone. This cultivar ripens midseason and it
bears well. It is unusually susceptible locally, in Florida, to damage by red-
banded thrips (Selenothrips rubricinctus (Giard.)), and may be killed by this pest
without adequate countermeasures (R.J. Campbell, personal communication,
1994; R.J. Knight, Jr, personal communication, 1995). It is moderately suscep-
tible to anthracnose and bacterial spot (Mayers et al., 1988).
Kent (Florida, USA)
The tree is large and vigorous with a dense, upright canopy; the fruit (Plate
20) is greenish yellow with a red or crimson blush, numerous small yellow
dots, oval, with rounded base, 1113 cm long by 9.511 cm broad by 9.9.5 cm
thick, weighing 600750 g; the skin is thick, tough and adherent; the esh is
rm, tender, melting and juicy with little bre, deep yellow to orange-yellow,
sweet with a rich avour and pleasant aroma, of excellent quality; the seed
is monoembryonic in a thick, woody stone. Fruit ripens late midseason to
late and bearing may be alternate. It is a seedling of Haden Brooks, which
is a seedling of Totapuri (Sandersha) (Schnell et al., 2006). Kent is not
commonly commercial in Florida because it is prone to storage disease, but
is a successful commercial cultivar in drier parts of Mexico, Central and
South America and West Africa (Campbell, 1992). It is highly susceptible to
bacterial black spot in Queensland (Mayers et al., 1988).
Khanefy (Egypt)
A cultivar of minor commercial importance. The fruit is large (475 g), green
with a yellow overlay and large, brown, smooth dots, ovate in shape (10.7 cm
long by 8.3 cm wide by 8.6 cm thick), with an adherent skin quite free of sur-
face disease; the esh is yellow, often with jelly seed, juicy, with no objection-
able bres and a bland avour unacceptable to many Western palates. The
stone is moderately large (53 g) (Knight and Sanford, 1998).
Mango Cultivars and Descriptors 55
Kyo Savoy (Thailand)
The tree is large, vigorous, with an open canopy made up of long branches;
the fruit is green when harvested (before the ripening process begins) turn-
ing to greenish yellow, oblong, 11.512.5 cm long by 5.56.5 cm broad by
56 cm thick, weighing 230340 g; the skin is thin, tender and adherent; the
esh is medium rm, tender and not very juicy with no bre, pale yellow,
very sweet with an insipid taste and a mild, pleasant aroma, of fair to good
quality; the seed is highly polyembryonic in a medium-thin stone. This is a
regular producer (C.W. Campbell, personal communication, 1995). The fruit
is often consumed green.
Langra (India)
Also called Darbhanga, David Ford, Hadialaziz, Hajipur Langra, Har-
doi Langra, Lan Garhi, Langra Faquirwala, Sylhet and Tikari. The tree
is moderately vigorous, forming a dense canopy; the fruit is greenish yellow
with medium to big dark-green dots, ovalish to oblong, 810.5 cm long by
6.57.5 cm broad by 67 cm thick, weighing 235375 g; the skin is medium
smooth, thick; the esh is rm to soft, breless, lemon yellow, very sweet
with a strong, pleasant aroma, juice moderately abundant; seed is monoem-
bryonic in a medium-sized, attened stone covered with dense, short and
soft bre; quality is very good. Fruit ripens early to midseason (Gangolly et al.,
1957; R.J. Knight, Jr, personal communication, 1995).
Mabrouka (Egypt)
A major commercial cultivar considered to have been originally introduced
from India. The tree is moderately vigorous; the fruit (Plate 21) is large (481 g),
yellow with an orange to red blush and small, light-yellow, smooth dots;
ovate-oblong in shape (13.7 cm long by 8.9 cm wide by 8.2 cm thick) with a
thick, non-adherent skin relatively free of surface disease; the esh is yellow,
rm and juicy with no objectionable bre, moderately agreeable in taste, of
acceptable quality. The monoembryonic seed is in a moderately large (51 g)
stone. Fruit ripens late midseason, ships well and has been marketed in
Poland (Singh, 1960; Knight and Sanford, 1998).
Madame Francis (Haiti)
The tree is moderately vigorous, medium sized, forming an open canopy; the
fruit (Plate 22) is greenish to bright yellow, with no blush and a few large rus-
set dots, oblong, sigmoid with rounded base, 1517 cm long by 8.511 cm
broad by 5.57.5 cm thick, weighing 370520 g; the skin is thin, tender and
adherent; the esh is soft and juicy with medium bre, orange, rich spicy and
sweet with a pleasant aroma, of fair to good quality; seed is polyembryonic
R.J. Knight et al. 56
in a thin, papery stone. This cultivar ripens early to midseason and bears
well. Shipped to North American markets from Haiti nearly 10 months of the
year (R.J. Campbell, personal communication, 2007).
Mallika (India)
The tree is a moderately vigorous dwarf with a dense canopy; the fruit (Plate
23) is bright yellow with no blush and numerous small, light-yellow dots,
oblong with rounded base, 1012 cm long by 6.57.5 cm broad by 55.5 cm
thick, weighing 280450 g; the skin is thick, tough and easily separating; the
esh is soft, tender and juicy with little bre, deep yellow to orange, rich,
strongly aromatic and sweet, of excellent quality; seed is monoembryonic in
a medium-thick and woody stone. This cultivar ripens midseason and is an
irregular producer. This cultivar came from crossing Neelum and Dashe-
hari (Singh et al., 1972; Campbell, 1992).
Manila (Mexico)
The tree is large, vigorous, with an upright, open canopy; the fruit (Plate 24)
is bright yellow, sometimes with a light-pink blush, a few small reddish dots,
long and slender with rounded base and bluntly pointed apex sometimes
with a small beak, 12.514 cm long by 5.56 cm broad by 55.5 cm thick,
weighing 180260 g; the skin is thin, medium tough and easily separating;
the esh is medium rm and juicy, with little to abundant bre, deep yellow,
sweet, rich and spicy in taste with a pleasant aroma, of good to very good
quality; seed is polyembryonic in a medium-thick and woody stone. This
cultivar ripens early midseason and crops fairly dependably. For a long time
Manila has been the most popular mango in Mexico.
Manzanillo (Mexico)
The tree is large, of medium vigour with an upright canopy; the fruit is yel-
lowish orange with 75% of the surface blushed an intense dark red with
numerous dots, oval with moderately attened base, averaging 12 cm long
by 10 cm broad by 7.5 cm thick, and 660 g in weight; the esh is low in bre,
slightly subacid and very palatable, quality high; seed is monoembryonic in
a relatively small stone. This cultivar ripens early in the season but spread
over a 60-day harvest period. It bears heavily without pronounced alterna-
tion and the fruit stores and ships well (Nez-Elisea, 1984).
Mesk (Egypt)
A major commercial cultivar. The tree is vigorous, the fruit small to medium
sized (312.5 g), yellow with a red blush, with small, corky yellow dots;
Mango Cultivars and Descriptors 57
ovate-oblong (11.3 cm long by 7.4 cm wide by 6.5 cm thick) with adherent
skin intermediate in thickness and fairly free of surface disease; the esh is
orange, frequently jelly-seeded, with no objectionable bres and a sweet,
agreeable taste of very good quality. The polyembryonic seed is in a moder-
ately large (52.5 g) stone. Fruit ripens late in the season (Knight and Sanford,
1998).
Mulgoa (India to Florida, USA)
Also spelled Mugoba and Mulgova. The tree is large, vigorous with open,
spreading canopy; the fruit (Plate 25) is bright yellow with a pink blush and
numerous large white dots, oval to ovate with attened base, 8.510.5 cm
long by 6.57.5 cm broad by 56 cm thick, weighing 340450 g; the skin
is thick, medium tough and adherent; the esh is soft, tender, melting and
juicy, with little bre, lemon yellow, rich spicy and sweet with strong, pleas-
ant aroma, of good to excellent quality; seed is monoembryonic in a thick,
woody stone. This cultivar ripens midseason to late and is a shy, irregular
bearer. Introduced to Florida in 1889 and called Mulgoba, this is the seed
parent of Haden, rst of a series of cultivars known as the Florida group. A
question exists whether the cultivar known in Florida is identical with the
Indian cultivar or is a seedling rootstock that survived after the scion was
killed by cold. In either case its superior quality ensured its retention and
propagation (Campbell, 1992). Literature serves to compound the nomencla-
tural confusion, as illustrated by Gangolly et al. (1957) whose Mulgoa fruit,
yellow overall and roundish oblique with a deeply depressed stem insertion,
does not resemble the cultivar introduced to Florida. Singh (1960), on the
other hand, portrays a rounded, lightly blushed greenish yellow fruit that
closely resembles the Florida mango. Furthermore, vegetative propagation
of selected chance seedlings has resulted in a variety of clonal types carried
under this name in India (Ratnam and Chellapa, no date, post-1954).
Nabeel (Egypt)
A minor commercial cultivar. The fruit is large (495 g), green with small yel-
low dots that are smooth; ovate-oblong (14 cm long by 9 cm wide by 7 cm
thick), with adherent skin relatively free of surface disease; the esh is
orange, rm and juicy, without objectionable bre, with a passable but not out-
standingly pleasing taste and acceptable quality. The seed is polyembryonic
in a large (56.6 g) stone (Knight and Sanford, 1998).
Nam Doc Mai (Thailand)
The tree is vigorous, medium sized with upright, dense canopy; the fruit
(Plate 26) is greenish to bright yellow with a slight pink blush and numerous
R.J. Knight et al. 58
small green dots, long and slender, sigmoid in shape with a rounded base,
1719 cm long by 7.58.5 cm broad by 6.57.5 cm thick, weighing 340580 g;
the skin is medium thick, tender and easily separated from the esh which is
soft, tender and juicy with no bre, lemon yellow, rich, spicy and very sweet
with pleasant aroma, of excellent quality; seed is polyembryonic in a thin,
papery stone. This cultivar ripens early midseason, fruits regularly and may
have multiple crops in one season (Campbell, 1992). It is highly resistant to
foliar infection, and resistant to fruit infection by bacterial black spot in
Queensland (Mayers et al., 1988).
Neelum (India)
The tree is moderately vigorous with a small, compact canopy; the fruit is
bright yellow with no blush and numerous small white dots, oval with at-
tened or slightly rounded base, 9.511 cm long by 7.58.5 cm broad by
66.5 cm thick, weighing 230300 g; the skin is thick, tender and easily sepa-
rating; the esh is soft, melting and juicy with no bre, deep yellow, mild and
sweet with a delightfully pleasant aroma, of good to excellent quality; seed is
monoembryonic in a medium-thick, woody stone. This cultivar is a late,
heavy bearer (Campbell, 1992).
Nuwun Chan (Thailand)
The tree is moderately vigorous, small, upright with a dense canopy; the fruit
(Plate 27) is greenish yellow with a pink to red blush, numerous small green
dots, long and slender with a attened base, 1618 cm long by 78 cm broad
by 66.5 cm thick, weighing 340500 g; the skin is thick, tough and easily
separating; the esh is soft, melting, juicy with little bre, pale yellow, mild
and sweet with a faint, pleasant aroma, of good eating quality; seed is poly-
embryonic in a thick, woody stone. This cultivar is an early, regular bearer.
Fruit is often eaten green (Campbell, 1992).
Okrung (Thailand)
The tree is moderately vigorous, medium sized and upright, forming a dense
canopy; the fruit (Plate 28) is green to greenish yellow with no blush and
numerous small white dots, oblong and sigmoid with a rounded base,
1113 cm long by 56 cm broad by 4.55.5 cm thick, weighing 160240 g; the
skin is thick, tough and medium adherent; the esh is soft and juicy with
abundant bre, yellow or greenish, mild, somewhat insipid and very sweet
with a pleasant aroma, of good quality; seed is polyembryonic in a thick,
woody stone. This cultivar ripens midseason, is a heavy producer and some-
times bears more than one crop/year (Campbell, 1992).
Mango Cultivars and Descriptors 59
Osteen (Florida, USA)
The tree is vigorous, medium sized, forming a dense canopy; the fruit (Plate
29) is yellow-orange with a purple or lavender blush and numerous small
white dots, oblong with rounded base, 1215.5 cm long by 910.5 cm broad by
8.69.5 cm thick, weighing 500760 g; the skin is thick, tough and easily sepa-
rating; the esh is rm and juicy, with little bre, lemon yellow, mild and sweet
with a pleasant aroma, of good quality; seed is monoembryonic in a thick and
woody stone. This cultivar ripens late midseason to late and is a regular
producer. It is a Haden seedling (Campbell, 1992; Schnell et al., 2006).
Pairi (India)
Also written Pairie, Paheri and Pirie; synonyms are said to be Peter,
Peter Pasand, Grape, Gohabunder, Nadusalai, Rasjuri and Yerra Goa.
The tree is moderately vigorous, forming a dense, rounded canopy; the fruit
(Plate 30) is medium sized, green to yellow with a bright red blush, roundish,
skin smooth, thick, esh golden yellow, slightly juicy, breless, with a deli-
cious subacid taste, of excellent quality; the thick stone covered with short,
bristly bre encloses monoembryonic seed (Popenoe, 1927; Singh, 1960). This
cultivar has long been popular as a dooryard fruit tree in Hawaii.
Palmer (Florida, USA)
The tree is moderately vigorous, forming a large, upright, tight canopy; the
fruit (Plate 31) is yellow-orange with a dark-red to crimson blush and a few
small white dots, oblong with rounded base, 1215 cm long by 8.510 cm
broad by 6.57.5 cm thick, weighing 510850 g; the skin is medium thick,
tough and adherent; the esh is rm and melting with little bre, orange-
yellow, mild and aromatic, of good quality; seed is monoembryonic in a
medium-thick woody stone. This is a late midseason cultivar and is a regular
bearer. It is a seedling of Haden (Schnell et al., 2006). In Florida it is of minor
commercial importance (Campbell, 1992). It is grown in Israel and is the seed
parent of Naomi. It is attracting increased attention in the western hemi-
sphere export market as a result of its superior eating quality.
Rosa (Brazil)
The tree is medium sized, of slow growth with a rounded canopy; the fruit
(Plate 32) is yellow to rose-red on the side exposed to sun, oblong-cordiform
and medium sized; the skin is thick and smooth; the esh is rm and mod-
erately juicy, brous, golden yellow, moderately sweet with a turpentine
aroma, of ordinary quality, susceptible to anthracnose disease; the seed is
polyembryonic in a small, oblong stone. This cultivar ripens midseason to
R.J. Knight et al. 60
late. It is one of the most important commercial cultivars in the Federal Dis-
trict of Brazil, used for juice as well as fresh consumption, and is one of the
most well-known cultivars in Brazil (Sampaio, 1980; L.C. Donadio, personal
communication, 1996; A. Pinto, personal communication, 1996).
Sensation (Florida, USA)
The tree is vigorous, with a moderately open, symmetrical canopy; the fruit
(Plate 33) is dark yellow with a prominent dark-red to purple blush that cov-
ers most of its surface, oval with rounded base and rounded apex, 911.5 cm
long by 78 cm broad by 6.57 cm thick, weighing 280340 g; the skin is
medium thick, tough and easily separating; the esh is rm and medium
juicy, breless, deep yellow, mild and sweet with a weak, pleasant aroma,
of fair to good quality; seed is monoembryonic in a thick, woody stone.
This cultivar ripens midseason to late (Campbell, 1992). It is a seedling of
Haden Brooks (Schnell et al., 2006), and the seed parent of B74. It alter-
nates severely, and in on years the fruit may be clustered so heavily that it
becomes diseased before maturity, thus Sensation is not of commercial
importance. It is highly resistant to bacterial black spot in Queensland (May-
ers et al., 1988), but often has severe internal breakdown (browning, water
soaking) (A.W. Whiley, personal communication, 1996).
Suvarnarekha (India)
Also called Swarnarekha and Sundri. The tree is moderately vigorous and
tall, with a rounded, dense, spreading canopy; the fruit is light cadmium yel-
low with a blush of jasper red and abundant small, light-coloured dots, ovate
oblong with a base slightly attened, of medium size, 11 cm long by 8.2 cm
broad, weighing 400 g; the skin is medium thick, easily separated, esh soft,
breless, primrose yellow with a pleasant aroma, sweet taste and abundant
juice, of medium to good quality; seed is monoembryonic in an oblong-oval
stone covered with soft, short bre. Ripens early in the season early and is
heavy bearing (Gangolly et al., 1957).
Tahar (Israel)
The tree is vigorous, medium sized, with an upright, dense canopy; the fruit
is bright yellow with a dark-red blush and numerous small white dots, ovate
with attened base, 11.513 cm long by 8.99.5 cm broad by 7.58 cm thick,
weighing 360520 g; the skin is thick, tough and easily separating; the esh is
soft and juicy with little bre, deep yellow, mild, aromatic and slightly insipid
with a strong odour not appreciated by many, of fair to good quality; seed is
monoembryonic in a medium-thick woody stone. This cultivar ripens in late
midseason and bears well in Israel (Campbell, 1992).
Mango Cultivars and Descriptors 61
Taimour (Egypt)
A major commercial cultivar. The tree is vigorous; the fruit (Plate 34) is large
(500 g), ripening late in the season, dark green with large light-brown dots,
smooth in texture, ovate-oblong (12.8 cm long by 8.4 cm wide by 8 cm thick),
with non-adherent skin of intermediate thickness, quite free from surface
disease; esh is orange, rm (free of jelly seed) and juicy with no objection-
able bre, of a delightfully rich, sweet taste and excellent quality. The seed is
polyembryonic in a medium-sized (50.8 g) stone (Knight and Sanford,
1998).
Tommy Atkins (Florida, USA)
The tree is vigorous, with a dense, rounded canopy; the fruit (Plate 35) is
orange-yellow, with a crimson or dark-red blush and numerous small, white
dots, oval to oblong, with broadly rounded base, 1214.5 cm long by 1013 cm
broad by 8.510 cm thick, weighing 450700 g; the skin is thick, tough and
adherent; the esh is rm and medium juicy; with a medium amount of bre,
lemon to deep yellow, mild and sweet with a strong pleasant aroma, of fair to
good quality; seed is monoembryonic in a thick, woody stone. This cultivar
ripens early to midseason. It is a Haden seedling (Schnell et al., 2006).
Tommy Atkins is the most important commercial cultivar in the western
hemisphere export mango market; it is highly resistant to anthracnose dis-
ease and handling and shipping stress, and a consistent, heavy producer
(Campbell, 1992). Jelly seed (internal breakdown) is a serious problem in
the moist subtropics and tropics outside Florida, where the mango is grown
on calcareous, well-drained soil (A.W. Whiley, personal communication,
1996).
Totapuri (India)
Also called Bangalora, Collector, Kallamai, Killi (Gilli), Mukku, Sand-
ersha and Thevadiyamuthi. The tree is of medium size, vigorous, spread-
ing with an open canopy; the fruit (Plate 36) is greenish yellow with a pink
blush and a few small, white dots, oblong, base rounded, apex rounded to
bluntly pointed with a large beak, 17.520 cm long by 911.5 cm broad by
8.510.5 cm thick, weighing 8001100 g; the skin is thick, tough and adherent;
the esh is rm and medium juicy with a weak, somewhat repugnant aroma,
of poor to fair quality; seed is monoembryonic in a thin, papery stone. This
cultivar ripens late midseason, is productive and regular bearing. Fruit cracks
when exposed to heavy rains at ripening time. Totapuri was imported to
Florida twice, as Sandersha in 1901 and as Totapuri in the early 1960s. It is
the seed parent of Anderson and Brooks, which is itself the parent of
Kent. It is called Totapuri in Bangalore, and Bangalora in much of the rest
of India (Gangolly et al., 1957; Campbell, 1992).
R.J. Knight et al. 62
Turpentine (West Indies)
The tree is vigorous, with a large, spreading, rounded canopy; the fruit (Plate
37) is bright yellow with a few large white dots, occasionally with a pink
blush, oval with a attened base, 7.58.5 cm long by 6.57.5 cm broad by
66.5 cm thick, weighing 140200 g; the skin is thick, tough and easily sepa-
rating; the esh is rm and juicy, with abundant coarse bre, lemon yellow,
rich, aromatic, spicy, resinous and sweet with a strong, pleasant aroma, of
poor to fair quality; seed is polyembryonic in a thick, woody stone. This cul-
tivar ripens early midseason to late midseason and is a heavy producer but
may alternate. It is commonly used as a grafting stock (Campbell, 1992).
Vallenato (Colombia)
The tree is vigorous, with an upright, dense canopy; the fruit (Plate 38) is
bright yellow, with a crimson blush, oblong with attened base, 89 cm long
by 78 cm broad by 67 cm thick, weighing 195340 g; the skin is thin, tough
and adherent; the esh is rm, juicy with abundant ne bre (not objection-
able), pale yellow, mild and sweet with a strong, pleasant aroma, of good to
excellent quality; seed is monoembryonic. This cultivar ripens in early mid-
season (R.J. Campbell, personal communication, 1995).
Van Dyke (Florida, USA)
The tree is moderately vigorous, with a large, open canopy; the fruit (Plate
39) is bright yellow with a bright red or crimson blush, oval with rounded
base, 911.5 cm long by 7.59.5 cm broad by 78 cm thick, weighing 250
520 g; the skin is thick, tough and easily separating; the esh is quite rm,
melting and juicy with little bre, orange-yellow, rich, spicy and sweet with
a strong, pleasant aroma, of good to excellent quality, but susceptible to inter-
nal breakdown; seed is monoembryonic in a medium-thick, woody stone.
This cultivar ripens in late midseason and is a regular, heavy producer. It is a
seedling of Haden (Campbell, 1992; Schnell et al., 2006).
White Succari (Egypt)
A cultivar of major importance. The tree is vigorous; the fruit is medium
large (410 g), greenish yellow with yellow overlay and small, brown dots of
smooth texture, ovate-oblong in shape (11.25 cm long by 8.3 cm wide by
8.0 cm thick), with thin adherent skin reasonably free of surface disease;
esh is orange, yielding and juicy with no objectionable bres, of an agree-
able sweet taste and very good quality. The seed is polyembryonic in a
moderate to large-sized stone (49 g), the season early (Knight and Sanford,
1998).
Mango Cultivars and Descriptors 63
Zebda (Egypt)
A cultivar of major importance. The tree is vigorous and regularly produc-
tive; the fruit (Plate 40) is large (660 g), green with no overlay and small,
brown dots of smooth texture, oblong-cylindrical in shape (14.6 cm long by
9.7 cm wide by 8.3 cm thick), with non-adherent skin quite free of surface
disease; esh is deep orange, rm and juicy, with no objectionable bre and
a mild, sweet taste, of acceptable quality. The seed is polyembryonic in a
moderately small (52 g) stone. This cultivar is of late-midseason maturity
(Knight and Sanford, 1998). It is highly tolerant of anthracnose and resistant
to malformation (R.C. Ploetz, personal communication, 2007).
3.4 Conclusion
The mango fruits nutritional value, aesthetic and gustatory appeal have
assured its growing importance in non-traditional markets since the late
1950s, as it has been introduced to consumers previously unacquainted with
it. Furthermore, the migration of large populations from South-east Asia and
other regions where this fruit is a traditional crop to metropolitan centres
where it has not been well known has created a permanent demand for it in
these new markets. An additional factor permitting market expansion has
been the growing mango production in areas previously unimportant in
world commerce such as Mexico, Brazil, Australia, West Africa, Israel, Flor-
ida and the Canary Islands. The fact that most new markets are remote from
areas of production has necessitated selection of cultivars for fresh market
sale that are dependably productive and resistant to harvest, handling and
shipping stress, with relatively long shelf life, for example Tommy Atkins,
Keitt and Madame Francis. The fruit quality of mango cultivars well suited
to packing and shipping has been a secondary consideration, and is gener-
ally not so high as that of cultivars acknowledged to be superior for eating.
Economic factors obviously must dictate what is grown for the fresh market.
The commercial market for processed mango products permits other
cultivars to be utilized, and these may vary with the product that is mar-
keted. Cultivars chosen for pure or juice preparation are likely to be quite
different from those used for manufacture of chutney or other products
requiring pulp that maintains its integrity after it is cooked. Totapuri (Sand-
ersha) or Turpentine, for example, considered mediocre for fresh consump-
tion, can be used to prepare excellent chutneys, as can many criollo types in
the West Indies. Tommy Atkins makes outstandingly good dried fruit sec-
tions, sweet and aromatic, even though its fresh-fruit quality is generally
conceded not to be high. Mango butter and mango leather are other products
that are appreciated by many who know them (see Raymundo et al., Chapter
17, this volume; Campbell and Campbell, 1983; Campbell and Smith, 1987).
As more fruit that is wholesome, but not of export quality, becomes available
in areas of increasing production, it is likely that processed mango products
will become more common.
R.J. Knight et al. 64
Despite the recognized high quality of many well-known mango cultivars,
considerable cultivar improvement is still needed in most regions of culture
before anything approaching perfection is likely to be achieved (Table 3.1).
For any given area, cultivars that combine adequate resistance to disease and
packing and shipping stress, regular heavy production, high quality, and
attractive appearance throughout a long bearing season are all requisites.
Production of seedlings from controlled crossing of different parents having
desired characters, followed by vigorous selection and evaluation of the
resultant selections, can produce such improved cultivars. Pursuit of com-
mon goals, including the cooperative exchange and testing of elite germ-
plasm in different regions of production, can accelerate progress towards this
objective (Lavi et al., 1989, 1993; Knight, 1993). Such interregional and inter-
national activities are to be encouraged because of their potential for advanc-
ing mango production and utilization in the world.
Acknowledgements
For their help in reviewing portions of this chapter and/or contributing vast
quantities of information on mango cultivar descriptors and attributes, the
authors are profoundly grateful to the following: Dr N. Balasundaram, Head,
Sugarcane Breeding Institute, Regional Centre, Karnal, India 132001; the late
Dr Carl W. Campbell, Tropical Research and Education Center, 18905 SW 280
Table 3.1. Ratings of selected mango cultivars grown in Florida (Source: Knight, 1993).
Cultivar Shape
a
Size
b
Firmness
c
Colour
d
Anthracnose
e
Fibre
f
Taste
g
Yield
h
Score
i
Alphonso 3 5 7 2 3 7 9 1
h
x
Boribo 3 8 8 4 7 9 5 6 x
Carabao 5 6 7 3 5 9 8 6
h
x
Haden 3 9 8 8 5 7 7 3
h
x
Keitt 4 10 9 6 8 9 8 8 ///
Kensington 3 8 7 7 7 8 7 6 /
Langra 2 6 8 3 5 8 8 3
h
x
Pope 3 9 5 7 2 8 8 1 x
Tommy
Atkins
3 9 9 9 9 6 6 7 ///
Van Dyke 3 7 10 9 7 8 7 6 ///
a
Ratings of 1 (round) to 5 (long) indicate fruit shape, not its desirability.
b
Ratings below 6 justify discard; those of 7 and above show size only, not merit.
c
Ratings of 110 where 1 = least and 10 = most.
d
Ratings of 110 where 1 = least and 10 = most.
e
Ratings of 110 where 1 = most and 10 = least susceptible.
f
Ratings of 110 where 1 = most and 10 = least.
g
Ratings of 110 where 1 = worst and 10 = best.
h
Trends markedly towards alternate bearing.
i
One or more checks (/) show overall value; (x) indicates no commercial acceptability.
Mango Cultivars and Descriptors 65
Street, Homestead, Florida 33031 USA; Luis C. Donadio, Universidade Estad-
ual Paulista, Rodavia Carlos Tonanni KM5, Jaboticabal, 14870, So Paulo,
Brazil; Dr Shmuel Gazit, Department of Horticulture, Hebrew University of
Jerusalem, Rehovot 76100, Israel; Mr Remy LePrette, Directeur, Interfel, 155
Rue F.G. Poissonierre, Paris 75009, France; Dr Alberto C. Pinto, Lider de Pro-
jeto/CPAC, EMBRAPA, Caixa Postal 08 223, CEP 73301-970, Brasilia, DF,
Brazil; Dr Eli Tomer, Department of Fruit Trees, the Volcani Centre, Institute
of Horticulture, PO Box 6, Bet Dagan 50-250, Israel; Dr Anthony W. Whiley,
Maroochy Horticultural Research Centre, PO Box 5093 SCMC, Nambour,
Queensland 4560, Australia.
References
Bondad, N.D. (1982) Mango and its relatives in the Philippines. Philippine Geographic
Journal 26, 88100.
Bondad, N.D. and Linsangan, E. (1979) Flowering in mango induced with potassium
nitrate. HortScience 14, 527528.
Campbell, B.A. and Campbell, C.W. (1983) Preservation of tropical fruits by drying.
Proceedings of the Florida State Horticultural Society 96, 229231.
Campbell, B.A. and Smith, J. (1987) An overview of tropical fruit uses in Florida. Pro-
ceedings of the Florida State Horticultural Society 100, 408411.
Campbell, C.W. and Campbell, R.J. (1995) Cogshall, a mango for the home garden.
Proceedings of the Florida State Horticultural Society 108, 369370.
Campbell, R.J. (ed.) (1992) A Guide to Mangos in Florida. Fairchild Tropical Garden,
Miami, Florida, USA.
Campbell, R.J., Ledesma, N. and Campbell, C.W. (2002) Tropical Mangos: How to Grow
the Worlds Most Delicious Fruit. Fairchild Tropical Garden, Miami, Florida, USA.
Gangolly, S.R., Singh, R., Katyal, S.L. and Singh, D. (1957) The Mango. Indian Council
of Agricultural Research, New Delhi, India.
International Board for Plant Genetic Resources (IBPGR) (1989) Descriptors for Mango.
International Board for Plant Genetic Resources, Rome.
Knight, R.J., Jr (1993) Evaluating important fruit characters in mango germplasm. Fruit
Varieties Journal 47, 2530.
Knight, R.J. and Sanford, R.L. (1998) Mango Cultivar Evaluation. Publication No. 42.
(USAID (United States Agency for International Development) Project No. 2630240).
ATUT (Agricultural Technology Utilization and Transfer) /Ronco, Giza, Egypt.
Knight, R.J. and Schnell, R.J. (1994) Mango introduction in Florida and the Haden
cultivars signicance to the modern industry. Economic Botany 48, 139145.
Lavi, U., Tomer, E. and Gait, S. (1989) Inheritance of agriculturally important traits in
mango. Euphytica 54, 510.
Lavi, U., Sharon, D., Tomer, E., Adato, A. and Gazit, S. (1993) Conventional and modern
breeding of mango cultivars and rootstocks. Acta Horticlturae 341, 146151.
Magallanes-Cedeo, R. (2004) Area-wide assessment of the Ataulfo mango cultivation
in the Soconusco region of Chiapas, Mexico. Acta Horticulturae 645, 361363.
Mayers, P.E., Whiley, A.W., Hutton, D.G. and Saranah, J.B. (1988) Integrated control of
bacterial black spot (Xanthomonas campestris pv. mangiferaeindicae) of mango. 1.
Evaluation of 23 cultivars of mango for foliar and fruit resistance to bacterial black
spot under orchard conditions at Childers, south east Queensland. Maroochy Hor-
ticultural Research Report 5, 100101.
R.J. Knight et al. 66
Mukherjee, S.K. (1985) Systematic and Ecogeographic Studies of Crop Gene Pools. 1.
Mangifera L. International Board for Plant Genetic Resources, Rome.
Naville, R. (1985) Les fruits tropicaux et subtropicaux sur le marche franais en 1984.
Fruits 40, 345356.
Naville, R. (1986) Les importations franaises de fruits tropicaux et subtropicaux en
1985. Fruits 41, 409420.
Nez-Elisea, R. (1984) Manzanillo-Nez: a new Mexican mango cultivar. Proceed-
ings of the Florida State Horticultural Society 97, 360363.
Pinto, A.C.Q., Andrade, S.R.M., Ramos, U.H.V. and Cordeiro, M.C.R. (2004) Intervari-
etal hybridization in mango (Mangica indica L.): techniques, main results and
their limitations. Acta Horticulturae 645, 327330.
Popenoe, W. (1927) Manual of Tropical and Subtropical Fruits. Macmillan, New York.
Prasad, A. (1977) Bearing behaviour and fruit quality of south Indian varieties of mango
in northern India. Indian Journal of Horticulture 34, 372376.
Ratnam, L.V. and Chellappa, T. (no date, post-1954) Mulgoas of Hyderabad. In: Rao,
B.U. (ed.) The Mango a Souvenir. Department of Agriculture, Neo Silver Jubilee
Press, Hyderabad, India.
Sampaio, J.M.M. (1980) Caractersticas gerais de algumas cultivares e tipos de manguei-
ras no Brasil. In: Donadio, L.C. (ed.) Anais do I Simposio Brasileiro sobre a Cultura
da Mangueira. Departamento de Fitotecnia, Faculdade de Ciencias Agrarias e Vet-
erinarias, Universidade Estadual de Jaboticabal, So Paulo, Brazil, pp. 3550.
Schnell, R.J., Brown, J.S., Campbell, R.J., Kuhn, D.N., Meerow, A.W. and Olano, C.T.
(2006) Mango genetic diversity analysis and pedigree inferences for Florida culti-
vars using microsatellite markers. Journal of the American Society for Horticultural
Science 131, 214224.
Singh, L.B. (1960) The Mango: Botany Cultivation and Utilization. Leonard Hill, London.
Singh, R.N., Majumder, P.K., Sharma, D.K. and Mukherkjee, S.K. (1972) Some promis-
ing mango hybrids. Acta Horticulturae 24, 117119.
Valmayor, R. (1962) The Mango: its Botany and Production. University of the Philip-
pines, Laguna, the Philippines.
Whiley, A.W. (2001) Mango (Mangifera indica) B74. Plant Varieties Journal 14, 4546.
Whiley, A.W. and Hofman, P.J. (2006) Calypso Best Practices Manual. (On CD).
Horticulture Ltd, Nambour, Australia.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 67
4 Breeding and Genetics
C.P.A. Iyer
1
and R.J. Schnell
2
1
Indian Institute of Horticultural Research, Bangalore, India
2
USDA ARS, National Germplasm Repository, Miami, Florida, USA
4.1 Introduction 68
4.2 Origin of Cultivars and Distribution 68
Mangifera species and mango 68
History of cultivation 69
Impact of Florida mangoes 70
4.3 Reproductive Mechanisms 70
Polyembryony 70
Floral biology and pollination 71
Incompatibility 72
Cytology 72
4.4 Inheritance of Characters 73
Dwarfness, regular bearing and precocity 73
Flesh colour 74
Skin colour 74
Flowering time 74
Beak 74
Disease resistance 74
Other horticultural traits 75
4.5 Breeding Objectives 75
General objectives 75
Specic objectives 76
4.6 Methods of Breeding 78
Selection from open-pollinated seedlings 78
Controlled pollination 79
4.7 Handling of Hybrid Populations and Selection 80
Criteria for initial selection 80
Pre-selection 81
Potential for marker assisted selection (MAS) 81
Molecular markers 81
4.8 Minimizing Problems in Breeding 83
Heavy fruit drop 83
C.P.A. Iyer and R.J. Schnell 68
Long juvenile phase 83
Polyembryony 84
4.9 Achievements of Conventional Breeding 84
India 85
Other countries 86
4.10 Mutations 87
Somatic mutations 87
Induced mutations 88
4.11 Breeding Potential of Wild Species 88
4.12 Conclusions 89
4.1 Introduction
Mango has been considered to be a difcult plant species to improve in
breeding programmes because of certain inherent characteristics including:
(i) a long juvenile phase; (ii) a high level of heterozygosity resulting in unpre-
dictable outcomes in hybridization; (iii) only one seed per fruit; (iv) heavy
fruit drop leading to low retention of crossed fruits; (v) polyembryony in
many cultivars; and (vi) the large area required for a meaningful assessment
of hybrids. Despite these drawbacks, mango breeding can be successful
because of its wide range of genetic variation and the ease with which a
selected hybrid can be vegetatively propagated. Barring a few hybrid variet-
ies resulting from planned hybridization programmes, which are now gain-
ing increased attention, almost all known cultivars have resulted from the
selection of chance seedlings from natural cross-pollinations. However, in
Florida, a number of cultivars have resulted from the screening of seedlings
from known mother plants. Most of the present-day mango cultivars were
selected on the Indian subcontinent; these selections were made based
mainly on fruit quality, with very little emphasis on modern horticultural
and industrial requirements. These requirements include precocity, dwarf-
ness, heavy and regular bearing, absence of physiological disorders, resis-
tance to disease and pests and good shipping qualities. With decreasing land
availability and the rising cost of labour, tree architecture requirements have
also changed. The need for new cultivars to meet these requisites pinpoints
the importance of planned hybridization rather than merely depending on
chance seedlings. Current knowledge of hybridization techniques, inheri-
tance patterns, management of hybrid populations and the development of
genetic markers have greatly reduced the uncertainty in mango breeding.
4.2 Origin of Cultivars and Distribution
Mangifera species and mango
Almost all the commercial cultivars belong to Mangifera indica. However,
a few commercial varieties of South-east Asia belong to other species, i.e.
M. altissima, M. caesia, M. foetida, M. grifthi, M. odorata, M. pentadra, M. sylvatica,
Breeding and Genetics 69
M. zeylanica, M. laurina, M. lagenifera, M. cochinchinensis, etc. The monoem-
bryonic mango (M. indica) originated in north-eastern India (Assam), the
Indo-Myanmar border region and Bangladesh (Chittagong Hill tract), where
it is still found as a wild tree, with very small fruits. It may also occur in the
lower Himalayan tract, near Nepal, Bhutan and Sikkim. Polyembryonic
mangoes are considered to have originated in South-east Asia. Wild man-
goes, representing different Mangifera species, can be found in tropical Asia,
particularly north-eastern India, Sri Lanka, Myanmar, Thailand, Indo-China,
southern China, Malaysia, Indonesia, Papua New Guinea, the Philippines
and as far as the Solomon and Caroline Islands in the east. There are more
than 60 species worldwide. The highest specic diversity is found in the
heart of the distribution area of the genus Mangifera; the Malay Peninsula,
Borneo and Sumatra (Bompard, 1993).
History of cultivation
Mango has been cultivated in India for at least 4000 years and over 1000 vari-
eties are recognized there (Mukherjee, 1953). Almost all of them are selec-
tions made from naturally occurring, open-pollinated seedlings. However,
based on random amplication of polymorphic DNA (RAPD) analysis, Rav-
ishankar et al. (2004) felt that the mono- and polyembryonic types of Indian
mango cultivars have a different genetic base, and that the polyembryonic
types might have been introduced from South-east Asia and are unlikely to
have originated in India. Mango culture gradually spread to tropical and
subtropical countries throughout the world, where selections were made
that were adapted to particular growing conditions. Thus, selection by man
has played the most signicant role in the development of new mango culti-
vars. The explorers who tasted the mango in the regions of its origin were
enchanted with its aromatic qualities, ambrosial avour and creamy, smooth
and silky texture, and introduced the fruit to other tropical regions. The
spread of Hinduism and Buddhism assisted in the distribution of mangoes in
South-east Asia. The Chinese traveller Hwen Tsang who visited India in the
rst half of the 7th century ad returned to China with the mango. The mango
was known in Baghdad in the 7th century.
The Persians or Omanis may have taken mangoes to East Africa around
the 10th century ad. The fruit was introduced to East and West Africa in the
early 16th century by the Portuguese and thence into Brazil. After being
established in Brazil, the mango was carried to the West Indies, being rst
planted in Barbados by about 1742 and later in the Dominican Republic and
Jamaica (about 1782). The mango was introduced into Mexico from the Phil-
ippines by the Spanish and also from the West Indies (Morton, 1987). Duval
et al. (2006) developed microsatellite markers for studying the genetic vari-
ability of Caribbean mangoes and concluded that there were two routes of
mango to the French West Indies, namely, cultivars grown in Central Amer-
ica (Mexico) and South America (Colombia) introduced from South-east Asia
and also from former French colonies in the Indian Ocean. As the mango
C.P.A. Iyer and R.J. Schnell 70
adapted to new locales, new cultivars were selected based on local adapta-
tion and fruit preferences.
Impact of Florida mangoes
The rst recorded successful introduction of mango into Florida (USA) was
made in 1861 (Knight, 1980). The earliest introductions were from the West
Indies and India, followed by the introduction of several hundred accessions
in the 20th century from South-east Asia, India and from other mango-growing
areas of the world (Florida Mango Forum, 1951). The introduction of mangoes
into Florida and subsequent development of a Florida group of mangoes has
been reviewed by Knight and Schnell (1994). The Florida mango cultivars are
unique in that they are hybrids between Indian cultivars (primarily mono-
embryonic) and the South-east Asian cultivars (primarily polyembryonic)
selected under south Florida conditions. The mango breeding system favours
out-crossing. Therefore, the proximity of numerous genotypes of disparate
geographical origins led to the production of many new seedlings by inter-
pollination in Florida (Knight and Schnell, 1993). Florida selections are
therefore not the result of a formal breeding programme. Early Florida selec-
tions were made by growers and enthusiasts and historical information is
often anecdotal. The Florida Mango Forum, established in 1938 for the
advancement of mango production, documented historical information on
the parentage of Florida cultivars in their proceedings. In addition to the
United States Department of Agriculture (USDA) Germplasm Resources In-
formation Network (GRIN) database, several sources compile information
on Florida mango selections and introduction of accessions to Florida ( Ruehle
and Ledin, 1956; Singh, 1960; Campbell and Campbell, 1993; Schnell et al.,
2006). With the exception of South-east Asia, Australia and some African
countries, which cultivate mostly locally selected varieties, the majority of
countries produce cultivars developed in Florida, i.e. Haden, Tommy
Atkins and Keitt (Galan Sauco, 1997). These Florida selections are now
widely grown commercial cultivars affording production stability across
many environments (see Mukherjee and Litz, Chapter 1, this volume).
4.3 Reproductive Mechanisms
Polyembryony
Nucellar embryos
Mangoes can be classied into two groups, monoembryonic and polyembry-
onic, based on their mode of reproduction from seeds. In general, monoem-
bryonic seeds are found in the sub-tropical group (Indian type) and the
polyembryonic seeds in the tropical group (South-east Asian). Monoembry-
onic mango seeds each contain a single zygotic embryo, and hence only one
seedling per seed that is of probable hybrid origin. Polyembryonic mango
Breeding and Genetics 71
seeds can contain one or more embryos, one of which is usually, but not
always zygotic. Adventitious embryos develop from the nucellus, a maternal
tissue surrounding the embryo sac, and consequently the seedlings of poly-
embryonic mangoes are genetically identical to the maternal parent. Adven-
titious embryos can also originate by direct budding from the cotyledons and
hypocotyls of other nucellar embryos (Juliano, 1934). According to Mahesh-
wari and Rangaswamy (1958), the nucellar cells destined to form adventi-
tious embryos are recognizable by their dense cytoplasm and starchy
contents. They gradually push into the embryo sac cavity where they divide
and differentiate into embryos.
Inheritance of polyembryony
Polyembryony is genetically determined. Leroy (1947) considered that adven-
tive embryony probably reects the effect of one or more recessive genes.
This view was supported by Sturrock (1968), whose study of the progenies of
monoembryonic mango hybridized with polyembryonic cultivars indicated
that monoembryony was possibly a dominant trait. In contrast, Aron et al.
(1998) and Brettell et al. (2004) observed that polyembryony in mango is con-
trolled by a single dominant gene. Schnell et al. (2006) reported that 58 of the
Florida cultivars had been classied with 50 being monoembryonic and eight
polyembryonic. Information from the Florida cultivars parentage analysis
using 25 microsatellite markers supported the ndings of Aron et al. (1998)
where polyembryony was found to be dominant. Haden is a cross of the
monoembryonic Mulgoba and the polyembryonic Turpentine. If we assume
that a single dominant gene controls this trait, all of the Indian cultivars in
Florida must be homozygous recessive and the Turpentine parent of Haden
must have been heterozygous. The evidence suggests that Haden inherited
the recessive allele from Turpentine, as all identied progeny of Haden are
monoembryonic with the exception of Winters. The most probable pollen
parent of Winters is Ono, a polyembryonic cultivar from Hawaii. The fre-
quency of this dominant allele is low in the Florida population and absent
from the Indian cultivars in Florida. In view of these interesting ndings, and
since a thorough knowledge of inheritance of polyembryony is essential for
speculating the origin of M. indica, more work on these lines is warranted.
Floral biology and pollination
The mango inorescence is primarily terminal, although axillary and multi-
ple panicles may also arise from axillary buds. Both perfect (hermaphrodite)
and staminate (male) owers occur in the same inorescence. The total num-
ber of owers in a panicle may vary from 1000 to 6000, depending on the
cultivar (Mukherjee, 1953). Initial fruit set in mango is directly related to the
proportion of perfect owers, although the nal fruit set does not necessarily
depend on this ratio (Iyer et al., 1989). It appears that the proportion of per-
fect owers in a cultivar becomes critical for optimum fruit set only when the
proportion drops to 1%.
C.P.A. Iyer and R.J. Schnell 72
Flowers begin to open early in the morning and anthesis has generally
been completed by noon. The greatest number of owers opens between 9
and 10 a.m. Although the receptivity of the stigma continues for 72 h after
anthesis, it is most receptive during the rst 6 h; however, there are reports
that the stigma can become receptive even before anthesis has occurred
(Singh, 1960). The minimum time required for pollen grains to germinate is
1.5 h (Sen et al., 1946; Singh, 1954; Spencer and Kennard, 1955). Singh and
Singh (1952) observed 98% pollen viability after 11 months in storage at 7C
and 25% relative humidity (RH), and 65.7% viability after 24 months of stor-
age at 0C and 25% RH.
Mango is cross-pollinated, which is carried out by insects such as the
common housey, honeybees and thrips, and possibly by other insects
al-though to a lesser extent. Pollination by wind and gravity has been sug-
gested to occur in mango (Popenoe, 1917; Maheshwari, 1934; Malik, 1951). In
nature, > 50% of owers do not receive any pollen. Some workers had sug-
gested that self-pollination in certain cultivars can also occur quite frequently
(Dijkman and Soule, 1951). Studies by Issarakraisila and Considine (1994)
have shown that for polyembryonic Kensington, a night temperature of
< 10C results in pollen grains with low viability (< 50%). The optimum

temperature for normal meiosis is between 15 and 33C with 7085%
viability.
Incompatibility
Although the existence of self-sterility in mango was suspected several years
ago (Ruehle and Lynch, 1948, cited in Sharma and Singh, 1970; Dijkman and
Soule, 1951), the prevalence of self-incompatibility was clearly established in
monoembryonic Dashehari by Singh et al. (1962). Subsequently, detailed
studies indicated that the four popular monoembryonic cultivars of northern
India (i.e. Dashehari, Langra, Chausa and Bombay Green) were self-
incompatible (Mukherjee et al., 1968; Sharma and Singh, 1970). Embryo-
logical studies have shown that although fertilization takes place after
self-pollination, degeneration of endosperm occurs 15 days after pollination
involving self-incompatible parents (Mukherjee et al., 1968). The self-
incompatibility system operating in mango appears to be of the sporophytic
type. Instances of cross-incompatibility among certain mango cultivars
have also been reported (Ram et al., 1976), necessitating the identication of
suitable pollinizers for mango.
Using an approach involving isozyme analysis, Dag et al. (2006) have
initiated studies in many commercial mango cultivars in Israel and con-
cluded that self-pollination is not a yield-limiting factor in monoembryonic
Maya and the practice of planting Maya in solid blocks is sound. They had
obtained similar results earlier with monoembryonic Tommy Atkins (Dag
et al., 1997).
Breeding and Genetics 73
Cytology
Chromosome number
Information on the cytology of mango is quite limited. Only a few Mangifera
species (i.e. M. indica, M. caloneura, M. sylvatica, M. foetida, M. caesia, M. odorata
and M. zeylancia) have been studied, and were found to have chromosome
numbers of 2n = 2x = 40 and n = x = 20 (Mukherjee, 1950, 1957; Roy and
Visweswariya, 1951). Chromosome numbers and ploidy status of other species
are yet to be studied. The only exception to this chromosome number that has
been reported to date (Roy and Visweswariya, 1951) involves Vallikolamban,
which was reported to be tetraploid (2n = 4x = 80), although subsequent stud-
ies have indicated that it is only a diploid (Majumder and Sharma, 1990).
Polyploidy
Mango has been referred to as an allopolyploid. Due to the presence of
secondary associations at metaphase of meiosis, Mukherjee (1950) suggested
that the basic chromosome number of Mangifera is n = 8. In addition, the high
number of somatic chromosomes and the correspondingly high number of
nucleolar chromosomes led him to conclude that mango is an allopolyploid.
However, the evidence used to arrive at this conclusion is not unequivocal.
In fact, the molecular marker evidence is antithetical to this conclusion.
Results from Duval et al. (2005), Viruel et al. (2005) and Schnell et al. (2005,
2006) using microsatellite markers all indicate that M. indica is diploid.
Although many wild Mangifera species are potentially valuable for crop
improvement, they are yet to be exploited. Mukherjee (1963) felt that the
different Mangifera species could intercross easily, based on the success ob-
tained with interspecic crosses between M. zeylanica and M. odorata.
4.4 Inheritance of Characters
High heterozygosity in the cultivars that are used in hybridization and the
inadequate number of hybrid progenies realized has made accurate genetic
analysis in mango very difcult. However, based on limited data, some indi-
cations are available which would be useful in selecting parents in breeding
programmes designed with specic objectives. In studies of the distribution
of different traits in seedlings derived from open-pollination (where the pol-
len parent is unknown), Lavi et al. (1989) observed: (i) there is no maternal
effect on juvenile period and fertility; (ii) there is a slight effect of the female
parent on fruit taste and size; (iii) there is a maternal parent effect on harvest
season and fruit colour.
Dwarfness, regular bearing and precocity
An analysis based on observations of more than 1000 hybrids, involving sev-
eral combinations, has revealed that dwarfness, regular bearing and precocity
C.P.A. Iyer and R.J. Schnell 74
are controlled by recessive genes (Sharma and Majumder, 1988a). Regularity
of bearing appears to be linked with precocity. Characters contributing to
biennial bearing are dominant over those governing regular bearing habit.
Flesh colour
Sharma (1987) considered that additive gene action may be involved in the
inheritance of esh colour; however, studies involving monoembryonic
Alphonso and Neelum have indicated that light yellow is dominant over
orange-yellow (Iyer, 1991).
Skin colour
With regard to skin colour of fruit, Sharma (1987) observed that when red
cultivars were crossed with green cultivars, the F
1
seedlings exhibited vari-
ous gradations of red. Iyer and Subramanyam (1987) also found a wide array
of colours in the hybrids when the coloured monoembryonic Janardhan
Pasand was crossed with green-fruited cultivars, indicating that colour is
mediated by a number of loci.
Flowering time
The owering response of mango cultivars in subtropical and tropical envi-
ronments differs greatly (see Davenport, Chapter 5 this volume). Trees can be
stimulated to ower under certain conditions in tropical environments using
ethephon; however, this is ineffective in subtropical environments. Schnell and
Knight (1998) investigated the repeatability of owering using eight cultivars
over six harvest cycles (years), collecting data weekly. Three characters were
evaluated: days to bloom (DTB) (from 1 November in each year), days in bloom
(DIB), and days in bloom and fruit (DIBF). Signicant differences were detected
for all three characters for both years and cultivars. Signicant differences were
not detected for replicate trees within cultivars. Repeatability (R) of the ower
phenology characters was high (R = 0.73, 0.88 and 0.77 for DTB, DIB and DIBF,
respectively). This indicates that much of the variation is heritable and useful
for extending owering times in subtropical environments.
Beak
The presence of a beak on mango fruit appears to be a dominant character
since most of the hybrid plants had this feature when monoembryonic Ban-
galora (Totapuri) was used as one of the parents in controlled crosses (Iyer
and Subramanyam, 1979). Bunch bearing was found to be a dominant char-
acter, as indicated in many crosses (Sharma et al., 1972) involving bunch-
bearing types with single-fruited cultivars.
Breeding and Genetics 75
Disease resistance
Bacterial canker (Xanthomonas campestris pv. mangiferaeindicae) resistance
appears to have cytoplasmic inheritance. Whenever Neelum, a susceptible
cultivar, is used as the female parent, susceptibility is transmitted to all the
hybrids, irrespective of the male parent (Sharma and Majumder, 1988a). It
has been suggested that internal breakdown (spongy tissue) is mediated by
recessive genes (Iyer, 1991). Susceptibility to mango malformation appears
to be dominant, since crosses involving resistant Bhadauran did not yield
any resistant hybrids (Sharma and Majumder, 1988a).
Other horticultural traits
The genetics of inheritance of various horticultural traits appears to be
unclear. More knowledge will be forthcoming only when large-scale con-
trolled hybridization experiments are undertaken at different mango research
centres. However, the information now available for some of the characters
is very useful in deciding which parental genotypes ought to be used in
hybridization programmes.
Brettell et al. (2004) subjected the large number of mango hybrids obtained
from the Australian National Mango Breeding Programme to a biometrical
analysis. Their data indicated that many of the important fruit quality aspects,
including fruit weight, fruit shape, ground skin colour, fruit width and pulp
depth have high heritabilities and can therefore be readily selected in a breed-
ing programme. Of particular interest is the observation that a high frequency
of hybrids with a red or burgundy blush can be recovered from crosses where
one parent has an intense red blush. Similarly, while the unique avour com-
pounds associated with Kensington Pride are also found in nearly 50% of
the hybrids involving Kensington Pride, leaf fragrance was not found to be
a reliable predictor of fruit avour.
4.5 Breeding Objectives
General objectives
Breeding objectives vary from region to region, depending on the specic
trait(s) for which improvement is sought. However, they can be broadly gen-
eralized to consist of the development of cultivars with: (i) regular bearing;
(ii) dwarf tree habit; (iii) precocity; (iv) attractive, good sized (300500 g),
good quality fruits (appealing avour and rm esh without bres); (v)
resistance to major diseases and pests; (vi) freedom from physiological dis-
orders; and (vii) good shipping qualities and shelf life. While it would be
hard to combine all these characteristics within a relatively short time, espe-
cially resistance to all major diseases and pests, all of these characteristics are
basic for commercial success.
C.P.A. Iyer and R.J. Schnell 76
With regard to the improvement of rootstocks by breeding, the main
desirable features are: (i) polyembryony; (ii) dwarng; (iii) tolerance of
adverse soil conditions (high pH, calcareous soil, etc.); and (iv) good scion-
compatibility.
Specic objectives
In addition to improving general characters such as yield and quality, breed-
ing has also been undertaken for certain specic purposes.
Dwarfness
Because of the obvious benets of comparatively dwarf trees for orchard man-
agement and fruit quality, attempts have been focused on obtaining hybrids
with a dwarf tree framework. Breeding for dwarfness is important in mango,
since a consistent dwarng effect of any rootstock has not been established to
date. The Indian cultivars that could be useful as a source for dwarfness
include Kerla Dwarf, Janardan Pasand, Manjeera, Amrapali, Creeping
(Iyer and Subramanyam, 1986) and Nileswar Dwarf (Singh, 1990).
Regular bearing
The causes of irregular bearing vary from region to region. In general, the
main reason for alternate bearing, particularly in subtropical India, is the
lack of initiation of vegetative growth soon after fruiting. However, two cul-
tivars, Neelum and Bangalora (Totapuri), which are regular bearers, have
been extensively used as either of the parents in a hybridization programme
to transfer the regular bearing habit to hybrids. Neelum has been observed
to be a good combiner and has contributed to the evolution of many regular-
bearing Indian hybrid cultivars. However, Bangalora is not a suitable par-
ent since the hybrids possess very prominent beaks and their fruit quality is
invariably poor. The regular bearing Florida cultivars (i.e. Tommy Atkins,
Keitt, etc.) also have potential as parents.
Fruit colour
Most of the commercial cultivars in South-east Asia possess green skin.
Efforts are underway to produce new hybrid cultivars that retain the good
qualities of these fruits together with attractive skin colour, so that they will
occupy a better position in international trade. Since good skin colour has
been shown to be transmissible to hybrids from suitably coloured parental
cultivars, a number of cultivars with coloured skin are being used for hybrid-
ization. In general, the attractively coloured Florida cultivars have been
found to be suitable parents. In addition, there are several Indian cultivars
(e.g. Janardan Pasand, Suvarnarekha, etc.) that would be suitable for use
as parents for this purpose.
In Florida, the skin colour of the mango is an important factor and red
skin is considered essential for mangoes shipped to northern markets. In the
past, the evaluation of mango colour has been subjective and based on visual
Breeding and Genetics 77
ratings. Large errors are associated with these types of ratings, which makes
evaluation based on fruit colour difcult. Ayala-Silva et al. (2005) used a colo-
rimeter to quantify fruit colour, quality and differentiation among cultivars.
Mango colour was measured with a Minolta Chroma Meter CR-400 (Osaka,
Japan) portable tristimulus colorimeter and fruit chromaticity was recorded
in Commission Internationale de lEclairage (CIE) L
*
, a
*
and b
*
colour space
coordinates. In this system of colour representation the values L
*
, a
*
and b
*

describe a uniform three-dimensional CIE colour space. If the L
*
, a
*
and b
*

coordinates are known, then the colour is not only described, but also located
in quantiable space. Maternal half-sib families (MHS) of the mango culti-
vars, Keitt, Tommy Atkins, Tyler Premier, Mamita, White Alfonso and
Sandersha were evaluated along with two parental check clones, Tommy
Atkins and Keitt. Signicant differences were found for each of the L
*
, a
*
and
b
*
colour space coordinates. Further work is underway to estimate the herita-
bility of these traits to estimate their usefulness for breeding and selection.
Disease resistance
MANGO MALFORMATION. Although no breeding work has been reported that spe-
cically addresses disease or pest resistance/tolerance, cultivars are known to
show varying degrees of susceptibility to biotic stress (see Ploetz and Freeman,
Chapter 8, this volume ). Mango malformation, caused by Fusarium subglutin-
ans, is a very serious disease that has threatened the very survival of the
mango industry in many subtropical mango-growing regions. As there are
no reliable cultural and chemical control measures available, breeding for
resistance/tolerance has been attempted using Bhadauran as the resistant
parent; however, all of the F
1
hybrids were susceptible to the disease (Sharma
and Majumder, 1988a). In this respect, the observations of Ram et al. (1987)
are very encouraging. Out of 102 cultivars screened, three of them, namely,
Bhydayam Dula, Samar Bahist Rampur and Mian Sahib, were free of
malformation and could be tried as one of the parents in hybridization.
BACTERIAL CANKER. Bacterial canker is a serious problem with many cultivars.
The only cultivar possessing true resistance to canker is Bombay Green
(Prakash and Srivastava, 1987) and hence could be a potential gene donor.
ANTHRACNOSE. Anthracnose, caused by Colletotrichum gloeosporioides Penz., is
the most widespread disease in all mango-growing countries, manifesting
itself in blossom blight, peduncle blight, leaf spot, twig blight, wither tip, fruit
russetting and fruit rot. Tommy Atkins is moderately tolerant of anthracnose
and coupled with its other desirable qualities (i.e., regular bearing, fruit colour,
etc.) should be a good parent in breeding programmes. In addition, Parish
and Fairchild have been reported to be relatively resistant (Yee, 1958).
POWDERY MILDEW. Powdery mildew caused by Oidium mangiferae Berthet, has
been reported to cause heavy loss of crops in years when RH is very high and
accompanied by cool nights during owering. Cultivar differences with respect
to susceptibility are recognized, and Pairi (Raspuri) is highly susceptible.
C.P.A. Iyer and R.J. Schnell 78
Gupta (1976) has listed those cultivars that are most tolerant of this disease
Neelum, Zardalu, Bangalora, Totapuri-Khurd and Janardan Pasand
and hence could be valuable in breeding programmes.
PEST RESISTANCE. Considerable variation is also known to occur among mango
cultivars with respect to their susceptibility to attack and injury by insect pests.
Although no resistant genotypes have been reported for the mango hopper
(Idiocerus spp.), the insect has been observed to avoid colonizing open plant
types where free movement of wind is possible, an observation that could be
useful in selection. Although complete resistance is not known to either fruit
y (Bactrocera spp.) or seed weevil (Stenochetus mangiferae), variation in the
degree of susceptibility has been reported (Iyer, 1991). Rossetto et al. (2006)
observed that resistance to fruit y is compatible with fruit quality and pro-
ductivity and advocated that resistance to fruit y should be one of the objec-
tives of all mango breeding programmes. Their results also indicated that the
main factors for resistance of mangoes to fruit ies lie in the fruit peel and not
in the fruit pulp.
4.6 Methods of Breeding
Selection from open-pollinated seedlings
In India, almost all cultivars are selections that were made from naturally occur-
ring open-pollinated seedlings. All of the Florida cultivars were selected from
open-pollinated seedling progenies; none has come from a controlled breeding
programme. Among the 64 Florida cultivars evaluated in the parentage analysis
by Schnell et al. (2006), the genetic background was found to be based on as few
as four Indian cultivars and the polyembryonic cultivar Turpentine. Two Indian
cultivars, Mulgoba and Sandersha, are in the background of most Florida
types with Amini, Bombay, Cambodiana, Long, Julie and Nam Doc Mai
making lesser contributions. In the parentage analysis Turpentine 10 was iden-
tied as a most probable paternal parent for Haden. The polyembryonic seed-
ling races of Cuba and Florida were considered the same by Popenoe (1920) who
called them the West Indian race (commonly known as Turpentine in Florida).
Haden was reported as the maternal parent for ten cultivars included in the
analysis, but based on the parentage analysis, 31 cultivars were found to have
Haden as one of the most likely parents. Likewise, the other important early
Florida selection Brooks is the parent of seven cultivars. Haden, Brooks and
seedlings of Haden and Brooks have contributed disproportionately to the
Florida group. In Florida, modern selection and breeding programmes for
mango have focused on cultivars with exceptional production, red skin, disease
resistance and extended shelf life. Methodology for crop improvement consists
of collecting seeds from selected maternal parents with desired characteristics
and growing them in close proximity to desirable male parents. Seedlings are
screened by leaf aroma and horticultural traits, leading to a eld population of
thousands of candidate seedlings (Campbell and Zill, 2006).
Breeding and Genetics 79
In the Canary Islands, Spain, breeding mainly involves selection of open-
pollinated seedlings of Lippens. Lavi et al. (2004) indicated that mother trees
should not be chosen entirely on the basis of their phenotypes and trees with
inferior performance could also be included since progeny performance is
quite unpredictable. They observed that most of the variance components of
the agriculturally interesting traits are non-additive (Lavi et al., 1998) and
most of these traits result from dominant and epistatic interactions.
The Israel mango-breeding programme therefore relies on open-pollination
involving many mango cultivars from various parts of the world and screen-
ing approximately 100 seedlings from each mother tree. The seedlings are
grown on their own roots in the nursery for about a year and then planted in
the eld at spacings of 2 6 m. Fruiting occurs after 36 years and rst selec-
tion is carried out based on eld and laboratory data. Fruit characteristics at
this stage are good skin colour, fruit weight of 400600 g and high fruit qual-
ity (good taste, absence of bres and small seed). Where a long harvest sea-
son is desired both early and late harvest seedlings are selected and a general
idea about shelf life of these seedlings is obtained. The second selection is
carried out under commercial conditions by several experienced farmers
using grafted plants. Plants that successfully pass this stage are planted in
semi-commercial plots for a nal assessment before recommendation to farm-
ers. The two selection stages are aimed at shortening the breeding programme
and minimizing both the false negatives (loss of interesting seedlings which
were not identied) and the false positives (wrong identication of interest-
ing seedlings which should actually be rejected). New mango cultivars have
also been selections made from open-pollinated seedlings. Maya and Nim-
rod are seedlings of the same mother tree (Oppenheimer, 1967); Tahar is a
seedling of Irwin (Slor and Gazit, 1982) and Naomi is a seedling of Palmer
(Tomer et al., 1993). The other promising selections include Shelly (late sea-
son), Tango (early season), Selection 20/1 (large fruit with aborted seeds)
and Selection 1/5 (shiny red colour).
The South African mango-breeding programme also places a major
emphasis on open-pollinated seedling selection. The screening of seedlings,
besides undergoing tests for various plant and fruit characters, also includes
shipping quality tests in which fruits are packed in 4 kg crates and stored at
11C for 28 days to simulate shipping conditions. After storage the fruits are
allowed to ripen at room temperature (25 2C) and screened thoroughly.
Promising seedlings are again eld-tested with Sabre as rootstock with a
row of 40 trees along with ten trees of commercial varieties. Two cultivars,
Joa (Palmer seedling) and Chene (Kent seedling), were released from
the breeding programme in 1996. Another high yielding selection A2-CD28
(Fascell seedling) is a midseason clone with an attractive pink blush (Human
et al., 2006) and has been recommended for Plant Breeders Rights in 2005.
The other promising selections are Osteen (Haden seedling) and Neldica
(Palmer seedling).
The 'R2E2' cultivar developed in Australia is a seedling progeny of the
Florida cultivar 'Kent', and is now the second most popular mango grown in
Australia.
C.P.A. Iyer and R.J. Schnell 80
Controlled pollination
Hand pollination
The traditional, cumbersome method involving the continued pollination of
owers on a panicle over several days when the owers are open has now
been replaced in India with more efcient methods. The current method
involves the pollination of a limited number of owers per panicle (maxi-
mum of ten), utilizing a larger number of panicles since it is very rare that a
panicle bears more than one fruit to maturity. Using this method, fruit set as
high as 3.85% can be achieved compared to the 0.231.57% efciency involv-
ing other methods (Mukherjee et al., 1968; Singh et al., 1980).
Caging
The enclosure of two desirable parents of synchronous owering in a screen
house with pollinating insects provides a more practical method of hybridiza-
tion. This method has been used in Israel (Degani et al., 1993), Brazil (Pinto and
Byrne, 1993) and South Africa (Cilliers et al., 1996). A standardized caging
technique for mango breeding was previously used in India following the
discovery of self-incompatibility in some of the most popular commercial culti-
vars (Sharma and Singh, 1970). This procedure involves the planting of self-in-
compatible (female) and male parents in specially prepared breeding plots, and
are enclosed in an insect-proof cage into which freshly reared houseies, or any
other suitable pollinator, are introduced to effect cross-pollination (Sharma
et al., 1972).
Polycross mating
A polycross is simply the use of a number of advanced selections or current
commercial clones planted in a design that maximizes the chance for
cross-pollination. The polycross design has been extensively used in
sugarcane breeding where small ower size and low numbers of seedlings
per cross make controlled pollinations difcult. At USDA Subtropical Horti-
culture Research Station (SHRS) in Miami the clones Haden, Tommy
Atkins, Kent, Keitt and Nam Doc Mai have been used to produce new
seedlings for selection. Five clones of each genotype were planted, with at
least one plant of each genotype next to all other genotypes. Over 1000 seed-
lings from known mother trees are planted as maternal half-sib families. The
pollen parent of superior selections is determined using microsatellite
markers.
4.7 Handling of Hybrid Populations and Selection
Criteria for initial selection
Primary selection from the hybrid progeny is based on: (i) precocity; (ii) fruit
size and shape; (iii) skin colour; (iv) fruit characteristics (high pulp to stone
ratio and freedom from bre and physiological disorders); and (v) fruit qual-
ity. Following this preliminary evaluation, selected hybrids are retained for
Breeding and Genetics 81
further screening. It is important to graft the hybrids onto proper rootstocks
as early as possible, as grafted plants are precocious. At least ten grafted
plants of each selected hybrid are used in the nal selection, which is based
on yield, regularity in bearing and response to diseases and pests, in addition
to other desirable fruit characters. At least 3 consecutive years performance
data should be collected before deciding on their suitability for release as
new cultivars.
Pre-selection
Trees have a long juvenile phase, and the development of pre-selection meth-
ods is important for discarding inferior seedlings at a very early stage, obvi-
ating the need for maintaining a large number of seedlings for long periods.
This can save time, land and labour. Leaf avour has been reported to be
directly correlated with fruit avour (Majumder et al., 1972; Whiley et al.,
1993). Emergence of new growth ushes, simultaneously with fruiting or
immediately after harvest, is indicative of regular bearing (Sharma et al.,
1972). A higher phloem to xylem ratio, associated with dwarng, has been
used effectively as a pre-selection criterion. Genotypes in which the ratio
exceeds 1.0 are least vigorous, those with a ratio between 0.6 and 1.0 are of
medium vigour and those with a ratio of less than 0.6 are most vigorous
(Kurian and Iyer, 1992). In addition, higher levels of phenolics in the apical
bud is associated with reduced vigour and dwarng (Iyer, 1991). Although
Majumder et al. (1981) indicated that low stomatal density is an indicator of
dwarfness this has not been conrmed by other workers (Iyer, 1991). Regular
bearing mango cultivars have low polyphenol oxidase (PPO) activity (cate-
cholase and cresolase) compared to alternate bearers (Sharma, 2003). Sharma
et al. (2000) observed that a strong positive correlation existed between the
incidence of oral malformation and both enzyme activity (catecholase and
cresolase) and phenolic content and speculated that PPO activity can be used
as a biochemical index for screening mango germplasm against malforma-
tion disease.
Potential for marker assisted selection (MAS)
More than 65 microsatellite markers have been developed for mango and
these are easily used to verify parentage using a software package such as
cervus (Marshall et al., 1998). When caging trees or using the polycross mat-
ing design it is possible to identify the male parent from a set of potential
male parents. This has been useful in cacao breeding where mistakes in pol-
lination have lead to the estimation of unreliable breeding values for parental
clones. The development of linkage maps and identication of quantitative
trait loci (QTL) for productivity and quality traits has led to a very successful
MAS in cacao (Schnell et al., 2007). This could serve as a model for future
mango breeding and selection efforts.
C.P.A. Iyer and R.J. Schnell 82
Molecular markers
Molecular markers can be used for estimating genetic relationships among
clones, for parentage analysis and for the development of a saturated linkage
map. Isozymes were the rst markers to be used for ngerprinting mango
cultivars, to determine self- versus cross-pollination and to estimate genetic
relationships (Degani et al., 1990; Knight and Schnell, 1994). RAPD markers
were also used to ngerprint cultivars and estimate genetic relationships in
mango (Schnell et al., 1995). A group of Haden seedlings and a random
group of seedlings were evaluated using 11 RAPD primers. This study sup-
ported the Haden parentage of Eldon, Lippens, Tommy Atkins and
Zill; however, the parentage of Glenn and Osteen was questioned. Adato
et al. (1995) used DNA ngerprinting (DFP) to evaluate genetic relationships
between 26 mango cultivars and 14 rootstocks. They provided a pedigree
that further conrmed the relationship between many of the Haden seed-
lings. Lopez-Valenzuela et al. (1997) used RAPD markers to estimate genetic
diversity among 15 rootstock cultivars using 13 markers, and identied a
specic RAPD band associated only with the polyembryonic types. Eiad-
thong et al. (1999) utilized anchored simple sequence repeat markers to anal-
yse 22 mango cultivars; they were able to distinguish genotypes, but were
unable to nd markers unique to either monoembryonic or polyembryonic
types, or for the Thai cultivars selected for green harvest (crispy mango) from
the cultivars selected for ripe fruit production. Kashkush et al. (2001) utilized
amplied fragment length polymorphisms (AFLP) to estimate genetic rela-
tionships between 16 cultivars and seven rootstock cultivars. They also anal-
ysed 29 progeny from a cross of Tommy-Atkins and Keitt and produced a
crude linkage map that identied 13 of the 20 linkage groups.
Viruel et al. (2005) developed the rst reported set of 16 microsatellite
markers for mango, of which 14 produced the expected one or two ampli-
cation products per genotype. These 14 microsatellites were used to evaluate
28 mango genotypes that included 14 Florida cultivars. Discrimination of all
28 genotypes was possible and the average number of alleles per locus was
5.3. Previously known pedigree information for the Haden family of man-
goes was conrmed and was in agreement with previously published RAPD
and DFP analyses (Adato et al., 1995; Schnell et al., 1995) with one exception.
Viruels clone of Zill was not resolved as a seedling of Haden. Schnell et al.
(2005) developed a second set of 15 microsatellite markers and analysed 59
Florida cultivars and four related species. Two of the microsatellites were
monomorphic among the Florida cultivars; the other 13 had an average num-
ber of alleles per locus of 4.2 with polymorphism information content (PIC)
values varying from 0.21 to 0.63.
Schnell et al. (2006) used 25 microsatellite loci to estimate genetic
diversity among 203 unique mangoes (M. indica L.), two M. grifthii Hook. f.
and three M. odorata Griff. accessions maintained at the National Germplasm
Repository (NGR) and by Fairchild Tropical Botanic Garden (FTBG) in Miami,
Florida. The 25 microsatellite loci had an average of 6.96 alleles per locus
and an average PIC value of 0.552 for the M. indica population. The total
Breeding and Genetics 83
propagation error in the collection (i.e. plants that had been incorrectly
labelled or grafted) was estimated to be 6.13%. When compared by origin,
the Florida cultivars were more closely related to Indian than to South-east
Asian cultivars. Unbiased gene diversity (H
nb
) of 0.600 and 0.582 was found
for Indian and South-east Asian cultivars, respectively, and both were higher
than H
nb
among Florida cultivars (0.538). When compared by horticultural
type, H
nb
was higher among the polyembryonic types (0.596) than in the
monoembryonic types (0.571).
To date 63 microsatellite markers have been developed for mango
(Duval et al., 2005; Honsho et al., 2005; Schnell et al., 2005; Viruel et al., 2005).
This number is more than adequate for genetic diversity studies and for par-
entage analysis as has been demonstrated by Schnell et al. (2006); however, it
is not enough to develop a saturated linkage map for the 20 linkage groups
of mango. Developing an additional 200 microsatellite or single nucleotide
polymorphic markers is a major objective of the USDA Agriculture Research
Service (ARS) programme in Miami over the next 2 years. Three experimen-
tal populations have been developed and planted in the eld as mapping
populations. The rst population is an F
2
population derived from self-polli-
nation of Tommy Atkins consisting of 168 seedlings that was planted in the
eld in 1995. The second population is an F
2
population derived from self-
pollination of Haden. A total of 224 seedlings from a single isolated Haden
tree have been in the eld for 3 years. Phenotypic data collection is in prog-
ress for both of these populations. The development of a saturated linkage
map and the identication of QTL for important traits are objectives for the
USDA-ARS programme in Miami for the next 5 years.
4.8 Minimizing Problems in Breeding
Heavy fruit drop
Heavy fruit drop ultimately results in few hybrid fruits, despite the large
number of owers used for cross-pollination. While many recommendations
are available to minimize mango fruit drop with growth regulators, these
have not been very useful in breeding programmes where the number of
owers remaining in a panicle is very low. Iyer and Subramanyam (1972)
suggested that embryo culture could be used to rescue hybrid embryos, and
Sahijram et al. (2005) developed in-vitro techniques to rescue immature mango
embryos from controlled crosses and recovered hybrid plants.
Long juvenile phase
Normally, mango seedlings require 310 years to ower, thereby prolonging
the breeding programme. Grafting individual hybrids on the proper
rootstocks at the earliest possible stage and growing them in a location
where climatic stress (particularly cold weather) prevails, induces precocious
C.P.A. Iyer and R.J. Schnell 84
owering. Iyer (1991) has reported signicant differences between seedlings
on their own roots and grafted plants of the same genotype with respect to
fruit size, quality and even colour in the early years. However, different
results have been reported in a recent study of the effect of rootstocks on the
performance of seedling scions with respect to ten horticultural traits (Lahav
et al., 1995). No difference of practical importance was found between the
original seedlings and their grafted duplicates.
Singh (1969) has suggested that young mango seedlings can be induced
to ower and fruit if they are grafted onto comparable shoots of a bearing
tree (a few days before owering). The scions are defoliated and girdled.
Using this technique, it has been reported that the fruit characteristics of F
1

hybrids can be determined within 2 years and F
2
hybrids within 4 years, thus
eliminating at least 10 years from the period required to raise and evaluate F
2

populations (Singh, 1963).
Ethephon (Chacko et al., 1974) and paclobutrazol (Anonymous, 1984)
have ower-inducing properties in mango and could also be utilized for
shortening the juvenile phase. However, they must be used with caution
since chemical induction of owering can alter fruit size and this could lead
to errors in judgement when making selections within the hybrid progeny.
Polyembryony
Seeds of polyembryonic mango cultivars characteristically contain several
nucellar embryos, and may also contain a zygotic embryo. While nucellar
seedlings are preferred as rootstocks for mango because of their uniformity,
the breeder, on the other hand, is generally interested in sexual seedlings for
the selection of improved rootstocks. Until recently, crosses involving poly-
embryonic cultivars as the maternal parents were generally not performed,
since reliable methods for identifying zygotic seedlings were not available.
The use of polymorphic enzyme systems (isozymes) (Degani et al., 1990,
1992) to identify zygotic seedlings (Schnell and Knight, 1992; Truscott, 1992;
Degani et al., 1993) is based on the fact that nucellar seedlings should have
the same isozyme alleles as the maternal parent. A variation at a locus coding
for an enzyme indicates that the plant has originated by sexual reproduction.
Zygotic seedlings arising from self-pollination are distinguished from nucel-
lar seedlings by being homozygous at one or more loci at which the female
parent is heterozygous. Statistically, when three or four heterozygous loci are
examined, up to 88 or 94%, respectively, of the selfed zygotic seedlings are
identiable (Moore and Castle, 1988). Cross-pollination by another cultivar
of the same genotype is equivalent to self-pollination. Zygotic seedlings from
cross-pollination are distinguishable from those resulting from self-pollination
if they express an allele not carried by the female parent.
The frequency of occurrence of zygotic seedlings varies among the poly-
embryonic mango cultivars, i.e. 22% with 13-1 (Degani et al., 1993), 20 and
24% with Turpentine (Degani et al., 1993 and Schnell and Knight, 1992,
respectively), 2 and 4% with Sabre (Truscott, 1992 and Schnell and Knight,
Breeding and Genetics 85
1992, respectively) and 36 and 64% with Madu and Golek, respectively
(Schnell and Knight, 1992).
4.9 Achievements of Conventional Breeding
Despite the many problems associated with mango breeding for cultivar devel-
opment, many useful hybrids have been released. The earliest attempts were
probably made in the West Indies to combine the good qualities of the Indian
mango with the indigenous types by controlled pollination (Brooks, 1912).
India
Intervarietal hybridization in India has resulted in the release of many culti-
vars. The work at Sabour initially yielded two promising hybrids: Mahmood
Bahar and Probashanker, both combinations of Bombay and Kalapady
(Roy et al., 1956). Subsequently, four more hybrids have been developed.
These are: Sundar Langra (Sardar Pasand Langra) having Langra
quality and regular bearing habit; Alfazli (Alphonso Fazli) with Fazli
quality and early ripening; Sabri (Gulabkhas Bombai) having Bombai
fruit shape and colour of Gulabkhas with regular bearing habit; and Jawa-
har (Gulabkhas Mahmood Bahar) having high pulp and early bearing
habit (Hoda and Ramkumar, 1993). Developed in Kodur, the hybrid Swarna-
jehangir, combining the high quality of Jehangir and the attractive colour of
Chinnaswarnarekha, is a prolic bearer and is the best of all hybrids devel-
oped at this centre. The other hybrids released from Kodur are Neeludin
(Neelum Himayuddin), Neelgoa (Neelum Yerra Mulgoa) and
Neeleshan (Neelum Baneshan).
Two excellent, regular-bearing hybrids, Mallika and Amrapali, were
developed and released by the Indian Agricultural Research Institute (IARI),
New Delhi (Singh et al., 1972). Mallika is a hybrid between Neelum and
Dashehari with a high total soluble solids (TSS) content, a higher percentage of
pulp, breless esh and a fruit size of about 300 g. Amrapali (Dashehari Nee-
lum) is precocious, distinctly dwarf and hence amenable to high-density
planting, a regular bearer with excellent quality and is also very rich in vitamin
A. Recently, two more cultivars, Arunima (Amrapali Sensation) and Pusa
Surya (a selection from Eldon) have been released from the IARI. A promising
mango hybrid Ambika, a cross between Amrapali and Janardhan Pasand,
having a yellow colour with red blush, rm esh and scanty bre was
released from the Central Institute of Sub-Tropical Horticulture, Lucknow.
Four hybrid cultivars were released from the Indian Institute of Horticul-
tural Research in Bangalore: Arka Aruna (Banganapalli Alphonso),
Arka Puneet (Alphonso Banganapalli), Arka Anmol (Alphonso
Janardhan Pasand) and Arka Neelkiran (Alphonso Neelum). Arka
Aruna is dwarf, and large fruited with a high percentage of pulp and high
TSS content. It is ideal for homesteads. Arka Puneet is very similar to
C.P.A. Iyer and R.J. Schnell 86
Alphonso but free of spongy tissue, has a good shelf life and is not suscep-
tible to fruit y attack. Arka Anmol is a heavy bearer with good keeping
quality (Iyer and Subramanyam, 1993). Arka Neelkiran is free of spongy
tissue and has excellent skin colour.
Ratna is a cross between Alphonso and Neelum that was carried out
at the Fruit Research Station, Vengurla, Maharashtra; it has a larger fruit size,
fruit quality similar to Alphonso and is free of spongy tissue (Salvi and
Gunjate, 1988). A parthenocarpic mango cultivar, Sindhu, has been devel-
oped at this station as a result of back-crossing Ratna with Alphonso (Gun-
jate and Burondkar, 1993).
Two hybrid cultivars were released from the Fruit Research Station in
Sangareddy, Andhra Pradesh. Au-Rumani (Rumani Mulgoa) is a regular
and prolic bearer with breless esh. Manjira (Rumani Neelum) is a
dwarf, regular and prolic bearer with good quality fruits.
The Paria Research Station in Gujarat developed three mango hybrids,
Neelphonso (Neelum Alphonso), Neeleshan Gujarat (Neelum Bane-
shan) and Neeleshwar (Neelum Dashehari). These hybrids are supe-
rior in TSS, total sugars and vitamin C, in addition to their dwarng habit,
with respect to their parents (Sachan et al., 1988).
Other countries
USA
Mango hybridization was reported from Hawaii in the 1920s, but no out-
standing problem appears to have been addressed or solved (Pope, 1929). A
number of crosses have been reported in Florida (Young and Ledin, 1954;
Sturrock, 1969), but all of the Florida cultivars are chance seedlings and none
came from controlled pollinations.
Israel
There is an extensive breeding programme in Israel aimed at producing
higher yielding cultivars with good quality, attractive fruit and with longer
harvest periods. Several hundred seedlings from open and controlled polli-
nations have been evaluated, and 14 of them have been identied as being of
interest (Lavi et al., 1993). The rootstock breeding programme is aimed at
developing rootstocks resistant to or tolerant of soil stresses, i.e. calcareous
soils, saline irrigation water and heavy non-aerated soils that predominate in
the mango-growing regions of Israel. Several interesting mono embryonic and
polyembryonic rootstocks have been selected (Lavi et al., 1993), but none has
performed better than 13-1, the currently preferred rootstock in Israel (Gazit
and Kadman, 1980).
Australia
A breeding programme to develop a new cultivar which retains the charac-
teristic avour of Kensington, but with improved productivity, greater dis-
ease resistance, enhanced skin colour and better postharvest performance,
Breeding and Genetics 87
was initiated in Queensland, Australia. These features are found in many
Florida cultivars (i.e. Irwin, Sensation and Tommy Atkins) which are
being used as maternal parents in crosses with Kensington (Whiley et al.,
1993). Promising hybrids have been identied in crosses involving Sensa-
tion, for example Calypso (see Knight et al., Chapter 3, this volume).
Calypso has increased shelf life, firmer fruit, extra blush for cosmetic
appeal, a higher flesh-to-seed ratio and consistent yields of high-quality fruit.
The Australian mango breeding programme was strengthened since 1994 by
launching a major effort involving various organizations located in different
agro-climatic zones in hybrid production, as well as regional testing.
Brazil
Breeding has been initiated in the tropical savannah of Brazil to develop cul-
tivars that are dwarf and with good quality fruit. Hybridizations have
involved local, Indian and Florida cultivars. Amrapali and Imperial were
good male parents to confer dwarng in the progeny (Pinto and Byrne, 1993).
Out of 2088 seedlings in the eld, 209 seedlings were selected in the rst
year and 42 of these were later identied as promising, from which four have
been released as new cultivars (Pinto et al., 2004). These four are: Alfa (Mal-
lika Van Dyke), which is semi-dwarf, high yielding and regular bearing;
Beta (Amrapali Winter), high yielding and moderately resistant to
anthracnose and Oidium; Roxa (Amrapali Tommy Atkins), with excel-
lent fruit quality; and Lita (Amrapali Tommy Atkins), high yielding
with excellent fruit quality.
South Africa
The South African breeding programme at the Citrus and Subtropical Fruit
Research Institute (CSFRI) is based on introductions, open-pollination and
mass selection. Four new cultivars have been released: Heidi, Neldawn,
Neldica and Ceriese. In addition, 12 promising selections have been iden-
tied for further evaluation (Marais, 1992).
4.10 Mutations
Somatic mutations
Asexual propagation enables the preservation of accumulated mutations
(macro and micro), which would normally be eliminated during sexual
propogation. In many fruit crops, bud mutations and chimeras occur rather
frequently and can provide an additional source of variability for selection.
However, such reported instances are relatively few in mango. Roy and
Visweswariya (1951) observed mutants of Puthi in which the number of
palisade cell layers differed from the original cultivar. Naik (1948) observed
signicant variation among trees of the same clone with respect to fruit shape,
size, colour and quality, which was ascribed to bud mutations. Davis Haden,
a sport of Haden, is larger than Haden and its season of maturity is about
C.P.A. Iyer and R.J. Schnell 88
a month earlier (Young and Ledin, 1954). Rosica from Peru, is a bud mutant
of Rosado de lca. Unlike its parent, Rosica is high yielding and regular
bearing, and does not produce seedless fruits (Medina, 1977).
Oppenheimer (1956), after a survey of many orchards in India, reported
wide variability in the performance of trees of the same clone within a single
orchard. Mukherjee et al. (1983) conducted a survey of mangoes in eastern
India and identied some superior clones. Singh and Chadha (1981), in a
study of orchards of Dashehari, located four clones which were superior in
performance. Singh et al. (1985) isolated two high-yielding clones from
orchards of Langra. Within Kensington, strains have also been identied
that show improved resistance to bacterial black spot (Whiley et al., 1993).
Roy (1950) observed a mutant of Alphonso with respect to fruit shape,
and suspected it to be a mericlinal chimera. Pandey (1998) has described
seven clones of Alphonso: Alphonso Behat and Alphonso Bihar from Bihar,
Alphonso Batli, Alphonso Black and Alphonso Bombay from Maharashtra,
Alphonso Punjab from Punjab and Alphonso White or Bili Ishada from
the North Canara district of Karnataka. Rajput et al. (1996) assembled several
Dashehari variants and after 14 years of observation, reported that the clone
Dashehari 51 was superior with respect to yield and regular bearing. Other
somatic mutants include: Cardozo Mankurad with large fruits of attractive
colour and high yields from Mankurad of Goa; dwarf selections from the
Rumani and Bangalora (Ramaswamy, 1989); development of Paiyur, a
dwarf selection from Neelum (Vijaya Kumar et al., 1991); Rati Bangana-
palli and Nuzuvid from Banganapalli (Anonymous, 1999); and MA-1,
regular bearing and high yielding with resistance to spongy tissue from
Alphonso (Mukunda, 2003).
In Thailand, Chaikiattiyos et al. (2000) selected clone SKoo7 (now known
as Kaew Sisaket) from 320 Kaew plants; SKoo7 has higher yield and
superior quality. Jintanawongse et al. (1999) also made superior selections for
yield and fruit quality from Nam Dok Mai, Khiew Sawoey, Rad and
Nang Klang Wau and DNA ngerprints of all these clones were made for
comparison with the parental clone.
For these studies, it is important to conduct a replicated cultivar evalua-
tion trial against standard commercial cultivars to establish that these varia-
tions are stable and not due to environmental responses. The use of genetic
markers should be explored to conrm that the new clones are genetically
distinct from the original cultivar.
Induced mutations
Mutation induction using ionizing radiation was attempted by Siddiqui et al.
(1966). Siddiqui (1985) irradiated dormant buds of Langra with high doses
of rays, and grafted them onto 1-year-old seedlings. A bud graft exposed to
3.0 kR bore fruits which were heavier, larger and had a more cream-yellow
pulp than the control. This variability was stable over three seasons. Sharma
and Majumder (1988b) irradiated bud sticks, topworked them onto 10-year-old
Breeding and Genetics 89
seedlings, and found that dosages above 5 kR are lethal for mango and that
the lethal dose required for 50% mortality (LD
50
) lies between 2 and 4 kR.
Effective dosages of the chemical mutagens, ethane methyl sulfonate (EMS)
and N-nitroso methyl urea (NMU), were 1.5 and 0.05%, respectively. The
spectrum of mutations induced by physical and chemical mutagens was
observed to be more or less the same, indicating the high sensitivity of certain
loci. The mutants included dwarfness, changes in shape and serration of
leaves and in TSS content in Dashehari. As in other perennial crops, muta-
genesis techniques that can allow useful traits to be targeted, as well as
isolating mutated sectors from a chimera, are essential.
4.11 Breeding Potential of Wild Species
Bompard (1993; see Bompard, Chapter 2, this volume) has made a compre-
hensive study of the wild Mangifera species and enumerated their potential
use in breeding. Mangifera laurina, which has subglabrous and laxly owered
panicles and is well adapted to areas with perpetual wet climates, is resistant
to anthracnose. Mangifera orophila from Malaysia and M. dongnaiensis from
Vietnam are both restricted to mountain forests 10001700 m above sea level
and their hybrids with mango could extend cultivation into temperate zones.
Mangifera magnica is completely free of bres; M. rufocostata and M.
swintonioides have an off-season bearing habit; M. pajang (endemic to Borneo)
and some strains of M. foetida have good quality fruits. Similarly Wani from
Bali and Borneo, the best variety of M. caesia, has a distinctive taste. Mangifera
casturi from South Kalimantan is a prolic bearer with small, black, sweet
fruits having good potential. Mangifera altissima is reportedly resistant to
mango pests, such as hoppers, tip borers and seed borers (Angeles, 1991).
Sharma and Choudhury (1976) observed that wild Mangifera trees identied
in Tripura State (north-eastern India) were free of mango malformation. The
wild species could also contribute to higher productivity. Fairchild (1948)
observed that crosses between ve-stamened mango and the Indian mango
(only one fertile stamen) could produce hybrids having better pollinating
quality. The interspecic compatibility of these species with M. indica must
be veried before they can be utilized in hybridization programmes (as sug-
gested by Bompard, 1993; Kostermans and Bompard, 1993; see Bompard,
Chapter 2, this volume).
4.12 Conclusions
Until recently, all mango cultivars arose as chance seedlings or as seedling
selections from known mother trees. Enthusiasm for controlled hybridiza-
tion by means of hand pollination waned because of the tedious nature of the
task and heavy fruit drop, resulting in only very few hybrids. This low hybrid
population was inadequate for selection and hence not many outstanding
hybrids were obtained. However, improvements in pollinating techniques
C.P.A. Iyer and R.J. Schnell 90
and more rapid screening of hybrid populations have enabled the release of
many hybrid mango cultivars of commercial value. Because of the world
markets demand for mangoes with specic qualities, the synthesis of new
cultivars has become imperative. Rapid strides in molecular biology and in
other aspects of biotechnology have opened up new approaches in plant
breeding. The development of polymerase chain reaction (PCR)-based
genetic markers, specically microsatellites, and their application to classical
breeding offer tremendous potential for mango improvement. The develop-
ment of a saturated linkage map and the identication of QTL for important
traits will allow the implementation of a MAS programme. The introduction
of specic genes for disease resistance from cultivars and wild species into
popular cultivars should soon be a reality. Without resorting to these new
technologies, mango breeding will continue to be a slow process.
References
Adato, A., Sharon, D. and Lavi, U. (1995) Application of DNA ngerprints for identica-
tion and genetic analyses of mango (Mangifera indica) genotypes. Journal of the
American Society for Horticultural Science 120, 259264.
Angeles, D.E. (1991) Mangifera altissima. In: Verheij, E.W.M. and Coronel, R.E. (eds)
Plant Resources of South-East Asia No.2: Edible Fruits and Nuts. Pudoc-DLO,
Wageningen, the Netherlands, pp. 206207.
Anonymous (1984) Paclobutrazol, Plant Growth Regulators for Fruits. ICI, Fernhurst,
West Sussex, UK.
Anonymous (1999) Improvement and management of horticultural crops. Department
of Agricultural Research and Education (DARE)/Indian Council of Agricultural Re-
search (ICAR) Annual Report 198999. ICAR, New Delhi, India, pp. 4460.
Aron, Y., Czosnek, H. and Gazit, S. (1998) Polyembryony in mango is controlled by a
single dominant gene. HortScience 33, 12411242.
Ayala-Silva, T., Schnell, R.J., Meerow, A.W., Winterstein, M., Cervantes, C. and Brown,
J.S. (2005) Determination of color and fruit traits of half-sib families of mango
(Mangifera indica L.). Proceedings of the Florida State Horticultural Society 118,
253257.
Bompard, J.M. (1993) The genus Mangifera rediscovered: the potential contribution of
wild species to mango cultivation. Acta Horticulturae 341, 6977.
Brettell, R.I.S., Johnson, P.R., Kulkarni, V.J., Muller, W. and Bally, I.S.E. (2004) Inheri-
tance of fruit characters in hybrid mangoes produced through controlled pollina-
tion. Acta Horticulturae 645, 319326.
Brooks, A.J. (1912) Articial cross fertilization of the mango. West Indies Bulletin 12,
567569.
Campbell, R.J. and Campbell, C.W. (1993) Commercial Florida mango cultivars. Acta
Horticulturae 341, 5559.
Campbell, R.J. and Zill, G. (2006) Mango selection and breeding for alternative markets
and uses. (Abstract). The Eighth International Mango Symposium, 510 February,
Sun City, South Africa, p. 19.
Chacko, E.K., Kohli, R.R., Doreswamy, R. and Randhawa, G.S. (1974) Effect of 2-
chloroethyl phosphonic acid on ower induction in juvenile mango seedlings.
Physiologia Plantarum 32, 188190.
Breeding and Genetics 91
Chaikiattiyos, S., Kurubunjerdjit, R., Akkaravessapong, P., Rattananukul, S., Chueychum,
P. and Anuput, P. (2000) Improvement and evaluation of the selected Kaew Sisaket
mango in Thailand. Acta Horticulturae 509, 185192.
Cilliers, B., Human, C.F., Snyman, J.C. and Carstens, K. (1996) Strategies, progress and
results from the South African mango breeding programme. Acta Horticulturae
445, 241244.
Dag, A., Eisenstein, D., Degani, C., El-Batsri, R., Zelig, M., Ziv, G. and Gazit, S. (1997)
Tommy Atkins mango as pollenizer for Lily. Acta Horticulturae 455, 209216.
Dag, A., Degani, C. and Gazit, S. (2006) Gene ow in mango orchards and its impact
on yield. (Abstract). The Eighth International Mango Symposium, 510 February,
Sun City, South Africa, p. 23.
Degani, C., El-Batsri, R. and Gazit, S. (1990) Enzyme polymorphism in mango. Journal
of the American Society for Horticultural Science 115, 844847.
Degani, C., Cohen, M., El-Batsri, R. and Gazit, S. (1992) PGI isozyme diversity and its
genetic control in mango. HortScience 27, 252254.
Degani, C., Cohen, M., Reuvani, O., El-Batsri, R. and Gazit, S. (1993) Frequency and
characteristic of zygotic seedlings from polyembryonic mango cultivars, deter-
mined using isozymes as genetic markers. Acta Horticulturae 341, 7885.
Dijkman, N.J. and Soule, M.J., Jr (1951) A tentative method of mango selection. Pro-
ceedings of the Florida State Horticultural Society 64, 257.
Duval, M., Bunel, F.J., Sitbon, C. and Risterucci, A.M. (2005) Development of microsatellite
markers for mango (Mangifera indica L.). Molecular Ecology Notes 5, 823826.
Duval, M., Bunel, J., Sitbon, C., Risterucci, A.M., Calcabre, C. and Le Bellec, F. (2006)
Genetic diversity of Caribbean mangoes using microsatellite markers. (Abstract). The
Eighth International Mango Symposium, 510 February, Sun City, South Africa, p. 29.
Eiadthong, W., Yonemori, K., Sugiura, A., Utsunomiya, N. and Subhadrabandhu, S.
(1999) Identication of mango cultivars of Thailand and evaluation of their genetic
variation using the amplied fragments by simple sequence repeat-(SSR-) anchored
primers. Scientia Horticulturae 82, 5766.
Fairchild, D. (1948) The mango relative of Cochin China: those with ve stamen owers.
Proceedings of the Florida State Horticultural Society 61, 250255.
Florida Mango Forum (1951) Mango Studies. Florida Mango Forum, Stuart, Florida.
Galan Sauco, V. (1997) Mango world production (outside Israel, Egypt and India). Acta
Horticulturae 455, 1522.
Gazit, S. and Kadman, A. (1980) 13-1 mango rootstock selection. HortScience 15, 669.
Gunjate, R.T. and Burondkar, M.M. (1993) Parthenocarpic mango developed through
hybridization. Acta Horticulturae 341, 107111.
Gupta, J.H. (1976) Reaction of mango varieties to powdery mildew (Oidium mangiferae
Berthet) in Uttar Pradesh. Progressive Horticulture 8, 6364.
Hoda, M.N. and Ramkumar (1993) Improvement of mango. In: Proceedings of the Na-
tional Seminar on Irregular Bearing in Mango Problems and Strategies (1213 July
1991). Kamal Printing Press, Muzafarpur, Bihar, India, pp. 3435.
Honsho, C., Nishiyama, K., Eiadthong, W. and Yonemori, K. (2005) Isolation and char-
acterisation of new microsatellite markers in mango. Molecular Ecology Notes 5,
152154.
Human, C.F., Rheeder, S. and Sippel, A.D. (2006) New cultivars and hybrid selections
from the mango breeding programme of the Agricultural Research Council in South
Africa. (Abstract). The Eighth International Mango Symposium, 510 February, Sun
City, South Africa, p. 27.
Issarakraisila, M. and Considine, J.A. (1994) Effects of temperature on pollen viability in
mango cv. Kensington. Annals of Botany 73, 231240.
C.P.A. Iyer and R.J. Schnell 92
Iyer, C.P.A. (1991) Recent advances in varietal improvement in mango. Acta Horticultu-
rae 291, 109132.
Iyer, C.P.A. and Subramanyam, M.D. (1972) Possible role of embryo culture in mango
breeding. Indian Journal of Horticulture 29, 135136.
Iyer, C.P.A. and Subramanyam, M.D. (1979) Improvement of mango by selection and
hybridization. Annual Report of the Indian Institute of Horticultural Research. In-
dian Institute of Horticultural Research, Bangalore, India, p. 16.
Iyer, C.P.A. and Subramanyam, M.D. (1986) Creeping, a promising genotype for intro-
duction of dwarfness in mango. Indian Journal of Horticulture 43, 221223.
Iyer, C.P.A. and Subramanyam, M.D. (1987) Improvement of mango by selection and
hybridization. Annual Report of the Indian Institute of Horticultural Research. In-
dian Institute of Horticultural Research, Bangalore, India, p. 11.
Iyer, C.P.A. and Subramanyam, M.D. (1993) Improvement of mango. In: Chadha, K.L.
and Pareek, O.P. (eds) Advances in Horticulture. Vol.1. Malhotra Publishing, New
Delhi, India, pp. 267278.
Iyer, C.P.A., Subbaiah, M.C., Subramanyam, M.D. and Prasada Rao, G.S. (1989) Screen-
ing of germplasm and correlation among certain characters in mango. Acta Horti-
culturae 231, 8390.
Jintanawongse, S., Chunwongse, J., Chumpong, S. and Hiranpradit, H. (1999) Improve-
ment of existing commercial mango cultivars in Thailand. (Abstract). The Sixth In-
ternational Mango Symposium, 69 April, Pattaya, Thailand.
Juliano, J.B. (1934) Origin of embryos in the strawberry mango. Philippine Journal of
Science 54, 553563.
Kashkush, K., Jinggui, F., Tomer, E., Hillel, J. and Lavi, U. (2001) Cultivar identication
and genetic map of mango (Mangifera indica). Euphytica 122, 129136.
Knight, R.J., Jr (1980) Origin and world importance of tropical and sub-tropical fruits. In:
Nagy, S. and Shaw, P.E. (eds) Tropical and Sub-tropical fruits. Composition, Proper-
ties and Uses. AVI Publishing, Westport, Conneticut, pp. 120.
Knight, R.J. and Schnell, R.J. (1993) Mango (Mangifera indica) introduction, and evaluation
in Florida and its impact on the world industry. Acta Horticulturae 341, 125135.
Knight, R.J. and Schnell, R.J. (1994) Mango (Mangifera indica L.) introduction and eval-
uation in Florida and its impact on the world industry. Acta Horticulturae 341,
125135.
Kostermans, A.J.G.H. and Bompard, J.M. (1993) The Mango: their Botany, Nomencla-
ture, Horticulture and Utilization. Academic Press, London.
Kurian, R.M. and Iyer, C.P.A. (1992) Stem anatomical characters in relation to tree vigour
in mango. Scientia Horticulturae 50, 245248.
Lahav, E., Tomer, E., Gazit, S. and Lavi, U. (1995) Performance of avocado (Persea
americana Mill.) and mango (Mangifera indica L.) seedlings compared with their
grafted trees. Journal of the American Society for Horticultural Science 120,
265269.
Lavi, U., Tomer, E. and Gazit, S. (1989) Inheritance of agriculturally important traits in
mango. Euphytica 44, 510.
Lavi, U., Sharon, D., Tomer, E., Adato, A. and Gazit, S. (1993) Conventional and modern
breeding of mango cultivars and rootstocks. Acta Horticulturae 341, 145151.
Lavi, U., Tomer, E. and Hillel, J. (1998) Components of genetic variation and genetic
correlations of mango traits. Scientia Horticulturae 75, 1125.
Lavi, U., Kashkush, K., Saada, D., Shats, H., Ravid, U. and Tomer, E. (2004) Mango
breeding and the potential of modern biology. Acta Horticulturae 645, 5159.
Leroy, J.F. (1947) La polyembryonie chez les Citrus son interet dans la culture et lame-
lioration. Revue Internationale de Botanique Appliquee Paris 27, 483495.
Breeding and Genetics 93
Lopez-Valenzuela, J.A., Martinez, O. and Paredes-Lopez, O. (1997) Geographic differ-
entiation and embryo type identication in Mangifera indica L. cultivars using
RAPD markers. HortScience 32, 11051108.
Maheshwari, P. (1934) The Indian mango. Current Science 3, 9798.
Maheshwari, P. and Rangaswamy, N.S. (1958) Polyembryony and in vitro culture of
embryos of Citrus and Mangifera. Indian Journal of Horticulture 15, 275282.
Majumder, P.K. and Sharma, D.K. (1990) Mango. In: Bose, T.K. and Mitra, S.K. (eds)
Fruits: Tropical and Subtropical. Naya Prokash, Calcutta, pp. 162.
Majumder, P.K., Singh, R.N., Sharma, D.K. and Mukerjee, S.K. (1972) Preliminary stud-
ies on inheritance in Mangifera indica L. Acta Horticulturae 24, 101106.
Majumder, P.K., Sharma, D.K. and Singh, R.N. (1981) Breeding for dwarfness in mango
(Mangifera indica L.). National Symposium on Tropical and Subtropical Fruit Crops,
Horticultural Society of India, Bangalore, p. 3.
Malik, P.C. (1951) Morphological and biology of the mango ower. Indian Journal of
Horticulture 14, 123.
Marais, Z. (1992) Mango evaluation for breeding. Citrus and Subtropical Fruit Research
Institute (CSFRI) Information Bulletin 234, 7.
Marshall, T.C., Slate, J., Kruuk, L. and Pemberton, J.M. (1998) Statistical condence for
likelihood-based paternity inference in natural populations. Molecular Ecology 7,
639655.
Medina, J.P. (1977) Rosica a new mango variety selected in Ica, Peru. Fruit Varieties
Journal 31, 8889.
Moore, G.A. and Castle, W.S. (1988) Morphological and isozymic analysis of open-
pollinated citrus rootstock population. Journal of Heredity 79, 5963.
Morton, J.F. (1987) Fruits of Warm Climates. Creative Resource Systems, Winterville,
North Carolina, pp. 221237.
Mukherjee, S.K. (1950) Mango; its allopolyploid nature. Nature 166, 196197.
Mukherjee, S.K. (1953) The mango its botany, cultivation, uses and future improve-
ments, especially as observed in India. Economic Botany 7, 130162.
Mukherjee, S.K. (1957) Cytology of some Malayan species of Mangifera. Cytologia 22,
239241.
Mukherjee, S.K. (1963) Cytology and breeding of mango. Punjab Horticultural Journal
3, 107115.
Mukherjee, S.K., Singh, R.N., Majumder, P.K. and Sharma, D.K. (1968) Present position
regarding breeding of mango (Mangifera indica L.) in India. Euphytica 17,
462467.
Mukherjee, S.K., Chakraborty, S., Sadhukhan, S.K. and Saha, P. (1983) Survey of man-
goes of West Bengal. Indian Journal of Horticulture 40, 713.
Mukunda, G.K. (2003) Studies on the performance of certain clones of mango cv. Al-
phonso. PhD thesis, University of Agricultural Sciences, Bangalore, India.
Naik, K.C. (1948) Improvement of the mango (Mangifera indica L.) by selection and
hybridization. Indian Journal of Agricultural Science 18, 3541.
Oppenheimer, C. (1956) Study tour report on subtropical fruit growing and research in
India and Ceylon. Special Bulletin No.3 State of Israel Ministry of Agriculture. Ag-
ricultural Research Station, Rehovot, Israel.
Oppenheimer, C. (1967) Nimrod a new mango variety selected in Israel. Proceedings
of the Florida State Horticultural Society 80, 358359.
Pandey, S.N. (1998) Mango cultivars. In: Srivastava, R.P. (ed.) Mango Cultivation. Inter-
national Book Distributing Co., Lucknow, India, pp. 3999.
Pinto, A.C.Q. and Byrne, D.H. (1993) Mango hybridization studies in tropical savannah
(Cerrados) of Brazil Central region. Acta Horticulturae 341, 98106.
C.P.A. Iyer and R.J. Schnell 94
Pinto, A.C.Q., Andrade, S.R.M., Ramos, V.H.V. and Cordeiro, M.C.R. (2004) Intervarietal
hybridization in mango: techniques, main results and their limitations. Acta Horti-
culturae 645, 327331.
Pope, W.T. (1929) Mango Culture in Hawaii. Hawaii Agricultural Experiment Station
Bulletin 58. Hawaii Agricultural Experiment Station, Hawaii.
Popenoe, W. (1917) The Pollination of the Mango. US Department of Agriculture (USDA)
Bulletin 542. USDA, Washington DC.
Popenoe, W. (1920) The mango. In: Manual of Tropical and Subtropical Fruits. The Mac-
millan Co., New York, p. 135.
Prakash, O. and Srivastava, K.C. (1987) Mango Diseases and their Management a
World Review. Today and Tomorrows Printers, New Delhi, India.
Rajput, M.S., Chadha, K.L. and Negi, S.S. (1996) Dashehari 51, a regular bearing and
high yielding clone of mango cv. Dashehari. (Abstract). The Fifth International
Mango Symposium, 16 September, Tel Aviv, Israel, p. 42.
Ram, N., Kamalwanshi, R.S. and Sachan, I.P. (1987) Studies on mango malformation.
Indian Journal of Mycology and Plant Pathology 17, 2933.
Ram, S., Bist, L.D., Lakhanpal, S.C. and Jamwal, I.S. (1976) Search of suitable pollinizer
for mango cultivars. Acta Horticulturae 57, 253263.
Ramaswamy, N. (1989) Survey and isolation of plus trees of mango. Acta Horticulturae
231, 9396.
Ravishankar, K.V., Chandrasekhar, P., Sreedhar, S.A., Dinesh, M.R., Anand, L. and Saip-
rasad, G.V.S. (2004) Diverse genetic bases of Indian polyembryonic and monoem-
bryonic mango cultivars. Current Science 87, 870871.
Rossetto, C.J., Bortoletto, N., Carvalho, C.R.L., Walder, J.M.M., Nogueira, N.L., Arthus,
V. and Lopes, L.A. (2006) Mango resistance to fruit ies: 1. Varietal selection and
mechanism of resistance. (Abstract). The Eighth International Mango Symposium,
510 February, Sun City, South Africa, p. 22.
Roy, B. (1950) A mango chimera. Current Science 19, 93.
Roy, B. and Visweswariya, S.S. (1951) Cytogenetics of mango and banana. Report of
Maharashtra Association for Cultivation of Science, Pune, India.
Roy, R.S., Mallik, P.C. and Sinha, R.P. (1956) Mango breeding in Bihar, India. Proceed-
ings of the American Society for Horticultural Science 68, 259264.
Ruehle, G.D. and Ledin, R.B. (1956) Mango Growing in Florida. Florida Agricultural
Experiment Station Bulletin 574. Florida Agricultural Experiment Station, Florida.
Sachan, S.C.P., Katrodia, J.S., Chundawat, B.S. and Patel, M.N. (1988) New mango hy-
brids from Gujarat. Acta Horticulturae 231, 103105.
Sahijram, L., Bollamma, K.T., Naren, A., Soneji, J.R., Dinesh, M.R. and Halesh, G.K.
(2005) In vitro hybrid embryo rescue in mango (Mangifera indica L.) breeding. In-
dian Journal of Horticulture 62, 235237.
Salvi, M.J. and Gunjate, R.T. (1988) Mango breeding work in Konkan region of Maha-
rashtra state. Acta Horticulturae 231, 100102.
Schnell, R.J. and Knight, R.J., Jr (1992) Frequency of zygotic seedlings from ve polyem-
bryonic mango rootstocks. HortScience 27, 174176.
Schnell, R.J. and Knight, R.J., Jr (1998) Phenology of owering among different
mango cultivars. Proceedings of the Florida State Horticultural Society 111,
320321.
Schnell, R.J., Ronning, C.M. and Knight, R.J., Jr (1995) Identication of cultivars and
validation of genetic relationships in Mangifera indica L. using RAPD markers.
Theoretical and Applied Genetics 90, 269271.
Schnell, R.J., Olano, C.T., Quintanilla, W.E. and Meerow, A.W. (2005) Isolation and
characterization of 15 microsatellite loci from mango (Mangifera indica L.) and
Breeding and Genetics 95
cross-species amplication in closely related taxa. Molecular Ecology Notes 5,
625627.
Schnell, R.J., Brown, J.S., Olano, C.T., Meerow, A.W., Campbell, R.J. and Kuhn, D.N.
(2006) Mango genetic diversity analysis and pedigree inferences for Florida culti-
vars using microsatellite markers. Journal of the American Society for Horticultural
Science 131, 214224.
Schnell, R.J., Brown, J.S., Kuhn, D.N., Cervantes-Martinez, C., Borrone, J.W., Olano, C.T.,
Motamayor, J.C., Phillips, W., Johnson, E., Monteverde-Penso, E.J., Amores, F. and
Lopes, U. (2007) Current challenges of tropical tree crop improvement: integrating ge-
nomics into an applied cacao breeding program. Acta Horticulturae 738, 129144.
Sen, P.K., Mallik, P.C. and Ganguly, B.D. (1946) Hybridization of the mango. Indian
Journal of Horticulture 4, 415.
Sharma, D.K. (1987) Mango breeding. Acta Horticulturae 196, 6167.
Sharma, D.K. and Choudhury, S.S. (1976) Occurrences of an unknown wild race of
Mangifera in Tripura. Current Science 45, 305306.
Sharma, D.K. and Majumder, P.K. (1988a) Further studies on inheritance in mango. Acta
Horticulturae 231, 106111.
Sharma, D.K. and Majumder, P.K. (1988b) Induction of variability in mango through
physical and chemical mutagens. Acta Horticulturae 231, 112116.
Sharma, D.K. and Singh, R.N. (1970) Self incompatibility in mango (Mangifera indica
L.). Horticultural Research 10, 108115.
Sharma, D.K. Majumder, P.K. and Singh, R.N. (1972) Inheritance pattern in mango
(Mangifera indica L.). In: Proceedings of the Symposium on Recent Advances in
Horticulture. Uttar Pradesh Institute of Agricultural Sciences, Kanpur, Uttar Pradesh,
India, pp. 6668.
Sharma, R.R. (2003) Catecholase and cresolase activity as biochemical index for screen-
ing mango seedlings at nursery stage for bearing behaviour in their future reproduc-
tive life. Plant Genetic Resources (PGR) Newsletter 133, 3134.
Sharma, R.R., Singh, C.N., Chbonkar, P., Goswami, A.M. and Singh, S.K. (2000) Poly-
phenol oxidase activity as an index for screening mango (Mangifera indica) germ-
plasm against malformation. Plant Genetic Resources (PGR) Newsletter 124, 4143.
Siddiqui, S.H. (1985) Induced somatic mutation in mango. Mangifera indica L. cv.
Langra. Pakistan Journal of Botany 17, 7579.
Siddiqui, S.H., Mujeeb, K.A. and Vati, S.M. (1966) Evolution of new varieties of mango
(Mangifera indica L.) through induced somatic mutations by ionizing radiations. In:
Proceedings of the First Agricultural Symposium. Atomic Energy Commission, Dac-
ca, Bangladesh, pp. 3437.
Singh, H. and Chadha, K.L. (1981) Improvement of Dashehari by clonal selection. (Ab-
stract). National Symposium on Tropical and Subtropical Fruit Crops. Horticultural
Society of India, Bangalore, p. 5.
Singh, L.B. (1960) The Mango Botany, Cultivation and Utilisation. Leonard Hill, London.
Singh, L.B. (1963) A new technique for inducing early fruiting in mango hybrids based
on movement of hormones. Proceedings of the Florida State Horticultural Society
75, 410412.
Singh, L.B. (1969) Mango. In: Ferwerda, F.P. and Wit, F. (eds) Outlines of Perennial
Crop Breeding in the Tropics. Veenen and Zonen, Wageningen, the Netherlands,
pp. 309327.
Singh, R.N. (1954) Studies on oral biology and subsequent developments of fruit in the
mango varieties, Dashehari and Langra. Indian Journal of Horticulture 11, 6988.
Singh, R.N. (1990) Mango. Series No. 3. Indian Council of Agricultural Research (ICAR),
New Delhi, India.
C.P.A. Iyer and R.J. Schnell 96
Singh, R.N., Majumder, P.K. and Sharma, D.K. (1962) Self-incompatibility in mango var.
Dashehari. Current Science 31, 209.
Singh, R.N., Majumder, P.K., Sharma, D.K. and Mukherjee, S.K. (1972) Some promising
mango hybrids. Acta Horticulturae 24, 117119.
Singh, R.N., Sharma, D.K. and Majumder, P.K. (1980) An efcient technique of mango
hybridization. Scientia Horticulturae 12, 299301.
Singh, R.N., Gorakh, S., Rao, O.P. and Mishra, J.S. (1985) Improvement of Banarsi
Langra through clonal selection. Progressive Horticulture 17, 273277.
Singh, S.N. and Singh, S.P. (1952) Studies on the storage and longevity of some fruits and
vegetables. Journal of Agriculture and Animal Husbandry Uttar Pradesh 2, 311.
Slor, E. and Gazit, S. (1982) Tahar, a new mango cultivar. Alon Honotea 36, 807. (In
Hebrew)
Spencer, J.L. and Kennard, W.C. (1955) Studies on mango fruit set in Puerto Rico. Tropi-
cal Agriculture 32, 323330.
Sturrock, D. (1969) Final report on some mango hybrids 1969. Proceedings of the
Florida State Horticultural Society 82, 318321.
Sturrock, T.T. (1968) Genetics of mango polyembryony. Proceedings of the Florida State
Horticultural Society 81, 311314.
Tomer, E., Lavi, U., Degani, C. and Gazit, S. (1993) Noami: a new mango cultivar.
HortScience 28, 755756.
Truscott, M. (1992) Biochemical screening of polyploidy mango seedlings. Citrus and
Subtropical Fruit Research Institute (CSFRI) Information Bulletin 237, 1718.
Vijaya Kumar, M., Ramaswamy, N. and Rajagopalan, R. (1991) Exploiting natural vari-
ability in mango. In: Proceedings of the National Seminar on Irregular Bearing in
Mango Problems and Strategy, Pusa, Bihar, India. Rajendra Agricultural Univer-
sity, Sabour, Bihar, India, pp. 5556.
Viruel, M.A., Escribano, P., Barbieri, M., Ferri, M. and Hormaza, J.I. (2005) Fingerprint-
ing, embryo type and geographic differentiation in mango (Mangifera indica L.,
Anacardiaceae) with microsatellites. Molecular Breeding 15, 383393.
Whiley, A.W., Mayers, P.E., Saranah, J. and Bartley, J.P. (1993) Breeding mangoes for
Australian conditions. Acta Horticulturae 341, 136145.
Yee, W. (1958) The Mango in Hawaii. Hawaii Agricultural Experiment Station Series
Circular No. 388. Hawaii Agricultural Experiment Station, Hawaii.
Young, T.W. and Ledin, R.B. (1954) Mango breeding. Proceedings of the Florida State
Horticultural Society 67, 241244.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 97
5 Reproductive Physiology
T.L. Davenport
University of Florida, Florida, USA
5.1 Introduction 98
5.2 Phenology 99
5.3 Shoot Development 100
Vegetative shoots 102
Reproductive shoots 104
5.4 Flowering Mechanisms 105
Shoot initiation 105
Induction 106
Florigenic promoter (FP) or stimulus 108
Vegetative promoter (VP) 110
5.5 Environmental Inuence on Vegetative and Reproductive Development 111
Temperature 111
Water relations 113
Effect of N on owering 114
Photoperiod 116
5.6 Hormonal Inuence on Flowering 116
Ethylene 116
Auxin 117
Cytokinins 118
Gibberellins 119
Plant growth retardants 121
5.7 Photoassimilate Inuence on Flowering 123
5.8 Horticultural Manipulation of Flowering 123
5.9 Conceptual Flowering Models 124
Carbohydrate-regulated owering models 124
Hormone-regulated owering models 127
5.10 Floral Management 133
5.11 Floral Biology 134
Sex ratio 134
Environmental determinants of sex ratio 134
Physiological determinants of sex ratio 135
T.L. Davenport 98
Anthesis and dehiscence 136
Pollen 136
Pollination 137
5.12 Fruit Development 139
5.13 Stenospermocarpy 139
5.14 Fruit Set and Retention 140
Sex ratio 140
Mineral nutrients 141
Hormonal control 141
5.15 Alternate Bearing 145
5.16 Conclusions 145
5.1 Introduction
Flowering and fruit set are the most critical of all events occurring after estab-
lishment of a tree crop. Given favourable growth conditions, the timing and
intensity of owering greatly determine when and how much fruit are pro-
duced. Many important details about owering are becoming clearer, espe-
cially in herbaceous plants, at the physiological, biochemical and molecular
levels (see reviews by Searle, 1965; Zeevaart, 1976, 2006; Bernier et al., 1981,
1993; Halevy, 19851986; Bernier, 1988; Kinet, 1993; Boss et al., 2004; Komeda,
2004; Putterill et al., 2004; Corbesier and Coupland, 2005).
Cool temperatures in the subtropics stimulate mango owering and age
of the last vegetative ush has an important bearing on its ability to ower in
marginally cool or warm temperatures of the tropics (van der Meulen et al.,
1971; Davenport, 2000, 2003). Consequently, mango owering can be en-
hanced during its normal season or manipulated to occur at other times of
the year in the tropics. For example, potassium nitrate (KNO
3
) can stimulate
out-of-season owering in mangoes in tropical latitudes (Barba, 1974; Nez-
Elisea, 1985; Davenport, 1993; Protacio, 2000), although this treatment has
not always been dependable. Various aspects of mango owering and/or
fruit set have been reviewed (Singh, 1958a, 1979; L.B. Singh, 1960, 1977;
Chacko, 1986, 1991; Chadha and Pal, 1986; Davenport, 1993, 2000, 2003; Dav-
enport and Nez-Elisea, 1997; Singh et al., 2005), and M.J. Soule (1950) pub-
lished an extensive annotated bibliography of the older literature related to
mango reproduction.
Understanding mango owering is essential to efciently utilize man-
agement systems that extend the owering and crop production seasons.
Recent studies of mango owering have resulted in conceptual models that
help explain the physiological basis of owering (Chacko, 1991; Cull, 1991;
Kulkarni, 1991, 2004; Whiley et al., 1991; Davenport and Nez-Elisea, 1997;
Davenport, 2000, 2003). Control of owering allows growers to harvest their
crops at the most protable times. Increasing the season of availability
improves competitiveness in the international marketplace, and promotes
the most efcient use of resources as costs of inputs continue to rise.
This chapter addresses the physiology of mango owering, early fruit set
and retention. Cultivar names and type of embryony (i.e. monoembryonic or
Reproductive Physiology 99
polyembryonic) are purposefully left out to focus on the physiological
aspects of reproduction regardless of whether they are tropically or subtrop-
ically adapted or from Indian or South-east Asian origin. Cultivars are
selected for their productivity in specic environments. Transfer of a cultivar
to a different environment often results in some alteration in performance. It
is reasonable to assume that the underlying mechanisms by which all culti-
vars respond to their environment within the framework of their genetic limi-
tations are similar. The concepts described herein, therefore, apply to all cultivars,
regardless of origin.
5.2 Phenology
Growth of mango and other tropical trees is not continuous (Nakasone et al.,
1955; Halle et al., 1978; Verheij, 1986; Davenport, 1993, 2000, 2003). Apical
buds spend most of the time in rest. Growth occurs as intermittent, ephem-
eral ushes of shoots from apical or lateral buds (Naik and Mohan Rao, 1942;
Singh, 1958a, b). Stems are quiescent or resting terminal vegetative structures
on branches from which shoot growth occurs. Shoots are elongating vegeta-
tive or reproductive structures that emerge from apical or lateral buds of
stems. Vegetative shoots develop a prescribed number of nodes during
growth before entering a resting state as a stem. Depending on environment,
periods of stem rest are generally short in young plants but usually last sev-
eral months between episodes of growth in mature trees. Vegetative growth
generally occurs up to three or four times a year on individual branches,
depending upon cultivar and growth conditions.
Development of the vegetative shoot from initiation of growth to full
elongation requires 36 weeks, depending on the cultivar and climatic condi-
tions (Whiley et al., 1991). During this period, 1020 new leaves are generally
produced before returning to a resting state. These rhythmic episodes of
extension growth are recorded on each branch as segments consisting of
compressed internodes interspersed with long internodes, that is articulate
growth (Tomlinson and Gill, 1973). Davenport (1992, 2003, 2006) referred to
regions of compressed internodes as intercalations and the entire segment of
long internodes terminating in an intercalation as an intercalary unit. The
number of intercalations between each branching point indicates the number
of vegetative growth episodes or ushes that have occurred between each
owering ush.
Flushes of vegetative growth occur on groups of stems borne on scaffold-
ing branches in isolated sections of tree canopy. Flushing stems are usually
connected at some common branch point within the tree limbs. Asynchro-
nous ushes of growth at various times in random portions of a tree canopy
may appear to be continuous growth but are simply ushes occurring in
various parts of the total canopy over time. Flowering ushes generally occur
after extended periods of stem rest in the low-latitude tropics or during cool
winter months in the high-latitude tropics and subtropics. Like vegetative
ushes, reproductive ushes are usually asynchronous in tropical climates
T.L. Davenport 100
(Verheij, 1986). In the subtropics, however, trees exposed to cold tempera-
tures (310C) display synchronized owering ushes throughout the tree
canopy approximately 1 month later. Subsequent vegetative ushes also tend
to be synchronous for one or two growth cycles depending upon the number
of retained fruit. Less intense, cool weather (1018C), however, results in
asynchronous reproductive ushes in responsive stems as is typical of trees
growing in the tropics. The timing of owering ushes of cultivars in various
locations has been reviewed by L.B. Singh (1960), Chadha and Pal (1986) and
Pandey (1989). Variations in owering patterns occur in all cultivars depend-
ing on their age and whether they are growing in dry or humid tropics or
subtropics (L.B. Singh, 1960).
5.3 Shoot Development
Flushes of vegetative extension growth of mango stems terminate with for-
mation of determinate panicles. Several weeks to a few months after separa-
tion of the last ower or fruit from these panicles are required for the central
axis of the panicle or rachis to dry and mechanically separate from the sup-
porting stem, depending on the longevity of attached fruit. Five to ten lateral
vegetative shoots typically develop from axillary buds located at the termi-
nal intercalation positioned in a compact whorl surrounding the panicle scar
of each stem (see Fig. 1 of Reece et al., 1949). These lateral shoots become the
branch points of stems. These branching shoots form 1015 leaves before the
apical buds return to a resting state to establish them as individual stems. Ini-
tiation of these lateral vegetative shoots may occur 23 months after desicca-
tion of panicles which fail to set fruit. Fruit-bearing stems do not initiate new
lateral shoots until several months after separation of fruit and rachis from the
stem (Kulkarni and Rameshwar, 1989). Such delayed vegetative growth can
reduce the potential for new shoots to ower during the next owering sea-
son (Singh and Khan, 1939; L.B. Singh, 1960, 1972; Monselise and Gold-
schmidt, 1982). The apical bud of stems is at rest for most of the year in mature
trees. Stems on centennial trees typically produce only one vegetative ush
during the year (N. Golez, personal communication, the Philippines, 1989).
The apical resting bud of each newly established lateral stem (intercalary
unit) is surrounded by a compact whorl of 1012 leaves with short inter-
nodes (intercalation) (Fig. 5.1). Protective bud scales are green but may be
brown at the tips due to desiccation (Sen and Mallik, 1941; Mustard and
Lynch, 1946; Singh, 1958b; Ravishankar et al., 1979). Resting buds possess a
number of pre-formed nodes, each of which contains a leaf bract or leaf pri-
mordium and a lateral meristem (Fig. 5.2; see Figs 712 in Chaikiattiyos et al.,
1994). The outermost, proximally located dried leaf bracts (bud scales) pro-
tect the more distal interior leaf bracts, leaf primordia and lateral meristems
from mechanical damage and desiccation. Leaf bracts are vestigial non-
developed leaves. Scales abort upon evocation of new shoots. Proximally
located bracts in apical buds fail to further develop beyond some enlarge-
ment and also abort with elongation of shoots.
Reproductive Physiology 101
If apical buds are initiated during vegetatively inductive conditions,
bracts develop as small leaves and the leaf primordia develop as the full-
sized leaves of vegetative shoots. Additional leaves result from nodes formed
by renewed activity of the apical meristem. The number of leaves (nodes) is
dependent upon the mean temperature during initiation, and increases as
temperatures rise (Whiley et al., 1989). The lateral meristems of the apical
bud develop as axillary buds at the base of petioles in the elongating vegeta-
tive shoot, each bearing protective bracts, leaf primordia and lateral mer-
istems (Fig. 5.3; see Fig. 1 of Reece et al., 1949).
In contrast, if shoot growth is initiated under oral inductive conditions,
the leaf bracts and primordia fail to fully develop, but the lateral meristems
Fig. 5.1. Apical bud of resting mango stem.
Leaf primordia (bracts)
Apical dome
(includes meristem)
Lateral meristematic
primordia
Fig. 5.2. Stylized cross-section of apical bud showing positions of apical meristem,
lateral meristems and leaf primordia.
T.L. Davenport 102
begin to elongate and branch at each node forming secondary, tertiary and
quaternary lateral meristems. Each branch point in the lateral inorescence
from the panicle axis to the oral pedicels bears a oral bract (i.e. partially
developed vestigial leaf primordium) (Fig. 5.4). The distal half of the panicle
structure is derived from newly formed nodes laid down by cell divisions in
the apical meristem prior to returning to a resting state. Mixed shoots, bear-
ing both leaves and inorescences at each node, result from development of
both the primary leaf primordia and the lateral meristems, which form the
inorescences in the same nodes as leaves.
Vegetative shoot induction, thus, involves stimulating development of
leaf primordia from resting buds while repressing development of lateral
meristems. Leaf primordia then follow a predetermined cascade of genetic
signals resulting in leaf development at each node. Because all shoots emerge
from resting buds, a vegetatively induced event does not involve simply
inhibition of owering. The putative inductive signal directing differentia-
tion of leaf primordia onto leaves upon initiation is termed a vegetative pro-
moter (VP) rather than a oral inhibitor.
Shoots bearing only inorescences (generative shoots) result from induc-
tive development of lateral meristems and suppression of leaf primordial
development. A predetermined cascade of owering gene signals is activated
in lateral meristems resulting in lateral cymose inorescences terminating
with owers. A distinct origenic promoter (FP) may be responsible for spe-
cic activation of the lateral meristems of mango. Mixed shoot induction
results in combined development of leaf primordia and lateral meristems.
Vegetative shoots
Vegetative shoots bear only leaves (Fig. 5.5). The anatomy of mango vege-
tative shoot development has been described (Singh, 1958b; Chaikiattiyos
Stem
Fig. 5.3. Axillary bud of resting mango stem. Leaf petioles (arrows).
Reproductive Physiology 103
et al., 1994). Vegetative shoots may arise either from axillary buds, if no apical
bud exists due to owering in the previous ush, or from the apical bud
when present. The latter is considered extension growth or addition of an
intercalary unit on the existing stem, but the developmental events during
shoot formation from either apical or lateral buds are basically the same.
Cells in the leaf primordia of initiating buds begin to form individual leaves
in the proximal portion of the vegetative shoot. Soon thereafter, the apical
meristem activates to form more nodes bearing leaf primordia and lateral
meristems. These newly formed leaf primordia develop as the distal portion
of the vegetative shoot if environmental conditions remain vegetatively
inductive (Nez-Elisea et al., 1996). Newly elongating vegetative shoots are
green in most cultivars but may be bronze or red in others. Fully expanded
1
Pedicel
1
Pedicel
1 Bract
2 Bract
3 Bract
2 Bract
1 Bract
2
Pedicel
1
Pedicel
1
1
Pedicel
1 Bract
4
2
3
5

1
Axis
Axis
Fig. 5.4. Diagram and photos of mango inorescence depicting the panicle axis and
primary (1), secondary (2) and succeeding levels of pedicel and cymose oral archi-
tecture. Vestigial leaf promorida (oral bracts) are depicted at the base of each level of
pedicel architecture.
T.L. Davenport 104
leaves are a shade of red, depending upon cultivar and cultural conditions
and are thin and limp from lack of lignication. The apical buds of vegetative
shoots generally become quiescent before completion of the limp, red-leaf
stage (Nez-Elisea and Davenport, 1995). Internodes are compressed at the
apex, and leaf development is arrested thereby forming a bud with protec-
tive outer scales, inner leaf primordia, lateral meristems and the apical mer-
istem. Fully expanded leaves become light green and stiff as they become
lignied and suberized. Vegetative shoots are mature when leaves become
dark green, which occurs when they are c.2 or 3 months old.
Reproductive shoots
Two types of reproductive shoots typically occur in mango. Generative
shoots display only owers and have oral bracts or non-developed leaves
at the base of each lateral inorescence (Fig. 5.5). Terminal inorescences,
i.e. panicles or thyrsoids (Weberling, 1989), develop from dormant apical
buds. The anatomy of panicle development has been described (Juliano
and Cuevas, 1932; Musahib-ud-din, 1946; Mustard and Lynch, 1946; Singh,
VEGETATIVE GENERATIVE MIXED CHIMERIC V/F TRANSITION F/V TRANSITION
GENERATIVE CHIMERIC V/F TRANSITION VEGETATIVE MIXED F/V TRANSITION
Fig. 5.5. Stylized diagrams and photomontage of shoot types found in mango. Transition
shoots shift from vegetative to oral (V/F) or oral to vegetative (F/V). Arrow ( )
represents individual leaves; oral diagram ( ) represents lateral inorescences.
Reproductive Physiology 105
1958b; L.B. Singh, 1960; Sturrock, 1966; Ravishankar et al., 1979; Scholeeld,
1982; Scholeeld et al., 1986). The complexes of primary to quaternary branch-
ing lateral structures of the inorescence each terminate with three cymose
owers. The terminal ower opens rst, followed by two subtending lateral
owers. These complexes form the lateral inorescence structures emerging
from the central axis of the panicle. The central axis extension also terminates
in a similar fashion. Morphological stages of oral buds and panicle develop-
ment were described by Shu (1981) and Oosthuyse (1991a). Reece et al. (1949)
described the development of inorescences initiated in lateral buds when
the terminal bud is missing. There are more nodes in dormant apical buds
and their bracts are more developed than in axillary buds; however, oral
evocation is indistinguishable.
Generative shoot development in apical buds initially involves swelling
of the lateral meristems and their bud scales. Each axillary meristem devel-
ops as an inorescence on a primary peduncle. The apical meristem then
forms new lateral meristems and leaf primordia for the distal portion of pan-
icle development if oral inductive conditions persist (Nez-Elisea et al.,
1996). Panicles may be open or compact, depending upon internode elonga-
tion, which is cultivar dependent (L.B. Singh, 1960), but the architecture gen-
erally conforms to that in Fig. 5.5. Mixed shoots develop under weak oral
inductive conditions (i.e. in the low-latitude tropics). Both leaves and pri-
mary pedunculate inorescences develop from the same nodes (Fig. 5.5).
Leaf primordia and lateral meristems develop as leaf and oral structures,
respectively.
5.4 Flowering Mechanisms
Mango stems undergo varying periods of rest between episodes of growth,
depending on tree age and environmental inuences. Resting mango buds
must, therefore, respond to two distinctly different signals for shoots to occur.
The rst signal initiates growth of the shoot and the second determines if it
will be vegetative or reproductive. The signals that regulate initiation of
shoot growth in resting buds differ from the inductive signals that regulate
shoot type.
Shoot initiation
Initiation is the onset of shoot development, regardless of the type of shoot
evoked. It involves cell division and elongation of cells in leaf primordia
(vegetative shoots), lateral meristems (generative shoots) or both (mixed
shoots) in the nodes of the resting buds, and is followed by cell divisions in
the apical meristem to form more nodes. Shoot initiation is stimulated by
pruning, defoliation and irrigation during dry conditions, or transition from
the dry to rainy season in the tropics. Application of nitrogen (N)-containing
fertilizers, exposure to ethylene, or a shift from cool to warm temperatures
T.L. Davenport 106
also stimulates shoot initiation. Reece et al. (1946, 1949), Mustard and Lynch
(1946), Nez-Elisea and Davenport (1992b), Nez-Elisea et al. (1996) and
Davenport et al. (2006a) observed that the vegetative or reproductive fate of
mango buds remains undetermined until after shoot growth is initiated.
Reece et al. (1949) proposed that a putative signal that triggers initiation of
shoot development is separate and different from the inductive signal, which
determines the fate of the shoot. Removal of apical buds by pruning stimu-
lates initiation of axillary shoots (Singh and Singh, 1956; Nez-Elisea and
Davenport, 1992b; Nez-Elisea et al., 1996; Davenport et al., 2006a). Defolia-
tion of the apical whorl of ve to ten leaves also stimulates shoot initiation
in dormant apical buds (Nez-Elisea et al., 1991; Nez-Elisea and Daven-
port, 1995). The fate of shoots that emerge in response to these initiation stim-
uli, however, is determined by other factors that are prevalent at the time of
initiation. Tip pruning, for example, during warm summer months results in
initiation of vegetative shoots from axillary buds, whereas pruning during
cool winter months usually results in initiation of axillary inorescences.
Induction
Induction in mango is the temporary commitment of buds to evoke a par-
ticular developmental pathway (i.e. vegetative shoot, generative shoot or
mixed shoot) when growth is initiated. Initiation of herbaceous plant ower-
ing refers to the onset of oral bud growth in actively growing vegetative
shoots after the oral inductive event (Bernier et al., 1981, 1993; Halevy, 1985
1986; Bernier, 1988; Huala and Sussex, 1993; Kinet, 1993). The inductive sig-
nal is formed in leaves, but the responsive buds are in continuous vegetative
growth at the time of oral induction in herbaceous plants and oral initia-
tion follows; whereas mango buds are in rest. Although the mango bud
must be initiated to grow, that growth is induced according to forces already
present.
Whereas the oral inductive signal in mango may be present prior to
bud initiation, it must be present at the time of initiation for owering to
occur (Kulkarni, 1988a; Nez-Elisea and Davenport, 1995; Nez-Elisea et al.,
1996; Davenport and Nez-Elisea, 1997; Davenport et al., 2006a). The induc-
tive signal can be shifted from oral (F) to vegetative (V) or vegetative to
oral, forming F/V or V/F transition shoots, by altering temperatures dur-
ing early shoot development (Batten and McConchie, 1995; Nez-Elisea
et al., 1996) (Fig. 5.5). This shift in morphogenic responses during shoot devel-
opment demonstrates the plasticity and temporal nature of induction, indi-
cating that cells of the apical meristem do not become irreversibly determined
under inductive conditions. These results demonstrate that, rather than being
irreversibly committed to a vegetative or reproductive fate at the onset of
shoot initiation, the mango apical meristem provides progenitor cells, some
of which differentiate into specic target cells at each node in the apex. The
apical meristem, therefore, may not be directly involved in the owering
process.
Reproductive Physiology 107
Target cells within leaf primordia and lateral meristems are competent to
respond to inductive signals; for example when initiated to grow under veg-
etatively inductive conditions, individual leaf primordia develop as leaves
and subtending lateral meristems associated with each developing leaf
develop as dormant axillary buds with protective bracts. These axillary buds
may develop in subsequent ushes as vegetative shoots when initiated in
vegetatively inductive conditions or as axillary inorescences under oral
inductive conditions. Under strongly oral-inductive conditions, leaf pri-
mordia fail to develop beyond the bract stage, become dormant, and lateral
meristems develop. Each lateral meristem forms nodes consisting of leaf pri-
mordia and meristems that are inuenced by the putative oral-inductive
stimulus, which suppresses development of newly formed leaf primordia.
Subsequently formed meristems form pedunculate structures that terminate
in cymose inorescences borne on each tertiary peduncle (Fig. 5.4). Forma-
tion of the primary, secondary, tertiary and quaternary peduncles, as well as
pedicels of inorescences are always accompanied by a subtending, aborted
bract or vestigial leaf at each node (Fig. 5.4). Such development is attributed
to a sequence of gene expression (Coen et al., 1990; Coen and Meyerowitz,
1991; Weigel et al., 1992; Coen and Carpenter, 1993; Lumsden, 1993; Yanofsky,
1995). Shoot initiation during weakly oral-inductive conditions activates
growth of leaf primordia to develop leaves and the lateral meristems to pro-
duce peduncles bearing lateral inorescences in each node of mixed shoots.
The bases of each pedicel branch within each lateral inorescence also bear a
vestigial leaf.
Upon termination of cell divisions in the apical meristem at the end of a
ushing period, no more nodes are formed. The apical bud of vegetative
shoots becomes quiescent, and the resting leaf primordia, bracts and lateral
meristems are poised to resume growth at a later date. When reproductive or
mixed shoots become quiescent, the lateral meristems ultimately develop
determinant cymose inorescences. The most distally located meristem is
possibly the determinant extension of the central axis forming the terminal
cymose oral group.
Chimeric shoots (Fig. 5.5) can occur in mango trees when shoot initiation
occurs during oral inductive conditions. They display inorescences on one
side of the longitudinally bisected shoot and leaves on the other. The shoot
axis is red on the oral side of red fruiting cultivars (typical of panicles) and
green on the vegetative side (typical of vegetative shoots). This difference
in the two sides extends to the apical bud, which bears an undeveloped
inorescence on the oral side and leaf bracts on the vegetative side. The
explanation for this spatial differentiation is that target nodes on each side of
the apical bud respond to the different inductive signals at the same time.
The apical meristem is not implicated except to form more nodes for the lat-
eral inductive responses on each side in the second portion of growth. Differ-
ences in inductive signals on each side of an existing shoot probably cause
the differential response. This phenomenon indicates that the fate of nodes
on each side of the shoot cannot be attributed to a single mother cell in the
apical meristem. The inductive response must involve cells formed in later
T.L. Davenport 108
cell divisions and would be determined by their location within nodes of
the bud.
Florigenic promoter (FP) or stimulus
Early owering work provided evidence for the presence of a graft transmis-
sible oral stimulus (i.e. origen) that was induced in leaves and was trans-
located to buds to stimulate oral development (Chailakhyan, 1936; Zeevaart
and Boyer, 1987). Florigen was functionally conserved across plant species
(Lang, 1965, 1984; Zeevaart, 1976; Lang et al., 1977). Floral induction in most
plants involves sensing of some environmental cue (i.e. daylength, water
stress or vernalizing temperature) in some organ (e.g. leaves). A putative o-
ral stimulus or alteration in the ratio of origenic to anti-origenic compo-
nents may be translocated to target cells in meristems (Bernier et al., 1981).
Photoassimilate movement from leaves in phloem facilitates its transport to
buds where it can interact to initiate owering (King and Zeevaart, 1973).
Until recently, a oral stimulus could not be identied. Alternative hypoth-
eses were proposed that nutrient diversion to the meristems could be
involved (Sachs and Hackett, 1983) or that oral induction might be con-
trolled by multiple factors, including the putative oral stimulus, photoas-
similates and phytohormones (Bernier et al., 1993).
Molecular biology of owering in the facultative, long-day, model plant,
Arabidopsis thaliana (reviewed in Zeevaart, 2006 and Aksenova et al., 2006),
has provided insight into the nature of the oral stimulus (FP). A network of
four interacting genetic signalling pathways may result in owering in
response to photoperiodic, vernalization, gibberellin and autonomous envi-
ronmental cues (Perilleux et al., 1994; Mouradov et al., 2002; Perilleux and
Bernier, 2002; Boss et al., 2004; Komeda, 2004; Putterill et al., 2004; Corbesier
and Coupland, 2005). The photoperiodic pathway involves activation of
the CONSTANS (CO) gene that encodes a zinc-nger protein, which in
turn induces expression of the FLOWERING LOCUS T (FT) gene in the
phloem tissue of leaves. FT is the terminal, integrating gene of the four path-
ways regulating owering in Arabidopsis. Its transcribed mRNA was initially
thought to be the FP that is transported in phloem to buds (Huang et al.,
2005); however, evidence indicates that the translated protein product of FT
is translocated to Arabidopsis buds (Corbesier et al., 2007). Analogous proteins
encoded by Hd3a, an ortholog of FT in rice (Tamaki et al., 2007), and the aspen
ortholog, PtFT1, which along with CO regulates the timing of owering and
growth cessation of Populus trichocarpa (Bohlenius et al., 2006), appear to be
the FP. In the buds, the protein product of FT is thought to combine with the
bZIP transcription factor (FD) protein to activate transcription of oral iden-
tity genes (i.e. APETALA1) to begin oral expression (Abe et al., 2005; Wigge
et al., 2005). Similar mechanisms are likely to exist in mango.
Zhang et al. (2005) and Davenport et al. (2006b) isolated a CONSTANS-
like gene (MiCOL) from mango leaf DNA. CO is a circadian expression gene
interacting with the photoperiodic pathway in Arabidopsis (Putterill et al.,
Reproductive Physiology 109
2004), and is central to activation of the FT gene in Arabidopsis during long
days. Its role in mango owering is unclear. The mango ortholog has 79%,
76% and 62% homology with two apple CO genes, MdCOL2 and MdCOL1,
and the Arabidopsis CO gene (AtCO), respectively. Isolation of the FT or
homologous gene responsible for synthesis of the FP has been unsuccessful.
Studies with mango indicate that a FP is synthesized in leaves during
exposure to cool, oral-inductive temperatures and moves to buds to induce
owering (Reece et al., 1946, 1949; Singh and Singh, 1956; L.B. Singh, 1959,
1962, 1977; R.N. Singh, 1961; Sen et al., 1972; Nez-Elisea and Davenport,
1989, 1992b; Davenport and Nez-Elisea, 1990; Davenport et al., 1995,
2006a). Unlike receptor sites in buds of Thlaspi arvense (Metzger, 1988) and
other plants requiring vernalization for oral induction (Zeevaart, 1976;
Bernier et al., 1981), mango leaves appear to be where the putative oral stim-
ulus is produced. Complete defoliation of girdled branches during inductive
conditions results in vegetative shoots instead of generative shoots (Reece
et al., 1949; Sen et al., 1972; Nez-Elisea and Davenport, 1989, 1992b; Nez-
Elisea et al., 1996; Davenport et al., 2006a). It appears to be transported over
long distances from leafy branches to defoliated branches (Sen et al., 1972;
Nez-Elisea et al., 1996).
The putative, temperature-regulated FP is short-lived in situ (Nez-
Elisea and Davenport, 1989, 1992b; Davenport et al., 1995; Nez-Elisea et al.,
1996). Leaess cuttings from trees during cool, oral inductive conditions
produce inorescences when stimulated to grow within 7 days of transfer to
warm, non-inductive conditions; the inuence of the removed leaves lasts
for 13 days when cuttings are stored at cool temperatures (Davenport et al.,
2001a). The same cuttings produce only vegetative shoots in both storage
conditions after the initial loss of reproductive shoot production. There are
more leaves on mango stems than are necessary for oral induction in cool
temperatures. Stems bearing as little as one-quarter of a cross-sectioned leaf
induce 95% generative shoots (Davenport et al., 2006a); the remaining shoots
are vegetative. Half of a leaf or more resulted in 100% generative shoots.
Thus, the limiting amount of leaf necessary for oral induction is less than a
quarter of a leaf per stem. Davenport et al. (2006a) demonstrated the quanti-
tative movement of mango FP from half to ve leaves on a donor stem to ve
leaess receiver stems located as far as 100 cm from the donor stem in isolated
branches during exposure to cool, oral inductive temperatures. The FP moves
with photoassimilates in phloem from donor leaves to buds in the receiver
stems.
The mango oral stimulus is graft transmissible (L.B. Singh, 1959, 1962;
Kulkarni, 1986, 1988b, 1991). Flowering of seedling stems is stimulated by
grafting onto mature trees or by grafting mature stems onto juvenile plants
(L.B. Singh, 1959, 1962). Some mango cultivars selected in the tropics can
ower at higher temperatures than others and are not restricted to winter
owering (Kulkarni, 1991). Transfer of the FP from tropical to subtropical selec-
tions was accomplished using reciprocal grafts between the two cultivar types
(Kulkarni, 1986, 1988b, 1991). Subtropical cultivars that seldom ower in warm
temperatures ower in the off season using these techniques. Three conditions
T.L. Davenport 110
were essential for summer owering to occur in the low-temperature-requiring
cultivars (receptors) when grafted to the summer owering type (donors): (i)
the summer-owering donor cultivar stocks or scions were in a owering
cycle; (ii) buds on the receptor scions or stocks of grafted plants had initi-
ated shoot growth during this cycle; and (iii) receptor stocks or scions had
been completely defoliated for transfer and/or expression of the oral stimu-
lus. The presence of any leaves on the receptor plants resulted in vegetative
shoots.
Girdling experiments to isolate treated mango branches from the rest of
the tree suggest that the FP is translocated via phloem to apical buds (King
and Zeevaart, 1973; Bernier et al., 1981; Nez-Elisea and Davenport, 1989,
1992b; Nez-Elisea et al., 1996; Davenport et al., 2006a). Shading experiments
to reduce photosynthate loading into the phloem also support this (Kulkarni,
1991). Reduced owering responses were observed in isolated leafy branches
that were provided with 90% and complete shading, which stopped photosyn-
thate production entirely, mimicked defoliation during cool, oral inductive
conditions, resulting in a vegetative growth response (R. Nez-Elisea, T.L.
Davenport and B. Schaffer, Florida, 1991, unpublished results).
Vegetative promoter (VP)
An independently regulated VP probably contributes to induction of vegeta-
tive shoots as opposed to a oral inhibitor or expression of a default vegeta-
tive status in the absence of sufcient FP at the time of shoot initiation.
Grafting studies (L.B. Singh, 1959, 1962; Kulkarni, 1986, 1988b, 1991, 2004)
demonstrated that complete removal of leaves from receptor stems is required
to express owering of those receptors when they are grafted to owering
donor stems. Kulkarni (1986, 1988b, 1991, 2004) considered that a putative
oral inhibitor in leaves of the non-induced receptor stems might antagonize
the inuence of the oral stimulus from donor leaves. Others have noted a
relationship between leaf age and the ability of shoots to be reproductive
(Singh et al., 1962a; Scholeeld et al., 1986). KNO
3
-stimulated early owering
in the tropics is successful only on stems that are at least 4 (Davenport, 2003)
to 7 months old (Astudillo and Bondad, 1978; Bondad and Apostol, 1979;
Nez-Elisea, 1985). Young stems often produce vegetative shoots when ini-
tiated under conditions that are oral inductive for more mature stems
(Nez-Elisea and Davenport, 1995; Davenport, 2003). The putative VP
appears to be most active in leaves of young stems and slowly dissipates
over time to allow expression of the FP when shoots are initiated to grow in
warm conditions.
The VP may be a gibberellin or closely associated with the gibberellin
synthesis pathway as indicated by enhanced owering responses of trees to
plant growth retardants. Mangoes growing in wet and humid, low-latitude
tropics tend to produce frequent vegetative ushes and ower sporadically,
perhaps due to higher levels of the VP in the young stems combined with
low levels of the putative FP when shoot initiation occurs. Paclobutrazol
Reproductive Physiology 111
(PBZ) reduces the time in rest necessary to allow oral induction during
warm temperature conditions by c.1 month (Davenport, 2003), thus increas-
ing the potential to produce reproductive shoots in younger stems when ini-
tiated to grow. PBZ and uniconazole, triazole compounds that inhibit kaurene
oxidase in the gibberellin-synthesis pathway (Dalziel and Lawrence, 1984;
Rademacher, 1991), stimulate production of owering shoots during weakly
inductive conditions (Burondkar and Gunjate, 1991, 1993; Tongumpai et al.,
1991a; Voon et al., 1991; Nartvaranant et al., 2000; Yeshitela et al., 2004a).
Application of PBZ to mango trees bearing 1-month-old stems produced
inorescences when bud break was initiated 3 months later by foliar applica-
tion of KNO
3
(Davenport, 2003).
Vegetative or reproductive induction at the time of shoot initiation is
governed by the ratio of the putative oral promotive to inhibitory compo-
nents (Lang et al., 1977; Lang, 1984; Kulkarni, 1988a; see Bernier et al., 1981 for
additional references). The mango oral inhibitor should be viewed as an
age-dependent VP. The presence of an age-regulated VP in mango leaves,
which moves with the temperature-regulated FP and photoassimilates in
phloem, may explain the induction of specic receptors by this promoter in
targeted leaf primordia to cause development of leaves in vegetative or
mixed shoots. A gradual decrease in the level or inuence of the VP may
cause vegetative shoots to develop when initiation occurs on 2-month-old
stems, and generative or mixed shoots when initiation occurs in stems from
4- to 7-month-old stems, given the constantly warm daily temperatures
maintaining a low level of FP in both situations.
5.5 Environmental Inuence on Vegetative and Reproductive
Development
The effects of temperature and water relations on determinating vegetative
and reproductive growth of mango have been addressed (Davenport and
Nez-Elisea, 1997; Davenport, 2000; Kulkarni, 2004; Bangerth, 2006). This
section focuses on the impacts of temperature, plant water relations, mineral
nutrition and photoperiod on shoot initiation and induction.
Temperature
The developmental fate of mango buds is strongly inuenced by tempera-
ture (Davenport and Nez-Elisea, 1997). Cool night temperatures < 15C in
combination with day temperatures < 20C typically induce owering if
shoot initiation occurs when plants are exposed to these conditions (Ou,
1980, 1982; Wolstenholme and Mullins, 1982a, b; Shu and Sheen, 1987; Whi-
ley et al., 1988, 1989, 1991; Nez-Elisea et al., 1993; Nez-Elisea, 1994;
Nez-Elisea and Davenport, 1994a, b). The physiological and molecular
basis for temperature perception in leaves with respect to oral induction is
not understood (Samach and Wigge, 2005). Whiley et al. (1988, 1989, 1991)
T.L. Davenport 112
described the vegetative growth and owering responses of several mo-
noembryonic and polyembryonic cultivars to four temperature regimes rang-
ing from vegetatively inductive (30C day/25C night) to oral inductive (15C
day/10C night). The effect of temperature on marcotted, container-grown
plants that were tip pruned or defoliated in order to stimulate shoot initia-
tion was also studied (Davenport, 1987; Nez-Elisea et al., 1991, 1993, 1996;
Nez-Elisea and Davenport, 1994b). Mango trees develop vegetative shoots
when shoot initiation occurs in warm temperatures (30C day/25C night),
whereas inorescences develop when shoots initiate growth in cool tempera-
ture conditions (18C day/10C night; or 15C day/10C night) (Whiley et al.
1989; Nez-Elisea and Davenport, 1991b, 1995; Nez-Elisea et al., 1993,
1996; Batten and McConchie, 1995). Bangerth et al. (2004) reported changes in
the major phytohormones in stems of containerized mango trees during
exposure to cool, oral inductive temperatures. The minimum leaf age and
time of exposure to a low temperature regime (18C day/10C night) required
by stems for oral induction was examined (Nez-Elisea and Davenport,
1995). Leaves are competent to respond to cool temperatures at 7 weeks,
forming a small percentage of generative shoots. As they age, higher propor-
tions of generative shoots are induced and warmer temperatures can stimu-
late oral induction. The response to temperature is moderated by age of the
previous ush. Stems that are 45 months beyond the limp, red-leaf stage of
development will be induced to form generative shoots if initiated to grow
at 2530C (Davenport, 2003).
Whiley et al. (1988, 1989, 1991) observed that at least 17 weeks are required
for initiation of reproductive shoots on non-clipped stems of trees maintained
at 15C day/10C night. In similar experiments with different cultivars with-
out previous clipping of distal leaves to stimulate initiation, inorescences
were observed after 5 weeks at 15C day/10C night (Chaikiattiyos et al.,
1994). Although inductive conditions were present in each of these studies,
shoot initiation was delayed by the presence of distal leaves. The earlier ini-
tiation of inorescence development in tip-pruned or tip-defoliated stems
compared to intact ones demonstrates that the oral stimulus may be pres-
ent, but the buds are not induced until initiation occurs. It demonstrates the
importance of stimulating initiation of stems by tip defoliation or pruning
at the onset of incubation in controlled environment conditions so that the
inductive response can be observed within a reasonable length of time.
The variable delays in shoot initiation in these studies occurred because the
experimental protocols depended on the plants internal initiation cycle to
initiate shoots. This cycle slows down when plants are exposed to lower tem-
peratures (Whiley et al., 1988, 1989, 1991).
Floral or vegetative induction occurs when shoots are initiated. Resting
buds of plants that are exposed to cool temperatures (18C day/10C night)
for > 3 weeks and then transferred to a warm temperature (30C day/25C
night) before initiation, produce only vegetative shoots (Nez-Elisea et al.,
1996). Thus, the stems do not remember that they had been exposed to oral
inductive conditions while still in rest. They responded to warm conditions
present when shoot initiation occurred.
Reproductive Physiology 113
This response to temperature conditions at the time of shoot initiation
extends to the formation of transition shoots if conditions change during
early shoot development. First reported by Naik and Mohan Rao (1943),
transition shoots are an unusual transition in expression of shoot type during
a single growth ush (Kulkarni, 1988b; Nez-Elisea and Davenport, 1989,
1992b; Batten and McConchie, 1995). The transition typically occurs near the
middle of the extending shoot. Resting buds possess preformed nodes, each
of which contains a primordial leaf or bract and a lateral meristem. The api-
cal meristem initiates cell division at the same time or soon after the nodal
target tissues begin development (Mustard and Lynch, 1946; L.B. Singh, 1960;
Nez-Elisea et al., 1996). Vegetative or inorescence development in the
pre-formed primordia is underway before the apical meristem begins to pro-
duce differentiating cells. Transfer from a warm, vegetatively inductive con-
dition to a cool, oral inductive environment at early bud break results in
formation of V/F transition shoots (Fig. 5.5). Transfer from cool to warm con-
ditions at the same stage of bud break results in formation of F/V transition
shoots (Batten and McConchie, 1995; Nez-Elisea et al., 1996).
The owering response to temperature occurs in mangoes growing in
subtropical latitudes where cool temperature is the dominant induction fac-
tor. Many cultivars ower erratically in the low-latitude tropics, providing
continuously warm temperatures with high soil and atmospheric moisture.
Under such conditions, the age of stems is the dominant inductive factor
(Buell, 1954; Nakasone et al., 1955; Ravishankar et al., 1979; Ou and Yen, 1985;
Issarakraisila et al., 1992), and occasional cool night temperatures in the upper
latitude tropics have a positive moderating effect (Davenport, 2003).
Water relations
In the absence of cool temperatures, mango trees in the tropics may ower in
response to irrigation or rain following periods of water stress lasting 612
weeks or more (Pongsomboon, 1991). Plant water stress has been presumed
to provide the stimulus for owering (reviewed in Whiley, 1993; Chaikiatti-
yos et al., 1994; Schaffer et al., 1994; Davenport and Nez-Elisea, 1997); how-
ever, most of these studies have failed to substantiate prolonged tree water
decit as a successful agent for oral induction.
Experiments with container-grown trees fail to produce inorescences
after 8 weeks of water decit (Wolstenholme and Hofmeyr, 1985). Under
glasshouse conditions (27C day/22C night; relative humidity (RH) 90%),
container-grown, monoembryonic cultivars were water stressed through
decit irrigation for 14 days, resulting in an average leaf xylem water poten-
tial of 3.9 MPa (Davenport, 1992; Nez-Elisea and Davenport, 1992a,
1994b). Following resumption of irrigation, all trees grew vegetatively. Sim-
ilarly, only vegetative growth was obtained when container-grown trees
were deprived of irrigation for 36 days during summer, although leaf xylem
water potentials of 3.78 MPa were attained (Nez-Elisea and Davenport,
1994b). Water stress imposed on plants during the cool autumn months
T.L. Davenport 114
(night temperatures < 15C) do not increase the proportion of apical buds
forming inorescences, but expedited shoot initiation after rewatering
(Nez-Elisea and Davenport, 1994b). These results demonstrated that cool
temperatures provide inductive conditions, whereas relief of water stress
accelerated shoot initiation under cool, inductive temperatures. Flowering
was delayed when container-grown monoembryonic mangoes were water-
stressed at 18C day/15C night (Chaikiattiyos et al., 1994). Water-stressed
trees held at 29C day/25C night did not ower.
Mango trees growing in the low-latitude tropics may ower after an
extended period of mild water stress (Harris, 1901; Collins, 1903; Kinman,
1918; Gangolly et al., 1957; Gangolly, 1960; L.B. Singh, 1960). Pongsomboon
et al. (1991) observed owering in eld-grown trees in the tropics following
6 weeks of withholding water. The primary impact of water stress appears to
be prevention of shoot initiation during stress. The accumulating age of stems
is greater in water-stressed trees than in trees maintained under well-watered
conditions that promote frequent vegetative ushes (Davenport, 1992, 1993;
Schaffer et al., 1994). This delay in ushing may provide more time for accu-
mulation of a putative FP (Schaffer et al., 1994) or reduction in the level of a
putative VP (Davenport and Nez-Elisea, 1997; Davenport, 2000). Some
cultivars appear to be better adapted to such delays in growth and perform
better in dry environments in the tropics.
Effect of N on owering
Subsequent to the discovery of ethephon to stimulate mango owering (Gon-
zalez, 1923; Alcala and San Pedro, 1935), Barba (1974), Bueno and Valmayor
(1974), Astudillo and Bondad (1978), Bondad et al. (1978), Bondad and Apos-
tol (1979), Pantastico and Manuel (1978) and Bondad and Linsangan (1979)
reported that KNO
3
could be used for the same purpose. This has been
exploited in the low- and mid-latitude tropics (Mosqueda-Vzquez and de
los Santos de la Rosa, 1981; Mosqueda-Vzquez and Avila-Resendiz, 1985;
Nez-Elisea, 1985, 1986; Ou and Yen, 1985; Winston and Wright, 1986;
Tongumpai et al., 1989; Goguey, 1993; Ravishankar et al., 1993; Sergent et al.,
1996; Yeshitela et al., 2004b, 2005). The nitrate (NO
3

) anion is the active


component of KNO
3
(Bueno and Valmayor, 1974), and ammonium nitrate
(NH
4
NO
3
) is twice as effective as KNO
3
(Nez-Elisea, 1988; Nez-Elisea
and Caldeira, 1988). In the low- and mid-latitude tropics, receptive trees
respond by developing visible reproductive buds within 2 weeks after
application. The effective spray concentration is 110% KNO
3
, depending on
the age of the trees and climate. Two to four per cent KNO
3
or calcium nitrate
(Ca(NO
3
)
2
) and 12% NH
4
NO
3
are effective for stimulating owering in
most conditions. The physiological and temporal timing of application is
important. Old trees, non-vigorous trees, and trees in which vegetative
ushes have been discouraged by low water potentials produce the best
response to NO
3

induction (N. Golez, personal communication, the Philip-


pines, 1989).
Reproductive Physiology 115
Chemical bud forcing is most effective in the tropics where distinct wet
and dry seasons prevail. The response to chemical bud forcing by NO
3

and
ethephon diminishes at latitudes > 22 N or S (Mosqueda-Vzquez and de
los Santos de la Rosa, 1981). Their effect may involve the decline of night
temperatures from 20C around the equator to 10C between 22 and 27
N or S latitude during winter months or by late summer vegetative ushes.
Trees in the wet or dry subtropics at 25 N or S have not responded to treat-
ments (Davenport, 1993).
Stems must be sufciently mature, dark green with a minimum age of 4
months since the previous limp, red-leaf stage in easily induced cultivars
and 5 months for more recalcitrant cultivars to obtain a reproductive shoot
response in the low-latitude tropics (Davenport, 2003). Bueno and Valmayor
(1974) indicated that leaves must be brittle when hand-crushed. Nez-
Elisea (1986, 1988) reported that stems must be at least 6 months old. Trees
that experience autumn dry periods become responsive to treatments as
early as October (northern hemisphere). Groups of stems within tree cano-
pies are produced through asynchronous ushes of growth, and vary in age;
only a few are responsive to the rst inductive spray. Subsequent biweekly
applications cause owering in canopy sectors as they reach the age-depen-
dent requirement for initiation. Early and out-of-season owering and fruit-
ing can thereby be achieved.
KNO
3
may be oral inductive in mango (Barba, 1974); however, trees in
the upper latitude tropics typically ush vegetatively rather than produce
bloom when either KNO
3
or NH
4
NO
3
is sprayed between June and Septem-
ber (N. Golez, personal communication, the Philippines, 1989). The warm,
rainy season producing frequent ushes of growth during this period is con-
ducive to a vegetative response to the sprays. These results indicate that
KNO
3
and NH
4
NO
3
stimulate shoot initiation but do not determine bud
morphogenesis. In buds released after KNO
3
or NH
4
NO
3
treatments, the
ratio of leaf-generated FP to VP and not NO
3

causes initiating buds to become


reproductive. Kulkarni (1988b, 2004) suggested that the oral stimulus is
present in stems when buds are forced in response to KNO
3
and suggested
that KNO
3
may also sensitize buds to the oral stimulus. Davenport (2003),
T.L. Davenport and J. Oleo (2006, unpublished data) and F. Ramirez and T.L.
Davenport (submitted for publication) observed 100% vegetative shoots
when 4% KNO
3
was foliar applied to 2-month-old stems; whereas, applica-
tion of the same spray treatment to 4.5-month-old stems on trees in the same
orchards resulted in 100% reproductive shoots.
Trees with high leaf N levels rarely ower in the tropics. Lack of ower-
ing is always due to frequent vegetative ushes of growth, especially during
the rainy season. Mango trees must have leaf N levels of 1.4% or less in order
to suppress frequent ushes of vegetative growth (Davenport, 2003). Leaf N
levels of < 1.1% suppress frequent ushes but also provide insufcient nutri-
tion to support good cropping. Thus, 1.11.4% N levels in leaves appear to be
optimum for good commercial production and control of owering time in a
managed orchard. The application of KNO
3
to the foliage of the resting stems
45 months after the limp, red-leaf stage will cause a owering response.
T.L. Davenport 116
Photoperiod
Flowering in most trees does not appear to be under photoperiodic control
(Kozlowski et al., 1991). Mango cultivation is concentrated between 27 N
and 27 S where the shortest annual photoperiod is c.10.5 h and the longest
photoperiod is c.13.5 h. Cultivars in the upper-latitude tropics and subtropics
ower during the winter when photoperiods are short; however, trees in the
low-latitude tropics, where a 12-h photoperiod is nearly constant, can ower
at any time of the year. Furthermore, owering on spring-initiated shoots in
the subtropics occurs during summer (Schaffer et al., 1994). Studies have
failed to demonstrate a correlation between 8-h photoperiods and owering
(Maiti, 1971; Maiti and Sen, 1978; Maiti et al., 1978). Nez-Elisea and Daven-
port (1995) studied the effects of 11-, 12-, 13- and 24-h photoperiods at 18C
day/10C night, or 11- and 13-h photoperiods at 30C day/25C night on
owering of container-grown trees. Photoperiod had no effect on the fate of
buds, and the promotive effect of cool temperatures on owering was inde-
pendent of photoperiod. Photoperiods of 11-, 12- or 13-h with 18C day/10C
night, caused owering in trees within 40 days. The 24-h photoperiod with
12-h thermoperiods of 18C and 10C caused owering of trees within 35
days. Photoperiods of 11- or 13-h at 30C day/25C night resulted in vegeta-
tive growth only. With warm temperatures, vegetative shoots were produced
in 17 days. These results conrm that oral induction is caused by cool tem-
peratures and not by short photoperiods and that warm temperature, not a
long photoperiod, caused vegetative induction.
5.6 Hormonal Inuence on Flowering
FP is a protein product of the FT gene in Arabidopsis (Corbesier et al., 2007)
and the Hd3a gene in rice (Tamaki et al., 2007) and moves in phloem from
leaves to buds; there is little evidence that phytohormones are directly
involved as the FP. Phytohormones appear to be responsible for shoot initia-
tion in conditions that are oral inductive.
Ethylene
Smudging has been utilized to stimulate mango owering in the Philippines.
Only branches that attain sufcient age respond to smudging by forming
reproductive shoots (Acala and San Pedro, 1935; Bueno and Valmayor, 1974).
Rodriguez (1932), investigating smoke-induced owering of pineapple, pro-
posed that ethylene, generated by burning material, may stimulate ower-
ing. Dutcher (1972) conrmed that smoke from smudge res contained
ethylene. Smudging and the use of ethephon in 1968 by F. Manuel (Barba,
1974) and others (Bondad, 1972, 1976) to promote mango owering sug-
gested that endogenous ethylene is integral for oral induction (Barba, 1974;
Bondad, 1976; Chadha and Pal, 1986). Ethephon effectively promotes owering
Reproductive Physiology 117
of mangoes under specic conditions in the low-latitude tropics (Davenport
and Nez-Elisea, 1997).
The involvement of endogenous ethylene in owering is supported by
observations that indirectly link it to symptoms of ethylene production.
Extrusion of latex from terminal buds occurs at the time of inorescence ini-
tiation, and epinasty of mature leaves near the apex during expansion of the
panicle has been observed (Davenport and Nez-Elisea, 1990, 1991). Both
are symptoms of plants exposed to high ethylene levels (Abeles, 1973). Indi-
rect support also comes from reports that KNO
3
-stimulated owering of
mango is mediated by increased levels of endogenous ethylene (Thuck-Thye,
1978; Lopez et al., 1984). Mosqueda-Vzquez and Avila-Resendiz (1985)
reported that the efcacy of KNO
3
was negated by cobalt chloride (CoCl
2
)
and silver nitrate (AgNO
3
), which inhibit the synthesis and action of ethyl-
ene, respectively, when sprayed 14 h after KNO
3
. Saidha et al. (1983) reported
a gradual increase in endogenous leaf ethylene production as the season of
oral initiation approached. Ethylene production by stems producing repro-
ductive shoots was up to vefold that of resting stems.
Inconsistent (Pandey et al., 1973; Sen et al., 1973; Winston and Wright,
1986) or non-responsive results with ethephon (Pandey and Narwadkar,
1984; Ou and Yen, 1985; Pandey, 1989) or smudging (Sen and Roy, 1935),
especially during warm, non-inductive conditions, have been reported. Dav-
enport and Nez-Elisea (1990, 1991) reported elevated ethylene production
in mango stems in response to ethephon sprays without an accompanying
oral response. Experiments were conducted during oral-inductive and
non-inductive periods. Unlike Saidha et al. (1983), they observed no increase
in ethylene production rates prior to or during panicle development.
The effect of ethylene on owering is unresolved. It is likely that ethylene
stimulates shoot initiation by inhibiting auxin transport from leaves to buds
and stems (Morgan and Gausman, 1966; Beyer and Morgan, 1971; Riov and
Goren, 1979, 1980; Ramina et al., 1986). This may increase the ratio of cytoki-
nin to auxin in buds and stimulate shoot initiation (Davenport, 2000). Other
factors (i.e. cool temperatures or aged leaves) may be responsible for oral
induction (Ona and de Guzman, 1982; Davenport, 1993).
Auxin
Although auxin may have a critical role in oral induction of mango (Chadha
and Pal, 1986; Hegele et al., 2006), there is little supporting evidence. The
application (L.B. Singh, 1961; Singh and Singh, 1963; Bakr et al., 1981; Pandey
and Narwadkar, 1984) and analysis of auxin in leaves (Paulas and Shanmu-
gavelu, 1989; Sivagami et al., 1989), stems (Chen, 1987) and shoots (Chacko et al.,
1972b) have been reported in relation to mango owering. These studies are
inconclusive due to inconsistencies in purication and analytical methodolo-
gies (Davenport and Nez-Elisea, 1997).
Auxin may indirectly stimulate root-produced cytokinins through initia-
tion of new root growth. Auxin is transported basipetally from growing
T.L. Davenport 118
shoots and leaves to roots (Goldsmith, 1968; Cane and Wilkins, 1970; Wilkins
and Cane, 1970; Goldsmith and Ray, 1973; Lomax et al., 1995) and stimulates
root initiation (Hassig, 1974; Wightman et al., 1980). The efcacy of various
auxins for stimulating adventitious rooting of mango marcots and cuttings
was reviewed by Davenport and Nez-Elisea (1997).
Auxin inhibits shoot initiation (Davies, 1995) and confers apical domi-
nance by preventing axillary bud break. Leaf-produced auxin and petiolar
auxin transport capacity declines as leaves age (Veen, 1969; Veen and Jacobs,
1969; Davenport et al., 1980). The interaction of decreasing auxin and accu-
mulating cytokinins in resting buds may explain the cyclic nature of shoot
initiation. The ratio of cytokinin to auxin levels in buds regulates shoot
initiation (Skoog and Miller, 1957; Bangerth, 1994; Cline et al., 1997; Beveridge
et al., 2003).
Cytokinins
Relationships between mango owering and the endogenous levels of cyto-
kinins in leaves (Paulas and Shanmugavelu, 1989; Kurian et al., 1992), stem
tips (Agrawal et al., 1980) and xylem sap (Chen, 1987) and the effect of cyto-
kinin applications on bud break and shoot development have been reported.
Chen (1985) described precocious owering of mango shoots in response to
early October application of 6-benzylaminopurine (BA). Flowering was
observed 1 month following application and 3 months later on non-treated
trees. Nez-Elisea et al. (1990) reported numerous reproductive shoots per
stem in response to the synthetic cytokinin, thidiazuron, during cool, oral
inductive conditions; however, numerous vegetative shoots per stem were
initiated when thidiazuron was applied during warm, vegetatively induc-
tive conditions. Early bud break was not achieved following foliar applica-
tion of Promalin (commercial formulation of BA and gibberellins A
4
+A
7
)
(Oosthuyse, 1991b), BA (A.K. Singh and Rajput, 1990) or kinetin (Singh and
Singh, 1974).
Chen (1987) reported the lowest levels of putative trans zeatin and its
riboside were translocated from roots during the vegetative shoot growth
and resting stages, whereas the highest levels occurred during early ower-
ing and full bloom. Paulas and Shanmugavelu (1989) observed no signicant
difference in cytokinin levels of the fourth and fth leaves during resting bud
and owering. Cytokinin levels in mango stem buds increased during expo-
sure to cool, oral inductive temperatures (Bangerth et al., 2004). Agrawal
et al. (1980) described 11 cytokinin-like substances isolated from stem tips of
an alternate-bearing cultivar in on and off years. Kurian et al. (1992) reported
a link between PBZ applications and reduction in cytokinins in mango leaves
with treatments, perhaps caused by reduction in feeder root development
and formation of thick, blunt roots (Bausher and Yelenosky, 1987; Peng et al.,
1991; Burrows et al., 1992; Yelenosky et al., 1993). Concurrent with this response
was suppression of bud initiation and reduced internode lengths for c.2
years.
Reproductive Physiology 119
The role of cytokinins in owering is unresolved due to sampling of dif-
ferent organs at non-comparable times or conditions. The elevated cytokinin
levels found prior to and during owering and the owering response to
applied BA led to the conclusion that cytokinins are involved in owering of
mango (Chen, 1985, 1987; Bangerth, 2006); however, such responses can be
explained if cytokinins are involved in stimulation of bud break (i.e. shoot
initiation) during oral inductive conditions.
A well-documented role for cytokinins in higher plants, especially evi-
dent in vitro, is bud organogenesis (Skoog and Miller, 1957; Miller, 1963;
Takahashi, 1986; Salisbury and Ross, 1992; Davies, 1995; Haberer and Kieber,
2002). The primary cytokinins in higher plants are trans zeatin, dihydrozeatin,
isopentenyl adenine and their ribosides. They are translocated from roots and
accumulate in resting buds (Hendry et al., 1982a, b) or can possibly be synthe-
sized in nearby tissues as regulated by auxin (Nordstrom et al., 2004; Tanaka
et al., 2006). Their rate of accumulation may relate to periodic root ushes that
alternate with shoot ushes (Krishnamurthi et al., 1960; Bevington and Castle,
1986; Cull, 1987, 1991; Parisot, 1988; Williamson and Coston, 1989).
Gibberellins
Gibberellins are tetracyclic diterpenoid compounds that vary in biological
activity according to the type and location of substituted side groups on a
basic ent-gibberellane skeleton. The number of known gibberellins is > 100
(Pearce et al., 1994). Reproductive shoot initiation is suppressed in many
woody angiosperms by gibberellic acid (GA
3
) (Pharis and King, 1985). GA
3

inhibits mango owering (older literature reviewed in Davenport and
Nez-Elisea, 1997; Nez-Elisea and Davenport, 1998).
GA
3
inhibition of mango owering is correlated with the applied con-
centration (Kachru et al., 1971, 1972) and may cause buds to develop vegeta-
tively under oral-inductive conditions. Nez-Elisea and Davenport (1991a,
1998) reported a delay in initiation of axillary shoots when GA
3
was foliar
applied to deblossomed stems during cool, oral inductive temperatures.
Higher concentrations caused longer delays in shoot initiation. GA
3
did not
inhibit oral induction, so long as cool, inductive temperatures were present
during axillary shoot initiation. Late initiating buds, which grew during
warm, spring temperatures, however, formed vegetative shoots. Similar
delays in reproductive shoot initiation in response to GA
3
application was
reported by Shawky et al. (1978) and Turnbull et al. (1996). Multiple applica-
tions, even at lower rates, are more effective than a single application (Tomer,
1984; Turnbull et al., 1996; Davenport and Smith, 1997). GA
3
treatment has
been recommended in the Canary Islands to delay owering until the danger
of frost has passed (Galn-Saco, 1990). In the subtropics of Australia, it is
used to prevent owering in newly planted trees during the spring so that the
full growing period can be utilized for vegetative growth, thereby hastening
orchard establishment (A.W. Whiley, personal communication, Queensland,
1996).
T.L. Davenport 120
Response to GA
3
varies among cultivars, growing conditions and timing
of application (Tomer, 1984; Oosthuyse, 1995a; Turnbull et al., 1996; Snchez-
Snchez et al., 2004). GA
3
can delay shoot initiation beyond the oral induc-
tive window, resulting in a vegetative ush when shoots develop in warm
weather (Kachru et al., 1971, 1972; Nez-Elisea and Davenport, 1991a, 1998;
S. Gazit, personal communication, Israel, 1993). The variable response to GA
3

may be related to levels of active gibberellin in buds at the time of applica-
tion, inconsistent uptake or differential sensitivity of buds, depending on
their position (apical versus axillary) or age (Nez-Elisea and Davenport,
1991a, 1998). Efcacy is related to the timing of application; immediately
prior to normal shoot initiation appears to be most effective (Davenport and
Smith, 1997).
Reports of endogenous gibberellins in mango tissues, especially in buds,
are difcult to interpret with respect to a regulatory role in bud break or
owering. Problems include sampling of tissues other than apical buds, i.e.
whole stems (Tongumpai et al., 1991b), leaves (Paulus and Shanmugavelu,
1989; Sivagami et al., 1989) and xylem sap (Chen, 1987), or at times when
developing shoots may contribute to the overall result (Chen, 1987). Pal and
Ram (1978) tentatively identied the presence of gibberellins A
1
, A
3
, A
4
, A
5
,
A
6
, A
7
and A
9
. Chen (1987) identied gibberellins A
1/3
, A
4/7
, A
5
, A
17
, A
20
and
A
29
. The estimated levels of gibberellins in apical buds for 6 months prior to
the owering season were reported to be higher in the off year than in the
on year of an alternate-bearing cultivar (Pal and Ram, 1978). Chen (1987)
reported the highest levels of gibberellins in xylem sap during leaf differen-
tiation and lower concentrations during rest, panicle emergence and full
owering. Tongumpai et al. (1991b) observed increasing levels of gibberellins
in whole stems over the 16 weeks prior to vegetative shoot emergence and
decreasing levels over the same period prior to panicle development. Gib-
berellins A
1
, epi-A
1
, A
3
, A
19
, A
20
and an unidentied gibberellin in buds and
leaves from shoot and stem tips of different ages have been quantied (Dav-
enport et al., 2001b). The detected gibberellins are members of the early
13-hydroxylation pathway of gibberellin synthesis (Takahashi, 1986; Pearce
et al., 1994). Gibberellins A
3
and A
19
were the most abundant gibberellins in
apical stem buds. The concentration of GA
3
increased within buds with
increasing age of stems, although concentrations of other GAs were variable.
The concentration of GA
3
did not change signicantly with age in leaves,
whereas that of most of the other GAs declined. Davenport et al. (2001b)
concluded that elevated GA
3
levels in buds may enhance or maintain the
synthesis or activity of endogenous auxin to maintain low cytokinin/auxin
ratios and enhance inhibition of shoot initiation (Jacobs and Case 1965; Scott
et al., 1967; Pharis et al., 1972; Ross et al., 1983; Law 1987; Law and Hamilton,
1989).
The roles of gibberellins and other phytohormones in shoot initiation
and induction is unclear. Endogenous levels in buds and leaves must be cor-
related with physiological events in individual stems. Experimental approaches
should include examination of resting buds up to both vegetative and repro-
ductive shoot initiation to avoid misinterpretation of results. Experiments
Reproductive Physiology 121
should utilize plants grown under dened conditions with specic environ-
mental controls for evaluation of cause and effect. Finally, extraction and
purication protocols should include quantiable internal standards and
use of sensitive unambiguous analytical techniques.
Plant growth retardants
Plant growth retardants have been evaluated to stimulate early or more
intense owering, especially in the off year of alternate-bearing cultivars
(Davenport and Nez-Elisea, 1997). They are in three main classes: (i) the
gibberellin transport inhibitor, daminozide (N-dimethylamino-succinamic
acid), known as alar or B-Nine; (ii) the onium type, chloremquat chloride
(2-chloroethyl trimethylammonium chloride), known as cycocel and CCC;
and (iii) the steroid-synthesis-inhibiting triazoles, for example PBZ (PP-333),
known as Cultar

, and uniconazole, known as XE-1019 or Sumagic (Rademacher,


1991, 2000a). The latter two classes of compounds inhibit ent-kaurene syn-
thetase, an enzyme in the gibberellin synthesis pathway (Nickell, 1983;
Dalziel and Lawrence, 1984; Rademacher, 1991, 2000a). Applying daminoz-
ide results in increased gibberellin levels, perhaps due to the inability to dis-
tribute it properly (Rademacher, 1991). Plant responses may depend upon
whether target tissues are near the site of gibberellin synthesis or sufciently
removed from it to be affected by the inhibited translocation.
Daminozide and cycocel
The efcacy of daminozide and cycocel for increasing owering in the off sea-
son of alternate-bearing cultivars has been studied (Maiti et al., 1972; Mukhopad-
hyaya, 1978; Rath and Das, 1979; Suryanarayana, 1980; Rath et al., 1982; Ou and
Yen, 1985), together with their ability to stimulate early owering (Suryanarayana
and Rao, 1977; Chen, 1985; Kurian and Iyer, 1993a, b). Enhanced, inconsistent
owering occurs in response to these compounds, especially cycocel.
Triazoles
PBZ is being used (except in the USA where it has not been cleared for use)
to stimulate enhanced or early owering. It is best applied to the soil due to
its low solubility, long residual activity and lack of efcient foliar uptake
(Rademacher, 2000b). PBZ applied as a soil drench (120 g active ingredient
(ai)/tree) reduces internode lengths and causes earlier and enhanced ower-
ing in mango trees (Hasdiseve and Tongumpai, 1986; Haw, 1986; Hongsb-
hanich, 1986). Depending on climate, residual activity lasts for c.2 years
(Kulkarni, 1988a). These results have been conrmed in different locations in
the tropics (Davenport and Nez-Elisea, 1997; Yeshitela et al., 2004a, b).
Nartvaranant et al. (2000) recommended soil application of PBZ at 11.5 g
ai/m of canopy diameter to achieve owering in 90120 days if the trees are
stimulated to ush. Davenport (2003) observed that such treatments allowed
a reduction of c.1 month in the time required for stem rest before stimulating
them to initiate reproductive shoots using KNO
3
. PBZ also reduces alternate
T.L. Davenport 122
bearing of some cultivars (Hillier and Rudge, 1991; Burondkar and Gunjate,
1993; Rao, 1997; Rao et al., 1997; Rao and Srihari, 1998; Vijayalakshmi and
Srinivasan, 1999). Cultivars that tend to ower with minimal inductive impe-
tus are more responsive and can be induced to ower out-of-season using
PBZ (Tongumpai et al., 1989). Nez-Elisea et al. (1993) demonstrated that
application of PBZ and uniconazole advanced bud break of containerized
trees in controlled environment chambers, but cool temperatures were neces-
sary to induce owering. Initiated shoots were induced to be vegetative in
warm temperatures. The greater proportion of purely reproductive panicles
in treated plants (compared with controls) suggests that triazoles impact the
level of a putative VP, probably a gibberellin. Whiley (1993) suggested a sec-
ondary mechanism for the oral promotive action of PBZ on mangoes, not-
ing inconsistent responses in the literature between cultivars, environments
and application times.
Application of PBZ reduces the number of panicles, despite increased
fruit set (Goguey, 1990). Davenport (1987, 1994) observed neither growth
inhibition nor enhanced or early owering in response to root drenches or
bark banding with uniconazole (15 g ai/tree) in trees growing in alkaline,
calcareous soil. He reported that new shoot growth was stunted with
extremely short internodes when trees were severely pruned soon after or as
long as 3 years after treatment. Yield was severely reduced due to the lack of
normal growth ushes. The growth stunting effect continued for 7 years after
pruning. Davenport (1994) warned that use of triazole plant growth retar-
dants for control of tree growth, owering or yield must be done with con-
siderable caution, especially if severe pruning of the trees is anticipated.
Residual uniconazole or PBZ applied as a soil drench or bark band is appar-
ently retained in high concentrations in main scaffolding branches. In Cen-
tral and South America, growers utilize PBZ annually to stimulate early
owering. A test tree should be severely pruned to determine if the trees are
affected by PBZ to anticipate the orchard response to later severe pruning.
Certain gibberellins (i.e. GA
1
) are necessary for shoot elongation. Inhibi-
tion of bud break and shoot elongation in response to application of the
growth retardants cycocel (Maiti et al., 1972) and triazoles (Kulkarni, 1988a;
Burondkar and Gunjate, 1991, 1993; Tongumpai et al., 1991a; Kurian et al.,
1992; Winston, 1992; Kurian and Iyer, 1993a, b; Nez-Elisea et al., 1993; Wer-
ner, 1993) have been reported. Elongation of panicles is inhibited, especially
by high levels of triazoles (Kulkarni, 1988b; Winston, 1992; Davenport, 1994;
Salomon and Reuveni, 1994). Inorescences in treated trees may become
compact, improving opportunities for disease and insect attack (Winston,
1992). Kurian et al. (1992) associated reduced cytokinin levels in leaves with
inhibition of shoot initiation in plants treated with soil drenches of PBZ. Ele-
vated, concentration-dependent levels of phenolic compounds were also
found in resting apical buds of PBZ-treated trees (Kurian et al., 1994). They
suggested that low cytokinin activity and high phenolic levels in buds con-
tributed to inhibition of shoot initiation.
The combined impact of the gibberellin synthesis-inhibiting triazoles on
shoot initiation, induction, and elongation implies that several different
Reproductive Physiology 123
gibberellins regulate specic activities in mango plants. This is supported by
the inhibitory effect of GA
3
on shoot initiation in contrast with early initia-
tion of owering in triazole-treated trees. Compression of reproductive and
vegetative shoot internodes may involve inhibition of GA
1
synthesis. Stimu-
lation of owering instead of vegetative growth during early initiation in
triazole-treated plants in marginal or non-oral inductive conditions, sug-
gests that the putative VP, a gibberellin other than GA
3
or GA
1
, is reduced
when gibberellin synthesis is inhibited.
5.7 Photoassimilate Inuence on Flowering
Flowering may be regulated by C:N ratios with high levels being conducive
to owering (Kraus and Kraybill, 1918). Photoassimilates reaching the apical
bud from leaves was central to several theories of oral induction (Sachs,
1977; Bernier and Sachs, 1979; Bernier et al., 1981, 1993; Bernier, 1988) includ-
ing mango (Mallik, 1951; L.B. Singh, 1960; Chacko and Ananthanarayanan,
1982; Rameshwar, 1989) and other species (Allsopp, 1965; Sachs, 1977; Mishra
and Dhillon, 1978; Ramina et al., 1979; Bernier et al., 1981; Sachs and Hackett,
1983). The theory of photoassimilate diversion to the apical bud (Sachs et al.,
1979) is the basis for the carbohydrate-regulated owering models (see
below). Sugars are utilized during panicle development (Ravishankar and
Mohan Rao, 1982). Starch reserves and C:N ratios have been correlated with
owering (Mishra and Dhillon, 1978; Suryanarayana, 1978a, b, c; Chacko and
Ananthanarayanan, 1982; Whiley et al., 1988, 1989, 1991; Robert and Wolsten-
holme, 1992; Shivashankara and Mathai, 1995), and the subject has been
reviewed (L.B. Singh, 1960, 1972; Singh, 1979; Chacko, 1986, 1991; Chadha
and Pal, 1986; Pandey, 1989). Starch accumulation during extended periods
of canopy rest prior to owering provides supportive evidence, but there is
little consensus regarding the role of carbohydrates and N in owering.
Photoassimilates may be necessary for oral induction. If a origenic
promoting gene product is synthesized in leaves in small amounts, it must be
able to move to those buds via phloem. Due to the requirement for high sol-
ute concentrations to motivate phloem ow, the low concentration of the FP
could not cause uid movement through sieve tubes of the phloem on its
own. The much higher concentrations of photoassimilated sugars carried by
water loading into the phloem in leaves passively transports the FP towards
the various sinks, including respiring buds, where they are utilized for oral
induction.
5.8 Horticultural Manipulation of Flowering
Mango owering can be stimulated by trunk or branch girdling, defoliation
and deblossoming (Pandey, 1989). Responses vary with cultivar and envi-
ronment. Trunk girdling of mango trees to promote owering is inconsis-
tently effective (Kinman, 1918; Gaskins, 1963; Winston and Wright, 1986) and
T.L. Davenport 124
can be detrimental to trees, especially if done in subsequent years. It has been
shown to increase owering in the off year of alternate-bearing cultivars;
however, it either has no effect or is only marginally benecial in the on year
(Mallik, 1951; Rath and Das, 1977, 1979; Rath et al., 1982; Rameshwar, 1989).
Girdling in late summer or early autumn usually results in less vegetative
ushing prior to owering, which is enhanced in trees exposed to marginally
inductive conditions. Tree response is dependent on the width of the girdle.
Narrow cuts result in either a short-term or no response; whereas, girdles that
are too wide can kill trees if they do not close within a reasonable time. Gir-
dling cuts phloem transport, starves roots of photoassimilates and interrupts
auxin transport to roots (Morris and Thomas, 1978; Hegele et al., 2004). These
are detrimental to root development and can alter the bud cytokinin:auxin
ratio due to reduced cytokinin translocation from roots. This results in delayed
shoot initiation, which can impact the level of the age-dependent, putative VP
when shoot initiation occurs. The delay in ushing, therefore, enhances
owering. Defoliation of trees stimulates ushing, possibly by altering the
cytokinin:auxin ratio in buds because leaves are the primary source of auxin.
Bloom delay is useful where recurring temperatures < 15C stimulate
owering, but continued low temperatures hamper pollination, fertilization
and early fruit development (Young and Sauls, 1979; Wolstenholme and Mul-
lins, 1982a, b; Whiley et al., 1988; Galn-Saco et al., 1991). Low temperatures
cause production of seedless, underdeveloped fruit. Deblossoming stimu-
lates growth of dormant axillary buds, which produce inorescences if initia-
tion occurs under conditions conducive to oral morphogenesis (Singh et al.,
1974). Late blooms can also be promoted with ethephon (Chadha and Pal,
1986; Galn-Saco et al., 1993) and cycloheximide (Pal and Chadha, 1982;
Shu, 1993). Sprays of these compounds cause abscission of apical panicles,
thereby releasing dormant axillary buds that will produce inorescences
under cool, oral-inductive conditions of the subtropics.
5.9 Conceptual Flowering Models
Several conceptual models have been proposed that attempt to explain the
physiological basis of mango owering. Each model should be viewed as a
collection of integrated ideas, which require rigorous testing for validity
within the context of the models. A useful model should explain how ower-
ing and vegetative growth is regulated in all cultivars and races in both
humid and dry climates in the tropics and subtropics. It should also be sup-
ported by the preponderance of research evidence. The owering models are
either carbohydrate-regulated or hormone-regulated.
Carbohydrate-regulated owering models
Cull (1987, 1991) presented a holistic approach for tree crop research and
management to maximize sustainable fruit production. This concept is based
Reproductive Physiology 125
on the axiom of genotype/environment adaptability expressed through the
annual phenological cycle and is an alternative to the traditional reductive-
based approach to crop research and development. He proposed that pro-
ductive cultivars follow normal phenological patterns from year to year
due to gene expression in specic environments. A signicant departure
from this pattern results in reduced or total crop failure. Annual variations in
climatic conditions that alter tree phenology can be countered by strategic
applications of nutrients, water, plant growth regulators and canopy manip-
ulation. The model does not attempt to explain the intricacies of shoot ini-
tiation or induction, but takes a broader approach in detailing temporal
relationships between reproductive and vegetative growth that lead to reli-
able cropping.
The fundamental principle underlying this model is that yield is the
product of photoassimilate (carbohydrate) accumulation and subsequent
redistribution during the annual growth cycle. Accumulated photoassimi-
lates would drive critical growth events that require higher levels of resources
than are available from current photoassimilate supplies. Cultivars that pro-
ceed with balanced reproductive, vegetative and rest phases are more likely
to have sufcient carbon resources to meet periods of critical demand and
therefore will sustain high yields. The model illustrates oral initiation as
occurring after a 2- to 3-month rest period during autumn/winter when a
critical threshold level of carbohydrate is reached in buds together with a
putative oral stimulus. Bud break during cool weather results in a high per-
centage of owering stems (> 90%; Searle et al., 1995) with fruit set and reten-
tion suppressing vegetative ushing on individual fruiting stems until after
they have matured and harvested. Shortly after harvest, vegetative buds are
released and a ush of growth occurs during the summer, which is followed
by a period of strong root growth. The regenerated canopy becomes a source
for rebuilding photoassimilate reserves that are stored in the roots, bark and
resting stems. In the tropics, growth events are less orderly, and cultivar and
management skills are of greater importance. The pre-owering rest period
is usually achieved by drought as temperatures remain above the critical
threshold for shoot growth (15C) (Whiley et al., 1989). Other practices used
with some success to enforce canopy quiescence are girdling and the applica-
tion of growth retardants.
The principles of phenological modelling have been advanced into work-
ing pheno/physiological models for avocado (Whiley, 1994) and mango
(Searle et al., 1995). The advantage of this approach is that the annual pro-
gression of growth cycles with associated physiological changes is studied
concurrently, adding a further dimension to our understanding of tree growth
and development. Information gathered in this way provides opportunities
to identify and assess critical yield-limiting events, which in the case of man-
goes largely relates to the success or failure of owering.
Chacko (1991) proposed a owering model driven by assimilate supply
and diversion to apical meristems (Fig. 5.6). Environmental conditions, such
as water stress, cool temperatures, high evaporative demand, ooding,
girdling and other events that inhibit vegetative growth result in a shift in

T
.
L
.

D
a
v
e
n
p
o
r
t
1
2
6
FLORAL
INDUCTION
KNO
3
(cultivar and location
specific)
High
starch
MISSING
INCREASED
ASSIMILATE supply
to SHOOT APEX
ASSIMILATE
DIVERSION
from SHOOT APEX
GROWTH
STIMULATION
and high
gibberellin
Exogenous
gibberellin
LINK
?
GROWTH
CHECK
Sugar
Water stress
Low temperature
High VPD
Flooding
High temperature
High humidity
High soil moisture
High
nitrogen
High
gibberellin
levels
JUVENILITY
HEREDITY
Over vigorous
cultivars
e.g. Kensington
Dwarf/precocious
cultivars
e.g. Irwin
Frequent
flushing of
roots and shoots
Stem girdling
Root pruning
High reserves
Efficient
assimilate
partitioning
Low reserves
More wood
formation
Mild nitrogen
stress
Growth
retardants
Inhibitors
FLORAL
INHIBITION
Fig. 5.6. Chackos Assimilate Supply and Diversion Flowering Model, a carbohydrate-regulated owering model (Source: Chacko, 1991).
Reproductive Physiology 127
carbohydrate partitioning and a diversion of soluble assimilates to stem api-
ces. The elevated carbohydrate status in buds, together with a oral stimulus,
results in oral induction. Vigorously growing cultivars and juvenile plants
have low starch reserves (Whiley et al., 1988, 1989, 1991) and a diversion of
soluble assimilates from stem apices results in oral inhibition. Conditions
that promote vegetative growth, i.e. high temperature and moisture, high
gibberellins and N, also lead to oral inhibition. Experiments involving
chemical girdling of trees are based on this model (Blaikie et al., 1999).
Hormone-regulated owering models
Tri-factor Hypothesis of Flowering
Extensive work on movement of the putative oral stimulus across grafts
from donor to receptor stems (Kulkarni, 1986, 1988b) and the inhibitory inu-
ence of fruit on subsequent owering (Kulkarni and Rameshwar, 1989) form
the basis of a owering model proposed by Kulkarni (1991): the Tri-factor
Hypothesis of Flowering in mango (Kulkarni, 2004). This theory (Fig. 5.7)
proposes an interactive role for a putative, cyclically produced oral stimu-
lus in leaves, a oral inhibitor in leaves and fruits, and bud activity during
the oral cycle. During dormancy following a vegetative cycle, genetic and
Flowering promoter
synthesized in the
leaves in the floral
cycle
Flowering inhibitor
and vegetative
promoter
synthesized in the
leaves and possibly
other organs
Bud activity
in synchrony with
the floral cycle
Pure panicles Mixed leafy panicles Vegetative flush
Genetic and Environmental Factors
Fig. 5.7. Kulkarnis Tri-factor Hypothesis of Flowering in mango, a hormone-regulated
owering model (Source: Kulkarni, 2004).
T.L. Davenport 128
environmental factors determine the level of synthesis of the putative oral
stimulus. Flowering occurs only if certain correlative factors are present, for
example if the receptor bud becomes active. If fruits are or have been recently
present on the stem, vegetative growth will result. Presence of the putative
oral inhibitor in leaves interferes with expression of the oral stimulus
resulting in vegetative growth. The level of the oral stimulus determines
the response: high levels give rise to normal panicles, intermediate levels
give rise to mixed panicles and low levels result in vegetative growth.
Comprehensive Conceptual Flowering Model
This is a model of owering involving the various classes of phytohormones
(Davenport, 1992, 1993, 2000, 2003; Davenport and Nez-Elisea, 1997)
(Fig. 5.8) based on many lines of experimental evidence as well as on research
of other tropical and subtropical fruit crops with similar phenological cycles
(Menzel, 1983; Davenport, 1990, 1992; Menzel et al., 1990; Menzel and Simp-
son, 1994; Davenport and Stern, 2005). Focusing on events occurring in indi-
vidual buds, it is applicable to monoembryonic and polyembryonic cultivars
in the tropics and subtropics and attempts to explain the physiological basis
for the annual progression of the phenological cycle.
SHOOT FORMATION. Two distinct events must occur for vegetative or repro-
ductive growth to occur in resting apical or lateral buds of mango: (i) the
Mango flowering model
PHOTOASSIMILATES
FRUIT
MIXED SHOOT GENERATIVE SHOOT
CHILLING TEMP.
OTHER FACTORS?
WATER STRESS
PROMOTER
IN LEAVES
INDUCTION
VEGETATIVE SHOOT
AUXIN
GIBBERELLINS
SHOOT INITIATION
ROOT INITIATION
ROOTS
CYTOKININS
GIRDLING
CHILLING TEMP.
STORAGE CARBOHYDRATES
PRUNING
DEFOLIATION
NITRATE SPRAY
ETHYLENE
FREQUENT
VEGETATIVE
GROWTH
AUXIN
GIBBERELLINS A
3
A
x
GA
1
GA
3
GA
x
Fig. 5.8. Davenports Comprehensive Conceptual Hormone-regulated Flowering Model
(Source: Davenport and Nez-Elisea, 1997; Davenport, 2000). Single lines indicate promotive
impact. Double lines indicate inhibitory impact.
Reproductive Physiology 129
bud(s) must be initiated to grow (shoot initiation); and (ii) at the time of ini-
tiation, shoot development (i.e. vegetative, mixed, or generative) is deter-
mined (induction). Although conditions for oral induction may be present
prior to shoot initiation, determination of that inductive condition in buds is
not made until initiation occurs. Initiation and induction events are regulated
by different signals and each may be manipulated by different stimuli.
Removing the apical whorl of leaves or tip pruning physiologically mature
stems stimulates shoot initiation in apical or lateral buds, respectively. If
containerized plants are maintained in warm temperatures (30C day/25C
night) following initiation, vegetative shoot growth is induced. If they are
kept under cool conditions (18C day/10C night), initiating shoots are
induced to be generative. In either of the two temperature regimes without
pruning, they do not initiate shoots until the natural ushing event occurs
much later. They become vegetative or reproductive according to the tem-
perature at the time of shoot initiation. If transferred from cool to warm tem-
peratures before shoot initiation, new shoot growth is induced to be vegetative.
Induction is therefore determined at the time of shoot initiation, and plants
rapidly lose their oral inductive potential when removed from the cool envi-
ronment. Determination of shoot type can be reversed during morphogenesis
by transferring containerized trees from warm-to-cool or cool-to-warm con-
ditions (Batten and McConchie, 1995; Nez-Elisea et al., 1996).
INITIATION CYCLE. The cyclic initiation of vegetative or reproductive shoots
is common to all mango cultivars. Developing vegetative shoots are rich
sources of auxins and gibberellins, which may be inhibitors in an internal
cycle that regulates shoot initiation. Auxins are actively transported basipe-
tally to roots from production sites in stems (Goldsmith, 1968; Cane and
Wilkins, 1970; Wilkins and Cane, 1970; Goldsmith and Ray, 1973), and they
stimulate adventitious root growth in mango and other crops (Hassig, 1974;
Wightman et al., 1980; Sadhu and Bose, 1988; Rajan and Ram, 1989; Nez-
Elisea et al., 1992). Elevated auxin synthesis in periodically ushing shoots is
likely to form a concentrated pulse of auxin, which inhibits recurring bud
break and moves basipetally to the roots. This pulse of elevated auxin may
stimulate initiation of new root ushes following each vegetative ush.
Alteration of root and shoot growth occurs in mango (Krishnamurthi et al.,
1960; Cull, 1987, 1991; Parisot, 1988) and other tropical and subtropical trees
(Bevington and Castle, 1986; Williamson and Coston, 1989; Ploetz et al., 1991,
1993). Timing of the root ush may depend on the distance from stems to
roots, the physiological condition of the transport path, and environmental
conditions (i.e. temperature or water relations).
New roots that develop following growth stimulation are a primary
source of cytokinins (Davies, 1995). Cytokinins are transported passively to
stems via the xylem sap in all plants and are active in bud break (Went, 1943;
Kende and Sitton, 1967; Sitton et al., 1967; Itai et al., 1973; Haberer and Kieber,
2002). Cytokinins stimulate shoot initiation in mango (Chen, 1985; Nez-
Elisea et al., 1990) and other plants (Oslund and Davenport, 1987; Belding and
Young, 1989; Williamson and Coston, 1989; Davenport, 1990; Davies, 1995;
T.L. Davenport 130
Henny, 1995). Auxin inhibits shoot initiation (Davies, 1995) and confers api-
cal dominance by preventing axillary bud break. Leaf-produced auxin and
petiolar auxin transport capacity declines as leaves age (Davenport et al.,
1980). Auxin and cytokinins may therefore be involved in the periodic cycle
of bud break.
A critical balance of these two phytohormones, possibly modulated by
GA
3
, may regulate shoot initiation. During a rest period, the inhibitory action
of auxin transported to buds decreases with time; whereas, cytokinin lev-
els in buds increase (Chen, 1987). When a critical cytokinin/auxin ratio is
achieved, the buds are stimulated to grow, thereby resetting the cycle with
initiation of new shoots. The interaction of auxin and cytokinin to control
bud break in plants is a concept that is supported by molecular studies (see
review by Nordstrom et al., 2004). Moreover, initiation of shoot growth occurs
following pruning, defoliation or the application of thidiazuron (Nez-
Elisea et al., 1990). Vigorous cultivars (Whiley et al., 1989) and young, small
trees under vegetatively promotive conditions ush frequently with only
short periods of rest; however, this cycle slows with age. Old centennial trees
ush infrequently (N. Golez, personal communication, the Philippines, 1989).
Foliar or soil-applied NO
3

stimulates initiation of reproductive shoots


only if applied after resting stems have attained an age to overcome any veg-
etatively inductive inuence. In contrast, high N in soils leads to high N lev-
els in leaves resulting in frequent vegetative ushes. The mechanism whereby
NO
3

stimulates shoot initiation is unknown.


Seeds are rich sources of auxin and gibberellins, which contribute to the
strong inhibition of bud break commonly observed on fruit-bearing mango
stems. The longer that fruit are attached to stems, the longer the postharvest
inhibition may last in the stem (Kulkarni and Rameshwar, 1989; Kulkarni,
1991).
Water stress inhibits shoot initiation by its direct impact on cell division
and elongation possibly by interfering with translocation of cytokinins from
roots. There is little evidence that water stress is directly involved in induc-
tive processes. During water stress, roots continue to grow and produce cyto-
kinins (Itai and Vaadia, 1965; Itai et al., 1968; Wu et al., 1994). Reduced xylem
ux due to limited soil hydration, and transpiration due to increased sto-
matal resistance during water stress may reduce the amount of cytokinins
reaching stems. After rewatering, the increased levels of cytokinins in roots
may translocate to and accumulate in buds. Auxin synthesis and transport
from leaves are reduced during water stress (Davenport et al., 1980) and may
require several days for correction after rewatering. This rapid shift in the
cytokinin/auxin ratio of buds may explain the shooting response that occurs
soon after relief of water stress. GA
3
may act with auxin to inhibit shoot ini-
tiation (Davenport et al., 2001b). Early owering in plants treated with PBZ
may be a response to lowered gibberellin levels, thus lowering the level of
initiation inhibitor.
This model could explain why sectors of tree canopies ush in the trop-
ics. Mango trees ush often and synchronously throughout the canopy when
they are young. With advancing age, the frequency of ushing is reduced
Reproductive Physiology 131
and synchrony is lost, resulting in sporadic ushes of vegetative or reproduc-
tive growth in sections of the canopy. As the distance between stems and
roots increases, the time required for transport of the putative pulses of ele-
vated auxin levels to roots, formed during a vegetative ush, is increased.
Groups of stems exhibiting simultaneous ushing ultimately connect to a
common branch. Dye trace studies indicate that water transport remains in
strict phylotaxic alignment from secondary roots to the canopy, even in large
trees (T.L. Davenport, unpublished results, Florida, 1991). Unless disturbed
by girdling or by pruning of branches or roots, specic branches in the can-
opy communicate only with those roots in phylotaxic alignment with them.
The hormone transport time may vary among sections of the canopy as the
tree grows. This generates individual initiation cycles in sections of the can-
opy that are separately maintained unless resynchronized with the rest of the
tree following a canopy-wide environmental trigger.
Synchronization of growth throughout trees occurs following exposure
to low temperature, water stress, light pruning of the entire tree and any
condition that would increase the postulated cytokinin/auxin ratio in buds
throughout the canopy. An increased ratio may occur by inhibiting auxin
transport from leaves to buds, or increasing cytokinin translocation from
roots to stems. Winter in the subtropics would reduce auxin transport;
whereas, water stress in the tropics may impact the availability of cytokinins
from roots and auxin from leaves. The intensity of the initiation response (i.e.
synchronization of ushes in the canopy) may be regulated by decreased
auxin transport at low temperatures, the base level of which may be deter-
mined by the age of individual stems. Passage of a strong, extended cold
front during subtropical winters produces synchronized owering. Milder
winters with weak cold fronts result in asynchronous owering in sections of
trees. The oldest sectors of canopies ower rst, followed by sectors bearing
sequentially younger ushes in subsequent cold fronts. Vegetative ushes
occur when night temperatures are > 18C for signicant periods between
cold fronts.
INDUCTION SWITCH. Floral or vegetative induction is possibly governed by the
interactive ratio of a FP that is up-regulated in low temperatures to an age-
regulated VP in leaves at the time of shoot initiation. High FP:VP ratios would
be conducive to induction of generative shoots, low ratios conducive to veg-
etative shoots and an intermediate ratio of the two would be conducive to
mixed shoots. Regardless of the endogenous levels of the two components
perceived in buds at the time of initiation, owering and vegetative growth
responses can best be explained by the ratio of the two.
Although the putative FP seems to be up-regulated during leaf exposure
to cool temperatures (< 18C), there appears to be a basal level present at all
times in leaves exposed to higher temperatures. Flowering of mango occurs
in low-latitude tropics lacking cool night temperatures when stems become
sufciently aged so that the ratio of the basal level of resident FP to decreas-
ing VP increases to a critical threshold to provide oral induction when
shoots are initiated. This could explain how owering on non-synchronized
T.L. Davenport 132
branches may occur at any time of the year in trees growing in low-latitude
tropics. High proportions of mixed shoots are commonly found in these con-
ditions, indicating the marginally oral-inductive ratios present under these
conditions. In contrast, owering in younger stems having higher levels of
VP is observed only when initiation occurs in cool, oral-inductive tempera-
tures. More owering occurs throughout the canopy when stems are exposed
to cool temperatures, attributable to the higher ratio of up-regulated FP to
resident VP.
Genetic differences in base levels of the putative FP and/or VP or the
receptors of these components could explain the range in owering responses
in tropical and subtropical cultivars and why a cultivar grown in an environ-
ment different from that in which it was selected is less productive. Cultivars
selected in the subtropics usually ower as well in the low-latitude tropics as
those selected in the tropics. Cool temperatures in the subtropics sometimes
cause earlier owering in tropical cultivars than those selected in the sub-
tropics. Kulkarni (1991) demonstrated that several multi-owering cultivars
can induce owering in receptor graft plants and cause a range of the ower-
ing response of the receivers to donors. Some cultivars may produce higher
base levels of putative FP than others. These are the same cultivars that read-
ily ower under warm temperatures and ower early during cool winter
months. The Comprehensive Conceptual Flowering Model suggests that
owering can occur at any time in any cultivar regardless of origin so long as
stems are sufciently old to reduce the VP level to below the critical FP/VP
ratio when initiation occurs.
Although the putative FP, perhaps a product of an ortholog of the Arabi-
dopsis FT gene, has not been identied, the VP may be a gibberellin. Triazoles
and other plant growth retardants that inhibit gibberellin synthesis, promote
strong and out-of-season owering under conditions that would normally
be marginally or non-oral inductive.
PHOTOASSIMILATES. Photoassimilates produced by leaves provide carbohy-
drates essential for development of roots and other plant organs, including
fruit. They are either used immediately by the nearest sinks (Finazzo et al.,
1994) or are stored in locations throughout the tree to be used when demand
for carbon resources exceeds the existing photosynthetic supply (Whiley
et al., 1988, 1989, 1991). A direct role for carbohydrates in shoot initiation or
induction is not part of this model, although they facilitate mass ow in
phloem from leaves to passively carry the FP to buds.
ALTERNATE BEARING. High levels of auxin and gibberellins produced in seeds
possibly inhibit shoot initiation on fruit-bearing stems for weeks or months
following fruit removal. Rapid production of new shoots following light
pruning of fruit-bearing stems after harvest indicates that residual levels of
auxin and gibberellins linger only in the rachis and last intercalary unit. If
fruit are not set on the lingering rachis, there is less inhibition. Heavy fruit set
in 1 year impacts the timing of subsequent shoot initiation on the large num-
ber of fruit-bearing branches. Substantial delays in subsequent vegetative
Reproductive Physiology 133
ushes until close to the normal owering period impact the owering abil-
ity of young shoots. This may explain the occurrence of chronic alternate
bearing in some cultivars.
5.10 Floral Management
The Comprehensive Conceptual Flowering Model is consistent with growth
and development patterns of mango trees in the tropics and subtropics. It
provides a reasonable explanation for the various events in the phenologi-
cal model of Cull, predicts what will happen under a dened set of cir-
cumstances and is being used to develop strategies for consistent mango
owering. Flowering can be obtained at any time of the year in a owering
management programme (Davenport, 2003).
The owering management programme begins each season with tip
pruning of the entire canopy of orchard trees. Tip pruning can be done imme-
diately after harvest to move production forward in the following year or
c.12 months after harvest, depending upon cultivar, in order to achieve har-
vest at the same time as the previous year. If sufcient soil water is available
at the time of pruning, a vegetative ush will occur on all pruned stems c.1
month later. The number of new shoots that will mature to become stems will
be ve- to eightfold greater than the original number of pre-pruned stems
due to initiation of many lateral vegetative shoots on each stem. This increase
in terminal stem number in the canopy will be reected in a concomitant
increase in yield. The frequent ushes that can cause an early second ush of
vegetative growth tend to be suppressed.
The new stems must not ush a second time until at least 5 months after
pruning (Davenport, 2006). If they ush within 34 months after pruning,
they will be induced to be vegetative. Pre-prune leaf N levels in the stems
must be 1.11.4% in order to suppress a second ush of vegetative growth
during the rainy season. Mild water stress after the post-prune ush during
the dry season will suppress a second, undesired vegetative ush when leaf
N levels are above the optimum range. Pruning near the end of the dry sea-
son in non-irrigated or furrow-irrigated trees should be avoided. Transition
from dry to wet season 23 months after pruning causes a rain-stimulated
vegetative ush prior to achieving sufcient age of stems from the last ush.
Test sprays of 4% KNO
3
on two to three representative trees should be
applied 5 months (for easily induced cultivars) and 6 months (for more dif-
cult to ower cultivars) after pruning. If no developing shoots occur within
2 weeks, the spray is repeated. A owering response is usually evident after
the second application. The other trees that were pruned on or near the same
date can then receive the foliar spray and will respond by synchronized ow-
ering. Although Davenport (2003) described the appropriate timing of PBZ
in a owering management programme, it is not recommended because
owering can be achieved without it. For orchard trees to be amenable to tip
pruning, efcient spray application of KNO
3
and easy harvesting, they
should be no taller than 4 m. Pruning to rejuvenate large mango trees and
T.L. Davenport 134
properly shape trees for the annual owering management programme is
recommended (Davenport, 2006).
5.11 Floral Biology
Detailed descriptions of generative shoots were reviewed in Davenport
and Nez-Elisea (1997). Juliano and Cuevas (1932) described ovary devel-
opment.
Sex ratio
Sex ratio (i.e. the proportion of perfect to staminate owers) is a variable
component within panicles, trees and among cultivars. This ratio varies with
cultivar, but is usually < 50% (Davenport and Nez-Elisea, 1997). Most per-
fect and staminate owers are borne in the proximal portion of panicles due
to their architecture (Musahib-ud-din and Dinsa, 1946; Cobin, 1950). The
variability in the perfect/staminate ower ratio may be governed by physi-
ological and environmental conditions. Most studies indicate that although
the total number of owers is substantially less in the distal half of panicles,
there is a greater proportion of perfect owers in this region (Davenport and
Nez-Elisea, 1997); however, this condition may be reversed in some culti-
vars (Hussein et al., 1989).
Perfect owers tend to form in the terminals of individual inorescences
while staminate owers are displayed in the earlier forming owers located
closer to the panicle axis. When panicles begin to elongate in the lower ino-
rescences, only staminate owers form and the perfect owers form at the
terminus of each lateral inorescence. As more distally located lateral ino-
rescences begin elongation and anthesis, they too rst display staminate
owers before perfect owers. These inorescences are progressively shorter
than previously formed proximal inorescences, and there are fewer stami-
nate owers. The nal vertical spike of the panicle is composed almost exclu-
sively of perfect owers. Flowers abscise soon after anthesis, thereby shifting
the sex ratio. The sex ratio should include sex type of all owers in each
panicle; however, sex ratios are normally determined at some arbitrary
moment during panicle development. Thus, the sex ratio is naturally vari-
able, increasing from extremely low to extremely high values so that timing
of observations during panicle development is critical.
Environmental determinants of sex ratio
Tropical cultivars yield poorly in the subtropics due to a small proportion of
perfect owers on inorescences (Singh and Singh, 1959; Singh et al., 1965;
Singh, 1971). Cool weather during inorescence development contributes to
fewer perfect owers (Naik and Mohan Rao, 1943; Singh et al., 1965, 1966).
Reproductive Physiology 135
Inorescences that emerge during the middle and end of the owering sea-
son produce two and seven times more perfect owers, respectively, than the
early breaking inorescences (Majumder and Mukherjee, 1961; Singh et al.,
1966). This response correlates with higher temperatures later in the ower-
ing season. In controlled-environment studies, low temperatures (15C
day/10C night) reduced the proportion of perfect owers, particularly in
tropical, polyembryonic cultivars relative to subtropical, monoembryonic
cultivars (Sukhvibul et al., 1999).
Physiological determinants of sex ratio
Endogenous factors affect the ratio of perfect to staminate owers. Bajwa
et al. (1956), Majumder and Mukherjee (1961) and Joubert et al. (1993) reported
that lateral inorescences on mixed shoots carried higher proportions of per-
fect owers. Inorescences on older trees produce higher proportions of per-
fect owers than those on young trees (Naik and Mohan Rao, 1943; Majumder
and Mukherjee, 1961; Chacko and Randhawa, 1971; Pandey, 1989). This also
occurs in inorescences borne on grafted compared with seedling trees
(Musahib-ud-din and Dinsa, 1946). The effect of tree maturity or rootstocks
on sex ratio of owers is not understood. Panicles carried within the canopy
of some cultivars (Majumder and Mukherjee, 1961; Singh et al., 1966) or on
particular sides of the canopy (Mukherjee, 1953; Majumder and Mukherjee,
1961) have been reported to have higher proportions of perfect owers.
Application of some hormones and growth regulators alters the sex ratio
of inorescences. GA
3
, applied at concentrations of 50100 mg/l just prior to
inorescence shoot initiation, substantially reduces the proportion of perfect
owers (Maiti, 1973), as do combination sprays of urea (0, 3 and 6%) and GA
3

(0, 15 and 30 mg/l) (Rajput and Singh, 1989). Soil-applied PBZ (10 g ai/tree)
signicantly increases the ratio of perfect/staminate owers (Kurian and Iyer,
1993a). Increases in oral ratio also occur with daminozide, whereas maleic
hydrazide either had no effect or lowered the ratio (Singh et al., 1965; Subhad-
rabandu, 1986). Foliar application of 50 mg/l BA with 2% calcium ion (Ca
2+
)
increased the proportion of perfect owers (Singh and Rajput, 1990). Naphtha-
lene acetic acid (NAA) at concentrations of 50, 100 and 200 mg/l increased the
perfect/staminate ower ratio (Mallik et al., 1959; Singh et al., 1965).
Other factors inuencing sex ratios of inorescences include stem age
and mineral nutrients. Gunjate et al. (1983), Desai et al. (1986) and Hussein
et al. (1989) reported that inorescences from stems that grew at different
times during the previous summer/autumn period had signicantly differ-
ent perfect/staminate ower ratios. Singh and Dhillon (1987) found that
boron (B) levels affect sex ratio.
Sex ratios can be manipulated with growth regulators, but has no com-
mercial advantage (A.W. Whiley, personal communication, Queensland,
1996). Increases in fruit yield resulting from chemically increased perfect/
staminate ower ratios have not been observed, suggesting that perfect
ower numbers are not the primary limitation to crop performance (Schaffer
T.L. Davenport 136
et al., 1994). If only one or two fruits were set on each terminal, the tree would
carry an unusually heavy crop. It is unlikely that reduced perfect ower
numbers due to cool temperatures during inorescence development is
directly responsible for poor fruit set and yields. Pollen viability, growth and
ovule fertilization are probably the main factors contributing to low fruit set
under these conditions.
Anthesis and dehiscence
Floral anthesis generally occurs at night in polyembryonic cultivars (Wagle,
1929; Torres, 1931; Galang and Lazo, 1937; Pimentel et al., 1984) and at night or
early morning in monoembryonic types (Popenoe, 1917; Musahib-ud-din and
Dinsa, 1946; Singh, 1954a; Randhawa and Damodaran, 1961a, b). Subsequent
dehiscence of the four-lobed anthers occurs during the daylight morning hours
revealing pale blue pollen grains (Torres, 1931; Galang and Lazo, 1937; Mallik,
1957; L.B. Singh, 1960). Anthesis and anther dehiscence are delayed by low
temperatures or overcast days (Singh, 1954a; De Wet and Robbertse, 1986a).
Dehiscence is also delayed by high RH, and pollination occurs primarily
around midday (Mallik, 1957; Randhawa and Damodaran, 1961a, b).
Stigmas are receptive from c.18 h prior to anthesis to at least 72 h after
anthesis with optimum receptivity within 3 h from anthesis (Popenoe, 1917;
Wagle, 1929; Sen et al., 1946; Singh, 1954a; Spencer and Kennard, 1956; Rand-
hawa and Damodaran, 1961a, b; Gunjate et al., 1983; Pimentel et al., 1984;
Robbertse et al., 1994). Receptive stigmas are shiny and white-green, whereas
non-receptive stigmas are desiccated and yellow-brown. Pollen germination
generally occurs within 90 min of deposition, although the percentage ger-
mination of pollen deposited on stigmas is relatively poor (Singh, 1954a; S.N.
Singh, 1961). Pollination is initiated by the formation of two unusual protu-
berances that meet to form a bridge or ponticulus connecting the dorsal side
of the ovule with the ovary wall in line with the base of the style (Joel and
Eisenstein, 1980). The ponticulus may guide the elongating pollen tube to the
ovule. Ovule fertilization occurs 4872 h after pollination (S.N. Singh, 1961;
Ram et al., 1976). Both zygote cell and endosperm nuclear division appear to
rest for about 2 weeks following pollination despite cell division and growth
of the ovary (Sharma and Singh, 1972; Ram et al., 1976). A description of
embryo development is presented by U.R. Singh (1961). Application of B
may improve stigma receptivity, pollen tube germination and growth and
ovule fertilization (De Wet and Robbertse, 1986b; Robbertse et al., 1988; De
Wet et al., 1989) as well as fruit development (Chen, 1979; Robbertse et al.,
1990); however, Jutamanee et al. (2002) could not verify the effect of B.
Pollen
Pollen grains are 2045 m long and are oblong when dry and more
spherical when hydrated (Popenoe, 1917; Jivanna Rao, 1923; Bijhouwer, 1937;
Reproductive Physiology 137
Mukherjee, 1950; Singh, 1954a; Randhawa and Damodaran, 1961b; S.N.
Singh, 1961). There are generally three equilateral, tapering furrows along
the longitudinal sides of dry pollen that give hydrated grains a roughly tri-
angular shape when viewed on end (Popenoe, 1917; Singh, 1954a; S.N. Singh,
1961; U.R. Singh and A.P. Singh, 1973). Each furrow has a germpore in its
centre (Mukherjee, 1950; S.N. Singh, 1961). Anthers produce c.250650 pollen
grains with a mean of 410 grains per anther (Popenoe, 1917, 1920; Spencer
and Kennard, 1955).
In vitro germination of mango pollen has been reported (Popenoe, 1917;
Spencer and Kennard, 1955; Young, 1958; Randhawa and Damodaran, 1961b;
S.N. Singh, 1961). Germination on stigmas was c.10% less than that on articial
media, that is 78% across cultivars (Spencer and Kennard, 1955), although
lower rates of germination have been reported (Mukherjee, 1950). Mango pol-
len is most viable soon after anther dehiscence and rapidly degrades (Sen et al.,
1946; Spencer and Kennard, 1955; Mallik, 1957; S.N. Singh, 1963). Although the
initial percentage of viable pollen is generally 90% during warm weather
(Popenoe, 1917; Mukherjee, 1949a, b; Singh, 1954a; S.N. Singh, 1961), cool tem-
peratures early in the owering season result in abnormal, non-viable pollen
grains (Popenoe, 1917; U.R. Singh and A.P. Singh, 1973; Shu et al., 1989; Gazit
et al., 1992; Issarakraisila et al., 1992). The pre-vacuolate stage of meiosis during
microsporogenisis is the most sensitive period to temperatures < 10C (Issara-
kraisila and Considine, 1994). Germination and pollen tube growth are reduced
by cool temperatures (S.N. Singh, 1961; Mullins, 1986; Robbertse et al., 1988;
Whiley et al., 1988; De Wet et al., 1989) and completely inhibited at tempera-
tures < 15C (Popenoe, 1917; Young, 1955; Sukhvibul et al., 2000).
Pollination
Pollination is a major yield-limiting constraint, due to the large number of
owers on trees and low fruit set. Unlike polyembryonic cultivars, which
produce nucellar embryos, pollination is necessary for fruit set with mo-
noembryonic cultivars (Popenoe, 1917; Young, 1942; Spencer and Kennard,
1955; Gunjate et al., 1983; Pimentel et al., 1984; Robbertse et al., 1994). Pollen
compatibility within and between cultivars has been widely investigated.
Complete or partial self-incompatibility has been reported (Mukherjee et al.,
1961; Singh et al., 1962b; Sharma and Singh, 1970, 1972; Ram et al., 1976; De
Wet et al., 1989; Robbertse et al., 1993). Incompatibility is evident by degen-
eration of embryonic and nucellar tissues and excessive loss of fruitlets. Cross
incompatibility between some cultivars has also been noted (Saha and Chhonkar,
1972; Ram et al., 1976; Robbertse et al., 1993).
Wind
Early investigators concluded that the species is wind pollinated (Hartless,
1914). Initially wet pollen dries to a powdery consistency on anthers soon
after anthesis in dry conditions (Pimentel et al., 1984), whence it is likely to be
liberated in moving air or via gravity to adjacent stigmas on the same and
T.L. Davenport 138
nearby owers (Naik and Mohan Rao, 1943; Mallik, 1957). Singh (1954a) and
S.N. Singh (1961) suggested, however, that the amount of pollen moving in
air streams was too low for wind to be a pollination vector. They did not
report the location of pollen-collecting slides or take into account the close
proximity of owers within inorescences or numbers of open owers in the
canopy. Panicles bagged to exclude pollinating insects were reported to set
fruit (Free and Williams, 1976), which were retained to maturity, thereby con-
rming that mango pollen can be transferred by air movement or gravity
(Bijhouwer, 1937; Mallik 1957). The tacit assumption that open-pollinated
owers are exclusively crossed is likely to be incorrect, although mango may
favour cross-pollination.
Insect
Popenoe (1917) reasoned that pollen transfer occurs primarily within owers
by insects. Panicles bagged to exclude insect visitation generally result in less
fruit set than on panicles in the open (Popenoe, 1917; Musahib-ud-din and
Dinsa, 1946; Mallik, 1957; Free and Williams, 1976; Jiron and Hedstrom, 1985).
Insects working mango owers include Diptera, Hymenoptera, Lepidoptera
and Coleoptera (Popenoe, 1917; Simao and Maranhao, 1959; Randhawa and
Damodaran, 1961b; McGregor, 1974; Anderson et al., 1982; Jiron and Hed-
strom, 1985). Flies of various genera are common on mango owers (Pope-
noe, 1917; Burns and Prayag, 1921; Bijhouwer, 1937; Singh, 1954a; Spencer
and Kennard, 1955; Eardley and Mansell, 1993). Polistes wasps are observed
on mango owers but are considered to be ineffectual for pollen transfer
(Spencer and Kennard, 1955; Free and Williams, 1976; Wolfenbarger, 1977).
Honeybees (Hymenoptera) are occasional visitors (Young, 1942; Simao and
Maranhao, 1959; Smith, 1960; Morton, 1964; Jiron and Hedstrom, 1985; Mac-
Millan, 1991; Du Toit and Swart, 1993, 1994; Eardley and Mansell, 1993, 1994),
but only if other more inviting owers are not present (Spencer and Kennard,
1955; Free and Williams, 1976; McGregor, 1976). They are assumed to be the
most effective pollinators of mango and may be more effective if hives are
placed in orchards during owering (Du Toit and Swart, 1993, 1994). Ander-
son et al. (1982) recorded actual pollen transfer on mango owers by insects
and found, in order of importance, the most efcient pollinators to be wasps,
bees, large ants and large ies.
With few exceptions (Mallik, 1957), pollen deposition rates are generally
low (Naik and Mohan Rao, 1943; Mukherjee, 1951). Differences in pollination
rates can be attributed to environmental conditions during owering, differ-
ing attraction of insects to specic cultivars, proximity of more attractive
owering species or a combination of the above. Young (1942) observed that
insects visit only 1012% of available owers. Depending on weather condi-
tions, insect activity on mango owers is usually continuous from early
morning to late afternoon, but nocturnal activity of some species has also
been reported (Jiron and Hedstrom, 1985).
The role of insects in cross-pollination is not understood. Anderson et al.
(1982) observed various insects carrying pollen to and from owers and
noted pollination subsequent to those visits; however, they made no distinction
Reproductive Physiology 139
between actual pollen depositions by visiting insects and pollen transferred
by other means. Wester (1920) considered that pollination is facilitated by
wind and to a lesser extent by insects, and this conclusion is probably correct
in most environments. Self-pollination within owers by insects while the
pollen is still damp is likely to occur. Use of isozyme (Degani et al., 1990; Rob-
bertse et al., 1993) and microsatellite DNA markers (Adato et al., 1995; Schnell
et al., 2005) to discern ratios of cross- versus self-pollinated fruitlets and off-
spring is the most accurate procedure to conrm self- and cross-pollination.
Initial fruit set of pollinated owers is inconsequential since most of these
fruitlets abscise before reaching maturity (Lynch and Mustard, 1950; Singh,
1954a; Randhawa and Demodaran, 1961a).
5.12 Fruit Development
Fruit growth is correlated with several growth regulating substances. Enlarge-
ment is sigmoidal reaching a constant size c.23 weeks before maturity
(Singh, 1954a; Randhawa and Damodaran, 1961a, b; Ram, 1983; Prakash and
Ram, 1984). The highest rates of fruit growth have been correlated with peak
levels of putative endogenous auxins found in seeds (Chacko et al., 1970a, b;
Singh and Singh, 1974; Chen, 1981; Ram, 1983; Prakash and Ram, 1984).
Baghel et al. (1987a, b) reported increased fruit mass with a combination
spray of NAA and urea to pre-anthesis panicles. Free and bound gibberellins,
especially in seeds (Ogawa, 1963; Ram, 1983), peak similarly to putative aux-
ins during fruit development (Chacko et al., 1970c, 1972a; Ram and Pal, 1979;
Chen, 1981). Cytokinins tentatively identied as zeatin and zeatin riboside
and other active fractions appear to vary in concentration in seeds and peri-
carp with two distinct peaks of activity. No particular relationship to growth
was evident (Ram, 1983; Ram et al., 1983). Seeds produced more cytokinin
activity than did the pericarp. In contrast, Chen (1983) found only one peak
of cytokinin activity in seeds and pulp occurring at about half full size of
both seed and pericarp. Seed tissues contain the highest cytokinin activity.
Levels of endogenous auxin, gibberellins and cytokinins in leaves during
fruit set were compared to production in leaves during other periods with-
out conclusive results (Paulas and Shanmugavelu, 1989).
5.13 Stenospermocarpy
Abscission of non-fertilized and fertilized owers is normal. Fruitlet abscis-
sion from pea size on is often associated with embryo abortion (Chandler,
1958; U.R. Singh, 1961; Singh, 1964; Lakshminarayana and Aguilar, 1975;
Ram et al., 1976) and is referred to as stenospermocarpy (Soule, 1985). Steno-
spermocarpy in mango is unusual (Chacko and Singh, 1969a) but occurs
regularly in some cultivars (Nez-Elisea and Davenport, 1983; Whiley et al.,
1988). Stenospermocarpic fruitlets have slower growth rates than seeded
fruit, generally become misshapen and fail to reach full size.
T.L. Davenport 140
Stenospermocarpy in some cultivars has been correlated with low tem-
peratures during owering and early fruit set in the subtropics (Lakshmina-
rayana and Aguilar, 1975; Young and Sauls, 1979; Whiley et al., 1988; Schaffer
et al., 1994). Nez-Elisea and Davenport (1983) reported that stenospermo-
carpic fruit often occur distal to seeded fruitlets within panicles and sug-
gested that embryo abortion is associated with high temperatures when these
latter fruit set. Secondary spring owering of some monoembryonic culti-
vars under high temperatures has resulted in high proportions of steno-
spermocarpic fruit (E.K. Chacko, personal communication, Australia, 1995).
Application of auxins, gibberellins and cytokinins produce seedless fruit in
some cultivars, suggesting that the abscission zone is protected by these hor-
mones despite the loss of the endogenous supply from the aborted seed (Ven-
kataratnam, 1949; Chacko and Singh, 1969a, b; Kulkarni and Rameshwar,
1978).
5.14 Fruit Set and Retention
Fruit set and retention of mango was recently reviewed by Singh et al. (2005).
Abscission of owers and fruitlets is accomplished by rapid formation of a
separation layer in the abscission zone in the pedicel-peduncle junction (Bar-
nell, 1939). U.R. Singh (1961) described formation of the abscission zone dur-
ing oral ontogeny and of the separation layer during abscission of owers
and fruitlets. The majority of panicles lose all fruitlets (Nez-Elisea and
Davenport, 1983). The pattern of fruitlet abscission is asymptotic with the
greatest losses occurring during the rst weeks after anthesis (Nez-Elisea
and Davenport, 1983; Prakash and Ram, 1984; Searle et al., 1995). Except for
the tendency to retain fruit in the distal portion of panicles, abscission of
owers and fruitlets is random. It can involve fruitlets regardless of size or
location.
Of the 813% of perfect owers setting fruit, < 1% reach maturity (Bijhou-
wer, 1937; Sen, 1939; Naik and Mohan Rao, 1943; Mukherjee, 1949b; U.R.
Singh, 1960; Randhawa and Damodaran, 1961a; Singh, 1978; Gunjate et al.,
1983; Prakash and Ram, 1984). Generally, most fruit are set on the most distal
spike portion of panicles (Chadha and Singh, 1963; Nez-Elisea and Daven-
port, 1983). Fruit loss has been associated with embryo abortion, resulting in
blackened or shrivelled embryos (Singh, 1954a, 1964; Chandler, 1958; U.R.
Singh, 1961; Sharma and Singh, 1972; Ram et al., 1976) after the fruit is sepa-
rated from the tree (Nez-Elisea and Davenport, 1983).
Sex ratio
The perfect/staminate oral ratio in panicles may inuence fruit set and pro-
ductivity (Naik and Mohan Rao, 1943; Singh, 1954b; Singh and Singh, 1959;
U.R. Singh, 1960). Mallik (1957) noted that more perfect owers are formed
in on than off years of alternate-bearing cultivars. Other studies, however,
Reproductive Physiology 141
have demonstrated that the number of perfect owers does not correlate
with subsequent yield (Randhawa and Damodaran, 1961a) so long as the
proportion of perfect owers is not < 4% (Singh, 1964, 1971). Most fruit are
borne in the distal portion of panicles (Shawky et al., 1977), which may be
correlated with the high ratio of perfect to staminate owers there. Schole-
eld and Oag (1984) estimated that one mature fruit is harvested for each 169
perfect owers in the distal half of the panicle; whereas 592 perfect owers
are required to produce one fruit in the proximal half. Therefore, intrinsic
factors other than sex ratio regulate fruit set.
Mineral nutrients
Boron is one of seven micronutrients required for normal plant growth. The
physiological function of B is unknown (Hu and Brown, 1994), although it is
essential for oral development, pollen germination, pollen tube growth,
embryo development and growth of organs (i.e. fruit) (Vasil, 1963; Agarwala
et al., 1981; Dell and Huang, 1997; Shorrocks, 1997). Decient soils are com-
monly found in mango-producing areas of Australia, Thailand, Central and
South America and Africa where symptoms are common (Aitken et al., 1987;
Singh et al., 2005). Boron applications to decient mango trees increase normal
fruit set (Robbertse et al., 1990; Raja et al., 2005). Fruitlet abscission in mangoes
has also been attributed to zinc (Zn) deciency (Jiron and Hedstrom, 1985).
Hormonal control
Auxin
Research demonstrating improved fruit set and retention following applica-
tion of several auxin analogues to pre-anthesis panicles or to panicles bearing
fruitlets of various sizes has been reviewed (Davenport and Nez-Elisea,
1997; Singh et al., 2005). NAA is the most effective auxin analogue for improv-
ing fruit retention (Prakash and Ram, 1986; Khan et al., 1993). Initial fruit set
was substantially increased when sprays of 200 mg/l indole acetic acid (IAA)
were applied to developing panicles (Singh et al., 1965). A 300400% increase
in fruit set resulted when NAA (40 or 50 mg/l) was sprayed at the pre-anthesis
stage (Ram, 1983; Singh and Ram, 1983; Prakash and Ram, 1986). Chen (1981)
reported no effect on fruit retention when 5 mg/l of either naphthaleneacet-
amide or -naphthoxoyacetic acid were applied three times at 2-week inter-
vals to panicles in which fruit had reached 4 mm in diameter.
Despite increased fruit retention of mango using exogenous applications
of auxins, few studies have examined endogenous auxins in fruit as related
to retention (Chacko et al., 1970a, b; Ram et al., 1983; Prakash and Ram, 1984).
Singh and Singh (1974) were unable to detect signicant differences in endog-
enous auxins or inhibitors when comparing alternate and regular bearing
cultivars. Chen (1981) observed lower levels of auxin-like substances in
T.L. Davenport 142
mesocarp and calyx tissues of abscised fruits than those of intact fruits. Simi-
lar decreases in auxin and gibberellins with an increase in abscisic acid as
fruitlets abscised were reported by Bains et al. (1999). The interaction of auxin
in fruit and abscission zones to maintain mango fruit retention is not clear.
Continuous auxin synthesis and basipetal transport to the abscission
zone is critical for maintenance of plant organs, including fruit (Crane, 1964;
Nitsch, 1965; Morgan et al., 1977; Davenport et al., 1980; Roberts and Osborne,
1981). Increased mango fruit set and retention in response to exogenously
applied auxins conrms this requirement; however, other hormonal factors
also appear to be involved. Developing seeds are rich sources of all the known
classes of phytohormones, including auxins (Crane, 1964; Nitsch, 1965; Chacko
et al., 1970a, b, c; Chen, 1981). Hence, exogenous enrichment of auxin in the
presence of other seed-produced phytohormones facilitates increased fruit
retention. In contrast, NAA (10 and 20 mg/l) spray-applied to bagged, self-
pollinated owers, does not result in development of stenospermocarpic
fruits beyond the marble size (Venkataratnam, 1949; Chacko and Singh,
1969a, b). Similarly, applications of 250 or 500 mg/l GA
3
or 250 mg/l BA
alone to panicles does not promote production of stenospermocarpic fruits
(Chacko and Singh, 1969a, b). Supplying exogenous -naphthoxyacetic acid
(10 mg/l), BA (250 mg/l) and GA
3
(250 and 500 mg/l) together in multiple
sprays until half grown, however, resulted in retention of several seedless
fruit to maturity. Chen (1983) and Oosthuyse (1995b) observed that gibberel-
lin, cytokinin and auxin reduce fruit drop of open-pollinated fruitlets of some
cultivars. Thus, although auxin is important for maintaining the abscission
zone, the presence of other phytohormones appears to be important for fruit-
let development (Chacko et al., 1970a, b; Ram, 1983; Ram et al., 1983).
Cytokinins
Although cytokinins are not generally thought to be associated directly with
abscission, Ram (1983) and Ram et al. (1983) concluded that low cytokinin
levels during fruit development might contribute to fruit loss. Chen (1983)
observed a correlation of low cytokinin levels in stenospermocarpic fruits
with abscission at the marble stage of growth. Application of 250 mg/l BA to
bagged panicles does not promote production of seedless fruits (Chacko and
Singh, 1969a, b). The synthetic cytokinin, N-(2-chloro-4-pyridyl)-N-phenylurea
(CPPU) also does not improve fruit set when applied alone at a rate of 10
mg/l to post-anthesis panicles (Oosthuyse, 1995b). The role of cytokinins in
separation events remains inconclusive.
Gibberellins
Gibberellins do not appear to be directly linked to the onset of abscission
(Chacko et al., 1970c, 1972a; Ram and Pal, 1979; Chen, 1981; Ram, 1983). Spray
applications of GA
3
to pre- and post-anthesis panicles to increase fruit set
and retention have been inconsistent. Increased yield (Teaotia et al., 1967;
Singh and Ram, 1983; Rajput and Singh, 1989) and production of seedless fruit
(Kulkarni and Rameshwar, 1978) have been reported from these treatments,
but Chacko and Singh (1969a, b) observed no such effects. Chen (1983) and
Reproductive Physiology 143
Oosthuyse (1995b) investigated the effects of several foliar applications of GA
3

starting at the 4 mm diameter stage, but were unable to improve fruit set.
Several classes of gibberellin-synthesis inhibitors have been tested for
reducing fruit drop. The growth retardants, daminozide and cycocel, increased
fruit set when applied to post-anthesis panicles (Singh and Ram, 1983). The
authors suggested that increased fruit retention might have been mediated
through increased cytokinin-like activity of the growth retardants. Although
initial fruit set was promoted by PBZ, yield was not improved (Kurian and
Iyer, 1993a). It is not clear whether the contrasting results of increased yield
(Kurian and Iyer, 1993b) were due to reduced fruit loss or more intense ow-
ering in response to treatment. Goguey (1990) reported increased fruit set
and retention using soil-applied PBZ at 5 g ai/tree. Spray application of uni-
conazole, a more biologically active triazole (5002000 mg/l), reportedly
increased fruit set and yield (Galila and El-Masry, 1991). It is difcult to
resolve the contradictory results demonstrating enhancement of fruit reten-
tion by GA
3
and inhibitors of its synthesis.
Inhibitors
Abscisic acid (ABA) is possibly involved in fruitlet abscission. Although cor-
relations exist between certain inhibitors and abscission of mango fruitlets,
no clear cause and effect relationships have been established. Fruit drop was
correlated with levels of an acidic inhibitor, possibly ABA (Chacko et al.,
1970b, 1972a; Singh and Singh, 1974; Ram, 1983; Prakash and Ram, 1984).
Chen (1981) reported similar changes in putative ABA with maximum levels
occurring during early fruit drop and with advancing age of fruits. Putative
ABA levels in abscised and retained fruits were compared and were highest
in the calyx and mesocarp of abscised fruitlets.
Ethylene
Ethylene has the greatest immediate impact on ower and fruitlet abscission.
Van Lelyveld and Nel (1982) reported higher levels of ethylene in abscised
fruitlets compared with those retained on trees. Nez-Elisea and Davenport
(1983, 1984, 1986) examined the dynamics of ethylene production in intact
and excised fruitlets from onset to separation. Increased production began in
explants about 26 h postharvest and increased logarithmically until fruit sep-
aration. Abscission of the fruitlets began 48 h after the onset of enhanced
ethylene production. Similar results with avocado fruitlet abscission experi-
ments (Davenport and Manners, 1982) indicate that the onset of ethylene
production in intact fruitlets is spontaneous in individual fruitlets followed
by abscission 48 h later. The pericarp provided the bulk of ethylene for induc-
tion of abscission processes; the pedicel produced no ethylene. There was
reduced fruit drop in response to inhibitors of ethylene production and action
(Singh and Ram, 1983; Naqvi et al., 1990, 1992). Whereas increased peroxi-
dase (Van Lelyveld, 1978) and polyphenol oxidase activities have been
reported in abscissed mango fruitlets (Van Lelyveld and Nel, 1982), Nez-
Elisea and Davenport (1984) observed no changes in peroxidase activity or
protein levels prior to separation of fruitlets.
T.L. Davenport 144
Abscission of stenospermocarpic fruits has been associated with small
increments in ethylene production (Nez-Elisea and Davenport, 1983). Sen-
sitivity to low levels of endogenous ethylene may reect the absence of seed-
produced auxins. Protection of the abscission zone depends on a constant
supply of auxin, and ethylene production levels in tissues correlate with
endogenous auxin levels (Roberts and Osborne, 1981).
Despite their roles in cell division, cell enlargement and maintenance of
the abscission zone in developing fruit, a specic recommendation for exog-
enous application of plant growth regulators, either alone or in combination,
to improve yield of mango has not been adopted. Phytohormones have little
residual effect on fruit development, and multiple applications of products
to counteract the short-term responses are prohibitively expensive.
Photoassimilates
Wolstenholme and Whiley (1995) discussed the ecophysiology of the mango
as a basis for preharvest management. They proposed that the adaptive sur-
vival strategies of the mango explain its notoriously poor cropping perfor-
mance. Mechanisms that impart tolerance to heat, drought and ood stresses,
which the tree has developed for survival in harsh environments, have come
at considerable carbon cost with the resultant diversion of photoassimilate
resources away from fruiting.
There is abundant evidence that heavy cropping in tree crops exhausts
stored reserves (Jones et al., 1975; Kaiser and Wolstenholme, 1994; Whiley et al.,
1996) and that current photosynthate is often unable to satisfy the demands
of fruit set and fruit growth after heavy and prolonged owering (Chacko
et al., 1982). There are signicant genotypic differences in photoassimilation
rates between low- and high-yielding cultivars growing in both the tropics
and the subtropics of Australia (Chacko et al., 1995; Searle et al., 1995). At each
location, photoassimilation rates were considerably greater on the higher-
yielding cultivar, and this difference was maintained from owering through
to fruit maturation.
14
C studies during the fruit set and abscission period also
demonstrated strong discrimination in the movement of assimilates, which
was dominated by randomly located fruit on panicles of the low-yielding
cultivar (Chacko et al., 1995). In contrast, assimilate discrimination to fruitlets
was less severe in the high-yielding cultivar with a more even distribution of
photoassimilates. It was concluded that the availability and distribution of
photoassimilates during the fruit set and establishment stages was largely
responsible for the yield differences between the cultivars.
Supporting evidence for the role of photoassimilates in fruit set and
retention also comes from enrichment studies (Schaffer et al., 1999). Container-
grown plants that owered in the open were transferred to controlled-
environment glasshouse rooms immediately after the completion of anthesis.
Temperatures were 28C day/20C night while the atmospheric carbon diox-
ide (CO
2
) concentrations were 350 or 600 mol/mol. Photoassimilation of
trees in the CO
2
-enriched rooms was approximately 60% greater than those
held at partial pressures of 350 mol/mol CO
2
. Fruit retention and nal yield
were signicantly higher on those trees grown at the partial pressure of
Reproductive Physiology 145
600 mol/mol CO
2
. Higher levels of available assimilates during the fruiting
cycle appear to benet fruit retention and yield.
5.15 Alternate Bearing
Alternate bearing of certain mango cultivars has plagued growers (Hartless,
1914; Rao, 1997). Lack of production in the off year is usually a result of lack
of oral initiation (R.N. Singh, 1959), and has been reviewed (Mukherjee,
1953; Gangolly et al., 1957; L.B. Singh, 1960, 1972; Chadha and Pal, 1986; Pan-
dey, 1989). Shoot initiation and induction described in this chapter perhaps
offers a clearer understanding of this phenomenon.
5.16 Conclusions
This chapter has provided a comprehensive review of investigations of vari-
ous factors potentially involved in mango owering, fruit set and retention.
Many of the reports cited are contradictory. Such variable results reect: (i)
the different experimental approaches utilized, especially in eld experi-
ments; (ii) the range of environments in which experiments have been con-
ducted; and (iii) differences in the responses of cultivars to treatments. As a
consequence, it is difcult to draw unambiguous conclusions with respect to
the role of specic factors on owering, fruit set and retention. More research
is clearly needed in these areas, particularly in controlled environments. For
example, although KNO
3
is utilized with great success to stimulate owering
in tropical conditions, confusion remains as to whether it effects initiation or
oral induction. Future studies on owering research should include results
of the proportion of stems that remain in rest and those that produce vegeta-
tive shoots as well as the proportion of reproductive shoots. Providing this
information allows an analysis of the impact of treatments on initiation and
inductive events.
This chapter also presents several hypothetical models for shoot devel-
opment and owering. Each of the proposed models consists of a series of
hypotheses that invite further study to test their validity. Future research
should challenge these models so that owering and crop yield can be better
understood in both the tropics and the subtropics.
References
Abe, M., Kobayashi, Y., Yamamoto, S., Daimon, Y., Yamaguchi, A., Ikeda, Y., Ichinoki, H.,
Notaguchi, M., Goto, K. and Araki, T. (2005) FD, a bZIP protein mediating signals
from the oral pathway integrator FT at the shoot apex. Science 309, 10521056.
Abeles, F.B. (1973) Ethylene in Plant Biology. Academic Press, New York, 302 pp.
Acala, P.E. and San Pedro, A. (1935) Bud differentiation in smudged mango trees. Philip-
pine Agriculture 24, 2748.
T.L. Davenport 146
Adato, A., Sharon, D., Lavi, U., Hillel, J. and Gazit, S. (1995) Application of DNA nger-
prints for identication and genetic analyses of mango (Mangifera indica) geno-
types. Journal of the American Society for Horticultural Science 120, 259264.
Agarwala, S.C., Sharma, P.N., Chatterjee, C. and Sharma, P.C. (1981) Development and
enzymatic changes during pollen development of boron decient maize. Journal of
Plant Nutrition 3, 329336.
Agrawal, A., Ram, S. and Garg, G.K. (1980) Endogenous cytokinins of mango (Mangifera
indica L.) shoot tips and their signicance in owering. Indian Journal of Experi-
mental Biology 18, 504509.
Aitken, R.L., Jeffery, A.J. and Compton, B.L. (1987) Evaluation of selected extractants for
boron in some Queensland soils. Australian Journal of Soil Research 25, 263273.
Aksenova, N.P., Milyaeva, E.L. and Romanov, G.A. (2006) Florigen goes molecular:
seventy years of the hormonal theory of owering regulation. Russian Journal of
Plant Physiology 53, 401406.
Alcala, P.E. and San Pedro, A. (1935) Bud differentiation in smudged mango trees. Phil-
ippine Agriculturist 24, 2740.
Allsopp, A. (1965) The signicance for development of water supply, osmotic relations
and nutrition. In: Lang, A. (ed.) Encyclopedia of Plant Physiology. XVII. Springer-
Verlag, Berlin, pp. 504552.
Anderson, D.L., Sedgley, M., Short, J.R.T. and Allwood, A.J. (1982) Insect pollination
of mango in northern Australia. Australian Journal of Agriculture Research 33,
541548.
Astudillo, E.O. and Bondad, N.D. (1978) Potassium nitrate-induced owering of Ca-
rabao mango shoots at different stages of maturity. Philippine Journal of Crop
Science 3, 147152.
Baghel, B.S., Sharma, R.K. and Nair, P.K.R. (1987a) Efcacy of preowering spray of urea
and NAA on physical standards of mango (Mangifera indica L.). Progressive Horti-
culture 19, 231234.
Baghel, B.S., Sharma, R.K. and Nair, P.K.R. (1987b) Inuence of preowering spray of
urea and NAA on fruit retention of mango (Mangifera indica L.). Progressive Horti-
culture 19, 200202.
Bains, K.S., Bajwa, G.S. and Singh, Z. (1999) Effects of indole-3-acetic acid, gibberellic
acid, and abscisic acid on abscission of mango fruitlets. Tropical Agriculture 76,
8892.
Bajwa, B.S., Bakshi, J.C. and Kocher, T.S. (1956) A note on the oral biology of Mangifera
indica L. var. Dashaheri. Indian Journal of Horticulture 13, 206209.
Bakr, E.I., Abdalla, K.M., Meligi, M.A. and Ismail, I.A. (1981) Floral differentiation in
mango as affected by growth regulators, ringing and defoliation. Egyptian Journal of
Horticulture 8, 161166.
Bangerth, F. (1994) Response of cytokinin concentration in the xylem exudate of bean
(Phaseolus vulgaris L.) plants to decapitation and auxin treatment, and relationship
to apical dominance. Planta 194, 439442.
Bangerth, F. (2006) Flower induction in perennial fruit trees: still an enigma? Acta Hor-
ticulturae 727, 177195.
Bangerth, F., Naphrom, D., Sruamsiri, P., Hegele, M., Boonplod, N. and Manochai, P.
(2004) Hormonal changes in various tissues of mango trees during ower induction
following cold temperature. Acta Horticulturae 645, 453457.
Barba, R.C. (1974) Induction of owering of the mango by chemical spray. Proceedings
of the Crop Science Society of the Philippines 5, 154160.
Barnell, E. (1939) Studies in tropical fruits. V. Some anatomical aspects of fruit fall in two
tropical arboreal plants. Annals of Botany 3, 7789.
Reproductive Physiology 147
Batten, D.J. and McConchie, C.A. (1995) Floral induction in growing buds of lychee
(Litchi chinensis) and mango (Mangifera indica). Australian Journal of Plant Physi-
ology 22, 783791.
Bausher, M.G. and Yelenosky, G. (1987) Morphological changes in citrus associated
with relatively high concentrations of paclobutrazol. Journal of Plant Growth Regu-
lation 5, 139147.
Belding, R.D. and Young, E. (1989) Shoot and root temperature effects on axillary cyto-
kinin levels during budbreak in young apple trees. HortScience 24, 115117.
Bernier, G. (1988) The control of oral evocation and morphogenesis. Annual Review of
Plant Physiology 39, 175219.
Bernier, G. and Sachs, R.M. (1979) Photosynthesis and owering. In: Marcelle, R.,
Clysters, H. and van Pouche, M. (eds) Photosynthesis and Plant Development. Dr. W.
Junk Publishers, The Hague, pp. 137150.
Bernier, G., Kinet, J.M. and Sachs, R.M. (1981) The Physiology of Flowering. Vols 1 and
2. CRC Press, Boca Raton, Florida.
Bernier, G., Havelange, A., Houssa, C., Petitjean, A. and Lejeune, P. (1993) Physiologi-
cal signals that induce owering. The Plant Cell 5, 11471155.
Beveridge, C.A., Weller, J.L., Singer, S.R. and Hofer, J.M.I. (2003) Axillary meristem de-
velopment. Budding relationships between networks controlling owering, branch-
ing, and photoperiod responsiveness. Plant Physiology 131, 927934.
Bevington, K.B. and Castle, W.S. (1986) Annual root growth pattern of young citrus trees
in relation to shoot growth, soil temperature, and soil water content. Journal of the
American Society for Horticultural Science 110, 840845.
Beyer, E.M., Jr and Morgan, P.W. (1971) Abscission: the role of ethylene modication of
auxin transport. Plant Physiology 48, 208212.
Bijhouwer, A.P.C. (1937) Een Bijdrage tot de Kennis Omtrent het Bloeien en het
Vruchtdragende, Vermogen van den Mangga (Mangifera indica L.). H. Veenman
and Zonen, Wageningen, the Netherlands, 106 pp.
Blaikie, S.J., Leonardi, J., Chacko, E.K., Muller, W.J. and Scott, N.S. (1999) Effect of cinc-
turing and chemical treatments on growth, owering and yield of mango. Austra-
lian Journal of Experimental Agriculture 39, 761770.
Bohlenius, H., Huang, T., Charbonnel-Campaa, L., Brunner, A.M., Jansson, S., Strauss,
S.H. and Nilsson, O. (2006) CO/FT regulatory module controls timing of owering
and seasonal growth cessation in trees. Science 312, 10401043.
Bondad, N.D. (1972) Effects of 2-chloroethane-phosphonic acid on some Philippine
horticultural crops. Philippine Journal of Geography 16, 3135.
Bondad, N.D. (1976) Response of some tropical and subtropical fruits to pre- and post-
harvest applications of ethephon. Economic Botany 30, 6780.
Bondad, N.D. and Apostol, C.J. (1979) Induction of owering and fruiting in immature
mango shoots with KNO
3
. Current Science 48, 591593.
Bondad, N.D. and Linsangan, E. (1979) Flowering in mango induced with potassium
nitrate. HortScience 14, 527528.
Bondad, N.D., Blanco, A.E. and Mercado, E.L. (1978) Foliar sprays of potassium nitrate
for ower induction in mango (Mangifera indica cultivar Pahutan) shoots. Philip-
pines Journal of Crop Science 3, 251256.
Boss, P.K., Bastow, R.M., Milne, J.S. and Dean, C. (2004) Multiple pathways in the deci-
sion to ower: enabling, promoting, and resetting. The Plant Cell 16, S18S31.
Buell, E.P. (1954) Flowering and fruiting habits of the mango in wet zone. Tropical Agri-
culturist 11, 280284.
Bueno, P.B. and Valmayor, R.V. (1974) Potassium nitrate: key to mango owering. Agri-
culture Los Banos 13, 416.
T.L. Davenport 148
Burns, W. and Prayag, S.H. (1921) The Book of the Mango. Bombay Department of
Agriculture Bulletin 103. Bombay Department of Agriculture, Bombay, 98 pp.
Burondkar, M.M. and Gunjate, R.T. (1991) Regulation of shoot growth and owering in
Alphonso mango with paclobutrazol. Acta Horticulture 291, 7984.
Burondkar, M.M. and Gunjate, R.T. (1993) Control of vegetative growth and induction
of regular and early cropping in Alphonso mango with paclobutrazol. Acta Horti-
culture 341, 206215.
Burrows, G.E., Boag, T.S. and Stewert, W.P. (1992) Changes in leaf, stem, and root anat-
omy of chrysanthemum cv. Lillian Hoek following paclobutrazol application. Jour-
nal of Plant Growth Regulation 11, 189194.
Cane, A.R. and Wilkins, M.B. (1970) Auxin transport in roots. Journal of Experimental
Botany 21(66), 212218.
Chacko, E.K. (1986) Physiology of vegetative and reproductive growth in mango (Mangifera
indica L.) trees. In: Proceedings of the First Australian Mango Research Workshop.
Commonwealth Scientic and Industrial Research Organization (CSIRO), Melbourne,
pp. 5470.
Chacko, E.K. (1991) Mango owering still an enigma. Acta Horticulture 291, 1221.
Chacko, E.K. and Ananthanarayanan, T.V. (1982) Accumulation of reserve substances
in Mangifera indica L. during ower initiation. Zeitschrift fr Panzenphysiologie
106, 281205.
Chacko, E.K. and Randhawa, G.S. (1971) Towards an understanding of the factors affect-
ing owering in mango. Andhra Agriculture Journal 18, 226236.
Chacko, E.K. and Singh, R.N. (1969a) Growth regulators induced parthenocarpy in
mango (Mangifera indica L.). Current Science 38, 249250.
Chacko, E.K. and Singh, R.N. (1969b) Induction of parthenocarpy in mango (Mangifera
indica L.) using plant growth regulators. HortScience 4, 121123.
Chacko, E.K., Singh, R.N. and Kachru, R.B. (1970a) Gibberellin-like substances in de-
veloping fruits of the mango (Mangifera indica L.). Journal of Horticultural Science
45, 371378.
Chacko, E.K., Singh, R.N. and Kachru, R.B. (1970b) Physiology of owering and fruit
growth in mango (Mangifera indica L.): partition, bioassay and characterization
of naturally occurring auxins and inhibitors in immature fruits. Indian Journal of
Experimental Biology 8, 135138.
Chacko, E.K., Kachru, R.B. and Singh, R.N. (1970c) Changes in the level of acidic and
neutral growth promoters during fruit development in Dashehari mango (Mangifera
indica L.). Journal of Horticulture Science 45, 341349.
Chacko, E.K., Kohli, R.R. and Randhawa, G.S. (1972a) Studies on the effect of 2-chloroethyl
phosphonic acid (Ethrel) on mango. I. Flower induction in an off year in Langra
trees. Indian Journal of Horticulture 29, 14.
Chacko, E.K., Singh, R.N. and Kachru, R.B. (1972b) Studies on the physiology of ower-
ing and fruit growth in mango (Mangifera indica L.). VII. Naturally occurring auxins
and inhibitors in the shoots of owering (on) and vegetative (off) mango trees. In-
dian Journal of Horticulture 29, 115125.
Chacko, E.K., Reddy, Y.T.N. and Ananthanarayanan, T.V. (1982) Studies on the relation-
ship between leaf number and area and fruit development in mango (Mangifera
indica L.). Journal of Horticultural Science 57, 483489.
Chacko, E.K., Lu, P. and Kohli, R.R. (1995) Photoassimilate production and distribution in
relation to productivity in mango (Mangifera indica L.). In: Mango 2000 Marketing
Seminar and Production Workshop Proceedings. Department of Primary Industries,
Brisbane, pp. 109124.
Chadha, K.L. and Pal, R.N. (1986) Mangifera indica. In: Halevy, A.C. (ed.) CRC Hand-
book of Flowering. Vol. 5. CRC Press, Boca Raton, Florida, pp. 211230.
Reproductive Physiology 149
Chadha, K.L. and Singh, K.K. (1963) Studies on fruit drop in mango. I. Fruit set, its reten-
tion and factors affecting it. Indian Journal of Horticulture 20, 172185.
Chaikiattiyos, S., Menzel, C.M. and Rasmussen, T.S. (1994) Floral induction in tropical
fruit trees: effects of temperature and water-supply. Journal of Horticultural Science
69, 397415.
Chailakhyan, M.K. (1936) New facts in support of the hormonal theory of plant develop-
ment. C. R. Acad. Sci. URSS 13, 7983.
Chandler, W.H. (1958) Evergreen Orchards, 2nd edn. Lea Febiger, Philadelphia, Penn-
sylvania, 535 pp.
Chen, W.S. (1979) Physiological studies of fruiting in mango trees. 1. Effect of calcium
and boron nutrition on fruiting. National Science Council Monthly (Republic of
China) 7, 12201230.
Chen, W.S. (1981) Physiological studies of fruiting in mango trees. II. Effect of endoge-
nous growth substances on fruiting. Proceedings National Science Council Part B,
Life Sciences (Republic of China) 5, 4955.
Chen, W.S. (1983) Cytokinins of the developing mango fruit. Plant Physiology 71, 356361.
Chen, W.S. (1985) Flower induction in mango (Mangifera indica L.) with plant growth
substances. Proceedings National Science Council Part B, Life Sciences (Republic
of China) 9, 912.
Chen, W.S. (1987) Endogenous growth substances in relation to shoot growth and ow-
er bud development of mango. Journal of the American Society for Horticulture
Science 112, 360363.
Cline, M., Wessel, T. and Iwamura, H. (1997) Cytokinin/auxin control of apical domi-
nance in Ipomoea nil. Plant and Cell Physiology 38, 659667.
Cobin, M. (1950) Mango selection, propagation and culture. University of Florida
Agricultural Experiment Station Annual Report. University of Florida, Gainesville,
Florida, pp. 257259.
Coen, E.S. and Carpenter, R. (1993) The metamorphosis of owers. The Plant Cell 5,
11751181.
Coen, E.S. and Meyerowitz, E.M. (1991) The war of the whorls: genetics interaction
controlling ower development. Nature 353, 3137.
Coen, E.S., Romero, J.M., Doyle, S., Elliott, R., Murphy, G. and Carpenter, R. (1990)
Floricaula: a homeotic gene required for ower development in Antirrhinum ma-
jus. Cell 63, 13111322.
Collins, G.N. (1903) The Mangoes in Puerto Rico. United States Department of Agricul-
ture (USDA) Bulletin No. 28. USDA, Washington, DC.
Corbesier, L. and Coupland, G. (2005) Photoperiodic owering of Arabidopsis: integrat-
ing genetic and physiological approaches to characterization of the oral stimulus.
Plant, Cell and Environment 28, 5466.
Corbesier, L., Vincent, C., Jang, S., Fornara, F., Fan, Q., Searle, I., Giakountis, A., Farrona,
S., Gissot, L., Turnbull, C.G.N. and Coupland, G. (2007) FT protein movement con-
tributes to long-distance signaling in oral induction of Arabidopsis. Science 316,
10301033.
Crane, J.C. (1964) Growth substances in fruit setting and development. Annual Review
of Plant Physiology 15, 303326.
Cull, B.W. (1987) A whole plant approach to productivity research for mango. In: Prense-
ley, R.T. and Tucker, G. (eds) Mangoes A Review. Commonwealth Science Coun-
cil, London, pp. 1928.
Cull, B.W. (1991) Mango crop management. Acta Horticulturae 291, 154173.
Dalziel, J. and Lawrence, D.K. (1984) Biochemical and biological effects of kaurene
oxidase inhibitors, such as paclobutrazol. British Plant Growth Regulator Group
Monograph 4, 114.
T.L. Davenport 150
Davenport, T.L. (1987) Opportunities for tropical fruit crop growth regulation. In: Pro-
ceedings of the 14th Annual Meeting of the Plant Growth Regulation Society of
America, Honolulu, Hawaii, pp. 406409.
Davenport, T.L. (1990) Citrus owering. Horticultural Reviews 12, 349408.
Davenport, T.L. (1992) Benecial effects of water stress. In: Davenport, T.L. and Har-
rington, H.M. (eds) Plant Stress in the Tropical Environment, Proceedings of a
Workshop held in Kailua-Kona, Hawaii. USDA/Co-operative States Research Ser-
vice/Caribbean Basin Administrative Group/Pacic Basin Administrative Group,
Gainesville, Florida, pp. 1620.
Davenport, T.L. (1993) Floral manipulation in mangos. In: Chia, L.E. and Evans, D.O.
(eds) Proceedings of the Conference on Mango in Hawaii. Cooperative Extension
Service, University of Hawaii, Honolulu, pp. 5460.
Davenport, T.L. (1994) Potential problems with use of uniconazole on mango (Mangifera
indica). PGRSA Quarterly (published by Plant Growth Regulation Society of Amer-
ica) 22, 143153.
Davenport, T.L. (2000) Processes inuencing oral initiation and bloom: the role of
phytohormones in a conceptual owering model. HortTechnology 10, 733739.
Davenport, T.L. (2003) Management of owering in three tropical and subtropical fruit
tree species. HortScience 38, 13311335.
Davenport, T.L. (2006) Pruning strategies to maximize tropical mango production from
the time of planting to restoration of old orchards. HortScience 41, 544548.
Davenport, T.L. and Manners, M.M. (1982) Nucellar senescence and ethylene produc-
tion as they relate to avocado fruitlet abscission. Journal of Experimental Botany 33,
815826.
Davenport, T.L. and Nez-Elisea, R. (1990) Ethylene and other endogenous factors pos-
sibly involved in mango owering. Acta Horticulturae 275, 441448.
Davenport, T.L. and Nez-Elisea, R. (1991) Is endogenous ethylene involved in ower-
ing of mango? Acta Horticulturae 291, 8594.
Davenport, T.L. and Nez-Elisea, R. (1997) Reproductive physiology. In: Litz, R.E. (ed.)
The Mango: Botany, Production and Uses. CAB International, Wallingford, UK,
pp. 69146.
Davenport, T.L. and Smith, S.M. (1997) Evaluation of potential treatments to stimulate
mango owering in sub-tropical latitudes. PGRSA Quarterly (published by Plant
Growth Regulation Society of America) 25, 114129.
Davenport, T.L. and Stern, R.A. (2005) Flowering. In: Menzel, C.M. and Waite, G.K.
(eds) Litchi and Longan: Botany, Cultivation and Uses. CAB International, Walling-
ford, UK, pp. 87113.
Davenport, T.L., Morgan, P.W. and Jordan, W.R. (1980) Reduction of auxin transport
capacity with age and internal water decits in cotton petioles. Plant Physiology
65, 10231025.
Davenport, T.L., Ramos, L. and Alani, A. (1995) Evidence for a transmissible origenic
promoter in mango. (Abstract). In: Proceedings of the 22nd Annual Meeting of the
Plant Growth Regulation Society of America, Minneapolis, Minnesota, p. 159.
Davenport, T.L., Kulkarni, V. and White, T.L. (2001a) Longevity of the origenic pro-
moter in mango. (Abstract). In: Proceedings of the 28th Annual Meeting of the Plant
Growth Regulation Society of America, Miami Beach, Florida, p. 53.
Davenport, T.L., Pearce, D.W. and Rood, S.B. (2001b) Correlation of endogenous
gibberellic acid with initiation of mango shoot growth. Journal of Plant Growth
Regulation 20, 308315.
Davenport, T.L., Ying, Z., Kulkarni, V. and White, T.L. (2006a) Evidence for a translocat-
able origenic promoter in mango. Scientia Horticulturae 110, 150159.
Reproductive Physiology 151
Davenport, T.L., Zhang, T. and Ying, Z. (2006b) Isolation of genes potentially regulating
mango owering. (Abstract). In: Proceedings of the 33rd Annual Meeting of the
Plant Growth Regulation Society of America, Quebec City, Quebec, pp. 109110.
Davies, P.J. (ed.) (1995) Plant Hormones Physiology, Biochemistry and Molecular Biol-
ogy. Kluwer Academic, Boston, 833 pp.
Degani, C., El-Batsri, R. and Gazit, S. (1990) Enzyme polymorphism in mango. Journal
of the American Society for Horticultural Science 115, 844847.
Dell, B. and Huang, L. (1997) Physiological response of plants to low boron. Plant and
Soil 193, 103120.
Desai, A.G., Limaye, V.P. and Gunjate, R.T. (1986) Floral biology of Alphonso, Goamankur
and Keser varieties of mango. Journal of Maharashtra Agriculture University 56, 3815.
De Wet, E. and Robbertse, P.J. (1986a) Pollen studies and stigma receptivity of Mangifera
indica L. cvs Haden and Sensation. South African Mango Growers Association
Yearbook 6, 2732.
De Wet, E. and Robbertse, P.J. (1986b) A preliminary study of the pollen of Mangifera
indica L. cv. Haden in South Africa. South African Journal of Plant and Soil 3, 8789.
De Wet, E., Robbertse, P.J. and Goeneveld, M.T. (1989) The inuence of temperature
and boron on pollen germination in Mangifera indica L. South African Journal of
Plant Soil 6, 228234.
Dutcher, R.D. (1972) Induction of early owering in Carabao mango in the Philippines
by smudging and ethephon application. HortScience 7, 343.
Du Toit, A.P. and Swart, D.J. (1993) Pollination of mango in the Letsitele Valley during
the 1992 owering season. First report. South African Mango Growers Association
Yearbook 13, 129130.
Du Toit, A.P. and Swart, D.J. (1994) Pollination of mango in the Letsitele Valley during
the 1993 owering season. Second report. South African Mango Growers Associa-
tion Yearbook 14, 6768.
Eardley, C.D. and Mansell, M.M. (1993) Preliminary report on the natural occurrence of
insect pollinators in a mango orchard. South African Mango Growers Association
Yearbook 13, 127128.
Eardley, C.D. and Mansell, M.W. (1994) Report on the natural occurrence of insect pol-
linators in a mango orchard. South African Mango Growers Association Yearbook
14, 6566.
Finazzo, S.F., Davenport, T.L. and Schaffer, B. (1994) Partitioning of photoassimilates in
avocado (Persea americana Mill.) during owering and fruit set. Tree Physiology
14, 153164.
Free, J.B. and Williams, I.H. (1976) Insect pollination of Anacardium occidentalis L.,
Mangifera indica L., Blighiaa sopida Koenig and Persea americana Mill. Tropical
Agriculture 53, 125139.
Galang, F.G. and Lazo, F.D. (1937) The setting of Carabao mango fruits as affected by
certain sprays. Philippine Journal of Agriculture 8, 187211.
Galn-Saco, V. (1990) Los Frutales Tropicales en los Subtropicos. I. Aguacate, Mango,
Litchi y Longan. Ediciones Mundi-Prensa, Madrid, 133 pp.
Galn-Saco, V., Fernndez, D.G. and Lpez, R.T. (1991) Effects of ethephon on mango
owering. Acta Horticulturae 291, 4350.
Galn-Saco, V., Fernndez, D.G., Hernndez, P. and Garca, V. (1993) Comparison of
manual and ethephon-induced deblossoming of mango cv. Keitt in the Canary Is-
lands. Acta Horticulturae 341, 248255.
Galila, A.S. and El-Masry, H.M. (1991) Effect of Ethrel and S-3307 D (a new plant growth
retardant) on owering, malformation, fruit set and yield of Ewais mango cultivar.
Annals of Agricultural Science (Cairo) 36, 645650.
T.L. Davenport 152
Gangolly, S.R. (1960) Some problems in mango research need an urgent solution. Pun-
jab Fruit Journal 23, 3946.
Gangolly, S.R., Singh, R., Katyal, S.L. and Singh, D. (1957) The Mango. Indian Council
of Agricultural Research (ICAR), New Delhi, 530 pp.
Gaskins, M.H. (1963) Girdling mango seedlings for inducing early fruit bearing. Pro-
ceedings of the Florida State Horticultural Society 50, 360363.
Gazit, S., Adato, I. and Roizman-Bustan, Y. (1992) Factors responsible for seasonal
changes in successful pollination rate of mango owers in Israel. (Abstract). IV In-
ternational Mango Symposium, Miami Beach, Florida, p. 36.
Goguey, T. (1990) Effets dapplications rptes de CULTAR (paclobutrazol) sur Mangifera
indica L. var. Valencia. Fruits 45, 599607.
Goguey, T. (1993) Study of the effects of three ower-inducing substances on Kent and
Zill mango. Acta Horticulturae 341, 216224.
Goldsmith, M.H.M. (1968) The transport of auxin. Annual Review of Plant Physiology
19, 347360.
Goldsmith, M.H.M. and Ray, P.M. (1973) Intracellular localization of the active process
in polar transport of auxin. Planta 111, 297314.
Gunjate, R.T., Jorwekar, D.P. and Lad, B.L. (1983) Pollination, fruit set and fruit drop in
Alphonso mango. Journal of Maharashtra Agricultural University 8, 168170.
Gonzalez, L.G. (1923) The smudging of mango trees and its effects. Philippine Agricul-
ture 12, 1527.
Haberer, G. and Kieber, J.J. (2002) Cytokinins. New insights into a classic phytohor-
mone. Plant Physiology 128, 354362.
Halevy, A.H. (ed.) (19851986) CRC Handbook of Flowering. Vols 15. CRC Press, Boca
Raton, Florida.
Halle, F., Oldeman, R.A.A. and Tomlinson, P.B. (1978) Tropical Trees and Forests: an
Architectural Analysis. Springer-Verlag, New York, 437 pp.
Harris, W. (1901) Historical notes on economic plants of Jamaica. Bulletin of the Botany
Department 8, 161178.
Hartless, A.C. (1914) Mango crops and some factors inuencing them. Agricultural Jour-
nal of India 9, 141159.
Hasdiseve, C. and Tongumpai, P. (1986) Effects of paclobutrazol on vegetative growth,
owering, and fruiting of mango Nam Doc Mai Twai #4. In: Proceedings of the
National Conference. Kasetsart University 24, Bangkok, Thailand, pp. 295302.
Hassig, B.E. (1974) Origins of adventitious roots. New Zealand Journal of Forest Science
4, 229310.
Haw, K.C. (1986) The potential use of PP 333 on fruits in Malaysia with particular refer-
ence to mangoes. In: Anon. Cultar, Tropical Crops Workshop, 2022 January 1986,
Imperial Chemical Industries (ICI), Fernhurst, Haslemere, Surrey, UK.
Hegele, M., Boonplod, N., Bangerth, F., Naphrom, D., Chattrakul, A., Sruamsiri, P. and
Manochai, P. (2004) Changes in photosynthesis, IAA export from leaves and cyto-
kinins in the xylem sap after girdling of young mango trees in combination with
different growth regulators and their possible signicance for ower induction.
Acta Horticulturae 645, 417424.
Hegele, M., Bangerth, F., Naphrom, D., Sruamsiri, P. and Manochai, P. (2006) Control of
ower induction in tropical/subtropical fruit trees by phytohormones using the ex-
ample of longan and mango. Acta Horticulturae 727, 217276.
Hendry, N.S., Van Staden, J. and Allan, P. (1982a) Cytokinins in citrus. I. Fluctuations in the
leaves during seasonal and developmental changes. Scientia Horticulturae 16, 916.
Hendry, N.S., Van Staden, J. and Allan, P. (1982b) Cytokinins in citrus. II. Fluctuations
during growth in juvenile and adult plants. Scientia Horticulturae 17, 247256.
Reproductive Physiology 153
Henny, R.J. (1995) Thidiazuron increases basal bud and shoot development in Spathip-
hyllum Petite. Plant Growth Regulation Society of America Quarterly 23, 1316.
Hillier, G.R. and Rudge, T.G. (1991) Promotion of regular fruit cropping in mango with
Cultar. Acta Horticulturae 291, 5159.
Hongsbhanich, N. (1986) Effects of paclobutrazol on vegetative growth, owering and
fruit setting in mango Nam Dok Mai Twai #4. In: Anon. Cultar, Tropical Crops
Workshop, January 2022, 1986, Imperial Chemical Industries (ICI), Fernhurst,
Haslemere, Surrey, UK.
Hu, H. and Brown, P.H. (1994) Localization of boron in cell walls of squash and to-
bacco and its association with pectin. Plant Physiology 105, 681689.
Huala, E. and Sussex, I.M. (1993) Determination and cell interactions in reproductive
meristems. The Plant Cell 5, 11571165.
Huang, T., Bohlenius, H., Erikson, S., Parcy, F. and Nilsson, O. (2005) The mRNA of the
Arabidopsis gene FT moves from leaf to shoot apex and induces owering. Science
309, 16941696.
Hussein, M.A., Mahmoud, H.M., Ahmed Amen, K.I. and Abo-El-Ez, A.T. (1989) Com-
parative studies on sex distribution of some mango varieties Mangifera indica L.
under Assuit conditions. Assuit Journal of Agricultural Science 20, 7982.
Issarakraisila, M. and Considine, J.A. (1994) Effects of temperature on microsporogene-
sis and pollen viability in mango cv. Kensington. Annals of Botany 73, 231234.
Issarakraisila, M., Considine, J.A. and Turner, D.W. (1992) Seasonal effects on oral biol-
ogy and fruit set of mangoes in a warm temperature region of Western Australia.
Acta Horticulturae 321, 626.
Itai, C. and Vaadia, Y. (1965) Kinetin-like activity in root exudate of water stressed sun-
ower plants. Physiologiea Plantarum 18, 941944.
Itai, C., Richmond, A. and Vaadia, Y. (1968) The role of root cytokinins during water and
salinity stress. Israel Journal of Botany 17, 187195.
Itai, C., Ben-Zioni, A. and Ordin, L. (1973) Correlative changes in endogenous hormone
levels and shoot growth induced by short heat treatments to the root. Physiologie
Plantarum 29, 355360.
Jacobs, W.P. and Case, D.B. (1965) Auxin transport, gibberellin, and apical dominance.
Science 148, 17291731.
Jiron, L.F. and Hedstrom, I. (1985) Pollination ecology of mango (Mangifera indica L.)
(Anacardiaceae) in the neotropic region. Turrialba 35, 269277.
Jivanna Rao, P.S. (1923) Pollen sterility in relation to vegetative propagation. Journal of
Madras Agricultural Student University 11, 419426.
Joel, D.M. and Eisenstein, D. (1980) A bridge between the ovule and ovary wall in
Mangifera indica L. (Anacardiaceae). Acta Botanica Neerlandica 29, 203206.
Jones, W.W., Embleton, T.W. and Coggins, C.W., Jr (1975) Starch content of roots of
Kinnow mandarin trees bearing fruits in alternate years. HortScience 10, 514.
Joubert, J.P., Robbertse, P.J., Coetzer, L.A. and Wishart, D.L. (1993) Inorescence char-
acteristics and ower sex ratio studies of container-grown mango trees. South African
Mango Growers Association Yearbook 13, 2733.
Juliano, J.B. and Cuevas, N.L. (1932) Floral morphology of the mango (Mangifera indica
Linn.) with special reference to the Pico variety from the Philippines. Philippine
Agriculturist 21, 449472.
Jutamanee, K., Eoomkham, S., Pichakum, A., Krisanapook, K. and Phavaphutanon, L.
(2002) Effects of calcium, boron and sorbitol on pollination and fruit set in mango
cv. Namdokmai. Acta Horticulturae 575, 829834.
Kachru, R.B., Singh, R.N. and Chacko, E.K. (1971) Inhibition of owering in mango
(Mangifera indica L.) by gibberellic acid. HortScience 6, 140141.
T.L. Davenport 154
Kachru, R.B., Singh, R.N. and Chacko, E.K. (1972) Inhibition of owering in Mangifera
indica L. by gibberellic acid. Acta Horticulturae 24, 206209.
Kaiser, C. and Wolstenholme, B.N. (1994) Aspects of delayed harvest of Hass avocado
(Persea americana Mill.) fruit in a cool subtropical climate. II. Fruit size, yield, phe-
nology and whole-tree starch cycling. Journal of Horticultural Science 69, 447457.
Kende, H. and Sitton, D. (1967) The physiological signicance of kinetin and gibberellin
like root hormones. Annals of New York Academy of Sciences 144, 235243.
Khan, M.A., Malik, A.B., Makhdoom, M.L. and Haq, A. (1993) Investigations on the ef-
ciency of exogenous synthetic growth regulators on fruit drop in mango (Mangifera
indica Linn.). Egyptian Journal of Horticulture 20, 114.
Kinet, J.M. (1993) Environmental, chemical, and genetic control of owering. Horticul-
tural Reviews 15, 279334.
King, R.W. and Zeevaart, J.A.D. (1973) Floral stimulus movement in Perilla and ower
inhibition by noninduced leaves. Plant Physiology 51, 727738.
Kinman, C.F. (1918) The Mango in Puerto Rico. Agricultural Experiment Station Bulletin
No. 24. Agricultural Experiment Station, Puerto Rico, 30 pp.
Komeda, Y. (2004) Genetic regulation of time to ower in Arabidopsis thaliana. Annual
Review of Plant Biology 55, 521535.
Kozlowski, T.T., Kramer, P.J. and Pallardy, S.G. (1991) The Physiological Ecology of
Woody Plants. Academic Press, New York, 657 pp.
Kraus, E.J. and Kraybill, H.R. (1918) Vegetation and Reproduction with Reference to the
Tomato. Bulletin of the Oregon Agricultural Experiment Station Number 149. Ore-
gon State University, Corvallis, Oregon.
Krishnamurthi, S., Randhawa, G.S. and Sivaraman Nair, P.C. (1960) Growth studies in
the mango (Mangifera indica L.) under Delhi (subtropical) conditions. Indian Journal
of Horticulture 17, 107118.
Kulkarni, V.J. (1986) Graft-induced off-season owering and fruiting in the mango
(Mangifera indica L.). Journal of Horticultural Science 61, 141145.
Kulkarni, V.J. (1988a) Chemical control of tree vigour and the promotion of owering
and fruiting in mango (Mangifera indica L.) using paclobutrazol. Journal of Horti-
cultural Science 63, 557566.
Kulkarni, V.J. (1988b) Further studies on graft-induced off-season owering and fruiting
in mango (Mangifera indica L.). Journal of Horticultural Science 63, 361367.
Kulkarni, V.J. (1991) Physiology of owering in mango studied by grafting. Acta Horti-
culturae 291, 95104.
Kulkarni, V.J. (2004) The tri-factor hypothesis of owering in mango. Acta Horticulturae
645, 6170.
Kulkarni, V.J. and Rameshwar, A. (1978) Natural and gibberellic acid induced partheno-
carpy in mango. Current Science 47, 353355.
Kulkarni, V.J. and Rameshwar, A. (1989) Effect of deblossoming and defruiting on off-
season owering and fruiting in mango (Mangifera indica L.). Scientia Horticulturae
39, 143148.
Kurian, R.M. and Iyer, C.P.A. (1993a) Chemical regulation of tree size in mango
(Mangifera indica L.) cv. Alphonso. II. Effects of growth retardants on owering and
fruit set. Journal of Horticultural Science 68, 355360.
Kurian, R.M. and Iyer, C.P.A. (1993b) Chemical regulation of tree size in mango
(Mangifera indica L.) cv. Alphonso. III. Effects of growth retardants on yield and
quality of fruits. Journal of Horticultural Science 68, 361364.
Kurian, R.M., Murti, G.S.R. and Iyer, C.P.A. (1992) Changes in cytokinin levels in mango
(Mangifera indica. L.) leaf extracts following soil drenches with paclobutrazol. Gar-
tenbauwissenschaft 57, 8486.
Reproductive Physiology 155
Kurian, R.M., Iyer, C.P.A. and Murti, G.S.R. (1994) Total phenols of stem apical bud in
relation to tree vigour in mango (Mangifera indica L.). Gartenbauwissenschaft 59,
268270.
Lakshminarayana, S. and Aguilar, P.H. (1975) Effect of orchard heating in reducing par-
thenocarpic fruits in Haden mango. Proceedings of the Horticultural Society 88,
502505.
Lang, A. (1965) Physiology of ower initiation. In: Ruhland, W. (ed.) Handbuch der
Panzenphysiologie XV/1. Springer Verlag, Berlin, pp. 13801536.
Lang, A. (1984) Die photoperiodische regulation von forderung und hemmung der
blutenbildung. Bericht Deutsch Botanik Gesellschaft 97, 293314.
Lang, A., Chailakhyan, M.K. and Frolova, I.A. (1977) Promotion and inhibition of ower
formation in a day neutral plant in grafts with a short-day plant and a long-day
plant. Proceedings of the National Academy of Sciences USA 74, 24122416.
Law, D.M. (1987) Gibberellin-enhanced indole-3-acetic acid biosynthesis: D-tryptophan
as the precursor of indole-3-acetic acid. Physiologia Plantarum 70, 626632.
Law, D.M. and Hamilton, R.H. (1989) Reduction in the free indole-3-acetic acid levels
in Alaska pea by the gibberellin biosynthsis inhibitor, uniconazol. Physiologia Plan-
tarum 76, 535538.
Lomax, T.L., Muday, G.K. and Rubery, P.H. (1995) Auxin transport. In: Davies, P.J. (ed.)
Plant Hormones. Kluwer Academic Publishers, Dordrecht, pp. 509530.
Lopez, M.R., Mosqueda-Vzquez, R. and Pimienta, B.E. (1984) El nitrato de potasio
como promotor de etileno endogeno y la induccion oral del mango (Mangifera
indica) cv. Manila. Resumes. X Congreso Nacional de Fitogenetica Instituto. Tecno-
logico Agropecuario No. 20 de Aguascalientes, p. 110.
Lumsden, P.J. (1993) Mechanisms of signal transduction. In: Jorden, B.R. (ed.) The Mo-
lecular Biology of Flowering. CAB International, Wallingford, UK, pp. 2145.
Lynch, S.J. and Mustard, M.J. (1950) Mangoes in Florida. Florida Department of Agricul-
ture, Florida, 82 pp.
MacMillan, H.E. (1991) Tropical Planting and Gardening, 6th edn. McMillan, London.
767 pp.
Maiti, S.K. (1971) Effect of photoperiod on growth and owering of mango (Mangifera
indica L.). Indian Agriculture 15, 213216.
Maiti, S.C. (1973) Effect of gibberellic acid (GA
3
) sprays on sex expression of mango
(Mangifera indica L.). Science and Culture 39, 150151.
Maiti, S.C. and Sen, P.K. (1978) Studies on photoperiodic response of biennial and regu-
lar bearing mangoes. Udyanika 2, 1116.
Maiti, S.C., Basu, R.N. and Sen, P.K. (1972) Chemical control of growth and owering
in Mangifera indica L. Acta Horticulturae 24, 192195.
Maiti, S.K., Maiti, S.C. and Sen, P.K. (1978) Effects of short day with and without ringing on
growth and owering of mango (Mangifera indica L.). Indian Agriculture 22, 159164.
Majumder, P.K. and Mukherjee, S.K. (1961) Studies on the variability of sex-expression
in mango (Mangifera indica L.). Indian Journal of Horticulture 18, 1219.
Mallik, P.C. (1951) Inducing owering in mango by ringing the bark. Indian Journal of
Horticulture 8, 110.
Mallik, P.C. (1957) Morphology and biology of the mango ower. Indian Journal of Hor-
ticulture 14, 123.
Mallik, P.C., Sahay, R.K. and Singh, D.L. (1959) Effect of hormones on the sex ratio in the
mango. Current Science 28, 410.
McGregor, S.E. (1974) Insect pollination of tropical crops. In: Compte-Rendue du III
Symposium International sur la Pollinisation. International Bee Research Associa-
tion, Prague, pp. 4755.
T.L. Davenport 156
McGregor, S.E. (1976) Flowering and fruiting of plants. Pollinating agents and their com-
parative value. Avocado. In: Insect Pollination of Cultivated Crop Plants. Agriculture
Handbook No. 496. United States Department of Agriculture (USDA), Washington,
DC, pp. 715, 1923, 9398.
Menzel, C.M. (1983) The control of oral initiation in lychee: a review. Scientia Horti-
culturae 21, 201215.
Menzel, C.M. and Simpson, D.R. (1994) Lychee. In: Schaffer, B. and Anderson, P.C. (eds)
Handbook of Environmental Physiology of Fruit Crops. Vol. II, Sub-Tropical and
Tropical Crops. CRC Press, Boca Raton, Florida, pp. 123145.
Menzel, C.M., Watson, B.J. and Simpson, D.R. (1990) Longan. In: Bose, T.K. and Mitra,
S.K. (eds) Fruits: Tropical and Subtropical. Naya Prokash, Calcutta, pp. 522546.
Metzger, J.D. (1988) Localization of the site of perception of thermoinductive tempera-
tures in Thlaspi arvense. Plant Physiology 88, 424428.
Miller, C.O. (1963) Kinin and kinin-like compounds. In: Linskens, H.F. and Tracey, M.V.
(eds) Modern Methods of Plant Analysis. Vol. 6. Springer, Berlin, pp. 194202.
Mishra, K.A. and Dhillon, B.S. (1978) Carbohydrates and mineral composition of leaves
in relation to fruit-bud differentiation in Langra mango. Indian Journal of Agricul-
tural Sciences 47, 4650.
Monselise, S.P. and Goldschmidt, E.E. (1982) Alternate bearing in fruit trees. Horticul-
tural Reviews 4, 128173.
Morgan, P.W. and Gausman, H.W. (1966) Effects of ethylene on auxin transport. Plant
Physiology 41, 4552.
Morgan, P.W., Jordan, W.R., Davenport, T.L. and Durham, J.L. (1977) Abscission re-
sponses to moisture stress, auxin transport inhibitors, and ethephon. Plant Physiol-
ogy 59, 710712.
Morris, D.A. and Thomas, T. (1978) A microautoradiographic study of auxin transport in
the stem of intact pea seedlings (Pisum sativum L.). Journal of Experimental Biology
29, 147157.
Morton, J.F. (1964) Honeybee plants of south Florida. Proceedings of the Florida State
Horticultural Society 77, 415436.
Mosqueda-Vzquez, R. and de los Santos de la Rosa, F. (1981) Aspersiones de nitrato de
potasio para adelantar e inducir la oracion del mango cv. Manila en Mexico. Pro-
ceedings of the Tropical Region of the American Society for Horticultural Science
25, 311316.
Mosqueda-Vzquez, R. and Avila-Resendiz, C. (1985) Induccion oral del mango con
aplicaciones de KNO
3
y su inhibicion al aplicar AgNO
3
o CoCl
2
. Horticultura Mex-
icana 1, 93101.
Mouradov, A., Cremer, F. and Coupland, G. (2002) Control of owering time: interacting
pathways as a basis for diversity. The Plant Cell 14, S111S130.
Mukherjee, S.K. (1949a) The mango and its relatives. Science and Culture 15, 59.
Mukherjee, S.K. (1949b) A monograph on the genus Mangifera L. Lloydia 12, 73136.
Mukherjee, S.K. (1950) Cytological investigations of the mango (Mangifera indica L.)
and the allied Indian species. Proceedings of the National Institute of Science in
India 16, 4.
Mukherjee, S.K. (1951) Pollen analysis in Mangifera in relation to fruit-set and taxono-
my. Journal of the Indian Botanical Society 30, 4955.
Mukherjee, S.K. (1953) The mango its botany, cultivation, uses and future improve-
ment, especially as observed in India. Economic Botany 7, 130162.
Mukherjee, S.K., Majumdar, P.K. and Chatterjee, S.S. (1961) An improved technique of
mango hybridization. Indian Journal of Horticulture 18, 302304.
Mukhopadhyaya, A.K. (1978) A note on the effect of growth retardants and L-methionine
on owering of mango (Mangifera indica L.). Horticulture Abstracts 48, 254.
Reproductive Physiology 157
Mullins, P.D.F. (1986) Effect of varying climatic regimes on ower behavior and pollina-
tion in Sensation and Haden mango trees. South African Mango Growers Asso-
ciation Research Report 6, 342.
Musahib-ud-din (1946) Fruit bud differentiation in mangoes in the Punjab. Punjab Fruit
Journal 10, 3031.
Musahib-ud-din and Dinsa, H.S. (1946) The oral count and fruit set studies in some of
the North Indian mango varieties. Punjab Fruit Journal 10, 3542.
Mustard, M.J. and Lynch, S.J. (1946) Flower-bud formation and development in Mangifera
indica. Botanical Gazette 108, 136140.
Naik, K.C. and Mohan Rao, M. (1942) Some factors governing fruit bud formation in
mangoes (Mangifera indica L.). Madras Agriculture Journal 30, 328335.
Naik, K.C. and Mohan Rao, M. (1943) Studies on blossom biology and pollination in
mangoes (Mangifera indica L.). Indian Journal of Horticulture 1, 107119.
Nakasone, H.Y., Bowers, F.A.I. and Beaumont, J.H. (1955) Terminal growth and ower-
ing behavior of the Pirie mango (Mangifera indica L.) in Hawaii. Proceedings of the
American Society for Horticultural Science 66, 183191.
Naqvi, S.S.M., Alam, S.M. and Mumtaz, S. (1990) Effect of cobalt and silver ions and
naphthaleneacetic acid on fruit retention in mango (Mangifera indica L.). Austra-
lian Journal of Experimental Agriculture 30, 433435.
Naqvi, S.S.M., Alam, S.M. and Mumtaz, S. (1992) Inuence of growth regulators, cobalt,
and silver ions on fruit drop in mango (Mangifera indica L.). Pakistan Journal of
Horticulture 24, 197200.
Nartvaranant, P., Subhadrabandhu, S. and Tongumpai, P. (2000) Practical aspect in pro-
ducing off-season mango in Thailand. Acta Horticulturae 509, 661668.
Nickell, L.G. (ed.) (1983) Plant Growth Regulating Chemicals. Vols 1 and 2. CRC Press,
Boca Raton, Florida.
Nitsch, J.P. (1965) Physiology of ower and fruit development. In: Ruhland, W. (ed.)
Handbuch der Panzenphysiologie XV/1. Springer Verlag, Berlin, pp. 15371646.
Nordstrom, A., Tarkowski, P., Tarkowska, D., Norbaek, R., Astot, C., Dolezal, K. and
Sandberg, G. (2004) Auxin regulation of cytokinin biosynthesis in Arabidopsis thal-
iana: a factor of potential importance for auxin-cytokinin-regulated development.
Proceedings of the National Academy of Science USA 101, 80398044.
Nez-Elisea, R. (1985) Flowering and fruit set of a monoembryonic and polyembry-
onic mango as inuenced by potassium nitrate sprays and shoot decapitation. Pro-
ceedings of the Florida State Horticultural Society 98, 179183.
Nez-Elisea, R. (1986) Produccin Temprana de Mango Haden y Manila con Asper-
siones de Nitrato de Potasio. Folleto para productores No. 8. Centro de Investiga-
ciones Agrcolas Pacco Centro (CIAPAC), Instituto Nacional de Investigaciones
Forestales, Agrcolas y Pecuarias (INIFAP), Secretara de Agricultura y Recursos
Hidrulicos (SARH), Campo Agrcola Experimental Tecomn, Tecomn, Mexico.
Nez-Elisea, R. (1988) Nitrato de Amonio: Nueva Alternativa para Adelantar la Flo-
racin y Cosecha del Mango. Desplegable para productores No. 4. Secretara de
Agricultura y Recursos Hidrulicos (SARH)-Instituto Nacional de Investigaciones
Forestales, Agrcolas y Pecuarias (INIFAP), Campo Experimental Tecomn, Tecomn,
Colima, Mexico.
Nez-Elisea, R. (1994) Environmental, developmental, and bioregulator control of
owering in mango. PhD thesis, University of Florida, Gainesville, Florida, USA.
Nez-Elisea, R. and Caldeira, M.L. (1988) Induction of owering in mango (Mangifera
indica L.) with ammonium nitrate sprays. HortScience 23, 833.
Nez-Elisea, R. and Davenport, T.L. (1983) Abscission and ethylene production of
mango (Mangifera indica L.) fruit cv Tommy Atkins. Proceedings of the Florida State
Horticultural Society 96, 185188.
T.L. Davenport 158
Nez-Elisea, R. and Davenport, T.L. (1984) Ethylene production by mango fruitlets and
its role in fruit abscission. (Abstract). In: Proceedings of the 11th Annual Meeting of
the Plant Growth Regulation Society of America, Boston, Massachusetts, p. 156.
Nez-Elisea, R. and Davenport, T.L. (1986) Abscission of mango fruitlets as inuenced
by enhanced ethylene biosynthesis. Plant Physiology 82, 991994.
Nez-Elisea, R. and Davenport, T.L. (1989) Expression of an endogenous owering
promoter in mango (Mangifera indica). (Abstract). In: Proceedings of the 16th An-
nual Meeting of the Plant Growth Regulation Society of America, Arlington, Vir-
ginia, pp. 245247.
NezElisea, R. and Davenport, T.L. (1991a) Flowering of Keitt mango in response to
deblossoming and gibberellic acid. Proceedings of the Florida State Horticultural
Society 104, 4143.
Nez-Elisea, R. and Davenport, T.L. (1991b) Effect of duration of low temperature treat-
ment on owering of containerized Tommy Atkins mango. (Abstract). In: Proceed-
ings of the 18th Annual Meeting of the Plant Growth Regulation Society of America,
Boston, Massachusetts, pp. 3941.
Nez-Elisea, R. and Davenport, T.L. (1992a) Inuence of water decit on vegetative
growth and owering of containerized mango. In: Davenport, T.L. and Harrington,
H.M. (eds) Plant Stress in the Tropical Environment, Proceedings of a Workshop
held in Kailua-Kona, Hawaii. USDA/Co-operative States Research Service/Carib-
bean Basin Administrative Group/Pacic Basin Administrative Group, Gainesville,
Florida, pp. 2425.
Nez-Elisea, R. and Davenport, T.L. (1992b) Requirement for mature leaves during
oral induction and oral transition in developing shoots of mango. Acta Horticul-
turae 296, 3337.
Nez-Elisea, R. and Davenport, T.L. (1994a) Effects of exogenous GA
3
and temperature on
mango inorescence formation. (Abstract). In: Proceedings of the 21st Annual Meeting
of the Plant Growth Regulation Society of America, Portland, Oregon, pp. 6061.
Nez-Elisea, R. and Davenport, T.L. (1994b) Flowering of mango trees in containers as
inuenced by seasonal temperature and water stress. Scientia Horticulturae 58,
5766.
Nez-Elisea, R. and Davenport, T.L. (1995) Effect of leaf age, duration of cool tempera-
ture treatment, and photoperiod on bud dormancy release and oral initiation in
mango. Scientia Horticulturae 62, 6373.
Nez-Elisea, R. and Davenport, T.L. (1998) Gibberellin and temperature effects on
dormancy release and shoot morphogenesis of mango (Mangifera indica L.). Scien-
tia Horticulturae 77, 1121.
Nez-Elisea, R., Caldeira, M.L. and Davenport, T.L. (1990) Thidiazuron effects on growth
initiation and expression in mango (Mangifera indica L.). HortScience 25, 1167.
Nez-Elisea, R., Davenport, T.L. and Caldeira, M.L. (1991) An experimental system to
study mango owering using containerized trees propagated by air-layering. Pro-
ceedings of the Florida State Horticultural Society 104, 3941.
Nez-Elisea, R., Caldeira, M.L., Ferreira, W. and Davenport, T.L. (1992) Adventitious
rooting of Tommy Atkins mango air layers induced with naphthaleneacetic acid.
HortScience 27, 926.
Nez-Elisea, R., Davenport, T.L. and Caldeira, M.L. (1993) Bud initiation and morpho-
genesis in Tommy Atkins mango as affected by temperature and triazole growth
retardants. Acta Horticulturae 341, 192198.
Nez-Elisea, R., Davenport, T.L. and Caldeira, M.L. (1996) Control of bud morphogen-
esis in mango (Mangifera indica L.) by girdling, defoliation and temperature modi-
cation. Journal of Horticultural Science 71, 2540.
Reproductive Physiology 159
Ogawa, Y. (1963) Studies on the conditions for gibberellin assay using rice seedling.
Plant and Cell Physiology 4, 227237.
Ona, M.L.D. and de Guzman, E.V. (1982) Floral gross morphology and histology of
Carabao mango (Mangifera indica L.) treated with a chemical owering inducer.
Philippine Agriculturist 65, 201203.
Oosthuyse, S.A. (1991a) Stages of development of the mango panicle. South African
Mango Growers Association Yearbook 11, 5961.
Oosthuyse, S.A. (1991b) Effect of Promalin on owering and lateral branching of young man-
go trees in the nursery. South African Mango Growers Association Yearbook 11, 54.
Oosthuyse, S.A. (1995a) Effect of aqueous application of GA
3
on owering of mango
trees: why in certain instances is owering prevented, and in others owering is
only delayed? In: Mango 2000 Marketing Seminar and Production Workshop
Proceedings. Department of Primary Industries, Brisbane, pp. 7585.
Oosthuyse, S.A. (1995b) Effect of post-bloom aqueous spray application of GA
3
, NAA,
and CPPU on fruit retention fruit size, and yield in Tommy Atkins and Heidi
mango. In: Mango 2000 Marketing Seminar and Production Workshop Proceed-
ings. Department of Primary Industries, Brisbane, pp. 105109.
Oslund, C.R. and Davenport, T.L. (1987) Seasonal enhancement of ower development
in Tahiti limes by marcottage. HortScience 22, 498501.
Ou, S.K. (1980) Temperature effect on owering and fruiting in the Irwin mango
(Mangifera indica L.). Journal of Agricultural Research in China 29, 301308.
Ou, S.K. (1982) Temperature effect on differential shoot development of mango during
owering period. Journal of Agricultural Research in China 31, 209212.
Ou, S.K. and Yen, C.R. (1985) Effect of growth regulators on the owering of Irwin
mango at high temperature. In: Forcing Culture of Fruit Trees. Taichung District
Agricultural Improvement Station, Taichung, Taiwan, pp. 137143.
Pal, R.N. and Chadha, K.L. (1982) Deblossoming mangoes with cycloheximide. Journal
of Horticultural Science 57, 331332.
Pal, S. and Ram, S. (1978) Endogenous gibberellins of mango shoot-tips and their sig-
nicance in owering. Scientia Horticulturae 9, 369379.
Pandey, R.M. (1989) Physiology of owering in mango. Acta Horticulturae 231, 361380.
Pandey, R.M. and Narwadkar, P.R. (1984) Studies on the induction of growth and ower-
ing in Dashehari mango. Indian Journal of Horticulture 41, 171176.
Pandey, R.M., Singh, R.N. and Sinha, G.C. (1973) Usefulness of ethrel in regulating
ower bearing in mango. Science and Culture 39, 148150.
Pantastico, E.B. and Manuel, F.C. (1978) The Philippines Recommends for Mango 1979.
Report for Philippines Council for Agriculture and Resources Research, the Philip-
pines, p. 43.
Parisot, E. (1988) Study of the growth rhythm in young mango plants (Mangifera indica
L.). III. Growth and development of young mango plants. Fruits 43, 235247.
Paulas, D. and Shanmugavelu, K.G. (1989) Physiological and biochemical changes in
the leaf tissues from quiescent to fruiting stages of mango. Acta Horticulturae 231,
394398.
Pearce, D.W., Koskioka, M. and Pharis, R.P. (1994) Review chromatography of gib-
berellins. Journal of Chromatography A 658, 91122.
Peng, L.Z., Tang, J., Xu, E., Song, N.J. and Lei, T. (1991) Effect of paclobutrazol on the
morphological and anatomical characteristics of the new organs of citrus trees.
China Citrus 20, 710.
Perilleux, C. and Bernier, G. (2002) The control of owering: do genetical and physio-
logical approaches converge? In: ONeill, S.D. and Roberts, J.A. (eds) Plant Repro-
duction, Annual Plant Reviews. Shefeld Academic Press, Shefeld, pp. 132.
T.L. Davenport 160
Perilleux, C., Bernier, G. and Kinet, J.M. (1994) Circadian rhythms and the induction of
owering in the long-day grass Lolium temulentum L. Plant Cell Environment 17,
755761.
Pharis, R.P. and King, R.W. (1985) Gibberellins and reproductive development in seed
plants. Annual Review of Plant Physiology 36, 517568.
Pharis, R.P., Kuo, C.C. and Glenn, J.L. (1972) Gibberellin, a primary determinant in the
expression of apical dominance, apical control and geotropic movement of conifer
shoots. In: Carr, D.J. (ed.) Plant Growth Substances, 1970. Springer-Verlag, New
York, pp. 441448.
Pimentel, R.B., Coronel, R.E. and Espino, R.F.C. (1984) Floral biology and fruit set in
mango (Mangifera indica L.) cultivars: Carabao, Pico and Kancha Mitha. Philip-
pine Journal of Crop Science 9, 4751.
Ploetz, R.C., Ramos, J.L., Parrado, E.S. and Shepard, E.S. (1991) Shoot and root growth
cycles of avocado in South Florida. Proceedings of the Florida State Horticultural
Society 104, 2124.
Ploetz, R.C., Ramos, J.L. and Parrado, J.L. (1993) Periodicity of shoot and root growth in
grafted avocado. Tropical Agriculture 70, 248251.
Pongsomboon, W. (1991) Effects of temperature and water stress on tree growth, ower-
ing, fruit growth and retention of mango (Mangifera indica L.). PhD thesis, Kasetsart
University, Bangkok, Thailand.
Pongsomboon, W., Stephenson, R.A., Whiley, A.W. and Subhadrabandu, S. (1991)
Development of water stress in juvenile mango (Mangifera indica L.) trees. Acta
Horticulturae 321, 496503.
Popenoe, W. (1917) The Pollination of the Mango. US Department of Agriculture (USDA)
Bulletin 542. USDA, Washington DC.
Popenoe, W. (1920) Manual of Tropical and Sub-tropical Fruits. Macmillan, New York,
474 pp.
Prakash, S. and Ram, S. (1984) Naturally occurring auxins and inhibitor and their role in
fruit growth and drop of mango Dashehari. Scientia Horticulturae 2, 241248.
Prakash, S. and Ram, S. (1986) Effect of various concentrations of auxins and their time
of application on fruit retention in mango (Mangifera indica L.) cv. Chausa. Proges-
sive Horticulture 18, 167174.
Protacio, C.M. (2000) A model for potassium nitrate-induced owering in mango. Acta
Horticulturae 509, 545552.
Putterill, J., Laurie, R. and Macknight, R. (2004) Its time to ower: the genetic control of
owering time. Bioessays 26, 363373.
Rademacher, W. (1991) Biochemical effects of plant growth retardants. In: Gausman,
H.W. (ed.) Plant Growth Regulators. Marcel Dekker, New York, pp. 169200.
Rademacher, W. (2000a) Growth retardants: effects on gibberellin biosynthesis and oth-
er metabolic pathways. Annual Review of Plant Physiology and Plant Molecular
Biology 51, 501531.
Rademacher, W. (2000b) Growth retardants in agriculture and horticulture. In: Proceed-
ings of the 27th Annual Meeting of the Plant Growth Regulation Society of America,
Kailua-Kona, Hawaii, pp. 8792.
Raja, M.E., Kumar, S.C.A. and Raju, S.Y. (2005) Boron deciency in mango (Mangifera
indica L.): a cause delineation study in acidic soils of Maharashtra, India. Soil Science
and Plant Nutrition 51, 751754.
Rajan, S. and Ram, S. (1989) Studies on the root regeneration in mango air-layers. Acta
Horticulturae 231, 192197.
Rajput, C.B.S. and Singh, J.N. (1989) Effects of urea and GA
3
sprays on the growth, ow-
ering and fruiting characters of mango. Acta Horticulturae 231, 301305.
Reproductive Physiology 161
Ram, S. (1983) Hormonal control of fruit growth and fruit drop in mango cv Dashehari.
Acta Horticulturae 134, 169178.
Ram, S. and Pal, S. (1979) Studies on the naturally occurring gibberellins in mango
(Mangifera indica L.) fruit. Journal of Horticultural Science 54, 209215.
Ram, S., Bist, L.D., Lakhanpal, S.C. and Jamwal, I.S. (1976) Search of suitable pollinizers
for mango cultivars. Acta Horticulturae 57, 253263.
Ram, S., Sirohi, S.C. and Rathore, V.S. (1983) Naturally occurring cytokinins in mango
(Mangifera indica L.) fruit. Australian Journal of Plant Physiology 10, 6573.
Rameshwar, A. (1989) Mango owering stress induced. Acta Horticulturae 231,
433439.
Ramina, A., Hackett, W.P. and Sachs, R.M. (1979) Flowering in Bougainvillea function
of assimilate supply and nutrient diversion. Plant Physiology 64, 810813.
Ramina, A., Masia, A. and Vizzoto, G. (1986) Ethylene and auxin transport and metabo-
lism in peach fruit abscission. Journal of the American Society for Horticultural
Science 111, 760764.
Randhawa, G.S. and Damodaran, V.K. (1961a) Studies on oral biology and sex-ratio in
mango (Mangifera indica L.) I. A review. Indian Journal of Horticulture 18, 935.
Randhawa, G.S. and Damodaran, V.K. (1961b) Studies on oral biology and sex ratio in
mango (Mangifera indica L.) var. Chausa, Dashehari and Krishanbhog. III. Anthesis,
dehiscence, receptivity of stigma, pollination, fruit set and fruit development. Indi-
an Journal of Horticulture 18, 5164.
Rao, M.M. (1997) Towards Making Mango a Regular Bearer. University of Agricultural
Sciences Technology Series No. 5, Dharwad, India, 94 pp.
Rao, M.M. and Srihari, D. (1998) Approaches for managing the problems of biennial bearing
in Alphonso mango trees. Journal of Maharashtra Agricultural University 23, 1921.
Rao, M.M., Srihari, D. and Patil, V.S. (1997) Further studies on chemical induction of
owering directly on fruited shoots in off phase Alphonso mango trees. Karnataka
Journal of Agricultural Science 10, 598601.
Rath, S. and Das, G.C. (1977) Effect of ringing, notching and defoliation in mango
shoots of Langra on ower induction. Orissa Journal of Horticulture 5, 1.
Rath, S. and Das, G.C. (1979) Effect of ringing and growth retardants on growth and
owering of mango. Scientia Horticulturae 10, 101104.
Rath, S., Das, G.C. and Singh, R.L. (1982) Manipulation of owering in mango by forc-
ing the dormant buds. Bangladesh Horticulture 10, 3941.
Ravishankar, H. and Mohan Rao, M. (1982) Changes in carbohydrate fractions and min-
erals in cultivar Alphonso mango (Mangifera indica L.) shoots. Journal of Maharash-
tra Agricultural University 7, 143145.
Ravishankar, H., Rao, M.M. and Bojappa, K.M. (1979) Fruit-bud differentiation in mango
Alphonso and Totapuri under mild tropical rainy conditions. Scientia Horticulturae
10, 9599.
Ravishankar, H., Nalawadi, U.G. and Hulamani, N.C. (1993) Use of growth regulators
to manipulate alternate bearing rhythm in mango (Mangifera indica L.). Karnataka
Journal of Agricultural Science 6, 712.
Reece, P.C., Furr, J.R. and Cooper, W.C. (1946) The inhibiting effect of the terminal bud
on ower formation in the axillary buds of the Haden mango. American Journal
of Botany 33, 209210.
Reece, P.C., Furr, J.R. and Cooper, W.C. (1949) Further studies of oral induction in the
Haden mango (Mangifera indica L.). American Journal of Botany 36, 734740.
Riov, J. and Goren, R. (1979) Effect of ethylene on auxin transport and metabolism in
midrib sections in relation to leaf abscission of woody plants. Plant, Cell and Envi-
ronment 2, 8389.
T.L. Davenport 162
Riov, J. and Goren, R. (1980) Effect of ethylene on auxin transport and metabolism in
midrib sections in relation to leaf abscission of woody plants. Israel Journal of Botany
28, 60.
Robbertse, P.J., De Wet, E. and Coetzer, L.A. (1988) The inuence of temperature and
boron on pollen tube growth in mango. South African Mango Growers Association
Yearbook 8, 46.
Robbertse, P.J., De Wet, E. and Coetzer, L.A. (1990) The inuence of boron application
on fruit set and yield of mangos. South African Mango Growers Association Year-
book 10, 2427.
Robbertse, P.J., Coetzer, L.A., Tomer, E. and Truscott, M. (1993) Sexual compatibility
between different mango cultivars. South African Mango Growers Association
Yearbook 13, 2426.
Robbertse, P.J., Coetzer, L.A., Tomer, E. and Smith, M.F. (1994) Sexual compatibility
between different mango cultivars. South African Mango Growers Association
Yearbook 14, 1820.
Robert, J.P. and Wolstenholme, B.N. (1992) Phenological cycles, carbohydrate, status,
and CPPU spray trials for four mango cultivars current research in the Nkwalini
Valley. South African Mango Growers Association Yearbook 12, 913.
Roberts, J.A. and Osborne, D.J. (1981) Auxin and the control of ethylene production
during the development and senescence of leaves and fruits. Journal of Experimen-
tal Botany 32, 875889.
Rodriguez, A.G. (1932) Inuence of smoke and ethylene on the fruiting of pineapple
(Ananas sativus Schult.). Journal of the Agricultural University of Puerto Rico
16, 518.
Ross, S.D., Pharis, R.P. and Binder, W.D. (1983) Growth regulators and conifers: their
physiology and potential uses in forestry. In: Nickell, L.G. (ed.) Plant Growth Regu-
lating Chemicals, Vol. II. CRC Press, Boca Raton, Florida, pp. 3578.
Sachs, R.M. (1977) Nutrient diversion: an hypothesis to explain the chemical control of
owering. HortScience 12, 220222.
Sachs, R.M. and Hackett, W.P. (1983) Source-sink relationships and owering. In: Meudt,
W.J. (ed.) Strategies of Plant Reproduction. Allenheld, Osmun, Ottawa, pp. 263272.
Sachs, R.M., Hackett, W.P., Ramina, A. and Maloof, C. (1979) Photosynthetic assimila-
tion and nutrient diversion as controlling factors in ower initiation in Bougainvil-
lea (San Diego Red) and Nicotiana tabacum cv. Wis. 38. In: Marcelle, R., Clysters,
H. and Van Poucke, M. (eds) Photosynthesis and Plant Development. Dr W. Junk
Publishers, The Hague, the Netherlands, pp. 95102.
Sadhu, M.K. and Bose, S. (1988) Some endogenous factors affecting rooting of mango
cuttings. Gartenbauwissenschaft 53, 137141.
Saha, B.P. and Chhonkar, V.S. (1972) Studies on the oral biology, pollen viability, and
intervarietal compatibility in mango (Mangifera indica L.). (Abstract). In: Program
and Abstracts of the Third International Symposium Sub-tropical and Tropical Hor-
ticulture, Horticultural Society of India, Bangalore, India, p. 5.
Saidha, T., Rao, V.N.M. and Santhanakrishnan, P. (1983) Internal leaf ethylene levels in
relation to owering in mango (Mangifera indica L.). Indian Journal of Horticulture
40, 139145.
Salisbury, F.B. and Ross, C.W. (1992) Plant Physiology. Wadsworth, Belmont, California,
540 pp.
Salomon, E. and Reuveni, O. (1994) Effect of paclobutrazol treatment on the growth and
rst owering of intact and autografted seedlings of mango. Scientia Horticulturae
60, 8187.
Samach, A. and Wigge, P.A. (2005) Ambient temperature perception in plants. Current
Opinion in Plant Biology 8, 483486.
Reproductive Physiology 163
Snchez-Snchez, E., Cabrera-Carbajal, F., Padilla-Valenzuela, I., Samaniego Russo, J.A.
and Aboytia-Mendivil, R. (2004) Gibberellic acid effect on sprouting and nutri-
tional balance of young trees of Keitt mango at the Mayo Valley, Sonora. Acta
Horticulturae 645, 447452.
Schaffer, B., Whiley, A.W. and Crane, J.H. (1994) Mango. In: Schaffer, B. and Andersen,
P.C. (eds) Handbook of Environmental Physiology of Fruit Crops Vol. II, Subtropical
and Tropical Crops. CRC Press, Boca Raton, Florida, pp. 165197.
Schaffer, B., Whiley, A.W. and Searle, C. (1999) Atmospheric CO
2
enrichment, root re-
striction, photosynthesis, and dry-matter partitioning in subtropical and tropical
fruit crops. HortScience 34, 10331037.
Schnell, R.J., Olano, C.T., Quintanilla, W.E. and Meerow, A.W. (2005) Isolation and
characterization of 15 microsatellite loci from mango (Mangifera indica L.) and
cross-species amplication in closely related taxa. Molecular Ecology Notes 5,
625627.
Scholeeld, P.B. (1982) Scanning electron microscope study of owers of avocado, li-
tchi, macadamia and mango. Scientia Horticulturae 16, 263272.
Scholeeld, P.B. and Oag, D.R. (1984) Flowering and fruit set of six cultivars of mango.
In: Proceedings of the First Australian Mango Research Workshop. Commonwealth
Scientic and Industrial Research Organization (CSIRO), Melbourne, pp. 96103.
Scholeeld, P.B., Oag, D.R. and Sedgley, M. (1986) The relationship between vegetative
and reproductive development in the mango in northern Australia. Australian Jour-
nal of Agricultural Research 37, 425433.
Scott, T.K., Case, D.B. and Jacobs, W.P. (1967) Auxin-gibberellin interaction in apical
dominance. Plant Physiology 42, 13291333.
Searle, N.E. (1965) Physiology of owering. Annual Review of Plant Physiology 16,
97118.
Searle, C., Whiley, A.W., Simpson, D.R. and Saranah, J.B. (1995) A preliminary pheno-
physiological model for Kensington mango in subtropical Australia. In: Mango
2000 Marketing Seminar and Production Workshop Proceedings. Department of
Primary Industries, Brisbane, pp. 127135.
Sen, P.K. (1939) Annual Report Fruit Research Station Sabour, India for 19381939. Fruit
Research Station, Sabour, India, pp. 1218.
Sen, P.K. and Mallik, P.C. (1941) The time of differentiation of the ower bud of the
mango. Indian Journal of Agricultural Sciences 11, 7479.
Sen, P.K. and Roy, R.S. (1935) Mineral nutrition of mango. Proceedings of the Indian
Science Congress 111, 123.
Sen, P.K., Mallik, P.C. and Ganguly, B.D. (1946) Hybridization of the mango. Indian
Journal of Horticulture 4, 415.
Sen, P.K., Maiti, S.K. and Maiti, S.C. (1972) Studies on inductions of axillary owering
in Mangifera indica L. Acta Horticulturae 24, 185188.
Sen, P.K., Bandopadhyay, M., Roy, S.S. and Basu, R.N. (1973) Use of ethrel in control-
ling non-uniform bearing of mango (Mangifera indica L.). Indian Agriculture 17,
285288.
Sergent, E., Ferrari, D. and Leal, F. (1996) Effects of potassium nitrate and paclobutrazol
on owering induction and yield of mango (Mangifera indica L.) cv. Haden. Acta
Horticulturae 455, 180187.
Sharma, D.K. and Singh, R.N. (1970) Self-incompatibility in mango (Mangifera indica
L.). Horticultural Research 10, 108118.
Sharma, D.K. and Singh, R.N. (1972) Investigations on self-incompatibility in Mangifera
indica L. Acta Horticulturae 24, 126130.
Shawky, I., Zidan, Z. and Dahshan, D.I. (1977) Sex distribution, fruit set and fruiting of
Zebda mango inorescence. Annals of Agricultural Science (Cairo) 20, 159.
T.L. Davenport 164
Shawky, I., Zidan, Z., El-Tomi, A. and Dahsan, D.I. (1978) Effect of GA
3
sprays on time
of blooming and owering malformation in Taimour mango. Egyptian Journal of
Horticulture 5, 123132.
Shivashankara, K.S. and Mathai, C.K. (1995) Physiological diversity among the poten-
tially productive branches of regular and irregular bearing mango cultivars. Photo-
synthetica 31, 135140.
Shorrocks, V.M. (1997) The occurrence and correction of boron deciency. Plant and
Soil 193, 121148.
Shu, Z.H. (1981) Flower differentiation of mango (Mangifera indica). National Science
Council Monograph 9, 865870.
Shu, Z.H. (1993) Chemical pruning and induction of panicles in mango. Acta Horticul-
turae 341, 199205.
Shu, Z.H. and Sheen, T.F. (1987) Floral induction in axillary buds of mango (Mangifera
indica L.) as affected by temperature. Scientia Horticulturae 31, 8187.
Shu, Z.H., Sheen, T.F. and Lee, K.C. (1989) Current researches on the unfruitfulness of
mango in Taiwan. Acta Horticulturae 231, 68.
Simao, S. and Maranhao, Z.C. (1959) Os insectos como agentes polinizadoses de
Mangueira. Anais Escola Superior de Agricultura, Luiz de Oucitoz, University of
So Paulo 76, 300304.
Singh, A.K. and Rajput, C.B.S. (1990) Effect of GA, BA, and calcium on vegetative growth
and owering in mango (Mangifera indica L.). Research and Development Report
7, 111.
Singh, L. and Khan, A.A. (1939) Relation of growth to fruit bearing in mangoes. Indian
Journal of Agricultural Sciences 9, 835867.
Singh, L.B. (1959) Movement of owering substances in the mango (Mangifera indica L.)
leaves. Horticultural Advance 3, 2027.
Singh, L.B. (1960) The Mango. Botany, Cultivation and Utilization. Leonard Hill, Lon-
don, 439 pp.
Singh, L.B. (1961) Biennial bearing in mango effect of gibberellic acid and maleic
hydrazide. Horticultural Advance 5, 96106.
Singh, L.B. (1962) A new technique for inducing early fruiting in mango hybrids based on
the movement of hormones. Proceedings of the Horticultural Society 75, 410412.
Singh, L.B. (1972) Biennial bearing in mango retrospect and prospect. Acta Horticul-
turae 24, 145148.
Singh, L.B. (1977) Mango. In: Alvim, P. de T. and Kozlowski, T.T. (eds) Ecophysiology of
Tropical Crops. Academic Press, New York, pp. 479485.
Singh, L.B. and Singh, R.N. (1956) Floral induction in axillary buds of mango shoots.
Proceedings of the American Society for Horticultural Science 68, 265269.
Singh, L.B. and Singh, R.N. (1959) Shy-bearing habit of south Indian mango varieties
and their sex ratio. Annual Report Fruit Research Station Saharanpur 19571859.
Fruit Research Station, Saharanpur, India, pp. 5457.
Singh, R. and Singh, R.N. (1974) Growth-promoting and growth-inhibiting substances in
developing fruits of biennial and regular bearing varieties of mango (Mangifera in-
dica L.). Indian Journal of Horticulture 31, 1622.
Singh, R.N. (1954a) Studies on the oral biology and subsequent development of fruits
in the mango (Mangifera indica L.) varieties Dashehari and Langra. Indian Journal
of Horticulture 11, 6988.
Singh, R.N. (1954b) The sex ratio and fruit set in three varieties of mango. Science 119,
389390.
Singh, R.N. (1958a) Studies in the differentiation and development of fruit buds in man-
go (Mangifera indica L.) I. Review of the literature. Horticultural Advance 2, 17.
Reproductive Physiology 165
Singh, R.N. (1958b) Studies in the differentiation and development of fruit buds in man-
go (Mangifera indica L.) II. Morphological and histological changes. Horticultural
Advance 2, 3743.
Singh, R.N. (1959) Studies in the differentiation and development of fruit buds in mango
(Mangifera indica L.) varieties III. Mango shoots and fruit bud differentiation. Horti-
cultural Advance 3, 2849.
Singh, R.N. (1961) Studies in the differentiation and development of fruit-buds in mango
(Mangifera indica L.). V. Effects of defoliation, decapitation and deblossoming on
fruit-bud differentiation. Indian Journal of Horticulture 18, 111.
Singh, R.N. (1964) Sex, pollination and post-fertilization problems in mango. World
Crops 16, 2426.
Singh, R.N. (1971) Biennial Bearing in Fruit-trees. Indian Council of Agricultural Re-
search (ICAR) Technical Bulletin (Agriculture) Number 30. ICAR, New Delhi.
Singh, R.N. (1978) Mango. Indian Council of Agricultural Research (ICAR), New Delhi,
99 pp.
Singh, R.N. (1979) Mango. Indian Council of Agricultural Research (ICAR) Low-priced
Books Series No. 3. ICAR, New Delhi.
Singh, R.N., Majumder, P.K. and Sharma, D.K. (1962a) Age of leaf as affecting fruit bud
formation in mango (Mangifera indica L.) var. Neelum. Science and Culture 28,
284285.
Singh, R.N., Majumdar, P.K. and Sharma, D.K. (1962b) Self-incompatibility in mango
(Mangifera indica L.) var. Dashehari. Current Science 31, 209.
Singh, R.N., Majumdar, P.K. and Sharma, D.K. (1965) Studies on the bearing behavior of
some south Indian varieties of mango (Mangifera indica L.) under north Indian
conditions. Tropical Agriculture 42, 171174.
Singh, R.N., Majumder, P.K. and Sharma, D.K. (1966) Sex-expression in mango
(Mangifera indica L.) with reference to prevailing temperature. Proceedings of the
American Society for Horticultural Science 89, 228229.
Singh, R.N., Majumder, P.K., Sharma, D.K., Sinha, G.C. and Bose, P.C. (1974) Effect of
de-blossoming on the productivity of mango. Scientia Horticulturae 2, 399403.
Singh, R.S. and Ram, S. (1983) Studies on the use of plant growth substances for fruit
retention in mango cv. Dashehari. Indian Journal of Horticulture 40, 188194.
Singh, S. and Singh, B. (1963) Alternate bearing in mango. II. Regulation of growth and
bearing with some plant growth regulators. Punjab Horticulture Journal 3, 137147.
Singh, S.N. (1961) Studies on the morphology and viability of the pollen grains of man-
go. Horticultural Advance 5, 121144.
Singh, S.N. (1963) Studies on the morphology, viability and preservation of pollen grains
of mango (Mangifera indica L.), litchi (Litchi chinensis Sonn.) and loquat (Eriobotrya
japonica Lindl.). Agra University Journal of Research Science 12, 317322.
Singh, U.R. (1960) Studies in the fruit-drop of mango (Mangifera indica L.). II. Nature
and extent of fruit-drop. Horticultural Advance 4, 142154.
Singh, U.R. (1961) Studies in the fruit-drop of mango. IV. Embryo development, its de-
generation and studies of fruit-pedicel and abscission zone. Horticultural Advance
5, 218227.
Singh, U.R. and Singh, A.P. (1973) Studies on the morphology and viability of pollen
grains of mango. Punjab Horticulture Journal 13, 237246.
Singh, Z. and Dhillon, B.S. (1987) Effect of foliar application of boron on vegetative and
panicle growth, sex expression, fruit retention and physicochemical characters of
fruits of mango (Mangifera indica L.) cv Dusheri. Tropical Agriculture 64, 305308.
Singh, Z., Malik, A.U. and Davenport, T.L. (2005) Fruit drop in mango. Horticultural
Reviews 31, 111154.
T.L. Davenport 166
Sitton, D., Itai, C. and Kende, K. (1967) Decreased cytokinin production in the roots as
a factor in shoot senescence. Planta 73, 296300.
Sivagami, S., Vijayan, K.P. and Natarajaratnam, N. (1989) Effect of nutrients and growth
regulating chemicals on biochemical aspects and hormonal balance with reference
to apical dominance in mango. Acta Horticulturae 231, 476482.
Skoog, F. and Miller, C.O. (1957) Chemical regulation of growth and organ formation in
plant tissues cultured in vitro. Symposium of the Society of Experimental Biology
54, 118130.
Smith, F.C. (1960) Beekeeping in the Tropics. Longmans, London, 265 pp.
Soule, J. (1985) Glossary for Horticultural Crops. Wiley, New York, 898 pp.
Soule, M.J. (1950) A Bibliography of the Mango (Mangifera indica L.). Florida Mango
Forum and University of Miami, Coral Gables, Florida, 89 pp.
Spencer, J.L. and Kennard, W.C. (1955) Studies on mango (Mangifera indica L.) fruit set
in Puerto Rico. Tropical Agriculture 32, 323330.
Spencer, J.L. and Kennard, W.C. (1956) Limited stigmatic receptivity may contribute to
low fruit-set in the mango. Proceedings of the American Society for Horticultural
Science 67, 287289.
Sturrock, T.T. (1966) The mango inorescence. Proceedings of the Horticultural Society
53, 366369.
Subhadrabandu, S. (1986) Studies of plant growth regulator effects on tropical and sub-
tropical tree fruits of Thailand. Acta Horticulturae 175, 291297.
Sukhvibul, N., Whiley, A.W., Smith, M.K., Hetherington, S.E. and Vithanage, V. (1999)
Effect of temperature on inorescence development and sex expression of mono-
and poly-embryonic mango (Mangifera indica L.) cultivars. Journal of Horticultural
Science and Biotechnology 74, 6468.
Sukhvibul, N., Hetherington, S.E., Whiley, A.W., Smith, M.K. and Vithanage, V. (2000)
Effect of temperature on pollen germination, pollen tube growth and seed develop-
ment in mango (Mangifera indica L.). Acta Horticulturae 509, 609616.
Suryanarayana, V. (1978a) Amino acid changes in mango shoots in relation to owering.
Plant Biochemistry Journal 5, 5057.
Suryanarayana, V. (1978b) Seasonal changes in ribonucleic acid and protein contents in
mango shoots in relation to owering. Plant Biochemistry 5, 913.
Suryanarayana, V. (1978c) Seasonal changes in sugars, starch, nitrogen and carbon-
nitrogen ratio in relation to owering in mango. Plant Biochemistry Journal 5, 108
117.
Suryanarayana, V. (1980) Amino acid changes in mango shoots as affected by growth
retardants in relation to owering. Plant Biochemistry Journal 7, 7882.
Suryanarayana, V. and Rao, V.N.M. (1977) Ascorbic acid changes in shoots of mango cv.
Mulgoa as affected by growth retardants in relation to owering. Indian Journal of
Plant Physiology 20, 8890.
Takahashi, N. (ed.) (1986) Chemistry of Plant Hormones. CRC Press, Boca Raton, Flori-
da, 277 pp.
Tamaki, S., Matsuo, S., Wong, H.L., Yokoi, S. and Shimamoto, K. (2007) Hd3a protein is
a mobile owering signal in rice. Science 316, 10331036.
Tanaka, M., Takei, K., Kojima, M., Sakakibara, H. and Mori, H. (2006) Auxin controls
local cytokinin biosynthesis in the nodal stem in apical dominance. The Plant Jour-
nal 45, 10281036.
Teaotia, S.S., Singh, R.N., Upadhyay, S.K. and Srivastava, V.S. (1967) Effect of growth
substances on fruit retention in mango (Mangifera indica L.) varieties Langra and
Dashehari. International Symposium on Sub-tropical and Tropical Horticulture.
New Delhi, pp. 224229.
Reproductive Physiology 167
Thuck-Thye, L. (1978) Ethylene and the induction of owering by potassium nitrate in
mango (Mangifera indica L.). MSc thesis, University of the Philippines, Los Banos,
the Philippines.
Tomer, E. (1984) Inhibition of owering in mango by gibberellic acid. Scientia Horticul-
turae 24, 299303.
Tomlinson, P.B. and Gill, A.M. (1973) Growth habits of tropical trees: some guiding
principles. In: Meggers, B.J., Ayensu, E.S. and Duckworth, W.D. (eds) Tropical For-
est Ecosystems in Africa and South America: a Comparative Review. Smithsonian
Institution, Washington, DC, pp. 124143.
Tongumpai, P., Hongsbhanich, N. and Voon, C.H. (1989) Cultar for owering regula-
tion of mango in Thailand. Acta Horticulturae 239, 375378.
Tongumpai, P., Jutamanee, K. and Subhadrabandhu, S. (1991a) Effect of paclobutrazol
on owering of mango cv. Khiew Sawoey. Acta Horticulturae 291, 6770.
Tongumpai, P., Jutamanee, K., Sethapakdi, R. and Subhandrabandu, S. (1991b) Variation
in level of gibberellin-like substances during vegetative growth and owering of
mango cv. Khiew Sawoey. Acta Horticulturae 291, 105107.
Torres, J.P. (1931) Some notes on Carabao mango owers. Philippine Journal of Agricul-
ture 2, 395398.
Turnbull, G.C.N., Anderson, K.L. and Winston, E.C. (1996) Inuence of gibberellin treat-
ment on owering and fruiting patterns in mango. Australian Journal of Experimen-
tal Agriculture 36, 603611.
van der Meulen, J.H.E., Smith, T., van den Boom, I.B., Kok, A., Schwartz, C. and Jacobs,
J. (1971) Mango Growing in South Africa. Leaet No. 48. Citrus and Subtropical
Fruit Research Institute, Nelspruit, South Africa, 40 pp.
Van Lelyveld, L.J. (1978) Peroxidase activity and isozymes in abscised and normal man-
go (Mangifera indica L.) fruits. Zeitschrift fr Panzenphysiologie 89, 453456.
Van Lelyveld, L.J. and Nel, E. (1982) Ethylene concentration and polyphenol oxidase
activity in mango (Mangifera indica L.) fruit abscission. Zeitschrift fr Panzenphys-
iologie 107, 179182.
Vasil, I.K. (1963) Effect of boron on pollen germination and pollen tube growth. In: Lin-
skens, H.F. (ed.) Pollen Physiology and Fertilization. North Holland, Amsterdam,
pp. 107119.
Veen, H. (1969) Auxin transport, auxin metabolism and ageing. Acta Botanica Neer-
landica 18, 447453.
Veen, H. and Jacobs, W.P. (1969) Transport and metabolism of indole-3-acetic acid in
coleus petiole segments of increasing age. Plant Physiology 44, 11571162.
Venkataratnam, L. (1949) Hormone induced set and parthenocarpy in mango. Current
Science 18, 409.
Verheij, E.W.M. (1986) Towards a classication of tropical tree fruits. Acta Horticulturae
175, 137150.
Vijayalakshmi, D. and Srinivasan, P.S. (1999) Morpho-physiological changes as inu-
enced by chemicals and growth regulators in alternate bearing mango cv. Alphonso.
Madras Agricicultural Journal 86, 485487.
Voon, C.H., Pitakpaivan, C. and Tan, S.J. (1991) Mango cropping manipulation with
Cultar. Acta Horticulturae 291, 219228.
Wagle, P.V. (1929) A preliminary study of the pollination of the Alphonso mango. Agri-
culture Journal of India 24, 259263.
Weberling, F. (1989) Morphology of Flowers and Inorescences. Cambridge University
Press, Cambridge, UK.
Weigel, D., Alvarez, J., Smyth, D.R., Yanofsky, M.F. and Meyerowitz, E.M. (1992) LEAFY
controls oral meristem identity in Arabidopsis. Cell 69, 843859.
T.L. Davenport 168
Went, F.W. (1943) Effect of the root system on tomato stem growth. Plant Physiology 18,
5164.
Werner, H. (1993) Inuence of paclobutrazol on growth and leaf nutrient content of
mango (cv. Blanco). Acta Horticulturae 341, 225231.
Wester, P.J. (1920) The mango. Philippine Bureau of Agriculture Bulletin 18, 370.
Whiley, A.W. (1993) Environmental effects on phenology and physiology of mango a
review. Acta Horticulturae 341, 168176.
Whiley, A.W. (1994) Ecophysiological studies and tree manipulation for maximisation of
yield potential in avocado (Persea americana Mill.). PhD thesis, The University of
Natal, Pietermaritzburg, South Africa.
Whiley, A.W., Saranah, J.B., Rasmussen, T.S., Winston, E.C. and Wolstenholme, B.N.
(1988) Effect of temperature on ten mango cultivars with relevance to production in
Australia. In: Proceedings of the Fourth Australasian Conference on Tree and Nut
Crops, Lismore, New South Wales, Australia, pp. 176185.
Whiley, A.W., Rasmussen, T.S., Saranah, J.B. and Wolstenholme, B.N. (1989) Effect of
temperature on growth, dry matter production and starch accumulation in ten man-
go (Mangifera indica L.) cultivars. Journal of Horticultural Science 64, 753765.
Whiley, A.W., Rasmussen, T.S., Wolstenholme, B.N., Saranah, J.B. and Cull, B.W. (1991)
Interpretation of growth responses of some mango cultivars grown under controlled
temperatures. Acta Horticulturae 291, 2231.
Whiley, A.W., Rasmussen, T.S., Saranah, J.B. and Wolstenholme, B.N. (1996) Delayed
harvest effects on yield, fruit size and starch cycling in avocado (Persea americana
Mill.) in two subtropical environments. II. The late-maturing cv. Hass. Scientia Hor-
ticulturae 66, 3549.
Wigge, P.A., Kim, M.C., Jaeger, K.E., Busch, W., Schmid, M., Lohmann, J.U. and Weigel,
D. (2005) Integration of spatial and temporal information during oral induction in
Arabidopsis. Science 309, 10561059.
Wightman, F., Schneider, E.A. and Thimann, K.V. (1980) Hormonal factors controlling
the initiation and development of lateral roots. II. Effects of exogenous growth fac-
tors on lateral root formation in pea roots. Physiologia Plantarum 49, 304314.
Wilkins, M.B. and Cane, A.R. (1970) Auxin transport in roots. Journal of Experimental
Botany 21, 195211.
Williamson, J.G. and Coston, D.C. (1989) The relationship among root growth, shoot
growth, and fruit growth of peach. Journal of the American Society for Horticultural
Science 114, 180183.
Winston, E.C. (1992) Evaluation of paclobutrazol on growth, owering and yield of man-
go cv. Kensington Pride. Australian Journal of Experimental Agriculture 32, 97104.
Winston, E.C. and Wright, R.M. (1986) Mango ower induction: ethephon, potassium
nitrate and cincturing. In: Proceedings of the First Australian Mango Research Work-
shop. Commonwealth Scientic and Industrial Research Organization (CSIRO),
Melbourne, pp. 202210.
Wolfenbarger, D.O. (1977) Comments on mango pollination. Proceedings of the Florida
State Horticultural Society 90, 240241.
Wolstenholme, B.N. and Hofmeyr, D. (1985) Effects of various oral induction treat-
ments on container-grown mango trees. South African Mango Growers Association
Research Yearbook 5, 3638.
Wolstenholme, B.N. and Mullins, P.C. (1982a) Flowering, pollination and fruit set of
mango a discussion of limiting factors. The Citrus and Subtropical Fruit Journal
October, 1122.
Wolstenholme, B.N. and Mullins, P.C. (1982b) Flowering, pollination and fruit set of
mango a discussion of limiting factors. South African Mango Growers Association
Yearbook 2, 511.
Reproductive Physiology 169
Wolstenholme, B.N. and Whiley, A.W. (1995) Ecophysiology of the mango tree as a basis
for pre-harvest management. In: Mango 2000 Marketing Seminar and Production
Workshop Proceedings. Department of Primary Industries, Brisbane, pp. 1017.
Wu, Y., Spollen, W.G., Sharp, R.E., Hetherington, P.R. and Fry, S.C. (1994) Root growth
maintenance at low water potentials: increased activity of xyloglucan endo-
transglycosylase and its possible regulation by abscisic acid. Plant Physiology 106,
607615.
Yanofsky, M.F. (1995) Floral meristems to oral organs: genes controlling early events in
Arabidopsis ower development. Annual Review of Plant Physiology and Plant Mo-
lecular Biology 46, 167188.
Yelenosky, G., Vu, C.V.J. and Bausher, M.G. (1993) Paclobutrazol-induced dwarng of
Valencia orange trees. Proceedings of the Florida State Horticultural Society 106,
329332.
Yeshitela, T., Robbertse, P.J. and Stassen, P.J.C. (2004a) Paclobutrazol suppressed vegeta-
tive growth and improved yield as well as fruit quality of Tommy Atkins mango
(Mangifera indica) in Ethiopia. New Zealand Journal of Crop and Horticultural Sci-
ence 32, 281293.
Yeshitela, T., Robbertse, P.J. and Stassen, P.J.C. (2004b) Effects of various inductive peri-
ods and chemicals on owering and vegetative growth of Tommy Atkins and Keitt
mango (Mangifera indica) cultivars. New Zealand Journal of Crop and Horticultural
Science 32, 209215.
Yeshitela, T., Robbertse, P.J. and Stassen, P.J.C. (2005) Potassium nitrate and urea sprays
affect owering and yields of Tommy Atkins (Mangifera indica) mango in Ethiopia.
South African Journal of Plant and Soil 22, 2832.
Young, T.W. (1942) Investigations of the unfruitfulness of the Haden mango in Florida.
Proceedings of the Florida State Horticultural Society 55, 106110.
Young, T.W. (1955) Inuence of temperature on growth of mango pollen. Proceedings of
the Florida State Horticultural Society 68, 308313.
Young, T.W. (1958) A technique for testing mango pollen viability on articial medium.
Horticultural Advances 2, 7275.
Young, T.W. and Sauls, J.W. (1979) The Mango Industry in Florida. Bulletin 189 Florida
Cooperative Extension Service. University of Florida, Gainesville, pp. 170.
Zeevaart, J. (1976) Physiology of ower formation. Annual Review of Plant Physiology
27, 321348.
Zeevaart, J.A.D. (2006) Florigen coming of age after 70 years. The Plant Cell 18, 1783
1789.
Zeevaart, J.A.D. and Boyer, G.L. (1987) Photoperiodic induction and the oral stimulus
in Perilla. In: Atherton, J.G. (ed.) Manipulation of Flowering. Butterworths, London,
pp. 269277.
Zhang, T., Ying, Z. and Davenport, T.L. (2005) Isolation and characterization of a Constans-
like gene in mango. (Abstract). In: Plant Biology 2005. Annual Meeting of the Amer-
ican Society of Plant Biologists. American Society of Plant Biologists, Seattle, Wash-
ington, p. 151.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
170 (ed. R.E. Litz)
6 Ecophysiology
B. Schaffer,
1
L. Urban,
2
P. Lu
3
and A.W. Whiley
4
1
University of Florida, Florida, USA
2
Centre INRA de Corse, San Giuliano, France
3
EWL Sciences, PO Box 39443, Winnellie, Northern Territory, Australia
4
Sunshine Coast Horticultural Services Pty Ltd, Nambour, Queensland, Australia
6.1 Introduction 170
6.2 Photosynthesis 172
Introduction 172
Light 177
Leaf temperature 179
Elevated atmospheric CO
2
concentration 181
Humidity 182
Flooding 182
Internal factors 184
Photosynthetic contributions by fruit 187
6.3 Plant Water Relations 188
6.4 Tree Growth and Development 190
Light 190
Temperature 192
Drought 193
Flooding 194
Wind 195
Salinity 196
Elevated atmospheric CO
2
concentration 197
6.5 Crop Production 198
Temperature limitations to crop production 198
Light interception and orchard design 199
6.6 Conclusions 200
6.1 Introduction
The genetic composition of mango cultivars is the primary determinant of
yield potential. However, actual yield, as well as tree growth and develop-
ment, are mediated by several endogenous factors including previous fruit
load, postharvest vegetative growth, preowering maturity of terminal shoots,
Ecophysiology 171
production and mobilization of carbohydrates, nutritional status, plant
growth substances and carbon to nitrogen ratios (Schaffer et al., 1994). These
factors are either directly or indirectly affected by environmental variables
such as light, temperature and water availability. Environmental conditions
outside the range for optimum growth may also impose stress which results
in physiological changes that reduce growth or cause permanent damage to
mango trees. For example, major climatic events (i.e. extended drought,
oods, wind storms, heat waves and freezes) can cause severe damage to
crops due to development of excessive stress. However, mediated stress and
the release from stress imposed by normal seasonal changes provide condi-
tions that result in the progression of cropping cycles due to phenological
changes in the plants. An example of benecial stress in mango is the
improved synchrony and reliability of owering in subtropical climates due
to cool winter temperatures. Thus, understanding the impact of the environ-
ment on tree physiology and growth, and the particular adaptive strategies
developed through the processes of evolution, can provide a framework to
manage the crop to maximize genetic yield potential (Schaffer et al., 1994).
Physiological responses of mango to environmental variables can be
related to the evolutionary centre of origin of a specic cultivar. Mango culti-
vars are classied into two ecotypes based on embryony (see Mukherjee and
Litz, Chapter 1, this volume). A race with a single zygotic seed, monoembry-
onic types, evolved in the dry subtropical, monsoonal regions of the Indian
subcontinent with very hot summers but cooler winters. The polyembryonic
types, produced through nucellar embryony, largely evolved in the consis-
tently hot, humid tropics of South-east Asia where the monsoonal pattern
still predominates but the dry season is shorter than that of the Indian sub-
continent (Mukherjee, 1972). Hybridization occurs freely within and between
the two ecotypes and has led to a proliferation of cultivars of widely varying
genetic composition. Differences in growth and owering responses to tem-
perature have been observed between the two embryonic ecotypes (Whiley
et al., 1989) and selection and breeding offer potential for increasing the crop-
ping performance of this notoriously low-yielding and recalcitrant tree across
a wider range of environmental conditions (Schaffer et al., 1994).
Inevitably, the many unique features of the mango tree represent its evo-
lutionary response to an indigenous environment that is particularly hostile,
with sustained extreme heat and high evaporative demand for much of the
year. This chapter provides an overview of the impact of environmental fac-
tors on physiology, growth and productivity of mango. Plant responses will
be considered in the context of the evolutionary origin and adaptability of
the mango tree. Responses to light, temperature and water are emphasized,
while the effects of atmospheric carbon dioxide (CO
2
) concentration, wind
and salinity are also discussed.
Photosynthesis and plant water relations are closely associated with
environmental conditions and directly affect plant growth and productivity.
The principles of leaf gas exchange and plant water relations are discussed to
provide a theoretical basis for interpreting physiological responses to the
environment. The impact of photosynthesis and tree water relations on
B. Schaffer et al. 172
growth and development under different environmental conditions is dis-
cussed. Pollination, fertilization, owering and fruit set, which are strongly
inuenced by environmental factors, have been addressed elsewhere (see
Davenport, Chapter 5, this volume; Schaffer et al., 1994). In the nal sec-
tion of this chapter, mango crop production is integrated with aspects of
ecophysiology.
6.2 Photosynthesis
Introduction
The net CO
2
assimilation rate (A
net
) in C
3
plants is a function of the carboxyla-
tion rate (V
c
), the oxygenation rate (V
o
) and the rate of CO
2
evolution in light
that results from processes other than photorespiration, sometimes called
day respiration (R
d
):
A
net
= V
c
0.5V
o
R
d
(6.1)
R
d
is usually inferred from measurements of leaf CO
2
exchanges after 5 min
in the dark (i.e. night respiration R
n
). However, it has been repeatedly
shown that R
d
is lower than R
n
(see Atkin et al. (2000) for review), so that light
is known to inhibit respiration, with a R
d
/R
n
value ranging from 30 to 100%
(see Peisker and Apel (2001) for review). Urban et al. (2008) established the
following linear regression for R
d
of mango leaves: R
d
= 0.35R
n
0.21, which
may be used to infer R
d
from R
n
for photosynthetic photon ux (Q) values
above 170 mol photons/m
2
/s.
Currently, modelling of A
net
often uses the Harley et al. (1992) version
of the Farquhar et al. (1980) model. According to this model, A
net
can be
expressed as:
A
net
= (1 0.5O/(C
i
))min(W
c
, W
j
, W
p
) R
d

(6.2)
where O represents the partial pressure of oxygen (O
2
) in the intercellular air
spaces (Pa), the specicity factor of ribulose-1,5-bisphosphate carboxylase/
oxygenase (Rubisco). C
i
is the partial pressure of CO
2
in the intercellular air
spaces (Pa), W
c
the carboxylation rate limited by the amount, activation state
or kinetic properties of Rubisco (mol CO
2
/m
2
/s), W
j
the carboxylation rate
limited by the rate of ribulose bisphosphate regeneration (mol CO
2
/m
2
/s),
and W
p
the carboxylation rate limited by triose phosphate utilization in
sucrose and starch synthesis (mol CO
2
/m
2
/s).
Usually O is set as 21 kPa (21%). The variable , which characterizes the
ratio of the afnities of CO
2
and O
2
for ribulose-1,5-bisphosphate in the active
site of Rubisco, can be calculated from the CO
2
compensation point * (the
CO
2
concentration at which photosynthesis equilibrates with respiration):
= 0.5O/* (6.3)
where = 2220 mol CO
2
/mol O
2
at 25C for Cogshall mango leaves (Urban
et al., 2008), which is lower than those given by Epron et al. (1995): 21002900
Ecophysiology 173
mol CO
2
/mol O
2
. Rubiscos large subunit is encoded by a single gene in the
chloroplast genome, and no post-transcriptional modications have been
discovered so far. It is thus very unlikely that can change in the short term
(Spreitzer and Salvucci, 2002).
The internal partial pressure of CO
2
(C
i
) is one of the two major variables
of photosynthesis (with the photosynthetically active photon ux density).
It may be calculated from the supply function:
C
i
= C
a
A
net
/g
b
A
net
/g
s
(6.4)
where C
a
is the partial pressure of CO
2
(Pa) in ambient air, g
b
represents the
leaf boundary layer conductance (mol H
2
O/m
2
/s), and g
s
is the stomatal
conductance of water (H
2
O) (mol H
2
O/m
2
/s).
Stomatal conductance is the major factor controlling A
net
. It ranges from
c.0.02 to c.0.4 mol H
2
O/m
2
/s in Cogshall mango leaves and may be linearly
related to A
net
(Urban et al., 2002, 2003, 2006). The slope of the relationship
between g
s
and A
net
however is affected by drought (Fig. 6.1). Variations in
the slope of this relationship reect changes in photosynthetic water use ef-
ciency and are not well understood.
It must be stressed that using C
i
as the driving variable of photosynthesis
is much debated. It has been advocated that C
c
, the partial pressure of CO
2
at
the site of carboxylation, should be utilized instead. Using C
i
implies that the
following assumptions have been made: C
c
= C
i
and g
m
= 0, where g
m
repre-
sents mesophyll conductance, also called liquid phase resistance, which
Dry
y = 0.009x + 0.024
R
2
= 0.695
0
0.05
0.10
0.15
0.20
0.25
0.30
0.35
0.40
0.45
0.50
0 5 10 15 20
A
net
(mol CO
2/
m
2
/
s)
g
s

(
m
o
l

H
2
O
/
m
2
/
s
)
Wet
y = 0.025x + 0.028
R
2
= 0.86
Fig. 6.1. The relationship between stomatal conductance (g
s
) and net photosynthesis
(A
net
) in mango leaves from well-irrigated () and drought-stressed () 12-year-old
Cogshall trees (Source: redrawn from Urban et al., 2006).
B. Schaffer et al. 174
encompasses diffusion from the intercellular leaf spaces to the carboxylation
sites in the chloroplasts. There is a growing body of evidence that g
m
is not
negligible in most species. The average value of g
m
in unstressed mango
leaves (0.21 mol CO
2
/m
2
/s) (Urban et al., 2008), calculated using the method
of Epron et al. (1995), is within the range of values for broadleaf species sur-
veyed by Ethier and Livingston (2004) and Manter and Kerrigan (2004). The
carboxylation rate (in Eqn 6.2) limited by the amount, activation state or
kinetic properties of Rubisco (W
c
) can be calculated as:
W
c
= V
cmax
C
i
/(C
i
+ K
c
(1 + O/K
o
)) (6.5)
where V
cmax
represents the maximum rate of carboxylation (mol CO
2
/
m
2
/s), and K
c
(Pa CO
2
) and K
o
(Pa O
2
) are the Michaelis constants of Rubisco
carboxylation and oxygenation, respectively. The V
cmax
values of well-
exposed mango leaves at a leaf temperature of 30C are typically in the range
of 80100 mol CO
2
/m
2
/s (Urban et al., 2006). Specic values of K
c
and K
o
for
mango leaves have not been estimated and are approximated using data
from other species (i.e. cotton or tobacco).
The carboxylation rate limited by the rate of ribulose bisphosphate regen-
eration (W
j
) is controlled by the rate of electron ow J (mol electrons/m
2
/s):
W
j
= JC
i
/(4(C
i
+ O/)) (6.6)
with
J = Q/(1 +
2

2
Q
2
/J
max
2
)
0.5
(6.7)
where Q is the photosynthetically active photon ux density (mol quanta/
m
2
/s), represents leaf absorbance (no units), is the apparent efciency of
light energy conversion (mol electrons/mol photons) and J
max
is the light-
saturated rate of electron transport (mol electrons/m
2
/s). Leaf absorbance
of mango leaves, measured from 390760 nm using an integrating sphere,
was found to be close to 0.81 (Urban et al., 2008) and is in the normal range of
values of the literature (Bauerle et al., 2004). Leaf absorbance, which is pos-
itively correlated with leaf chlorophyll content, may increase as a conse-
quence of paclobutrazol treatments (Gonzalez and Blaikie, 2003). The apparent
efciency of light energy conversion in mango reaches 0.32 mol electrons/
mol photons (Urban et al., 2004b), in the absence of photoinhibition or pho-
todamage. This value corresponds to the mean value of operational (Sin-
gaas et al., 2001). The J
max
values of well-exposed mango leaves at a leaf
temperature of 30C are typically in the 120150 mol CO
2
/m
2
/s range. The
J
max
as well as the V
cmax
values are rather low when compared to values from
other species and partly explain why maximal rates of leaf photosynthesis
(A
max
) are rather low, typically 1215 mol CO
2
/m
2
/s.
The carboxylation rate limited by triose phosphate utilization during
sucrose and starch synthesis (W
p
in Equation 2), can be calculated by:
W
p
= 3TPU + V
o
/2 = 3TPU + V
c
0.5C
i
/(C
i
) (6.8)
where TPU is the rate of phosphate release in triose phosphate utilization
during starch and sucrose production. The TPU is usually not included in
Ecophysiology 175
most studies on photosynthetic capacity because of methodological difcul-
ties. However, Urban et al. (2003) found that TPU = 812 mol CO
2
/m
2
/s at
a leaf temperature of 30C in well-exposed Cogshall mango leaves.
The variables V
cmax
and J
max
are temperature dependent and their depen-
dency is described by:
Parameter (V
cmax
, J
max
, TPU) = exp(c H
a
/(RT
l
))
/(1+exp((ST
l
H
d
)/(RT
l
))) (6.9)
where c is a scaling factor, H
a
(J/mol) the activation energy of the given
parameter, R the gas constant (8.3143 J/K/mol), T
l
(K) the leaf temperature,
S (J/mol) an entropy term and H
d
(J/mol) the deactivation energy of the
given parameter.
Similarly, the temperature dependency of R
d
, , K
c
and K
o
is described
by:
Parameter (R
d
, , K
c
, K
o
) = exp(c H
a
/(RT
l
)) (6.10)
Proteins of the Calvin cycle and thylakoids represent the majority of leaf
nitrogen (N). Therefore, photosynthetic capacity is strongly related to leaf N
content expressed on an area basis (N
a
) (Field and Mooney, 1986; Evans, 1989;
Kellomki and Wang, 1997; Walcroft et al., 1997). To account for the relation-
ship commonly observed between the parameters dening photosynthetic
capacity (V
cmax
, J
max
, TPU and R
d
mainly) and N
a
(Field and Mooney, 1983;
Harley et al., 1992) (Fig. 6.2), scaling factors c of V
cmax
, J
max
, TPU and R
d
may
be related to N
a
, either linearly or slightly non-linearly.
In summary, leaf net photosynthesis depends on ve major classes of
factors, either variables (external or internal factors) or parameters (more or
less constant factors), provided that plants are not exposed to too extreme
conditions; we may consider the internal factors as genetic factors. The ve
classes of factors are:
The photosynthetically active photon ux density ( 1. Q), which is the major
driving variable of photosynthesis. Gross photosynthesis is determined by
Q while C
i
determines the proportion of photorespiration, and thus net
photosynthesis. One of the major environmental factors affecting C
i
is water
availability in the root zone through its effect on g
s
.
Leaf nitrogen concentration ( 2. N
a
), which is not a rate-determining factor
of photosynthesis, unlike Q, but may be considered as a rate-limiting factor.
In other words, N
a
sets the photosynthetic potential of a leaf (i.e. photosyn-
thetic capacity). We shall see below which factors inuence N
a
in mango
leaves.
Leaf temperature inuences leaf photosynthesis. Net photosynthesis is 3.
positively correlated with leaf temperature in a normal range. Leaf tempera-
ture (T
l
) is not a driving variable of photosynthesis but it is the single most
important rate-determining factor after Q. In addition, extreme temperatures
may inuence photosynthesis through their damaging effects. Kinetics of
enzymes involved with photosynthetic reactions collectively comprise an
additional set of factors that inuence leaf net photosynthesis.
B. Schaffer et al. 176
Several parameters related to enzymes include the specicity factor of 4.
Rubisco (), the Michaelis constants of Rubisco carboxylation and oxygen-
ation, K
c
and K
o
, the activation and deactivation energies of the different pa-
rameters H
a
and H
d
, the entropy terms S, c factors and leaf absorbance
(). With the exception of K
c
and K
o
, the specic values of all these parame-
ters have been estimated for Cogshall mango (Urban et al., 2003).
The apparent efciency of light energy conversion ( 5. ). This factor be-
longs to a category of its own since it should theoretically not differ from one
y = 41.52x 15.52
R
2
= 0.87
y = 201.64x
1
+ 173.41
R
2
= 0.88
0
20
40
60
80
100
120
140
(a)
(b)
1.0 1.5 2.0 2.5 3.0 3.5 4.0
N
a
(g N/m
2
)
V
c
m
a
x


(

m
o
l

C
O
2
/
m
2
/
s
)
0
50
100
150
200
250
1.0 1.5 2.0 2.5 3.0 3.5 4.0
y = 66.94x 15.40
R
2
= 0.83
y = 330.44x
1
+ 291.55
R
2
= 0.86
J
m
a
x


(

m
o
l
/
m
2
/
s
)
N
a
(g N/m
2
)
Fig. 6.2. Relationship between (a) the maximum rate of carboxylation (V
cmax
) and (b)
the light-saturated rate of electron transport (J
max
), and nitrogen concentration per unit
leaf area (N
a
). Measurements were performed on mango leaves of 3-year-old Cogshall
trees (), standard leaves () and leaves close to developing fruits () of 11-year-old
Cogshall trees. Best t lines for pooled data correspond to the linear (
_
) and the ax
1
+
b (

) models (Source: redrawn from Urban et al., 2003).


Ecophysiology 177
species to another and may be considered as a constant in the absence of
photoinhibition and photodamage.
Light
Light exposure
Plants allocate nitrogen resources within the canopy to enhance photosyn-
thetic capacity at locations exposed to high incident light levels, thus maxi-
mizing whole plant carbon gain (Field and Mooney, 1983; Hollinger, 1996;
Carswell et al., 2000). For leaves of a given age and for a given nitrogen sup-
ply, leaf N per unit leaf area appears to be strongly related with light expo-
sure (DeJong and Doyle, 1985; Le Roux et al., 1999, 2001; Rosati et al., 1999,
2000). Photosynthetic light acclimation of leaves may result from changes in
either leaf nitrogen concentration (N
m
) or mass-to-area ratio (M
a
) because
N
a
= M
a
N
m
. Lynch and Gonzlez (1993) observed a negative correlation
between N
m
and light exposure in the tropical fruit tree Borojoa patinoi, but
such a behaviour is rare; positive correlations between N
m
and light exposure
are more commonly observed. In addition, photosynthetic light acclimation
of leaves may result from changes in partitioning of total leaf N among the
different pools of the photosynthetic machinery (Evans, 1989). In mango,
light acclimation of photosynthesis results mainly from changes in M
a
, and
to a lesser extent from changes in allocation of total leaf N at low irradiance;
whereas changes in N
m
play only a minor role (Fig. 6.3). Light acclimation of
mango leaves thus follows a pattern similar to peach leaves (Le Roux et al.,
1999; Walcroft et al., 2002).
Light intensity
Photosynthesis of Cogshall mango trees increases with increasing levels of
light intensity to reach a maximum at Q = 1200 mol photons/m
2
/s (L.
Urban, unpublished data). Whiley et al. (1999) measured Q at 1284 mol pho-
tons/m
2
/s for eld-grown Kensington Pride trees growing in subtropical
Queensland, Australia, which is well below full sunlight (full sunlight 2000
mol photons/m
2
/s). Such a high threshold is a typical feature of sun plants.
Individual leaves are rarely able to utilize full sunlight; whole trees consist of
many leaves that shade each other, so that only a small fraction of a trees
leaves are exposed to full sun at any given time of the day, while the rest of
the leaves receive subsaturating photon uxes in the form of small patches of
light that penetrate through gaps of the leaf canopy. Because the photosyn-
thetic response of whole trees is the sum of the photosynthetic activity of all
the leaves, only rarely is photosynthesis saturated with light at the whole-
tree level.
While most leaves experience subsaturating light intensities, well-exposed
leaves of the upper-crown may receive excessive quantities of light. Those
leaves must dissipate the absorbed light energy in excess to prevent damage to
the photosynthetic apparatus. Moderate decreases in maximal quantum
efciency (i.e. quantum efciency of dark-adapted leaves F
v
/F
mPredawn
) are
B. Schaffer et al. 178
typical features of moderate photoinhibition and should be interpreted in
terms of non-photochemical quenching, an adaptative mechanism involving
the xanthophyll cycle and allowing excess energy to be dissipated in the form
of heat (Adams et al., 2005). Such small decreases in F
v
/F
mPredawn
are com-
monly observed in mango leaves even from well-irrigated trees (Urban and
Alphonsout, 2007).
When temperature (30C) and water vapour pressure decit (VPD < 1
kPa) are non-limiting, and in the absence of photoinhibition, maximal rates
of net leaf photosynthesis (A
max
) may reach 1215 mol CO
2
/m
2
/s at saturating
0.0
1.0
2.0
3.0
0.0 0.2 0.4 0.6 0.8 1.0
Gap fraction
N
m

(
g

N
/
g

d
r
y

m
a
t
t
e
r
)
Tree # 1
y = 1.42x + 1.43
R
2
= 0.90
0.0
0.5
1.0
1.5
2.0
2.5
3.0
0.0 0.2 0.4 0.6 0.8 1.0
Gap fraction
N
a

(
g
/
m
2
)Tree # 2
y = 1.33x + 1.44
R
2
= 0.79
(a)
(b)
Fig. 6.3. Relationship between (a) leaf nitrogen concentration per unit mass (N
m
) and
(b) leaf nitrogen concentration per unit leaf area (N
a
) and the gap fraction for mango
leaves measured in the crown of two 3-year-old Cogshall trees. Gap fractions were
measured as an indicator of light exposure. Measurements were performed on leaves
< 2 months old (), 8 months old (), 1214 months old () and 1720 months old
() (Source: Urban et al., 2003).
Ecophysiology 179
Q, on Cogshall trees. Whiley et al. (1999) measured A
max
of 15.2 mol CO
2
/
m
2
/s for Kensington Pride trees growing in a subtropical climate in Queen-
sland, Australia. However, values > 16 mol CO
2
/m
2
/s were observed on
eld-grown trees of Tommy Atkins, Haden and Irwin on sunny days
during the wet season in tropical regions of Australia (P. Lu, unpublished
data). This is much higher than citrus (< 10 mol CO
2
/m
2
/s), but substan-
tially lower than plum (approx. 26 mol CO
2
/m
2
/s). Whiley et al. (1999) esti-
mated the light compensation point to be 29 mol photons/m
2
/s for leaves
of non-stressed, eld-grown mango trees, which is much higher than that
attributed to shade-tolerant species (< 10 mol photons/m
2
/s) (Harvey,
1979). The data show that mango trees are basically sun-adapted plants.
Leaf temperature
Effect of temperatures in a normal range on leaf photosynthetic capacity
Medlyn et al. (2002) calculated optimal temperatures for V
cmax
and J
max
to be
3541C and 3038C, respectively. Tree species native to cold climates had
the lowest temperature optima for both V
cmax
and J
max
. For Cogshall mango
trees, calculated temperature optima for V
cmax
and J
max
are 44 and 45.5C,
respectively. They demonstrated that mango photosynthesis increases with
temperature well above 40C (Fig. 6.4). Although there is a clear lack of refer-
ences for other tropical trees, it is tempting to attribute the temperature
response of mango photosynthesis to its tropical origin.
Estimates of H
a
, H
d
and S are 7.0695, 17.0799 and 536 J/mol for V
cmax
,
and 3.8782, 10.2211 and 317 J/mol for J
max
, respectively. Both H
d
and S are
within the range of published values for V
cmax
and J
max
(Dreyer et al., 2001).
In addition, H
a
for V
cmax
is within the 6080 kJ/mol range for many species,
including crop species as well as deciduous and evergreen trees (Medlyn
et al., 2002). For mango, H
a
for J
max
is consistent with data published for
evergreen species (Medlyn et al., 2002), which is consistent with the fact that
mango leaves commonly are 24 years old before senescence and abscision.
However, the J
max
/V
cmax
at 25C for mango is about 1.86, approximately 11%
higher than the mean value calculated over the whole range of species stud-
ied by Medlyn et al. (2002). Lower activation energies for J
max
than V
cmax

result in a temperature-induced decrease in J
max
/V
cmax
, conrming previous
observations by Walcroft et al. (1997) and Dreyer et al. (2001). The estimate of
H
a
for R
d
is 4.5710 J/mol. H
a
is higher than the range of published values
for R
d
(Dreyer et al., 2001).
Chilling temperatures
F
v
/F
mPredawn
decreases in mango leaves with decreasing temperature, while
chilling reduces quantum efciency (Whiley et al., 1999; Sukhvibul et al.,
2000; Weng et al., 2006a, b). The decrease in F
v
/F
mPredawn
may be interpreted
to reect sustained engagement of zeaxanthin in photoprotective energy dis-
sipation. Decreases in quantum efciency correspond to a decrease in the
rate of electron ow. It may be argued that sustained zeaxanthin-dependent
B. Schaffer et al. 180
energy dissipation reduces the risk of formation of singlet oxygen
1
O
2

in the
antennae, while decreases in J lower the risk of electrons reducing O
2
to anion
superoxide O
2

in the photosynthetic electron transport chain (Adams et al.,


2005). In other words, decreases in F
v
/F
mPredawn
and quantum efciency cor-
respond to adaptative mechanisms against the effect of cold, when photosyn-
thesis is low and there is an imbalance between the quantity of light energy
absorbed and the quantity of energy used in the photochemical reactions of
0.0
0.5
1.0
1.5
2.0
2.5
3.0
3.5
4.0
(a)
(b)
15 20 25 30 35 40 45 50
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
1.6
1.8
2.0
15 20 25 30 35 40 45 50
T
l
(C)
J
m
a
x
/
J
m
a
x

a
t

2
5

C
V
c
m
a
x
/
V
c
m
a
x

a
t

2
5

C
T
l
(C)
Fig. 6.4. Temperature response functions adjusted to the (a) maximal rate of car-
boxylation (V
cmax
) and (b) the light-saturated rate of photosynthetic electron ux (J
max
),
normalized to the mean value at 25C in leaves from Cogshall mango seedlings.
The data scatter represents the real scatter at each temperature. Reference values at
25C were computed for each of eight leaves, taken from young trees from two
origins ( and ), and a unique temperature response was adjusted over the range of
normalized data. T
l
, the leaf temperature; the dotted lines correspond to Equation 9;
the solid lines correspond to Equation 10 (no deactivation energy component).
Ecophysiology 181
photosynthesis (Adams et al., 2005). Interestingly, Weng et al. (2006b) found
that mango leaves transferred from warm and dark to chilling conditions
showed only slight down-regulation of PSII efciency when compared to
leaves moved from dim light to chilling conditions. Of course, long-term
exposure to cold and very low temperatures ( 10C) may eventually
result in true photodamage, not just photoinhibition. Very low values of F
v
/
F
mPredawn
, decreases in chlorophyll content and slow recovery kinetics are all
indicators of photodamage. Sukhvibul et al. (2000) observed that susceptibil-
ity to cold-induced photodamage was more pronounced in polyembryonic
cultivars than in monoembryonic cultivars, possibly reecting their different
eco-evolutionary development.
Elevated atmospheric CO
2
concentration
The CO
2
concentration in the earths atmosphere has been increasing rapidly
since the early 20th century and is continuing to rise, primarily due to burn-
ing of fossil fuels (Houghton, 2005). Earths atmospheric CO
2
concentration
is currently about 370 mol CO
2
/mol (Houghton, 2005) and is projected to
reach 600 mol CO
2
/mol by 2050. Elevated ambient CO
2
levels will undoubt-
edly affect cropping systems since atmospheric CO
2
concentrations can sig-
nicantly affect plant growth and productivity (Idso and Kimball, 1991;
Houghton, 2005). There is little published information concerning the effects
of elevated ambient CO
2
levels on physiology, growth and production of
tropical fruit trees, including mango.
Schaffer et al. (1997) exposed leaves of eld- and container-grown Kens-
ington (syn. Kensington Pride) trees to short durations (several minutes) of
varying ambient CO
2
concentrations. They found that under saturating light
levels for photosynthesis, net photosynthesis increased as ambient CO
2
con-
centration increased up to 1200 mol CO
2
/mol. At ambient CO
2
concentra-
tions > 1200 mol CO
2
/mol, net photosynthesis stabilized, probably due to
leaves reaching their maximum biochemical capacity to x carbon. Studies
with Cogshall mango trees indicated that when C
a
increases, stomata close
swiftly and C
i
may become very unpredictable (L. Urban, unpublished data).
Therefore, using C
i
may be preferable to C
a
for quantifying short-term effects
of elevated CO
2
concentations on A
net
of mango. Saturating CO
2
levels may
often be reached at C
i
= 800 mol CO
2
/mol air.
Long-term (612 months) exposure of Kensington mango trees to an
atmospheric CO
2
concentration of 700 mol/mol resulted in higher net CO
2

assimilation rates than in leaves of plants grown at atmospheric CO
2
concen-
trations of 350 mol/mol when net CO
2
assimilation was measured at the
same CO
2
concentration as the growth environment. However, carboxylation
efciency (the amount of CO
2
xed per mole of ambient CO
2
) was lower for
plants in the CO
2
-enriched environment compared to plants in the ambient
(350 mol CO
2
/mol) environment (Schaffer et al., 1997). Although further
studies are needed to determine the effects of long-term exposure to elevated
CO
2
concentrations on mango growth and productivity, it appears that
B. Schaffer et al. 182
mango will benet from increases in atmospheric CO
2
concentrations. How-
ever, the effects of increased atmospheric CO
2
concentrations associated with
global warming on mango production may be offset by higher respiratory
losses and increased assimilate partitioning to shoot growth in highly vegeta-
tive cultivars (Schaffer et al., 1999). Therefore, responses of mango cultivars
to elevated atmospheric CO
2
concentrations need to be evaluated over a
range of temperatures to ascertain their likely performance under changing
atmospheric conditions.
Humidity
Although mango production occurs in the tropics and subtropics in areas of
high and low relative humidity (RH) (Campbell, 1984), there are very few
published reports on the effects of RH or VPD on physiology and tree
growth.
In a study with container-grown Kensington plants, Pongsomboon et al.
(1992) reported that stomatal conductance was inversely correlated with
VPD. Differences between cultivar responses to VPD have been observed in
eld-grown Irwin (monoembryonic) and Kensington (polyembryonic)
mango trees during the wet and dry season in tropical Australia. During the
wet season and for well-irrigated trees during the dry season, both Irwin
and Kensington showed decreasing stomatal conductance with increasing
leaf-to-air vapour pressure decit (LAVPD) but Kensington showed a more
rapid decrease than Irwin (Fig. 6.5) (P. Lu, unpublished data). It was also
observed that daytime leaf xylem water potential was lower in Irwin than
in Kensington while predawn water potentials were similar for both culti-
vars (P. Lu, unpublished data). These results indicate that under similar soil
water conditions, Kensington tends to close stomata much more rapidly
than Irwin to conserve water under dry atmospheric conditions. This water
conservation strategy is probably a reection of Kensingtons adaptive
responses to the hot and dry seasonal tropical environment under which it
evolved (Wolstenholme and Whiley, 1995). Other studies in tropical Austra-
lia revealed that polyembryonic Nam Doc Mai behaved like Irwin (P. Lu,
unpublished data). However, Nam Doc Mai in Thailand has comparatively
low vigour compared to Kensington when grown in the tropics.
Further research is required to determine if differences in photosynthetic
or stomatal responses of mango to VPD are indeed based on embryonal char-
acteristics. Clarication of the reasons for variation would undoubtedly facil-
itate breeding and selection of cultivars for dry and humid areas.
Flooding
The primary effect of ooding on plants is due to a reduction in soil oxygen
concentration. Oxygen levels in the soil can decrease from 20% to < 5% within
12 days of ooding (Crane and Davies, 1988) and soils eventually become
Ecophysiology 183
anoxic (no oxygen). Mango is considered to be a moderately ood-tolerant
species (Schaffer et al., 1994, 2006) and waterlogging or ooding of trees peri-
odically occurs in many of the regions where the crop is grown (Plate 41).
Mango trees have evolved a mechanism to cope with temporary ooding (see
Flooding section under Tree Growth and Development, this chapter).
Typically, the rst easily measurable responses of fruit trees to ooding
are reductions in A
max
, g
s
and transpiration, which occur within 23 days fol-
lowing ooding (Larson et al., 1991c; Schaffer et al., 1992, 2006). Short-term
anoxia results in a decrease in net photosynthesis which cannot be related to
a g
s
-associated decrease in C
i
(Zude-Sasse et al., 2001). Removing trees from
ooded conditions after 28 days reversed the ooding-induced decrease in
leaf gas exchange, resulting in a gradual increase in photosynthesis and tran-
spiration to preooded rates.
Although ooding adversely affects mango trees, short-term ooding of
trees in limestone soils can result in increased micronutrient availability with
improved plant nutritional status. In calcareous soils of south Florida, in
which iron (Fe) was withheld from the fertilizer programme, short-term
ooding (1020 days) of polyembryonic Peach mango trees resulted in an
increase in net photosynthetic rates to above preooding levels following the
release of trees from ooding (Larson et al., 1992). This increase in photosyn-
thesis has been correlated with improved Fe and manganese (Mn) uptake as
1 2 3 4 5 6 7 8
0.0
0.1
0.2
0.3
0.4
0.5
0.6
Kensington: R
2
= 0.635
Irwin: R
2
= 0.594
S
t
o
m
a
t
a
l

c
o
n
d
u
c
t
a
n
c
e

(
m
o
l
/
m
2
/
s
)
LAVPD (kPa)
Fig. 6.5. Correlation between leaf stomatal conductance and leaf-to-air vapour
pressure decit (LAVPD) during the dry and wet season for Irwin (closed circles)
and Kensington mango trees (open circles). Trees were well irrigated during the dry
season and all measurements were taken when the Q > 300 mol photons/m
2
/s
(n = 12) (Source: P. Lu, unpublished data).
B. Schaffer et al. 184
a result of these elements becoming more soluble when calcareous soils are
ooded (Larson et al., 1991b, 1992).
Internal factors
Leaf age
Leaf characteristics (i.e. photosynthetic capacity and the amount of N per
unit area) are generally strongly inuenced by leaf age, with maximum val-
ues being observed when leaves have just completed full expansion (Con-
stable and Rawson, 1980; Marshall and Biscoe, 1980; Dwyer and Stewart,
1986; Field, 1987; Wilson et al., 2000; Frak et al., 2001). Chlorophyll content is
three to four times lower in young than in mature mango leaves (Zude and
Ludders, 1997). Similarly, the concentration of Rubisco is lower in young
than in mature, green leaves (Nii et al., 1995). In contrast to many other plant
species, once mango leaves are mature the relationship between N
a
and irra-
diance does not seem to be affected by leaf age (Urban et al., 2003). The N
a

values may remain high in old leaves experiencing high irradiance. This
indicates that changes in N
a
in mango leaves are inuenced by irradiance
and not age, at least during the rst year.
Carbohydrate accumulation and source-sink balance
Source-sink imbalances can exert feedback down-regulation or repression of
leaf photosynthesis through carbohydrate accumulation in leaves (Azcon-
Bieto, 1983; Foyer, 1988; Koch, 1996; Schaffer et al., 1997; Whiley et al., 1999;
Paul and Foyer, 2001; Paul and Pellny, 2003). Transient accumulations of car-
bohydrates in leaves, as they have been observed during the diurnal period,
may impair the rate of electron transport (Pammenter et al., 1993). Changes
in photosynthetic capacity, not just assimilation rates, are more likely to be
observed in association with lasting source-sink imbalances. One hypotheti-
cal mechanism is that high levels of carbohydrates repress the expression of
genes coding for several photosynthetic enzymes (Krapp and Stitt, 1995;
Koch, 1996; Drake et al., 1997). Alternatively, carbohydrates may interact with
hormonal signals to control gene expression (Thomas and Rodriguez, 1994).
There is also some evidence that photosynthetic capacity is related to leaf
carbohydrate status through the effect of the latter on phosphate availability
(Riesmeier et al., 1993; Sun et al., 1999). In the long term, carbohydrate accu-
mulation may eventually lead to cell death. High sugar concentration has
been associated with senescence in leaves of several species (Noodn et al.,
1997; Wingler et al., 1998; Quirino et al., 2001). Reduced energy utilization by
CO
2
assimilation, like the one resulting from carbohydrate accumulation, in
combination with high energy capture is potentially dangerous and can
result in over-reduction of the electron transport chain, photoinhibition and
oxidative stress caused by photoreduction of oxygen to superoxide O
2

in the
Mehler-ascorbate peroxidase reaction (Badger, 1985). Moreover, reactive sin-
glet oxygen
1
O
2
can be formed through reaction of oxygen with triplet chlo-
rophyll released by the breakdown of the chlorophyll-protein complexes in
Ecophysiology 185
thylakoids (Merzlyak and Hendry, 1994). Formation of reactive oxygen spe-
cies can lead to membrane damage and eventually cell death.
Whiley et al. (1999) observed that A
max
, which is closely related to photo-
synthetic capacity, the quantum yield and F
v
/F
mPredawn
are substantially
lower in mango trees grown in containers (root-restricted) when compared
to eld-grown trees. These observations were conrmed by Urban and
Alphonsout (2007) who studied the effect of the removal of a 1 cm-wide band
of bark on leaf photosynthesis and leaf N content of 3-year-old and 11-year-
old Cogshall mango trees. Girdling is a common horticultural practice used
to manipulate tree growth and development in many fruit species. Its most
immediate effect is to stop the basipetal movement of assimilates through the
phloem, which results in an accumulation of carbohydrates above the girdle
(Roper and Williams, 1989; Schaper and Chacko, 1993; Di Vaio et al., 2001).
Girdling can promote oral induction in mango (Chacko, 1991), but it has
also been shown to reduce net photosynthesis. The major effect of girdling is
a dramatic increase in leaf carbohydrate concentration and a concomitant
decrease in photosynthetic electron transport and net photosynthesis (Fig. 6.6)
(Gonzalez and Blaikie, 2003; Urban and Alphonsout, 2007). Urban and
Alphonsout (2007) observed that A
net
was reduced by 77% within 28 days
from girdling and remained at about 2 mol CO
2
/m
2
/s until the beginning
of owering. The decrease in photosynthetic electron transport rate (J) and
sustained photoprotection (reected by the decrease in F
v
/F
mPredawn
) pro-
tected leaves of girdled branches effectively from photodamage, as shown by
the vigorous recovery of A
net
and J observed immediately after the appear-
ance of inorescences. This increase in A
net
and J was associated with no
Q = 2000 mol photons/m
2
/s
y = 155.4e
0.0401x
R
2
= 0.74
0
50
100
150
200
250
0 10 20 30 40
Starch (g/m
2
)
J
(

m
o
l

e
l
e
c
t
r
o
n
s
/
m
2
/
s
)
A
Q = 1200 mol photons/m
2
/s
y = 120.3e
0.0405x
R
2
= 0.74
Q = 400 mol photons/m
2
/s
y = 84.5e
0.0313x
R
2
= 0.69
Fig. 6.6. The relationship between the total photosynthetic electron ux (J) measured
at photosynthetic photon ux (Q) = 400 (), 1200 () and 2000 () mol photons/
m
2
/s, and the amount of starch per unit leaf area. Best t lines at each Q were assessed
from measurements performed on both girdled and non-girdled mango leaves before
owering. Data were used to establish the following relationship: J = (0.0434Q +
72.8)*e
0.0412[starch]a
(Source: Urban and Alphonsout, 2007).
B. Schaffer et al. 186
decrease in leaf carbohydrate content during the rst month following the
onset of owering, suggesting that the effect of carbohydrate accumulation
on photosynthesis is mediated by sink activity. Apart from its negative effect
on the carbon budget of mango trees, girdling appeared to be rather harm-
less. However, leaf N concentration decreased, which indicates that there
may indeed exist long-term negative effects of girdling on photosynthetic
capacity. The width of bark (phloem) removed may be critical with respect to
the intensity of the effect of girdling on the tree. Whiley et al. (2006) girdled
the trunks of B74 (Calypso) mango trees in the Northern Territory of
Australia in autumn (as soon as they had come out of the wet season). The
girdles were no more than the thickness of a pruning saw (about 1 mm) and
healed within 6 weeks. In the rst and third years after girdling, the trees had
signicantly higher yields than non-girdled trees on which, coincidentally,
fruit matured early, thus giving market advantage. There was no signicant
difference in yield in the second year of treatment between girdled and non-
girdled trees. The rst and third years had strong natural induction while the
second year gave poor owering across all varieties in the district. Thus, dur-
ing years of strong induction, this type of girdling most likely provided extra
carbohydrate reserves to drive owering and support fruit set and retention
while in the off-owering year there was sufcient carbohydrate reserves to
support reproductive activity. In contrast to observations with Cogshall
mango trees (Urban and Alphonsout, 2007), there was no evidence of long-term
effects of narrow girdles on leaf N of B74 mango trees (Whiley et al., 2006).
However, when wider girdles are made, tree recovery may take much longer
leading to sustained physiological disruption.
Proximity of inorescences
While the effects of water stress and high light, temperature and atmospheric
CO
2
concentration on photosynthesis are increasingly well described, very
little is known about the effect of phenology, and especially of owering on
photosynthesis of mango. There is some evidence that owering may have
an effect on photosynthesis. Flowering-associated decreases in A
net
and g
s

were observed in sweet cherry (Roper et al., 1988) and mango (Shivashankara
and Mahai, 2000; Urban et al., 2004a). Lack of precise knowledge about the
effect of owering on photosynthesis may impair our ability to adequately
simulate photosynthesis, especially for tropical fruit trees for which ower-
ing often extends over a long period of time. Mango owering can last for >
2 months. Therefore, its effect on photosynthesis should not be overlooked.
Urban et al. (2004a) showed that the decrease in A
net
in mango leaves close to
inorescences is not attributable to a g
s
-associated decrease in C
i
or to an
increase in R
d
. R
d
was lower in leaves close to inorescences than in standard
leaves. If any, the effect of R
d
on A
net
was a positive one. This study suggested
strongly that the decrease in A
net
was due to a decrease in the electron ow in
photosystem II, but failed to provide direct evidence for it as well as the ele-
ments for interpretation. Using a modelling approach, Urban et al. (2008)
conrmed that there is a decrease in the total light-driven photosynthetic
electron ux in leaves close to inorescences and showed that the decrease
Ecophysiology 187
in A
net
is also attributable to an increase in photorespiration. The latter
appears to be the consequence of a g
m
-associated decrease in C
c
, while the
former results from an increase in electron ow towards alternative sinks, a
decrease in the amount of leaf N per unit leaf area, and, hypothetically, either
a decrease in leaf N allocation to the bioenergetic pool of the photosynthetic
machinery, inorganic phosphorus depletion in leaves, or feedback inhibition
of photosynthesis. The latter hypothesis is least probable in the absence of
carbohydrate accumulation in leaves close to inorescences. Both of these
hypotheses need to be tested to further our understanding of the inhibiting
effect of inorescences on photosynthesis of nearby leaves. Interestingly, net
photosynthesis measured on leaves close to panicles bearing set fruits are
intermediary between those measured on standard leaves and those mea-
sured on leaves close to inorescences, suggesting that changes in photosyn-
thesis associated with owering are reversible. Urban et al. (2008) also showed
that processes other than temperature or light acclimation, and acclimation
to reduced sink activity, may cause leaf N concentration and photosynthetic
capacity to vary in mango. Whiley (unpublished data) observed that the
mango leaves immediately adjacent to inorescences lose colour intensity as
the inorescence grows out. Although leaf N over this period was not mea-
sured in mango, it has been measured in avocado (Whiley, 1994) which has a
similar intense burst of owering in which a large biomass is produced in a
short time. Leaf N declines rapidly in avocado leaves as the inorescences
break from buds and grow and then N stabilizes (at a lower concentration)
by mid-bloom. This can be reversed by N applications during owering and
the additional application of a growth retardant (paclobutrazol (PBZ)) giving
leaf N and A a signicant boost. Similarly to avocado, it is likely that the
reduction in A close to mango inoresences is related to reduced leaf N.
Photosynthetic contributions by fruit
Fruit of many species have chlorophyll and photosynthetic activity, particu-
larly during the early stages of growth (Jones, 1981; Whiley et al., 1991). How-
ever, for most crops, respiratory losses from fruit exceed photosynthetic gains
throughout ontogeny (Kriedemann, 1968; Whiley et al., 1991). An exception
to this is blueberry (Vaccinium spp.) fruit in which there is a net photosyn-
thetic gain from petal fall through to colour break, with an estimated 15% of
the total fruit carbon requirement contributed from fruit photosynthesis
(Birkhold et al., 1992). Studies with monoembryonic Dashehari mangoes
showed that when fruit were approximately 10 mm in diameter, the photo-
synthetic rate of fruit was 2.7% that of leaves and declined to 1.2% of leaf
photosynthesis at fruit maturity (Chauhan and Pandey, 1984). However, even
this comparatively small carbon contribution may be important during the
critical fruit set period when trees rely on stored carbohydrates and a rela-
tively inefcient canopy to supply current photosynthates. Further studies
with mangoes to establish optimum light regimes for fruit photosynthesis at
different stages of ontogeny are warranted.
B. Schaffer et al. 188
6.3 Plant Water Relations
In this section, theoretical concepts of plant water relations are briey out-
lined to help interpret the effects of environmental factors on mango water
relations (see also Nobel (1983) and Baker (1984)).
An important concept in plant water relations is water potential (),
which is a measure of the free energy of water. For pure water, = 0. As sol-
utes are added to water, its free energy decreases and becomes more nega-
tive. Water moves along a gradient from higher to lower (more negative) .
can be expressed as:
=

+
p
+
m
(6.11)
where is osmotic (or solute) potential which refers to the effect of solutes
on the change in free energy of water;
p
is the hydrostatic or pressure poten-
tial also referred to as the turgor pressure; and
m
is the matric potential,
which is generally negligible in plant cells.
In plant cells,
p
is generally positive or equal to 0. However, in xylem
tissue of transpiring plants,
p
is negative (under tension). The driving force
for transpiration is the vapour pressure difference between the leaf (consid-
ered to be water saturated) and the surrounding air. Thus, water moves from
a greater to a lower (or more negative)
p
and hence along a decreasing
gradient. The cohesive forces of the H
2
O molecules allows the xylem water to
remain in a continuous column even though there is a negative
p
.
Plant water stress can be determined from Eqn 6.11 and from changes in
. The components of , such as

and
p
, can often be used to dene the
sources of water stress. The drought tolerance of mango highlights some
unique aspects of physiology of this tree with respect to its water manage-
ment. Typical mango environments in the tropics impose extreme water
stress and high evaporative demand for prolonged periods. Adaptive strate-
gies of mango trees include a deep root system (Sukonthasing et al., 1991),
desiccation-tolerant surface feeder roots and drought avoidance mechanisms
thought to be mediated by a comprehensive system of resin canals distrib-
uted throughout the tree (Venning, 1948; Joel, 1980; Joel and Fahn, 1980a, b;
Pongsomboon, 1991) and rapid stomatal closure. Plants with laticfers or resin
ducts/canals have been reported to be drought tolerant due to extended
maintenance of turgor following the withdrawal of water (Downton, 1981;
Kramer, 1983; Kallarackal et al., 1990). While the mechanism of turgor main-
tenance remains unresolved, it is believed that the latex or resin is probably
involved in the modulation of plant water status (Kallarackal et al., 1990).
The differentiation, structure and distribution of resin canals in mango has
been described by Venning (1948), Joel (1980) and Joel and Fahn (1980a, b, c).
Resin canals are present in trunks, shoots, leaves and fruit (exocarp) of mango
in close association with the vascular tissues. The resin contains mainly ter-
penes, but phenols and protein-carbohydrate mucilage are also present (Joel
and Fahn, 1980c). In well-watered trees, the resin is under positive pressure
and freely exudes from damaged or cut surfaces (Pongsomboon, 1991). In
studies of the development of water decit in container-grown Kensington
Ecophysiology 189
mango trees, loss of turgor occurred in expanding leaves when leaf water
potential (
l
) reached 1.2 Mpa. In mature leaves turgor was not lost until
l

reached 1.75 MPa (Pongsomboon, 1991). Necrotic leaf areas appeared when

l
reached approximately 3.2 MPa with permanent wilting developing at
3.45 MPa. This is high (less negative) compared with a
l
of 6.6, 5.0 MPa
for orange (Citrus sinensis) and macadamia (Macadamia integrifolia), respec-
tively (Fereres et al., 1979; Stephenson et al., 1989). Thus, mango leaves toler-
ate less internal water stress than woody perennial fruit trees from more
mesic environments. With mango, however, the permanent wilting point
was reached 36 days after withholding water compared to 10 days for simi-
larly sized macadamia trees (Stephenson et al., 1989). The higher critical
threshold of
l
and the longer period of survival for Kensington mango
indicates that drought tolerance is based on more effective water regulation
to prevent desiccation and on maintenance of leaf turgor rather than resis-
tance by tissues to damage.
Reich and Borchert (1988) observed that stomatal regulation in mango
signicantly reduced the rate of development of internal water decit when
compared with four other tropical tree species. In water-withholding studies,
the radial expansion of mango trunks continued when most of the other
species were shrinking, indicating that mango trees could better tolerate
drought conditions and maintain photoassimilation rates. This is consistent
with the decrease in g
s
/A
net
observed by Urban et al. (2006) as a consequence
of drought (Fig. 6.1). A decrease in g
s
/A
net
indicates that there is an increase
in photosynthetic water use efciency.
In containerized Kensington mango trees, there was a linear correlation
between stomatal conductance and
l
during the development of water
stress (Pongsomboon, 1991). In contrast, with avocado and macadamia,
stomatal response was much more rapid with a curvilinear relationship
between
l
and stomatal conductance, and stomatal closure reached at 1.2
and 3.0 MPa, respectively. The slower response of stomatal conductance to

l
in

mango trees appears to be related to a more effective mechanism for
the mediation of water decit development compared with avocado and
macadamia.
Pongsomboon (1991) monitored leaf water potential, osmotic potential of
resin (
r
) and osmotic potential of the whole leaf tissue (

) in container-
grown Kensington mango trees when water was withheld for a 45-day
period. When tissues were fully hydrated,
l
and
r
were higher than

. For
40 days of the drying cycle,
l
and
r
declined at a similar rate; however,


declined to about 1.2 MPa within 4 days where it remained stable until 18
days into the drying cycle. There was a subsequent decline in

for 28 days
after withholding water when it stabilized at 2.0 MPa, remaining constant
for another 12 days. Pongsomboon (1991) suggested that osmotic adjustment
occurred, probably mediated through the resin as water decit developed. It
appears, therefore, that the energy investment by the tree in a resin canal sys-
tem is justied by the vital drought-avoidance benets conferred by main-
taining turgor and preventing wilting under prolonged periods of water
stress. Further investigations are required to substantiate these conclusions.
B. Schaffer et al. 190
In a recent experiment of 2-year-old potted Cogshall mango trees
grafted on Maison Rouge rootstock, midday minimum leaf water potential
remained relatively constant, about 1.0 MPa, for the range of predawn leaf
water potential (a surrogate of soil water potential) from 0 to 0.5 MPa, but
when the predawn leaf water potential dropped below 0.5 MPa, leaf water
potential fell rapidly to about 1.8 MPa (Gaelle Damour, Centre de coopra-
tion Internationale en Recherche Agronomique pour le Dveloppement
(CIRAD), personal communication). This isohydric behaviour seemed to be
associated with rapid stomatal closure in response to decrease in water avail-
ability (Urban and Jannoyer, 2004).
Development of novel techniques to study xylem integrity have pro-
vided some unique insights into stomatal function (Tyree and Sperry, 1988;
Cochard et al., 2002). Xylem sap is transported under negative pressures in
plants and therefore is susceptible to cavitation events that render xylem
conduits non-conductive. Cavitation occurs when the negative sap pressure
exceeds a threshold value dened by anatomical characteristics (Tyree and
Sperry, 1988). Many species function very close to the point of embolism.
Therefore, stomata control both plant water losses and sap pressure and thus
may actively control the risk of xylem embolism (Jones and Sutherland,
1991). Xylem vulnerability to cavitation was studied in twigs of 13-year-old
Cogshall mango trees. Xylem vessels started to cavitate at a xylem water
potential of about of 1.5 MPa and underwent a rapid and substantial loss
(90%) of hydraulic conductivity when xylem water potential dropped from
2.0 to 3.0 MPa (Fig. 6.7) (H. Cochard, unpublished data). This result is con-
sistent with the observation that leaves lose turgor when
l
reaches 1.75
MPa and permanent wilting appears when
l
is above 3.0 MPa (Pongsom-
boon, 1991) when Fig. 6.7 predicts 90% of xylem hydraulic conductivity is
lost. It also conrms that mango, like many other trees, operates close to the
point of embolism via effective stomatal control.
Direct measurement of xylem water potential using a pressure chamber
is problematic due to the presence of latex (Castro Neto et al., 2004). It is more
problematic in some cultivars, for example Kensington, than in others, such
as Irwin. Alternative indicators of plant water status such as measurements
of whole-tree water use (sap ow) and stem/branch shrinkage were found to
be sensitive and integrated indicators of whole-tree water status in mango
and were successfully used to schedule irrigation in a mango orchard in the
seasonal dry-wet tropical region of northern Australia (Lu, 2002, 2006).
6.4 Tree Growth and Development
Light
In an orchard, light distribution within and between tree canopies can have
a profound effect on growth and development of the fruit. We have previ-
ously discussed the effect of light on photosynthesis and dened the opti-
mum light levels required for mango leaves. When light levels fall below the
Ecophysiology 191
threshold required for light saturation of photosynthesis, the subsequent
reduction in available photoassimilates will affect growth of the tree. In many
tree fruit crops, ower-bud induction, fruit size and fruit colour are reduced
when low light levels occur due to crowding within and between tree cano-
pies (Jackson, 1980; Flore, 1994; Whiley and Schaffer, 1994). There is no pub-
lished information on the effect of light levels on mango fruit size, although
fruit are photosynthetically active and a reduction in size under low Q could
be expected.
Fruit skin colour is an important feature of mango with fruit of many
cultivars developing attractive pink to red coloration. Fruit colour is geneti-
cally determined and the reddish blush is generally more developed in
monoembryonic cultivars, while fruit from most polyembryonic cultivars
remain green/yellow at maturity. Skin coloration of mature fruit is partly
due to anthocyanins which develop when tissues are exposed to light. While
this subject is well researched in other fruit crops (Proctor and Creasey, 1971),
light levels required for skin coloration of mango fruit have not been quanti-
ed. Studies in Australia with the polyembryonic cultivar Kensington,
which develops a blush only on the exposed side of the fruit, indicated that
the position of fruit on trees had a signicant effect on the development of
colour due to differences in the penetration of light into the canopy during
fruit ontogeny (Schaffer et al., 1994). The intensity of redness was greatest on
fruit from the eastern side of the tree followed by fruit from the south-western
100
80
60
40
P
L
C
20
0
4 3 2
Xylem pressure (MPa)
1 0
2006 light
2006 shade
2005
Fig. 6.7. Vulnerability of xylem of 13-year-old Cogshall mango trees to cavitation
in twig segments from sun-exposed (light) or shaded sides of the tree. Cavitation is
expressed as percentage loss of conductivity (PLC) with decreasing xylem water
potential. Symbols represent means and error bars represent one standard error.
Vulnerability curves were obtained with the centrifuge technique (Source: Cochard
et al, 2005; and from H. Cochard, unpublished data, with permission).
B. Schaffer et al. 192
and northern sides of the tree. This information establishes an important con-
cept with respect to the light regime but does not quantify the absolute light
levels required for anthocyanin development. Further research is necessary
to establish physiological parameters from which pruning and orchard man-
agement strategies can be developed.
Temperature
Mango is a predominantly tropical species although the tree will usually
grow and produce more successfully in frost-free subtropical latitudes with
a marked dry season and high heat accumulation. Under optimum tempera-
tures with non-limiting nutrients and water, the tree remains vegetative with
growth ushes occurring at regular intervals. The large size and poor crop-
ping of trees in the humid lowland tropics are well known, and there is a
direct relationship between temperature and the frequency of vegetative
ushes. Trees grown at 20C days/15C nights (20/15C) required 20 weeks
(mean of ten cultivars) to complete a growth/dormancy cycle while at
30/25C the same cycle was completed in 6 weeks (Whiley et al., 1989). There
are marked differences between cultivars with respect to their tendency
towards vegetative growth. For instance, in controlled temperature studies
over a 20-week period, at 30/25C, Irwin produced 2.0 growth ushes with
approximately 45 days of dormancy between active growth periods while
Kensington produced 4.7 growth ushes with only 5 days of quiescence
between ushes (Whiley et al., 1989). Dry matter accumulation over the 20
weeks was similar for the two cultivars; however, starch accumulation in the
woody trunk tissues of Irwin and Kensington was 13 and 3.6% of dry mat-
ter, respectively. The response differences between these two cultivars may
be the contributing factor to their performance at tropical latitudes where
temperature is non-limiting for growth. Under these conditions Irwin has
more reliable cropping than Kensington, suggesting that the genetically
determined low-vigour trait is more sensitive to environmentally precipi-
tated stresses that induce owering.
The number and size of leaves which develop on each growth ush are
also inuenced by temperature. Whiley et al. (1989) reported that on trees
growing at 20/15C an average of 7.1 leaves per ush were produced while
at 30/25C there were 13.6 leaves on each growth ush (data are mean values
from ten cultivars). At 30/25C the mean leaf size was 300% greater than
those on trees growing at 20/15C. Soil temperatures have also been reported
to have a strong effect on the growth of mangoes. In studies with Irwin
grafted on Turpentine rootstocks, episodic shoot growth occurred when
soil temperatures were held at 27C or 32C for 120 days but an extended
dormant period developed when soil temperatures were held at 21C (Yusof
et al., 1969). These results indicate that environmental control over shoot
growth of mangoes may in part be related to soil temperatures.
From controlled temperature studies it has been calculated that the median
daily temperature (mean of the maximum and minimum daily temperatures)
Ecophysiology 193
at which shoot growth ceases is approximately 15C (mean value for ten cul-
tivars) (Whiley et al., 1989). Subsequent studies (Issarakraisila et al., 1991)
have conrmed that 15C is the critical minimum growth temperature for
shoots of Kensington.
Stress-inducing temperatures which prevent shoot growth have been
shown to promote oral induction in mangoes, but this is outside the scope
of this discussion. For further information of the effects of temperature on
pollination, oral initiation and fruit development, see Davenport, Chapter
5, this volume and Schaffer et al. (1994). We again emphasize that although
mango is a heat-loving crop well adapted to the hot, semi-arid subtropics
and monsoonal tropics, in these environments it experiences extremes of heat,
drought and evaporative demand that may cumulatively reduce potential
production capacity.
Drought
Although mango is considered to be drought tolerant and may survive with-
out rain or irrigation for > 8 months (Gandhi, 1955), water decits during the
reproductive cycle can have severe effects on the retention and early growth
of mango fruit. In studies with bearing, container-grown Irwin trees, pre-
dawn
l
levels were maintained at either less than 0.3 MPa (non-stressed)
or 1.2 MPa (water stressed) for the rst 2 months after fruit set. For the rst
5 days following fruit set, all trees lost a similar percentage of fruit, but there-
after fruit abscission was greater on water-stressed trees. After 1 month,
drought-stressed trees had retained approximately 4% of their initial fruit set
compared with approximately 8% on non-stressed trees. During the rst 30
days following fruit set, the rate of fruit growth for non-stressed trees was
twice that of drought-stressed trees, and nal fruit size (measured 60 days
after fruit set) of non-stressed trees was 20% greater than on water-stressed
trees. In a separate study, another group of Irwin trees was maintained
stress-free (pre-dawn water potential above 0.3 MPa) during the rst month
following fruit set with water-stress (1.2 MPa) imposed on some trees dur-
ing the second month of fruit development (Pongsomboon, 1991). There was
no effect of drought on fruit retention but nal fruit size was 34% smaller for
stressed compared with non-stressed trees.
In eld studies, Singh and Arora (1965) compared fruit drop of monoem-
bryonic Dashehari trees irrigated at 1-week intervals with trees irrigated at
3-week intervals. During the rst 6 weeks of fruit growth, weekly irrigation
reduced fruit drop compared with the irrigation at 3-week intervals. During
the latter stages of fruit development, these gains were lost as more fruit
dropped from the weekly irrigated trees. In another study, eld-grown
monoembryonic Tommy Atkins trees were managed under different irriga-
tion regimes from early fruit set until the start of the rainy season (approxi-
mately 43 days) (Larson et al., 1989). Trees were irrigated on a 7- or 14-day
schedule or received no irrigation. Pre-dawn
l
was 0.3 MPa for trees irri-
gated on the 7-day schedule and decreased to 0.5 MPa for the non-irrigated
B. Schaffer et al. 194
trees. Irrigation at 7-day intervals resulted in the greatest yield with the larg-
est fruit, especially during the early harvest period (Larson et al., 1989).
Irrigation, therefore, particularly during the rst 46 weeks following
fruit set, increases individual fruit size and yield. This is a critical period of
fruit development since it is when cell division is most rapid and cell walls
are developed. Even slight reductions in plant water status during this period
can have adverse effects on fruit growth and retention (Pongsomboon, 1991).
Although drought tolerance of the mango tree is well known, this comes at
considerable cost to tree performance, particularly in areas with prolonged
dry seasons that extend through owering and fruiting. Irrigation is there-
fore one of the most powerful tools to alleviate non-lethal yet potentially
yield-reducing drought stress.
Flooding
Studies with container-grown mango trees have reported variable responses
with respect to tree survival. Larson et al. (1991c) observed that as many as
45% of trees died after their roots were submerged in water for 410 days, but
in the surviving population no further mortality occurred when ooding
was extended for up to 110 days. In other experiments, there was no tree
mortality after container-grown mango trees were ooded from 1 to several
months although tree growth was signicantly reduced (Larson et al., 1991c;
B. Schaffer unpublished data, Homestead, Florida, 1993).
The ability of mango trees to survive prolonged ooding appears to be
dependent on the development of hypertrophic (swollen) stem lenticels
immediately above the water line (Plate 42). The initial stages of lenticel
hypertrophy are characterized by the development of intercellular spaces in
the phellem tissue and production of additional phellem tissue by increased
phellogen activity. Later stages of hypertrophy are characterized by the
development of intercellular spaces in the phellem tissue and cortex (Larson
et al., 1991a). Observations vary among studies whether or not trees devel-
oped hypertrophic lenticels or how quickly after ooding they formed. These
anomalies have been attributed to environmental differences at the time of
root submersion (Larson et al., 1991c). In trees that died as a result of ooding
stress there was no lenticel hypertrophy; however, stem lenticels hypertro-
phied within 410 days on mango trees that survived ooding (Larson et al.,
1991a, c, 1993). Sealing hypertrophic lenticels of mango trees with silicone
grease or petroleum jelly resulted in tree death within 3 days of ooding,
thereby demonstrating their necessity for tree survival. The role of hypertro-
phic lenticels in ood-tolerant species is not clear, although they are thought
to eliminate potentially toxic metabolites such as ethanol, acetaldehyde and
ethylene which result from anaerobic respiration in the roots (Chirkova and
Gutman, 1972; Larson et al., 1993). They may also confer ood tolerance by
enhancing O
2
diffusion to the roots (Kozlowski, 1984).
In some instances, adventitious roots have developed above the water
line when container-grown mango trees have been ooded for long periods
Ecophysiology 195
(Schaffer et al., 1994). It is likely that these roots facilitate the absorption and
translocation of O
2
to submerged roots and their development is a common
morphological response of many woody plants to root anoxia. The develop-
ment of adventitious roots has not been reported for ooded, eld-grown
trees and they may only form on young trunks after extended ooding peri-
ods, which usually do not occur under normal production conditions.
Vegetative growth of mango trees generally declines when trees become
ooded for > 23 days. When trees in a limestone soil in containers were
ooded for > 110 days, there was a 94% reduction in shoot extension growth,
while ooding for approximately 10 days resulted in a 57% reduction in
shoot extension growth (Larson et al., 1991c). In a subsequent study, the stem
radial growth (a more sensitive indicator of tree growth than shoot extension
growth) of mango trees decreased 2 weeks after roots were submerged.
Flooding for > 14 days also signicantly reduced root dry weight, resulting
in an increased shoot to root ratio (Larson et al., 1991c). These adverse effects
of ooding on the growth of mango trees are expected as reduced net photo-
synthesis and presumably higher root respiration limit the availability of
carbon-based assimilates required for growth.
Wind
Most fruit trees benet from wind protection, particularly during the estab-
lishment years when the disruption of physiological processes results in a
signicant depression of growth in young trees. In addition, wind also causes
abrasions to the skin of fruits, particularly when they are small, which
develop into unsightly blemishes by the time they are fully grown thereby
reducing quality and market value. However, the cost of windbreaks may
not be offset by higher returns. In some mango-producing regions, winds are
not sufciently strong to justify the cost of wind protection. Until recently,
wind protection in South Africa was not recommended for mangoes due to
the loss of potential cropping space by living windbreaks, their potential to
create frost pockets, and the likelihood of promoting the incidence of ower
and fruit diseases through increased humidity (Van der Meulen et al., 1971);
however, the value of windbreaks is well appreciated today in South Africa
(B.N Wolstenholme, personal communication, Pietermaritzburg, South
Africa, 1995).
In studies with Kensington mangoes in Australia, where articial wind-
breaks were constructed to shelter trees from the prevailing summer south-
easterly winds, a 600% increase in yield was recorded in the rst year
following wind protection (Mayers et al., 1984). This signicant improvement
in tree performance was a result of better growth of trees which set and held
more fruit per panicle, suffered less damage to leaves (cuticle fracturing) and
had reduced fruit loss from bacterial black spot (caused by Xanthomonas camp-
estris pv. mangiferaeindicae) compared to the wind-exposed trees. These results
indicate that wind can have a signicant effect on mango productivity from
the reduction of both growth and yield through undisclosed physiological
B. Schaffer et al. 196
mechanisms, and a decreased level of bacterial black spot infection. The pro-
vision of windbreaks in orchards is expensive with decisions to be made on
the use of either living or articial shelters. Requirements for wind protec-
tion will vary depending on site circumstances, and all factors pertaining to
crop performance will require careful consideration.
Salinity
Salt stress in mango trees produces symptoms similar to those described for
other species (Harding et al., 1958; Ehlig, 1960; Kadman, 1964; Bingham et al.,
1968). Mild symptoms of chloride toxicity are scorched leaf tips and margins
and leaf curling, while in more severe cases growth ceases, leaves abscise and
trees die. Necrotic areas develop on leaves of trees exposed to high sodium
levels (Jindal et al., 1976; Kadman et al., 1976; Gazit and Kadman, 1980). Irri-
gation water concentrations of 2060 mM sodium chloride (NaCl) or sodium
sulfate (Na
2
SO
4
) reduced leaf area and changed the branching structure of
container-grown mango trees, suggesting that salinity resulted in reduced
leaf cell elongation, and affected the activity of the terminal meristem
(Schmutz and Ldders, 1993). As the duration of the exposure to saline con-
ditions increased, transpiration decreased exponentially (Schmutz and Ld-
ders, 1993). In a later study, Schmutz (2000) found that following a gradual
increase in salinity of the nutrient solution applied to potted polyembryonic
13-1 rootstocks (from 0 to 120 mM NaCl over 15 days), A
max
signicantly
declined despite there being no visible leaf symptoms of salt toxicity. The
decline in A
max
occurred within 6 days of beginning the salinity treatment,
which was 15 mM NaCl for 3 days followed by 30 mM NaCl for 3 days. This
indicates that photoassimilation in mango is quite sensitive to exposure of
trees to salinity.
There is considerable variation in salinity stress of mango, both within
and between populations of mono- or polyembryonic mango ecotypes. Based
on the results of limited studies, there appears to be greater salt tolerance in
polyembryonic than in monoembryonic populations (Jindal et al., 1975; Kad-
man et al., 1976). In seedling populations from mono- and polyembryonic
cultivars irrigated for 2 years with water containing approximately 10 mM
chloride, most plants developed leaf scorching after 6 months which gradu-
ally became more severe, culminating in degeneration and death. However,
some seedlings which had no damage or only slight toxicity symptoms were
mostly of the polyembryonic 13-1 rootstock cutivar or related types (Kad-
man et al., 1976). Leaf analyses revealed that the chloride concentration in
tolerant seedlings (0.680.77%) was greater than in susceptible seedlings
(0.430.55%). In addition, tolerant plants had lower leaf concentrations of
potassium, calcium and magnesium than saline-sensitive seedlings, possibly
a result of comparative nutrient dilution since vegetative growth was greater
for saline-tolerant than for saline-sensitive seedlings. Kadman et al. (1976)
also suggested that the mechanism of chloride tolerance in 13-1 was based
on greater physiological tolerance of chloride concentrations in leaf tissues,
Ecophysiology 197
rather than ion exclusion or a selective uptake mechanism common in other
species (Collander, 1941; Walker, 1986). However, the relative sodium toler-
ance of 13-1 was due to exclusion of sodium from shoots and its accumula-
tion in root cell vacuoles (Schmutz and Ldder, 1993). More recently Hoult
et al. (1996) reported signicant cultivar differences within a population of 21
polyembryonic mango cultivars exposed to saline (480 mg/l NaCl) irrigation
water for 10 months. Differences were measured in leaf Na (0.371.34%) and
Cl (0.391.07%) concentrations but these were poorly correlated with toxicity
symptoms on leaves.
Salinization of agricultural land is increasing and in many areas salinity
management is critical. There appears to be sufcient genetic diversity within
Mangifera indica to enable the selection and development of saline-tolerant
rootstocks (Kadman et al., 1976; Gazit and Kadman, 1980; Hoult et al., 1996).
However, quantitative data on the critical limits of soil and water salinity
which mango trees will tolerate without reductions in yield and fruit quality
are needed.
Elevated atmospheric CO
2
concentration
Growing Kensington mango trees for 6 months in a controlled atmosphere
glasshouse with an ambient CO
2
concentration of 600 mol/mol resulted in
more dry matter partitioned to the roots compared to plants grown in an
ambient CO
2
environment of 350 mol/mol (Schaffer et al., 1999) (Fig. 6.8).
Fruit dry weight was greater for mango trees grown in an atmospheric CO
2

concentration of 600 mol/mol compared to trees grown at 350 mol/mol
(Schaffer et al., 1999) (Fig. 6.9). Most of the increased total fruit dry weight at
Old
leaves
Old
branches
New
leaves
Plant part
New
branches
Trunk Roots
0
100
200
300
400
500
D
r
y

w
e
i
g
h
t

(
g
)
600 mol CO
2
/mol
350 mol CO
2
/mol
Fig. 6.8. Partitioning of dry matter in Kensington mango trees grown for 6 months in
atmospheric CO
2
concentrations of 350 or 600 mol/mol. Bars represent means (n = 6
trees) standard error (Source: redrawn from Schaffer et al., 1999).
B. Schaffer et al. 198
the higher atmospheric CO
2
concentration was a result of increased amount
of pulp; whereas, there were no signicant effects of increased atmospheric
CO
2
concentration on dry matter accumulation in the skin, testa or seed
(Schaffer et al., 1999) (Fig. 6.9). Thus, increasing the atmospheric CO
2
assimi-
lation rate increased growth and partitioning to the mesocarp. Therefore,
increased atmospheric CO
2
concentration (at least to 600 mol/mol) as a
result of global climate change may increase the economic yield of mango
(Schaffer et al., 1997). However, under actual environmental conditions result-
ing from global climate change, water and nutrient availability may be limit-
ing and is likely to offset any increases in biomass or economic crop yield
resulting from elevated ambient CO
2
concentrations (Gitay et al., 2001).
6.5 Crop Production
Temperature limitations to crop production
Temperature is probably the most important environmental variable to con-
sider when selecting mango cultivars for particular sites. The mean tempera-
ture range for optimum growth of mango is about 2430C (Mukherjee, 1953;
Whiley et al., 1989). However, mango trees can tolerate temperatures up to
48C for short periods (Mukherjee, 1953). Mango trees have limited tolerance
to cold and trees are usually severely damaged or killed after a few hours at
temperatures < 0C (Carmichael, 1958; Campbell et al., 1977). Although
mature trees have withstood temperatures of 4C for a few hours with lim-
ited damage, juvenile trees were killed after 13 h at 4C to 6C (Campbell et al.,
1977).
Fruit tissues
Skin Flesh Testa Seed Total fruit
D
r
y

w
e
i
g
h
t

(
g
)
0
10
20
30
40
50
600 mol CO
2
/mol
350 mol CO
2
/mol
Fig. 6.9. Partitioning of dry matter in Kensington fruit of trees grown for 6 months
in atmospheric CO
2
concentrations of 350 or 500 mol/mol. Bars represent means
(n = 3349 fruit) standard error (Source: redrawn from Schaffer et al., 1999).
Ecophysiology 199
Monoembryonic mango cultivars tend to be more cold tolerant than
polyembryonic cultivars, probably due their probable subtropical origin.
There are also differences in fruit setting capacity between mono- and poly-
embryonic cultivars at subtropical latitudes, where monoembryonic culti-
vars crop more successfully when minimum temperatures fall below 12C
during owering (Searle et al., 1995). Differences in low temperature toler-
ance have important implications for the selection of suitable cultivars for
production under specic climatic conditions.
Light interception and orchard design
Light interception and utilization within tree canopies is a primary consider-
ation in orchard design. Thus, tree spacing as well as pruning practices in
orchards are primarily based on maximizing light for photosynthesis (see
previous section on Light under Photosynthesis, this chapter). The follow-
ing equation describes the relationship between light interception, photosyn-
thesis and yield in fruit tree orchards:
Biological Yield = (Light Available) (% Light Intercepted) (Photosynthe-
sis) Respiration (6.12)
where biological yield refers to dry matter production, including that of fruit
(Lakso, 1994). The amount of light available is a function of climate and can-
not be manipulated, and potential net photosynthetic efciency of a crop is
inherent and cannot be altered without genetic manipulation. Thus, optimiz-
ing biological yield is based on maximizing the percentage of light inter-
cepted by the orchard canopy and minimizing stress so that photosynthetic
potential is not compromised. With respect to light, the mango tree presents
special problems due to long-lived leaves, dense canopies, and the potential
for vigorous growth in some cultivars.
Maximizing light interception by the photosynthetic surface of an orchard
is a function of tree spacing, canopy density and height. For some fruit tree
trees, for example apple (Malus domestica) and citrus (Citrus spp.), the use of
low vigour rootstocks and pruning provides opportunities for an array of
management options. However, the lack of dwarng rootstocks and the com-
plications in pruning due to the oral morphology of mango (inorescences
are predominantly borne terminally on the most recently produced shoots)
limit the efcient harvest of light with respect to maintenance of productiv-
ity. Improved productivity can be obtained by grafting, which either short-
ens the juvenility phase and/or exerts some control over vigour. Compared
to temperate fruit orchards, canopies of commercial mango orchards have a
higher proportion of shade to sun leaves (Schaffer et al., 1994). Mango trees
are rarely selectively pruned (Young and Sauls, 1981), although novel ideas
of heading back cuts after harvest have been investigated (Oosthuyse, 1992).
In some areas, trees are mechanically topped and hedged to control tree size.
Hedging encourages increased complexity (ramication), resulting in a dense
outer canopy. Where costs are not prohibitive, strategically timed, selective
B. Schaffer et al. 200
pruning to increase the percentage of leaves exposed to > 60% of full sunlight
will increase the photosynthetic efciency of the canopy with a potential
improvement in yield. Schaffer and Gaye (1989) increased light interception
of mango by removing 25% of the canopy. This resulted in higher chlorophyll
concentrations in leaves of pruned canopies later in the year. Although net
photosynthesis was not measured in that study, it is likely that the higher leaf
chlorophyll concentrations in the pruned canopies resulted in higher photo-
synthetic rates.
Light utilization of mango can be enhanced by pruning, but the timing of
such treatments is critical. For example, in the subtropics, shoots produced
following harvest generally ower 35 months after being exposed to induc-
tive (cool) temperatures. Therefore, trees should be pruned immediately after
harvest to improve light penetration and contain tree size. However, in the
tropics there is a shorter period between the cessation of summer growth and
owering and summer-grown shoots of many cultivars fail to induct that
year (Scholeeld et al., 1986; see Davenport, Chapter 5, this volume). There-
fore, due to the removal of potential owering sites, summer pruning of
mango trees in the tropics generally reduces yield in the following season.
Growth control of mango trees through the development of dwarng
rootstocks, low-vigour cultivars and pruning strategies to increase light pen-
etration within tree canopies and by entire orchards, may improve yield per-
formance of this crop. Improved information on light interception, critical
leaf to fruit ratios and relationships between shoot maturity and oral induc-
tion with respect to genotype/environmental interactions will enhance the
development of improved cultural practices for mango production. It is per-
tinent to emphasize that few evergreen fruit trees are as precocious as mango,
or as large at maturity. Orchardists should take advantage of the precocity
and of light interception principles by initial high-density planting, with
hedgerows perhaps the best option. As trees become crowded, however, ef-
ciency of light interception is compromised and remedial action before this
occurs is required to maintain productivity.
6.6 Conclusions
Although much literature in the past stressed problems associated with low
temperature on mango growth and production, sustained high temperatures
with associated soil moisture stress during fruit ontogeny and high evapora-
tive demand are perhaps the major reasons for relatively low yields in mango
orchards worldwide, especially in the tropics. Although mango trees have a
number of survival mechanisms that allow them to cope with stressful envi-
ronments, these come at a considerable energy cost thereby potentially reduc-
ing the availability of carbon-based inputs for fruiting. It is likely that annual
assimilation gains and resource availability during critical developmental
periods are inadequate for sustained high yields of quality fruit. These prob-
lems can be alleviated by development of improved germplasm with adapta-
tion to specic environments (Whiley et al., 2006). In the future, greater effort
Ecophysiology 201
is required for rootstock development and understanding the manipulative
effect on whole tree physiology. Expansion of human populations and cli-
mate change scenarios predict lesser and poorer quality water available for
cropping systems across tropical and subtropical latitudes. Hence, greater
salinity tolerance within the species should be identied. Knowledge of
mango physiology, particularly in relation to the trees responses to varying
environmental conditions remains basic and must be expanded. A greater
understanding of these principles together with their application will assist
in the development of more productive cropping systems.
References
Adams, W.W. III, Zarter, C.R., Mueh, K.E., Amiard, V.S.E. and Demmig-Adams, B. (2005)
Energy dissipation and photoinhibition: a continuum of photoprotection. In: Demming-
Adams, B., Adams, W.W. III and Mattoo, A.K. (eds) Photoprotection, Photoinhibi-
tion, Gene Regulation and Environment. Springer Verlag, Berlin, pp. 4964.
Atkin, O.L.K., Millar, A.H., Grdestrom, P. and Day, B. (2000) Photosynthesis, carbohy-
drate metabolism and respiration in leaves of higher plants. In: Leegoood, R.C.,
Sharkey, T.D. and von Caemmerer, S. (eds) Photosynthesis, Physiology and Metabo-
lism. Kluwer Academic, London, pp. 153175.
Azcon-Bieto, J. (1983) Inhibition of photosynthesis by carbohydrates in wheat leaves.
Plant Physiology 73, 681686.
Badger, M.R. (1985) Photosynthetic oxygen exchange. Annual Review of Plant Physiology
36, 2753.
Baker, D.A. (1984) Water relations. In: Wilkins, M.G. (ed.) Advanced Plant Physiology.
Pitman, London, pp. 297316.
Bauerle, W.L., Weston, D.J., Bowden, J.D., Dudley, J.B. and Toler, B. (2004) Leaf absorp-
tance of photosynthetically active radiation in relation to chlorophyll meter estimates
among woody species. Scientia Horticulturae 10, 169178.
Bingham, F.T., Fenn, L.B. and Oertli, J.J. (1968) A sand culture study of chloride toxicity
to mature avocado trees. Proceedings of the Soil Science Society of America 32,
249252.
Birkhold, K.T., Koch, K.E. and Darnell, R.L. (1992) Carbon and nitrogen economy of
developing rabbiteye blueberry fruit. Journal of the American Society for Horticul-
tural Science 117, 139145.
Campbell, C.W. (1984) Tropical fruits and nuts. In: Martin, F.W. (ed.) CRC Handbook of
Tropical Food Crops. CRC Press, Boca Raton, Florida, pp. 235274.
Campbell, C.W., Knight, R.J. and Zareski, N.L. (1977) Freeze damage to tropical fruits in
southern Florida in 1977. Proceedings of the Florida State Horticultural Society 90,
254257.
Carmichael, W.W. (1958) Observations of cold damage to mangos in Dade County and
the lower west coast. Proceedings of the Florida State Horticultural Society 71,
333335.
Carswell, F.E., Meir, P., Wandell, E.V., Bonates, L.C.M., Kruijt, B., Barbosa, E.M., Nobre,
A.D., Grace, J. and Jarvis, P.G. (2000) Photosynthetic capacity in a central Amazo-
nian rain forest. Tree Physiology 20, 179186.
Castro Neto, M.T., Reinhardt, D.H. and da S.Ledo, C.A. (2004) Determination of water
potential on mango trees by pressure chamber. Acta Horticulturae 645, 425427.
Chacko, E.K. (1991) Mango owering still an enigma. Acta Horticulturae 291, 1221.
B. Schaffer et al. 202
Chauhan, P.S. and Pandey, R.M. (1984) Relative
14
CO
2
xation by leaves and fruits, and
translocation of
14
C-sucrose in mango. Scientia Horticulturae 22, 121128.
Chirkova, T.V. and Gutman, T.S. (1972) Physiological role of branch lenticels in willow and
poplar under conditions of root anaerobiosis. Soviet Plant Physiology 19, 289295.
Cochard, H., Coll, L., Le Roux, X. and Amglio, T. (2002) Unraveling the effects of plant
hydraulics on stomatal conductance during water stress in walnut. Plant Physiology
128, 282290.
Cochard, H., Damour, G., Bodet, C., Tharwat, I., Poirier, M. and Amglio, T. (2005)
Evaluation of a new centrifuge technique for rapid generation of xylem vulnerabil-
ity curves. Physiologia Plantarum 124, 410418.
Collander, R. (1941) Selective absorption of cations by higher plants. Plant Physiology
16, 691720.
Constable, G.A. and Rawson, H.M. (1980) Effect of leaf position, expansion and age on
photosynthesis, transpiration and water use efciency of cotton. Australian Journal
of Plant Physiology 7, 89100.
Crane, J.H. and Davies, F.S. (1988) Periodic and seasonal ooding effects on survival,
growth, and stomatal conductance of young rabbiteye blueberry plants. Journal of
the American Society for Horticultural Science 113, 488493.
DeJong, T.M. and Doyle, J.F. (1985) Seasonal relationships between leaf nitrogen con-
tent (photosynthetic capacity) and leaf canopy light exposure in peach (Prunus per-
sica). Plant Cell and Environment 8, 701706.
Di Vaio, C., Petito, A. and Buccheri, M. (2001) Effect of girdling on gas exchanges and
leaf mineral content in the Independence nectarine. Journal of Plant Nutrition 24,
10471060.
Downton, W.J.S. (1981) Water relations in Nerium oleander. Australian Journal of Plant
Physiology 8, 329334.
Drake, B.G., Gonzales-Meler, M. and Long, S.P. (1997) More efcient plants: a conse-
quence of rising atmospheric CO
2
? Annual Review of Plant Physiology and Plant
Molecular Biology 48, 609639.
Dreyer, E., Le Roux, X., Montpied, P., Daudet, F.A. and Masson, F. (2001) Temperature
response of leaf photosynthetic capacity in seedlings from seven temperate species.
Tree Physiology 21, 223232.
Dwyer, L. and Stewart, D. (1986) Effect of leaf age and position on net photosynthetic
rates in maize (Zea mays L.). Agricultural and Forest Meteorology 37, 2946.
Ehlig, C.F. (1960) Effects of salinity on four varieties of table grapes grown in sand cul-
ture. Proceedings of the American Society for Horticultural Science 76, 323331.
Epron, D., Godard, C., Cornic, G. and Genty, B. (1995) Limitation of net CO
2
assimila-
tion rate by internal resistances to CO
2
transfer in the leaves of two tree species
(Fagus sylvatica L. and Castanea sativa Mill.). Plant Cell and Environment 18, 4351.
Ethier, G.J. and Livingston, N.J. (2004) On the need to incorporate sensitivity to CO
2
transfer conductance into the Farquhar-von Caemmerer-Berry leaf photosynthesis
model. Plant Cell and Environment 27, 137153.
Evans, J.R. (1989) Photosynthesis and nitrogen relationships in leaves of C3 plants.
Oecologia 78, 819.
Farquhar, G.D., von Caemmerer, S. and Berry, J.A. (1980) A biochemical model of pho-
tosynthetic CO
2
assimilation in leaves of C3 species. Planta 149, 7890.
Fereres, E., Cruz-Romero, G., Hoffmann, G.J. and Rawlins, S.L. (1979) Recovery of or-
ange trees following severe water stress. Journal of Applied Ecology 16, 833842.
Field, C.B. (1987) Leaf-age effects on stomatal conductance. In: Zeiger, E., Farquhar, G.D.
and Cowan, I. (eds) Stomatal Function. Stanford University Press, Palo Alto, Califor-
nia, pp. 367384.
Ecophysiology 203
Field, C.B. and Mooney, H. (1983) Leaf age and seasonal effects on light, water, and
nitrogen use efciency in a California shrub. Oecologia 56, 348355.
Field, C.B. and Mooney, H. (1986) The photosynthesis-nitrogen relationship in wild
plants. In: Givnish, G.T. (ed.) On the Economy of Plant Form and Function. Cam-
bridge University Press, Cambridge, pp. 2555.
Flore, J.A. (1994) Stone fruit. In: Schaffer, B. and Andersen, P.C. (eds) Handbook of Envi-
ronmental Physiology of Fruit Crops. Vol. 1, Temperate Crops. CRC Press, Boca
Raton, Florida, pp. 233270.
Foyer, C.H. (1988) Feedback inhibition of photosynthesis through source-sink regulation
in leaves. Plant Physiology 26, 483492.
Frak, E., Le Roux, X., Millard, P., Dreyer, E., Vandame, M. and Wendler, R. (2001) Chang-
es in total leaf nitrogen and partitioning of leaf nitrogen drive photosynthetic ac-
climation to light in fully developed walnut leaves. Plant Cell and Environment 24,
12791288.
Gandhi, S.R. (1955) The Mango in India. Farm Bulletin 6. Indian Council for Agricul-
tural Research, New Delhi, 64 pp.
Gazit, S. and Kadman, A. (1980) 13-1 mango rootstock selection. HortScience 15,
669.
Gitay, H., Brown, S., Easterlin, W. and Jallow, B. (2001) Ecosystems and their goods and
services. In: McCarthy, J.J., Osvaldo, F.C., Leary, N.A., Dokken, D.J. and Whilte,
K.S. (eds) Climate Change 2001: Impacts, Adaptation and Vulnerability. Cambridge
University Press, Cambridge, UK, pp. 237242.
Gonzalez, A. and Blaikie, S.J. (2003) Seasonal variation of carbon assimilation in mango
(cv. Kensington Pride): effect of owering treatments. Australian Journal of Agricul-
tural Research 54, 309321.
Harding, R.B., Miller, M.P. and Fireman, M. (1958) Absorption of salts by citrus leaves
during sprinkling with water soluble surface irrigation. Proceedings of the Ameri-
can Society for Horticultural Science 71, 248256.
Harley, P.C., Thomas, R.B., Reynolds, J.F. and Strain, B.R. (1992) Modelling photosyn-
thesis of cotton grown in elevated CO
2
. Plant Cell and Environment 15, 271282.
Harvey, G.W. (1979) Photosynthetic performance of isolated leaf cells from sun and
shade plants. Carnegie Institute Washington Yearbook 79, 161164.
Hollinger, D.Y. (1996) Optimality and nitrogen allocation in a tree canopy. Tree Physiol-
ogy 16, 627634.
Houghton, J. (2005) Global warming. Reports on Progress in Physics 68, 13431403.
Hoult, M.D., Donnelly, M.M. and Smith, M.W. (1996) Salt exclusion varies amongst
polyembryonic mango cultivar seedlings. Acta Horticulturae 455, 455458.
Idso, S.B. and Kimball, B.A. (1991) Downward regulation of photosynthesis and growth
at high CO
2
levels. Plant Physiology 96, 990992.
Issarakraisila, M., Considine, J.A. and Turner, D.W. (1991) Pattern of vegetative and re-
productive growth of mango trees in a warm temperate region of Western Australia.
Acta Horticulturae 291, 188197.
Jackson, J.E. (1980) Light interception and utilization by orchard systems. Horticultural
Reviews 2, 208287.
Jindal, P.C., Singh, J.P. and Gupta, O.P. (1975) Screening of mango seedlings for salt
tolerance. Haryana Journal of Horticultural Science 4, 112115.
Jindal, P.C., Singh, J.P. and Gupta, O.P. (1976) Studies on salt tolerance in mango
injurious effects of salts on young mango seedlings. Progressive Horticulture 8,
6571.
Joel, D.M. (1980) Resin ducts in the mango fruit: a defense system. Journal of Experi-
mental Botany 31, 17071718.
B. Schaffer et al. 204
Joel, D.M. and Fahn, A. (1980a) Ultrastructure of the resin ducts of the Mangifera indica
L. (Anacardiaceae). 1. Differentiation and senescence of the shoot ducts. Annals of
Botany 46, 225223.
Joel, D.M. and Fahn, A. (1980b) Ultrastructure of the resin ducts of the Mangifera indica
L. (Anacardiaceae). 2. Resin secretion in the primary stem ducts. Annals of Botany
46, 779783.
Joel, D.M. and Fahn, A. (1980c) Ultrastructure of the resin ducts of the Mangifera indica
L. (Anacardiaceae). 3. Secretion of the protein-polysaccharide mucilage in the fruit.
Annals of Botany 46, 785790.
Jones, H.G. (1981) Carbon dioxide exchange of developing apple (Malus pumila Mill.)
fruits. Journal of Experimental Botany 32, 12031210.
Jones, H.G. and Sutherland, R.A. (1991) Stomatal control of xylem embolism. Plant Cell
and Environment 14, 607612.
Kadman, A. (1964) The uptake and accumulation of sodium in avocado seedlings. Pro-
ceedings of the American Society for Horticultural Science 85, 179182.
Kadman, A., Gazit, S. and Ziv, G. (1976) Selection of mango rootstocks for adverse
water and soil conditions in arid areas. Acta Horticulturae 57, 8188.
Kallarackal, J., Milburn, J.A. and Baker, D.A. (1990) Water relations of the banana. III.
Effects of controlled water stress on water potential, transpiration, photosynthesis
and leaf growth. Australian Journal of Plant Physiology 17, 7990.
Kellomki, S. and Wang, K.-Y. (1997) Photosynthetic responses of Scots pine to elevated
CO
2
and nitrogen supply: results of a branch-in-bag experiment. Tree Physiology
17, 231240.
Koch, K.E. (1996) Carbohydrated-modulated gene expression in plants. Annual Review
of Plant Physiology and Plant Molecular Biology 47, 509540.
Kozlowski, T.T. (1984) Responses of woody plants to ooding. In: Kozlowski, T.T. (ed.)
Flooding and Plant Growth. Academic Press, New York, pp. 129163.
Kramer, P.J. (1983) Water Relations of Plants. Academic Press, London.
Krapp, A. and Stitt, M. (1995) An evaluation of direct and indirect mechanisms for the
sink-regulation of photosynthesis in spinach: changes in gas exchanges, carbohy-
drates, metabolites, enzyme activities and steady-state transcript levels after cold-
girdling source leaves. Planta 195, 313323.
Kriedemann, P.E. (1968) Observations on gas exchange in the developing sultana berry.
Australian Journal of Biological Science 21, 907916.
Lakso, A.N. (1994) Apple. In: Schaffer, B. and Andersen, P.C. (eds) Handbook of Environ-
mental Physiology of Fruit Crops. Vol. I. Temperate Crops. CRC Press, Boca Raton,
Florida, pp. 343.
Larson, K.D., Schaffer, B. and Davies, F.S. (1989) Effect of irrigation on leaf water poten-
tial, growth and yield of mango trees. Proceedings of the Florida State Horticultural
Society 102, 226228.
Larson, K.D., Davies, F.S. and Schaffer, B. (1991a) Floodwater temperature and lenticel
hypertrophy of Mangifera indica L. American Journal of Botany 78, 13971403.
Larson, K.D., Graetz, D.A. and Schaffer, B. (1991b) Flood-induced chemical transforma-
tions in calcareous agricultural soils of South Florida. Soil Science 152, 3340.
Larson, K.D., Schaffer, B. and Davies, F.S. (1991c) Flooding, leaf gas exchange and
growth of mango trees in containers. Journal of the American Society of Horticul-
tural Science 116, 156160.
Larson, K.D., Schaffer, B. and Davies, F.S. (1992) Flooding, mineral nutrition and gas
exchange of mango trees. Scientia Horticulturae 52, 113124.
Larson, K.D., Schaffer, B. and Davies, F.S. (1993) Floodwater oxygen content, lenticel
hypertrophy, and ethylene evolution from mango (Mangifera indica L.) trees. Jour-
nal of Experimental Botany 44, 665671.
Ecophysiology 205
Le Roux, X., Sinoquet, H. and Vandame, M. (1999) Spatial distribution of leaf weight per
area and leaf nitrogen concentration in relation to local radiation regime within an
isolated tree crown. Tree Physiology 19, 181188.
Le Roux, X., Walcroft, A.S., Daudet, F.A., Sinoquet, H., Chaves, M.M., Rodrigues, A. and
Osorio, L. (2001) Photosynthetic light acclimation in peach leaves: importance of
changes in mass:area ratio, nitrogen concentration, and leaf nitrogen partitioning.
Tree Physiology 21, 377386.
Lu, P. (2002) Measurement of whole-tree water use of some tropical and subtropical
tree crops and its application in irrigation management. Acta Horticulturae 575,
781789.
Lu, P. (2006) Mango water relations and irrigation schedule. In: Malik, A.U. (ed.) Pro-
ceedings of the First International Conference on Mango and Date Palm: Culture
and Export. University of Agriculture, Faisalabad, Pakistan, 2023 June 2005,
pp. 111.
Lynch, J. and Gonzlez, A. (1993) Canopy nutrient allocation in relation to incident light
in the tropical fruit tree Borojoa patinoi (Cuatr.). Journal of the American Society for
Horticultural Science 118, 777785.
Manter, D.K. and Kerrigan, J. (2004) A/C
i
curve analysis across a range of woody plant
species: inuence of regression analysis parameters and mesophyll conductance.
Journal of Experimental Botany 55, 25812588.
Marshall, B. and Biscoe, P.V. (1980) A model for C
3
leaves describing the dependence of
net photosynthesis on irradiance. II. Application to the analysis of ag leaf photo-
synthesis. Journal of Experimental Botany 31, 4148.
Mayers, P.E., Hutton, D.G. and Saranah, J. (1984) Integrated control of bacterial black
spot of mangoes in south east Queensland. In: Chapman, G.R. (ed.) Proceedings of
the First Australian Mango Research Workshop. Commonwealth Scientic and
Industrial Research Organization (CSIRO), Melbourne, pp. 258260.
Medlyn, B.E., Dreyer, E., Ellsworth, D., Forstreuter, M., Harley, P.C., Kirschbaum, M.U.F.,
Le Roux, X., Loustau, D., Montpied, P., Strassemeyer, J., Walcroft, A.S. and Wang,
K. (2002) Temperature response of parameters of a biochemically-based model of
photosynthesis: a review of experimental data. Plant Cell and Environment 25,
11691176.
Merzlyak, M.N. and Hendry, G.A.F. (1994) Free radical metabolism, pigment degrada-
tion and lipid peroxidation in leaves during senescence. Proceedings of the Royal
Society of Edinburgh 102B, 459471.
Mukherjee, S.K. (1953) The mango its botany, cultivation, uses, and future improve-
ment, especially as observed in India. Economic Botany 7, 130160.
Mukherjee, S.K. (1972) Origin of the mango (Mangifera indica). Economic Botany 26,
260266.
Nii, N., Watanabe, T., Yamaguchi, K. and Nishimura, M. (1995) Changes of anatomical
features, photosynthesis and ribulose-bisphosphate carboxylase-oxygenase content
of mango leaves. Annals of Botany 76, 649656.
Nobel, P.S. (1983) Biophysical Plant Physiology and Ecology. W.H. Freeman, New
York.
Noodn, L.D., Guiamt, J.J. and John, J. (1997) Senescence mechanisms. Physiologia
Plantarum 101, 746753.
Oosthuyse, S.A. (1992) Ideas on pruning of mango trees. South African Mango Growers
Association Yearbook 12, 18.
Pammenter, N.W., Loreto, F. and Sharkey, T.D. (1993) End product feedback effects on
photosynthetic electron transport. Photosynthesis Research 35, 514.
Paul, M.J. and Foyer, C. (2001) Sink regulation of photosynthesis. Journal of Experimen-
tal Botany 360, 13831400.
B. Schaffer et al. 206
Paul, M.J. and Pellny, T.L. (2003) Carbon metabolite feedback regulation of leaf photo-
synthesis and development. Journal of Experimental Botany 382, 539547.
Peisker, M. and Apel, H. (2001) Inhibition by light of CO
2
evolution from dark evolution:
comparison of two gas exchange methods. Photosynthesis Research 70, 291298.
Pongsomboon, W. (1991) Effects of temperature and water stress on tree growth, ower-
ing, fruit growth and retention of mango (Mangifera indica L.). PhD thesis, Kasetsart
University, Bangkok, Thailand.
Pongsomboon, W., Whiley, A.W., Stephenson, R.A. and Subhadrabandhu, S. (1992) Ef-
fect of air temperatures on diurnal variation of water potential, conductance and
CO
2
assimilation of mango (Mangifera indica L.) leaves. Acta Horticulturae 321,
472481.
Proctor, J.T.A. and Creasey, L.L. (1971) Effect of supplementary light on anthocyanin
synthesis in McIntosh apples. Journal of the American Society for Horticultural
Science 96, 523526.
Quirino, B.F., Reiter, W.D. and Amasino, R.D. (2001) One of two tandem Arabidopsis
genes homologous to monosaccharide transporters is senescence-associated. Plant
Molecular Biology 46, 447457.
Reich, P.B. and Borchert, R. (1988) Changes with leaf age in stomatal function and water
status of several tropical tree species. Biotropica 20, 6069.
Riesmeier, J.W., Flugge, U.-I., Schulz, B., Heineke, D., Heldt, H.-W., Willmitzer, L. and
Frommer, W.B. (1993) Antisense repression of the chloroplast triose phosphate
translocator affects carbon partitioning in transgenic potato plants. Proceedings of
the National Academy of Science of the USA 90, 61606164.
Roper, T.R. and Williams, L. (1989) Net CO
2
assimilation and carbohydrate partitioning
of grapevine leaves in response to trunk girdling and gibberellic acid application.
Plant Physiology 89, 11361140.
Roper, T.R., Keller, J.D., Loescher, W.H. and Rom, C.R. (1988) Photosynthesis and car-
bohydrate partitioning in sweet cherry: fruiting effects. Physiologia Plantarum 72,
4247.
Rosati, A., Esparza, G., DeJong, T.M. and Pearcy, R.W. (1999) Inuence of canopy light
environment and nitrogen availability on leaf photosynthetic characteristics and
photosynthetic nitrogen-use efciency of eld-grown nectarine trees. Tree Physiol-
ogy 19, 173180.
Rosati, A., Day, K.R. and DeJong, T.M. (2000) Distribution of leaf mass per unit leaf area
and leaf nitrogen concentration determine partitioning of leaf nitrogen within cano-
pies. Tree Physiology 20, 271276.
Schaffer, B. and Gaye, G.O. (1989) Seasonal effects of pruning on light penetration,
specic leaf density, and chlorophyll content of mango. Scientia Horticulturae 41,
5561.
Schaffer, B., Andersen, P.C. and Ploetz, R.C. (1992) Responses of fruit crops to ooding.
Horticultural Reviews 13, 257313.
Schaffer, B., Whiley, A.W. and Crane, J.H. (1994) Mango. In: Schaffer, B. and Andersen,
P.C. (eds) Handbook of Environmental Physiology of Fruit Crops. Vol. 2, Sub-tropical
and Tropical Crops. CRC Press, Boca Raton, Florida, pp. 165197.
Schaffer, B., Whiley, A.W., Searle, C. and Nissen, R.J. (1997) Leaf gas exchange, dry mat-
ter partitioning, and mineral element concentrations in mango (Mangifera indica L.)
as inuenced by elevated atmospheric CO
2
concentration and root restriction.
Journal of the American Society for Horticulture Science 122, 849855.
Schaffer, B., Whiley, A.W. and Searle, C. (1999) Atmospheric CO
2
enrichment, root re-
striction, photosynthesis, and dry-matter partitioning in subtropical and tropical
fruit crops. HortScience 34, 10331037.
Ecophysiology 207
Schaffer, B., Davies, F.S. and Crane, J.H. (2006) Responses of subtropical and tropical
fruit trees to ooding in calcareous soil a review. HortScience 41, 549555.
Schaper, H. and Chacko, E.K. (1993) Effect of irradiance, leaf age, chlorophyll content
and branch-girdling on gas exchange of cashew (Anacardium occidentale L.)
leaves. Journal of Horticultural Science 68, 541550.
Schmutz, U. (2000) Effect of salt stress (NaCl) on whole plant CO
2
-gas exchange in
mango. Acta Horticulturae 509, 269276.
Schmutz, U. and Ldders, P. (1993) Physiology of saline stress in one mango (Mangifera
indica L.) rootstock. Acta Horticulturae 341, 160167.
Scholeeld, P.B., Oag, D.R. and Sedgley, M. (1986) The relationship between vegetative
and reproductive development in the mango in northern Australia. Australian Journal
of Agricultural Research 37, 425433.
Searle, C., Whiley, A.W., Simpson, D.R. and Saranah, J.B. (1995) A preliminary phenol-
physiological model for Kensington in subtropical Australia. In: Mango 2000
Marketing Seminar and Production Workshop Proceedings. Department of Primary
Industries, Brisbane, pp. 124127.
Shivashankara, K.S. and Mahai, C.K. (2000) Inhibition of photosynthesis by owering in
mango (Mangifera indica L.). A study by gas exchange methods. Scientia Horticul-
turae 83, 205212.
Singaas, E.L., Ort, D.R. and deLucia, E.H. (2001) Variation in measured values of photo-
synthetic quantum yield in ecophysiological studies. Oecologia 128, 1523.
Singh, R.N. and Arora, K.S. (1965) Some factors affecting fruit drop in mango (Mangifera
indica L.). Indian Journal of Agricultural Science 35, 196205.
Spreitzer, R.J. and Salvucci, M.E. (2002) Rubisco: structure, regulatory interactions, and
possibilities for a better enzyme. Annual Review of Plant Physiology and Plant
Molecular Biology 53, 449475.
Stephenson, R.A., Ko, H.L. and Gallagher, E.C. (1989) Plant water relations of stressed,
non-bearing macadamia trees. Scientia Horticulturae 39, 4153.
Sukhvibul, N., Whiley, A.W., Smith, M.K. and Hetherington, S.E. (2000) Susceptibility of
mango (Mangifera indica L.) to cold-induced photoinhibition and recovery at
different temperatures. Australian Journal of Agricultural Research 51, 503513.
Sukonthasing, S., Wongrakpanich, M. and Verheij, E.W.M. (1991) Mangifera indica L.
In: Verheij, E.W.M. and Coronel, R.E. (eds) Plant Resources of South-East Asia, No.
2. Edible Fruits and Nuts. Pudoc, Wageningen, the Netherlands, pp. 211216.
Sun, J., Okita, T.W. and Edwards, G.E. (1999) Modication of carbon partitioning, pho-
tosynthetic capacity, and O
2
sensitivity in Arabidopsis plants with low ADP-glucose
pyrophosphorylase activity. Plant Physiology 119, 267276.
Thomas, B.R. and Rodriguez, R.L. (1994) Metabolite signals regulate gene expression
and source/sink relations in cereal seedlings. Plant Physiology 54, 201207.
Tyree, M.T. and Sperry, J.S. (1988) Do woody plants operate near the point of cata-
strophic xylem dysfunction caused by dynamic water stress? Answers from a model.
Plant Physiology 88, 574580.
Urban, L. and Alphonsout, L. (2007) Girdling decreases photosynthetic electron uxes and
induces sustained photoprotection in mango leaves. Tree Physiology 27, 345352.
Urban, L. and Jannoyer, M. (2004) Functioning and role of stomata in mango leaves.
Acta Horticulturae 645, 441446.
Urban, L., Berteuil, F. and Lchaudel, M. (2002) A coupled photosynthesis and stomatal
conductance model for mango leaves. Acta Horticulturae 584, 8188.
Urban, L., Le Roux, X., Sinoquet, H., Jaffuel, S. and Jannoyer, M. (2003) A biochemical
model of photosynthesis for mango leaves: evidence for an effect of the local
source/sink balance on photosynthetic capacity. Tree Physiology 23, 289300.
B. Schaffer et al. 208
Urban, L., Lu, P. and Thibaud, R. (2004a) Inhibitory effect of owering on leaf photosyn-
thesis in mango. Tree Physiology 24, 387399.
Urban, L., Lchaudel, M. and Lu, P. (2004b) Interpreting the effect of fruit load and
girdling on leaf net photosynthesis in mango. Journal of Experimental Botany 405,
20752085.
Urban, L., Montpied, P. and Normand, F. (2006) Season effects on leaf nitrogen partition-
ing and photosynthetic water use efciency in mango. Journal of Plant Physiology
163, 4857.
Urban, L., Jgouzo, L., Damour, G., Vandame, M. and Franois, C. (2008) Interpreting
the decrease in leaf photosynthesis during owering in mango. Tree Physiology 28,
10251036.
Van der Meulen, A., Smith, J.H.E., Kok, J.B., Schwartz, A. and Jacobs, C.J. (1971) Mango
growing in South Africa. Citrus and Subtropical Fruit Research Institute Leaet No.
48. Department of Agriculture and Technical, Pretoria.
Venning, F.D. (1948) The ontogeny of the laticiferous canals in the Anacardiacae. Amer-
ican Journal of Botany 35, 637644.
Walcroft, A.S., Whitehead, D., Silvester, W.B. and Kelliher, F.M. (1997) The response of
photosynthetic model parameters to temperature and nitrogen concentration in Pi-
nus radiata D. Don. Plant Cell and Environment 20, 13381348.
Walcroft, A., Le Roux, X., Diaz-Espejo, A., Dones, N. and Sinoquet, H. (2002) Spatial
and temporal variability of photosynthetic capacity within a peach tree crown. Tree
Physiology 22, 929938.
Walker, R.R. (1986) Sodium exclusion and potassium-sodium selectivity in salt-treated
trifoliate orange (Poncirus trifoliata) and Cleopatra mandarin (Citrus reticulata)
plants. Australian Journal of Plant Physiology 13, 293303.
Weng, J.H., Chen, Y.N. and Liao, T.S. (2006a) Relationships between chlorophyll uo-
rescence parameters and photochemical reectance index of tree species adapated
to different temperature regimes. Functional Plant Biology 33, 241246.
Weng, J.H., Jhaung, L.H., Jiang, J.Y., Lai, G.M. and Liao, T.S. (2006b) Down-regulation
of photosystem 2 efciency and spectral reectance in mango leaves under very
low irradiance and varied chilling treatments. Photosynthetica 44, 248254.
Whiley, A.W. (1994) Ecophysiological studies and tree manipulation for maximisation of
yield potential in avocado (Persea americana Mill.). PhD thesis, University of Natal,
Pietermaritzburg, South Africa, 174 pp.
Whiley, A.W. and Schaffer, B. (1994) Avocado. In: Schaffer, B. and Andersen, P.C. (eds)
Handbook of Environmental Physiology of Fruit Crops, Vol. 2. Sub-Tropical and
Tropical Crops. CRC Press, Boca Raton, Florida, pp. 335.
Whiley, A.W., Rasmussen, T.S., Saranah, J.B. and Wolstenholme, B.N. (1989) Effect of
temperature on growth, dry matter production and starch accumulation in ten man-
go (Mangifera indica L.) cultivars. Journal of Horticultural Science 64, 753765.
Whiley, A.W., Schaffer, B. and Lara, S.P. (1991) Carbon dioxide exchange of developing
avocado (Persea americana Mill.) fruit. Tree Physiology 11, 8594.
Whiley, A.W., Searle, C., Schaffer, B. and Wolstenholme, B.N. (1999) Cool orchard tem-
peratures or growing trees in containers can inhibit leaf gas exchanges of avocado
and mango. Journal of the American Society for Horticultural Science 124, 4651.
Whiley, A.W., Hofman, P.J., Marques, J.R., Stubbings, B. and Whiley, D.G. (2006) Devel-
opment of best practice pre- and postharvest protocols for production of Calypso
mango. Final Project Report, Horticulture Australia Ltd (FR02049), Sydney, Austra-
lia, pp. 157.
Wilson, K.B., Baldocchi, D.D. and Hanson, P.J. (2000) Quantifying stomatal and non-
stomatal limitations to carbon assimilation resulting from leaf aging and drought in
mature deciduous tree species. Tree Physiology 20, 787797.
Ecophysiology 209
Wingler, A., von Schaewen, R.C., Leegood, P.J. and Quick, W.P. (1998) Regulation of
leaf senescence by cytokinin, sugars and light. Effects on NADH-dependent hy-
droxypyruvate reductase. Planta 116, 329335.
Wolstenholme, B.N. and Whiley, A.W. (1995) Ecophysiology of the mango tree as a
basis for pre-harvest management. In: Mango 2000 Marketing Seminar and Pro-
duction Workshop, Department of Primary Industries, Brisbane, pp. 519.
Young, T.W. and Sauls, J.W. (1981) The Mango Industry in Florida. Florida Cooperative
Extension Services Bulletin 189, University of Florida, Gainesville, Florida.
Yusof, I.M., Buchanan, D.W. and Gerber, J.F. (1969) The response of avocado and mango
to soil temperature. Journal of the American Society for Horticultural Science 94,
619621.
Zude, M. and Ludders, P. (1997) Vegetative growth cycles and comparison of chloro-
phyll and phenol contents, gas exchange and water regime from young to old
leaves. Journal of Applied Botany 71, 1013.
Zude-Sasse, M., Hartmond, U., Ebert, G. and Ludders, P. (2001) Pyridine nucleotide
charge reduces photosynthesis under short-term oxygen deciency. Journal of the
American Society for Horticultural Science 126, 703709.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
210 (ed. R.E. Litz)
7 Fruit Diseases
D. Prusky, I. Kobiler, I. Miyara and N. Alkan
Agricultural Research Organization, Bet Dagan, Israel
7.1 Introduction 210
7.2 Anthracnose 211
The pathogenesis strategy 212
Epidemiology the disease cycle 213
Mechanisms of fungal pathogenicity and fruit resistance 214
Management 216
7.3 Alternaria Rot (Black Spot) 219
Pathogenesis 219
Mechanisms of fungal pathogenicity and fruit resistance 220
Management 220
7.4 Stem-end Rots 221
Pathogenesis 221
Epidemiology 222
Management 223
7.5 Other Minor Diseases 224
7.6 Conclusions 224
7.1 Introduction
Postharvest diseases can reduce fruit quality and cause severe economic
losses, due to decay resulting in completely unmarketable and blemished
fruits that are often sold in less demanding local markets, where the prices
are considerably lower than export prices. It is clear to the producer that
quality at the time of harvest cannot be improved but merely maintained for
a limited period of time. Harvesting fruits at the optimal stage, with respect
to size and maturity, can, therefore, ensure peak quality and maximum shelf-
life potential. Thus, managing total tree health can contribute to reducing
postharvest losses. It is known that older and neglected orchards may become
a profuse inoculum source for postharvest diseases. Furthermore, preharvest
Fruit Diseases 211
stress factors such as excess or shortage of water, uctuating or extreme envi-
ronmental conditions and high nitrogen levels (Hawthorne, 1989) can render
fruit more susceptible to postharvest diseases. After harvest, improper treat-
ment of fruits through storage at non-optimal temperatures accelerates fruit
deterioration as a result of enhancement of normal physiological processes
such as respiration and ethylene production (Thompson et al., 2002). More-
over, excessive temperatures in the eld during harvesting, in transit and in
the packing house can render tropical fruit more susceptible to chilling injury
and can contribute to the development of disease. Combinations of high tem-
peratures and high relative humidity (RH) favour the growth of postharvest
pathogens, and can contribute to the development of disease at the retail end
(Banik et al., 1998). Proper, uninterrupted cooling, therefore, protects quality
and extends the shelf life of produce. Precooling of products (Thompson et
al., 2002), and application of top or liquid icing, vacuum, hydrocooling and
forced air cooling (Thompson et al., 2002) are examples of effective alterna-
tive methods that can be used to remove eld heat and to restrict pathogen
growth. Effective cold-chain management is crucial to ensuring product
integrity and preventing postharvest pathogens from spoiling produce in
transit or shortly after arrival (Lizada, 1993). Furthermore, low-temperature
storage conditions are generally not conducive to disease development, a
fact that can be exploited to ensure quality and extend shelf life (Banik et al.,
1998). However, improper cooling during shipping, or interrupted cooling
will also promote microbial growth, resulting ultimately in product spoilage.
In practice, several breaks in the cold chain might have signicant impacts on
decay development.
Among the postharvest diseases of mango, anthracnose is the most prev-
alent in humid growing regions. The incidence of this disease can reach
almost 100% in fruit produced under wet or very humid conditions. Other
common postharvest diseases of mango in humid areas are stem-end rot,
caused by Lasiodiplodia theobromae (Pat.) Griffon & Maubl. (Sangchote, 1998)
and Dothiorella dominicana Petrak. et Cif. (Johnson and Sangchote, 1994), and
black mould rot, caused by Aspergillus niger van Tieghem. Mango black spot,
caused by Alternaria alternata (Fr.:Fr.) Keissl. (Prusky et al., 1983), is prevalent
in dry countries.
7.2 Anthracnose
Mango anthracnose is caused by Glomerella cingulata (Stoneman) Spauld. &
H. Schrenk (anamorph: Colletotrichum gloeosporioides (Penz.) Penz. & Sacc. In
Penz.), C. gloeosporioides Penz. var. minor J.H. Simmonds (Fitzell and Peak,
1984) and Colletotrichum acutatum J.H. Simmonds (Freeman et al., 1998). The
pathogen also causes blossom blight, leaf blight and, in severe cases, tree
dieback (Ploetz, 1994; Ploetz and Prakash, 1997). The form-genus Colletotri-
chum Corda (form-order Melanconiales; form-class Coelomycetes; subdivi-
sion Deuteromycotina) comprises imperfect fungal species which exist as
Glomerella (subdivision Ascomycotina) in their sexual, teleomorphic or perfect
D. Prusky et al. 212
state. These fungal pathogens occur worldwide, and the genus is synony-
mous with anthracnose. Leaf anthracnose appears as irregular black necrotic
spots on both sides of the mango leaves. Lesions often coalesce and form
large necrotic areas, frequently along the leaf margins. Lesions develop pri-
marily on young tissue, and conidia are formed in lesions of all ages. Under
favourable conditions, the fungus can invade the twigs and cause dieback
(Ploetz et al., 1996). Panicle anthracnose or blossom blight can affect both the
inorescence stalk and the individual owers; in the stalk, elongated dark
grey to black lesions appear; the blighted owers are dry, and their colour
ranges from brown to black. Fruits smaller than pea-size can be infected and
abort; whereas, larger fruits that are aborted because of normal self-thinning
or other physiological causes are usually mummied. The resulting mummi-
ed fruit are invaded saprophytically by C. gloeosporioides, and the fungus
sporulates abundantly on them.
Although eld anthracnose causes considerable damage, the vast losses
inicted by postharvest anthracnose are of far greater economic importance.
Postharvest anthracnose appears on the fruit surface, as rounded brown to
black lesions with an indenite border. Lesions > 2 cm in diameter are fairly
common. Lesions of various sizes can coalesce and cover extensive areas of
the fruit, typically in a tear-drop pattern that develops from the basal towards
the distal end of the fruit. Lesions are usually restricted to the peel, but in
severe cases the fungus can invade the pulp. In advanced stages of the dis-
ease, the fungus produces acervuli, and abundant orange to salmon-pink
masses of conidia appear on the lesions.
The pathogenesis strategy
Colletotrichum gloeosporioides is regarded as a hemibiotrophic species that
commences its invasion with a transient post-penetrative asymptomatic bio-
trophy, characterized by a temporary connement with a localized mode of
intracellular hemibiotrophy. This is succeeded by a phase of destructive
necrotrophy that culminates in the appearance of disease symptoms and pro-
duction of conidiomata by the pathogens (Prusky and Plumbley, 1992; Latunde-
Dada et al., 1999). Quiescent species pass through a prolonged phase of
pre-penetrative growth that is arrested in synchrony with the physiological
state of the infected organ.
Following its landing on the fruit or host tissue, the ungerminated
aseptate conidium differentiates to a melanized appressorium. In the case of
Colletotrichum lindemuthianum, appressorial adhesion is mediated by the man-
nose- and galactose-rich glycoproteins of the extracellular matrices, which
are secreted during appressorial differentiation (Pain et al., 1996). Both the
surface wax of the specic fruit and the hard surface of the tissue are recog-
nized as host signals that selectively trigger germination and appresso-
rium formation solely by the conidia of C. gloeosporioides (Podila et al., 1993;
Liu and Kolattukudy, 1998). The melanin of the melanized appressoria pro-
tects the fungus and behaves as a permeability barrier, and the appressoria
Fruit Diseases 213
contain osmolytes (i.e. glycerol) that are needed for generating the osmoti-
cum from which high turgor pressure will drive the invasive forces of the
penetrating hyphae through the small appressorial pores (0.5 m in Col-
letotrichum sublineolum). The osmolyte is generated by the metabolism of
stored glycogen, trehalose and lipids and, at an in vivo concentration of at
least 3.3 M in mature-melanized appressoria (de Jong et al., 1997), this osmo-
lyte is capable of generating turgor pressures as high as 8 MPa (Howard et al.,
1991; Money and Howard, 1996). Melanized appressoria appear to be quite
capable of a forcible, non-enzymatic penetration of an intact host surface.
However, Dickman et al. (1982) suggested that attack on papaya by C. gloeo-
sporioides depended on cutinase production by the pathogen. The germi-
nated appressoria in Colletotrichum develop single infection hyphae that
grow and extend into the waxy cuticle, reach the rst layers of pericarp
cells, and then become quiescent for long periods of time (Coates et al.,
1993; Prusky, 1996).
In recent years, the taxonomy of Colletotrichum has been claried by the
adoption of molecular biological techniques that involve the use of poly-
merase chain reaction amplication, alignment of nucleotide sequences, and
the construction of dendograms, phylogenetic trees and similarity matrices
from the data generated. For example, Sherriff et al. (1994) and Sreenivasap-
rasad et al. (1996) used homologies between the nucleotide sequences from
amplied non-transcribed regions (internally transcribed spacers 1 and 2
(ITS1, ITS2)) and the large subunits (domains 1 and 2) of ribosomal DNA
extracted from a wide range of isolates to both justify and resolve the taxo-
nomic status of Colletotrichum species. Bailey (1997) proposed the adoption of
species aggregates based on these and other data.
Epidemiology the disease cycle
Conidia are formed abundantly in the mango canopy (Fitzell and Peak, 1984)
which, therefore, is considered to be the primary source of inoculum. In the
eld, C. gloeosporioides produces conidia on lesions on leaves, twigs, panicles
and mummied fruits (Arauz, 2000). Conidia can be rain-splashed onto other
leaves or owers, where they can cause secondary infections. Developing
fruit can be infected, and some isolates can cause preharvest fruit loss (Gan-
totti and Davis, 1993). In the case of postharvest anthracnose, developing
fruits are infected in the eld, but the infections remain quiescent until the
onset of ripening, which occurs after harvest. Once the climacteric period of
the fruit starts, lesions begin to develop but there is no fruit-to-fruit infection.
In this context Prusky and co-workers (Guetsky et al., 2005) suggested that
the capability of C. gloeosporioides to cause early disease symptoms in unripe
fruits depends on the activation of laccases by specic strains of the fungus.
Details of these systems are discussed in the following section.
Colletotrichum gloeosporioides requires free water or RH > 95% to enable
conidial germination and appressorium formation (Fitzell et al., 1984; Dodd
et al., 1991). However, conidia can survive for 12 weeks under low RH and
D. Prusky et al. 214
then germinate if exposed to 100% RH (Estrada et al., 1993). In general, infec-
tion is favoured at temperatures ranging from 20 to 30C.
Mechanisms of fungal pathogenicity and fruit resistance
Unripe mango fruits are reservoirs of extremely high concentrations of pre-
formed antimicrobial compounds. This arsenal of constitutive resistance
weapons may accumulate in the immature pericarp, but not in the mesocarp,
at concentrations in fruit fresh weight of up to 220 g/g. They include mix-
tures of 5-substituted resorcinols such as resorcinol-5-(12-heptadecadienyl)
and resorcinol-5-(pentadecyl) (Droby et al., 1986, 1987; Prusky and Plumbley,
1992; Prusky, 1996) (Fig. 7.1). Faced by such a chemically adverse environ-
ment, the fungus usually postpones its development until the concentrations
of the compounds in the host decline. The capability to penetrate the host
without encountering the arsenal of passive chemical resistance or triggering
the weapons of active resistance is inherent in a number of fruit-infecting
Colletotrichum species such as C. gloeosporioides, C. acutatum and A. alternata
(Prusky and Keen, 1993). The conidia, melanized appressoria and penetra-
tion pegs of these quiescent species have evolved mechanisms of physiologi-
cal inactivity that enable them to pause until the host-ripening process and
the decline of antifungal compounds starts, thus enabling the avoidance of
host defences that exist within unripe, pre-climacteric fruits. Colletotrichum
gloeosporioides is a eld-to-store pathogen whose conidia infect during the
preharvest growth of fruits. In fruits such as mango and avocado, termina-
tion of fungal quiescence is associated jointly with the climacteric production
of ethylene during fruit ripening and the onset of dramatic reductions in the
levels of preformed fungitoxic resorcinols (Droby et al., 1986, 1987). Flaishman
HO HO HO HO
(CH
2
)
11
CH
CH
(CH
2
)
3
CH
3
(CH
2
)
14
CH
2
CH
3
Resorcinol-5-(pentadecyl) Resorcinol-5-(12-heptadecenyl)
Fig. 7.1. Chemical structure of the 5-substituted resorcinols isolated from mango
fruits.
Fruit Diseases 215
and Kolattukudy (1994) hypothesized that ethylene is involved in terminat-
ing quiescence, by inducing appressoria formation and hyphal growth,
which strongly suggests that Colletotrichum spp. must have coevolved with
their hosts, to develop a mechanism that uses the hosts ripening hormone as
a signal to reactivate the infection process. However, when Prusky and co-
workers (Prusky et al., 1996) applied ethylene to unripe fruits they could not
enhance the termination of quiescence as had been suggested, possibly indi-
cating that ethylene is a signal that induces appressoria formation in vitro,
but that does not enhance fungal growth in the fruit. Furthermore, since fun-
gal infection and appressoria formation occur in the orchard, it is difcult to
conceive how ethylene produced much later, during ripening and storage,
could enhance appressoria formation on the fruit. Regardless of the mecha-
nisms that may be involved in the onset and termination of quiescence, qui-
escent infection appears to be a case of coevolution; it is advantageous to
both pathogen and host in a natural ecosystem to allocate chemical defences
to the immature fruit but not to the ripe fruit (Prusky and Keen, 1993).
Quiescence also may result from a localized host response that is often
associated with an oxidative burst, i.e. production of reactive oxygen species
(ROS). Localized generation of ROS during quiescence was found to be one
of the earliest (within 23 h) detectable cytological defence responses to
attempted penetration of unripe, resistant avocado fruits by C. gloeosporioides.
Beno-Moualem and Prusky (2000) proposed that fungal infection of unripe
fruits leads to production of ROS during quiescence, at the infection loci
surrounding the germinated appressoria and their penetration hyphae. This
localized ROS production would activate the synthesis of antifungal com-
pounds or compounds that inhibit fungal metabolism (Ardi et al., 1998) at the
infection loci, thereby enhancing and/or preserving the levels of antifungal
compounds and, in turn, inhibiting fungal development and imposing
quiescence.
Guetsky et al. (2005) suggested that initiation of quiescence could result
from the fungal capability to induce laccase activity that modulates the metab-
olism of preformed antifungal compounds and the activation of the quies-
cent C. gloeosporioides that occurs during fruit ripening. Early activity of
aggressive isolates in unripe fruits included increased laccase activities that
resulted in early metabolism of preformed antifungal compounds leading to
early appearance of symptoms.
When the levels of toxic compounds in the fruit peel decline, C. gloeospo-
rioides can also enhance its colonization of ripening fruits dynamically by
locally altering the pH of the fruit at the infection site to suit the increased
expression of pathogenicity factors and the enzymatic arsenal (Yakoby et al.,
2000; Prusky et al., 2001; Eshel et al., 2002; Prusky and Yakoby, 2003). In the
pathogen C. gloeosporioides, the gene pelB encoding for the enzyme pectate
lyase, a key factor for virulence, is expressed when the pH is > 6.0, a value at
which decay is initiated in the tissue (Yakoby et al., 2000, 2001). Also in the
case of A. alternata, another pathogen of mango fruits, the expression of the
endoglucanase gene AaK1 is maximal at pH levels > 6.0 (i.e. values that are
characteristic of decayed tissue). AaK1 is not expressed at the lower pH
D. Prusky et al. 216
values at which the pathogen is quiescent (Eshel et al., 2002). Ambient alkal-
ization by Colletotrichum is achieved by active secretion of ammonia, which is
produced as a result of protease activity and deamination of amino acids.
The pathogenicity of C. gloeosporioides and expression of the virulence factor
PL-B both depend on raising the ambient pH (Drori et al., 2003). This modu-
lation of environmental pH has been used as the basis for a new approach to
disease control in mango fruits, and is discussed below.
Management
Control of postharvest anthracnose can be achieved by eld management,
postharvest treatments or a combination of both. Management of mango
anthracnose in the eld involves cultural and chemical practices, as well as
cultivar selection.
Cultural control
Since the development of mango anthracnose is dependent on moisture or
high RH, orchards ideally should be established in areas with a well-dened
dry season, to allow for fruit development under conditions unfavourable
for disease development. In the tropics, mango owering usually occurs dur-
ing dry seasons, but anthracnose incidence of > 90% is common in fruits that
develop during the rainy season (Arauz, 1999). In contrast, the incidence and
severity of mango anthracnose can be close to zero in fruits that develop
completely in the dry season, without the application of any other control
measure (Arauz, 2000).
Considerable effort has been invested in understanding and managing
mango owering. Flowering can be advanced by several weeks by applying
potassium nitrate sprays to mature foliage (Nez-Elisea, 1985). The growth
retardant paclobutrazol, alone or followed by potassium nitrate sprays, can
also be used to advance owering (Nez-Elisea et al., 1993). Both treatments
could contribute to the manipulation of the owering season to a less sensi-
tive period. Field sanitation of the tree itself is difcult to practise. Elimina-
tion of dry panicles and mummied fruits is time consuming. Bagging results
in reduced anthracnose severity, but it also reduces the red colour of the fruit,
which could reduce consumer appeal (Hofman et al., 1997).
Resistant cultivars
Although all commercial mango cultivars are susceptible to anthracnose,
some are less susceptible than others; Tommy Atkins and Keitt are less
susceptible than Irwin, Kent, Haden and Edward (Campbell, 1992).
Chemical control
In extreme situations, in which fruit develops entirely under disease-favouring
conditions, seasonal applications of up to 25 sprays of protective and sys-
temic fungicides have been reported (Dodd et al., 1997). However, few fungi-
cides are approved in importing countries for use on mango. Therefore, the
Fruit Diseases 217
choice of fungicides depends on the intended destination of the fruit. Dithio-
carbamate fungicides are highly effective for anthracnose control but can be
used for only a few specic countries, whereas copper fungicides are recom-
mended, but their efcacy is lower (Arauz, 2000). Fungicides with erradicant
activity for mango anthracnose include benzimidazoles and the imidazole
prochloraz. Benomyl has been used under calendar-based schedules, usually
in a mix with protectant fungicides, to delay the build-up of resistance in the
pathogen population. Prochloraz has been used as a protectant or as an erad-
icant spray (Estrada et al., 1996), but is mainly used only as a postharvest
treatment.
Biological control
A number of microorganisms with in vitro or in vivo activity against C. gloeo-
sporioides have been isolated (Jeffries and Koomen, 1992), but few examples
had been used commercially in the eld until Korsten (2004) isolated Bacillus
spp. from leaf and fruit surfaces, and effectively controlled anthracnose of
mango. Postharvest control was achieved with semi-commercial preharvest
sprays or postharvest packing house dips, sprays or ultra-low-volume appli-
cations. Integrated treatments involving antagonists combined with quarter-
strength or recommended dosages of fungicides such as prochloraz or sodium
hypochlorite also effectively suppressed postharvest anthracnose of mango.
Commercializing the antagonists in South Africa (Korsten, 2004) and in the
Philippines proved to be difcult because of the limitations set by the local
registration guidelines, and the effect of product formulation on antagonist
performance in commercial applications.
Postharvest control
Traditionally, postharvest control of mango anthracnose has aimed at eradi-
cation of quiescent infections on the fruit. Such eradication is achieved com-
mercially by thermal and chemical treatments, or a combination of both
(McMillan, 1987). Dipping fruit in hot water alone is moderately efcient;
temperatures of 5055C for 315 min have been recommended, with the
higher temperatures corresponding to the shorter exposures. Fruit from Latin
America entering the USA market must undergo a quarantine hot water
treatment to eliminate fruit y (Ceratitis capitata and Anastrepha spp.) larvae;
the fruit is immersed in water at 46C for 90120 min, depending on variety
and fruit size. The efcacy of the fruit y quarantine treatment varies from 60
to 85% for elimination of anthracnose infections (McGuire, 1991). Hot water
treatments leave no chemical residue on the fruit and could be a good anthra-
cnose control option for organically-produced mangoes or for mangoes tar-
geted for markets in the USA, where no fungicides are currently approved
for postharvest use. Temperature and time controls are critical, because fruits
can easily be damaged by overexposure to heat. Several fungicides have been
applied after harvest to control mango anthracnose. Benomyl, at rates vary-
ing from 500 to 1000 g/ml, was used as a postharvest dip in the past, but its
use is no longer permitted. Thiabendazole at 10002000 g/ml is also effec-
tive, and there is interest in its registration for postharvest use with mango,
D. Prusky et al. 218
as it is currently used on other fruits such as citrus. Prochloraz can be used

with fruit shipped to the European Union (EU), at rates up to 1000 g/ml; its
efcacy ranges from 65% under very high disease pressure to 94% under
moderate disease pressure (Arauz, 2000). One advantage of benzimidazole
fungicides (i.e. benomyl or thiabendazole) is that they are also effective in
controlling stem-end rot caused by L. theobromae, which is the second most
important postharvest disease of mango in tropical areas. Imidazoles such as
prochloraz and imazalil are not effective against L. theobromae on mango
(Estrada et al., 1996). The combination of hot water and fungicides is the most
effective commercial postharvest treatment for the control of mango anthra-
cnose; the rate of fungicide application and the duration of exposure to hot
water are both lower, and efcacy is higher, than either treatment used sepa-
rately. The hot water and the fungicides can be applied sequentially or together.
Irradiation of fruit to control anthracnose has been attempted: gamma irra-
diation was not successful (Spalding and Reeder, 1986), but a short-wave
infrared radiation treatment developed in South Africa resulted in anthra-
cnose levels similar to those resulting from the commercial hot-water treat-
ment, and was much faster and less expensive (Saaiman, 1996a).
Prusky and Keen (1993) suggested that it might be possible to prolong
the period of fruit resistance and to delay the onset of anthracnose develop-
ment until the fruit ripens. The climacteric rise occurs simultaneously with
the production of ethylene and with changes in external and internal colour,
avour, aroma and rmness, and with a reduction in fruit resistance to fun-
gal attack (Prusky and Keen, 1993). Postharvest practices such as cold stor-
age and controlled atmosphere storage maintain resistance to decay by
delaying the ripening process, but there are two limitations to the potential
benet of this approach in mango. First, mangoes, similarly to many other
tropical and subtropical fruits, are sensitive to chilling and are injured at tem-
peratures < 1013C, depending on the cultivar and the duration of expo-
sure. Second, once the fruits are allowed to ripen under ambient conditions,
disease develops normally. Nevertheless, some progress has been made
towards anthracnose control through maintaining fruit resistance beyond
the onset of ripening. In the last decade, a better understanding of the physi-
ological basis of quiescent infections on tropical and subtropical climacteric
fruits has been achieved, mainly through the work of D. Prusky and his co-
workers (1993). The decline in antifungal compounds, which is brought
about by oxidative processes, can be delayed so that it occurs closer to full
ripeness. In avocadoes and mangoes, the concentrations of antifungal com-
pounds were enhanced and, consequently, the decline in concentration was
delayed, by exposure of fruit to an atmosphere containing 30% carbon diox-
ide (CO
2
) for 24 h, or by treatment of fruit with the antioxidant compound
butylated hydroxy anisole (Prusky, 1988; Kobiler et al., 1998). The delay
resulted in less disease in ripe fruit.
Postharvest biological control of anthracnose in mangoes has been
attempted. In an investigation with a strain of a Bacillus sp. that exhibited in
vitro activity against C. gloeosporioides, it was found that disease control in
vivo was obtained when fruits were inoculated with the bacterium 24 h prior
Fruit Diseases 219
to inoculation with the fungus, but not when fruit were inoculated with the
pathogen rst (Korsten et al., 1992), which indicated that the quiescent phase
of the fungus was not affected by the antagonist. Other approaches to disease
control using biological methods included the use of a non-pathogenic strain
of Colletotrichum magna that colonizes the fruit endophytically and prevents
infection by C. gloeosporioides (Prusky et al., 1993); and the expression of an
antifungal peptide in the yeast Saccharomyces, which controlled postharvest
diseases caused by C. coccodes (Jones and Prusky, 2001).
7.3 Alternaria Rot (Black Spot)
Alternaria alternata (Fr.:Fr.) Keissl. causes black spot of mango. Conidiophores,
arising singly or in small groups, are simple or branched, straight or bent,
and sometimes geniculate, and pale- to mid-olivaceous or gold in colour, and
smooth in texture. The conidia are oboclavata; they are borne in long chains
in culture, and most have three to ve septa.
Pathogenesis
Germinated conidia penetrate mainly through wounds and specically
through lenticels of the fruits, and then become quiescent.
Symptoms
Alternaria rot of mango has been increasingly reported as an important
pathogen that causes blossom disease and postharvest fruit rot in ripening
fruits in Australia, Egypt, India, Israel and South Africa. The symptoms are
small, black circular spots that develop around the lenticels. Initially, the
spots are concentrated around the stem end of the fruits, where there are
large numbers of lenticels. The spots can grow and coalesce to become a sin-
gle spot that covers a signicant part of the fruit surface. At rst, the decay is
rm and does not penetrate the pulp more than 12 mm, but later the disease
progresses into the esh, which darkens and becomes soft (Prusky et al.,
1983). Symptoms of alternaria rot are more limited, darker and rmer than
those of anthracnose. The former pathogen also attacks mango leaves, and
symptoms can be observed throughout the year. The pathogen may also
attack mango inorescences, resulting in a signicant decrease in fruit set.
Epidemiology
The main sources of inoculum are conidia released from infected leaves,
twigs and inorescences; however, Alternaria spores easily can be found in all
the dry tissues of mango trees in the orchard. Conidia are transferred to the
fruit by air currents and in dew runoff (Ploetz et al., 1994). Germination of
conidia depends on the RH in the orchard during fruit growth. The area of
quiescent Alternaria infection on mango fruit at harvest increased as the num-
ber of hours of exposure to RH 80% increased over 320 h (Prusky et al.,
D. Prusky et al. 220
1983). Regions with the highest potential for disease incidence are located
close to the 8590% RH isolines during the fruit growth period. The interme-
diate regions lie between 75 and 85% RH, and the lowest potential risk is in
the dry regions, where the prevailing RH is < 75% (Prusky et al., 1992).
Mechanisms of fungal pathogenicity and fruit resistance
As in the case of anthracnose, the mechanism of resistance against Alternaria
is also related to the presence of high concentrations of preformed antimicro-
bial compounds (Droby et al., 1986, 1987). Susceptibility was found to depend
on the decline of the compound to subfungitoxic concentrations, which
occurred faster in Tommy Atkins than in Keitt.
Alternaria alternata can also alter the local fruit pH at the infection site
dynamically, to match the increased expression of pathogenicity factors and
the enzymatic arsenal (Eshel et al., 2002; Niem et al., 2007). In the case of A.
alternata, the expression of the endoglucanase gene AaK1 was found to be
maximal at pH levels > 6.0, which are characteristic of decayed tissue; it was
not expressed at the lower pH values at which the pathogen was quiescent
(Eshel et al., 2002). Alkalization of the ambient environment by Alternaria is
achieved by active secretion of ammonia (Eshel et al., 2002), probably as a
result of protease activity and deamination of amino acids. This modulation
of environmental pH has been used as the basis of a new approach to disease
control in mango fruits, and is discussed below.
Management
The losses caused by alternaria rot can be minimized by a regular eld-
spraying programme and by postharvest fungicide treatments.
Field and postharvest chemical control
Preharvest treatments with dithiocarbamate fungicides inhibit the develop-
ment of latent infection. Three sprays with the protectant fungicide maneb,
starting 2 weeks after fruit set, are most effective for disease control (Prusky
et al., 1983). However, since quiescent infections do not develop until after
harvest and ripening, the application of a postharvest treatment by spraying
the fruits on the packing line with 900 g/ml prochloraz is simpler and more
efcient than the preharvest fungicide treatment.
Control of alternaria rot is signicantly improved by a combination of
physical and chemical treatments. The physical treatment includes a 1520 s
hot water spraying and brushing (HWB) treatment at temperatures between
50 and 55C (Prusky et al., 1999). This new approach improved fruit quality
at the same time as it reduced disease incidence. If this physical treatment is
followed by a 250 g/ml prochloraz spray it can further improve disease
control. Prusky et al. (1999) concluded that the type and strength of the post-
harvest treatment should be varied according to the level of quiescent infection
Fruit Diseases 221
of A. alternata at the time of harvest. Although the fungicide prochloraz is
very effective for the control of postharvest disease, a milder postharvest treat-
ment, such as chlorine at 500 g/ml, can be applied to fruit in which a low
incidence of quiescent infections is found at harvest (Prusky et al., 2002).
This postharvest physical-chemical treatment has been further improved
in light of the recent nding that A. alternata pathogenicity may modulate the
pH of the host environment to promote colonization (Eshel et al., 2002; Prusky
and Yakoby, 2003; Prusky and Lichter, 2007). Application of a combination of
HWB for 1520 s, followed by spraying with 50 mM hydrochloric acid (HCl),
effectively controlled alternaria rot in stored mango fruit. Similar HWB treat-
ments followed by spraying with reduced concentrations of prochloraz at
45 g/ml in 50 mM HCl inhibited alternaria rot development better than
treatment with HCl alone (Prusky et al., 2006). The enhancement of prochlo-
raz activity in acidied solutions was attributed to its enhanced solubility
under acidic conditions, which resulted in an increase in the fungicidally
active ingredient in the solution. This technology represents the latest devel-
opment in the control of postharvest diseases, and provides a simple treat-
ment for the control of diseases that alkalinize the host environment,
including both alternaria rot and anthracnose.
7.4 Stem-end Rots
Stem-end rots of mango fruit present one of

the most serious postharvest
problems that affect this industry

worldwide. They become more prominent
as orchards become older. Losses increase when fruits are stored for pro-
longed periods at low temperatures or when fruits ripen at temperatures
> 28C. The stem-end rot diseases are caused

by a variety of fungal patho-
gens including: D. dominicana (anamorph of Botryosphaeria dothidea), Dothi-
orella mangiferae, L. theobromae (Botryodiplodia theobromae), Phomopsis mangiferae
and Pestalotiopsis mangiferae, among which Botryosphaeria

is the dominant
pathogen (Darvas 1991; Johnson et al., 1992; Sangchote, 1998). Stem-end rot
diseases cause heavy losses in mangoes during storage (Prusky et al., 1992;
Mitra, 1997; Kobiler et al., 2001).
Pathogenesis
Johnson and co-workers (1993) have suggested that spores of the pathogen
may germinate and penetrate into the host tissue through wounds, and
remain as endophytes in the branches of mango trees.
Symptoms
Depending on the fungus involved, a variety of symptoms may develop at
the stem and as the fruit ripens. Infections by L. theobromae (B. theobromae), D.
dominicana synonym Fusicoccum aesculi (anamorph of B. dothidea) and D.
mangiferae, cause the formation of diffuse, water-soaked tissue that spreads
D. Prusky et al. 222
from the stem end as ngerlike projections, which darken and coalesce into
circumpedicular lesions or lobed margins (Johnson et al.,

1992; Slippers et al.,
2004, 2005). Necrosis remains beneath the cuticle and may penetrate through-
out the fruit esh. Supercial mycelia may appear around the pedicel and
ruptures of the skin or, in some cases, may penetrate through the epidermis.
The ascomata of D. mangiferae are initially embedded, either separately or
grouped in complex stromata, and they nally erupt through the epidermis
and open. The spore wall is dark and thick, and becomes thinner towards the
end. Conidiophores are hyaline, cylindrical, smooth and branched at the
base. A watery uid may drain from the stem or from surface ruptures (Kor-
sten, 2004). Diseased fruit could infect a whole box of fruits by direct contact,
and thereby spread the pathogen in the box. In the case of injured fruit,
lesions could appear at some distance from the stem. Stem-end rot diseases
also have a major effect on the avour of the fruit.
The disease may also cause tip and branch dieback and cankers on mango
trees (Ramos et al., 1991). Anamorph morphology is commonly used to iden-
tify Botryosphaeria spp. (Jacobs and Rehner, 1998; Slippers et al., 2004), but the
morphological

distinctions among the anamorphs of some of the closely
related

species are not clear. Recent studies based on DNA sequence

data have
highlighted taxonomic groups and relationships in

Botryosphaeria (Jacobs and
Rehner, 1998; Slippers et al., 2004). These data, combined with morphological

characteristics, could clarify the current taxonomic confusion.
Phomopsis mangiferae is a weak parasite of less economic importance than
the species above; it produces a dark, circumpeduncular lesion with dened
edges that spreads relatively slowly but penetrates deeply into the esh.
Fruiting bodies may develop on the affected surface. Phomopsis mangiferae
can also be distinguished by a dark pinhead-size pycnidial fruiting body
(Johnson et al., 1992). The lesion caused by Phomopsis may be similar to the
stem-end symptoms cause by C. gloeosporioides and A. alternata. However, the
lesions of the latter two diseases penetrate only to a depth of 1020 mm.
Lesions of L. theobromae can be distinguished by their wrinkled black
edges which have a velvety appearance. In the dark zone, pycnidial masses
are formed (Johnson et al., 1992). In affected plants, twigs die back from the
tips and into old wood, giving a scorched appearance to the limb with abun-
dant gum secretions from branches, stems and the main trunk. Pestalotiopsis
mangiferae appears as silvery grey spots that vary in size, and are usually
sharply outlined by a dark border. The fungus may appear as a member of
the complex of stem-rot pathogens.
Epidemiology
The pathogens causing stem-end rots initiate inoculation in the orchards as
the trees age. They are enabled to do so by mismanagement or neglect of
orchards, and are affected by periods of rain and high RH at the beginning
and end of the dry season (Johnson et al., 1991; Lonsdale, 1993; Johnson and
Sangchote, 1994; Gonzlez et al., 1999). Hyphae of the fungi colonize the
Fruit Diseases 223
oral parts of mango trees, develop endophytically in healthy tissue of all
parts of mango plants, especially in the fruit pedicel, and remain quiescent
until the fruit matures, at which stage the parasite is ready to infect through
the stem end by developing in the xylem vessels of the maturing fruit (John-
son et al., 1992). Pathogens can colonize ower parts, remain inactive pend-
ing button abscission and then penetrate the stem end, as in the case of
Diplodia natalensis in citrus (Nadel, 1944), of which no sexual stage has been
found in nature. Fruits can be also infected at the stem by the soilborne patho-
gen L. theobromae (anamorph of B. theobromae), if the fruits are placed on the
ground (Johnson et al., 1992) after harvest. It is also possible that infection can
result from transfer of spores by insects; decayed fruits produce volatiles,
which are hypothesized to be attractants of an insect that could be a vector of
the fungal spores (Nago and Matsumoto, 1994).
The range of pathogens that cause stem-end rot is inuenced by tempera-
ture, moisture stress and the nutrition status of the host. The initial systemic
infection plays a crucial role in establishing blossom-blight infection. How-
ever, secondary infection is apparently an even more important factor in devel-
opment and incidence of soft stem-end rots. Secondary infections occur when
rain washes spores away from various inoculum sources such as leaves and
stems (Saaiman, 1996b). Endophytic Botryosphaeria spp. were found to be espe-
cially prominent in trees continually exposed to water shortage and salt stress
(Grobler et al., 2002), and Botryosphaeria spp. were found to move into develop-
ing fruit, resulting in postharvest stem-end rot development (Lonsdale, 1993).
Management
Field and postharvest control
A variety of postharvest practices can be used to delay disease develop-
ment. These include low-temperature storage and modied- or controlled-
atmosphere storage; however, the management of stem-end rots is far from
being perfected.
Cultural control
Johnson et al. (1992) demonstrated that infection of mango fruit before har-
vest occurred through endophytic colonization of pedicel tissue by Botry-
osphaeria spp. present from previous growth ushes, and the possibility of
pruning to promote growth ushing was tested as a means to reduce inocu-
lum in the stem tissue from which new-season inorescences emerged. Cooke
et al. (1998) reported that the levels of endophytic organisms such as Botry-
osphaeria spp. were reduced signicantly when a pruning programme was
implemented in mango orchards as a preharvest control measure. Korsten
(2006) found that prevention of water stress during fruit development and
maturation, and avoidance of placing fruits on the ground suppressed dis-
ease development. He also suggested that harvesting of immature fruit
should be avoided; fruit should be cooled to 13C immediately after harvest
and chemical treatment, and stored in a well-ventilated place.
D. Prusky et al. 224
Chemical control
Postharvest control of Botryosphaeria spp. was achieved by postharvest dip-
ping, spraying or ultra-low-volume application of benomyl (where possible).
Prochloraz or sodium hypochlorite also effectively suppressed postharvest
stem-end rot of mango (Plan et al., 2002; Korsten, 2006). A combined treat-
ment of wax and hot water (55C) provide very effective control of most
postharvest pathogens (Sangchote, 1998), but in some cases partial-vacuum
inltration improved disease control, which suggests that control efciency
may have been reduced because the fungicide did not reach the pathogen
(Plan et al., 2002).
A treatment consisting of HWB and prochloraz followed by 2,4-dichloro-
phenoxyacetic acid (2,4-D) diluted in wax, reduced side and stem-end decay
by 5070%, and improved fruit quality during prolonged storage (Kobiler et al.,
2001). The best control was obtained by concentrations of 2,4-D ranging from
75 to 175 g/ml; this efciently controlled side rots caused by A. alternata and
the stem-end rots caused by A. alternata, Phomopsis spp. and Lasiodiplodia
spp. The combination of HWB, prochloraz application and 2,4-D treatment
reduced the incidence of stem-end rot after 4 weeks of storage at 14C and 7
days of shelf life at 20C from 86 to 10% in Tommy Atkins and from 63 to
12% in Keith (Kobiler et al., 2001).
Biological control
Bacillus licheniformis, on its own or alternated with copper oxychloride, has
been evaluated as a preharvest spray treatment to control mango fruit dis-
eases. Preharvest applications of B. licheniformis at 3-week intervals from
owering until harvest controlled moderate levels of anthracnose, and of soft
rot caused by Botryosphaeria, which suggests a potential treatment for com-
mercial preharvest applications (Silimela and Korsten, 2006).
7.5 Other Minor Diseases
Other pathogens may attack mango fruits after harvest through occasional
wounds, and cause severe diseases, such as black mould caused by Aspergillus
spp. and transit rot caused by Rhizopus spp. Disease control begins in the eld,
and is followed by postharvest sanitation, and avoidance of latex burn (stain)
and mechanical injury. A hot water treatment consisting of 46C for 60120
min and fungicides can be used, depending on the cultivar (Spalding and
Reeder, 1986). HWB at 55C for 20 s shows good control (Prusky et al., 1999).
7.6 Conclusions
Long periods in transit, new marketing approaches for mangoes (e.g. Ready
to Eat) and stringent international standards and requirements have raised
the need for improved approaches to disease control, in order to preserve
fruit quality. Integrated postharvest treatment has provided a more durable,
Fruit Diseases 225
consistent, sustainable and practical solution than previously available to
enable producers to control postharvest diseases. Ensuring product quality
and safety starts in the orchard, with proper handling conditions and is fol-
lowed by optimal postharvest treatments; however, the combination of post-
harvest treatments with the proper use of the cold-chain concept may provide
the ultimate in favourable conditions for preserving fruit quality.
References
Arauz, L.F. (1999) Optimizacin econmica del combate de enfermedades en cultivos:
un ejemplo basado en la antracnosis del fruto del mango. (Abstract). In: XI Con-
greso Nacional Agron. Recursos Naturales, V Congreso Nacional Entomol., IV
Congreso Nacional Fitopatol., III Congreso Nacional Suelos I Congreso Nacional
Ext. Agrc. Forestal. Vol. II. San Jos, Costa Rica, p. 61.
Arauz, L.F. (2000) Mango anthracnose: economic impact and current options for inte-
grated management. Plant Disease 84, 600611.
Ardi, R., Kobiler, I., Jacoby, B., Keen, N.T. and Prusky, D. (1998) Involvement of epicat-
echin biosynthesis in the activation of the mechanism of resistance of avocado fruits
to Colletotrichum gloeosporioides. Physiological and Molecular Plant Pathology
53, 269285.
Bailey, J.A. (1997) rDNA sequencing: an essential guide to the taxonomy and diagnosis
of Colletotrichum. In: Dehne, H.-W., Adam, G., Diekmann, M., Frahm, J., Mauler-
Machnik, A. and van Halteren, P. (eds) Diagnosis and Identication of Plant Patho-
gens, Vol. 11, Developments in Plant Pathology. Kluwer Academic, Dordrecht, the
Netherlands, pp. 4751.
Banik, A.K., Kaiser, S.A.K.M. and Dhua, R.S. (1998) Inuence of temperature and hu-
midity on the growth and development of Colletotrichum gloeosporioides and
Diplodia natalensis causing postharvest fruit rot of mango. Advances in Plant Sci-
ences 11, 5157.
Beno-Moualem, D. and Prusky, D. (2000) Early events during quiescent infection devel-
opment by Colletotrichum gloeosporioides in unripe avocado fruits. Phytopathology
90, 553559.
Campbell, R.J. (ed.) (1992) A Guide to Mangoes in Florida. Fairchild Tropical Garden,
Miami, Florida.
Coates, L.M., Muirhead, I.F., Irwin, J.A.G. and Gowanlock, D.H. (1993) Initial infection
processes by Colletotrichum gloeosporioides on avocado fruit. Mycological Re-
search 97, 13631370.
Cooke, A.W., van der Kruyssen, A. and Johnson, G.I. (1998) The effect of commercial
pruning on colonization of mango by endophytic Dothiorella species. In: Champ,
B.R., Highley, E. and Johnson, G.I. (eds) Post-harvest Handling of Tropical Fruit.
Australian Centre for International Agricultural Research (ACIAR), Canberra,
pp. 128130.
Darvas, J.M. (1991) Dothiorella dominicana, a new mango pathogen in South Africa.
Phytophylactica 23, 295298.
de Jong, J.C., McCormark, B.J., Smirnoff, N. and Talbot, N.J. (1997) Glycerol generates
turgor in rice blast. Nature 389, 244245.
Dickman, M.B., Patil, S.S. and Kolattukudy, P.E. (1982) Purication, characterization and
role in infection of an extracellular cutinolytic enzyme from Colletotrichum gloeo-
sporioides Penz. on Carica papaya L. Physiological Plant Pathology 20, 333347.
D. Prusky et al. 226
Dodd, J.C., Estrada, A.B., Matcham, J., Jeffries, P. and Jeger, M.J. (1991) The effect of
climatic factors on Colletotrichum gloeosporioides, causal agent of mango anthra-
cnose, in the Philippines. Plant Pathology 40, 568575.
Dodd, J.C., Prusky, D. and Jeffries, P. (1997) Fruit diseases. In: Litz, R.E. (ed.) The Mango:
Botany, Production and Uses. CAB International, Wallingford, UK, pp. 257280.
Droby, S., Prusky, D., Jacoby, B. and Goldman, A. (1986) Presence of an antifungal com-
pound and its relation in the latency of Alternaria alternata in unripe peel of mango
fruits. Physiological and Molecular Plant Pathology 29, 173183.
Droby, S., Prusky, D. and Jacoby, D. (1987) Induction of an antifungal agent in unripe
mango fruits to demonstrate their involvement in latent infections of Alternaria al-
ternata. Physiological and Molecular Plant Pathology 30, 285292.
Drori, N., Kramer-Haimovich, H., Rollins, J., Dinoor, A., Okon, Y., Pines, O. and Prusky,
D. (2003) A combination of external pH and nitrogen assimilation affect secretion
of the virulence factor pectate lyase by C. gloeosporioides. Applied and Environ-
mental Microbiology 69, 32583262.
Eshel, D., Miara, I., Ailing, T., Dinoor, A. and Prusky, D. (2002) pH regulates endogluca-
nase expression and virulence of Alternaria alternata in persimmon fruits. Molecular
Plant Microbe Interactions 15, 774779.
Estrada, A.B., Dodd, J.C. and Jeffries, P. (1993) Effect of environment on the in vitro
growth and development of Colletotrichum gloeosporioides isolates from the Phil-
ippines. Acta Horticulturae 341, 360370.
Estrada, A.B., Jeffries, P. and Dodd, J.C. (1996) Field evaluation of a predictive model
to control anthracnose disease of mango in the Philippines. Plant Pathology 45,
294301.
Fitzell, R.D. and Peak, C.M. (1984) The epidemiology of anthracnose disease of mango:
inoculum sources, spore production and dispersal. Annals of Applied Biology 104,
5359.
Fitzell, R.D., Peak, C.M. and Darnell, R.E. (1984) A model for estimating infection levels
of anthracnose disease of mango. Annals of Applied Biology 104, 451458.
Flaishman, M.A. and Kolattukudy, P.E. (1994) Timing of fungal invasion using hosts rip-
ening hormone as a signal. Proceedings of the National Academy of Sciences USA
91, 65796583.
Freeman, S., Katan, T. and Shabi, E. (1998) Characterization of Colletotrichum species
responsible for anthracnose diseases of various fruits. Plant Disease 82, 596605.
Gantotti, B.V. and Davis, M.J. (1993) Pectic zymogram analysis for characterizing
genetic diversity of the mango anthracnose pathogen. Acta Horticulturae 341,
353359.
Gonzlez, E., Umaa, G. and Arauz, L.F. (1999) Fluctuacin poblacional de Botryodi-
plodia theobromae Pat. en mango. Agronoma Costarricense 23, 2129.
Grobler, L., Swart, S.H. and Korsten, L. (2002) Preliminary survey of a mango nursery
and associated orchards to establish the status of endophytic Botryosphaeria
spp. South African Mango Growers Association Yearbook 22, 104106.
Guetsky, R., Kobiler, I., Wang, X., Perlman, N., Gollop, N., Avila-Quezada, G., Hadar, I.
and Prusky, D. (2005) Metabolism of the avonoid epicatechin by laccase of Col-
letotrichum gloeosporioides and its effect on pathogenicity on avocado fruits. Phy-
topathology 95, 13411348.
Hawthorne, B.T. (1989) Effects of cultural practices on the incidence of storage rots in
Cucurbita spp. New Zealand Journal of Crop and Horticultural Science 17, 4954.
Hofman, P.J., Smith, L.G., Joyce, D.C., Johnson, G.I. and Meiburg, G.F. (1997) Bagging
of mango (Mangifera indica cv. Keitt) fruit inuences fruit quality and mineral
composition. Postharvest Biology and Technology 12, 8391.
Fruit Diseases 227
Howard, R.J., Ferrari, M.A., Roach, D.H. and Money, N.P. (1991) Penetration of hard
substrates by a fungus employing enormous turgor pressures. Proceedings of the
National Academy of Sciences USA 88, 1128111284.
Jacobs, K.A. and Rehner, S.A. (1998) Comparison of cultural and morphological charac-
ters and ITS sequences in anamorphs of Botryosphaeria and related taxa. Mycolo-
gia 90, 601610.
Jeffries, P. and Koomen, I. (1992) Strategies and prospects for biological control of dis-
eases caused by Colletotrichum. In: Bailey, J.A. and Jeger, M.J. (eds) Colletotrichum:
Biology, Pathology and Control. CAB International, Wallingford, UK, pp. 337357.
Johnson, G.I. and Sangchote, S. (1994) Control of post-harvest diseases of tropical fruits:
challenges for the 21st century. In: Champ, B.R., Highley, E. and Johnson, G.I. (eds)
Post-harvest Handling of Tropical Fruit. Australian Centre for International Agricul-
tural Research (ACIAR), Canberra, pp. 140161.
Johnson, G.I., Mead, A.J., Cooke, A.W. and Dean, J.R. (1991) Mango stem end rot patho-
gens infection levels between owering and harvest. Annals of Applied Biology
119, 465473.
Johnson, G.I., Mead, A.J., Cooke, A.W. and Dean, J.R. (1992) Mango stem end rot patho-
gens fruit infection by endophytic colonization of the inorescence and pedicel.
Annals of Applied Biology 120, 225234.
Johnson, G.I., Cooke, T. and Mead, A.J. (1993) Infection and quiescence of mango stem-
end rot pathogens. Acta Horticulturae 341, 329336.
Jones, R.W. and Prusky, D. (2001) Expression of an antifungal peptide in Saccharomy-
ces: a new approach for biocontrol of the postharvest disease caused by Colletotri-
chum coccodes. Phytopathology 92, 3337.
Kobiler, I., Reved, R., Artez, L. and Prusky, D. (1998) Antifungal compounds regulating
quiescent disease in mango. 1998. In: Johnson, G.I., Higley, E. and Joyce, D.C.
(eds) Australian Centre for International Agricultural Research (ACIAR) Proceedings
80. ACIAR, Canberra, pp. 109114.
Kobiler, I., Shalom, Y., Roth, I., Akerman, M., Vinokour, Y., Fuchs, Y. and Prusky, D.
(2001) Effect of 2,4-dichlorophenoxyacetic acid on the incidence of side and stem-
end rots in mango fruits. Postharvest Biology and Technology 23, 2332.
Korsten, L. (2004) Biological control in South Africa: can it provide a sustainable solu-
tion for control of fruit diseases? South African Journal of Botany 70, 128139.
Korsten, L. (2006) Advances in control of postharvest diseases in tropical fresh produce.
International Journal of Postharvest Technology and Innovation 1, 4861.
Korsten, L., Lonsdale, J.H., De Villiers, E.E. and De Jager, E.S. (1992) Pre-harvest con-
trol of mango diseases. South African Mango Growers Association Yearbook 12,
7278.
Latunde-Dada, A.O., OConnell, R.J., Nash, C. and Lucas, J.A. (1999) Stomatal penetra-
tion of cowpea (Vigna unguiculata) leaves by a Colletotrichum species causing la-
tent anthracnose. Plant Pathology 48, 777785.
Liu, Z.M. and Kolattukudy, P.E. (1998) Identication of a gene product induced by hard-
surface contact of Colletotrichum gloeosporioides conidia as a ubiquitin-conjugating
enzyme by yeast complementation. Journal of Bacteriology 180, 35923597.
Lizada, C. (1993) Fruit handling systems in developing countries. In: Postharvest Han-
dling of Tropical Fruits. Australian Centre for International Agricultural Research
(ACIAR) Publication 50. ACIAR, Canberra, p. 102.
Lonsdale, J.H. (1993) Preliminary results on the mode of infection of Nattrassia mangifer-
ae in mango. South African Mango Growers Association Yearbook 13, 9799.
McGuire, R.G. (1991) Concomitant decay reductions when mangoes are treated with
heat to control infestations of Caribbean fruit ies. Plant Disease 75, 946949.
D. Prusky et al. 228
McMillan, R.T., Jr (1987) Effectiveness of various postharvest treatments for mango de-
cay control. Proceedings of the Florida State Horticultural Society 100, 79.
Mitra, S. (1997) Mango. In: Mitra, S.K. and Baldwin, E.A. (eds) Postharvest Physiology
and Storage of Tropical and Subtropical Fruits. CAB International, Wallingford, UK,
pp. 122185.
Money, N.P. and Howard, R.J. (1996) Conrmation of a link between fungal pigmenta-
tion, turgor pressure and pathogenicity using a new method of turgor measurement.
Fungal Genetics and Biology 20, 217227.
Nadel, M. (1944) Anatomical study of the button of Shamouti oranges in relation to
stem-end rot. Palestine Journal of Botany, Rehovot Series 4, 166170.
Nago, H. and Matsumoto, M. (1994) An ecological role of volatiles produced by Lasio-
diplodia theobromae. Bioscience Biotechnology Biochemistry 58, 12671272.
Niem, J., Miyara, I., Reuveni, M., Ettedy, Y., Flaishman, M. and Prusky, D. (2007) Core
rot development in Red Delicious apples is affected by locule susceptibility to Al-
ternaria alternata. Phytopathology 97, 14151421.
Nez-Elisea, R. (1985) Flowering and fruit set of a monembryonic and a polyembry-
onic mango as inuenced by potassium nitrate sprays and shoot decapitation. Pro-
ceedings of the Florida State Horticultural Society 98, 179183.
Nez-Elisea, R., Davenport, T.L. and Caldeira, M.L. (1993) Bud initiation and morpho-
genesis in Tommy Atkins mango as affected by temperature and triazole growth
retardants. Acta Horticulturae 341, 192198.
Pain, N.A., Green, J.R., Jones, G.L. and OConnell, R.J. (1996) Composition and organi-
sation of extracellular matrices around germ tubes and appressoria of Colletotri-
chum lindemuthianum. Protoplasma 190, 119130.
Plan, M.R.R., Joyce, D.C., Ogle, H.J. and Johnson, G.I. (2002) Mango stem-end rot
(Botryosphaeria dothidea) disease control by partial-pressure inltration of fungi-
cides. Australian Journal of Experimental Agriculture 42, 625629.
Ploetz, R.C. (1994) Anthracnose. In: Ploetz, R.C., Zentmyer, G.A., Nishijima, W.T.,
Rohrbach, K.G. and Ohr, H.D. (eds) Compendium of Tropical Fruit Diseases. Amer-
ican Phytopathological Society (APS) Press, St Paul, Minnesota, pp. 3536.
Ploetz, R.C. and Prakash, O. (1997) Foliar, oral and soilborne diseases. In: Litz, R.E.
(ed.) The Mango: Botany, Production and Uses. CAB International, Wallingford,
UK, pp. 281325.
Ploetz, R.C., Zentmyer, G.A., Nishijima, W.T., Rohrbach, K.G. and Ohr, H.D. (1994)
Mango. In: Ploetz, R.C., Zentmyer, G.A., Nishijima, W.T., Rohrbach, K.G. and Ohr,
H.D. (eds) Compendium of Tropical Fruit Diseases. American Phytopathological
Society (APS) Press, St Paul, Minnesota, pp. 3340.
Ploetz, R.C., Benscher, D., Vzquez, A., Colls, A., Nagel, J. and Schaffer, B. (1996) Man-
go decline: research in Florida on an apparently wide-spread disease complex.
Acta Horticulturae 455, 547553.
Podila, G.K., Rogers, L.M. and Kolattukudy, P.E. (1993) Chemical signals from avocado
surface wax trigger germination and appressorium formation in Colletotrichum
gloeosporioides. Plant Physiology 103, 267272.
Prusky, D. (1988) The use of antioxidants to delay the onset of anthracnose and stem end
rot in avocado fruits after harvest. Plant Disease 72, 381384.
Prusky, D. (1996) Pathogen quiescence in postharvest diseases. Annual Reviews of Phy-
topathology 34, 413434.
Prusky, D. and Keen, N.T. (1993) Involvement of preformed antifungal compounds in
the resistance of subtropical fruits to fungal decay. Plant Disease 77, 114119.
Prusky, D. and Lichter, A. (2007) Activation of quiescent infections by postharvest patho-
gens during transition from the biotrophic to the necrotrophic stage. FEMS Micro-
biological Letters 268, 18.
Fruit Diseases 229
Prusky, D. and Plumbley, R.A. (1992) Quiescent infections of Colletotrichum in tropical
and subtropical fruits. In: Bailey, J.A. and Jeger, M.J. (eds) Colletotrichum: Biology,
Pathology and Control. CAB International, Wallingford, UK, pp. 289307.
Prusky, D. and Yakoby, N. (2003) Pathogenic fungi: leading or led by ambient pH?
Molecular Plant Pathology 4, 509516.
Prusky, D., Fuchs, Y. and Yanko, U. (1983) Assessment of latent infections as a basis for
control of postharvest disease of mango. Plant Disease 67, 816818.
Prusky, D., Plumbley, R. and Kobiler, I. (1991) Modulation of natural resistance of avo-
cado fruits to Colletotrichum gloeosporioides by CO
2
. Physiological and Molecular
Plant Pathology 39, 325334.
Prusky, D., Gat, Z. and Burd, P. (1992) Effect of relative humidity during mango growth
on the incidence of quiescent infections of Alternaria alternata. Plant Disease 77,
249252.
Prusky, D., Freeman, S., Rodriguez, R.J. and Keen, N.T. (1993) A nonpathogenic mutant
strain of Colletotrichum magna induces resistance to C. gloeosporioides in avocado
fruits. Molecular Plant-Microbe Interactions 7, 326333.
Prusky, D., Wattad, C. and Koliber, I. (1996) Effect of ethylene on the activation of qui-
escent infections of Colletotrichum gloeosporioides in avocado fruits. Molecular
Plant-Microbe Interactions 9, 864868.
Prusky, D., Fuchs, I., Kobiler, I., Roth, I., Weksler, A., Shalom, Y., Falik, E., Zauberman,
G., Pesis, E., Akerman, M., Yekutiely, O., Weisblum, A., Regev, R. and Artes, L.
(1999) Effect of hot water brushing, prochloraz treatment and waxing on the inci-
dence of black spot decay caused by Alternaria alternata in mango fruit. Postharvest
Biology and Technology 15, 165174.
Prusky, D., McEvoy, J.L., Leverentz, B. and Conway, L.S. (2001) Local modulation of host
pH by Colletotrichum species as a mechanism to increase virulence. Molecular
Plant Microbe Interactions 14, 11051113.
Prusky, D., Shalom, Y., Kobiler, I., Akerman, M. and Fuchs, Y. (2002) The level of quies-
cent infection of Alternaria alternata in mango fruits at harvest determines the post-
harvest treatment applied for the control of rots during storage. Postharvest Biology
and Technology 25, 339347.
Prusky, D., Kobiler, I., Akerman, M. and Miyara, I. (2006) Effect of acidic solutions and
acidic prochloraz on the control of postharvest decay caused by Alternaria alter-
nata in mango and persimmon fruits. Postharvest Biology and Technology 42,
134141.
Ramos, L.J., Lara, S.P., McMillan, R.T. Jr and Narayanan, K.R. (1991) Tip dieback of mango
(Mangifera indica) caused by Botryosphaeria ribis. Plant Disease 75, 315318.
Saaiman, W.C. (1996a) Short wave infra-red radiation as an alternative to the hot water
bath to control postharvest decay in mangoes. Acta Horticulturae 455, 773779.
Saaiman, W.C. (1996b) Preliminary report on the time of infection of the soft brown rot
pathogen Nattrassia mangiferae in mango. South African Mango Growers Associa-
tion Yearbook 16, 5557.
Sangchote, S. (1998) Fruit rots of mangosteen and their control. In: Champ, B.R., High-
ley, E. and Johnson, G.I. (eds) Post-harvest Handling of Tropical Fruit. Australian
Centre for International Agricultural Research (ACIAR), Canberra, pp. 8186.
Sherriff, C., Whelan, M.J., Arnold, G.M., Lafay, J.-F., Brygoo, Y. and Bailey, J.A. (1994)
Ribosomal DNA sequence analysis reveals new species groupings in the genus Col-
letotrichum. Experimental Mycology 18, 121138.
Silimela, M. and Korsten, L. (2006) Evaluation of pre-harvest Bacillus licheniformis
sprays to control mango fruit diseases. Crop Protection 26, 14711481.
Slippers, B., Crous, P.W., Denman, S., Coutinho, T.A., Wingeld, B.D. and Wingeld, M.J.
(2004) Combined multiple gene genealogies and phenotypic characters differentiate
D. Prusky et al. 230
several species previously identied as Botryosphaeria dothidea. Mycologia 96,
83101.
Slippers, B., Johnson, G.I., Crous, P.W., Coutinho, T.A., Wingeld, B.D. and Wingeld,
M.J. (2005) Phylogenetic and morphological re-evaluation of the Botryosphaeria
species causing diseases of Mangifera indica. Mycologia 97, 99110.
Spalding, D.H. and Reeder, W.F. (1986) Decay and acceptability of mangoes treated
with combinations of hot water, imazalil, and g-radiation. Plant Disease 70, 1149
1151.
Sreenivasaprasad, S., Mills, P.R., Meehan, B.M. and Brown, A.E. (1996) Phylogeny and
systematics of 18 Colletotrichum species based on ribosomal DNA spacer sequenc-
es. Genome 39, 499512.
Thompson, J.F., Mitchell, F.G. and Kasmire, R.F. (2002) Cooling horticultural commodi-
ties. In: Kader, A.A. (ed.) Postharvest Technology of Horticultural Crops. University
of California Press, Davis, California, pp. 97112.
Yakoby, N., Kobiler, I., Dinoor, A. and Prusky D. (2000) pH regulation of pectate lyase
secretion modulates the attack of Colletotrichum gloeosporioides on avocado fruits.
Applied and Environmental Microbiology 66, 10261030.
Yakoby, N., Zhou, R., Kobiler, I., Dinoor, A. and Prusky, D. (2001) Development of Col-
letotrichum gloeosporioides restriction enzyme-mediated integration mutants as
biocontrol agents against anthracnose disease in avocado fruits. Phytopathology
91, 143148.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 231
8 Foliar, Floral and Soilborne Diseases
R.C. Ploetz
1
and S. Freeman
2
1
University of Florida, Florida, USA
2
Agricultural Research Organization, Bet Dagan, Israel
8.1 Introduction 232
8.2 Foliar and Floral Diseases 232
Algal leaf spot 232
Alternaria leafspot 233
Anthracnose 235
Apical necrosis 240
Bacterial black spot (black canker) 241
Black-banded disease 245
Black mildew, sooty blotch and sooty mould 246
Blossom blight 248
Decline disorders 249
Galls and scaly bark 256
Grey leafspot 259
Leaf blight 260
Malformation 261
Parasitic plants 270
Phoma blight 270
Phoma leafspot 270
Pink disease 271
Powdery mildew 271
Scab 274
Seca and sudden decline 274
Stigmina leafspot 279
8.3 Soilborne Diseases 281
Black root rot 281
Nematode damage 281
Phytophthora diseases 282
Root rot and damping off 284
Sclerotium rot 285
Verticillium wilt 286
White root disease 286
8.4 Conclusions 289
R.C. Ploetz and S. Freeman 232
8.1 Introduction
Diseases affect all tissues and developmental phases of mango. Mango dis-
eases have been reviewed in the rst edition of this book (Litz, 1997) and by
Singh (1968), Cook (1975), Snowdon (1990), Ploetz et al. (1994) and Ploetz
(2003). Lim and Khoo (1985), Prakash and Srivastava (1987) and Ridgeway
(1989) have also reviewed this subject. This chapter focuses on foliar, oral
and soilborne diseases of mango. Each is discussed with respect to signi-
cance, geographical distribution, history, symptoms, causal agent(s), epide-
miology and management. These diseases are caused mainly by eukaryotes
(Domain Eukaryota) among which the true fungi, Eumycota (Ascomycota
and Basidiomycota), predominate. Other less important eukaryotes include
the fungus-like oomycetes (Oomycota), nematodes (Metazoa), and parasitic
plants and green algae (Plantae). With one exception, relatively minor pathogens
of mango are prokaryotes in the Domain Eubacteria; all are Gram-negative
-proteobacteria. None of these diseases is caused by other plant-pathogenic
eukaryotes (protozoa), prokaryotes (- and -proteobacteria, Mollicutes and
Firmicutes), or the nucleic acid-based pathogens, the viruses and viroids.
8.2 Foliar and Floral Diseases
Diseases above ground are the most important and conspicuous problems on
mango. Since many of the pathogens that incite foliar diseases also affect
panicles, diseases on each are combined in this section, as are a few diseases
that affect the branches and trunks of mature trees.
Algal leaf spot
Algal leaf spot, also known as red rust, is caused by a parasitic green alga,
Cephaleuros virescens Kunze (synonyms: Cephaleuros parasiticus Karst, Ce-
phaleuros mycoidea Karst) (family Trentepohliaceae, division Chlorophyta)
(Lim and Khoo, 1985). It is a common problem on mango and many other
tropical and subtropical plants (Joubert and Rijkenberg, 1971). Although
algal leaf spot can cause serious problems on tea Camellia sinensis (L.) O.
Kuntz, black pepper, Piper nigrum L., and other important crops, it is usually
debilitating on mango only in poorly managed orchards (Lim and Khoo,
1985). In the latter cases, abiotic or biotic stresses, such as mites, insects and
other foliar diseases, can increase the severity of this disease.
Conspicuous symptoms of algal leafspot are orange to rust-coloured,
velutinous patches on both leaf surfaces (Plate 43) (Lim and Khoo, 1985).
They are initially 58 mm in diameter, but can merge to involve large, irregu-
lar sections of the leaf. They later assume a dull, greyish green colour, and
eventually become bleached patches. The alga can also affect twigs and
branches, causing the bark to crack as the pathogens laments extend into
the host cortical tissues. The orange tufts produced by C. virescens are the
algal thallus located beneath the host cuticle. They produce erect cells, some
Foliar, Floral and Soilborne Diseases 233
of which enlarge to produce stalked, terminal or ovoid, 30 24 m, sporangia
(Fig. 8.1). Sporangia produce biagellate zoospores that are dispersed by
rainsplash and wind and are the primary infective propagules. Flask-shaped
gametangia in the thallus are responsible for sexual reproduction. In free
water, they release 832 biagellate gametes, pairs of which fuse and give
rise to dwarf sporophytes. The sporophytes produce microsporangia that
bear quadriagellate zoospores; their role is not known.
Cephaleuros virescens requires a humid environment to establish and
spread (Lim and Khoo, 1985). Pruning the canopy, mowing beneath trees and
wider row spacings to increase air circulation and sunlight penetration all
help combat the disease. Proper fertilization and irrigation, and control of
insect pests, mites and other foliar diseases increase the trees ability to cope
with algal leafspot. Algacides or fungicides (i.e. fentin acetate or those con-
taining copper (Cu)) can be used to control the disease in severe cases.
Alternaria leafspot
Alternaria alternata (Fr.) Kreissler (synonyms: Alternaria fasciculata (Cooke
and Ellis) L. Jones and Grout, Alternaria tenuis Nees, and Macosporium
Fig. 8.1. Branch of the thallus of Cephaleuros virescens that has terminated in several
oval sporangia (Photograph courtesy of T.-K. Lim).
R.C. Ploetz and S. Freeman 234
Fig. 8.2. Conidia and conidiophores of Alternaria alternata (Source: Ellis, 1971).
fasciculatum Cooke and Ellis; no teleomorph is known) causes black spot on
mango, alternaria leaf spot and lesions on inorescences (Prusky et al., 1983;
Cronje et al., 1990). Although the fungus is cosmopolitan and has a large
number of host plants (Neergaard, 1945; Domsch et al., 1980), the diseases it
causes on mango are most prevalent in arid environments. In Israel, it is a
more important disease on fruit than leaves.
Symptoms on leaves are round, dark spots, 13 mm in diameter; they are
most prevalent on the underside of leaves (Prusky, 1994). Similar spots that
occur on the rachis of inorescences can reduce fruit set (Cronje et al., 1990).
Conidiophores of the causal fungus originate alone or in small groups, and
are smooth and pale to mid-olivaceous or golden brown. They are sometimes
geniculate, and vary between simple and branched, and straight to exuous.
Conidia are obclavate, 2036 99.5 m, three- to ve-septate and borne in
long chains (Fig. 8.2).
Foliar, Floral and Soilborne Diseases 235
Certain antifungal compounds are associated with the latency of A. alter-
nata infections on fruit (Droby et al., 1986, 1987). Whether the same com-
pounds are formed in leaves and inorescences has not been reported. Infected
leaves, twigs, inorescences and leaf litter are signicant sources of inocu-
lum for fruit infection. Conidia of A. alternata are dispersed by air currents
(Prusky, 1994). Several different fungicides and combined treatments are
effective against the diseases that are caused by this pathogen. Postharvest
development of alternaria fruit rot during storage is usually prevented by a
combination of hot water brushing in combination with prochloraz at
225 g/ml (Prusky et al., 2002).
The nding that A. alternata alkalinizes the host pH (Eshel et al., 2002)
prompted investigations into modulating environmental pH with acid treat-
ments to reduce fungal colonization. Spore germination and germ-tube elon-
gation of A. alternata in vitro were inhibited by, respectively, 95 and 65% by
exposure to 1.25 mM hydrochloric acid (HCl). Germination was completely
inhibited by 2.5 mM HCl (Prusky et al., 2006). Acidied solutions containing
45 g/ml prochloraz inhibited alternaria rot development better than treat-
ment with HCl alone (Eshel et al., 2002). These results have not been extended
to control foliar and oral diseases that are caused by this pathogen.
Anthracnose
Anthracnose is the most important disease of mango (Cook, 1975; Lim and
Khoo, 1985; Ploetz, 2003), particularly on fruit in humid, high rainfall envi-
ronments. It is usually replaced by, and is less important than, other diseases
in dry production areas. Anthracnose can also be a serious problem on foli-
age and owers. Crowded and moist conditions in nurseries can result in
signicant damage to young leaves and, in extreme cases, new orchards have
been devastated (Bose et al., 1973). Blossom blight, which has been attributed
to one of the anthracnose agents but is also caused by other fungi, is covered
separately below.
Symptoms
On panicles, necrotic owers abscise leaving persistent peduncles. Small, cir-
cular dark spots also develop on pedicles and peduncles. Lesions may enlarge
and coalesce to form large patches of necrotic, dark brown tissue. With suf-
cient rainfall, salmon- to orange-coloured fructications of the pathogen
develop on the affected tissues. On leaves, lesions are dark brown and sur-
rounded by chlorotic haloes, have irregular, rounded margins, and are not
delimited by veins (Fig. 8.3). Lesions are 0.51.0 cm in diameter on mature
leaves, but can expand on young leaves. Eventually, large, necrotic patches
can develop that deteriorate and fall from the leaf giving it a tattered appear-
ance. Although different mango cultivars are known to vary considerably in
their resistance to anthracnose on fruit, reports on the foliar and oral resis-
tance of different cultivars and whether resistances of the various organs are
related have not been published.
R.C. Ploetz and S. Freeman 236
Aetiology
In most production regions, anthracnose is caused by the ascomycete fungus,
Colletotrichum gloeosporioides Penz. (teleomorph: Glomerella cingulata (Stonem.)
Spauld. and Schrenk.) (Cook, 1975; Snowdon, 1990; Ploetz, 2003). In Austra-
lia, Florida USA, India, Japan and Taiwan, Colletotrichum acutatumSimmonds
(teleomorph: Glomerella acutata) plays a minor role (Fitzell, 1979; Prakash,
1990; Weng and Chuang, 1995; Taba et al., 2004; Tarnowski, unpublished). In
Colombia, Colletotrichum boninense J. Moriwaki, Toy. Sato and T. Tsukiboshi
has been reported (Afanador-Kafuri et al., 2003).
Colletotrichum spp. cause diseases on several subtropical and tropical
hosts (Jeffries et al., 1990; Freeman et al., 1998). Cultures of the fungus on
potato dextrose agar (PDA) are greyish white to dark grey and usually pro-
duce an aerial mycelium ranging from a thick mat to sparse tufts (Holliday,
1980). Conidia are hyaline, unicellular and either cylindrical with obtuse ends
or ellipsoidal with a rounded apex and a narrow, truncate base. They are
720 2.55 m, and are formed on hyaline to faintly brown conidiophores in
Fig. 8.3. Foliar symptoms of anthracnose (Photograph courtesy of S. Freeman).
Foliar, Floral and Soilborne Diseases 237
acervuli that are irregular in shape and about 500 m in diameter. Setae are
48 200 m, one- to four-septate, brown and slightly swollen at the base and
tapered at the apex. Hyphopodia have been used to distinguish isolates of C.
gloeosporioides and C. acutatum (Du et al., 2005), but provided ambiguous
results in Florida (Palmateer et al., 2007).
Characterization and taxonomic identication of Colletotrichum spp. has
relied on morphology and host range (Freeman et al., 1998; Du et al., 2005). In
general, C. gloeosporioides (Fig. 8.4) produces longer and narrower conidia
than C. acutatum (Fig. 8.5), as well as circular vs lobed hyphopodia. However,
variability in culture and host range has made these criteria unreliable for
species identication (Adaskaveg and Hartin, 1997; Freeman et al., 1998). Col-
letotrichum gloeosporioides and C. acutatum are species complexes with a large
number of hosts that are so broadly dened that the names are of limited use
to the taxonomist or plant health practitioner (Du et al., 2005). Several molec-
ular tools have been implemented to differentiate within and among Col-
letotrichum spp., including: species-specic polymerase chain reaction (PCR)
primers, random amplied polymorphic DNA (RAPD) and arbitrarily
primed PCR (Freeman and Rodriguez, 1995; Afanador-Kafuri et al., 2003);
A+T-rich DNA analyses (Freeman et al., 1993); sequence analyses of the inter-
nal transcribed spacer (ITS) regions of ribosomal DNA (rDNA) (Sreenivasap-
rasad et al., 1996; Freeman et al., 2001; Afanador-Kafuri et al., 2003) and
MAT1-2 mating type sequences (Du et al., 2005).
(a)
(b)
(c)
(d)
(g)
(e)
(f )
10
50
Fig. 8.4. (a) Acervulus and emergent setae, (b) conidiophores, (c) conidia, (d) perithecium, (e)
asci, (f) ascospores and (g) appressoria at hyphal termini of Glomerella cingulata (anamorph:
Colletotrichum gloeosporioides) (Source: from Commonwealth Mycological Institute (CMI)
description no. 315).
R.C. Ploetz and S. Freeman 238
Relatively few studies have examined isolates of C. gloeosporioides from
mango. Gantotti and Davis (1993) reported that isolates from different organs
produced different pectin-degrading enzymes, whereas Alahakoon et al.
(1994b) indicated that > 80% of isolates from mango leaves, inorescences
and fruit in Sri Lanka were identical, based on restriction fragment length
polymorphisms (RFLPs). Rivera-Vargas et al. (2006) reported that isolates of
C. gloeosporioides from leaves, fruit, owers and panicles caused similar dis-
ease on detached leaves. Work is needed to clarify whether, and the extent to
which, isolates from different mango organs differ, and whether there is a
relationship among disease reactions on various organs.
Analyses of RFLPs and RAPDs indicated that isolates from mango are
genetically uniform and differ from those recovered from avocado, caram-
bola and papaya (Hodson et al., 1993; Alahakoon et al., 1994a; Hayden et al.,
1994). Isolates genetically similar to those from the latter crops occur infre-
quently on mango, and mango isolates were not found on the other plants
and were usually virulent only on mango. Alahakoon et al. (1994b) suggested
(c)
(d)
10
(a)
25
(b)
10
Fig. 8.5. (a) Acervulus, (b) conidiogenous cells and setae, (c) conidia and (d) appres-
soria of Colletotrichum acutatum, anamorph of Glomerella acutata (Source: from CMI
description no. 630).
Foliar, Floral and Soilborne Diseases 239
that the mango isolates comprise a C. gloeosporioides population that was dis-
seminated worldwide from a single source, perhaps as an endophyte.
A recent study identied Colletotrichum spp. that infect mango, passion-
fruit (Passiora spp.) and tamarillo (Solanum betacea cav. Sendt.) in Colombia
and assessed whether cross-infection between the host species occurred
(Afanador-Kafuri et al., 2003). With species-specic PCR primers, most of the
mango isolates were identied as C. gloeosporioides. However, DNA of the
passionfruit isolates and single isolates from tamarillo and mango were not
amplied by either C. acutatum- or C. gloeosporioides-specic primers; they
were identied later as C. boninense (Freeman, unpublished data). Further
molecular analyses determined that the isolates of C. gloeosporioides from
mango were heterogeneous, but that the population of C. boninense from pas-
sionfruit, tamarillo and mango was uniform; it may not be host specic.
The origins and diversity of C. gloeosporioides on mango require more
study. Furthermore, whether distinct populations of this diverse species
occur on different mango cultivars and organs, and whether disease reac-
tions on one mango organ could be used to predict those on another should
be determined. These results would be relevant to host resistance and
improvement of the crop, as well as the chemical and cultural control of this
important disease.
Epidemiology and management
Fitzell and Peak (1984) determined that conidia were the most important
inoculum in Australia. They are produced on branch terminals, mummied
inorescences, ower bracts and leaves. New leaf ushes are the most signi-
cant source of inoculum, and this was corroborated by Dodd et al. (1991) in the
Philippines. Optimum production of conidia occurred between 25 and 30C
when free moisture was available, but also formed at 9597% relative humid-
ity (RH). The pathogens teleomorph played no apparent role in the spread of
the disease (Fitzell and Peak, 1984). The threat posed by C. gloeosporioides to
fruit production is greatest in areas with heavy rainfall. In general, this begins
with the onset of owering (Lim and Khoo, 1985; Jeffries et al., 1990). During
commercial production these diseases are managed with diverse chemicals.
Registration of the various pesticides varies in different production areas
(Jeffries et al., 1990).
Many of the most effective fungicides are systemic. They have selective
modes of action, but are prone to lost or reduced efcacy due to the develop-
ment of resistance in the targeted pathogens. Resistance in C. gloeosporioides
to benomyl is now common (Maymon et al., 2006), and there is inherent toler-
ance in C. acutatum to this fungicide (Freeman et al., 1998; Peres et al., 2002).
Newer strobilurin fungicides that are also susceptible to resistance problems
must be used properly (no more than three applications per season and alter-
nated with fungicides with different modes of action) to ensure that their
efcacy is not lost.
Although fungicide applications usually focus on reducing damage to
fruit, disease control on foliage is indicated in some, and on inorescences
in most situations. Trees in nurseries usually require protection if they are
R.C. Ploetz and S. Freeman 240
crowded or receive overhead irrigation. Since infected foliage and branch
terminals represent important reservoirs of inoculum for blossom and fruit
infection, fruit set and anthracnose control on fruit are enhanced if disease
control is exercised prior to owering (Jeffries et al., 1990). Off-season control
measures are benecial in production environments that receive signicant
rainfall.
Apical necrosis
Apical necrosis was rst reported in Spain in 1991, and now occurs in Israel,
Italy, Portugal and possibly Egypt (Cazorla et al., 1998, 2006; F.M. Cazorla,
Mlaga, 2007, personal communication). The disease can be quite damaging,
and limits production when panicles are affected. Apical buds, leaves and
panicles are susceptible (Fig. 8.6ac), but fruit are not (Cazorla et al., 1998).
Vegetative and oral apices are affected by a dark-brown to blackish necrosis
(Fig. 8.6a, c). Necrosis on leaves begins as blackened, water-soaked lesions,
13 mm in diameter, that can coalesce and expand to cover large areas (Fig.
8.6c). Necrosis also extends from affected buds to petioles, through the leaf
midrib, and can cover large areas (Fig. 8.6a). A milky bacterial exudate often
develops on affected apical buds, but infrequently on petioles (Fig. 8.6b).
Large portions of the canopy and high numbers of owers can be killed.
Fig. 8.6. Symptoms caused by the apical necrosis pathogen, Pseudomonas syringae
pv. syringae. (a) Extensive necrosis on young stem, apical bud, petioles and leaves;
(b) bacterial exudate on necrotic stem; and (c) death of a developing oral panicle and
associated leaf necrosis (Photographs courtesy of F.M. Cazorla).
(a) (b) (c)
Foliar, Floral and Soilborne Diseases 241
The causal bacterium, Pseudomonas syringae pv. syringae van Hall, affects
many perennial fruit crops (Hirano and Upper, 1983; Kennelly et al., 2007). It
is an epiphyte and is generally not an aggressive pathogen. Disease usually
develops after high populations of the bacterium develop in host tissues as a
result of host predisposition. Strains from mango produce an antimetabolite
toxin, mangotoxin, which plays a role in pathogen virulence and symptom
development (Arrebola et al., 2007).
Cold, wet weather and host genotype are primary factors in the develop-
ment of apical necrosis (Cazorla et al., 1998, 2006). Tommy Atkins, Lippens
and Manzanillo are very susceptible whereas Keitt and Sensation are less
so. Apical necrosis is managed in commercial orchards with Cu-containing
pesticides, although there has been a recent increase in control failures and
Cu resistance in Spain and Portugal (Cazorla et al., 2002, 2006). These out-
breaks have been associated with several different Cu-resistance plasmids in
the causal bacterium. Carzorla et al. (2006) determined that the plant resis-
tance activator acibenzolar-S-methyl and the phosphonate derivative fosetyl-Al
provided control comparable to Bordeaux mixture, and that the latter treat-
ment might protect plants due to the protective lm it provides against
wound entry for the pathogen.
Bacterial black spot (black canker)
Bacterial black spot is a destructive leaf, stem and fruit disease in many pro-
duction areas (Gagnevin and Pruvost, 2001). In India, the disease is called
bacterial canker or black canker due to the cankers it causes on the stems of
some cultivars (Prakash et al., 1994). It can be the most important disease
where fungal-induced diseases are well controlled (Gagnevin and Pruvost,
2001). Bacterial black spot has been identied in Australia, Myanmar, the
Comoros, India, Japan, Kenya, Malaysia, Mauritius, New Caledonia, Paki-
stan, the Philippines, Runion, Rodrigues, South Africa, Taiwan, Thailand
and the United Arab Emirates (UAE) (Fukuda et al., 1990; Pruvost et al., 1992;
Prakash et al., 1994; Gagnevin and Pruvost, 1995, 2001; Kishun, 1995; Ah-You
et al., 2007b). Given the ease with which the pathogen is disseminated in
propagation materials and its wide, conrmed distribution, bacterial black
spot may be more widely spread than is currently recognized.
Symptoms
Mango leaves, stems and fruit are affected (Manicom and Pruvost, 1994;
Gagnevin and Pruvost, 2001). On leaves, water-soaked spots are initially 13
mm in diameter. As they enlarge, they become raised, black and angular, are
limited by veins and surrounded by chlorotic haloes (Fig. 8.7). These lesions
are larger and more conspicuously raised than those caused by other xan-
thomonads that have been recovered from other species in the Anacardiaceae
(Ah-You et al., 2007a). Lesions can merge to form large necrotic patches, and
bacteria may ooze from lesions during wet conditions. Old lesions dry out,
turn white or grey, and crack. Defoliation occurs in severe cases. Anthracnose
R.C. Ploetz and S. Freeman 242
lesions are not raised or as black and angular as those caused by bacterial
black spot. On branches, bacterial black spot lesions are dark and cracked
along the long axis (Fig. 8.8). They develop only on highly susceptible culti-
vars and are often associated with wounds. Conspicuous, star-shaped lesions
are produced on fruit.
Aetiology
Diverse xanthomonads have been recovered from mango and other hosts in
the Anacardiaceae (Gagnevin and Pruvost, 2001; Ah-You et al., 2007a). Only
some of these cause symptoms of bacterial black spot. Early reports that this
disease is caused by Pseudomonas mangiferae-indicae (Patel et al., 1948) and
(a)
(b)
Fig. 8.7. (a) Symptoms of bacterial black spot on the undersurface of a mango leaf,
caused by Xanthomonas axonopodis pv. mangiferaeindicae (Photograph courtesy
of O. Pruvost). (b) Symptoms of bacterial spot on the undersurface of a mango leaf,
caused by a yellow-pigmented xanthomonad (Photograph courtesy of R.C. Ploetz).
Bacterial black spot lesions are larger and more raised than those of bacterial spot.
Foliar, Floral and Soilborne Diseases 243
Erwinia mangiferae (Steyn et al., 1974) are erroneous. The pathogens place-
ment in Pseudomonas was probably due to its production of non-pigmented
colonies in culture (P. syringae pv. syringae causes a different disease of mango,
apical necrosis see above). Cook (1975) indicated that E. mangiferae is a

saprophyte that reached high populations in old lesions.
Pathological, cultural, biochemical, physiological, serological and genetic
data indicate that strains of the pathogen from different production areas are
diverse (Sanders et al., 1994; Gagnevin and Pruvost, 1995, 2001; Kishun, 1995;
Pruvost et al., 2005). Genetic diversity is greatest among strains from South-
east Asia, suggesting that this region of host diversity is also a centre of
pathogen diversication (Gagnevin and Pruvost, 1995).
The pathogen has a single agellum, is Gram-negative, rod shaped and
0.40.5 1.01.5 m (Manicom and Wallis, 1984). It is oxidase negative and
does not reduce nitrates to nitrites. It cannot use asparagine as a sole carbon
and nitrogen source, but is able to hydrolyse starch, esculin, gelatin and
casein. On articial media, colonies are cream coloured. (The latter trait is
atypical for xanthomonads, which are usually yellow in culture.) Yellow-
pigmented xanthomonads have been recovered from mango in Brazil,
Runion, South Africa and Florida USA. These strains cause non-raised leaf
lesions (Fig. 8.7b), do not cause fruit or stem lesions, and should not be
Fig. 8.8. A twig canker caused by X. axonopodis pv. mangiferaeindicae that is
associated with a wound (Photograph courtesy of O. Pruvost).
R.C. Ploetz and S. Freeman 244
classied as pv. mangiferaeindicae (Ah-You et al., 2007a). Three genetically and
pathologically distinct groups were identied from different geographic
regions and hosts in the Anacardiaceae by Ah-You et al. (2007a). Group I
strains were from the Old World, multiplied in mango and cashew (Anacar-
dium occidentale L.), fell in amplied fragment length polymorphism (AFLP)
group 9.5 of Xanthomonas axonopodis, and contained strains that produced
typical bacterial black spot symptoms on mango. These strains of Xanthomo-
nas campestris pv. mangiferaeindicae sensu lato were redescribed as X. axonopo-
dis pv. mangiferaeindicae sensu novo (s.n.) (Ah-You et al., 2007a). In contrast,
Group II strains were from Brazil, multiplied in cashew, but not mango, and
fell in AFLP group 9.6 of X. axonopodis. They are associated with symptoms
on mango that differ from those of bacterial black spot, including brown, at
leaf lesions, and black, depressed lesions on fruit of only a few cultivars that
is sometimes associated with pulp rot. These strains were responsible for
previous reports of bacterial black spot in Brazil (Gagnevin and Pruvost,
2001; Ah-You et al., 2007a). Group III strains are responsible for a unique syn-
drome on Spondias dulcis and Spondias mombin in the French West Indies; they
fell in AFLP group 9.4. Groups II and III were described, respectively, as X.
axonopodis pv. anacardii and X. axonopodis pv. spondiae (Ah-You et al., 2007a).
Epidemiology and management
The pathogen is disseminated by wind-driven rain (Gagnevin and Pruvost,
2001). Long-distance dissemination occurs via infected propagation material
and, less frequently, in infected seed. True seed are not infected, but surface
contamination is known. Insects may play a role in dissemination, but these
interactions are little studied. The pathogen is an epiphytic colonist of leaves
(Manicom, 1986; Pruvost et al., 1990), buds (Pruvost et al., 1993) and fruit
(Pruvost and Luisetti, 1991b). Infection occurs through wounds and, less
often, stomata on old leaves (young leaves are thought to be resistant due to
their non-functional stomata) (Gagnevin and Pruvost, 2001). High RH
(> 90%) and moderate temperatures (2530C) favour disease development
(Kishun and Sohi, 1983; Pruvost and Luisetti, 1991b). There is a direct rela-
tionship between the level of disease that develops on leaves and fruit (Man-
icom, 1986; Pruvost et al., 1990). Pruvost and Luisetti (1991b) considered that
leaf susceptibility was an important criterion when cultivars were selected
for lower fruit susceptibility.
Resistance to bacterial black spot varies among mango cultivars, and
resistant cultivars should be used where disease pressure is high (Manicom
and Pruvost, 1994). When new orchards are established, pathogen-free plant-
ing material should be utilized. Since the pathogen moves short distances in
wind-blown aerosols (usually within orchards), the long-distance spread of
the pathogen depends almost entirely upon the movement of infected plants
(Manicom and Pruvost, 1994). Windbreaks can be used to reduce wounding
and infected twigs (Fig. 8.8) should be pruned to reduce inoculum in the
canopy.
Bacterial black spot can be quite difcult to control, particularly on sus-
ceptible cultivars. Available chemicals may be only marginally effective
Foliar, Floral and Soilborne Diseases 245
under high disease pressure (Pruvost et al., 1989). During rainy weather,
applications of Cu-based bactericides are recommended. The application
schedules for these compounds focus on protecting fruit, and vary according
to the length of time fruit are exposed to wet conditions (Manicom and Pru-
vost, 1994). Although agricultural antibiotics (e.g. various formulations of
streptomycin sulfate or nitrate) have been reported to be effective (Misra and
Prakash, 1992; Viljoen and Kotz, 1972), resistance that develops to these
products after continuous use limits their long-term effectiveness against
this disease. Biological control measures have not been widely researched.
Pruvost and Luisetti (1991a) reported little success. In India, Kishun (1994)
indicated that a strain of Bacillus coagulans from the phylloplane of mango
was effective against strains of the pathogen, although control of bacterial
black spot in the eld was not reported.
Black-banded disease
This is a relatively unimportant disease that affects mango leaves and
branches (Reddy et al., 1961). The causal fungus, Rhinocladium corticola Mas-
see (described as corticolum) (teleomorph: Peziotrichum corticolum (Massee)
Subrumanian), was described on the bark of trees in Poona, India (Hughes,
1980). It produces repent, intricately branched, septate, olivaceous hyphae
57 m in diameter. Erect hyphae bear globose, olivaceous, densely and
minutely tuberculate, 1518 m conidia. Hughes (1980) questioned whether
this was a species of Rhinocladium since it was quite different from other spe-
cies in the genus. It forms a black, velvety mass of hyphae on affected sur-
faces in conspicuous blotches or bands. The fungus is restricted to the outer
portions of bark. Bordeaux mixture controls the disease, but is not required
in most situations.
Black mildew, sooty blotch and sooty mould
Several ascomycetes produce dark-coloured, usually supercial growths
on the surfaces of stems, leaves and fruit (Lim and Khoo, 1985). These
range from thin, diffuse webs of dark hyphae to opaque, felty layers; in
extreme cases, a thick crust of hyphae forms. The variations in appearance
result, presumably, from the different species of fungi that are involved.
These are usually not important problems in well-maintained orchards.
However, layers of hyphae that the black mildew and sooty mould fungi
form may be thick enough to block sunlight and inhibit photosynthesis.
These blemishes also detract from the appearance and marketability of
fruit.
Sooty moulds develop in the presence of aphids, mealybugs, scales and
other sucking insects that produce honeydew (excreta) when feeding. Hon-
eydew is a required food source for these fungi, and the problems they cause
dissipate if the associated insects are controlled. In contrast, black mildews
R.C. Ploetz and S. Freeman 246
and sooty blotch are not dependent on honeydew and grow directly on host
surfaces.
Aetiology
Black mildew and sooty mould are similar in appearance, but their respec-
tive causal agents are distinct (Lim and Khoo, 1985). The black or dark mil-
dews are a group of mostly tropical obligate plant pathogens that produce
two types of hyphopodia (Alexopoulos et al., 1996). Capitate hyphopodia are
lobed appressoria from which infection haustoria are formed, whereas
mucronate hyphopodia function as conidiogenous cells. Black mildew of
mango is caused by Meliola mangiferae Earle (Sordariomycetes, Ascomycota)
(Fig. 8.9).
In contrast, the fungi that cause sooty moulds are diverse saprophytes
that require honeydew to colonize plant surfaces. In Malaysia, Lim and Khoo
(1985) listed coelomycetes (Polychaeton), hyphomycetes (Tripospermum) and
loculoascomycetes (Antennulariella, Chaetothyrium, Limacinula and Scorias),
whereas the reported agents in India were hyphomycetes (Leptoxyphium,
(a)
(c)
(b)
10 mm
10 m
(d)
250 m
1 mm
Fig. 8.9. (ac) Signs of black mildew, and (d) microscopic features of the causal
agent, Meliola mangiferae (Source: from CMI description no. 1355).
Foliar, Floral and Soilborne Diseases 247
Microxyphium and Tripospermum) and loculoascomycetes (Capnodium) (Butler
and Bisby, 1931; Prakash, 1988). In Pakistan, 18 species in eight genera were
associated with sooty mould, including the foliar pathogens Aspergillus,
Alternaria, Botryodiplodia, Capnodium, Cladosporium, Curvularia, Fusarium and
Helminthosporium spp. (Hamid and Jalaluddin, 2006). Since some of the sooty
mould fungi may not sporulate on plants and because they are often found
in combination with one another, it is usually difcult to identify the specic
species that are involved.
Although sooty blotch has been used as a synonym for sooty mould
(Singh, 1968), sooty blotch refers specically to disease complexes on tem-
perate and tropical plants that are not associated with honeydew and are
caused by a diverse group of Dothidiomycetes (Ascomycota) (Johnson et al.,
1997; Ploetz et al., 2000; Batzer et al., 2005). The specic agents that are associ-
ated with sooty blotch on mango are not known, but resemble those from
apple, carambola and pear (Plate 44).
Epidemiology and management
Sooty moulds develop on honeydew that is produced on plant surfaces by
aphids, mealybugs, scales and other sucking insects. They are managed by
controlling the associated insects with oils and insecticides (Lim and Khoo,
1985). In Pakistan, spraying of fungicides (sulfur (S) and mancozeb) and
insecticides (malathion, diazinon and coal tar at 1 kg/tree) separately reduced
the incidence of sooty mould on foliage, whereas a mixture of fungicides and
insecticides further decreased sooty mould incidence (Hamid and Jalalud-
din, 2006). In India, sooty mould, caused by species of Microxyphium, Lep-
toxyphium and Tripospermum, was best controlled with a spray of S and
parathion-methyl (Prakash, 1991).
Sooty blotch management has not been investigated on mango;
however, signs of these fungi have been removed from apple with various
postharvest washes (Batzer et al., 2002), and managed in apple orchards
with diverse contact and systemic fungicides (Williamson and Sutton,
2000).
Blossom blight
Blossom blight can reduce fruit set and production considerably since ow-
ers and large areas of the panicle can be killed. When this disease was con-
trolled with fungicides in the Philippines, a 5580% increase in fruit set
occurred (Pordesimo, 1982).
Symptoms
Blossom blight starts as a wilt of the affected part of the inorescence that is
often curved, the shepherds crook symptom (Fig. 8.10). The peduncle
blackens and dies back from the tip. Internally, discoloration advances
beyond the surface lesion. Large black lesions can develop lower on the
peduncle, and once it is girdled the apex dies.
R.C. Ploetz and S. Freeman 248
Aetiology
The cause of blossom blight is confused. Colletotrichum gloeosporioides has
been reported most frequently as the responsible fungus (Fitzell et al., 1984;
Jeffries et al., 1990), and A. alternata has also been reported to attack panicles
and reduce fruit set (Cronje et al., 1990). Powdery mildew (see section below)
also damages panicles, but its symptoms are distinct from those of blossom
blight. In South Africa, symptoms caused by A. alternata and C. gloeosporioides
are small, mainly supercial black spots, 12 25 mm, on the peduncle
(Darvas, 1993; Lonsdale and Kotz, 1993). Rather than blossom blight, Lonsdale
and Kotz (1993) indicated that these pathogens caused blossom spot. In con-
trast, Lonsdale and Kotz (1993) reported that Dothiorella mangiferae caused
extensive, systemic damage, and Darvas (1993) indicated that Dothiorella domini-
cana is the only fungus that caused typical symptoms of blossom blight.
Crous et al. (2006) placed these fungi in a new genus, Neofusicoccum (see
Decline disorders section below). Work is needed to determine the distribu-
tion of Neofusicoccum-incited panicle disease, and the identity of the most
important blossom blight pathogens worldwide.
Epidemiology and management
Little is known about the epidemiology of Neofusicoccum-incited panicle dis-
ease. Studies on the stem-end rot diseases have shown that the causal fungi
are endophytes. The roles of internal and external sources of inoculum in
the development of panicle disease are unknown. For optimal fruit set and
Fig. 8.10. Symptoms of blossom blight on panicles of mango. Note the blighted,
curved terminals and almost complete lack of fruit set (Photograph courtesy of
D. Benscher).
Foliar, Floral and Soilborne Diseases 249
development, blossom blight must be controlled. Once owering begins,
early and frequent fungicide applications are necessary in many areas,
depending on rainfall. Previously published infection models can be used to
time applications appropriately (Fitzell et al., 1984; Dodd et al., 1991). Fitzell
et al. (1984) investigated environmental conditions that were conducive to
infection by C. gloeosporioides, and indicated that temperature and free mois-
ture were important determinants of infection. They developed a model for
scheduling fungicide application, which reduced the number of applications
that were needed to control blossom blight. Presumably, systemic fungicides
would be needed to control disease caused by endophytic agents.
Decline disorders
Several diseases of mango have been variously termed blight, canker, decline,
gummosis, twig blight, tip dieback and stem bleeding. They have similar
symptoms and aetiologies.
Symptoms
These widespread problems are not well understood. Symptoms include: (i)
marginal scorching of leaf lamina; (ii) foliar symptoms of nutritional de-
ciencies, particularly of iron (Fe) and manganese (Mn); (iii) vascular discolor-
ation (Fig. 8.11a); (iv) dieback of small branches basipetally from the terminal
that may or may not progress to defoliation (Fig. 8.11b and c); (v) gummosis,
an oozing of a clear or cloudy exudate either from terminal buds or from
branches, scaffold limbs or trunks (Plate 45); and (vi) root degeneration (Lim
and Khoo, 1985; Ploetz et al., 1996a; Ploetz and Prakash, 1997).
Aetiology
Diverse biotic and abiotic factors may be primary causes of decline symp-
toms or predisposing agents (McSorley et al., 1980; Kadman and Gazit, 1984;
Schaffer et al., 1988; Ploetz and Prakash, 1997). Fungi are the most common
agents. They are endophytes that also cause stem-end rots on mango fruit,
and are usually secondary pathogens that cause disease on weakened, pre-
disposed hosts (Johnson et al., 1992; Ploetz and Prakash, 1997; Slippers et al.,
2005; Slippers and Wingeld, 2007). Several species cause all or some of the
above symptoms when used to individually inoculate plants (Ploetz et al.,
1996a). Their frequent association with one another in affected tissues may
indicate that these symptoms usually develop, or develop more severely,
after multiple infections.
Several of these pathogens are in the Botryosphaeriaceae (Dothidiomy-
cetes, Ascomycota). The taxonomy and nomenclature of these fungi has been
confused, and phylogenetic understanding of major groups within Botry-
osphaeria remains poor (Crous et al., 2006). With 28S rDNA sequence data,
Crous et al. (2006) examined natural relationships among available members
of the family. Ten lineages were distinguished, most of which contained ana-
morphs with distinct morphological features. New relationships were revealed
R.C. Ploetz and S. Freeman 250
in some of the lineages that necessitated the renaming of several taxa (Crous
et al., 2006). These new names and holomorphs are used below when discuss-
ing the taxa that occur on mango.
Lasiodiplodia theobromae (Pat.) Griffon and Maubl.) (synonyms: Botryodi-
plodia theobromae Pat., Diplodia natalensis Pole-Evans, and Diplodia theobromae
(Pat.) W. Nowell) is the most common and widespread cause of decline dis-
eases of mango (Ploetz and Prakash, 1997), and affects many other host plants
in the tropics (Punithalingam, 1976). Crous and Palm (1999) declared B. theo-
bromae, a nomen dubium. Denman et al. (2000) reduced D. natalensis and L.
theobromae to synonymy with D. theobromae. However, adopting this change
was questioned by Burgess et al. (2006), who noted that ve species of Lasio-
diplodia fell in a phylogenetic clade that had 100% bootstrap support; it was
distinct from a clade that included species of Diplodia and Dothiorella. The
teleomorph of L. theobromae, formerly Botryosphaeria rhodina (Cooke) Arx
(synonym: Physalospora rhodina Cooke), is usually not found in nature. In the
study of Crous et al. (2006), the genus Botryosphaeria was reserved for the type
species Botryosphaeria dothidea (Moug.:Fr.) (anamorph: Fusicoccum aesculi
Corda), which was in a different clade than L. theobromae. However, Crous et
al. (2006) refrained from renaming B. rhodina until the poorly resolved clade
in which it resided could be claried with work with additional isolates and
analyses.
Lasiodiplodia theobromae attacks weakened trees that are predisposed by:
high and low temperatures; drought; high RH; hardpan soils; sun scorch;
and tar and tanglefoot (Muller, 1940; Das Gupta and Zacchariah, 1945;
Alvarez-Garca and Lpez-Graca, 1971; Acua and Waite, 1977; Ploetz et al.,
1996a). It is often an endophyte, infects wounded plants, and is found in soil,
on dead twigs, mummied fruit and on organic debris beneath trees (Johnson
et al., 1992).
(a) (b) (c)
Fig. 8.11. Among the symptoms that are associated with mango decline are:
(a) internal/vascular discoloration and branch terminal death (tip dieback) that may not
(b), or may be associated with defoliation (c) (Photographs courtesy of D. Benscher).
(a) (b) (c)
Foliar, Floral and Soilborne Diseases 251
Dieback caused by L. theobromae has been recognized as a signicant dis-
ease in India since the 1940s. It was the most serious disease of mango in the
Jaipur district (Verma and Singh, 1970), and affected 3040% of the planta-
tions in the Moradabad region of Uttar Pradesh (Prakash and Srivastava,
1987). Das Gupta and Zacchariah (1945) indicated that only L. theobromae
caused dieback; Phoma sp. and two Fusariumspp. were not pathogenic. Lasio-
diplodia theobromae caused a canker on mango in Indonesia (Muller, 1940) and
Malaysia (Lim and Khoo, 1985), dieback in Egypt and the Sonsonate area of
El Salvador (Acua and Waite, 1977), and gummosis and dieback in Puerto
Rico (Alvarez-Garca and Lpez-Graca, 1971).
Lasiodiplodia theobromae produces uffy, grey-black mycelium on oatmeal
agar (OA) and PDA (Johnson, 1994b). Conidiomata may be simple or develop
into aggregated stromatic bodies (Burgess et al., 2006). Cirri of conidia may
ooze from ostioles. Conidia are initially hyaline, aseptate, granular, ovoid
to ellipsoid and thick-walled (Fig. 8.12). Mature conidia are two celled,
1733 1015 m, and brown walled with numerous longitudinal striations.
Paraphyses are usually present. The teleomorph was produced when indi-
vidual isolates were cultured on caimito fruits (Chrysophyllum cainito L.) or
papaya (Carica papaya L.) stems, suggesting that it was homothallic (Alvarez-
Garca and Lpez-Graca, 1971).
One of the most important mango pathogens causes stem-end rot on
fruit, dieback and blossom blight. Crous et al. (2006) refer to it as Neofusicoccum
parvum (Pennycook and Samuels) Crous, Slippers and A.J.L. Phillips (formerly
Fusicoccum parvum Pennycook and Samuels) (teleomorph: Botryosphaeria-like;
formerly Botryosphaeria parva Pennycook and Samuels). Slippers et al. (2005)
argued that D. dominicana Petro and Cif. may be synonymous with this fun-
gus. Johnson (1992), an author of the Slippers et al. (2005) paper, had placed
D. dominicana in synonymy with F. aesculi Corda (teleomorph B. dothidea
(Moug.:Fr.) Ces. and De Not.). They indicated that this fungus had been mis-
identied as Botryosphaeria ribis Gross. and Duggar (anamorph: Fusicoccum
sp.) and B. dothidea (anamorph: F. aesculi), due to overlapping host ranges
and spore dimensions. They felt that the tip dieback fungus reported by
Ramos et al. (1991) as B. ribis was probably N. parvum (= B. parva).
Neofusicoccum parvum produces cottony grey mycelium and discrete pyc-
nidia or stromatic multilocular fruiting bodies on, respectively, PDA and
OA (Johnson et al., 1991). Discrete, immersed pycnidia in a subcuticular
pseudostroma are produced on mango. Conidia are fusiform to navicular,
14.725.5 (19) 4.57 (5.2) m, hyaline and unicellular (Slippers et al., 2005).
Sometimes, brown, biseptate conidia are observed. The teleomorph develops
infrequently on OA, and has been found on mango twigs in tree litter in
Australia (Johnson et al., 1991). On twigs, pseudothecia are subglobose to
pyriform, 210 120 m, and form beneath the epidermis. Ascostromata are
hemi-lenticular and up to 10 mm wide on OA. Asci are eight spored, bituni-
cate and irregularly biseriate. Ascospores are hyaline, single celled, fusiform
and 1625 4.59.5 m.
Neofusicoccum mangiferum (Syd. and P. Syd.) Crous, Slippers and A.J.L.
Phillips (basionym: Dothiorella mangiferae Syd. and P. Syd.; synonyms:
R.C. Ploetz and S. Freeman 252
Nattrassia mangiferae (Syd. and P. Syd.) B. Sutton and Dyko; Fusicoccum
mangiferum(Syd. and P. Syd.) Johnson, Slippers and M.J. Wingf.) causes blos-
som blight and postharvest fruit rot in South Africa (Lonsdale and Kotz,
1993; Saaiman, 1996). On PDA, N. mangiferum produces grey, felted myce-
lium with gregarious, partly immersed, discrete conidiomata, pepper-spot
patterns of pycnidial initials, and dark grey mycelium that lacks the white
tufts found in similar species (i.e. N. parvum) (Johnson, 1994b; Slippers et al.,
2005). On mango, the fungus produces unilocular conidiomata in subcuticu-
lar pseudostroma. The conidia of N. mangiferum differ from those produced
by other Neofusicoccum spp. by their shorter average length (1314 m) and
smaller length/width ratio (22.5) (Slippers et al., 2005). They are usually
unicellular, ellipsoid to ovoid, 13.6 5.4 m and hyaline, although conidia
often become one or two septate, light brown with distinctly darker middle
cells. The teleomorph (not identied) resembles B. parva, and develops
occasionally on OA.
(b)
(d)
(c)
(a)
500
10
Fig. 8.12. Pycnidia (a and b), conidiogenous cells and immature conidia (c) and
mature and immature conidia (d) of Lasiodiplodia theobromae (Source: from CMI
description no. 519).
Foliar, Floral and Soilborne Diseases 253
A dieback disease of mango has been recognized in Niger (Reckhaus
and Adamou, 1987). Neoscytalidium dimidiatum (Penz.) Crous and Slippers
(basionym: Torula dimidiata Penz.; synonyms: Scytalidium dimidiatum (Penz.)
B. Sutton and Dyko (Fig. 8.13); Fusicoccum dimidiatum (Penz.) D.F. Farr;
Hendersonula toruloidea Natrass) causes sudden wilting of shoots to large
branches, ring of leaves and trunk cankers from which a clear exudate orig-
inates. Reckhaus and Adamou (1987) believed that water stress was a pri-
mary, predisposing factor in the development of this disease.
Botryosphaeria dothidea (anamorph: F. aesculi) is an uncommon mango
pathogen (Ploetz and Prakash, 1997; Slippers et al., 2005). It produces a uffy
grey mycelium with discrete pycnidia on PDA or stromatic multilocular
fruiting bodies on OA. Discrete, immersed pycnidia are produced on mango.
(e)
(d)
(c)
(a)
(b)
10
50
Fig. 8.13. (a) Vertical section of stroma, (b) part of pycnidial wall and conidiophores,
(c) conidiophores, (d) conidia, and (e) the Scytalidium-like synanamorph of Neoscyta-
lidium dimidiatum (Source: from CMI description no. 274).
R.C. Ploetz and S. Freeman 254
Conidia are 18.830.4 4.57 m, hyaline, and single celled (Fig. 8.14). The
teleomorph is occasionally produced on OA and has been found in litter
beneath avocado and mango trees (Johnson, 1994b; Michailides et al., 1999).
On twigs, pseudothecia are subglobose to pyriform, 210 120 m, and im-
mersed beneath the epidermis. On OA, ascostromata are hemi-lenticular and
up to 10 mm wide. Asci are eight spored, bitunicate and irregularly biseriate.
Ascospores are hyaline, single celled, fusiform and 1625 4.59.5 m.
Mango decline is an important disorder in Florida USA, Israel and
other areas that have calcareous soils (Schaffer, 1994; Ploetz et al., 1996a).
Symptoms include interveinal chlorosis and marginal necrosis of leaves,
(a)
(c)
(b)
10
500
Fig. 8.14. (a) Conidia, (b) conidiophores and (c) a vertical section of a conidioma of
Fusicoccum aesculi, anamorph of Botryosphaeria dothidea (Source: Sutton, 1980).
Foliar, Floral and Soilborne Diseases 255
dieback of young twigs that progresses to larger branches, reduced growth of
secondary roots, gummosis and vascular discoloration. Several different
factors have been associated with mango decline in Florida. Schaffer et al.
(1988) used the Diagnosis and Recommendation Integrated System (DRIS) to
assess the nutritional status of declining and healthy Tommy Atkins trees.
The nutrient imbalance index, an indication of the overall nutrient balance in
trees, was greatest for declining trees. DRIS identied Mn, Fe or both ele-
ments as the most decient in declining trees, and in two of the three declin-
ing orchards that they investigated, concentrations of these elements were
below the critical range. Mineral deciencies may be predisposing factors in
the development of mango decline, since pathogenic fungi are recovered
from symptomatic trees (see below).
McSorley et al. (1980) detected the nematode Hemicriconemoides mangiferae
Siddiqi at low, but consistent levels on declining mango trees. They sug-
gested that it may have been responsible for the reduced root growth in
affected trees, and could play a role in the development of the disorder.
Smith and Scudder (1951) reported that Diplodia sp. caused a dieback
of mango. Additional species of fungi were examined by Ploetz et al.
(1996a). Alternaria alternata, C. gloeosporioides, N. parvum (D. dominicana), L.
theobromae and two Phomopsis spp. were recovered from trees with diverse
decline symptoms, and caused one or more of these symptoms on Keitt and
Tommy Atkins. Colletotrichum gloeosporioides, N. parvum and L. theobromae
were most damaging, and caused signicant bud necrosis, tip dieback,
gummosis and vascular discoloration (Fig. 8.15); these symptoms were
distinguishable only when C. gloeosporioides sporulated on inoculated
branches.
In summary, several different fungi cause, or are associated with, decline
symptoms worldwide; most are endophytes that have Botryosphaeria or
Botryosphaeria-like teleomorphs (Botryosphaeriaceae). Stress and wound pre-
disposition are usually associated with symptom development.
Management
Controlling decline disorders of mango is difcult. Techniques that could
detect these pathogens in plants would be useful to identify pathogen-free
propagation materials. The internal location and the diversity of fungi that
are involved decrease the opportunities for controlling these disorders with
fungicides (Peterson et al., 1991). Because signicant movement of some of
these pathogens may occur via wind and rainsplash, strategic applications of
broad-spectrum protectant fungicides may be effective at certain times of the
year (Lonsdale and Kotz, 1993), but have not been tested. In India, dieback
was managed by pruning affected portions of the canopy and treating the
wounded areas with a 5:5:50 Bordeaux mix (Prakash and Raoof, 1989). Man-
agement of the controllable predisposing factors, such as drought stress, may
also be benecial. A better understanding of the epidemiology of these dis-
eases would assist these efforts. Pruning to force synchronous ushes of
foliar growth might enable the avoidance of windows of infection for certain
pathogens (Johnson, 1994b).
R.C. Ploetz and S. Freeman 256
Galls and scaly bark
Gall and scaly bark disorders of mango are known in several producing
regions. These diseases are usually minor problems.
Symptoms
In India, bark scaling develops as deep cracks along the entire rootstock por-
tion of the plant, and cracks may penetrate the phloem and become necrotic
(Prakash and Srivastava, 1987). These symptoms resemble those of a scaly
bark disorder, cuarteado, in Colombia (Cook, 1975). In Hawaii, similar
symptoms developed on mango seedlings (Cook et al., 1971). The bark from
the soil line to the rst branches was rough and scaly, and xylem pegs, 5 mm
long, were evident when the bark was removed around leaf scars and
secondary branches.
In Mexico, a disorder known as nanahuate, bolas or buba of mango,
causes galls, 510 cm in diameter, which resemble a cauliower, are initially
light green, but become dark brown as they die (Fig. 8.16) (Angulo and
Villapudua, 1982). The galls remain attached to trees for many years, and
severely affected branches die. Similar symptoms are found in Florida USA,
and are associated with pruning injuries. Larger galls have also been noted
in Puerto Rico, as well as the US Department of Agriculture (USDA)
Agriculture Research Service (ARS) in Miami and University of Florida
in Homestead (Fig. 8.17) (Ploetz et al., 1996b; R. Rodriguez, personal
communication). Some of the latter galls are large, have rough, scaly exteri-
ors, and are usually found on the main trunk or scaffold limbs of affected
trees. Cracks may penetrate the phloem and become necrotic, but the branch
death that is associated with galls in Mexico and India has not been
observed.
(a) (b) (c)
Fig. 8.15. Decline symptoms induced on Tommy Atkins plants articially inoculated
with isolates of: (a) C. gloeosporioides; (b) Neofusicoccum parvum; and (c) L. theobro-
mae (Photographs courtesy of D. Benscher).
(a) (b) (c)
Foliar, Floral and Soilborne Diseases 257
Aetiology
Fusarium decemcellulare C. Brick (synonym: Fusarium rigidiuscula (Brick) Snyd.
and Hans.) causes these diseases in Florida USA, Mexico and Venezuela
(Malaguti and de Reyes, 1964; Angulo and Villapudua, 1982; Ploetz et al.,
1996b). Colonies on PDA are dark carmine-red on the underside. The fungus
produces microconidia in false heads or chains on branched and non-
branched monophialides (Fig. 8.18). Large macroconidia, 9255 75.5 m,
are produced in slimy yellow sporodochia, c.1 mm in diameter. The funguss
Fig. 8.16. Galls of the buba type in Haiti (Photograph courtesy of Carolyn Cohen,
USDA, Animal and Plant Health Inspection Service (APHIS)).
Fig. 8.17. Large, 30-year-old gall on Langra in the USDA-ARS mango germplasm
repository in Miami (Photograph courtesy of R.C. Ploetz).
R.C. Ploetz and S. Freeman 258
teleomorph, Albonectria rigidiuscula (synonyms: Nectria rigidiuscula Berk. and
Broome, and Calonectria rigidiuscula) (Rossman et al., 1998), has not been
observed on mango.
Fusarium decemcellulare causes corky bark, gall, canker and dieback dis-
eases on diverse woody hosts in the subtropics and tropics (Holliday, 1980;
Farr et al., 1989; Aleri et al., 1994). It causes an important disease of cacao
(Theobroma cacao L.), cushion gall, as well as a stem gall on loquat (Eriobotrya
japonica (Thunb.) Lindl.) and scaly bark of pongam (Pongamia pinnata (L.)
Pierre). Host specialization in the fungus has not been reported. Fusarium
decemcellulare has not been reported to cause gall and scaly bark disorders of
mango in other areas. The possible role of Agrobacterium tumefaciens was
examined in Miami; however, the bacterium could not be recovered from
affected tissues (R. McGuire, Miami, 1993, personal communication). In
Hawaii, microorganisms were not recovered from affected plants (Cook,
1975).
Epidemiology
Isolates of F. decemcellulare from mango are only mildly aggressive (Ploetz
et al., 1996b), and require wounding in order to infect. A recent outbreak of
scaly bark in a commercial mango orchard in Florida USA was associated
with pruning wounds. In the cushion gall disease on cacao, F. decemcellulare
interacts with several different insect pests and pathogenic agents (Holliday,
(b)
(c)
(a)
20
Fig. 8.18. (a) Ascus and ascospores of Albonectria rigidiuscula, and (b) micro-
conidia and conidiophores, and (c) macroconidia and conidiophores of its anamorph,
Fusarium decemcellulare (Source: from CMI description no. 21).
Foliar, Floral and Soilborne Diseases 259
1980; Ploetz, 2007). These insects may facilitate infection and disseminate the
pathogen. Insect-F. decemcellulare interactions have not been investigated on
mango.
Management
No pesticides have been identied to control this problem. Measures that
should be helpful include the removal and destruction of affected branches
and trees in the orchard, disinfestation of pruning equipment to ensure that
the pathogen is not spread during pruning operations, and the use of healthy
planting material in new orchards.
Grey leafspot
Pestalotiopsis mangiferae (Henn.) Steyaert (synonym: Pestalolia mangiferae
Henn.; no teleomorph of the fungus is known) causes grey leafspot and stem-
end rot of mango fruit (Lim and Khoo, 1985; Johnson, 1994b). It is a weak
pathogen that usually requires wounding in order to infect mango. Grey
leafspot is usually unimportant and occurs mainly on unhealthy or poorly
maintained trees.
Pestalotiopsis mangiferae produces abundant conidia in acervuli that
develop in grey leafspot lesions and necrotic areas on fruit (Lim and Khoo,
1985). As lesions age, black columns of spores emanate through ruptures in
the host epidermis. Conidia are produced that have three thick-walled,
brownish, concolorous median cells and thin-walled, hyaline apical and
basal cells; the apical cells bear three characteristic appendages (Fig. 8.19).
Conidia are 20 5 m, fusiform and straight to slightly curved. Two other
species of Pestalotiopsis that occur on mango produce larger conidia, Pestalo-
tiopsis mangifolia Guba and Pestalotiopsis versicolor Speg. (synonyms: Pestaloti-
opsis cliftoniae Tracy and Earle and Pestalotiopsis coccolobae Ellis and Everh.).
On leaves, symptoms are light grey spots, usually 220 mm in diameter
(Lim and Khoo, 1985). These may coalesce to form large patches of necrotic
tissue on leaves. Lesions are surrounded by dark, raised margins, and as they
mature, raised black dots (which are columns of the pathogens conidia) are
evident in lesion centres. Although most cultivars are susceptible, specic
control measures are usually not required. Dithiocarbamate fungicides con-
trol this disease.
Leaf blight
This disease has been reported in India and Nigeria (Hingorani et al., 1960;
Cook, 1975; Okigbo, 2001; Okigbo and Osuinde, 2003), and the causal fun-
gus, Macrophoma mangiferae Hingorani and Sharma (Ascomycota), has also
been intercepted in shipments to the USA from Mexico (Systematic Mycol-
ogy and Microbiology Laboratory, USDA-ARS, Beltsville). This is not a
serious problem.
R.C. Ploetz and S. Freeman 260
Macrophoma mangiferae produces subepidermal, globose pycnidia, 77231
m in diameter. Hyaline conidiophores, 1.52 811 m, produce unicellular
conidia, 5.37 10.524.5 m. No teleomorph is known. Since the genus Mac-
rophoma has been placed in synonymy with Sphaeropsis, this species should
be redescribed.
Leaves, twigs and fruit are affected, especially when the latter are stored.
Small, yellow spots gradually enlarge to become brown with wide purple
margins. The lesions are initially circular, but develop an irregular appear-
ance and may encompass large portions of the leaf surface. Pycnidia form
most frequently on the underside of leaves. Elliptical stem lesions are infre-
quent but can girdle stems. In India, the disease is most serious during the
rainy season (Verma and Singh, 1996b). Macrophoma mangiferae survives in
pycnidia that develop on bark of twigs of young mango plants, blighted
leaves and as dormant mycelium in wood (Verma and Singh, 1996a). Okigbo
(2001) reported that the fungus survived adverse conditions best in stems
and branches.
Four applications of captan, Bordeaux mixture, captafol, carbendazim
and thiophanate-methyl were effective on young plants (Verma and Singh,
1996a). The bacterium Bacillus subtilis NCIB 3610, isolated from soil under a
mango tree, inhibited M. mangiferae on agar plates, and symptoms were
reduced in the eld by the application of the antagonist (Okigbo and Osuinde,
2003).
(a)
(b)
50 m
(c)
10 m
Fig. 8.19. (a) Vertical section of an acervulus, (b) mature conidia, and (c) conidiog-
enous cells and developing conidia of Pestalotiopsis mangiferae (Source: from CMI
description no. 676).
Foliar, Floral and Soilborne Diseases 261
Malformation
Malformation is one of the most destructive mango diseases (Ploetz, 2001).
Although trees are not killed, the vegetative phase of the disease impedes
canopy development and the oral phase dramatically reduces fruit yield.
Based on the incidence and severity of malformation in Egypt, an estimated
US$15 million of fruit were lost in 1998 (Ploetz et al., 2002). Losses in more
important producing countries (i.e. India) are undoubtedly much greater.
Malformation was rst reported in India in 1891 (Kumar and Beniwal,
1991), and has subsequently been observed in Brazil, Myanmar, Egypt, El
Salvador, India, Israel, Malaysia, Mexico, Nicaragua, Oman, Pakistan, South
Africa, Sudan, Spain, Swaziland, Uganda and the USA (Flechtmann et al.,
1973; Crookes and Rijkenberg, 1985; Lim and Khoo, 1985; Kumar and Beni-
wal, 1991; Ploetz, 2001; Kvas et al., 2007; S. Freeman, Bet Dagan, 2007, unpub-
lished data; G.I. Johnson, Jamison, Australia, 2007, personal communication;
J.F. Leslie, Manhattan, Kansas, 2007, personal communication). Since the
pathogen is easily disseminated in infected germplasm and there are con-
spicuous gaps in the disjunct distribution (note African and American records),
malformation is probably more widely spread.
Symptoms
Malformation affects vegetative and oral meristematic tissues (Fig. 8.20)
(Ploetz, 2001). Vegetative malformation is most serious on seedlings and
small plants in nurseries, especially where seedlings are grown beneath
affected trees, a common practice in the Middle East (Ploetz et al., 2002;
Youssef et al., 2007). Vegetative malformation also occurs on mature trees.
Apical and axillary buds produce misshapen shoots with shortened inter-
nodes and dwarfed leaves that are brittle and recurve towards the sup-
porting stem (Fig. 8.20). Shoots may not expand fully, resulting in a bunched
appearance (i.e. the bunchy-top symptom of the disease).
Floral malformation is most important. Affected inorescences usually
do not set fruit or fruit are aborted. Primary or secondary axes on affected
panicles are shortened, thickened and highly branched (Fig. 8.20). Malformed
panicles produce up to three times the normal number of owers, range from
one-half to two times the normal size, and have an increased proportion of
male to perfect owers. Malformed panicles may also produce dwarfed and
distorted leaves (exhibit phyllody).
Aetiology
The aetiology of malformation has been controversial for almost as long as
the disease has been recognized (Ploetz, 2001). Suggested causes include
mites (Narasimhan, 1954), nutritional problems (Prasad et al., 1965), physio-
logical or hormonal imbalances (Dang and Daulta, 1982; Singh and Dhillon,
1989), viruses (Kauser, 1959) and unknown causes (Kumar and Beniwal,
1991). Summanwar et al. (1966) demonstrated that a fungus, identied then
as Fusarium moniliforme Sheld., was responsible for the oral phase of this
disease. Varma et al. (1974) later showed that F. moniliforme also caused
R.C. Ploetz and S. Freeman 262
(a)
(b)
Fig. 8.20. Among the symptoms caused by malformation are: (a) in panicles, an in-
crease in the size and number of owers and interspersed oral and vegetative organs
(phylody); and (b) in vegetative shoots, compact or retarded growth of buds and brittle,
dwarfed and recurved leaves. Symptoms in (a) are on Haden in Michoacan, Mexico
and are associated with an undescribed species in the Gibberella fujikuroi species
complex, whereas (b) is on a Van Dyke plant articially inoculated with an isolate of
Fusarium mangiferae (Photographs courtesy of R.C. Ploetz).
(a)
(b)
Foliar, Floral and Soilborne Diseases 263
vegetative malformation. This pathogen has had several synonyms in
the literature, including: Fusarium subglutinans (Wollenweb. and Reinking)
Nelson, Toussoun and Marasas; F. moniliforme Sheldon var. subglutinans
Wollenweb. and Reinking; and F. moniliforme Sheldon emend. Snyd and
Hans. Subglutinans sensu Snyd., Hans. and Oswald.
In 2002, 29 strains of the pathogen from Egypt, Florida USA, Israel,
Malaysia and South Africa were redescribed as a new species in the Gibberella
fujikuroi species complex (GFSC), Fusarium mangiferae Britz, Wingeld and
Marasas (Steenkamp et al., 2000; Britz et al., 2002). Fusarium mangiferae resem-
bles morphologically other members of the GFSC. It was established based
on -tubulin and histone H3 DNA sequences, subtle morphological differ-
ences, and because most of the examined strains had been shown in previous
studies to cause malformation on articially inoculated mango. The presence
of F. mangiferae has been veried in India (ODonnell et al., 1998; Zheng and
Ploetz, 2002), Oman (Kvas et al., 2007) and Spain (S. Freeman, Bet Dagan,
2007, unpublished results). Although a recent report from Pakistan mentions
F. mangiferae, the identity of the pathogen there is not clear since the authors
only discussed morphological characteristics of the pathogen (Iqbal et al.,
2006).
Based on DNA sequence data (ODonnell et al., 1998, 2000; Steenkamp
et al., 1999, 2000), F. mangiferae is related to a lineage that includes Fusarium
fujikuroi Nirenberg, Fusarium proliferatum (Matsushima) Nirenberg, and
Fusarium sacchari (E. J. Butler) W. Gams (Marasas et al., 2006), and corresponds
to the Asian Clade of ODonnell et al. (1998). Based on combined sequence
data for ve genes, the closest known relative of F. mangiferae is an isolate
from tropical rainforest soil in Papua New Guinea (Marasas et al., 2006).
Fusarium mangiferae produces white, occose mycelium on PDA with
light- to dark-purple pigments in the agar (Leslie and Summerell, 2006).
Cream-coloured sporodochia on carnation leaf agar (CLA) produce abun-
dant thin-walled, long, slender and straight to slightly curved, three- to ve-
septate macroconidia with curved apical cells and foot-shaped basal cells
(Fig. 8.21). Single celled, rarely single septate, obovoid microconidia are
produced in false heads on polyphialides with two to ve conidiogenous
openings and on monophialides. Microconidial chains and chlamydospores
are absent.
A second species, Fusarium sterilihyphosum Britz, Wingeld and Marasas,
was described originally for isolates from a small area in South Africa (Britz
et al., 2002). In subsequent work, it was detected in Brazil (Ploetz, 2003; K.
ODonnell, unpublished results), and was recently shown to cause malfor-
mation in Brazil after articial inoculation (Lima et al., 2006b). On PDA, colo-
nies of F. sterilihyphosum produce white, occose mycelium with rose to light
purple pigmentation in the agar (Leslie and Summerell, 2006). Uncommon,
cream- to orange-coloured sporodochia are produced on CLA that produce
rare, long, slender, three to ve-septate macroconidia (Fig. 8.22). On mono-
and polyphialides, obovoid, oval to allantoid microconidia that are usually
single celled are produced on false heads. Distinctive sterile coiled hyphae
are produced by some isolates of this species.
R.C. Ploetz and S. Freeman 264
Fusarium sterilihyphosum is relatively uncommon in South Africa and
Brazil where, respectively, F. mangiferae and a third, unnamed species that
resembles F. sterilihyphosum morphologically, predominate. The latter taxon
is phylogenetically distinct from F. mangiferae and F. sterilihyphosum, produces
a unique teleomorph in the GFSC, and has been shown to cause malforma-
tion (Lima et al., 2006a, b).
Fusarium mangiferae has not been found in either Brazil (Lima et al., 2006a,
b) or Mexico (Rodrguez-Alvarado et al., 2006, 2008). In the latter studies,
isolates that were recovered from malformed trees resembled F. sterilihypho-
sum in that they induced malformation symptoms, formed sterile coiled
hyphae and produced a PCR fragment that is also produced by isolates of F.
sterilihyphosum (see below). Translation elongation factor-1 DNA sequences
for isolates from several areas in Mexico are identical, but differ signi-
cantly from other taxa in the GFSC; they probably represent a new species
(Rodrguez-Alvarado et al., 2006, 2008; K. ODonnell, Peoria, 2007, personal
communication). Additional work is needed to clarify relationships among the
strains in Brazil and Mexico, and whether they are found elsewhere in the Amer-
icas. Likewise, whether F. mangiferae is found outside Florida USA in the western
hemisphere should be determined; it predominates in the eastern hemisphere.
PCR primer pairs have been used to diagnose some of the above taxa.
Zheng and Ploetz (2002) developed a pair, 1-3F/R, that amplies a 608 bp
fragment for F. mangiferae. It has been used extensively for diagnostic pur-
poses (Youssef et al., 2007). Another pair, 61-2F/R, developed to diagnose
Fusarium verticilloides (published as F. moniliforme in Mller et al., 1999), did
not amplify F. mangiferae DNA, but when amplication protocols were mod-
ied, amplied a 445 bp-fragment for strains of F. sterilihyphosum and the
new species from Mexico (Zheng and Ploetz, 2002; Rodrguez-Alvarado et al.,
2008). It has not been tested with the unnamed pathogen from Brazil.
(a)
(b)
(c)
(d)
(e)
(f )
Fig. 8.21. Microscopic features of Fusarium mangiferae: (a) and (b), macroconidia;
(c) and (d), microconidia, and (e) and (f), microconidia in situ on carnation leaf agar.
Scale bars for (a)(d) = 25 m, and (e)(f) = 50 m (Photographs courtesy of Suzanne
Bullock).
Foliar, Floral and Soilborne Diseases 265
Three other taxa have been associated with mango malformation. Fusar-
ium oxysporum Schlecht emend. Snyder and Hansen (Fig. 8.23) was reported
in Egypt and Mexico (Bhatnagar and Beniwal, 1977; Kumar and Beniwal,
1991), but these reports appear to be erroneous since bona de, vouchered
specimens have neither been described nor shown to cause the disease (Plo-
etz, 2001; Rodrguez-Alvarado et al., 2008). Fusariumsp. nov. (Britz et al., 2002)
and F. proliferatum (Gibberella intermedia (Kuhlman) Samuels, Nirenberg and
Seifert) were recovered from malformed mango trees in Malaysia (Leslie,
1995), but their pathogenicity has not been determined.
Epidemiology
Although malformation has been reproduced with F. sterilihyphosum and the
unnamed taxa from Brazil and Mexico, no work has been conducted on the
epidemiology of disease that is caused by these pathogens. Thus, results
below are from work on F. mangiferae or what is presumed to be this species.
Fusarium mangiferae is spread by grafting and in infected nursery stock
(Prakash and Srivastava, 1987). Since seed do not appear to harbour the
fungus (Saeed and Schlosser, 1972; Youssef et al., 2007), seedlings should be
disease free. Microconidia of F. mangiferae are probable infective propagules
since they are the primary spores that are produced by the fungus and form
profusely on dead malformed tissues. The disease spreads slowly in orchards,
perhaps because conidia of the pathogen die quickly when exposed to sun-
light (Manicom, 1989). Experimentally, populations of F. mangiferae in infected
panicles in Egypt and Israel declined rapidly during the summer (Youssef
et al., 2007). Wounding enhances infection and subsequent disease develop-
ment (Ploetz, 2001).
(a) (c) (e)
(b) (d) (f )
(g)
(h)
Fig. 8.22. Microscopic features of Fusarium sterilihyphosum: (a) and (b), macroconidia; (c) and
(d), microconidia; (e) and (f), coiled hyphae; and (g) and (h), microconidia in situ on carnation
leaf agar. Scale bars for (a)(d) = 25 m and (e)(f) = 50 m (Photographs courtesy of Suzanne
Bullock).
R.C. Ploetz and S. Freeman 266
The mango bud mite, Aceria (Eriophyes) mangiferae Sayed, is often
observed in high numbers on malformed trees and has been indicted as the
cause, or a factor in the development, of this disease (Narasimhan, 1954,
1959; Nariani and Seth, 1962). Circumstantial evidence indicates that the mite
does not cause malformation (Ploetz and Prakash, 1997); for example, A.
mangiferae is present in Australia, but the disease is not (Ridgeway, 1989).
However, A. mangiferae probably vectors the pathogen. It has been recovered
from the mites body on culture media (Crookes and Rijkenberg, 1985; Sattar,
Ismailia, 2006, personal communication), and was recently shown to adhere
to its body (Gamliel-Atinsky, Freeman and Palevsky, unpublished data) (Fig.
8.24). The mite cannot ingest the pathogen, due to its small mouthparts.
However, it was able to move spores of F. mangiferae to infection courts in
mango buds via external contamination of its body, and increased infection
Fig. 8.23. Microscopic features of Fusarium oxysporum: (a) sporodochia, (b) macro-
conidia, (c) microconidia in false head on monophialide, (d) terminal and intercalary
chlamydospores, and (e) macroconidia and microconidia (Photographs courtesy of
K. ODonnell).
(a) (b)
(c)
(d) (e)
Foliar, Floral and Soilborne Diseases 267
of buds by the pathogen (Gamliel-Atinsky et al., 2007). Presumably, the mites
feeding on buds facilitated infection. Aceria mangiferae does not appear to
play a signicant role in disseminating the pathogen among trees. In Israel,
mites were not found in traps that were designed to monitor their movement
in a malformed mango orchard, although high numbers of F. mangiferae
conidia were recorded in the air in this orchard (Gamliel-Atinsky et al.,
2007).
The distribution of F. mangiferae in affected trees suggests that vegetative
and oral buds are the primary sites of infection and that systemic coloniza-
tion of older, subtending tissues does not occur. Freeman et al. (1999) trans-
formed isolates of F. mangiferae from mango with the uidA reporter gene
(-glucuronidase), and used them to articially inoculate mango. Their results
veried that bud and ower tissues of the host are primary infection sites,
and that wounds provide points of entry for the pathogen. In Florida USA, F.
mangiferae was restricted almost entirely to malformed oral and vegetative
tissues (Ploetz, 1994). Levels of infection were highest in malformed owers
and vegetative shoots (c.6585%), were much lower or non-existent in asymp-
tomatic tissues (011%), and were rare in branches (04%) even when they
supported malformed owers or shoots. Remnant infections of F. mangiferae
in scaffold branches and trunks were restricted almost exclusively to branch
scars or dormant apices (Freeman, unpublished data). Reports of root
infection by F. oxysporum (Kumar and Beniwal, 1991) or F. mangiferae (Abdel-
Sattar, 1973; Kumar and Beniwal, 1991) causing malformation in seedling
plants have not been corroborated. Although roots can be infected by
GFP-labelled microconida of
Fusarium mangiferae
20.0 m
Aceria mangiferae
Fig. 8.24. Body of the mango bud mite, Aceria mangiferae, to which green
uorescent protein (gfp)-labelled microconidia of Fusarium mangiferae have adhered
(Photograph courtesy of E. Gamiel-Atinsky).
R.C. Ploetz and S. Freeman 268
F. mangiferae, these infections are not systemic and do not appear to result in
symptom development (Youssef et al., 2007).
The localized and variable levels of infection by F. mangiferae that have
been noted in diseased and non-symptomatic tissue (Ploetz, 1994; Youssef
et al., 2007), suggest that there are thresholds of infection, whereby malfor-
mation develops only after a sufcient proportion of a host meristem is colo-
nized by the pathogen. This hypothesis is supported by the long latent period
that exists before symptoms develop in articially inoculated plants and the
hormonal perturbations that probably occur when meristematic tissues are
infected with this pathogen (van Staden et al., 1989; van Staden and Nichol-
son, 1989; Ploetz, 2001).
Management
Management of malformation can be difcult. New plantings should be
established with pathogen-free nursery stock. Scion material should never be
taken from an affected orchard, and any affected plants that are observed in
the nursery should be removed and burned immediately. Nurseries should
not be established in orchards that are affected by malformation. Once the
disease is found in an orchard, control is possible, but time consuming. In
these cases, cultural management has been most effective (Narisimhan, 1959;
Singh et al., 1974; Manicom, 1989). All affected terminals and the subtending
three nodes are cut from trees, removed from the eld and burned. Unfortu-
nately, producers may be unwilling to devote the effort that is required to
ensure that this approach succeeds. In addition, it may be difcult to impose
this treatment on large trees.
A diverse array of pesticides, hormones and growth regulators has been
tested against malformation, but these measures have been marginally effec-
tive. Singh et al. (1994) obtained moderate control with sulfates of cobalt (Co),
cadmium (Cd) and nickel (Ni) in India, but it is unlikely that these toxic com-
pounds could be used safely. Darvas (1987) reduced the percentage of mal-
formed inorescences from 96% to 48% by injecting Keitt trees with the
fungicide fosetyl-Al. This reduction was signicant (P < 0.05), but the increase
in fruit yield, 4695 kg of fruit/tree, was not. Other fungicidal compounds
have been generally less effective (Diekman et al., 1982; Chakrabarti and
Ghosal, 1989). In general, the protected, internal location of the pathogen in
affected trees makes it difcult to control this disease. When applied as foliar
sprays or via continuous drip irrigation, prochloraz reduced the severity of
malformation signicantly in Israel, but this was dependent on the timing of
application (Freeman et al., unpublished data). Although disease was not
completely controlled, this and other systemic fungicides might be useful in
future integrated management programmes that would incorporate other
measures such as removal of symptomatic terminals and use of tolerant
cultivars.
Prakash and Srivistava (1987) indicated that There is great variation in
the susceptibility of existing varieties. Unfortunately, controlled inoculations
have not been used to determine cultivar resistance, and these reports have
come from non-replicated tests; cultivars listed as resistant may have come
Foliar, Floral and Soilborne Diseases 269
from healthy nursery stock or may have escaped infection once planted in
the eld (Ploetz, 2001). For example, Bastawros (1996) reported that two
newly introduced cultivars in Egypt, Kent and Keitt, were immune (0%
disease); however, these cultivars are susceptible in Florida USA (Ploetz,
unpublished data). Controlled inoculations with grafted plants of different
genotypes and quantied levels of virulent isolates of F. mangiferae have not
been reported.
Recently, Ploetz (unpublished data) utilized previously described proto-
cols (Freeman et al., 1999) to assess malformation development on grafted
plants of diverse genotypes. Disease development was affected by: the length
of time after inoculation and inoculated apical buds remained dormant after
inoculation; the isolate of F. mangiferae that was used for inoculation; and
mango genotype. Virulent isolates, patience (latent period ranged from 40 to
210 days), and sufcient replication were needed to successfully conduct
screenings for response to malformation. Future work is warranted to investi-
gate attributes that are related, and might predict resistance, to this disease.
The symptoms of malformation suggest that a hormone imbalance
occurs in affected tissues. Singh and Dhillon (1989) assayed levels of indole
acetic acid (IAA), gibberellic acid (GA
3
) and zeatin in malformed and healthy
mango seedlings. Whereas IAA and GA
3
levels were, respectively, about ten
and ve times lower in malformed plants, levels of zeatin were about ve
times higher. Van Staden and colleagues (Nicholson and van Staden, 1988;
van Staden and Nicholson, 1989; van Staden et al., 1989) examined specic
cytokinins produced by mango and F. moniliforme (presumably F. mangiferae).
They determined that the cytokinin complements in healthy and malformed
panicles differed qualitatively and quantitatively, and that the pathogen pro-
duced some of the hormones and metabolites that were implicated in dis-
ease development. However, it was impossible to assign unequivocal roles
for production of hormones by the host and pathogen and the subsequent
development of symptoms. For example, whether production of hormones
by the pathogen directly caused the noted changes or whether hormone pro-
duction by the host was somehow altered in the presence of the pathogen
was not clear. Additional work would be needed to clarify these interactions,
and whether F. sterilihyphosum and the unnamed pathogens in Brazil and
Mexico also produce cytokinins or other hormones in affected plants.
Parasitic plants
The family Loranthaceae contains several parasitic plant species that affect
mango. In Malaysia, Dendrophthoe (fomerly Loranthus) pentandra Linn. is
the most important species (Lim and Khoo, 1985). Other Dendrophthoe spp.,
Elytranthe spp. and Viscum spp. are also known in Malaysia, but are less
important. In India, Dendrophthoe falcata (formerly Loranthus longiorus) is
common, and other, less frequently encountered, species include Macrosolen
cochinchinensis, Helicanthes elasticus and Elytranthe capitellata (Majumder
and Sharma, 1990). These parasites are usually only important in neglected
R.C. Ploetz and S. Freeman 270
orchards. Although they are photosynthetically self-sufcient, the plants
obtain water and minerals from the host plant via haustoria that penetrate
and colonize the host vascular system. In severe cases, the removal of water
and minerals from parasitized branches is sufcient to reduce the vitality and
yields of trees.
Since the appearance of these plants is distinct from the mango host, they are
easily distinguished in infected trees (Lim and Khoo, 1985). The points at which
the mango host is penetrated are usually characterized by swollen growths
called burrs. Lim and Khoo (1985) and Majumder and Sharma (1990) indicated
that affected portions of trees should be removed far enough below burrs to
remove haustoria of the parasite. After affected tissues are removed, cut sur-
faces can be treated with creosote or other wound dressings. These plants can
also be treated with herbicides, such as 2,4-dichlorophenoxyacetic acid (2,4-D),
but these are dosage sensitive treatments and pose a risk to the host plant.
Phoma blight
Phoma blight is widespread in India (Prakash and Singh, 1977). It occurs only on
old leaves. Initially, lesions are minute and yellow-brown (Prakash and Singh,
1977). As they expand they darken to brown or cinnamon, become irregular, and
may ultimately develop dark margins and dull-grey centres. In severe cases,
necrotic patches as large as 13 cm in diameter may form that cause defoliation.
The disease is caused by Phoma glomerata (Corda) Wollenw. and Hochapf
(Prakash and Singh, 1977). It forms globose to obpyriform, light-coloured to car-
bonaceous pycnidia that average 30400 m in diameter. Pycnidia have one to
three ostioles, form singly or in clusters, and have short necks. On PDA, pyc-
nidia and conidia are abundant. Conidia are hyaline to dark coloured, ovoid to
ellipsoid, unicellular or occasionally bicellular, and average 8.3 3.2 m.
Phoma leafspot
Another Phoma sp. causes a leafspot in India (Prakash and Singh, 1976b), and
is referred to as phoma leafspot. On young leaves, Phoma sorghina (Sacc.)
Boerema. Doren. and Vankest causes irregular to roughly circular, water-
soaked spots, up to 2.5 mm in diameter. Lesions are brown with a yellow to
brown margin. Lesions on midribs are elongated and more conspicuous, and
may coalesce to up to 14 cm in diameter. They can be confused with those
caused by anthracnose.
Pink disease
A basidiomycete, Erythricium salmonicolor (Berk. and Broome) Burdsall (syn-
onyms: Corticium salmonicolor Berk. and Broome, and Phanerocbaete salmoni-
color (Berk. and Broome) Jlich; anamorph: Necator decretus Massee) causes
Foliar, Floral and Soilborne Diseases 271
pink disease. Pink disease affects many economically important woody
plants in the humid tropics, where it is one of the most destructive diseases
of mango (Holliday, 1980). The disease is also known as cobweb, rubellosis
and thread blight (Prakash and Srivistava, 1987). It has been most thor-
oughly studied on rubber, Hevea brasiliensis, an important host in South-east
Asia (Rao, 1975). On mango, pink disease can signicantly reduce tree vigour
and yield, especially in 6- to 15-year-old trees (Lim and Khoo, 1985).
Symptoms rst appear as white, felty mycelial threads on twigs and
branch crotches (Lim and Khoo, 1985). If favourable conditions persist, the
mycelial threads coalesce to form a rough, pink crust on the bark surface. This
stage usually takes 1 to several months to develop and coincides with the
penetration of bark and internal colonization of the tree. Affected bark often
cracks. As the fungus kills the vascular and cambial areas beneath the bark,
branches above the colonized areas die, resulting in a sparse, patchy canopy.
Two types of sporulation occur (Holliday, 1980; Lim and Khoo, 1985).
Erythricium salmonicolor produces a smooth, clammy, pinkish white hyme-
nium over the pink crust it forms on bark. Basidiospores form on the hyme-
nium and are borne on sterigmata on narrowly clavate to cylindrical basidia
(Fig. 8.25). Basidiospores are hyaline, broadly ellipsoidal and 810 57 m.
Conidia of N. decretus, which are produced in reddish-orange sporodochia,
are hyaline, ellipsoidal, unicellular and 1018 612 m. Although the infec-
tion process has apparently not been studied in mango, basidiospores can
infect rubber trees (Holliday, 1980). The anamorph and teleomorph are
(c)
(a)
(b)
(f )
(e)
(d)
100
20
Fig. 8.25. (a) Conidium-bearing pustule, and (f) conidiogenous cells and conidia
of Necator decretus, and (b) hymenium, (c) basal hyphae, (d) immature and mature
basidia, and (e) basidiospores of its teleomorph, Erythricium salmonicolor (Source:
from CMI description no. 511).
R.C. Ploetz and S. Freeman 272
formed during wet conditions, and conidia and basidiospores are dispersed
by rainsplash and wind.
Pink disease management relies on early detection and removal of
affected tissues from orchards. When removal of syptomatic tissues is imprac-
tical, control depends upon treatment with fungicides. Several protectant
and systemic fungicides are effective (Lim, 1994). They should be used as
soon as symptoms are evident, and as long as the disease is present. All cul-
tivars of mango that have been tested in Malaysia are susceptible (Lim and
Khoo, 1985).
Powdery mildew
Powdery mildew is a widespread and important disease of leaves, panicles
and fruit. The disease can result in yield reductions as high as 90%, due
mainly to its effect on fruit set and development (Schoeman et al., 1995).
Symptoms
Mango cultivars vary in their response to powdery mildew (Palti et al., 1974).
On the most susceptible cultivars, virtually all foliar, oral and fruit parts of
the plant are affected (Plate 46). Powdery growth can cover all tissues on
panicles, resulting in a brown, shrivelled necrosis. Since fruit set and reten-
tion can be affected, the disease can have a profound impact on yield. Foliage
can also be damaged signicantly, and young leaves are most susceptible.
White, powdery coatings of conidia develop on either side or both sides of
leaves, depending on the cultivar. When damage occurs on the undersides of
leaves it is often restricted to the mid-rib. In all cases, leaves become dis-
torted, and affected areas turn purple and ultimately necrotic.
Aetiology
Powdery mildew is caused by Oidium mangiferae Berthet, a host-specic fun-
gus (Prakash and Srivistava, 1987; Ploetz and Prakash, 1997). It was rst
described in Brazil (Berthet, 1914), and is now recognized in most mango-
producing regions (Palti et al., 1974). Conidium and haustorium traits indi-
cate that O. mangiferae belongs to the Erysiphe polygoni group (Johnson, 1994a).
Although the pathogen was originally classied as Erysiphe cichoracearum by
Wagle (1928), Uppal et al. (1941) noted that it produced saccate and lobed
appressoria, which are not characteristic of E. cichoracearum. The pathogen
has no known teleomorph, a common trait for powdery mildew fungi in the
tropics (Holliday, 1980). Conidia of O. mangiferae are unicellular, hyaline,
elliptical to barrel-shaped and measure 3343 1828 m (Uppal et al., 1941;
Palti et al., 1974). They are produced in large numbers on host surfaces, and
impart a powdery appearance to affected tissues (Plate 46). The lengths of
germ tubes vary depending upon RH, and they terminate in appressoria. Glob-
ular haustoria form in host epidermal cells. Conidiophores are of the pseudoid-
ium type, with two to four septa and a straight basal cell (Boesewinkel,
1980).
Foliar, Floral and Soilborne Diseases 273
Epidemiology
Powdery mildew is most severe during cool, dry weather. Conidia are dis-
seminated by wind and are released on a diurnal basis (Schoeman et al.,
1995). Peak spore release, between 11:00 to 16:00 h, was positively correlated
with hourly temperature and negatively correlated with RH, vapour pres-
sure decit and leaf wetness (all P < 0.01). Conidia germinate at temperatures
ranging from 9 to 32C (23C is optimal), and at RH as low as 20% (Palti et al.,
1974). Since germination occurs in such a wide range of RH, disease develop-
ment is usually independent of this parameter. Infection can occur within
57 h, and conidia are produced within 5 days of infection. Disease develop-
ment occurs within a very broad range of temperatures, 1031C. Gupta
(1989) reported that dry weather encouraged disease development.
Management
Mango cultivars vary in their resistance to powdery mildew (Palti et al.,
1974). Zill, Kent, Alphonso, Seddek and Nam Doc Mai are very sus-
ceptible; Haden, Glenn, Carrie, Zebda, Hindi be Sennara, Ewaise and
Keitt are moderately susceptible; and Sensation, Tommy Atkins and
Kensington are slightly susceptible (Ploetz et al., 1994; Nofal and Haggag,
2006). In India, Tiwari et al. (2006) reported that Baigan Phalli, Barbalia,
Dabari, Dilpasand, Khirama, Nagarideeh, Oloor and Totapari were
highly resistant and Amrapali was most susceptible.
Schoeman et al. (1995) recommended that the rst fungicide application
to control this disease should occur when panicles begin to change colour.
Assuming an effective period of 3 weeks for a given application, they con-
cluded that applications should continue every third week until panicle sus-
ceptibility decreased at the end of fruit set. Powdery mildew is easily
controlled with S (Johnson, 1994a). Other fungicides are effective, but many
have negative environmental impacts (Ray, 2003; Tavares et al., 2004). Foliar
sprays of di-potassium hydrogen orthophosphate (K
2
HPO
4
) and potassium
di-hydrogen orthophosphate (KH
2
PO
4
), systemic fungicides, and an alterna-
tion of fertilizer and systemic fungicides inhibited powdery mildew on pan-
icles (Reuveni et al., 1998; Nofal and Haggag, 2006). Treatments of the
fertilizers with half or a quarter of the recommended rate of sterol-inhibitor
fungicides and Kresoxym-methyl provided protection comparable with or
superior to that of standard fungicides alone (Oosthuyse, 1998; Reuveni et al.,
1998). Sulfur can burn owers and young fruit during warm, sunny condi-
tions (Johnson, 1994a), and three fungicides used during bloom, dinocap,
fenbuconazole and hexaconazole, can reduce pollen germination (Dag et al.,
2001).
Scab
Elsino mangiferae Bitancourt and Jenkins (anamorph: Sphaceloma mangiferae
Bitancourt and Jenkins) causes scab on mango (Bitancourt and Jenkins, 1943;
Cook, 1975). The disease was rst recognized in Cuba and Florida USA in the
R.C. Ploetz and S. Freeman 274
1940s and is now widespread in the western hemisphere. Scab is important
in nurseries since young host tissues are most susceptible, and because moist
environments aid infection (Ruehle and Ledin, 1955). Lesions, usually rst
observed on the underside of leaves, are dark brown to black, and 12 mm in
diameter. They may enlarge to 5 mm in diameter and become light grey with
narrow, dark borders. Affected foliage develops a distorted appearance, and
greyish blotches are produced on stems.
Elsino mangiferae produces brownish ascocarps, 3048 80160 m, in
the host epidermis. Globular asci, 1015 m in diameter, are dispersed in
ascocarps, and contain one to eight hyaline ascospores. Ascospores are
46 1013 m, three septate and constricted at the median septum; the sub-
apical cell is longitudinally septate. Conidiophores of S. mangiferae are erect,
sinuous, 2.53.5 1235 m, and wider at the base. Conidia are brown, one
or two celled, and 24 629 m.
Young host tissues are most susceptible. Rainy weather promotes sporu-
lation of the fungus, but specic information is not available on the epidemi-
ology of scab. Whether conidia and ascospores are infectious is not known.
Seca and sudden decline
This is a disease that is known by several different names in Brazil and the
Middle East and is the only one that routinely kills mango trees. Seca (dry-
ing), murcha (withering), branch blight and Recife sickness in Brazil, was
rst recognized in Pernambuco in 1938, and is now also found in Bahia,
Goias, the Federal District, Rio de Janiero and So Paulo (Ribeiro, 1997; Colo-
simo et al., 2000; Silveira et al., 2006). It threatens neighbouring states due to
its efcient movement in infected propagation materials, on pruning equip-
ment, and via a mobile beetle vector.
In 1998, a disease termed sudden decline began to kill trees in Oman
(Al Adawi et al., 2003, 2006), about the same time a similar problem (i.e. quick
decline or sudden death) was observed in Pakistan (Malik et al., 2005). In
many ways these diseases resembled seca. Circumstantial evidence sug-
gested that the disease was introduced from Brazil by a producer with
orchards in Oman and Pakistan (M. Deadman, Muscat, 2005, personal com-
munication). By 2007, many mango-producing areas in Oman and Pakistan
were affected and uncontrolled dissemination of infected germplasm may
have spread the disease elsewhere in the region. Its spread into India should
be investigated (A.W. Cooke, Indooropilly, 2007, personal communication).
Symptoms
Symptoms include: discoloration of the vascular cambium; exudation of an
amber-coloured gum from the trunk and branches, particularly from galler-
ies of the putative beetle vector of the pathogen; wilting; rapid death of
branches and entire trees without defoliation; and a scorched appearance of
dead trees (Plate 47) (Junqueira et al., 2002; Al Adawi et al., 2006). On grafted
trees, scions, rootstocks or both may be susceptible and exhibit vascular
Foliar, Floral and Soilborne Diseases 275
symptoms. In Oman, where susceptible Omani seedlings are used as root-
stocks, the disease is frequently a problem of rootstocks (Al Adawi et al.,
2006), whereas in Brazil, the disease is associated with the scion (P < 0.01)
(Colosimo et al., 2000); rootstock cultivar had an insignicant impact on dis-
ease development in the latter work (P > 0.05). When disease development
begins in the canopy, symptoms may initiate in a branch or portion of a tree,
but death of the entire plant usually follows. Where root/rootstock infection
is involved, sudden death of the entire tree usually occurs.
Aetiology
Ceratocystis mbriata Ellis and Halst. sensu lato (s.l.) (anamorph: Thielaviopsis
sp.) was reported in Brazil in the 1930s (Viegas, 1960; Ribiero, 1980; Silveira
et al., 2006), and is recognized as the primary cause of seca. Diplodia reciensis
Batista (= Lasiodiplodia theobromae?) was indicted as the cause of Recife sick-
ness in Brazil (Batista, 1947), but it probably plays no role or a secondary role
in the development of this disease (see below). In Oman, C. mbriata s.l.
causes sudden decline, but L. theobromae and Ceratocystis omanensis Al Subhi,
M.J. Wingf., M. van Wyk and Deadman are also associated with the disease
(Al Adawi et al., 2006; van Wyk et al., 2007). The ease with which L. theobromae
and the difculty with which C. mbriata s.l. are recovered from affected trees
may have been responsible for previous assumptions that D. reciensis
caused Recife sickness in Brazil and L. theobromae caused sudden decline in
Oman (Batista, 1947; Ploetz and Prakash, 1997; Al Adawi et al., 2003, 2006).
Ceratocystis contains many pathogens, particularly of trees (Kile, 1993).
The wide host range of C. mbriata s.l. led Webster and Butler (1967) to
hypothesize that it was a species complex, and DNA sequences have begun
to delineate some of the host-specic, often morphologically indistinct, taxa
in the species (van Wyk et al., 2007). A contemporary view is that C. mbriata
sensu stricto (s.s.) specically refers to the cause of black rot of sweet potato
(Ipomoea batatas L.) on which it was rst described (Halsted and Fairchild,
1891). Other cryptic, monophyletic lineages of C. mbriata s.l. have been
described as distinct species (Engelbrecht and Harrington, 2005; Johnson
et al., 2005; van Wyk et al., 2005, 2007), and more will likely follow.
Two new Ceratocystis spp. have been described on mango in the Oman
Gulf region. Ceratocystis omanensis belongs to the Ceratocystis moniliformis
Hedgc. s.l. species complex (Al Subhi et al., 2006), which are typically not
primary pathogens. Ceratocystis omanensis is a minor pathogen on mango.
The primary sudden decline agent in Oman and Pakistan, C. mbriata s.l.,
represents a monophyletic lineage based on ITS, -tubulin and translation
elongation factor (TEF) 1- DNA sequence comparisons, and it has unique
morphological characteristics; it was renamed Ceratocystis manginecans M.
van Wyk, A Al Adawi and M.J. Wingf. sp. nov. (van Wyk et al., 2007).
On 2% malt extract agar (MEA), colonies of C. manginecans are greyish
olive and have a banana odour (van Wyk et al., 2007). Hyphae are smooth
and segmented (Fig. 8.26). Ascomatal bases are globose, black and (153)192
254(281) m in diameter; ascomatal necks are dark brown, lighter towards
the apices (514)557635(673) m long, (25)3242(48) m, wide at the
R.C. Ploetz and S. Freeman 276
base, and (14)1622(26) m wide at the tip; and ostiolar hyphae are hya-
line, divergent and (42)4559(69) m long. Asci are evanescent, and
ascospores are hyaline, hat-shaped, 34 m long, and 45 m wide without,
and 78 m wide within the sheath. Primary conidiophores are phialidic,
lageniform, hyaline, (72)81109(144) m long, 57(9) m wide at the base,
68(9) m wide at the broadest point, and 36 m wide at the tip. Secondary
conidiophores are tube like, ared at the mouth, short, hyaline, (59)65
77(84) m long, 58 m wide at the base and (5)68 m wide at the tip.
Primary conidia are hyaline, cylindrical, (15)2329(33) m long, and 36
m wide; and secondary conidia are hyaline, barrel shaped, (8)911(12) m
long, and 57(8) m wide. Chlamydospores are brown, thick-walled, glo-
bose to subglobose, (11)1214 m long and 911(12) m wide.
Two isolates of C. mbriata s.l. from mango in Brazil (CBS 114721 and CBS
600.70) have been compared to isolates of C. manginecans (van Wyk et al.,
2005, 2007). They are similar to, but differ signicantly from, C. manginecans.
They reside with C. manginecans in a clade that contains other New World
(a)
(d) (g)
(c)
(e) (b) (f)
Fig. 8.26. Microscopic features of Ceratocystis manginecans: (a) globose ascoma,
(b) divergent ostiolar hyphae, (c) hat-shaped ascospore, (d) segmented hypha,
(e) primary phialidic conidiophore with emerging cylindrical conidia, (f) secondary
conidiophore with emerging chain of barrel-shaped conidia, and (g) dematiaceous
chlamydospores and cylindrical- and barrel-shaped conidia. Scale bars: (a) = 100 m,
(b) = 20 m, (c) = 5 m, (d) = 20 m, (e) = 20 m, (f) = 20 m, (g) = 20 m.
(Source: van Wyk et al., 2007).
Foliar, Floral and Soilborne Diseases 277
species, Ceratocystis cacaofunesta and Ceratocystis platani. Research is needed
to: (i) examine additional isolates of C. mbriata s.l. from mango in Brazil; (ii)
describe the putative, new species; (iii) determine whether C. manginecans is
present in Brazil; and (iv) clarify pathogenic variation in the agent(s) in Oman
and Brazil. At least two pathotypes of C. mbriata s.l. are evident in Brazil
(Rossetto et al., 1996; Junqueira et al., 2002; Silveira et al., 2006).
Rossetto et al. (1996) evaluated 15 cultivars against two isolates of the
pathogen in Brazil. Eight-year-old trees were inoculated c.40 cm beneath
branch apices with IAC FITO 4905, which is pathogenic to Jasmim, and IAC
FITO 334-1, which is not. So Quirino, Irwin, Edwards and Van Dyke
were resistant, and IAC 100 Bourbon was moderately resistant. Glenn, Joe
Welch, Zill and Haden were susceptible, and Kent responded as Jas-
mim, resisting IAC FITO 334-1 and succumbing to IAC FITO 4905.
Epidemiology
Genotype has a profound impact on disease development, and severe epi-
demics occur wherever susceptible rootstocks and/or scions are used. Greater
disease develops when trees are stressed, although it is not clear whether this
results from an increased attraction of the vector to stressed trees or reduced
disease resistance in the host. The associated pathogens are moved easily in
infected germplasm, the route by which the diseases have spread in Brazil
and probably to Oman. Pruning implements also move the pathogen, and soil,
once infested with chlamydospores of the pathogen, can be a long-term reser-
voir of inoculum. Insect dissemination plays a particularly insidious role.
Beetles (Coleoptera: Scolytidae) are closely associated with seca in Brazil
(Batista, 1947; Viegas, 1960; Piza, 1966; Ribiero, 1980). Batista (1947) indicated
that Xyleborus afnis was the sole vector of D. reciensis. In contrast, Ribiero
(1980) reported that Hypocryphalus mangiferae Stebbing was the primary vec-
tor of C. mbriata s.l. (Fig. 8.27). It produced galleries in the cambium of
affected trees (Plate 47a), and was the only scolytid found on healthy, as well
as diseased, trees. Hypocryphalus mangiferae is also associated with the dis-
eases in Oman and Pakistan, where C. manginecans is recovered from the
insect and trees are commonly found with insect probing damage before dis-
ease develops (Al Adawi et al., 2006; van Wyk et al., 2007).
The interactions between H. mangiferae and the Ceratocystis pathogens
of mango are incompletely understood. In olfactometer tests in Brazil,
H. mangiferae was attracted to cultures of C. mbriata s.l., and larvae of the
insect were raised to adulthood on the fungus (Ribiero, 1980). Several other
species, many of which are in the genus Xyleborus, were also associated with
seca, but because they were found only in diseased trees they appeared to be
opportunistic feeders on C. mbriata s.l. rather than primary vectors. Although
the sequence of events in Brazil and the Oman Gulf has not been studied, it
is probable that H. mangiferae contaminates its body with these pathogens
while feeding in diseased trees and subsequently disseminates the pathogen
to healthy trees.
Hypocryphalus mangiferae is thought to be native to some of the same
areas in southern Asia where mango evolved (Wood, 1982; Butani, 1993;
R.C. Ploetz and S. Freeman 278
Atkinson and Peck, 1994; Mukherjee, 1997). Thus, the insect would have
been introduced into Brazil and would have been a new encounter, rather
than coevolved, relationship with C. mbriata s.l. (van Wyk et al., 2007). In
contrast, if C. manginecans were introduced into Oman and Pakistan from
Brazil, it may have then established a relationship with a native insect.
Although the available information suggests that the H. mangiferae
Ceratocystis interactions on mango were recent, opportunistic encounters in
the New World, additional work is needed.
Management
Given the ease with which these pathogens are moved and their destructive
impact, preventing their dissemination to new areas must be a high priority.
Pathogen-free propagation material should be used whenever new plantings
are established and germplasm is moved. Clean pruning implements should
be used in affected areas, and should be frequently disinfested with bleach,
formalin or other disinfestants (Junqueira et al., 2002). Trees that have been
killed by the disease must be removed and destroyed as they are signicant
reservoirs of inoculum and infested vectors. Where partially resistant culti-
vars are grown, the removal and burning of affected branches and treatment
of the exposed branch stubs with Cu fungicides is recommended (Ribeiro
et al., 1995; Ribeiro, 1997).
Managing these diseases with fungicides on susceptible cultivars would
be a challenge. External applications of protectant or systemic fungicides
would probably be ineffective, given the internal, protected location of the
pathogen. Injecting fungicides, as is done to control Dutch elm disease, might
be effective. However, this would probably not be allowed where concerns
Fig. 8.27. Hypocryphalus mangiferae, vector of the seca and sudden decline
pathogens (Photograph courtesy of R.C. Ploetz).
Foliar, Floral and Soilborne Diseases 279
with pesticide contamination of fruit exist. Treatment of germplasm collec-
tions and young, non-bearing trees might be the only situations in which
fungicide injection would be possible.
Genetic resistance offers the best hope for managing these diseases. Var-
ious levels of tolerance have been observed in Brazil and resistant clones
have been developed. However, pathogenic variation in the causal fungus in
Brazil has hindered progress (Rossetto et al., 1996; Junqueira et al., 2002; Sil-
veira et al., 2006). Although disease responses of some genotypes vary in dif-
ferent production areas in the country, Manga Dagua, Pico, IAC 101,
IAC 102, Edwards, Van Dyke and Carabao resist two races of the patho-
gen, and Rosa, Sabina, Sao Quirino, Oliveira Neto, Jasmim, Sensation,
Irwin and Tommy Atkins are generally tolerant (Ribiero, 1997; Junqueira
et al., 2002). Kent and Jasmim respond differentially (see above), and
Coquinho, Glenn, Joe Welch, Zill and Haden are susceptible. Although
Espada is also reported to be tolerant, old trees are frequently attacked. In
commercial orchards, the disease on Espada is managed by grafting onto
resistant rootstocks and pruning diseased branches. Colosimo et al. (2000)
worked with other scion cutivars, although in a single location (and against
a single pathotype?). They reported that Oliveira was most resistant, Car-
lota, Imperial, Extrema and Pahiri had intermediate resistance, and
Bourbon was most susceptible.
One must also recognize the impact of other diseases on different
cultivars. Carvalho et al. (2004) described two new cultivars, IAC 103 Espada
Vermelha and IAC 109 Votupa, which had moderate resistance to seca.
IAC 103 Espada Vermelha also had moderate resistance to powdery mil-
dew but was susceptible to anthracnose. Both cultivars were susceptible to
malformation.
Stigmina leafspot
Stigmina leafspot is caused by Stigmina mangiferae (Koorders) Ellis (synonym:
Cercospora mangiferae Koorders; a teleomorph for the fungus is not known).
Lim and Khoo (1985) indicated that the disease occurs on a range of cultivars,
and is most severe during rainy weather. Both young and old leaves are
affected. Dark-brown spots, 12 mm in diameter, are formed initially by the
fungus. These can enlarge and coalesce to 1 cm or larger, and are surrounded
by conspicuous chlorotic haloes that aid diagnosis of this disease. The fungus
produces large, olive-grey conidia, 3060 3.55.0 m, usually on the lower
leaf surface (Fig. 8.28). Conidia are wider at the base than the apex, are
straight to curved, have three to seven septa, and are borne in subglobular,
dark stromata that are 2060 m in diameter.
Although the fungus sporulates sparsely on articial media, it produces
copious conidia on necrotic host tissue, especially in leaf litter. Thus, Lim and
Khoo (1985) suggested that removing such debris from orchards and burning
it would assist control efforts. Several different fungicides are effective (Lim
and Khoo, 1985).
R.C. Ploetz and S. Freeman 280
8.3 Soilborne Diseases
Although soilborne diseases of mango are relatively less important than
foliar and oral diseases, they can cause signicant damage to seedlings,
nursery stock and mature trees. In general, the pathogens that are involved
are different from those that cause problems above ground. Chemical
Fig. 8.28. Conidia of Stigmina mangiferae, cause of stigmina leaf spot (Source: from
CMI description no. 097).
Foliar, Floral and Soilborne Diseases 281
management is indicated rarely for these diseases; sanitation and other
cultural measures are used most often.
Black root rot
Black root rot is reported to be an uncommon problem on young mango trees
(Lim and Khoo, 1985). Canopies of affected plants wilt suddenly and subse-
quently defoliate. Roots exhibit a water-soaked, blackened decay, and have
an unpleasant, putrid odour. Although black root rot is associated with pro-
longed ooding, its precise aetiology is not known. Several species of fungi
have been recovered from affected plants, including Fusarium solani, F. oxyspo-
rum and L. theobromae, but these were thought to be secondary colonizers of
roots (Lim and Khoo, 1985). Although mango is generally considered to be
ood intolerant, its ood tolerance is actually variable (Larson, 1991). Varia-
tion in the responses of individual trees in orchards is evident after ooding,
and when potted plants are ooded experimentally, they usually adapt by
forming hypertrophied lenticels (intumescence) (Larson et al., 1993). Plants
that do not adapt in this manner succumb fairly rapidly. Roots of the latter
plants have symptoms of black root rot (R.C. Ploetz, Homestead, Florida,
1988, personal observations). Although ood tolerance is environmentally
and biochemically complex (Larson et al., 1993), some of the fungi reported
by Lim and Khoo (1985) may interact with ood-induced stress in this host
to cause black root rot.
Nematode damage
Decline of mango trees due to nematodes has been reported in various
regions (Khan et al., 1971, 2005; McSorley et al., 1980; Anita and Chaubey,
2003). Infestations occur in areas where warm temperatures and sandy,
moist, well-drained soils predominate (Ploetz et al., 1994). Many nematode
species have been recovered from mango roots, including: Helicotylenchus
dihystera (Cobb) Sher, Quinisulcius acutus (Allen) Siddiqi, Rotylenchulus reniformis
Linford and Oliveira, Criconemella sp., Pratylenchus brachyurus (Godfrey) Fil-
ipjev and Schuurmans Stekhoven, Xiphinema sp., Meloidogyne sp., Praty-
lenchus sp. and Hoplolaimus sp. However, only Hemicriconemoides mangiferae
Siddiqi is pathogenic (Powers and McSorley, 1994). Although high popula-
tions of R. reniformis are often found on mango trees, no correlation has
been shown between their density and tree health.
Populations of H. mangiferae vary according to soil moisture and tem-
perature (Khan et al., 1971). Soil moisture < 10% and > 30%, as well as tem-
peratures < 15C and > 35C are detrimental to nematode survival and are
likely to reduce populations. In addition, tree age appears to be a signicant
factor, since H. mangiferae is found more frequently on old (> 10 years) than
young trees (< 3 years). Management is difcult and may depend on pre-
plant chemical applications plus cultural control measures (McSorley et al.,
R.C. Ploetz and S. Freeman 282
1981). Phenamiphos and 1,2-dibromo-3-chloropropane reduce levels of
H. mangiferae after planting; however, neither are registered for use in the
USA. Anita and Chaubey (2004) reported that high organic matter content in
the rhizosphere had a detrimental effect on nematode populations in the
eld.
During orchard establishment, nematode-free nursery stock should be
used. Since H. mangiferae is partially endoparasitic, it is moved easily to clean
eld sites. The use of clean planting material in infested elds should also be
avoided. If such land must be used, soil fumigation prior to planting should
be conducted. Fruit yields may still be maintained if infected trees are well
irrigated and fertilized.
Phytophthora diseases
Phytophthora palmivora (E.E. Butler) (Oomycota) causes diseases of mango in
several areas. It caused wilt, crown rot, root rot and the death of nursery trees
in Arizona USA, the Philippines and Thailand (Kueprakone et al., 1986;
Matheron and Matejka, 1988; Tsao et al., 1994). Gumming and conspicuous
bark lesions develop above ground on these plants, whereas root and crown
rots are evident at or below the ground level. Crowded conditions and exces-
sive irrigation and rainfall exacerbate these diseases. Sanitation, the use of
less-crowded conditions and reduced irrigation would be benecial.
Damage has also been recorded on the trunks of eld-grown, mature
trees in the Ivory Coast (Lourd and Keuli, 1975), and on fruit in Australia,
Malaysia and West Africa (Turner, 1960 cited in Chee, 1969; Cooke, 2007).
Mortality of trees is not observed, but substantial stem cracking and bleed-
ing does occur. The symptoms that occur on fruit have not been recorded in
Malaysia and West Africa, but on Calypso fruit in Australia, a rm chocolate-
brown decay is produced that has a sweet odour. Fruit isolates in Australia
caused leaf blight and crown canker on mango seedlings (Fig. 8.29).
Phytophthora palmivora has coenocytic hyphae, up to 7 m in diameter, papil-
late sporangia, 3156.4 20.736.7 m, which germinate either directly with
germ tubes or indirectly with motile zoospores (Fig. 8.30) (Waterhouse, 1970;
Erwin and Ribiero, 1996). Zoospores are the primary infective propagule,
and require free water for movement. Phytophthora palmivora is heterothallic.
Antheridia are amphigynous and oogonia are spherical. Chlamydospores,
c.35 m in diameter, are also formed.
Recently, a Phytophthora sp. was isolated in Spain from mango trees that
were wilted, chlorotic and had sparse canopies and cracked bark (Zea-
Bonilla et al., 2007). On V8 agar, sporangia were semi-papillate, obovoid
and 51 (2852) 36 (2237) m. Paragynous antheridia, spherical oogonia
and oospores of 28 (1932) m in diameter were produced homothalli-
cally. Ribosomal DNA sequences (ITS1, 5.8S rDNA and ITS2) (GenBank
Accession No. AM235209) were compared with those in GenBank; the
closest match, 99% identity, was with Phytophthora citricola, corroborating
the above morphological identication (Fig. 8.31). An isolate deposited in
Foliar, Floral and Soilborne Diseases 283
Fig. 8.29. Symptoms induced by Phytophthora palmivora after articial inoculation
of (a) stems and (b) foliage of mango seedlings (Photographs courtesy of A.W. Cooke).
Fig. 8.30. (a) Sporangia, (b) oogonia with amphygynous antheridia and oospores,
and (c) chlamydospore of Phytophthora palmivora (Source: from CMI description
no. 831).
(a) (b)
R.C. Ploetz and S. Freeman 284
the Spanish Type Culture Collection, CECT 20567, caused root rot on
Florida and lesions on leaves and stems of seedlings of Gomera 3.
Root rot and damping off
The oomycete, Pythium vexans de Bary, can cause root rot and wilt of seed-
lings (Lim and Khoo, 1985). In Malaysia, this pathogen caused seedling losses
of up to 30% in nurseries. Symptoms included wilting of foliage, which
initially becomes pale green, but later develops necrotic patches. Roots
develop a wet, blackened necrosis that begins in ne roots and progresses to
larger roots and the root collar. Death of seedlings often occurs. Lim and
Khoo (1985) indicated that overcrowding, excessive moisture and the use of
polybags favoured this disease.
Prakash and Singh (1980) reported that the basidiomycete Rhizoctonia
solani Kuhn (teleomorph: Thanatephorus cucumeris (Frank) Donk) caused root
and damping off of seedlings in India (Fig. 8.32). Affected tissues become
soft, dark brown or black, and seedlings may ultimately become completely
girdled and collapse. Mycelia and sclerotia of the pathogen form conspicu-
ously on affected tissues.
Sclerotium rot
This disease has been reported in Brazil (Almeida et al., 1979), India (Prakash
and Singh, 1976a) and the Philippines (Palo, 1933). The causal fungus, Sclero-
tium rolfsii Sacc. (teleomorph: Athelia rolfsii (Curzi) Tu and Kimbrough;
Fig. 8.31. (a) Semipapillate sporangia and (b) oogonia of Phytophthora citricola
(Source: from CMI description no. 114).
Foliar, Floral and Soilborne Diseases 285
synonyms: Corticium rolfsii Curzi and Pellicularia rolftii E. West), produces
globular, brown sclerotia, 1.02.6 mm in diameter. Sclerotia are resilient struc-
tures that enable the pathogen to survive adverse environmental conditions.
Symptoms begin with the formation of felty white tufts of mycelium of the
pathogen around the base of seedlings. The fungus can girdle the entire stem
to a height of 5 cm or more above the soil line. It eventually forms conspicu-
ous sclerotia in high numbers. Ultimately, seedlings wilt and die. Seed may
also rot prior to germination. The disease is controlled via sanitation and the
disinfestation of seedbeds.
Verticillium wilt
Verticillium wilt of mango was rst reported in Florida USA (Marlatt et al.,
1970). The disease was originally attributed to Verticillium albo-atrum Reinke
and Berth., but this was before Verticillium dahliae Kleb. was recognized as a
distinct species. Since the causal fungus described by Marlatt et al. (1970)
formed microsclerotia, it is clear that V. dahliae was involved (Fig. 8.33).
Symptoms of the disease include a ring and necrosis of leaves, usually
in a portion of the canopy. Sectoral development of the disease often does not
progress to other portions of the trees, which may recover. Killed leaves usu-
ally remain attached to the tree, and the xylem of affected branches is dis-
coloured brown (Fig. 8.34). Verticillium wilt is relatively uncommon, and is
(a)
(b)
(c)
20
(d)
Fig. 8.32. (a) Sclerotial cells, (b) mycelium and (c) monilioid cells of Rhizoctonia
solani, and (d) basidia and basidiospores of its teleomorph, Thanatephorus cucumeris
(Source: from CMI description no. 406).
R.C. Ploetz and S. Freeman 286
found on land where susceptible vegetable crops (i.e. potato, tomato and
aubergine) were recently grown (Pohronezny and Marlatt, 1982). New mango
orchards should not be planted on such sites.
White root disease
Rigidoporus lignosus (Klotzsch) Imazeki, the basidiomycete that causes white
root disease, is a common soil inhabitant in the humid tropics of Africa and
Asia (Holliday, 1980). It has also been reported in the western hemisphere,
but the identity of the fungus there is unclear. Rigidoporus lignosus has a wide
host range on woody perennials, including rubber, the host on which the
pathogen was rst reported (1904 in Malaysia). The signicant losses in rub-
ber plantations in the eastern hemisphere have resulted in considerable
research on this pathogen (Nandris et al., 1987).
Rigidoporus lignosus produces white rhizomorphs on the surfaces of roots
and root crowns that later darken to a yellowish and then reddish colour
(Lim and Khoo, 1985; Nandris et al., 1987). The leading edge of the rhizo-
morph is well dened and seldom appears above ground. It undergoes a
morphogenic change to produce infectious hyphae that penetrate the host
epidermis and subsequently degrade wood. Rigidoporus lignosus produces a
non-differentiated white rot that affects lignin in host cell walls.
(a)
(b)
10
(c) (d)
Fig. 8.33. (a) Verticilliate conidiophore, (b) phialospores and (c) immature and (d) mature
microsclerotia of Verticillium dahliae (Source: from CMI description no. 256).
Foliar, Floral and Soilborne Diseases 287
The fungus is most damaging on mango if orchards are established in
old rubber plantations or newly cleared jungle sites (Lim and Khoo, 1985).
Previously colonized stumps and debris from rubber and other hosts are pri-
mary sources of inoculum. Orange-yellow, bracket-like sporophores are pro-
duced during the rainy season on the root collar, trunk or exposed roots
(Fig. 8.35). Basidiospores are viable, but may play a secondary role in dis-
seminating the disease. Rhizomorphs are more signicant epidemiologically,
since they grow rapidly and can advance great distances in soil in the absence
of woody substrates.
Fig. 8.34. Vascular discoloration caused by V. dahliae (Photograph courtesy of
R.C. Ploetz).
R.C. Ploetz and S. Freeman 288
White root disease is managed by eliminating or avoiding colonized
woody debris when new orchards are established (Lim and Khoo, 1985).
Unfortunately, this is difcult and often impractical. Alternative measures
include: (i) treating planting holes with S to promote the growth of antago-
nistic microorganisms; (ii) treating stumps after clearing operations with
chemicals that discourage their colonization by basidiospores; and (iii) estab-
lishment of leguminous cover crops that promote growth of the pathogen
and the eventual exhaustion of its energy reserves.
8.4 Conclusions
As mango production continues to increase in different regions, so will the
scope and types of disease problems. Although new fungicides and bacteri-
cides will be developed in the future, it is probable that reliance on these
tools will diminish. In recent years, the regulation of pesticides has increased
while the number of new compounds that have been registered for use has
decreased. Since it appears certain that this trend will continue, the problems
posed by diseases must be solved increasingly with alternative disease
control strategies.
10 (c)
(d)
(a) (b)
20
Fig. 8.35. (a) Upper and (b) lower surface of sporophore, (c) basidiospores, and (d) hyphae of
Rigidoporus lignosus (Source: from CMI description no. 198).
Foliar, Floral and Soilborne Diseases 289
A willingness among producers to utilize new technologies will play a
role in this process. New methods to detect important pathogens have been
developed (Zheng and Ploetz, 2002), but more work is needed. Effective
detection protocols, especially for those pathogens that can colonize host tis-
sues without causing symptoms, could be used to interdict important, exotic
pathogens and identify pathogen-free propagation materials. Detection pro-
tocols could also indirectly assist disease control efforts through their use in
epidemiological studies, research on disease resistance, or to clarify portions
of disease cycles.
Acknowledgements
The authors thank Francisco Cazorla for comments on apical necrosis; Ber-
nard Slippers and Mike Wingeld for comments on the decline section; Syl-
via Fernandez, John Leslie, Christiano Lima, Kerry ODonnell and Gerardo
Rodriquez for information on malformation; Ali Obaid Al-Adawi and Mike
Wingeld for comments on seca and sudden decline; and Robert Knight for
translating Portuguese articles on seca. The following individuals are thanked
for pictures: Francisco Cazorla, T.-K. Lim, Oliver Pruvost, Carolyn Cohen,
Suzanne Bullock, Efrat Gamliel-Atinsky, Eric Palevsky and Tony Cooke.
Kevin Hyde, editor of Fungal Diversity, is thanked for permission to use
micrographs of Ceratocystis manginecans.
References
Abdel-Sattar, M.M.A. (1973) Histopathology of mango malformation. MSc. thesis, Al-
Azhar University, Cairo.
Acua, H.E. and Waite, B.H. (1977) La muerte regresiva del mango (Mangifera indica L.)
en El Salvador. Proceedings of the American Society of Horticultural Sciences, Trop-
ical Region 21, 1516.
Adaskaveg, J.E. and Hartin, R.J. (1997) Characterization of Colletotrichum acutatum
isolates causing anthracnose of almond and peach in California. Phytopathology
87, 979987.
Afanador-Kafuri, L., Minz, D., Maymon, M. and Freeman, S. (2003) Characterization of
Colletotrichum isolates from tamarillo, passiora and mango in Colombia and
identication of a unique species from the genus. Phytopathology 93, 579587.
Ah-You, N., Gagnevin, L., Chiroleu, F., Jouen, E., Rodrigues Neto, J. and Pruvost, O.
(2007a) Pathological variations within Xanthomonas campestris pv. mangiferae-
indicae support its separation into three distinct pathovars that can be
distinguished by amplied fragment length polymorphism. Phytopathology 97,
15681577.
Ah-You, N., Gagnevin, L., Pruvost, O., Myint, N.T. and Johnson, G.I. (2007b) First report
in Myanmar of Xanthomonas axonopodis pv. mangiferaeindicae causing mango
bacterial canker on Mangifera indica. Plant Disease 91, 1686.
Al Adawi, A.O., Deadman, M.L., Al Rawahi, A.K., Khan, A.J. and Al Maqbali, Y.M. (2003)
Diplodia theobromae associated with sudden decline of mango in the Sultanate of
Oman. Plant Pathology 52, 419.
R.C. Ploetz and S. Freeman 290
Al Adawi, A.O., Deadman, M.L., Al Rawahi, A.K., Al Maqbali, Y.M., Al Jahwari, A.A., Al
Saadi, B.A., Al Amri, I.S. and Wingeld, M.J. (2006) Aetiology and causal agents of
mango sudden decline disease in the Sultanate of Oman. European Journal of Plant
Pathology 116, 247254.
Al Subhi, A.M., Al Adawi, A.O., Van Wyk, M., Deadman, M.L. and Wingeld, M.J.
(2006) Ceratocystis omanensis, a new species from diseased mango trees in Oman.
Mycological Research 110, 237245.
Alahakoon, P.W., Brown, A.E. and Sreenivasaprasad, S. (1994a) Cross-infection potential
of genetic groups of Colletotrichum gloeosporioides on tropical fruits. Physiological
and Molecular Plant Pathology 44, 93103.
Alahakoon, P.W., Brown, A.E. and Sreenivasaprasad, S. (1994b) Genetic characteriza-
tion of Colletotrichum gloeosporioides isolates obtained from mango. International
Journal of Pest Management 40, 225229.
Alexopoulos, C.J., Mims, C.W. and Blackwell, M. (1996) Introductory Mycology, 4th
edn. John Wiley and Sons, New York.
Aleri, SA., Jr, Langdon, K.R., Kimbrough, J.W., El-Gholl, N.E. and Wehlburg, C. (1994)
Diseases and Disorders of Plants in Florida. Bulletin No. 14. Florida Department of
Agricultural and Consumer Services, Tallahassee, Florida.
Almeida, R.T. de, Landim, C.M.U. and Teixeira, L.M.S. (1979) Seedling blight (Sclero-
tium rolfsii Sacc.) of cashew and mango in the state of Ceara, Brazil. Fitossanidade
3, 4041.
Alvarez-Garca, L.A. and Lpez-Graca, J. (1971) Gummosis, dieback, and fruit rot
disease of mango (Mangifera indica L.) caused by Physalospora rhodina (B. and
C.) Cke. in Puerto Rico. Journal of Agriculture University of Puerto Rico 55,
435450.
Angulo, S.M. and Villapudua, J.R. (1982) Buba of mango (Mangifera indica L.) in the
state of Sinaloa, Mexico. (Abstract). Phytopathology 72, 171.
Anita and Chaubey, A.K. (2003) Inuence of soil temperature and moisture on popula-
tion dynamics of ectoparasitic nematodes infesting Mangifera indica. Annals of
Plant Protection Sciences 11, 181183.
Anita and Chaubey, A.K. (2004) Impact of organic matter on the population dynamics of
Hemicriconemoides mangiferae infesting Mangifera indica. Biological Memoirs 30,
108111.
Arrebola, E., Cazorla, F.M., Romero, D., Prez-Garca, A. and de Vicente, A. (2007) A
nonribosomal peptide synthetase gene (mgoA) of Pseudomonas syringae pv. syrin-
gae is involved in mangotoxin biosynthesis and is required for full virulence. Mo-
lecular Plant-Microbe Interactions 20, 500509.
Atkinson, T.H. and Peck, S.B. (1994) Annotated checklist of the bark and ambrosia bee-
tles (Coleoptera: Platypodidae and Scolytidae) of tropical Southern Florida. The
Florida Entomologist 77, 313329.
Bastawros, M.K. (1996) Mango malformation in Egypt. Acta Horticulturae 455, 566574.
Batista, A.C. (1947) Mal do recife (grave doenca da manggueira). Unpublished thesis,
Pernambuco College of Agriculture, Pernambuco, Brazil (Abstract in Review of Ap-
plied Mycology (1948) 27, 7778).
Batzer, J.C., Gleason, M.L., Weldon, B., Dixon, P.M. and Nutter, F.W., Jr (2002) Evalua-
tion of postharvest removal of sooty blotch and yspeck on apples using sodium
hypochlorite, hydrogen peroxide with peroxyacetic acid, and soap. Plant Disease
86, 13251332.
Batzer, J.C., Gleason, M.L., Harrington, T.C. and Tiffany, L.H. (2005) Expansion of the
sooty blotch and yspeck complex on apples based on analysis of ribosomal DNA
gene sequences and morphology. Mycologia 97, 12681286.
Foliar, Floral and Soilborne Diseases 291
Berthet, J.A. (1914) Molestia cia mangueira. Bolm Agriculturae (So Paolo) 15,
818819.
Bhatnagar, S.S. and Beniwal, S.P.S. (1977) Involvement of Fusarium oxysporum in causa-
tion of mango malformation. Plant Disease Reporter 61, 894898.
Bitancourt, A.A. and Jenkins, A.E. (1943) Scab of mango caused by Elsinoe. (Abstract).
Phytopathology 33, 1.
Boesewinkel, H.J. (1980) The identity of mango mildew, Oidium mangiferae. Phyto-
pathologische Zeitschrift 99, 126130.
Bose, S.K., Sindhan, G.S. and Pandey, B.H. (1973) Studies on dieback disease of mango
in the Tarai region of Kuaon. Progressive Horticulture 70, 557584.
Britz, H., Steenkamp, E.T., Coutinho, T.A., Wingeld, B.D., Marasas, W.F.O. and Wing-
eld, M.J. (2002) Two new species of Fusarium section Liseola associated with man-
go malformation. Mycologia 94, 722730.
Burgess, T.I., Barber, P.A., Mohali, S., Pegg, G., de Beer, W. and Wingeld, M.J. (2006)
Three new Lasiodiplodia spp. from the tropics, recognized based on DNA sequence
comparisons and morphology. Mycologia 98, 423435.
Butani, D.K. (1993) Mango: Pest Problems. Periodical Expert Book. Agency D-42, Vivek
Vihar, Delhi, India.
Butler, E.J. and Bisby, G.R. (1931) The Fungi of India. Scientic Monograph No.1. The
Imperial Council of Agricultural Research in India, Calcutta.
Carvalho, C.R.L., Rossetto, C.J., Mantovani, D.M.B., Morgano, M.A., de Castro, J.V. and
Bortoletto, N. (2004) Avalia de cultivares de mangueira selectionadas pelo Insiti-
tuto Agrontmico de Campinas comparadas a outras de importncia comercial. Re-
vista Brasil Fruticultura 26, 264271.
Cazorla, F.M., Tors, J.A., Olalla, L., Prez-Garca, A., Farr, J.M. and de Vicente, A.
(1998) Bacterial apical necrosis of mango in southern Spain: a disease caused by
Pseudomonas syringae pv. syringae. Phytopathology 88, 614620.
Cazorla, F.M., Arrebola, E., Sesma, A., Prez-Garca, A., Codina, J.C., Murillo, J. and de
Vicente, A. (2002) Copper resistance in Pseudomonas syringae strains isolated from
mango is encoded mainly by plasmids. Phytopathology 92, 909916.
Cazorla, F.M., Arrebola, E., Olea, F., Velasco, L., Hermoso, J.M., Prez-Garca, A., Tors,
J.A., Farr, J.M. and deVicente, A. (2006) Field evaluation of treatments for the con-
trol of the bacterial apical necrosis of mango (Mangifera indica) caused by Pseudomo-
nas syringae pv. syringae. European Journal of Plant Pathology 116, 279288.
Chakrabarti, D.K. and Ghosal, S. (1989) The disease cycle of mango malformation in-
duced by Fusarium moniliforme var. subglutinans and the curative effects of
mangiferin metal chelates. Journal of Phytopathology 125, 238246.
Chee, K.H. (1969) Hosts of Phytophthora palmivora. Review of Applied Mycology 48,
337344.
Colosimo, E.A., Chalita, L.V.A.S. and Demetrio, C.G.B. (2000) Tests of proportional
hazards and proportional odds models for grouped survival data. Biometrics 56,
12331240.
Cook, A.A. (1975) Diseases of Tropical and Subtropical Fruits and Nuts. Hafner Press,
New York.
Cook, A.A., Milbrath, G.M. and Hamilton, R.A. (1971) Woody gall and scaly bark of
Mangifera indica in Hawaii. (Abstract). Phytopathology 61, 1320.
Cooke, A.W. (2007) Phytophthora postharvest fruit rot in mango. Advisory poster pro-
duced by the Queensland Government Department of Primary Industries and Fish-
eries, Indooropilly, Queensland, Australia.
Cronje, C., Wehnwe, F.C. and Kotz, J.M. (1990) Alternaria alternata as a lesion patho-
gen of mango inorescences in South Africa. Phytophylactica 22, 117118.
R.C. Ploetz and S. Freeman 292
Crookes, C.A. and Rijkenberg, F.H.J. (1985) A literature review of the distribution, symp-
tomatology, cause and control of mango blossom malformation. South African
Mango Growers Association Yearbook 5, 1524.
Crous, P.W. and Palm, M.E. (1999) Reassessment of the anamorph genera of Botryoplo-
dia, Dothiorella and Fusicoccum. Sydowia 52, 167175.
Crous, P.W., Slippers, B., Wingeld, M.J., Rheeder, J., Marasas, W.F.O., Philips, A.J.L.,
Alves, A., Burgess, T., Barber, P. and Groenewald, J.Z. (2006) Phylogenetic lineages
in the Botryosphaeriaceae. Studies in Mycology 55, 235253.
Dag, A., Eisenstein, D. and Gazit, S. (2001) Effects of three fungicides used to control
powdery mildew in mango on pollen germination and pollen-tube growth. Pro-
ceedings of the InterAmerican Society for Tropical Horticulture 43, 123126.
Dang, J.K. and Daulta, B.S. (1982) Mango malformation a review. Pesticides 16, 511.
Darvas, J.M. (1987) Control of mango blossom malformation with trunk injection. South
African Mango Growers Association Yearbook 7, 2124.
Darvas, J.M. (1993) Dothiorella dominicana, an important mango pathogen in South
Africa. Acta Horticulturae 341, 321328.
Das Gupta, S.N. and Zacchariah, H.T. (1945) Studies on the diseases of Mangifera in-
dica. Part V. On the die-back of mango trees. Journal of the Indian Botanical Society
24, 101108. (Abstract in Review of Applied Mycology (1946) 25, 267268).
Denman, S., Crous, P.W., Taylor, J.E., Kang, J.-C., Pascoe, I. and Wingeld, M.J. (2000)
An overview of the taxonomic history of Botryosphaeria, and a re-evaluation of its
anamorphs based on morphology and ITS and rDNA phylogeny. Studies in Mycol-
ogy 45, 129140.
Diekman, F., Manicom, B.Q. and Coetzee, K. (1982) An attempt to control blossom
malformation of mangoes with chemical sprays. Subtropica 3, 1516.
Dodd, J.C., Estrada, A.B., Matcham, J., Jefferies, P. and Jeger, M.J. (1991) The effect of
climatic factors on Colletotrichum gloeosporioides, causal agent of mango anthra-
cnose, in the Philippines. Plant Pathology 40, 568575.
Domsch, K.H., Gams, W. and Anderson, T.-H. (1980) Compendium of Soil Fungi. Aca-
demic Press, London.
Droby, S., Prusky, D., Jacoby, B. and Goldman, A. (1986) Presence of antifungal com-
pounds in the peel of mango fruits and their relation to latent infections by Alter-
naria alternata. Physiological and Molecular Plant Pathology 29, 173183.
Droby, S., Prusky, D., Jacoby, B. and Goldman, A. (1987) Induction of antifungal resor-
cinols in esh of unripe mango fruits and its relation to latent infection by Alter-
naria alternata. Physiological and Molecular Plant Pathology 30, 285292.
Du, M., Schardl, C.L., Nuckels, E.M. and Vaillancourt, L.J. (2005) Using mating-type
gene sequences for improved phylogenetic resolution of Collectotrichum species
complexes. Mycologia 97, 641658.
Ellis, M.B. (1971) Dematiaceous Hypomycetes. Commonwealth Mycological Institute,
Kew, Surrey, UK, 608 pp.
Engelbrecht, C.J. and Harrington, T.C. (2005) Intersterility, morphology and taxonomy of
Ceratocystis mbriata on sweet potato, cacao and sycamore. Mycologia 97, 5769.
Erwin, D.C. and Ribiero, O.K. (1996) Phytophthora Diseases Worldwide. American Phy-
topathological Society (APS) Press, St Paul, Minnesota.
Eshel, D., Miyara, I., Ailinng, T., Dinoor, A. and Prusky, D. (2002) pH regulates endoglu-
canase expression and virulence of Alternaria alternata in persimmon fruits. Mo-
lecular Plant-Microbe Interactions 15, 774779.
Farr, D.F., Bills, G.F., Chamuris, G.P. and Rossman, A.Y. (1989) Fungi on Plant and Plant
Products in the United States. American Phytopathological Society (APS) Press, St
Paul, Minnesota.
Foliar, Floral and Soilborne Diseases 293
Fitzell, R.D. (1979) Colletotrichum acutatum as a cause of anthracnose of mango in
New South Wales. Plant Disease Reporter 63, 10671070.
Fitzell, R.D. and Peak, C.M. (1984) The epidemiology of anthracnose disease of mango:
inoculum sources, spore production and dispersal. Annals of Applied Biology 104,
5359.
Fitzell, R.D., Peak, C.M. and Darnell, R.E. (1984) A model for estimating infection levels
of anthracnose disease of mango. Annals of Applied Biology 104, 451458.
Flechtmann, C.H.W., Kimati, H., Medcalf, J.C. and Ferr, J. (1973) Preliminary observa-
tions on malformation in mango (Mangifera indica L.) inorescences and the fun-
gus, insects and mite found on them. Review of Plant Pathology 52, 162163.
Freeman, S. and Rodriguez, R.J. (1995) Differentiation of Colletotrichum species respon-
sible for anthracnose of strawberry by arbitrarily primed PCR. Mycological Research
99, 501504.
Freeman, S., Pham, M. and Rodriguez, R.J. (1993) Molecular genotyping of Colletotri-
chum species based on arbitrarily primed PCR, A+T-rich DNA, and nuclear DNA
analyses. Experimental Mycology 17, 309322.
Freeman, S., Katan, T. and Shabi, E. (1998) Characterization of Colletotrichum species
responsible for anthracnose diseases of various fruits. Plant Disease 82, 596605.
Freeman, S., Maimon, M. and Pinkas, Y. (1999) Use of GUS transformants of Fusarium
subglutinans for determining etiology of mango malformation disease. Phytopathol-
ogy 89, 456461.
Freeman, S., Minz, D., Maymon, M. and Zveibil, A. (2001) Genetic diversity within Col-
letotrichum acutatum sensu Simmonds. Phytopathology 91, 586592.
Fukuda, T., Uehara, K., Azegami, K., Tabei, H. and Nishiyama, K. (1990) Bacterial can-
ker of mango in Japan caused by Xanthomonas campestris pv. mangiferaeindicae.
Annals of the Phytopathological Society of Japan 56, 474480.
Gagnevin, L. and Pruvost, O. (1995) Assessment of genomic variability in Xanthomonas
campestris pv. mangiferaeindicae. (Abstract). Phytopathology 85, 1163.
Gagnevin, L. and Pruvost, O. (2001) Epidemiology and control of mango bacterial black
spot. Plant Disease 85, 928935.
Gamliel-Atinsky, E., Sztejnberg, A., Maymon, M., Belausov, E., Palevsky, E. and Free-
man, S. (2007) Interaction of the mango bud mite (Aceria mangiferae) with Fusari-
um mangiferae, causal agent of mango malformation disease. Phytoparasitica 35,
198199.
Gantotti, B.V. and Davis, M.J. (1993) Pectic zymogram analysis for characterizing
genetic diversity of the mango anthracnose pathogen. Acta Horticulturae 341,
353359.
Gupta, J.H. (1989) Perpetuation and epidemiology of powdery mildew of mango. Acta
Horticulturae 231, 528533.
Halsted, B.D. and Fairchild, D.G. (1891) Sweet-potato black rot. Journal of Mycology 7,
111.
Hamid, M. and Jalaluddin, M. (2006) Recurring incidence of sooty mould of mango
in Karachi and its control. International Journal of Biology and Biotechnology 3,
561565.
Hayden, H.L., Pegg, K.G., Aitken, E.A.B. and Irwin, J.A.G. (1994) Genetic relationships
as assessed by molecular markers and cross-infection among strains of Colletotri-
chum gloeosporioides. Australian Journal of Botany 42, 918.
Hingorani, M.K., Sharma, O.P. and Sohi, H.S. (1960) Studies on blight disease of mango
caused by Macrophoma mangiferae. Indian Phytopathology 13, 137143.
Hirano, S.S. and Upper, C.D. (1983) Ecology and epidemiology of foliar bacterial plant
pathogens. Annual Review of Phytopathology 21, 243269.
R.C. Ploetz and S. Freeman 294
Hodson, A., Mill, P.R. and Brown, A.E. (1993) Ribosomal and mitochondrial DNA poly-
morphisms in Colletotrichum gloeosporioides isolated from tropical fruits. Myco-
logical Research 97, 329335.
Holliday, P. (1980) Fungus Diseases of Tropical Crops. Cambridge University Press,
Cambridge.
Hughes, S.J. (1980) New Zealand Fungi 29. Rhinocladium Sacc. et March. New Zealand
Journal of Botany 18, 163172.
Iqbal, Z., Mehboob-ur-Rahman, Dasti, A.A., Saleem, A. and Zafar, Y. (2006) RAPD anal-
ysis of Fusarium isolates causing Mango Malformation disease in Pakistan. World
Journal of Microbiology and Biotechnology 22, 11611167.
Jeffries, P., Dodd, J.C., Jeger, M.J. and Plumbley, R.A. (1990) The biology and control of
Colletotrichum species on tropical fruit crops. Plant Pathology 39, 343366.
Johnson, E.M., Sutton, T.B. and Hodges, C.S. (1997) Etiology of apple sooty blotch dis-
ease in North Carolina. Phytopathology 87, 8895.
Johnson, G.I. (1992) Biology and control of stem-end rot pathogens of mango. PhD
thesis, University of Queensland, Brisbane, Australia.
Johnson, G.I. (1994a) Powdery mildew. In: Ploetz, R.C., Zentmyer, G.A., Nishijima, W.,
Rohrbach, K. and Ohr, H.D. (eds) Compendium of Tropical Fruit Diseases. American
Phytopathological Society (APS) Press, St Paul, Minnesota, pp. 3839.
Johnson, G.I. (1994b) Stem-end rot. In: Ploetz, R.C., Zentmyer, G.A., Nishijima, W.,
Rohrbach, K. and Ohr, R.D. (eds) Compendium of Tropical Fruit Diseases. American
Phytopathological Society (APS) Press, St Paul, Minnesota, pp. 3941.
Johnson, G.I., Cooke, A.W., Mead, A.J. and Wells, L.A. (1991) Stem end rot of mango in
Australia: causes and control. Acta Horticulturae 219, 288295.
Johnson, G.I., Mead, A.J., Cooke, A.W. and Dean, J.R. (1992) Mango stem end rot patho-
gens fruit infection by endophytic colonisation of the inorescence and pedicel.
Annals of Applied Biology 120, 225234.
Johnson, J.A., Harrington, T.C. and Engelbrecht, C.J.B. (2005) Phylogeny and taxonomy
of the North American clade of the Ceratocystis mbriata complex. Mycologia 97,
10671092.
Joubert, J.J. and Rijkenberg, F.H.J. (1971) Parasitic green algae. Annual Review of Phyto-
pathology 9, 4564.
Junqueira, N.T.V., Pinto, A.C. de Q., da Cunha, M.M. and Ramos, V.H.V. (2002) Controle
das doenas da mangueira. In: Zambolin, L., do Vale, F.X.R., Monteiro, A.J.A. and
Costa, H. (eds) Controle de Doenas de Plantas Fruteiras. Vol. 1. Universidade Fed-
eral de Viosa, Viosa, Brazil, pp. 323403.
Kadman, A. and Gazit, S. (1984) The problem of iron deciency in mango trees and
experiments to cure it in Israel. Journal of Plant Nutrition 7, 282290.
Kauser, A.G. (1959) Malformation of inorescence in mango. Punjab Fruit Journal 22,
1921.
Kennelly, M.M., Cazorla, F.M., de Vicente, A., Ramos, C. and Sundin, G.W. (2007)
Pseudomonas syringae diseases of fruit trees: progress toward understanding and
control. Plant Disease 91, 417.
Khan, A.M., Adhami, A. and Saxena, S.K. (1971) Population changes of some stylet-
bearing nematodes associated with mango (Mangifera indica L.). Indian Journal of
Nematology 1, 99105.
Khan, A., Sayed, M. and Shaukat, S.S. (2005) Nematodes associated with mango in
Sindh. International Journal of Biology and Biotechnology 2, 917919.
Kile, G.A. (1993) Plant diseases caused by species of Ceratocystis sensu stricto and
Chalara. In: Wingeld, M.J., Seifert, K.A. and Webber, J.F. (eds) Ceratocystis and
Foliar, Floral and Soilborne Diseases 295
Ophiostoma: Taxonomy, Ecology and Pathogenicity. American Phytopathological
Society (APS) Press, St Paul, Minnesota, pp. 173183.
Kishun, R. (1994) Evaluation of phylloplane micro-organisms from mango against Xan-
thomonas campestris pv. mangiferaeindicae. Indian Phytopathology 47, 313.
Kishun, R. (1995) Detection and management of Xanthomonas campestris pv.
mangiferaeindicae. In: Varma, J.P., Verma, A. and Kumar, D. (eds) Detection of
Plant Pathogens and their Management. Angkar Publishers, New Delhi, India,
pp. 173182.
Kishun, R. and Sohi, H.S. (1983) Bacterial canker in mangoes. Indian Farmers Digest
14, 2123.
Kueprakone, U., Saengkong, S., Pienpuck, K. and Choobumroong, W. (1986) Phytophtho-
ra palmivora (Butl.) Butl., the causal organism of black rot of mango seedlings. Thai
Agricultural Research Journal 4, 6773. (in Thai with an English abstract)
Kumar, J. and Beniwal, S.P.S. (1991) Mango malformation. In: Kumar, J., Chaube, H.S.,
Singh, U.S. and Mukhopadhyay, A.N. (eds) Plant Diseases of International Impor-
tance. Vol. III: Diseases of Fruit Crops. Prentice Hall, Englewood Cliffs, New Jersey,
pp. 357393.
Kvas, M., Steenkamp, E.T., Al Adawi, A.O., Deadman, M.L., Al Jahwari, A.A., Marasas,
W.F.O., Wingeld, B.D., Ploetz, R.C. and Wingeld, M.J. (2007) Fusarium mangifer-
ae associated with mango malformation in the Sultanate of Oman. European Jour-
nal of Plant Pathology 121, 195199.
Larson, K.D. (1991) Physiological, anatomical and growth responses of mango (Mangifera
indica L.) trees to ooding. PhD thesis, University of Florida, Gainesville, Florida.
Larson, K.D., Schaffer, B. and Davies, F.S. (1993) Floodwater oxygen content, ethylene
production and lenticel hypertrophy in ooded mango (Mangifera indica L.) trees.
Journal of Experimental Botany 44, 665671.
Leslie, J.F. (1995) Gibberella fujikuroi: available populations and variable traits. Cana-
dian Journal of Botany 73, S282S291.
Leslie, J.F. and Summerell, B.A. (2006) The Fusarium Lab Manual. Blackwell, Ames, Iowa.
Lim, T.K. (1994) Pink disease. In: Ploetz, R.C., Zentmyer, G.A., Nishijima, W.T., Rohr-
bach, K.G. and Ohr, H.D. (eds) Compendium of Tropical Fruit Diseases. American
Phytopathological Society (APS) Press, St Paul, Minnesota, pp. 3738.
Lim, T.K. and Khoo, K.C. (1985) Diseases and Disorders of Mango in Malaysia. Tropical
Press, Kuala Lumpur, Malaysia.
Lima, C.S., Costa, S.S., Campos, M.A. and Pfenning, L.H. (2006a) A new Fusarium spe-
cies associated with mango malformation in Brazil. In: XXXIX Annual Meeting of
the Brazilian Phytopathological Society, 2006, Salvador. Fitopatologia Brasileira
(S)31, 191.
Lima, C.S., Pfenning, L.H., Costa, S.S., Campos, M.A. and Leslie, J.F. (2006b) A new
biological species of the Gibberella fujikuroi complex associated with mango mal-
formation in Brazil. In: XXXIX Annual Meeting of the Brazilian Phytopathological
Society, 2006, Salvador. Fitopatologia Brasileira (S) 31, 191.
Litz, R.E. (ed.) (1997) The Mango: Botany, Production and Uses. CAB International,
Wallingford, UK.
Lonsdale, J.H. and Kotz, J.M. (1993) Chemical control of mango blossom diseases and
the effect on fruit set and yield. Plant Disease 77, 558562.
Lourd, M. and Keuli, S.D. (1975) Note sur un chancre a Phytophthora du manguier en
Cote dlvoire. Fruits 30, 541544.
Majumder, P.K. and Sharma, D.K. (1990) Mango. In: Bose, T.K. and Mitra, S.K. (eds)
Fruits: Tropical and Subtropical. Naya Prokash, Calcutta, pp. 4062.
R.C. Ploetz and S. Freeman 296
Malaguti, G. and de Reyes, C. (1964) A gall disease of cacao and mango in Venezuela
caused by Calonectria rigidiuscula. (Abstract). Phytopathology 54, 499.
Malik, M.T., Khan, S.M., Dasti, A.A. and Kazmi, M.R. (2005) First record of Ceratocystis
pimbriata (sic) causal organism of mango sudden death in Pakistan. Pakistan Jour-
nal of Phytopathology 17, 187191.
Manicom, B.Q. (1986) Factors affecting bacterial black spot of mangoes caused by
Xanthomonas campestris pv. mangiferaeindicae. Annals of Applied Biology 109,
129135.
Manicom, B.Q. (1989) Blossom malformation of mango. South African Mango Growers
Association Yearbook 10, 1112.
Manicom, B.Q. and Pruvost, O.P. (1994) Bacterial black spot. In: Ploetz, R.C., Zentmyer,
G.A., Nishijima, W.T., Rohrbach, K.G. and Ohr, H.D. (eds) Compendium of Tropical
Fruit Diseases. American Phytopathological Society (APS) Press, St Paul, Minnesota,
pp. 4142.
Manicom, B.Q. and Wallis, F.M. (1984) Further characterization of Xanthomonas camp-
estris pv. mangiferaeindicae. International Journal of Systematic Bacteriology 34,
7779.
Marasas, W.F.O., Ploetz, R.C., Wingeld, M.J., Wingeld, B.D. and Steenkamp, E.T.
(2006) Mango malformation disease and the associated Fusarium species. Phytopa-
thology 96, 667672.
Marlatt, R.B., Knight, R.J., Jr and Goldweber, S. (1970) Verticillium wilt of mango
(Mangifera indica) in Florida. Plant Disease Reporter 54, 569571.
Matheron, M.E. and Matejka, J.C. (1988) Phytophthora crown and root rot on nursery-
grown mango trees delivered to Arizona. (Abstract). Phytopathology 78, 1572.
Maymon, M., Zveibil, A., Pivonia, S., Minz, D. and Freeman, S. (2006) Identication and
characterization of benomyl-resistant and -sensitive eld isolates of Colletotrichum
gloeosporioides from statice (Limonium spp.). Phytopathology 96, 542548.
McSorley, R., Parrado, J.L. and Goldweber, S. (1980) Observations on a mango decline in
South Florida. Proceedings of the Florida State Horticultural Society 93, 132133.
McSorley, R., Parrado, J.L. and Goldweber, S. (1981) Plant-parasitic nematodes associ-
ated with mango and relationship to tree condition. Nematropica 11, 19.
Michailides, T.J., Morgan, D.P. and Felts, D. (1999) Other hosts of Botryosphaeria
dothidea and their relation to Botryosphaeria pistachio blight. California Pistachio
Production Reports. California Pistachio Commission, Fresno, California, USA.
Misra, A.K. and Prakash, O. (1992) Control of bacterial canker of mango by chemicals.
Pesticides 18, 3233.
Mukherjee, S.K. (1997) Introduction: botany and importance. In: Litz, R.E. (ed.) The
Mango: Botany, Production and Uses. CAB International, Wallingford, UK,
pp. 119.
Mller, E.M., Chelkowski, J. and Geiger, H.H. (1999) Species-specic PCR assays for the
fungal pathogens Fusarium moniliforme and Fusarium subglutinans and their
application to diagnose maize ear rot disease. Journal of Phytopathology 147,
497508.
Muller, H.R.A. (1940) Overzicht van de belangrijkste mangga-ziekten in Nederlandsch
Indie. (English abstract in Review of Applied Mycology (1940) 19, 355.)
Nandris, D., Nicole, M. and Geiger, J.P. (1987) Root rot diseases of rubber trees. Plant
Disease 71, 298306.
Narasimhan, M.J. (1954) Malformation of panicles in mango incited by spp. of Erio-
phyes. Current Science 23, 297298.
Narasimhan, M.J. (1959) Control of mango malformation disease. Current Science 28,
254255.
Foliar, Floral and Soilborne Diseases 297
Nariani, T.K. and Seth, M.L. (1962) Role of eriophyid mites in causing malformation
disease of mango. Indian Phytopathology 15, 231234.
Neergaard, P. (1945) Danish Species of Alternaria and Stemphylium. Taxonomy, Parasiti-
cism, Economical Signicance. Einar Munksgaard, Copenhagen, Denmark.
Nicholson, R.I.D. and van Staden, J. (1988) Cytokinins and mango ower malformation.
I. Tentative identication of the complement in healthy and malformed inores-
cences. Journal of Plant Physiology 132, 720724.
Nofal, M.A. and Haggag, W.M. (2006) Integrated management of powdery mildew of
mango in Egypt. Crop Protection 25, 480486.
ODonnell, K., Cigelnik, E. and Nirenberg, H.I. (1998) Molecular systematics and phy-
logeography of the Gibberella fujikuroi species complex. Mycologia 90, 465493.
ODonnell, K., Nirenberg, H.I., Aoki, T. and Cigelnik, E. (2000) A multigene phylogeny
of the Gibberella fujikuroi species complex: detection of additional phylogeneti-
cally distinct species. Mycoscience 41, 6178.
Okigbo, R.N. (2001) Occurrence, pathogenicity and survival of Macrophoma mangifer-
ae in leaves, branches and stems of mango (Mangifera indica L.). Plant Protection
Science 37, 138144.
Okigbo, R.N. and Osuinde, M.I. (2003) Fungal leaf spot diseases of mango (Mangifera
indica L.) in southeastern Nigeria and biological control with Bacillus subtilis. Plant
Protection Science 39, 7077.
Oosthuyse, S.A. (1998) Cost reduction of powdery mildew control in mango with mono
potassium phosphate. South African Mango Growers Association Yearbook 18,
4042.
Palmateer, A.J., Ploetz, R.C., van Santer, E. and Correll, J.C. (2007) First occurrence of anthra-
cnose caused by Colletotrichum gloeosporioides on pitahaya. Plant Disease 91, 631.
Palo, M.A. (1933) Sclerotium seed rot and seedling rot of mango. Philippine Journal of
Science 52, 237261.
Palti, J., Pinkas, Y. and Chorin, M. (1974) Powdery mildew of mango. Plant Disease Re-
porter 58, 4549.
Patel, M.K., Kulkarni, Y.S. and Moniz, L. (1948) Pseudomonas mangiferae-indicae,
pathogenic on mango. Indian Phytopathology 1, 147152.
Peres, N.A.R., Souza, N.L., Zitko, S.E. and Timmer, L.W. (2002) Activity of benomyl for
control of postbloom fruit drop of citrus caused by Colletotrichum acutatum. Plant
Disease 86, 620624.
Peterson, R.A., Johnson, G.I., Schipke, L.G. and Cooke, A.W. (1991) Chemical control
of stem end rot. Acta Horticulturae 291, 304307.
Piza, C. de T. (ed.) (1966) Anais do simposio sobre a seca da mangueira. (Abstract in
Review of Applied Mycology (1967) 46, 378.)
Ploetz, R.C. (1994) Distribution and prevalence of Fusarium subglutinans in mango trees
affected by malformation. Canadian Journal of Botany 72, 79.
Ploetz, R.C. (2001) Malformation: a unique and important disease of mango, Mangifera
indica L. In: Summerell, B.A., Leslie, J.F., Backhouse, D. and Bryden, W.L. (eds)
Fusarium: Paul E. Nelson Memorial Symposium. American Phytopathological Soci-
ety (APS) Press, St Paul, Minnesota, pp. 233247.
Ploetz, R.C. (2003) Diseases of mango. In: Ploetz, R.C. (ed.) Diseases of Tropical Fruit
Crops. CAB International, Wallingford, UK, pp. 327363.
Ploetz, R.C. (2007) Cacao diseases: important threats to chocolate production world-
wide. Phytopathology 97, 16341639.
Ploetz, R.C. and Prakash, O. (1997) Foliar, oral and soilborne diseases of mango. In:
Litz, R.E. (ed.) The Mango: Botany, Production and Uses. CAB International, Wall-
ingford, UK, pp. 281325.
R.C. Ploetz and S. Freeman 298
Ploetz, R.C., Zentmyer, G.A., Nishijima, W., Rohrbach, K. and Ohr, H.D. (eds) (1994)
Compendium of Tropical Fruit Diseases. American Phytopathological Society (APS)
Press, St Paul, Minnesota.
Ploetz, R.C., Benscher, D., Vasquez, A., Colls, A., Nagel, J. and Schaffer, B. (1996a) A
re-evaluation of mango decline in Florida. Plant Disease 80, 664668.
Ploetz, R., Vazquez, A. and Benscher, D. (1996b) First report of Fusarium decemcellulare
as a pathogen of mango in the United States. Plant Disease 80, 1207.
Ploetz, R.C., Benscher, D., Dorey, A.J. and Vazquez, A. (2000) The epidemiology, control
and cause of sooty blotch of carambola, Averrhoa carambola L., in South Florida.
Fruits 55, 241252.
Ploetz, R., Zheng, Q.I., Vazquez, A. and Abdel Sattar, M.A. (2002) Current status and
impact of mango malformation in Egypt. International Journal of Pest Management
48, 279285.
Pohronezney, K. and Marlatt, R.B. (1982) Some Common Diseases of Mango in Florida.
Plant Pathology Fact Sheet PP-23. University of Florida, Gainesville, Florida.
Pordesimo, A.N. (1982) Development of programmed spray application for the control
of anthracnose. In: PCARRD Research Highlights 1982. Philippine Council for Ag-
riculture, Forestry and Natural Resources Research and Development (PCARRD),
Los Baos, the Philippines, pp. 3839.
Powers, L.E. and McSorley, R. (1994) Nematodes. In: Ploetz, R.C., Zentmyer, G.A.,
Nishijima, W., Rohrbach, K. and Ohr, H.D. (eds) Compendium of Tropical Fruit
Diseases. American Phytopathological Society (APS) Press, St Paul, Minnesota, pp.
4243.
Prakash, O. (1988) Sooty mould disease of mango and its control. International Journal
of Tropical Plant Disease 9, 277280.
Prakash, O. (1990) Report of the plant pathologist. Annual Report of CIHNP (CMRS),
Lucknow for 19891990.
Prakash, O. (1991) Sooty mould disease of mango and its control. International Journal
of Tropical Plant Diseases 9, 277280.
Prakash, O. and Raoof, M.A. (1989) Die back disease of mango (Mangifera indica),
its distribution, incidence, cause and management. Fitopatologia Brasiliensis 14,
207215.
Prakash, O. and Singh, U.N. (1976a) Basal rot of mango seedling caused by Sclerotium
rolfsii. Indian Journal of Mycology and Plant Pathology 6, 75.
Prakash, O. and Singh, U.N. (1976b) New disease of mango. In: Proceedings of AICFRP
Workshop, 2428 May, Hyderabad, India.
Prakash, O. and Singh, U.N. (1977) Phoma blight, a new disease of mango. Plant Dis-
ease Reporter 61, 419421.
Prakash, O. and Singh, U.N. (1980) Root rot and damping off of mango seedlings caused
by Rhizoctonia solani. Indian Journal of Mycology and Plant Pathology 10, 69.
Prakash, O. and Srivastava, K.C. (1987) Mango Diseases and their Management a
World Review. Tommorows Printer, New Delhi, India.
Prakash, O., Misra, A.K. and Raoof, M.A. (1994) Studies on mango bacterial canker
disease. Biological Memoirs 20, 95107.
Prasad, A., Singh, H. and Shukla, T.N. (1965) Present status of mango malformation
disease. Indian Journal of Horticulture 22, 254265.
Prusky, D. (1994) Alternaria rot (black spot). In: Ploetz, R.C., Zentmyer, G.A., Nishijima,
W., Rohrbach, K. and Ohr, H.D. (eds) Compendium of Tropical Fruit Diseases.
American Phytopathological Society (APS) Press, St Paul, Minnesota, pp. 3435.
Prusky, D., Fuch, Y. and Yanko, U. (1983) Assessment of latent infections as a basis for
control of post harvest disease of mango. Plant Disease 67, 816818.
Foliar, Floral and Soilborne Diseases 299
Prusky, D., Shalom, Y., Kobiler, I., Akerman, M. and Fuchs, Y. (2002) The level of quies-
cent infection of Alternaria alternata in mango fruits at harvest determines the post-
harvest treatment applied for the control of rots during storage. Postharvest Biology
and Technology 25, 339347.
Prusky, D., Kobiler, I., Akerman, M. and Miyara, I. (2006) Effect of acidic solutions and
acidic prochloraz on the control of postharvest decay caused by Alternaria alternata
in mango and persimmon fruit. Postharvest Biology and Technology 42, 134141.
Pruvost, O. and Luisetti, J. (1991a) Attempts to develop a biological control of bacterial
black spot of mangoes. Acta Horticulturae 291, 324337.
Pruvost, O. and Luisetti, J. (1991b) Effect of time of inoculation with Xanthomonas
campestris pv. mangiferaeindicae on mango fruits susceptibility. Epiphytic survival
of X. c. pv. mangiferaeindicae on mango fruits in relation to disease development.
Journal of Phytopathology 133, 139151.
Pruvost, O., Couteau, A. and Luisetti, J. (1989) Efcacit de diffrentes formulations
chemiques pour Jutter contre la maladie des taches noires de la mangue (Xan-
thomonas campestris pv. mangiferaeindicae). Fruits 44, 343350.
Pruvost, O., Couteau, A. and Luisetti, J. (1990) Development of bacterial black spot of
mangoes and epiphytic populations of the pathogen (Xanthomonas campestris pv.
mangiferaeindicae) under natural conditions in Reunion Island. Fruits 45, 125140.
Pruvost, O., Couteau, A. and Luisetti, J. (1992) Pepper tree (Schinus terebenthifolius
Radii), a new host plant for Xanthomonas campestris pv. mangiferaeindicae. Journal
of Phytopathology 135, 289298.
Pruvost, O., Couteau, A., Verniere, C. and Luisetti, J. (1993) Epiphytic survival of Xan-
thomonas campestris pv. mangiferaeindicae on mango buds. Acta Horticulturae
341, 337344.
Pruvost, O., Roumagnac, P., Gaube, C., Chiroleu, F. and Gagnevin, L. (2005) New me-
dia for the semiselective isolation and enumeration of Xanthomonas campestris pv.
mangiferaeindicae, the causal agent of mango bacterial black spot. Journal of Ap-
plied Microbiology 99, 803815.
Punithalingam, E. (1976) Botryodiplodia theobromae. Commonwealth Mycological In-
stitute (CMI) Descriptions of Pathogenic Fungi and Bacteria No. 519. CMI, Kew,
Surrey, UK.
Ramos, L.J., Lara, S.P., McMillan, R.T. and Narayanan, K.R. (1991) Tip dieback of mango
(Mangifera indica) caused by Botyrosphaeria ribis. Plant Disease 75, 315318.
Rao, B.S. (1975) Maladies of Hevea in Malaysia. Rubber Research Institute of Malaysia,
Kuala Lumpur, Malaysia.
Ray, S.K. (2003) Efcacy of different fungicides for management of powdery mildew of
mango in West Bengal. Journal of Mycopathological Research 41, 153155.
Reckhaus, P. and Adamou, I. (1987) Hendersonula dieback of mango in Niger. (Disease
note). Plant Disease 71, 1045.
Reddy, G.S., Govinda, R.P. and Paparao, A. (1961) Black banded disease of mango.
Andhra Agricultural Journal 8, 120123.
Reuveni, M., Harpaz, M. and Reuveni, R. (1998) Integrated control of powdery mildew
on eld-grown mango trees by foliar sprays of mono-potassium phosphate fertilizer,
sterol inhibitor fungicides and the strobilurin Kresoxym-methyl. European Journal
of Plant Pathology 104, 853860.
Ribiero, I.J.A. (1980) Seca de mangueira. Agentes causais e estudio da molesta. Anais do
I Simposio Brasiliero Sobre a Cultura de Mangueira, 2428 Nov. 1980. Sociedad
Brasileira de Fruticultura, Jaboticobal, Brazil, pp. 123130.
Ribiero, I.J.A. (1997) Doenas da mangueira (Mangifera indica L.). In: Kimati, H., Amo-
rim, A., Bergamin-Filho, A., Camargo, L.E.A. and Rezende, J.A.M. (eds) Manual de
R.C. Ploetz and S. Freeman 300
Fitopatologia. Vol. 2: Doenas das Plantas Cultivadas. Agronmica Ceres, So
Paulo, Brazil, pp. 511524.
Ribiero, I.J.A., Rosetto, C.J., Donadio, L.C., Sabino, J.C. Martins, A.L.M. and Gallo, P.B.
(1995) Mango wilt. XIV. Selection of mango (Mangifera indica L.) rootstocks resis-
tant to the mango wilt fungus Ceratocystis mbriata Ell. and Halst. Acta Horticultu-
rae 370, 159161.
Ridgeway, R. (ed.) (1989) Mango Pests and Disorders. Department of Primary Industries,
Brisbane, Queensland, Australia.
Rivera-Vargas, L., Lugo-Noel, Y., McGovern, R.J., Sijo, T. and Davis, M.J. (2006) Occur-
rence and distribution of Colletotrichum spp. on mango (Mangifera indica L.) in
Puerto Rico and Florida, USA. Plant Pathology Journal 5, 191198.
Rodrguez-Alvarado, G., Fernndez-Pava, S.P. and Ploetz, R.C. (2006) Fusarium sp.
characterization causing mango malformation in Michoacan Mexico. Phytopathol-
ogy 96, S99.
Rodrguez-Alvarado, G., Fernndez-Pava, S.P., Ploetz, R.C. and Valenzuela-Vzquez,
M. (2008) A Fusarium sp., different from Fusarium oxysporum and F. mangiferae, is
associated with mango malformation in Michoacn, Mexico. New Disease
Reports. Plant Pathology 57, 781.
Rossetto, C.J., Ribeiro, I.J.A., Igue, T. and Gallo, P.B. (1996) Mango wilt: XV. Varietal re-
sistance against two isolates of Ceratocystis mbriata. Bragantia 55, 117121.
Rossman, A.Y., Samuels, G.J., Rogerson, C.T. and Lowen, R. (1998) Genera of Bionectri-
aceae, Hypocreaceae and Nectriaceae (Hypocreales, Ascomycetes). Studies in My-
cology 42, 1248.
Ruehle, G.D. and Ledin, R.B. (1955) Mango Growing in Florida. Bulletin 574. Agricul-
tural Experiment Station, University of Florida, Gainesville, Florida.
Saaiman, W.C. (1996) Biology and control of Nattrassia mangiferae. Acta Horticulturae
455, 558564.
Saeed, A. and Schlosser, E. (1972) Effect of some cultural practices on the incidence of
mango malformation. Zeitschrift fr Panzenkrankheiten und Panzenschutz 79,
349351.
Sanders, G.M., Verschoor, J.A., van Wyngaard, S., Korsten, L. and Kotz, J.M. (1994)
Production of monoclonal antibodies against Xanthomonas campestris pv. mangifer-
aeindicae and their use to investigate differences in virulence. Journal of Applied
Bacteriology 77, 509518.
Schaffer, B. (1994) Decline. In: Ploetz, R.C., Zentmyer, G.A., Nishijima, W., Rohrbach,
K. and Ohr, H.D. (eds) Compendium of Tropical Fruit Diseases. American Phyto-
pathological Society (APS) Press, St Paul, Minnesota, p. 43.
Schaffer, B., Larson, K.D., Snyder, G.H. and Sanchez, C.A. (1988) Identication of mineral
deciencies associated with mango decline by DRIS. HortScience 23, 617619.
Schoeman, M.H., Manicom, B.Q. and Wingeld, M.J. (1995) Epidemiology of powdery
mildew on mango blossoms. Plant Disease 79, 524528.
Silveira, S.F., Harrington, T.C., Mussi-Dias, V., Engelbrecht, C.J.B., Alfenas, A.C. and Sil-
va, C.R. (2006) Annona squamosa, a new host of Ceratocystis mbriata. Fitopatolo-
gia Brazileira 31, 394397.
Singh, L.B. (1968) The Mango: Botany, Cultivation, and Utilization. Leonard Hill,
London.
Singh, R.N., Majumder, P.K., Sharma, D.K., Sinha, G.C. and Bose, P.C. (1974) Effect of
de-blossoming on the productivity of mango. Scientia Horticulturae 2, 399403.
Singh, Z. and Dhillon, B.S. (1989) Hormonal changes associated with vegetative malfor-
mation of mango (Mangifera indica L.). Journal of Phytopathology 125, 193197.
Foliar, Floral and Soilborne Diseases 301
Singh, Z., Singh, L., Arora, C.L. and Dhillon, B.S. (1994) Effect of cobalt, cadmium, and
nickel as inhibitors of ethylene biosynthesis on oral malformation, yield, and fruit
quality of mango. Journal of Plant Nutrition 17, 16591670.
Slippers, B. and Wingeld, M.J. (2007) Botryosphaeriaceae as endophytes and latent
pathogens of woody plants: diversity, ecology and impact. Fungal Biology Reviews
21, 90106.
Slippers, B., Johnson, G.I., Crous, P.W., Coutinho, T.A., Wingeld, B.D. and Wingeld,
M.J. (2005) Phylogenetic and morphological re-evaluation of the Botryosphaeria
species causing diseases of Mangifera indica. Mycologia 97, 99110.
Smith, P.F. and Scudder, G.K. (1951) Some studies of mineral deciency symptoms in
mango. Proceedings of the Florida State Horticultural Society 64, 243248.
Snowdon, A.L. (1990) Color Atlas of Post-Harvest Diseases and Disorders of Fruits and
Vegetables. Vol. 1. General Introduction and Fruits. CRC Press, Boca Raton,
Florida.
Sreenivasaprasad, S., Mills, P.R., Meehan, B.M. and Brown, A.E. (1996) Phylogeny and
systematics of 18 Colletotrichum species based on ribosomal DNA spacer sequenc-
es. Genome 39, 499512.
Steenkamp, E.T., Wingeld, B.D., Coutinho, T.A., Wingeld, M.J. and Marasas, W.F.O.
(1999) Differentiation of Fusarium subglutinans f. sp. pini by histone gene sequence
data. Applied and Environmental Microbiology 65, 34013406.
Steenkamp, E.T., Britz, H., Coutinho, T.A., Wingeld, B.D., Marasas, W.F.O., Wingeld,
B.D., Marasas, W.F.O. and Wingeld, M.J. (2000) Molecular characterization of
Fusarium subglutinans associated with mango malformation. Molecular Plant
Pathology 1, 187193.
Steyn, P.L., Viljoen, N.M. and Kotz, J.M. (1974) The causal organism of bacterial black-
spot of mangoes. Phytopathology 64, 14001404.
Summanwar, A.S, Raychaudhuri, S.P. and Phatak, S.C. (1966) Association of the fungus
Fusarium moniliforme Sheld. with the malformation in mango (Mangifera indica L.).
Indian Phytopathology 19, 227228.
Sutton, B.C. (1980) The Coelomycetes. Commonwealth Mycological Institute, Kew, Sur-
rey, UK, 696 pp.
Taba, S., Takaesu, K., Ooshiro, A., Moromizato, Z. and Takushi, T. (2004) Anthracnose of
mango caused by Colletotrichum acutatum. Japanese Journal of Tropical Agricul-
ture 48, 5761.
Tavares, S.C.C. de H., da Cruz, S.C., de Siqueira, J.G., Lima, M.L.C., Silva, P.C.G.C., de
Neves, R.A.F. and Menezes, C.A. (2004) Alternative control to powdery mildew
(Erysiphe poligoni d. c. Sin) on mango in submedio Sao Francisco river in Brazilian
semi-arid region. Acta Horticulturae 645, 471474.
Tiwari, R.K.S., Ashok, S., Rajput, M.L. and Bisen, R.K. (2006) Relative susceptibility of
mango varieties to powdery mildew caused by Oidium mangiferae. Advances in
Plant Sciences 19, 181183.
Tsao, P.H., Luzaran, P.B., de los Santos, A.B., Portales, L.A., Gochango, A.M. and Gru-
ber, L.C. (1994) Phytophthora crown and foot rot of mango detected in Philippine
nurseries. (Disease note). Plant Disease 78, 100.
Uppal, B.N., Patel, M.K. and Kamat, M.N. (1941) Powdery mildew of mango. Journal of
the University of Bombay 9, 1216.
van Staden, J. and Nicholson, R.J.D. (1989) Cytokinins and mango ower malformation.
The cytokinin complement produced by Fusarium moniliforme and the ability of
the fungus to incorporate (8-14C) adenine into cytokinins. Physiological and Mo-
lecular Plant Pathology 35, 423431.
R.C. Ploetz and S. Freeman 302
van Staden, J., Bayley, A.D. and Macrae, S. (1989) Cytokinins and mango ower malfor-
mation. III. The metabolism of (3H) iso-pentenyladenine and (8-14C) zeatin by
Fusarium moniliforme. Physiological and Molecular Plant Pathology 35, 433438.
van Wyk, M., Al Adawi, A.O., Wingeld, B.D., Al Subhi, A.M., Deadman, M.L. and
Wingeld, M.J. (2005) DNA based characterization of Ceratocystis mbriata iso-
lates associated with mango decline in Oman. Australasian Plant Pathology 34,
587590.
van Wyk, M., Al Adawi, A.O., Khan, I.A., Deadman, M.L., Wingeld, B.D., Ploetz, R.C.
and Wingeld, M.J. (2007) Ceratocystis manginecans sp. nov., causal agent of the
destructive mango die-back disease in Oman and Pakistan. Fungal Diversity 27,
213230.
Varma, A., Lele, V.C., Raychauduri, S.P., Ram, A. and Sang, A. (1974) Mango malforma-
tion: a fungal disease. Phytopathologische Zeitschrift 70, 254257.
Verma, K.S. and Singh, J. (1996a) Perpetuation and management of Macrophoma
mangiferae causing blight of mango. Indian Journal of Mycology and Plant Pathol-
ogy 26, 7578.
Verma, K.S. and Singh, J. (1996b) Pathology and epidemiology of Macrophoma blight of
mango. Indian Journal of Mycology and Plant Pathology 26, 98100.
Verma, O.P. and Singh, R.D. (1970) Epidemiology of mango dieback caused by Botyo-
diplodia theobromae Pat. Indian Journal of Agricultural Science 40, 813818.
Viegas, A.P. (1960) Mango blight. Bragantia 19, 163182. (Abstract in Review of Applied
Mycology (1963) 42, 696.)
Viljoen, N.M. and Kotz, J.M. (1972) Bacterial black spot of mango. Citrus Grower and
Sub-Tropical Fruit Journal June, pp. 58.
Wagle, P.V. (1928) Studies in the shedding of mango owers and fruits. Part I, Memoirs
of the Department of Agriculture. India Botanical Sciences 8, 219249.
Waterhouse, G.M. (1970) The Genus Phytophthora de Bary, 2nd edn. Mycological
Papers No. 122. Commonwealth Mycological Institute, Kew, Surrey, UK.
Webster, R.K. and Butler, E.E. (1967) A morphological and biological concept of the
species Ceratocystis mbriata. Canadian Journal of Botany 45, 14571468.
Weng, F.J. and Chuang, T.Y. (1995) Grouping of mango anthracnose fungus in Taiwan.
Plant Protection Bulletin 3, 295309.
Williamson, S.M. and Sutton, T.B. (2000) Sooty blotch and yspeck of apple: etiology,
biology, and control. Plant Disease 84, 714724.
Wood, S.L. (1982) The bark and ambrosia beetles of North and Central America (Co-
leoptera: Scolytidae), a taxonomic monograph. Great Basin Naturalist, Memoirs 6,
11356.
Youssef, S.A., Maymon, M., Zveibil, A., Klein-Gueta, D., Sztejnberg, A., Shalaby, A.A.
and Freeman, S. (2007) Epidemiological aspects of mango malformation disease
caused by Fusarium mangiferae and source of infection in seedlings cultivated in
orchards in Egypt. Plant Pathology 56, 257263.
Zea-Bonilla, T., Martn-Snchez, P.M., Hermoso, J.M., Carmona, M.P., Segundo, E. and
Prez-Jimenz, R.M. (2007) First report of Phytophthora citricola on Mangifera in-
dica in Spain. New Disease Reports. Plant Pathology 56, 356.
Zheng, Q. and Ploetz, R. (2002) Genetic diversity in, and development of a PCR assay
for identifying, the mango malformation pathogen. Plant Pathology 51, 208216.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 303
9 Physiological Disorders
V. Galn Saco
Instituto Canario de Investigaciones Agrarias, Tenerife (Canary Islands), Spain
9.1 Introduction 303
9.2 Internal Fruit Breakdown 304
Symptoms 305
Histology 305
Altered physiology 306
Causes 306
Cultivar susceptibility 307
Control 308
9.3 Lumpy Tissue 309
9.4 Ricey Tissue 310
9.5 Fruit Cracking 310
9.6 Black Tip Disorder 310
9.7 Lenticel Spot 311
9.8 Other Fruit Disorders 311
9.9 Stem Disorders 312
9.1 Introduction
Physiological disorders can be dened as ailments that have not been caused
by infecting organisms. Unlike mango malformation, for example, which has
been considered as a physiological disorder in the past (IBPGR, 1989) but has
a phytopathological origin, true physiological disorders cannot be transmit-
ted from plant to plant, mechanically or by insect bites. For the most part,
they are a result of some form of physical damage or of an altered physiology
of the tree or fruit. In the particular case of fruit disorders the most preva-
lent and important physiological disorders of mango according to Subra-
manyan et al. (1971), they are essentially the result of imbalances in metabolism
induced by some factor or factors in the pre- or postharvest environment
that leads to cell collapse and, typically, the appearance of waterlogged or
V. Galn Saco 304
brown areas on some part of the fruit. The complexity of events leading to
the occurrence of physiological disorders makes it more difcult to pinpoint
the causal factors than is the case of symptoms caused by pathogens or pests.
Physiological disorders occur relatively frequently in many fruit and
vegetable species. Some well-known examples include bitter pit, water core
and internal breakdown in apple (Atkinson et al., 1980) and in watermelon
(Singh et al., 1975; Cirulli and Ciccarese, 1981), blossom end rot in tomato and
pepper (Winsor and Adams, 1987), mesocarp discoloration and pulp spot in
avocado (Van Lelyveld et al., 1984; Bower and Cutting, 1987) and yellow pulp
in banana (Melin and Aubert, 1973).
Although physiological disorders of annual crops and temperate fruits
have been studied extensively, tropical fruits have only recently been stud-
ied in order to understand and control physiological disorders. This is largely
due to the rapid expansion of these crops during the past few decades.
Preharvest factors that predispose mango fruit to physiological disorders
include growing location, orchard condition, tree nutrition and condition at
the time of harvest. Postharvest storage conditions, such as temperature,
oxygen and carbon dioxide levels, packaging and surface coating treatments,
are also potential contributing factors to the occurrence of these disorders
(Subramanyan et al., 1971).
Evidence of specic metabolic changes that occur in mango fruits that
express this type of disorder is scarce. Probably due to this lack of knowl-
edge, disorders are usually named for the associated environmental factor,
for example chilling injury, or by the altered appearance that the physiologi-
cal disorder confers to the fruit. Clear examples of the latter are internal fruit
breakdown, which is by far the most prevalent physiological disorder in
mango, lenticel spot, fruit cracking and black tip disorder.
9.2 Internal Fruit Breakdown
Most opinion (Winston, 1984; Schaffer, 1994) is that several physiological dis-
orders currently listed as separate disorders are in fact different degrees or
aspects of the same problem, internal fruit breakdown (IFB). These include
tip pulp, soft nose (i.e. waterlogging of the esh near the distal end), jelly
seed (i.e. overripe esh around the seed, surrounded by rm esh), stem end
breakdown (i.e. an open cavity in the pulp at the stem end), spongy tissue
(i.e. areas of the esh that appear spongy and have a greyish black discolor-
ation), soft centre and soft esh. This aggregation is still open to some debate.
Raymond et al. (1998) asserted that temporal and spatial differences in symp-
tom development within the fruit have been found between soft nose, jelly
seed and stem end cavity, validating their return to separate disorder status,
but they recognized that no major microscopic differences could be found.
None the less, irrespective of the disparity of symptoms, all of these disor-
ders have a common denominator: calcium imbalance. With this dening
criterion, IFB would also include the physiopathy described in Malaysia as
yeasty or insidious fruit rot, which shows a marked interdependence with
Physiological Disorders 305
calcium (Lim and Khoo, 1985) and also what Velasco Crdenas (1974) calls
ablandamiento del pico (literally peak softening).
Cheema and Dani (1934) rst reported an IFB disorder, but perhaps the
most comprehensive work published on physiological disorders of the mango
fruit was that of Malo and Campbell (1978), who described the different
degrees of tissue decomposition in fruit of Tommy Atkins. The comprehen-
sive work of Wainwright and Burbage (1989) reviews the symptomatol-
ogy, chemical changes and causes of IFB, illustrating its occurrence in many
cultivars in virtually all the producing areas of the world. Losses from IFB
vary geographically and also among cultivars, and can affect 100% of the
harvested fruit (Subramanyan et al., 1971; Malo and Campbell, 1978; Bro-
drick and Thord-Gray, 1982; Galn Saco et al., 1984; Santos Filho et al., 2002;
Cracknell Torres et al., 2003a).
Symptoms
Symptoms usually begin to appear early during fruit development and
advance rapidly, eventually making the fruit inedible (see Plates 4852). The
process commences while the fruit is still hanging on the panicle, with an
interruption occurring in the vascular tissues of the peduncle and endocarp,
and usually followed by the formation of a cavity close to the funiculum
(stem end breakdown). In advanced stages, the affected tissues around the
cavity become grey or blackish. As the disorder progresses, the proximal end
of the fruit becomes mushy to the touch (soft nose). In some cases, a yellow-
ing of the skin between the apex and the stigmatic end of the fruit can also
occur. The bre surrounding the endocarp may fully disintegrate and in
severe cases an accumulation of a transparent, gelatinous substance devel-
ops around the seed (jelly seed). The affected mesocarp matures more rap-
idly than the healthy esh and acquires a characteristic deeper yellow
colour; the affected tissue may be so extensive that greyish, watery tissues
appear over the whole mesocarp (spongy tissue) (Malo and Campbell, 1978;
Winston, 1984; Joshi and Roy, 1985; Wainwright and Burbage, 1989; Katrodia
and Chuva, 1993). The absence of latex and the lack of rmness in the proxi-
mal end of the fruit at harvest can be clear signs of the disorder (Chaplin,
1986), although the lack of latex alone has also been observed in black tip
disorder (see below).
Histology
Histological differences have been found between fruits of different cultivars
affected by IFB. For example, in the very susceptible Tommy Atkins the xylem
elements in the mesocarp and the mesocarp cells are all highly deteriorated. In
the case of a less sensitive cultivar (i.e. Lippens) the affected mesocarp cells
have intact cell walls, even though the tissue is obviously affected, showing
a translucent mesocarp (Cracknell Torres and Galn Saco, 2003).
V. Galn Saco 306
Altered physiology
A reduction in rmness, total soluble solids, total pectin and pectinase activ-
ity was observed by these researchers in the affected mesocarp tissue, con-
rming the results of Van Lelyveld and Smith (1979) and Roe and Bruemmer
(1981), both of whom also observed greater activity of both pectinase and
malic enzyme in affected pulp of Alphonso. On the other hand, the higher
level of nitrogen, the lower levels of calcium, and the higher nitrogen to cal-
cium ratio observed by Cracknell Torres and Galn Saco (2003) in the
affected tissue of Tommy Atkins and Lippens is in agreement with the
studies of Young (1960) and Young and Miner (1961) but in conict with
results of Krishnamurthy (1981), who found no differences among calcium
levels between affected and unaffected mesocarp tissues. Other differences
in chemical composition have been detected in affected and unaffected tis-
sues (Subramanyan et al., 1971; Patkar et al., 1983; Nuevo et al., 1984; Gupta
et al., 1985) and further research should be devoted to ascertain the true
nature of this disorder.
Causes
Despite many attempts to isolate a pathogen from affected tissues, no patho-
gen has yet been linked with IFB. In Malaysia, spongy tissue has been found
in Harumanis fruit which has been affected by the fruit-piercing moth
Othreis spp. and in Fan Siamese fruit, which has been associated with dam-
age caused by phytotoxic sprays of some fungicides (Lim and Khoo, 1985).
Postharvest vapour heat treatment at 46C for 10 min has also been reported
to induce IFB in Carabao fruit (Esquerra et al., 1990).
Several environmental factors have been linked to IFB. For example, in
India the disorder appears to be more prevalent in coastal areas (Subraman-
yan et al., 1971). Gunjate et al. (1982) observed that it is associated with fruit
that remains exposed to the sun following harvest. IFB has also been
attributed to heat convection from the soil at air temperatures around
55C (Katrodia and Rane, 1989). In several experiments with the cultivar
Alphonso in India, a positive correlation was observed between IFB inci-
dence and relative density. Mangoes that were harvested with a relative den-
sity between 1.00 and 1.20 showed no IFB, while fruits greater than 1.2 had a
30% incidence of IFB (Krishnamurthy, 1980). Fruit weight also seems to be an
important factor at least in the case of some cultivars like Alphonso in India
(Subramanyan et al., 1971) and in Spain with Sensation (Hermoso et al.,
1996), with a positive correlation in both cases between fruit weight and
incidence of IFB.
Cultural practices such as excessive irrigation have also been considered
to be possible causes of IFB. Katrodia (1988) observed a higher incidence of
IFB following periods of rainfall just prior to harvesting of Alphonso in India;
however, trials done elsewhere with Tommy Atkins (Malo and Camp-
bell, 1978) or Sensation (Farr and Hermoso, 1993) showed no correlation
Physiological Disorders 307
between excessive irrigation and IFB. Furthermore, in Malaysia the incidence
of IFB (insidious or yeasty fruit rot) has been reported to be higher during the
drier months of the year (Lim and Khoo, 1985).
Most observations and eld studies associate IFB with calcium deciency,
which is also the main cause of similar disorders in other fruits (Bangerth,
1979; Winsor and Adams, 1987). In general, low levels of leaf calcium at har-
vest coincide with high IFB incidence. In Florida, IFB appears to occur more
in acid and sandy soils with low calcium content than in basic, calcareous
soils (Young, 1957; Schaffer, 1994), but in Harumanis yeasty or insidious
fruit rot occurs irrespective of various types of soils, ranging from acidic to
alkaline. It has been suggested that the degree of IFB is aggravated in the
presence of excess nitrogen fertilization (Young, 1960; Young and Miner,
1961; Young et al., 1962, 1965).
The direct relationship between IFB and calcium and nitrogen content
has been conrmed by analysis of affected and non-affected fruits of
orchard-grown Harumanis (Ahmad Tarmizi et al., 1993), and in soilless cul-
tivation trials involving Tommy Atkins (Cracknell Torres et al., 2003b). In
the latter study, positive correlations were observed between mesocarpic
nitrogen levels and IFB and between the nitrogen:calcium ratio and IFB, but
there was a negative relationship between calcium and IFB.
Cultivar susceptibility
Although IFB has been observed in at least 65 cultivars in 23 countries (Galn
Saco, 1999), it is clear that varietal differences play a role in the intensity and
expression of the symptoms, as these have been described in different degrees
of intensity according to countries and cultivars.
According to Young (1957), cultivars of Indian origin and their offspring
show a higher incidence of IFB, for example Alphonso whose susceptibility
has been well documented (Subramanyan et al., 1971; Gunjate et al., 1979,
1982; Krishnamurthy, 1980, 1981; Patkar et al., 1983; Gupta et al., 1985; Lim
and Khoo, 1985; Katrodia and Rane, 1989; Lad et al., 1992). There is, however,
conicting evidence; some Indian cultivars, for example Rajapuri (Katrodia,
1988), Banganapalli, Kalapardy, Janradhan Pasand (Iyer and Subraman-
yan, 1992), Pairi, Kesar and Neelum (Lad et al., 1992), rarely demonstrate
these disorders. All Florida cultivars are affected due to their Indian parent-
age (Schaffer, 1994); Haden, Kent, Keitt, Sensation, Tommy Atkins and
Van Dyke are repeatedly mentioned as particularly susceptible (De Larous-
silhe, 1980; Galn Saco and Fernndez Galvn, 1987; Campbell, 1992; Oost-
huyse, 1993; Knight, 1997; Galn Saco, 1999; Cracknell Torres et al., 2003a).
IFB problems are also cited for the Malaysian cultivar Harumanis or Aru-
manis (Lim and Khoo, 1985; Tengku Ab.Malik, 1992a, b; Ahmad Tarmizi et al.,
1993).
IFB incidence in some cultivars is so low as to be virtually non-existent,
particularly the polyembryonic and brous-eshed types like Turpentine
in Florida (Malo and Campbell, 1978), Turpentine, Coquinho and Espada
V. Galn Saco 308
in Brazil (Ferreira, 1989), and Gomera 1 in the Canary Islands (Cracknell Tor-
res et al., 2003a). There is at least one documented case of IFB in Australia
involving fruit of the polyembryonic Kamerunga White (Winston, 1984).
Cultivar-related differences with respect to IFB incidence have been
described by Cracknell Torres et al. (2003a). They studied 28 cultivars and
observed that Edward, Gomera 1, Irwin, Valencia Pride, Mabroka, Ah
Ping and Heidi were almost free of this disorder. Modern Israeli cultivars,
such as Shelly and Tango have also exhibited low frequency of IFB (Lavi et
al., 1997a, b). Observations of the extent of spongy tissue in various hybrids
and selfed progenies of Alphonso clearly indicate that this disorder is
genetically determined, and Alphonso is apparently homozygous recessive
for this character (Iyer and Subramanyan, 1992).
Control
Despite the close relationship that exists between nutrition and IFB, very few
practical recommendations have been proposed for controlling this problem.
There is general agreement with the observations of Young (1960) and Young
and Miner (1961) that maintaining a low leaf content of nitrogen (<1.2%) and
a calcium level 2.5% minimizes the amount of affected fruits. High nitrogen
levels enhance vegetative growth and rapid fruit development. Calcium
moves only in the xylem. Calcium concentration in organs such as fruits,
which have a low rate of transpiration and which are preferably supplied via
the phloem, can result in calcium levels falling below the critical level
required for membrane integrity and cell wall stability (Marschner, 1995).
The antagonistic effect of nitrogen fertilization, especially when ammonium
salts are applied, and calcium uptake and accumulation on leaves and fruits,
is well documented in many fruit species (Shear, 1975; Lewis et al., 1977; Lud-
ders, 1979) and has been clearly demonstrated for mangoes (Young et al.,
1962, 1965).
Lime application has also been recommended to increase the cation
exchange percentage to values 7.0 (Ferreira, 1989). According to Schaffer
(1994), IFB has been corrected in Keitt by calcium application either to the soil
as calcium carbonate (CaCO
3
) or as a foliar calcium nitrate (Ca (NO
3
)
2
) spray.
Cultural practices such as mulching have been cited as being benecial
for reducing IFB. Mulching lowers the soil temperature and thereby miti-
gates the reected or rising heat that eventually affects the fruit (Katrodia,
1988; Lad et al., 1992). The accompanying reduction in the transpiration
stream of the tree and the consequent minor mobilization of calcium to the
fruit (Bangerth, 1979) may also be important.
Certain rootstocks can improve calcium uptake and accumulation and
provides an important new procedure for controlling IFB. Fieldwork was
initiated in Malaysia at the end of the last century (Tengku Ab.Malik, 1996),
but further studies remain unreported.
Early harvesting at the green-ripe stage has been recommended as a
measure to reduce IFB (Young, 1957; Young and Miner, 1961; Galn Saco
Physiological Disorders 309
et al., 1984; Winston, 1984), but for some cultivars this practice results in
lower quality fruits. The elimination at harvesting of fruits that do not exude
latex (Mead and Winston, 1991) and avoiding the exposure of fruit to direct
sunlight during harvest (Gunjate et al., 1982) have been recommended as prac-
tical methods to reduce the incidence of IFB in mangoes in the market. Accord-
ing to Santos Filho et al. (2002), postharvest hydrothermal treatments should
be avoided in fruits coming from orchards with a history of IFB incidence.
Preharvest dips of 0.52.0% of calcium chloride (CaCl
2
), applied from the
second month after fruit set until harvest, can increase calcium content in the
fruit, and has been reported to be an effective control measure for spongy
tissue in fruit of Alphonso (Gunjate et al., 1979). In contrast, foliar sprays
with calcium have been reported to be ineffective for increasing leaf cal-
cium content (McKenzie, 1995), and IFB has even been increased by applica-
tion of calcium, which indicates that a nutrition imbalance within the fruit
was induced (Oosthuyse, 1997).
An integrated approach to control insidious fruit rot incidence in Haru-
manis has been developed in Malaysia. This includes maintaining soil pH
close to 6.0, late pruning, application of calcium sprays (2% CaCl
2
) to the
fruits 46 weeks after owering, supplemental irrigation, monitoring nitro-
gen and potassium fertilization and anticipated harvesting (Tengku Ab.
Malik, 1992a, b). Galn Saco (1999) suggested the following general inte-
grated management measures: avoid planting sensitive cultivars (i.e. Sensa-
tion, Tommy Atkins, etc.); use appropriate rootstocks (i.e. Tangkai Panjang
or others with high calcium absorption capacity); harvest at the green-ripe
stage; use an appropriate fertilization programme that is rich in calcium and
poor in nitrogen; avoid excessive irrigation close to fruit maturity; and
mulch soil in regions with hot summers.
There are notable differences in IFB resistance among cultivars. Because
of the widespread incidence of this problem, breeding for rootstocks and sci-
ons for IFB resistance is critical for resolving this problem (Iyer and Degani,
1997). Iyer and Subramanyan (1992) have described the progress of breeding
studies to overcome IFB in India, obtaining hybrids of Alphonso by other
Indian cultivars free from spongy tissue.
Biotechnological approaches also merit attention. Litz and Lavi (1997),
based on work done in tomatoes to control jelly seed (Smith et al., 1990;
Oeller et al., 1991), have suggested that control of IFB could be achieved by
manipulating ripening using the antisense strategy, either by blocking ethyl-
ene biosynthesis or with polygalacturonase, but no work has been reported
in mango so far to test this suggestion.
9.3 Lumpy Tissue
This disorder, of unknown etiology, occurs frequently in Thailand, where it
occurs mainly in the cultivars Namta-an, Parg-grabong and Pinsen Dang,
and in the Philippines in Pico. The disorder becomes apparent as fruit
ripening begins, with indentations or scoring of the peel, which become
V. Galn Saco 310
increasingly marked as ripening progresses. Opened fruits display a meso-
carp full of white lumps of intact, starch-lled cells (Lizada et al., 1984).
9.4 Ricey Tissue
This disorder is of unknown origin, but is common in Carabao in the Philip-
pines. No external symptoms are visible, but internal symptoms are found in
ripening fruit and even at physiological maturity, and consist of small
lesions. The lesions resemble grains of rice in size and aspect (hence the
name) on the mesocarp, surrounded by cotton wool-like tissue. Except for
the small changes in the texture of the affected area, no other organoleptic
changes have been detected (Lizada et al., 1984).
9.5 Fruit Cracking
The only symptom of this disorder is that the fruit crack suddenly while still
on the tree, the cracks forming clean, knifelike cuts breaking the skin and
into the esh. Secondary infections by Colletotrichum gloeosporioides (the
causal agent of anthracnose) or other fungal pathogens can occur. The cause
of fruit cracking appears to be related to water tension differences in the skin
during periods of high relative humidity and it occurs when trees are heavily
irrigated after a prolonged dry period, and if heavy rains are intermixed with
dry spells. Frequently almost all of the fruits on the tree are affected, although
those near maturity are the most susceptible. The incidence of fruit cracking
seems to be higher in cultivars with little or no bre. Control measures may
include regular watering and mulching, but the ultimate solution may lie in
breeding as in IFB (Lim and Khoo, 1985).
Another cause for fruit splitting is infection by bacterial black spot
(Xanthomonas campestris), which is considered to be a major cause of fruit
splitting. Consequently, prevention is the only solution to the problem by
ensuring adequate wind protection, which is also benecial if the damage is
caused by differences in water tension of the skin cells, and/or appropriate
chemical sprays to prevent infection (Meurant et al., 1997).
9.6 Black Tip Disorder
This disorder is widespread in India (Prakash and Srivastava, 1987). Ram
(1988) indicated that it was rst described in 1909. Three disorders frequently
cited prior to 1958, namely taper tip, tip pulp and girdle necrosis, are in fact
variants of black tip. The rst symptom of this disorder is the etiolation and
yellowing of the distal end of fruit, which then turns black and hardens,
causing the fruit to ripen prematurely and making it unmarketable (Plate 53).
The vascular bundles in the pedicel may turn brown and decay. Affected
fruits do not secrete latex at harvest.
Physiological Disorders 311
Outside India, the only known occurrence of black tip is from the Guang-
dong province of China (Zhang et al., 1995). In both India and China, the
disorder occurs in areas close to brick-making kilns, particularly where trees
are more exposed to fume-laden winds coming from the brick works. Although
not fully assessed, it seems that uorine is the causal agent of this disorder
(Zhang et al., 1995), although a dry hot climate may enhance the effect of
gases. Differences in susceptibility among Indian cultivars have been reported
by Ram (1988), who also suggested the possibility of varietal selection for
orchards in the vicinity of brick kilns. Majumder and Sharma (1985) recom-
mend a minimum distance between kiln and trees of 1.6 km in the path of
prevailing winds and 0.8 km on the other sides, as well as recommending
prevention by spraying three times with borax (0.6%) and caustic soda (0.8%),
for example prior to owering, during owering and at fruit set.
9.7 Lenticel Spot
This disorder is characterized by the development of small spots of corky
tissue in the skin lenticels that darken as the fruit changes colour during
ripening, which makes the fruit unmarketable. The causes are not entirely
uncertain, but it is most often associated with incorrect postharvest practices,
including low storage temperatures, high humidity of the storage atmos-
phere, excessive immersion time in postharvest dips and excessive detergent
in wash water (Oosthuyse, 1993; Meurant et al., 1997). Some growers link it
to other causes, such as delayed harvesting or wet conditions during picking.
Differences among cultivars with respect to the severity of this problem have
been observed. It is very common in the late harvest of Keitt in the subtrop-
ics at the beginning of winter under cooler and wetter conditions.
9.8 Other Fruit Disorders
Spots and marks on green fruit can have several different causes: cold dam-
age or chilling injury, usually seen as small raised brown or discoloured spots
on the skin, or pitting surrounded by sunken areas affecting the epidermis
and the endocarp (Lizada et al., 1984). Storage temperatures below 13C have
been identied as major causes of chilling injury, and must be avoided (Meurant
et al., 1997). High temperatures or sudden exposure to sunlight can cause
sunburn, which is characterized by bleached or yellow patches on the skin,
which may become leathery, yellow-brown to black, and lightly sunken.
Trees should be well irrigated during fruit lling and harvested fruit must
always be kept (even briey) in full shade.
Wind damage, resulting from prolonged rubbing of the fruit against
leaves or dead twigs, requires adequate windbreaks and timely pruning of
dead wood. If hail damage is likely to be a frequent problem, the whole
orchard will need to be protected by nets. Damage caused during harvest,
storage and transport, can only be reduced by careful handling. Sapburn,
V. Galn Saco 312
caused by prolonged contact with latex leaking onto the skin from a cut stem,
can be minimized by suitable harvesting and packing operations. High levels
of carbon dioxide (CO
2
) during storage provoke the development of off-
avours and small internal lesions, as well as exudating brown tissues
(Chaplin, 1986) and inhibition of ripening (Thompson, 1971).
A disorder named pulpa negra, literally black esh, which speaks quite
clearly as to the symptoms, has been reported in Mexico, especially in Haden
(Mora et al., 1998). Although the problem was detected in fruits that had been
stored at 13C for more than 20 days, it is not entirely clear that low tempera-
ture is the only cause as it also occurred in fruits reportedly stored only at
room temperature.
9.9 Stem Disorders
While some slight cracking of the bark around the trunk may be normal,
some cultivars, like Manzanillo or Gomera 4, are prone to ssuring, with
an unusually high degree of suberication. This disorder is common in the
Canary Islands, Spain, and it is more pronounced on the sides of the trunk
that are wetted by sprinklers than on the less exposed sides (Fernndez
Galvn and Galn Saco, unpublished). This phenomenon of stem cracking
has been also associated with bad orchard management, particularly with
bad irrigation practices (Mora et al., 1998). Some rootstocks, including
Gomera 1 in the Canary Islands, frequently exhibit gall-like growths along
the branches, thought to be linked to climatic factors which facilitate
swelling of lateral buds without being enough to provoke ushing and the
corresponding shoot elongation.
References
Ahmad Tarmizi, S., Tengku Ab.Malik, T.B., Pauziah, M. and Zahrah, T. (1993) Incidence
of insidious fruit rot as related to mineral nutrients in Harumanis mangoes. Malay-
sian Agricultural Research and Development Institute (MARDI) Research Journal
21, 4349.
Atkinson, D., Jackson, J.E., Sharples, R.O. and Wallery, W.M. (1980) Mineral Nutrition of
Fruit Trees. Butterworths, London.
Bangerth, F. (1979) Calcium related disorders of plants. Annual Review of Phytopathol-
ogy 17, 97122.
Bower, J.P. and Cutting, J.G.M. (1987) Some factors affecting pre-harvest quality in avo-
cado fruit. South African Avocado Growers Association Yearbook 10, 143146.
Brodrick, H.T. and Thord-Gray, R. (1982) Irradiation in perspective the signicance for
the mango industry. South African Mango Growers Association Research Report 2,
2336.
Campbell, R.F. (1992) A Guide to Mangos in Florida. Fairchild Tropical Garden, Miami,
Florida.
Chaplin, G.R. (1986) Postharvest physiology of mango fruit. In: Proceedings of the First
Australian Mango Research Workshop. Commonwealth Scientic and Industrial
Research Organization (CSIRO), Melbourne, Australia, pp. 261270.
Physiological Disorders 313
Cheema, G.S. and Dani, P.G. (1934) Report on the export of mangoes to Europe in 1932
and 1933. Bulletin of the Department of Agriculture, Bombay 170, 131.
Cirulli, M. and Ciccarese, F. (1981) Effect of mineral fertilizers on the incidence of
blossom-end-rot of watermelon. Phytopathology 71, 5053.
Cracknell Torres, A. and Galn Saco, V. (2003) The study of the problem of mango
(Mangifera indica L.) internal breakdown. Acta Horticulturae 645, 167174.
Cracknell Torres, A., Cid Ballarn, M.C., Socorro Monzn, A.R., Fernndez Galvn, D.
and Galn Saco, V. (2003a) Incidence of internal fruit breakdown in various man-
go (Mangifera indica L.) cultivars. Acta Horticulturae 645, 315318.
Cracknell Torres, A., Cid Ballarn, M.C., Socorro Monzn, A.R., Fernndez Galvn, D.,
Rosell Garca, P. and Galn Saco, V. (2003b) Effects of nitrogen and calcium supply
on the incidence of internal fruit breakdown in Tommy Atkins mangoes (Mangifera
indica L.) grown in a soilless system. Acta Horticulturae 645, 387394.
De Laroussilhe, F. (1980) Le Manguier. Techniques Agricoles et Productions Tropicales.
Maisonneuve et Larose, Paris, France.
Esquerra, E.B., Brena, S.R., Reyes, M.U. and Lizada, M.C.C. (1990) Physiological break-
down in vapour heat-treated Carabao mango. Acta Horticulturae 269, 425434.
Farr, J.M. and Hermoso, J.M. (1993) Mulching and irrigation effects on growth,
cropping and fruit quality of the mango cv. Sensation. Acta Horticulturae 341,
295302.
Ferreira, F.R. (1989) Colapso interno do fruto. Simposio Brasileiro Sobre A Cultura Da
Mangueira 2. Faculdade de Ciencias Agrarias e Veterinarias; Sociedade Brasileira
de Fruticultura, Jaboticabal, Brazil, pp. 149155.
Galn Saco, V. (1999) El Cultivo del Mango. Mundi-Prensa, Madrid, Spain.
Galn Saco, V. and Fernndez Galvn, D. (1987) El cultivo del mango en Canarias.
Cuadernos de Divulgacin 1/87. Consejera de Agricultura Ganadera y Pesca,
Gobierno de Canarias, Spain.
Galn Saco, V., Fernndez Galvn, D. and Calvo, R. (1984) The incidence of soft-nose
in mangos in the Canary Islands. Proceedings of the Florida State Horticultural
Society 97, 358360.
Gunjate, R.T., Tare, S.J., Rangwala, A.D. and Limaye, V.P. (1979) Effect of pre-harvest and
post-harvest calcium treatments on calcium content and occurrence of spongy
tissue in Alphonso mango fruits. Indian Journal of Horticulture 37, 140144.
Gunjate, R.T., Walimbe, B.P., Lad, B.L. and Limaye, V.P. (1982) Development of internal
breakdown in Alphonso mango by post-harvest exposure of fruits to sunlight.
Science and Culture 48, 188190.
Gupta, D.N., Lad, B.L., Chavan, A.S. and Salvi, M.J. (1985) Enzyme studies on spongy
tissue: a physiological ripening disorder in Alphonso mango. Journal of Maharash-
tra Agricultural University 10, 280282.
Hermoso, J.M., Guirado, E., Gmez, R., Castilla, A., Velasco, J.M. and Farr, J.M. (1996)
Effects of nutrients and growth substances on internal breakdown of Sensation
mango fruits. Acta Horticulturae 455, 9299.
International Board for Plant Genetic Resources (IBPGR) (1989) Descriptors for Mango.
IBPGR, Rome.
Iyer, C.P.A. and Degani, C. (1997) Classical breeding and genetics. In: Litz, R.E. (ed.)
The Mango, Botany, Production and Uses. CAB International, Wallingford, UK,
pp. 4968.
Iyer, C.P.A. and Subramanyan, M.D. (1992) Possibilities of overcoming physiological
disorders in mango by breeding. Acta Horticulturae 317, 241244.
Joshi, G.D. and Roy, S.K. (1985) Spongy tissue in mango, a physiological disorder. Indian
Journal of Horticulture 29, 2122.
V. Galn Saco 314
Katrodia, J.S. (1988) Spongy tissue in mango causes and control measures. Acta Hor-
ticulturae 23, 814826.
Katrodia, J.S. and Chuva, H.P. (1993) Spongy tissue in mango. In: Chadha, K.L. and Pa-
reek, O.P. (eds) Advances in Horticulture Vol. 4 Fruit Crops: Part 4. Malhotra
Publishing House, New Delhi, India, pp. 20692080.
Katrodia, J.S. and Rane, D.A. (1989) The pattern of distribution of spongy tissue in the
affected Alphonso fruits at different locations. Acta Horticulturae 231, 873877.
Knight, R.J., Jr (1997) Important mango cultivars and their descriptors. In: Litz, R.E. (ed.)
The Mango, Botany, Production and Uses. CAB International, Wallingford, UK,
pp. 545566.
Krishnamurthy, S. (1980) Internal breakdown during ripening of Alphonso mango
(Mangifera indica Linn.) in relation to specic gravity of the fruit. Journal of Food
Science and Technology 17, 198199.
Krishnamurthy, S. (1981) Chemical studies on internal fruit breakdown in Alphonso
mango (Mangifera indica Linn). Journal of Horticultural Science 56, 247250.
Lad, B.L., Gunjate, R.T. and Salvi, M.J. (1992) Causes and control measures of spongy
tissue disorder in Alphonso mango fruit: an integrated approach. Maharashtra
Journal of Horticulture 6, 2532.
Lavi, U., Kaufman, D., Sharon, D. and Tomer, E. (1997a) Tango: a new mango cultivar.
HortScience 32, 137.
Lavi, U., Kaufman, D., Sharon, D. and Tomer, E. (1997b) Shelly: a new mango cultivar.
HortScience 32, 138.
Lewis, T.L., Martin, D., Cerny, J. and Ratkowsky, D.A. (1977) The effect of increasing the
supply of nitrogen, phosphorous, calcium and potassium to the roots of Morton
Worcester apple trees on leaf and fruit composition and on the incidence of bitter
pit at harvest. Journal of Horticulture Science 57, 409419.
Lim, T.K. and Khoo, K.C. (1985) Diseases and Disorders of Mango in Malaysia. Tropical
Press SDN. BDH. Kuala Lumpur, Malaysia.
Litz, R.E. and Lavi, U. (1997) Biotechnology. In: Litz, R.E. (ed.) The Mango, Botany,
Production and Uses. CAB International, Wallingford, UK, pp. 401424.
Lizada, M.C.C., Kosiyachinda, S. and Mendoza, D.B. (1984) Physiological disorders of
mango. In: Mendoza, D.B. and Willis, R.B.H. (eds) Mango. Association of South-east
Asian nations (ASEAN) Food Handling Bureau, Kuala Lumpur, Malaysia, pp. 6874.
Ludders, P. (1979) The effect of nitrogen nutrition on bitter pit in apple. Communications
in Soil Science and Plant Analysis 6, 261272.
Majumder, P.K. and Sharma, D.K. (1985) Mango. In: Bose, T.K. (ed.) Fruits of India.
Tropical and Subtropical. Naya Prokash, Calcutta, India, pp. 69153.
Malo, S.E. and Campbell, C.W. (1978) Studies on mango fruit breakdown in Florida. Pro-
ceedings of the American Society for Horticultural Science Tropical Region 22, 115.
Marschner, H. (1995) Mineral Nutrition of Higher Plants. Academic Press. London.
McKenzie, C.B. (1995) The effect of calcium and potassium sprays on Sensation mango
leaf nutrients concentration and fruit quality. South Africa Mango Growers Asso-
ciation Yearbook 15, 4447.
Mead, A.J. and Winston, E.C. (1991) Description of the disorder stem-end cavity, possi-
ble causes and suggestions for reducing its incidence in packing sheds. Acta Horti-
culturae 291, 265271.
Melin, P. and Aubert, B. (1973) Observations on a type of abnormal maturation of banana
(yellow pulp) before harvesting. Fruits 28, 831842.
Meurant, N., Holmes, R., MacLeod, N., Fullelove, G., Bally, I. and Kernot, I. (1997)
Mango Information Kit. Agrilink Series. Queensland Horticulture Institute, Depart-
ment of Primary Industries, Brisbane, Queensland, Australia.
Physiological Disorders 315
Mora, A.A., Vega, P.A. and Tliz, O.D. (1998) Enfermedades del mango. In: Tliz, D.
(ed.) El Mango y su Manejo Integrado en Michoacn. GUIM (Grupo Interdisciplin-
ario de Investigacin en Mango). Colegio de Postgraduados, Montecillo, Texcoco,
Mxico, pp. 1844.
Nuevo, P.A., Cua, A.U. and Lizada, M.C.C. (1984) Internal breakdown in Carabao
mango subjected to modied atmospheres. III. Starch in the spongy tissue. Posthar-
vest Research Notes 1, 3435.
Oeller, P.W., Lu, M.W., Taylor, L.P., Pike, D.A. and Theologis, A. (1991) Reversible inhi-
bition of tomato fruit senescense by antisense RNA. Science 254, 437439.
Oosthuyse, S.A. (1993) Disorders of breless mangos grown in South Africa for export.
South Africa Mango Growers Association Yearbook 13, 8088.
Oosthuyse, S.A. (1997) The effect of calcium and magnesium chelate sprays at owering
on fruit quality and physiological disorders in mango. South Africa Mango Growers
Association Yearbook 17, 2932.
Patkar, V.R., Gunjate, R.T. and Lad, B.L. (1983) Effect of spongy tissue on chemical com-
position of ripe Alphonso fruit. South Indian Horticulture 32, 8385.
Prakash, O. and Srivastava, K.C. (1987) Mango Diseases and their Management. A
World Review. Today and Tomorrows Publishers, New Delhi, India.
Ram, S. (1988) Factors associated with black tip and internal necrosis in mango and
their control. Acta Horticulturae 231, 797804.
Raymond, L., Schaffer, B., Brecht, J.K. and Crane, J.H. (1998) Internal breakdown in
mango fruit: symptomatology and histology of jelly seed, soft nose and stem-end
cavity. Postharvest Biology and Technology 13, 5970.
Roe, B. and Bruemmer, J.H. (1981) Changes in pectic substances and enzymes during
ripening and storage of Keitt mangoes. Journal of Food Science 46, 186189.
Santos Filho, H.P., Cavalcanti Cruz de Holanda Tavares, S., Pires de Matos, A., De Ol-
iveira Costa, V.S., Antnio, W. and Ferreira dos Santos, C. (2002) Doenas. Monito-
ramento e controle. In: De Carvalho Gen, P.J. and De Queiroz Pinto, A.C.A. (eds)
Cultura da Mangueira. Embrapa Informaao Tecnolgica, Brasilia, pp. 299352.
Schaffer, B. (1994) Mango disorders caused by abiotic factors. In: Ploetz, R.C., Zent-
myer, G.A., Nishijima, W.T., Rohrbach, K.G. and Ohr, H.D. (eds) Compendium of
Tropical Fruit Diseases. APS Press, The American Phytopathological Society, Min-
nesota, USA, pp. 4344.
Shear, C.B. (1975) Calcium nutrition and quality in fruit crops. Communications in Soil
Science and Plant Analysis 6, 233244.
Singh, R., Kumar, J.C. and Nandpuri, K.S. (1975) A study on the inuence of the struc-
tural chemical constituents of the skin of watermelon (Citrullus lanatus Sch.) fruit
on the incidence of its blossom-end-rot and cracking (physiological disorders). In-
dian Journal of Horticulture 32, 98101.
Smith, C.J.S.,Watson, C.F. and Morris, P.C. (1990) Inheritance and effect on ripening of
antisense polygalacturonase genes in transgenic tomatoes. Plant Molecular Biology
14, 369374.
Subramanyan, H., Krishnamurthy, S., Subhadra, N.V., Dalal, V.B., Rahdhawa, G.S. and
Chacko, E.K. (1971) Studies on internal breakdown, a physiological ripening disor-
der in Alphonso mangoes (Mangifera indica L.). Tropical Science 13, 203210.
Tengku Ab.Malik, T.M. (1992a) Pattern of calcium accumulation in Harumanis mango
fruits at different stages of maturity. In: Proceedings of the International Radiation
Protection Association (IRPA) Seminar (Agriculture Sector). Vol.1. Crops and Plants,
Kuala Lumpur, Malaysia, pp. 399400.
Tengku Ab.Malik, T.M. (1992b) Measures to control and prevent Insidious Fruit Rot (IFR)
in Harumanis mangoes. In: Proceedings of the International Radiation Protection
V. Galn Saco 316
Association (IRPA) Seminar (Agriculture Sector). Vol. 1. Crops and Plants, Kuala
Lumpur, Malaysia, pp. 405406.
Tengku Ab.Malik, T.M. (1996) Screening for mango rootstocks with high calcium uptake.
In: Vijaysegaran, S., Pauziah, M., Mohamed, M.S. and Ahmad Tarmizi, S. (comps.)
Proceedings of the International Conference on Tropical Fruits. Vol. III. Supplemen-
tary, 2326 July, Kuala Lumpur, Malaysia. Malaysian Agricultural Research and
Development Institute (MARDI), Kuala Lumpur, Malaysia, pp. 189194.
Thompson, A.K. (1971) The storage of mango fruits. Tropical Agriculture, 48, 6370.
Van Lelyveld, L.J. and Smith, J.H.E. (1979) Physiological factors in the maturation and
ripening of mango (Mangifera indica L.) fruit in relation to the jelly-seed physiolog-
ical disorder. Journal of Horticultural Science 54, 283287.
Van Lelyveld, L.J., Van Gerrish, C. and Dixos, R.A. (1984) Enzyme activities and poly-
phenols related to mesocarp discolouration of avocado fruit. Phytochemistry 23,
15311534.
Velasco Crdenas, J. (1974) El Mango en Mxico. Descripcin, Cultivo, Mejoramiento
y Utilizacin. Comisin Nacional de Fruticultura, SAG, Mexico.
Wainwright, H. and Burbage, M.B. (1989) Physiological disorders in mango (Mangifera
indica L.) fruit. Journal of Horticultural Science 64, 125135.
Winsor, G.W. and Adams, P. (1987) Diagnosis of Mineral Disorders in Plants. Vol. 3.
Glasshouse Crops. Her Majestys Stationery Ofce (HMSO), London.
Winston, E.C. (1984) Observations of internal mango esh breakdown: need for stan-
dardization of terminology. In: Proceedings of the First Australian Mango Research
Workshop. Commonwealth Scientic and Industrial Research Organization (CSIRO)
Press, Melbourne, Australia, pp. 7782.
Young, T.W. (1957) Soft-nose, a physiological disorder in mango fruits. Proceedings of
the Florida State Horticultural Society 70, 280283.
Young, T.W. (1960) Response of Kent mangoes to nitrogen fertilization. Proceedings of
the Florida State Horticultural Society 73, 334336.
Young, T.W. and Miner, J.T. (1961) Relationship of nitrogen and calcium to soft-nose
disorder in mango fruits. Journal of the American Society for Horticultural Science
78, 201208.
Young, T.W., Koo, R.C.J. and Miner, J.T. (1962) Effects of nitrogen, potassium and calci-
um fertilization on Kent mangoes in deep, acid, and sandy soil. Proceedings of the
Florida State Horticultural Society 75, 364371.
Young, T.W., Koo, R.C.J. and Miner, J.T. (1965) Fertilizer trials with Kent mangoes. Pro-
ceedings of the Florida State Horticultural Society 78, 369375.
Zhang, Ch., Huang, H. and Kuang, Y. (1995) A study of the cause of the mango black tip
disorder. Scientia Horticulturae 64, 4954.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 317
10 Pests
J.E. Pea,
1
M. Aluja
2
and M. Wysoki
3
1
University of Florida, Florida, USA
2
Instituto de Ecologa, AC, Xalapa, Veracruz, Mexico
3
Institute of Plant Protection, Bet Dagan, Israel
10.1 Introduction 317
10.2 Mango Fruit Pests 318
Fruit ies 318
Fruit y control: brief overview 322
Mango seed weevils 328
Mango seed borer (Lepidoptera: Pyralidae) 331
Fruitspotting bugs (Hemiptera; Coreidae) 332
Thrips 333
Blossom pests 334
Pests of buds and leaves 338
Pests of trunks, twigs and roots 344
10.3 Discussion 345
10.1 Introduction
Mango, like most fruit trees, is usually attacked by two or three key pests,
several secondary pests and by a large number of occasional pests in local-
ized areas where it is grown. Worldwide lists of pests of mango have been
published by Laroussilhe (1980), Tandon and Verghese (1985) and Veeresh
(1989). The pests of mango in India (Srivastava, 1998; Anonymous, 2006),
Australia (Anonymous, 1989), Pakistan (Mohyuddin, 1981), Israel (Wysoki et al.,
1993; Swirski et al., 2002), the USA (Pea, 1993), West Africa (Vannire et al.,
2004), Brazil (Assis and Rabelo, 2005), Central America (Coto et al., 1995) and
Puerto Rico (Martorell, 1975) have also been described. Some publications
contain check lists of mango pests and most contain details of life histories and
control of mango pests (Morin, 1967; Golez, 1991; Murray, 1991).
Of c.322 species of insects and mites that have been recorded as minor
and major pests of mango, 127 (39%) are foliage feeders, 87 (27%) are fruit
feeders, 36 (12%) feed on the inorescence, 33 (10%) inhabit buds and 39
J.E. Pea et al. 318
(12%) feed on branches, the trunk and roots. The four key pests (fruit ies,
seed weevils, tree borers and mango hoppers) require annual control mea-
sures. Secondary pests generally occur at sub-economic levels, but can
become serious pests as a result of changes in cultural practices and cultivar
or because of indiscriminate use of pesticides against a key pest. For exam-
ple, Mohyuddin and Mahmood (1993) reported that scale insects became
serious pests following non-judicious use of insecticides against fruit ies.
Similarly, mites, Oligonychus spp., are secondary pests of mango, which can
become serious because of human intervention. Occasional or incidental
pests also can cause economic damage only in localized areas at certain times.
The majority of pests reported here are of this category.
10.2 Mango Fruit Pests
With current world emphasis on quality fruit for local consumption and
export, insects that blemish fruit by feeding, scratching or ovipositing in the
pulp or seed can cause high losses. Only fruit ies, seed weevils and lepi-
dopterous larvae actually penetrate the fruit pulp and seed. The feeding of
other pests (e.g. Othreis materna (L.), Gonodonta pyrgo (Cram.), Gonodonta clo-
tilda (Stoll) and Leptoglossus stigmai (Herbest)) often extends only into the
pulp of ripening mangoes (Angeles and Requena, 1966).
Fruit ies
Most mango-producing countries are in fruit-y infested areas, and produc-
ers suffer signicant direct economic losses (larval feeding renders fruit
unmarketable) and indirect economic losses (quarantine restrictions hinder-
ing exports), resulting from the presence of fruit ies (Hill, 1975; Umeya and
Hirao, 1975; Anonymous, 1987; Yee, 1987; Singh, 1991; Aluja, 1993; Aluja
et al., 1996; Vannire et al., 2004; Aluja and Mangan, 2008). Few insects have a
greater impact on international marketing and world trade in agricultural
produce than tephritid fruit ies (Hendrichs, 1996; Aluja and Mangan, 2008).
Approximately 60 species of fruit ies are reported to attack mango and a
related species, Mangifera foetida (White and Elson-Harris, 1992; Aluja et al.,
1996; Malavasi and Zucchi, 2000; Clarke et al., 2001, 2005; Norrbom, 2004;
Vayssires et al., 2005). Fruit ies attacking mango belong to the genera
Anastrepha (c.12 species), Bactrocera (c.33 species), Ceratitis (eight species) and
Dirioxa (two species) (White and Elson-Harris, 1992; Vayssires and Kala-
bane, 2000; Lux et al., 2003; Norrbom, 2004; Vayssires et al., 2004, 2005; Sec-
retariat of the Pacic Community, 2005). Some of these species are referred to
as the mango fruit y: Anastrepha obliqua Macquart, Bactrocera frauenfeldi
(Shiner), Ceratitis cosyra (Walker) (Aluja, 1993; Leblanc and Allwood, 1997;
Lux et al., 2003; Steck, 2003). All Dacus species that attack mango have been
recently placed in the genus Bactrocera (Drew) (Thomson, 2005).
Pests 319
The following discussion is mainly restricted to the biology, ecology and
management of mango-infesting fruit ies. Regulatory issues, such as risk
analysis and postharvest treatments are discussed by Johnson and Hofman
in Chapter 15, this volume. Postharvest treatments specically related to
mangoes and fruit ies have been discussed by Sharp et al. (1988, 1989a, b),
Hallman and Sharp (1990), Nascimento et al. (1992), Mangan and Hallman
(1998), Shellie and Mangan (2002a, b) and Bustos et al. (2004). Broad regula-
tory issues were recently reviewed by Follet and Neven (2006).
General reviews on biology, ecology and behaviour of economically
important and non-pestiferous fruit ies, many of them infesting mangoes,
were written or edited by Christenson and Foote (1960), Bateman (1972),
Fletcher (1987), Robinson and Hooper (1989), Aluja (1994), Aluja and Norrbom
(2000) and Malavasi and Zucchi (2000).
Anastrepha
Anastrepha spp. are endemic to the western hemisphere and their range
extends from the southern USA to northern Argentina and includes the
Caribbean islands (Aluja, 1994) (Plate 54). Twelve Anastrepha species have
been purportedly associated with mango (Norrbom, 2004). Of these, A. obli-
qua, A. ludens (Loew) and A. suspensa (Loew) stand out as economically
important pests of mangoes (Aluja et al., 1996; Norrbom, 2004). Anastrepha
obliqua is reportedly the most common fruit y pest in the Americas (Jirn
and Hedstrm, 1988; Nascimento et al., 1992). This species is the most com-
mon fruit y pest of mangoes in Mexico, Costa Rica, Honduras and Guate-
mala (Soto-Maniti et al., 1987; Jirn and Hedstrm, 1991; Aluja et al., 1996;
Camargo et al., 1996; Sponagel et al., 1996), but it also attacks mangoes in
Cuba, Puerto Rico, Jamaica, El Salvador and Venezuela (Norrbom, 2004). In
Mexico, A. ludens commonly attacks mangoes at higher elevations, while A.
obliqua dominates at lower altitudes (Aluja et al., 1996). In Brazil and Ecuador,
mangoes are mainly attacked by A. fraterculus (Wiedemann), but A. turpiniae
Stone, A. serpentina Wiedemann, A. pseudoparallela (Loew) and A. zuelaniae
Stone have been reported (Zucchi, 1988; Rebouas et al., 1996; Arias and Jines,
2004; Norrbom, 2004; Barbosa et al., 2005; A. Malavasi, January 2008, So
Paulo, personal communication).
BIOLOGY OF ANASTREPHA FRUIT FLIES. Anastrepha fruit y biology today is based on
basic studies carried out between 1900 and 1944 (Aluja, 1994, 1999). The basic
life cycle is very similar among most pestiferous Anastrepha species. For
example, egg incubation of A. ludens in mango requires 3.8 days, larval devel-
opment requires 14.2 days and pupal development has been known to need
14.2 days at 27 2C (Leyva et al., 1991). Larvae pass through three instars
before emerging from the fruit and burrowing into the ground to pupate
(Aluja, 1994). Clutch size varies between one egg/clutch in A. obliqua and >100
eggs in A. grandis Mcquart (Aluja, 1994). Importantly, in species laying sev-
eral eggs per clutch (e.g. A. ludens), clutch size is determined by females on a
case-by-case basis and is greatly inuenced by degree of ripeness and the
concomitant degree of epicarp hardness (Daz-Fleischer and Aluja, 2003;
J.E. Pea et al. 320
Birke et al., 2006). Life expectancy varies greatly depending on the host on
which the larvae developed and environmental conditions (for example see
Toledo and Lara 1996, and Malavasi and Zucchi, 2000), but some adults can
live for >150 days (Aluja et al., 2000).
Abundance of A. obliqua populations has been positively correlated with
temperature and negatively correlated with relative humidity (RH) (Herrera
and Vias, 1977). However, studies by Celedonio-Hurtado et al. (1995) and
Aluja et al. (1996) demonstrated the lack of a clear relationship between rain-
fall and Anastrepha y captures in mango orchards in Mexico. They indicated
that overall population uctuation patterns can vary greatly among orchards
within a fairly small geographic region.
Bactrocera
Bactrocera spp. are pests of mango in Africa and Australasia (Drew, 1989;
Leblanc and Allwood, 1997; Leblanc et al., 1997; Tenakanai, 1997; Hancock
et al., 2000; Hollingsworth et al., 2003; Clarke et al., 2005) (Plate 55). The
common species reported on mango include B. tryoni (Frogatt), B. zonata
(Saunders), B. dorsalis (Hendel), B. neohumeralis (Hardy), B. jarvisi (Tryon), B.
papayae Drew and B. frauenfeldi (Schiner) (Umeya and Hirao, 1975; Drew and
Hancock, 1994; Hollingsworth et al., 2003). Two species, B. phillippiensis Drew
& Hancock and B. occipitalis Bezzi, have been recorded for Palau, Pacic
Islands (Secretariat of the Pacic Community, 2005), and recently, a new spe-
cies, B. invadens Drew, Tsuruta and White, was reported for West Africa
(Kenya, Benin) (Lux et al., 2003; Vayssires et al., 2005). Bactrocera correcta
(Bezzi), B. caryeae (Kapoor), B. curcubitae (Coquillett), B. diversa (Coquillett)
and B. tau (Walker) have been reported in India (Australian Government,
2004).
BIOLOGY OF BACTROCERA FRUIT FLIES. The biology of dacine fruit ies was most
recently reviewed by Fletcher (1987); additional details can be found in
Christenson and Foote (1960), Bateman (1972), Robinson and Hooper (1989),
White and Elson-Harris (1992) and Aluja and Norrbom (2000). As with most
pestiferous ies, females within Bactrocera insert their eggs beneath the fruit
skin, especially in ripening fruit; white banana-shaped eggs are usually
deposited in clusters, hatching after 1.520 days (White and Elson-Harris,
1992; Messing, 1999). A single female can lay >1000 eggs over her lifetime
(White and Elson-Harris, 1992). One generation requires c.37 days with a
period of 19 days for egg to adult transformation; eggs hatch 38 h after ovi-
position; larvae develop in 78 days and adults emerge in 1011 days (Mess-
ing, 1999). There are usually three larval instars. The larvae tunnel into the
fruit, contaminating it with frass and providing entry for fungi and bacteria.
Depending on factors such as temperature conditions and type of host, larval
development can be completed in 78 days (Messing, 1999) but can take up
to 2 weeks (White and Elson-Harris, 1992). When the infested fruit is imma-
ture, the fruit ripens prematurely and is unt for marketing. Fully-grown
larvae c.7 mm long drop to the ground and enter the soil where they pupate.
Pests 321
After emergence, the females require a protein source for egg maturation
(White and Elson-Harris, 1992). Studies with B. dorsalis in India (Singh, 1991)
indicated that pupal period was longest (18 days) at 15C and shortest (6
days) at 35C. Warm, humid weather is favourable for Bactrocera fruit ies
and pest populations build up as mango ripening occurs. Bactrocera popula-
tions decrease during dry periods.
Syed et al. (1970) reported that up to 30% of mango fruit were attacked by
B. dorsalis in July and August. Mohyuddin and Mahmood (1993) reported
that mango fruit are heavily attacked in Central Punjab during July and
August, with up to 35% of the fruit being damaged by B. dorsalis and B. zonata.
Vayssires et al. (2005) reported the presence of B. invadens after the rst sig-
nicant rains in mid-April, reaching >900 males captured/trap/week. Trap
captures peaked at 1800 ies/trap/week in mid-June when the presence of
B. invadens was related to ripening of different mango cultivars. Ekesi et al.
(2006) observed that B. invadens shared mango fruit with Ceratitis cosyra in
Africa and suggested that B. invadens is predominantly a lowland pest.
Ceratitis
Eight Ceratitis spp. have been reported to attack mango fruit. The Mediter-
ranean fruit y C. capitata (Wiedemann) is a common polyphagous pest in
mango-growing areas of Hawaii USA, Israel, Australia, Spain, Mexico,
Runion and Brazil and elsewhere in South America (Etienne, 1966; Morin,
1967; Galn-Saco, 1990; Harris et al., 1993; Barbosa et al., 2005; Woods et al.,
2005) (Plate 56). In Africa the most common species are C. cosyra (Walker), C.
fasciventris (Bezzi), C. rosa (Karrsch), C. anonae (Graham) and less frequently
C. capitata (Wiedemann) (Lux et al., 2003); whereas, C. catoirii Guer. occurs in
Runion (Etienne, 1968). Ceratitis quinaria and C. silvestrii are considered of
economic importance in Benin (Vayssires et al., 2005). Ceratitis cosyra is
broadly distributed across Africa and causes enormous damage, which can
result in total loss of the crop. On average about 2030% of mango produc-
tion is lost due to this y species in various African countries (Lux et al.,
2003).
BIOLOGY OF CERATITIS FRUIT FLIES. Flies within Ceratitis, particularly C. capitata,
are quite cosmopolitan, and their basic life cycle varies greatly according to
site (Papadopoulos et al., 1996). In Israel females seek suitable sites for ovipo-
sition and puncture mango fruit early in the season, before the fruit has rip-
ened. According to Wysoki et al. (1993) these barren punctures damage the
fruit, due to the leakage of resins from the fruit. The female can oviposit all
over the fruit, with no preference for any part. Later, when fruit development
is suitable for maggot development, the oviposition sites become light in
colour and the tissue softens. The fully-grown maggots leave the fruit and
pupate in the soil. The developmental period is c.34 weeks and 810 genera-
tions/year can occur depending on temperature and other factors intrinsic to
the y population (Hill, 1975).
J.E. Pea et al. 322
Fruit y control: brief overview
The current trends in fruit y control call for coordinated, area-wide
approaches (Hendrichs, 1996, 2001; Tan, 2000; Huang et al., 2006; Enkerlin,
2007; Orankanok et al., 2007) whose major objective is to overcome the often
ineffective and environmentally unsustainable control schemes resulting
from uncoordinated actions by individual producers. Aluja et al. (1996) pro-
pose that since fruit ies that attack mango also attack other fruit crops in the
same area, their management must be based on mango, wild hosts and other
commercially grown host plants. Thus, to improve the efciency of fruit y
management, host plant blooming and fruiting factors need to be elucidated.
Hendrichs (1996) stated that when fruit growers pursue a concerted fruit y
population management strategy over signicantly large areas, the number
of fruit ies moving into orchards from neighbouring orchards is largely
reduced. Aluja (1993, 1996) and Aluja et al. (1996) suggest a fruit y manage-
ment scheme based on border trapping, enhancement of host-plant resistance
through use of plant growth regulators, use of the sterile insect technique
and bait stations and augmentative parasitoid releases (Sivinski, 1996; Sivin-
ski et al., 1996; Malavasi and Zucchi, 2000; Montoya et al., 2000; Tan, 2000;
Dyck et al., 2005; Mangan and Moreno, 2007). Aluja (1993) and Aluja and
Liedo (1986) state that accomplishment of these goals depends on grower
status (rich versus poor), access to technology, cost, scale (single orchard,
regional level, national level), globalization of markets and local regulations
restricting impact on the environment.
Monitoring and sampling
Monitoring fruit ies attacking mango serves different purposes: (i) to apply
a control or management tactic after the presence of the fruit y is noticed;
and (ii) to verify if fruit y species will attack mango under natural condi-
tions. In general, thresholds for adult fruit ies are quarantine-mediated
(Beers et al., 1993). These thresholds vary from location to location, but de-
pending on the fruit y species they are typically based on the capture of a
single fruit y. In other fruit crops, a threshold of ve ies/trap is recom-
mended resulting in a reduction from four chemical sprays to 1.5 sprays/
season (Beers et al., 1993). Sampling for fruit ies in mango is mostly per-
formed using adult traps, because eggs and young larvae are often difcult
to see in the fruit and because the primary aim of management programmes
is to prevent fruit damage.
In the case of pestiferous species within Anastrepha and some Bactrocera
species, the most widely used traps since the early 1970s for monitoring and
controlling populations are glass and plastic versions of the McPhail trap,
which is baited with a mixture of protein (occasionally hydrolysed cotton
seed together with borax, molasses or fermented juices) and water (Balock
and Lopez, 1969; Jirn, 1995). More recently, human urine has been success-
fully tested as bait for McPhail and McPhail-type traps for resource-poor
farmers in tropical countries (Piero et al., 2003; Aluja and Piero, 2004). The
McPhail trap has provided different results in mango orchards. Balock and
Pests 323
Lopez (1969) reported that high concentrations of McPhail traps reduced the
build-up of y populations and protected mangoes from severe injury dur-
ing certain periods of the year. However, the McPhail trap has several draw-
backs. It is expensive, breaks easily, is cumbersome to service and, most
importantly, is quite inefcient. Aluja et al. (1989), working in a mixed mango
orchard in Chiapas, Mexico, found that only 31.1% of Anastrepha spp. ies
landing on the McPhail trap were caught with many ies entering the trap
but then escaping. Due to the low efciency of the McPhail trap it is being
replaced with Multi-Lure

traps, which provide new trap designs. Dry


synthetic-food-based lures have also been developed, i.e. BioLure

(Suterra
LLC, Inc., Bend, Oregon) (Heath et al., 1995, 1997; Epsky et al., 1999) and
Nu-Lure

(Advanced Pheromone Technologies) (Robacker and Wareld,


1993; Robacker et al., 1997; Robacker, 2001).
Fruit y presence has also been monitored in Australia using Dakpot


fruit y traps hung beneath the tree canopy (Anonymous, 1989). Methyl
eugenol is considered the most powerful male lure for oriental fruit ies.
Methyl eugenol was used for successful monitoring, control and erradication
of B. dorsalis in Oahu Hawaii USA (Steiner and Lee, 1955), Rota Island (Steiner
et al., 1965) and Okinawa, Kume, Miyako and Uaekama Islands (Japan) (Iwa-
hashi, 1984). It has been used for monitoring B. umbrosa (F.) in the Philippines
(Umeya and Hirao, 1975), and is used to lure B. invadens in Africa, which is
unlike other African Dacini species that are attracted to Cue-lures (Lux et al.,
2003; Anonymous, 2005). In Palau, Pacic Islands, two lures are used to
attract mango ies: Bactrocera fraeunfeldi (Schiner) is attracted to Cue-lure,
and B. occipitalis and B. philippinensis Drew and Hancock to methyl eugenol
(Secretariat of the Pacic Community, 2005). Bactrocera dorsalis and B. umbrosa
were monitored and controlled by mass trapping of males with methyl
eugenol and infestations were brought to sub-economic levels in Pakistan
(Mohyuddin and Mahmood, 1993). However, concern over the carcinogenic-
ity of methyl eugenol (Waddell et al., 2004) calls for the development of other
para-pheromones to attract Bactrocera fruit ies. Trimedlure is still consid-
ered an important para-pheromone for the Mediterranean fruit y, with the
exception of C. cosyra adults, which are attracted to terpinyl acetate and not
to trimedlure (Steck, 2003). The attractiveness of mango compounds is
currently being investigated. For example some of the volatiles emitted by
Tommy Atkins mangoes, i.e. terpenes (p-cymene and limonene), are
attractive to C. capitata adults (Hernndez-Snchez et al., 2001).
Many questions linger with respect to the optimal trap number and time
for trap placement in mango groves. In Naru, to produce mango free of B.
frauenfeldi, 300400 traps baited with methyl eugenol plus a toxicant were
placed every square kilometre and trapping density was increased around
mango plantings (Anonymous, 2002). Even though large numbers of traps
can be utilized to increase detection sensitivity, the cumulative costs and
logistical considerations do not make this option practical. Traps with spe-
cic and effective lures that can detect the F
1
generation at low trap densities
(510 traps/km
2
) would t the description of a good detection and monitor-
ing device (Tween, 1993).
J.E. Pea et al. 324
Sampling for earlier fruit y stages can be used to demonstrate that the
fruit is not susceptible to fruit y attack. For instance, the Caribbean fruit y,
A. suspensa, may not attack green mango fruit (Pea et al., 2006b). Pea et al.
(2006b) initiated research to determine if the Caribbean fruit y will attack
green Tommy Atkins mangoes and infest it under eld and forced labora-
tory conditions. Through a sequential collection of fruit from fruit-y infested
mango orchards, fruit were dissected for eggs and larvae. At the same time,
fruit were stored and held for puparia emergence. In addition fruit were
exposed in cages to wild fruit ies and traps were placed to verify the pres-
ence of fruit ies. Estimating the time that Tommy Atkins fruit remain
immature and therefore non-hosts for fruit ies, may provide a window for
mango exports in some fruit y-infested areas. Lux et al. (2003) also mention
that small growers tend to harvest fruit before maturation as a strategy to
evade fruit infestation.
Chemical control
Mango plantations account for major insecticide use in the tropics (Cunning-
ham, 1984). From the late 1960s to date, the conventional control of fruit ies
was through toxic bait sprays that combine proteinaceous bait (e.g. hydroly-
sed protein) with an insecticide (Lpez et al., 1969; Soto-Manati et al., 1987;
Mangan et al., 2006; Mangan and Moreno, 2007). For many years the insecti-
cide of choice has been malathion (Peck and McQuate, 2000; Burns et al.,
2001). Fruit ies are highly susceptible to any insecticide, and other com-
pounds have also been widely used. For example, Singh (1991) reported that
5% aldrin dust, when mixed in soil, provided the highest residual toxicity to
falling mature larvae (23.4% after 15 days), compared to BHC endosulfan
and quinolphos. Vergherse et al. (2004), working on the control of B. dorsalis
in India, used a rotation of fenthion (0.05%), deltamethrin (0.0028%), carbaryl
(0.2%) and dimethoate (0.06%) to reduce the risk of resistance development.
Yee (1987) concluded that weekly applications of malathion for 3 months can
also provide effective control.
Since the late 1990s, there has been a concerted effort to nd environmen-
tally friendly alternatives to malathion (Peck and McQuate, 2000). Cyromaz-
ine, imidacloprid (organochlorinated compound), spinosad (bacteria-derived
insecticide) and phototoxic dyes (Phloxine B) have been successfully tested
against various fruit y species (Daz-Fleischer et al., 1996; King and Hen-
nessey, 1996; Peck and McQuate, 2000; Vargas et al., 2002; Liburd et al., 2004;
McQuate et al., 2005). Despite their success, and as is typical with insecticides
intensively applied on a large scale, resistance has already been documented
in the case of spinosad (Wang et al., 2005; Hsu and Feng, 2006) or collateral
damage (i.e. negative impact on natural enemies) (Stark et al., 2004). In
Pakistan, application of pesticides caused reduction of fruit y infestations,
but their use has created scale insect problems by eliminating their natural
enemies (Mohyuddin and Mahmood, 1993). Such an effect had been reported
by Ehler and Endicott (1984) with pests of olive, citrus and walnut. Another
recent development with respect to chemical control of fruit ies has been the
renement of the bait-station concept (Mangan and Moreno, 2007).
Pests 325
Without chemical control, the damage from C. capitata as a result of bar-
ren and fertile punctures can be as high as 60%. Control is achieved by aerial
bait spraying, ground cover spraying and spot spraying of trees. In Spain,
chemical control has been achieved by applying organophosphates and
hydrolysed albumen (Khanzada and Naqvi, 1985), but this approach is under
pressure because of environmental concerns. Decisions on when to apply
insecticides are based on the appearance of the rst trapped males (trimedlure-
baited traps are used). When applying insecticides directly, dimethoate (0.1%)
and fenthion (0.15%) are used (Khanzada and Naqvi, 1985). Bait sprays are
based on naziman (1:1 protein hydrolysate: malathion/4 l water; Wysoki
et al., 1993). Removal of fallen fruit can also prevent build-up of Mediterranean
fruit y populations.
Pestiferous Anastrepha spp. are susceptible to most insecticides (Shaw
and Spisshakoff, 1958; Shaw, 1961). Bait sprays applied from the ground and
from the air are successful, but they can cause environmental damage, surges
of secondary pest populations and reductions in parasitoid populations
(Lpez et al., 1969; Soto-Manitiu et al., 1987). In Peru, control measures against
Anastrepha in mango begin when McPhail trap catches average two adults/
trap/week (Herrera and Vias, 1977). In Mexico, control starts when the fruit
is 85-days-old and is suspended 2 weeks before harvest (Cabrera et al., 1993).
In Costa Rica, dipterex and malathion are sprayed weekly and reduce mango
damage up to 40% (Soto-Manitiu et al., 1987). In Brazil, malathion with protein
and sugarcane bagasse is used (Carvalho and De Queiroz, 2002). In Ecuador,
Arias and Jines (2004) recommend a spray of malathion (1%) with protein (4%)
once the fruit y population reaches 0.14 fruit ies/trap/day (FTD).
Biological control
PARASITOIDS. Classical biological control and repeated augmentative releases
of mass-reared parasitoids have been used to suppress Anastrepha, Ceratitis
and Bactrocera populations (Wharton, 1978; Sivinski, 1996; Sivinski et al.,
1996, 1997, 2000; Montoya et al., 2000). In Florida USA, Mexico, Costa Rica,
Brazil, Colombia and Peru, parasitoid species (i.e. Diachasmimorpha longicau-
data (Ashmead), Fopius vandenboschi (Fullaway) and Aceratoneuromyia indica
(Silvestri)) have been imported and released for the control of A. suspensa, A.
ludens and A. fraterculus (Ovruski et al., 2000). Despite the widespread use of
exotic parasitoids over the past 80100 years, the current trend is to resort to
native species as they pose less of an environmental threat to local fauna
(Garca-Medel et al., 2007; Aluja et al., 2009).
Use of parasitoids with mango is hindered by the fact that fruit are very
large and therefore provide larvae a refuge from parasitism (Lpez et al.,
1999). As a consequence, Aluja (1993) and Montoya et al. (2000) recommended
that parasitoids should be released outside the mango orchards to attack y
larvae in their much smaller native hosts and thereby signicantly reduce the
size of populations entering mango orchards.
Several parasitoids, for example Opius fullawayi (= Diachasmimorpha
fullawayi (Silvestri)), Diachasmimorpha kraussi, D. Fullaway, D. tryoni (Cam-
eron), Opius bellus Gahan, Biosteres longicaudatus Ashmead (= D. longicaudata),
J.E. Pea et al. 326
B. tryoni (Couron) (= D. tryoni (Cameron)) and Biosteres oophilus Fullaway
(= Fopius arisanus (Sonan)), parasitize C. capitata (Beardsley, 1961; Wharton
and Marsh, 1978). Bess et al. (1961) reported that the most important parasi-
toids collected from C. capitata in Hawaii USA were F. vandenboschi, B. oophilus
(= Opius oophilus) (= F. arisanus) and B. longicaudatus. In Brazil, mainly Doryc-
tobracon areolatus (Szpligeti) (97%) and D. longicaudata (3%) parasitize fruit
y larvae attacking mango (Carvalho and De Queiroz, 2002). In Kenya, Ghana,
Tanzania, Uganda and Cote dIvoire, the most important parasitoids obtained
from Ceratitis spp. emerging from mangoes were D. fullawayi, Fopius cauda-
tus (Szpligeti), Psyttalia cosyrae (Wilkinson) and Tetrastichus giffardianus Sil-
vestri (Lux et al., 2003). In Mexico and other parts of Latin America, the most
common parasitoids attacking fruit ies that infest mangoes (A. obliqua, A.
ludens, A. pseudoparallela, A. turpiniae) are D. areolatus, Doryctobracon brasilien-
sis (Szpligeti), Doryctobracon crawfordi (Viereck), Doryctobracon uminensis
(Lima) and Utetes anastrephae (Viereck) (Lpez et al., 1999; Ovruski et al., 2000;
Zucchi, 2000). In Pakistan, the parasitoids attacking B. zonata include Opius
longicaudatus (= D. longicaudata), Dirhinus giffardii Silvestri, and Bracon sp.; O.
longicaudatus (= D. longicaudata), D. giffardii and Spalangia grotiusi Girault
were reported to attack B. dorsalis, albeit in small numbers (Syed et al., 1970).
Microbial control
Use of pathogens/disease agents (fungi, bacteria, nematodes) has been
attempted with varying degrees of success. For example, Metarhizium
anisopliae has been evaluated in small-scale mango orchards in Kenya using
bait stations laced with the pathogen. Results do not show differences
between use of pathogens and use of insecticides (malathion) (Lux et al.,
2003). Lezama-Gutierrez et al. (2000) also evaluated isolates of M. anisopliae
against larvae of A. ludens. They suggested that M. anisopliae can cause a
2243% reduction in adult emergence, depending on the soil where the lar-
vae pupariates. De la Rosa et al. (2002) evaluated the fungus Beauveria bassi-
ana (Bals.) under laboratory conditions and concluded that highest mortality
was achieved at the adult stage, while Dimbi et al. (2003) reported on the
pathogenicity of M. anisopliae and B. bassiana on different species of Ceratitis.
Poinar and Hislop (1981), Lindegren and Vail (1986) and Toledo et al. (2006)
have investigated the use of various nematodes, Heterorhabditis bacteriophora,
Heterorhabditis heliothidis (Khan, Brooks and Hirschmann) and Steinernema
feltiae Filipjev, against Anastrepha, Bactrocera and Ceratitis. Finally, Robacker
et al. (1996) and Toledo et al. (1999) have tested various strains/isolates of
Bacillus thuringiensis (Berliner) against larvae of A. ludens, A. obliqua and A.
serpentina. For additional details on microbial control of pestiferous fruit ies,
we recommend the recent review by Dolinski and Lacey (2007).
Predators
In addition to parasitoids, pathogens and nematodes, ants have been used to
control fruit ies in mango orchards. Peng and Christian (2006) used the
weaver ant, Oecophylla smaragdina (Fabricius) for control of the Jarvis fruit y,
B. jarvisi, in mango orchards in Australia. Van Mele et al. (2007 and references
Pests 327
therein) in Benin used an African weaver ant (Oecophylla longinoda). Aluja
and colleagues (Aluja et al., 2005) investigated the potential of ants as possi-
ble biological control agents in various tropical orchards and backyard gar-
dens in which mango trees were growing with other fruit trees.
Cultural fruit y control
Fruit bagging is one of the best solutions to prevent fruit y attack of mango
and other tropical fruits (Aluja, 1996; Pea et al., 1999; Paderes and Orden,
2004). Success with mangoes can be quite high, but Bondad (1985) demon-
strated that bagging materials do not always resist the effect of rain/wind.
Therefore, while bagged mangoes tend to produce a greater amount of mar-
ketable fruit than those not bagged, more research is needed to determine the
type of bags to use for different mango varieties and the best time to bag fruit
(Love et al., 2003).
Jirn (1995) reported that A. obliqua populations could be reduced by
increasing planting distances in order to reduce RH and increase solar radia-
tion within orchards. In India, cultural control practices include removal of
fallen fruit and inter-tree ploughing and raking (followed by insecticide
cover sprays). Such practices can reduce fruit y infestation between 77%
and 100% (Verghese et al., 2004).
A cultural practice that can impinge on the success of management pro-
grammes is the widespread use of potassium nitrate (KNO
3
) sprays to accel-
erate and synchronize owering of mango trees. As a result, fruit harvests
can be advanced and synchronized. Such a procedure can, under certain cir-
cumstances, help control fruit ies, but can also exacerbate the problem. For
example, in the case of the Mexican fruit y (A. ludens), a notorius pest of
citrus that also attacks mangoes, advancing the mango harvest offers ideal
conditions for adult pests to move from citrus groves to mango orchards.
This would not occur if mango trees owered naturally since fruit ripen sev-
eral months after the citrus harvest (Martin Aluja, personal observation).
Host resistance
Yee (1987) reported that B. dorsalis does not attack all mango cultivars to the
same extent. The most susceptible cultivars in Hawaii USA are Hawaiian,
Pirie and Sandersha. Singh (1991) indicated that the frequency of Bactro-
cera injury to physiologically mature fruit of Dashehari ranged from 3.6 to
10%, while in fully ripe fruit the frequency of injured fruit ranged from 10 to
25.9%. Highest damage was reported in fully ripe fruit of Mallika followed
by Totapari.
Susceptibility of different mango cultivars to attack by A. obliqua was
measured by Carvalho et al. (1996) who observed that Espada showed no
infestation by A. obliqua, whereas Carlota was highly infested. In this study,
the survival of adults of A. obliqua was lower when the larvae were fed on
Espada compared to Carlota. Furthermore, Espada had an adverse effect
on the longevity of A. obliqua females, possibly due to the presence of toxic
substances (Carvalho and De Queiroz, 2002) or absence of essential nutri-
ents. Jirn and Soto-Manitiu (1987) also observed that susceptibility of
J.E. Pea et al. 328
mangoes to A. obliqua differed among cultivars. Rosinha, Coquinho and
Espada were resistant to A. obliqua attack, whereas Smith and Pope were
highly susceptible. According to Joel (1980), mangoes contain resin ducts in
the exocarp that confer protection against the vertical movement of the ovi-
positor and larval movement. Other studies have shown that resistance is
related to degree of maturity (Daz-Fleischer and Aluja, 2003; Aluja and
Mangan, 2008); immature mango fruit are less susceptible to A. suspensa
than mature mangoes when infested articially (Hennesey and Schnell,
2001). In Brazil, use of gibberellic acid (GA
3
)

reduces the susceptibility of
Tommy Atkins mangoes to attack by C. capitata based on articial delay of
fruit maturity (de Macedo, 1988). Differences on attack by A. ludens to mango
might be inuenced by volatiles from green or yellow fruits (Garcia-Ramirez
et al., 2004).
Quarantine treatments
Quarantine treatments have been reviewed by Johnston and Hofman (Chap-
ter 15, this volume). Several quarantine treatments have been developed for
harvested mangoes: irradiation, hot water or hot water followed by immer-
sion cooling are widely used (Sharp et al., 1988, 1989a, b, c; Hallman and
Sharp, 1990; Nascimento et al., 1992; Mangan and Sharp, 1994; Mangan and
Hallman, 1998; Shellie and Mangan, 2002a, b; Bustos et al., 2004; additional
references in reviews by Mangan and Hallman, 1998 and Follet and Neven,
2006).
Mango seed weevils
The mango seed weevil, Sternochetus mangifereae (Fabricius), and the mango
pulp weevil, Sternochetus frigidus (Fabricius) (Coleoptera: Curculionidae) are
important pests of mango (Plates 5759). Quarantine restrictions prevent the
export of fresh weevil-infested mangoes into uninfested areas. The esh of
ripe fruit is damaged when mango seed weevil adults emerge from the seed,
and weevil-damaged seed may limit plant propagation in nurseries and
orchards (Johnson, 1989). Early fruit drop may be caused by severe weevil
infestations (Subramanian, 1925). The mango seed weevil occurs from India
through South-east Asia to Australia, on tropical Pacic Islands, in parts of
Africa, in the Caribbean region and in northern South America (Balock and
Kozuma, 1964; Shukla and Tandon, 1985; Johnson, 1989; Schotman, 1989).
Biology
Srivastava (1998) reports that S. mangiferae is a greyish brown weevil, 8 mm
long and about 4 mm wide; its habits are nocturnal, usually feeding and ovi-
positing at dusk. After emergence, adults enter diapause and the duration
varies with the geographic range (Schotman, 1989). According to Shukla and
Tandon (1985), all adults emerging in southern India during June enter dia-
pause from July until late February in the following year. The beginning and
the end of diapause seem to be associated with long-day and short-day
Pests 329
photoperiods, respectively (Balock and Kozuma, 1964). The mango seed
weevil produces only a single generation each year. In Tamil Nadu, India,
adults feed on leaves and tender mango shoots in March and April (Subra-
manian, 1925). Shukla and Tandon (1985) report that females began oviposit-
ing 34 days after mating when fruit reaches a marble-size. Oviposition
varied from 3 to 5 weeks (Subramanian, 1925; Shukla and Tandon, 1985;
Hansen et al., 1989). The female uses its snout to make a cavity in the fruit,
lays a single egg and then covers it with a secretion (Pradhan, 1969).
According to Srivastava (1998), about six larvae can be found within a
mango seed. Generally, only a single larva completes development within
each fruit. Larval development and pupation occurs within the seed. Adults
are formed 1 week later; however, adults generally emerge from the seed
c.12 months after fruit drop (Balock and Kozuma, 1964). The weevils over-
season under bark and stone walls, where they remain dormant until the
next owering season (Van Dine, 1906; Balock and Kozuma, 1964; Shukla
and Tandon, 1985).
Sternochetus frigidus attacks Mangifera indica, M. foetida and M. sylvatica.
The weevil occurs in Bangladesh, Brunei, India, Indonesia, Myanmar, Paki-
stan, Papua New Guinea, the Philippines, Singapore and Thailand (CAB
International, 2003). Sternochetus frigidus lays eggs on mango fruit with a
minimum diameter of 6 cm (De and Pande, 1988). Newly hatched larvae tun-
nel directly through the fruit pulp. Pupation takes place in a brown cocoon
within a chamber adjacent to the kernel. The weevils leave the ripe fruit
through a hole in the peel. This pest is likely to survive storage and transpor-
tation (Australian Government, 2004). Reproductively immature adult wee-
vils overwinter inside seed or other protective places from May until February
in India (De and Pande, 1988).
Sampling
Shukla et al. (1988) reported the intra-tree distribution of eggs of S. mangiferae
on Baganpalli mango; the highest number of eggs per fruit occurred on fruit
in the lower region of the tree. With increasing tree height, egg deposition on
fruit decreases. No statistical differences on fruit infestation were observed
on north, south, east or west directional quadrats of the tree. Eggs were
deposited in the lower region of the fruit rather than the pedicel. Weevils
enter diapause in crevices in the tree trunk. Most of the weevils (87%) are at
a height of 02 m in the trunk compared to 7% at 24 m and only 4% above
4 m. Emery (2002) considered that since both the mango pulp weevil (S. frigi-
dus) and the mango seed weevil (S. mangiferae) infest fruit at an early stage,
any fruit is a viable sample; however, as infested fruit all ripen precociously,
the sensitivity of surveys is enhanced by seeking out nearly ripe and fallen
fruit prior to harvest. If the survey coincides with the mango harvest, rejected
or fallen fruit should be inspected. Fruit should be sampled by longitudinal
dissection of fruit through the seed to expose the kernel. If the fruit is ripe, it
should be struck along the longitudinal axis with a hammer, and the seed
should be opened with pliers. The random sampling of 600 fruit from ran-
domly selected properties in each area provides a 9% chance of detecting a
J.E. Pea et al. 330
0.5% infestation of fruit. Sample size can be determined from the following
formula:
Probability of > 1 infested fruit = 1 probability of no infected fruit in
total sample = 1 (1 0.5%)
600
= 1 (1 0.005)
600
= 1 (0.995)
600
= 95%
Economic damage
Follett and Gabbard (2000) report that germination rates for infested seed of
polyembryonic Common are equal to those of uninfested seed. Germina-
tion is signicantly reduced for infested seed of monoembryonic Haden
compared with uninfested control seed, although germination of infested
seed was >70%. Direct feeding damage to the pulp was found in only 0.11%
of 3602 mango fruit, which suggests that S. mangiferae is a less serious pest of
mangoes than previously considered.
Cultural control
Field sanitation, i.e. the removal of all fallen fruit and seed, is very labour
intensive, and demands complete removal and disposal of fallen fruit. This
procedure has been inconsistent in demonstrating pest control. In India, eld
sanitation reduced infestation of the mango nut weevil, Sternochetus gravis
(Fabricius), by only 22% (De and Pande, 1987). In Hawaii USA, eld sanita-
tion failed to reduce infestation rates (Hansen and Amstrong, 1990).
Chemical control
Various insecticides have been evaluated for controlling adult weevils, par-
ticularly during oviposition (Balock and Kozuma, 1964; Shukla and Tandon,
1985). The most effective control was provided by the organophosphate fen-
thion, which reduced infestation to <17%. In another eld test, the pyrethroid
deltamethrin and the carbamate carbaryl were most effective, both resulting
in <15% infestation rates. Spot application of diazinon on tree trunks was
recommended based on cost, efciency and least environmental damage.
Verghese et al. (2004) reported that commercially available azadirachtin was
not effective for management of S. mangiferae in India.
Resistant cultivars
Mango cultivars resistant to the mango weevil would be benecial. Potential
mechanisms of resistance are seedless cultivars, those that form seed early or
those that fruit off-season. Most cultivars grown in Hawaii and India are
equally susceptible (Bagle and Prasad, 1984; Hansen et al., 1989), although
Itamaraca has shown some resistance (Balock and Kozuma, 1964).
Biological control
The mango weevil has few natural enemies. Parasitoids are unknown, prob-
ably because of the concealed nature of most life stages. Adults may be sus-
ceptible to predation by ants, rodents, lizards and birds (Hansen, 1993). A
baculovirus has been reported that affects the larvae of S. mangiferae (Shukla
et al., 1984).
Pests 331
Mango fruit infested with seed weevil do not show any visible external
symptoms and cause considerable quality control problems and economic
loss to the mango-processing industry as well as restriction in export of fresh
fruit. A non-destructive X-ray inspection method has been developed to
detect weevil-infested fruit. X-ray radiographs of infested mangoes show
dark areas in the seed corresponding to disintegrated kernel tissue as a con-
sequence of feeding by developing grubs. Non-infested mangoes show a uni-
formly light-grey area representing healthy kernel. There is a close agreement
between fruit showing weevil infestation based on their X-ray images and
physical examination of cut fruit, indicating the reliability of the technique.
X-ray imaging has good potential for application in the processing industry
and the export trade as a quality control measure (Thomas et al., 1995).
Mango seed borer (Lepidoptera: Pyralidae)
Distribution and biology
The red banded mango caterpillar or mango seed borer, Deanolis sublimbalis
Snellen, also referred to as Noorda albizonalis Hampson (Waterhouse, 1998), is
an important pest of mangoes in the Philippines (Anonymous, 1984), India
(Zaheruddeen and Sujatha, 1993), Vietnam (Nguyen et al., 1998; van Mele et al.,
2001), China (Li et al., 1997), Thailand, Indonesia and Papua New Guinea
(Cunningham, 1984). The oval white eggs are laid in groups at the fruit apex
and take 34 days to hatch. The larvae develop through ve instars in 1420
days and they pupate in cocoons in the soil. The period from egg to adult
requires 2840 days. The insect apparently prefers mango, but M. odorata and
M. minor have also been recorded as hosts. Adults are generally nocturnal
and spend most of the day under leaves on the tree. A shorter developmental
period has been observed when larvae develop on pulp rather than in seed
of Carabao fruit, although those reared on the seed were larger and lived
longer (Waterhouse, 1998).
Damage
Mango fruit in all stages of development are susceptible to attack (Water-
house, 1998). The rst larval instars feed on tissues beneath the skin, and
bore through the mango pulp to the seed, which is consumed. Up to 11 larvae
have been recorded in a single fruit, but usually there is only one. Infested
fruit split and rot, and fall to the ground (Anonymous, 1984). In the Guimaras
Islands, the Philippines, Golez (1991) recorded 12.5% fruit infestation and in
serious outbreak years, 4050% yield reductions are possible. Waterhouse
(1998) considered that since D. sublimbalis is capable of causing such levels of
damage, it might be a more important pest of mangoes than has generally
been realized. It may have been overlooked as a pest or has recently spread
to new areas and has become evident as a pest there. Van Mele et al. (2001)
suggested that damage caused by D. sublimbalis in the Mekong Delta has
been wrongly attributed to fruit ies; however, Waterhouse (1998) states that
soon after boring by D. sublimbalis, secondary infestations with fruit ies
J.E. Pea et al. 332
(Bactrocera ferrugineous, B. frauenfeldi) occur, together with infections by bac-
teria and fungi.
Biological control
According to Waterhouse (1998) no parasitoids were detected in Java, Indo-
nesia. However, in the Guimaras Islands of the Philippines, the vespid wasp,
Rychium attrisimum, preys on the larvae as they exit the fruit to pupate. Lar-
vae are used to stock the wasps nests as food for their young. The egg para-
sitoids Trichogramma chilonis Ishii and Trichogramma chilotreae attack the pest
in Luzon (Golez, 1991). Leefmans and van der Vecht (1930) reported that an
entomopathogenic fungus infected the larvae in Indonesia.
Monitoring and control
Infested fruit can be detected by the presence of a dark-brown ring and cat-
erpillar frass at the point of entry (CAB International, 2003). Mango fruit
become susceptible to the seed borer c.60 days post-induction, and insecti-
cide applications should commence then. Further sprays at 75, 90 and 105
days post-induction are required to fully protect the fruit. The most effective
chemicals are deltamethrin and cyuthrin (Golez, 1991). Infested fruit should
be removed from trees before the larvae can leave them to attack neighbour-
ing fruit; fruit should be wrapped in protective bags at 5565 days after pol-
lination, and fallen fruit should be destroyed (Anonymous, 1984).
Other Lepidoptera that can attack fruit have been reported in India and
the Philippines (Australian Government, 2004). The pomegranate fruit borer,
Deudores isocrates (Fabricius) (Lepidoptera: Lycaenidae), which is also a pest
of loquat, lychee, guava and pear, could attack mango by laying single eggs
on shoots; the emerging larva bore into the fruit (Srivastava, 1998). In the
Philippines, the larvae of the cocoa tussock moth, Orgyia postica (Walker)
(Lepidoptera: Lymantriidae), regularly attack leaves of cocoa, but its larva
also attack mango fruit and panicles (Fasih et al., 1989).
Fruitspotting bugs (Hemiptera; Coreidae)
The yellowish green coreid bugs, Amblypelta lutescens lutescens (Distant) and
Amblypelta nitida Stl occur along the coast of Queensland, Australia, and
attack most of the tropical and subtropical fruit crops there (Waite and Huwer,
1998). They prefer to feed on young, green fruit, but A. l. lutescens also dam-
ages the terminals of a number of hosts. In tropical north Queensland, A. l.
lutescens is the dominant species and feeds on young fruit causing black
lesions to develop and the fruit to fall. It also feeds on the terminals and leaf
petioles, causing wilting and dieback (Cunningham, 1989). In the subtropical
south, both species attack mango, but A. nitida connes its attention to the
fruit, while A. l. lutescens also attacks fruit and terminal growth (G.K. Waite,
1995, unpublished results). The bugs breed in natural rainforest areas, and
y into the orchards to feed on the fruit and terminals. Female bugs lay
individual, opalescent green eggs under the leaves. There are ve nymphal
instars and a generation takes c.40 days.
Pests 333
The main predators of fruitspotting bugs are spiders, particularly mem-
bers of the family Thomisidae. Several species of egg parasitoids have been
recorded. In north Queensland, Ooencyrtus sp. (Encyrtidae), Anastatus sp.
(Eupelmidae) and Gryon sp. (Scelionidae) parasitized 37.591.6% of eggs of
A. l. lutescens (Fay and Huwer, 1994). In south Queensland, Anastatus sp. and
Gryon meridianum (Dodd) parasitize eggs of A. nitida and A. l. lutescens to a
similar extent (Waite and Petzl, 1994).
Because fruitspotting bugs continuously migrate into orchards, more
than one insecticide spray may be required to protect the young fruit. How-
ever, the fruit are safe from attack once they are c.50 mm long, and two or
three sprays of endosulfan at intervals of 2 weeks are generally sufcient to
protect them.
The coconut bug, Pseudotheraptus wayi Brown, was rst recorded on man-
goes in South Africa in 1977, and now also attacks guavas, pecans, macada-
mias, avocados and loquats. It causes damage similar to that of Amblypelta
spp. (De Villiers, 1990).
Helopeltis spp. (Miridae) are minor pests of mango, cashew and cacao in
the Philippines and in northern Australia, where they feed on fruit and cause
supercial corky blemishes. Insecticides are used to control them, but in the
Philippines, bagging is also effective (Anonymous, 1984).
Plant bugs within the Lygaeidae and Pyrrhocoridae, i.e. the Indian milk-
weed bug, Spilostethus pandurus (Scopoli) and the red cotton bug, Dysdercus
koenigri (Fabricius), can injure fruit, inorescences and leaves of mangoes in
India (Australian Government, 2004). However, D. koenigri is an important
pest of cotton rather than mango (Schaefer and Ahmad, 2000), while S. pan-
durus feeds preferentially on members of the Asclepediaceae (i.e. Calotropis)
(Sweet, 2000).
A hymenopterous parasitoid complex attacking the eggs of the banana-
spotting bug, A. l. lutescens, was rst reported in north Queensland. It includes
an Anastatus sp. (Eupelmidae), Ooencyrtus sp. nov. (Encyrtidae) and a Gryon
sp. (Scelionidae), and is similar to other complexes known to attack eggs of
related coreids in Africa, Indonesia and Papua New Guinea. Parasitism
ranged from 37.591.6% in eggs collected at three sites from orange jessa-
mine, Murraya paniculata (L.) Jack. Anastatus sp. was the dominant parasitoid
(Fay and Huwer, 1994).
Thrips
Grove et al. (2001) reported that in South Africa, the citrus thrips, Scirtothrips
aurantii Faure and the red banded thrips Selenothrips rubrocinctus (Giard) are
the only thrips that caused lesions on fruit (Plates 6063). However, Grove
et al. (2001) considered S. aurantii to have more economic importance than S.
rubrocinctus. Grove and Pringle (2000), using a two-stage sampling system to
determine population levels of S. aurantii, showed that S. aurantii has a clumped
distribution, and recommended that 50 fruit per orchard should be examined
in order to obtain accurate population estimates for pest management.
J.E. Pea et al. 334
Blossom pests
Midges, caterpillars, hoppers, thrips and mites are the most important pests
attacking mango inorescences.
Midges
The mango gall midge or mango blister midge, Erosomya mangiferae Felt, is a
major pest, destroying owers and up to 70% of set fruit (Plate 64). It was
rst described by Felt (1911) in St Vincent (West Indies). Barnes (1948) recog-
nized nine gall midges from mango; two of these, Asynapta sp. and E. man-
giferae, are from the West Indies. Butani (1979) reported ve cecidomyiid
species on mango blossoms, including Erosomya indica (Grover and Prasad).
Dasyneura mangiferae (Felt) was reported in Hawaii USA (Vannire et al.,
2004). In recent times, Gagne and Etienne (2006) reported the species Gephy-
raulus mangiferae (Felt), n.comb. infesting mango owers on the island of
Guadeloupe, French West Indies. Male adults of E. mangiferae are 1.61 mm
and females 1.32 mm long. Eggs are deposited in folds between sepals and
petals of ower buds. The larval stage has four instars. Young larvae are
cream coloured and late instar larvae are yellowish. Larval feeding prevents
ower opening and consequently development of the fruit does not occur.
Infested buds develop as long pointed galls, in which pupation occurs (Van-
nire et al., 2004). Studies of population uctuation of Erosomya sp. have been
conducted in India by Grover (1986a), who reported that emergence of adults
was higher at 24C and 6082% RH compared to lower temperatures and
RH. Abbas (1985) described systematic surveys to determine the percentage
of infestation of E. indica, and showed that infestation follows a negative
binomial infestation. The midge infests the newly emerged panicles by ovi-
positing at bud burst stage, and the rst instar maggots bore into the grow-
ing panicle. The second generation then infests very young fruit, which drop
before the marble stage. Sampling of mango midges needs to include affected
tissue, different trapping devices, pheromones, etc. On citrus, use of coloured
sticky traps placed in the tree canopy provides a more efcient method of
sampling the citrus midge, Prodiplosis longila Cagn, than ground emergence
traps and collection of larval samples (Pea and Duncan, 1992).
In a survey of parasitoids of cecidomyiid pests of mango in India, Grover
(1986b) reported that Platygaster sp., Systasis sp. and Eupelmus sp. were asso-
ciated with Dasineura sp., and Tetrastychus sp. was associated with E. indica.
An external parasitoid, the pteromalid, Pirene sp., attacked Procystiphora
mangiferae (Felt). Predators of the cecidomyiids include Formicai sp., Oeco-
phila sp. and Camponotus sp.
Mango hoppers
Approximately 18 species of leaf hoppers have been reported as pests of
mango. Of these, Idioscopus clypealis Leth., Idioscopus niveosparsus Leth., Idiosco-
pus magpurensis Pruthi and Amritodus atkinsoni Leth., are important (Virakta-
math and Viraktamath, 1985; Viraktamath, 1997; Fletcher and Dangereld,
2002) (Plate 65). The females deposit their eggs in panicles or midribs of tender
Pests 335
leaves. The adults and nymphs preferentially feed on young leaves and ow-
ers or shoots, and excrete honeydew upon which sooty mould develops
(Ahmed et al., 1981). This interferes with photosynthesis, adversely affecting
plant growth and yield (Godase et al., 2004). Affected inorescences turn
brown, become dehydrated and fruit set does not occur.
There has been no systematic study of the biology of most of the leaf
hoppers that attack mango; however, biology of A. atkinsoni, I. clypealis and I.
niveosparus has been studied by Sohi and Sohi (1990). Both A. atkinsoni and I.
niveosparsus are multivoltine. In A. atkinsoni, the egg, nymphal (ve instars)
and adult stages require 79, 1517 and 34 days, respectively (Patel et al.,
1977). Development from egg to adult is normally complete in 2530 days.
There can be between one and six generations of A. atkinsoni in different areas
of India. In Pakistan there are four to ve generations in the Central Punjab
(Mohyuddin, unpublished data). In the Philippines, I. clypealis is reported to
have one to four generations, whereas it has ve or six generations in India.
Idioscopus nagpurensis is univoltine. In Pakistan, it normally oviposits in
mango inorescences during March. Nymphs feed on inorescences during
March and April. From May to February of the following year, only aestivat-
ing adults are found (Mohyuddin, unpublished data). Most of these species
are quite fecund. Amritodus atkinsoni reportedly lays 200 eggs during its life-
time (Rahman, 1940) and I. clypealis lays 100190 eggs in the Punjab (Husain
and Pruthi, 1924). Amritodus atkinsoni eggs are laid in the midribs of tender
leaves, ower buds and inorescences (Babu et al., 2002). Idioscopus niveospar-
sus lays c.238 eggs in 9 weeks under laboratory conditions (Mohyuddin,
unpublished data). Mohyuddin and Mahmood (1993) reported that A. atkin-
soni and I. niveosparsus occur in upper portions of mango trees during differ-
ent times of the year. Amritodus brevistilus and I. niveosparus populations
increase from February to peak in MarchApril in Sri Lanka, while peaks of
I. clypealis occur in March and September (Kudagamage et al., 2001). Idiosco-
pus clypealis populations peak in south-eastern India during March and April
(Tandon et al., 1983). Idioscopus nivesoparus and I. clypealis peaks coincided
with major and minor owering periods while population peaks for A. bre-
vistilus coincide with the occurrence of vegetative ushing (Kudagamage
et al., 2001).
Azizur Rahman and Singh (2004) demonstrated that A. atkinsoni popula-
tions on panicles of Langra mango were negatively correlated with high
RH; whereas no signicant relationships were observed with rainfall, sun-
shine and wind velocity.
SAMPLING. Very few sampling studies have been reported for hoppers on
mango. A sequential sampling plan for mango hoppers was recommended
by Verghese et al. (1985) in India. Mohyuddin and Mahmood (1993) reported
sampling by direct visual examination: A. atkinsoni and I. niveosparsus were
found on upper portions of mango trees during different times of the year.
They moved to the lower parts of the stems and the leaves during summer.
Tandon et al. (1989) reported that distribution of I. niveosparsus was aggre-
gated and was best explained by Iwaos patchiness regression. To assess
J.E. Pea et al. 336
damage, they recommended a sampling size of 5998 panicles/tree. Verghese
et al. (1985) developed a sequential sampling plan classifying infestations of
adults and nymphs of I. clypealis as light, moderate and severe.
BIOLOGICAL CONTROL. Several natural enemies have been described from west
and South-east Asia. Mohyuddin and Mahmood (1993) reported the egg par-
asitoids, Gonatocentrus sp., Miurfens sp. nr. mangiferae Viggiani and Hayat,
Centrodora sp. nr. scolypopae Valentine, Aprostocetus sp. and Quadrastichus sp.,
and the adult ectoparasitoid Epipyrops fuliginosa Tames in Pakistan. Fasih and
Srivastava (1990) reported that Aprostocetus sp., Gonatocerus sp. and Polynema
sp. parasitize eggs. Five species of predators, including Chrysopa lacciperda
(Kimmins), Mallada boninensis (Okomote), Bochartia sp. and two unindenti-
ed species (one each of Mantidae and Lygaeidae) prey on nymphs (Fasih
and Srivastava, 1990). In India, Sadana and Kumari (1991) studied the ef-
cacy of the lyssomanid spider, Lyssomanes sikkimensis on I. clypealis. Classical
biological control of mango hoppers has not been attempted. Whitwell (1993)
described four genera of parasitoids from Dominica, the most common being
Aprostocetus sp., followed by Platygaster sp., Synopeas sp. and Zatropis sp.
Peng and Christian (2005a, b) reported that the weaver ant, Oecophylla smarag-
dina (Hymenoptera: Formicidae) is an efcient biocontrol agent of I. nididulus
in northern Australia. The entomopathogens, Verticillium lecanii (Zimmer-
man) Viegas, Beauveria bassiana Balsamo (Vuillemin) and Isaria tax, infect
I. clypealis in India (Kumar et al., 1993; Srivastava and Tandon, 1986) while
the effectiveness of Metarhizium anisopliae var. anisopliae was tested under
laboratory conditions against A. atkinsoni (Vyas et al., 1993).
CHEMICAL CONTROL. Several pesticides have been tried for controlling mango
hoppers (Tandon and Lai, 1979; Yazdani and Mehto, 1980; Shah et al., 1983;
Shukla and Prassad, 1984; Islam and Elegio, 1997; Kudagamage et al., 2001).
Khanzada and Naqvi (1985) reported that six sprays of fenitrothion/year
were effective for controlling mango hoppers in Pakistan. Nachiappan and
Baskaran (1986) tested eight insecticides: phasalone, endosulfan, carbaryl,
penthoate, fenitrothion, monocrotophos, quinalphos and phosphamidom.
Endosulfan provided the best control when spraying was done 1 week after
owering and then 14 days later. Mohyuddin and Mahmood (1993) reported
that monitor applied at 5 m on tree trunks and leaves in May provided control
of mango hoppers. Jhala et al. (1989) considered that sprays of carbaryl during
the off-season maintained the hopper population at low-density levels.
Godase et al. (2004) demonstrated that sprays of 0.05% monocrotophos at
the rst panicle emergence and a second spray 15 days later are essential to
prevent yield loss. Kudagamage et al. (2001) found that imidacloprid (Admire
SL 200) controlled mango hoppers if applied just after owering and again
10 days later.
Lepidoptera
The lepidopteran ower feeders are the second most important inorescence
pests of mango. Geometrids, for example Chloropteryx glauciptera Hampson and
Pests 337
Oxydia vesulia (Cramer) infestation was reported in Dominica by Whitwell
(1993). Infestations increase during the owering season, averaging three
larvae/inorescence (87% infestation) to c.100% infestation later in the ow-
ering season. Eggs of the noctuidae Penicillaria jocosatrix Guene are laid pre-
dominantly on or near the inorescences or new leaves. The microlepidoptera
complex attacking mango in Florida USA consists of Pococera attramentalis
Lederer, Pleuroprucha insulsaria (Guene) (Plate 66), Platynota rostrana
(Walker), Tallula spp. and Racheospila gerularia (Hbner) (Plate 67). Most of
the damage to inorescences (Plate 68) is caused by P. attramentalis and P.
rostrana. Pococera attramentalis and P. attramentalis are also common pests of
sorghum (Kring et al., 1987) and other tropical fruit trees. The larvae of both
species feed on the axis of the inorescence, petals and ovaries late in the
owering season; dried fallen owers are webbed together and fastened to
ower clusters to form nests (Patel et al., 1977). The Lepidoptera complex
attacking mango owers in Australia consists of several species from the
families Geometridae, Lymanthridae, Noctuidae, Pyralidae and Tortricidae.
In Brazil, Barbosa (2005) and Barbosa et al. (2005) reported Pleuroprucha asth-
enaria (Walker) (Lepidoptera: Geometridae) and Cryptoblabes gnidiella (Milliere)
(Lepidoptera: Pyralidae) affecting mango inorescences. The Pleuroprucha
asthenaria life cycle from egg to adult is 17 days and the C. gnidiella life cycle
is 36 days (Barbosa, 2005). Pleuroprucha asthenaria can cause premature ripen-
ing of injured fruit while C. gnidiella infests inorescences that are compacted
from paclobutrazol applications (Barbosa, 2005).
According to Schreiner (1987), Dipel

reduced caterpillar damage, but


careful monitoring or constant spraying was necessary to prevent signicant
damage. In Brazil, the pesticides Bacillus thuringiensis, trichlorfon and lamb-
dacyhalothrin provided 6659% mortality (Barbosa, 2005). Control with pes-
ticides is mostly unjustiable in Florida USA and Australia, but regular
monitoring is needed for early detection of population increases (Cunning-
ham, 1984; Pea, 1993).
Classical biological control of lepidopteran insects attacking mango in
Dominica was initiated with the introduction of the wasps, Aleiodes sp. and
Euplectrus sp., and the y Blepharella lateralis Macquart. Populations of the
pest were reduced to 25% of pre-release levels; parasitization rates were
2099%, with Euplectrus sp. being the most abundant parasitoid (Nafus,
1991). The parasitoid Macrocentrus prob. delicatus attacks P. attramentalis;
however, the parasitism rate is unknown (Pea, 1993). In Brazil, C. gnidiella is
parasitized by Brachymeria pseudoovata Blanch (Hymenoptera: Chalcididae).
Thrips
The western ower thrips, Frankliniella occidentalis (Pergande) damages ow-
ers and fruit in Israel (Wysoki et al., 1993). The developmental time of F. occi-
dentalis from egg to egg at 25C occurs between 14.8 and 16.65 days. The
duration of development of F. occidentalis from egg to adult is closely related
to environmental conditions, especially temperature. Frankliniella (possibly
cubensis) is present in mango owers during the dry season in Costa Rica,
requiring several applications of systemic insecticides (Jirn, 1993). In Florida,
J.E. Pea et al. 338
the thrips complex consisting of Frankliniella bispinosa (Morgan) and F. kelliae
(Sakimura) is the most frequently observed blossom pest on owers and
causes damage by ovipositing in the panicle and feeding on the oral necta-
ries and anthers, which may result in premature loss of pollen. The biology
of F. bispinosa has been reviewed by Watson (1917), and Sakimura (1981)
studied the taxonomy of F. kelliae. In South Africa, Grove et al. (2001) reported
that Thrips acaciae Thybom, Thrips tenellus Trybom and S. aurantii were the
most abundant species collected from mango owers.
SAMPLING. Thrips density is related to ower phenology and the prevalent
dry season in Florida USA. Pea (1993) suggested that aerial trapping is
superior to ower inspection, but because there is no method for determin-
ing the true population size, the aerial trap method cannot be shown to be an
unbiased estimator. In India, Verghese et al. (1985) determined that the lower
mango canopy is better for sampling, and recommended a sample size of 55
panicles/tree for surveying. Verghese et al. (1988) indicated that distribution
of Thrips palmi (Karny) on mango panicles is better explained by Iwaos
patchiness regression, which indicated an aggregated distribution and
suggested that the lower canopy should be sampled. Sample sizes should
be 55 panicles/tree for control and survey studies, with a 20% error of the
mean and 92 panicles/tree to obtain a lower (5%) percentage error of the mean.
In Florida, Pea et al. (2006a) showed that a cumulative number of 400700
thrips/panicle during 4 weeks causes 3350% yield reduction of Keitt
mangoes.
CHEMICAL CONTROL. The efcacy of different pesticides (acetamiprid, fenpro-
patrin, milbemectin and zeta-cypermethrin, novaluron) was tested by Pea
et al. (2006a) against mango ower thrips. All treatments, except milbemec-
tin, reduced thrips densities 5 days after application of the rst spray. Danitol
and zeta-cypermethrin had the lowest numbers of thrips 12 and 20 days after
the rst spray. All treatments had lower thrips densities compared to the
control at 28, 35 and 43 days after the second application.
BIOLOGICAL CONTROL. Several parasitoids and predators, such as Ceranisus
menes (Walker) in Israel (Rubin and Kuslitizky, 1992) and the predators, Orius
sp., Anystis agilis Banks and Hypoaspis aculifer (Canestrini) (Loomans et al.,
1995), are candidates for biological control of F. occidentalis, while Dasyscapus
parvipennis Gahan is considered to have good potential for the biological con-
trol of ower thrips in Puerto Rico (Bartlet, 1938).
Pests of buds and leaves
Foliage feeders is one of the largest groups of injurious insects of mango.
Pests of mango buds and foliage may cause damage by reducing the photo-
synthetic area of the plant, thereby reducing the quantity of photosynthates
translocated to the fruit. The most destructive mango leaf feeders are thrips,
Pests 339
midges, mites, scales, whiteies, mealybugs, weevils, ants, locusts and cater-
pillars (Jeppson et al., 1975; Bhole et al., 1987; Jadhav and Dalvi, 1987; Tigvatt-
nanont, 1988). Formation of leaf galls in India is caused by the Eurytomidae,
Mangoma spinidorsum (Subba Rao, 1986), but there is little information on
their importance as foliage pests.
Thrips
The Mediterranean mango thrips, Scirtothrips mangiferae Priesner, is a severe
pest of mango in Israel, causing the young leaves to curl along the midrib,
distorting their shape and leading to premature drop (Wysoki et al., 1993).
The twigs of infested shoots are much shorter than those of uninfested ones.
The population of the thrips is low during winter, increases in early spring
and reaches its peak during summer (Wysoki et al., 1993). Yellow sticky traps can
be used for monitoring thrips densities. Ganz et al. (1990) established that an
average population of ten Mediterranean mango thrips per young shoot was the
threshold above which chemical control is required. Efcient control has been
achieved by spray application of uvalinate or acephate (Ganz et al., 1990).
The red banded thrips, Selenothrips rubrocinctus (Giard), is an important
pest of cacao in the Caribbean islands and attacks mango and avocado in
Australia and Florida and Hawaii USA. The adults feed on the underside of
leaves, causing necrosis and subsequent leaf drop. According to Hill (1975),
S. rubrocinctus is only a pest in mango nurseries, and rarely damages mature
trees. Its biology was reviewed by Moznette (1922). Adult thrips are dark
bodied with a red band on the rst abdominal segment. The immature stages
are light orange with abdominal segments one and two and the anal seg-
ments bright red. The population of this species peaks during the dry season
and declines during the rainy season. According to Yee (1987), the thrips are
controlled by malathion (25% v/w).
The weaver ant, O. smaragdina (Hymenoptera: Formicidae) is considered
an effective biological control of S. rubrocinctus in the Northern Territory of
Australia (Peng and Christian, 2004).
Midges
Mango leaves are attacked by different Cecidomyiidae species, especially in
Asia, but also in the Caribbean region. Two genera, Procontarinia Kieffer and
Cecconi and Erosomyia Felt, are particularly associated with mango and all
known species have been reared from mango (USDA, 1981; Schreiner, 1991;
Harris and Schreiner, 1992; Uechi et al., 2002; Gagne and Medina, 2004).
Prasad (1971) described the biology of the main species attacking mango in
India. A new species of gall midge, Procontarinia schreineri Harris, which
attacks mango foliage in Guam, lays eggs on young mango leaves and larvae
develop rapidly over c.5 days and induce blister galls. According to Harris
and Schreiner (1992), the main factors affecting populations of this midge are
rainfall and location. More galls are present during rainy periods, possibly
because RH improves larval and pupal survival. No differences were
observed in gall densities collected from lower and top portions of the tree.
Askari and Radjabi (2003) observed overlapping generations of Procontarinia
J.E. Pea et al. 340
matteiana in Iran related to the different leaf ushing patterns and found that
the optimum pest temperatures were 1026C. Differences on susceptibility
of mango cultivars to P. matteiana might indicate that susceptible cultivars
should not be grown in areas infested by this gall midge (Jhala et al., 1987;
Githure et al., 1998). Daneel et al. (2000) suggested that products to control P.
matteiana should be applied after harvest, coinciding with the rst major
ush and a second spray 6 weeks later. Austin (1984) and Sankaran and Mjeni
(1989) have reported several platygastrid species parasitizing Procontarinia
spp. and their prospects for biological control of the pest.
Mites
The mango bud mite, Aceria mangiferae (Sayed) (Acari: Eriophyidae), attacks
buds and inorescences (Keifer et al., 1982; Ochoa et al., 1994) (Plates 6971).
According to Jeppson et al. (1975) this mite stunts and brooms twigs, causing
bud proliferation and appears to be responsible for necrosis of bud tissue
cells (Varma et al., 1974). In Hawaii, Tegonotus mangiferae (Acari: Eriophyidae)
feeds on the underside of leaves (Jeppson et al., 1975), while another species,
Metaculus mangiferae (Attiah) (Acari: Eriophyidae) causes russeting of termi-
nal leaves, buds and inorescences. The latter is an important pest in Egypt,
India, Palestine and Angola (Jeppson et al., 1975). The puncture wounds of
several acarines (Acari: Tetranychidae) cause serious damage to leaves, which
may dry and fall. The main pest in Mauritius, India, Egypt, Israel and Peru is
Oligonychus mangiferus (Rahman and Sapra); in Israel, the spider mite Tet-
ranychus cinnabarinus (Boisduval), which lives on the underside of the leaves,
causes bronzing around the puncture wounds. The adult life span of O.
mangiferus is 10.11 days for females and 4.21 days for males (Rai et al., 1988).
The avocado red mites, Oligonychus yothersi McGregor and Oligonychus puni-
cae (Hirst), feed on the upper surface of leaves and cause considerable sti-
pling around the midrib at high population densities (Andrews and Poe,
1980). If the tetranychid mites are sufciently abundant, infested leaves may
drop. There is little or no information on sampling techniques or for their
economic thresholds.
Aceria mangiferae occurs wherever mango is grown (Denmark, 1983;
Doreste, 1984). There has been controversy regarding a possible association
between this mite and oral and foliar galls, i.e. mango malformation (Sayed,
1946; Narasimhan, 1959; Summanwar and Raychoudhury, 1968; Denmark,
1983; Ochoa et al., 1994). However, A. mangiferae does not cause mango mal-
formation, but may be a carrier of Fusarium mangiferae, which is recognized
as the causal agent of mango malformation (Varma et al., 1974; Freeman et al.,
2004). Aceria mangiferae life history has been described by Abou-Awad (1981);
it is completed in 15 days at 2527C.
DAMAGE. According to Keifer (cited in Jeppson et al., 1975), A. mangiferae
infestation of buds results in arrested growth, with the stunted, short, young
stems close together at the terminal branch. When the leaves fall, the overall
effect is scanty growth of twiggy branches, with a few stubby, short branch-
lets and discoloured buds. The mite appears to be responsible for necrosis of
Pests 341
bud tissue cells, which is initiated externally at the edge of the bud and pro-
gresses toward the centre and internal areas of the bud (Pea et al., 2005).
SAMPLING. Pea et al. (2005) reported preliminary results of the distribution
and sampling techniques for A. mangiferae. Taylors power law and Iwaos
patchiness regression were used to analyse spatial distributions of the mango
bud mite in mango orchards. Taylors power law generally provided a better
description of variance-mean relationships for the species than did Iwaos
patchiness regression. The species exhibited aggregated patterns of spatial
distribution. More mites were found on apical buds than on lateral-latent
buds. Sample size requirements for xed levels of precision were determined
by using variance-mean relationships (Pea et al., 2005). For a given mean,
and desired precision, different numbers of samples are required. For
instance, for a 10% precision and at 0.5 mites/bud, c.220 samples are required,
whereas for a 30% precision at 0.5 mites/bud, only 25 samples are required.
Sampling small arthropods (i.e. mites) is operationally difcult and often
time consuming. To ease this burden, presence-absence, or binomial, sam-
pling was tested in place of complete counts for estimating or classifying
densities of these organisms. Binomial sampling is based on dening the pro-
portion of one or more individuals the percent of incidence (the incidence
P(I)) and the density of animals (m) per sampling unit. At densities of three
and ten mites/bud, 71% and 42% buds, respectively, are considered unin-
fested. According to the equation, P(I) = 0.28 + 0.01 (mites/bud) an infesta-
tion level of more than ten mites/bud or a P(I) > 38% could be used as the
nominal threshold (Pea et al., 2005).
BIOLOGICAL CONTROL. The phytoseiid Amblyseius swirski Athias Henriot is
associated with A. mangiferae (Abou-Awad, 1981). In Florida USA, several
unidentied phytoseiids occur on buds infested with A. mangiferae. Tenuipal-
pid (Brevipalpus phoenicis), Tydeid and Tarsonemid (Tarsonemus confusus
(Ewing)) mites also inhabit mango buds. Therefore, it is difcult to deter-
mine the mite species that is the host prey (Pea, personal observation).
CHEMICAL CONTROL. Osman (1979) reported that applications of four full cov-
erage sprays of dichlorvos were effective for controlling A. mangiferae in
Egypt. Rai et al. (1966) cautioned that chemical control should be directed to
apparently healthy and not malformed tissues. In Florida USA, agrimek plus
citrus oil, fenproximate and fenpropathrin resulted in the lowest mite densi-
ties 12 days after application. Agrimek plus citrus oil, and acequinocyl
resulted in the lowest mite densities 26 days after treatment (Pea et al., 2005).
In Brazil, Nascimento et al. (2002) recommended sulfur applications.
PLANT RESISTANCE. In Florida USA, the densities of A. mangiferae were mea-
sured on 22 mango cultivars from December 1997 to June 1998. Keenan, an
unknown cultivar, cv. 9819, Brander and Bombay Green had signicantly
more mites than Joellen, Duncan, Red Itamaraca, Smith, Wally and
Hindi (Pea et al., 2005).
J.E. Pea et al. 342
Scales
ARMORED SCALES. At least 26 species of diaspidids attack mangoes worldwide
(Chua and Wood, 1990). In India, Aspidiotus destructor Signoret causes serious
damage, while Parlatoria pergandii Comstock and Lepidosaphes gloverii (Pack-
ard) damage 3-year-old plants (cited in Chua and Wood, 1990). Radionaspis
indica (Marlatt) (= Leucaspis indica) encourages growth of black mould, which
covers young branches (Dekle, 1976). Several diaspidids, e.g. Aulacaspis
mangiferae (tubercularis) Newstead, attack shoots and leaves; the oleander
scale (Plate 72) in Florida USA (Miller and Davidson, 2005) and the mango
scale in Ghana (van Halteren, 1970) cause similar damage. They are damag-
ing not only because they feed on sap, but also because of the toxicity of their
saliva (Singh, 1991). Scales inhabit both leaf surfaces and also are on fruit.
Van Halteren (1970) concluded that A. mangiferae development is completed
in 3540 days for females and 2328 days for males.
SOFT SCALES. Other species of Coccidae, Coccus viridis (Green), Coccus longu-
lus, Ceroplastes actiniformis, Philephedra tuberculosa Nakahara and Gill and the
mango shield scales Milviscutulus mangiferae (Green) and Viusonia stellifera
(Westwood) in Asia, Africa, Australia, Israel and the Americas cause similar
damage. These coccids are generally polyphagous, attacking different genera
and species. They are mobile and injure mango because of the production of
honeydew and the subsequent accumulation of sooty mould on the honey-
dew (Escalante, 1974; Silva and Cavalcante, 1977). Most of these scales can be
suppressed at sub-economic levels, either by application of selective pesti-
cides (i.e. oils) or by biological control agents.
SAMPLING. Because of their small size, it is laborious to sample all stages of
scales. Pheromones in tent-style traps have been used with other fruit crops,
as well as monitoring crawler movement using sticky bands close to infested
leaves. Either double-sided sticky tape, or tape coated with Vaseline

is effec-
tive for trapping crawlers. Bands are removed and examined under a micro-
scope to determine crawler numbers. Dark-coloured tape provides a better
contrast to detect crawlers (Beers et al., 1993).
BIOLOGICAL CONTROL. In a survey of mango-producing areas in South Africa,
Labuschagne (1993) determined that the predatory thrips Auleurodothrips
fasciapennis Franklin and the parasitoid Aspidiotiphagus citrinus (Crawford)
are the most important biocontrol agents of A. tubercularis. In South Africa,
Joubert et al. (2000) obtained 46% parasitism of A. tubercularis using an
unidentied species of the parasitoid, Aphytis sp. Arias et al. (2004a) observed
Coccidophilus spp. (Coleoptera: Coccinellidae) and Chrysopa spp. preying on
A. tubercularis in Ecuador; the exotic predator Cybocephalus nipponicus
(Coleoptera: Nitidulidae) was introduced to supplement predation of the
former scale (Arias et al., 2004b). Several parasites have been recorded in
Israel parasitizing the mango shield scale: Coccophagus lycimnia (Walker),
C. eritraensis Compere, C. scutellaris (Dalman), C. bivittatus (Compere),
Pests 343
Microterys avus (Howard) and Metaphicus avus Howard. Usually no chem-
ical control is required for this scale in Israel due to the activity of natural
enemies (Kr and Rosen, 1980). Natural enemies for the control of the pink
wax scale Ceroplastes rubens Maskell in Australia include the parasitic wasps
Anicetus benecus Ishii and Yasumatsu, Aenasioidea varia Girault and Rhopal-
encyrtoidea dubia Girault (Cunningham, 1984).
Whiteies and blackies
The two y species of economic importance are the whitey, Aleurodicus dis-
persus Russel and the blacky, Aleurocanthus woglumi Ashby. The whiteies
suck cell sap from leaves, which wilt when whitey populations are high.
High infestations can almost blacken entire trees, reducing photosynthetic
efciency and causing defoliation (Angeles et al., 1971; Pea, 1993). A number
of parasitoids, e.g. Encarsia opulenta (Silvestri) and Amitus hesperidus (Silves-
tri), attack the immature stages and provide good control.
Mealybugs
Mealybugs injure mango by sucking sap through their stylets, and excreting
large amounts of honeydew onto fruit and leaves. Sooty mould fungus
growth on the honeydew can render the fruit unmarketable, and reduce the
photosynthetic efciency of leaves and cause leaf drop (CAB International,
2003). The margarodid mango mealybug Drossicha stebbingi (Green) is a seri-
ous pest of mango in India and Pakistan (Prasad and Singh, 1976; Mohyud-
din, 1981; Mohyuddin and Mahmood, 1993). It is univoltine. After mating,
females enter the soil in June and die after laying eggs at a depth of up to
15 cm. These begin to hatch at the end of December or early January. The
nymphs emerge from the soil and move to tender shoots where they settle.
Prasad and Singh (1976) reported that the intensity of attack varied with
respect to year and locality in India, probably because of soil and environ-
mental conditions. Moderate rainfall (5560 mm) during oviposition and dry
conditions during hatching appear to favour development. Adults develop
in April. They mate, and males die soon afterwards. The females enter the
soil in May for oviposition, and the diapausing eggs remain in the soil until
the end of December.
The pseudococcid fruit tree mealybug, Rastrococcus invadens Williams, is
a serious pest of several crops, including mango in West Africa (Agounk et al.,
1988). Mealybugs feed on leaves and fruit. Females have three moults and
males have four. The entire life cycle can be completed in 3184 days. The
mealybugs weaken plants by puncturing the tissues and consuming sap, but
the major damage is caused by the production of large amounts of honeydew
upon which saprophitic fungi develop. The resultant thick black layer of
sooty mould causes a drastic reduction in photosynthetic efciency, resulting
in premature leaf drop. Rastrococcus invadens severely reduces fruit produc-
tion in some areas of Africa (Moore and Cross, 1992).
SAMPLING. Boavida et al. (1992) devised sampling plans for R. invadens, but
advised that the sampling strategy was only practical for estimating medium
to high mealybug populations in the eld. Narasimham and Chacko (1991)
J.E. Pea et al. 344
determined that densities of R. invadens Williams, Rastrococcus iceryodes
(Green) and Rastrococcus mangiferae (Green) were signicantly higher on the
abaxial than the adaxial surface. Mealybug density was also higher from
ground level to 2 m compared with >2 m; they also observed that spatial
distribution of R. iceryodes did not differ among internal and external cano-
pies, whereas densities of R. invadens and R. mangiferae were higher in the
external canopy. They also determined that there were statistical differences
causing some inter-tree variation, but did not determine if the mealybugs
followed a random or contagious distribution.
CONTROL. Various control methods, including banding tree trunks with vari-
ous materials to prevent D. stebbingi nymphs from climbing (Lakra et al.,
1980; Srivastava, 1981) and dusting chlorinated hydrocarbons on the soil
(Srivastava, 1981), have been tried with little success. In Pakistan the mango
mealybug was controlled by hoeing or ploughing the soil and conservation
of the predator, Sumnius renardi Weise, by wrapping burlap around the trunks
of the trees (Mohyuddin and Mahmood, 1993).
Pests of trunks, twigs and roots
Coleopterans and scales
Stem-boring Coleoptera and scales as a group of injurious mango insects
have not been studied in great detail (van Whervin, 1968; Woodruff, 1985).
The wide host range of borers and overlapping borer species has compli-
cated their study. Tunnelling of borer larvae in branches and trunks of mango
and the slow feeding of some scale species in certain seasons and regions
may cause serious reductions in yields and might also contribute to mango
decline. However, borer occurrence and injury tends to be sporadic and
below levels requiring direct action. Few natural enemies have been reported
for suppression of borer populations in mango. Detailed coverage of stem-
boring species is beyond the scope of this chapter; however, Hypocryphalus
mangiferae (Stebbing), Apate monachus Fabricius and Batocera rubus L. are pests
within this group.
Infestations of the mango scale, Radionaspis indica (Marlatt), and plumose
scale, Morganella longispina, commonly occur on the trunk, branches and
buds. Severe infestations include cracking of bark, exudation of sap and
decline of the upper branches. Pea (1993) demonstrated that branches with
both species of scale showed more decline symptoms than branches with
low-scale density. Research on the bionomics and control of these scales is
necessary to conrm their role in mango decline symptomatology.
The scolytids, Hypocryphalus mangiferae (Stebbing) and Xylosandrus com-
pactus (Eichoff) directly attack the main stem and branches (Silveira, 1960;
Wysoki et al., 1993). Growth of fungal mycelia can extend terminally and
basally from the beetle gallery in the mango tree and can kill the affected
branches. The insects prefer trees that have been weakened by pathogens,
wind, etc., but after a population has been established, the infestation spreads
Pests 345
to healthy trees. Hypocryphalus mangiferae has been associated with mango
wilt disease in Brazil and Oman (van Wyk et al., 2007). Following initial trap-
ping results from Berti Filho and Fletchman (1986), Rocha da Silva (2006)
established that H. mangiferae is attracted to trees where the fungus is pres-
ent. Scolytid beetles are attracted to mango trees in response to visual stimuli,
to host-specic chemicals and to species-specic aggregation pheromones
(Lindgreen et al., 1982). The evaluation of traps as tools for managing ambro-
sia beetles on mangoes in Florida USA is necessary in order to reduce their
damage in newly established groves.
Termites
Mango orchards are becoming more common in dry and semi-arid areas
with vast termite populations. Mango growing in infested areas often results
in plant growth suppression as a result of reduced root establishment, inva-
sion and pruning of roots (Rogers et al., 1999). For example, six termite spe-
cies (Odontotermes indicus Thakur, O. lokanandi Chatterjee and Thakur, O.
obesus (Rambur), O. giriensis (Roonwal and Chhotani), O. bhagwatii Chatter-
jee and Thakur and Microtermes obesi Holm) were recorded from mango
orchards in India (Srivastava and Singh, 2004). More species of termites were
observed in winter than during the summer and rainy season. Veeresh et al.
(1989) observed that O. wallonensis, O. horni and O. obesus constructed earthen
sheeting on the stem of small mango trees. Singh (1960), cited by Srivastava
(1998), reports Eutermes (Nasutitermes) costali, Calutermes (Cryptotermes) bre-
bis, Heterotermes tenuis, Coptotermes gestroi, Neotermes (Kelatermes) bosei (Gard-
neri) Synder, Microcereoutermes perofnis, Calotermes (Neotermes) greeni and
Coptotermes curvignathus also affecting mango in India. According to Srivas-
tava (1998), the most important termite species affecting mango are O. obesus
and O. wallonensis; O. wallonensis nests in the root zone in Uttar Pradesh,
India. The workers feed on roots, stems and branches.
Colonies of the subterranean termite, Coptotermes curvignathus Holmgren,
were monitored in Malaysia using bait matrices containing 0.5% hexau-
muron (Said Sajap et al., 2000). In Florida, urban dwellings infested with the
Formosan termite, C. formosanus Shiraki, were treated with baits containing
0.5% weight/weight noviumuron (Cabrera and Thoms, 2006). These baits
might be useful when mango orchards are planned for areas infested with
termites. In India, termite infestations are controlled with a combination of
monthly irrigation, hoeing and application of neem cake (Singh and Singh,
2003).
10.3 Discussion
In general, most mango pests also occur on other fruit crop species. Fruit
ies, scales, mites, thrips, lepidopteran ower feeders, mirids, weevils and
beetles are mostly generalists, and some of their management schemes need
to be implemented with this in mind. In the case of fruit ies, Aluja (1996)
suggests surveying vegetation adjacent to infested mango orchards as
J.E. Pea et al. 346
populations are sustained and multiplied in these locations and from them
adult ies move into commercial orchards to attack ripening fruit (Aluja et
al., 1996). Management of key pests (i.e. fruit ies, seed weevils, etc.) must be
mandatory, in order to have an effect on a large region. The use of some mea-
sures (i.e. quarantine, etc.) must involve neighbouring producing countries
in order to have a positive effect on sanitation. The most progressive exam-
ples in management of mango pests are in Australia, Mexico and Israel, while
other producing countries, such as South Africa, Brazil and Ecuador, are tak-
ing serious steps to reconcile the opposing forces of globalization of markets
and sustainibility. For other areas where maximizing yields and blemish-free
fruit is not a priority, the emphasis should be biological control. Management
tactics that can be improved include the following:
Selective pesticides. Pesticides that are used in integrated pest man- 1.
agement programmes must have selective toxicity. The current trend is the
development of chemicals that are highly effective for a limited group of in-
sects. Daz-Fleischer et al. (1996) suggested the use of cyromazine to reduce
fertility of A. obliqua. Cunningham (1989) suggested that oils could be uti-
lized for control of scales in mango; however, most of the recommendations
are based on highly toxic or illegal, non-registered persistent chemicals
(Singh, 1991; van Mele et al., 2001; de Bie, 2004). In South Africa, Joubert et al.
(2004) tested kaolin, which is the active ingredient of Surround, a non-toxic
natural clay mineral, against the mango seed weevil, mango scale and citrus
thrips. Surround was effective against citrus thrips, mango weevil and co-
conut bug, P. wayi, but caused outbreaks of mango scale and long-tailed
mealybug, Pseudococcus longispinus. However, producers of export fruit rely
on calendar-based chemical control when trees are heavily infested with
mango scale (Joubert et al., 2004).
Biological control. Biological control has great potential as a tactic for
regulating pest populations in integrated pest management programmes in
mango orchards. However, it will be difcult for biological control alone to
reduce a pest from an economic to a completely non-economic status for
pests attacking fruit. A combination of augmentative releases of parasi-
toids and the use of sterile insects, at least from a theoretical perspective,
has been considered to be more effective for fruit ies than either method
applied alone (Barclay, 1987). Biological control should be highly effective
for indirect pests. Indeed, numerous studies have been conducted in many
mango-producing countries to promote the use of parasites and predators
for this type of pest (Cunningham, 1989; Mohyuddin and Mahmood, 1993;
Moore and Cross, 1993; Whitwell, 1993; Wysoki et al., 1993; Labuschagne
et al., 1995).
Host plant resistance. Tolerance of mango to pests is mentioned for 2. Noorda
sp. and Idioscopus sp. (Bagle and Prasad, 1984; Cunningham, 1989), while man-
go resistance to Sternochetum mangiferae is mentioned by Hansen (1993).
Carvalho et al. (1996) have also demonstrated the different degrees of suscepti-
bility of mango cultivars to A. obliqua. Most of this research, however, needs
Pests 347
to be assessed further. Angeles (1991) reported that Mangifera altissima does
not seem to be affected by leaf hoppers, tip borers and seed borers in the
Philippines. There is little doubt that wild mangoes have potential in breed-
ing. Determining the tolerance or insect resistance of mango cultivars and
related species should be done in natural stands or in established germplasm
collections. After the initial selection has been made, evidence for the pattern
of resistance must be established and changes in the environment, whether
geographic or temporal, should not disrupt or decrease the resistance to any
great extent. Therefore, tests for resistance in mango to insects should in-
clude provision for exposure to insects under varying conditions whenever
possible.
Pheromones and trapping devices. Developments in the identication 3.
and synthesis of sex pheromones have resulted in their possible use for pest
management in mango orchards (Chu et al., 1994; Khan et al., 2002, 2005;
Sheikh et al., 2008). Food attractants, however, remain the most common
monitoring tools. Trapping techniques can be utilized to reduce pesticide use
by improving timing of sprays as a result of better monitoring of pest popu-
lations. It remains uncertain if trapping techniques can be used to predict
infestations by fruit-feeding pests and if they can be used for direct control
(by mass trapping) over several years.
Cultural and physical control. Use of cultural and physical techniques 4.
(i.e. pruning, bagging, etc.) depends on costs of control, availability of techni-
cal assistance and market purposes. Paderes and Orden (2004) observed that
bagging and pruning of Manila in the Philippines is mostly inuenced by
the availability of technical assistance for growers.
Greatly increased regulation of pesticides, heightened public aware-
ness of environmental contamination, pesticide resistance problems in
pests and the high cost of chemical pest control has resulted in increasing
reliance on integrated pest control as an important strategy in sustainable
agriculture.
Acknowledgements
We are particularly indebted to Walther Enkerlin (Joint IAEA/FAO Division,
Vienna), Andrea Birke-Biewendt, Alberto Anzures-Dadda and Larissa Guilln-
Conde (Instituto de Ecologa, AC) and Aldo Malavasi (private consultant) for
their assistance and for providing many valuable references. M. Aluja
acknowledges the nancial support of the Mexican Campaa Nacional Contra
Moscas de la Fruta (Convenio SAGARPA-IICA-INECOL) and the Instituto
de Ecologa, AC. He also acknowledges support from CONACyT through a
Sabbatical Year Fellowship (Ref. 79449) and thanks Benno Graf and Jrg
Samietz (Forschungsanstalt Agroscope Changins-Wdenswil ACW), for pro-
viding ideal working conditions during the nal phase of the publication
process of this chapter.
J.E. Pea et al. 348
References
Abbas, S. (1985) Studies on the mango inorescence midge, Erosomyia indica Grover.
Acta Horticulturae 231, 593596.
Abou-Awad, B. (1981) Ecological and biological studies on the mango bud mite, Erio-
phyes mangiferae (Sayed), with description of immature stages Eriophyoidea: Erio-
phyidae. Acarologia 22, 145150.
Agounk, D., Agricola, U. and Bokonon-Ganta, H. (1988) Rastrococcus invadens Wil-
liams (Hemiptera: Pseudococcidae), a serious exotic pest of fruit trees and other
plants in West Africa. Bulletin of Entomology Research 78, 695702.
Ahmed, M.U., Ahmed, M., Baluch, M.A. and Naheed, R. (1981) Sooty mold on mango
plants and its relationship with leafhoppers and climatic factors in Karachi-Pakistan
during 197879. Pakistan Journal of Science and Industrial Research 24, 140144.
Aluja, M. (1993) Manejo Integrado de la Mosca de la Fruta. Editorial Trillas, Mexico
Distrito Federal, Mexico.
Aluja, M. (1994) Bionomics and management of Anastrepha. Annual Review of Ento-
mology 39, 155178.
Aluja, M. (1996) Future trends in fruit y management. In: McPheron, B. and Steck, G.
(eds) Fruit Fly Pests: a World Assessment of their Biology and Management. St Lu-
cie Press, Delray Beach, Florida, pp. 309320.
Aluja, M. (1999) Fruit y (Diptera: Tephritidae) research in Latin America: myths, reali-
ties and dreams. Anais da Sociedade Entomologica do Brasil 28, 565594.
Aluja, M.R. and Liedo, P.F. (1986) Perspectives on future integrated management of fruit
ies in Mexico. In: Mangel, M., Carey, J.R. and Plant, R.E. (eds) Pest Control: Op-
erations and Systems Analysis in Fruit Fly Management. NATO ASI Series Vol. G11.
Springer-Verlag, Berlin, pp. 942.
Aluja, M. and Mangan, R.L. (2008) Host status determination for pestiferous fruit y
(Diptera: Tephritidae) species: conceptual, methodological and regulatory consid-
erations. Annual Review of Entomology 53, 473502.
Aluja, M. and Norrbom, A.L. (eds) (2000) Fruit Flies (Tephritidae): Phylogeny and Evolu-
tion of Behavior. CRC Press, Boca Raton, Florida.
Aluja, M. and Piero, J. (2004) Testing human urine as a low-tech bait for Anastrepha
spp. (Diptera: Tephritidae) in small guava, mango, sapodilla and grapefruit orchards.
Florida Entomologist 87, 4150.
Aluja, M., Cabrera, M., Guillen, J., Celedonio, H. and Ayora, F. (1989) Behavior of Anas-
trepha ludens, A. obliqua and A. serpentina (Diptera: Tephritidae) on a wild mango
tree (Mangifera indica) harbouring three McPhail traps. Insect Science and its Ap-
plication 10, 309318.
Aluja, M., Celedonio-Hurtado, H., Liedo, P., Cabrera, M., Castillo, F., Guillen, J. and
Rios, E. (1996) Seasonal population uctuations and ecological implications for
management of Anastrepha fruit ies (Diptera: Tachinidae) in commercial mango
orchards in southern Mexico. Journal of Economic Entomology 89, 654667.
Aluja, M., Piero, J., Jcome, I., Daz-Fleischer, F. and Sivinski, J. (2000) Behavior of ies
in the genus Anastrepha (Trypetinae: Toxotrypanini). In: Aluja, M. and Norrbom, A.
(eds) Fruit Flies (Tephritidae): Phylogeny and Evolution of Behavior. CRC Press,
Boca Raton, Florida, pp. 375406.
Aluja, M., Sivinski, J., Rull, J. and Hodgson, P.J. (2005) Behavior and predation of fruit
y larvae (Anastrepha spp.) (Diptera: Tephritidae) after exiting fruit in four types of
habitats in tropical Veracruz, Mexico. Environmental Entomology 34, 15071516.
Aluja, M., Sivinski, J., Ovruski, S., Guilln, L., Lpez, M., Cancino, J., Torres-Anaya,
A., Gallegos-Chan, G. and Ruz, L. (2009) Colonization and domestication of
Pests 349
seven species of native New World Hymenopterous larval-prepupal and pupal fruit
y (Diptera: Tephritidae) parasitoids. Biocontrol Science and Technology (in press).
Andrews, K. and Poe, S. (1980) Spider mites of El Salvador, Central America (Acari:
Tetranychidae). Florida Entomologist 63, 502505.
Angeles, D.E. (1991) Mangifera atissima. In: Coronel, R.E. and Verheij, E.W.M. (eds)
Plant Resources of South-east Asia No.2: Edible Fruits and Nuts. Pudoc-DLO,
Wageningen, the Netherlands, pp. 206207.
Angeles, N. and Requena, J. (1966) Moth injury of mangoes in Venezuela. Plant Protec-
tion Bulletin FAO 14, 8687.
Angeles, N., Dedordy, N., Paredes, P. and Requena, J. (1971) Citrus blacky in Venezu-
ela. Agronomia Tropical 21, 7175.
Anonymous (1984) Major insect pests and diseases of mango and their control. In: The
Mango Technical committee. The Philippines recommends for Mango. Philippines
Agriculture and Resources Research Foundation, 124 pp.
Anonymous (1987) Red-banded mango caterpillar Noorda albizonalis Hampsoni (Lepi-
doptera: Pyralidae). Plant Quarantine Leaet-Australia 51.
Anonymous (1989) Mango Pests and Disorders. Information Series QI89007. Queen-
sland Department of Primary Industries, Brisbane, Australia.
Anonymous (2002) Fruit ies eradicated from Nauru. 3pp. Available at: http://www.
paciy.org/Success_stories/Nauru_eradl.htm (accessed 10 September 2008).
Anonymous (2005) Malathion (CAS Number 121-75-5). Toxicology information about
insecticides used for eradicating mosquitoes. Available at: http://www.atsdr.cdc.
gov/consultations/malathion.html (accessed 20 July 2007).
Anonymous (2006) Importation of Fresh Mango Fruit (Mangifera indica) from India into
the United States, a Qualitative Pest Risk Assessment. Animal and Plant Health In-
spection Service, Plant Protection Quarantine, Raleigh, North Carolina.
Arias, M. and Jines, A. (2004) Manejo Integrado de Moscas de la Fruta en el Litoral Ecu-
atoriano. Instituto Nacional de Investigaciones Agropecuarias Programa de Mod-
ernizacin de Servicios Agropecuarios, Guayaquil, Equador.
Arias, M., Jines, A., Bustos, P., Pluas, M. and Carrera, C. (2004a) Enemigos Naturales
de Aulacaspis Tubercularis (Homoptera: Diaspididae) en Mango. Instituto Na-
cional Autonomo de Investigaciones Agropecuarias (INIAP), Boliche, Guayas,
Ecuador.
Arias, M., Jines, A., Maldonado, E. and Pluas, M. (2004b) Cybocephalus nipponicus
Endrody-Younga (Coleoptera: Nitidulidae) un Predator Extico Para el Control del
Aulacaspis Tubercularis en Mango de Exportacin. Instituto Nacional Autonomo de
Investigaciones Agropecuarias (INIAP), Boliche, Guayas, Ecuador.
Askari, M. and Radjabi, G. (2003) Study on the biology and population uctuations of
mango midge gall Procontarinia mattiana (Diptera: Cecidomyiidae) in Homozogan
province. Applied Entomology and Phytopathology 70, 121135.
Assis, E. and Rabelo, F. (2005) Pragas da Mangueira, Monitoramento, Nivel de Ao e
Controle. Empresa Brasileira de Pesquisa Agropecuaria, Embrapa Semi-Arido, Min-
isterio da Agricultura, Pecuaria e Abastecimiento, Petrolina, Brazil.
Austin, A.D. (1984) New species of Platygastridae (Hymenoptera) from India which
parasitise pests of mango, particularly Procontarinia spp. (Diptera: Cecidomyiidae).
Bulletin of Entomological Research 74, 549557.
Australian Government (2004) Mangoes from India, Draft Revised Import Policy. De-
partment of Agriculture, Fisheries and Forestry, Canberra, Australia.
Azizur Rahman, S.K. and Singh, G. (2004) Population dynamics of mango hopper (Am-
ritodus atkinsoni) on Langra mango (Mangifera indica) and its relationship with
abiotic factors. Indian Journal of Agricultural Science 74, 566569.
J.E. Pea et al. 350
Babu, L.B., Maheswari, T.U. and Rao, N.V. (2002) Seasonal incidence and biology of the
mango hopper Amritodus atkinsoni Lethier (Homoptera: Cicadellidae). Entomon
27, 3542.
Bagle, B. and Prasad, V. (1984) Varietal incidence and control of stone weevil Sternoche-
tus mangiferae Fabricius. Indian Journal of Entomology 46, 389392.
Balock, J.W. and Kozuma, T. (1964) Notes on the biology and economic importance of
the mango weevil, Sternochetus mangiferae (Fabricius), in Hawaii (Coleoptera:
Curculionidae). Proceedings of the Hawaiian Entomological Society 8, 353364.
Balock, J.W. and Lopez, D.F. (1969) Trapping for control of the Mexican fruit y in
mango and citrus groves. Journal of Economic Entomology 62, 5356.
Barbosa, F.R. (2005) Arthropodes-pragas associados a cultura da mangueira no Brasil e
seu controle. In: Menezes, E. and Barbosa, F.R. (eds) Pragas da Mangueira, Embra-
pa Semi-arido. Ministerio da Agricultura, Pecuaria e Abastecimiento, Petrolina, Bra-
zil, pp. 150.
Barbosa, F.R., de Gonalves, M.E., Moreira, W.A., de Alencar, J.A., de Souza E.A., da
Silva, C.S.B., Souza, A. and da Miranda, I. (2005) Arthrpodes-Praga e Predatores
(Arthropoda) Associados Cultura da Manguerira no Vale do So Francisco, Nor-
deste do Brasil. Neotropical Entomology 34, 471474.
Barclay, H.J. (1987) Models for pest control: complementary effects of periodic releases
of sterile pests and parasitoids. Theoretical Population Biology 32, 7689.
Barnes, H.F. (1948) Gall Midges of Economic Importance. Vol. 3: Gall Midges of Fruit.
Crosby Lockwood and Sons, London.
Bartlet, K. (1938) The introduction and colonization in Puerto Rico of Dasyscapus parvi-
pennis Gahan, a parasite of thrips. Agricultural Notes Puerto Rico Experiment Sta-
tion US Department of Agriculture (USDA) 87, 6.
Bateman, M.A. (1972) The ecology of fruit ies. Annual Review of Entomology 17,
493518.
Beardsley, J.W. (1961) A new review of the Hawaiian Braconidae (Hymenoptera). Pro-
ceedings of the Hawaiian Entomology Society 27, 333366.
Beers, E., Hull, L. and Jones, V. (1993) Sampling pest and benecial arthropods of apples.
In: Pedigo, L. and Buntin, D. (eds) Handbook of Sampling Methods for Arthropods in
Agriculture. Preface. CRC Press, Boca Raton, Florida, pp. 383416.
Berti Filho, E. and Flechtman, C. (1986) A model of ethanol trap to collect scolytidae
and platypodidae (Insecta, Coleoptera). Instituto de Pesquisas e Estudios Florestais
34, 5356.
Bess, H.A., van den Bosch, R. and Haramoto, F.H. (1961) Fruit y parasites and their ac-
tivities in Hawaii. Proceedings of the Hawaiian Entomology Society 27, 367378.
Bhole, S., Jadhav, V., Dumbre, R. and Dalvi, C. (1987) Seasonal incidence and chemical
control of mango nursery pests in the Konkan region. Journal of the Maharashtra
Agricultural University 12, 387388.
Birke, A., Aluja, M., Greany, P., Bigurra, E., Prez-Staples, D. and McDonald, R. (2006) Long
aculeus and behavior of Anastrepha ludens renders gibberellic acid ineffective as an
agent to reduce Ruby Red grapefruit susceptibility to the attack of this pestiferous fruit
y in commercial citrus orchards. Journal of Economic Entomology 99, 11841193.
Boavida, C., Neuenschwander, P. and Schulthess, F. (1992) Spatial distribution of Rastro-
coccus invadens Williams (Homoptera: Pseudococcidae) in mango trees. Journal of
Applied Entomology 114, 381391.
Bondad, N.D. (1985) Mango bagging. Animal Husbandry and Agricultural Journal June
Issue, 1114.
Burns, R.E., Harris, D.L., Moreno, D.S. and Eger, J.E. (2001) Efcacy of spinosad bait
sprays to control Mediterranean and Caribbean fruit ies (Diptera: Tephritidae) in
commercial citrus in Florida. Florida Entomologist 84, 672678.
Pests 351
Bustos, M.E., Enkerlin, W., Reyes, J. and Toledo, J. (2004) Irradiation of mangoes as a
postharvest quarantine treatment for fruit ies (Diptera: Tephritidae). Journal of Eco-
nomic Entomology 97, 286292.
Butani, D.K. (1979) Insects and Fruits. Periodical Expert Book Agency, Delhi, India.
CAB International (2003) Crop Protection Compendium. CAB International, Walling-
ford, UK.
Cabrera, B.J. and Thoms, E. (2006) Versatility of baits containing noviumuron for con-
trol of structural infestations. Florida Entomologist 89, 2031.
Cabrera, H., Villanueva, J. and Ortega, D. (1993) Fruit y infestation timing on Manila
mango fruit at Veracruz, Mexico. (Abstract). IV International Mango Symposium,
p. 118.
Camargo, C.A., Odell, E. and Jirn, L.F. (1996) Interspecic interactions and host prefer-
ence of Anastrepha obliqua and Ceratitis capitata (Diptera: Tephritidae), two pests
of mango in Central America. Florida Entomologist 79, 266268.
Carvalho, P.J. and De Queiroz, A.C. (eds) (2002) A Cultura da Mangueira. CIP-Brasil,
Brasilia.
Carvalho, R. da S., Nascimento, A., Morgante, J. and Fonseca, N. (1996) Susceptibility
of different mango varieties to the attack of the fruit y, Anastrepha obliqua. In:
McPheron, B. and Steck, G. (eds) Fruit Fly Pests: a World Assessment of their Biol-
ogy and Management. St Lucie Press, Delray Beach, Florida, pp. 325331.
Celedonio-Hurtado, H., Aluja, M. and Liedo, P. (1995) Adult population uctuations of
Anastrepha species (Diptera: Tephritidae) in tropical orchard habitats of Chiapas,
Mexico. Environmental Entomology 24, 861869.
Christenson, L.D. and Foote, R.H. (1960) Biology of fruit ies. Annual Review of Ento-
mology 5, 171192.
Chu, Y., Lee, K.T. and Tseng, Y.H. (1994) Occurrence of melon and oriental fruit y in
Republic of Naru. Plant Protection Bulletin 36, 131140.
Chua, T.H. and Wood, B.J. (1990) Other tropical fruit trees and shrubs. In: Rosen, D.
(ed.) Armored Scale Insects, their Biology, Natural Enemies and Control, 4B. El-
sevier, New York, pp. 543552.
Clarke, A.R., Allwood, A., Chinajariyawong, A., Drew, R.A.I., Hengsawad, C., Jirasurat,
M., Kong-Krong, C., Kritsaneepaibon, S. and Vijaysegaran, S. (2001) Seasonal abun-
dance and host use patterns of seven Bactrocera Macquart species (Diptera: Te-
phritidae) in Thailand and Peninsular Malaysia. The Rafes Bulletin of Zoology 49,
207220.
Clarke, A.R., Armstrong, K.F., Carmichael, A.E., Milne, J.R., Raghu, S., Roderick, G.K.
and Yeates, D.K. (2005) Invasive phytophagous pests arising through a recent tropi-
cal evolutionary radiation: the Bactrocera dorsalis complex of fruit ies. Annual
Review of Entomology 50, 293319.
Coto, D., Saunders, J.L., Vargas, C.L. and King, A.B.S. (1995) Plagas invertebradas de
cultivos tropicales con enfasis en America central. Centro Agronmico Tropical de
Investigacin y Enseanza. Serie Tecnica, Manual Tcnico 12.
Cunningham, I.C. (1984) Mango insect pests. In: Australian Mango Research Workshop,
Commonwealth Scientic and Industrial Research Organization (CSIRO), Mel-
bourne, pp. 211224.
Cunningham, I.C. (1989) Pests. In: Bagshaw, J. (ed.) Mango Pests and Disorders. Depart-
ment of Primary Industries Information Series, Q189007, pp. 1021.
Daneel, M.S., Jager, K. de, Steyn, W. and Husselman, J. (2000) Efcacy of different in-
secticides against gall y on mangoes. South African Mango Growers Association
Yearbook 19, 20.
de Bie, C.A. (2004) The yield map of mango in Phrao, Thailand, as investigated through
comparative performance evaluation. Scientia Horticulturae 102, 3752.
J.E. Pea et al. 352
De, K. and Pande, Y.D. (1987) Evaluation of certain non-insecticidal methods of reduc-
ing infestation of the mango nut weevil, Sternochetus gravis (F.) in India. Tropical
Pest Management 33, 2728.
De, K. and Pande, Y.D. (1988) Bionomics and some behavioural aspects of the mango
stone weevil, Sternochetus gravis (Fabricius) (Coleoptera: Curculionidae). Entomon
13, 1724.
De la Rosa, W., Lpez, F.L. and Liedo, P. (2002) Beauveria bassiana as a pathogen of the
Mexican fruit y (Diptera: Tephritidae) under laboratory conditions. Journal of Eco-
nomic Entomology 95, 3643.
De Macedo, F.P. (1998) Efeitos da aplicao de cido giberlico em manga (Mangifera
indica L.) cultivar Tommy Atkins: aspectos horticulturais e entomolgicos. Tesis de
doctorado, Universidade de So Paulo, Brazil, 120 pp.
De Villiers, E.A. (1990) Coconut bugs in avocados. Farming in South Africa. Avocados
H.5, 2pp.
Dekle, G.W. (1976) Florida armored scale insects. Arthropods of Florida and neighbor-
ing land areas, Florida Department of Agriculture and Consumer Services, Gaines-
ville, Florida, 3, 345 pp.
Denmark, H. (1983) Eriophyes mangiferae (Sayed) a pest of mango (Acarina: Eriophyidae).
Entomology Circular Florida Department of Agriculture and Consumer Services 254.
Daz-Fleischer, F. and Aluja, M. (2003) Clutch size in frugivorous insects as a function of
host rmness: the case of the tephritid y Anastrepha ludens. Ecological Entomol-
ogy 28, 268277.
Daz-Fleischer, F., Toledo, J., Enkerlin, W. and Fernandez, J. (1996) Cyromazine: effects
on three species of Anastrepha. In: McPheron, B. and Steck, G. (eds) Fruit Fly Pests:
a World Assessment of their Biology and Management. St Lucie Press, Delray
Beach, Florida, pp. 336337.
Dimbi, S., Maniania, N.K., Lux, S., Ekesi, S. and Mueke, J.K. (2003) Pathogenicity of Metar-
hizium anisopliae (Metsch.) Sorokin and Beauveria bassiana (Balsamo) Vuillemin, to
three adult fruit y species: Ceratitis capitata (Wiedemann), C. rosa var. fasciventris
Karsch and C. cosyra (Walker) (Diptera: Tephritidae). Mycopathologia 156, 375382.
Dolinski, C. and Lacey, L.A. (2007) Microbial control of arthropod pests of tropical tree
fruits. Neotropical Entomology 36, 161179.
Doreste, E. (1984) Acarologia. Instituto Interamericano de Cooperacion para la Agricul-
tura. San Jose, Costa Rica, 391 pp.
Drew, R.A.I. (1989) The tropical fruit ies (Diptera: Tephritidae: Dacinae) of the Austral-
asian and Oceanian regions. Memoirs of the Queensland Museum 26, 521.
Drew, R.A.I. and Hancock, D.L. (1994) The Bactrocera dorsalis complex of fruit ies
(Diptera: Tephritidae: Dacinae) in Asia. Bulletin of Entomological Research Supple-
mental Series, No.2, 68 pp.
Dyck, V.A., Hendrichs, J. and Robinson, A.S. (eds) (2005) Sterile Insect Technique. Prin-
ciples and Practice in Area-Wide Integrated Pest Management. Springer, Dordrecht,
the Netherlands.
Ehler, I.E. and Endicott, P.C. (1984) Effect of malathion-bait sprays on biological control
of insect pests of olive, citrus and walnut. Hilgardia 52, 147.
Ekesi, S., Nderitu, P.W. and Rwomushana, I. (2006) Field infestation, life history and
demographic parameters of the fruit y Bactrocera invadens (Diptera: Tephritidae)
in Africa. Bulletin of Entomological Research 96, 379386.
Emery, R. (2002) Mango seed weevil and mango pulp weevil. Available at: http://www.
agric.wa.gov.au/ento/Su...%20seed%20and%20pulp%20weevil.html (accessed 20
July 2007).
Enkerlin, W.R. (2007) Impact of fruit y control programmes using the sterile insect
technique. In: Dyck, V.A., Hendrichs, J. and Robinson, A.S. (eds) Sterile Insect Tech-
Pests 353
nique. Principles and Practice in Area-Wide Integrated Pest Management. Springer,
Dordrecht, the Netherlands, pp. 651676.
Epsky, N., Hendrichs, J., Katsoyannos, B., Vasquez, A., Ros, J., Zumreolu, A., Bakri, A.,
Seewooruthum, S. and Heath, R. (1999) Field evaluation of female-targeted trapping
systems for Ceratitis capitata (Diptera: Tephritidae) in seven countries. Journal of
Economic Entomology 92, 156164.
Escalante, J.A. (1974) Insects of economic importance in Quillabamba, Cusco. Revista
Peruana de Entomologia 17, 5153.
Etienne, J. (1966) Lutte contre les mouches des fruits. Rapporte de Institute de Recher-
ches Agronomiques Tropicales Runion. Institute National de la Recherche
Agronomique, Linne de Paradis, Reunion, France, pp. 112113.
Etienne, J. (1968) Lutte contre les mouches des fruits. Rapporte de Institute de Recherches
Agronomiques Tropicales Runion. Institute National de la Recherche Agronomique,
Linne de Paradis, Reunion, France, pp. 117124.
Faih, H.A. and Huwer, R. (1994) Egg parasitoids collected from Amblypelta lutescens
lutescens (Distant) (Hemiptera: Coreidae) in north Queensland. Journal of the Aus-
tralian Entomology Society 32, 365367.
Fasih, M., and Srivastava, R.P. (1990) Parasites and predators of insect pests of mango.
Integrated Pest Control 32, 3941.
Fasih, M., Srivastava, R.P., Abbas, S. and Sharma, S. (1989) Outbreak of Orgyia postica
Walker (Lymantriidae: Lepidoptera), a new pest on mango in Uttar Pradesh. Current
Science 58, 12581260.
Fay, H.A. and Huwer, R. (1994) Egg parasitoids collected from Amblypelta lutescens
(Distant) (Hemiptera: Coreidae) in north Queensland. Journal of the Australian En-
tomological Society 32, 365367.
Fletcher, B.S. (1987) The biology of dacine fruit ies. Annual Review of Entomology 32, 115144.
Fletcher, M.J. and Dangereld, P.C. (2002) Idioscopus clypealis (Lethierry), a second
new leafhopper pest of mango in Australia (Hemiptera: Cicadellidae: Idiocerinae).
Australian Journal of Entomology 41, 3538.
Follett, P. and Gabbard, Z. (2000) Effect of mango weevil (Coleoptera: Curculionidae) damage
on mango seed viability in Hawaii. Journal of Economic Entomology 93, 12371240.
Follett, P. and Neven, L.G. (2006) Current trends in quarantine entomology. Annual Re-
view of Entomology 51, 359385.
Freeman, S., Klein-Gueta, D., Korolev, D. and Sztenjnberg, A. (2004) Epidemiology and
survival of Fusarium mangiferae, the causal agent of mango malformation disease.
Acta Horticulturae 645, 487491.
Gagne, R.J. and Etienne, J. (2006) Gephyraulus mangiferae (Felt) N. Comb. (Diptera:
Cecidomyiidae): a mango pest from India newly recorded from the western hemi-
sphere. Proceedings of the Entomology Society of Washington 108, 930937.
Gagne, R.J. and Medina, C.D. (2004) A new species of Procontarinia (Diptera: Cecid-
omyiidae), an important new pest of mango in the Philippines. Proceedings of the
Entomological Society of Washington 106, 1925.
Galn-Saco, V. (1990) Los Frutales Tropicales en los Subtropicos. MundiPrensa, Ma-
drid, Spain, pp. 8384.
Ganz, S., Hamieri, Y. and Nakach, Y. (1990) Control trials on the mango thrips. Annual Report
of the Eden Experimental Station Israel. Institute of Plant Protection, Bet Sheam, Israel.
Garca-Medel, D., Sivinski, J., Daz-Fleischer, F., Ramirez-Romero, R. and Aluja, M.
(2007) Foraging behavior by six fruit y parasitoids (Hymenoptera: Braconidae) re-
leased as single- or multiple-species cohorts in eld cages: inuence of fruit loca-
tion and host density. Biological Control 43, 1222.
Garca-Ramrez, M.J., Cibrain-Tovat, J., Arzuf-Barrera, R., Lpez-Collado, J. and Soto-
Hernndez, M. (2004) Preferencia de Anastrepha ludens (Loew) (Diptera: Tephritidae)
J.E. Pea et al. 354
por voltiles de frutos verdes o amarillos de mango y naranja. Agrociencia 38,
423430.
Githure, C.W., Schoeman, A.S. and McGeoch, M.A. (1998) Differential susceptibility of
mango cultivars in South Africa to galling by the mango gall y, Procontarinia mat-
teiana Kieffer & Cecconi (Diptera: Cecidoyiidae). African Entomology 6, 3340.
Godase, S.K., Bhole, S.R., Shivpuje, P.R. and Patil, B.P. (2004) Assessment of yield loss
in mango (Mangifera indica) due to mango hopper (Idioscopus niveoparvasus) (Ho-
moptera: Cicadelidae). Indian Journal of Agricultural Sciences 74, 370372.
Golez, H.G. (1991) Bionomics and control of the mango seed borer, Noorda albizonalis
Hampson (Pyralidae: Lepidoptera). Acta Horticulturae 291, 418424.
Grove, T. and Pringle, K. (2000) A sampling system for estimating population levels of
the citrus thrips, Scirtothrips aurantii faure (Thysanoptera: Thripidae), mango or-
chards. African Entomology 8, 223226.
Grove, T., Giliomee, J.H. and Pringle, K. (2001) Thrips (Thysanoptera) species associated
with mango trees in South Africa. African Entomology 9, 153162.
Grover, P. (1986a) Integrated control of midge pests. Cecidologia Internationale 7, 128.
Grover, P. (1986b) Population uctuation of Erosomya indica and Dasineura amaraman-
jarae and co-related extent of damage. Cecidologia Internationale 7,4357.
Hallman, G.J. and Sharp, J.L. (1990) Mortality of Caribbean fruit y (Diptera: Tephriti-
dae) larvae infesting mangoes subjected to hot-water treatment, then immersion
cooling. Journal of Economic Entomology 83, 23202323.
Hancock, D.L., Hamacek, E.L., Lloyd, A.C. and Elson-Harris, M.M. (2000) The Distribu-
tion and Host Plants of Fruit Flies (Diptera: Tephritidae) in Australia. Department of
Primary Industries, Brisbane, Australia.
Hansen, J.D. (1993) Dynamics and control of the mango seed weevil. Acta Horticulturae
341, 415420.
Hansen, J.D. and Amstrong, J.W. (1990) The failure of eld sanitation to reduce infesta-
tion by the mango nut weevil, Cryptorhynchus mangiferae (F.)(Coleoptera: Curcu-
lionidae). Tropical Pest Management 36, 359361.
Hansen, J.D., Armstrong, J.W. and Brown, S. (1989) The distribution of the mango wee-
vil, Cryptorhynchus mangiferae (Coleoptera: Curculionidae), in Hawaii. Proceed-
ings of the Hawaiian Entomological Society 29, 3139.
Harris, E.J., Vargas, R.I. and Gilmore, J.E. (1993) Seasonality in occurrence and distribu-
tion of Mediterranean fruit y (Diptera: Tephritidae) in upland and lowland areas of
Kauai, Hawaii. Environmental Entomology 22, 404410.
Harris, K.M. and Schreiner, I.H. (1992) A new species of the gall midge (Diptera: Ceci-
domyiidae) attacking mango foliage in Guam, with observations on its pest status
and biology. Bulletin of Entomological Research 82, 4148.
Heath, R.R., Epsky, N.D., Guzman, A., Dueben, B.D., Manukian, A. and Meyer, W.L.
(1995) Development of a dry plastic insect trap with food-based synthetic attractant
for the Mediterranean and Mexican fruit ies (Diptera: Tephritidae). Journal of Eco-
nomic Entomology 88, 13071315.
Heath, R.R., Epsky, N.D., Dueben, B.D., Rizo, J. and Jernimo, F. (1997) Adding methyl
substitute ammonia derivates to food-based synthetic attractant on capture of the
Mediterranean and Mexican fruit ies (Diptera: Tephritidae). Journal of Economic
Entomology 90, 15841589.
Hendrichs, J. (1996) Action programs against fruit ies of economic importance: session
overview. In: McPheron, B. and Steck, G. (eds) Fruit Fly Pests: a World Assessment of
their Biology and Management. St Lucie Press, Delray Beach, Florida, pp. 513519.
Hendrichs, J. (2001) Use of the Sterile Insect Technique against Key Insect Pests. Food
and Agriculture Organization (FAO)/International Atomic Energy Agency Division
Pests 355
of Nuclear techniques in Food and Agriculture. Available at: http://www.sustdev.
org/journals/edition.02/download/sdi2_2_2.pdf (accessed 20 July 2007).
Hennesey, M. and Schnell, R.J. (2001) Resistance of immature mango fruits to Caribbe-
an fruit y (Diptera: Tephritidae). Florida Entomologist 84, 318319.
Hernndez-Snchez, G., Sanz-Barbosa, I., Casaa-Giner, V. and Primo-Yfera, E. (2001)
Attractiveness for Ceratitis capitata (Wiedemann) (Dipt., Tephritidae) of mango
(Mangifera indica, cv. Tommy Atkins) airborne terpenes. Journal of Applied Ento-
mology 125, 189192.
Herrera, A.J. and Vias, L.E. (1977) Fruit ies (Diptera: Tephritidae) on mangoes in Chu-
lucanas, Piura. Revista Peruana de Entomoogia 20, 107114.
Hill, D. (1975) Agricultural Insect Pests of the Tropics and their Control. Cambridge
University Press, New York.
Hollingsworth, R.G., Drew, R., Allwood, A.J., Roming, M., Vagalo, M. and Tsatsia, F.
(2003) Host plants and relative abundance of fruit y (Diptera: Tephritidae) species
in the Solomon Islands. Australian Journal of Entomology 42, 95108.
Hsu, J.-C. and Feng, H.-T. (2006) Development of resistance to Sinosad in oriental fruit
y (Diptera: Tephritidae) in laboratory selection and cross-resistance. Journal of
Economic Entomology 99, 931936.
Huang, T.-Ch., Tze-chung, H., Cheng, E.Y., Kao, Ch., Hwang, Y. and Chiang, M. (2006)
Area-wide Control of the Oriental Fruit Fly and Melon Fly in Taiwan. Available at:
www.agnet.org/activities/sw/2006/954198798/paper-868546184.pdf (accessed 20
July 2007).
Husain, M.A. and Pruthi, H.S. (1924) A short note on the life history of the mango hop-
per (Idiocerus spp.) in the Punjab. In: Report Proceedings V of the Entomology
Meeting, Pusa, pp. 252260.
Islam, M.S. and Elegio, D.T. (1997) Effect of time and frequency of insecticide spraying
on the control of mango leafhoppers (Idioscopus spp.). Bangladesh Journal of Ento-
mology 7, 9399.
Iwahashi, O. (1984) The control project of the oriental fruit y in Okinawa. Chinese
Journal of Entomology 4, 107120. (in Chinese)
Jadhav, V. and Dalvi, C. (1987) Bionomics of mango nursery leaf eating caterpillar. Jour-
nal of Maharashtra Agricultural University 12, 384385.
Jeppson, L., Keiffer, H. and Baker, E. (1975) Mites Injurious to Economic Plants. Univer-
sity of California Press, Berkeley, California.
Jhala, R.C., Patel, Z. and Shah, A. (1987) Studies on the relative occurrence of the leaf-
gall midge (Procontarinia matteiana Keiffer and Cecconi) on different varieties of
mango in south Gujarat, India. Tropical Pest Management 33, 277279.
Jhala, R.C., Shah, A.H., Patel, Z.P. and Patel, R.L. (1989) Studies on population dynamics
of mango hopper and scope of spraying in integrated pest management programme.
Acta Horticulturae 231, 597601.
Jirn, L.F. (1993) Plagas del follaje y de paniculas asociadas con el mango en Costa Rica.
In: Memorias Ier Seminario Nacional de Mango. Punta Arenas, Chile, pp. 2226.
Jirn, L.F. (1995) Opciones al uso de insecticidas en mango. In: Garcia, J., Fuentes, G.
and Monge, J. (eds) Opciones al uso Unilateral de Plaguicidas en Costa Rica. Vol.
2. Editorial Euned, San Jos, Costa Rica, pp. 129155.
Jirn, L.F. and Hedstrm, I. (1988) Occurrence of fruit ies of the genera Anastrepha and
Ceratitis (Diptera: Tephritidae), and their host plant availability. Florida Entomolo-
gist 71, 6273.
Jirn, L.F. and Hedstrm, I. (1991) Population uctuations of economic species of Anas-
trepha (Diptera: Tephritidae) related to mango fruiting phenology in Costa Rica.
Florida Entomologist 74, 98105.
J.E. Pea et al. 356
Jirn, L.F. and Soto-Manitiu, J. (1987) Las Moscas de las frutas (Diptera: Tephritidae) en
Costa Rica: situacion actual, analisis y comentario. Agronomia Costarricense 11,
255261.
Joel, D.M. (1980) Resin ducts in the mango fruit: a defense system. Journal of Experimen-
tal Botany 30, 17071718.
Johnson, P.J. (1989) Mango seed weevil (Sternochetus mangiferae F.): identication, biology
and management. University of Idaho Pip Tip 6, 37.
Joubert, P.H., Daneel, M.S. and Grove, T. (2000) Progress towards integrated pest man-
agement (ipm) on mangoes in South Africa. Acta Horticulturae 509, 811817.
Joubert, P., Grove, T., de Beer, M.S. and Steyn, W.P. (2004) Evaluation of kaolin (Surround WP)
in an ipm program on mangoes in South Africa. Acta Horticulturae 645, 493499.
Keifer, H.H., Baker, E.W., Kono, T., Delnado, M. and Styer, W. (1982) An illustrated
guide to plant abnormalities caused by Eriophyid mites in North America. US De-
partment of Agriculture, Agriculture Handbook No. 573, 7475.
Kr, R. and Rosen, D. (1980) Parasites of soft scales (Homoptera: Coccidae) in Israel: an
annotated list. Journal of the Entomological Society of South Africa 43, 113128.
Khan, M.A., Ashfaq, M., Khaliq, A. and Ahmad, A. (2002) Effect of pheromone traps on
the population capture of fruit y species and their infestation in mango orchards at
Multan. Pakistan Entomologist 24, 153157.
Khan, M.A., Ashfaq, M., Akram, W. and Lee, J.-J. (2005) Management of fruit ies (Dip-
tera: Tephritidae) of the most perishable fruits. Entomological Research 35, 7984.
Khanzada, A.G. and Naqvi, K.M. (1985) The optimum time for the control of mango
hopper Idioscopus spp. (Hemiptera: Cicadellidae). Proceedings of the Entomologi-
cal Society of Karachi 14/15, 149156.
King, J.R. and Hennessey, M.K. (1996) Spinosad bait for the Caribbean fruit y (Diptera:
Tephritidae). Florida Entomologist 79, 526531.
Kring, T.J., Yearian, W.C. and Steward, V.B. (1987) Pleuroprucha insulsaria (Lepidoptera:
Geometridae): a potential pest of grain sorghum. Journal of the Kansas Entomo-
logical Society 60, 476477.
Kudagamage, C., de Z. Rajapakse, H.L., Rajapakse, R.H. and Ratnasekara, D. (2001) The
population dynamics and insecticidal control of three leaf hoppers, Amrytodus brevis-
tilus Leth., Idioscopus niveosparus Leth., and Idioscopus clipealis Leth. (Homoptera:
Cicadellidae) in mango. Journal of Entomological Research 25, 121125.
Kumar, S., Rai, A., Patel, C. and Bhatl, R. (1993) Studies on spatial dsitribution of Amri-
todes atckinsoni (Leth.) infesting mango in south Gujarat, India. (Abstract). IV Inter-
national Mango Symposium, p. 122.
Labuschagne, T.I. (1993) Progress with integrated control of the mango scale Aulacaspis
tubercularis Newstead. South African Mango Growers Association Yearbook 13,
134135.
Labuschagne, T.I., Steyn, W.P. and De Beer, M.S. (1995) Voorkoms en moontlike alterna-
tive beheermetodes van vrugtevliee opp mangos. South African Mango Growers
Association Yearbook 15, 102105.
Lakra, R.K., Kharub, W. and Singh, Z. (1980) Pest management systems for the mango
mealybug Drosicha mangifera Green a polyphagous pest of fruit trees in Haryan.
Indian Journal of Entomology 42, 153165.
Laroussilhe, F. de (1980) Le Manguier. G.P. Maisonneuve and Larouse, Paris.
Leblanc, L. and Allwood, A.J. (1997) Mango fruit y (Bactrocera frauenfeldi): why so many
in federated states in Micronesia? In: Allwood, A.J. and Drew, R.A.I. (eds) Manage-
ment of Fruit Flies in the Pacic. Australian Centre for International Agricultural
Research (ACIAR) Proceedings No. 76. ACIAR, Canberra, Australia, pp. 125130.
Leblanc, L., Leweniqila, L., Tau, D., Tumukon, T., Kassim, A. and Hollingsworth, R.
(1997) Can fruit ies be controlled in a village with a mixed orchard? Pacic Island
Pests 357
experiences. In: Allwood, A.J. and Drew, R.A.I. (eds) Management of Fruit Flies in
the Pacic. Australian Centre for International Agricultural Research (ACIAR) Pro-
ceedings No. 76. ACIAR, Canberra, Australia, pp.187191.
Leefmans, S. and van der Vecht, J. (1930) The red-ringed mango caterpillar (in Dutch, English
summary). Korte Mededeelingen van het lustituut voor Plantenziekten, 14 pp.
Leyva, J.L., Browning, H. and Gilstrap, F. (1991) Development of Anastrepha ludens (Dip-
tera: Tephritidae) in several host fruit. Environmental Entomology 20, 11601165.
Lezama-Gutierrez, R., Trujillo, A., Molina-Ochoa, J., Rebolledo-Dominguez, O., Pesca-
dor, A., Lopez-Edwards, M. and Aluja, M. (2000) Virulence of Metarrizium anisopliae
(Deuteromycotina: Hypomycetes) on Anastrepha ludens (Diptera: Tephritidae): labo-
ratory and eld trials. Journal of Economic Entomology 93, 10801084.
Li, L.Y., Wang, R. and Waterhouse, D.F. (1997) The distribution and importance of
major insect pests and weeds of agriculture and plantation forests in southern
China. Chinese Academy of Agricultural Sciences and ACIAR, 185 pp.
Liburd, O.E., Holler, T.C. and Moses, A.L. (2004) Toxicity of Imidacloprid-treated spheres
to Caribbean fruit y, Anastrepha suspensa (Diptera: Tephritidae) and its parasitoid
Diachasmimorpha longicaudata (Hymenoptera: Braconidae) in the laboratory.
Journal of Economic Entomology 97, 525529.
Lindegren, J. and Vail, P. (1986) Susceptibility of Mediterranean fruit y, melon y and
oriental fruit y (Diptera: Tephritidae) to the entomogenous nematode Steinernema
feltiae in laboratory tests. Environmental Entomology 4, 465468.
Lindgreen, S., Borden, J., Gray, D., Lee, P., Palmer, D. and Chong, L. (1982) Evaluation
of two trap log techniques for ambrosia beetles (Coleoptera: Scolytidae) in timber
processing areas. Journal of Economic Entomology 75, 577586.
Loomans, A.J.M., van Lenteren, J.C., Tommasini, M.G., Maini, S. and Riudavets, J. (1995)
Biological control of thrips pests. Wageningen Agricultural University Papers No. 95.
Lpez, F., Chambers, D., Sanchez, M. and Kamasaki, H. (1969) Control of the Mexican
fruit y by bait sprays concentrated at discrete locations. Journal of Economic Ento-
mology 62, 12551257.
Lpez, M., Aluja, M. and Sivinski, J. (1999) Hymenopterous larval-pupal and pupal
parasitoids of Anastrepha ies (Diptera: Tephritidae) in Mexico. Biological Control
15, 119129.
Love, K., Bowen, R. and Paull, R. (2003) Increasing marketable production of exotic trop-
ical fruit with protective covering. Western Region, Sustainable Agriculture Research
and Education, Project Report, Farmer/Rancher, Sustainable Agriculture Research and
Education, Utah State University, Captain Cook, Hawaii, USA.
Lux, S.A., Ekesi, S., Dimbi, S., Mohamed, S. and Billah, M. (2003) Mango-infesting fruit
ies in Africa: perspectives and limitations of biological approaches to their man-
agement. In: Neuenschwander, P., Borgemeister, C. and Langewald, J. (eds) Bio-
logical Control in IPM Systems in Africa. CAB International, Wallingford, UK,
pp. 277293.
Malavasi, A. and Zucchi, R.A. (eds) (2000) Moscas-Das-Frutas de Importncia Econmica
no Brasil: Conhecimento Bsico e Aplicado. Holos Editora, Ribeiro Preto, Brazil.
Mangan, R.L. and Hallman, G.H. (1998) Temperature treatments for quarantine security:
new approaches for fresh commodities. In: Hallman, G.H. and Denlinger, D.L.
(eds) Temperature Sensitivity in Insects and Application in Integrated Pest Manage-
ment. Westview Press, Boulder, Colorado, pp. 201234.
Mangan, R.L. and Moreno, D.S. (2007) Development of bait stations for fruit y popula-
tion suppression. Journal of Economic Entomology 100, 440450.
Mangan, R.L. and Sharp, J.L. (1994) Combination treatments. In: Sharp, J.L. and Hall-
man, G.H. (eds) Quarantine Treatments for Pests of Food Plants. Westview Press,
Boulder, Colorado, pp. 239247.
J.E. Pea et al. 358
Mangan, R.L., Moreno, D.S. and Thompson, G.D. (2006) Bait dilution, spinosad concen-
tration, and efcacy of GF-120 based fruit y sprays. Crop Protection 25, 125133.
Martorell, L.F. (1975) Annotated Food Plant Catalog of the Insects of Puerto Rico. Agri-
cultural Experiment Station, University of Puerto Rico, Ro Piedras, Puerto Rico.
McQuate, G.T., Peck, S.L., Barr, P.G. and Sylva, C.D. (2005) Comparative evaluation of
spinosad and phloxine B as toxicants in protein baits for suppression of three fruit
y (Diptera: Tephritidae) species. Journal of Economic Entomology 98, 11701178.
Messing, R. (1999) Managing Fruit Flies on Farms in Hawaii. Cooperative Extension
Service, College of Tropical Agriculture and Human Resources, University of Hawaii
at Manoa, Hawaii.
Miller, D.R. and Davidson, J. (2005) Armored Scale Insect Pests of Trees and Shrubs
(Hemiptera: Diaspididae). Comstock Publishing Associates, Ithaca, New York.
Mohyuddin, A.I. (1981) A review of biological control in Pakistan. In: Proceedings of the
Second Pakistan Congress of Zoology, Tandojam, Pakistan, pp. 3179.
Mohyuddin, A.I. and Mahmood, R. (1993) Integrated control of mango pests in Pakistan.
Acta Horticulturae 341, 467483.
Montoya, P., Liedo, P., Benrey, B., Cancino, J., Barrera, J.F., Sivinski, J. and Aluja, M.
(2000) Biological control of Anastrepha spp. (Diptera: Tephritidae) in mango or-
chards through augmentative releases of Diachasmimorpha longicaudata (Ash-
mead) (Hymenoptera: Braconidae). Biological Control 18, 216224.
Moore, D. and Cross, A.E. (1992) Competition between two primary parasitoids, Gyranu-
soidea tebygi Noyes and Anagyrus mangicola Noyes, attacking the mealybug Ras-
trococcus invadens Williams and the inuence of a hyperparasitoid Chartocerus
hyalipennis Hayat. Biocontrol Science and Technology 2, 225234.
Moore, D. and Cross, A.E. (1993) Biological control of the fruit tree mealybug, Rastro-
coccus invadens Williams; single or multiple introduction. Acta Horticulturae 341,
433440.
Morin, C. (1967) Cultivo de Frutales Tropicales. Librerias ABC, Lima, Peru, pp. 225229.
Moznette, G. (1922) Insects injurious to the mango in Florida and how to combat them.
Farmers Bulletin 1257.
Murray, R.C. (1991) Arthropod pests of mango in the Caribbean Islands. Review of Crop
Protection. University of the West Indies, Kingston, Jamaica.
Nachiappan, R.M. and Baskaran, P. (1986) Field evaluation of certain insecticidal sprays
against mango leaf-hoppers. Pesticides 20, 4144.
Nafus, D. (1991) Biological control of Penicilllaria jocosatrix (Lepidoptera: Noctuidae)
on mango on Guam with notes on the biology of its parasitoids. Environmental
Entomology 20, 17251731.
Narasinham, M. (1959) Control of mango malformation disease. Current Science
28, 254.
Narasimham, U.A. and Chacko, M.J. (1991) The distribution of some Rastrococcus spp.,
(Homoptera: Pseudococcidae) on mango in India. Bulletin of Entomological Research
81, 445448.
Nascimento, A., Malavasi, A., Morgante, J. and Duarte, A.L. (1992) Hot-water immer-
sion treatment for mango infested with Anastrepha fraterculus, A. obliqua, and
Ceratitis capitata (Diptera: Tephritidae) in Brazil. Journal of Economic Entomology
85, 456460.
Nascimento do, A.S., da Silva Carvalho, da Costa Mendonca, M. and Braga, R. (2002)
Pragas e seu controle. In: de Carvalho, P.J. and de Queiroz Pinto, A.C. (eds) A Cultura
da Mangueira. Embrapa Informao Tecnolgica, Braslia DF, Brazil, pp. 279297.
Nguyen, T.N., Nguyen, V. and Le van, D. (1998) Insect pests on citrus, durian, longan
and mango in selected provinces of the Meking Delta of Vietnam. In: Van Mele, P.
and Nguyen, V.H. (eds) Proceedings of the First Symposium on Fruit Production in
Pests 359
the Mekong Delta Focusing on Integrated Pest Management. Cautho, Vietnam,
pp. 7176 (in Vietnamese).
Norrbom, A.L. (2004) Host Plant Database for Anastrepha and Toxotrypana (Diptera:
Tephritidae: Toxotrypanini). Diptera Data Dissemination Disk (CD-ROM), North
American Dipterists Society, Washington, DC.
Ochoa, R., Aguilar, H. and Vargas, C. (1994) Phytophagous Mites of Central America: an
Illustrated Guide. Centro Agronomico de Investigacion y Enseanza Technical Ser-
vice 6, Turrialba, Costa Rica.
Orankanok, W., Chinvinijkul, S., Thanaphum, S., Sutantawong, M. and Enkerlin, W.R.
(2007) Area-wide integrated control of oriental fruit y (Bactrocera dorsalis, Hen-
del) and Guava fruit y (B. correcta, Bezzii) in Thailand using the sterile insect
technique (SIT). In: Vreysen, M.J.B., Robinson, A.S. and Hendrichs, J. (eds) Area-
wide Control of Insect Pests: From Research to Field Experimentation. Springer,
Dordrecht, the Netherlands, pp. 517526.
Osman, A.A. (1979) Notes on the control of the bud mite Aceria mangiferae (Sayed) in
Egypt (Acarine: Eriophyidae). Bulletin of the Entomological Society of Egypt Eco-
nomic Services 9, 119126.
Ovruski, S., Aluja, M., Sivinski, J. and Wharton, R. (2000) Hymenopteran parasitoids on
fruit infesting Tephritidae (Diptera) in Latin America and the southern United States:
diversity, distribution, taxonomic status and their use in fruit y biological control.
Integrated Pest Management Reviews 5, 81107.
Paderes, A. and Orden, M.E. (2004) Factors Determinating the Adoption of Pruning
and Bagging by Major Mango Producers in the Philippines. Available at: http://
www.kmitl.ac.th/science/Journal/jan2004/MANGOAdopt.pdf (accessed 12 March
2005).
Papadopoulos, N.K., Carey, J.R., Katsoyannos, B.I. and Kouloussis, N.A. (1996) Over-
wintering of the Mediterranean fruit y (Diptera: Tephritidae) in Northern Greece.
Annals of the Entomological Society of America 89, 526534.
Patel, R.K., Patel, S.R. and Shah, S. (1977) Biology of mango hopper, Amritodus atkin-
soni (Leth.) (Jassidae: Hemiptera) in South Gujarat. Indian Journal of Entomology
37, 150153.
Peck, S.L. and McQuate, G.T. (2000) Field tests of environmentally friendly malathion
replacements to suppress wild Mediterranean fruit y (Diptera: Tephritidae) popula-
tions. Journal of Economic Entomology 93, 280289.
Pea, J.E. (1993) Pests of mango in Florida. Acta Horticulturae 341, 395406.
Pea, J.E. and Duncan, R. (1992) Sampling methods for Prodiplosis longila (Diptera:
Cecidomyiidae) in limes. Environmental Entomology 21, 9961001.
Pea, J.E., Duncan, R., Vasquez, T. and Hennessey, M. (1999) Guava arthropod season-
ality and control of fruit ies in south Florida. Proceedings of the Florida State
Horticultural Society 112, 206209.
Pea, J.E., Palevsky, E., Otero-Colina, G., Ochoa, R. and Meister, C.W. (2005) Mango
bud mite, Aceria mangiferae bionomics and control under Florida conditions. Pro-
ceedings of the Florida State Horticultural Society 118, 228234.
Pea, J.E., Duncan, R. and Meister, C. (2006a) Chemical control of owering thrips in-
festing tropical fruits. Proceedings of the Florida State Horticultural Society 119,
2124.
Pea, J.E., Gould, W., Hennessey, M.K., Hallman, G. and Crane, J.H. (2006b) Labora-
tory and eld infestation studies on immature green Tommy Atkins and Keitt
mangoes to determine host status to the Caribbean fruit y (Diptera: Tephritidae).
Proceedings of the Florida State Horticultural Society 119, 1620.
Peng, R.K. and Christian, K. (2004) The weaver ant, Oecophylla smaragdina (Hy-
menoptera: Formicidae), an effective biological control agent of the red banded thrips
J.E. Pea et al. 360
Selenothrips rubrocinctus (Thysanoptera: Thripidae) in mango crops in the Northern
Territory of Australia. International Journal of Pest Management 50, 107114.
Peng, R.K. and Christian, K. (2005a) The control efcacy of the weaver ant, Oecophylla
smaragdina (Hymenoptera: Formicidae), on the mango leafhopper, Idioscopus niti-
dulus (Hemiptera: Cicadellidae) in mango orchards in the Northern Territory. Inter-
national Journal of Pest Management 51, 297304.
Peng, R. and Christian, K. (2005b) Integrated pest management in mango orchards in the
Northern Territory Australia, using the weaver ant Oecophylla smaragdina (Hymenoptera:
Formicidae) as a key element. International Journal of Pest Management 51, 149155.
Peng, R.K. and Christian, K. (2006) Effective control of Jarviss fruit y Bactrocera jarvisi
(Diptera: Tephritidae), by weaver ant, Oecophyla smaragdina (Hymenoptera: For-
micidae), in mango orchards in the Northern Territory of Australia. International
Journal of Pest Management 52, 275282.
Piero, J., Aluja, M., Vzquez, A., Equihua, M. and Varn, J. (2003) Human urine and
chicken feces as fruit y (Diptera: Tephritidae) attractants for resource-poor fruit
growers. Journal of Economic Entomology 96, 334340.
Poinar, G.O., Jr and Hislop, R.G. (1981) Mortality of Mediterranean fruit y adults (Cer-
atitis capitata) from parasitic nematodes (Neoaplectana and Heterorhabditis spp.).
Institute for Research and Cognitive Science, Medical Science. Microbiology, Para-
sitology and Infectious Disease 9, 641.
Prasad, S.N. (1971) The Mango Midge Pests. Cecidological Society, Allahabad, India.
Prasad, V. and Singh, R.K. (1976) Prevalence and control of mango mealy bug Drosicha
stebbingi (Green) in Bihar. Indian Journal of Entomology 38, 214224.
Rahman, K.A. (1940) Important insect pests of the mango and how to control them.
Punjab Fruit Journal 3, 6.
Rai, B., Verma, S. and Kumar, K. (1966) Evaluation of pesticides for control of mango
bud mite, Aceria mangiferae Sayed (Acarina: Eriophyidae). Indian Journal of Ento-
mology 28, 176180.
Rai, B., Shah, A. and Patel, R. (1988) Biology of Oligonychus mangiferus (Tetranychidae:
Acarina), a pest of mango in Gujarat. Gujarat Agricultural University Research Journal
14, 510.
Rebouas, A., Bas, I.V., Martins, J. and Magalhes, O. (1996) Manga, Tecnologia de
Produo e Mercado. Universidade Estadual do Sudoeste da Bahia, Vitria da
Conquista, Brazil.
Robacker, D.C. (2001) Roles of putrescine and 1-pyrroline in attractiveness of technical-
grade putrescine to the Mexican fruit y (Diptera: Tephritidae). Florida Entomologist
84, 679685.
Robacker, D.C. and Wareld, W.C. (1993) Attraction of both sexes of Mexican fruit y,
Anastrepha ludens, to a mixture of ammonia, methylamine, and putrescine. Journal
of Chemical Ecology 19, 29993016.
Robacker, D.C., Martnez, A.J., Garca, J.A., Daz, M. and Romero, C. (1996) Toxicity of
Bacillus thuringiensis to Mexican fruit y (Diptera: Tephritidae). Journal of Eco-
nomic Entomology 89, 104110.
Robacker, D.C., DeMilo, A.B. and Voaden, D.J. (1997) Mexican fruit y attractants: ef-
fects of 1-pyrroline and other amines on attractiveness of a mixture of ammonia,
methylamine, and putrescine. Journal of Chemical Ecology 23, 12631280.
Robinson, A.S. and Hooper, G. (eds) (1989) Fruit Flies. Their Biology, Natural Enemies
and Control. Volumes A and B. Elsevier, Amsterdam, the Netherlands.
Rocha da Silva, C. (2006) Comportamento e comunicaao quimica da broca da manguei-
ra (Hypocryphalus mangiferae Stebb) (Coleoptera: Curculionidae: Scolytinae). Uni-
versidade Estadual do Norte Fluminense, Camposdos Goytacazes, Rio do Janeiro,
Theses, 59 pp.
Pests 361
Rogers, L., French, J. and Elgar, M. (1999) Suppression of plant growth on the mounds of
the termite Coptotermes lacteus Froggatt (Isoptera: Rhinotermitidae). Insects Socieux
46, 366371.
Rubin, A. and Kuslitizky, W. (1992) First record of Ceranisus menes (Hymenoptera:
Eulophidae) in Israel. Phytoparasitica 20, 123124.
Sadana, G.L. and Kumari, M. (1991) Predatory potential of the lyssomanid spider, Lys-
somanes sikkimeri Tikader on the mango hopper, Ideoscopus clipealis (Lethierry).
Entomon 16, 283285.
Said Sajap, A., Amit, S. and Welker, J. (2000) Evaluation of hexaumuron for controlling
the subterranean termite Coptotrermes curvignathus (Isoptera: Rhinotermitidae) in
Malaysia. Journal of Economic Entomology 93, 429433.
Sakimura, K. (1981) A review of Frankliniella bruneri Watson and description of F. kel-
liae, n. sp. (Thysanoptera: Thripidae). Florida Entomologist 64, 483490.
Sankaran, T. and Mjeni, A.M. (1989) Recent studies on the mango leaf-midge Procon-
tarinia matteiana Kieffer and Cecconi (Diptera: Cecidomyiidae) and its parasites in
India and on prospects for biological control of the pest in Oman. Acta Horticultu-
rae 231, 587592.
Sayed, M. (1946) Aceria mangiferae. Bulletin of the Society of Entomology 30, 710.
Schaefer, C.W. and Ahmad, I. (2000) Cotton strainers and their relatives (Pyrrhocoroi-
dea: Pyrrhocoridae and Largidae). In: Schaefer, C.W. and Panizzi, A.R. (eds) Het-
eroptera of Economic Importance. CRC Press, Boca Raton, Florida, pp. 271307.
Schotman, L. (1989) FAO/RLAC Plant Quarantine Programme. Data sheet on mango
seed weevil. FAO Accession XF8988239, 23 pp. http://www.fao.org/agris/search
(accessed 1 September 2008).
Schreiner, I. (1987) Mango shoot caterpillar control on mango owers, 1985. Insecticide
and Acaricide Tests 12, 94.
Schreiner, I. (1991) Procontarinia sp. nov.: a mango pest long misdiagnosed as anthra-
cnose. In: Agricultural Development American Pacic, Crop Protection Conference
Proceedings 1989, Hawaii Institute Tropical Agriculture and Human Resources,
Research Extension Series 134, 5355.
Secretariat of the Pacic Community (2005) Fruit Flies in Palau. Pest Advisory Leaet
No. 44, Palau, Pacic Islands.
Shah, A.H., Jhala, R.C., Patel, G.M. and Patel, S.H. (1983) Evaluation of effective dose
of monocrotophos for the control of mango hopper Amritodus atkinsoni Lethierry
(Cicadellidae: Homoptera) by injection method and its comparison with foliar
application. Gujarat Agricultural University Research Journal 9, 1418.
Sharp, J.L., Ouye, M.T., Thalman, R., Hart, W.G., Ingle, S.J. and Chew, V. (1988) Submer-
sion of Francis mango in hot water as a quarantine treatment for the West Indian
fruit y and the Caribbean fruit y (Diptera: Tephritidae). Journal of Economic Ento-
mology 81, 14311436.
Sharp, J.L., Ouye, M.T., Ingle, S.J., Hart, W.G., Enkerlin, E., Celedonio, H., Toledo, J.,
Stevens, L., Quintero, E., Reyes, J. and Schwarz, A. (1989a) Hot-water quarantine
treatment for mangoes from the State of Chiapas, Mexico, infested with Mediterra-
nean fruit y and Anastrepha serpentina (Wiedemann) (Diptera: Tephritidae). Jour-
nal of Economic Entomology 82, 16631666.
Sharp, J.L., Ouye, M.T., Hart, W.G., Ingle, S.J., Hallman, G., Gould, W. and Chew, V.
(1989b) Immersion of Florida mangoes in hot water as a quarantine treatment
for Caribbean fruit y (Diptera: Tephritidae). Journal of Economic Entomology 82,
186188.
Sharp, J.L., Ouye, M.T., Ingle, S.J. and Hart, W.G. (1989c) Hot water quarantine treat-
ment for mangoes from Mexico infested with Mexican fruit y and West Indian fruit
y (Diptera: Tephritidae). Journal of Economic Entomology 82, 16571662.
J.E. Pea et al. 362
Shaw, J. (1961) Airplane applications of malathion bait spray for Mexican fruit y con-
trol. Journal of Economic Entomology 54, 600601.
Shaw, J. and Spisshakoff, L.M. (1958) Insecticidas para el control de la mosca Mexicana
de la fruta en citricos y mango. Memorias Primer Congreso Naciona de Entomolo-
gia Fitopatologia Escuela Nacional Agricola Chapingo. Escuela Nacional Agricola
de Chapingo, Texcoco, Mexico, pp. 205217.
Sheikh, A.D., Bashir, A. and Jilani, G. (2008) Incidence and Effects of Fruit Fly on Mango
Quality in Punjab. Technology and Transfer Institute, Faisalabad, 3 pp.
Shellie, K.C. and Mangan, R.L. (2002a) Hot water immersion as a quarantine treatment
for large mangoes: articial versus cage infestation. Journal of the American Society
for Horticultural Science 127, 430434.
Shellie, K.C. and Mangan, R.L. (2002b) Cooling method and fruit weight: efcacy of hot
water quarantine treatment for Mexican fruit y (Diptera: Tephritidae) in mango.
HortScience 37, 910913.
Shukla, R.P. and Prassad, V.G. (1984) Evaluation of some insecticides against mango
hopper, Idioscopus clypealis (Lethierry)(Homoptera: Cicadellidae). Indian Journal
of Agricultural Science 54, 677681.
Shukla, R.P. and Tandon, P.L. (1985) Bio-ecology and management of the mango weevil,
Sternochetus mangiferae (Fabricius) (Coleoptera: Curculionidae). International
Journal of Tropical Agriculture 3, 293303.
Shukla, R.P., Tandon, P.L. and Singh, S.J. (1984) Baculovirus a new pathogen of mango
nut weevil, Sternochetus mangiferae (Fabricius) (Coleoptera: Curculionidae). Cur-
rent Science 53, 593599.
Shukla, R.P., Tandon, P.L. and Suman, C.L. (1988) Intra-tree distribution of eggs and dia-
pausing adults of the stone weevil. Acta Horticulturae 231, 566570.
Silva, Q.M. and Cavalcante, R.D. (1977) Vinsonia stellifera (Westwood 1871),
(Hom. Coccidae), on various plants in the state of Cear, Brazil. Fitossanidade 2,
2526.
Silveira, da, R. (1960) Contribucao ao studo do Hypocryphalus mangiferae (Coleoptera:
Scolytidae). PhD thesis, Escuela Agricultura de So Paulo, So Paulo, Brazil.
Singh, G. (1991) Loss assessment, ecology and management of mango fruit y, Dacus
sp. Acta Horticulturae 291, 425436.
Singh, S.K. and Singh, G.B. (2003) Management of termite infestation in mango orchard
by cultural practices and organic amendments. Indian Journal of Agricultural Re-
search 37, 148150.
Sivinski, J.M. (1996) The past and potential of biological control of fruit ies. In: McPher-
on, B.A. and Steck, G.J. (eds) Fruit Fly Pests. A World Assessment of their Biology
and Management. St Lucie Press, Delray Beach, Florida, pp. 365375.
Sivinski, J.M., Calkins, C.O., Baranowski, R.M., Harris, D., Brambila, J., Diaz, J., Bums,
R.E., Holler, T. and Dodson, D. (1996) Suppression of Caribbean fruit y (Anas-
trepha suspensa (Loew) Diptera: Tephritidae) population through releases of the
parasitoid Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae).
Biological Control 6, 177185.
Sivinski, J., Aluja, M. and Lpez, M. (1997) Spatial and temporal distributions of parasi-
toids of Mexican Anastrepha species (Diptera: Tephritidae) within the canopies of
fruit trees. Annals of the Entomological Society of America 90, 604618.
Sivinski, J., Piero, J. and Aluja, M. (2000) The distributions of parasitoids (Hymenoptera)
of Anastrepha fruit ies (Diptera: Tephritidae) along an altitudinal gradient in Vera-
cruz, Mexico. Biological Control 18, 258269.
Sohi, A.S. and Sohi, A.S. (1990) Mango leafhoppers (Homoptera: Cicadellidae) a
review. Journal of Insect Science 3, 112.
Pests 363
Soto-Maniti, J., Jirn, L. and Hernndez, R. (1987) Chemical control and ecological
observations of fruit ies of the genus Anastrepha Schiner (Diptera: Tephritidae) on
mango. Turrialba 37, 245251.
Srivastava, R.P. (1981) Comparative efcacy of various insecticidal dusts against mango
mealybug eggs. Indian Journal of Entomology 43, 225229.
Srivastava, R.P. (1998) Pests of mango. In: Srivastava, R.P. (ed.) Mango Cultivation. Inter-
national Bood Distributing, Charbagh, Lucknow, India, pp.175299.
Srivastava, R. and Tandon, P. (1986) Natural occurrence of two entomogenous fungi
pathogenic to mango hopper, Idioscopus clypealis Leth. Indian Journal of Plant
Pathology 4, 121123.
Srivastava, S. and Singh, N.B. (2004) Survey of termites infesting mango orchards at four
locations in Bareilly, Uttar Pradesh. Biological Memoirs 30, 4851.
Stark, J.D., Vargas, R. and Miller, N. (2004) Toxicity of spinosad in protein bait to three
economically important tephritid fruit y species (Diptera: Tephritidae) and their par-
asitoids (Hymenoptera: Braconidae). Journal of Economic Entomology 97, 911915.
Steck, G.J. (2003) Mango Fruit Fly, Marula Fruit Fly, Ceratitis cosyra (Walker) (Insecta: Dip-
tera: Tephritidae). Available at: http://edis.ifas.u.edu/IN563 (accessed 20 July 2007).
Steiner, L.F. and Lee, R.K. (1955) Large area tests of male annihilation method for orien-
tal fruity control. Journal of Economic Entomology 48, 311317.
Steiner, L.F., Mitchell, W.C., Harris, E., Kozuma, T. and Fujimoto, M. (1965) Oriental fruit
y eradication by male annihilation. Journal of Economic Entomology 58, 961964.
Subba Rao, B.R. (1986) Mangoma spinidorsum gen. et sp. n. (Hymneoptera: Eurytomi-
dae) associated with leaf galls. Bulletin of Entomological Research 76, 389392.
Subramanian, C.K. (1925) A note on the life history of Cryptorhynchus mangiferae Fab.
Madras Agricultural Department Yearbook. Madras Agricultural Department, Ma-
dras, India, pp. 2936.
Summanwar, A. and Raychoudhury, S. (1968) The role of the eriophyid mite (Aceria
mangiferae) in the causation of mango malformation. Indian Phytopathology 21,
463464.
Sweet, M.H. (2000) Seed and chinch bugs (Lygaeoidea). In: Schaefer, C.W. and Panizzi,
A. (eds) Heteroptera of Economic Importance. CRC Press, Boca Raton, Florida,
pp. 143264.
Swirski, E., Wysoki, M. and Izhar, Y. (2002) Subtropical pests of Israel. Fruit Board of
Israel and Institute of Plant Protection, The Volcani Center, Tel Aviv, 284 pp.
Syed, R.A., Ghani, M.A. and Murtaza, M. (1970) Studies on trypetids and their natural
enemies in West Pakistan. III. Dacus (Strumeta) zonatus (Saunders). Technical Bul-
letin of the Commonwealth Institute of Biological Control 13, 116.
Tan, K.-H. (2000) Area-Wide Control of Fruit Flies and Other Insect Pests. Penerbit Uni-
versiti Sains Malaysia, Pulau Pinang, Malaysia.
Tandon, P.L. and Lai, B. (1979) Studies on the chemical control of the mango hopper,
Idioscopus clypealis Leth. (Cicadellidae: Homoptera). International Pest Control
21, 69.
Tandon, P.L. and Verghese, A. (1985) World List of Insect, Mite and Other Pests of Man-
go. Technical Document of the Indian Institute for Horticultural Research 5. Indian
Institute for Horticultural Research, Bangalore, India.
Tandon, P.L., Lal, B. and Rao, G.S. (1983) Prediction of mango hopper Idioscopus cly-
pealis (Leth.) population in relation to physical environmental factors. Journal of
Entomology 8, 257261.
Tandon, P.L., Verghese, A. and Prasada, G.S. (1989) Spatial distribution, sampling plan
and appropriate transformation of the mango hopper, Idioscopus niveoparsus (Lethi-
erry) (Homoptera: Cicadellidae). Giornale Italiano di Entomologia 4, 235242.
J.E. Pea et al. 364
Tenakanai, D. (1997) Fruit y fauna in Papua New Guinea. In: Allwood, A.J. and Drew,
R.A.I. (eds) Management of Fruit Flies in the Pacic. Australian Centre for Interna-
tional Agricultural Research (ACIAR) Proceedings No. 76. ACIAR, Canberra, Austra-
lia, pp. 8794.
Thomas, P., Kannan, A., Degwekar, V.H. and Ramamurthy, M.S. (1995) Non-destructive
detection of seed weevil-infested mango fruits by X-ray imaging. Post Harvest Biol-
ogy and Technology 5, 161165.
Thomson, F.C. (ed.) (2005) BioSystematic Database of World Diptera, Version 7.5. Avail-
able at: http://www.diptera.org/biosys.htm (accessed 5 November 2007).
Tigvattnanont, S. (1988) Biological and autoecological studies of the mango leaf-cutting
weevil, Deporaus marginatus Pascos (Coleoptera: Attelabidae). Khon Kaen Agricul-
tural Journal 16, 5162.
Toledo, J. and Lara, J. (1996) Comparison of the biology of Anastrepha obliqua reared in
mango (Mangifera indica L.) and in mombin (Spondias mombin) infested under
eld conditions. In: McPheron, B. and Steck, G. (eds) Fruit Fly Pests: a World Assess-
ment of their Biology and Management. St Lucie Press, Delray Beach, Florida,
pp. 359362.
Toledo, J., Liedo, P., Williams, T. and Ibarra, J. (1999) Toxicity of Bacillus thuringiensis
-exotoxin to three species of fruit ies (Diptera: Tephritidae). Journal of Economic
Entomology 92, 10521056.
Toledo, J., Rasgado, M.A., Ibarra, J.E., Gmez, A., Liedo, P. and Williams, T. (2006)
Infection of Anastrepha ludens following soil applications of Heterorhabditis
bacteriophora in a mango orchard. Entomologia Experimentalis et Applicata 119,
155162.
Tween, G. (1993) Fruit y control and eradication program management: factors inu-
encing action criteria and program design. In: Aluja, M. and Liedo, P. (eds) Fruit
Flies: Biology and Management. Springer, New York, pp. 308310.
Uechi, N., Kawamura, F., Tokuda, M. and Yukawa, J. (2002) A mango pest, Procon-
tarinia mangicola (shi) comb. Nov. (Diptera: Cecidomyiidae), recently found in
Okinawa, Japan. Applied Entomology and Zoology 37, 589593.
Umeya, K. and Hirao, J. (1975) Attraction of the jackfruity, Dacus umbrosus F. (Diptera:
Tephritidae) and lace wing, Chrysopa sp. (Neuroptera: Chrysopidae) to lure traps
baited with methyl eugenol and cuelure in the Philippines. Applied Entomology
and Zoology 10, 6062.
United States Department of Agriculture (USDA) (1981) Plant Pest News 1.
van Dine, D.L. (1906) The mango weevil (Cryptorhynchus mangiferae Falor.). Hawaii
Agricultural Experiment Station Bulletin 17, 111.
van Halteren, P. (1970) Notes on the biology of the scale insect Aulacaspis mangiferae
Newst. (Diaspididae: Hemiptera) on mango. Ghana Journal of Agricultural Science
3, 8385.
van Mele, P., Cuc, T.T. and van Huis, A. (2001) Farmers knowledge, perceptions and
practices in mango pest management in the Mekong Delta, Vietnam. International
Journal of Pest Management 47, 716.
van Mele, P., Vayssires, J.-F., van Tellingen, E. and Vrolijks, J. (2007) Effects of an African
weaver ant, Oecophylla longinoda, in controlling mango fruit ies (Diptera:
Tephritidae) in Benin. Journal of Economic Entomology 100, 695701.
van Whervin, L.W. (1968) Citrus Weevils of Jamaica and Some of their Parasites. Techni-
cal Bulletin No. 1. Citrus Research Unit, University of the West Indies, St Augus-
tine.
van Wyk, M., Al Adawi, A., Khan, I., Deadman, M., Al Jahwari, A., Wingeld, B., Ploetz,
R. and Wingeld, M. (2007) Ceratocystis manginecans sp. nov., causal agent of a
Pests 365
destructive mango wilt disease in Oman and Pakistan. Fungal Diversity 27, 213
230.
Van Zyl, E., Kotze, J.M. and Steyn, P.L. (1988) Isolation of Xanthomonas campestris pv.
Mangifera indica from gall y induced lesions on mango leaves. Phytophylactica
20, 8990.
Vannire, H., Didier, C., Rey, J.Y., Diallo, T.M., Kita, S. and Sangar, M. (2004) La
mangue en Afrique de lOuest francophone: les systmes de production et les iti-
nraires techniques. Fruits 59, 383398.
Vargas, R.I., Miller, N.W. and Prokopy, R.J. (2002) Attraction and feeding responses of Med-
iterranean fruit y and a natural enemy to protein baits laced with two novel toxins,
phloxine B and spinosad. Entomologia Experimentalis et Applicata 102, 273282.
Varma, A., Lele, V.C., Raychoudhuri, S.P., Ram, A. and Sang, A. (1974) Mango malforma-
tion: a fungal disease. Phytopathologische Zeitschrift 79, 254257.
Vayssires, J.-F. and Kalabane, S. (2000) Inventory and uctuations of the catches of Dip-
tera Tephritidae associated with mangoes in Coastal Guinea. Fruits 55, 259270.
Vayssires, J.-F., Sanogo, F. and Noussourou, M. (2004) Inventaire des espces de
mouches des fruits (Diptera: Tephritidae) infodes au manguier au Mali et essais
de lutte raisonne. Fruits 59, 316.
Vayssires, J.-F., Goergen, G., Lokossou, O., Dossa, P. and Akponon, C. (2005) A new
Bactrocera species in Benin among mango fruit y (Diptera: Tephritidae) species.
Fruits 60, 371377.
Veeresh, G.K. (1989) Pest problems in mango world situation. Acta Horticulturae 231,
551565.
Veeresh, G.K., Rajagopal, D. and Kumar, N.G. (1989) Management of termites in mango
orchard. Acta Horticulturae 231, 633638.
Verghese, A., Rao, G. and Prasada-Rao, G. (1985) Sequential plan for mango leaf hop-
per, Idioscopus clypealis Lethierry. Entomology 10, 285290.
Verghese, A., Tandon, P. and Prasada-Rao, G. (1988) Ecological studies relevant to the
management of Thrips palmi Karny on mango in India. Tropical Pest Management
34, 5558.
Verghese, A., Tandon, P.L. and Stonehouse, J.M. (2003) Economic evaluation of inte-
grated management of oriental fruit y Bactrocera dorsalis (Diptera: Tephritidae) in
mango in India. Crop Protection 23, 6163.
Verghese, A., Nagaraju, D.K., Jayanthi, P.D.K., Vasudev, V. and Madhura, H.S. (2004)
Is azadirachtin useful in the management of the mango stone weevil (Sternoche-
tus mangiferae Fabricius)? Journal of Food Agriculture and Environment 2,
213216.
Viraktamath, C.A. (1997) A revision of the Idiocerine leafhopper genus Amritodus
(Hemiptera:Cicadellidae) breeding on mango. Entomon 22, 111117.
Viraktamath, S. and Viraktamath, C.A. (1985) New species of Busoniominus and
Idioscopus (Homoptera: Cicadelidae: Idiocerinae) breeding on mango in south India.
Entomon 10, 305311.
Vyas, R.V., Patel, J.J., Godhani, P.H. and Yadav, D.N. (1993) Evaluation of green musca-
rdine fungus (Metarrhizium anisopliae var. anisopliae) for control of mango hopper
(Amritodus atkinsoni). Indian Journal of Agricultural Sciences 63, 602603.
Waddell, W.J., Crooks, N.H. and Carmichael, P.L. (2004) Correlation of tumours with
DNA adducts from methyl eugenol and tamoxifen in rats. Toxicological Sciences
79, 3840.
Waite, G.K. and Huwer, R.K. (1998) Host plants and their role in the ecology of the
fruitspotting bugs Amblypelta nitida Stl and Amblypelta lutescens (Hemiptera: Co-
reidae). Australian Journal of Entomology 37, 340349.
J.E. Pea et al. 366
Waite, G.K. and Petzl, N. (1994) A search for egg parasites of Amblypelta spp. (Hemiptera:
Coreidae) in south-east Queensland. In: Menzel, C. and Croswell, A. (eds) Ma-
roochy Research Station Report No. 7, pp. 1314.
Wang, X.G., Jarjees, E.A., McGraw, B.K., Bokonon-Ganta, A.H., Messing, R.H. and
Johnson, M.W. (2005) Effects of spinosad-based fruit y bait GF-120 on tephritid
fruit y and aphid parasitoids. Biological Control 35, 155162.
Waterhouse, D.F. (1998) Biological control of insect pests: South-East Asian prospects.
Australian Centre for International Agricultural Research Monograph No. 51, Can-
berra, Australia, 548 pp.
Watson, J.R. (1917) Thysanoptera of Florida. The Florida Buggist 1, 5377.
Wharton, R.A. (1978) Classical biological control of fruit-infesting Tephritidae. In: Rob-
inson, A.S. and Hooper, G. (eds) Fruit Flies. Their Biology, Natural Enemies and
Control. Vol. B. Elsevier, Amsterdam, the Netherlands, pp. 303313.
Wharton, R.A. and Marsh, P.M. (1978) New world Opiinae (Hymenoptera: Braconidae)
parasitic on Tephritidae (Diptera). Journal of Washington Academy of Science 68,
147165.
White, I.M. and Elson-Harris, M. (1992) Fruit Flies of Economic Signicance. CAB Inter-
national, Wallingford, UK.
Whitwell, A.C. (1993) The pest/predator/parasitoid complex on mango inorescences in
Dominica. Acta Horticulturae 341, 421432.
Woodruff, R.E. (1985) Citrus weevils of Florida and the West Indies: preliminary report
on systematics, biology and distribution (Coleoptera: Curculionidae). Florida Ento-
mologist 68, 370379.
Woods, B., Lacey, I.B., Brockway, C.A. and Johnstone, C.P. (2005) Hosts of Mediterra-
nean fruit y Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) from Broome
and the Broome Peninsula, Western Australia. Australian Journal of Entomology 44,
437441.
Wysoki, M., Ben-Dov, Y., Swirski, E. and Izhar, Y. (1993) The arthropod pests of mango
in Israel. Acta Horticulturae 341, 452466.
Yazdani, S.S. and Mehto, D.N. (1980) A note on the control of mango hopper, Amrito-
dus atkinsoni Leth. at Dholi. Indian Journal of Entomology 42, 275276.
Yee, W. (1987) The mango in Hawaii. Cooperative Extension Service, University of
Hawaii Circular 388, 1922.
Zaheruddeen, S.M. and Sujatha, A. (1993) Record of Deanolis albizonalis (Hampson)
(Pyralidae: Odontinae) as mango fruit borer in Andhra Pradesh. Journal of the Bom-
bay Natural History Society 90, 528.
Zucchi, R.A. (1988) Moscas-das-frutas (Diptera: Tephritidae) no Brasil: Taxonomia, Dis-
tribuio Geogrca e Hospedeiros. Fundao Cargill, Campinas, Brazil.
Zucchi, R.A. (2000) Espcies de Anastrepha, sinonmias, plantas hospedeiras e para-
sitides. In: Malavasi, A. and Zucchi, R.A. (eds) Moscas-das-frutas de Importncia
Econmica no Brasil: Conhecimento Bsico e Aplicado. Holos Editora, Ribeiro
Preto, Brazil.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 367
11 Crop Production: Propagation
S. Ram
1
and R.E. Litz
2
1
GB Pant University of Agriculture and Technology, Pantnagar, India
2
University of Florida, Florida, USA
11.1 Introduction 367
11.2 Seed Propagation 369
Monoembryonic seed 369
Polyembryonic seed 369
11.3 Vegetative Propagation 374
Preparation of rootstock 376
Attached methods 378
Detached methods of grafting 380
Effect of rootstock 385
Top and double working 386
Rooting 387
Micropropagation 391
11.4 Comparative Performance of Trees Propagated by Different Methods 391
11.5 Conclusions 392
11.1 Introduction
Mango reproduces naturally by seed, although this is rarely a horticultural
practice, particularly for monoembryonic cultivars. Instead, vegetative prop-
agation is utilized to preserve the unique phenotypes of superior selections,
and has been based upon grafting and rooting methods, growing plants from
nucellar seedlings of polyembryonic mangoes and micropropagation (Fig.
11.1). Grafting of mango can be either attached or detached. In the former, the
scion is not severed from the mother plant until its union with the rootstock
is complete, i.e. approach grafting, tongue, saddle and root grafting. In the
latter, the scion is removed from the mother tree and then joined with the
rootstock, and both are allowed to grow prior to cutting of the rootstock
above the graft union. Detached methods include rind or crown grafting,
cleft or wedge grafting, whip or splice grafting, side grafting, veneer grafting,

S
.

R
a
m

a
n
d

R
.
E
.

L
i
t
z
3
6
8
Propagation
Sexual Asexual
Monoembryonic
seed
Polyembryonic
seed
Grafting Rooting Micropropagation
Attached Detached
Approach
Tongue
Saddle
Root
Cleft or wedge
Rind or crown
Veneer
Notch or inlaying
Side
Splice or whip
Budding
Layering Cutting
Ground
Air
Pot
Organogenesis
Shoot tip culture
Somatic
embryogenesis
Fig. 11.1. Methods of mango propagation.
Crop Production: Propagation 369
notch or inlaying, T or shield budding, forkert budding, modied forkert
budding, patch budding, modied patch budding, chip budding, etc. Root-
ing methods include layering and cutting techniques. Although in vitro meth-
ods for regenerating mango have been reported via somatic embryogenesis,
organogenesis and shoot tip culture (see Litz et al., Chapter 18, this volume),
their practical application for propagation has not yet been demonstrated.
The various methods of mango propagation are shown in Figs 11.211.8.
Mango propagation has been discussed recently by Singh and Singh (1998),
Galn-Saco (1999) and Neto et al. (2002).
11.2 Seed Propagation
Monoembryonic seed
Seed propagation does not ensure true-to-type plant reproduction of
monoembryonic mango selections; however, it was extensively used before
vegetative methods for mango propagation were known (Singh, 1960). Large
seedling orchards were planted during the medieval period of Indian history
until vegetative propagation was introduced into India by the Portuguese in
the late 15th century. Monoembryonic seeds contain only a single sexual
embryo, and a single plant grows from a seed of a monoembryonic cultivar.
Polyembryonic seed
Polyembryony is common in mango cultivars that originated and are grown
in the moist tropics. In Knight et al. (Chapter 3, this volume), there is a survey
of the major polyembryonic and monoembryonic mango cultivars. Trees
from nucellar seedlings are identical to the mother plant (see Mukherjee and
Litz, Chapter 1, this volume). In polyembryonic seed, only one embryo is
zygotic in origin; it either degenerates or produces weak and stunted seed-
lings (Maheshwari et al., 1955; Sachar and Chopra, 1957; Singh, 1960; Stur-
rock, 1968). Approximately three to eight seedlings normally originate from
a single polyembryonic seed (Garner and Chaudhri, 1976), although 30 or
more embryos have been recorded in a single polyembryonic mango seed
(Juliano, 1934, 1937). Nucellar seedlings can be distinguished from the zygotic
seedling on the basis of their greater vigour c.1 month after germination.
Polyembryonic mango cultivars have traditionally been seed propagated
in South-east Asia and in some tropical Latin American countries. In many
mango-growing areas, polyembryonic and monoembryonic mango selec-
tions are grafted onto uniform, nucellar seedling rootstock. Zygotic and
nucellar seedlings can be morphologically similar. Therefore, detection and
separation of nucellar seedlings is important in breeding programmes involv-
ing controlled pollinations. Different protein and DNA markers have been
used. Degani et al. (1993) and Schnell and Knight (1993) demonstrated that
isozymes and RAPD markers can be used to distinguish zygotic embryos from
S. Ram and R.E. Litz 370
(a) Approach grafting
(b) Tongue grafting
(c) Saddle grafting
(d) Root grafting
Fig. 11.2. Grafting methods of mango propagation: (a) approach grafting; (b) tongue
grafting; (c) saddle grafting; (d) root grafting.
Crop Production: Propagation 371
(a) Cleft grafting
(b) Soft wood grafting
(c) Epicotyl grafting
(d) Splice (whip) grafting
Fig. 11.3. Grafting methods of mango propagation: (a) cleft grafting; (b) soft wood
grafting; (c) epicotyl grafting; (d) splice or whip grafting.
S. Ram and R.E. Litz 372
nucellar seedlings. Eiadthong et al. (1999), using single sequence repeats (SSRs)
could distinguish among mango cultivars, and separate them according to
their geographic origin; however, monoembryonic and polyembryonic geno-
types could not be differentiated. Amplied fragment length polymorphism
(AFLP) markers have been used for cultivar identication (Kashkush et al.,
(a) Side grafting
(b) Notch grafting
(c) Veneer grafting
(d) Rind or crown grafting
Fig. 11.4. Grafting methods of mango propagation: (a) side grafting; (b) notch grafting
or inlaying; (c) veneer grafting; (d) rind or crown grafting.
Crop Production: Propagation 373
(a) Shield budding
(b) Patch budding
(c) Modified patch budding
(d) Forkert budding
Fig. 11.5. Budding methods of mango propagation: (a) shield budding; (b) patch
budding; (c) modied patch budding; (d) forkert budding.
S. Ram and R.E. Litz 374
2001) and Gonzalez et al. (2002) identied a range of inter-simple-sequence-
repeats (ISSR) primer sequences with potential for cultivar identication.
11.3 Vegetative Propagation
Vegetative or asexual propagation preserves the genotype and phenotype of
elite selections by means of grafting, rooting and micropropagation. Trees
(a) Modified forkert budding
(b) H-budding
(c) Chip budding
Fig. 11.6. Budding methods of mango propagation: (a) modied forkert budding;
(b) H-budding; (c) chip budding.
Crop Production: Propagation 375
propagated by grafting onto seedling rootstocks usually ower after 3 or 4
years in comparison to the 510 years for seedling trees and they are gener-
ally smaller than seedling trees because they begin to bear fruit earlier. The
oldest method for vegetatively propagating mango is probably the attached
method, also known as inarching or approach grafting.
(a) Modified chip budding
(b) Flap budding
(c) Window budding
Fig. 11.7. Budding methods of mango propagation: (a) modied chip budding;
(b) ap budding; (c) window budding.
S. Ram and R.E. Litz 376
Preparation of rootstock
Rootstocks are either zygotic or nucellar seedlings. Nucellar seedling root-
stocks are clonal, and have many advantages over heterogeneous monoem-
bryonic seedlings as rootstocks (see Crane et al., Chapter 13, this volume).
Clonal rootstocks have generally been selected for specic soil types and
(a) Top working
(b) Double working
Fig. 11.8. Top working (a) and double working (b) of mango.
Crop Production: Propagation 377
stress tolerance, and behaviour of the scion cultivar on clonal rootstock is highly
predictable. Monoembryonic seedlings are generally used as rootstocks in
India. Polyembryonic Turpentine and Kensington Pride seedlings are used
as rootstocks in Florida and Australia, respectively, whereas polyembryonic
Sabre, 13-1 and 4-9 seedlings are used for rootstocks in Israel. Throughout
South-east Asia, polyembryonic seedlings are used for rootstocks, e.g. poly-
embryonic Saing and Thalapt in Myanmar (Grant and William, 1949). In
Senegal, polyembryonic Amelioree, De Cameron and Madame Francis are
used as rootstocks (Furon, 1966). In Brazil, seedlings of polyembryonic
Carabao, Mangua dAgua and Pico are used for rootstock; they are consid-
ered to have resistance to Ceratocystis mbriata (Neto et al., 2002); the IAC series
(IAC100-104) are used to control seca de mangueira (Neto et al., 2002).
Freshly extracted mango seeds from ripe fruits germinate with higher
frequency (7691%) than those from overripe, rm or green fruits (Shant and
Saproo, 1974). Seeds with large cotyledons from seedling trees germinate
earlier, store better and seedling growth is more vigorous than seeds with
small cotyledons (Naik, 1949; Simao, 1960). Naik (1941) noted differences in
germination and vigour in monoembryonic seedling progenies of different
parentages. Seedlings of Chausa reportedly produced better height and
girth than Langra and seedling vigour was closely related to the weight of
the seed stone (Giri and Chaudhery, 1966). Mango seeds are recalcitrant, and
lose viability (Ledin and Ruehle, 1954; Ruehle and Ledin, 1955; Singh, 1960)
within 30 days. Therefore, seed should be collected and sown within 1 week
after collection. About 80% of seed germination occurs if they are sown
within 1 month of extraction (Stephens, 1960; Sturrock, 1968; Chacko and
Singh, 1971). Parisot (1988) recorded that seeds that are free of pulp could be
stored for up to 84 days at 15C on sterile cotton with deionized water. Husk-
ing or decortication of the hard endocarp of the seed improves germination
(Paul and Guneratnam, 1938; Simao, 1960; Chauran et al., 1979; Abdel-Galil,
1992). At the time of sowing, treatment with fungicides also improves germi-
nation, particularly of dehusked seeds (Chauran et al., 1979). Seeds should be
sown without injuring the cotyledons and preferably early in the season.
Seedbed preparation
The standard practice for seed germination varies. In India, seedbeds are
used, whereas in most other countries, seeds are planted individually in pots
or plastic bags. Nurseries are usually under partial shade to prevent exces-
sive evaporation from the plant and soil and to protect seedlings from inclem-
ent weather; shade is particularly useful in areas with dry, hot climates.
Seedbeds can also be used in the open provided the soil is adequately moist.
Highly sandy or loamy soils are generally unsuitable for mango seed germi-
nation. Soil should be well drained with plenty of organic matter, and seeds
should be sown c.15 cm apart in a special seedbed in which 25 cm top soil of
the bed is excavated and the bed bottom is hardened with concrete or lined
with an iron or polyethylene sheet to prevent elongation of the taproot and
to encourage brous root development (Stephens, 1948). Germination can be
expedited if seeds are soaked in 20100 mg/l gibberellic acid (GA
3
) for 2448 h
S. Ram and R.E. Litz 378
or sprayed with 50300 mg/l GA
3
; however, these seedlings are usually
unsatisfactory for grafting. Seeds should be planted with convex edge up at
a depth of 57 cm (Singh, 1960; Bakhshi, 1963) or with a small portion exposed
above the ground level (Pope, 1929; Ruehle and Ledin, 1955; Bakhshi, 1963;
Anonymous, 1975). Soil moisture should be maintained. Seeds germinate
23 weeks after planting (Ruehle and Ledin, 1955; Ahmed and Rashid, 1961;
Singh and Jawanda, 1962). Seedlings that are 1-month-old with coppery
leaves and with their stones attached survive transplanting better than older
plants without much damage to their root system (Ruehle and Ledin, 1955).
The taproot should be slightly pruned at the time of planting.
Nursery
Seedlings should be transplanted without disturbing the roots to nursery
beds or pots at least 1 month before grafting. Either the crown of the seed-
lings should be pruned or the distal half of the leaves should be removed to
reduce transpiration. Seedbed-grown plants generally have better germina-
tion rates and lower production costs than those grown directly in nursery
beds. Seedlings should be actively growing at the time of grafting. Success in
grafting depends on several factors, such as age and vigour of rootstock and
scion, diseases, insect pests, environment and the skill of the grafter.
Attached methods
Approach grafting is expensive, cumbersome, requires a long time to pro-
duce grafts and the success rate is often low. Other methods are now used
because they are cheaper and easier.
Approach grafting
A scion shoot, while still attached to the parent plant, can be grafted onto the
seedling rootstock by making a long cut (57 cm) of similar length and depth
on one side of both rootstock and scion through the cambium and slightly
into the wood (Fig. 11.2a). The cut surfaces of rootstock and scion are pressed
rmly together and secured with waxed string, rafa or grafting tape. After
union, the scion is severed below the graft union and the rootstock above the
union. Several years ago, approach grafting was standard practice for propa-
gating mango in many countries, but has been largely replaced by detached
grafting methods, except for parts of India. For approach grafting, seedlings
4560 cm long with a diameter of 1.25 cm are utilized (Asadullah and Khan,
1960; Singh, 1960). They are planted in pots or plastic bags containing soil
mixture, and are kept beneath the mother tree. Alternatively, the ball of earth
around the root system of the seedling grown for rootstock is tied in jutties
made of dried grass, paddy straw or plastic bags and planted under the
mother tree or raised to the shoot height of the mother tree on a raised plat-
form or directly suspended from branches of the mother tree. The seedling
rootstock can be a few months- to 2-years-old (Naik, 1949; Garg, 1954; Ahmed,
1960; Singh, 1960). Juvenile seedlings whose leaves are copper coloured and
Crop Production: Propagation 379
with the seed still attached to the seedling are often used. The root system is
covered with moist sphagnum moss or similar material and wrapped in
polyethylene to prevent drying. Production costs are less with young seed-
lings than with older material.
In the subtropics, approach grafting in spring can achieve 88100% suc-
cess with some cultivars (Asadullah and Khan, 1960; Majhail and Singh,
1962a, b; Talukdar and Ahmed, 1965). Majhail and Singh (1962b) observed
that seedling size and age do not affect grafting success in spring, but that
larger seedlings yielded better success from July to September. Approach
grafting should be avoided during winter months when plants are dormant.
In the tropics, mangoes can be approach grafted year-round (Ahmed, 1960;
Singh, 1960). Approach grafting should be done when the trees are in active
growth (Singh, 1960; Rao, 1967). In low rainfall areas, approach grafting
should be done at the time of the onset of rains, while in regions of heavy
rainfall, it should be done soon after the rainy season is over, provided tem-
peratures are not <15C, the critical temperature for the growth of mango
trees (Whiley, 1993). Healthy trees should be used for scion wood. The suc-
cess of approach grafting is cultivar dependent, for example Langra scions
establishes better than either Dashehari or Chausa (Asadullah and Khan,
1960). Better success has also been reported with seedlings of 1.31.6 cm
diameter than with smaller shoots (Giri, 1966). Variations of approach graft-
ing include multistock, inorescence, top working and adjuvant grafting. At
one time, south Indian nurserymen would approach graft a single large scion
with two to ve rootstocks (Patwardhan and Deshmukh, 1931; Singh, 1960).
Grafts with large and old scions having several branches become top heavy, and
fail to thrive. New growth from inorescences that are approach grafted is veg-
etative. Approach grafting has also been used to top work old seedling trees with
cultivars (Singh, 1960). Adjuvant grafting refers to approach grafting of diseased
or damaged trees in order to rejuvenate them on new rootstocks.
Tongue grafting
Tongue grafting is practised with thicker rootstock than scion. The rootstock
is rst cut as in approach grafting; a second cut is made into the wood of the
stem approximately halfway down the rst cut in a sloping direction point-
ing tongue upwards. A similar cut is made in the scion shoot. Both cuts are
made in such a manner that one ts into the other tightly (Fig. 11.2b). Root-
stock and scion are tied together with grafting tape and a graft union is com-
plete in c.12 months. The scion is then severed from the mother tree and the
rootstock is decapitated above the graft union. Tongue grafting is slightly
more complicated than simple approach grafting; however, the success rate
is better because three surfaces of the cambium layer are involved in the graft
union (Burns and Prayag, 1921).
Saddle grafting
With saddle grafting, the rootstock is decapitated c.25 cm above ground level,
and two upward sloping cuts are made on the sides of the rootstock to form
a 57 cm long wedge on its cut end. On the stem of the scion shoot, a tongue
S. Ram and R.E. Litz 380
is made and the outer surface of the tongue is not pared. The wedge of the
rootstock is then tted into the groove of the tongue (Fig. 11.2c). Subsequently,
the joint is secured with grafting tape and the scion shoot is separated from the
mother tree after the union is complete (Singh, 1960). Saddle grafting is easier
to perform than tongue grafting but is more difcult than approach grafting.
Root grafting
Root grafting is similar to bench grafting. A seedling plant (c.l year old) is
potted in a U-shaped pot with a notched edge (78 cm deep and 5 cm wide)
(Singh, 1960). The 710 cm taproot is projected through the notch and the pot
itself contains the root. The seedling above the collar is staked, and placed in
the shade for 11.5 months for establishment. Seedlings are approach grafted
with the scion (Fig. 11.2d), and the graft is separated from the mother tree
after the union is complete. Grafts are maintained in the shade and watered
regularly. Naik (1941, 1949) reported 100% success with root grafting.
Detached methods of grafting
Cleft or wedge grafting
Singh (1960) suggested that cleft grafting can be used with rootstocks of
greater diameter than the scion and also used for top working (Fig. 11.3a).
Rootstock of 57 cm or more in diameter should be used and cleft grafted
after decapitating the stock c.45 cm above the ground. The decapitated root-
stock is split to a depth of about 5 cm through the centre of the stem with a
knife and a mallet. After the knife is removed, a hard wooden wedge is
inserted to keep it open for the subsequent insertion of the scion. Scions
(1520 cm long) from a terminal shoot >3 months old is wedged securely (to
a depth of 67 cm). The cleft of the scion is slipped into the split of the stock
that has been forced open with the wooden wedge. The thicker edge of the
cleft is placed towards the outer edge of the rootstock in such a way that the
back of the rootstock and scion meet rmly. Another scion is also inserted
diametrically opposite the rst one. The wooden wedge is removed and cut
surfaces are sealed with grafting wax. The graft union requires c.2 months.
Cleft or wedge grafting is appropriate for replacing the crown of young trees;
however, with young seedling rootstock, this has also been used for large-scale
propagation. In Brazil, Pinheiro et al. (1970) reported that cleft grafting was
more successful (97%) than four other grafting methods tried. Cleft grafting is
easier to use than veneer grafting and has a similar rate of success (Ram, 1993).
Success can be improved when leaves are retained below the point where the
rootstock is severed, when the grafting portion of the rootstock is a new ush
and when the stem is pinkish green; however, the scion should be mature.
Soft wood grafting
Insertion of the scion into the bronze-coloured stem of the rootstock has been
referred to as soft wood grafting (Fig. 11.3b). Amin (1974, 1978) reported 100%
success if leaves are not removed below the graft union. Seedling rootstocks
Crop Production: Propagation 381
of various ages can be used; however, success decreases with age, i.e. 80%
with 1-year-old rootstocks and less with 6-month-old seedlings (Gaur, 1984;
Reddy and Melanta, 1988; Kulwal and Tayde, 1989; Panickar and Desai,
1989; Gandhoke, 1993). This technique is easy and is effective in dry, hot
weather and in areas with low precipitation.
Epicotyl or stone grafting
Epicotyl grafting involves cleft grafting of scions onto 2-week-old seedling
rootstock (Traub and Auchter, 1934). Predefoliation of the scion is not essen-
tial, although it is widely practised (Fig. 11.3c). Moderate temperature and
high relative humidity (RH) are important for success. A 23 cm long slant-
ing cut is made in the epicotyl with a matching cut on the proximal portion
of the scion in the splice method for grafting. For wedge grafting, a vertical
cut 45 cm long is made in the decapitated epicotyl to receive the wedge-
shaped scion. The scion and rootstock are tied together with grafting tape,
and the small grafted plant is planted immediately and watered. The scions
sprout within a month. The success rates of splice and wedge grafting meth-
ods are reported to be >50% (Bhan et al., 1969; Gupta et al., 1988; Dhunaga
et al., 1989; Gunjate, 1989; Jinturkar and Narwadkar, 1989; Kulwal and Tayde,
1989; Madalgeri et al., 1989; Narwadkar and Anserwadekar, 1989; Radhamony
et al., 1989; Singh et al., 1989; Srivastava, 1989; Patil et al., 1991); under green-
house conditions in the subtropics, success can be >90% (Ram, unpublished).
Grafted plants are maintained under partial shade in a moderate and moist
environment. In the subtropics, growth is initially slow and can require c.2
years to attain the size of a veneer graft; however, in the tropics, growth is
much more rapid.
Splice or whip grafting
This is the easiest method for propagating mango and is widely used in the
subtropics. The rootstock should be c.12 years old, the diameter of both
rootstock and scion should be 0.61.0 cm. The rootstock is severed c.1020 cm
above the soil and a diagonal cut (34 cm long) is made at the distal end of
the rootstock (Fig. 11.3d). A similar slanting cut is made on the proximal end
of the scion and the cut surfaces of both the rootstock and the scion are bound
together with grafting tape. The union (usually 90% successful) takes place
in c.2 months (Ahmed, 1974). Torres (1949, 1960) used this method with 3- to
9-month-old seedling rootstock with a high success rate. Majumder and
Rathore (1970) reported 50% success with splice grafting with 2-week-old
seedlings and up to 60% success with 30-day-old seedlings; however, sur-
vival of the grafts was poor.
Side grafting
Side grafting, also known as the Nakamura method (Fig. 11.4a), was for-
merly popular (Burns and Prayag, 1921; Pope, 1929; Pope and Storey, 1933;
Walters, 1932; Tanaka, 1939; Lynch and Mustard, 1950; Thrower, 1954; Ruehle
and Ledin, 1955; Lynch and Nelson, 1956; Ahmed, 1960; Singh, 1960; Serpa,
1964; Rao, 1967). This method is supposed to be highly effective in mild
S. Ram and R.E. Litz 382
weather in the absence of strong winds, intense heat and heavy rain (Rao,
1967), and success has also varied (50100%) with different cultivars (Veera-
raghavan, 1945). In India, side grafting is generally used in humid, coastal
areas with 1.01.5 cm diameter rootstock. Scions should be c.10 cm long and
defoliated c.1 week prior to grafting. The rootstock age has varied from 4 to
36 months with 7290% success (Mulat, 1959; Kashyap et al., 1972; Faruque
and Fakir, 1973; Kanwar and Bhajwa, 1974; Ben-yaacov, 1976). The scion is
inserted into the wedge and tied with grafting tape. The union is complete
after 23 months. The top of the rootstock can be removed above the graft
union after new scion growth occurs.
Notch grafting or inlaying
This method is used in the humid tropics with >8 cm diameter rootstock
(Singh, 1960). The rootstock is decapitated c.45 cm above the ground level.
V-shaped notches (45 cm) are made at equal distances, depending upon the
thickness of the rootstock, from the periphery of the cut surface. The proxi-
mal end of each scion is tted into each notch of the rootstock (Fig. 11.4b) and
scions are secured to the rootstock with grafting wax. The union is normally
complete in c.23 months (Singh, 1960).
Veneer grafting
This grafting technique was rst described by Lynch (1941), and has been
widely adopted (Cooper and Furr, 1945; Ruehle and Ledin, 1955; Mukherjee
and Majumder, 1961, 1962, 1964; Wolfe, 1963; Ahmed, 1964; Serpa, 1964; Jagirdar
et al., 1968; Bhambota et al., 1971; Prasad et al., 1973; Ramirez Diaz, 1973; Gun-
jate and Limaye, 1978; Ram and Bist, 1982; Pinto et al., 2004). Veneer grafting
is usually more effective than other methods of grafting (Bhambota et al.,
1971; Prasad et al., 1973), with approximately 90% success (Ram and Bist,
1982). For veneer grafting, 3- to 6-month-old 1.01.5 cm diameter scion shoots
with lush green leaves should be defoliated 315 days prior to grafting
(depending upon the season), leaving the petioles attached (Mukherjee and
Majumder, 1961; Singh and Srivastava, 1979a, b; Ram and Bist, 1982; Rajan
and Sinha, 1987; Bajpai et al., 1989). Prior defoliation may not be required under
humid conditions (Gunjate et al., 1976) or when extremes of temperature and
RH do not occur. The scion should be 1015 cm long, although smaller scions
(510 cm) have also been used (Ram and Bist, 1982). Older shoots can also be
used (Mukherjee and Majumder, 1961; Jagirdar et al., 1968; Ram and Bist,
1982). Scions stored for 8 days at ambient temperature (2535C) in moist
sphagnum moss covered with polyethylene can still be grafted with a 50%
success rate during March through to July (Ram and Bist, 1982). Cultivar
differences may be important.
The use of seedling rootstock at different ages (3 months to 2 years) has
resulted in 40100% success, depending upon the season, scion maturity,
predefoliation period, storage condition of scion, etc. (Zill, 1951; Subra, 1954;
Mukherjee and Majumder, 1961, 1962, 1964; Ahmed, 1964; Serpa, 1964;
Jagirdar and Bhatti, 1968; Bhambota et al., 1971; Prasad et al., 1973; Ramirez Diaz,
1973; Gunjate and Limaye, 1978; Ram and Bist, 1982). Dipping or smearing
Crop Production: Propagation 383
growth regulator onto the cut surfaces of the scion and rootstock has been
ineffective (Kulkarni et al., 1989). The rootstock is prepared for veneer grafting
by making a slanting cut (5 cm long); an oblique cut is then made at the base of
the rst cut, so that a piece of wood along with bark is removed (Fig. 11.4c).
The base of the scion wood is then tted into the rootstock so that the cut
surfaces with the cambium layers of scion and rootstock are in contact with
each other. The rootstock and scion are secured together with grafting tape.
The union takes place after 1.52.0 months. After scion growth begins, the
rootstock is removed above the graft union. Some workers have recom-
mended rst ringing and then removing the rootstock shoot 12 weeks later,
while others have partially cut through the rootstock shoot before severing
the shoot completely a few days later. After new leaves of the scion turn
green, the rootstock may be removed completely together with grafting tape
(Mukherjee and Majumder, 1961; Ram and Bist, 1982).
Rind or crown grafting
Rind or crown grafting has been used in the Americas and India. Its success
depends upon several factors, such as season, rootstock, scion maturity, mod-
erate temperature and high RH (Gangolly et al., 1957; Popenoe, 1959; Singh,
1960). The rootstock is decapitated 30 cm above the soil. A longitudinal cut
(8 cm) is made in the bark from the top of the decapitated rootstock, down-
wards, without injuring the wood below the bark. The bark is loosened from
the wood with a wooden wedge or spatula and a freshly prepared 3-month-
old scion is inserted after creating a wedge c.6 cm long at the proximal end of
the scion. The ap of bark is then secured over the inserted scion with grafting
tape. Several scions may be inserted into a single thick rootstock (Fig. 11.4d).
High RH around the graft is important for success. The graft union occurs in
c.1 month, but the grafting tape should be removed from the grafts only after
the leaves of the scion turn green. Rind or crown grafting is a cumbersome
method and is not often used in commercial nurseries; however, it can be
useful for top working.
Budding
Budding involves the grafting of a single bud, with or without wood attached,
with the rootstock. Budding methods are referred to by the shape of the bud,
ap of the rootstock covering the bud before insertion, etc. The various meth-
ods for budding are illustrated in Figs 11.511.7. The inserted bud eventually
develops as the crown. Rootstock above the bud is severed after bud estab-
lishment. Budding provides a stronger union and the graft union occurs ear-
lier than with other grafting methods. Both rootstock and scion should be
actively growing, and excision of the bud is easy. Stimulation of the bud prior
to its excision by predefoliation, application of growth regulators and gir-
dling can improve the efciency of the procedure. A bud is normally selected
from a 3- to 4-month-old scion shoot, and is budded onto rootstock stems of
1.01.25 cm diameter c.2530 cm above ground level where the rootstock is
greyish green, except in cases of very juvenile rootstock. Generally 1- and
2-year-old rootstocks are better for grafting than younger ones. The bud is
S. Ram and R.E. Litz 384
completely covered by grafting tape for a few weeks. Bud growth can be
stimulated by removing grafting tape c.2 weeks after grafting.
Shield or T budding
Shield or T budding of mango (Fig. 11.5a) occurs with varying success,
ranging from 3294% (Singh and Khan, 1943, 1946; Singh and Ram, 1946;
Singh, 1946; Singh, 1960; Singh and Srivastava, 1961). In northern India and
Pakistan, shield or T budding is generally most effective from March to July
(India) and MarchApril (Pakistan). Defoliating the scion shoot for 1015
days before budding (Pope, 1929; Parsons, 1931; Khan, 1960; Rao, 1967; Teaotia
and Maurya, 1970) and girdling of the scion shoot several weeks prior to
budding can increase bud take (Anonymous, 1950). Scion cultivar differences
also appear to be important, for example success is better with Langra than
with Aman Dashehari and Chausa (Ahmed and Chatha, 1960); however,
success is higher with Sindhri than with Langra and Banganpalli (Jagirdar
and Ali, 1968). Marked differences in success have also been reported for dif-
ferent agroecological zones in the same season: 45% in the dry hot plains of
the Punjab and 90% in relatively cool areas (Bajwa and Ram, 1946).
Temperature, RH and storage media inuence bud storage (Walters,
1932; Sundara Rao, 1956; Srivastava, 1963a, b; Rao, 1967). Storage of bud-
wood in moist sphagnum moss for 10 days is generally satisfactory. Sealing
the cut ends of budwood with melted wax and storage in a thermos ask
with ice for 48 h does not adversely affect budding (Singh and Khan, 1943).
Buds can be inserted in inverted T cuts using 2- to 5-year-old rootstock
(Higgins, 1910; Ruehle and Ledin, 1955). T budding in situ is also successful
(Singh and Khan, 1942, 1946; Bajwa and Ram, 1946; Khan, 1960) in March
April and AugustOctober (Ullah and Ali, 1955). The bud should be secured
with grafting tape. Soule (1971) described four stages in the formation of the
bud union: (i) pre-callus with wound periderm up to 4 days after budding;
(ii) callus proliferation 8 days after budding; (iii) cambial bridge between
rootstock and scion and vascular tissue differentiation 12 days after budding;
and (iv) the healed union after 24 weeks. If the buds are green up to 15 days
after budding, the grafting tape should be removed and the rootstock should
be girdled 10 cm above the union to stimulate the bud to sprout. The root-
stock is nally removed above the budding point after a growth ush of the
scion (Luthra and Sharma, 1946; Ruehle and Ledin, 1955; Hayes, 1957). Budded
plants are transplanted only after at least one seasons growth.
Patch budding
Patch budding (Figs 11.5b and c) (Verma, 1942; Ullah and Ali, 1955; Singh,
1960; Moreuil, 1963; Teaotia, 1963; Jauhari and Singh, 1970; Teaotia and
Maurya, 1970; Prasad and Singh, 1972; Prasad et al., 1973) is used as an alterna-
tive to approach grafting (Garner and Chaudhri, 1976). Under subtropical
north Indian conditions, March and May through to July is the optimum period
for patch budding. In tropical Malaysia, patch budding is successful from
December through to February. The scion is defoliated 2 weeks prior to budding.
Rootstocks are severed above the budding point 12 weeks after budding.
Crop Production: Propagation 385
Patch budding is apparently superior to forkert budding with Langra and
Dashehari (Teaotia, 1963).
Forkert budding
Over 90% grafting success has been reported with forkert budding (Fig.
11.5d) in tropical South-east Asia and Sri Lanka (Paul and Guneratnam,
1937a, b). In Maharashtra, India during July and August, success with this
technique using 1-year-old rootstock was 6070% (Gandhi, 1942), and 100%
with Langra and Dashehari (Singh and Srivastava, 1962; Teaotia, 1963). A
modied forkert budding method (Fig. 11.6a) can be more effective than
shield budding (Garner and Chaudhri, 1976). H-budding is another modi-
cation of the forkert method (Singh, 1960), and derives its name from an
H-shaped cut made in the rootstock (Fig. 11.6b).
Chip budding
For chip budding, 2- to 3-month-old seedlings are used (Fig. 11.6c). The bud
is excised with a piece of wood attached to it. Graft union can occur 34
weeks after budding because rootstock tissues are partially undifferentiated
and possess a broadly exposed cambium. The method is very efcient, and is
widely used (Lynch and Nelson, 1949, 1956; Soule, 1954; Subra, 1954; Ruehle
and Ledin, 1955; Ahmed, 1960; Bhan et al., 1969; Rajput and Haribabu, 1971),
often with modications (Lynch and Mustard, 1950; Singh, 1960) (Fig.
11.7a).
Flap budding
The excised bud shape is boat-shaped instead of rectangular (Fig. 11.7b). This
procedure has been used in South-east Asia (Naik, 1949; Garner and Cha-
udhri, 1976).
Window budding
This technique is similar to ap budding except that a window is created in
the ap for the bud so that ap does not press the bud while being tied (Fig.
11.7c). This method has been used in Queensland, Australia. The bud union
occurs in c.4 weeks (Stephens, 1948).
Effect of rootstock
Relative advantages of polyembryonic rootstocks
Swamy et al. (1972) described the results of a rootstock trial involving six
polyembryonic rootstocks for Baneshan and four for Neelum. Based on
growth and fruit yield from 1942 to 1965, Pahutan and Goa rootstocks were
recommended for Neelum and Olour rootstock was recommended for
Baneshan. Oppenheimer (1960) reported the highest yield for Haden mango
occurred on polyembryonic Sabre rootstock among three different rootstocks
tried. George and Nair (1969) conducted a rootstock trial using polyembryonic
Chandrakaran and Bappakai and monoembryonic Puliyan rootstocks for
S. Ram and R.E. Litz 386
Bennett, Alphonso and Baneshan. Chandrakaran and Bappakai grew
more vigorously and produced higher yields during the rst 6 years of
growth compared to monoembryonic Puliyan rootstock. In south India,
polyembryonic Olour and Bappakai and monoembryonic rootstocks were
evaluated using Neelum as a scion (Gowder and Irulappan, 1970). Bap-
pakai was recommended as a rootstock based on fruit yield and uniform
growth of Neelum.
Performance of Dashehari on 24 rootstocks (ten polyembryonic and 14
monoembryonic) was reported by Rajan and Pandey (1991a, b). Rootstocks
strongly affected tree vigour with tree height ranging from c.4.0 to 6.0 m after 14
years of growth. ST-9 and Latra rootstocks stimulated the most vigorous
growth. Ambalvi, Pahutan, Olour, Nakkare, Mylepelian, Rumani, Willard,
Mundappa, Pahutan, Ambalvi and Moovandan caused dwarng. Stem
swelling was recorded with Mahmuda Vikarabad rootstock; however, the scion
girth varied with rootstock. Rumani and Ambalvi rootstocks caused less scion
girth (0.750.8 m) than the vigorous ST-9 and Latra rootstocks (0.790.84 m).
Crown spread was also greater with vigorous rootstocks than with dwarng
rootstocks. The canopy of Dashehari trees on ST-9 and Latra rootstock was
2.15 and 2.25 times greater than on Rumani, respectively. Fruit yield varied
from 8 kg to 25 kg/tree. Dwarng rootstock resulted in higher fruit yield per unit
canopy volume without much change in fruit quality. Polyembryonic rootstocks
Ambalvi, Mylepelian, Olour and Villaikolamban were compared with
Dashehari seedling rootstock using Dashehari as the scion (Jauhari et al., 1972;
Singh and Singh, 1976). The Dashehari seedling rootstock was most vigorous
and produced higher yields, whereas Villaikolamban resulted in the least
vigour and yield. Villaikolamban caused dwarng of Alphonso and low
yields (Anonymous, 1994), and has been recommended as a rootstock for high
density Alphonso orchards, based upon higher yield/m
3
of canopy.
Neelum fruit quality on Bappakai rootstock was better compared to
Neelum grown on Olour and monoembryonic seedlings (Gowder and Iru-
lappan, 1970). Larger Neelum fruits having high total soluble solids devel-
oped on Pahutan rootstock in comparison with Goa, Olour and Salem
rootstocks (Swamy et al., 1972). Madu and Gurih rootstocks delayed fruit-
ing in Indonesia compared to Gurung, Kopjor, Budadaja, Nanas and
Saigon (Kusumo et al., 1971). Polyembryonic rootstock 13-1 has been dem-
onstrated to tolerate calcareous soil containing 20% calcium carbonate and
saline irrigation water containing >600 ppm chloride (Stoler, 1976; Kadman
et al., 1978). Gazit and Kadman (1980) grew Maya on 13-l rootstock on cal-
careous soils with up to 20% lime and 250 ppm chloride. Ann and Gomera
polyembryonic rootstocks also show salt tolerance (Galn-Saco, 1993).
Top and double working
Top working is used to rejuvenate unproductive trees and for grafting on
seedling trees (Fig. 11.8a). Two methods are used for top working mango.
Branches or main limbs are cut back to 30 cm during winter months or with
Crop Production: Propagation 387
the onset of spring. The new shoots are budded or grafted in the following
summer or autumn, usually by shield budding or veneer grafting. Singh
(1990) suggested that half of a tree should be top worked in the rst year and
the other half in the second year, although top working of a complete tree in
a single year has been highly successful (Lynch, 1941; Ruehle and Ledin,
1955; Ahmed, 1960; Singh, 1960). Cutting back of limbs in October or early
November and veneer grafting in MarchApril has been recommended in
Florida (Carmichael, 1957/58; Miner, 1962); in Uttar Pradesh (India), top
working on 8-year-old mango trees was more successful during June (Singh,
1960). After top working, trees come into bearing within 2 years (Furon and
Plaud, 1972).
Grafting on an already grafted or budded tree is referred to as double
working (Singh, 1960), and double-worked trees therefore consist of three
genetically different parts, i.e. the rootstock, interstock and crown (Fig. 11.8b).
In Florida, old plantings of Haden have been double worked with Tommy
Atkins (Mitchell, 1971). Double working can also be used to repair trees
affected by disease or cold injury and to develop a strong framework. Kala-
pady has reportedly been used as an interstock in order to dwarf Langra
(Sen, 1939). In one trial, 12 interstocks were grafted on two rootstocks for
Dashehari; rootstock-interstock combinations signicantly affected tree
height and vigour. Fruit yield was also inuenced by rootstock in all the com-
binations. Kurukkan interstock on ST-9 rootstock yielded more fruit in
comparison with Ambalvi on the same rootstock. The maximum yield was
obtained with Nakkare/Chandrakaran and Ambalvi/Chandrakaran;
whereas the lowest yield was obtained with Ambalvi/ST-9. Iyer et al.
(1990) reported varying success (2073%) with double-worked Langra.
Rooting
Mango air layers have been induced to root either while they are still attached
to the parent tree or by cutting shoots into pieces containing one to several
buds. Several rooting methods have been successful with mango.
Layering
Layering is used chiey to propagate monoembryonic mango cultivars on
their own roots. Rooting methods involve the initial training of the main
stem. Shoots close to ground level are ringed and covered with soil, while
other shoots are covered with soil mixture or wet sphagnum moss at the
ringed portion and wrapped with polyethylene to maintain moisture (Singh,
1960; Majumder and Mukherjee, 1961; Mukherjee and Majumder, 1963).
Auxins such as -naphthalene acetic acid (NAA) and indole butyric acid
(IBA) are generally essential in order to induce rooting even after ringing.
Ground layering
Vigorous, 1- or 2-year-old shoots are selected near the base of the parent tree.
The greyish-green bark portion of the shoot (35 cm length) is ringed without
S. Ram and R.E. Litz 388
injuring the wood. On the proximal end of the shoot just above the ring,
growth regulators are applied and the ringed portion is buried in the soil.
Depending upon the growth regulators and their concentration, ringed
shoots (or layers) generate roots within 23 months. The rooted shoot can
then be detached from the mother tree and planted.
Mound layering or stooling
This technique involves the use of juvenile shoots or shoots with induced
juvenility (through heading back and etiolation followed by use of growth
regulators), and is more efcient than simple layering (Majumder and
Mukherjee, 1961). One-year-old mango seedlings should be decapitated
2530 cm above ground level in FebruaryMarch, and several new shoots
will develop from the stem by May. When leaves of the new shoots turn
green, the shoots are ringed at their proximal end and treated with 5000 ppm
IBA in lanolin paste just above the ring. A soil mound is created around the
base of ringed shoots. Rooting occurs after 12 months and rooted layers can
be detached from the parent plant and established in individual pots. Each
layer can generate between four and seven roots. Stooling has been success-
ful with many cultivars using growth regulators and rooting cofactors in
stool beds (Ram et al., 1981; Sirohi and Ram, unpublished data). Dwarf culti-
vars root poorly, but can be stimulated to form thin and brous roots by
applying auxin, for example IBA (15,00030,000 ppm) and NAA (250010,000
ppm) mixtures in lanolin paste with or without rooting cofactors, such as caf-
feic acid (0.046M) and catechol (0.046M). Caffeic acid appears to be highly
synergistic to IBA.
Pot layering
With pot layering, the ringed portion of the shoot is maintained in a soil mix-
ture in special pots at a convenient height within the tree canopy. This can be
done only during periods of high RH, but the success is low (c.1520%)
(Singh, 1960).
Air layering
Although air layering or marcottage is one of the oldest methods for propagat-
ing mango, good success has been elusive (Grove, 1947; Garg, 1954; Rangacha-
rlu and Rao, 1956; Rao and Rao, 1956; Choudhury, 1959; Jauhari and Nigam,
1962; Rao et al., 1963; Mukherjee and Majumder, 1963; Srivastava, 1963a, b;
Mukherjee and Bid, 1965; Sen and Bose, 1966; Bid and Mukherjee, 1969; Basu
et al., 1972; Prasad and Singh, 1972; Nez-Elisea et al., 1992). Juvenility, high
RH and moderate temperature are important factors for air layering mango
(Singh, 1954; Rao et al., 1963; Rao, 1967; Chhonkar and Singh, 1972; Singh
et al., 1979; Ram and Sirohi, 1982a). Cultivar effects on air layering are also
pronounced; and vigorous cultivars root better than dwarf cultivars (Garg,
1954; Rao and Rao, 1956; Rao, 1967; Basu et al., 1972; Ram and Sirohi,
1982b).
Layering success can be improved with auxins, such as 23% indole ace-
tic acid (IAA) and 0.52.0% NAA (Thakurta and Dutt, 1941; Singh and Teotia,
Crop Production: Propagation 389
1951; Srivastava, 1963a, b; Nez-Elisea et al., 1992). Although Nez-Elisea
et al. (1992) used NAA alone, NAA and IBA mixtures not only improve root-
ing but also accelerate the production of brous roots, which increases sur-
vival. Etiolated layers just above the girdle require less auxin than non-etiolated
branches. Root regeneration in air layers requires up to 90 days, but emer-
gence of roots is rst noticeable a month after layering. Sen et al. (1961) and
Mukjerjee and Bid (1965) observed that IBA depletes sugar in the bark and
wood of the rooting zones. Ram and Sirohi (1982a) studied the interaction of
rooting cofactors involved with air layering of mango. Application of 0.01
0.08M IBA in lanolin paste increased root numbers from 2.2 to 2.8 with
Dashehari, but root number could be increased from 2.8 to 6.0 with 0.01
0.10M catechol. Survival was correlated with increased numbers of roots;
however, IBA and catechol did not show any additive or synergistic effect on
air layering. July was the best month for rooting of layers (>90% success) and
winter months were the worst.
Rajan and Ram (1989) showed that 0.0573M NAA and 0.05 10
2
M chlo-
rogenic acid had a synergistic effect on rooting of layers and the number of
roots formed during March when applied separately with 0.0735M IBA. In
October, NAA and chlorogenic acid showed synergism with respect to num-
ber of roots produced, but their effect on rooting was additive. Pyrogallol (0.06
10
2
M) and 6-benzylaminopurine (0.22 10
3
M) also had a synergistic effect on
root regeneration during October and March. IBA increases endogenous root-
ing cofactors and lowers rooting inhibitors, resulting in early rooting.
Cuttings
The potential of mango cuttings to form roots depends on maturity of the
cutting, rooting medium, temperature, RH, etc. Girdling or etiolation of
stems for a few weeks and use of growth regulators have improved rooting
frequency (Thakurta and Dutt, 1941; Gardner and Piper, 1943; Van Overbeek
and Gregory, 1945; Said and Shoushan, 1945; Khalifa et al., 1964; Dijkman,
1950; Ledin and Ruehle, 1954; Gowda, 1962; Ahmed, 1964; Sen and Bose,
1964; Mukherjee et al., 1966, 1967; Maurice, 1969; Sen et al., 1969; Ali and
Nazir, 1970; Prasad and Pathak, 1970; Bid and Mukherjee, 1972; Bid et al.,
1972). Rooting is improved with cuttings from the lower bole of the tree
rather than from the upper bole. Mukherjee et al. (1966) reported that cut-
tings from 4-year-old trees root more readily than those from 10-year-old
trees, alhough Sen et al. (1968) had better success with older shoots. Hard-
wood cuttings reportedly root better (Hussain, 1953; Benincasi, 1970) than
semi-hardwood cuttings (Singh and Teaotia, 1951). Mango cuttings should
be c.15 cm long and 1.21.8 cm girth with between three and ve buds (Garner
and Chaudhri, 1976). Retention of one to two leaves at the apex of the cut-
tings has helped rooting, and is attributed to the presence of rooting cofactors
in the leaves.
In India and Pakistan, cuttings are generally made during the spring
months (Hussain, 1953). Cuttings can be stored for a short time before apply-
ing the rooting treatment. Khalifa et al. (1964) rinsed cuttings for 24 h with
running tap water, which increased their shelf life. Longevity of cuttings can
S. Ram and R.E. Litz 390
also be increased by inserting them in a solution consisting of 100 ppm IBA,
10 mg/litre vitamin B
1
, 2% ammonium sulphate ((NH
4
)
2
SO
4
) and 2% sucrose.
Cuttings can be stored at 4C for 20 days. Rooting of hardwood cuttings
under mist can be improved with IBA and NAA (Lambourne, 1935; Cooper,
1936; Thakurta and Dutt, 1941; Cooper and Stoutemyer, 1945; Dijkman, 1950;
Jauhari and Nigam, 1958; Sen and Bose, 1964; Mukherjee et al., 1965; Basu
et al., 1966; Sen et al., 1969; Ali and Nazir, 1970; Rajan and Ram, 1983a, b). IBA
increases reducing and non-reducing sugars, and stimulates a favourable
carbon to nitrogen (C:N) ratio in ungirdled cuttings comparable to girdled
cuttings without IBA. Application of NAA to the ring 1 week before cuttings
are made and dipping their basal ends in an IBA solution at the time of
detaching, increases rooting efciency (Van Overbeek and Gregory, 1945; Sen
et al., 1969). Low pH potting mixtures (pH 4.57.0) reportedly are better for
rooting than higher pH mixtures (Kains and McQuestan, 1955). Reddy and
Majumder (1975) and Mitra and Bose (1986) used bottom heat at 30C 2
and 35C, respectively, to stimulate root formation. Reuveni et al. (1991) and
Reuveni and Castoriano (1993) treated semi-hardwood cuttings of Turpen-
tine, Gomera, Sabre and 13-1 with bottom heat at 20, 25, 30 and 35C
under mist, and observed most rooting occurred at 25 and 30C. The percent-
age of rooting in cultivars varied from 6095% with between four and ten
roots/cutting. Wounding of the proximal ends of cuttings from 7-month-old
seedlings together with bottom heat increases the number of roots per cut-
ting (Reddy, 1970).
The involvement of phytochrome in root regeneration of mango cuttings
was proposed by Reddy et al. (1975), who observed 92% rooting with etio-
lated and red-light-treated cuttings of 7-month-old seedlings, whereas cut-
tings in a hot bed treated with 5000 ppm IBA and planted under normal light
resulted in 41% rooting.
Reddy and Majumder (1975) achieved 100% rooting of cuttings from
7-month-old mango seedlings treated with 5000 ppm IBA combined with
2000 ppm each of umbelliferone, ruten or quercetin. Phenols and avonoids
acted as rooting cofactors of IBA with juvenile cuttings. Sadhu et al. (1978)
and Reddy and Majumder (1978) observed a synergistic effect of some phe-
nolic compounds (e.g. p-hydroxybenzoic acid, p-coumaric acid and ferulic
acid) used as a preplanting treatment with auxin for induction of rooting
in hardwood mango cuttings. Synergism was more pronounced with IBA
than IAA.
Basu et al. (1970) observed no signicant difference in the levels of endog-
enous rooting substances (i.e. p-hydroxybenzoic acid, p-coumaric acid and
abscisic acid) between juvenile and non-juvenile cuttings. Rajan and Ram
(1983a, b) studied the levels of endogenous rooting hormones of mango cut-
tings with bottom heat under mist and measured increased levels of rooting
hormones and low levels of rooting inhibitors. Fayek et al. (1981) reported
that shoots of 35-year-old Madu contained more rooting inhibitors than
shoots of 1-year-old plants. Using 15,000 ppm IBA and 10,000 ppm NAA,
Rajan and Ram (1983a, b) obtained c.70% rooting with mature cuttings; about
eight brous roots were regenerated as a result of each treatment, and all
Crop Production: Propagation 391
cuttings survived in the nursery. Sadhu (1979) and Sadhu and Bose (1980a, b)
found that 10,000 ppm cycocel pretreatments of 10-year-old Langra resulted
in 41% rooting with 2.2 roots/cutting.
Micropropagation
Somatic embryogenesis from cultured nucellar explants of polyembryonic
cultivars of Mangifera indica was rst reported by Litz et al. (1982, 1984). Sub-
sequently, somatic embryogenesis was induced from the cultured nucellus of
monoembryonic cultivars (Litz, 1984). DeWald et al. (1989a, b) optimized the
protocols for large-scale production of embryogenic cultures in suspension
and for somatic embryo maturation and germination. Shoot tip culture of
Turpentine, Gomera and 13-1 seedlings has been described by Yang and
Ludders (1993); the rate of multiplication was low, and performance of plants
in the eld has not been tested. The current status of somatic embryogenesis
and organogenesis of mango is reviewed by Litz et al., Chapter 18, this volume.
11.4 Comparative Performance of Trees Propagated by Different
Methods
Ram and Sirohi (1989) compared the performance of Dashehari propagated
by cleft grafting, approach grafting, veneer grafting, epicotyl grafting, stool-
ing and air layering. The trees were of uniform size at the time of planting.
Approach-grafted plants did not establish in the eld as well as the other
methods. During the rst 12 years, epicotyl grafts grew more rapidly than
cleft-grafted, veneer-grafted, approach-grafted, stooled and air-layered trees,
respectively (Ram, 1993). The development of the crown also followed the
same pattern. The trees propagated by rooting did not develop an erect main
stem to an adequate height (1.2 m), which hindered cultural operations under
the tree. Trees propagated by approach grafting, veneer grafting and epicotyl
grafting all produced branches close to the ground. Maximum yields were
obtained in trees propagated by epicotyl and cleft grafting followed by
veneer grafting, stooling, air layering and approach grafting, respectively.
The architecture of cleft-grafted plants was much better than trees propa-
gated by other methods; whereas, rooting methods produced twiggy, spread-
ing and dwarf trees (Ram, 1993). After 18 years, cleft-grafted trees were
superior with respect to architecture and yield, followed by epicotyl and
veneer-grafted trees. Rajan and Pandey (1991a, b) conducted experiments
with Dashehari and Langra that were propagated by different vegetative
methods using monoembryonic seedling rootstock. Approach-grafted
Dashehari attained the greatest height followed by epicotyl-grafted, bud-
ded and air-layered trees after 7 years. Budded Langra grew most vigor-
ously, followed by approach-grafted, veneer-grafted, epicotyl-grafted and
air-layered trees. Similar growth of scion girth and crown spread was
recorded with both cultivars. The type of shoot used for approach grafting
S. Ram and R.E. Litz 392
affected tree growth of Dashehari. Grafting height should be minimized
and close contact should be maximized to achieve faster growth when vigorous
rootstocks are used.
11.5 Conclusions
Standard methods are widely utilized for propagating mango scion cultivars
with increasing efcacy. In many regions, including India and Mexico, scion
cultivars are still being propagated on heterogeneous monoembryonic seed-
ling rootstocks (see Crane et al., Chapter 13, this volume), despite the demon-
strated advantages of clonal nucellar rootstocks. The potential of clonally
propagated monoembryonic mango rootstock has not been properly investi-
gated. The genetic heterogeneity of monoembryonic mangoes has been
explored neither for stress tolerance nor for their effects on scion growth and
development. Somatic embryogenesis could play an important role in such
investigations as an alternative propagation method (see Litz et al., Chapter
18, this volume). Other Mangifera species also have interesting attributes, and
should be screened for graft compatibility with mango (see Bompard, Chap-
ter 2, this volume). The species that could be tested as rootstock for mango
might extend mango cultivation to areas where abiotic and biotic stresses
currently limit production and could provide a better source for dwarng
rootstock for high-density orchards. Mango species growing in swamps or
seasonally inundated areas (i.e. Mangifera decandra, Mangifera gedebe, Mangifera
inicarpoides, Mangifera grifthii and Mangifera quadrida) represent a promis-
ing source of rootstock for the development of mango cultivation on poorly
drained soils and inundated lands. In West Kalimantan, Mangifera laurina is
occasionally used as a rootstock for commercial mango cultivars on periodi-
cally inundated riverbeds, and is now being tried at Sabah (Bompard, 1993).
Mangifera zeylanica has been tried as a rootstock for several mango cultivars
in Sri Lanka (Parsons, 1931). Interspecic hybridization of other Mangifera
species with the common mango could also increase the available genetic
variability for rootstock development.
The potential for using species in other genera in the Anacardiaceae as
rootstock has scarcely been explored. Burns and Prayag (1921) investigated
the use of Semecarpus anacardium, Spondias mangifera (Spondias pinnate), Spondias
acuminata and Horigarna grahami as rootstocks for common mango without
any success. Anacardium occidentale (cashew) seedlings have been reported to
be graft compatible with mango scions and mango fruit on cashew rootstock
were reportedly almost double the size of normal fruit with smaller seeds and
free from bre (Garner and Chaudhri, 1976). Mango rootstock development
should receive as much attention as breeding of scion cultivars.
Classical breeding and grafting studies should be greatly expanded to
include the enormous genetic variability within the genus. Emerging bio-
technological approaches also should not be overlooked as alternative prop-
agation methods and as the means to engineer specic horticultural traits in
candidate rootstocks.
Crop Production: Propagation 393
References
Abdel-Galil, H.A. (1992) Evaluation of certain techniques for germination of mango
seed stones. Assiut Journal of Agricultural Sciences 23, 134151.
Ahmed, S. (1960) Propagation of mangoes. Punjab Fruit Journal 23, 4953.
Ahmed, S. (1964) Propagation of mango by veneer grafting. West Pakistan Journal of
Agriculture Research 2, 3244.
Ahmed, S. (1974) Selection and evaluation of local mango cultivars and strains in North-
east Brazil. In: 6th Report of the UNDP/FAO/CEPED Project Brazil 71/555, Rome,
Italy, pp. 13.
Ahmed, S. and Chatha, G.A. (1960) Some morphological and anatomical studies of
scion buds and bud union zone in mangoes. Punjab Fruit Journal 23, 105111.
Ahmed, S. and Rashid, A. (1961) Importance of seed in the fruit industry. Agriculture
Pakistan 12, 252257.
Ali, N. and Nazir, M. (1970) Effect of auxins on carbohydrate and nitrogen contents of
mango cuttings. Pakistan Journal of Science 22, 1221.
Amin, R.S. (1974) A study on the establishment of mango orchard with wedge graft on
in-situ grown mango seedling in dry region of the Gujarat State. Haryana Horticul-
tural Journal 3, 160167.
Amin, R.S. (1978) In-situ soft wood grafting in mango. Indian Horticulture 23, 710.
Anonymous (1950) Annual Report of the Florida Agricultural Experiment Station for the
Year ending June 30. University of Florida, Gainesville, Florida, p. 263.
Anonymous (1975) The Philippines Recommends for Mango 1975. Philippine Council
for Agricultural Research, Manila, the Philippines.
Anonymous (1994) Production Technology Mango. Information Bulletin No. 12. Indian
Institute of Horticulture Research, Hessarghatta, Bangalore, India, p. 47.
Asadullah, M. and Khan, M.D. (1960) Studies of various factors affecting success in
grafting by approach (inarching) in mangoes. Punjab Fruit Journal 23, 5970.
Bajpai, P.N., Singh, A.R., Yati, V. and Chaturvedi, O.P. (1989) Effect of cultivars and age
of rootstocks on the performance of veneer grafting in mango. Acta Horticulturae
231, 259262.
Bajwa, B.S. and Ram, A. (1946) Budding in mango seedlings in-situ. Punjab Fruit Jour-
nal 10, 1013.
Bakhshi, I.C. (1963) Propagation studies with monoembryonic and polyembryonic vari-
eties of mango. Punjab Horticultural Journal 3, 185193.
Basu, R.N., Roychoudhury, N., Bose, T.K. and Sen, P.K. (1966) Rooting of mango cutting.
Indian Agriculturist 10, 147151.
Basu, R.N., Roychoudhary, N., Gosh, B. and Sen, P.K. (1970) Endogenous rooting fac-
tors in the mango (Mangifera indica L.) seedling and mature shoots. Horticulture
Science 2, 9198.
Basu, R.N., Roychoudhury, N., Lahiri, B. and Sen, P.K. (1972) Biochemical changes
during root formation in Mangifera indica L. air layers. Acta Horticulturae 24,
6471.
Benincasi, R. (1970) Tentative di applicazione di tecniche di propagazione agamica in
pi ante da frutto tropicali presso la stazione sperimentate di Mvuazi (Basso Congo)
(Kinshasha). Revista di Agricoltura Subtropicale e Tropicale 64, 182195.
Ben-yaacov, A. (1976) Recovery of frost-destroyed mango trees by regrafting at the root
shoot transition zone. Proceedings of the American Society for Horticulture Science
Tropical Region 24, 151156.
Bhambota, J.R., Rajput, M.S. and Sandhu, K.S. (1971) Veneer grafting a successful method
of propagation. Punjab Horticultural Journal 11, 4043.
S. Ram and R.E. Litz 394
Bhan, K.C., Samaddar, R.N. and Yadav, P.C. (1969) Chip budding and stone-grafting of
mangoes in India. Tropical Agriculture Trinidad 46, 247253.
Bid, N.N. and Mukherjee, S.K. (1969) Varietal response to etiolation and growth-regulator
treatment in air-layering of mango (Mangifera indica L.). Indian Journal of Agricul-
tural Science 39, 10131019.
Bid, N.N. and Mukherjee, S.K. (1972) Studies into the effects of forced shoot etiolation
and different media on the rootage of Mangifera indica L. cuttings. Acta Horticultu-
rae 24, 7781.
Bid, N.N., Mukherjee, S.K. and Majumder, P.K. (1972) Studies on different methods of
propagation of Mangifera indica with respect to success, survival, growth and their
clonal multiplication by stooling. Acta Horticulturae 24, 8588.
Bompard, J.M. (1993) The genus Mangifera rediscovered. The potential contribution of
wild species to mango cultivation. Acta Horticulturae 341, 6977.
Burns, W. and Prayag, S.H. (1921) The Book of the Mango. Department of Agriculture
Bulletin, Bombay, India.
Carmichael, W.W. (1957/58) Rejuvenation of old mango groves by hedging or top-
working to new varieties. Proceedings of the Florida State Horticultural Society
70, 290291.
Chacko, E.K. and Singh, R.N. (1971) Studies on the longevity of papaya, phalsa, guava and
mango seeds. Proceedings of the International Seed Testing Association 36, 147158.
Chauran, O.R., Manica, I., Pinheiro, R.V.R., Conde, A.R. and Chaves, J.R.P. (1979) The
effect of storage time, seed treatment and fungicide on seed germination and growth
of mango seedlings. Revista Ceres 26, 112.
Chhonkar, V.S. and Singh, R.K. (1972) Propagation of Mangifera indica L. by air layering.
Acta Horticulturae 24, 8992.
Choudhury, K.R. (1959) A short note on substitutes for moss as plastering material in air
layering of mango. Science and Culture 25, 152153.
Cooper, W.C. (1936) Transport of root-forming hormones in woody cuttings. Plant Phys-
iology 11, 779.
Cooper, W.C. and Furr, J.R. (1945) The cinchona veneer graft method of propagating
subtropical fruit trees. Proceedings of the Florida State Horticultural Society 58,
176180.
Cooper, W.C. and Stoutemyer, V.T. (1945) Suggestion for the use of growth substances in
the propagation of tropical plants. Tropical Agriculture Trinidad 22, 21.
Degani, C., Cohen, M., Reuveni, O., El-Batsri, R. and Gazit, S. (1993) Frequency and
characteristics of zygotic seedlings from polyembryonic mango cultivars, deter-
mined using isozymes as genetic markers. Acta Horticulturae 341, 7881.
DeWald, S.G., Litz, R.E. and Moore, G.A. (1989a) Optimizing somatic embyro pro-
duction in mango. Journal of the American Society for Horticultural Science 114,
712716.
DeWald, S.G., Litz, R.E. and Moore, G.A. (1989b) Maturation and germination of man-
go somatic embryos. Journal of American Society for Horticultural Science 114,
837841.
Dhunaga, D.B., Arvindakshan, M. and Gopikumar, K. (1989) Standardization of stone
grafting in mango. Acta Horticulturae 231, 170174.
Dijkman, M.J. (1950) Rooting of Haden mango (Mangifera indica L.) leaf bud cuttings.
Science 111, 663665.
Eiadthong, W., Yonemori, K., Sugiura, A., Utsunomiya, N. and Subhadrabandhu, S.
(1999) Identication of mango cultivars of Thailand and evaluation of their genetic
variation using the amplied fragments by simple sequence repeat-(SSR-) anchored
primers. Scientia Horticulturae 82, 5766.
Crop Production: Propagation 395
Faruque, A.H.M. and Fakir, M.M.A.S. (1973) Propagation of mango by different methods
of grafting. Bangladesh Horticulture 1, 2528.
Fayek, M.A., Salim, H.H., Ibrahi, F.A. and Menibary, M.A. (1981) Tentative identication
of the major inhibiting compound in mango shoots. Annals of Agricultural Science
16, 209216.
Furon, V. (1966) La multiplication du manguier au Senegal. Fruits 21, 189193.
Furon, V. and Plaud, G. (1972) Topworking mango trees in Senegal. Fruits 27, 293296.
Galn-Saco, V. (1993) The situation of mango culture in the world. Acta Horticulturae
341, 3141.
Galn-Saco, V. (1999) El Cultivo del Mango. Ediciones Mundi-Prensa, Madrid, Spain.
Gandhi, S.R. (1942) Recent advances in horticultural practices. Poona Agriculture Col-
lege Magazine 34, 86.
Gandhoke, M.M.S. (1993) Flush grafting in mango way to sure success. (Abstract).
Golden Jubilee Symposium Horticultural Research Changing Scenario. Horticul-
tural Society of India, Bangalore, India, p. 445.
Gangolly, S.R., Singh, R., Katyal, S.L. and Singh, D. (1957) The Mango. Indian Council
of Agriculture Research, New Delhi, India.
Gardner, D.E. and Piper, R.B. (1943) Base of propagation of some sub-tropical fruits by
cuttings from young seedlings. Proceedings of the Florida State Horticultural Soci-
ety 56, 124126.
Garg, M.L. (1954) Mango grafting made easy and cheap. Indian Journal of Horticulture
11, 145146.
Garner, R.J. and Chaudhri, S.A. (1976) Mangifera indica mango. In: Garner, R.J. and
Chaudhri, S.A. (eds) The Propagation of Tropical Fruit Trees. Food and Agriculture
Organization of the United Nations and the Commonwealth Agricultural Bureau,
Wallingford, UK, pp. 403474.
Gaur, N.V.S. (1984) Comparative evaluation of selected methods of mango propagation.
Punjab Horticultural Journal 24, 16.
Gazit, S. and Kadman, A. (1980) 13-1 mango rootstock selection. HortScience 15,
669.
George, P.V. and Nair, T.N.K. (1969) On the performance of mono- and polyembryonic
rootstocks in mango grafts. Agriculture Research Journal Kerala 7, 79.
Giri, A. (1966) Transplanting success with varying stem girths of mango seedlings. Agri-
culture Pakistan 17, 195200.
Giri, A. and Chaudhery, M.Y. (1966) Relation of mango stone weight to its germination
and seedling vigor. Pakistan Journal of Science 18, 148150.
Gonzalez, A., Coulson, M. and Brettell, R. (2002) Development of DNA markers (ISSRs)
in mango. Acta Horticulturae 575, 139143.
Gowda, M.H.M. (1962) Some methods of propagation of mango. In: Fruit Nursery Prac-
tices in India. Directorate of Extension, Ministry of Food and Agriculture, New
Delhi, India, pp. 5464.
Gowder, R.B. and Irulappan, I. (1970) Performance of Neelum variety of mango (Mangifera
indica L.) on polyembryonic rootstocks. Madras Agriculture Journal 57, 29.
Grant, J.W. and William, A.N.P. (1949) Burma Fruits and their Cultivation. Burma Bul-
letin 30. Department of Agriculture, Rangoon, Burma.
Grove, W.R. (1947) Wrapping air layers with rubber. Proceedings of the Florida State
Horticultural Society 60, 184187.
Gunjate, R.T. (1989) Standardization of stone grafting for the Konkan region. Acta Hor-
ticulturae 231, 164167.
Gunjate, R.T. and Limaye, Y.P. (1978) Veneer grafting of mango seedlings raised in earth-
en pots. Journal of Maharashtra Agricultural University 3, 6162.
S. Ram and R.E. Litz 396
Gunjate, R.T., Uradya, V.S. and Limaye, V.P. (1976) Effect of season and defoliation of
scion shoots on success in veneer grafting in Alphonso mango. Journal of Maha-
rashtra Agricultural University 3, 5152.
Gupta, O.P., Jawanda, J.S. and Sharma, K.C. (I988) Stone grafting in mango under Jammu
conditions. Progressive Horticulture 20, 1114.
Hayes, W.B. (1957) Fruit Growing in India, 3rd edn. Kitabistan, Allahabad, India.
Higgins, J.E. (1910) Shield budding the mango. Bulletin of the Hawaii Agriculture
Experiment Station 20, 1.
Hussain, I. (1953) Some studies on the effect of hormones on the rooting of cuttings of
horticultural plants. PhD thesis, University of Punjab, Lahore, Pakistan.
Iyer, C.P.A., Subramanyam, M.D. and Dinesh, M.R. (1990) A simple procedure for dou-
ble working in mango. Journal of Maharashtra Agricultural University 15, 244
245.
Jagirdar, S.A.P. and Ali, S.K. (1968) Effect on scion variety and methods of raising stock
on bud-take in mango. West Pakistan Journal of Agricultural Research 6, 8387.
Jagirdar, S.A.P. and Bhatti, M.S. (1968) Effect of type of scion wood and age of stock on
the success of veneer grafting in mango (Mangifera indica). West Pakistan Journal
of Agricultural Research 6, 8897.
Jagirdar, S.A.P., Nizamani, M.H. and Shaikh, M.A. (1968) Varietal response of veneer
grafting in mango. West Pakistan Journal of Agricultural Research 6, 8689.
Jauhari, O.S. and Nigam, Y.N. (1958) Effect of hormones on rooting in cuttings of mango.
Kanpur Agriculture College Journal 17, 13.
Jauhari, O.S. and Nigam, V.N. (1962) Air layering of mango. In: Fruit Nursery Practices
in India. Directorate of Extension, Ministry of Food and Agriculture, New Delhi,
India, pp. 7781.
Jauhari, O.S. and Singh, R.D. (1970) Effect of bud activation, lopping and wrapping
materials on budding in mango (Mangifera indica) variety Langra. Punjab Horticul-
tural Journal 10, 198202.
Jauhari, O.S., Teotia, S.S. and Upadhyay, S.K. (1972) Rootstock studies in Mangifera
indica L. Acta Horticulturae 24, 107109.
Jinturkar, S.P. and Narwadkar, P.R. (1989) Effect of environmental conditions on the
success of epicotyl grafting in mango. Acta Horticulturae 231, 252255.
Juliano, J.B. (1934) Origin of embryos in the strawberry mango. Philippine Journal of
Science 54, 553556.
Juliano, J.B. (1937) Embryos of carabao mango (Mangifera indica L.). Philippines Agri-
culture 25, 749760.
Kadman, A., Gazit, S. and Ziv, G. (1978) Experiments with the selection of mango root-
stocks for the Southern Arava. Israel Journal of Botany 27, 3435.
Kains, M.G. and McQuestan, L.M. (1955) Propagation of Plants. Orange Judd, New York.
Kanwar, J.S. and Bhajwa, M.S. (1974) Propagation of mango by side grafting. Indian
Journal of Agricultural Science 44, 270272.
Kashkush, K., Jinggui, F., Tomer, E., Hillel, J. and Lavi, U. (2001) Cultivar identication
and genetic map of mango (Mangifera indica). Euphytica 122, 129136.
Kashyap, R., Jyotishi, R.P. and Sharma, A.B. (1972) Techniques of side-grafting in mango.
Acta Horticulturae 24, 99100.
Khalifa, A.S., EI-Azzouni, M.M. and Wah, Y.A. (1964) Physiological studies on mango
cuttings. Indian Journal of Horticulture 21, 179185.
Khan, A.A. (1960) Mango budding in-situ. A new technique likely to revolutionise the
mango industry. Punjab Fruit Journal 23, 113116.
Kulkarni, V.I., Ratnam, K.K. and Ramkrishna, G. (1989) Propagation studies in mango.
Acta Horticulturae 231, 186191.
Crop Production: Propagation 397
Kulwal, L.V. and Tayde, G.S. (1989) Studies on propagation of mango varieties by soft
wood grafting under Akola condition. Acta Horticulturae 231, 256258.
Kusumo, S., Soehendro, R. and Rijanto, B. (1971) Mango rootstock experiment at
Poh-djentrek. Bulletin Hortikultura No.3. Tjaban Lembaga Penelitian Horticulture,
Indonesia from Ringkasan. Publikasi dan Laporan Penelitian Pertanian 2, 333.
Lamboume, J. (1935) The etiolation shoot method of fruit propagation. Malaysian Agri-
culture Journal 23, 514527.
Ledin, R.B. and Ruehle, G.D. (1954) Mango selection, propagation and culture. Annual
Report of Florida Agricultural Experimental Station. University of Florida, Gainesville,
Florida, pp. 291292.
Litz, R.E. (1984) In vitro somatic embryogenesis from nucellar callus of monoembryonic
mango. HortScience 19, 715717.
Litz, R.E., Knight, R.J. and Gazit, S. (1982) Somatic embryos from cultured ovules of
Mangifera indica L. Plant Cell Report 1, 264266.
Litz, R.E., Knight, R.J, Jr and Gazit, S. (1984) In vitro somatic embryogenesis from
Mangifera indica L. callus. Scientia Horticulturae 22, 233240.
Luthra, J.C. and Sharma, M.M.L. (1946) Some studies on the conductability and histol-
ogy of grafted mango shoots. Horticultural Botany 35, 4.
Lynch, S.J. (1941) Nursery Propagation and Topworking of Mangoes. Bulletin of Florida
Agriculture Experimental Station 560. University of Florida, Gainesville, Florida, 4 pp.
Lynch, S.J. and Mustard, M.J. (1950) Mangoes in Florida. Bulletin of the Florida Depart-
ment of Agriculture 135. University of Florida, Gainesville, Florida.
Lynch, S.J. and Nelson, R.O. (1949) Mango budding. Proceedings of the Florida State
Horticultural Society 62, 207209.
Lynch, S.J. and Nelson, R.O. (1956) Current methods of vegetative propagation of avo-
cado, mango, lychee and guava in Florida. Ceiba 4, 315337.
Madalgeri, M.B., Hulamani, N.C. and Patil, V.R. (1989) Response of mango varieties and
hybrids to epicotyl grafting. Progressive Horticulture 21, 173175.
Maheshwari, P., Sachar, R.C. and Chopra, R.N. (1955) Embryological studies in mango
(Mangifera indica L.). In: Proceedings of Indian Science Congress, Baroda, Gujarat,
India, p. 233.
Majhail, H.S. and Singh, K.K. (1962a) Inarching in mango. I. The effect of Alkathene wrapper,
time of inarching and size of stock seedling. Punjab Horticultural Journal 2, 3343.
Majhail, H.S. and Singh, K.K. (1962b) Inarching in mango II. The optimum period of
grafting and age of stock seedling. Punjab Horticultural Journal 2, 109113.
Majumder, P.K and Mukherjee, S.K. (1961) A note on the propagation of root-stocks in
mango. Indian Journal of Horticulture 18, 167168.
Majumder, P.K. and Rathore, D.S. (1970) Epicotyl grafting in mango (Mangifera indica
L.). Current Science 39, 142.
Maurice, J. (1969) Section-grafting and rooting of tropical fruit trees. World Crops 21,
265267.
Miner, J.T. (1962) Top working mango trees in Florida. Proceedings of the Tropical
Region of the American Society for Horticultural Science 6, 3133.
Mitchell, E.F. (1971) Mango production and marketing practices Florida. Proceedings
of the Florida State Horticultural Society 84, 307310.
Mitra, S.K. and Bose, T.K. (1986) Mango. In: Bose, T.K. (ed.) Propagation of Tropical and
Subtropical Horticultural Crops. Naya Prokash, Calcutta, India, pp. 206207.
Moreuil, C. (1963) Le manguier au Congo. Fruits 18, 295301.
Mukherjee, S.K. and Bid, N.N. (1965) Propagation of mango (Mangifera indica L.) II.
Effect of etiolation and growth regulator treatment on the success of air-layering.
Indian Journal of Agricultural Science 35, 305314.
S. Ram and R.E. Litz 398
Mukherjee, S.K. and Majumder, P.K. (1961) Veneer grafting in mango has its own
advantages. Indian Horticulture 6, 3.
Mukherjee, S.K. and Majumder, P.K. (1962) Veneer grafting in mango. In: Fruit Nursery
Practices in India. Directorate of Extension, Ministry of Food and Agriculture, New
Delhi, India, pp. 6566.
Mukherjee, S.K. and Majumder, P.K. (1963) Standardization of root-stocks of mango. I.
Studies on the propagation of clonal root-stocks by stooling and layering. Indian
Journal of Horticulture 20, 204209.
Mukherjee, S.K. and Majumder, P.K. (1964) Effect of different factors on the success of
veneer grafting. Indian Journal of Horticulture 21, 4650.
Mukherjee, S.K., Majumder, P.K., Bid, N.N. and Goswami, A.N. (1965) Clonal propaga-
tion of mango (Mangifera indica L.) through cuttings. Current Science 34, 434435.
Mukherjee, S.K., Majumder, P.K. and Goswami, A.M. (1966) Effect of position of the
shoots on the rooting of mango cuttings. Science and Culture 32, 377378.
Mukherjee, S.K., Majumder, P.K., Bid, N.N. and Goswami, A.M. (1967) Standardization
of rootstocks of mango (Mangifera indica L.) II. Studies on the effects of source,
invigoration and etiolation on the rooting of mango cuttings. Journal of Horticul-
tural Science 42, 8387.
Mulat, B. (1959) Mango grafting. Fruits 14, 219223.
Naik, K.C. (1941) Studies on the propagation of the mango. Mangifera indica L. Indian
Journal of Agriculture Science 11, 736768.
Naik, K.C. (1949) South Indian Fruits and their Culture. P. Varadachary and Company,
Madras, India.
Narwadkar, P.R. and Anserwadekar, K.W. (1989) Effect of growth regulator on the suc-
cess of epicotyl grafting in mango. Acta Horticulturae 231, 175178.
Neto, M.T. de C., Fonseca, N., Filho, H.P.S. and Juior, A.T.C. (2002) Propagacao e padrao
da muda. In: Genu, P.J. de C. and Pinto, A.C. de Q. (eds) A Cultura da Mangueira.
Embrapa Informao Tecnolgica, Braslia, Brazil, pp. 117136.
Nez-Elisea, R., Caldeira, M.L., Ferreira, W. and Davenport, T.L. (1992) Adventitious
rooting of Tommy Atkins mango air layers induced with naphthaleneacetic acid.
HortScience 27, 926.
Oppenheimer, C. (1960) The relationship between tree size and yield in mango and
avocado. Horticulture Advance 4, 615.
Panickar, P. and Desai, A.G. (1989) Effect of age of scion mother tree, different ushes of
rootstock and in-situ grafting on success and growth of soft wood grafts of Alphonso
mango. Progressive Horticulture 21, 141144.
Parisot, E. (1988) Study of the growth rhythm in young mango plants. 1. Description,
germination and storage of polyembryonic mango seeds. Fruits 43, 97105.
Parsons, T.H. (1931) The mango in Ceylon. Tropical Horticulturist 76, 119211.
Patil, A.A., Vadigeni, B.G. and Nalawadi, U.G. (1991) Response of mango varieties to
stone grafting. Current Research of University of Agricultural Sciences (Bangalore)
20, 135136.
Patwardhan, G.B. and Deshmukh, G.B. (1931) A Handbook of Horticultural Practices.
Patwardhan, Poona, India.
Paul, W.R.C. and Guneratnam, S.C. (1937a) The propagation of mango in Jaffna. I. Trop-
ical Agriculturist 88, 8691.
Paul, W.R.C. and Guneratnam, S.C. (1937b) The propagation of mango in Jaffna. II.
Tropical Agriculturist 88, 331337.
Paul, W.R.C. and Guneratnam, S.C. (1938) Mango stocks. Tropical Agriculturist 90, 3435.
Pinheiro, R.V.R., Anderson, O. and Fortes, J.M. (1970) Comparacao de modalidades de enx-
ertia na propagacao da mangueira (Mangifera indica L.). Review Ceres 17, 264273.
Crop Production: Propagation 399
Pinto, A.C.Q., Andrade, S.R.M., Amaro, A.A. and Gomes, U. (2004) Mango industry in
Brazil. Acta Horticulturae 645, 3750.
Pope, W.T. (1929) Mango Culture in Hawaii. Hawaii Agricultural Experiment Station
Bulletin 58. Hawaii Agricultural Experiment Station, Hawaii, 58 pp.
Pope, W.T. and Storey, W.B. (1933) Grafting Tropical Fruit Trees in Hawaii. Hawaii Agricul-
tural Experimental Station Circular 6. Hawaii Agricultural Experiment Station, Hawaii.
Popenoe, W. (1959) EI mango. Suelo Tico 11, 2327.
Prasad, A. and Pathak, R.A. (1970) Response of plant growth regulators to mango
(Mangifera indica L.). Tropical Agriculturist 126, 95106.
Prasad, A. and Singh, A.R. (1972) Mango propagation studies at Government Horticul-
tural Research Institute, Saharanpur, India. Acta Horticulturae 24, 9398.
Prasad, A., Singh, R.D. and Sirohi, R.S. (1973) Comparative study of veneer grafting and
patch budding in Mangifera indica L. cv Dashehari. Punjab Horticultural Journal
13, 3034.
Radhamony, P.S., Gopikumar, K. and Valsalakumari, P.K. (1989) Varietal responses of
scion to stone grafting in mango for commercial propagation. South Indian Horti-
culture 37, 298299.
Rajan, S. and Pandey, D. (1991a) Studies on propagational techniques in mango. Central
Institute of Horticulture for Northern Plains Annual Report 198990. Central Insti-
tute of Horticulture for Northern Plains, Lucknow, India, pp. 1920.
Rajan, S. and Pandey, D. (1991b) Standardization of root stock on mango. Central Insti-
tute of Horticulture for Northern Plains Annual Report 198990. Central Institute of
Horticulture for Northern Plains, Lucknow, India, pp. 1729.
Rajan, S. and Ram, S. (1983a) Some factors affecting root regeneration in mango cutting
in mist and hot bed. Progressive Horticulture 15, 1116.
Rajan, S. and Ram, S. (1983b) Some new concepts in rootage of mango. Progressive
Horticulture 15, 155156.
Rajan, S. and Ram, S. (1989) Studies on the root regeneration in mango air layers. Acta
Horticulturae 231, 192197.
Rajan, S. and Sinha, G.C. (1987) Use of aluminium foil for increasing veneer grafting
success under adverse conditions. Progressive Horticulture 19, 141142.
Rajput, C.B.S. and Haribabu, R.S. (1971) Recent techniques of mango propagation.
World Crops 23, 146148.
Ram, S. (1993) Factors affecting mango tree architecture. Acta Horticulturae 341, 177191.
Ram, S. and Bist, L.D. (1982) Studies on veneer grafting of mango in Tarai. Punjab Hor-
ticultural Journal 22, 6471.
Ram, S. and Sirohi, S.C. (1982a) Effect of growth regulators on air layering in mango.
(Abstract). National Seminar on Plant Propagation. Department of Horticulture Bidhan
Chandra Krishi Vishwavidyalaya and Society for Advancement of Horticulture,
Kalyani, West Bengal, pp. 7273.
Ram, S. and Sirohi, S.C. (1982b) Performance of mango cultivar Dashehari trees propa-
gated by different vegetative methods. (Abstract). National Seminar on Plant Propa-
gation. Department of Horticulture Bidhan Chandra Krishi Vishwavidyalaya and
Society for Advancement of Horticulture, Kalyani, West Bengal, pp. 9394.
Ram, S. and Sirohi, S.C. (1989) Performance of Dashehari mango trees propagated by
different vegetative methods. Acta Horticulturae 231, 210215.
Ram, S., Sirohi, S.C. and Bist, L.D. (1981) Interaction of diphenolics with auxin in stool-
ing of mango. Punjab Horticultural Joumal 21, 4652.
Ramirez Diaz, J.M. (1973) The effect of cutting back root stock on the take and initial
growth of the Kent mango scion and its relationship with apical dominance. Agri-
cultura Technica en Mexico 3, 245252.
S. Ram and R.E. Litz 400
Rangacharlu, V.S. and Rao, M.V. (1956) Air grafting of mango. (A new technique in the
grafting of mango without the aid of soil or water). Andhra Agricultural Journal 1,
105109.
Rao, S.N., Swamy, G.S., Seeramulu, P. and Reddy, R.N. (1963) Growth regulators in
propagation of mango by air layering. Punjab Horticultural Journal 3, 175179.
Rao, V.N.M. (1967) Propagation practices. In: The Mango Handbook. Indian Council for
Agricultural Research (ICAR), New Delhi, India, pp. 3269.
Rao, V.N.M. and Rao, I.K.S. (1956) Mango grafting with the aid of plastic lm wrapper.
South Indian Horticulture 4, 5455.
Reddy, C.V. and Melanta, K.R. (1988) Effect of age of root stocks on the success of soft wood
grafting of mango in container and in situ. South Indian Horticulture 36, 143145.
Reddy, Y.N. (1970) Physiological and biochemical studies on the regeneration of mango
(Mangifera indica L.) and guava (Psidium guajava L.) cuttings. PhD thesis, Indian
Agricultural Research Institute, New Delhi, India.
Reddy, Y.N. and Majumder, P.K. (1975) Bottom heat a new technique for rooting of
hard wood cuttings of tropical fruits. Current Science 44, 444445.
Reddy, Y.N. and Majumder, P.K. (1978) Synergism of phenols and avonoids with IBA in
regeneration of mango and guava cuttings. Vatika 1, 3744.
Reddy, Y.N., Majumder, P.K., Pandey, R.M. and Singh, R.N. (1975) Phytochrome medi-
ated root generation in mung (Phaseolus aureus Roxb) and mango (Mangifera indica
L.) cuttings. Current Science 44, 509510.
Reuveni, O. and Castoriano, M. (1993) Effect of bottom heat temperature on rooting of
mango cuttings of different cultivars. Acta Horticulturae 341, 288291.
Reuveni, O., Kuepper, W. and Wiebel, J. (1991) Rooting of mango cuttings under inter-
mittent mist. Acta Horticulturae 291, 174181.
Ruehle, G.D. and Ledin, R.B. (1955) Mango Growing in Florida. Bulletin of Florida Agri-
culture Experimental Station 574. University of Florida, Gainesville, Florida, 90 pp.
Sachar, R.C. and Chopra, R.N. (1957) A study of the endosperm and embryo in Mangifera.
Indian Journal of Agricultural Science 27, 219228.
Sadhu, M.K. (1979) Effect of pretreatment of stock plants of mango with cycocel, ethrel
and morphactin on the rooting of cutting and air layers. Scientia Horticulturae 10,
363368.
Sadhu, M.K. and Bose, S. (1980a) Effect of ethylene on rooting of cuttings and air layers
of mango, guava and water-apple. Indian Journal of Horticulture 37, 335337.
Sadhu, M.K. and Bose, S. (1980b) Note on promotion of rooting in difcult-to-root fruit-
tree cuttings with 2-chloroethyl phosphonic acid and acetylene in the presence of
auxins. Indian Journal of Agriculture Science 50, 448450.
Sadhu, M.K., Bose, S. and Saha, L. (1978) Auxin synergist in rooting of mango cuttings.
Scientia Horticulturae 9, 381387.
Said, H. and Shoushan, A.A. (1945) Root formation of cuttings of plants which normally
do not root. Nature 155, 791.
Schnell, R.J. and Knight, R.J. (1993) Genetic relationships among Mangifera spp. based
upon RAPD markers. Acta Horticulturae 341, 8692.
Sen, P.K. (1939) Annual Report of the Fruit Research Station, Sabour Bihar. Superinten-
dent Government Printing, Patna, Bihar, India.
Sen, P.K. and Bose, T.K. (1964) Physiological studies on regeneration of roots. VI: Effects
of growth substances on root formation in justicia, mulberry and mango stem cut-
tings and concomitant changes in carbohydrate and nitrogenous substances in the
rooting tissues. Indian Agriculturist 8, 102119.
Sen, P.K. and Bose, T.K. (1966) Physiological studies on regeneration of roots. VII. Rela-
tion of chemical composition to the regeneration of roots in airlayers of mango
Crop Production: Propagation 401
(Mangifera indica L.) and Litchi (Litchi chinensis Sam.) VIII. Role of leaves in rooting
of stem cuttings. Indian Agriculturist 10, 7586.
Sen, P.K., Bose, T.K. and Sasibhushan, T. (1961) Propagation of mango, Mangifera indica
L., by air-layering with growth substances. Indian Agriculturist 5, 167172.
Sen, P.K., Basu, R.N., Bose, T.K. and Roychoudhary, N. (1968) Rooting of mango cut-
tings under mist. Current Science 37, 144146.
Sen, P.K., Basu, R.N., Bose, T.K. and Roychoudhary, N. (1969) Mist propagation of fruit
tree cuttings. Plant Science 1, 216219.
Serpa, D. (1964) Propagacion del mango. Faculty of Agronomy, Universidad Central de
Venezuela 2, 24.
Shant, P.S. and Saproo, B.L. (1974) Effects of various stages of fruit maturity and different
levels of pulp extraction on the germination of mango stones. Bangladesh Horticul-
ture 2, 4950.
Simao, S. (1960) Seeds of green mangoes for rootstock production. Review Agriculture
Piracicaba 35, 183188.
Singh, B. and Ram, A. (1946) Budding of mango seedlings in situ, a brief rsum of work
done during the years 194345. Punjab Fruit Journal 10, 10.
Singh, J.R. and Srivastava, R.P. (1961) Budding as a commercial method of mango prop-
agation. Gardening 3, 1921.
Singh, J.R. and Srivastava, R.P. (1962) Studies in budding of mango. I. Indian Journal of
Horticulture 19, 130134.
Singh, K.K. and Jawanda, J.S. (1962) Some promising seedling mango trees of Punjab.
Punjab Horticultural Journal 2, 133138.
Singh, L. and Khan, A.A. (1942) Mango budding in situ. Punjab Fruit Journal 6, 195206.
Singh, L. and Khan, A.A. (1943) How to prolong the life of mango bud-wood. Punjab
Fruit Journal 7, 12641265.
Singh, L. and Khan, A.A. (1946) Mango budding in situ, a new technique likely to revo-
lutionise the mango industry. Punjab Fruit Journal 10, 14.
Singh, L.B. (1954) Propagation of mango by air layering for root-stocks. Proceedings of
the American Society of Horticultural Science 63, 128130.
Singh, L.B. (1960) The Mango. Leonard Hill (Books) Limited, London.
Singh, M.P., Gill, S.S. and Khajuria, R.N. (1989) Standardization of propagation tech-
niques in mango. Acta Horticulturae 231, 179181.
Singh, N.P. and Srivastava, R.P. (1979a) Studies on the different aspects involved in
veneer grafting in mango. Progressive Horticulture 11, 6774.
Singh, N.P. and Srivastava, R.P. (1979b) Practical hints on veneer grafting in mango.
Indian Horitculture 23, 68.
Singh, N.P., Srivastava, R.P., Singh, R. and Rajput, M.S. (1979) Seasonal effect on success in
different methods of mango propagation. Indian Journal of Horticulture 36, 135139.
Singh, R. and Singh, C.P. (1998) Planting and care, propagation, rootstock and irrigation
in mango crops. In: Srivastava, R.P. (ed.) Mango Cultivation. International Book
Distributing Co., Lucknow, India, pp. 101127.
Singh, R.N. (1990) Mango. Indian Council of Agricultural Research, New Delhi, India.
Singh, S. (1946) Propagation of the mango in the Punjab. Punjab Fruit Journal 10, 8.
Singh, S.N. and Teaotia, S.S. (1951) Effect of some hormones on rootage of mango. Sci-
ence and Culture 17, 207210.
Singh, U.R. and Singh, A.P. (1976) Rootstock studies in mango (Mangifera indica). Pro-
gressive Horticulture 8, 1319.
Soule, M.J., Jr (1954) Inuence of type of graftage, age of plant, and certain environmen-
tal factors on the loss of water by young mango buddings. PhD thesis, University of
Florida, Gainesville, Florida.
S. Ram and R.E. Litz 402
Soule, M.J., Jr (1971) Anatomy of the bud union in mango (Mangifera indica L.). Journal
of the American Society of Horticultural Science 96, 380383.
Srivastava, R.P. (1963a) Studies in the response of plant growth regulators in air-layered
shoots of mango (Mangifera indica L.). Journal of Science Research, Banaras Hindu
University 11, 13.
Srivastava, R.P. (1963b) Propagation of mango and guava by transported buds. Science
and Culture 29, 145146.
Srivastava, R.P. (1989) Propagation of mango by newer techniques. Acta Horticulturae
231, 266267.
Stephens, S.E. (1948) The mango in Queensland. Queensland Agricultural Journal 68,
7181; 146153; 208215.
Stephens, S.E. (1960) Mango growing in Queensland. Queensland Agriculture Journal
86, 761766.
Stoler, S. (1976) Mango rootstock trial in the Jordan valley. Special Publication. Agricul-
tural Research Organisation Israel 65, 9598.
Sturrock, T.T. (1968) Nucellar embryos of the mango. Proceedings of the Florida State
Horticultural Society 80, 350354.
Subra, P. (1954) Greffage du manguier au Cameroun. Fruits 9, 498501.
Sundara Rao, Y.R. (1956) Preservation of bud wood. South Indian Horticulture 4, 312.
Swamy, G.S., Rao, E.Y.R. and Raju, D.S. (1972) Polyembryonic rootstocks for mango.
Acta Horticulturae 24, 110113.
Talukdar, M.R. and Ahmed, S. (1965) Success of in arching done on three varieties of
mango on young stocks at Lyallpur. Pakistan Journal of Science 17, 7274.
Tanaka, T. (1939) A new method in mango propagation. Proceedings of the Agriculture
Society of Trinidad 39, 232287.
Teaotia, S.S. (1963) Budding in mango (Mangifera indica L.). Kanpur Agriculture
College Journal 23, 4950.
Teaotia, S.S. and Maurya, V.N.M. (1970) Studies on the propagation of mango by bud-
ding. Progressive Horticulture 2, 3544.
Thakurta, A.G. and Dutt, B.K. (1941) Vegetative propagation of mango from gootee
(marcottage) and cuttings by treatment of high concentration of auxin. Current Sci-
ence 10, 297.
Thrower, L.B. (1954) Commercial horticulture in the Sudan. 2. Mango cultivation.
Bulletin Ministry of Agriculture Sudan 11, 66.
Torres, J.P. (1949) Splice grafting of mango. Philippines Journal of Agriculture 14, 247
255.
Torres, J.P. (1960) Splice grafting of mango. Punjab Fruit Journal 23, 7376.
Traub, H.P. and Auchter, B.C. (1934) Propagation experiments with avocado, mango
and papaya. Proceedings of the American Society of Horticulture Science 30,
382385.
Ullah, B. and Ali, S. (1955) Review of mango budding in situ in the Punjab. Pakistan
Review of Agriculture 2, 7476, 96.
Van Overbeek, J. and Gregory, L.E. (1945) Investigations on the fundamentals of plant
propagation by means of cuttings. Second Annual Report of Director Institute of
Tropical Agriculture, Mayaguez, Puerto Rico for the Fiscal Year of 194344. Insti-
tute of Tropical Agriculture, Mayaguez, Puerto Rico, pp. 9299.
Veeraraghavan, R. (1945) Side grafting in mangoes. Indian Journal of Horticulture 3, 45.
Verma, S.R. (1942) A novel mango graft. Punjab Fruit Journal 6, 1183.
Walters, E.A. (1932) Plant propagation. Tropical Agriculture, Trinidad 9, 3539.
Whiley, A.W. (1993) Environmental effects on phenology and physiology of mango. A
review. Acta Horticulturae 341, 168176.
Crop Production: Propagation 403
Wolfe, B.S. (1963) The mango in Florida 1887 to 1962. Proceedings of the Florida
State Horticultural Society 75, 387391.
Yang, Z. and Ludders, P. (1993) Effect of growth regulator and media on in vitro shoot tip
culture of different cultivars of mango (Mangifera indica L.) rootstocks. Acta Horti-
culturae 341, 240247.
Zill, L. (1951) Top working of mango. In: Proceedings of the Florida Mango Forum.
Florida Mango Forum, Florida, p. 147.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
404 (ed. R.E. Litz)
12 Crop Production: Mineral Nutrition
I.S.E. Bally
Department of Primary Industries and Fisheries, Queensland, Australia
12.1 Introduction 405
12.2 Soils, Mineral Diagnosis and Sampling 405
12.3 Tissue Mineral Diagnosis and Sampling 406
12.4 Interpreting Soil and Leaf Analyses 406
12.5 Nitrogen 407
Sources, uptake and translocation of N 409
Nitrogen deciency and toxicity 409
Effect of N on crop production and fruit quality 410
Nitrogen management 411
12.6 Phosphorus 411
Sources, uptake and translocation of P 411
Phosphorus deciency and toxicity 412
Effect of P on crop production and fruit quality 412
12.7 Potassium 412
Sources, uptake and translocation of K 413
Potassium deciency and toxicity 413
Effects of K on crop production and fruit quality 414
12.8 Magnesium 414
Sources, uptake and translocation of Mg 414
Magnesium deciency and toxicity 414
12.9 Sulfur 415
Sources, uptake and translocation of S 415
Sulfur deciency and toxicity 415
12.10 Zinc 416
Sources, uptake and translocation of Zn 416
Zinc deciency and toxicity 416
Effect of Zn on crop productivity 417
12.11 Manganese 417
Role of Mn 417
Sources, uptake and translocation of Mn 417
Manganese deciency and toxicity 418
Crop Production: Mineral Nutrition 405
Effect of Mn on crop production 418
12.12 Iron 418
Sources, uptake and translocation of Fe 418
Iron deciency and toxicity 419
12.13 Calcium 419
Sources, uptake and translocation of Ca 419
Calcium deciency and toxicity 420
Effects of Ca on crop production and fruit quality 420
Calcium in leaves and fruit 421
12.14 Boron 422
Sources, uptake and translocation of B 422
Boron deciency and toxicity 423
Effect of B on crop production and fruit quality 423
12.15 Conclusion 424
12.1 Introduction
Assessment of the mineral status of mango trees is not without its challenges.
Like many other tropical woody perennial tree crop species, mangoes have
complicated and variable phenological cycles that inuence the trees uptake
and translocation of minerals. Their extensive root system enables them to
exploit unevenly distributed minerals throughout the soil prole; these min-
erals are often not assessed during routine soil analysis. The leaves, trunk,
bark and roots act as mineral reserves that buffer many short-term mineral
shortages (Robinson, 1986b). Soil and leaf mineral analyses used as short-
term indicators of tree mineral status, tree productivity or fruit quality are
therefore difcult and unreliable (Catchpoole and Bally, 1995).
12.2 Soils, Mineral Diagnosis and Sampling
Statistical relationships between the soil and tree mineral status have been
difcult to establish because of the large mineral reserves in trees and the
variable distribution of minerals in the soil prole (Robinson, 1986b). How-
ever, soil mineral analyses can be useful for determining the availability of
the essential minerals, and if they are within the optimal range for mango.
Soil analysis also provides valuable information on other soil properties that
can inuence mineral availability to the tree, i.e. pH, electrical conductivity
(Ec) and concentrations of organic matter and clays. Regular soil analysis in
conjunction with tissue analysis in mature cropping orchards can provide
useful information on the changes in soil minerals over time and on the
inuences of fertilizer programmes on soil and tree status.
There are several publications that outline the techniques that can be uti-
lized for obtaining representative soil samples for mineral analysis (Dow
et al., 1991; Peverill et al., 1999; Anonymous, 2007). Before a new orchard is
planted, the soil should be sampled from several depths and analysed to
capture a full picture of the mineral status throughout the soil prole. This
I.S.E. Bally 406
will facilitate the adjustment of pH and deep placement of minerals before
the trees are planted. In established orchards the sites of soil sampling will
vary according to the age of the trees and the soil type. In sandy or duplex
soil types where minerals are easily leached from the upper to the lower pro-
le, deep (11.5 m) monitoring will indicate any mineral build-up and help
avoid unnecessary fertilizer applications. In lighter sandy soils, surface sam-
pling to depths of 1015 cm can underestimate soil mineral concentrations as
these layers are often leached and are low in clay colloids.
12.3 Tissue Mineral Diagnosis and Sampling
Leaf mineral analysis is commonly used to assess mango tree mineral status,
and is useful for developing and monitoring tree fertilizer programmes.
Leaves often display visual symptoms of toxic and decient concentrations
of many minerals. Sampling mango leaves for mineral analysis should be
done when the tree is at its most phenologically quiescent stage, i.e. when
leaf mineral concentrations are most stable. One of the most stable periods in
the mango phenological cycle is the dormant phase, which occurs after the
completion of summer ushing and approximately 2 weeks before the emer-
gence of ower panicles. The common practice of withholding irrigation
water leading up to owering makes this period the most inactive of the year
and an ideal time for leaf sampling. At other times of the year, leaf mineral
concentrations sharply decrease when the tree is owering and fruiting and
increase in the months following harvesting (Catchpoole and Bally, 1995;
Oosthuyse, 2000b). Sampling at an inactive growth stage reduces variability
between leaf samples and provides a stable reference point for annual com-
parisons. Guides with respect to the most appropriate leaves for sampling
have been variously reported (Kumar and Nauriyal, 1979; Chadha et al., 1980;
Smith, 1992; Catchpoole and Bally, 1995), and generally concur that the most
appropriate leaves to sample are the third or fourth leaf behind the apical
bud, or the rst full-size leaf of the most recently matured dormant ush
where leaves are fully expanded and hardened off.
Leaves recently sprayed with foliar nutrients or fungicides should be
avoided or noted, as analyses of manganese (Mn), zinc (Zn), boron (B) and
copper (Cu) are commonly affected by mineral residues on the outside of
leaves or imbedded in the cuticle, and which are not available to the tree.
Some publications recommend that sampled leaves should be washed with
deionized water to reduce residues from sprays and dust from the operators
hands (Reuter et al., 1986; Shu et al., 1992).
12.4 Interpreting Soil and Leaf Analyses
Several published soil and leaf mineral standards are available to assist in
interpretating analysis results (Smith and Scudder, 1951; Young and Koo,
1969; Kumar and Nauriyal, 1977; Robinson, 1986a; Peverill et al., 1999; Stassen
Crop Production: Mineral Nutrition 407
et al., 1999) (Table 12.1). Most of these standards are in reasonable agreement
with respect to the optimal ranges of individual minerals (Samra and Arora,
1997). Soil and leaf analyses should be interpreted by comparing with these
standards and with previous soil and leaf analyses and past seasons crop-
ping and fertilizer application history. The recommended optimal ranges of
leaf mineral concentrations should be considered as general guides only, as
high-yielding trees producing good quality fruit have been found to vary
widely in leaf mineral composition (Catchpoole and Bally, 1995; Oosthuyse,
2000b; Medeiros et al., 2004).
Positive relationships between leaf minerals and tree productivity have
been reported. Oosthuyse (1997) determined that leaf concentrations of nitro-
gen (N), phosphorus (P), potassium (K), magnesium (Mg) and Zn inuenced
the number of fruit retained, and that leaf Zn and Mg also inuenced fruit
size. Rao and Mukherjee (1989) observed positive correlations between tree
yields and leaf N and K from non-fruiting terminals in ve Indian mango
cultivars with generally low N and K concentrations. Although these rela-
tionships have been observed, there are many factors that can inuence pro-
ductivity. Therefore, predicting productivity based on leaf analysis alone is
difcult. However, soil and leaf analyses are useful for identifying major
mineral imbalances and long-term trends in tree nutrition, and can be used
to adjust fertilizer programmes.
Leaf age can affect the mineral concentration in assays. Minerals that are
mobile within the plant, for example N, P, K and Mg, generally decrease during
leaf aging and the less mobile minerals, such as calcium (Ca), sulfur (S), B and
Mn, accumulate in leaves with age (Chadha et al., 1980; Medeiros et al., 2004).
Mango leaves that have been sampled from fruiting terminals generally
display increasing concentrations of N, K, Ca, Mn, iron (Fe), Cu and Zn dur-
ing early fruit development and decreasing concentrations during late fruit
development and maturation. These minerals, with the exception of Mg and
P, also vary greatly among fruiting terminals (Oosthuyse, 1997, 2000b).
The Diagnosis and Recommendation Integrated System (DRIS) for inter-
preting leaf mineral analysis, uses ratios of mineral concentrations rather
than the absolute mineral concentration to identify limiting minerals in order
of their effect on the tree (Beauls, 1973). With mango, DRIS has been used
with varying success. Raghupathi et al. (2004) observed that DRIS was unable
to diagnose the nutrient imbalance of a particular nutrient in isolation; how-
ever, others have used the technique more successfully. Schaffer et al. (1988)
used DRIS to identify Mn and Fe as the most decient elements in orchards
with tree decline, a disorder of unknown aetiology, and Hundal et al. (2005),
Raj and Rao (2006) and Wadt et al. (2007) utilized DRIS to identify yield-
limiting mineral combinations in mangoes grown in India and Brazil.
12.5 Nitrogen
Nitrogen is one of the most important elements for crop production, and has
a signicant role in mango growth, yield and fruit quality. Nitrogen is an

I
.
S
.
E
.

B
a
l
l
y
4
0
8
Table 12.1. Suggested optimal mango leaf mineral concentration ranges according to different sources.
Mineral
a
References
Unit
Robinson
et al. (1997)
Smith and
Scudder
(1951)
Young and
Koo (1969,
1971), Young and
Sauls (1981)
Crane
et al.
(1997)
Catchpoole
and Bally
(1995)
Kumar
and Nauriyal
(1977)
Pimplasker
and Bhargava
(2003)
Stassen
et al.
(1999)
Bhargava
and Chadha
(1988)
N 1.01.5 1.54 1.01.5 1.21.6 0.81.9 1.0 0.891.93 1.25 1.23
P % 0.080.18 0.05 0.090.18 0.090.12 0.121.3 0.10 0.060.11 1.45 0.06
K % 0.31.2 0.97 0.51.0 0.40.8 0.42.5 0.50 1.022.01 0.1 0.54
Ca % 2.03.5 0.91 3.05.0 2.03.5 1.52.8 1.50 0.81.05 1.71
Mg % 0.150.4 0.26 0.150.47 0.250.35 0.20.4 0.15 2.8 0.91
S % 0.50.6 0.10.23 0.50 0.110.17 0.3 0.12
B % 5080 2484 20140 50
Fe mg/kg 7200 38120 70100 30120 80 1.71
Mn mg/kg 60500 92182 160980 80 66
Zn mg/kg 20150 10119 2040 2063 1126 40 25
Cu mg/kg 1020 2835 10150 20 12
Mo mg/kg 0.20.4 50
a
N, nitrogen; P, phosphorus; K, potassium; Ca, calcium; Mg, magnesium; S, sulfur; B, boron; Fe, iron; Mn, manganese; Zn, zinc; Cu, copper;
Mo, molybdenum.
Crop Production: Mineral Nutrition 409
essential component of many plant tissues, and occurs in chlorophyll, amino
acids, proteins, enzymes and growth hormones and is a major driver of plant
growth, having a direct effect on tree vigour (Marschner, 1995). Nitrogen at
100 g/tree/year has been shown to be sufcient for mango tree growth
(Kanwar et al., 1987), and N concentrations inuence concentrations of other
elements when N is either low or in excess (Sen et al., 1947).
Sources, uptake and translocation of N
Nitrogen in the soil occurs in many forms, but for the most part as large and
complex organic molecules that comprise the organic matter. These mole-
cules are too large for roots to absorb, and are broken down to nitrate (NO
3

)
and ammonium (NH
4
+
). The concentration of N in soil is dependent on the
concentrations of soil organic matter and mineral N. Nitrogen that is avail-
able to the plant is determined by the processes of mineralization, immobili-
zation, de-nitrication, volatilization and leaching, which are inuenced by
temperature, moisture, pH and aeration. Nitrogen is easily leached from the
soil by rain and irrigation, and consequently, soil N often limits tree growth,
especially in sandy soils (Strong and Mason, 1999).
Common N fertilizers include: urea (CO(NH
2
)
2
, 46% N), potassium
nitrate (KNO
3
, 13% N, 38% K), calcium nitrate (CaNO
3
, 15.5% N, 19% Ca),
ammonium nitrate (NH
4
NO
3
, 35% N), sulfate of ammonia ((NH
4
)
2
SO
4
, 21%
N, 23.6% S). Nitrogen is taken up by mango trees primarily through the roots
as NO
3

and NH
4
+
; NO
3

is the preferred source. Nitrogen can also be adsorbed


through leaves as ammonia (NH
3
), urea and amino acids. Nitrate, after being
adsorbed by the roots, is either reduced to NH
4
+
in the roots or translocated
to the leaves, stems or other tissues, where it is reduced to NH
4
+
in a two-step
process in which NO
3

is reduced to nitrite (NO


2

) which is in turn reduced to


NH
4
+
(Marschner, 1995). Ammonium metabolism is complex, and several
pathways are necessary to produce amino acids, proteins, enzymes and hor-
mones. Within the tree, N concentrations vary among tissues and are depen-
dent on N availability in the soil and demand within the tree. If N is limiting,
the plant can translocate N from older tissues to new growing tissues where
it is required (Marschner, 1995).
Nitrogen deciency and toxicity
Nitrogen deciency symptoms appear in leaves as yellowing or chlorosis
initially of the older leaves, slow growth and lack of vigour. Severe N de-
ciency can cause complete leaf yellowing, leaf and fruit abscission and death
of twigs and branches (Sen et al., 1947; Smith and Scudder, 1951). Nitrogen
toxicity is not often seen in mango; however, trees with high N concentra-
tions have dark green leaves and excessive vegetative vigour, often at the
expense of owering and cropping (Tiwari and Rajput, 1976). High N con-
centrations are also associated with internal fruit disorders, e.g. jelly seed
I.S.E. Bally 410
and internal breakdown in many cultivars (Tarmizi et al., 1993; Cracknell
Torres et al., 2004; see Galn Saco, Chapter 9, this volume).
Effect of N on crop production and fruit quality
Nitrogen has a major effect on mango tree vigour, stimulating both vegeta-
tive and oral growth. Increases in shoot length, leaves per shoot and leaf
area have been demonstrated by Singh et al. (1973), Tiwari and Rajput (1975),
Syamal and Mishra (1989), Reddy et al. (2000) and Sergent et al. (2000).
Yeshitela et al. (2005) reported that N in combination with K, as KNO
3

and urea, improved the percentage of terminal shoots that ower; however,
excessive N can stimulate vegetative growth at the expense of owering,
fruit set and fruit quality (Scholeeld et al., 1986; Monselise and Goren, 1987;
Silva et al., 2002). Nitrogen can cause increased fruit set and retention (Singh
et al., 1973; Oosthuyse, 1997) and fruit weight and yield (Alvian, 1974; Tiwari
and Rajput, 1975; Young and Koo, 1975; Alvian and Figueroa, 1977; Syamal
and Mishra, 1989; Reddy et al., 2003). Kanwar et al. (1987) demonstrated that
N at 100 g/tree/year-of-age was sufcient for tree growth and Young and
Koo (1975) reported that maximum yields were increased by N applications
of 0.81.1 kg/tree/year. Many reports of the effect of N on increased mango
fruit size and yields are based on data of N in combination with other ele-
ments (i.e. P and K) and may be partly attributed to these combinations;
however, N has a major inuence on productivity in mango.
High N application rates that stimulate yield increases can also have
negative effects on fruit quality. Nitrogen has been negatively associated
with fruit colour in mango (Oosthuyse, 1993; McKenzie, 1994, 1995). Nguyen
et al. (2004) demonstrated that high N applications during fruit growth
inhibited the de-greening of ripening fruit, causing green skin at ripeness.
Bally (2007) also reported a negative relationship between N and fruit colour,
demonstrating that high leaf N concentrations reduce the percentage of
yellow skin in ripe fruit, reduced the lightness and chroma (vividness) of the
yellow colour, the percentage of skin covered with blush and the intensity of
the blush colour (Plate 73). He also identied a signicant exponential rela-
tionship between the severity of sunburn and skin N concentrations; the
effects of sunburn were reduced as N concentrations increased in the fruit
skin.
Early indications of the negative effect of N on postharvest rots in mango
were demonstrated by Weng and Chuang (1997), who observed that N posi-
tively affected the germination rate, hyphal growth and appressoria forma-
tion of Colletotrichum gloeosporioides. When Nguyen et al. (2004) investigated
the effect of N on fruit quality, they found that 300 g/tree applied as a
foliar spray signicantly increased the severity of anthracnose caused by C.
gloeosporioides. These observations were conrmed by Bally (2007), who found
that the severity of fruit anthracnose during ripening increased with applied
N (Plate 74). He also demonstrated that the N effect was because N enhanced
the decline of antifungal resorcinol compounds in the fruit exocarp during
Crop Production: Mineral Nutrition 411
ripening. Nitrogen has been a recognized factor in determining internal fruit
quality of mangoes, with imbalances of N and other minerals, principally Ca,
being implicated as a major factor causing the various forms of internal
breakdown (Young and Miner, 1961; Subramanyam et al., 1971; Malo and
Campbell, 1978; Burdon et al., 1992; Tarmizi et al., 1993; Cracknell Torres et al.,
2004; Torres et al., 2004; see Galn Saco, Chapter 9, this volume).
Nitrogen management
Leaf N concentrations between 1 to 1.5% dry weight (DW) are generally con-
sidered to be in the optimal range (Robinson et al., 1997). Optimum concen-
trations for fruit skin N have not been published, although Catchpoole and
Bally (1995) reported skin N concentrations of 0.069% DW in mature Kens-
ington Pride fruit. Leaf N concentrations vary throughout the year, and are
inuenced by the growth events during the phenological cycle. Nitrogen
concentrations are generally greatest at the end of the summer vegetative
ushing period and decrease during panicle growth, owering and fruiting
(Catchpoole and Bally, 1995; Medeiros et al., 2004).
Because N is highly mobile within the tree and a primary driver of
growth and fruit quality, the general practice of monitoring leaf and soil N
annually is inadequate for assessing tree N status at all stages of tree phenol-
ogy. A cheap, rapid test for N is needed to provide closer monitoring of
changes in leaf N status and allow the rates and timing of supplementary N
applications to be matched to tree and fruit demands. Bally and Still (per-
sonal communication) have calibrated the Konica-Minolta Soil Plant Analy-
sis Diagnostic (SPAD-502) meter to measure the chlorophyll content of leaves,
which is directly related to their N status (Gonzalez et al., 2005).
12.6 Phosphorus
Phosphorus is important for cell division and growth and is a component of
many essential plant molecules such as sugar-phosphates that are involved
in respiration, photosynthesis and other metabolic pathways, nucleotides
such as DNA and RNA, phospholipids in membranes and as pryophosphate
(PPi) in ATP and other cellular energy metabolism molecules (Salisbury and
Ross, 1992). Phosphorus is also involved in root tip elongation, fruit ripening
and leaf expansion.
Sources, uptake and translocation of P
Phosphorus that is available for plants occurs in two forms in soil, primarily
as the monovalent phosphate anion (H
2
OP
4

) and secondly as the divalent


anion (HPO
4
2
) in the soil water solution. The balance of these anions is
dependent on soil pH, with HPO
4
2
favoured in soils >pH 7 and H
2
OP
4

I.S.E. Bally 412
favoured in soils <pH 7 and most readily available in soils with pH between
6 and 7. In higher pH soils, calcium phosphate compounds tie up P from
plant availability, and in lower pH soils, P oxidizes with Fe, aluminium (Al)
and Mn to become unavailable to plants. Other forms of soil P include liable
P that is bound on clays and organic matter, which can become available to
trees when dissolved in water.
Phosphorus is most rapidly taken up by the tree as HPO
4
2
and slower as
H
2
OP
4

and often enhanced by chemical association with N. After entry into


the roots, the anions are either converted to organic phosphates immediately
or after transport to other tissues. Within the tree, P is easily translocated
between tissues, with redistribution usually occurring from older leaves to
younger actively growing tissues. Phosphorus concentrations within the
mango tree are highest in the roots, wood and bark and lowest in the leaves
(Vuuren and Stassen, 1997). Leaf P concentrations are generally at their low-
est during fruit development and at their highest in the vegetative growth
period (Medeiros et al., 2004).
Phosphorus deciency and toxicity
As P is a mobile element, deciencies are initially seen in the older leaves.
Phosphorous deciency and toxicity are rare in mango trees, with toxicity
symptoms occasionally seen in nursery stock where trees are grown in pot-
ting media based on sugarcane mill/press mud. Symptoms appear as stunted
seedling growth with bronzing of the leaves. Phosphorus deciency appears
rst in older leaves as marginal necrosis with brown taints, tip necrosis, pre-
mature abscission and stem dieback (Smith and Scudder, 1951; Singh and
Saxena, 1994). Sen et al. (1947) describe deciency symptoms in leaves as
developing a reddish-purple colour on the undersides of leaves that spreads
over the entire leaf, eventually encompassing the veins, with leaves becom-
ing thick and stiff. Phosphorus-decient trees have restricted root development
that reduces tree and fruit size (Stassen et al., 1999).
Effect of P on crop production and fruit quality
There are several reports of P applied in combination with N and/or K and
resulting in increased yields (Samra and Arora, 1997); however, these reports
do not separate the P from the other elements. Reddy and Majmudar (1985)
reported an association between higher concentrations of P in terminals and
oral induction (Narwadkar and Pandey, 1989).
12.7 Potassium
Potassium in the cytoplasm is an activator of enzymes involved in photosyn-
thesis, respiration and starch and protein synthesis (Bhandal and Malik, 1988;
Crop Production: Mineral Nutrition 413
Marschner, 1995). Potassium is important for cell growth due to its role in cell
expansion and development of thick epidermal cell walls that increase the
resistance of trees to pathogens and insect pests. Potassium is involved in
tree water status by regulating water uptake by the roots and water loss
through the leaf stomata (Salisbury and Ross, 1992).
Sources, uptake and translocation of K
Most of the K in soils (9098%) is in the form of insoluble crystalline minerals
that are unavailable to plants. Available K occurs in the soil solution as K
+
ions and attached to cation exchange sites on clays (Gourley, 1999). Potas-
sium moves readily between the exchange sites and the soil solution and is
inuenced by moisture and temperature. Heavy applications of Ca and Mg
fertilizers compete with K for exchange sites, thereby reducing the availability
and uptake of K (Lim and Khoo, 1985). Potassium concentrations are lower
in soils with low cation exchange capacities, e.g. granites, sands, highly
leached and acidic soils, and are higher in clay soils (Lim and Khoo, 1985;
Gourley, 1999).
The main K fertilizers used in mango production are potassium chloride
(KCl) also known as muriate of potash, potassium sulfate (K
2
SO
4
) and potas-
sium nitrate (KNO
3
). Potassium sulfate is generally preferred to KCl because
it has a neutral effect on pH and because mangoes are very sensitive to chlo-
rides. Mango trees take up K as the K
+
ion from the soil solution. Potassium
is readily redistributed, typically from old leaves to young growing tissues.
The highest concentrations of K occur in leaves, fruit, roots and bark (Vuuren
and Stassen, 1997). Leaf K concentrations are generally at their highest at the
end of the postharvest summer vegetative ushing period, decreasing with
owering and fruiting (Catchpoole and Bally, 1995; Medeiros et al., 2004).
Mango trees can absorb more K than is required for maximum tree growth;
excessive uptake of K can cause imbalances with other minerals in the tree
(Gourley, 1999).
Potassium deciency and toxicity
Potassium deciency rst appears in older mango leaves as the tree redistrib-
utes K to young growing tissues. Sen et al. (1947) induced K deciency by
growing mangoes in sand culture and observed brown necrosis on the leaf
margins extending from the leaf tip towards the base. Smith and Scudder
(1951) also generated K deciency symptoms in sand culture and described
them as irregularly distributed yellow spots and necrotic areas along the
margins of small, thin attenuated leaves which are very persistent. Lim and
Khoo (1985) described the non-necrotic areas of the leaf as dull, yellowish
green to light green; symptoms usually develop in the dry season or when
irrigation has been stopped.
I.S.E. Bally 414
Effects of K on crop production and fruit quality
Potassium nitrate applications just prior to and at the owering stage promote
owering, increase fruit set and fruit retention (Sergent and Leal, 1989; Lyan-
naz, 1994; Ferrari and Sergent, 1996; Oosthuyse, 1997; Rojas and Leal, 1997;
Sergent et al., 1997; Saleh and El-Monem, 2003; Shinde et al., 2006). In the low
and mid-latitude tropics, KNO
3
is used to stimulate out-of-season owering;
however, this effect is lost in higher latitudes (Davenport and Nez- Elisea,
1997; see Davenport, Chapter 5, this volume). Shongwe and Roberts-Nkrumah
(1997) suggested the KNO
3
effect on owering is a result of lowering the tran-
spiration rate and increasing water use efciency. Protacio (2000) suggests the
KNO
3
effect on owering is primarily due to N stimulation rather than K and
he postulated that KNO
3
overcomes the inhibitory effects of gibberellic acid
(GA
3
) on starch accumulation by elevating the N concentrations over the N
threshold to synchronize bud break from apices with an existing oral initial.
Potassium inuences fruit quality of many species (Marschner, 1995);
however, in mango there are only a few studies that link K nutrition with
increased fruit quality. Shinde et al. (2006) observed that increased K fertiliza-
tion increased fruit weight (5.15%), ascorbic acid (26.99%), organoleptic score
for texture, avour, colour and shelf life and reduced physiological weight loss
(22.79%) and spongy tissue (68.08%). Potassium in the form of monopotassium
phosphate (KH
2
PO
4
) suppresses powdery mildew on mango (Reuveni et al.,
1998; Oosthuyse, 2000a), but it is not clear if the effect is due to P or K.
12.8 Magnesium
Magnesium is a component of enzymes that transport P, develop green col-
oration in chlorophyll, activate other enzymes and is involved in carbohydrate
metabolism and nucleic synthesis (Marschner, 1995; Stassen et al., 1999).
Sources, uptake and translocation of Mg
Magnesium is present in soil as the minerals biotite, serpentine, olivine and
hornblende, in organic matter, and as exchangeable Mg
2+
in the soil solution.
Mg
2+
is the second most dominant cation after Ca
2+
and successfully com-
petes with K
+
and Na
+
, replacing them on exchange sites (Aitken and Scott,
1999). Magnesium is a mobile element that is taken up by the plant as Mg
2+
.
Magnesium concentrations in the tree can be depressed by high concentra-
tions of other cations, such as Mn
2+
, Ca
2+
, K
+
, NH
4
+
and Al
3+
, in the root
environment (Aitken and Scott, 1999).
Magnesium deciency and toxicity
Magnesium deciency rst appears in older leaves, due to its high mobility
within the tree, as pale green or yellow leaves with the inner vein areas of the
Crop Production: Mineral Nutrition 415
leaf lamina becoming mottled and necrotic while the leaf veins remain green.
Young trees are stunted by Mg deciency. Smith and Scudder (1951) gener-
ated symptoms of Mg deciency in sand culture and described them as a
green wedge pattern formed by the lateral intrusion of a bronzed chlorosis
along the leaf margin (Plate 75). Deciency symptoms are more likely to
occur in textured, sandy or highly leached soils due to their low cation
exchange capacity or when heavy applications of liming products, K fertil-
izers or green manuring have been applied (Stassen et al., 1999). Magne-
sium deciency has been associated with a browning skin discoloration in
Kensington Pride fruit.
12.9 Sulfur
Sulfur is a macro element required in large amounts by the tree. Sulfur is an
essential component of the amino acids cystine and methionine that make up
photosynthetic proteins, the vitamins thiamine and biotin and coenzyme-A used
in the synthesis and breakdown of fatty acids (Salisbury and Ross, 1992).
Sources, uptake and translocation of S
Up to 7090% of S in soil is present in organic matter (Marschner, 1995), and
sulfates that are available for plants are mainly found in the surface layers
and the soil solution with only small amounts adsorbed to soil colloids. As a
result, S is very prone to leaching. High rainfall, light-textured and dry soils
are adverse to S accumulation and uptake (Anonymous, 1988).
Supplementary sources of S include sulfates as minor components of fer-
tilizers, i.e. superphosphate, sulfate of ammonia, potassium sulfate and gyp-
sum as well as some fungicides and miticides. Organic-based sulfate
compounds are not available to plants until they have been converted to the
sulfate ion (SO
4
2
) by bacteria in the mineralization process. The divalent sul-
fate anions (SO
4
2
) are actively taken up by the roots and translocated to
shoots or growing tissues. Sulfur can also be adsorbed through the leaves as
sulfur dioxide (SO
2
), usually from pollution from burning fossil fuels. Sulfur
is not readily re-translocated from one tissue to another. The competitive
effects of other anions such as phosphates and nitrates can reduce the uptake
of S (Marschner, 1995).
Sulfur deciency and toxicity
Sulfur deciency in mango is uncommon as it is generally present in ade-
quate quantities in most soils and is a component of many fertilizers, fungi-
cides and miticides. Smith and Scudder (1951) describe S deciency in mango
as lateral necrotic spots that occur on the vascular bundles and lamina on a
very deep-green leaf and premature defoliation. In other species symptoms
I.S.E. Bally 416
often appear to be similar to N deciency, but are distributed more evenly
between young and old leaves, with the younger tissues developing symp-
toms rst as S is not easily redistributed from older to younger tissues
(Salisbury and Ross, 1992; Marschner, 1995).
Sulfur toxicity occurs when pollutants high in SO
2
content are converted
to bisulfate (HSO
3
) when combined with water, and is further oxidized to
sulfuric acid (H
2
SO
4
), often known as acid rain. Sulfur dioxide adsorbed by
leaves reacts with water to form HSO
3
that is toxic to the leaf, inhibiting
photosynthesis and degrading chlorophyll (Marschner, 1995).
Klumpp et al. (2003) reported soil S concentrations of between 53.2 and
86.0 mg/kg DW in contaminated soils in the vicinity of a copper smelter in Brazil
compared with 3348 mg/kg DW in uncontaminated soils. Leaves from mango
trees growing in the contaminated soils had S concentrations of 3.8 mg/g DW,
double that of trees growing in uncontaminated soils (1.9 mg/g DW). At these
concentrations no obvious leaf symptoms were visible, indicating that mango
has a high tolerance of S in the soil and atmosphere. Information on mango leaf
S concentrations is scarce; however, a wide range of concentrations are consid-
ered to be optimal for mango growth (Table 12.1): 0.66.4 mg/g DW S (Marchal
et al., 1991), 1.071.69 mg/g DW S (Pimplasker and Bhargava, 2003). Chaudhary
and Nauriyal (1988) induced leaf deciency symptoms in 1-year-old seedlings in
sand culture when leaf S concentrations were between 0.32% and 0.74% DW.
12.10 Zinc
Zinc is chemically bound to Fe and Mn to form components of chlorophyll, and
is essential for photosynthesis (Weir and Cresswell, 1995). Zinc is essential for the
synthesis of proteins and hormones, including auxins, and is required for the
maintenance of biomembranes (Salisbury and Ross, 1992; Marschner, 1995).
Sources, uptake and translocation of Zn
Zinc is present in soil in a range of inorganic minerals and organic matter
with the solubility of the inorganic forms decreasing as soil pH increases.
Common Zn fertilizers include zinc oxide (ZnO, 80% Zn), zinc chelates
(ZnNa
2
EDTA and Zn(NH
4
)
2
EDTA, 6.514% Zn), zinc sulfates (ZnSO
4
, 23%
Zn). Zinc is taken up by the tree in the form of the zinc ion (Zn
2+
), and trans-
located to regions of growth such as shoot tips. Zinc is not readily translo-
cated to other tissues. Zinc is readily taken up through the leaves, and foliar
Zn application is a common method of managing Zn in mango orchards.
Zinc deciency and toxicity
Zinc deciency is often referred to as little leaf because leaves fail to reach
full size. Symptoms rst appear in immature leaves in the coloured stage, as
Crop Production: Mineral Nutrition 417
a thickening of the leaf and failure to fully expand. Leaf expansion is often
stunted on one side of the blade, causing it to form a sickle shape (Dilly et al.,
1997) (Plate 76). Symptoms in fully mature leaves are prominent light-yellow
or olive-green veins on the upper surface that are thick and brittle. Internode
length is reduced, thereby causing a rosetting effect (Lim and Khoo, 1985;
Agarwala et al., 1988; Marschner, 1995). Zinc and Fe deciency often occur
together because of their association with calcareous high pH soils.
Effect of Zn on crop productivity
Singh and Rajput (1977) reported that foliar sprays of 2.08.0% ZnSO
4
prior
to owering increased the number of perfect owers in panicles and later
increased fruit yield, fruit sugar, ascorbic acid and total soluble solids (TSS).
Daulta et al. (1981) also observed that foliar Zn sprays increased fruit set and
improved fruit quality, but had no effect on sex ratio. Kumar and Kumar
(1989) demonstrated that 1% ZnSO
4
sprays applied to Dashehari trees
improved postharvest fruit life by reducing weight loss, spoilage, increasing
sugars, reducing acidity and slightly increasing vitamin A. Littlemore et al.
(1991) observed that 1% ZnSO
4
foliar sprays applied quarterly to Kensing-
ton Pride was sufcient to maintain leaf Zn concentrations above critical
concentrations and avoid symptoms of little leaf. Although leaf Zn concen-
trations were improved and symptoms cured, no effect on tree yields was
observed. Soil applications are often ineffective due to adverse soil conditions
making it unavailable to trees.
12.11 Manganese
Role of Mn
Manganese is a trace element that is required in small amounts by the tree; it
is important in the redox processes and is a cofactor, activating many enzymes
that catalyse oxidation, reduction, decarboxylation and hydrolytic reactions.
Manganese is an essential mineral for photosynthesis and the biosynthesis of
proteins, carbohydrates and lipids (Marschner, 1995).
Sources, uptake and translocation of Mn
Manganese is found in soil in the insoluble form of manganese oxide (MnO
2
)
which is reduced to the water soluble exchangeable ion (Mn
2+
) that is taken
up by tree roots. Mn
2+
is mainly found in the soil solution but can also be
loosely bound to organic matter and clay colloids. Reduction of MnO
2
to
Mn
2+
is carried out by organic reducing agents in aerobic soils and is favoured
by reduced pH. As the soil pH increases, microbial oxidation of Mn
2+
increases.
Both reduction and oxidation processes occur in the soil simultaneously, but
I.S.E. Bally 418
the balance of available Mn is governed by soil pH, with low pH favouring
reduction and high pH favouring oxidation (Peverill et al., 1999).
Manganese deciency and toxicity
Manganese deciency in mango is initially expressed by light necrosis
between the main veins of middle and younger leaves that later become
necrotic. Buff-coloured spots appear on the primary leaf veins at the margins
of the leaves and coalesce to form necrotic patches before leaf abscission. In
severe Mn deciency, necrosis begins at the leaf tip, spreading to the base of
the lamina (Smith and Scudder, 1951; Agarwala et al., 1988).
Effect of Mn on crop production
There are few reports on the effect of Mn on crop production or fruit quality.
Schaffer (1994) suggested that mango decline was associated with trees that
are decient in Mn and Fe.
12.12 Iron
Iron is a trace element that is required in small quantities, and is a component
of several enzymes and molecules. It is a component of chlorophyll and leaf
Fe concentrations directly impact photosynthesis. Iron-containing enzymes
participate in oxidation processes that release energy from sugars and
starches, the conversion of nitrates to ammonia and the biosynthesis of eth-
ylene (Marschner, 1995). In proteins, Fe acts as an electron carrier by alternate
oxidation and reduction between Fe
2+
to Fe
3+
(Salisbury and Ross, 1992).
Sources, uptake and translocation of Fe
Available Fe occurs in the soil solution in very low concentrations (~10
15
M)
as the ionic forms Fe
2+
and Fe
3+
. Fe
2+
is far more soluble and more easily
taken up than Fe
3+
; however, in well-aerated soil, Fe
2+
is readily oxidized to
Fe
3+
and precipitated, becoming unavailable. Fe
2+
can also be displaced by
other cations or by alkaline soils (Salisbury and Ross, 1992). Plants partly
overcome Fe unavailability by producing ligands that bind to Fe
3+
to form
soluble chelates that maintain Fe in solution. On the root surface the Fe
3+
is
reduced to Fe
2+
which is immediately adsorbed by the roots. This process is
often inhibited in high pH soils and in calcareous soils. Iron deciency is com-
mon in mango (Schmidt, 1999). Iron is readily absorbed through leaves,
which is a useful method for management of Fe when soil conditions are not
conducive to uptake.
Crop Production: Mineral Nutrition 419
Iron deciency and toxicity
Iron deciency initially appears in young leaves as reduced concentrations of
chlorophyll. Iron-decient leaves are pasty yellow/green with developing
chlorosis of the whole leaf as severity increases (Plate 77). Leaves fail to
develop to full size and necrosis of the leaf tips occurs in severe cases. Iron
deciency is often seen on calcareous soils with >20% calcium carbonate
(CaCO
3
), with poor aeration and drainage in the coolest part of the year
(Marschner, 1995). Iron deciency reduces the ability of mango trees to
photosynthesize, thereby stunting tree growth and reducing yields.
12.13 Calcium
A major role of Ca is membrane stability, which is achieved when Ca
2+
binds
to phosphate and carboxylate groups of phospholipids and proteins on the
surface of the plasmalemma, and thereby preventing leaking of solutes to the
cytoplasm (Kirkby and Pilbeam, 1984). Calcium protects cells from toxins,
slowing the aging of plant tissues and promoting longer shelf life of many
fruits. Calcium is important for pectin polymers that strengthen cell walls
and provide defence from pathogens (Ferguson, 1984). Most Ca in trees is
xed in cell walls and is not easily translocated. Calcium is essential for new
root hair and leaf development.
Sources, uptake and translocation of Ca
Soil Ca occurs as exchangeable, non-exchangeable and soluble forms.
Exchangeable Ca makes up 6585% of the total exchange capacity of normal
soils, and is weakly bonded, allowing rapid exchange with the soil solution,
and is thus prone to leaching (Kirkby, 1979; Mengel and Kirkby, 1987). Non-
exchangeable Ca is bound to minerals, for example feldspars, amphiboles,
phosphates and carbonates. Soil Ca concentrations are often up to ten times
higher than other cations such as K
+
and Mg
2+
and usually similar or slightly
higher than sodium ions (Na
+
) (Mengel and Kirkby, 1987).
Calcium is taken up passively through limited unsuberized tips of
actively growing roots and moves from the root cortex to the stele mainly
through the apoplast or free space of the root tips. It then enters the xylem,
moving upwards with the mass transpiration ow (Bangerth, 1979; Mengel
and Kirkby, 1987; Ho and Adams, 1989; Marschner, 1995). Root Ca concentra-
tions are lower than other cations such as K
+
and Mg
2+
because it is passively
taken up and easily replaced from the exterior surface of the plasma mem-
brane by other cations (Marschner, 1995). Root Ca concentrations can there-
fore be inuenced by the relative concentrations of other cations (K
+
, Mg
2+
,
NH
4
+
) in the soil solution. The suppression of Ca uptake by high soil concen-
trations of NH
4
+
may account for the link between high N nutrition and many
Ca-related disorders. On the other hand, high NO
3

can stimulate the uptake


I.S.E. Bally 420
of cations by stimulating organic anion synthesis, which attracts Ca
2+
(Kirkby,
1979; Mengel and Kirkby, 1987).
Calcium has low mobility in the phloem (Bangerth, 1979; Kirkby and
Pilbeam, 1984), limiting translocation within the tree. Ca required by devel-
oping organs is supplied via the xylem. Calcium deciencies are often caused
by low availability, poor uptake and/or poor translocation within the plant.
Root pressure can also contribute to the mass ow, but the extent to which
root pressure contributes to Ca ow in mango is unclear.
The intensity of evaporation from any organ will directly inuence the
amount of Ca the organ receives (Marschner, 1995). Low transpiring plant
organs, such as fruit, are more prone to Ca deciencies than actively transpir-
ing leaves. This is especially the case if the organ is fast growing and requires
high amounts of Ca (Mengel and Kirkby, 1987; Marschner, 1995). Limited Ca
translocation occurs in the absence of mass ow through exchange adsorp-
tion in the xylem vessels, similar to an exchange column (Mengel and Kirkby,
1987). As tissues grow, the cation exchange sites of the new cell walls act as a
sink for the Ca in the xylem exchange column, drawing the Ca up the column
(Clarkson, 1984). This exchange movement of Ca may be important for lower
transpiring organs such as the shoot apex and fruit (Mengel and Kirkby, 1987;
Marcelle et al., 1990). The translocation of Ca is induced by the polar move-
ment of indole-3-acetic acid (IAA) that is produced in the shoot apex and
rapidly growing organs (Banuelos et al., 1987; Ho and Adams, 1989).
Calcium deciency and toxicity
Calcium deciency symptoms reect the low mobility of Ca within the tree,
rst appearing in young, actively growing tissues. Ca-decient plants generally
display symptoms of membrane degeneration, which have been likened to
senescence and fruit ripening (Fallahi et al., 1977; Bangerth, 1979). Many symp-
toms of internal disorders in mango appear to be associated with premature
ripening or cell degeneration (Burdon et al., 1991, 1992). Raymond et al. (1998a)
observed that cell disruption and rupture of the cell walls were the rst micro-
scopic indicators of soft nose, stem-end cavity and jelly seed. High Ca concen-
trations reduce the binding sites for enzymatic degradation (Rhodes, 1980). No
direct symptoms of Ca toxicity have been recorded in mango, but high concen-
trations of Ca in soils can displace other minerals such as Mg, Zn, B, Cu and P.
Effects of Ca on crop production and fruit quality
In mango, fruit Ca has been implicated as a factor in fruit quality, for example
with regards to internal physiological disorders, shelf life and disease resis-
tance. Many symptoms of internal physiological disorders appear to be asso-
ciated with low Ca-related membrane degeneration, i.e. premature ripening
(Burdon et al., 1992; Raymond et al., 1998a). Links between internal break-
down and Ca nutrition in mango were reported by Young (1957), who found
Crop Production: Mineral Nutrition 421
that increasing Ca fertilization generally resulted in decreased incidence of
soft nose. Young and Miner (1961) and Shear (1975) suggested a link existed
between Ca and N with respect to physiological disorders by demonstrating
the negative effect of N on internal breakdown could be partly overcome by
high Ca concentrations in the fruit. When Malo and Campbell (1978) were
unable to alter internal breakdown in Tommy Atkins by N or K fertilization
over a 4-year period, the high Ca in the calcareous soils of Florida was
suggested as a possible reason.
Recent investigations have monitored Ca concentrations in fruit as well
as in soil. Subramanyam et al. (1971) observed that fruit with spongy-tissue
disorder had Ca concentrations of 74 mg/100 g DW, compared with
85 mg/100 g DW for healthy fruit. Rane et al. (1976), Kadiyala (1995) and
Tarmizi et al. (1993) also reported signicantly lower Ca concentrations in
fruit with physiological disorders. Burdon et al. (1991, 1992) measured lower
Ca concentrations (5.85 mg/100 g fresh weight (FW)) in the distal portions of
Kent fruit with soft-nose symptoms than in healthy fruit (8.02 mg/100 g
FW), and concluded that physiological disorders are associated with Ca de-
ciency, but that Ca may be only one of many causal factors and not necessar-
ily the primary cause. There have been other reports that increased Ca
concentration is associated with physiological disorders; however, these have
been attributed to mineral redistribution after tissue breakdown (Burdon et al.,
1991; Raymond et al., 1998b).
Comparing fruit Ca data from different publications is difcult, as some
quote Ca concentrations on a FW basis and others on a DW basis, and many
authors do not specify whether they measured whole fruit or fruit pulp.
None the less, evidence allows the conclusion that low Ca concentrations are
a causal factor of physiological disorders in mango fruit.
There are mixed reports of the effect of fruit Ca on postharvest fruit
breakdown in mango. Singh et al. (1987) reported that spray-to-runoff appli-
cations of calcium chloride (CaCl
2
, 36% Ca) and calcium nitrate (Ca(NO
3
)
2
,
18% Ca) at rates between 5 and 20 ml/l a week before harvest, increased shelf
life of Alphonso mangoes by 10 days. However, the authors did not discuss
whether the extended shelf life was due to delayed ripening or to reduced
postharvest disease. When Duttaray et al. (1993) attempted to repeat this
experiment to control stem-end rot caused by Diplodiia natalensis Pole Evans
using 25005000 l/l Ca as CaCl
2
and 450900 l/l Ca as Ca(NO
3
)
2
, they sig-
nicantly increased the incidence of stem-end rot. Simmons et al. (1997) used
preharvest Ca sprays and manipulation of leaf:fruit ratios to increase fruit
Ca, but were unable to signicantly reduce postharvest disease.
Calcium in leaves and fruit
Leaf concentrations of Ca increase with age, and are considered to be ade-
quate if they are between 23.5% DW in acid soils and 3.05.0% in alkaline
soils (Reuter and Robinson, 1997). Leaf Ca concentrations generally do not
correlate well with fruit Ca concentrations (Bally, 2007).
I.S.E. Bally 422
Calcium concentrations in mango fruit can vary between and within
fruit. Gunjate et al. (1979) measured a gradient in Ca concentration from the
stem end (122 mg/100 g DW) to the apex (110 mg/100 g DW) of Alphonso
fruit, and from the skin (142 mg/100 g DW) to the centre (130.4 mg/100 g DW)
and to the seed (128.4 mg/100 g DW). Gradients of Ca concentrations from the
inner-mesocarp to the outer-mesocarp, and from the stem end to the fruit apex
have been reported (Burdon et al., 1991; Tarmizi et al., 1993; Shorter and Joyce,
1998). Variation in Ca concentrations within fruit tissues may be related to the
position of the vascular tissue and the transpirational properties of the tissue
that inuence the diffusion path of Ca (Shorter and Joyce, 1998).
Concentrations of Ca on a DW basis increase rapidly during early fruit
development (cell division stage) followed by a gradual reduction later (cell
expansion stage); however, concentrations vary between reports. Gunjate
et al. (1979) measured Ca concentrations in Alphonso of 241 mg/100 g DW
at fruit set decreasing to 68 mg/100 g DW at harvest. Chattopadhyay and
Sarkar (1990) also observed a similar decline, but measured higher concen-
trations of 600700 mg/100 g DW Ca at fruit set and 350450 mg/100 g DW
Ca at harvest in four cultivars. More recently, Bally (2007) measured fruit
mesocarp Ca between 0.22% and 0.33% 30 days after fruit set and between
0.08% and 0.14% at harvest (155 days after fruit set) in Keitt. Both Gunjate
et al. (1979) and Bally (2007) also noted large uctuations in Ca concentra-
tions in the rst half of fruit development that reected imbalances in supply
and demand of developing fruit.
12.14 Boron
Boron binds as cis-diol borate complexes to mannose and certain other sug-
ars in cell wall polysaccharides and also has a role in sugar movement in the
tree. Boron also is important in nucleic acid synthesis (Salisbury and Ross,
1992), and is essential for pollen and ower development, pollen germina-
tion and pollen tube growth (Stanley and Lichtenberg, 1963; Gupta et al.,
1985) and is thus essential for mango fruit set. Boron also is important in the
synthesis of proteins that translocate sugars (Gupta et al., 1985).
Sources, uptake and translocation of B
Most B in soil occurs as axenite, tourmaline, ulexite, colemenite and kermite;
they are relatively insoluble and are released very slowly so that small
amounts are available to plants (Gupta et al., 1985). Plant-available B is mainly
associated with soil organic matter and soil solution and is dependent on soil
physical and chemical properties, with the highest concentrations found in
soils derived from marine sediments, and the lowest concentrations in
light sandy soils derived from granites (Weir and Cresswell, 1995; Peverill
et al., 1999). Low B concentrations are associated with low organic matter,
high rainfall or irrigation, high pH, high Ca and dry soils (Gupta et al., 1985).
Crop Production: Mineral Nutrition 423
Supplementary forms of B include sodium borate as borax (Na
2
B
4
O
7
, 10% B)
and as solubor (Na
2
B
8
O
13
4H
2
O, 20% B), boric acid (HBO
3-
, 18% B), calcium
borate (Ca
3
(BO
3
)
2
, 12% B) and calcium-sodium borate (CaNa
3
B
5
O
10
, 18% B).
Boron is primarily taken up by the roots as un-dissociated boric acid
(B(OH)
3
) and transported in the xylem vessels by mass ow, accumulating in
organs with the highest transpiration rates, that is leaves and growing shoots.
There is only limited redistribution of B via the phloem (Raven, 1980); how-
ever, this limited redistribution and xylem supply is usually sufcient for
normal tree health (Shelp et al., 1995).
Boron deciency and toxicity
Boron deciency is expressed in tissues that are rapidly expanding and have
low transpirational rates, for example roots, fruits and shoots (Shelp et al.,
1995). In mango, B deciency can result in poor owering, pollination and
reduced fruit set. It is expressed in growing shoots by uneven cell division,
causing leaves to grow lop-sided with a curved sickle shape and deformed
lamina and margins (Plate 78). Leaves often have shot-holes that are sur-
rounded by a light-green halo and ragged margins. Apical dominance can be
lost with swelling of the internodes. The main raceme of panicles can develop
a slight bend or kink towards the tip and in some cultivars the bark splits and
oozes black gummy sap from the cracks, known as gummosis (Plate 78)
(Nartvaranant et al., 2002). Agarwala et al. (1988) generated B deciency in
1-year-old seedlings using sand culture, and described mild deciency symp-
toms, as mild chlorosis with a marked reduction in length and width of the
leaves. In more severe cases, older leaves become chocolate-brown at the base,
spreading to the tip before becoming completely chlorotic. Stems turned black,
lost apical dominance and eventually stopped growing. Boron deciency can
be aggravated by high N status of trees (Ram et al., 1989; Raja et al., 2005).
In mango fruit, B deciency causes cracks that split open; there is brown
discoloration of the mesocarp. Lumpy, deformed fruit may also be a symp-
tom of B deciency. Meneses et al. (1994) examined normal and deformed
fruit using neutron capture radiography, and suggested that the deformities
were due to B toxicity. Boron toxicity is common in mango with typical symp-
toms appearing in leaves as dark spots on leaf margins that coalesce, eventu-
ally leading to marginal leaf necrosis in more severe cases (Plate 79). Boron
toxicity often occurs after excessive application of B fertilizers. Symptoms can
be amended through leaching of fertilizer from the root zone, raising soil pH
with applications of lime or stimulating growth through application of N;
however, these measures may have other implications for crop production.
Effect of B on crop production and fruit quality
Boron deciency symptoms of trunk gummosis in Kyo Savoy and Nam
Doc Mai in Thailand were successfully remedied by soil applications of 20 or
I.S.E. Bally 424
25 g/m borax (11% B) during the summer wet season; however, the response
time and effect on gummosis of the two cultivars differed (Nartvaranant
et al., 2002). Foliar application of B solutions at the pre-owering stage
increased yield and fruit quality in several studies. Dutta (2004) observed
that 3000 ppm boric acid is optimal for maximizing yield and quality of Him-
sagar in West Bengal, India. Coetzer et al. (1991) reported that foliar applica-
tion of B at owering raised leaf B concentrations to 60 mg/kg and increased
yields from 14 to 22 kg/tree as well as improving fruit quality. When Lora-
Meneses et al. (1992) applied boric acid to the skin of developing fruit they
found it was not signicantly translocated into the mesocarp and remained
in the surface layers of the skin around secretary glands. The response of
mango trees to soil applications of B varies between cultivars. Rossetto et al.
(2000) observed that Winter was least sensitive, whereas, Tommy Atkins,
Haden and Van Dyke in declining order were more sensitive.
12.15 Conclusion
Our understanding of mineral nutrition in mango is incomplete and lacks
detailed knowledge of the effects of many minerals, as demonstrated above.
A better understanding has generally come from observations of the effects
of minerals on fruit quality rather than yield. Studying the effects of minerals
on mango production and fruit quality is difcult because of their delayed
response to applied minerals and large tree reserves. The effects of minerals
on productivity and fruit quality can be greatly enhanced by also determin-
ing their effects on other phenological events that contribute to productivity
and fruit quality. More regular and targeted assessment of mango tree min-
eral status will facilitate improved management of mango trees. This may
require the determination of fruit mineral concentration standards for differ-
ent stages of fruit development. Development and employment of low-cost,
rapid and non-destructive techniques to measure minerals will enable mineral
uxes within trees to be monitored closely.
References
Agarwala, S.C., Nautiyal, B.D., Chatterjee, C. and Sharma, C.P. (1988) Manganese, zinc
and boron deciency in mango. Scientia Horticulturae 35, 99107.
Aitken, R.L. and Scott, B.J. (1999) Magnesium. In: Peverill, K.I., Sparrow, L.A. and Reuter,
D.J. (eds) Soil Analysis: an Interpretation Manual. Commonwealth Scientic and
Industrial Research Organization (CSIRO), Melbourne, Australia, pp. 255262.
Alvian, R.L. (1974) Mango fertilization on soils of the Maracay series. Agronomia Tropi-
cal 22, 535539.
Alvian, R.L. and Figueroa, M. (1977) Timing of nitrogen application in mangoes grown
on Maracay soil in Aragua (Argenti Venezuela). Agronomia Tropical 27, 491501.
Anonymous (1988) Sulfur. Consolidated Fertiliser Ltd (CFL), Brisbane, Australia.
Anonymous (2007) A Guide to Soil Sampling. Spectrum Analytic. Available at: http://
www.spectrumanalytic.com/support/library/ff/soil_sampling_instructons.htm (accessed
16 March 2008).
Crop Production: Mineral Nutrition 425
Bally, I.S.E. (2007) The effect of preharvest nutrition and crop load on fruit quality and
postharvest disease in mango (Mangifera indica L.). PhD dissertation, University of
Queensland, Brisbane, Australia.
Bangerth, F. (1979) Calcium-related physiological disorders of plants. Annual Review of
Phytopathology 17, 97122.
Banuelos, G.S., Bangerth, F. and Marschner, H. (1987) Relationship between polar
basipetal auxin transport and acropetal Ca
2+
transport in tomato fruit. Physiologia
Plantarum 71, 321327.
Beauls, E.R. (1973) Diagnosis and recommendation integrated system (DRIS): a gen-
eral scheme of experimentation and calibration based on principles developed
from research in plant nutrition. Soil Science Bulletin University of Natal 1, 131.
Bhandal, I.S. and Malik, C.P. (1988) Potassium estimation, uptake and its role in the
physiology and metabolism of owering plants. International Review of Cytology
110, 205245.
Bhargava, B.S. and Chadha, K.L. (1988) Leaf nutrient guide for fruit and plantation crops.
Fertiliser News 33, 2129.
Burdon, J.N., Moore, K.G. and Wainwright, H. (1991) Mineral distribution in mango
fruit susceptible to the physiological disorder soft-nose. Scientia Horticulturae 48,
329336.
Burdon, J.N., Moore, K.G. and Wainwright, H. (1992) A preliminary examination of the
physiological disorder soft-nose in mango fruit. Acta Horticulturae 296, 1522.
Catchpoole, D.W. and Bally, I.S.E. (1995) Nutrition of mango trees: a study of the rela-
tionship between applied fertiliser, leaf elemental composition, and tree perfor-
mance (owering and fruit yield). In: Proceedings of the Mango 2000 Marketing
Seminar and Production Workshop. Queensland Department of Primary Industries
and Fisheries, Townsville, Queensland, Australia, pp. 91104.
Chadha, K.L., Samra, J.S. and Thakur, R.S. (1980) Standardisation of leaf-sampling
technique for mineral composition of leaves of mango cultivar Chausa. Scientia
Horticulturae 13, 323329.
Chattopadhyay, P.K. and Sarkar, S.K. (1990) Studies on the mineral composition of
inorescence and fruits at different stages of growth in four commercial culti-
vars of mango in West Bengal. Haryana Journal of Horticultural Sciences 19,
268272.
Chaudhary, S.K. and Nauriyal, J.P. (1988) Effect of deciency of calcium, magnesium
and sulphur on the uptake of other nutrients in mango. Acta Horticulturae 231,
296300.
Clarkson, D.T. (1984) Calcium transport between tissue and its distribution in the plant.
Plant, Cell and Environment 7, 449465.
Coetzer, L.A., Robbertse, P.J. and de Wet, E. (1991) The inuence of boron applications
on fruit production and cold storage of mangoes. South African Mango Growers
Association Yearbook 11, 2931.
Cracknell Torres, A., Cid Ballarim, M.C., Socorro Monzon, A.R., Fernandez Galvan, D.,
Rosell Garcia, P. and Galan Sauco, V. (2004) Effects of nitrogen and calcium supply
on the incidence of internal fruit breakdown in Tommy Atkins mangoes (Mangifera
indica L.) grown in a soilless system. Acta Horticulturae 645, 387401.
Crane, J.H., Bally, I.S.E., Tomer, E. and Mosqueada-Vazqez, R. (1997) Crop production.
In: Litz, R.E. (ed.) The Mango: Botany, Production and Uses. CAB International,
Wallingford, UK, pp. 203256.
Daulta, B.S., Singh, H.K. and Chauhan, K.S. (1981) Effect of zinc and CCC sprays on
owering fruiting and physio-chemical composition of fruit in mango (Mangifera
indica L.) cv. Dashehari. Haryana Journal of Horticultural Science 10, 161165.
I.S.E. Bally 426
Davenport, T.L. and Nez-Elisea, R. (1997) Reproductive physiology of mango. In: Litz,
R.E. (ed.) The Mango: Botany, Production and Uses. CAB International, Wallingford,
UK, pp. 69146.
Dilly, O., Romheld, V., Marschner, H. and Chen, Y. (1997) Nutritional disorder in mango
trees on calcareous soil in Israel. Acta Horticulturae 448, 360.
Dow, A.I., Rushmore, F.A., Halvorson, A.R. and Tukey, R.B. (1991) Orchard Soil
Sampling. Washington State University. Available at: http://cru.cahe.wsu.edu/
CEPublications/eb1595/eb1595.html (accessed 16 March 2008).
Dutta, P. (2004) Effect of foliar boron application on panicle growth, fruit retention and
physio-chemical characters of mango cv. Himsagar. Indian Journal of Horticulture
61, 265266.
Duttaray, S.K., Singh, R.D.N., Kabir, J., Dhua, R.S. and Kaiser, S. (1993) Effect of pre-
harvest treatment of calcium compounds on the incidence of Diplodia stem-end rot
of mango in storage. Crop Research Hisar 63, 419422.
Fallahi, E., Conway, W.S., Hickey, K.D. and Sams, C.E. (1977) The role of calcium and
nitrogen in postharvest quality and disease resistance of apples. HortScience 32,
831835.
Ferguson, I.B. (1984) Calcium in plant senescence and fruit ripening. Plant, Cell and
Environment 7, 477489.
Gonzalez, A., Azam, G., Madddern, T. and Renfree, R. (2005) Evaluating the Use of the
Spad-502 Meter to Manage Nitrogen in Trees (Kensington Pride) in the Northern
Territory. Department of Primary Industry, Fisheries and Mines Technical Annual
Report 323. Department of Primary Industry, Fisheries and Mines, Darwin, Northern
Territory, Australia.
Gourley, C.J.P. (1999) Potassium. In: Peverill, K.I., Sparrow, L.A. and Reuter, D.J. (eds)
Soil Analysis: an Interpretation Manual. Commonwealth Scientic and Industrial
Research Organization (CSIRO), Melbourne, Australia, pp. 229245.
Gunjate, R.T., Tare, S.J., Rangawala, A.D. and Limaye, V.P. (1979) Changes in calcium
content in Alphonso mango fruits and leaves from fruit-set to harvesting. Indian
Journal of Horticulture 36, 383386.
Gupta, U.C., Jame, Y.W., Campbell, C.A., Leyshon, A.J. and Nicholaichuk, W. (1985)
Boron toxicity and deciency: a review. Canadian Journal of Soil Science 65,
318409.
Ho, L.C. and Adams, P. (1989) Calcium deciency a matter of inadequate transport to
rapidly growing organs. Plants Today 2, 202207.
Hundal, H.S., Singh, D. and Brar, J.S. (2005) Diagnosis and recommendation integrated
system for monitoring nutrient status of mango trees in submountainous area of
Punjab, India. Communications in Soil Science and Plant Analysis 36, 2085
2099.
Kadiyala, H.B. (1995) Composition and distribution of minerals in mango fruit during
ripening and storage with reference to internal breakdown, a postharvest disorder.
In: Proceedings of the International Symposium on Postharvest Science and
Technology of Horticultural Crops. China Agricultural Scientech Press, Beijing,
pp. 1017.
Kanwar, J.S., Nijjar, G.S. and Kahlon, G.S. (1987) Effect of nitrogen, phosphorus and
potassium fertiliser on the growth and productivity of Dashehari mango (Mangifera
indica). Journal of Research of the Punjab Agricultural University 24, 411422.
Kirkby, E.A. (1979) Maximising calcium uptake by plants. Communications in Soil Science
and Plant Analysis 10, 89113.
Kirkby, E.A. and Pilbeam, D.J. (1984) Calcium as a plant nutrient. Plant, Cell and Envi-
ronment 7, 397405.
Crop Production: Mineral Nutrition 427
Klumpp, A., Hintemann, T., Lima, J.S. and Kandeler, E. (2003) Bioindication of air pollution
effects near a copper smelter in Brazil using mango trees and soil microbiological
properties. Environmental Pollution 126, 313321.
Kumar, O.V. and Kumar, G. (1989) Effect of pre-harvest foliar sprays of zinc on post-
harvest changes in the quality of mango cv. Dashehari. Acta Horticulturae 231,
763770.
Kumar, S. and Nauriyal, J.P. (1977) Nutritional studies on mango tentative leaf analysis
standards. Indian Journal of Horticulture 34, 100106.
Kumar, S. and Nauriyal, J.P. (1979) Foliar sampling technique in mango. Punjab Horti-
cultural Journal 19, 1015.
Lim, T.K. and Khoo, K.C. (1985) Diseases and Disorders of Mango in Malaysia. Tropical
Press, Kuala Lumpur, Malaysia.
Littlemore, J., Winston, E.C., Howitt, C.J., Farrell, P.O. and Wiffen, D.C. (1991) Improved
methods for zinc and boron applications to mango (Mangifera indica L.) cv.
Kensington Pride in the Mareeba-Dimbulah district of North Queensland. Austra-
lian Journal of Experimental Agriculture 31, 117121.
Lora-Meneses, L.G., Jimnez-Dam, R., Gallardo-Badilla, M., Martini, F. and Theller, M.
(1992) A study of the distribution and uptake of boron in mango fruits using neutron-
capture-radiography. Comtes Rendus de lAcademie des Sciences, Paris 315, 255264.
Malo, S.E. and Campbell, C.W. (1978) Studies on mango fruit breakdown in Florida.
Proceedings of the Tropical Region of the American Society for Horticultural Science
22, 119.
Marcelle, R., Simon, P. and Lennes, G. (1990) The effects of IAA and TIBA on calcium
transport and accumulation in fruits. Acta Horticulturae 120, 193198.
Marchal, J., Goguly, T. and Didier, C. (1991) A mineral assessment of the mango tree,
variety Amelie. Fruits 46, 477487.
Marschner, H. (1995) Mineral Nutrition of Higher Plants, 2nd edn. Academic Press,
London.
McKenzie, C.B. (1994) The background skin colour of exported mango fruit in relation
to tree nitrogen status. South African Mango Growers Association Yearbook 14,
2628.
McKenzie, C.B. (1995) The effect of calcium and potassium foliar and fruit sprays on
Sensation mango leaf nutrient concentrations and fruit quality. South African
Mango Growers Association Yearbook 15, 4447.
Medeiros, A.A., Amorim, J.A.R., Silva, D.J., Dantas, J.A. and Guerra, A.G. (2004) Min-
eral composition of leaves and fruits of irrigated mango trees in Rio Grande do
norte state Brazil. Acta Horticulturae 645, 403408.
Meneses, L.G.L., Dam, R.J. and Martini, F. (1994) Neutron-capture radiography of the
distribution of boron in normal and deformed mango fruits. Journal of Trace and
Microprobe Techniques 12, 97102.
Mengel, K. and Kirkby, E.A. (1987) Principles of Plant Nutrition, 4th edn. International
Potash Institute, Worblaufen-Bern, Switzerland.
Monselise, S.P. and Goren, R. (1987) Preharvest growing conditions and postharvest
behaviour of subtropical and temperate-zone fruits. HortScience 22, 11851189.
Nartvaranant, P., Subhadrabandhu, S. and Whiley, A.W. (2002) Effect of soil boron
application on gummosis and leaf boron content of mango (Mangifera indica L.)
cvs. Khieo Sawoei and Nam Dok Mai. Acta Horticulturae 575, 875879.
Narwadkar, P.R. and Pandey, R.M. (1989) Studies on the phosphorus translocation in mango
with special reference to alternate bearing. Acta Horticulturae 231, 440447.
Nguyen, H., Hofman, P., Holmes, R., Bally, I., Stubbings, B. and McConchie, R. (2004)
Effect of nitrogen on the skin colour and other quality attributes of ripe Kensington
I.S.E. Bally 428
Pride mango (Mangifera indica L.) fruit. Journal of Horticultural Science and Bio-
technology 79, 204210.
Oosthuyse, S.A. (1993) Effect of spray application of KNO
3
, urea and growth regulators
on the yield of Tommy Atkins mango. South African Mango Growers Association
Yearbook 13, 5862.
Oosthuyse, S.A. (1997) Effect of KNO
3
sprays to owering mango trees on fruit reten-
tion, fruit size, tree yield, and fruit quality. Acta Horticulturae 455, 359366.
Oosthuyse, S.A. (2000a) Cost reduction of powdery mildew control in mango with
mono potassium phosphate. Acta Horticulturae 509, 719723.
Oosthuyse, S.A. (2000b) Variation of leaf nutrition status in relation to fruit growth in
mango. Acta Horticulturae 509, 375378.
Peverill, K.I., Sparrow, L.A. and Reuter, D.J. (eds) (1999) Soil Analysis: an Interpretation
Manual. Commonwealth Scientic and Industrial Research Organization (CSIRO),
Melbourne, Australia.
Pimplasker, M. and Bhargava, B.S. (2003) Leaf and soil nutrient norms in mango
(Mangifera indica L.) grown in tribal belt of Southern Gujarat. Journal of the Indian
Society of Soil Science 51, 268272.
Protacio, C.M. (2000) A model for potassium nitrate-induced owering in mango. Acta
Horticulturae 509, 545552.
Raghupathi, H.B., Reddy, Y.T.N., Rein, M.K. and Bhargava, B.S. (2004) Diagnosis of
nutrient imbalance in mango by DRIS and PCA approaches. Journal of Plant Nutri-
tion 27, 11311148.
Raj, G.B. and Rao, A.P. (2006) Identication of yield-limiting nutrients in mango through
DRIS indices. Communications in Soil Science and Plant Analysis 37, 17611774.
Raja, M.E., Kumar, S.C.A. and Raju, S.Y. (2005) Boron deciency in mango (Mangifera
indica L.): a cause delineation study in acidic soils of Maharashtra, India. Soil Sci-
ence and Plant Nutrition 51, 751754.
Ram, S., Bist, L.D. and Sirohi, S.C. (1989) Internal fruit necrosis of mango and its control.
Acta Horticulturae 231, 805813.
Rane, D.A., Katordia, J.S. and Kulkarni, D.N. (1976) Problem of spongy tissue develop-
ment in mango (Mangifera indica L.) cv. Alphonso, a review. Journal of Maharashtra
Agricultural University 1, 8994.
Rao, D.P. and Mukherjee, S.K. (1989) Nutrient status in leaf and soil of some cultivars of
mango in relation to yield. Acta Horticulturae 231, 286295.
Raven, J.A. (1980) Short- and long-distance transport of boric acid in plants. New Phy-
tologist 84, 231249.
Raymond, L., Schaffer, B., Brecht, J.K. and Crane, J.H. (1998a) Internal breakdown in
mango fruit: symptomology and histology of jelly seed, soft nose and stem-end cav-
ity. Postharvest Biology and Technology 13, 5970.
Raymond, L., Schaffer, B., Brecht, J.K. and Hanlon, E.A. (1998b) Internal breakdown,
mineral element concentration and weight of mango fruit. Journal of Plant Nutrition
21, 871889.
Reddy, S.E. and Majmudar, A.M. (1985) Tracking phosphorus patterns in mango
(Mangifera indica L.) and possible relations to oral induction. Nutrient Cycling in
Agroecosystems 6, 225234.
Reddy, Y.T.N., Kurian, R.M., Kohli, R.R. and Singh, G. (2000) Effect of nitrogen, phos-
phorus and potassium on growth, yield and fruit quality of Totapuri mango
(Mangifera indica). Indian Journal of Agricultural Sciences 70, 475478.
Reddy, Y.T.N., Kumar, R.M., Singh, G. and Raghupati, H.B. (2003) Long-term effects of
nitrogen on growth, leaf-nutrient status and fruit yield of Totapuri mango (Mangifera
indica). Indian Journal of Agricultural Sciences 73, 206208.
Crop Production: Mineral Nutrition 429
Reuter, D.J. and Robinson, J.B. (eds) (1997) Plant Analysis: an Interpretation Manual, 2nd
edn. Commonwealth Scientic and Industrial Research Organization (CSIRO),
Melbourne, Australia.
Reuter, D.J., Robinson, J.B., Peverill, K.I. and Price, G.H. (1986) Guidelines for collecting,
handling, and analysing plant materials. In: Reuter, D.J. and Robinson, J.B. (eds) Plant
Analysis: an Interpretation Manual. Inkata, Melbourne, Australia, pp. 2032.
Reuveni, M., Harpaz, M. and Reuveni, R. (1998) Integrated control of powdery mildew
on eld-grown mango trees by foliar sprays of mono-potassium phosphate fertilizer,
sterol inhibitor fungicides and the strobilurin Kresoxym-methyl. European Journal
of Plant Pathology 104, 853860.
Rhodes, K.J.C. (1980) The maturation and ripening of fruits. In: Thimann, K.V. (ed.) Senes-
cence in Plants. CRC Press, Boca Raton, Florida, pp. 157205.
Robinson, J.B. (1986a) Fruits, vines and nuts. In: Reuter, D.J. and Robinson, J.B. (eds) Plant
Analysis: an Interpretation Manual. Inkata, Melbourne, Australia, pp. 120147.
Robinson, J.B. (1986b) Tree mineral nutrition. Acta Horticulturae 175, 163172.
Robinson, J.B., Treeby, M.T. and Stephenson, R.A. (1997) Fruits, vines and nuts. In: Reuter,
D.J. and Robinson, J.B. (eds) Plant Analysis: an Interpretation Manual. Inkata, Mel-
bourne, Australia, pp. 349382.
Rojas, E. and Leal, F. (1997) Effects of pruning and potassium nitrate spray on oral and
vegetative bud break of mango cv. Haden. Acta Horticulturae 455, 522529.
Rossetto, C.J., Furlani, P.R., Bortoletto, N., Quaggio, J.A. and Igue, T. (2000) Differential
response of mango varieties to boron. Acta Horticulturae 509, 259264.
Saleh, M.M.S. and El-Monem, E. (2003) Improving the productivity of Fagri Kalan man-
go trees grown under sandy soil conditions using potassium, boron and sucrose as
foliar spray. Annals of Agricultural Science (Cairo) 48, 747756.
Salisbury, F.B. and Ross, C.W. (1992) Plant Physiology. Wadsworth, Belmont,
California.
Samra, J.S. and Arora, Y.K. (1997) Mineral nutrition. In: Litz, R.E. (ed.) The Mango:
Botany, Production and Uses. CAB International, Wallingford, UK, pp. 175201.
Schaffer, B. (1994) Mango disorders caused by abiotic factors. In: Ploetz, R.C.,
Zentmyer, G.A., Nishijima, W.T., Rohrbach, K.G. and Ohr, H.D. (eds) Compendium
of Tropical Fruit Diseases. American Phytopathological Society, St Paul, Minnesota,
pp. 4344.
Schaffer, B., Larson, K.D., Snyder, G.H. and Sanchez, C.A. (1988) Identication of mineral
deciencies associated with mango decline by DRIS. HortScience 23, 617619.
Schmidt, W. (1999) Mechanisms and regulation of reduction-based iron uptake in plants.
New Phytologist 141, 126.
Scholeeld, P.B., Oag, D.R. and Sedgley, M. (1986) The relationship between vegetative
and reproductive development in the mango in Northern Australia. Australian Jour-
nal of Agricultural Research 37, 425433.
Sen, P.K., Roy, P.K. and De, B.N. (1947) Hunger signs on mango. Indian Journal of Hor-
ticulture. 5, 3544.
Sergent, E. and Leal, F. (1989) Flowering induction in mango (Mangifera indica L.) with
KNO
3
. Revista de la Facultad de Agronomia, Universidad Central de Venezuela 15,
1732.
Sergent, E., Ferrari, D. and Leal, F. (1997) Effects of potassium nitrate and paclobutrazol
on owering induction and yield of mango (Mangifera indica L.) cv. Haden. Acta
Horticulturae 455, 180187.
Sergent, E., Leal, F. and Anez, M. (2000) Potassium thiosulphate, urea and potassium
nitrate applications on vegetative and oral growth in mango Haden. Acta Horti-
culturae 509, 653659.
I.S.E. Bally 430
Shear, C.B. (1975) Calcium related disorders of fruits and vegetables. HortScience 10,
316365.
Shelp, B.J., Marentes, E., Kitheka, A.M. and Vivekanandan, P. (1995) Boron mobility in
plants. Physiologia Plantarum 94, 356361.
Shinde, A.K., Dabke, D.J., Jadhav, B.B., Kandalkar, M.P. and Burondkar, M.M. (2006)
Effect of dose and source of potassium on yield and quality of Alphonso mango
(Mangifera indica). Indian Journal of Agricultural Science 76, 213217.
Shongwe, V. and Roberts-Nkrumah, L.B. (1997) Physiological and growth responses of
mango (Mangifera indica L.) to methanol and potassium nitrate application. Acta
Horticulturae 455, 6470.
Shorter, A.J. and Joyce, D.C. (1998) Effect of partial pressure inltration of calcium into
Kensington mango fruit. Australian Journal of Experimental Agriculture 38, 287294.
Shu, Z., Sheen, T.F., Lin, S.L. and Lee, K.C. (1992) Effects of microelement-containing
pesticides on nutrient concentrations of mango leaves. Acta Horticulturae 321,
553560.
Silva, D.J., Quaggio, J.A., Pinto, J.A.C., Pinto, A.C.Q. and Megalhaes, A.F.J. (2002)
Nutrio e Adubao. In: A Cultura da Mangueira. Embrapa Informao Tecnolgi-
ca, Braslia, Brazil, pp. 191221.
Simmons, S.L., Hofman, P.J., Whiley, A.W. and Hetherington, S.E. (1997) Effects of pre-
harvest calcium sprays and fertilisers, leaf:fruit ratios and water stress on mango
fruit quality. In: Coates, L.M., Hofman, P.J. and Johnson, G.I. (eds) Proceedings of
the Disease Control and Storage Life Extension in Fruit. Australian Centre for Inter-
national Agricultural Research (ACIAR), Canberra, Australia, pp. 1926.
Singh, B.P., Singh, V.B., Shrivastava, P.K. and Singh, P.R. (1973) Effects of urea applica-
tion on growth performance and mineral content of mangoes (Mangifera indica L.)
variety Langra and Chausa. Balwamt Vidyapeeth Journal of Agricultural and Scien-
tic Research 15, 5458.
Singh, M.P. and Saxena, P.K. (1994) Deciency and excess feeding signs of macro-
elements on mango (Mangifera indica L) cv. Dashehari. Recent Horticulture 1,
3035.
Singh, R.N., Singh, G., Mishra, J.S. and Rao, O.P. (1987) Studies on the effect of pre- and
post-harvest treatment of calcium nitrate and calcium chloride on the storage life of
Amrapali mango. Progressive Horticulture 19, 19.
Singh, R.R. and Rajput, C.B.S. (1977) Effects of various concentrations of zinc on vegeta-
tive growth characters, owering, fruiting and physico-chemical composition of
fruits in mango (Mangifera indica L.). Haryana Journal of Horticultural Science 6,
1014.
Smith, B.L. (1992) Time of leaf sampling and analysis norms for mangoes (cv Sensation).
South African Mango Growers Association Yearbook 12, 5456.
Smith, P.F. and Scudder, G.K. (1951) Some studies of mineral deciency symptoms in
mango. Proceedings of the Florida State Horticultural Society 64, 243248.
Stanley, R.G. and Lichtenberg, E.A. (1963) The effect of various boron compounds on in
vitro germination of pollen. Physiologia Plantarum 16, 337346.
Stassen, P.J.C., Mostert, P.G. and Smith, B.L. (1999) Mango tree nutrition. A crop per-
spective. Neltropika Bulletin 303, 4151.
Strong, W.M. and Mason, M.G. (1999) Nitrogen. In: Peverill, K.I., Sparrow, L.A. and
Reuter, D.J. (eds) Soil Analysis: an Interpretation Manual. Commonwealth Scientic
and Industrial Research Organization (CSIRO), Melbourne, Australia, pp. 171184.
Subramanyam, H., Krishnamurthy, S., Subhadra, N.V., Dalal, V.B., Randhawa, G. and
Chacko, E.K. (1971) Studies on mineral breakdown, a physiological ripening disor-
der in Alphonso mangoes (Mangifera indica L.). Tropical Sciences 13, 203211.
Crop Production: Mineral Nutrition 431
Syamal, M.M. and Mishra, K.A. (1989) Effect of NPK on growth, owering, fruiting and
quality of mango. Acta Horticulturae 231, 276281.
Tarmizi, A.S., Tengku, A.B.T.M., Pauziah, M. and Zahrah, T. (1993) Incidence of insidi-
ous fruit rot as related to mineral nutrients in Harumanis mangoes. Journal of the
Malaysian Agricultural Research and Development Institute 21, 4349.
Tiwari, J.P. and Rajput, C.B.S. (1975) Effects of urea sprays on vegetative growth and
fruit. Bangladesh Horticulture 3, 3136.
Tiwari, J.P. and Rajput, C.B.S. (1976) Signicance of nitrogen on the growth, owering
and fruiting of mango cultivars. Acta Horticulturae 57, 2936.
Torres, M.D., Farre, J.M. and Hermoso, J.M. (2004) Nitrogen and calcium fertilisation
on productivity and fruit quality of mango cv. Sensation. Acta Horticulturae 645,
394402.
Vuuren, B.P.H.J. and Stassen, P.J.C. (1997) Seasonal uptake of macro elements by young
bearing Sensation mango trees. Acta Horticulturae 455, 167174.
Wadt, P.G.S., Silva, D.J., Maia, C.E., Tome, J.B., Pinto, P.A.D. and Machado, P. (2007)
Modelling of functions in calculating DRIS indices. Pesquisa Agropecuaria Brasile-
ira 42, 5764.
Weir, R.G. and Cresswell, G.C. (1995) Plant Nutrient Disorders. 2 Tropical Fruit and Nut
Crops. Inkata, Melbourne, Australia.
Weng, F.Y. and Chuang, T.Y. (1997) Nutrient requirement of conidial germination and
characteristics of spore matrix for mango anthracnose fungus. Plant Pathology Bul-
letin 6, 1724.
Yeshitela, T., Robbertse, P.J. and Stassen, P.J.C. (2005) Potassium nitrate and urea sprays
affect owering and yields of Tommy Atkins (Mangifera indica) mango in Ethiopia.
South African Journal of Plant and Soil 22, 2832.
Young, T.W. (1957) Soft nose a physiological disorder in mango fruits. Proceedings of
the Florida State Horticultural Society 70, 280283.
Young, T.W. and Koo, R.C.J. (1969) Mineral composition of Florida mango leaves. Pro-
ceedings of the Florida State Horticultural Society 82, 324328.
Young, T.W. and Koo, R.C. (1971) Variations in mineral content of Florida mango leaves.
Journal of the Florida State Horticultural Society 84, 298303.
Young, T.W. and Koo, R.C.J. (1975) Increasing yield of Parvin and Kent mangoes on
Lakewood sands by increased nitrogen and potassium fertilisation. Proceedings of
the Florida State Horticultural Society 87, 380384.
Young, T.W. and Miner, J.T. (1961) Relationship of nitrogen and calcium to soft nose
disorder in mango fruits. Journal of the American Society for Horticultural Science
78, 201208.
Young, T.W. and Sauls, J.W. (1981) The mango industry in Florida. Florida Cooperative
Extension Service Bulletin 189, 170.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
432 (ed. R.E. Litz)
13 Crop Production: Management
J.H. Crane,
1
S. Salazar-Garca,
2
T.-S. Lin,
3
A.C. de Queiroz Pinto
4
and Z.-H. Sh
5
1
University of Florida, Florida, USA
2
Instituto Nacional de Investigaciones Forestales, Agrcolas y Pecuarias, Santiago
Ixcuintla, Mexico
3
National Taiwan University, Taipei, Taiwan
4
Private Consultant on Tropical Fruits, Brasilia, Brazil
5
Meiho Institute of Technology, Pingtung, Taiwan
13.1 Introduction 432
13.2 Production Areas and Yields 434
13.3 Climate of Production Areas 436
13.4 Soils and Soil Preparation 440
13.5 Plant Propagation and Rootstocks 443
13.6 Major Cultivars 445
13.7 Plant Spacing 449
13.8 Fertilizer Practices 452
13.9 Irrigation Practices 460
13.10 Vegetative Growth and Reproduction Manipulation 463
13.11 Environmental Stress Management 468
13.12 Harvesting Practices 470
13.13 Conclusions 472
13.1 Introduction
Mangoes have been in cultivation for several thousand years (Mukherjee,
1953, 1972; Kostermans and Bompard, 1993), and crop production practices
have continually improved (Singh, 1960, 1978; Crane et al., 1997). A detailed
understanding of mango plant physiology and behaviour in relation to cli-
matic and edaphic conditions, genetics and cultural practices dates back to
the late 1950s (Mukherjee, 1953; Chadha and Pal, 1986; Chacko, 1989; Whiley,
1993; Schaffer et al., 1994; Kulkarni, 2004; Davenport, 2006). Recent reviews
(Cull, 1991; Whiley, 1993; Schaffer et al., 1994; Davenport and Nez-Elisea,
1997; Kulkarni, 2004; Davenport, 2006) on mango crop management and
physiology illustrate that the best prospect for improving mango production
must involve a holistic approach to cultural practices that considers the spe-
cic climatic and edaphic environment, cultivar and tree phenology for a
Crop Production: Management 433
given production area. Most importantly, the development of a phenological
approach to understanding and managing mango orchards is very promis-
ing from a practical standpoint (Cull, 1991). The phenological approach to
crop management is based on monitoring environmental conditions and
tree-growth stages and conditions, and manipulating the tree through cul-
tural practices for optimum production (Fig. 13.1). Published research can be
used to develop a holistic crop management approach.
Mango production in India has been reviewed extensively (Singh 1960,
1978; Majumder and Sharma, 1990; Kostermans and Bompard, 1993; Negi,
2000). The purpose of this chapter is to discuss current concepts, strategies
and innovations for mango culture and to describe current production prac-
tices in four commercial mango-producing areas: Brazil, Mexico, Taiwan and
the USA. Crane et al. (1997) has described the cultural practices in Australia,
Israel, Mexico and the USA.
The Brazilian and Mexican mango industries now account for about 62%
and 8% of exports to the USA, respectively (Perez and Pollack, 2007). Improve-
ments in cultural practices in both countries have increased production ef-
ciency and exports. Taiwans mango industry, although relatively small, is
innovative and provides top-quality mangoes for the local and Asian mar-
kets. Mango production in the USA is small, and production is geared
towards national speciality markets.
Apr Mar Feb Jan Dec Nov Oct Sept Aug July Jun May
Oct Sep Aug Jul Jun May Apr Mar Feb Jan Dec Nov
n
s
A
m
o
u
n
t

o
f

d
e
v
e
l
o
p
m
e
n
t
Flower bud
development
Flowering and
fruit set
Fruit
development
Premature
fruit drop
Harvest
Root flush
Unwanted
vegetative
flush
Postharvest
vegetative flush
Fig.13.1. Theoretical mango phenological cycle (Source: after Cull, 1991). s, southern
hemisphere; n, northern hemisphere.
J.H. Crane et al. 434
13.2 Production Areas and Yields
The mango industry of Brazil is extensive due to favourable soil and climatic
conditions. Bahia and So Paulo are the most important production areas of
Brazil with 34,600 and 28,800 ha, respectively, of the estimated 70,000 ha of
mango production in 2001 (Table 13.1). The north and south of Brazil account
for only 6.0% of the total mango-production area (Souza et al., 2002). The
most recent mango census (2005 data) estimated mango production in Brazil
to be c.970,700 t (Gazeta, 2006; Agrianual, 2007), which is a slight increase
since 2001. Bahia and So Paulo also have the largest production, respec-
tively (Table 13.1). Brazil exports 133,300 t of mangoes, 13.7% of the total
mango production. Although pest and disease problems and the highly com-
petitive internal and external markets have limited the expansion of the
mango industry in some Brazilian regions, important technologies have been
Table 13.1. Regions of mango production in Brazil, Mexico, Taiwan and the USA
(Source: see table footnotes).
Country State Region Hectares Production (t)
Brazil
a
Bahia North-east 19,000 306,000
Pernambuco 8,000 146,000
So Paulo South-east 18,000 245,000
Minas Gerais 7,000 130,000
Miscellaneous Other regions 18,000 143,000
Mexico
b
Sinaloa Nayarit Northern and Central
Pacic region
106,740 1,138,361
Jalisco Colima
Michoacn
Guerrero
Oaxaca Southern Pacic region 42,180 367,257
Chiapas
Tamaulipas Gulf of Mexico region 28,778 191,996
Veracruz
Compeche
Miscellaneous Other states 7,827 37,151
Taiwan
c
Tainan Central-South 18,200 191,332
Pingtung South
Miscellaneous Other prefectures
USA
d
Florida Southern region 324 5,986
Puerto Rico South, South-West Coast 1,079 14,206
California Coachella Valley 105 1,905
Hawaii Islands of Hawaii, Oahu,
Maui, Kauai
142 317
a
Source of data on Brazil: Agrianual (2006, 2007); Anonymous (2006).
b
Source of data on Mexico: SIIAP (2007).
c
Source of data on Taiwan: Agriculture and Forestry Department (1996).
d
Source of data on USA: Linden (2006); Anonymous (2007); J.H. Crane, personal communication.
Crop Production: Management 435
developed to improve and support the mango industry (Santos Filho et al.,
2002). This has resulted in an increase of 58% in mango exports from 1998 to
2005 (Souza et al., 2002; Agrianual, 2007).
There are c.181,525 ha of mangoes in 23 of Mexicos 32 states (SIIAP,
2007); there are four large mango-producing regions (Table 13.1), ranging
from 1433N to 27N latitude. They are distinguished by differences in cli-
mate, soils and cultivars. There are 106,740 ha of mango in the Northern and
Central Pacic region; the major cultivars are Tommy Atkins, Ataulfo,
Kent, Haden and Keitt. The Southern Pacic region has 42,180 ha and
produces Manila (Carabao), Ataulfo, Tommy Atkins, Haden and
Kent. The leading cultivars of the Gulf of Mexico region (28,778 ha) are
Manila followed by Tommy Atkins. Mango production in Mexico was
about 1.7 million t in 2006 and the states of Sinaloa, Guerrero, Nayarit, Oax-
aca, Chiapas, Veracruz and Michoacn accounted for 92% of the crop (SIIAP,
2007). All these states except Veracruz (which abuts the Gulf of Mexico) are
situated along the Pacic coastal and inland areas.
In 2006 Mexico exported 196,120 t to the USA valued at >US$132 million
(EMEX, 2007). Mexico supplies 87.5% of the mangoes consumed in the USA
and mango exports in 2006 were 17.9% higher than in the 2005. The major
export cultivars are Tommy Atkins (33%), Kent (23%), Ataulfo (19%), Keitt
(14%) and Haden (11%). None the less, most mango production (c.85%) in
Mexico is for the domestic market. The most important mango warehouses
and distributors are in Mexico City, Guadalajara and Monterrey.
The 7-month mango season in Mexico is due to differences in latitude,
phenological cycles, rainfall and soil moisture patterns and the use of growth
regulators. Chiapas, along the South Pacic coast, is usually the rst to har-
vest mangoes (Ataulfo). Full bloom of Ataulfo occurs from mid-November
to mid-February and fruit is harvested from the end of January to the end of
May. Mango harvesting continues northwards from Chiapas to the states of
Oaxaca, Guerrero, Michoacn, Colima, Jalisco, Nayarit, Sinaloa and Sonora.
Sinaloa is a major mango producer with the latest harvest season for Manila
(July), Haden (July), Tommy Atkins (July), Kent (JulyAugust) and Keitt
(AugustSeptember). The Manila mango is produced mainly in Veracruz
and Guerrero. Full bloom occurs in JanuaryFebruary and fruit is harvested
in MayJune.
The production area in Taiwan increased from 17,000 ha in 1987 to 21,000
ha in 1992 and stabilized at c.20,000 ha for c.10 years when the yield was
17 kg/tree; however, as production has increased to c.27 kg/tree, the plant-
ing area has decreased to c.18,200 ha in 2006 (Table 13.1). There are six
important mango-growing prefectures in Taiwan: Nantou, Taichung, Chiayi,
Tainan, Kaohsiung and Pingtung from north to south in western and eastern
Taiwan. Tainan and Pingtung are the most important production areas, com-
prising about 90% of the total harvested area.
Mangoes are grown commercially in four areas of the USA: Florida
(2528N), Puerto Rico (18185N), Hawaii (2230N) and California
(33335N) (Table 13.1). Puerto Rico has the largest area (1079 ha) (Alvarado-
Ortiz and Acin, 2004), followed by Florida (324 ha) (FASS, 2003; Anonymous,
J.H. Crane et al. 436
2007; J.H. Crane, personal communication), Hawaii (142 ha) (NASS-Hawaii,
2006; Anonymous, 2007) and California (105 ha) (Linden, 2006; Anonymous,
2007). Mango production in the USA is estimated to be only 22,414 t with
Puerto Rico accounting for >63% of the production (Alvarado-Ortiz and
Acin, 2004; Linden, 2006; Anonymous, 2007). The four areas have different
climates which present different production opportunities and challenges.
The main production areas of Florida are along the south-east coast in
Miami-Dade County (2527N), the east (Palm Beach, Broward and Indian
River Counties) and west (Lee County) coasts. The season in Florida is from
May through to September. Puerto Rico is a Caribbean island (18N, 67W)
(Espenshade, 1992; Bahr and Johnston, 1993c). Most of the commercial pro-
duction is near Ponce on the south coast and Mayaguez in the south-west
(23.9 ha). The main mango season in Puerto Rico is May through to Septem-
ber, although some mangoes are harvested as late as mid-November. In
Hawaii (1922N), production occurs on the islands of Hawaii, Oahu, Maui
and Kauai. The season in Hawaii is from May to August, although some pro-
duction occurs year-round. The California industry is on the western side of
the Salton Sea in the Coachella Valley of southern California (33N) (Scott,
1990). Mango production is from mid-August to October (Linden, 2006).
13.3 Climate of Production Areas
Mangoes are mainly cultivated in the tropics (25N, 25S) and subtropics
(35N, 35S), although limited production also occurs in warm temperate/
subtropical, i.e. Mediterranean-type areas, for example Israel, southern Spain
and the Canary Islands and California. The ideal areas for mango production
have a cool and/or dry period prior to owering followed by moderate soil
moisture and moderately hot temperatures (3033C) (Chacko, 1986). The
diversity of climates and soils in mango-production areas reects the adapt-
ability of the species and improvements in cultural practices.
Temperature and availability of water are the most signicant environ-
mental factors that inuence commercial mango production by affecting the
frequency, intensity, duration and time of vegetative growth and owering
(Chacko, 1986, 1989; Whiley, 1993; Schaffer et al., 1994; Nez-Elisea and
Davenport, 1995; Davenport, 2006) and disease incidence (Johnson et al.,
1989; Ploetz, 1994, 2003; Ploetz et al., 1994). Temperatures <15C and >30C
inhibit pollen tube germination, thereby impeding fertilization and resulting
in embryo abortion and fruit abscission.
Mango production in Brazil extends from 3N, close to the Amazon
region, where a humid, hot tropical climate predominates almost year-round
to approximately 25S in Paran, which has a subtropical climate. Between
these two extremes, there are various soil and climatic conditions where
mango trees are grown intensively. The semi-arid tropical climate of the
north-east region of Petrolina in Pernambuco (2113N and 4750W) is more
appropriate for high-quality mango production than Votuporanga in So
Paulo (2113S and 4750W), which has a subtropical climate (Table 13.2).
C
r
o
p

P
r
o
d
u
c
t
i
o
n
:

M
a
n
a
g
e
m
e
n
t
4
3
7
Table 13.2. Climatic parameters for Brazil, Mexico, Taiwan and the USA.
Country State Region Climatic category
Temperature (C)
a
Annual mean rainfall (mm)
a
Annual Summer Winter Annual Season
Yearly
mean Mean Mean
Yearly
mean Wet Dry
Brazil
b
Par North region Tropical hot humid 26.0 31.5 22.0 2893 436.2 111.8
Gois Central region Savannah dry tropics 29.8 31.9 17.7 1576 270.3 6.2
Bahia North-east region Tropical semi-arid 27.9 33.8 22.0 545 139.6 1.7
Pernambuco Tropical semi-arid 26.2 32.0 20.5 570 136.2 4.8
Piau Tropical hot subhumid 27.5 32.9 22.1 1449 286.3 11.6
So Paulo South-east region Subtropical cool 21.3 25.1 17.6 1623 127.1 40.0
Minas Gerais Tropical mesothermic 26.5 35.0 18.0 840 105.2 6.0
Paran South region Subtropical very cool 17.6 22.7 12.4 1408 165.0 74.5
Mexico
c
Sinaloa Northern and
Central Pacic
region
Warm subhumid
tropics
24.7 27.5 (2238) 22.0 (1329) 922 799 123
Nayarit Warm subhumid tropics 24.7 26.3 (2438) 23.1 (1330) 1396 1346 50
Colima Warm semi-arid tropics 26.5 27.6 (2538) 25.4 (1430) 660 605 55
Michoacn Very warm semi-arid
tropics
28.3 29.6 (2840) 27.0 (1832) 709 601 108
Guerrero Warm subhumid tropics 28.0 29.0 (2840) 26.9 (1832) 1011 910 101
Oaxaca Southern Pacic
region
Isothermal warm
humid tropics
25.9 26.9 (2635) 25.0 (1828) 1779 1731 48
Chiapas 24.7 24.9 (2635) 24.4 (1828) 3732 3661 71
Central
Veracruz
Gulf of Mexico
region
Isothermal warm
subhumid tropics
24.8 26.7 (2638) 22.9 (1430) 1026 898 128
Southern
Veracruz
Isothermal warm
humid tropics
25.5 27.6 (2638) 23.4 (1430) 2093 1807 286
Taiwan
d
Tainan Central-South Subtropical 23.6 26.8 (23.928.3) 19.1 (17.120.4) 1733 264 (71415) 25 (1541)
Pingtung South Subtropical 24.7 26.8 (24.927.9) 21.8 (20.422.7) 2158 329 (170511) 30 (1947)
Average Whole island Subtropical 25.0 25.8 (23.127.5) 19.1 (17.321.6) 1885 273 (92392) 42 (2460)
(Continued)

J
.
H
.

C
r
a
n
e

e
t

a
l
.
4
3
8
Table 13.2. Continued
Country State Region Climatic category
Temperature (C)
a
Annual mean rainfall (mm)
a
Annual Summer Winter Annual Season
Yearly
mean Mean Mean
Yearly
mean Wet Dry
USA
e
Florida Southern coasts Marine subtropics
east coast
23.2 26.8 (2627) 18.9 (1819) 1643 1313 333
Marine subtropics
west coast
23.3 27.1 (2428) 18.0 (1718) 1354 1064 290
Puerto Rico South Coastal area Marine tropical 26.3 27.7 (2332) 24.8 (1930) 904 632 272
California Coachella Valley Dry warm subtropical 22.5 31.8 (3033) 13.3 (1115) 79.7 55.1 24.6
Hawaii Island of Hawaii Marine tropical 23.3 24.2 (1633) 22.5 (1333) 3279 1926 1353
Island of Oahu Marine tropical 25.1 30.1 (1834) 24.0 (1332) 559 414 145
Island of Maui Marine tropical 24.2 25.6 (1535) 22.9 (1132) 531 442 89
Island of Kauai Marine tropical 24.2 25.6 (1732) 23.1 (1231) 1091 698 393
a
Numbers in parentheses are ranges.
b
Brazil: rainfall gures for the wet and dry season are per month.
c
Source of data on Mexico: Garca (1973).
d
Taiwan: temperature ranges are averages for lowest and highest summer (AprilOctober) and winter (NovemberMarch) months and the yearly mean;
precipitation is averages for wettest (MayOctober in Pintung; AprilSeptember in Tainan and island average) and driest (OctoberMarch) months and yearly
annual. Available at http://www.cwb.gov.tw/
e
Source of data on USA: Butson and Prine (1968); Getz (1979); E.E. Toro, personal communication; C.L. Chia, personal communication; Quayle et al. (1995);
Garczynski (1995); SERCC (2007).
Crop Production: Management 439
Some cultivars, for example Haden in north-eastern Brazil, have poor fruit
set when temperatures are >35C.
Solar radiation is very important for fruit development, since its dura-
tion and intensity are directly related to photosynthesis and carbohydrate
production (Mukherjee, 1953). Light incidence depends on the season (Allen
et al., 1998). According to Lima Filho (2000), mango production in the semi-
arid region of Brazil is where maximum solar radiation occurs in summer
(October southern hemisphere; 528 cal/cm
2
/day) and the minimum in win-
ter (June southern hemisphere; 363 cal/cm
2
/day), which corresponds to the
owering and fruit development stages, respectively.
Mango production in Mexico is in the tropics. Most rainfall occurs dur-
ing the summer and may be accompanied by hurricanes (Table 13.2). Climate
in the Northern and Central Pacic region (1727N) ranges from warm sub-
humid to warm/very warm semi-arid tropics with a 7-month dry season
(Garca, 1973). In Colima and Michoacn, mangoes are irrigated due to low
annual precipitation (semi-arid condition with a 6- to 8-month dry season;
the mean annual temperature is 26.528.3C). The Southern Pacic and Gulf
of Mexico coastal regions have isothermal (< 5C monthly temperature oscil-
lation) warm, subhumid to humid climates with a 68-month dry season
(Garca, 1973).
The Gulf of Mexico coastal region is affected by winds from the north
(1128 m/s) from October to April. These winds cause direct damage, such as
limbs breaking and ower and fruit drop, and also increase plant respiration
and transpiration rates that stress the trees. In the Pacic coastal regions hur-
ricanes usually occur from August to October. Recent, unusually warm win-
ters have resulted in undesirable late autumn or winter vegetative ushes and
poor owering. Normal bloom occurs in late January and February; how-
ever, late autumn or winter shoots delay anthesis during hot spring tempera-
tures (April to early May) which result either in low fruit set or parthenocarpic
fruit.
The primary production areas in Taiwan are from 22N (Fungshan, Ping-
tung) to 24N (Nanto and Taichung prefectures). The average temperature of
these areas is c.23C, with a mean of 20C during owering (December
March) and c.18.6C during development (AprilAugust) (Table 13.2). Flower
induction in mango is not a problem in Taiwan because of its subtropical
climate; however, fruit set can be affected by low temperature, rainfall, etc. A
monsoon-type climate prevails in southern Taiwan with precipitation
occurring mostly from May through to September. There is little rainfall
(usually <50 mm) during ower bud formation and owering (autumn-winter)
(Table 13.2).
The production areas of the USA have different climates. South Florida
has a marine subtropical climate (Table 13.2). South-east Florida has a mean
annual temperature of 23C, a mean annual rainfall of 1643 mm and 62%
relative humidity (RH) (Butson and Prine, 1968; Getz, 1979; Barrick and
Black, 1980). Constant ocean-spawned winds of 49 km/h buffet the region
from February through to October. The wet season (two-thirds annual rain-
fall) occurs during the late spring into autumn (MayOctober, northern
J.H. Crane et al. 440
hemisphere) and the dry season occurs during winter and early spring
(NovemberApril). Lowest temperatures (-4 to -6C) occur from December
through to February with a 70% probability of 0C at least once each year
(Bradley, 1975).
The Puerto Rican industry is mostly along the south and south-west
coast at elevations between sea level and 50 m (Table 13.2) (Aponte-Morn
et al., 1977). The La Cordillera central mountain range divides the island from
north to south and has a major impact on the islands climate, with annual
rainfall of 1550 mm in the north and 904 mm in the south (Espenshade, 1992;
SERCC, 2007). In the main production areas, the dry season is from Decem-
ber to May, although July and August are also dry (Aponte-Morn et al.,
1977). Mean maximum and minimum temperatures vary with altitude; how-
ever, along the coastal production areas, 2429C is normal (Espenshade,
1992; SERCC, 2007). The lowest temperatures (1820C) occur along the coast
during January and February.
The Hawaii mango industry is located mostly on the leeward coasts of
Oahu, Hawaii and Maui islands; however, limited production occurs on the
windward side of Kauai Island (Table 13.2). Each commercial planting has a
distinct climatological niche that varies with altitude and location with
respect to mountains and predominant north-east trade winds. Annual rain-
fall is 531 mm on Maui, 559 mm on Oahu and 3279 mm near Hilo (east coast
of Hawaii) (Table 13.2; C.L. Chia, personal communication). Temperatures,
like rainfall, vary with elevation and location but generally range between 21
and 27C (Bahr and Johnston, 1993b). Lower (13C) and higher temperatures
(32C) occur occasionally.
The Coachela Valley of California is in the Salton Trough Desert area
(Bahr and Johnston, 1993a) (Table 13.2). The valley ranges from 80 m below
sea level to 488 m above sea level (Espenshade, 1992; Aslan et al., 1993) and
the climate is arid. Temperatures in the valley vary considerably depending
upon season, elevation and exposure. Some data reported herein represent
the average for the area and may not accurately reect the particular micro-
climate of the mango orchards in the valley (Garczynski, 1995). The mean
annual temperature is 22.5C; the average summer temperature is 31.8C,
and the average winter temperature is 13.3C (Aslan et al., 1991; Garczynski,
1995). Winter temperatures range from 22.1C to 4.5C (Garczynski, 1995);
however, Schacht (1992) reported a low of -4.4C for 2 h. Summer tempera-
ture extremes range from a low of 11C to 50C (Garczynski, 1995). The aver-
age annual rainfall is 79.8 mm with two-thirds of this occurring during the
winter (DecemberMarch) (Aslan et al., 1991; Garczynski, 1995). The valley is
buffeted by wind and sandstorms during late spring, and these can cause
severe damage (Schacht, 1992; Aslan et al., 1993).
13.4 Soils and Soil Preparation
Mangoes tolerate many soil types (Majumder and Sharma, 1990; Crane and
Campbell, 1991; Kostermans and Bompard, 1993). Trees grow most vigorously
Crop Production: Management 441
in deep, fertile, moderately acid to neutral pH, loam-type soils. They tolerate
infertile sands, volcanic ash and limestone-based soils, excessively drained
and or periodically ooded soils and soils with acid (pH 4.57) to alkaline pH
(pH 78.5). Mangoes are sensitive to saline and sodic soil conditions and
proper irrigation practices and the use of salt-tolerant rootstocks is imperative
for successful crop production in some areas (Schaffer et al., 1994).
Land preparation includes clearing virgin or existing orchard trees, disk-
ing, destruction of subsurface hardpans (slip ploughing), mixing of upper
and lower soil proles, crushing supercial bedrock and mixing rock and soil
layers, formation of land contours to facilitate drainage and/or ood irriga-
tion, bedding and amending soils with organic matter and inorganic chemi-
cals, for example hydrated lime (Ca(OH)
2
), dolomite and preplant fertilizers.
Mango is grown on a wide range of soil types, from latossols with a high
percentage of sand in the north-east of Brazil to loamy oxysols in the south-
east. Some areas in the north-east have shallow soils that need improved
drainage. Soils of the central region are chemically poor and acid (pH 3.74.7)
and require lime and/or gypsum application prior to orchard establishment.
Soil analysis is used for determining the suitability of soil for production and
potential fruit quality, especially in areas cultivated for export fruit. Typically
soil analysis is conducted prior to land clearing and ploughing to determine
nutrient levels and the need for lime and/or gypsum application. Soil sam-
ples are taken from 020 cm and 2040 cm depth of the soil prole. Soil sam-
ples are usually taken at a 030 cm and 3060 cm soil depth for mature and
established orchards. Ploughing, harrowing and lime and/or gypsum incor-
poration are recommended at 30 cm depth and at least 30 days prior to rainy
weather (Pinto and Ramos, 1998). Liming is very important for acid soils
(<pH 4.0) in the central region of Brazil in order to increase soil pH to 6.06.5.
It also improves the soil base saturation to 6070% and results in better
soil conditions for growth and production (Pinto, 2000). The amount of lime
to be applied is based on a basis saturation equation (where hydrogen (H),
aluminium (Al), calcium (Ca), magnesium (Mg) and potassium (K)):
Lime rate (t/ha): (T 0.5) S where T = (H + Al) + S and S = Ca + Mg + K
Gypsum applications are recommended for acid subsoils with >20% Al
saturation and <0.5 cmol/dm
3
Ca at soil depth of 60 cm (Andrade, 2004).
This has been shown to signicantly reduce fruit physiological disorders,
such as soft nose.
Production areas of Mexico vary in soil characteristics, and include at
coastal areas and steep mountain slopes from sea level to >600 m above sea
level. Orchards on sloped land commonly utilize individual tree terraces.
Soil depth ranges from 30 cm to >3 m (alluvial soils). Shallow and eroded
soils are common on hilly terrain. The presence of medium to large (1050 kg)
boulders can impede the use of machinery but they reduce erosion and
increase soil-water storage capacity. In general, poor soil drainage is not a
problem; however, areas with clay soils have drainage problems during peri-
ods of heavy rain, especially if there is a shallow water table. A range of soil
types is used for mango production in Mexico. In the Northern Pacic region,
J.H. Crane et al. 442
mangoes are planted in cambisols (pH 6.07.0), luvisols (pH 6.57.5) and
feozem (pH 5.06.0) soils with loamy textures (Anonymous, 1970, 1982).
These soils are well drained, with 23% organic matter, moderate to high
water holding capacity and cation exchange capacity (CEC) of 1020+
meq/100 g soil. In the Central Pacic region mangoes are planted on cambi-
sols, feozem and regosols with light textures (Anonymous, 1970, 1982). The
regosols (pH 6.57.5) are well drained, with <2% organic matter content, low
water holding capacity and CEC <10 meq/100 g soil. Soils planted to man-
goes in the Southern Pacic region include nitosols, luvisols and feozem
(Anonymous, 1970, 1982). The nitosols (pH 5.06.0) are well drained, with
>3% organic matter content, low to moderate water holding capacity and
low CEC (<1020 meq/100 g soil). Soils in the Gulf of Mexico region include
uvisols, cambisols and vertisols of loamy and clayey textures (Anonymous,
1970, 1982). The uvisols and vertisols are moderately and slowly drained
soils, respectively. Soil pH for uvisols ranges from 5.5 to 7.5 with <2%
organic matter content. Soil pH for the vertisols is alkaline (7.58.0) and the
organic matter content is moderate (2.13%). The water holding capacity and
CEC is low to moderate for the uvisols (<1020 meq/100 g soil) and high
for the vertisols (1020+ meq/100 g soil).
Soil tillage is used in at lands before planting. Dimensions of planting
holes range from 4060 cm depth and 3050 cm diameter in light-textured
soils. Planting holes can be larger in heavy-textured or stony soils. On steep
hills only planting holes are made and individual terraces are built up to
hold soil, water, organic matter, fertilizers or soil amendments. Preplant soil
analyses are rarely taken; however, chemical or organic fertilizers are com-
monly applied at planting time. With rapid expansion of the mango industry,
orchards have been planted in shallow soils (0.751.2 m depth) underlain
with a hardpan. These soils are poorly drained and root growth and exten-
sion are limited. Trees growing in such soils are prone to drought stress dur-
ing dry periods, ooding stress after rains and nutritional deciencies. The
weakened trees appear to be more susceptible to pathogens, for example
Botryodiplodia theobromae, which causes cankers or stem dieback and eventu-
ally tree death (Ponce-Gonzlez and Salazar-Garca, 1992). Cultural practices
are not available to ameliorate these problems and planting in such soils is
not recommended.
Mangoes are grown in Taiwan on sandy loam, loamy sands, clay and
coarse sandy soils. Soil pH ranges from 5.0 to >7.8. Most trees are grown on
sloping land. Silt clay loam with pH 5.06.0 is mostly found in the Pingtung
area, while the Tainan area has clay soils with pH 7.37.8. Acid soils are
amended with Ca(OH)
2
or dolomite and alkaline soils are amended with
sulfur (S) or acid-based fertilizers.
The topography in the mango-production areas of Florida is at and
ranges from sea level to c.6.1 m above sea level. Soils in Florida include vari-
ous sands, muck and oolitic limestone. In the main production area (Miami-
Dade County), mangoes are planted in extensively scaried (crushed) and
trenched oolitic limestone rock (Krome and Chekika very gravelly loam)
(Colburn and Goldweber, 1961; Noble et al., 1996). The limestone soil is very
Crop Production: Management 443
permeable (1.55.1 cm/h), with high pH (7.48.4), low organic matter con-
tent (310%), low water holding capacity (0.20.3 cm/cm of soil) and low
CEC (16.037.0 meq/100 g soil) (Calhoun et al., 1974; Anonymous, 1989). In
areas subject to ooding, crushed rock is formed into beds (0.61.0 m high
and 1.01.5 m wide) before planting. The sandy soils in other Florida produc-
tion areas are poorly to well drained with or without an undulation hard-
pan 0.53.0 m down in the soil prole (Henderson et al., 1984). The highly
organic muck soils in Palm Beach County are poorly drained and underlain
by dense limestone bedrock. Beds of varying dimensions are made in the
sandy and muck areas to increase the proportion of the root system above
ood levels. The sand and muck soils are characterized by an acid to alka-
line pH (3.68.4) and low cation exchange capacities (Carlisle et al., 1978;
Henderson et al., 1984).
In Puerto Rico, mangoes are grown on at and gently sloping land con-
sisting of alluvial fans and terraces level with or slightly above the ood
plain (Bierbolini et al., 1979). Some orchards in western Puerto Rico are
located on moderately steep to very steep slopes (1260% slope) and rounded
hill tops that are somewhat eroded and supercial (Bierbolini et al., 1975).
Soils in southern Puerto Rico are very deep, moderately well to well drained
and consist of ne-textured sediment of sandy and clayey loam over gravelly
ne-textured sediment. The pH ranges from moderately acid (pH 6) to alka-
line (pH 8 or above). Soils have good to very good native fertility and good
water holding capacity. Soils in the west are well drained and moderately
acid (pH 5.06.0).
Mango production in Hawaii occurs on volcanic ash soils varying from
recent to highly weathered (R. Yost, personal communication). They tend to
be well drained; some soils must be slip ploughed to break up the hardpans
in preparation for planting. The pH ranges from 5 to 8 and the fertility varies
substantially. Mangoes in California are on lacustrine deposits consisting of
ne-textured sediments that are highly stratied sandy loam soils with clay
lenses (S. Aslan, personal communication). The soil is alkaline (pH 78.4) and
calcareous (Aslan et al., 1991). Land is slip ploughed to 1.51.8 m depth to
break up and mix the stratied soil layers.
13.5 Plant Propagation and Rootstocks
Mango production areas rely on traditional propagation, i.e. seedage, graft-
ing and budding. Marcottage (air layering) methods have been tried for
many years (Rajan and Ram, 1988; Majumder and Sharma, 1990) and although
recently improved (Nez-Elisea et al., 1989, 1991, 1992; Lambe et al., 1991;
Nez-Elisea and Davenport, 1995), it is still not used commercially. In vitro
techniques hold great promise for rootstock propagation and cultivar
improvement, and are currently being perfected and investigated (Litz, 1984,
1986, 2005; Litz et al., 1984; see Litz et al., Chapter 18, this volume).
The monoembryonic Florida cultivars in Brazil are vegetatively propa-
gated by grafting and budding on polyembryonic rootstocks. Rootstock and
J.H. Crane et al. 444
grafted mangoes are established in nurseries by sowing seeds in a soil sub-
strate consisting of a mixture of organic matter and semi-sterilized soil plus
3 kg of superphosphate (3Ca(H
2
PO
4
)
2
) and 0.5 kg of potassium nitrate (KNO
3
)
for each m
3
of mixture (Pinto et al., 1981; Castro Neto et al., 2002). In the south-
east, the indirect seeding technique is used: seed is sown in the soil to germi-
nate and then transplanted to small softwood containers or jacs. Subsequently,
seedlings are patch budded. In the central and north-east regions, seedlings
are grown in substrate in plastic bags and subsequently whip or veneer grafted.
The endocarp is removed prior to sowing to accelerate germination and
improve uniformity of rootstock growth (Pinto and Gen, 1981).
The most important characteristics for mango rootstocks are vigour, high
yield potential, compatibility with the scion, environmental adaptability and
resistance to pests and diseases (Rosetto et al., 1996; Nascimento et al., 2002).
Polyembryonic cultivars, that is Espada and Fiapo in the north-east and
Rosinha and Coquinho in the south-east, are the most important root-
stocks. The Agencia Paulista de Tecnologia Agropecuria (APTA) has
developed root-rot-resistant rootstocks, including IAC-101, IAC-102 and
IAC-106; however, they are not widely used.
There are no specic mango rootstocks in Mexico. Before the mid-20th
century, seedling plants were used; however, uniform plantings did exist
because polyembryonic Manila was grown in the Gulf of Mexico region.
Elsewhere, there were monoembryonic trees, resulting in a heterogeneous
mix of trees. Since the late 1950s and early 1960s, polyembryonic criollo root-
stocks have been used for rootstocks; however, due to their vigour, mono-
embryonic rootstocks are still in use in some areas. Commercial production is
done in partially shaded nurseries. A seedbed is prepared using sterilized
substrate (loamy soil, composted organic matter or a mixture). The brous
seedcoat is removed to expose the smooth, bean-shaped seed, which is soaked
for about 1030 min in 50 mg/l gibberellic acid (GA
3
) to reduce germination
time. Seeds are placed on the seedbed (usually in MayJuly) with their hump
up and watered every other day. After 35 weeks, vigorous and healthy seed-
lings are transferred to black plastic bags 20 cm wide 40 cm long lled with
a sterilized mixture of sandy loam soil and organic matter (3:1). Rootstocks
are veneer grafted when they are 46 months old. Grafted plants are ready for
planting by MayJuly of the next year, during the rainy summer season.
Some experienced mango growers initially plant seedling trees in the
orchard to ensure a vigorous tree. Grafting is then performed 36 months
after planting. In some cases a low vigour interstock like Esmeralda is top-
worked onto the seedling rootstock and then grafted to the desired scion.
The use of Esmeralda interstocks has reduced tree canopy size by 28% and
33% for Manila and Ataulfo, respectively (Mosqueda-Vzquez et al., 1996;
Vzquez-Valdivia et al., 2000).
Mangoes in Taiwan are generally splice or veneer grafted; however, a
combination of the two methods is used when the rootstock and scion are
of different diameters. Polyembryonic Tsar-swain seedlings are pre-
ferred as clonal rootstocks although seedlings of monoembryonic Irwin
are sometimes used.
Crop Production: Management 445
The main commercial cultivars in the USA, Keitt, Tommy Atkins, Van
Dyke, Haden and Brooks, are monoembryonic. Orchards are planted with
grafted or budded trees (Aponte-Morn et al., 1977; Chia et al., 1988; Crane
and Campbell, 1991; Hamilton et al., 1992). The most common rootstocks in
Florida are seedlings of polyembryonic Turpentine, Number 11 and
Criollo. In Hawaii, any mango seedlings are used. In California, Turpen-
tine or Criollo are used. In Puerto Rico, polyembryonic Mangotino and
Pasote seedlings are used in the Ponce region and Mayaguezano, Turpen-
tine and Colombo Kidney in the Mayaguez region (Aponte-Morn et al.,
1977; Toro, 1988; E.E. Toro, personal communication). Regardless of the seed
source, roguing the zygotic seedlings is important for obtaining uniform tree
size, growth characteristics and production (Schnell and Knight, 1991, 1992;
Degani et al., 1993; Schnell et al., 1994). Seed is removed from mature fruit
along with the husk, and the seed is planted in well-drained container media
(Sauls and Campbell, 1980; Young and Sauls, 1989). Seedlings are large
enough to graft or bud after 26 months. The most common vegetative prop-
agation methods in Florida are veneer and cleft grafting (Sauls and Camp-
bell, 1980; Crane and Campbell, 1991) and chip and T or inverted T
budding. The trees are usually ready for eld planting in 612 months. Graft-
ing may be done at any time of year when suitable rootstocks are available,
but it is more successful during warm weather. In Puerto Rico the most
common propagation method is cleft grafting (Aponte-Morn et al., 1977;
Toro, 1988). Grafting is most successful during the spring and from September
through to November.
Topworking of established trees is common. Trees are cut back to several
major limbs or the trunk. In Florida, several sprouts emerging from the
pruned limbs are allowed to develop to 1.37.6 cm diameter, and are veneer
grafted with the new cultivar (Sauls and Campbell, 1980). In Puerto Rico,
several sprouts are allowed to develop, and larger sprouts (2.57.6 cm diam-
eter) are cleft grafted and smaller diameter sprouts (1.01.5 cm) are bark
grafted (Toro, 1988).
13.6 Major Cultivars
For detailed information about cultivars, refer to Knight et al., Chapter 3, this
volume.
Mango seeds were introduced by Portuguese and Spanish colonizers to
Brazil during the 16th century. Subsequently, mango seeds were dispersed
into the interior of the country. There are >120 local mango cultivars, many of
which are the result of open pollinations among Indian and Philippine mango
germplasm. There is a large amount of variability among the cultivars with
respect to colour, shape and size of the fruit. Most of these cultivars are for
the fresh fruit market (e.g. Espada, Bourbon, Ub, Rosa, Coit,
Coquinho and Lira), or for juice, jellies and other value-added products
(e.g. Ub, Coit, Mamo) (Table 13.3). Some polyembryonic criollo culti-
vars (e.g. Espada, Coquinho Comum and Fiapo) are used as rootstocks.

J
.
H
.

C
r
a
n
e

e
t

a
l
.
4
4
6
Table 13.3. Major commercial mango cultivars in Brazil, Mexico, Taiwan and the USA.
Country Cultivar Origin Seed type Production season
a
Tree growth habit
b
Brazil Espada Brazil Polyembryonic Oct.Dec.
Bourbon Brazil Monoembryonic Oct.Dec.
Ub Brazil Monoembryonic Oct.Dec.
Rosa Brazil Polyembryonic Oct.Dec.
Coit Brazil Monoembryonic Oct.Dec.
Mamo Brazil Monoembryonic Oct.Dec.
Coquinho Brazil Polyembryonic Oct.Dec.
Van Dyke USA Monoembryonic Nov.Jan. Large/spreading/open
Tommy Atkins USA Monoembryonic Nov.Jan. Large/dense
Haden USA Monoembryonic Nov.Jan. Large/spreading
Keitt USA Monoembryonic Feb.Mar. Small/spreading/open
Palmer USA Monoembryonic Jan.Feb. Small/spreading
Mexico Manila Mexico Polyembryonic Jan.Aug. Large/dense
Tommy Atkins USA Monoembryonic Feb.Aug. Large/dense
Haden USA Monoembryonic MayJuly Large/spreading
Kent USA Monoembryonic Feb.Aug. Large/dense
Keitt USA Monoembryonic MaySep. Medium/spreading/open
Ataulfo Mexico Polyembryonic Jan.June Large/spreading/upright
Taiwan Tsar-swain Taiwan Polyembryonic Mar.July
Jin-hwung Taiwan Monoembryonic JuneSept.
Tainong No. 1 Taiwan Monoembryonic MayJune
Irwin USA Monoembryonic AprilAug.
Keitt USA Monoembryonic Aug.Oct. Small/spreading
USA Tommy Atkins USA Monoembryonic JuneAug. Large/dense
Keitt USA Monoembryonic Aug.Oct. Large/spreading
Kent USA Monoembryonic JulyAug. Large/upright/dense
Van Dyke USA Monoembryonic JulyAug. Large/spreading/open
Parvin USA Monoembryonic JulyAug. Large/spreading/dense
Haden USA Monoembryonic Junemid-July Large/spreading
Brooks USA Monoembryonic Aug.Oct. Small/open
a
Natural fruit production season for country in which cultivar is being grown.
b
Growth habit for country in which cultivar is being grown.
Crop Production: Management 447
Florida selections (i.e. Haden, Keitt, Tommy Atkins, Van Dyke and Palmer)
are grown for the export trade. Most Florida cultivars produce from Decem-
ber to January except Palmer, which is harvested between January and Feb-
ruary and Keitt, which is harvested between February and March. Tommy
Atkins represents 80% of the commercial export volume in Brazil (Pinto,
2004); Palmer is increasing in demand in the national market.
Mango production in Mexico relies on the following cultivars (from early
to late beginning of harvest season): Manila, Ataulfo, Kent, Haden,
Tommy Atkins and Keitt (Table 13.3). Zill, Irwin, Sensation, Diplomtico,
Manililla, Oro, Ovo and criollos are also grown. Mango harvest seasons
are no longer inexible as they may be modied by pruning, water manage-
ment and growth regulators. The most widely planted cultivar in Mexico is
Manila (45,396 ha); no export gures have been reported. Although Manila
is grown all over the country it dominates in Veracruz (24,908 ha) and
Guerrero (9198 ha). It is an alternate bearer and the harvest season is from
May to June in Veracruz and July in Sinaloa (De los Santos and Mosqueda-
Vzquez, 198889; C. Guzmn, personal communication). There are 22,890
ha of Tommy Atkins and it is the major export cultivar to the USA (33% of
total exports). Fruit is harvested from February (Michoacn) to August
(Colima, Jalisco and Sinaloa) (V. Medina and C. Guzmn, personal commu-
nications). Kent is cultivated on 13,366 ha and accounts for 23% of exports
to the USA. Sinaloa is the major producer of Kent (9710 ha) and the begin-
ning of harvest season is similar to Tommy Atkins (Campbell, 1992); however,
it can be harvested as late as September in Colima, Jalisco and Nayarit.
Ataulfo is from Chiapas and is now cultivated nationwide on >34,000
ha. The major producing states are Chiapas (south) and Nayarit (north) with
18,334 ha and 7156 ha, respectively. The harvest season is FebruaryMay in
Chiapas and JuneJuly in Sinaloa (V. Palacio and C. Guzmn, personal com-
munications). In some years this cultivar is the third most important export
to the USA, and accounted for 19% of exports in 2006 (EMEX, 2007). Keitt
(6828 ha) is the fourth most important export cultivar (14% exports). Most
Keitt orchards are located in Sinaloa (5198 ha). Keitt is harvested from May
(Nayarit) to September (Colima, Jalisco and Sinaloa). Large fruit size and
fungal lesions sometimes cause marketing problems because harvest occurs
during the rainy season. Haden (27,768 ha) is still an important mango
although exports to the USA have decreased to 11% (EMEX, 2007); the major
producing states are Michoacn (15,573 ha), Guerrero (5053 ha) and
Sinaloa (3518 ha). Fruit are harvested from February/March (Michoacn and
Guerrero) to August/September in Colima, Jalisco and Michoacn. Alternate
bearing and mango malformation are problems and Haden orchards are
being replaced by or topworked to cultivars like Ataulfo.
The mango was introduced into Taiwan from South-east Asia by the
Dutch in the 16th century (Chen, 1991). The introductions were all polyem-
bryonic and included Tsar-swain, Pung-swain, Va-swain and Kee-gway-
swain. Tsar-swain is still very important (Table 13.3). From 1954 to 1973, 35
monoembryonic cultivars were introduced from the USA, and Irwin and
Keitt have been major cultivars in Taiwan since 1960. Jin-hwung, which is
J.H. Crane et al. 448
derived from a chance seedling of White and Keitt, was selected in 1980.
Tainong No. 1 and Tainong No. 2 are derived from controlled pollinations
and were released in 1985 by the Fengshan Tropical Horticultural Experiment
Station, Taiwan Agricultural Research Institute. Only Tsar-swain, Jin-hwung,
Irwin and Keitt and Tainong No. 1 are commercially important today.
Fruit in the southern prefectures or in lower elevation orchards are har-
vested earlier than the northern prefectures or orchards at higher elevations.
Tsar-swain comprises about 40% of total production; its season is from
March to August, depending on the prefecture or location of the orchard. The
fruit of Tsar-swain is 154 g, 100 mm long and 64 mm wide, total soluble
solids (TSS) value is 17 Brix, total titratable acidity 2.2% and seed/pulp
weight ratio 0.84 g. Irwin comprises 40% of the production and is harvested
from Pingtung in April and in Tainan in August. Jin-hwung comprises
about 10% of production and is harvested from June to September. The fruit
of Jin-hwung is 965 g, 144 mm long and 99 mm wide, TSS value is 15 Brix,
total titrateable acidity 2.3% and seed/pulp weight ratio 0.94 g. Keitt makes
up c.5% of production and is harvested from August to October. The fruit of
Tainong No. 1 is 237 g, 99 mm long and 69 mm wide, TSS value is 20 Brix,
total titrateable acidity 4.0% and seed/pulp weight ratio 0.85 g.
The major cultivars in Florida are Keitt and Tommy Atkins, which
account for c.70% and 20% of the hectarage, respectively (Table 13.3). Small
commercial hectarages of Van Dyke, Palmer, Irwin, Raposa and Kent
are also grown. In Puerto Rico, the major export cultivars are Keitt, which
makes up c.60% of the hectarage, Parvin (20%), Irwin (10%), Tommy
Atkins (5%) and Haden (< 5%). Other cultivars (i.e. Davis-Haden, Palmer,
Kent, Mayaguezano, Poste and Cubano) are grown on a small scale. The
local cultivars are grown for the domestic market (Toro, 1988). The commer-
cial hectarage of California is mostly Keitt, although other cultivars have
been evaluated (Scott, 1990; Linden, 2006).
Immature Keitt is the major early season cultivar (picked green) and
mature Tommy Atkins is the major early season cultivar in Florida (Table 13.3)
(J.H. Crane, personal communication). Mature Keitt and Kent mangoes
are the major late season cultivars. Six- to 8-year-old Tommy Atkins trees
produce 75150 kg/tree and older trees may produce up to 300 kg/tree.
Internal breakdown varies from year to year and may be aggravated by
over-fertilization with nitrogen (N). Fruit are considered moderately
resistant to anthracnose. The harvest season is June through to August.
Keitt trees are precocious and produce large crops regularly during July
through to September.
Palmer, Irwin and Van Dyke are grown commercially on a limited
scale in Florida and Puerto Rico (except Van Dyke). In Florida, Palmer is
harvested from July to early September, Irwin from June to early July, and
Van Dyke from July to August. In the recent past, Baileys Marvel, Brooks
and Haden were grown commercially; however, their importance has
declined due to natural disasters and susceptibility to anthracnose. Haden
is no longer grown commercially in Florida (Crane and Campbell, 1991;
Campbell, 1992) but, Glenn, a seedling of Haden, has been recommended
Crop Production: Management 449
(Campbell and Campbell, 1996). Harders is grown in Manoa, Oahu Island,
Hawaii (Hamilton et al., 1992; Hamilton, 1993). Trees bear regularly and often
produce off-season good-quality fruit in the late autumn and winter. Rapoza,
a seedling of Irwin, was selected in Hawaii in 1984 (Hamilton et al., 1992);
its harvest season is from mid-July to October.
13.7 Plant Spacing
Plant spacing and density are inuenced by climate, soil type and depth,
rootstock and scion vigour, growth habit and the targeted tree size. Cultural
practices including tree-size control, fertilizer and irrigation availability,
methods, rates, timing and frequencies; current technology and the necessity
for orchard access by farm machinery also inuence plant spacing and con-
guration. The cost of borrowed capital, land, water, irrigation, orchard
maintenance and net returns will dictate what cultivars are grown and how
they are managed (see Evans and Mendoza, Chapter 16, this volume).
Initial plant spacing and density should be planned to maximize early
yield and returns from young orchards and to maintain high yields from
mature orchards. Trees in overcrowded orchards compete for water, nutri-
ents and light and eventually lose production in the lower canopy. The ef-
cacy of foliar sprays (nutrients, pesticides and growth regulators) is reduced
and harvest is more difcult in crowded orchards.
Plant spacing among trees and rows has decreased in recent years to
optimize returns on investments in land, equipment and orchard infrastruc-
ture. This has been possible because of more insight into the physiology of
mango trees, improvements in irrigation system design and efciency, and
better fertilizer delivery systems and pruning (i.e. intense hand and/or
mechanical pruning) and the use of plant growth regulators. Closer plant
spacings require more tree-size control and expertise on the part of the pro-
ducer. Various training systems are advocated for new trees to force complex
branching and increase bearing surface volume and fruit production poten-
tial (Fig. 13.2). Topping and hedging and/or hand pruning (selective prun-
ing and/or heading back) is used to maintain mature tree size and fruit
production and improve fruit colour (Fig. 13.3).
In Brazil, the traditional spacing of 10 m 10 m in a rectangular or qua-
dratic format, with a density of 100 plants/ha, has been replaced by plant-
ings of 8 m (between rows) 5 m (within rows) to 5 m 5 m with higher plant
densities, which vary from 250 to 400 plants/ha (Cunha and Castro Neto,
2000; Mouco et al., 2002). In general, growers use two types of pruning: for-
mation and production pruning (Albuquerque et al., 2002). Formation prun-
ing is used to establish the intial tree architecture and trees are pruned ve
times for several years leaving c.243 branches prior to starting mango pro-
duction (Fig. 13.2). Pruning systems include cleaning, skirting of the lower
canopy, lateral, central and top-canopy pruning, and pruning to correct
poor canopy development and maintain adequate canopy after production
commences (Albuquerque et al., 2002).
J.H. Crane et al. 450
In most of Brazil, holes for plantings are 60 60 60 cm and simple or
triple superphosphate, lime and manure are usually mixed with the exca-
vated soil c.20 days prior to planting (Pinto and Ramos, 1998; Albuquerque
et al., 1999). In the north-east, planting is during the rainy season and new
plantings are mulched to reduce soil evaporation where solar irradiation and
temperature are very intense, which can lead to death of newly planted
mango trees. Intercropping with crops such as beans, maize, papaya and
pineapple improves incomes during the rst 3 years of orchard establishment
Planted
single
stem
Headed
to ~80 cm
First
branching
Tip prune
shoots 10 mm
diameter
After each flush
repeat tip pruning
of shoots 10 mm in
diameter once they
have matured
Fig. 13.2. Tree training system for young trees to increase branching and productive
canopy volume (Source: after Oosthuyse, 1995; Campbell and Wasielewski, 2000).
Central leader
Rectangle Rectangle with roof Open vase
Central leader topped Closed vase
Fig. 13.3. Various mechanical topping and hedging schemes that may include
selective hand pruning or shoot tipping to maintain tree size and regular bearing.
Crop Production: Management 451
(Mouco et al., 2002). Windbreaks, consisting of pine trees, elephant grass and
three rows of banana plants, are often used during the rst 2 years of orchard
establishment.
In Mexico, orchards were originally established at 10 m 10 m to 16 m
16 m, and trees were not pruned, which resulted in enormous and very pro-
ductive trees; however, care and harvest from very large trees is problematic.
Plant spacing is based on cultivar vigour (Ataulfo and Manila are most
vigorous), land slope (wider spaces as slope increases), farm machinery, cli-
matic conditions and soil fertility (wider distances for warmer climates and
more fertile soils) and water availability (Cruzaley-Sarabia et al., 2006). Irri-
gated orchards may handle closer spacings if pruning is practised. Under
rainfed conditions, soil moisture availability may have an important impact
on tree size. For example, 10-year-old orchards may have a few big trees
(c.69100 trees/ha) or if trees are spaced more closely (300600 trees/ha), tree
size is reduced.
Currently, both square and rectangular planting patterns are common in
Mexico although hexagonal planting systems are also used. For square
designs, tree spacing ranges from 5 m (400 trees/ha) to 10 m (100 trees/ha).
Rectangular planting systems have more options and the most common
arrangements are 6 m 5 m (333 trees/ha), 8 m 5 m (250 trees/ha), 8 m 6 m
(205 trees/ha) and 10 m 5 m (200 trees/ha) (Chvez-Contreras et al., 2001).
In Taiwan, plant spacing for monoembryonic cultivars (i.e. Tsar-swain)
ranges from 6 m 6 m to 10 m 10 m (100256 trees/ha). In contrast, plant
spacing for polyembryonic cultivars ranges from 4 m 5 m to 5 m 6 m,
depending on the topography and soil fertility. Orchards on sloped lands
and infertile soils are less vigorous and are planted at higher densities than
orchards on lowlands and fertile soils.
Recommended plant spacings in Hawaii and Florida are similar. The
cooler climate of south-eastern California allows close spacing, i.e. 3 m
in-row by 5.4 m between-rows (617 trees/ha). Traditional plant spacings in
Florida were as high as 11 m in-rows and 1114 m between-rows (6482
trees/ha) (Young and Sauls, 1989). Plant spacings in Florida currently include
3.5 m 6.1 m (468 trees/ha), 4.5 m 6.1 m (364 trees/ha), 4.5 m 7.6 m (292
trees/ha), 6.1 m 6.1 m (268 trees/ha), 6.1 m 7.6 m (215 trees/ha) and 7.6
m 7.6 m (173 trees/ha) (Young and Sauls, 1989; Crane and Campbell, 1991).
In Puerto Rico, plant spacings are 3.7 m 5.5 m (491 trees/ha), 4.6 m 9.1 m
(238 trees/ha), 6.1 m 9.1 m (180 trees/ha), 7.6 m 7.6 m (173 trees/ha) and
9.1 m 9.1 m (120 trees/ha) (Toro, 1988). Interplanting mango trees at moder-
ate to wide plant spacings (i.e. 5.47.5 m in-row) with banana or plantain
(Musa sp.), and papaya (Carica papaya L.) in Florida and plantains in Puerto
Rico is widely practised. Mango trees have also been planted at close spacing
(e.g. 3.04.5 m) and every other tree is removed if crowding becomes a prob-
lem. Controlling tree size and maintaining crop productivity is important,
otherwise, competition among trees will reduce yields and fruit quality (Toro,
1988; Crane and Campbell, 1991; Oosthuyse, 1995). Annual or biennial hand
pruning and/or mechanical hedging and topping is necessary and should
begin several years before trees begin to crowd and should continue after
J.H. Crane et al. 452
trees grow to their desired size based on plant spacing. Mature trees are
topped at 3.55 m, and hedged trees are cut on a slight angle (510) to leave
a 2.53.5 m row middle (J.H. Crane, personal communication). Trees can be
maintained at in-row spacings of 23 m; however, this involves intense tree
training and hand pruning, which most producers have been unwilling to
adopt (Fig. 13.3) (Oothuyse, 1995; Oosthuyse and Jacobs, 1995; Stassen et al.,
1999; Campbell and Wasielewski, 2000). Severe hedging is utilized to increase
light penetration and re-establish inner productive canopy but this can result
in little to no production in the following year. Some producers utilize a com-
bination of mechanical pruning followed by selective pruning to open the
inner canopy to light and air movement, improving fruit colour and reducing
disease pressure.
13.8 Fertilizer Practices
Fertilizer practices are inuenced by availability and cost of organic and
inorganic materials, application costs, soil type and depth, irrigation prac-
tices and rainfall, cultivar and production objectives. The response to fertil-
izer is inuenced by fertilizer source, rate, timing and method of application,
tree-growth stage, climate, edaphic conditions, soil moisture status and cul-
tivar vigour. The objective of fertilizer practices for young trees during years
1 and 2 is to establish the tree and maintain constant canopy growth and
health while building a structurally strong tree. Objectives for bearing trees
include maintenance of tree health, avoidance of alternate bearing and allow-
ance for vegetative dormancy which results in ower induction and initiation.
Over-fertilization of bearing trees leads to continuous vegetative growth
under some climatic conditions, reduced owering and fruit yields and
increased occurrence of physiological disorders of the fruit (Schaffer et al.,
1994; Coelho et al., 2002; Nguyen et al., 2004). Likewise, improper fertilization
may lead to nutrient deciencies and/or toxicities which result in reduce tree
growth, yields and fruit quality. Fertilizer practices will continue to be rened
and developed as our understanding of the physiological needs and responses
to essential plant nutrients increases and as concerns increase about the effect
of excess and/or leached nutrients on the natural environment.
Fertilizer management is based on soil and leaf analysis in Brazil (Tables
13.413.6; Santos et al., 2002). The quantity and type of fertilizer application
varies from the orchard establishment period (mostly vegetative plant
growth) to the orchard production period. At the time of planting, 2030 l/
hole of cattle manure is usually mixed with the native soil. Subsequently, N,
phosphorus (P) and potassium (K) are applied 23 times annually. During
young tree establishment, P and K are recommended only if these elements
are decient (Tables 13.5 and 13.6). These fertilizers are broadcast mechanically
in the planting rows, and then incorporated into the top soil layer (Andrade,
2004; Sousa et al., 2004). The quantities of macronutrients applied at the time of
planting and at various growth stages in south-eastern Brazil (mainly So
Paulo) and the central regions are based on plant age and soil P and K content
Crop Production: Management 453
Table 13.4. Standard leaf nutrient content ranges for mature mango trees in Brazil,
Mexico, Taiwan and Florida USA.
Mineral Unit
Range for mature trees
a
Brazil Mexico
b
Taiwan
c
USA
d
N % 1.21.4 1.21.5 1.41.7 1.01.5
P % 0.080.16 0.070.13 0.10.15 0.090.18
K % 0.51.6 0.60.7 0.91.2 0.51.0
Ca % 2.03.5 2.33.4 1.01.8 3.05.0
Mg % 0.250.5 0.140.19 0.20.35 0.150.47
B ppm 50100 NR
e
NR 2454
Fe ppm 50200 97114 60120 38120
Mn ppm 50100 191802 30200 92182
Zn ppm 2040 1420 20100 101119
Cu ppm 1050 512 520 2835
a
Recommended leaf nutrient levels based on research and the literature.
b
Source of data for Mexico: Guzmn-Estrada (2001, 2004, 2006). Range shown for all cultivars
tested (see Table 13.8 for detail by cultivar).
c
Values are for Irwin leaves. Source of data for Taiwan: Job (1989).
d
Source of data for USA: Young and Koo (1969, 1971); Young and Sauls (1989).
e
NR, not reported.
Table 13.5. Soil phosphate and potash corrective fertilization rates based on soil
analysis in Brazil.
(a) Rate of P
2
O
5
(kg/ha) application.
Clay content (%)
Level of P availability in the soil
Low Medium Adequate
15 60 30 0
1635 100 50 0
3660 200 100 0
>60 280 140 0
(b) Rate of K
2
O (kg/ha) application.
Soil K content
(mg/dm
3
)
Level of K availability in the soil
Low Medium Adequate
Cation exchange capacity = <4.0 cmol/dm
3
at pH 7 and <20% clay
<15 50
1640 25
>40 0
Cation exchange capacity = >4.0 cmol/dm
3
at pH 7 and >20% clay
<25 100
2580 50
>80 0
J.H. Crane et al. 454
(Table 13.5); however, the quantities of N, P and K for mango production are
based mainly on soil and leaf analysis. Leaves used for analyses are 68
months old, from the mid-canopy and from branches with fruits and from all
four sides of the canopy to reduce variation in analysis results. The proce-
dure for sampling leaves is: (i) divide the orchard into separate areas of no
more than 10 ha with trees of the same age and productivity and growing on
similar soil; (ii) collect healthy leaves from the middle of the tree canopy,
from the four cardinal points on normal branches with recently matured
leaves from the previous ush of growth, with leaves not less than 4 months
old. Remove four leaves per plant, from a total of 20 plants selected ran-
domly, and take the leaves prior to the application of nitrates or other foliar
fertilizers that are applied to break the dormancy of the oral buds.
There are two distinct periods of mango fertilization in Brazil: pre- and
postharvest fertilizations (Alves et al., 2002). In the preharvest fertilization of
non-irrigated orchards, P should be applied in a single dose, before ower-
ing, and incorporated with a medium-weight plough. At the beginning of the
rainy period 40% of the N and K should be applied and the remainder after
owering, during fruit development. In irrigated orchards c.40% of the P
should be applied before owering and 60% postharvest. For N, 50% is
applied preharvest (i.e. after the start of fruit set) and 50% postharvest. Potas-
sium applications should be distributed throughout the production cycle but
more during fruit development and c.25% postharvest. In So Paulo and cen-
tral Brazil c.40% and 20% of the N and K should be applied after harvest and
at the end of the rainy season (i.e. beginning of March), respectively.
When the productivity of orchards in north-east Brazil is <10 t/ha and
leaf N concentrations exceed 16 g/kg and the P and K concentrations in the
soil prole are >40 mg/dm
3
and 0.45 cmol/dm
3
, respectively, application of
N, P and K is unnecessary (Table 13.7). On the other hand, if the expected
productivity is >50 t/ha and leaf N concentrations are <12 g/kg and the P
and K concentrations in the soil prole are <10 mg/dm
3
and <0.16 cmol/
dm
3
, respectively, 120 kg/ha of N, 150 kg/ha of phosphate (P
2
O
5
) and 250 kg/
ha of K
2
O should be applied.
Table 13.6. Quantity of P
2
O
5
and K
2
O applications based on young tree plant age, and
P and K soil content of oxisoils for So Paulo and the Brazilian central regions.
Tree age
(years)
P soil content (mg/dm
3
) Exchangeable K soil content (mmol/dm
3
)
Trace <6 612 1330 >30 <0.8 0.81.5 1.63.0 >3.0
Rate of P
2
O
5
application (g/plant) Rate of K
2
O application (g/plant)
01 30 0 0 0 0 40 0 0 0
12 60 160 120 80 0 80 40 0 0
23 120 240 160 100 0 160 120 80 40
34 160 320 240 120 0 240 180 120 80
C
r
o
p

P
r
o
d
u
c
t
i
o
n
:

M
a
n
a
g
e
m
e
n
t
4
5
5
Table 13.7. Quantity of N, P
2
O
5
and K
2
O applications based on fruit productivity, leaf N content, and P and K soil content for the semi-arid
regions of Brazil.
Fruit
production
(t/ha)
N leaf content (g/kg) Soil P content (mg/dm
3
) K soil content (mmol
c
dm
-3
)
<12 1214 1416 >16 <10 1020 2140 >40 <1.6 1.63.0 3.04.5 >4.5
N application rate (kg/ha) P
2
O
5
application rate (kg/ha) K
2
O application rate (kg/ha)
1520 60 40 20 0 45 30 15 0 80 40 20 0
2030 75 50 25 0 65 45 20 0 120 60 30 0
3040 90 60 30 0 85 60 30 0 160 80 45 0
J.H. Crane et al. 456
Deciencies of zinc (Zn) and boron (B) are common in orchards in Brazil.
Zinc sulfate and borax are normally used to correct these deciencies; the
rates depend upon leaf analyses (Silva et al., 2002). Lime is applied when base
saturation is <60%. Gypsum at a rate of 2 t/ha for sand and 3 t/ha for clay
soils is recommended to reduce the incidence of internal breakdown (Silva
et al., 2002). Gypsum (280 g/m
2
) incorporated at a 30 cm soil depth before
orchard establishment under Cerrados conditions (pH 3.7 and very poor Ca
levels) reduces internal breakdown from 60 to 3% in Tommy Atkins (Pinto
et al., 1994).
Fertilization is not a common practice in most mango-producing regions
of Mexico. In Colima (Central Pacic region) <30% of orchards are fertilized.
However, in the Soconusco area of Chiapas (Southern Pacic region), 86% of
Ataulfo orchards are fertilized; 56.7% orchards are fertilized once a year,
39.5% twice a year, and 2.7% receive three applications per year. In fertilized
orchards, there is a high variation in amounts, materials, timing and forms of
fertilizer applications. Recommendations are empirical since there are few
published guidelines. Standard leaf nutrient levels have been proposed for
several cultivars grown in Sinaloa (Table 13.4, Table 13.8). Only a few growers
use leaf or soil analyses. Fertigation is rarely employed. In the Soconusco area
of Chiapas, Ataulfo mango trees have K, Ca, Mg, Zn and B deciencies. In
southern Sinaloa, most mango cultivars (Ataulfo, Haden, Tommy Atkins,
Kent, Keitt) are decient in N, P, K, Mg, S, copper (Cu), Zn and B, normal in
Ca and iron (Fe) and high in manganese (Mn) (Guzmn-Estrada, 2001). In
Colima, where mango is cultivated on sandy soils with pH 78.4, Fe and Zn are
the most common deciencies. In Nayarit, irrigated and non-irrigated Haden
and Tommy Atkins trees are decient in K, P and Ca (in this order) and have
excess Mg (Salazar-Garca et al., 1993). No Al toxicity has been reported.
Table 13.8. Standard leaf nutrient content values for selected mango cultivars in Mexico
(Source: Guzmn-Estrada, 2001, 2004, 2006).
Mineral Unit
Cultivar
Ataulfo Haden Keitt Kent Manila Tommy Atkins
N % 1.31 1.20 1.28 1.16 1.46 1.28
P % 0.07 0.09 0.15 0.08 0.09 0.13
K % 0.60 0.59 0.56 0.57 0.66 0.62
Ca % 2.53 3.40 2.87 3.23 2.31 2.77
Mg % 0.14 0.16 0.19 0.19 0.18 0.15
S % 0.23 0.17 0.20 0.15 0.15 0.17
B ppm NR
a
NR NR NR NR NR
Fe ppm 113.6 101.3 99.4 96.6 112.7 96.2
Mn ppm 801.9 328.1 191.1 243.8 236.3 219.3
Zn ppm 20.2 16.6 21.4 13.6 17.7 16.4
Cu ppm 5.40 5.43 11.70 6.70 6.83 6.96
a
NR, not reported.
Crop Production: Management 457
Except for N, visual symptoms of nutrient deciencies are uncommon in
commercial mango orchards; however, fertilized trees have signicantly
increased yield and fruit size. Nitrogen is most commonly applied, followed
by P and K. Current fertilization practices vary with the mango-producing
region, soil type, cultivar and tree age. The NPK recommendations for
Manila trees at 14 years, 510 years, 1015 years, 1620 years and >20 years
old in the Gulf of Mexico region are: 0.2-0.1-0.1 kg/tree/year, 0.4-0.2-0.4 kg/
tree/year, 0.6-0.3-0.6 kg/tree/year, 0.8-0.4-0.8 kg/tree/year and 1.0-0.5-
1.0 kg/tree/year, respectively.
Young trees in the Southern Pacic region receive NPK at 0.4-0.2-0.2 kg/
tree/year, respectively, from year 1 to year 5, and at 0.7-0.7-0.7 kg/tree/year
thereafter. In Michoacn (Central Pacic region), mature Haden and Tommy
Atkins trees receive NPK at 1.1-0.4-0.9 kg/tree/year (Chvez-Contreras
et al., 2001). The NPK recommendation in the Northern Pacic region at 14
years, 510 years, and 1015 years of age is 0.4-0.2-0.2 kg/tree/year, 1.3-0.55-
0.85 kg/tree/year and 2.8-0.9-1.8 kg/tree/year, respectively. Although soil
amendments are needed in regions with low or high pH soils (i.e. Chiapas
and Colima) they are not used.
Micronutrients are not usually included in fertilization programmes;
however, circumstantial evidence suggests there is an interaction between
ower B deciency and extreme high temperature as a major factor causing
stenocarpic fruit. Foliar B applications at either pre- or during full bloom
have become a routine practice for Ataulfo orchards. Zinc, Mn and Cu, are
included with fungicide sprays during bloom and the rainy season.
General fertilizer recommendations for mangoes of different ages vary in
Taiwan (Table 13.9). Fertilizers are applied in two equal rates: (i) after har-
vest; and (ii) when fruit are at the marble stage of development. Fertilizer is
applied by broadcasting, banding, side dressing and hole-application. Leaf
analysis is used for nutrient diagnosis and fertilizer recommendations (Table
13.4). The rates of N, P and K increase with tree age (Table 13.9). Dolomite is
commonly used to adjust soil acidity and as a Mg supplement. The applica-
tion rates are 1 t/ha/year for sandy soils, 1.5 t/ha/year for loamy or silty
soils and 2 t/ha/year for clay soils. Sulfur is applied as ammonium sulfate
((NH
4
)
2
SO
4
), magnesium sulfate (MgSO
4
) or calcium sulfate (CaSO
4
). Boron
is applied at a rate of 50 g/tree or as a 0.3 % spray 23 times a year. No addi-
tional micronutrient supplements are recommended.
Nutritional needs and fertilizer requirements under Florida conditions
have been well studied, although similar studies have not been done in
Puerto Rico (Aponte-Morn et al., 1977; Toro, 1988), Hawaii and California.
The sandy and calcareous soils in Florida are very low in native nutrient
content and cation exchange capacity. These soils require inorganic and
organic fertilizers for optimum growth and production. The leaf mineral con-
tent and deciency symptoms of young containerized Haden and Zill
trees in sand culture have been described for N, P, K, Mg, Mn and S (Smith
and Scudder, 1951). Obvious leaf symptoms of Ca, Cu, Zn and B were not
observed during a 3-year investigation, although classical Zn deciency
symptoms were observed in the eld.
J.H. Crane et al. 458
Leaf nutrient levels in mature mango trees on calcareous and sandy soils
have been investigated (Young and Koo, 1969, 1971; Koo and Young, 1972).
Signicant variation in mineral content was due to cultivar, leaf age, position
of sample leaf on the twig, presence or absence of fruit and soil type. Ranges
of mineral nutrient levels for maintaining productive trees under Florida
conditions were developed (Table 13.4). Fertilizer frequency, rates and timing
in Florida are based on observation, leaf and soil nutrient content and experi-
ence. Leaves are sampled at least once a year between December and Febru-
ary; mature leaves are collected from at least 30 trees at 0.92.4 m and from
all sides of the trees, and before the trees have been sprayed with nutrients
(Young and Koo, 1971; Koo and Young, 1972; Young and Sauls, 1989). Sam-
ples should be taken for trees showing nutrient deciencies, different culti-
vars and from groves under different cultural programmes and growing in
different soil types.
Nitrogen has the greatest effect on tree growth and yield and is used as
the basis for determining the amount of fertilizer to apply; however, some
caution must be used in recommendations from the past because of the
change from highly to less soluble fertilizer ingredients that has occurred
over the last 30 years (J.H. Crane, personal communication). The aim of the
programme for the rst 2 or 3 years is to produce a strong, healthy canopy so
that when production begins, trees will bear regularly and heavily. Young
non-bearing trees (13-years-old) are fertilized with 100225 g/tree of a
210% N and K (K
2
O) source (e.g. 4-2-8, 4-8-12, 8-2-8-2, 8-3-9; N-P
2
O
5
-
K
2
O-Mg) or similar material also containing P (P
2
O
5
) and Mg, at 68 week
intervals during the rst year (Young, 1974; Young and Sauls, 1989). About
2550% of N should be in an organic or slow-release form. The amount of
fertilizer is gradually increased (up to 1.4 kg application/tree) and the fre-
quency decreased (24 times/year) during years 2 to 4. Sources of N recom-
mended for mangoes in Florida soils include ammonium nitrate (NH
4
NO
3
)
and potassium nitrate (KNO
3
), some in a slow-release and/or organic form.
Urea is not recommended for calcareous soils because it volatilizes as ammo-
nia gas, but S-coated urea is used. Currently, low N analysis fertilizers are
recommended because excessive vegetative growth and increased fruit
physiological disorders (e.g. internal breakdown) are caused by moderate to
high N applications (Young and Miner, 1960, 1961; Nguyen et al., 2004). Cal-
cium applications may be necessary to raise the pH of acid sandy soils in
Table 13.9. General nitrogen, phosphate and potash fertilizer recommendations
(g of nutrient/tree/year) for mango trees in Taiwan.
Nutrient
Tree age (years)
12 34 57 810 >11
Nitrogen (N) 150 225 240 300 360
Phosphorus (P
2
O
5
) 20 75 160 200 240
Potassium (K
2
O) 120 225 360 450 540
Crop Production: Management 459
other areas of Florida. Recommended Ca sources include dolomite, gypsum,
calcium carbonate (CaCO
3
) and calcium nitrate (Ca(NO
3
)
2
).
Microelement application methods depend upon soil type, leaf analysis
and other cultural practices (Young, 1974; Young and Sauls, 1989; Crane and
Campbell, 1991). Foliar applications of Mn, Zn and Cu are made to trees
growing in calcareous soils, because soil applications are ineffective due to
high Ca content and pH. Soil applications of chelated Mn and Zn materials
formulated for trees growing in calcareous soils are recommended. Foliar or
soil applications of non-chelated Mn and Zn are appropriate for trees grow-
ing in acid sands, although foliar applications may be more efcacious and
less expensive. Usually Cu is applied as a fungicide and additional applica-
tions are unnecessary. Magnesium is frequently applied foliarly together
with Mn and Zn. Typical sources of Mn include manganese sulfate (MnSO
4
),
manganese nitrate (Mn(NO
3
)
2
) and manganese oxide (MnO), and of Zn
include zinc sulfate (ZnSO
4
), zinc oxide (ZnO) and zinc nitrate (Zn(NO
3
)
2
).
Recommended rates for foliar applications include 3.465.7 kg each of a Zn
and an Mn sulfate per 937/l of water/ha. However, commercial microele-
ment formulations containing Mn, Zn, Cu, molybdenum (Mo), S and B are
used because of their convenience. Microelements are applied 24 times/
year.
Iron is applied to soil as a drench with chelated materials formulated
for calcareous soils; however, these are expensive (Young and Sauls, 1989).
Typically, chelated iron as EDDHA (sodium ferric ethylenediamine
di-(o-hydroxyphenylacetate)) and EDDTA (sodium ferric diethylnetriamine
pentaacetate) can prevent and correct Fe deciency in calcareous and acid
sandy soils, respectively. For young trees, 7.114.2 g/tree applied as a soil
drench 24 times/year is recommended. Trees are irrigated for 12 days
before applying a mixture of chelated iron and water to the soil around the
base of the tree and then irrigated briey afterwards to ensure the material
has moved into the upper soil prole where most brous roots are located.
Foliar applications of ferrous sulphate (FeSO
4
) and other Fe compounds
(including chelated materials) with or without adjuvants have been ineffec-
tive for maintaining, preventing or correcting Fe deciency (Leonard and
Stewart, 1953; Leonard and Calvert, 1971; Green et al., 1999). Studies with
mild acids plus organosilicone adjuvant and FeSO
4
show promise as an
inexpensive foliar iron application method (Crane et al., 2007, 2008).
The fertilizer programme for fruit-producing trees involves maintenance
of tree health and fruit production and quality. The early recommendations
for mature mango trees growing in calcareous and acid sandy soils were
111168 kg of N and K/ha/year (Young and Miner, 1960, 1961; Young et al.,
1962, 1965; Young, 1974; Young and Koo, 1974); however, the recommended
N rate has been reduced to 4060% of earlier levels (J.H. Crane, personal
communication) due to the slow-release component of modern mixed fertil-
izer formulations. Nitrogen applications >225 kg/ha/year reduce fruit qual-
ity, cause excessive vegetative growth and increase internal breakdown
(Young and Miner, 1960, 1961; Raymond et al., 1998); leaf tissue analysis for
N should dictate application (Davenport, 2006). Leaf N levels >1.5% can
J.H. Crane et al. 460
result in little or no owering. Fertilizer mixtures containing 210% N and
416% K are used. Fertilizer rates may be reduced if leaf N and K levels are
within acceptable ranges and tree performance is acceptable. Phosphorus
applications may be reduced if leaf nutrient levels are within the acceptable
range; fertilizer mixtures with 35% P are normal. Additional Ca applications,
e.g. gypsum, dolomite, gypsum and Ca(NO
3
)
2
, on acid sandy soils increase
Ca leaf levels and reduce the incidence of internal fruit breakdown (Young
et al., 1962).
Microelements are applied to mature trees regardless of leaf analysis as a
preventative measure. This is especially true for Fe and Zn, because correct-
ing deciency of these elements is sometimes difcult and expensive. Rates
for foliar sprays of Mg, Zn and Mn are similar to those for young trees with
24 applications/year. The rates for chelated iron for mature trees range from
14 to 113 g/tree 12 times/year.
Fertilizer may be applied by fertigation. Mixed fertilizers containing P
are not recommended because phosphates can precipitate and clog the irri-
gation system. Phosphate can be applied alone as dilute H
3
PO
4
. Iron can also
be applied through microsprinklers or a drip system at increased frequency
and reduced rates.
In Puerto Rico, fertilizers are also applied in dry form and through low-
volume irrigation systems. Young non-bearing trees (13 years old) are fertil-
ized with 4541591 g/tree of a 1215% N, 56% P (P
2
O
5
) and 810% K (K
2
O)
source (e.g. 12-6-8, 12-6-10, 15-5-10; N-P
2
O
5
-K
2
O), split into two applications
(December/February and April/May) (Toro, 1988). When trees begin to bear
(after 34 years), 10-5-15, 12-6-16, or 10-5-20 is applied in split applications
which increase in amount with tree age (3.46.8 kg/tree maximum). Some
orchards are fertilized through the irrigation system at 715 day intervals
(E.E. Toro, personal communication). In these plantings, (NH
4
)
2
SO
4
, urea,
phosphoric acid (H
3
PO
4
) and KNO
3
or potassium sulfate (K
2
SO
4
) are used.
Micronutrients are applied either to the soil or to foliage in Puerto Rico; rec-
ommendations for young trees are foliar applications at 2130 day intervals
during the growing season (Toro, 1988). Micronutrients are applied as needed
in mature orchards and may be applied once during the spring and autumn.
Sources of micronutrients are sulfate forms of Zn, Cu and Mn.
13.9 Irrigation Practices
Water requirements for mango production are not precisely known and cur-
rent irrigation practices are based on experience and climatic and limited
edaphic measurements. Irrigation practices are inuenced by available tech-
nology and cost, soil type and depth, rainfall amount and distribution, fertil-
izer practices and production objectives. The response to irrigation is
inuenced by the rate, timing and method of application, tree-growth stage,
climate, edaphic conditions and cultivar.
Irrigation of young trees assists tree establishment, prevents drought
stress, and maintains constant canopy and root growth. Objectives for bearing
Crop Production: Management 461
trees are maintenance of tree health, avoidance of severe drought stress and
enhancement of vegetative dormancy. Mangoes are drought tolerant (Schaffer
et al., 1994); however, in some production areas under non-irrigated condi-
tions, fruit production and quality may be reduced. Excessive irrigation can
cause reduced tree growth and tree decline (Larson et al., 1989a, 1991d; J.H.
Crane, personal communication). Mango production has been displaced
onto marginal lands that possess low water and/or nutrient holding capacity
and high pH. Saline water for irrigation is a concern. Irrigation methods
include ood and furrow, high-pressure volume guns, high-volume under- and
over-tree sprinkler and low-pressure microsprinkler and drip systems.
In north-eastern Brazil, under semi-arid tropical conditions, irrigation is
essential throughout the hot and dry season (Albuquerque et al., 1999). Sev-
eral irrigation systems are used, with c.41% of orchards using microsprinkler
systems. About 21% use other types of irrigation (e.g. furrows, drip, basin,
etc.) and 33% of orchards use no irrigation (Gomes et al., 2002). Mean produc-
tivity of irrigated orchards may be as high as 25 t/ha compared with only 12
t/ha for non-irrigated orchards (Coelho et al., 2002). Irrigation is used on
14,500 ha (74% of the cultivated area) of commercial mango in the north-east.
Microsprinkler irrigation is the most common method (30.3% of the irrigated
area) (Gomes et al., 2002). Commercial mango orchards in the south-east,
mainly in So Paulo, are not irrigated.
Fertigated orchards require well-trained people. Some nutrients can
cause corrosion of irrigation pipes, and proper management is essential to
prevent environmental damage. Proper selection of nutrients is critical due
to problems of solubility and compatibility. Urea, (NH
4
)
2
SO
4
and KNO
3
are
the primary N fertilizers; they are highly soluble and are compatible with
most nutrients. However, the SO
4
2

is incompatible with Ca and their mixture
causes precipitation and clogging of emitters.
The oldest commercial orchards in Mexico were rainfed and established
in areas with deep soils and annual rainfall >900 mm (subhumid and humid
tropics). However, growth of the mango industry has forced the planting of
semi-arid tropical areas (annual rainfall 600700 mm) (Colima and Micho-
acn) (Table 13.2). Currently, 67% of orchards are rainfed (SIIAP, 2007) with
annual rainfalls that uctuate from 900 to 3700 mm; 80% of the rainfall occurs
in JuneOctober. Irrespective of the amount of annual rainfall, since the late
1980s a signicant proportion of the new and old mango orchards (60,000 ha)
have installed irrigation systems. This is to prevent water decits and to
increase yield and fruit size and quality. Sources of irrigation water include
rivers or deep wells and irrigation may be applied either by gravity or by
electrical or diesel engines.
Irrigation management involves either furrow or pressurized systems.
The furrow soil surface system (FSSS) can be simple, crossed, furrow-basin
and sh spine. Low-pressure drip and microsprinkler systems are most com-
mon, particularly the latter. Two microsprinklers (90120 l/h) or 1015 drip
emitters (812 l/h) are used per mature tree in a low-pressure system. In
semi-arid regions, mango potential evapotranspiration from October to June
each year is 16,000 m
3
/ha. The FSSS system applies >25,000 m
3
/ha of water;
J.H. Crane et al. 462
however, low-pressure systems can reduce water use to 5000 m
3
/ha with no
negative effect on yield or fruit quality (L.M. Tapia, personal communica-
tion). Water requirements in the semi-arid region of Michoacn are calculated
by using evapotranspiration measured with a Class A pan and multiplied by
the crop coefcient Kc, which for practical purposes is considered as 0.4 for
vegetative growth and 0.8 for fruit growth and development (Chvez-
Contreras et al., 2001). The FSSS water schedule for obtaining maximum
yield and fruit quality in Michoacn is: September (pre-bloom), one 20 cm
irrigation; OctoberDecember (bloom), two 10 cm irrigations spaced 20 days
apart; JanuaryApril (fruit growth), eight 10 cm irrigations at 1517 day inter-
vals; MayJuly (vegetative growth), three 50 cm irrigations at 2125 day inter-
vals. The amount of water used for drip irrigation in mature mango trees
during fruit growth and development is 1350 l/tree/week and for microsprin-
klers it is 1560 l/tree/week. After the rst irrigation, growers water every 8
days in low-pressure irrigation systems and every 1520 days in FSSS.
Soil-water content inuences the frequency of owering, the number
and length of panicles, yield and quality of mango fruits in Taiwan (Chang
and Lu, 1995). Two types of irrigation system are used: (i) furrow ooding in
the low lands; and (ii) microsprinklers, overhead sprinklers and drip (trickle)
irrigation on sloped lands. Microsprinklers have become the most important
irrigation system in recent years and fertigation is utilized by many produc-
ers. Irrigation management is based on producer experience and observation
of the soil, current weather and tree phenology.
In Florida, rainfall is not evenly distributed through the year, with the
dry season occurring during the autumnwinter (OctoberApril). During the
wet springsummer season (MaySeptember), dry periods of 3 days or more
can occur. Most orchards in Florida are irrigated only during prolonged dry
periods. Several systems are common. High-volume overhead or under-tree
sprinklers which run at high pressures (28,12170,303 kg/m
2
) and distribute
large amounts of water (0.510.89 cm/ha) and microsprinkler systems which
run at low pressure (703028,121 kg/m
2
) and distribute lower volumes of
water (37.91113.6 l/tree/ha). Some growers use both systems: the microsprin-
kler system for irrigation and fertigation and the high-volume system for
cold protection during freezing weather.
Well-established trees require little to no irrigation. During prolonged
drought conditions (>30 days with no signicant rainfall), irrigation may be
applied. Irrigation is not recommended during the cool autumn and winter
months, to enhance the vegetative dormancy period induced by cool tem-
peratures, to enhance synchronization of apical stems and to intensify the
owering response after growth commences.
Currently, irrigation recommendations and practices are based on obser-
vation and experience with c.6.35 cm water/application/ha. Preliminary
research suggests that irrigation at 7-day intervals during the period of fruit
development increases fruit size, earliness and yield (Larson et al., 1989b);
however, this has not been implemented by growers. Most Florida producers
rarely irrigate their mango orchards during the spring and summer as rain-
fall during this period coincides with fruit development and is sufcient for
Crop Production: Management 463
fruit production. In Puerto Rico, Hawaii and California, microirrigation is
used. In Hawaii, where temperatures are consistently warm in the mango-
growing areas, mature trees are more productive if irrigation is withheld for
at least 2 months before owering (Chia et al., 1988). Drip and microsprinkler
irrigation is used in California, but irrigation regimes have not been reported.
In Puerto Rico, drip irrigation is more common than microsprinklers (E.E.
Toro, personal communication). The drippers apply 3.797.75 l/tree/ha and
trees are irrigated 23 times/week. When microsprinklers are used, one
microsprinkler is placed between two trees in-row. Trees may be irrigated
from 911 months of the year; water is withheld for 13 months prior to oral
induction or owering.
13.10 Vegetative Growth and Reproduction Manipulation
Mango trees require new vegetative growth in order to produce fruit each year.
The optimum temperature for vegetative growth is 2430C (Krishnamurthi
et al., 1961; Sh and Sheen, 1987; Whiley et al., 1989) and when levels of essen-
tial plant nutrients and water are not limiting, vigorous growth results.
Mature leaves and a period of cessation of vegetative growth (i.e. mature api-
cal and subtending meristems) are required for the transformation from veg-
etative to reproductive growth (Nez-Elisea and Davenport, 1992; Kulkarni,
2004; Davenport, 2006).
Canopy management and reproductive manipulation vary according to
climatic conditions, cultivar and available technology (Davenport, 1993).
Ultimate mango-tree size depends upon the climactic and edaphic condi-
tions and cultural practices. Under optimum conditions, trees can reach
heights and canopy diameters of 30 m or more (Kostermans and Bompard,
1993); however, it is difcult to protect large trees from insects, diseases and
strong winds, and harvesting is difcult and costly. With the increase in costs
for orchard establishment and maintenance, the number of trees per unit
area has increased and tree size has decreased, whereas production of high
quality fruit per unit area has increased. Growers in some production areas
manipulate the period of owering and fruit production. Tree size can be
controlled to maximize the number of trees per unit area and maintain
productive tree canopy and yields.
The natural period of mango production in Brazil was originally from
October to January. Production has increased from September through to
April through the use of precocious and late-bearing cultivars along with
oral induction techniques. There is potential that whole-year harvesting
and mango supply can be achieved, although there are some months with
low mango yields. There are three types of oral induction and their use
depends upon phenology and season. In general, the steps are: (i) pruning of
apical internodes after harvesting; (ii) application of paclobutrazol (PBZ) to
stimulate owering by inhibiting gibberellin biosynthesis; (iii) spraying with
K
2
SO
4
(22.5%) to increase carbohydrate levels in the tree; (iv) drought stress
to facilitate growth cessation; (v) applications of ethylene (optional); and
J.H. Crane et al. 464
(vi) three applications of 34% KNO
3
and Ca(NO
3
)
2
to trigger owering
(Albuquerque et al., 2002). This ower initiation protocol results in off-season
owering and fruit production when mangoes are scarce and prices are high
(between April and August) in the domestic market.
Tree-size control and pruning strategies appropriate for the various cul-
tivars and diverse climatic regions in Mexico are under development.
Researchers in conjuction with producers are investigating appropriate tech-
nologies that take into account the current tree size, tree spacing and congu-
ration in the orchards, the available technology, and the goal of the pruning
programme. In young trees the central leader is pruned (5 mm above previ-
ous growth scar) a year after planting to promote lateral branching and pre-
cocious owering. Once new branches have matured the best-positioned 25
lateral branches around the main stem are selected. Trees are pruned for 23
years after every 23 vegetative ushes by heading back the wood of the last
two growths (Guzmn-Estrada and Vzquez-Valdivia, 2006). Young tree
training is important for Ataulfo and Keitt and results in strong canopies
that will resist heavy fruit loads and strong winds. Tree training is unnecessary
for Manila, Haden, Tommy Atkins and Kent.
Usually mature trees are pruned every year after harvest to promote
light penetration into the canopy and to remove weak, diseased, broken and
poorly positioned branches. Mechanical topping and hedging is not widely
practised. However, this pruning system was recently introduced in orchards
in Nayarit to control the pink hibiscus mealybug (Maconellicoccus hirsutus
Green). In Chiapas, 8-year-old trees and older are pruned by hand (machete)
in alternate years to delay or avoid canopy crowding; one side of the canopy
is pruned in one year and the other side is pruned the next year.
To rejuvenate old, very large mango trees and/or overcrowed orchards,
trees are cut back to main limbs at 12 m above the soil level. This technique
is common in the subhumid and semi-arid production areas of Chiapas and
Veracruz. However, in the warm humid tropical areas of Chiapas and Veracruz
this technique is not widely used because of the subsequent loss of fruit pro-
duction and excessive regrowth (34 vegetative ushes/year) that occur.
Selected trees can be removed to avoid overcrowding.
Climatic conditions are conducive to early bloom and harvest from the
Chiapas to Michoacn production regions, where forced blooming and early
fruit production is used by >80% of producers to obtain higher prices. Even
when a signicant amount of advanced bloom is induced, blooming during
the normal owering period may occur. The intensity of the normal bloom is
inuenced by the intensity of the fruit set by the advanced bloom. The most
common method to advance bloom involves canopy sprays of KNO
3
or
NH
4
NO
3
(Mosqueda-Vzquez and De los Santos, 1982; Nez-Elisea, 1986,
1988; Guzmn-Estrada, 1991; Sandoval-Esquivez et al., 1993). The effect of
these compounds is inuenced by cultivar and environmental conditions
(probably temperature). In the Gulf of Mexico region, one to two sprays of
2% KNO
3
or 1% NH
4
NO
3
solution are applied to Manila trees any time
from 15 October to 30 November to stimulate early owering. Manila and
Ataulfo are sprayed with 2% KNO
3
during the same period in the Southern
Crop Production: Management 465
Pacic region. In the Central Pacic region, Haden, Manila and Ataulfo
trees are treated with one or two sprays of 24% KNO
3
or 12% NH
4
NO
3
at
any time during the rst half of November. Similarly, Haden and Manila
are treated during November with 8% KNO
3
or 4% NH
4
NO
3
in the Northern
Pacic region. Tommy Atkins does not respond to foliar nitrate treatments
for promoting early owering. This is due to delayed oral initiation in this
cultivar so that when nitrate treatments are applied the buds are not irrevers-
ibly committed to owering (Prez-Barraza et al., 2000). Consequently, veg-
etative growth is produced in response to treatments that stimulate bud break
(Prez-Barraza et al., 2006a). However, application of PBZ promotes early
bloom in Tommy Atkins (Salazar-Garca and Vzquez-Valdivia, 1997).
Currently, soil applications of Cultar

(25% a.i.) close to the tree trunk is


used for most cultivars in dosages that range from 1 ml/m canopy diameter
for Manila in Veracruz to 24 ml for Tommy Atkins, Haden and Ataulfo
in Michoacn, applied at 12 year intervals. The response to PBZ treatment is
enhanced by 3045 days water stress and canopy sprays with nitrates
(Chvez-Contreras et al., 2001). In Nayarit and Sinaloa late bloom and harvest
are protable because they are the last two production areas to be harvested
in Mexico. Two canopy sprays of 50 mg/l GA
3
(15 and 30 November) cause
delayed bloom and shift 86% of the harvest to 1 month later (Prez-Barraza
et al., 2006b).
In Taiwan, cultivar, latitude, elevation and cultural practices are used
for off-season production (Sh et al., 2000). Several strategies are utilized to
stimulate owering and fruit set:
After harvest the last one or two vegetative ushes on each shoot are cut 1.
back to control tree size. Subsequently, two ushes of healthy shoots are
allowed to grow to serve as fruiting shoots for the next year. Weak or crowd-
ed shoots are removed to facilitate ventilation and light penetration.
In general, owering is not a problem in subtropical Taiwan; however, 2.
poor fruit set due to bad weather and lack of pollinators occurs occasionally.
A recent programme to increase the population of pollinators, mainly the
greenbottle ies (Chrysomyia megacephala Fabricius) in mango orchards has
been very successful. Increased yield has been noted and the practice has
been exploited commercially throughout the island.
Most of the mango fruit harvest goes to the domestic market within a 3.
short time period, causing the price to decline rapidly. Off-season fruit pro-
duction is thus very important to avoid this sharp drop in price.
Physical trunk damage by girdling and ringing and application of ethrel 4.
is used to promote early owering of the early season cultivar Tsar-swain
(Liu, 1996). Foliar applications of KNO
3
can promote early owering but
only if applied to ower-bud-induced shoots; PBZ is not recommended for
use on edible crops in Taiwan.
Panicle removal has been used to postpone owering and fruiting (Sh 5.
and Sheen, 1987; Sh, 1993). Emerging terminal panicles are removed by
hand. Chemical removal of terminal panicles with hydrogen cyanamide
(CH
2
N
2
) or calcium cyanamide (CaCN
2
) causes leaf damage and is not as
J.H. Crane et al. 466
effective as pruning (Hwang et al., 2004). Axillary panicle induction has some
advantages, because owering can be timed to avoid frost or cold tempera-
tures or a period of excessive rainfall during the normal owering period
(Singh et al., 1974; Shen and Huang, 1980). Axillary panicle removal also
reduces mango malformation and reduces alternate bearing (Majumder et al.,
1976; Pal and Chadha, 1982).
Control of owering and tree size in the USA varies with respect to differ-
ent climatic, edaphic and soil conditions as well as cultural practices (Daven-
port, 1993). In Florida, cool temperatures during the winter months (December
through to February) are usually adequate to arrest vegetative growth and
induce ower bud differentiation. In some years, the duration of cool tempera-
tures prohibits oral expression until late winter/early spring (February/
March) and when continuous warm temperatures begin (March), profuse, syn-
chronized owering occurs. In some years, warm and cool periods may occur
for a few days to weeks during the winter and partial owering may occur 24
times during the winter. This prolongs the owering and harvest season.
Flowering and fruiting of trees are not actively manipulated in Florida.
For young trees, vegetative growth is encouraged during the rst 23 years
and panicles may be removed by hand or natural pathogens, i.e. powdery
mildew or anthracnose are allowed to kill panicles and owers. Tipping and
selective pruning of young trees is recommended to improve tree structure,
control tree size and enhance early fruit production (Oosthuyse and Jacobs,
1995; Campbell and Wasielewski, 2000); however, selective pruning and
mechanical topping and hedging are used to control mature-tree size. Selec-
tive pruning usually involves removal of selected scaffold limbs to open up
the tree canopy to light and to remove dead wood. Mature trees may or may
not be allowed to grow together in the tree row to form hedgerows. Periodic
mechanical topping at 3.55 m and hedging to leave a 2.53.5 m row middle is
common (Crane and Campbell, 1991; J.H. Crane, personal communication).
Limiting the between-row spread of the trees to 2.53 m improves light pen-
etration into the tree canopy. In some orchards, hedging the inner sides of the
canopy of adjacent rows every 24 years and/or topping every third or fourth
row every 24 years is recommended. Trees are mechanically pruned immedi-
ately after harvest. Timing the pruning to selected rows each year ensures that
most of the planting will always be productive if continuous ushing of
pruned parts of the canopy prohibits reproductive growth the following
spring. Pruning trees shoots of 210 cm diameter immediately after harvest
and then tip pruning 34 times to force multiple lateral growths has been
advocated (Davenport, 2006). This strategy: (i) reduces or controls tree size;
(ii) shapes trees to facilitate subsequent tip pruning; (iii) synchronizes the veg-
etative ushing; and (iv) inhibits continuous vegetative ushing and prolongs
the period of vegetative dormancy. After vegetative growth has ceased for 5 or
more months, trees will synchronously ower when regrowth occurs.
In Puerto Rico, continuous vegetative growth of 12-year-old trees is
promoted. During this time panicles may be removed by hand to promote
vegetative growth, and this encourages rapid development of large trees.
Crop Production: Management 467
Temperatures are insufcient to inhibit vegetative growth and induce reli-
able and consistent bud differentiation, causing erratic or poor owering and
poorly synchronized fruit yields. The most common method for synchroniz-
ing and enhancing the time of owering involves a combination of drought
stress and timed application of KNO
3
. This method depends on cultivar and
the desired time of fruit harvest. To slow (or stop) vegetative growth and to
stress the trees, irrigation is withheld for 13 months prior to owering. The
drought stress is prolonged until leaves become dark green and show slight
signs of wilting. A 15% solution of KNO
3
is applied to the foliage, which
induces owering 34 weeks later. Regular irrigation is resumed when c.75%
of the panicles have set fruit. The timing of the drought stress and KNO
3

spray varies with cultivar and when fruit harvest is desired.
PBZ has been used to control vegetative growth after trees ush in
response to pruning following harvest. A soil drench of PBZ (710 g/tree) is
applied and trees are irrigated for 812 h. Irrigation is then withheld for 3090
days until trees show signs of drought stress. A foliar application of 12%
KNO
3
is applied, and owering occurs 3045 days later. The amount of tree
training depends upon the cultivar and is practised on young trees when
they are c.11.5 m high. Keitt and Palmer tend to have long branches of
various lengths and benet from training to create a stronger limb structure
and more compact tree. Trees are generally trained to a modied central
leader system, and some selective pruning of older trees is practised to open
the canopy to more light and air movement. Mechanical topping and hedging
are used to shape mature trees. Usually trees are topped to 34.5 m immedi-
ately after harvest.
In Hawaii, seasons with heavy crops are commonly followed by light or
no crops for 12 years (Nagao and Nishina, 1993). The small variation in
warm temperatures and evenly distributed rainfall encourage vegetative
growth, which reduces the potential for fruit production. This is less prob-
lematic on the drier leeward sides of the islands. Diseases (i.e. anthracnose
and powdery mildew) reduce production by infecting panicles and owers.
Preliminary trials with 2 and 4% KNO
3
applications during the winter (Feb-
ruary) resulted in 6684% owering 5 weeks after treatment (Nagao and
Nishina, 1993); however, this procedure is not utilized commercially.
Young Keitt mango trees are extensively trained by hand (usually twice
a year) during the rst 45 years to improve the structural strength of scaf-
fold limbs, increase the number of terminals and to control tree size in
California. Panicles are removed by hand during the rst 4 years. Training is
necessary because non-trained Keitt trees tend to have a few very long scaf-
fold limbs, little branching and are structurally weak. Bearing trees are
pruned annually to maintain trees at 3.14.6 m. In California, cool tempera-
tures during the winter months (November through to February) are ade-
quate to arrest vegetative growth and induce bud differentiation. The
duration of cool temperatures may inhibit oral expression until early spring
(March and April), when warm temperatures allow profuse, synchronized
owering. Warm periods during winter may stimulate early owering,
which may be damaged by subsequent cold temperatures (Schacht, 1992).
J.H. Crane et al. 468
Early season owering when cold temperatures and dry windy conditions
prevail (DecemberFebruary) results in poor fruit set and abnormal fruit.
Therefore, pruning of mature trees just before April (early spring) delays
owering and induces synchronous axillary owering after the danger of
cold has passed.
13.11 Environmental Stress Management
Mango is an adaptable species that withstands a range of subtropical and
tropical climates and soils. Physiological responses are related to the evolu-
tionary history of mangoes, with monoembryonic cultivars better adapted to
the subtropics and polyembryonic cultivars better adapted to the tropics.
In Brazil, the important commercial mango areas are in the tropical semi-
arid climate of the north-east. Flooding and freezing rarely occur there, but
wind and drought stress are very common and negatively affect growth of
young mango trees and increase fruit drop. Windbreaks reduce wind stress;
compact rows of elephant grass and/or rows of banana trees are utilized
(Mouco et al., 2002). In general, 34 rows of banana trees are planted around
or in perpendicular rows in the orchard against the main wind ow.
Drought stress, particularly in north-eastern Brazil, can suppress mango
growth and production, and irrigation is used to ameliorate this problem.
Drought stress has been used to increase endogenous ethylene concentra-
tions and trigger oral induction 7090 days after PBZ application. Drought
stress should be avoided during fruit development. Coelho et al. (2002)
reported the crop coefcient (Kc) for mango increased from 0.39 at owering
to 0.85 during fruit development.
In Mexico, the Gulf of Mexico coastal region may experience strong, dry
northerly winds (1128 m/s) from October to April and cause limb breakage
and increase ower and fruit drop by desiccation. To ameliorate this prob-
lem, growers have planted east-west oriented bamboo (Bambusa vulgaris)
and Australian pine (Casuarina equisetifolia) windbreaks. Bamboo is planted
outside the Australian pine trees and two rows of pines are planted in a stag-
gered arrangement. All coastal mango-producing regions in Mexico are
potential targets of hurricanes during the summer rainy season, even in late
October. Broken limbs are pruned, damaged trees are pruned back to sound
wood, and some trees are replaced or top-worked.
Flooding for 23 weeks may occur during the summer rains. This is
mainly a problem in low-lying areas of Chiapas and Nayarit that have loamy
soils and a high water table. Drainage canals have been constructed in such
areas to minimize the problem. Drought stress is common as 67% of Mexicos
mango orchards are not irrigated (SIIAP, 2007). Annual rainfall ranges from
900 to 3700 mm in Mexico and is concentrated in June to October. The dry
season begins in October (close to the period of oral initiation) and contin-
ues through the harvest period for early and some mid-season cultivars.
Drought stress in areas of low rainfall causes intense fruit drop, which reduces
yield and fruit size especially in heavily bearing trees. More that 60,000 ha of
Crop Production: Management 469
mangoes are irrigated in low rainfall areas. In Veracruz and Chiapas most
orchards are adjacent to riverbanks and the deep soils provide root access to
the water table. No data are available supporting the benet of irrigation in
medium-high precipitation areas.
Most mango-producing regions in Mexico are frost free. The only report
of frost damage to 24-year-old mango orchards was in Sonora where freez-
ing events may occur every 68 years (E. Snchez, personal communication).
Low temperatures (10C) during bloom reduce fruit set, especially in
Ataulfo. Treatment with GA
3
to delay the bloom has been suggested as a
method to avoid low-temperature damage during owering but owering
under high-temperature conditions is also detrimental to fruit set and crop
yield (Prez-Barraza et al., 2006b).
In Taiwan, damage from typhoons occurs periodically and trees are reset
and either pruned to recover or replanted. In general, freezing temperatures
are not a problem in the mango-production areas but cool temperatures can
reduce fruit set. To avoid cool or cold temperatures during the normal ow-
ering period, shoot tips may be pruned to delay and force owering from
lateral buds. Flooding in production areas is uncommon, and drought stress
is not an issue because most producers irrigate during dry periods.
The production areas of Hawaii and Puerto Rico are frost free; however,
Florida and California may experience temperatures at and below freezing
during the winter months (December through to February). In Florida, freez-
ing temperatures (0 to 6C) may occur for a few hours for 14 nights/year,
although freezing temperatures for 1315 h within a 24 h period have been
reported (Johnson, 1970; Campbell et al., 1977). In California, temperatures as
low as 6.6C are common and frosts may occur 1015 times/year during
January and February (Aslan et al., 1993). Mango trees do not acclimate to
cold temperatures (McKellar et al., 1983), although differences in cold toler-
ance and recovery from cold damage have been observed (Carmichael, 1958).
In Florida, high-volume overhead and under-tree irrigation is used to protect
trees during freezing weather. At least 0.6 cm of water/ha is distributed.
Overhead systems are designed for complete coverage (overlapping spray
patterns) of the trees and under-tree systems are designed to spray 0.92.4 m
into the tree canopy. Irrigation commences before freezing temperatures are
reached (usually c.23C) and continues until ice has melted. These systems
are powered by diesel or gas engines as electrical power is unreliable during
freezing weather conditions. In California, microsprinklers, wind machines
and helicopters are used to raise the air temperature of plantings (Schacht,
1992).
Mango trees are relatively tolerant of wind stress (Schaffer et al., 1994;
Crane and Balerdi, 2005); however, newly planted trees are commonly staked
at planting in the calcareous soils of Florida to prevent damage to the bark
and cambium caused by constant movement and rubbing against the rocky
soil. Staking also stabilizes the tree against toppling during hurricanes. In
contrast, tolerance to mature trees depends on tree size, with larger trees
being more vulnerable to wind damage than pruned trees (Crane et al., 1993,
1994, 2001; Crane and Balerdi, 1996; NASS, 2006).
J.H. Crane et al. 470
In the mid-20th century, windbreaks of C. equisetifolia were planted
around many mango orchards in Florida. However, intrusion into the orchard,
shading and damage to mango trees when trees toppled into orchards during
hurricanes has stopped this practice. Some producers have topped remain-
ing pine windbreaks at 4.96.7 m to reduce orchard shading and their potential
to topple (Crane et al., 1993). In Hawaii, natural windbreaks are recommended
for some areas where constant winds are a problem for establishing young
trees (C.L. Chia, personal communication). In California, constant winds
during spring may damage panicles and young fruit; man-made windscreens
are used to protect trees (Scott, 1990; Schacht, 1992). The windscreens can be
raised to prevent trapping of cold air within the plantings. Puerto Rico is
affected by hurricanes but no specic ameliorating recommendations have
been reported (Toro, 1988).
Flooding is not common in production areas of Puerto Rico, California
and Hawaii; however, periodic ooding (121 days) is typical during the
summer in Florida. Trees have been planted on beds of crushed limestone
rock, 0.61.0 m high and 1.01.5 m wide, which allows part of the root system
to be above water. Planting of orchards at sites at or below 2 m above sea level
is not recommended. Mango trees are moderately ood tolerant (Larson et al.,
1991c; Schaffer et al., 1994, 2006), although this is affected by oodwater tem-
perature, oxygen content and anatomical adaptations of the rootstock (Lar-
son et al., 1991a, 1993a, b). Periodic ooding of the limestone-based soils in
Florida increases the availability of soil Mn and Fe (Larson et al., 1991b, 1991d,
1992), alleviating plant deciencies of these elements. Furthermore, rhizo-
sphere anoxia increases reduction of Fe
3+
to Fe
2+
by nicotinamide adenine
dinucleotide (NADH) in mango root tissue and Fe uptake by mango roots in
oxygen-depleted media (Zude-Sasse and Schaffer, 2000).
13.12 Harvesting Practices
Harvesting is done by hand in Brazil, Mexico, Taiwan and the USA (Evans,
2007) and is one of the most expensive operations in mango production because
fruit do not mature synchronously, and trees require multiple pickings. The
technology to mechanize mango harvest is difcult because of differences in
fruit colour, size and weight among cultivars, difculty in determining fruit
maturity, requirement for multiple pickings, lack of tree-size-controlling
rootstocks and tree training and moderate to large tree canopies.
Prior to harvesting in Brazil, excessive set fruit and injured and diseased
fruit are removed, part of the rachis is removed to prevent scarring and bruis-
ing of the fruit, and some leaves that shade fruit may be removed to allow for
better peel colour development (Alves et al., 2002). Fruit in the lower part of
the canopy are harvested from the ground; however, ladders are required for
picking fruit high in the tree canopy. Peel damage due to latex exudation at
the stem end of the fruit during harvest is a very common problem, and
>50% of harvested fruit may be affected. This problem occurs mainly when
fruit are picked high in the canopy with a picking pole by severing the fruit
Crop Production: Management 471
near the stem end of the pedicel. An improved picking pole that cuts the
petiole c.2 cm above the pedicel reduces latex burn to <10%. Before trans-
porting to the packing house, fruit containers are placed in the shade to avoid
increasing the pulp temperature.
In Mexico, harvesting is done by hand when fruit have reached physio-
logical maturity. Picking poles with bamboo baskets (Gulf of Mexico and
Southern Pacic regions) or nylon net bags or cotton bags (Central and
Northern Pacic regions) and a cutting blade at the distal end are used to
reach fruit high in the canopy. Ladders may be used, although climbing trees
is more common. Fruit is usually placed in 2030 kg wooden or plastic boxes.
The inner walls of the picking crates are covered by newspaper to absorb
latex and decrease sap peel damage.
Several fruit maturity indexes exist for mango: pulp TSS content and/or
acidity, pulp rmness, skin or pulp colour, calendar days or heat units from
bloom to harvest. No index alone is successful because of differences among
cultivars and signicant variability in fruit maturity within trees and orchards
and among orchards. Pickers are trained to distinguish the following charac-
teristics: (i) size, form and fruit colour; (ii) shoulder development (higher
than the base of peduncle); (iii) cavity formation at the base of the peduncle;
and (iv) increased lenticel size (Chvez-Contreras et al., 2001). The Associa-
tion of Mango Packers and Exporters (EMEX; Empacadoras de Mango de
Exportacin) have established some minimal physical and chemical stan-
dards to dene maturity for Haden, Tommy Atkins, Kent, Keitt and
Ataulfo. A photographic maturity index chart is used by growers, pickers,
companies and packing houses. Mango fruit-peel damage by latex is common
when tree sap pressure is high. To minimize this damage, irrigation is stopped
at least 2 weeks prior to harvest; fruit are also picked with a long peduncle
and are washed immediately after the harvest bin is full.
Mature-green mangoes are the rst fruit to be harvested and all the cri-
ollo, Oro and Florida cultivars t this category. These mangoes are for the
domestic market and usually sell for high prices. Manila mangoes are
picked when their colour changes from pale green to greyish green. Ataulfo
is harvested when the green peel shows a yellow colour break. The Florida
mangoes are picked mature-green and at colour break.
The harvest season in Taiwan begins in March with Tsar-swain and
ends in September or October with Keitt. Depending on the cultivar and
market, fruits are harvested at 7080% maturity. Tsar-swain is harvested
either at the green stage for pickles or at a mature ripe stage for domestic
markets. Irwin fruit are harvested at either 80% maturity for export or at
>90% maturity for local markets. Green mature fruits may be triggered to
ripen with calcium carbide, ethephon or ethrel at 3040C, depending on the
cultivar. Mature fruits are stored at at 812C for several days to several
weeks. Fruit destined for export are disinfested using the vapour heat method;
the fruit core temperature must reach and be held at 46.5C for 30 min.
There are two markets for mango producers in the USA: the green mar-
ket for non-ripe fruit and the tree-ripened market. The green and tree-ripened
markets are speciality, niche markets where the green fruit are used as a
J.H. Crane et al. 472
component of processed foods (e.g. chutneys, preserved pieces), whereas the
tree-ripened fruit target the demand for ready-to-eat mangoes. Green mangoes
are picked before the fruit is mature, whereas the tree-ripened mangoes are
allowed to develop almost full colour and ripeness before being picked.
Florida mangoes are more expensive than imports and the volume of fruit is
very limited and cannot supply the demand of the national market. Usually
multiple pickings are required to harvest the crop but this is inuenced by
market demand and prices, the number of blooms producing the crop and
weather conditions.
Harvesting in the USA is by hand. A long picking pole with a canvas or
nylon bag attached to a metal ring with a cutting blade at the distal end is
commonly used in Florida (Aponte Morn et al., 1977; Crane and Campbell,
1991). Other picking aids such as ladders and mobile hydraulic lifts are also
used. Time of harvesting depends upon cultivar, the intended market and
market demand. In Florida, green mangoes may be picked after March, while
fruit-picking time for the fresh market depends upon the cultivar reaching
the mature stage desired (Crane and Campbell, 1991). In Puerto Rico, man-
goes are harvested from March to November, depending upon cultivar, mar-
ket price and the date when owering was induced. The mango season in
California is restricted to September/November, and in Hawaii the main sea-
son is May to August. Approximately 50% of mangoes produced in California
are certied organic (Linden, 2006).
13.13 Conclusions
Differences in mango culture are due to the climatic and edaphic conditions,
available information and technology, and tradition in each production area.
The interaction of climate and cultural practices, for example irrigation, fer-
tilizer, pruning, etc., that are essential for optimizing crop yields and quality
is not completely understood. None the less, horticultural systems can be
developed and tested that can impact fruit production and quality. It is
important that area- and, in many cases, site- and cultivar-specic cultural
information and practices need to be developed to optimize production and
fruit quality. This chapter has addressed the current state of mango culture in
four different production areas, and has emphasized improvements that are
essential for a prosperous industry.
Ackowledgements
The Mexican co-author acknowledges the following INIFAP researchers,
based at several Research Stations (CE) and states: Ernesto Snchez-Snchez,
CE-Valle del Yaqui (Sonora); Camerino Guzmn-Estrada, CE-Sur de Sinaloa
(Sinaloa); R. Mosqueda-Vzquez (deceased) and Enrique N. Becerra-Leor,
CE-Cotaxtla (Veracruz); Fulgencio M. Tucuch-Cauich, CE-EDZNA (Campeche);
Crop Production: Management 473
Rubn Cruzaley-Sarabia, CE-Iguala (Guerrero); Vctor Medina-Urrutia,
CE-Tecomn (Colima); Vctor Palacio-Martnez, CE-Rosario Izapa (Chiapas).
References
Agrianual (2006) Gazeta, Santa Cruz, Rio Grande do Sul, Brazil.
Agrianual (2007) Manga, mercado and perspectivas. Anurio da Agricultura Brasileira.
Instituto FNP, AGRA FNP Pesquisas Ltd, So Paulo, Brazil, pp. 378386.
Agriculture and Forestry Department (1996) Taiwan Agriculture Yearbook. Economic
Section, Agriculture and Forestry Department, Taiwan Provincial Government,
Nantou, Taiwan.
Albuquerque, J.A.S. de, Mouco, M.A. do C., Medina, V.D., Santos, C.R. dos and Tavares,
S.C.C. de H.O. (1999) Cultivo da Mangueira Irrigada no Semi-rido Brasileiro.
Embrapa Semi-rido, Petrolina, Brazil.
Albuquerque, J.A.S. de, Mouco, M.A. do C., Medina V.D. and Vasconcelos, L.F.L. (2002)
Sistemas de poda. In: Genu, P.J.C. and Pinto, A.C. de Q. (eds) A Cultura da Mangueira.
Embrapa Informao Tecnolgica, Braslia, Brazil, pp. 244257.
Allen, R.G., Perrira, L.S., Raes, D. and Smith, M. (1998) Crop Evaporation, Guidelines
for Computing Crop Water Requirements. Food and Agriculture Organization (FAO)
Irrigation and Drainage Paper 56. FAO, Rome.
Alvarado-Ortiz, A.N. and Acin, N. (2004) Crop prole for mangoes in Puerto Rico. Avail-
able at: http://www.ipmcenters.org/cropproles/docs/PRmango.html (accessed 19
May 2008).
Alves, R.E., Filgueiras, H.A.C., Menezes, J.B., Assis, J.S. de, Lima, M.A.C. de, Amorim,
T.B.F. and Martins, A.G. (2002) Colheita e ps-colheita. In: Genu, P.J.C. and Pinto,
A.C. de Q. (eds) A Cultura da Mangueira. Embrapa Informao Tecnolgica, Bras-
lia, Brazil, pp. 382405.
Andrade, L.R.M. de (2004) Corretivos e fertilizantes para culturas perenes e semiper-
enes. In: Sousa, D.M.G. and Lobato, E. (eds) Cerrado, Correo do Solo e Adubao.
Embrapa Informao Tecnolgica, Brasilia, Brazil, pp. 317366.
Anonymous (1970) Clasicacin del Suelo. Food and Agriculture Organization (FAO)/United
Nations Educational, Scientic and Cultural Organization (UNESCO), Mxico.
Anonymous (1982) Inventarios de reas Erosionadas, Rangos de Pendiente y Unidades
de Suelo del Estado de Veracruz. Direccin General de Conservacin del Suelo y
Agua. Secretara de Agricultura y Recursos Hidrulicos (SARH), Mxico.
Anonymous (1989) Interim Report. South Dade Soil and Water Conservation
District and Soil Conservation Service. United States Department of Agriculture
(USDA), Soil Conservation Service, United States Government Printing Ofce,
Washington, DC.
Anonymous (2007) Importation of mangos from India. Federal Register 72, 10902
10907.
Aponte-Morn, C.E., Ayala-Almodvar, A., Cibes-Viad, H., Jackson, G., Liu, L.J.,
Llorns, A., Lpez-Garca, J., Orengo-Santiago, E., Prez-Escolar, M.E., Prez-Lpez,
A., Reyes-Soto, I., Toro-Toro, E. and Torres-Seplveda, A. (1977) Conjunto Tecno-
logico para la Produccion de Mango. Publicacion 114. Universidad de Puerto Rico,
Estacion Experimental Agricola, Ro Peidras, Puerto Rico.
Aslan, S., Neja, R., Estrada, D., Dignon, W. and Mitchell, S. (1991) Soil, Water, and Cli-
matic Considerations in Selecting Date Palm Planting Sites in the Coachella Valley.
Coachella Valley Resource Conservation District, Coachella Valley Water District,
Riverside County Farm Bureau, Coachella Valley Center, Indio, California, pp. 620.
J.H. Crane et al. 474
Aslan, S., Bergan, M., Baetz, R., Slayback, R.D., Dyer, D. and Dignon, W. (1993) Ten-
year Research Findings on Water-efcient Ornamental Plants for the Coachella Val-
ley. Desert Water Agency and Coachella Valley Resource Conservation District,
Palm Springs, California, pp. 26.
Bahr, L.S. and Johnston, B. (eds) (1993a) California. Colliers Encyclopedia. Vol. 5. P.F.
Collier, New York.
Bahr, L.S. and Johnston, B. (eds) (1993b) Hawaii. Colliers Encyclopedia. Vol. 11. P.F.
Collier, New York.
Bahr, L.S. and Johnston, B. (eds) (1993c) Puerto Rico. Colliers Encyclopedia. Vol. 19. P.F.
Collier, New York.
Barrick, W.E. and Black, R.J. (1980) Florida Climate Data. Florida Energy Extension Ser-
vice, EES-5. Florida Cooperative Extension Service, University of Florida, Institute of
Food and Agricultural Sciences, Gainesville, Florida.
Bierbolini, R.E., Ramos, G.A., Mas, J.T., Torres, E.O., Trigo, J.E., Rivera, W.F., Brunet, J.E.
and Rivera, L.H. (1975) Soil Survey of Magaguez Area of Western Puerto Rico.
USDA-Soil Conservation Service and University of Puerto Rico, Cartographic Divi-
sion, Washington, DC, pp. 1311.
Bierbolini, R.E., Acevedo, G., Mas, J.T., Torres, E.O. and Rivera, L.H. (1979) Soil Survey
of the Area of Southern Puerto Rico. United States Department of Agriculture
(USDA), Soil Conservation Service and University of Puerto Rico, US Government
Printing Ofce, Washington, DC, pp. 187.
Bradley, J.T. (1975) Freeze Probabilities in Florida. Bulletin 777. Florida Cooperative
Extension Service, University of Florida, Institute of Food and Agricultural Sciences,
Gainesville, Florida.
Butson, K.D. and Prine, G.M. (1968) Weekly Rainfall Frequencies in Florida. Circular
5-187. Florida Cooperative Extension Service, University of Florida, Institute of
Food and Agricultural Sciences, Gainesville, Florida.
Calhoun, F.G., Carlisle, V.W., Caldwell, R.E., Zelazny, L.W., Hammond, L.C. and
Breland, H.L. (1974) Characterization Data for Selected Florida Soils. Soil Science
Research Report Number 74. United States Department of Agriculture (USDA), Soil
Conservation Service, Washington, DC.
Campbell, C.W. and Campbell, R.J. (1996) The Glenn mango, an early-maturing culti-
var. Proceedings of the Florida State Horticultural Society 109, 233234.
Campbell, C.W., Knight, R.J. Jr and Zareski, N.L. (1977) Freeze damage to tropical fruits
in southern Florida in 1977. Proceedings of the Florida State Horticulture Society
90, 254257.
Campbell, R.J. (1992) A Guide to Mangos in Florida. Fairchild Tropical Garden, Miami,
Florida.
Campbell, R.J. and Wasielewski, J. (2000) Mango tree training techniques for the hot
tropics. Acta Horticulturae 509, 641651.
Carlisle, V.W., Caldwell, R.E., Sodek III, F., Hammond, L.C., Calhoun, F.G., Granger,
M.A. and Breland, H.L. (1978) Characterization Data for Selected Florida Soils. Soil
Science Research Report Number 78-1. United States Department of Agriculture
(USDA), Soil Conservation Service, Washington, DC.
Carmichael, W.W. (1958) Observations of cold damage to mangos in Dade County and the
lower east coast. Proceedings of the Florida State Horticulture Society 71, 333335.
Castro Neto, M.T., Fonseca, N., Santos Filho, H.P. and Cavalcante Junior, A.T. (2002)
Propagao e padro da muda. In: Genu, P.J.C. and Pinto, A.C. de Q. (eds) A Cultura
da Mangueira. Embrapa Informao Tecnolgica, Braslia, Brazil, pp. 118136.
Chacko, E.K. (1986) Physiology of vegetative and reproductive growth in mango (Mangifera
indica L.) trees. In: Proceedings of the First Australian Mango Research Workshop.
Crop Production: Management 475
Commonwealth Scientic and Industrial Research Organization (CSIRO), Melbourne,
pp. 5470.
Chacko, E.K. (1989) Mango owering still an enigma. Acta Horticulturae 291, 1221.
Chadha, K.L. and Pal, R.N. (1986) Mangifera indica. In: CRC Handbook of Flowering,
Vol. 5. CRC Press, Boca Raton, Florida, pp. 211230.
Chang, M.-C. and Lu, C.-J. (1995) Effects of soil water content on the development, yield
and quality of mango (Mangifera indica). Research Bulletin Tainan, District Agricul-
tural Improvement Station 32, 4555.
Chvez-Contreras, X., Vega-Pia, A., Tapia-Vargas, L.M. and Miranda-Salcedo, M.A.
(2001) Mango, su Manejo y Produccin en el Trpico Seco de Mxico. Libro
Tcnico 1. Secretara de Agricultura, Ganadera, Desarrollo Rural, Pesca y Aliment-
acin (SAGARPA)-Instituto Nacional de Investigaciones Forestales, Agrcolas y
Pecuarias (INIFAP), Campo Experimental Valle de Apatzingn. Michoacn, Mexico,
pp. 108.
Chen, M.-S. (1991) Current status and prospect of mango production in Taiwan. In: Pro-
ceedings of Symposium on Production Research of Fruit Crops, Chiayi City, Taiwan,
pp. 317332.
Chia, C.L., Hamilton, R.A. and Evans, D.O. (1988) Mango. Commodity Fact Sheet MAN-
3(A). Hawaii Cooperative Extension Service, Hawaii Institute of Tropical Agriculture
and Human Resources, University of Hawaii at Manoa, Honolulu, Hawaii.
Coelho, E.F., Oliveira, A.S. de, Netto, A.O.A., Teixeira, A.H. de C., Arajo, E.C.E. and
Bassoi, L.H. (2002) Irrigao. In: Genu, P.J.C. and Pinto, A.C. de Q. (eds) A Cultura
da Mangueira. Embrapa Informao Tecnolgica, Braslia, Brazil, pp. 167189.
Colburn, B. and Goldweber, S. (1961) Preparation of oolitic limestone soil for agricul-
tural use. Proceedings of the Florida State Horticultural Society 74, 343345.
Crane, J.H. and Balerdi, C.F. (1996) Effect of Hurricane Andrew on mango trees in Florida
and their recovery. Acta Horticulturae 455, 323330.
Crane, J.H. and Balerdi, C.F. (2005) Preparing for and Recovery from Hurricane and
Tropical Storm Damage to Tropical Fruit Groves in Florida. HS1022. Florida Coop-
erative Extension Service, University of Florida, Institute of Food and Agricultural
Sciences, Gainesville, Florida, pp. 18.
Crane, J.H. and Campbell, C.W. (1991) The Mango. Fact Sheet FC-2. Florida Coopera-
tive Extension Service, University of Florida, Institute of Food and Agricultural Sci-
ences, Gainesville, Florida.
Crane, J.H., Campbell, R.J. and Balerdi, C.F. (1993) Effect of Hurricane Andrew on tropical
fruit trees. Proceedings of the Florida State Horticultural Society 106, 139144.
Crane, J., Balerdi, C., Campbell, R., Campbell, C. and Goldweber, S. (1994) Managing
fruit orchards to minimize hurricane damage. HortTechnology 4, 2127.
Crane, J.H., Bally, I.S.E., Mosqueda-Vzquez, R.V. and Tomer, E. (1997) Crop produc-
tion. In: Litz, R.E. (ed.) The Mango: Botany, Production, and Uses. CAB Inter national,
Wallingford, UK, pp. 203256.
Crane, J.H., Schaffer, B. and Campbell, R.J. (2001) Recovery from hurricanes and the
long-term impacts on perennial tropical fruit crops in south Florida. HortScience
36, 16.
Crane, J., Schaffer, B., Li, Y., Evans, E., Montas, W. and Li, C. (2007) Effect of foliarly-
applied acids and ferrous sulfate on iron nutrition of avocado trees. In: Proceedings
of the VI Congreso Mundial Palta, Via del Mar, Chile, pp. 113.
Crane, J.H., Schaffer, B., Li, Y.C., Evans, E.A., Montas, W. and Li, C. (2008) Effect of
foliarly-applied acids and ferrous sulfate on leaf ferrous iron content and leaf
greenness of lychee trees. Proceedings of the Florida State Horticultural Society 121
(in press).
J.H. Crane et al. 476
Cruzaley-Sarabia, R., Ariza-Flores, R., Romero Gomezcaa, N.R., Noriega-Cant, D.H.
and Barrios-Ayala, A. (2006) Manual para Cultivar Mango en el Estado de Guerrero.
Libro Tcnico 3. Secretara de Agricultura, Ganadera, Desarrollo Rural, Pesca y
Alimentacin (SAGARPA)-Instituto Nacional de Investigaciones Forestales, Agrco-
las y Pecuarias (INIFAP), Campo Experimental Iguala, Iguala, Guerrero, Mxico,
96 pp.
Cull, B.W. (1991) Mango crop management. Acta Horticulturae 291, 154173.
Cunha, G.A.P. da and Castro Neto, M.T. (2000) Implantao do pomar. In: Manga
Produo, Aspectos Tcnicos. Embrapa Informaes Tecnolgicas, Braslia, Brazil,
pp. 2930.
Davenport, T. (1993) Floral manipulation in mangos. In: Proceedings of a Conference on
Mango in Hawaii. University of Hawaii at Manoa, College of Tropical Agriculture
and Human Resources, Honolulu, Hawaii, pp. 5460.
Davenport, T.L. (2006) Pruning strategies to maximize tropical mango production from
the time of planting to restoration of old orchards. HortScience 41, 544548.
Davenport, T.L. and Nez-Elisea, R. (1997) Reproductive physiology. In: Litz, R.E. (ed.)
The Mango: Botany, Production, and Uses, CAB International, Wallingford, UK,
pp. 69146.
Degani, C., Cohen, M., Reuveni, O., El-Batsri, R. and Gazit, S. (1993) Frequency and
characteristics of zygotic seedlings from polyembryonic mango cultivars deter-
mined using isozymes as genetic markers. Acta Horticulturae 341, 7885.
De los Santos, R.F. and Mosqueda-Vzquez, R. (198889) Comparacin de 21 cultiva-
res y 12 selecciones Mexicanas de mango Mangifera indica L. en la zona central
del estado de Veracruz. Revista Chapingo 6263, 6368.
Empacadoras de Mango de Exportacin (EMEX) (2007) Cierre de Temporada 2006. Em-
pacadoras de Mango de Exportacin (EMEX, Asociacin Civil). Zapopan, Jalisco,
Mexico.
Espenshade, E.W. Jr (ed.) (1992) Goodes World Atlas, 18th edn. Rand McNally, New
York.
Evans, E. (2007) Mangoes: Estimated Production Costs in the Miami-Dade County,
20032004. Commodity Budgets. Agricultural Economics Extension. Available at:
http://agecon-trec.ifas.u.edu/documents/mango0304.pdf (accessed 19 May
2008).
Florida Agricultural Statistics Service (FASS) (2003) Tropical Fruit, Production and Value,
2003. FASS, Orlando, Florida, pp. 12.
Garca, E. (1973) Modicaciones al Sistema de Clasicacin Climtica de Kppen.
Instituto de Geografa, Universidad Nacional Autonoma de Mexico, Mexico City.
Garczynski, C.J. (1995) National Weather Service, Station: 048892, 19511993.
National Weather Service, University of California, Riverside, California.
Gazeta Grupo de Comunicaes (2006) Anurio Brasileiro da Fruticultura. Gazeta Santa
Cruz, Rio Grande do Sul, Brazil.
Getz, R. (1979) Florida Daily Temperature Normals. Circular 464. Florida Cooperative
Extension Service, University of Florida, Institute of Food and Agricultural Sciences,
Gainesville, Florida.
Gomes, U., Tozetto, L.J. and Pinto, A.C. de Q. (2002) Censo Frutcola do Nordeste Brasil-
eiro: Situao da Cultura de Manga. Boletim Tcnico. CODEVASF, Braslia, Brazil,
p. 6.
Green, J., Crane, J.H., Li, Y., Sanford, R. and Rodriguez, O. (1999) Preliminary results on
the effectiveness of two organosilicone-based adjuvants plus iron to correct leaf
iron chlorosis of containerized carambola (Averrhoa carambola) trees. Proceedings
of the Florida State Horticultural Society 112, 176178.
Crop Production: Management 477
Guzmn-Estrada, C. (1991) Aplicacin foliar de agroqumicos para inducir la oracin
en Mangifera indica L. cv. Manila y Haden en el Sur de Sinaloa. Memoria del XXIV
Congreso Nacional de la Ciencia del Suelo, Pachuca, Hidalgo, Mexico.
Guzmn-Estrada, C. (2001) Evaluacin nutrimental en los huertos comerciales de man-
go en el sur de Sinaloa. Horticultura Mexicana 3, 178.
Guzmn-Estrada, C. (2004) Evaluacin y determinacin del abastecimiento nutrimental
en mango del sur de Sinaloa. Informes Anuales 2000, 20012003 y 20032004.
Campo Experimental Sur de Sinaloa, Instituto Nacional de Investigaciones Forestales,
Agrcolas y Pecuarias (INIFAP), Sur de Sinaloa, Mexico, 12 pp.
Guzmn-Estrada, C. (2006) Nutrient supply of mango in Southern Sinaloa, Mxico.
(Abstract). In: The Eighth International Mango Symposium, Sun City, Johannesburg,
South Africa, 510 February.
Guzmn-Estrada, C. and Vzquez-Valdivia, V. (2006) La poda del mango. In: Vzquez-
Valdivia, V. and Prez-Barraza, M.H. (eds) El Cultivo del Mango: Principios y Tec-
nologa de Produccin. Libro Tcnico No. 1. Instituto Nacional de Investigaciones
Forestales, Agrcolas y Pecuarias (INIFAP), Centro de Investigacin Regional Pac-
co Centro (CIRPAC), Campo Experimental Santiago Ixcuintla, Santiago Ixcuintla,
Mexico, pp. 87120.
Hamilton, R.A. (1993) Origin and classication of mango varieties in Hawaii. In: Pro-
ceedings of a Conference on Mango in Hawaii. University of Hawaii at Manoa,
College of Tropical Agriculture and Human Resources, Honolulu, pp. 2833.
Hamilton, R.A., Chia, C.L. and Evans, D.O. (1992) Mango Cultivars in Hawaii. Informa-
tion Text Series 042. Hawaii Cooperative Extension Service, Hawaii Institute of
Tropical Agriculture and Human Resources, University of Hawaii at Manoa, Hono-
lulu, Hawaii.
Henderson, W.G., Jr, Carter, L.J., Moore, A.L., Stein, R.A., Wettstein, C.A. and Yamataki,
H. (1984) Soil Survey of Lee County, Florida. United States Department of Agricul-
ture (USDA), Natural Resource Conservation Service, US Government Printing
Ofce, Washington, DC, pp. 1185.
Hwang, S.F., Chen, D.L., Hu, W.S. and Sh, S.H. (2004) Axillary panicle induction by
chemicals in mango tree (Mangifera indica L.). Acta Horticulturae 645, 177182.
Job, J.-R. (1989) Fertilizer application rates, soil fertilities, yields and qualities of man-
goes in Taiwan. Soil and fertilizer experiment report. Department of Agriculture and
Forest, Taiwan Province, Taichung, pp. 201229.
Johnson, G., Muirhead, I., Mayers, P. and Cooke, T. (1989) Diseases. In: Bagshaw, J., Brown,
B., Cooke, T., Cunningham, I., Johnson, G., Mayers, P. and Muirhead, I. (eds) Mango
Pests and Disorders. Department of Primary Industries, Brisbane, Australia, pp. 19.
Johnson, W.O. (1970) Minimum Temperatures in the Agricultural Areas of Peninsular
Florida, Summary of Seasons 19371967. Institute of Food and Agricultural Sci-
ences (IFAS) Publication No. 9. IFAS, University of Florida, Gainesville, Florida.
Koo, R.J.C. and Young, T.W. (1972) Effects of age and position on mineral composition of
mango leaves. Journal of the American Society for Horticultural Science 97, 792794.
Kostermans, A.J.G.H. and Bompard, J. (1993) The Mangoes, their Botany, Nomencla-
ture, Horticulture and Utilization. Academic Press, New York.
Krishnamurthi, S., Randhawa, G.S. and Sivaraman Nair, P.C. (1961) Growth studies in
the mango (Mangifera indica L.) under Delhi (sub-tropical) conditions. Indian Jour-
nal of Horticulture 18, 106118.
Kulkarni, V.J. (2004) The tri-factor hypothesis of owering in mango. Acta Horticulturae
645, 6170.
Lambe, A., Nez-Elisea, R. and Davenport, T. (1991) New developments in marcotting
mangos. Tropical Fruit World 1, 8082.
J.H. Crane et al. 478
Larson, K.D., Schaffer, B. and Davies, F.S. (1989a) Flooding, carbon assimilation and
growth of mango trees. (Abstract). In: Program and Abstracts of the American Soci-
ety for Horticultural Science, p. 126.
Larson, K.D., Schaffer, B. and Davies, F.S. (1989b) Effect of irrigation on leaf water
potential, growth, and yield of mango trees. Proceedings of the Florida State Horti-
culture Society 102, 226228.
Larson, K.D., Davies, F.S. and Schaffer, B. (1991a) Floodwater temperature and stem
lenticel hypertrophy in Mangifera indica (Anacardiaceae). American Journal of
Botany 78, 13971403.
Larson, K.D., Graetz, D.A. and Schaffer, B. (1991b) Flood-induced chemical transforma-
tions in calcareous agricultural soils of south Florida. Soil Science 152, 3340.
Larson, K.D., Schaffer, B. and Davies, F.S. (1991c) Flooding, leaf gas exchange, and
growth of mango in containers. Journal of the American Society for Horticultural
Science 116, 156160.
Larson, K.D., Schaffer, B. and Davies, F.S. (1991d) Mango responses to ooding in lime-
stone soil. Proceedings of the Florida State Horticultural Society 104, 3339.
Larson, K.D., Schaffer, B., Davies, F.S. and Sanchez, C.A. (1992) Flooding, mineral nutri-
tion, and gas exchange of mango trees. Scientia Horticulturae 52, 113124.
Larson, K.D., Schaffer, B. and Davies, F.S. (1993a) Floodwater oxygen content, ethylene
production and lenticel hypertrophy in ooded mango (Mangifera indica L.) trees.
Journal of Experimental Botany 22, 665671.
Larson, K.D., Schaffer, B. and Davies, F.S. (1993b) Physiological, morphological and
growth responses of mango trees to ooding. Acta Horticulturae 341, 152159.
Leonard, C.D. and Calvert, D.V. (1971) Field tests with new iron chelates on citrus
growing on calcareous soils. Proceedings of the Florida State Horticultural Society
84, 2431.
Leonard, C.D. and Stewart, I. (1953) Chelated iron as a corrective for lime-induced chlo-
rosis in citrus. Proceedings of the Florida State Horticultural Society 66, 4954.
Lima Filho, J.M.P. (2000) Determinao do potencial hdrico da mangueira utilizando-
se a cmara de presso. Anais do XVI Congresso Brasileiro de Fruticultura. (Resu-
mos em CD ROM). Embrapa Agroindstria Tropical, Fortaleza, Brazil.
Linden, T. (2006) Organic Mangos Now Coming out of California. The Produce News.
Available at: http://www.producenews.com/storydetail.cfm?ID=6216 (accessed 8
December 2007).
Litz, R.E. (1984) In vitro somatic embryogenisis from nucellar callus of monoembryonic
mango. HortScience 19, 715717.
Litz, R.E. (1986) Mango. In: Bajaj, Y.P.S. (ed.) Biotechnology in Agriculture and Forestry.
Vol. 1: Trees. Springer-Verlag, Berlin, pp. 267273.
Litz, R.E. (ed.) (2005) Biotechnology of Perennial Fruit and Nut Crops. CAB International,
Wallingford, UK.
Litz, R.E., Knight, R.J., Jr and Gazit, S. (1984) In vitro somatic embryogenesis from
Mangifera indica L. callus. Scientia Horticulturae 222, 233240.
Liu, M.-F. (1996) Mangoes. Extension Bulletin. Agriculture and Forestry Department,
Taiwan Provincial Government. Nantou, Taiwan.
Majumder, P.K. and Sharma, D.K. (1990) Mango. In: Bose, T.K. and Mitra, S.K. (eds)
Fruits: Tropical and Subtropical. Naya Prokash, Calcutta, India, pp. 162.
Majumder, P.K., Sharma, D.K., Singh, M.P. and Singh, R.N. (1976) Improve productivity
of malformed mango trees. Indian Horticulture 20, 78.
McKellar, M., Buchanan, D.W. and Campbell, C.W. (1983) Cold hardiness of two culti-
vars of avocado and a mango. Proceedings of the Florida State Horticultural Society
96, 212215.
Crop Production: Management 479
Mosqueda-Vzquez, R. and De los Santos, R.F. (1982) Aspersiones de nitrato de potasio
para adelantar e inducir la oracin del mango cv. Manila in Mexico. Proceedings
of the Tropical Region, American Society for Horticultural Science 25, 311331.
Mosqueda-Vzquez, R., Avila-Resndiz, C., Garca, P.E., De los Santos, R.F. and Ireta-
Ojeda, A. (1996) Esmeralda un Clon Para Usarse Como Interinjerto y Reducir el
Tamao del rbol de Mango Cultivar Manila. Technical Bulletin No. 12. Instituto
Nacional de Investigaciones Forestales, Agrcolas y Pecuarias (INIFAP)-Colegio de
Postgraduados, Veracruz, Mxico, 21 pp.
Mouco, M.A. do C., Albuquerque, J.A.S., Pinto, A.C. de Q., Castro Neto, M.T. de and
Barbosa, F.R. (2002) Implantao do pomar. In: Genu, P.J.C. and Pinto, A.C. de Q.
(eds) A Cultura da Mangueira. Embrapa Informao Tecnolgica, Braslia, Brazil,
pp. 138143.
Mukherjee, S.K. (1953) The mango its botany, cultivation, uses and future improve-
ment, especially as observed in India. Economic Botany 7, 130162.
Mukherjee, S.K. (1972) Origin of mango (Mangifera indica). Economic Botany 26, 260
264.
Nagao, M.A. and Nishina, M.S. (1993) Use of potassium nitrate on mango owering. In:
Proceedings of a Conference on Mango in Hawaii. University of Hawaii at Manoa,
College of Tropical Agriculture and Human Resources, Honolulu, Hawaii, pp. 61
66.
Nascimento, A.S. de, Carvalho, R. da S., Mendona, M. da C. and Sobrinho, R.B. (2002)
Pragas e seu controle In: Genu, P.J.C. and Pinto, A.C. de Q. (eds) A Cultura da
Mangueira. Embrapa Tecnolgica, Braslia, Brazil, pp. 277297.
National Agricultural Statistics Service (NASS)-Hawaii (2006) Hawaii Tropical Specialty
Fruits. NASS, United States Department of Agriculture (USDA), Hawaii Field Ofce
and Hawaii Department of Agriculture, Agricultural Development Division, Wash-
ington, DC, pp. 14.
Negi, S.S. (2000) Mango production in India. Acta Horticulturae 509, 6978.
Nguyen, H., Hofman, P., Holmes, R., Bally, I., Stubbings, B. and McConchie, R. (2004)
Effect of nitrogen on the skin colour and other quality attributes of ripe Kensington
Pride mango (Mangifera indica L.) fruit. Journal of Horticultural Science and Bio-
technology 79, 204210.
Noble, C.V., Drew, R.W. and Slabaugh, J.D. (1996) Soil Survey of Dade County Area,
Florida. United States Department of Agriculture (USDA) Natural Resource Conser-
vation Service, US Government Printing Ofce, Washington, DC, pp. 1116.
Nez-Elisea, R. (1986) Produccin Temprana de Mango Haden y Manila con Asper-
siones de Nitrato de Potasio. Folleto para productores No. 8. Centro de Investiga-
ciones Agrcolas Pacco Centro (CIAPAC), Instituto Nacional de Investigaciones
Forestales, Agrcolas y Pecuarias (INIFAP), Secretara de Agricultura y Recursos
Hidrulicos (SARH), Campo Agrcola Experimental Tecomn, Tecomn, Mexico.
Nez-Elisea, R. (1988) Nitrato de Amonio: Nueva Alternativa para Adelantar la Flo-
racin y Cosecha del Mango. Desplegable para productores No. 4. Secretara de
Agricultura y Recursos Hidrulicos (SARH)-Instituto Nacional de Investigaciones
Forestales, Agrcolas y Pecuarias (INIFAP), Campo Experimental Tecomn, Tecomn,
Colima, Mexico.
Nez-Elisea, R. and Davenport, T.L. (1992) Requirement for mature leaves during oral
induction and oral transition in developing shoots of mango. Acta Horticulturae
296, 3337.
Nez-Elisea, R. and Davenport, T.L. (1995) Effect of leaf age, duration of cool tempera-
ture treatment, and photoperiod on bud dormancy release and fruit initiation in
mango. Scientia Horticulturae 62, 6373.
J.H. Crane et al. 480
Nez-Elisea, R., Ferreira, W., Caldeira, M.L. and Davenport, T.L. (1989) Marcottage of
mango: a useful tool for physiological studies of ower induction. (Abstract). In:
Proceedings of the 86th Annual Meeting of the American Society for Hortcultural
Science, Ames, Iowa, p. 115.
Nez-Elisea, R., Davenport, T.L. and Caldiera, M.L. (1991) An experimental system to
study mango owering using containerized trees propagated by air-layering. Pro-
ceedings of the Florida State Horticultural Society 104, 3941.
Nez-Elisea, R., Ferreira, W., Caldeira, M.L. and Davenport, T.L. (1992) Adventitious
rooting of Tommy Atkins mango air layers induced with naphthaleneacetic acid.
HortScience 27, 926.
Oosthuyse, S.A. (1995) Pruning of mango trees: an update. South African Mango Grow-
ers Association Yearbook 15, 115.
Oosthuyse, S.A. and Jacobs, G. (1995) Relationship between branching frequency, and
growth, cropping and structural strength of 2-year-old mango trees. Scientia Horti-
culturae 64, 8593.
Pal, R.N. and Chadha, K.L. (1982) Deblossoming mangoes with cycloheximide. Journal
of Horticultural Science 57, 331332.
Perez, A. and Pollack, S. (2007) Fruit and Tree Nuts Outlook. FTS-327. United States
Department of Agriculture (USDA) Economic Research Service, Washington, DC,
pp. 1314.
Prez-Barraza, M.H., Salazar-Garca, S. and Vzquez-Valdivia, V. (2000) Delayed ino-
rescence bud initiation, a clue for the lack of response of the Tommy Atkins man-
go to promoters of owering. Acta Horticulturae 509, 567572.
Prez-Barraza, M.H., Vzquez-Valdivia, V. and Salazar-Garca, S. (2006a) Defoliacin
de brotes apicales y su efecto sobre la diferenciacin oral del mango Tommy At-
kins. Fitotecnia Mexicana 29, 313319.
Prez-Barraza, M.H., Vzquez-Valdivia, V. and Salazar-Garca, S. (2006b) Manipulacin
de la oracin y cosecha. In: Vzquez-Valdivia, V. and Prez-Barraza, M.H. (eds) El
Cultivo del Mango: Principios y Tecnologa de Produccin. Libro Tcnico No. 1.
Instituto Nacional de Investigaciones Forestales, Agrcolas y Pecuarias (INIFAP),
Centro de Investigacin Regional Pacco Centro (CIRPAC), Campo Experimental
Santiago Ixcuintla, Santiago Ixcuintla, Mexico, pp. 187209.
Pinto, A.C. de Q. (2000) A teortica no Cultivo da Manga: Sinopse. Embrapa Cerrados,
Planaltina, Brasilia, Brazil.
Pinto, A.C. de Q. (2004) Melhoramento gentico da manga (Mangifera indica L.) no
Brasil. In: Rozane, D.E., Darezzo, R.J., Aguiar, R.L., Aguilera, G.H.A. and Zam-
bolim, L. (eds) Manga, Produo Integrada, Industrilalizao e Comercializao.
UFV, Viosa, Brazil, pp. 1778.
Pinto, A.C. de Q. and Gen, P.J. de C. (1981) Inuncia do adubo orgnico e de se-
mentes sem endocarpo sobre a germinao e vigor de porta-enxertos de manguei-
ra. Pesquisa Agropecuria Brasileira 16, 111115.
Pinto, A.C. de Q. and Ramos, V.H.V. (1998) Formao do pomar. Guia Tcnico do
Produtor Rural 18, Embrapa Cerrados, Brasilia, Brazil, p. 2.
Pinto, A.C. de Q., Ramos, V.H.V., Junqueira, N.T.V., Lobato, E. and de Souza, D.M.
(1994) Relao Ca/N nas folhas e seu efeito na produo e qualidade da manga
Tommy Atkins sob condies de Cerrados. (Abstract). Congresso Brasileiro de Fru-
ticultura 13, Salvador, Bahia, Brazil, p. 763.
Ploetz, R.C. (1994) Mango disease caused by fungi. In: Ploetz, R.C., Zentmyer, G.A.,
Nishijima, W., Rohrbach, K. and Ohr, H.D. (eds) Compendium of Tropical Fruit
Diseases. American Phytopathological Society (APS) Press, St Paul, Minnesota,
pp. 3440.
Crop Production: Management 481
Ploetz, R.C. (2003) Diseases of mango. In: Ploetz, R.C. (ed.) Diseases of Tropical Fruit
Crops. CAB International, Wallingford, UK, pp. 327363.
Ponce-Gonzlez, F. and Salazar-Garca, S. (1992) Etiologa del cancro del mango
(Mangifera indica L.) cv. Manila en las Varas, Nayarit, Mxico. Revista Mexicana de
Fitopatologia 10, 153155.
Quayle, R.G., Cram, R.S. and Burgin, M.G. (1995) Climatic Averages and Extremes for
US Cities. Historical Climatology Series 6-3. US Department of Commerce, National
Oceanic and Atmospheric Administration (NOAA), Ashville, North Carolina,
pp. 7679.
Rajan, S. and Ram, S. (1988) Studies on the root regeneration in mango air-layers. Acta
Horticulturae 231, 192197.
Raymond, L., Schaffer, B., Brecht, J.K. and Hanlon, E.A. (1998) Internal breakdown,
mineral element concentration, and weight of mango fruit. Journal of Plant Nutri-
tion 21, 871889.
Rossetto, C.J., Ribeiro, I.J.A., Gallo, P.B., Soares, N.B., Bortoletto, N. and Paulo, E.M.
(1996) Mango breeding for resistance to diseases and pests. Acta Horticulturae
455, 299304.
Salazar-Garca, S. and Vzquez-Valdivia, V. (1997) Physiological persistence of
paclobutrazol on the Tommy Atkins mango (Mangifera indica L.) under rainfed
conditions. Journal of Horticultural Science 72, 339345.
Salazar-Garca, S., Gutirrez-Camacho, G., Becerra-Bernal, E. and Gmez-Aguilar, R.
(1993) Diagnstico nutricional del mango en San Blas, Nayarit. Revista Fitotecnia
Mexicana 16, 190202.
Sandoval-Esquivez, A., Hernndez, O.J., Montecillo, T.J.L. and Quilantan, C.J. (1993)
Manual de Produccin de Mango en la Costa de Chiapas. Publicacin especial No.
1. Campo Experimental Rosario Izapa, Centro de Investigacin Regional Pacco
Sur (CIRPS), Instituto Nacional de Investigaciones Forestales, Agrcolas y Pecuarias
(INIFAP), Secretara de Agricultura y Recursos Hidrulicos (SARH), Tapachula,
Chiapas, Mexico.
Santos Filho, H.P., Tavares, S.C.C. de H., Matos, A.P., Costa, V.S. de O., Moreira, W.A.
and Santos, C.C.F dos (2002) Doenas, monitoramento e controle. In: Genu, P.J.C.
and Pinto, A.C. de Q. (eds) A Cultura da Mangueira. Embrapa Informao
Tecnolgica, Braslia, Brazil, pp. 300352.
Sauls, J.W. and Campbell, C.W. (1980) Mango Propagation. Fact Sheet FC-58. Florida
Cooperative Extension Service, University of Florida, Institute of Food and Agricul-
tural Sciences, Gainesville, Florida.
Schacht, H. (1992) Desert green. California Farmer 275, 1011.
Schaffer, B., Whiley, A.W. and Crane, J.H. (1994) Mango. In: Schaffer, B. and Andersen, P.C.
(eds) Handbook of Environmental Physiology of Fruit Crops, Vol. II, Sub-tropical and
Tropical Crops. CRC Press, Boca Raton, Florida, pp. 165197.
Schaffer, B., Davies, F.S. and Crane, J.H. (2006) Responses of subtropical and tropical
fruit trees to ooding in calcareous soil. HortScience 41, 549555.
Schnell, R.J. and Knight, R.J., Jr (1991) Are polyembryonic mangos dependable sources
of nucellar seedlings for rootstocks? Proceedings of the Florida State Horticultural
Society 104, 4447.
Schnell, R.J. and Knight, R.J., Jr (1992) Frequency of zygotic seedlings from ve polyem-
bryonic mango rootstocks. HortScience 27, 174176.
Schnell, R.J., Knight, R.J., Jr and Harkins, D.M. (1994) Eliminating zygotic seedlings in
Turpentine mango rootstock populations by visual roguing. HortScience 29,
319320.
Scott, L. (1990) A Tropical Oasis. The Press-Enterprise, Riverside County, California.
J.H. Crane et al. 482
Shen, T.-M. and Huang, P.-C. (1980) Effect of chemical treatments and inorescence
pinching on the regulation of owering date and fruit setting of mango tree
(Mangifera indica L.). Journal of the Chinese Society for Horticultural Science 26,
6170.
Sh, Z.H. (1993) Chemical pruning and induction of panicles in mango. Acta Horticul-
turae 341, 199205.
Sh, Z.H. and Sheen, T.F. (1987) Floral induction in axillary buds of mango (Mangifera
indica L.) as affected by temperature. Scientia Horticulturae 31, 8187.
Sh, Z.H., Yen, C.R., Ke, L.S., Lin, T.S., Shiesh, C.C., Wang, D.N. and Liu, M.F. (2000)
Mango production in Taiwan. Acta Horticulturae 509, 8794.
Silva, D.J., Quaggio, J.A., Pinto, P.A. da C., Pinto, A.C. de Q. and Magalhes, A.F. de J.
(2002) Nutrio e adubao In: Genu, P.J.C. and Pinto, A.C. de Q. (eds) A Cultura
da Mangueira. Embrapa Informao Tecnolgica, Braslia, Brazil, pp. 192221.
Singh, R.N., Majumder, P.K., Sharma, D.K., Sinha, G.C. and Bose, P.C. (1974) Effect of
de-blossoming on the productivity of mango. Scientia Horticulturae 2, 399403.
Singh, L.B. (1960) The Mango: Botany, Cultivation and Utilization. Leonard Hill, Lon-
don.
Singh, R.N. (1978) Mango. Indian Council for Agricultural Research, New Delhi, India.
Sistema Integral de Informacin Agroalimentaria y Pesquera (SIIAP) (2007) Anuario Es-
tadstico de la Produccin Agrcola. SAGARPA-Mxico. Available at: http://www.
oeidrus-portal.gob.mx/aagricola_siap/icultivo/index.jsp (accessed 10 September
2007).
Smith, P.F. and Scudder, G.K., Jr (1951) Some studies of mineral deciency symptoms in
mango. Proceedings of the Florida State Horticultural Society 64, 243248.
Sousa, D.M.G. de, Lobato, E. and Rein, T.A. (2004) Adubao com fsforo. In: Sousa,
D.MG. and Lobato, E. (eds) Cerrado, Correo do Solo e Adubao. Embrapa Infor-
mao Tecnolgica, Brasilia, Brazil, pp. 147168.
Southeast Regional Climate Center (SERCC) (2007) Ponce 4E, Puerto Rico: Period of
Record General Climate Summary Precipitation. Available at: http://radar.meas.
ncsu.edu/cgi-bin/sercc/cliMAIN.pl?pr7292 (accessed 22 August 2007).
Souza, J. da S., Almeida, C.O., Arajo, J.L.P. and Cardoso, C.E.L. (2002) Aspectos scio-
econmicos. In: Genu, P.J.C. and Pinto, A.C. de Q. (eds) A Cultura da Mangueira.
Embrapa Informao Tecnolgica, Braslia, Brazil, pp. 2029.
Stassen, P.J.C., Grove, H.G. and Davie, S.J. (1999) Tree shaping strategies for higher
density mango orchards. Journal of Applied Horticulture 1, 14.
Toro, E.E. (1988) Cultivo de Mangos en Puerto Rico. H-125. Universidad de Puerto Rico,
Recinto de Mayaguez, Colegio de Ciencias Agricolas, Mayaguez, Puerto Rico,
pp. 195.
Vzquez-Valdivia, V., Salazar-Garca, S. and Prez-Barraza, M.H. (2000) Esmeralda in-
terstocks reduce Ataulfo mango tree size with no reduction in yield: results of the
rst ve years. Acta Horticulturae 509, 291296.
Whiley, A.W. (1993) Environmental effects on phenology and physiology of mango a
review. Acta Horticulturae 341, 168176.
Whiley, A.W., Rasmussen, T.S., Saranah, J.B. and Wolstenholme, B.N. (1989) Effect of
temperature on growth, dry matter production and starch accumulation in ten man-
go (Mangifera indica L.) cultivars. Journal of Horticultural Science 64, 753765.
Young, T.W. (1974) Fertilizing Mango Trees in Florida. Report SUB 68-1. University of
Florida, Institute of Food and Agricultural Sciences, Agricultural Research and Edu-
cation Center, Homestead, Florida.
Young, T.W. and Koo, R.C.J. (1969) Mineral composition of Florida mango leaves. Pro-
ceedings of the Florida State Horticulture Society 82, 324328.
Crop Production: Management 483
Young, T.W. and Koo, R.C.J. (1971) Variations in mineral content of Florida mango
leaves. Proceedings of the Florida State Horticulture Society 84, 298303.
Young, T.W. and Koo, R.C.J. (1974) Increasing yield of Parvin and Kent mango on
Lakewood sand by increased nitrogen and potassium fertilization. Proceedings of
the Florida State Horticulture Society 87, 380384.
Young, T.W. and Miner, J.T. (1960) Response of Kent mangos to nitrogen fertilization.
Proceedings of the Florida State Horticulture Society 73, 334336.
Young, T.W. and Miner, J.T. (1961) Relationship of nitrogen and calcium to soft-nose
disorder in mango fruits. Proceedings of the Florida State Horticulture Society 78,
201208.
Young, T.W. and Sauls, J.W. (1989) The Mango Industry of Florida. Bulletin 189. Florida
Cooperative Extension Service, University of Florida, Institute of Food and Agricul-
tural Sciences, Gainesville, Florida.
Young, T.W., Koo, R.C.J. and Miner, J.T. (1962) Effects of nitrogen, potassium and calcium
fertilization on Kent mangos on deep, acid, sandy soil. Proceedings of the Florida
State Horticulture Society 75, 364371.
Young, T.W., Koo, R.C.J. and Miner, J.T. (1965) Fertilizer trials with Kent mangos.
Proceedings of the Florida State Horticulture Society 78, 369375.
Zude-Sasse, M. and Schaffer, B. (2000) Inuence of soil oxygen depletion on iron uptake
and reduction in mango (Mangifera indica L.) root. Proceedings of the Florida State
Horticultural Society 113, 14.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
484 (ed. R.E. Litz)
14 Postharvest Physiology
J.K. Brecht
1
and E.M. Yahia
2
1
University of Florida, Florida, USA
2
Universidad Autnoma de Quertaro, Quertaro, Mexico
14.1 Introduction 484
14.2 Contribution of Mango Fruit to Human Nutrition and Health 485
14.3 Mango Ripening Physiology 491
Climacteric behaviour 491
Ethylene production and responses 493
14.4 Compositional Changes during Fruit Maturation and Ripening 494
Organic acids 494
Soluble sugars 495
Structural polysaccharides 496
Pigments and colour 498
Phenolic compounds 502
Flavour (taste, aroma) 504
14.5 Transpiration and Water Loss 507
14.6 Physical Damage and Physiological Disorders 507
Chilling injury (CI) 507
Heat injury 508
14.7 Modied Atmospheres (MA) and Controlled Atmospheres (CA) 508
Injuries associated with MA and CA 510
Modied atmosphere packaging (MAP) 511
Semipermeable coatings 513
Insecticidal CA 514
14.8 Manipulation of Mango Postharvest Physiology by Molecular Biology 515
14.9 Conclusions 516
14.1 Introduction
Successful postharvest handling of mangoes requires knowledge of the post-
harvest physiology of the fruit and how the fruit physiology determines the
best handling practices to maintain and develop high fruit quality. For example,
mango, like banana, tomato and avocado, is a climacteric fruit, which means
Postharvest Physiology 485
that it may be picked when mature but before ripening has commenced, and
subsequently ripened postharvest. As mango fruit mature on the tree and
begin to ripen, eating quality improves, but potential marketable life de-
creases due to the difculty of controlling the ripening changes once they
have been initiated, increased bruising susceptibility and increased decay.
Susceptible mango cultivars tend to develop more internal breakdown (jelly
seed, soft nose and stem-end cavity) the longer that harvesting is delayed
(Raymond et al., 1998; see Galn Saco, Chapter 9, this volume). As a tropical
species, mangoes are subject to chilling injury (CI), which limits the use of
refrigeration to maintain postharvest quality. Mangoes are also subject to
other physiological disorders, physical damage and decay, the symptoms of
which may make the fruit unmarketable (Yahia et al., 2006a).
Mangoes harvested at a mature but unripe stage of development (mature-
green) can be stored in the unripe state as long as the initiation of ethylene
production and hence ripening is avoided. The initiation of ripening can be
avoided by prompt cooling and storage at a low temperature at which ripen-
ing does not occur or, more effectively, by changing the composition of the
storage atmosphere so that the oxygen (O
2
) level is reduced and carbon diox-
ide (CO
2
) level is raised. This latter approach is called either modied atmo-
sphere (MA) or controlled atmosphere (CA) storage, depending on the degree
of control. These technologies slow fruit metabolism and specically inhibit
the initiation of ethylene production. With MA or CA transport or storage,
mangoes can typically be maintained in a rm, green condition for several
days longer than can be achieved with normal refrigerated air storage. How-
ever, there are limits to the levels of O
2
and CO
2
that can be tolerated by
mangoes and these limits are affected by several factors, including cultivar,
maturity or ripeness stage, storage temperature and storage time (Yahia,
1998).
Mango postharvest physiology and technology have been described in
previous reports, book chapters and reviews (Subramanyam et al., 1975; Lak-
shminarayana, 1980; Ledger, 1986; Peacock, 1986; Lizada, 1991; Coates and
Johnson, 1993; Johnson and Coates, 1993; Lizada, 1993; Heather, 1994; Jacobi
et al., 1994; Johnson et al., 1997; Mitra and Baldwin, 1997; Tharanathan et al.,
2006).
14.2 Contribution of Mango Fruit to Human Nutrition and Health
Consumers are becoming aware of the nutritional and health benets of fresh
fruits and vegetables. Mango fruit are a rich source of vitamin C (Table 14.1),
although the content decreases during ripening (Thomas, 1975; Vinci et al.,
1995). Raspuri mango is rich in vitamin C (300 mg/100 g fresh fruit) during
the early stages of development, but the concentration is less (39.169.5
mg/100 g) at maturity (Siddappa and Bhatia, 1954). The content of vitamin C
was between 13 and 178 mg/100 g in the ripe fruit of 50 cultivars surveyed
by Singh (1960). The vitamin C content in fully grown mango fruit of culti-
vars in Puerto Rico ranged between 6 and 63 mg/100 g (Iguina de George
J.K. Brecht and E.M. Yahia 486
Table 14.1. Composition of the edible portion of mango fruit (Source: USDA/ARS,
2007).
Nutrient Unit Value per 100 g edible portion
Water g 81.71
Energy kcal 65
Energy kJ 272
Protein g 0.51
Total lipid (fat) g 0.27
Ash g 0.50
Carbohydrate, by difference g 17.00
Fibre, total dietary g 1.8
Sugars, total g 14.80
Minerals
Calcium mg 10
Iron mg 0.13
Magnesium mg 9
Phosphorus mg 11
Potassium mg 156
Sodium mg 2
Zinc mg 0.04
Copper mg 0.110
Manganese mg 0.027
Selenium g 0.6
Vitamins
Vitamin C (total ascorbic acid) mg 27.7
Thiamine mg 0.058
Riboavin mg 0.057
Niacin mg 0.584
Pantothenic acid mg 0.160
Vitamin B
6
mg 0.134
Folate, total g 14
Folic acid g 0
Folate, food g 14
Vitamin B
12
g 0.00
Vitamin A IU 765
Retinol g 0
Vitamin E (-tocopherol) mg 1.12
Vitamin K (phylloquinone) g 4.2
Lipids
Fatty acids, total saturated g 0.066
4:0 g 0.000
6:0 g 0.000
8:0 g 0.000
10:0 g 0.000
12:0 g 0.001
14:0 g 0.009
16:0 g 0.052
18:0 g 0.003
Fatty acids, total monounsaturated g 0.101
(Continued)
Postharvest Physiology 487
et al., 1969). Vitamin C content was 105.2, 65.7 and 17.3 mg/100 g in Langra,
Ashwini and Fazli mangoes, respectively (Gofur et al., 1994), and decreased
rapidly 57 weeks after fruit set, and when ripe fruit were stored at room
temperature. Vitamin B
1
(thiamine) in two mango cultivars was 3560 g/100 g,
Table 14.1. Continued
Nutrient Unit Value per 100 g edible portion
16:1 undifferentiated g 0.048
18:1 undifferentiated g 0.054
20:1 g 0.000
22:1 undifferentiated g 0.000
Fatty acids, total polyunsaturated g 0.051
18:2 undifferentiated g 0.014
18:3 undifferentiated g 0.037
18:4 g 0.000
20:4 undifferentiated g 0.000
20:5 n-3 g 0.000
22:5 n-3 g 0.000
22:6 n-3 g 0.000
Cholesterol g 0
Amino acids
Tryptophan g 0.008
Threonine g 0.019
Isoleucine g 0.018
Leucine g 0.031
Lysine g 0.041
Methionine g 0.005
Phenylalanine g 0.017
Tyrosine g 0.010
Valine g 0.026
Arginine g 0.019
Histidine g 0.012
Alanine g 0.051
Aspartic acid g 0.042
Glutamic acid g 0.060
Glycine g 0.021
Proline g 0.018
Serine g 0.022
Other
Ethanol g 0.0
Caffeine mg 0
Theobromine mg 0
-Carotene g 445
-Carotene g 17
-Cryptoxanthin g 11
Lycopene g 0
Lutein + zeaxanthin g 0
J.K. Brecht and E.M. Yahia 488
and vitamin B
2
(riboavin) in three cultivars was 4555 g/100 g

(Stahl,
1935). Thiamine content of four Philippine cultivars was 57600 g/100 g,
and riboavin content of three cultivars was 37730 g/100 g (Quinones
et al., 1944). Folic acid in green mangoes was 36 mg/100 g (Gosh, 1960).
The mango fruit is a rich source of carotenoids, some of which function
as provitamin A: -carotene (all-trans), -cryptoxthanin (all-trans and cis),
zeaxanthin (all-trans), luteoxanthin isomers, violaxanthin (all-trans and cis)
and neoxanthin (all-trans and cis) (Mercadante et al., 1997; Yahia et al., 2006b;
Ornelas-Paz et al., 2007, 2008). Total carotenoid content rose from 12.3 to 38.0
g/g in Keitt and from 17.0 to 51.2 g/g in Tommy Atkins from the mature-
green to the ripe stage (Mercadante and Rodriguez-Amaya, 1998), and ripen-
ing alterations occurred principally in the major carotenoids, violaxantin
and -carotene. With Keitt, all-trans--carotene, all-trans-violaxanthin and
9-cis-violaxanthin increased from 1.7, 5.4 and 1.7 g/g, respectively, in the
mature-green fruit to 6.7, 18.0 and 7.2 g/g in the ripe fruit (Mercadante and
Rodriguez-Amaya, 1998). In Tommy Atkins these carotenoids increased
from 2.0, 6.9 and 3.3 g/g to 5.8, 22.4 and 14.5 g/g, respectively, during
ripening. Geographic effects were reported to be substantial (Mercadante
and Rodriguez-Amaya, 1998). Some of the cis and trans isomers of provitamin
A reported in Haden and Tommy Atkins mangoes include 13-cis--carotene
(trace amounts), trans--carotene (12.515.5 g/g) and trans--cryptoxanthin
(0.30.4 g/g) (Godoy and Rodriguez-Amaya, 1994). In processed mango
juice, violaxanthin was not detected, auroxanthin appeared at an appre-
ciable level, and -carotene was the principal carotenoid (Mercadante and
Rodriguez-Amaya, 1998). The major carotenoid in Bourbon, Haden,
Extreme, Golden and Tommy Atkins mangoes is -carotene (4884% of
the total), while epoxycarotenoids (violaxanthin, luteoxanthin and mutatox-
anthin) constitute 1349% of the total (Godoy and Rodriguez-Amaya, 1989).
Mean vitamin A in these mangoes (retinol equivalents/100 g) ranges from
115.3 (Haden) to 430.5 (Extreme).
Children in Senegal with normal cytology had higher serum retinol and
-carotene levels than those with abnormal cytology after massive oral doses
of vitamin A and consumption of mangoes (Carlier et al., 1992). Mango retinol
is highly bioavailable (82% efciency) by estimating vitamin A and carotene
reserves in the liver and plasma of rats (Yuyama et al., 1991). During mango
fruit ripening, vitamin A increases ripe mangoes are tenfold richer in caro-
tene than partially ripe fruit, while unripe green mangoes contain only trace
amounts (Modi and Reddy, 1967). Mevalonic acid, a precursor of carotenoids,
increases progressively during mango ripening (Modi and Reddy, 1967).
Vitamin A equivalents in 100 g of mango fruit are 1000 to 6000 IU (Singh,
1960). The -carotene content of the fruit of 30 mango cultivars in Puerto Rico
ranged from 400 to 800 IU/100 g fresh fruit (Iguina de George et al., 1969).
The development of -carotene in mangoes held at 1621C was lower than
that at 2028C (Vazquez-Salinas and Lakshminarayana, 1985). Jungalwala
and Cama (1963) identied 16 different carotenoids in Alphonso mangoes, and
-carotene accounted for 60% of the total. Of the oxycarotenoids, luteoxan-
thin, violaxanthin and cis-violaxanthin were present in signicant amounts.
Postharvest Physiology 489
All the oxycarotenoids were present as -carotene derivatives, mostly as
epoxides of zeaxanthin. Variation in carotenoid content, as in many other
constituents, is due to several factors, including cultivar, geography, climate,
storage/processing conditions and analytical procedures employed.
Several carotenoids occur in fruit of different mango cultivars (Cano and
de Ancos, 1994; Ben-Amotz and Fishler, 1998; Chen et al., 2004), but only a
few of them occur in signicant concentrations (Ornelas-Paz et al., 2007).
Mercadante et al. (1997) quantied several carotenoids in Keitt mangoes;
the most predominant ones were all-trans--carotene, all-trans-violaxanthin
and 9-cis-violaxanthin, accounting for 27, 38 and 18% of the total carotenoid
content, respectively. Similar ndings have been reported for crude extracts
from other mango cultivars (Mercadante and Rodrguez-Amaya, 1998; Pott
et al., 2003a, b). Carotenoids are responsible for the yellow-orange colour of
mango mesocarp (Vzquez-Caicedo et al., 2004). All-trans--carotene and the
dibutyrates of all-trans-violaxanthin and 9-cis-violaxanthin are the main car-
otenoids in Ataulfo and Manila mangoes (Yahia et al., 2006b; Ornelas-Paz
et al., 2008; Fig.14.1). The content of these carotenoids during fruit ripening
increased exponentially in Ataulfo and exponentially or in a second order
polynomial manner in Manila, and the highest correlation coefcients were
obtained for the relationships between the internal and external a* and h
colour values and the content of the evaluated carotenoids in both mango
Cultivar
Haden Ataulfo Tommy
Atkins
Manila Criollo Kent Paraso
P
u
l
p

c
a
r
o
t
e
n
o
i
d

c
o
n
t
e
n
t

(
m
g
/
1
0
0

g
)
0.0
0.5
1.0
1.5
2.0
2.5
3.0
3.5
All-trans-violaxanthin
9-cis-Violaxanthin
All-trans--carotene
Fig. 14.1. Content of selected carotenoids in pulp of several mango cultivars. Data
represent the mean of eight individual observations for each cultivar standard error
(Source: Ornelas-Paz et al., 2007).
J.K. Brecht and E.M. Yahia 490
cultivars (R = 0.810.94). Equations to predict the content of the most impor-
tant carotenoids in Manila and Ataulfo mangoes on the basis of their inter-
nal and external colour values were obtained by Ornelas-Paz et al. (2008).
The content of -tocopherol is approx. 0.5 mg/100 g in an unidentied
cultivar from Costa Rica (Burns et al., 2003), while the United States Depart-
ment of Agriculture (USDA) Nutrient Database (USDA/ARS, 2007) indicates
an -tocopherol content of 1.12 mg/100 g. Ornelas-Paz et al. (2007) found that
-tocopherol is the only detectable tocopherol in seven mango cultivars
(Fig. 14.2); Haden and Tommy Atkins mangoes had the highest amounts (380
and 470 g/100 g, respectively), with c.200250 g/100 g in the other cultivars.
Mango fruit are rich in several types of antioxidant phytochemicals, that
is carotenoids and phenolics (Ornelas-Paz et al., 2007; Rocha-Ribeiro et al.,
2007). Botting et al. (1999), showed that mango fruit have antimutagens and
the heterocyclic amine 2-amino-3-methylimidazo[4,5-f]quinoline. Percival
et al. (2006) observed that whole mango juice inhibited cell proliferation in
the leukaemic cell line HL-60 and also inhibited the neoplastic transforma-
tion of BALB/3T3 cells. Garca-Sols et al. (2008) studied the effect of Ataulfo
mango consumption on chemically induced mammary carcinogenesis and
plasma antioxidant capacity in rats treated with N-methyl-N-nitrosourea
(MNU). Mango was administered in the drinking water (0.020.06 g/ml)
during both short-term and long-term (LT) periods to rats treated or not with
Cultivar
Haden Ataulfo Tommy
Atkins
Manila Criollo Kent Paraso

-
T
o
c
o
p
h
e
r
o
l

(

g
/
1
0
0

g
)
0
100
200
300
400
500
600
Fig. 14.2. The content of -tocopherol in the pulp of several mango cultivars. Data
represent the mean of eight individual observations for each cultivar standard error
(Source: Ornelas-Paz et al., 2008).
Postharvest Physiology 491
MNU. Rats treated with MNU showed no differences in mammary carcino-
genesis or in plasma antioxidant capacity measured by both ferric reducing/
antioxidant power (FRAP) and total oxyradical scavenging capacity assays.
However, in animals not treated with MNU, but with an LT intake of mango,
the plasma antioxidant capacity as measured by the FRAP assay tended to
increase in a dose-dependent manner. This suggests that mango consump-
tion by healthy subjects may increase antioxidants in plasma.
14.3 Mango Ripening Physiology
Ripening is part of the natural senescence of mango fruit. It is an irreversible
process that contributes to organelle disruption and changes in chemical con-
stituents, avour and texture. While ripening improves the eating quality of
mango fruit, the postharvest life of the fruit is reduced. Natural senescence,
and thus ripening, is aggravated and promoted by ethylene, mechanical
injury and high temperature. This process can be delayed by lower tempera-
ture, elimination of mechanical damage and reducing ethylene production
(Yahia et al., 2006a). Ripening of mango is inhibited while fruit are attached
to the tree, and respiration and ripening are stimulated upon detachment
(Lakshminarayana, 1973). Burg and Burg (1962) reported that ethylene levels
in the tissues of mature-green, attached mango fruit were relatively high
(1.87 l/l) and suggested that ethylene was ineffective for promoting ripen-
ing due to a ripening inhibitor supplied by the tree.
Changes associated with mango fruit ripening include: (i) esh colour
from greenish yellow to yellow to orange in all cultivars (Plate 80a); (ii) skin
colour from green to yellow in some cultivars (Plate 80b); (iii) chlorophyll
decreases and carotenoid content increases; (iv) esh rmness decreases and
juiciness increases; (v) starch is converted into sugars; (vi) total soluble solids
(TSS) content increases; (vii) titratable acidity decreases; (viii) characteristic
aroma volatiles increase; (ix) CO
2
production rate increases from 4050 to
160200 mg/kg/h at 20C; and (x) ethylene production rate increases from
0.10.2 to 13 l/kg/h at 20C. Gowda and Huddar (2000) found the changes
in eight mango selections during ripening included reductions in fruit
weight, volume, length, thickness, rmness, pulp content, pulp:peel ratio,
starch and vitamin C, and increases in TSS, pH, total sugars, sugar:acid ratio,
pulp carotenoid content and peel colour.
Climacteric behaviour
Mango is a climacteric fruit, exhibiting a climacteric pattern of respiration
and an increase in ethylene production during ripening (Cua and Lizada,
1990; Reddy and Srivastava, 1999; Lalel et al., 2003; Fig. 14.3). The initiation of
ethylene production within the fruit triggers and coordinates the changes
that occur during ripening. These changes include colour changes in the peel
and esh, softening of the esh, and development of sweet avour and
J.K. Brecht and E.M. Yahia 492
aroma. Mangoes can be ripened after harvest when picked at physiological
maturity (mature-green), when they are fully sized, but before ripening has
been initiated. Maturity indices are chosen to predict fruit quality potential
and postharvest behaviour (Peacock et al., 1986; Medlicott et al., 1988). After
harvest, the fruit is then cooled and isolated from possible sources of ethyl-
ene (ripening fruit, engine exhaust, smoke, etc.) during storage or shipping.
This is the primary strategy used to control ripening and thus extend shelf
life. Respiration patterns and ripening behaviour vary among cultivars, with
different climatic conditions and growing locations (Krishnamurthy and
Subramanyam, 1970). Respiration is very high after fruit set and then declines
and is maintained at a low rate until fruit ripening begins.
The rise in respiration and ethylene production during the climacteric is
related to fruit ripening. The respiratory peak in Alphonso mangoes har-
vested mature-green occurs 5 days after harvest, and the fruit ripens within
7 or 8 days (Karmarkar and Joshi, 1941), while in Kent and Haden mangoes
the peak occurs on days 9 and 11, respectively (Burg and Burg, 1962), and in
Pairi mangoes on day 9 (Krishnamurthy and Subramanyam, 1970). These dif-
ferences are normal due to differences in location, climatic conditions, orchard
and tree conditions, and postharvest temperature. The rise in the climacteric
respiration in Dashehari, Amrapali and Rataul mangoes coincides with the
highest level of sucrose and polygalacturonase (PG; EC 3.2.1.15) activity in
ripening fruit (Kalra and Tandon, 1983). Respiration and ethylene production
are excellent maturity indices, but require considerable expense to measure.
The expression of alternative oxidase (Aox) and uncoupling proteins
(Ucp) has been investigated during mango ripening and compared with the
expression of peroxisomal thiolase (EC 2.3.1.16), a ripening marker in mango
(Considine et al., 2001). The multigene family for Aox in mango is expressed
differentially during mango fruit ripening. Abundance of Aox message and
protein peaks at the ripe stage, while expression of the single gene for the
Ucp peaks at the turning or half-ripe stage, and the protein abundance peaks
at the ripe stage. Proteins of the cytochrome chain peak at the mature-green
0
0
1
2
3
4
5
6
1 2 3
Hard green Sprung green
Ripe Half ripe
Ripening period (days)
C
O
2

(
m
m
o
l
/
k
g
/
h
)
E
t
h
y
l
e
n
e

(
m
m
o
l
/
k
g
/
h
)
4 5 6 7 8 9 10 0
0
10
20
30
40
50
1 2 3
Ripening period (days)
4 5 6 7 8 9 10
Hard green
Sprung green
Ripe
Half ripe
Fig. 14.3. The climacteric pattern of respiration and ethylene production during mango fruit
ripening (Source: Lalel et al., 2003).
Postharvest Physiology 493
stage, suggesting that increases in cytochrome chain components are impor-
tant for facilitating the climacteric burst of respiration and that Aox and Ucp
are important in postclimacteric senescence processes (Considine et al., 2001).
Because both message and protein for the Aox and Ucp increase in a similar
pattern, their expression is not controlled in a reciprocal manner but may be
active simultaneously.
Fruit slicing affects respiration rate (Allong et al., 2001). Slicing of mature-
green Julia and Graham mangoes increased respiration rate immediately
after cutting, but it decreased signicantly within the rst 12 h of storage at 5
or 10C, yet still remained at levels above that of the intact fruit throughout the
storage period. The effect of slicing on half-ripe and rm-ripe fruit is an initial
increase in respiration followed by a decline to levels of the intact fruit.
Ethylene production and responses
Mangoes have a moderate ethylene production peak of 13 l/kg/h during
ripening at 20C. Ethylene, applied directly or as ethrel, induces faster and
more uniform fruit softening (Lakshminarayana, 1973; Barmore, 1974; Lak-
shminarayana et al., 1974; Sornsrivichai and Waru-Aswapti, 1989). Ethylene
treatment can be prior to shipping (Barmore and Mitchell, 1975). There is
disagreement regarding the effect of ethylene treatment on quality (Chaplin,
1988), and this may be related to maturity when treated. Treatment of imma-
ture fruit leads to softening, but the fruit have poor avour.
Mango fruit ripening is accompanied by increased ethylene production,
which coordinates the ripening process. Mango expresses an autocatalytic
increase in ethylene production during ripening (Mattoo and Modi, 1969b).
Ethylene production starts before full ripeness is reached (Burg and Burg,
1962; Cua and Lizada, 1990). Ethylene production in unripe mango fruit is
very low (<0.1 l/kg/h) (Burdon et al., 1996). Ethylene production decreases
as the fruit matures, is then undetectable for a time and reappears upon ini-
tiation of ripening (Akamine and Goo, 1973). Kent and Haden mango fruit
have internal ethylene concentrations of c.0.01 l/l during the preclimacteric
phase, increasing to c.0.08 l/l at the initiation of the climacteric, and up to
3.0 l/l at the climacteric peak. Burg and Burg (1962) reported that ethylene
production rises when or before CO
2
production rises in ripening mangoes,
while Biale and Young (1981) included mangoes among fruits in which eth-
ylene rises after CO
2
production rises.
Only a small concentration of exogenous ethylene (0.005 l/l) is needed
to initiate mango ripening (Wills et al., 2001). The small amount of ethylene
in the fruit at harvest is sufcient to initiate ripening (Burg and Burg, 1962).
While fruit of Amrapali and Deshehari mangoes produce a measurable
amount of ethylene during ripening (Reddy and Srivastava, 1999), ethylene
production does not follow a climacteric pattern and two ethylene peaks (at
the mature-green and full-ripe stages) were recorded. This is probably due to
the way that ethylene was measured in the different fruit, and the lack of
control exerted on maturity stages of fruit. In Carabao mangoes, the peak of
J.K. Brecht and E.M. Yahia 494
ethylene production occurs 110 days after ower initiation, and declines
as fruit approached full maturity (Cua and Lizada, 1990). The content of
1-aminocyclopropane-1-carboxylic acid (ACC), the immediate precursor of
ethylene, increases in different tissues (peel, outer and inner mesocarp) dur-
ing ripening in both cultivars, Amrapali and Deshehari, while ACC oxi-
dase (ACO; EC 1.14.17.4), which catalyse the conversion of ACC to ethylene
and ethylene production, decline (Reddy and Srivastava, 1999). Fruit peel
has the highest levels of ethylene and ACO and less ACC accumulation than
the outer and inner mesocarp at the mature-green stage. The inner mesocarp
has less ACO activity and high ACC accumulation during the ripening pro-
cess compared to peel; levels in the outer mesocarp are intermediate between
those in the peel and inner mesocarp. Changes in the ability to convert ACC
to ethylene in the peel are not related to changes in ripening parameters in
the fruit pulp (Lederman et al., 1997). Mango seed also produces ethylene
(Reddy and Srivastava, 1999). Fruit slicing has no measurable effect on ethyl-
ene production in Julia and Graham mangoes (Allong et al., 2001).
Treatment of mango fruit with acetaldehyde or ethanol (0.1, 0.5 or 1%
ethanol or acetaldehyde vapour) has concentration-dependent inhibitory
effects on ethylene production (Burdon et al., 1996). Application of ACC to
acetaldehyde- or ethanol-treated fruit discs indicates that acetaldehyde can
completely eliminate increased ACO activity, whereas ethanol cannot (Bur-
don et al., 1996). Accordingly, Burdon et al. (1996) suggested that acetalde-
hyde can either inhibit ACO activity directly or prevent the increase in the
enzyme, thereby providing a possible mechanism for retarding fruit ripening.
14.4 Compositional Changes during Fruit Maturation and Ripening
Several important metabolic changes occur during the maturation and ripen-
ing of mangoes, and some of those are useful as maturity indices (Ketsa et al.,
1991). The ripening changes are irreversible senescence processes that are
related to degradation of organelles or changes in chemical constituents, and
thus relate to the quality and postharvest life of the fruit. Natural senescence
is aggravated and promoted by ethylene, mechanical injury and high tem-
perature, and can be delayed by low temperature, elimination of mechanical
damage and reduction of ethylene production.
Organic acids
Organic acids are important for respiratory activity and as avour constitu-
ents. During maturation and ripening, mango fruit experience a substantial
loss of organic acids. The predominant acids in mature mango fruit are citric,
succinic, malic and tartaric acids; citric acid has the highest concentration
and tartaric acid the lowest (Shashirekha and Patwardhan, 1976; Sarker and
Muhsi, 1981; Medlicott and Thompson, 1985). Citric acid content increases
steadily during fruit development in Irwin, reaching a maximum at the
beginning of the endocarp-hardening period, and then decreases steadily
Postharvest Physiology 495
(Ito et al., 1997). In Keitt the predominant organic acids are citric and malic
acids, but tartaric, oxalic, ascorbic, and -ketoglutaric acids also are present,
and the initial loss in acidity is due to a substantial loss in citric acid and a
small loss in malic acid (Medlicott and Thompson, 1985). In Badami man-
goes, citric acid is the major organic acid, but malic and succinic acids are
also present (Shashirekha and Patwardhan, 1976). In Fazli mangoes, oxalic,
citric, malic, pyruvic and succinic acids have been detected and tartaric acid
has been detected in Zardalu mangoes (Kumar et al., 1993). In general, citric
and succinic acids decrease during ripening while malic acid shows different
changes with different cultivars (Lizada, 1993).
Mango fruit contain organic acids involved in tricarboxylic acid cycle
reactions (i.e. oxalic, succinic, pyruvic, oxaloacetic and -ketoglutaric acids).
In Pairi mangoes, maximum concentration of -oxoglutaric and pyruvic acids
occur before the climacteric peak. Aspartic and glutamic acid concentrations
increase for c.3 days after harvest and then decrease as the climacteric maxi-
mum is reached (Krishnamurthy et al., 1971). Malic enzyme (EC 1.1.1.40), which
catalyses the oxidative decarboxylation of L-malic to pyruvic acid, occurs in the
three-quarter-ripe and ripe stages and the activity pattern during ripening is
similar in Alphonso, Banganpalli, Dasheri, Fazli, Langra and Suvar-
narekha (Selvaraj and Kumar, 1994). In Alfonso (sic), the levels of malic dehy-
drogenase (EC 1.1.1.37) and succinic dehydrogenase (EC 1.3.5.1) increase with
the onset of ripening; whereas, the level of citrate synthase (EC 2.3.3.1) increases
several-fold during maturation but decreases markedly during ripening (Baqui
et al., 1974). The activity of malic enzyme increases during ripening, reaching a
maximum immediately after the climacteric peak, and then declines (Dubery
et al., 1984). The activity patterns of phosphoenol pyruvate carboxylase (PEPC;
EC 4.1.1.49) and pyruvate decarboxylase (EC 4.1.1.1) during ripening vary
among different cultivars, while malic enzyme activity increases during ripen-
ing. PEPC activity is relatively high in Alphonso and Langara, but low in
Dashehari and Totapuri during ripening (Selvaraj and Kumar, 1994).
Soluble sugars
The increase in soluble sugars is a major change during mango fruit ripening,
and sweetness is the most important compositional change related to mango
avour. While starch content increases in chloroplasts during mango fruit
development, it is almost completely hydrolysed to simple sugars during
ripening (Medlicott et al., 1986; Selvaraj et al., 1989; Kumar et al., 1994; Ito
et al., 1997). In Alphonso, starch content is 14% (by weight) at the immature
stage and c.0.3% at the ripe stage. Similarly, starch is almost undetectable in
Irwin mangoes after ripening, whereas sucrose increases signicantly and
fructose increases slightly (Ito et al., 1997). Starch content decreases slightly
during ripening of Haden, but is insufcient to account for the observed
increase in the level of sucrose (Castrillo et al., 1992).
Ripe mango contains up to 1020% total sugars on a fresh weight (FW)
basis, depending on the cultivar and the stage of ripeness. At the beginning
J.K. Brecht and E.M. Yahia 496
of ripening, reducing sugars make up most of the sugar content, while there
are more non-reducing (c.17%) than reducing (3%) sugars in completely ripe
fruit. Sucrose contributes 57% of the total sugar in ripe Keitt mangoes, with
fructose and glucose making up 28% and 15%, respectively (Medlicott and
Thompson, 1985). Although Krishnamurthy et al. (1971), Lakshminarayana
(1973, 1975) and Shashirekha and Patwardhan (1976) reported a simultane-
ous increase of glucose, fructose and sucrose during ripening, Vazquez-Salinas
and Lakshminarayana (1985) observed a gradual reduction in glucose and
fructose and a continuous increase of sucrose during ripening in Haden,
Irwin, Kent and Keitt. Medlicott and Thompson (1985) and Vazquez-Salinas
and Lackshminarayana (1985) identied the main reducing sugar as fruc-
tose, while Selvaraj et al. (1989) reported that glucose is predominant. Con-
icting reports on the relative concentrations of individual sugars in mango
fruit during ripening is cultivar-dependent and due to different storage and
handling conditions (Medlicott and Thompson, 1985).
Sucrose content increases during ripening as a result of starch hydrolysis
from increased amylase (EC 3.2.1.1) activity (Mattoo and Modi, 1969a; Fuchs
et al., 1980; Tandon and Kalra, 1983). The high activities of sucrose synthase (EC
2.4.1.13) and invertase (EC 3.2.1.26) in the mesocarp during ripening indicate
active sucrose metabolism (Kumar et al., 1994). Hexoses and hexose phosphates
can be formed from pyruvate by gluconeogenesis (Selvaraj and Kumar, 1994).
The activity of glucose-6-phosphatase (EC 3.1.3.9) reportedly increases up to
the three-quarter-ripe stage; whereas, fructose-1,6-diphosphatase (EC 3.1.3.11)
activity increases as the fruit ripens from the three-quarter-ripe to full-
ripe stage (Kumar and Selvaraj, 1990). The glycolytic enzyme hexokinase
(6-phosphofructokinase; EC 2.7.1.11) has maximum activity at the ripe stage,
while pyruvate kinase (EC 2.7.1.40) activity increases until the three-quarter-ripe
stage and declines at ripening (Selvaraj and Kumar, 1994). The pattern of
activity changes in hexokinase/phosphofructokinase and pyruvate kinase
demonstrates that glycolysis is activated during mango fruit ripening.
Reducing sugars, mainly fructose, increase slightly during ripening, and
sucrose synthase (EC 2.4.1.13) activity increases approximately ten times
during the phase of rapid sucrose accumulation (Castrillo et al., 1992). This
activity accounts for the maximum rate of sucrose synthesis. The proportion
of sucrose phosphate synthase (EC 2.4.1.14) activity that is sensitive to inhibi-
tion by inorganic phosphate changes during ripening (Castrillo et al., 1992).
Maximum catalytic activity of sucrose synthase is constant throughout the
ripening period and contributes signicantly to sucrose metabolism. The
activities of neutral and acid invertases (EC 3.2.1.26) are very low in com-
parison with the other enzymes of sucrose synthesis. Acid invertase activity
increases and later decreases during ripening.
Structural polysaccharides
Pulp rmness is important for the evaluation of fruit maturity potential for
transport and storage, and as a quality characteristic. Fruit softening and cell
Postharvest Physiology 497
wall changes are principal changes associated with fruit ripening. Fruit tex-
ture changes are due to changes in cell walls and pectic substances in the
middle lamella, and these are cultivar-related (Selvaraj and Kumar, 1989).
Softening of mango fruit is characterized by increased solubility of cell wall
pectins (Roe and Bruemmer, 1981; Tandon and Kalra, 1984; Lazan et al., 1986;
Nasrijal, 1993). In general, water-soluble polysaccharides increase during
ripening (Lazan et al., 1986; Brinson et al., 1988), but water- and alkali-soluble
pectins decline in Keitt mangoes, and ammonium oxalate-soluble pectins
increase as the fruit become soft (Roe and Bruemmer, 1981). There is an over-
all loss of galactosyl and deoxyhexosyl residues during mango fruit ripen-
ing, the latter indicating degradation of the pectin component of the wall
(Muda et al., 1995). The loss of galactose appears to be restricted to the chela-
tor soluble fraction of the wall pectin, while loss of deoxyhexose seems to be
more evenly distributed among the pectin.
Pectinesterase (PE; EC 3.1.1.11), which catalyses the deesterication of
methyl groups from acidic pectins, has been detected in ripening mangoes
(Tahir and Malik, 1977; Roe and Bruemmer, 1981; Ali et al., 1990, 1995; Abu-
Sarra and Abu-Goukh, 1992). Physiological maturity in ripened mangoes is
associated with lower PE activity (van Lelyveld and Smith, 1979) and peel
has higher PE activity than pulp (Ashraf et al., 1981). Endo-polygalacturonase
(PG; EC 3.2.1.15), which is responsible for degrading the 1-4-linked galactur-
onic acid residues, occurs in ripening fruit (Lazan et al., 1986, 1993; Abu-Sarra
and Abu-Goukh, 1992). Enzymatic and/or non-enzymatic processes, in addi-
tion to PG activity, are involved in the extensive softening of fruit (Mitcham
and McDonald, 1992). Other cell wall hydrolases can be detected in ripening
fruit, including cellulases (EC 3.2.1.4; Lazan et al., 1986; Abu-Sarra and Abu-
Goukh, 1992), -galactosidase (EC 3.2.1.23; Ali et al., 1990, 1995; Lazan et al.,
1993), galactanase (EC 3.2.1.145; Ali et al., 1990) and xylanase (EC 3.2.1.8; Ali
et al., 1990).
Ripening in mangoes, as characterized by decreased tissue rmness, is
initiated in inner mesocarp tissue close to the seed, and progresses outwards
(Lazan et al., 1993). Pectin solubilization in inner and outer mesocarp tissues
is comparable, but pectin solubilization begins earlier in the inner than in the
outer mesocarp (Lazan et al., 1993). The outer mesocarp of Keitt remains
rm longer than Tommy Atkins, and the inner is softer than the outer meso-
carp at each stage of ripening in both cultivars (Mitcham and McDonald,
1992). Cell wall neutral sugars, particularly arabinosyl, rhamnosyl and galac-
tosyl residues, decrease with ripening in both cultivars. Keitt has more
loosely associated, chelator-soluble pectin, accumulates more soluble poly-
uronides and retains more total pectin at the ripe stage than Tommy Atkins.
Both cultivars have similar PG activity, which increases with ripening. The
amount and molecular weight (MW) of cell wall hemicellulose decreases
with ripening in both cultivars. Galactose is the only cell wall neutral sugar
to show a signicant decrease during ripening of Sensation mangoes (Sey-
mour et al., 1990). Losses of neutral sugars can be due to hydrolysis of galac-
tans and arabinogalactans by -galactosidase having galactanase activity.
-Galactosidase activity shows a parallel increase to tissue softening during
J.K. Brecht and E.M. Yahia 498
ripening. The close correlations between changes in -galactosidase during rip-
ening, and between changes in -galactosidase activity with tissue softening,
and increased pectin solubility and degradation suggests that -galactosidase
has an important role in cell-wall pectin modication and mango fruit soft-
ening during ripening (Ali et al., 1995).
Postharvest treatments, such as refrigeration, packaging, application of
fruit coatings, etc., can retard mango fruit softening and activity of pectinases
(Lazan et al., 1990; Nasrijal, 1993). Calcium (Ca) joins free carboxyl groups
resulting from PE-catalysed hydrolysis of methyl ester bonds to form Ca-
bridges between adjacent pectin molecules. Calcium application to Haden
mangoes by inltration or dipping extends their storage life by 1 week (Zam-
brano and Manzano, 1995). Postharvest vacuum application of Ca to mango
has also been tried (Tirmazi and Wills, 1981; Wills et al., 1988; van Eeden,
1992; Yuen et al., 1993). Vacuum (300 mm Hg) inltration of 14% calcium
chloride (CaCl
2
) into Amrapali and Deashehari mangoes ripened at 25C
inhibits PG activity, while ethylene treatment (1 l/l) markedly increases its
activity (Reddy and Srivastava, 1999). Pressure (115 kPa for 2 min) or vacuum
inltration (32 kPa) with 18% CaCl
2
delays ripening of Kensington Pride
mangoes by 12 or 8 days, respectively (Yuen et al., 1993). Yuen et al. (1993)
reported that vacuum inltration of CaCl
2
causes some peel injury, which
can be reduced by: (i) increasing the temperature of the fruit esh or the
CaCl
2
solution during pressure inltration; (ii) packaging the fruit in sealed
polyethylene during pressure inltration; and (iii) packaging the fruit in
sealed polyethylene bags or cling or shrink wraps after CaCl
2
treatment. Cal-
cium chloride inltration of Keitt mangoes reduces ethylene production,
respiration rate and the incidence of storage decay (van Eeden, 1992).
Pigments and colour
Mango skin colour is important for its role in the perception of overall qual-
ity (Gonzlez-Aguilar et al., 2001) and can be important for determining the
appropriate maturity for harvesting (Cocozza et al., 2004; Jha et al., 2007), pro-
cessing (Mahayothee et al., 2004) and consumption (Cocozza et al., 2004; Jha
et al., 2007). The loss of green colour is an obvious sign of fruit ripening in
many mango cultivars. The development of the optimum skin colour usually
denes mango quality. Some mango cultivars retain green colour in ripe
fruit. Depending on the cultivar, skin colour can change from dark to olive-
green; sometimes reddish, orange-yellow or yellowish hues appear from the
base colour. Some cultivars develop a reddish blush, which has been attrib-
uted to anthocyanins. Colour changes in mango fruit are due to the disap-
pearance of chlorophyll and the appearance of other pigments (Fig. 14.4).
Chloroplasts are transformed to chromoplasts containing yellow or red pig-
ments (John et al., 1970; Lakshminarayana, 1980; Parikh et al., 1990; Lizada,
1993). Well-arranged grana and osmiophilic globules occur in chloroplasts of
cells in the peel of unripe mangoes (Parikh et al., 1990), and lose integrity
during ripening. Osmiophilic globules appear, indicating the transformation
Postharvest Physiology 499
of the chloroplast to a chromoplast. In yellow cultivars, carotenoids and
xanthophylls are the predominant pigments. The anthocyanin paenoidin-
3-galactoside occurs in the skin of some cultivars (Proctor and Creasy, 1969).
During fruit ripening, chlorophyll concentration decreases substantially in
Keitt, while carotenoid concentration increases and anthocyanin decreases
gradually in Tommy Atkins (Medlicott et al., 1986). In Keitt, a substantial loss
of chlorophyll in the peel occurs after the fruit begin to soften. Peel colour is not
an adequate maturity index, since the fruit is already soft when the colour
change occurs. Tommy Atkins mangoes develop more red and yellow pigmen-
tation in the peel and mesocarp than Keitt (Mitcham and McDonald, 1992).
Mango fruit pulp contains high concentrations of carotenoids (up to 9
mg/100 g), causing the development of an intense yellow to orange colour.
Mango is a good source of vitamin A. The pulp carotenoid level is cultivar-
dependent. In Alphonso, 16 fractions of carotenoids have been reported:
50% of those are -carotene (Jungalwala and Cama, 1963; John et al., 1970). No
qualitative changes in carotenoid composition have been reported for Keitt and
Tommy Atkins mangoes from mature-green to the ripe stage, although quan-
titative changes occur during ripening (Mercadante and Rodriguez-Amaya,
6
5
15
Green Green/yellow Yellow/green Yellow
Anthocyanin
LSD
(P = 0.05)
LSD
(P = 0.05)
LSD
(P = 0.05)
Carotenoid
Chlorophyll
10
5
4
3
C
a
r
o
t
e
n
o
i
d
s

(

g
/
c
m
2
)
C
h
l
o
r
o
p
h
y
l
l
s

(

g
/
c
m
2
)
A
n
t
h
o
c
y
a
n
i
n
s

(

g
/
c
m
2
)
2
1
10
8
6
4
2
0 3 6
Storage time (days)
9 12 15
Fig. 14.4. Carotenoid, anthocyanin and chlorophyll concentrations in the peel of
Tommy Atkins mango during ripening at 22C (Source: Medlicott et al., 1986).
LSD, least signicant difference.
J.K. Brecht and E.M. Yahia 500
1998). However, John et al. (1970) detected 15, 14 and 17 carotenoids in Bad-
ami mangoes at mature-green, partially ripe and fully ripe stages of fruit,
respectively. Variation with respect to pigment types and quantities is due to
cultivar differences, geography and climate, different maturity stages and
treatments after harvest; discrepancies in results are probably due to differ-
ent analytical procedures.
Mango skin colour can be used to estimate the content of all-trans--
carotene (Vzquez-Caicedo et al., 2004), the most important provitamin A
carotenoid (Wolf, 1984). Ornelas-Paz et al. (2007) demonstrated that the val-
ues of external and internal colour are similar in Manila and Ataulfo man-
goes (non-blushed) in contrast to blushed cultivars (Criollo, Paraso and
Kent). The carotenoids in fruit skin of some mango cultivars can be corre-
lated with some non-destructive colour measurements (Table 14.2; Figs.
14.514.8 (Ornelas-Paz et al., 2008).
The most abundant carotene of mango is all-trans--carotene, while the
most important xanthophylls are violaxanthin and its isomers (Wilberg and
Rodriguez-Amaya 1995; Chen et al., 2004). Mercadante et al. (1997) quantied
many cartenoids of Keitt mangoes and concluded that the most predomi-
nant xanthophylls were all-trans-violaxanthin and 9-cis-violaxanthin, account-
ing for 38% and 18% of total carotenoid content, respectively, although other
xanthophylls are important in other cultivars (Ben-Amotz and Fishler, 1998;
Setiawan et al., 2001).
Modi and Reddy (1967) reported an increase during mango ripening of the
carotene precursors, mevalonic acid (MVA) and geraniol, with a concomitant
increase in carotene content. The geraniol concentration of unripe Alphonso
Table 14.2. Correlation coefcients (R) for the relationships between the content of the
main carotenoids in mesocarp and the internal/external colour values in Ataulfo and Manila
mango fruit. The correlation analysis was performed using = 0.5 (Source: Ornelas-Paz
et al., 2008).
Cultivar Colour value
a
All-trans-violaxanthin 9-cis-Violaxanthin All-trans--carotene
Ataulfo a* 0.84/0.90 0.83/0.87 0.90/0.90
b* 0.05/0.41 0.00/0.41 0.05/0.45
L* 0.75/0.19 0.75/0.21 0.80/0.27
C* 0.31/0.71 0.34/0.70 0.33/0.72
h 0.88/0.89 0.86/0.87 0.94/0.90
Manila a* 0.92/0.87 0.93/0.89 0.86/0.81
b* 0.76/0.69 0.75/0.67 0.67/0.54
L* 0.86/0.35 0.86/0.32 0.74/0.18
C* 0.81/0.74 0.81/0.73 0.73/0.61
h 0.90/0.89 0.92/0.91 0.82/0.82
a
Colour was recorded using the Commission Internationale de lEclairage (CIE) L*a*b* uniform colour
space, where L* indicates lightness on a scale of 0 (black) to 100 (white), a* indicates chromaticity on a
green () to red (+) axis, and b* indicates chromaticity on a blue () to yellow (+) axis. Numerical values
of a* and b* were converted into chroma (C) and hue angle (h), which represent colour purity and the
shade of colour, respectively.
P
o
s
t
h
a
r
v
e
s
t

P
h
y
s
i
o
l
o
g
y
5
0
1
30 35 40 45
0
10
20
30
40
0 5 10 15 20 25 30 40 45 50 55 60 60 65 70 75 80 85
0
10
20
30
40
5 0 5 10 15 20 25
P
i
g
m
e
n
t

c
o
n
t
e
n
t
(
1
0

3

g
/
k
g
)
P
i
g
m
e
n
t

c
o
n
t
e
n
t
(
1
0

3

g
/
k
g
)
60 62 64 66 68
Mesocarp Mesocarp Mesocarp
Peel Peel Peel
a* value b* value L* value
Fig. 14.5. Relationships between the content of all-trans--carotene (), all-trans-violaxanthin (as dibutyrate, ), 9-cis-violaxanthin (as dibu-
tyrate, ) in mesocarp and the a*, b* and L* values, measured in mesocarp or peel of Ataulfo mango fruit during ripening. Each point repre-
sents the mean of two independent measurements the standard error (vertical bars). The continuous line represents an exponential regression
(Source: Ornelas-Paz et al., 2008).
J.K. Brecht and E.M. Yahia 502
mangoes varies from 0.5 to 3.0 mol with 0.0 to 0.5 mol MVA; in ripe mangoes
the corresponding levels are 510 and 15 mol, respectively. The increase in
free geraniol and MVA indicates that these compounds are dephosphorylated
during ripening. Acid phosphatase (EC 3.1.3.2) may regulate carotenogenesis
in ripe mangoes (Mattoo et al., 1968). Mangoes stored at low temperatures and
then ripened at room temperature fail to synthesize as much carotenoids as
fruit held at room temperature (Krishnamurthy and Subramanyam, 1973;
Thomas, 1975). Hot water treatments increase the colour intensity of the pulp
(Medlicott et al., 1986) and the peel (Esguerra and Lizada, 1990).
Tongdum mangoes, which ripen without changing colour, have three-
fold more chlorophyll and slightly more -carotene in the peel and have higher
rates of ethylene production compared with Nam Dok Mai mangoes, which
change from green to yellow upon ripening (Ketsa et al., 1999). Activities of
chlorophyllase (EC 3.1.1.14) and peroxidase (EC 1.11.1.7) in the peel of ripe
Tongdum fruit are about half of that in Nam Dok Mai fruit. Changes in the
peel of ripe green mangoes are due to either or both a lower activity of chloro-
phyllase or peroxidase activity and are not a result of low ethylene production.
Phenolic compounds
The phenolic content of mangoes is high early during development, then
decreases and remains fairly steady during ripening (Lakshminarayana et al.,
0
10
20
30
40
40 45 50 55 60 65 55 60 65 70 75 80 85
0
10
20
30
40
30 35 40 45 55 60 65 70 75 80 85 90 95
Mesocarp Mesocarp
C* value
Peel
Peel
h value
P
i
g
m
e
n
t

c
o
n
t
e
n
t
(
1
0

3

g
/
k
g
)
P
i
g
m
e
n
t

c
o
n
t
e
n
t
(
1
0

3

g
/
k
g
)
Fig. 14.6. Relationships between the content of all-trans--carotene (), all-trans-violaxanthin
(as dibutyrate, ), 9-cis-violaxanthin (as dibutyrate, ) in mesocarp and the C* and h values,
measured in mesocarp or peel of Ataulfo mango fruit during ripening. Each point represents
the mean of two independent measurements the standard error (vertical bars). The continuous
line represents an exponential regression (Source: Ornelas-Paz et al., 2008).
P
o
s
t
h
a
r
v
e
s
t

P
h
y
s
i
o
l
o
g
y
5
0
3
0
10
20
30
40
-5 0 5 10 15 20 25 20 30 40 50 60 50 55 60 65 70 75 80 85
0
10
20
30
40
-10 -5 0 5 10 15 20 20 25 30 35 40 55 60 65 70 75
Mesocarp Mesocarp
Peel
a* value b* value L* value
Mesocarp
Peel Peel
P
i
g
m
e
n
t

c
o
n
t
e
n
t
(
1
0

3

g
/
k
g
)
P
i
g
m
e
n
t

c
o
n
t
e
n
t
(
1
0

3

g
/
k
g
)
Fig. 14.7. Relationships between the content of all-trans--carotene (), all-trans-violaxanthin (as dibutyrate, ), 9-cis-violaxanthin
(as dibutyrate, ) in mesocarp and the a*, b* and L* values, measured in mesocarp or peel of Manila mango fruit during ripening. Each point
represents the mean of two independent measurements the standard error (vertical bars). The continuous line represents an exponential or
second order polynomial regression (Source: Ornelas-Paz et al., 2008).
J.K. Brecht and E.M. Yahia 504
1970). This is associated with loss of astringency (Selvaraj and Kumar, 1989).
The peel of mango fruit has a higher phenolic content than the pulp at all
stages of fruit development (Jain, 1961; Lakshminarayana et al., 1970).
Polyphenol oxidase (PPO; EC 1.14.18.1) catalyses the oxidation of mono-
and diphenols to o-quinones, which polymerize to produce brown pigments.
PPO activity increases slightly from harvest maturity to the half-ripe stage and
then declines in Banganapalli, Dashehari, Fazli and Langra mangoes, and
decreases in Alphonso, Suvarnarekha and Totapuri mangoes (Selvaraj and
Kumar, 1989). The PPO isolated from Haden mango is active towards the
o-diphenolic compounds, showing higher activity in the presence of catechol,
followed by chlorogenic acid, but not with monophenols (Park et al., 1980).
Flavour (taste, aroma)
Sugar changes are very important for organoleptic attributes in the mango
fruit. Fruit avour is mostly a balance between the content of sugars and
organic acids (Medlicott and Thompson, 1985) as well as aromatic volatiles.
Kapse et al. (1989) determined that increasing TSS and decreasing acidity
C* value
60 65 70 75 80 85 90 95
0
10
20
30
40
20 25 30 35 40 45 60 65 70 75 80 85 90 95 100 105 110
Mesocarp
0
10
20
30
40
20 25 30 35 40 45 50 55 60 65
Mesocarp
Peel
Peel
h value
P
i
g
m
e
n
t

c
o
n
t
e
n
t
(
1
0

3

g
/
k
g
)
P
i
g
m
e
n
t

c
o
n
t
e
n
t
(
1
0

3

g
/
k
g
)
Fig. 14.8. Relationships between the content of all-trans--carotene (), all-trans-violaxanthin
(as dibutyrate, ), 9-cis-violaxanthin (as dibutyrate, ) in mesocarp and the C* and h values,
measured in mesocarp or peel of Manila mango fruit during ripening. Each point represents
the mean of two independent measurements the standard error (vertical bars). The continu-
ous line represents an exponential or second order polynomial regression (Source: Ornelas-Paz
et al., 2008).
Postharvest Physiology 505
increases avour ratings of mango fruit. Sucrose is the predominant sugar in
ripe mango fruit (Tandon and Kalra, 1983; Medlicott and Thomson, 1985;
Vazquez-Salinas and Lakshminarayana, 1985). The predominant acid in mango
fruit is citric (Medlicott and Thompson, 1985; Lizada, 1993). Several factors
affect sugar and acid contents in mango, including cultivar (Kapse et al., 1989;
Kundu and Ghosh, 1992; Gowda et al., 1994), stage of maturity at harvest
(Shashirekha and Patwardhan, 1976; Morga et al., 1979; Tandon and Kalra,
1983), postharvest treatments (Kumar et al., 1993) and storage conditions
(Vazquez-Salinas and Lakshminarayana, 1985).
Ripe mangoes contain >300 volatiles (Pino et al., 2005), but not all of them
are odour-active and thus do not contribute signicantly to aroma. Studies
have identied the volatiles of mango, but not their aromatic activity. The
predominant volatiles in some cultivars are monoterpenes and sesquiter-
penes (MacLeod and De Troconis, 1982; Engel and Tressl, 1983; Pino et al.,
2005), as well as lactones and some fatty acids (MacLeod and Pieris, 1984;
MacLeod and Snyder, 1985; Wilson et al., 1990). However, there is no indica-
tion of the presence of a single avour impact component (Engel and Tressl,
1983). Some mango cultivars have a peach-like avour that may be related to
the presence of lactones, which contribute to the avour of peaches (Prunus
persica) (Lakshminarayana, 1980; MacLeod et al., 1988, Wilson et al., 1990).
MacLeod et al. (1988) detected four lactones in Kensington Pride that are
also the major volatiles of peach. Monoterpene hydrocarbons represent about
49% (w/w) of the total volatiles in Kensington Pride, with -terpinolene
being the most abundant (26%) and 16 esters representing 33% (MacLeod
et al., 1988). The esters, together with some of the lactones, contribute to the
avour of Kensington Pride mangoes.
Indian mangoes have a unique avour, which has been attributed to (Z)-
ocimine (Engel and Tressl, 1983; Lizada, 1993). Pino et al. (1989) detected 83
volatiles in Corazon, Bizcochuelo and Super Haden mangoes, and total
volatiles ranged between 39 mg/kg in Bizcochuelo to 70 mg/kg in Corazon.
The identied volatiles include -cubebene, -maaliene, ethyl(Z)-9-hexade-
canoate, ethyl(Z)-9,12-octadecanoate, ethyl(Z)(Z)(Z)-6,9,12-octadecanoate,
cucarvone, 2-methylpropane-2-ol, 3-methylepentan-ol, thymol and carvacrol
(Pino et al., 1989). MacLeod and Snyder (1985) listed the volatile components
of several mango cultivars, including Willard and Parrot from Sri Lanka;
levels of -terpinolene were similar to Kensington Pride.
Kostermans and Bompard (1993) considered that lack of bre was linked
to an absence of aroma and at taste and smell, but some cultivars such as
Kensington Pride are low in bre and have a distinctive avour and aroma
prole, and a high level of -terpinolene (Bartley and Schwede, 1987; MacLeod
et al., 1988). Lipid content of the pulp is correlated with the avour character-
istics of some mango cultivars (Bandyopadhyay and Gholap, 1973a; Gholap
and Bandyopadhyay, 1975b, 1976). The ripening of Alphonso mangoes at
ambient temperature is accompanied by a sharp increase in triglyceride con-
tent, together with the development of a strong aroma and avour (Gholap
and Bandyopadhyay, 1975a, 1976), but ripening at 10C results in a bland
aroma and avour (Bandyopadhyay and Gholap, 1973b). Totapuri mangoes,
J.K. Brecht and E.M. Yahia 506
a bland cultivar, showed no change in the development of aroma or in the
pulp lipid content (Gholap and Bandyopadhyay, 1975b). During ripening at
ambient temperature, palmitoleic acid content is higher than that of palmitic
acid in Alphonso, whereas ripening at low temperature does not affect the
proportions of these two fatty acids (Bandyopadhyay and Gholap, 1973b).
The relative proportions of palmitoleic and palmitic acids in Totapuri mango
pulp are constant irrespective of the ripening conditions (Gholap and Ban-
dyopadhyay, 1975b). Gholap and Bandyopadhyay (1976, 1980) suggested
that the relative contents of palmitic and palmitoleic acids determine the a-
vour quality of mango fruit.
The absence of lactones having coconut-like odour notes in Totapuri
mangoes may be signicant for differentiating its aroma characteristics from
Alphonso, together with the presence of certain similar and dissimilar com-
ponents (Bandyopadhyay, 1983). The aroma of green mangoes has been
attributed to cis-ocimine in Alphonso and -myrcene in Batali mangoes
(Gholap and Bandyopadhyay, 1976; Bandyopadhyay, 1983). Table 14.3 lists
characteristic aromas of Alphonso and Totapuri mangoes and their possi-
ble chemical identities.
In almost all fruits, aromatic volatiles are produced at later stages of rip-
ening (Yahia, 1994). Tree-ripe Tommy Atkins mangoes produce higher lev-
els of all aroma volatiles except hexanal than do mature-green fruit (Bender
et al., 2000a). Both mature-green and tree-ripe mangoes stored in 25 kPa CO
2

tend to have lower terpene (especially p-cymene) and hexanal concentra-
tions than those stored in 10 kPa CO
2
and air-stored fruit. Acetaldehyde and
ethanol levels tend to be higher in tree-ripe mangoes held in 25 kPa CO
2
than
in those from 10 kPa CO
2
or air storage, especially at 8C. Inhibition of volatile
production by 25 kPa CO
2
is greater in mature-green than in tree-ripe
Table 14.3. Characteristic aromas in Alphonso and Totapuri mangoes and their possible
chemical causes (Source: Bandyopadhyay, 1983).
Aroma Alphonso Totapuri
Fruit, estery Acetaldehyde, methyl acetate,
ethyl acetate, n-butyl acetate
Propionaldehyde
Methyl acetate
Green-mango-like cis-Ocimine -Myrcene
Camphoraceous Not detected Detected, but not identied
Earthy Caryophyllene-pinene Not detected
Almond-like Benzaldehyde Not detected
Burnt-sugar-like Benzonitrile Not detected
Spicy Not detected -Terpinene
Sweet, sugar-like Detected, but not identied Not detected
Coconut oil-like -Caprolactone, -octalactone,
-undecalactone
Not detected
Postharvest Physiology 507
mangoes, and at 8C compared to 12C for tree-ripe fruit. However, aroma
volatile levels in tree-ripe mangoes from 25 kPa CO
2
are equal to or greater
than those in mature-green fruit treatments. Atmospheres that prolong
mango shelf life by slowing ripening processes can allow tree-ripe mangoes
to be stored or shipped without sacricing their aroma quality.
Quality enhancement has been used to determine properties critical to a-
vour acceptability of mangoes, and focus group interviews have been conducted
to determine sensory attributes important to the purchase and consumption
of mangoes (Malundo, 1996). Sugars and acids enhance perception of specic
avour notes in mango, including aromatics (Malundo et al., 2001).
14.5 Transpiration and Water Loss
Water loss lowers fruit weight, resulting in shrivelling, and may further reduce
quality by causing poor colour development and uneven ripening. Water is lost
from mango fruit through stomata, lenticels and other openings. Relative
humidity (RH) inside the fruit is 100% and water is lost when RH in the envi-
ronment surrounding the fruit is <100%. Water loss is also greatly inuenced by
temperature. With constant RH and air movement, water loss increases signi-
cantly with any increase in temperature. Transpiration rate is inuenced by cul-
tivar and ripeness stage. It is correlated with skin thickness, morphological
structure, epidermal cells and surface wax coating. For example, waxes usually
develop on the epidermis of fruit in the later stages of development and thus it
is common for fruit harvested early to shrivel faster compared with those har-
vested at a more advanced stage of development (Yahia et al., 2006a).
14.6 Physical Damage and Physiological Disorders
Mangoes are susceptible to physical damage at every step of the postharvest
handling chain (see Johnson and Hofman, Chapter 15, this volume) and
reduction/elimination of mechanical injury is essential to reduce losses in
quality and postharvest life. Mango fruit are susceptible to various physio-
logical disorders that inuence fruit quality (see Galn Saco, Chapter 9, this
volume). These disorders are either induced or inherent, and several of them
become apparent during fruit ripening. Disorders, i.e. chilling injury (CI) and
heat injury, may be induced after harvest. Inherent physiological disorders
include spongy stem end in Kensington Pride (Brown et al., 1981), soft nose
in Florida mangoes (Young, 1957) and internal breakdown, spongy tissue or
soft nose in Indian Alphonso mangoes (Subramanyam et al., 1971).
Chilling injury (CI)
Low storage temperatures can injure mature-green mangoes if storage exceeds
a day or so at or near 0C to a few weeks at just below 12C. This problem
limits the use of low storage temperature to manage postharvest ripening
J.K. Brecht and E.M. Yahia 508
and seriously affects the ability of handlers to store mangoes or transport
them over long distances, because temperatures that are low enough to delay
ripening, decay and senescence may damage the fruit. The symptoms of CI
include greyish, scald-like discoloration on the skin, followed by pitting,
uneven ripening, and poor avour, aroma and colour development (Hatton
et al., 1965; Medlicott et al., 1990). The symptoms often are not apparent at the
low temperature, but develop later, when the fruit are brought to warmer
temperatures for ripening or are displayed for sale. Other symptoms in
mango fruit held at room temperature for 12 days after low temperature
storage were described as discoloured and with pitted areas on the surface
(Srivastava, 1967; Kane, 1977) followed by increased susceptibility to micro-
bial spoilage (Sadasivam et al., 1971; Subramanyam et al., 1975).
Chilling susceptibility varies with cultivar (Farooqui et al., 1985); Haden
and Keitt are particularly susceptible. Sensation developed more skin
symptoms than Sammar Bahisht mangoes (Farooqui et al., 1985). While CI
has generally been reported to occur in mango fruit at temperatures below
c.1013C (Mukherjee, 1958; Akamine, 1963; Hatton et al., 1965; Musa, 1974;
Couey, 1986), some cultivars (i.e. Dashehari and Langara) were reported to
be safely stored at 78C for up to 25 days (Mann and Singh, 1976). While
most cultivars show injury at <10C if fruit have just reached maturity, toler-
ance of CI increases as fruit ripen (Medlicott et al., 1990; Mohammed and
Brecht, 2002). Tolerance of Keitt mango fruit of CI was induced by pre-storage
heat treatments (McCollum et al., 1993).
Heat injury
Mango is highly tolerant of heat (Yahia et al., 2000; Jacobi et al., 2001b). Man-
goes that are not stored in refrigerated conditions after harvest may be
exposed to extremely high ambient temperatures in many production areas.
This may lead to heat injury, especially if the fruit are exposed to >30C for
>10 days, but injury can also occur more rapidly at higher temperatures. The
heat disinfestation treatments of mangoes that are required for insect quar-
antine security may injure fruit that are not fully mature (Jacobi and Giles,
1997; Jacobi et al., 2001a).
External symptoms of heat injury include lenticel spotting and skin
browning (scald) with secondary disease development, while internal
symptoms include mesocarp browning, tissue cavitation and starch spots
(Jacobi and Wong, 1992; Jacobi and Giles, 1997; Mitcham and McDonald,
1997; Jacobi et al., 2001a, b). Ripening of heat-injured mangoes may also be
inhibited (Jacobi et al., 2001a, b).
14.7 Modied Atmospheres (MA) and Controlled Atmospheres (CA)
Long-term marine shipping in MA and CA has been used for transit from
several countries (Yahia, 1993). Research results are very contradictory due to
Postharvest Physiology 509
the different cultivars and maturity stages of mangoes used, different atmo-
spheres implemented and lack of experimental controls. Optimum condition
for prolonged shipping or storage is reported to be 35 kPa O
2
plus 510 kPa
CO
2
, which can delay ripening, but the benets are not very signicant. Use
of CA and MA would most likely be benecial in delaying fruit ripening
during marine transport for 2 weeks or more.
Bender et al. (2000b) determined the tolerance of preclimacteric Haden
and Tommy Atkins to reduced O
2
levels for storage times in typical marine
shipments. They reported that mangoes can tolerate 3 kPa O
2
for 23 weeks
at 1215C and that tolerance of low O
2
decreases as mangoes ripen. All low
O
2
treatments reduced mature-green mango respiration; however, elevated
ethanol production occurred in 2 and 3 kPa O
2
storage, with the levels two to
threefold higher in Tommy Atkins than in Haden. Haden fruit at the
onset of the climacteric accumulated ethanol in 4 kPa O
2
and produced 1020
times more ethanol in 2 and 3 kPa O
2
than preclimacteric fruit. There were no
visible injury symptoms, but off-avour developed in mature-green fruit at 2
kPa O
2
and in ripening-initiated fruit at 2 and 3 kPa O
2
. Ethanol production
was not affected by storage in 25 kPa CO
2
. Ethylene production was reduced
slightly by low O
2
; however, Haden fruit also showed a residual inhibitory
effect on ethylene production at 2 or 3 kPa O
2
storage, while Tommy Atkins
fruit stored in 2 kPa O
2
produced a burst of ethylene upon transfer to air at
20C. Fruit rmness, total sugars and starch levels did not differ among treat-
ments, but 2, 3 or 4 kPa O
2
and 25 kPa CO
2
maintained signicantly higher
acidity than 5 kPa O
2
or air. The epidermal ground colour responded differ-
ently to low O
2
and high CO
2
in the two cultivars. Only 2 kPa O
2
maintained
Haden colour better than air, while all low O
2
levels maintained Tommy
Atkins colour better than air. High CO
2
was more effective than low O
2
in
maintaining Haden colour, but had about the same effect as low O
2
on
Tommy Atkins.
Properly selected atmospheres, which prolong mango shelf life by slow-
ing ripening processes, can allow tree-ripe mangoes to be stored or shipped
without sacricing their superior aroma. Mature-green and tree-ripe Tommy
Atkins mangoes were stored for 21 days in air or in a CA (5 kPa O
2
+ 10 kPa
or 25 kPa CO
2
) at 12C (mature-green) and at either 8 or 12C (tree-ripe)
(Bender et al., 2000a). Tree-ripe mangoes produced much higher levels of all
aroma volatiles except hexanal than mature-green fruit after ripening for 2
days. Both mature-green and tree-ripe mangoes stored in 25 kPa CO
2
had
lower terpene (especially p-cymene) and hexanal levels than those stored in
10 kPa CO
2
and air-stored fruit. Acetaldehyde and ethanol levels were higher
in tree-ripe mangoes from 25 kPa CO
2
than in those from 10 kPa CO
2
or air
storage, especially at 8C. Inhibition of volatile production by 25 kPa CO
2

was greater in mature-green than in tree-ripe mangoes, and at 8C com-
pared to 12C for tree-ripe fruit. Aroma volatile levels in tree-ripe mangoes
from the 25 kPa CO
2
treatment equalled or exceeded those in mature-green
fruit treatments.
Mangoes have high tolerance of short-term elevated CO
2
atmospheres
(Yahia, 1998). Mangoes can tolerate CO
2
atmospheres of up to 25 kPa for
J.K. Brecht and E.M. Yahia 510
2 weeks at 12C (Bender et al., 2000b). High (25 kPa) CO
2
inhibits ethylene
production, but increases ethanol production. Aroma volatiles are reduced
following 25 kPa CO
2
treatment, while 10 kPa CO
2
, low O
2
atmospheres and
storage temperature did not signicantly inuence production of terpene
hydrocarbons, which are characteristic of Florida-type mangoes. Mature-
green Tommy Atkins mangoes can be stored for 21 days in CA (5 kPa O
2
+
10 kPa or 25 kPa CO
2
) at 12C, while tree-ripe fruit can be stored for 21 days
in the same atmospheres at either 8 or 12C (Bender et al., 2000a).
The quality of Keitt mangoes was evaluated during storage for 6 days
at 20C in an extremely low O
2
(LO) CA (approximately 0.3 kPa) before stor-
age in modied atmosphere packaging (MAP) made from three, low-density
polyethylene (LDPE) lms with different gas permeability characteristics
(Gonzlez-Aguilar et al., 1997). Both LO and MA treatments delayed the
losses of colour, weight and rmness. Fruit maintained good appearance
with a signicant delay of ripening. Mangoes are very tolerant of LO treat-
ment; however, some MAP fruit developed a fermented taste after 10 and 20
days at 20C. Short duration (6-day) storage of mangoes in LO did not other-
wise have any deleterious effect on fruit quality during subsequent storage
under MA or normal atmosphere. Properly selected atmospheres, which pro-
long mango shelf life by slowing ripening, permit fruit to be shipped without
sacricing superior aroma.
Beaulieu

and Lea (2003) studied Keitt and Palmer mangoes without
heat treatment to assess volatile and quality changes in stored fresh-cut man-
goes prepared from rm-ripe (FR) and soft-ripe (SR) fruit, and to assess what
effect MAP may have on cut fruit physiology, overall quality and volatile
retention or loss. Subjective appraisals of fresh-cut mangoes based on aroma
and cut edge or tissue damage indicated that most SR cubes are unmarket-
able by day 7 at 4C. Both cultivars stored in MAP at 4C had almost identical
O
2
consumption, which is independent of ripeness. The CO
2
and O
2
concen-
trations measured for cubes stored in passive MAP indicated that the system
is inadequate to prevent potential anaerobic respiration after 7 days storage.
Injuries associated with MA and CA
A 10 kPa CO
2
atmosphere alleviates chilling symptoms in Kensington Pride
fruit, but higher concentrations are injurious; low O
2
(5 kPa) has no signi-
cant effect (OHare and Prasad, 1993). Higher concentrations of CO
2
(>10
kPa) are ineffective for alleviating CI at 7C, and tend to cause tissue injury
and high levels of ethanol in the pulp. Injury in Kensington Pride caused by
higher levels of CO
2
appears to be more severe at lower temperatures (OHare
and Prasad 1993; Bender et al., 1994, 1995), which could be a result of either
compounding injury (chilling + CO
2
) or reduced sensitivity of ripe mango
to CO
2
.
Rad mangoes develop internal browning and off-avour in atmo-
spheres containing 6 and 8 kPa CO
2
(Noomhorm and Tiasuwan, 1995). The
presence of starchy mesocarp in Carabao mangoes, which is characteristic
Postharvest Physiology 511
of internal breakdown, increases during storage in MA (Gautam and Lizada,
1984). Fruit stored for 45 days have severe symptoms, including air pockets
in the mesocarp resulting in spongy tissue (Nuevo et al., 1984a, b). Paren-
chyma cells of affected tissues have c.18 starch granules per cell, compared to
c.2 starch granules in healthy adjacent cells. However, no difference in starch
granule shape was detected between the spongy and healthy tissues. The
spongy tissue, which usually occurs in the inner mesocarp near the seed and
becomes evident during ripening, has almost ten times the starch content of
healthy tissue in the same fruit. External symptoms of internal browning due
to MA include failure of the peel to develop colour beyond the half-yellow
stage.
Carabao mangoes stored in polyethylene bags (0.04 mm thickness) for
1 day at 2531C had a faint fermented odour that disappeared during sub-
sequent ripening outside the bags (Gautam and Lizada, 1984). The fermented
odour increases with time, and persists throughout ripening when the fruit
are stored for 25 days. The respiratory quotient of this cultivar ranged from
0.59 at 21 kPa O
2
to 6.03 at 2.4 kPa O
2
, which indicates a progressively anaer-
obic metabolism (Sy and Mendoza, 1984). CO
2
production decreases as O
2

decreases from 21 to 3 kPa, but increases at <3 kPa O
2
. Fermentative decar-
boxylation could explain the odour (Lakshminarayana and Subramanyam,
1970).
Pronounced decay appears after storage of Rad mangoes for 20 days in
atmospheres containing 46 kPa O
2
with 48 kPa CO
2
at 13C and 94% RH,
and severe incidence of decay appears after 25 days (Noomhorm and Tiasu-
wan, 1995). Greater incidence of decay (stem-end rot and anthracnose) occurs
in Carabao mango stored in MA for 25 days at 2531C (Gautam and
Lizada, 1984).
Modied atmosphere packaging (MAP)
Modied atmosphere packaging is used to create a benecial MA around a
packaged product using semipermeable lm to restrict the movement of
respiratory gases into and out of the package; at equilibrium, the respiration
rate of fruit in MAP is equal to the diffusion of the respiratory gases through
the lm. Fruit wrapped in 0.08 mm thick polyethylene bags, with and with-
out perlite-potassium permanganate (KMnO
4
) and stored for 3 weeks at 10C
before treatment with ethylene developed normal colour, texture and avour
(Esguerra et al., 1978). Individually sealed Keitt in low-density (LDPE) and
high-density (HDPE) polyethylene lms for 4 weeks at 20C exhibited
delayed ripening, reduced weight loss and did not develop any off-avours
(Gonzalez et al., 1990). The LDPE had a thickness of 0.010 mm and permea-
bilities of 700 cm
3
O
2
/m
2
/h/atm and 0.257 g H
2
O/m
2
/h/atm. The HDPE
lm had a thickness of 0.020 mm and permeabilities of 800 cm
3
O
2
/m
2
/h/
atm and 0.166 g H
2
O/m
2
/h/atm.
In a study to model MAP for mango, Keitt fruit were individually vacuum
packaged in LDPE lm (0.0245 mm thick, 25.0 g/m
2
) and stored at 7C/8090%
J.K. Brecht and E.M. Yahia 512
RH, 12C/7585% RH, 17C/7080% RH, 22C/6575% RH or 25C/6575%
RH (Yamashita et al., 1997). After mass transfer had reached steady state,
respiration rates, moisture loss, permeability of peel and lm to water vapour,
and composition of atmosphere around the fruit were determined for 33
days. Daily rates of weight loss increased from 4.1 g/kg of fruit at 7C to 10.9
g/kg at 25C. Respiration rates also increased with storage temperature for
both packaged and unpackaged mangoes, and were 21, 38 and 43% less in
packaged fruit at 12, 17 and 22C, respectively. Permeability of peel was 600-
fold greater than that of the plastic lm. The in-package CO
2
levels increased
and O
2
decreased with time; concentration changes were greatest during the
rst 1015 days of storage and were more marked at the higher tempera-
tures. Experimental and calculated values for CO
2
levels differed by 29%,
depending on temperature.
Tommy Atkins mangoes individually sealed in heat-shrinkable lms
and stored for 2 weeks at 12.8C and then ripened at 21C had less weight
loss, but did not show differences in rmness, skin colour development,
decay development or time to fruit ripening, and had more off-avours than
unwrapped fruit (Miller et al., 1983). Polyethylene lms used were: Clysar
EH-60 lm of 0.01 mm nominal thickness, Clysar EHC-50 copolymer lm of
0.013 mm nominal thickness, and Clysar EHC-100 copolymer lm of 0.025
mm nominal thickness. Individual mature fruit of the same cultivar were
later sealed in Clysar EHC-50 copolymer lm with 0.013 mm thickness, and
Cryovac D955 with 0.015 mm thickness, and stored at 21C and 8590% RH
(Miller et al., 1986). The O
2
permeabilities of the lms were 620 cm
3
/24 h/
m
2
/atm and 9833 cm
3
/24 h/m
2
/atm, respectively. Water permeability was
1.5 g/24 h/m
2
and 2.0 g/24 h/m
2
at 23C, respectively. Fruit in MAP had less
weight loss, but higher incidence of decay and off-avour at soft-ripeness
than unsealed fruit. The authors concluded that there were no practical ben-
ets from wrapping this fruit in these lms and storage at 21C or even at
lower temperatures. Film wrapping mangoes at various stages of ripeness
after harvestwill not improve the maintenance of mango quality during
storage or ripening.
Keitt mangoes were individually sealed in LDPE lms and in a heat-
shrinkable copolymer (Cryovac D-955) lm with non-sealed mangoes as the
control and stored for up to 5 weeks at 12C, 17C or 22C (Yamashita et al.,
1999). MAP reduced the rate constant of vitamin C degradation at all tem-
peratures and vitamin C content of individually packaged mangoes was less
affected by storage temperature than the control. Values for Q10 (the ratio of
CO
2
production to O
2
consumption in respiration) were 1.3 and 1.0 for man-
goes wrapped with the heat-shrinkable copolymer and the LDPE lms,
respectively, and 2.8 for the non-sealed control.
The combined effect of hot benomyl (1000 ppm) at 55C for 5 min and
lm packaging in 0.01 mm PVC extended the storage life of mature-green
Nam Dok Mai mangoes stored at 13C (Sornsrivichai et al., 1992). Fruit qual-
ity was not affected by lm packaging after 4 weeks, but fruit showed infe-
rior quality after 6 weeks. The inhibition of carotene pigmentation in the peel
of this cultivar may be related to O
2
concentration inside the package and not
Postharvest Physiology 513
to CO
2
concentration (Yantarasri et al., 1994). At least 16 kPa O
2
is essential for
development of peel colour to the marketable stage (greenish).
Kensington Pride mangoes treated with heated benomyl (0.5 g/l at
51.5C for 5 min) and sealed in polyethylene bags (0.04 mm thickness) for
various durations at 20C, had off-avour and lacked normal skin colour
when ripened, but ripened satisfactorily in perforated bags (Chaplin et al.,
1982). The postharvest life of these fruit was not consistently longer than the
control. The CO
2
concentration in the bags was >20 kPa while the O
2
concen-
tration was <5 kPa. The incidence of off-avours was reduced by including
ethylene-absorbent blocks in the bags. The authors concluded that mangoes
cannot be stored satisfactorily at ambient temperature by such technique;
however, Stead and Chithambo (1980) reported that fruit ripening at 2030C
was delayed 5 days by sealing in polyethylene bags (0.02 mm thickness) with
ethylene-absorbent blocks without any abnormal avour.
Tommy Atkins and Keitt mangoes were individually sealed in shrink-
able Cryovac polyolen lms (0.015 or 0.019 mm thickness), either non-
perforated (MD lm) or perforated with eight holes of 1.7 mm diameter/
inch
2
(MPY) or eight holes of 0.4 mm diameter/inch
2
(SM60M) (Rodov et al.,
1994). After 23 weeks at 14C and an additional week at 17C, mangoes
packaged in perforated polyolen lms ripened normally. Optimum results
were achieved when the lm with 0.4 mm perforations was combined with
increased free volume inside the package by sealing the fruit within polysty-
rene trays. After 3 weeks of storage and 1 week of shelf life, sealed Keitt
mangoes were inferior to the control; they were less ripe, but beyond 4 weeks
(up to 6 weeks) sealed fruit had better quality scores because they were less
overripe. Sealing did not reduce decay of fruit stored for long periods.
Non-perforated PVC lm packaging of Nam Doc Mai mangoes was not
sufciently permeable for O
2
exchange to allow proper ripening (Yantarasri
et al., 1995). Therefore, a perforated MA was used in which fruit were
wrapped in polystyrene trays (three fruit/pack) at 20C with perforation
area of 0.004 cm
2
. Fruit ripened normally with no off-avours. Colour devel-
opment in the peel required a higher concentration of O
2
than the esh. A
lm of pore area 0.008 cm
2
allowed fruit colour to develop after 3 weeks
while a pore area of 0.39 cm
2
allowed the fruit to colour within 2 weeks.
Semipermeable coatings
Some fruit coatings can create an internal MA within the fruit due to semi-
permeable restriction of O
2
and CO
2
movement in and out of the fruit. Bald-
win et al. (1999) tested two types of fruit coatings polysaccharide-based and
carnauba wax-based for their effect on external and internal mango fruit
atmospheres and quality factors during simulated commercial storage at 10
or 15C with 9099% RH, followed by simulated marketing conditions at
20C and 56% RH. The coatings exhibited markedly different O
2
permeabil-
ity characteristics under laboratory conditions. Polysaccharide coatings were
less permeable to respiratory gases (i.e. O
2
and CO
2
) and more permeable to
J.K. Brecht and E.M. Yahia 514
water vapour compared to carnauba wax. When applied to fruit under simu-
lated commercial conditions, however, the differences between the coatings
with regards to their permeability to respiratory gases were much reduced,
most likely due to high RH during cold storage. Both coatings created a MA
within the fruit, reduced decay and improved appearance by imparting a sub-
tle shine; but only the polysaccharide coating delayed ripening and increased
concentrations of avour volatiles. The carnauba wax coating signicantly
reduced water loss compared to uncoated and polysaccharide-coating treat-
ments.
Julie mangoes treated with 0.75% w/v aqueous solution of Pro-long
semipermeable fruit coating (a mixture of sucrose esters of fatty acids and
sodium salt of carboxy methyl cellulose) and stored at 25C and 8595% RH
exhibited reduced weight loss, retarded ripening and increased storage life
(6 days longer) without evidence of adverse effects on quality (Dhalla and
Hanson, 1988). Treatment with 1.0% Pro-long could increase ethanol concen-
tration in the pulp. Treatment with Pro-long (0.82.4%) also delayed ripening
of Haden (Carrillo-Lpez et al., 1996).
Insecticidal CA
Mangoes are very tolerant of insecticidal atmospheres, and thus a potential
commercial application is feasible, especially in combination with other treat-
ments (i.e. heat). Keitt mango tolerated as low as 0.2 kPa O
2
and as high as
80 kPa CO
2
for up to 5 days without any injury upon ripening, although fer-
mentative odours could be noted while the fruit were under atmosphere
stress (Yahia, 1993, 1994, 1995, 1997; Ortega and Yahia, 2000). Other mango
cultivars were also evaluated and were very tolerant of extreme atmospheres
(Yahia, 1998).
Storage of Keitt mangoes in an insecticidal MA (0.030.26 kPa O
2
, 7279
kPa CO
2
, balance nitrogen (N
2
)) and CA (0.2 kPa O
2
, balance N
2
; or 2 kPa O
2

+ 50 kPa CO
2
, balance N
2
) for up to 5 days at 20C delayed fruit ripening as
indicated by respiration, esh rmness and colour development (Yahia et al.,
1989; Yahia, 1993; Yahia and Tiznado, 1993; Yahia and Vazquez, 1993). The
activities of phosphofructokinase, alcohol dehydrogenase (EC 1.1.1.1) and
pyruvate decarboxylase were enhanced but activity of pyruvate kinase, suc-
cinate dehydrogenase and -ketoglutarate dehydrogenase (EC 1.2.4.2) was
unaffected. Although these atmospheres caused changes in glycolysis and
tricarboxylic acid cycle, there was no indication of injury and the fruit rip-
ened normally in air. Sensory evaluation conducted after fruit ripening
showed no off-avours, and there were no differences between fruit main-
tained in the MA or CA and those maintained continuously in air. Keitt
mango is therefore very tolerant of insecticidal atmospheres, and 5 days
exposure is sufcient to control many insects (Rojas-Villegas et al., 1996).
Storage of Keitt and Tommy Atkins mangoes for 21 days at 12C in
atmospheres containing 25, 45, 50 or 70 kPa CO
2
plus either 3 kPa O
2
or air,
induced ethanol production of 0.183.84 ml/kg/h after transfer to air at 20C
Postharvest Physiology 515
for 5 days (Bender et al., 1994). Atmospheres containing 50 or 70 kPa CO
2

caused fruit injury, and resulted in the highest ethanol production rates.
Enclosure of Haden and Tommy Atkins mangoes in sealed 20 l jars with an
initial atmosphere of 90 kPa CO
2
in air or 97 kPa N
2
+ 3 kPa O
2
for 24 h prior
to storage delayed their ripening, and no injury was reported (Pesis et al.,
1994).
14.8 Manipulation of Mango Postharvest Physiology by Molecular
Biology
Mango fruit quality is compromised when harvest occurs before the fruit are
fully mature since they are unable to achieve the full complement of avour
and aroma during the postharvest handling period compared with fruit
harvested at a fully mature or ripening-initiated stage of development. As a
climacteric fruit, maturity in mango corresponds to attainment of ripening
competence. The presence of ethylene is required for the progression and com-
pletion of mango ripening. Thus, strategies for prolonging the postharvest
life and maintaining postharvest quality of mango other than disease control
are focused on reducing the effects of ethylene. This situation provides an
excellent opportunity to utilize genetic transformation to improve mango
postharvest quality by manipulating the role of ethylene (see Litz et al., Chap-
ter 18, this volume). Cruz-Hernndez et al. (1997) transformed Hindi mango
with mango ACO and ACC synthase (EC 4.4.1.14) in the antisense orienta-
tion. A cDNA that codes for mango ACO was also isolated and characterized
by Zainal et al. (1999). Suppression of mango ethylene biosynthesis should
allow harvesting of advanced maturity fruit that contain high levels of sug-
ars and possess enhanced capacity to produce ripe aroma volatiles after
exposure to exogenous ethylene. Progression of ripening in such fruit can be
easily halted at the most desirable and convenient time by simply removing
exogenous ethylene.
A cDNA homologue of the ethylene receptor gene ETR-1, referred to as
METR1, which codes for a polypeptide of 802 amino acids with a predicted
89 kDa MW has been isolated (Gutierrez-Martnez et al., 2001). Two or more
ETR homologues exist in mango. The level of METR1 mRNA in the mesocarp
increases transiently during wounding. Repression of genes involved in eth-
ylene action in mango fruit should result in ethylene-insensitive fruit that are
minimally affected by exposure to ethylene in the postharvest environment,
resulting in better control of ripening and senescence to maintain mango
postharvest quality.
Internal breakdown in mango fruit is essentially a problem of premature
ripening; the longer harvest of susceptible varieties is delayed, the greater
the incidence of internal breakdown. Using molecular approaches to reduce
ethylene production and action in mature fruit could reduce internal break-
down or premature ripening and lead to greatly improved quality. Another
approach to mitigating internal breakdown would be to target genes involved
in the maintenance of cell wall integrity. Vasanthaiah et al. (2006) demonstrated
J.K. Brecht and E.M. Yahia 516
differential expression of several genes in tissue showing internal breakdown
symptoms compared with healthy tissue. They suggested that oxidative
stress may be one of the causes of the disorder.
Sane et al. (2005) have isolated and characterized an ethylene-dependent
a-expansin gene, MiExpA1, which is correlated with softening in mango.
Expression of MiExpA1 increases with the progression of ripening and treat-
ment with 1-methyl cyclopropene inhibits both ripening/softening as well
as MiExpA1 transcript and protein accumulation. Recently, a pectate lyase
(EC 4.2.2.2) homologue from ripening mango (MiPel1) has been cloned (Chour-
asia et al., 2006). A progressive increase in transcript accumulation was observed
during ripening but expression was delayed signicantly in fruit in air without
exogenous ethylene. The expression was specic to fruit and was triggered
only during ripening. Increased transcript accumulation during ripening was
associated with pectin solubilization. Pectate lyase may be closely associated
with pectin degradation and have an important role in mango softening.
14.9 Conclusions
Mango fruit have the potential to develop extremely desirable texture, taste
and aroma that make this fruit highly appreciated and desirable. Strategies
used to extend mango shelf life are based on control of ripening, ethylene
action and ethylene production. Therefore, fruit are usually harvested at the
mature-green stage, prior to ripening initiation, and stored and transported
at low temperatures at or near the threshold for induction of chilling injury.
These practices result in poor quality, immature and chill-injured mangoes
on the market. Successful handling of ripening-initiated mangoes is prob-
lematic due to the fruits short shelf life and the increased incidence of inter-
nal breakdown that accompanies delayed harvests makes international
transport of ripening mangoes almost impossible. Consequently, the export
market for fresh mangoes, which expanded rapidly in the 1990s, has not con-
tinued its rapid expansion in recent years.
Future expansion of mango consumption will require an understanding
of mango postharvest physiology in order to overcome the problems of CI,
internal breakdown, and premature and uneven ripening. This may involve
increased transport of tree-ripe mangoes in CA-equipped marine containers
or in MAP. It may involve development of improved procedures for storage
and ripening to offer preconditioned, ripening-initiated, ready-to-eat man-
goes to consumers. It may also involve genetic transformation of mango to
manipulate the progression and uniformity of ripening and softening.
References
Abu-Sarra, A.F. and Abu-Goukh, A.A. (1992) Changes in pectinesterase, polygalactu-
ronase and cellulase activity during mango fruit ripening. Journal of Horticultural
Science 67, 561568.
Postharvest Physiology 517
Akamine, E.K. (1963) Haden mango storage. Hawaii Farm Science 12, 6.
Akamine, E.K. and Goo, T. (1973) Respiration and ethylene production during ontogeny
of fruit. Journal of the American Society for Horticultural Science 98, 381383.
Ali, Z.M., Abu, S. and Lazan, H. (1990) Glycosidases in ripening mango. Transactions
of the Malaysian Society of Plant Physiology 1, 1114.
Ali, Z.M., Armugam, S. and Lazan, H. (1995) -Galactosidase and its signicance in
ripening mango fruit. Phytochemistry 38, 11091114.
Allong, R., Wickham, L.D. and Mohammed, M. (2001) Effect of slicing on the rate of
respiration, ethylene production and ripening of mango fruit. Journal of Food Qual-
ity 24, 405419.
Ashraf, M., Khan, N., Ahmed, M. and Elahi, M. (1981) Studies on the pectinesterase
activity and some chemical constituents of some Pakistani mango varieties
during storage and ripening. Journal of Agricultural and Food Chemistry 29,
526528.
Baldwin, E.A., Burns, J.K., Kazokas, W., Brecht, J.K., Hagenmaier, R.D., Bender, R.J. and
Pesis, E. (1999) Effect of two edible coatings with different permeability character-
istics on mango (Mangifera indica L.) ripening during storage. Postharvest Biology
and Technology 17, 215226.
Bandyopadhyay, C. (1983) Contribution of gas chromatography to food avour research.
Pafai Journal 5, 2630.
Bandyopadhyay, C. and Gholap, A.S. (1973a) Changes in fatty acids in ripening mango
pulp (variety Alphonso). Journal of Agricultural and Food Chemistry 21, 496497.
Bandyopadhyay, C. and Gholap, A.S. (1973b) Relationship of aroma and avour charac-
ters of mango (Mangifera indica L.) to fatty acid composition. Journal of the Science
of Food and Agriculture 24, 14971503.
Baqui, S.M., Mattoo, A.K. and Modi, V.V. (1974) Mitochondrial enzymes in mango fruit
during ripening. Phytochemistry 13, 20492055.
Barmore, C.R. (1974) Ripening of mangos with ethylene and ethephon. Proceedings of
the Florida State Horticultural Society 87, 331334.
Barmore, C.R. and Mitchell, E.F. (1975) Ethylene pre-ripening of mangoes prior to ship-
ment. Proceedings of the Florida State Horticultural Society 88, 469471.
Bartley, J.P. and Schwede, A. (1987) Volatile avour components in the headspace of the
Australian or Bowen mango. Journal of Food Science 52, 353360.
Beaulieu, J.C. and Lea, J.M. (2003) Volatile and quality changes in fresh-cut mangos
prepared from rm-ripe and soft-ripe fruit, stored in clamshell containers and pas-
sive MAP. Postharvest Biology and Technology 30, 1528.
Ben-Amotz, A. and Fishler, R. (1998) Analysis of carotenoids with emphasis on 9-cis--
carotene in vegetables and fruits commonly consumed in Israel. Food Chemistry
62, 515520.
Bender, R.J., Brecht, J.K. and Campbell, C.A. (1994) Response of Kent and Tommy
Atkins mangoes to reduced O
2
and elevated CO
2
. Proceedings of the Florida State
Horticultural Society 107, 274277.
Bender, R.J., Brecht, J.K. and Sargent, S.A. (1995) Inhibition of ethylene production in
mango fruit by elevated CO
2
and recovery during subsequent air storage. Proceed-
ings of the Florida State Horticultural Society 108, 279285.
Bender, R.J., Brecht, J.K., Baldwin, E.A. and Malundo, T.M.M. (2000a) Aroma volatiles
of mature-green and tree-ripe Tommy Atkins mangoes after controlled atmosphere
vs. air storage. HortScience 35, 684686.
Bender, R.J., Brecht, J.K., Sargent, S.A. and Huber, D.J. (2000b) Mango tolerance to re-
duced oxygen levels in controlled atmosphere storage. Journal of the American
Society for Horticultural Science 125, 707713.
J.K. Brecht and E.M. Yahia 518
Biale, J.B. and Young, R.E. (1981) The avocado pear. In: Hulme, A.C. (ed.) The Biochem-
istry of Fruits and their Products. Vol. 2. Academic Press, New York, pp. 163.
Botting, K.J., Yong, M.M., Pearson, A.E., Harris, P.J. and Ferguson, L.R. (1999) Antimuta-
gens in food plants eaten by Polynesians: micronutrients, phytochemicals and pro-
tection against bacterial mutagenicity of heterocyclic amine 2-amino-3-
methylimidazo[4,5-f]quinoline. Food and Chemical Toxicology 37, 95103.
Brinson, K., Dey, P.M., John, M.A. and Pridham, J.B. (1988) Post-harvest changes in
Mangifera indica mesocarp cell walls and cytoplasmic polysaccharides. Phy-
tochemistry 27, 719723.
Brown, B.I., Scott, K.J. and Chaplin, G.R. (1981) Spongy stem end in mango. Queen-
sland Fruit and Vegetable News 52, 497.
Burdon, J., Dori, S., Marinansky, R. and Pesis, E. (1996) Acetaldehyde inhibition of ethyl-
ene biosynthesis in mango fruit. Postharvest Biology and Technology 8, 153161.
Burg, S.P. and Burg, E.A. (1962) Role of ethylene in fruit ripening. Plant Physiology 37,
179189.
Burns, J., Fraser, P.D. and Bramley, P.M. (2003) Identication and quantication of caro-
tenoids, tocopherols and chlorophylls in commonly consumed fruits and vegeta-
bles. Phytochemistry 62, 939947.
Cano, M.P. and de Ancos, B. (1994) Carotenoid and carotenoid ester composition in
mango fruit as inuenced by processing method. Journal of Agricultural and Food
Chemistry 42, 27372742.
Carlier, C., Etchepare, M., Ceccon, J.F., Mourey, M.S. and Amedee-Manesme, O. (1992)
Efcacy of massive oral doses of retinyl palmitate and mango (Mangifera indica L.)
consumption to correct an existing vitamin A deciency in Senegalese children.
British Journal of Nutrition 68, 529540.
Carrillo-Lpez, A., Rojas-Villagas, R. and Yahia, E.M. (1996) Ripening and quality of
mango fruit as affected by coating with Semperfresh. Acta Horticulturae 370,
206216.
Castrillo, M., Kruger, N.J. and Whatley, F.R. (1992) Sucrose metabolism in mango fruit
during ripening. Plant Science 84, 4551.
Chaplin, G.R. (1988) Advances in postharvest physiology of mango. Acta Horticulturae
231, 639648.
Chaplin, G.R., Scott, K.L. and Brown, B.I. (1982) Effect of storing mangoes in polyethylene
bags at ambient temperature. Singapore Journal of Primary Industries 10, 8488.
Chen, J.P., Tai, C.Y. and Chen, B.H. (2004) Improved liquid chromatographic method for
determination of carotenoids in Taiwanese mango (Mangifera indica L.). Journal of
Chromatography 1054, 261268.
Chourasia, A., Sane, V.A. and Nath, P. (2006) Differential expression of pectate lyase
during ethylene-induced postharvest softening of mango (Mangifera indica var.
Dashehari). Physiologia Plantarum 128, 546555.
Coates, L.M. and Johnson, G.I. (1993) Effective disease control in heat-disinfected fruit.
Postharvest News and Information 4, 35N40N.
Cocozza, F.M., Jorge, J.T., Alves, R.E., Filgueiras, H.A.C., Garruti, D.S. and Pereira,
M.E.C. (2004) Sensory and physical evaluations of cold stored Tommy Atkins
mangoes inuenced by 1-MCP and modied atmosphere packaging. Acta Horticul-
turae 645, 655661.
Considine, M.J., Daley, D.O. and Whelan, J. (2001) The expression of alternative oxi-
dase and uncoupling protein during fruit ripening in mango. Plant Physiology 126,
16191629.
Couey, M.H. (1986) Chilling injury in fruits of tropical and subtropical origin. Hort-
Science 17, 162165.
Postharvest Physiology 519
Cruz-Hernndez, A., Gmez-Lim, M.A. and Litz, R.E. (1997) Transformation of mango
somatic embryos. Acta Horticulturae 455, 292298.
Cua, A.U. and Lizada, M.C.C. (1990) Ethylene production in the Carabao mango
(Mangifera indica L.) fruit during maturation and ripening. Acta Horticulturae 269,
169179.
Dhalla, R. and Hanson, S.W. (1988) Effect of permeable coatings on the storage life of
fruits. II. Pro-long treatment of mangoes (Mangifera indica L., cv Julia). International
Journal of Food Science and Technology 23, 107112.
Dubery, I.A., Van Rensburg, L.J. and Schabort, J.C. (1984) Malic enzyme activity and
related biochemical aspects during ripening of -irradiated mango fruit. Phytochem-
istry 23, 13831386.
Engel, K.H. and Tressl, R. (1983) Studies on the volatile components of two mango vari-
eties. Journal of Agricultural and Food Chemistry 31, 796801.
Esguerra, E.B. and Lizada, M.C.C. (1990) The postharvest behavior and quality of Cara-
bao mangoes subjected to vapor heat treatment. ASEAN Food Journal 5, 612.
Esguerra, E.B., Mendoza, D.B., Jr and Pantastico, E.R.B. (1978) Regulation of fruit ripen-
ing. II. Use of perlite-KMnO
4
insert as an ethylene absorbent. Philippine Journal of
Science 107, 2331.
Farooqui, Q.A., Sattar, A., Daud, K. and Hussain, M. (1985) Studies on the postharvest
chilling sensitivity of mango fruit (Mangifera indica L.). Proceedings of the Florida
State Horticultural Society 98, 220221.
Fuchs, Y., Pesis, E. and Zauberman, C. (1980) Changes in amylase activity, starch and
sugar contents in mango fruit pulp. Scientia Horticulturae 13, 155160.
Garca-Sols, P., Yahia, E.M. and Aceves, C. (2008) Study of the effect of Ataulfo mango
(Mangifera indica L.) intake on mammary carcinogenesis and antioxidant capacity
in plasma of N-methyl-N-nitrosourea (MNU)-treated rats. Food Chemistry 111,
309315.
Gautam, D.M. and Lizada, M.C.C. (1984) Internal breakdown in Carabao mango sub-
jected to modied atmospheres. Storage duration and severity symptoms. Posthar-
vest Research Notes 1, 2830.
Gholap, A.S. and Bandyopadhyay, C. (1975a) Comparative assessment of aromatic prin-
ciples in ripe Alphonso and Langra mangoes. Journal of Agricultural and Food
Science 12, 262263.
Gholap, A.S. and Bandyopadhyay, C. (1975b) Contribution of lipid to aroma of ripening
mango. Journal of the American Oil Chemistry Society 52, 514516.
Gholap, A.S. and Bandyopadhyay, C. (1976) Fatty acid composition as a quality index of
ripe mango (Mangifera indica L.) pulp. Indian Food Packer 30, 6364.
Gholap, A.S. and Bandyopadhyay, C. (1980) Fatty acid biogenesis in ripening mango
(Mangifera indica cv. Alphonso). Journal of Agricultural and Food Chemistry 28,
839841.
Godoy, H.T. and Rodriguez-Amaya, D.B. (1989) Carotenoid composition of commercial
mangoes from Brazil. Lebensmittel-Wissenschaft und Technologie 22, 100103.
Godoy, H.T. and Rodriguez-Amaya, D.B. (1994) Occurrence of cis-isomers of provitamin
A in Brazilian fruits. Journal of Agricultural and Food Chemistry 42, 13061313.
Gofur, M.A., Shaque, M. and Helali, O.H. (1994) Effect of various factors on the vita-
min C (ascorbic acid) content of some mango varieties grown in Rajshahi region.
Bangladesh Journal of Scientic and Industrial Research 29, 163171.
Gonzalez, G., Yahia, E.M. and Higuera, I. (1990) Modied atmosphere packaging (MAP)
of mango and avocado fruit. Acta Horticulturae 269, 335344.
Gonzlez-Aguilar, G., Gardea, A., Martinez-Tellez, M.A., Baez, R. and Felix, L. (1997)
Low oxygen treatment before storage in normal or modied atmosphere packaging
J.K. Brecht and E.M. Yahia 520
of mangoes to extend shelf life. Journal of Food Science and Technology 34,
399404.
Gonzlez-Aguilar, G.A., Buta, J.G. and Wang, C.Y. (2001) Methyl jasmonate reduces
chilling injury symptoms and enhances colour development of Kent mangoes.
Journal of the Science of Food and Agriculture 81, 12441249.
Gosh, S. (1960) The content of folic acid and its conjugates in some common Indian
fruits. Science and Culture 26, 287292.
Gowda, I.N.D. and Huddar, A.G. (2000) Evaluation of mango hybrids for storage behav-
iour and sensory qualities. Journal of Food Science and Technology (Mysore) 37,
620623.
Gowda, I.N.D., Ramanjaneya, K.H., Iyer, C.P.A., Subramanyam, M.D. and Dinesh, M.R.
(1994) Physico-chemical and processing quality of four new mango hybrids in
comparison to two commercial cultivars. Journal of Food Science and Technology
(Mysore) 31, 385388.
Gutierrez-Martnez, P., Lpez Gmez, R. and Gmez-Lim, M.A. (2001) Identication of
an ETR1-homologue from mango fruit expressing during fruit ripening and wound-
ing. Journal of Plant Physiology 158, 101108.
Hatton, T.T., Jr, Reeder, W.F. and Campbell, C.W. (1965) Ripening and Storage of Florida
Mangoes. Marketing Research Report 725. Agricultural Research Service, United
States Department of Agriculture (USDA), Washington, DC.
Heather, N.W. (1994) Quarantine disinfestation of tropical fruits: non-chemical options.
In: Champ, B.C., Highley, E. and Johnson, G.I. (eds) Postharvest Handling of Tropi-
cal Fruits. Australian Centre for International Agricultural Research (ACIAR) Pro-
ceedings 50. ACIAR, Canberra, pp. 272279.
Iguina de George, L.M., Collazo de Rivera, A.L., Benero, J.R. and Pennock, W. (1969)
Provitamin A and vitamin C contents of several varieties of mango (Mangifera in-
dica L.) grown in Puerto Rico. Journal of Agriculture of the University of Puerto Rico
53, 100105.
Ito, T., Sasaki, K. and Yoshida, Y. (1997) Changes in respiration rate, saccharide and or-
ganic acid content during the development and ripening of mango fruit (Mangifera
indica L. Irwin) cultured in a plastic house. Journal of the Japanese Society for
Horticultural Science 66, 629635.
Jacobi, K.K. and Giles, J.E. (1997) Quality of Kensington mango (Mangifera indica
Linn.) fruit following combined vapour heat disinfestation and hot water disease
control treatments. Postharvest Biology and Technology 12, 285292.
Jacobi, K.K. and Wong, L.S. (1992) Quality of Kensigton mango (Mangifera indica L.)
following hot water and vapour-heat treatments. Postharvest Biology and Technol-
ogy 1, 349359.
Jacobi, K.K., Coates, L.M. and Wong, L.S. (1994) Heat disinfestation of mangoes: effect
on fruit quality and disease control. In: Champ, B.R., Haghley, E. and Johnson, G.I.
(eds) Postharvest Handling of Tropical Fruits. Australian Centre for International
Agricultural Research (ACIAR) Proceedings 50. ACIAR, Canberra, pp. 280287.
Jacobi, K.K., MacRae, E.A. and Hetherington, S.E. (2001a) Effect of fruit maturity on the
response of Kensington mango fruit to heat treatment. Australian Journal of Ex-
perimental Agriculture 41, 793803.
Jacobi, K.K., MacRae, E.A. and Hetherington, S.E. (2001b) Postharvest heat disinfesta-
tion treatments of mango fruit. Scientia Horticulturae 89, 171193.
Jain, N.L. (1961) Chemistry and technology of mango. Review of Food Technology 3,
131162.
Jha, S.N., Chopra, S. and Kingsly, A.R.P. (2007) Modeling of color values for non-
destructive evaluation of maturity of mango. Journal of Food Engineering 78, 2226.
Postharvest Physiology 521
John, J., Subbarayan, C. and Cama, H.R. (1970) Carotenoids in three stages of ripening
of mango. Journal of Food Science 35, 262265.
Johnson, G.I. and Coates, L.M. (1993) Postharvest diseases of mango. Postharvest News
and Information 4, 27N34N.
Johnson, G.I., Sharp, J.L., Milne, D.L. and Oosthuyse, S.A. (1997) Postharvest technol-
ogy and quarantine treatments. In: Litz, R. (ed.) The Mango: Botany, Production and
Uses. CAB International, Wallingford, UK, pp. 447507.
Jungalwala, F.B. and Cama, H.R. (1963) Carotenoids in mango (Mangifera indica) fruit.
Indian Journal of Chemistry 1, 3640.
Kalra, S.K. and Tandon, D.K. (1983) Ripening behavior of Dashehari mango in relation
to harvest period. Scientia Horticulturae 19, 263269.
Kane, O. (1977) Recherches sur lvolution normale ou pathologique de mangues
(Mangifera indica L.) conserves au froid dans lair ou en atmosphre contrle.
Thse doct 3me cycle. Universit Pierre et Marie Curie, Paris, 124 pp.
Kapse, B.M., Rane, D.A. and Khedkar, D.M. (1989) Correlation between bio-chemical
parameters and organoleptic evaluation in mango varieties. Acta Horticulturae
231, 756762.
Karmarkar, D.V. and Joshi, B.M. (1941) Respiration studies of the Alphonso mango.
Indian Journal of Agricultural Science 11, 9931005.
Ketsa, S., Rattanamalee, S. and Babprasert, C. (1991) Growth, development, biochemi-
cal changes and harvesting index of mango (Mangifera indica L.) cv. Tongdum.
Kasetsart Journal (Natural Science) 25, 391399.
Ketsa, S., Phakawatmongkol, W. and Subhadrabhandhu, S. (1999) Peel enzymatic activ-
ity and colour changes in ripening mango fruit. Journal of Plant Physiology 154,
363366.
Kostermans, A.J.G.H. and Bompard, J.M. (1993) The Mangoes: Botany, Nomenclature,
Horticulture and Utilization. Academic Press, London, 249 pp.
Krishnamurthy, S. and Subramanyam, H. (1970) Respiratory climacteric and chemical
changes in the mango fruit Mangifera indica L. Journal of the American Society for
Horticultural Science 95, 333337.
Krishnamurthy, S. and Subramanyam, H. (1973) Pre and postharvest physiology of the
mango fruit. Tropical Science 15, 167193.
Krishnamurthy, S., Patwardhan, M.V. and Subramanyan, H. (1971) Biochemical changes
during ripening of the mango fruit. Phytochemistry 10, 25772581.
Kumar, R. and Selvaraj, Y. (1990) Fructose-1,6-biophosphatase in ripening mango
(Mangifera indica L.) fruit. Indian Journal of Experimental Biology 28, 284287.
Kumar, S., Das, D.K., Singh, A.K. and Prasad, U.S. (1993) Changes in non-volatile or-
ganic acid composition and pH during maturation and ripening of two mango cul-
tivars. Indian Journal of Plant Physiology 36, 107111.
Kumar, S., Das, D.K., Singh, A.K. and Prasad, U.S. (1994) Sucrose metabolism during matu-
ration and ripening of mango cultivars. Plant Physiology and Biochemistry 21, 2732.
Kundu, S. and Ghosh, S.N. (1992) Studies on physico-chemical characteristics of mango
cultivars grown in the laterite tract of West Bengal. Haryana Journal of Horticul-
tural Sciences 21(3/4), 129139.
Lakshminarayana, S. (1973) Respiration and ripening patterns in the life cycle of the
mango fruit. Journal of Horticultural Science 48, 227233.
Lakshminarayana, S. (1975) Relation of time of harvest on respiration, chemical con-
stituents and storage life of mangoes. Proceedings of the Florida State Horticultural
Society 88, 477480.
Lakshminarayana, S. (1980) Mango. In: Nagy, S. and Shaw, P.E. (eds) Tropical and Sub-
tropical Fruits. AVI Publishing Co., Westport, Connecticut, pp. 184257.
J.K. Brecht and E.M. Yahia 522
Lakshminarayana, S. and Subramanyam, H. (1970) Carbon dioxide injury and fermenta-
tive decarboxylation in mango fruit at low temperature storage. Journal of Food
Science and Technology 7, 148152.
Lakshminarayana, S., Subhadra, N.V. and Subramanyam, H. (1970) Some aspects of the
developmental physiology of the mango fruit. Journal of Horticultural Science 45,
133142.
Lakshminarayana, S., Shety, M.S. and Krishnaprasad, C.A. (1974) Accelerated ripening
of Alphonso mangos by the application of ethrel. Tropical Science 17, 95101.
Lalel, H.J.D., Singh, Z. and Tan, S.C. (2003) Maturity stage at harvest affects fruit ripen-
ing, quality and biosynthesis of aroma volatile compounds in Kensington Pride
mango. Journal of Horticultural Science and Biotechnology 78, 225233.
Lazan, H., Ali, Z.M., Lee, K.W. and Woon, J. (1986) The potential role of polygalactu-
ronase in pectin degradation of softening of mango. ASEAN Food Journal 2, 9398.
Lazan, H., Ali, Z.M. and Sani, H.A. (1990) Effect of Vaporgard on polygalacturonase,
malic enzyme and ripening in mango. Acta Horticulturae 269, 359366.
Lazan, H., Ali, Z.M., Soh, J. and Talkan, Z. (1993) The biochemical basis of differential
ripening in mango. Acta Horticulturae 341, 500509.
Lederman, I.E., Zauberman, G., Weksler, A., Rot, I. and Fuchs, Y. (1997) Ethylene-forming
capacity during cold storage and chilling injury development in Keitt mango fruit.
Postharvest Biology and Technology 10, 107112.
Ledger, S.N. (1986) Mango packaging. In: Proceedings of the First Australian Mango
Research Workshop. Commonwealth Scientic and Industrial Research Organiza-
tion (CSIRO), Melbourne, pp. 314319.
Lizada, C. (1993) Mango. In: Seymour, G.B., Taylor, J.E. and Tucker, G.A. (eds) Biochem-
istry of Fruit Ripening. Chapman and Hill, London, pp. 255271.
Lizada, M.C. (1991) Postharvest physiology of the mango a review. Acta Horticulturae
219, 437452.
MacLeod, A.J. and De Troconis, N.G. (1982) Volatile components of some mango fruit.
Phytochemistry 21, 25232526.
MacLeod, A.J. and Pieris, N.M. (1984) Comparison of volatile components in some
mango cultivars. Phytochemistry 23, 361366.
MacLeod, A.J. and Snyder, C.H. (1985) Volatile components of two cultivars of mango
from Florida. Journal of Agricultural and Food Chemistry 33, 380384.
MacLeod, A.J., MacLeod, G. and Snyder, C.H. (1988) Volatile aroma constituents of
mango (cv. Kensington). Phytochemistry 27, 21892193.
Mahayothee, B., Muhlbauer, W., Neidhart, S. and Carle, R. (2004) Inuence of posthar-
vest ripening process on appropriate maturity for drying mangoes Nam Dokmai
and Kaew. Acta Horticulturae 645, 241248.
Malundo, T.M.M. (1996) Application of the quality enhancement (QE) approach to man-
go (Mangifera indica L.) avor research. PhD dissertation, Department of Food Sci-
ence and Technology, University of Georgia, Athens, Georgia, USA, 134 pp.
Malundo, T.M.M., Shewfelt, R.L., Ware, G.O. and Baldwin, E.A. (2001) Sugars and acids
inuence avor properties of mango (Mangifera indica). Journal of the American
Society for Horticultural Science 126, 115121.
Mann, S.S. and Singh, R.N. (1976) The cold storage life of Deshari mangoes. Scientic
Horticulture 5, 249254.
Mattoo, A.K. and Modi, V.V. (1969a) Biochemical aspects of ripening and chilling injury
in mango fruit. Proceedings of the International Conference on Tropical and Sub-
tropical Fruit. Tropical Products Institute, London, pp. 111115.
Mattoo, A.K. and Modi, V.V. (1969b) Ethylene and ripening of mangos. Plant Physiology
44, 308310.
Postharvest Physiology 523
Mattoo, A.K., Modi, V.V. and Reddy, V.V.R. (1968) Oxidation and carotenogenesis regu-
lating factors in mangoes. Indian Journal of Biochemistry 5, 111114.
McCollum, T.G., DAquino, S. and McDonald, R.E. (1993) Heat treatment inhibits man-
go chilling injury. HortScience 28, 197198.
Medlicott, A.P. and Thompson, A.K. (1985) Analysis of sugar and organic acids in ripen-
ing mango fruit (Mangifera indica var. Keitt) by high performance liquid chromatog-
raphy. Journal of the Science of Food and Agriculture 36, 561566.
Medlicott, A.P., Bhogal, M. and Reynolds, S.B. (1986) Changes in peel pigmentation
during ripening of mango fruit (Mangifera indica var. Tommy Atkins). Annals of Ap-
plied Biology 109, 651656.
Medlicott, A.P., Reynolds, S.B., New, S.W. and Thompson, A.K. (1988) Harvest maturity
effects on mango fruit ripening. Tropical Agriculture 65, 153157.
Medlicott, A.P., Sigrist, J.M.M. and Sy, O. (1990) Ripening of mangoes following low
temperature storage. Journal of the American Society for Horticultural Science 115,
430434.
Mercadante, A.Z. and Rodriguez-Amaya, D.B. (1998) Effects of ripening, cultivar differ-
ences and processing on the carotenoid composition of mango. Journal of Agricul-
tural and Food Chemistry 46, 128130.
Mercadante, A.Z., Rodriguez-Amaya, D.B. and Britton, G. (1997) HPLC and mass spec-
trometric analysis of carotenoids from mango. Journal of Agricultural and Food
Chemistry 45, 120123.
Miller, W.R., Hale, P.W., Spalding, D.H. and Davis, P. (1983) Quality and decay of
mango fruit wrapped in heat-shrinkable lm. HortScience 18, 957958.
Miller, W.R., Spalding, D.R. and Hale, P.W. (1986) Film wrapping mangoes at advancing
stages of postharvest ripening. Tropical Science 26, 917.
Mitcham, E.J. and McDonald, R.E. (1992) Cell wall modication during ripening of
Keitt and Tommy Atkins mango fruit. Journal of the American Society for Horti-
cultural Science 117, 919924.
Mitcham, E.J. and McDonald, R.E. (1997) Effects of postharvest heat treatments on inner
and outer tissue of mango fruit. Tropical Science 37, 193205.
Mitra, S.K. and Baldwin, E.A. (1997) Mango. In: Mitra, S.K. (ed.) Postharvest Physiology
and Storage of Tropical and Subtropical Fruits. CAB International, Wallingford, UK,
pp. 85122.
Modi, V.V. and Reddy, V.V.R. (1967) Carotenogenesis in ripening mangoes. Indian Jour-
nal of Experimental Biology 5, 233235.
Mohammed, M. and Brecht, J.K. (2002) Reduction of chilling injury in Tommy Atkins
mangoes during ripening. Scientia Horticulturae 95, 297308.
Morga, N.S., Lustre, A.O., Tunac, M.M., Balagot, A.H. and Soriano, M.R. (1979) Physi-
cochemical changes in Philippine Carabao mangoes during ripening. Food Chem-
istry 4, 225234.
Muda, P., Seymour, G.B., Errington, N. and Tucker, G.A. (1995) Compositional changes in
cell wall polymers during mango fruit ripening. Carbohydrate Polymers 26, 255260.
Mukherjee, S.K. (1958) The origin of mango. Indian Journal of Horticulture 15, 129134.
Musa, S.K. (1974) Preliminary investigations on the storage and ripening of Totapuri
mangoes in the Sudan. Tropical Science 16, 6573.
Nasrijal, N.H. (1993) Effect of storage temperature on pectin depolymerization and soft-
ening of mango fruit. BSc thesis, Department of Botany, Universiti Kebangsaan,
Bangi, Malaysia.
Noomhorm, A. and Tiasuwan, N. (1995) Controlled atmosphere storage of mango
fruit, Mangifera indica L. cv. Rad. Journal of Food Processing and Preservation 19,
271281.
J.K. Brecht and E.M. Yahia 524
Nuevo, P.A., Cua, A.U. and Lizada, M.C.C. (1984a) Internal breakdown in Carabao
mango subjected to modied atmospheres. III. Starch in the spongy tissue. Posthar-
vest Research Notes 1, 6364.
Nuevo, P.A., Pantastico, E.R.B. and Mendosa, D.B. (1984b) Gas diffusion factors in fruits
I. Anatomical structure in mango fruit. Postharvest Research Notes 1, 15.
OHare, T.J. and Prasad, A. (1993) The effect of temperature and carbon dioxide on chill-
ing symptoms in mango. Acta Horticulturae 343, 244250.
Ornelas-Paz, J. de J., Yahia, E.M. and Gardea, A. (2007) Identication and quantication
of xanthophyll esters, carotenes and tocopherols in the fruit of seven Mexican man-
go cultivars by liquid chromatography-APcI
+
-time of ight mass spectrometry. Jour-
nal of Agricultural and Food Chemistry 55, 66286635.
Ornelas-Paz, J. de J., Yahia, E.M. and Gardea, A.A. (2008) Changes in external and
internal color during postharvest ripening of Manila and Ataulfo mango fruit
and relationship with carotenoid content determined by liquid chromatography
APcI+-time-of-flight mass spectrometry. Postharvest Biology and Technology
50, 145152.
Ortega, D. and Yahia, E.M. (2000) Tolerance and quality of mango fruit exposed to con-
trolled atmospheres at high temperatures. Postharvest Biology and Technology 20,
195201.
Parikh, H.R., Nair, G.M. and Modi, V.V. (1990) Some structural changes during ripening
of mangoes (Mangifera indica var. Alphonso) by abscisic acid treatment. Annals of
Botany 65, 121127.
Park, Y.K., Sato, H.H., Almeida, T.D. and Moretti, R.H. (1980) Polyphenoloxidase of
mango (Mangifera indica var. Haden). Journal of Food Science 45, 16191621.
Peacock, B.C. (1986) Postharvest handling of mangoes. In: Proceedings of the First
Australian Mango Research Workshop. Commonwealth Scientic and Industrial
Research Organization (CSIRO), Melbourne, pp. 295313.
Peacock, B.C., Murray, C., Kosiyachinda, S., Kasittrakakul, M. and Tansiriyakul, S. (1986)
Inuence of harvest maturity of mangoes on storage potential and ripe fruit quality.
ASEAN Food Journal 2, 99103.
Percival, S.S., Talcott, S.T., Chin, S.T., Mallak, A.C., Lounds-Singleton, A. and Pettit-
Moore, J. (2006) Neoplastic transformation of BALB/3T3 cells and cell cycle of
HL-60 cells are inhibited by mango (Mangifera indica L.) juice and mango juice
extract. Journal of Nutrition 136, 13001304.
Pesis, E., Marinansky, R., Rot, I. and Weksler, A. (1994) Effect of postharvest application
of acetaldehyde or a short period of anaerobiosis on fruit senescence. Proceedings
of the Royal Society Edinburgh 102b, 473478.
Pino, J., Rosado, A. and Sanchez, R. (1989) Volatile components of three cultivars of
mango from Cuba. Die Nahrung 33, 709715.
Pino, J., Mesa, J., Muoz, Y., Mart, M.P. and Marbot, R. (2005) Volatile components
from mango (Mangifera indica L.) cultivars. Journal of Agricultural and Food Chem-
istry 53, 22132223.
Pott, I., Breithaupt, D.E. and Carle, R. (2003a) Detection of unusual carotenoid esters in
fresh mango (Mangifera indica L. cv. Kent). Phytochemistry 64, 825829.
Pott, I., Marx, M., Neidhart, S., Mhlbauer, W. and Carle, R. (2003b) Quantitative deter-
mination of -carotene stereoisomers in fresh, dried, and solar-dried mango (Mangifera
indica L.). Journal of Agricultural and Food Chemistry 51, 45274531.
Proctor, J.T.A. and Creasy, L.L. (1969) The anthocyanins of the mango fruit. Phytochem-
istry 8, 2108.
Quinones, V.L., Guerrant, N.B. and Adams Dutcher, R. (1944) Vitamin content of some
tropical fruits, their juices and nectars. Journal of Food Science 9, 415417.
Postharvest Physiology 525
Raymond, L., Schaffer, B., Brecht, J.K. and Crane, J.H. (1998) Internal breakdown in
mango fruit: symptomology and histology of jelly seed, soft nose and stem-end cav-
ity. Postharvest Biology and Technology 13, 5970.
Reddy, Y.V. and Srivastava, G.C. (1999) Ethylene biosynthesis and respiration in mango
fruits during ripening. Indian Journal of Plant Physiology 4, 3235.
Rocha-Ribeiro, S.M., Queiroz, J.H., Lopes, M., Campos, F.M. and Pinheiro, H.M. (2007)
Antioxidant in mango (Mangifera indica L.) pulp. Plant Foods for Human Nutrition
62, 1317.
Rodov, V., Ben Yehoshua, S., Fishman, S., Gotlieb, S., Fierman, T. and Fang, D.Q. (1994)
Reducing decay and extending shelf life of bell peppers and mangoes by modied
atmosphere packaging. In: Champ, B.C., Highley, E. and Johnson, G.I. (eds) Post-
harvest Handling of Tropical Fruits. Australian Centre for International Agricultural
Research (ACIAR) Proceedings 50. ACIAR, Canberra, pp. 416418.
Roe, B. and Bruemmer, J.H. (1981) Changes in pectic substances and enzymes during
ripening and storage of Keitt mangos. Journal of Food Science 46, 186189.
Rojas-Villegas, A., Carrillo-Lopez, A., Silveira, M., Avena-Bustillos, R. and Yahia, E.M.
(1996) Effects of insecticidal atmospheres on the mortality of fruits ies in mango.
Acta Horticulturae 370, 8992.
Sadasivam, R., Muthuswany, S., Sundaraj, J.S. and Vasudevan, V. (1971) Note on chilling
injury in mango (Mangifera indica L.) fruits in refrigerated storage. Indian Journal of
Agricultural Science 41, 715716.
Sane, V.A., Chourasia, A. and Nath, P. (2005) Softening in mango (Mangifera indica cv.
Dashehari) is correlated with the expression of an early ethylene responsive, rip-
ening related expansin gene, MiExpA1. Postharvest Biology and Technology 38,
223230.
Sarker, S. and Muhsi, A.A. (1981) A study on the content and interconversions of or-
ganic acids of mango (Mangifera indica L.) at various stages of fruit development.
Bangladesh Journal of Agricultural Science 8, 6975.
Selvaraj, Y. and Kumar, R. (1989) Studies on fruit softening enzyme and polyphenoloxi-
dase activity in ripening mango (Mangifera indica L.) fruit. Journal of Food Science
and Technology 26, 218222.
Selvaraj, Y. and Kumar, R. (1994) Enzymatic regulation in ripening mango fruit. Indian
Journal of Horticulture 51, 316323.
Selvaraj, Y., Kumar, R. and Pal, D.K. (1989) Changes in sugars, organics acids, amino
acids, lipid constituents and aroma characteristics of ripening mango (Mangifera
indica L.) fruit. Journal of Food Science and Technology 26, 306311.
Setiawan, B., Sulaeman, A., Giraud, D.W. and Driskell, J.A. (2001) Carotenoid content of
selected Indonesian fruits. Journal of Food Composition and Analysis 14, 169176.
Seymour, G.B., NDiaye, M., Wainwright, H. and Tucker, G.A. (1990) Effect of cultivar
and harvest maturity on ripening of mangoes during storage. Journal of Horticul-
tural Science 65, 479483.
Shashirekha, M.S. and Patwardhan, M.V. (1976) Changes in amino acids, sugars and
nonvolatile organic acids in a ripening mango fruit (Mangifera indica L. Badami
variety). Lebensmittel-Wissenschaft und Technologie 9, 369370.
Siddappa, G.S. and Bhatia, B.S. (1954) Tender green mangoes as source of vitamin C.
Indian Journal for Horticulture Science 11, 104106.
Singh, L.B. (1960) The Mango. Leonard Hill, London.
Sornsrivichai, J. and Waru-Aswapti, O. (1989) The effect of maturity and ripening tem-
perature on fruit quality of ethrel treated mangoes. (Abstract). In: Proceedings of
the Third International Mango Symposium, Darwin, Northern Territory, Australia,
p. 25.
J.K. Brecht and E.M. Yahia 526
Sornsrivichai, S., Gomolmanee, S., Boonyakiat, D., Uthaibutra, J., Boon-Long, P. and
Gemma, H. (1992) Seal packaging by plastic lm as a technique for limiting fungal
decay of mangoes. Acta Horticulturae 296, 2332.
Srivastava, H.C. (1967) Grading, storage and marketing. In: The Mango: a Handbook.
Indian Council of Agricultural Research, New Delhi, India, pp. 99109.
Stahl, A.L. (1935) Composition of Miscellaneous Tropical and Subtropical Florida Fruits.
Bulletin 283. University of Florida Agricultural Experiment Station, Gainesville,
Florida.
Stead, D.E. and Chithambo, G.S.G. (1980) Studies on the storage of tropical fruits in
polyethylene bags. Luso: Journal of Science and Technology (Malawi) 1, 39.
Subramanyam, H., Krishnamurthy, S., Subhadra, N.V., Dalal, V.B., Randhawa, G.S. and
Chacko, E.K. (1971) Studies on internal breakdown, a physiological ripening disor-
der in Alphonso mangoes (Mangifera indica L.). Tropical Science 13, 203210.
Subramanyam, H., Krishnamurthy, S. and Parpia, H.A.B. (1975) Physiology and bio-
chemistry of mango fruit. Advances in Food Research 21, 223305.
Sy, D.A. and Mendoza, D.B., Jr (1984) Respiratory responses of Carabao mango to dif-
ferent levels of oxygen. Postharvest Research Notes 1, 34.
Tahir, M.A. and Malik, A.A. (1977) The activity of pectic enzymes (pectinesterase, polyga-
lacturonase) in various mango products. Pakistan Journal of Research 29, 7174.
Tandon, D.K. and Kalra, S.K. (1983) Changes in sugars, starch and amylase activity dur-
ing development of mango fruit cv. Dashehari. Journal of Horticultural Science 58,
449453.
Tandon, D.K. and Kalra, K. (1984) Pectin changes during the development of mango
fruit cv. Dashehari. Journal of Horticultural Science 59, 283286.
Tharanathan, R.N., Yashoda, H.M. and Prabha, T.N. (2006) Mango (Mangifera indica L.),
the king of fruits an overview. Food Reviews International 22, 95123.
Thomas, P. (1975) Effect of postharvest temperatures on quality, carotenoids and ascor-
bic acid contents in Alphonso mangos on ripening. Journal of Food Science 40,
704706.
Tirmazi, S.I.H. and Wills, R.B.H. (1981) Retardation of ripening of mangoes by posthar-
vest application of calcium. Tropical Agriculture 58, 137141.
United States Department of Agriculture (USDA)/Agricultural Research Service (ARS)
(2007) USDA National Nutrient Database for Standard Reference, Release 20.
Nutrient Data Laboratory Home Page. Available at: http://www.ars.usda.gov/nutri-
entdata (accessed 28 August 2008).
van Eeden, S.J. (1992) Calcium inltration as a possible postharvest treatment to in-
crease storage potential of mango fruit. South African Mango Growers Association
Yearbook 12, 2627.
van Lelyveld, L.J. and Smith, J.H.E. (1979) Physiological factor in maturation and ripen-
ing of mango (Mangifera indica) fruit in relation to the jelly seed physiological dis-
order. Journal of Horticultural Science 54, 283287.
Vasanthaiah, H.K.N., Ravishankar, K.V., Shivashankara, K.S., Kodthalu, S., Anand, L.,
Narayanaswamy, P., Mukunda, G. and Prasad, T.G. (2006) Cloning and character-
ization of differentially expressed genes of internal breakdown in mango fruit
(Mangifera indica). Journal of Plant Physiology 163, 671679.
Vzquez-Caicedo, A.L., Neidhart, S. and Carle, R. (2004) Postharvest ripening behav-
iour of nine Thai mango cultivars and their suitability for industrial applications.
Acta Horticulturae 645, 617625.
Vazquez-Salinas, C. and Lakshminarayana, D. (1985) Compositional changes in mango
fruit during ripening at different storage temperatures. Journal of Food Science 50,
16461648.
Postharvest Physiology 527
Vinci, G., Botre, F. and Mele, G. (1995) Ascorbic acid in exotic fruits: a liquid chromato-
graphic investigation. Food Chemistry 53, 211214.
Wilberg, V.C. and Rodriguez-Amaya, D.B. (1995) HPLC quantitation of major carote-
noids of fresh and processed guava, mango and papaya. Lebensmittel-Wissenschaft
und Technologie 28, 474480.
Wills, R.B.H., Yuen, C.M.C., Sabri Laksmi, L.D.S. and Suyanti (1988) Effect of calcium
inltration on delayed ripening of three mango cvs in Indonesia. ASEAN Food Jour-
nal 4, 6758.
Wills, R.B.H., Warton, M.A., Mussa, D.M.D.N. and Chew, L.P. (2001) Ripening of cli-
macteric fruits initiated at low ethylene levels. Australian Journal of Experimental
Agriculture 41, 8992.
Wilson, C.W., Shaw, P.E. and Knight, R.J. (1990) Importance of some lactones and 2,5
dimethyl-4-hydroxy-3(2H)-furanone to mango (Mangifera indica L.) aroma. Journal
of Agricultural and Food Chemistry 38, 15561559.
Wolf, G. (1984) Multiple functions of vitamin A. Physiological Reviews 64, 873937.
Yahia, E.M. (1993) Responses of some tropical fruits to insecticidal atmospheres. Acta
Horticulturae 343, 371376.
Yahia, E.M. (1994) The potential use of insecticidal atmospheres for mango, avocado,
and papaya fruits. In: Champ, B.C., Highley, E. and Johnson, G.I. (eds) Postharvest
Handling of Tropical Fruits. Australian Centre for International Agricultural Re-
search (ACIAR) Proceedings 50. ACIAR, Canberra, pp. 373374.
Yahia, E.M. (1995) Application of differential scanning calorimetry in the study of avo-
cado and mango fruit responses to hypoxia. In: Ait-Oubahou, A. and El-Otmani, M.
(eds) Postharvest Physiology, Pathology and Technologies for Horticultural Crops:
Recent Advances. Institut Agronomique and Vtrinaire Hassan II, Agadir, Moroc-
co, pp. 206209.
Yahia, E.M. (1997) Modied/controlled atmospheres for mango. In: Kader, A.A. (ed.)
Proceedings of CA97, Vol. 3: Fruits Other Than Apples and Pears. University of
California, Davis, California, pp. 110116.
Yahia, E.M. (1998) Modied and controlled atmospheres for tropical fruits. Horticultural
Reviews, 22, 123183.
Yahia, E.M. and Tiznado, M. (1993) Tolerance and responses of harvested mango to in-
secticidal oxygen atmospheres. HortScience 28, 10311033.
Yahia, E.M. and Vazquez, L. (1993) Tolerance and responses of mango to insecticidal
oxygen and carbon dioxide atmospheres. Food Science and Technology (Lebensmittel-
Wissenschaft und Technologie) 26, 4248.
Yahia, E.M., Medina, F. and Rivera, M. (1989) The tolerance of mango and papaya to
atmospheres containing very high levels of CO
2
and/or very low levels of O
2
as a
possible insect control treatment. In: Fellman, J.K. (ed.) Fifth Proceedings of Interna-
tional CA Research Conference, Vol. 2 Other Commodities and Storage Recom-
mendations, Wenatchee, Washington, pp. 7789.
Yahia, E.M., Villagomez, G. and Juarez, A. (2000) Tolerance and responses of different
fruits and vegetables to hot air treatments at 43C or 48C and 50% RH for 160
minutes. In: Artes, F., Gil, M.I. and Conesa, M.A. (eds) Improving Postharvest Tech-
nologies of Fruits, Vegetables and Ornamentals. Vol. 2. International Institute of
Refrigeration, Paris, pp. 714718.
Yahia, E.M., Ornelas-Paz, J. de J. and Ariza, F.R. (2006a) The Mango. Editorial Trillas,
Mxico City, Mexico, 224 pp. (in Spanish)
Yahia, E.M., Ornelas-Paz, J. de J. and Gardea, A. (2006b) Extraction, separation and
partial identication of Ataulfo mango fruit carotenoids. Acta Horticulturae 712,
333338.
J.K. Brecht and E.M. Yahia 528
Yamashita, F., Benassi, M.T. and Kieckbusch, T.G. (1997) Shelf life extension of individu-
ally lm-wrapped mangoes. Tropical Science 37, 249255.
Yamashita, F., Benassi, M.T. and Kieckbusch, T.G. (1999) Effect of modied atmosphere
packaging on the kinetics of vitamin C degradation in mangos. Brazilian Journal of
Food Technology 2, 127130.
Yantarasri, T., Uthaibutra, J., Sornrivichai, J., Kumpuan, W., Sardsud, V. and Kanathum,
N. (1994) Modied atmosphere packaging by perforated polymeric lm and its
effect on physical properties of mango fruit. In: Champ, B.R., Highley, E. and
Johnson, G.I. (eds) Postharvest Handling of Tropical Fruits. Australian Centre for
International Agricultural Research (ACIAR) Proceedings 50. ACIAR, Canberra,
pp. 438440.
Yantarasri, T., Ben Yehoshua, S., Rodov, V., Kumpuan, W., Uthaibutra, J. and Sornsrivichal,
J. (1995) Development of perforated modied atmosphere package for mango. Acta
Horticulturae 398, 8191.
Young, T.W. (1957) Soft nose a physiological disorder in mango fruit. Proceedings of
the Florida State Horticultural Society 70, 280283.
Yuen, C.M.C., Tan, S.C., Joyce, D. and Chettri, P. (1993) Effect of postharvest calcium
and polymeric lms on ripening and peel injury in Kensington Pride mangoes.
ASEAN Food Journal 8, 110113.
Yuyama, L.K.O., Duarte Favaro, R.M., Yuyama, K. and Vannucchi, H. (1991) Bioavail-
ability of vitamin A from peach palm (Bactris gasipaes H.B.K.) and from mango
(Mangifera indica L.) in rats. Nutrition Research 11, 11671175.
Zainal, Z., Tucker, G.A. and Lycett, G.W. (1999) Isolation and characterisation of a
cDNA encoding 1-aminocyclopropane-1-carboxylate oxidase from mango
(Mangifera indica L.). Asia-Pacic Journal of Molecular Biology and Biotechnology
7, 5359.
Zambrano, J. and Manzano, J. (1995) Efecto de la aplicacin de sales de calcio sobre la
maduracin de frutos de mango. Agronoma Tropical 45, 407413.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 529
15 Postharvest Technology and
Quarantine Treatments
G.I. Johnson
1
and P.J. Hofman
2
1
Horticulture 4 Development, Jamison, Australian Capital Territory, Australia
2
Department of Primary Industries and Fisheries, Nambour, Queensland, Australia
15.1 Introduction 530
15.2 Considerations Inuencing Postharvest Requirements 531
Market and consumer research 531
Quality assurance (QA) and Good Agricultural Practice (GAP) 531
15.3 Preharvest Management 535
Maturity 535
Adjusting maturation time 538
Skin colour and lenticel damage 538
Storage life and physiological disorders 540
Pests and diseases 542
Weather conditions 544
15.4 Flavour and Aroma 544
15.5 Harvesting and Transport to the Packhouse 544
Timing 544
Sapburn 545
Harvesting and desapping 546
Transport to the packhouse 548
15.6 Packhouse Measures 548
Delivery inspection and traceability 550
Desapping and washing 550
Disease control 550
Brushing 554
Grading and sizing 554
Grade standards 555
Disinfestation 557
15.7 Preparing Fruit for Market 566
Surface coatings 566
Packaging 567
Inspection 568
Palletizing 568
Precooling 569
G.I. Johnson and P.J. Hofman 530
Ethylene and ripening 569
15.8 Pre- and Post-shipping Storage 571
Cool storage 571
Controlled and modied atmosphere storage 573
15.9 Transport 575
15.10 Marketing 578
Networks and cooperatives 578
Promotion and consumer education 578
15.11 Conclusions 579
15.1 Introduction
Postharvest handling of mangoes is the last phase (from the tree to mouth) of
an agribusiness venture. To optimize productivity, protable uses for all
grades of fruit should be sought, and stable employment provided for key
skilled staff. Sustainable land and water management, and compliance with
health, safety and nancial obligations to employees are also necessary.
Increasingly, Good Agricultural Practice (GAP) protocols need to be observed
(GAP, 2003; FFTC-GAP, 2007).
This chapter reviews the technology and quarantine treatments that have
been developed for the postharvest handling of mangoes. Peacock (1986), Led-
ger (1986, 1991b) and Opara and Nguyen (1999) have previously reviewed
postharvest handling of mangoes, while several others have reviewed spe-
cic aspects of the topic (Johnson and Coates, 1993; Heather, 1994; Jacobi
et al., 2001b; Singh et al., 2004; Yahia, 2006). Less than 10% of total world
mango production is exported. Export markets for mangoes have expanded
because of social changes and rising demand, increased international air
cargo space for some sectors and promotion of export fruit production in
developing countries (Procter and Cropley, 1994; FAOSTAT, 2008). Expan-
sion of production to meet the supply requirements of export and distant
domestic markets has been possible because of successful integrated man-
agement strategies and disinfestation technologies to control diseases and
insects (Johnson and Coates, 1993; Johnson and Heather, 1995; Ploetz, 2007),
and increased land availability due to deforestation and diversication away
from rice. Market development has also been facilitated through harmoniza-
tion of the rules of trade between nations and regions at global and near
global levels (WTO, 2008), agreement on pest and disease risk management
under the International Plant Protection Convention (IPPC), various bilateral
and regional Free Trade Agreements (FTA, 2007), and the global expansion of
supermarkets. Simultaneously, knowledge of and concerns about exotic pest
risks to domestic fruit production, socio-political concerns about chemical
residues on food, environmental management and labour conditions, and
rising production and marketing costs, have impinged upon market access
and stimulated international dialogue and research initiatives which address
these concerns (Buchanan, 1994; Gullino and Kuijpers, 1994; Ploetz, 2003,
2007).
Postharvest Technology 531
15.2 Considerations Inuencing Postharvest Requirements
Market and consumer research
Strategies and procedures for horticultural market research have been out-
lined by Minnis (1993, 1994), Hall et al. (2001) and Kitinoja and Kader (2003).
Increasingly, a supply chain approach is being taken (Johnson and Hofman,
2004). Hofman and Ledger (2006) proposed that the supply chain approach
should be used to guide research and development and that there needs to be
a champion in the supply chain with signicant inuence and a desire for
improving chain status and performance. Key features are identication of
market demand, through-put, price and prot ows, seasonal uctuations in
availability and demands, supply competitors (and their commodity statis-
tics), importer-buyer requirements and relationships, options for value-adding,
and consumer expectations and sales promotions. Point-of-sale transaction
data from supermarkets and other sales outlets in target markets can be an
invaluable source of intelligence and can be purchased from marketing infor-
mation specialists. Detailed supply chain assessments can guide options for
innovation and improvement (Johnson and Hofman, 2004).
The volume, grade and quality of product available for export requires
analysis in relation to buyer specications, retail customer and provedore
preferences, technological and regulatory requirements for supplying the
market, and the production, packaging, cooling and transportation proto-
cols/options that are needed/available or specied under agreed codes of
practice (NRI, 2008a). Procter and Cropley (1994), Mahendra et al. (2002) and
the Natural Resources Institute (NRI) (2008b) provide some perspectives on
these issues.
Quality assurance (QA) and Good Agricultural Practice (GAP)
Postharvest handling assures timely delivery of a product that closely matches
buyer specications and complies with mandatory regulatory require-
ments. Satisfying customers underpins quality assurance (QA) and obser-
vance of GAP (Box 15.1), which aims to produce a product of the desired
standard, compliant with regional or internationally agreed standards in
production and handling, and encourage regular, larger and more frequent
purchases, and brand loyalty. As export markets become increasingly com-
petitive, responsive observance of QA and GAP certication can be vital for
maintaining and expanding market niche (Johnson and Coates, 1991; Askar
and Treptow, 1993; Ledger and Premier, 2006). Pieiro et al. (2004) provide
generic guidelines for the development of quality and safety guidelines for
fresh produce, and Ledger and Premier (2006) provide guidelines for a
Mango Quality Plan.
Components of a postharvest handling system may be developed in
commercial condence to enhance brand-name reputation and increase mar-
ket share. Increasingly they are also agreed under contractual agreements
G.I. Johnson and P.J. Hofman 532
with supermarkets or exporters. No regulations govern the pursuit of supe-
rior but legal agribusiness practices that may enhance grower and seller rep-
utations or provide additional advantages. Regulations govern chemical
residues, pests, diseases, product and packaging specications. The sections
that follow need to be considered in developing the postharvest components
of GAP protocols and dynamic QA systems.
Regulatory restrictions and quarantine treatments
Quarantine treatments disinfest produce of target pests. They are a critical
component of protocols designed to satisfy market-stipulated prohibitions
against pest entry to countries, or regions within countries. Protocols stipulate
procedures for monitoring, detecting, eliminating and handling pest-affected
Box 15.1. What is Good Agricultural Practice (GAP)?
GAP enhances trader-buyer relationships and the reputation of producers and
traders for the suitability of produce and environmental care. A key feature of
GAP is the role of independent certifying authorities (usually private sector) in
accreditation and compliance monitoring, to provide additional assurance to
buyers of certication creditability. GAP can be applied to any farming system
or scale. Key elements include: sustainable practices such as integrated pest
management (IPM), responsible fertilizer use and care for the environment. It
relies on four key principles (Wikipedia, 2007; NRI, 2008b):
Economic and efcient production of adequate supplies of safe and nutri-
tious foods or other agricultural commodities.
Compliance with sustainable practices which improve natural resource
availability.
Assuring the economic and environmental sustainability of the farming
enterprise.
Meeting social and cultural norms and expectations in production and
marketing.
GAP has gained global prominence because of the trend to making it a com-
pulsory standard for exporters to Europe. A private sector initiative, EurepGAP
sets voluntary standards for agricultural product certication anywhere. It involves
partnership between farmers and retailers wanting to establish and implement
procedures and standards for certifying application and compliance in GAP. It is
applied before the farm-gate (from planting to market dispatch), and is not nec-
essarily a standard that is displayed to customers. EurepGAP involves an array
of documentation including general regulations, control points, compliance cri-
teria and checklists (EurepGAP, 2007).
In 2007, EurepGAP transformed into GLOBALGAP (GLOBALGAP, 2007) that
could be tailored and adopted anywhere in the world. In parallel with the Eurep-
GAP process, the South-east Asian countries have been working towards
Association of South-east Asian Nations (ASEAN) GAP standards suited to
fruit marketing within ASEAN (ASEAN-GAP, 2007; FFTC-GAP, 2007; Johnson
et al., 2008).
Postharvest Technology 533
produce. Frequently, a detailed pest risk analysis (PRA) is required, protocol
efcacy against pests of quarantine concern must be demonstrated, appli-
cator integrity scrutinized and market-access requirements and rights ap-
proved by regulatory and agricultural authorities in the importing and
exporting country or region. Increasingly, systems approaches are being
applied. These minimize the need for reliance on quarantine treatments by
including PRA, production-based pest management systems, consideration
of establishing pest-free areas or identifying commodities which are non-
hosts of quarantine pests.
Restrictions or limitations on pesticide residues and other contaminants
in marketed produce are also considerations in the choice of quarantine treat-
ment and market access. Pesticide residue monitoring protocols are often
required (Johnson and Heather, 1995; Sharp and Heather, 2002; McMaugh,
2005). By restricting or preventing market access, quarantine and pesticide
residue restrictions can unintentionally operate as quasi-trade barriers, effec-
tively reducing competition with domestic production of the same or alter-
native products. However, this is decreasing because of better adherence to
science-based decision making as the foundation for quarantine restrictions
(WTO, 2008). Government authority web sites and export agencies can pro-
vide information on quarantine and tariff barriers, pesticide use restrictions,
inspection, packaging and labelling regulations (PPQ, 2007; IPPC, 2008) (see
Disinfestation section under 15.6 Packhouse Measures, this chapter). Phyto-
sanitary requirements for fresh mango exports to some markets are shown in
Table 15.1. Marketing through reputable exporters, known to the importing
country as suppliers of produce that comply with regulatory restrictions, can
encourage producer and importer condence.
Limitations of the product
Mangoes ripen rapidly and have low tolerance of temperatures <10C. Post-
harvest handling procedures for mangoes aim to optimize quality and mini-
mize premature ripening and fruit damage. Precise maintenance of fruit
quality and the storage environment demands inputs at every stage from
picking to the consumer (Ledger, 1991b; Milne, 1994; Ledger et al., 2002b).
Supply, logistics and transport
Market development cannot succeed without a reliable supply of suitable or
market-compliant produce. Production seasons are usually short (28
weeks), cultivar appearance and avour diverse, maturation time variable, and
orchards sometimes small and scattered. Practical solutions to these limita-
tions (i.e. sourcing from a range of ecoclimates with differing maturation
dates, or ower-induction technologies) are critical for industry develop-
ment. Competitive air-freight rates and rapid road transport, combined with
cool-chain handling and atmosphere management can make nearby export
markets almost as accessible as distant domestic markets. Special perishable
produce rates may be negotiated or subsidized by government, especially
during industry establishment. Sea freight is necessary for moving large
volumes when air freight is uneconomical. Out-turn problems often arise,
G.I. Johnson and P.J. Hofman 534
especially during market development (Snowdon, 1994). Insurance should
be arranged to cover losses. Specialist inputs may be required to identify
causes of the out-turn problems and in the apportionment of liability amongst
producer, shipper and marketer (Snowdon, 1990, 1994).
Personnel
QA systems and GAP protocols must encompass the human component of
an organization or business (Bunt and Piccone, 1994; Rolle, 2006; Sonneveld,
2006; FFTC-GAP, 2007). Market agents, exporters, farm/packhouse suppli-
ers, nance providers and transport personnel and companies need to be
selected and worked with from the quality perspective. Human resource
development, training and education have been of major signicance in the
success of many industries.
Table 15.1. Typical phytosanitary requirements for mangoes for some countries.
Country
Phytosanitary requirements
a
(usually for mangoes from individual
countries or regions on a case-by-case basis)
Australia Approved treatment for fruit y, area free of pulp weevil
(Sternochetus gravis (F.))
Canada No phytosanitary certicate required
China Phytosanitary certicate required. Field management measures for
specic pests of quarantine concern to China plus approved
disinfestation treatment for fruit ies
EU Phytosanitary certicate required
Indonesia Phytosanitary certicate required plus grown in area free of Queensland
and Mediterranean fruit y
Japan Phytosanitary certicate required plus disinfestation schedule approved
for nominated mango cultivars and fruit y species and inspection of
approved quantity of fruit (25%)
Korea Phytosanitary certicate required, combined with eld surveys
Malaysia Must be free of seed weevil on inspection
New Zealand Phytosanitary certicate required plus approved disinfestation schedule
for nominated fruit y species
Saudi Arabia Phytosanitary certicate required. Require destructive test of 2% of
consignment for seed weevil, or eld survey verication of block freedom
Singapore No restrictions
United Arab
emirates
Phytosanitary certicate required. Require destructive test of 2% of
consignment for seed weevil, or eld survey verication of block freedom
USA Phytosanitary certicate required plus disinfestation approved for
nominated mango cultivars and fruit y species
a
General requirements: Prior approval to import is required to access the market of many countries.
Packhouse and disinfestation treatment/facility inspections may be required by exporting and importing
regulatory authorities. Import permits may cover multiple importations but usually require renewal every
312 months. Phytosanitary certicates must be issued by a government authority. Fruit must be free
of soil and debris and packed in clean, new containers. Timber packaging and pallets will be subject to
additional requirements. Consignments found to contain quarantinable pests will be rejected, and either
re-exported or destroyed.
Postharvest Technology 535
Ledger and Bagshaw (1994) refer to three general styles of management:
(i) defect detection; (ii) defect prevention; and (iii) continuous improvement.
They consider that quality management of horticulture has traditionally been
the defect detection style, but it is now moving to the continuous improve-
ment style via the implementation of QA, total quality management and
supply-chain improvement systems. Key ingredients of successful imple-
mentation of quality systems and supply chains are: (i) unqualied commit-
ment by the owner and senior management of the human, material and other
resources needed to introduce and maintain the system; and (ii) employee
understanding and active participation in the process (Ledger and Bagshaw,
1994).
Protability and sustainability
Higher freight rates, tariffs and taxes, pesticide-use monitoring and quaran-
tine and security clearance times can affect sales negotiations and prot mar-
gins, as can socio-cultural differences among retail buyer, importer, exporter
and producer during marketing negotiations and exporter and buyer per-
ceptions and consumer expectations of quality. Intended markets, retailers
and regional trade fairs should be visited to make personal contacts and to
assess suitability of the market and retailing facilities, conditions and pros-
pects. Ongoing market monitoring is vital, with regular out-turn inspections
by trained personnel, and contingency arrangements for product regrading
at destination if required. Prompt, personal attention to client concerns or
product problems can be essential for continuing success (Johnson, 1997;
Vinning and Young, 2006).
15.3 Preharvest Management
The effects of production practices on fruit quality have been reviewed by
Arpaia (1994), Hofman and Smith (1994), Hofman (1998) and Hewett (2006).
The fruit characteristics inuenced by preharvest factors include internal and
external colour, shape, size, sweetness, vitality (the inherent capacity to
maintain quality after harvest) (Hofman et al., 1997b), cleanliness and residue
levels, and the occurrence of pest or disease infestations or biotic/abiotic
damage. The main production factors inuencing at-harvest quality apart
from genotype include chemical treatment regimes and orchard hygiene,
weather conditions before and at-harvest, irrigation, pruning systems and
tree nutrition.
Maturity
Peacock (1986) considered that fruit maturity referred to its stage of ontog-
eny, with fruit of different maturities being at different stages of ontogeny.
Fully mature mango fruit are strictly those which have produced a fully
developed seed and which have reached their full physiological potential in
G.I. Johnson and P.J. Hofman 536
relation to size increase and dry matter accumulation within the constraints
of the growth environment. When fruit size and dry matter concentration
reach a plateau, climacteric fruit such as mango can undergo ripening, where
colour, texture, avour and aroma may change (Watada et al., 1984). In these
fruit, a sharp rise, followed by a decline in respiration also accompanies the
transformation from not-ready-to eat (unripe), to edible (ripe), to senescent
(overripe). Ripening signals the completion of seed ontogeny, and encour-
ages dispersal of the seed by attracting vertebrate fructivores (Cipollini and
Stiles, 1992). If fruit are not harvested, maturation and ripening occur on the
tree. Ripe fruit fall to the ground, or are consumed by bats, primates, pha-
langers, birds or humans, either on the tree or after detachment.
Softening and sweetening of fruit esh and colour changes can occur at
any stage of ontogeny, even in pea-sized fruit (Oosthuyse, 1995). Fruit drop
at any stage of development is preceded by these events, and the likelihood
of their occurrence increases as fruit size and dry matter levels approach
their maxima (Singh et al., 2004). Although the changes constitute some of
the components of ripening, they can only be regarded as such if the fruit
have attained physiological maturity, i.e. the stage of development when a
plant or plant part will continue ontogeny even if detached (Watada et al.,
1984; Yashoda et al., 2006).
When fruit are removed from the tree several days before the onset of
ripening, they are initially hard and green. The fruit progressively soften,
change colour and develop aroma at a rate determined by cultivar, storage
environment and at-harvest maturity. Ideally, fruit are picked, treated, packed
and transported while hard-green, and arrive at retail markets at some pre-
determined stage of colour development (usually more yellow or red, than
green, and sprung, but still rm). The rate at which ripening will occur
under particular storage conditions depends upon the stage of ontogeny at
harvest. More mature fruit will ripen more rapidly than less mature fruit.
Accurately estimating when the fruit are ready for harvest is critical to
consistently meet customer expectations. This is called horticultural matu-
rity, and several criteria of horticultural maturity are possible. One is a legal
minimum or buyer-specied standard of maturity, which conrms that the
fruit would be acceptable for consumption or processing when ripe or ready
to eat for green-eating types. Maturity estimation often relies on visual or
calendar-based (days from owering) assessment or in some cases the appli-
cation of a simple test (e.g. dry matter or esh colour assessment). In more
exacting cases, more accurate estimates of horticultural maturity may be
required to assess product suitability for more stringent or narrower quality
specications, as may be required for contract sales or sea-export consign-
ments.
In both cases, easy to assess harvest indices relying on visual, chemo-
sensory or fruit-age attributes are needed, and they must correlate with the
commercially relevant fruit characteristics measured in prescribed tests. Pea-
cock (1986), Harvey (1987) and Askar and Treptow (1993) reviewed methods
for assessing fruit maturity. In mango, dry matter, esh colour, skin colour,
fruit shape, Brix, specic gravity or days from owering or heat accumulation
Postharvest Technology 537
units (e.g. degree-days) have been used (Baker, 1986; Hofman and Ledger,
2006). In South Africa esh colour is favoured, while in Australia, skin colour,
dry matter and accumulated heat units are considered as well (Fig. 15.1).
Using several maturity indicators will usually increase the accuracy of pre-
dicting the rst acceptable harvest date. Information would be cultivar-
specic. In some cultivars there may be few reliable visible indicators of
maturity that allow picking of the most mature fruit on the tree, particularly
when owering has occurred over many weeks. In these instances it is very
difcult for pickers to spot pick the more mature fruit in a cost-effective way.
Variation in maturity between fruit can also be inuenced by where fruit
develop on the tree. In the cooler subtropical areas of the southern hemi-
sphere, fruit on the northern or western side mature more quickly than fruit
on the southern side (Hofman et al., 1995; Oosthuyse, 1995; Hofman et al.,
1998). In these situations, it may be more feasible for growers to map the
maturity of their blocks or orchards to identify groups of trees that have more
mature fruit, and/or identify parts of the tree (e.g. the northern/western side
in the southern hemisphere) that generally hold more mature fruit. Pickers
can then be instructed to pick all the fruit on a specied canopy position and
from specied areas of the orchard in order to harvest more mature fruit.
This can be more cost effective than selectively picking from individual trees.
New technologies such as portable near infrared spectroscopy (NIRS) units
may assist in non-destructively mapping maturity proles across orchards
(Subedi et al., 2007).
The maturity of fruit at harvest is important for determining fruit quality
at out-turn in overseas markets (see 15.8 Pre- and Post-shipping Storage sec-
tion, this chapter). If fruit in a carton or pallet are of uneven maturity, it may
be impossible to nd an effective storage regime(s) which will ensure good
quality of all fruit on arrival. One fruit of more advanced maturity in a box or
pallet can accelerate the ripening of other fruit, which can then arrive with
Flesh colour at harvest (chart)
2 4 6 8 10 12
F
l
a
v
o
u
r

(
1

9
)
3
4
5
6
7
r = 0.86**
Heat units
1200 1400 1600 1800 2000
F
l
a
v
o
u
r

(
1

9
)
3
4
5
6
7
Early harvest
Mid-harvest
Late harvest
r = 0.83
Fig. 15.1. Relation between esh colour (1 = white; 15 = very yellow) and accumulated
heat units (>10C) with avour of ripe B74 mango grown under Australian conditions.
**= Signicant to P = 0.01. (Source: Hofman and Marques, unpublished data).
G.I. Johnson and P.J. Hofman 538
disease symptoms and with little or no shelf life. Variable maturity within
treatment lots can also adversely affect product quality after heat disinfesta-
tion (Jacobi et al., 1995).
On-farm record keeping and analysis of date of owering, seasonal prod-
uct maturity, orchard management schedules, environmental data, transport
regimes, market destinations and out-turn problems may enable some pre-
diction of at-market quality and better selection of the appropriate destina-
tion market. Recent research is focusing on non-destructive methods that
could be used for checking fruit maturity in automated grading systems
(Joyce et al., 1993; Subedi et al., 2007). Improvements in product quality and
performance resulting from the effective use of such systems over several
seasons, can provide considerable competitive advantages when developing
buyer relationships or consumer brand/country-of-origin loyalty.
Adjusting maturation time
Flowering may be hastened or delayed by pruning, owering inducers or
growth regulators (Davenport and Nez-Elisea, 1997), to move the fruit-set
period into a different time period, and bring forward or delay the harvest
date to coincide with higher demand and market prices. Disadvantages
could include higher at-harvest temperatures or greater risk of rainfall prior
to harvest adversely affecting fruit quality.
Some cultivars and growing conditions are more favourable for manipu-
lating owering date than others. In the Philippines, manipulation of Cara-
bao owering with potassium nitrate (KNO
3
) helps spread production and
market supplies year-round (Bondad and Linsagan, 1979). The treatment
also increases Kent owering but does not alter timing (Goguey, 1993). By
contrast, chemical manipulation of owering has not been effective on Kens-
ington Pride (Barba, 1974).
Evenness in owering within and between tree/block/farm lots can
contribute to product uniformity and increased customer condence. Treat-
ments that increase evenness of owering can have commercial benet in
this regard also.
Skin colour and lenticel damage
Skin colour can affect sales, with markets preferring colour (green, yellow,
orange, red blush) familiar to past purchase experience, known use and cul-
tivar knowledge, or ethnic-group preferences. Fruit position on the tree af-
fects red colour development, since sun exposure is important for anthocyanin
development. Likewise, bagging of fruit can decrease red and green skin
colour on ripe fruit (Hofman et al., 1997a). Nitrogen (N) can increase the pro-
portion of the ripe fruit with green skin (Fig. 15.2) by retaining skin chloro-
phyll (McKenzie, 1994; Nguyen, 2003; Nguyen et al., 2004; Bally, 2007). In
cultivars susceptible to green skin at ripeness, N fertilization rates should be
Postharvest Technology 539
balanced between improving yield and reducing quality. Applying N to trees
soon after harvest can minimize these negative effects in the subsequent
crop. Additional N can be applied just before owering as long as leaf N
concentrations are below a certain level (Whiley and Hofman, 2007). This
level may vary with cultivar. Excessive fruit calcium (Ca) concentrations in
mango will also retard green colour loss during ripening (Wills et al., 1988).
Some cultivars are marketed green (e.g. Keow Savoey), which are consumed
mature-green before softening and colour development occur.
Enhanced prominence or damage to lenticels on the skin can affect visual
appearance (Plate 81). Various terms have been used, including lenticel dam-
age, discoloration and spotting. Symptoms can be caused by darkening of
the cells immediately around the lenticel producing a brown or black spot, or
by a red or green halo around the lenticel with or without the black or brown
spot in the centre (Bezuidenhout et al., 2005; Self et al., 2006). Recent studies
suggest that the discoloration is not primarily caused by loss of cellular func-
tion, but rather by the deposition of phenolic pigments in the cell wall (du
Plooy et al., 2006). It is possible that leakage of precursors (elicitors) from
adjacent resin canals into the cell wall next to the lenticels contributes to pig-
ment formation. Lenticel discoloration may be a stress-related self-defence
Total N application (g/tree)
0 75 150 300 Fol. 0 150 300 450 Fol. 0 150 300 450 Fol.
HG orchard LG1 orchard LG2 orchard
G
r
e
e
n

c
o
l
o
u
r

(
%
)
0
1
2
5
10
20
30
40
Fig. 15.2. Percentage of green skin colour on ripe Kensington Pride mango fruit
from trees in three different orchards (coded HG, LG1 and LG2) following application
of N (0450 g/tree). (Source: H. Nguyen et al., 2004). Fol. = foliar N sprays to a total of
50 g/tree. Bars represent least signicant difference (LSD) at 5%.
G.I. Johnson and P.J. Hofman 540
mechanism against foreign particles and infection entering through the len-
ticels (Bezuidenhout et al., 2005; du Plooy et al., 2006).
The severity of lenticel damage is often difcult to control, but strategies
for minimizing damage exist. There are cultivar differences in susceptibility
(Oosthuyse, 1999), possibly related to differences in lenticel, wax and/or
cuticle structure or composition (du Plooy et al., 2004). Strong negative cor-
relations have been found with maximum and minimum temperature and
Class A pan evaporation, and strong positive correlations with maximum
relative humidity (RH) and rain at harvest (Oosthuyse, 1998). These results
suggest that cool, humid and wet conditions around harvest increase the risk
of lenticel damage. Increased damage following excess irrigation during the
latter stages of fruit growth (Simmons, 1998) support the above conclusions.
Lenticel damage can also be more severe in larger fruit obtained from
branches with higher leaf:fruit ratios (Table 15.2), possibly because of greater
damage to the lenticels during fruit growth (Simmons et al., 1998).
Storage life and physiological disorders
Physiological disorders include a range of symptoms that affect shelf life and
marketability (Johnson et al., 1996). These generally result in either prema-
ture ripening of parts of the fruit (e.g. soft nose and jelly seed) or tissue break-
down (i.e. stem-end cavity) (Winston, 1986; Mead and Winston, 1991; Whiley,
1999) and tissue breakdown in Keitt (Bally, 2007). These disorders are more
severe in more mature fruit (Young, 1957; Katrodia, 1989; Mead and Winston,
1991) and are often evident on the tree or after ripening without storage.
Mango disorders are affected by growing conditions (Young, 1957; Young
and Miner, 1961). Production away from the coast, and higher altitude and/
or lower temperature are associated with lower incidence of spongy tissue
(Subramanayam et al., 1980; Katrodia, 1989), and susceptibility to the disor-
der is also affected by rootstock (Joshi and Roy, 1985). Stem-end cavity ap-
pears to be more severe in wet conditions near harvest (Wainwright and
Table 15.2. Effects of leaf:fruit ratios on quality of Kensington Pride mango fruit after ripen-
ing at 22C (Source: Simmons et al., 1998).
a
Leaf:fruit
ratio
Fruit
mass (g)
Dry
matter (%)
Lenticel spotting
(15)
Disease
Severity (15) Incidence (%)
Control 441.4
c
13.0
b
3.5
c
1.2
c
13.3
b
30 363.2
d
12.0
c
3.5
c
1.8
b
40.0
ab
60 532.5
b
13.7
b
3.9
b
2.0
b
43.3
ab
120 696.6
a
15.1
a
4.2
a
2.7
a
63.3
a
a
Treatments were applied by girdling individual branches. Control fruit were from non-girdled branches.
Values are means of 30 fruit per treatment. Values with different letters within columns are signicantly
different at P < 0.05.
Postharvest Technology 541
Burbage, 1989; Mead and Winston, 1991). Incidence of spongy tissue has been
reduced by mulches that decrease radiated and reected eld heat (Katrodia
and Sheth, 1989). Severity of watery pulp breakdown in Keitt is lower with
higher crop loads from similar size trees (Bally, 2007), possibly because of
smaller fruit size at high crop loads.
Several reports suggest links between low fruit Ca and mango disorders.
High leaf Ca has been related to reduced soft nose (Young and Miner, 1961)
and reduced stem-end cavity (Mead and Winston, 1991). Soil Ca applications
can reduce stem-end cavity incidence (Whiley, 1999), but these responses are
not always consistent. Applications of Ca to Keitt from just before owering
onwards did not increase leaf or fruit Ca during fruit growth or at commer-
cial harvest (Bally, 2007). However, soil characteristics may also affect responses
to soil Ca applications. In the sandy soils typical of Australian orchards,
and even in the heavier clay subtropical soils with low cation exchange
capacity, Ca can be rapidly removed from the top soil prole, resulting in
little long-term increase in soil solution Ca (Hofman and Mullen, 2005;
Bally, 2007). Regular (two/week), small applications are required to consis-
tently increase solution Ca (Hofman and Mullen, 2005). Other factors (e.g.
vegetative vigour and water status) can inuence fruit Ca uptake (Hofman
and Smith, 1994).
Foliar Ca treatments have produced inconsistent effects, in some cases
having little or no effect on fruit Ca concentrations or quality (Singh et al.,
1987; Simmons et al., 1995), and in other instances reducing internal disor-
ders (Chitarra et al., 2001). Postharvest dips have also had mixed results, with
both positive (Wills et al., 1988; Singh et al., 2000) and nil or very small effects
(Joyce et al., 2001) reported. As a result there is little commercial use of foliar
or postharvest Ca treatments in mango. Future development of more labile
Ca formulations may provide more consistent results.
Other nutrients have also been associated with fruit quality (Hofman
and Smith, 1994). There are relatively few reports of potassium (K) and mag-
nesium (Mg) effects on mango fruit quality. Higher Ca and Mg, and a ten-
dency towards lower K in mango fruit, have been noted in fruit from orchards
with no incidence of soft nose, compared with fruit from orchards with high
incidence of the disorder (Burdon and Moore, 1991). Conversely, K levels are
positively correlated with disease resistance; Karunanayake (2007) reported
reductions in disease on mango fruit from trees receiving additional K.
Application of triple the recommended level of K signicantly reduced stem
end rot caused by Lasiodiplodia theobromae, while the severity of anthracnose
was most reduced by application of the recommended level of K compared
to nil and triple rate treatments.
Excess N can reduce storage life and quality, and excessive levels cause
deterioration of avocado fruit quality (Wolstenholme, 2004) where its effect
may be mediated through vegetative:reproductive balance and crop load
(Hofman and Mullen, 2005). High N has been associated with increased dis-
orders in mango (Young and Miner, 1961; Mead and Winston, 1991), possibly
through the dilution effect of increased fruit size on Ca concentrations; how-
ever, soil N applications later during fruit growth do not affect watery pulp
G.I. Johnson and P.J. Hofman 542
breakdown in fruit Keitt fruit (Bally, 2007). Heavy rain late during fruit
development can release soil N previously unavailable to trees due to low
soil moisture, potentially resulting in high fruit levels. Boron (B) deciency
has been related to abnormal fruit development (Lahav and Whiley, 2002)
and increased fruit storage disorders (Yogaratnam and Johnson, 1982; Smith
et al., 1997). It may also be important in mango (Coetzer et al., 1991). The effect of
larger fruit size and maturity on shelf and storage life (Seymour et al., 1990) can
also be mediated through production factors inuencing fruit set and leaf:fruit
ratio. Fruit position in the canopy may also play a role here (see above).
Pests and diseases
Postharvest diseases and pests are reduced by various preharvest control
measures including orchard hygiene, manipulation of owering, integrated
management and the use of chemical and biological controls (Johnson et al.,
1989a; Johnson, 1997; Fonseca et al., 2004b; Ploetz, 2004; Akem, 2006; Astridge
and Baron, 2007a, b, c; Chin et al., 2007; Diedhiou et al., 2007). Prusky et al.
(Chapter 7, this volume) and Ploetz and Freeman (see Chapter 8, this vol-
ume) reviewed preharvest management of several postharvest diseases,
while Dann et al. (2005, 2007) reviewed novel treatment options. Under the
range of subtropical to tropical and dry to wet ecoclimates, combinations of
treatments have been recommended to protect vegetative growth ushes,
ower panicles and developing fruit from infections that lead to anthracnose,
bacterial spot, powdery mildew, scab and stem-end-rot symptoms on fruit
during development or after harvest (Pofey et al., 1999; Ledger, 2004; Sto-
volt and Dirou, 2004; Akem, 2006).
Lonsdale (1993) found that monthly applications of copper oxychloride
(CuCl
2
3Cu(OH)
2
) in combination with mancozeb controlled most mango
postharvest diseases. Copper (Cu) alone was less effective in controlling
anthracnose. Copper sprays also provide protection against bacterial spot,
while mancozeb can provide protection against scab (Pofey et al., 1999; Led-
ger, 2004). Timmer and Zitko (1996) and Hardy et al. (2004) discussed the
application of copper treatments to citrus for disease control. Formulations
differ in their weather hardiness and indicated that retention of copper oxide
(CuO) is superior to retention of copper chloride (CuCl
2
) or oxychloride, and
application of Cu with the pH <76 can damage fruit and leaves. Lonsdale
(1993) considered there was no disease control benet on mango by alternat-
ing Cu with prochloraz sprays, but Ledger (2004) recommended their strate-
gic application every 34 weeks in rotation with mancozeb and Cu when
rainy conditions favoured anthracnose on developing fruit.
Azoxystrobin (Amistar

) and other strobilurin fungicides effectively


control anthracnose, alternaria and powdery mildew on mango (Reuveni
et al., 1998; Willingham et al., 1999; Reuveni, 2000; Sundravadana et al., 2006,
2007), Botryosphaera parva (Syn., Dothiorella dominicana) and Phomopsis sp.
causing stem end rot (Everett et al., 2005), and Cercospora leaf spot (Ane-
siadis et al., 2003) on other hosts. In Australia, no more than three strategic
Postharvest Technology 543
applications of azoxystrobin are recommended for eld control of anthra-
cnose on mango in alternation with Cu, prochloraz and mancozeb schedules.
Azoxystrobin should be appled as one or two applications at owering and/
or early fruit set at no less than 14-day intervals, and again at 21 and 7 days
before harvest (Stovolt and Dirou, 2004). Application of azoxystrobin to control
anthracnose on mango leaves signicantly reduces subsequent fruit disease
and boosts yield (Sunarharum, 2007).
Recent research has also focused on the potential of defence-boosting
treatments, applied before or after harvest to reduce disease impacts on yield
and shelf life (Terry and Joyce, 2004; Dann et al., 2007; Karunanayake, 2007).
Anthracnose on mango fruit can be reduced by salicylic acid, its functional
analogue benzothiadiazole (BTH) (= acibenzolar-S-methyl = Bion

), and
ultraviolet (UV-C) irradiation with variable results, including phytotoxic
effects (Zainuri et al., 2001; Zeng and Waibo, 2005; Zainuri, 2006; Zeng et al.,
2006; Karunanayake, 2007).
Orchard hygiene, including reduction of inoculum by removal of old
fruit and branches, and removing prunings from the orchard, can reduce
anthracnose and stem end rots on fruit after harvest (Johnson, 1994; Ledger,
2004; Ploetz, 2004). Some eld diseases can disgure fruit. Following heavy
rain close to harvest, bacterial spot can appear as discrete raised lesions, fol-
lowed by fruit cracking or rupture and fruit drop. Black spot orchard man-
agement practices (i.e. windbreaks, Cu sprays and use of resistant cultivars)
can reduce the risk of damage on fruit (Johnson et al., 1996; Dodd et al., 1997;
Ploetz and Prakash, 1997). In cool conditions, powdery mildew infection on
young fruit can cause a ghosting symptom on mature fruit similar to that
caused by powdery mildew on apples. Scab and sooty moulds can disgure
fruit. Generally, spray programmes for anthracnose will control scab (Cond
and Pitkethly, 2007), while sooty moulds are managed through integrated
management of scale insects and by postharvest brushing (Johnson and
Coates, 1993).
Tree nutrition can affect fruit disease incidence and severity. High Ca can
reduce fruit diseases in many fruit crops (Hofman and Smith, 1994). Nitro-
gen application before owering can increase mango fruit disease (H. Nguyen
et al., 2004); applications up to 6 weeks after owering can increase anthra-
cnose severity (Bally, 2007). Negative correlations occur between exocarp N
percentage and antifungal resorcinol concentrations.
Pofey et al. (1999), Pea (2004) and Pea et al. (see Chapter 10, this vol-
ume) discussed integrated pest management (IPM) for reducing pest dam-
age and quarantine hazards associated with fruit. Preharvest control measures
for fruit ies, seed weevils, scales and other skin defect-causing pests con-
tribute signicantly to product quality improvement. IPM strategies can
adequately control orchard pests while reducing reliance on pesticides (Cun-
ningham, 1986, 1991a, b, c; Vijaysegaran, 1994; Pea, 2004; Pea et al., Chap-
ter 10, this volume). Bagging of developing fruit can reduce or eliminate
disease infection (Hofman et al., 1997a) and fruit y infestation (Kitagawa
et al., 1992). However, for export market access, and regardless of effective
eld control, postharvest disinfestation measures are often mandatory.
G.I. Johnson and P.J. Hofman 544
Weather conditions
Rain before harvest, and high RH and temperatures can increase disease lev-
els, fruit susceptibility to heat and brush damage, lenticel damage and reduce
storage life (Dodd et al., 1991a; Estrada et al., 1993; Prusky et al., 1993a, b; Cooke
and Johnson, 1994; Oosthuyse, 1998; Jacobi et al., 2001b). Disease risk predic-
tion based on the monitoring of environmental variables to determine fungi-
cide application frequency, can reduce pesticide residues (Fitzell and Peak,
1984; Fitzell et al., 1984; Peak et al., 1986; Dodd et al., 1991a, b, 1992; Prusky et al.,
1993b). Irrigation frequency and water availability for tree growth can signi-
cantly impact postharvest diseases and disorders (Simmons, 1998).
15.4 Flavour and Aroma
Flavour is largely determined by sugars and volatiles in the ripe fruit, both of
which increase in more mature fruit. The aroma produced by ripening and
ripe mango can help attract customers, and provide some indication of a-
vour development. In mango fruits, more than 280 different aroma volatile
compounds have been reported (Singh et al., 2004). Variation in the constitu-
ent aromatic compounds in mango cultivars results in aroma and avour
diversity (MacLeod and Snyder, 1985; MacLeod et al., 1988; Torres et al., 2007).
The high fruit levels of -terpinolene contribute to the characteristic avour
of stronger-avoured cultivars such as Kensington Pride (Bartley and
Schwede, 1987; MacLeod et al., 1988). Kensington Pride harvested at the
green-sprung stage have higher concentrations of total aroma volatiles com-
pared with fruit harvested at the hard-green or coloured stages (Lalel et al.,
2003b). Most of the glycosidally bound aroma compounds increase in the
pulp as the fruit matures, which contribute to improved avour. During the
rst 7 days of ripening, -turpinolene is the major volatile compound, but in
the later stages of ripening ethyl octanoate dominates (Lalel et al., 2003a).
15.5 Harvesting and Transport to the Packhouse
Harvesting of mango is determined according to attainment of acceptable/
required maturity: (i) for arrival at market during the time of peak demand/
highest price to maximize the chance of early sale; or (ii) to minimize loading
wait at the shipping port. Fruit is generally picked into eld crates or bins,
with or without the use of mechanical picking platforms.
Timing
Maturity is determined by assessing variables such as days from owering,
accumulated heat units, esh dry matter percentage, esh colour or fruit
shape/skin colour (see Maturity section under 15.3 Preharvest Management,
Postharvest Technology 545
this chapter). Fruit water potential uctuates diurnally, and can affect fruit
quality. The water potential of fruit at harvest can affect susceptibility to han-
dling, heat damage and product storage potential (Joyce and Patterson, 1994).
In hot weather, fruit should be harvested in the coolest part of the day to
reduce fruit overheating and energy requirements for postharvest cooling,
and to minimize worker discomfort. Harvest during rain can reduce fruit
quality (see Weather conditions section under 15.3 Preharvest Management,
this chapter).
Sapburn
Severing the stem from the fruit causes relatively large volumes of latex to
spurt or ooze from the cut stem. The sap is of low pH and high oil content
and can burn the surface of the fruit (Bagshaw and Brown, 1989). The oil frac-
tion contains terpinolene and resorcinols and is the fraction of the latex that
causes the damage. Skin damage is particularly severe with Kensington
Pride (OHare, 1994), and less serious in Florida cultivars. In Pakistan,
Chausa is more susceptible than Sindhri and Dashehari (Maqbool et al.,
2007). OHare (1994) observed that latex levels are lower and less phytotoxic
in Nam Doc Mai, Nang Klang Wun, Tong Dum and Keow Savoey (0.16
0.48 ml/fruit), than in Kensington Pride (1.67 ml/fruit). The oil compo-
nent of the latex of Thai cultivars is much lower than that of Kensington
(OHare, 1994; Hassan, 2007). The concentrations and ratio of the two main
resorcinols, 5-n-pentadecylresorcinol and 5-n-heptadecenylresorcinol, differ
among cultivars (Hassan, 2007).
Factors affecting the potential of latex to cause sapburn are not well
understood. It appears that both the terpene in the oil fraction of the sap, and
adequate polyphenol oxidase (PPO)/peroxidise concentrations in the skin
are required to develop sapburn; PPO and resorcinols in sap are less signi-
cant (Loveys et al., 1992; Robinson et al., 1993; John et al., 2002). Rain near
harvest and high N in fruit result in more severe sapburn in Kensington
Pride. However, negative relationships have been observed between exo-
carp N concentration and alkyl resorcinols (Hassan, 2007). Sap from fruit
harvested early in the day causes less sapburn than sap from fruit harvested
later in the day, although early-harvested fruit exude more sap than late-
harvested fruit (Maqbool et al., 2007).
Latex may provide protection against infestation by fruit y larvae (Joel,
1978, 1980) and may also contribute to disease tolerance. The 5-substituted
resorcinols have antifungal properties (Cojocaru et al., 1985; Droby et al., 1986,
1987; Prusky and Plumbley, 1992). Karunanayake (2007) extracted a resorci-
nol and a resorcinol derivative from the dichloromethane phase of Sri Lankan
mango peel extracts that had antifungal properties. Differing relationships
occur between resorcinol levels and relative susceptibilities of different
mango cultivars to anthracnose (Karunanayake, 2007; Hassan et al., 2007).
Strong positive relationships occur between resorcinol concentration in the
peel and latex, and fruit resistance to articially inoculated anthracnose.
G.I. Johnson and P.J. Hofman 546
Kensington Pride has more non-aqueous latex, higher concentrations of resor-
cinols and greater tolerance of anthracnose than Nam Doc Mai (Hassan, 2007).
Less anthracnose occurs on fruit ripened with an intact stem, compared with
de-sapped fruit. Hegnauer (1994) reviewed the phytochemistry of Mangifera.
Cultivars that are prone to sapburn can be harvested with 1020 mm
stems attached, and re-trimmed at the packhouse. Latex does not usually
exude from longer stems because there is no continuity between the fruit and
stem resin ducts (Joel, 1980), and the fruit lactifers are not severed; however,
stems can break in transit to the packhouse, resulting in latex leakage and
sapburn. Long stems left on the fruit to reduce sapburn has variable effects
on disease depending on disease type and storage time. With shorter storage
periods, anthracnose and stem-end-rot incidence and severity can be lower
in fruit with stems attached due to higher levels of resorcinols (Hassan, 2007),
but during longer storage stem-end-rot levels can increase due to the higher
levels of inoculum associated with the retained stalk (Johnson et al., 1993).
Mango latex can cause skin disorders in humans (Keil et al., 1946; Oka
et al., 2004). Bandyopadhyay et al. (1985) noted that resorcinol derivatives are
allergens in the Anacardiaceae, and suggested that the 5-substituted resorci-
nol in mango latex causes dermatitis. Both heptadec(adi)enyl resorcinols and
pentadecylresorcinol can elicite an allergic reaction in sensitive patients (Oka
et al., 2004). Harvesting and packhouse personnel must avoid contact with
the latex of high-risk cultivars.
Harvesting and desapping
Rough handling at harvest can cause skin damage and internal fracturing or
bruising. Using hooked sticks or shaking the tree to detach fruit causes skin
damage and esh fracturing (Ledger, 1991a; Abu-Goukh and Mohamed,
2004) and sapburn. Mechanical damage during harvest also causes soft,
darkened areas and bruises on fruit following hot water treatment. Mangoes
should be handled as if they were eggs. Long-handled secateurs cut and grip
the stem, allowing the fruit to be carefully lowered to the picking bin. Contact
with soil and soilborne pathogens should be avoided (Johnson et al., 1993).
In cultivars where sapburn is a problem, latex should be drained from
the fruit (desapping or bleeding) to minimize the incidence and severity of
sapburn. Several systems have been assessed for reducing damage (Brown
et al., 1986; Ledger, 1991b; Holmes et al., 1993; Lim and Kuppelweiser, 1993;
OHare and Prasad, 1993; OHare, 1994; Shorter and Joyce, 1994). In Austra-
lia, the main commercial practices (Plate 82) are:
Desapping in the eld with harvest aids using detergent. The basic de-
sign characteristics include detergent spraying onto a tarpaulin, a trough
with the same detergent and a nal spray before fruit are placed in a eld
bin. The fruit are either hooked from the tree in the direction of the harvest
aid and onto the catching surface or the fruit are snapped directly off the
tree and placed onto the tarpaulin. The fruit roll from the tarpaulin into
Postharvest Technology 547
the trough containing detergent and then into 300400 kg eld bins.
Alkaline detergents that deactivate damaging sap components are most
effective; high concentrations of surfactant in the detergent are not re-
quired. The crucial factors are that fruit should be exposed to detergent
for at least 90 s, the detergent is either not recycled, or replaced before
sap accumulation in the detergent causes other damage such as skin
browning (Bally et al., 1997; OHare et al., 1999).
Another design includes a motorized hydraulic ladder (cherry picker)
with the fruit desapped for 12 s before placing in a basket containing a
spray of alkaline detergent. This system is particularly effective for tall
trees, but care must be taken that fruit are covered by the detergent for at
least 90 s.
Picking fruit with long stems into small 18 kg crates and desapping in
the shed. The fruit are dipped into detergent before desapping and plac-
ing on a long conveyor system that holds the fruit inverted and provides
detergent/water sprays for a few minutes. The fruit are inverted for c.20
min before drying and packing.
In these systems, the detergent is not strongly alkaline, but the surfactant
should be of sufcient concentration to provide a protective coating around
the fruit before desapping. Stem breakage must be minimized in the crates,
as this can cause sapburn and quality loss (Holmes et al., 1993; Holmes and
Bally, 1994). Latex must not spray or drip onto fruit being desapped. Workers
who are sensitive to mango sap should wear hand protectants, aprons and
footwear to minimize skin contact. The detergent must reduce sapburn and
skin browning without causing other damage (i.e. lenticel spotting) (Fig.
15.3). Desapping in the eld by inverting fruit directly onto racks without
detergents has been used for sensitive cultivars and when particular growing
conditions have increased fruit susceptibility to lenticel spotting or other
damage from detergents; however, labour costs are becoming prohibitive,
requiring compromises between cost and quality. Desapping by inverting
and placing on the ground signicantly increases the incidence and earlier
appearance of stem end rot caused by soilborne L. theobromae, and is not
recommended (Johnson et al., 1993).
Harvest aids have reduced in-shed desapping. Holmes et al. (1993) found
916% of fruit in eld crates were affected by sapburn when harvest aids
were not used. Harvest aids provide the greatest reduction in total sapburn
(from 69% to 1518%). While harvest aids can signicantly reduce sapburn,
inappropriate use can increase some forms of skin browning (Bally et al.,
1997). Underhill and Dahler (1995) described four types of skin browning
which produce symptoms distinct from the sapburn caused by the oil phase
of latex. Several forms of skin browning involve tissue reactions with sap/
detergent mixtures. Symptoms vary if latex enters fruit through micro-cracks
in the cuticle or the lenticels (OHare et al., 1999). Holmes (2003) developed
guidelines for the use of harvest aids.
Cost savings associated with the use of harvest aids can be lost if fruit-
to-fruit or fruit-to-ground impact is not minimized during harvest. Rough
G.I. Johnson and P.J. Hofman 548
harvesting can increase the incidence of bruising and internal fracturing, and
lower wholesale returns. Thorough training of picking crews and supervi-
sion of their performance is required to maintain good practice.
Transport to the packhouse
Harvested fruit should be transported to the packhouse as soon as possible,
with no prolonged exposure to the sun. Rough handling and transport must
be minimized. Roads/tracks from orchard to packhouse should be smooth,
with transport vehicle tyres correctly inated, and special suspensions to
reduce vibration and damage.
15.6 Packhouse Measures
Harvested fruit are transported to a central packhouse which provides shel-
ter from rain and sun, and facilities for cleaning, treating, packing, cooling
and storing fruit until consignment to market (Schoorl and Holt, 1982, 1985).
Mechanized packhouse systems can offer labour savings and increased re-
turns (Murray and George, 1994). When manual handling was reduced from
ve to two steps, fruit appearance improved, disease losses were lowered,
sizing accuracy improved, packing rate increased and space, labour and
supervisory requirements were reduced (Murray and George, 1994).
A typical packhouse sequence is shown in Fig. 15.4. In packhouses that
include a disinfestation facility, the sequence must be modied to allow for
Detergent
C
o
n
t
r
o
l
L
O
C
M
a
n
g
o

C
l
e
a
r
M
a
n
g
o

G
l
o
M
a
n
g
o

M
a
g
i
c
M
a
n
g
o

P
l
u
s
M
a
n
g
o

W
a
s
h
S
u
p
e
r
c
o
n
c
e
n
t
r
a
t
e
L
e
n
t
i
c
e
l

s
p
o
t
t
i
n
g

(
7
d
a
y
s

a
f
t
e
r

h
a
r
v
e
s
t
)
1
2
3
Block 1
Block 2
Block 3
Fig. 15.3. Effect of detergents on lenticel spotting on skin of B74 mango (0 = no
lenticel spotting; 4 = severe spotting). Fruit were obtained from three different blocks
in two orchards, dipped in detergent for 2 min, then held at 20C for 7 days (Source:
Hofman and Ledger, 2006). The bar represents the least signicant difference (LSD)
at 5%.
Postharvest Technology 549
1. Deliver 18. Air or sea freight
2. Weigh 17. Cool chain transport
3. Cold wash and brush 16. Palletizing
4. Hot water/fungicide bath 15. Outer boxing
5. Fungicide application 14. Cool/ethylene gas
6. Dry 13. Pack
7. Wax and brush (optional) 12. Disinfestation
8. Dry 11. Crates
9. Grade 10. Size
Processing
grade
Second
grade
First grade
(domestic/export)
Extra class
(export)
Processing Local marketing
Fig. 15.4. Packhouse and marketing activities for mango. Waxes are not applied in some
countries because of abnormal ripening and off-avour development. When disinfestation is
not required, steps 11, 12 and 15 would be omitted. When fruit are heat disinfested, they may
be packed into an inner box prior to cooling. For some markets accessible by air, the fruit may
be treated with ethylene and stored until near ripe. The boxed fruit may then be packed into
an outer carton prior to palletizing. Pre-ripening allows fruit to be rechecked prior to despatch,
with fruit of unsatisfactory appearance (e.g. skin damage) redirected to domestic market or
processing.
G.I. Johnson and P.J. Hofman 550
sizing before disinfestation, with tray packing after treatment. Packhouse
design and installation consultants can provide substantial savings by elimi-
nating bottlenecks and minimizing product damage points.
Delivery inspection and traceability
Harvested fruit in eld crates should be treated, packed and cooled as soon
as possible. Quality and contaminant management systems may require that
a record system tracks the block or trees from which each bin of fruit is har-
vested. This enables records to be kept of tree or block yield, quality perfor-
mance and defect levels as well as labour performance rates and pesticide
residues. Individual fruit in a tray can be traced back to the tree or block from
which it was harvested. Block/tree-to-tray traceability systems and pack-out
records allow problems (i.e. excessive or unapproved pesticide detections) to
be traced to the site of the problem and relevant action taken. They can also
be used to motivate producers or picking teams to deliver high quality
produce.
At the packhouse, samples of fruit should be evaluated immediately for
maturity, blemishes and disease and pest incidence and recorded in an appro-
priately designed computerized system. Preharvest orchard inspections can
reveal the defects that can be anticipated in the packhouse. Some degree of
in-eld sorting can occur at the point of harvest, and soft or damaged fruit
collected separately and discarded.
Desapping and washing
Unloading should avoid dropping, damaging and wounding of fruit. Fruit
are normally unloaded from eld bins into bin dumps if desapping is unnec-
essary or removed manually from the crates for desapping (see Harvesting
and desapping section under 15.5 Harvesting and Transport to Packhouse,
this chapter). Detergents and sanitizers are sometimes added to washing
water. Their use requires careful consideration. Some may cause fruit dam-
age, or promote early fruit disease expression (Korsten et al., 1993). Chlorine
is added and carefully regulated to wash and/or rinse water in some pack-
houses, but this is not essential. Quaternary ammonium disinfectants should
not be added to wash water as their direct application to foodstuffs is gener-
ally not permitted.
Disease control
Hot water and fungicide application
Hot water dips, or sprays over brushes, with or without fungicide, and fun-
gicide sprays or dips, can eradicate quiescent fungal infections that have
been established on and beneath the cuticle and within the pedicel prior to
Postharvest Technology 551
harvest (Johnson et al., 1989a, b, 1991, 1992; Kernot et al., 1999; Pofey et al.,
1999; Plan et al., 2002). Suslow (2000) provides generic recommendations for
the postharvest handling of produce. Postharvest disease treatment efcacy
varies with infection level, cultivar, ripening status and storage regime. Hot
water treatment also cleans fruit, but can contribute to increased skin dam-
age (Cooke and Johnson, 1994). Anthracnose caused by Colletotrichum gloeo-
sporioides Penz. and Colletotrichum acutatum Simmonds, is controlled more
readily than stem end rots (or soft brown rot) caused by anamorphs of Botry-
osphaeria spp. (Fusicossum spp., Neofusicoccum spp., L. theobromae (Pat.) (Griff.
and Maubl.)) (Johnson, 1994; Slippers et al., 2005; Crous et al., 2006) and Pho-
mopsis mangiferae Ahmad, and alternaria rot caused by Alternaria alternata
(Fr.) Keissler. The latter is generally only a problem in fruit from dry regions
or in fruit from more humid areas during storage for 3 weeks or more (Prusky
et al., 1980, 1993a, b, and Chapter 7, this volume; Johnson et al., 1990b).
Fruit are moved through a water bath for 5 min at 4850C for less mature
fruit and hot-water-damage-susceptible cultivars (e.g. Zill and Irwin) and
at 5055C for mature fruit and less susceptible cultivars (Anonymous, 1994b).
Treatment for 3 min may be adequate for control of anthracnose, while
immersion for up to 7 min may enhance control of stem end rot (Muirhead
and Grattidge, 1986; Sepiah, 1986; Johnson et al., 1989b). In large-scale facili-
ties, dip tanks may range from 3000 to 5000 l, with fruit immersed and moved
through the tank by a series of paddles. In tank construction, non-corrodible
materials such as stainless steel and breglass are preferred, and the con-
veyor system that contains the paddles should travel along the bottom of the
tank to reduce damage to fruit that sink. Accuracy in temperature control,
efcacy of the heating unit and timing of fruit ow through the bath are
critical. Temperature probe placement at pump inlet and outlet and thorough
water circulation to ensure accurate temperature reading and to minimize
hot spots are critical. Impurities (e.g. minerals, sediment and debris) in dip
water can affect fungicide performance and stain or damage the fruit. In-line
lters in the inlet and pump circulation systems should be installed and
cleaned regularly.
Where acceptable, carbendazim can be added to the hot water at the rec-
ommended rate, and topped up and replaced regularly, to provide improved
control of stem end rot and anthracnose at lower temperatures (52C). Beno-
myl has been withdrawn for postharvest use, but much of the benomyl use
information (from earlier research) is relevant for carbendazim (Johnson et al.,
1997). Also, hot thiabendazole (TBZ) is generally as effective as hot benomyl
for controlling stem end rot, but may provide inferior control of anthracnose
(Coates et al., 1993). The active component of benomyl and TBZ in plants,
carbendazim (MBC), is identical (Erwin, 1973; Muirhead, 1976); however,
TBZ also contains sulfur (S) which affects its rate of breakdown and spec-
trum of activity (D. Guest, personal communication, Melbourne, 1995). Beno-
myl penetrates plant tissue more effectively than TBZ, carbendazim or
thiophanate methyl (Eckert, 1983).
Dipping fruit in hot, dirty, latex-contaminated water can increase phyto-
toxicity and lenticel damage. Hot fungicide dips lose efcacy due to sap
G.I. Johnson and P.J. Hofman 552
build-up in the dip tank and stripping out of fungicide (Wells and Littlemore,
1989). Ledger (2004) optimized dip:fruit ratio and dipping practices. Prochlo-
raz provides good control of anthracnose and alternaria rot, but does not pro-
vide control for stem end rot (Johnson et al., 1990b; Johnson and Coates, 1993).
In South Africa, for local markets prochloraz is added at 405 ppm of active
ingredient (ai) of a 45% emulsiable concentrate (ec) formulation; for export
markets, 810 ppm prochloraz is used. Fruit are immersed for 20 s. In Australia,
prochloraz at 250 ppm is applied by overhead spray, and fruit require 1520 s
to pass through the prochloraz spray race on a roller conveyer system.
A maximum residue level (MRL) of 7.0 mg/kg for prochloraz for assorted
tropical and subtropical fruits with an inedible peel is recommended (CODEX-
MRL, 2008). This group MRL replaces individual fruit commodity MRLs,
and takes into account the lower residues in the esh compared to the skin
(Muller and Burt, 1989). Some registered use-rates for postharvest applica-
tion of prochloraz are listed in Table 15.3.
Hot water sprays over brushes (55C for 1520 s) is an effective alterna-
tive to hot water dips containing prochloraz for controlling alternaria rot
(Prusky et al., 1999, 2006). Application of hot water spraying and brushing for
1520 s (HWB) followed by a spray of 50 mM hydrochloric acid (HCl), alone
or in combination with prochloraz, also improved control of alternaria rot
(Prusky et al., 2006). These treatments have not been tested for anthracnose.
Using 2,4-dichlorophenoxyacetic acid (2,4-D) diluted in wax after HWB and
prochloraz reduces stem end rot (Kobiler et al., 2001). A hot water and beno-
myl combination treatment followed by a prochloraz spray provides effec-
tive control of anthracnose, stem end rot and alternaria rot during longer
storage (Johnson et al., 1989a, 1990b). Similar benets are now attributed to
hot TBZ dip and cold prochloraz spray (Ledger, 2004).
When fungicides are used in the packhouse, spent dip suspensions and
fungicide containers must be disposed using approved methods, often
included with supplier recommendations. Carbendazim suspensions can be
drained into a trench lled with stones, but runoff must be avoided. Car-
bendazim and other benzimidazole fungicides are toxic to earthworms (Wright
and Stringer, 1973).
Table 15.3. Registered uses of prochloraz for postharvest treatment of mango
(Source: adapted from Lunn, 2004).
Country Form
a
Method Rate (kg ai/100 l)
Australia ec 30 s spray 0.025
Brazil ec 2 min dip 0.05
China ec/wp 1 min dip 0.050.1
Colombia ec Not specied 0.025
Peru ec Not specied 0.020.045
South Africa ec 20 s dip 0.08
a
ai, active ingredient; ec, emulsiable concentrate; wp, wettable powder.
Postharvest Technology 553
Heat
Pest disinfestation treatments involving heat provide some control of anthra-
cnose, but do not adequately control mango fruit pathogens for export. Tem-
perature and time combinations suitable for non-deleterious fruit disinfestation
are sublethal to a signicant percentage of quiescent infections beneath the
fruit cuticle and pedicel tissues (Coates and Johnson, 1993). In many regions,
fruit skin temperatures frequently approach the mid-40C range during pre-
harvest development, a natural selection pressure favouring heat-tolerant
fungal infection structures. For Kensington Pride, hot benomyl in combina-
tion with either prochloraz or vapour heat at 46.5C for 20 min controls stem
end rot more effectively than hot benomyl alone and TBZ, alone or in combi-
nation with vapour heat, during storage at 23C for 15 days (Coates et al.,
1993). Disease control in combination with heat disinfestation has been
reviewed by Coates and Johnson (1993) and Jacobi et al. (1994, 2001a).
Future options
Heat is an ideal disease control treatment, since it is environmentally safe and
non-chemical. Its effectiveness would be enhanced if fruit tolerance could be
increased by genetic manipulation or the development of pre-conditioning
treatments. Pre-treatments to render quiescent structures of pathogens more
susceptible to heat would also improve disease control. Measures to increase
efcacy could include other energy sources, chemicals, adjuvants, fumigants
or microorganisms to damage or soften fungal wall structures.
Treatments to delay fruit ripening also limit or reduce disease losses. Storage
quality would benet from the development of cultivars or pre-conditioning
treatments to improve tolerance of fruit for cool storage or controlled atmo-
sphere (CA) and modied atmosphere (MA) storage (Brecht and Yahia,
Chapter 14, this volume). With increasing concerns about the use of chemi-
cals on food (Gullino and Kuijpers, 1994), and in view of current limitations
on heat treatment and storage regime disease control efcacy, non-deleteri-
ous alternatives to synthetic fungicides are required. Alternatives to fungi-
cides for controlling postharvest diseases have been reviewed by Johnson
and Sangchote (1994) and Korsten (2006). Options include: (i) biological con-
trol, i.e. the use of microorganisms to control pathogens (Wilson and Pusey,
1985; Jeffries and Koomen, 1992; Korsten et al., 1993, 1994; Korsten 2006); (ii)
enhanced exploitation of naturally occurring antifungal compounds in fruit
(Prusky et al., 1982; Johnson et al., 1998; Joyce et al., 1999; Zainuri et al., 2003;
Hassan et al., 2007); (iii) application of fruit coatings such as chitosan with
both MA and antifungal effects (El Ghaouth et al., 1992a, b; Wilson et al., 1994;
El Ghaouth and Wilson, 1995); (iv) exposure to UV-C light (wavelength <280
nm) (Chalutz et al., 1992; Wilson et al., 1994). Zainuri (2006) reported some
promise in the use of UV-C radiation for control of anthracnose, but fruit
damage risks and treatment dose accuracy were critical; (v) containment of
fruit in atmospheres containing high levels of carbon dioxide (CO
2
) for 2448
h after harvest (ushing) (Prusky et al., 1992, 1993c); (vi) regulation of fruit
ripening (Brady, 1994); and (vii) application of naturally occurring plant
products (Fallik and Grinberg, 1992; Wagner and Flores, 1994). Many of these
G.I. Johnson and P.J. Hofman 554
options may delay disease development by eliciting increases in antifungal
compounds in the fruit (Prusky and Keen, 1993; Wilson et al., 1994; Zainuri,
2006).
What are the alternatives to synthetic fungicides for controlling mango dis-
eases? Bacteria active against mango isolates of C. gloeosporioides, stem end and
soft brown rot pathogens have been evaluated (Koomen et al., 1990; Korsten
et al., 1991, 1992, 1993; Jeffries and Koomen, 1992; Coates et al., 1995; Korsten,
2006). Antifungal resorcinols in the peel of mango fruit interfere with the devel-
opment of anthracnose and alternaria rot (Cojocaru et al., 1985; Droby et al., 1986,
1987; Prusky and Keen, 1993; Zainuri, 2006; Hassan, 2007), with higher levels
present in some cultivars (Hassan, 2007; Karunanayake, 2007; Hassan et al., 2007).
Lonsdale (1992, 1993) found that enclosure of Keitt in high-density polyethyl-
ene bags with 30% CO
2
and 15% oxygen (O
2
) for 24 h at 11C prior to storage,
improved control of anthracnose. However, 24 h exposure to 20% CO
2
signi-
cantly increased the incidence of soft brown rot (stem end rot) in Keitt and
Kent compared to untreated fruit, especially in the absence of O
2
. UV irradia-
tion of fruit for 1030 s in combination with wax prior to storage is similar to hot
water in reducing the incidence of soft brown rot compared to untreated fruit.
Brushing
Brushing on mango packing lines can occur after, or at the same time as, hot
water and fungicide treatments. Hot water treatment washes sap away, and
loosens supercial debris, scale insect carapaces and sooty mould, which
are removed as the fruit pass over rotating brushes. Brushing also removes
supercial deposits of fungicides that accumulate on fruit from orchard
application of Cu fungicides (Lonsdale, 1993) and incorrect mixing or sedi-
mentation of benzimidazole fungicides resulting from sap accumulation in
dip tanks (Wells and Littlemore, 1989). Soft, non-damaging brushes should
be used, washed every day and replaced seasonally.
For Kensington Pride mangoes harvested after rain, skin marking, fruit
shrivel and weight loss increase signicantly on fruit treated with a hot water
and fungicide dip or a hot water and fungicide dip followed by treatment with
prochloraz, when fruit brushing followed either or both treatments relative to
untreated and untreated/brushed fruit. Prochloraz before brushing resulted
in fruit quality similar to untreated or brushing only (Cooke and Johnson, 1994).
Brushing can increase lenticel spotting (Oosthuyse, 1999). Therefore, the num-
ber and type of brushes must remove foreign matter and polish the fruit, while
not increasing risk of brush and lenticel damage, especially during wet weather
and with heat treatments (see Weather conditions and Skin colour and lenticel
damage sections, both under 15.3 Preharvest Management, this chapter).
Grading and sizing
The purposes of grading are to sort fruit into dened categories of unifor-
mity and to divert out-of-grade fruit from the pack line to either a second
Postharvest Technology 555
grade, processing or reject line. Mangoes with defects outside acceptable lim-
its as dened in a grade schedule or chart are manually removed and trans-
ferred (by conveyer belt) to seconds or processing lines as appropriate. The
purpose of sizing is to categorize fruit into size or weight groups for packing.
Fruit must be sized prior to disinfestation with hot water or vapour heat to
ensure consistent treatment responses. Typical systems include automatic
graders that separate fruit by weight into groupings that correspond to pre-
determined categories (Schoorl and Holt, 1982, 1985). Camera vision systems
can separate for colour, defects and shape. Fruit usually accumulate in sepa-
rate bins for packing into cartons or into bulk containers for processing or
disinfestation. The fruit are packed manually into single-layer trays, with
plastic or cardboard liners that have depressions designed to accommodate
fruit of a particular size. The depressions provide some support for individ-
ual fruit during packing, while the cardboard liners also provide some
additional buffering against impact damage. The pattern of the depressions
facilitates most efcient utilization of carton space. Mango tray liners com-
monly accommodate 1225 fruit for 6.5 kg trays. Some tray liners may be
inappropriate for sea export due to interference with vertical airow.
Organic materials (i.e. paper, leaves or shredded wood) have been
used to cushion individual fruit in cartons. These materials can harbour
pathogens, for example Rhizopus stolonifer (Ehrenb. Fr. Lind.), which
causes transit rot of mangoes and has been detected in shredded wood
used in mango packaging. Shredded wood creates micro-wounds in the
fruit skin, providing points of entry for hyphae growing on the wood.
Losses are more severe when fruit have been removed from cold storage,
allowing condensation to develop on the fruit and shredded wood
( Muirhead and Grattidge, 1986).
Grade standards
The International Standardisation Organisation (ISO) is a non-governmental
organization (NGO), and is a network of national standards institutes (157
countries). ISO is the global leader for development and publication of stan-
dards. ISO publishes a range of standards for fruit and vegetables, testing,
crop and postharvest management procedures and food safety, system
auditing and nutrient and water testing that are relevant to mango systems
benchmarking and improvement. A range of standards may also be dened
within GAP certication protocols and nationally developed marketing
arrangements. Agreed grade standards provide a reference point for produc-
ers and traders in production and marketing (EurepGAP, 2007; ISO, 2008).
The CODEX Alimentarius Commission also oversees the development of
standards for fresh and processed fruit with the CODEX Committee on Fresh
Fruit and Vegetables (CODEX, 2008a). CODEX standards for fresh fruit spec-
ify provisions for quality, sizing, tolerances, presentation, marking and label-
ling, and contaminants (CODEX, 2008a). There are CODEX standards for
fresh mangoes, canned mangoes and mango chutney (CODEX, 2008b).
G.I. Johnson and P.J. Hofman 556
Minimum requirements for grade standards specify that fruit intended
for international trade should be intact, rm, fresh in appearance, sound,
clean, free from black stains and bruising, free from damage caused by low
temperatures and free from pests and pest damage. Fruit should be carefully
picked at the stage of physiological development which will allow transport
and handling and continuation of the ripening process so that fruit will ripen
to consumer expectations. Class standards can depend on customer speci-
cations, and can be based on fruit size and appearance. Colour illustrations
in Anonymous (1993) and Amesbury et al. (2002) are indicative of some of the
quality standards that can be specied for appearance, shape and colour, and
tolerance levels for supercial skin defects. Similar charts are often available
for individual cultivars and are produced during the development of QA
systems for specic marketing groups and customers.
Packing-line QA inspections
Packing-line control inspections are used to monitor grading efcacy and
packing-line damage. Packing-line inspection samples are taken soon after
fruit pass points in the line at which defects are most likely to be overlooked
and/or induced. For start and end-of-line pack-out checks, and out-turn
inspections, randomly selected cartons are unpacked, and all fruit are checked
for compliance with preset quality parameters. Most value is gained from
quality control checks if records are kept and evaluated, with feedback/trou-
ble shooting as necessary, to constantly improve the system (Ledger and Bag-
shaw, 1994; Ledger and Premier, 2006). Computer analysis of such information
provides a seasonal benchmarking record of QA improvement, and high-
lights areas for attention in packing-line improvement and personnel train-
ing. Record keeping is mandatory under GAP certication systems.
Future options
Greater automation of grading and packing will become necessary as pro-
duction and labour costs increase, and as customers become more demanding
(Hilton, 1994). Recent advances in computing have made possible high-speed
sorting using visual systems for colour, shape and externally visible defects.
Also, NIRS systems can now be used in-line to sort for esh characteristics
that inuence avour. In mango, percentage dry matter and esh colour are
related to ripe fruit avour, and can be estimated using NIRS (Saranwong
et al., 2004; Subedi et al., 2007). Estimation is sufciently accurate to allow
acceptable separation into several categories for nal avour. NIRS may also
be useful for predicting ripening behaviour and weight loss during ripening
(Mahayothee et al., 2004). Given the inuence of weight loss in chilling injury
(CI) development during cold storage (Bower et al., 2003), NIRS may also be
able to estimate potential for CI during cold storage.
There is interest in other non-destructive quality assessment for the pack-
house. Joyce et al. (1993) noted that future innovation could lead to proton
magnetic resonance imaging (MRI) technology suitable for packing-line
applications to allow non-destructive detection of internal disorders and pest
infestations. X-ray imaging may have potential for detecting seed weevil
Postharvest Technology 557
damage in mangoes (Thomas et al., 1995; Reyes et al., 2000), but recent inves-
tigations suggest that neither X-ray imaging nor MRI is sufciently reliable
for quarantine purposes, particularly where larvae are small (R.A. Jordan,
personal communication, 2007). New methods of nuclear magnetic reso-
nance (NMR) (Marigheto et al., 2008) may distinguish internal disorders such
as jelly seed. Acoustic/ultrasonic methods can sort for fruit rmness (Miz-
rach, 2008) and may help identify softening fruit with internal disorders and
reduced storage life. Robotics in sorting and packing will be used increas-
ingly where labour costs and availability are high.
Disinfestation
Disinfestation treatments, backed by integrated eld control programmes
and/or area freedom stipulations under market access approvals, provide
assurance to authorities of an importing country that the commodity will be
free of target pests and not pose a quarantine threat (Johnson and Heather,
1995; Follett and Nevin, 2005). Market access application and approval ar-
rangements for most countries operate under national legislation and regula-
tions formulated under the framework of the IPPC and the World Trade
Organisation (WTO) Agreement on the Application of Sanitary and Phyto-
sanitary Measures (IPPC, 2008; SPS, 2008). Key aspects of quarantine regula-
tion of crop pests are covered by International Standards for Phytosanitary
Measures (ISPM), developed and agreed in consultation with signatory cou-
ntries to the IPPC under the Food and Agriculture Organization (FAO) (ISPM,
2007a). ISPM (2007b) cover guidelines on the phytosanitary measures for
regulated pests. Under the IPPC, a technical panel on phytosanitary mea-
sures has oversight of international guidance on phytosanitary treatments
including assessment and recommendations for use as international stan-
dards. In addition to ISPM, a range of regional plant protection measures
has been established. It has been agreed internationally (ISPM, 2007b) that
phytosanitary treatments should full the following requirements:
Provide effective destruction, inactivation or removal of pests or render
them infertile or devitalized. Normally, stipulation of the destruction efca-
cy level, with quantication or statistical benchmarking, is required. When
experimental data are unavailable or inadequate, other evidence is needed
to support a claim of efcacy, for example historical/practical experience.
The technology and treatment regime used should be: (i) clearly delin-
eated, with evidence to conrm adequate adherence to scientic meth-
ods in generating data (experimental designs). Data in support of the treat-
ment efcacy should be veriable, replicable and statistically valid, and
preferably published in a peer-reviewed journal; (ii) appropriate for use in
international or regional trade and research; and (iii) safe to apply, with no
undesirable impacts on treated commodities or the environment.
Approvals for disinfestation treatments and market access are obtained on
a country-by-country basis. A starting point for intending exporters is to
G.I. Johnson and P.J. Hofman 558
determine the disinfestation requirements for mangoes entering target mar-
kets, for example the import health standards for New Zealand (BANZ, 2008)
and the Plant Protection and Quarantine (PPQ) manuals for the USA (PPQ,
2007). Australian exporters can use the Australian Quarantine and Inspection
Service (AQIS) phyto exports database (AQIS, 2008b). The AQIS web site plant
product export database provides summary information (Table 15.4) and
detailed information on phytosanitary requirements for potential exporters.
Follett and Nevin (2005) noted that increased trade has increased exotic
pest threats and attention to quarantine and regulatory issues. Risk-based
alternatives were replacing the probit 9 standard for quarantine efcacy. Cul-
tivar testing was seen as necessary only for some treatments and commodi-
ties, and generic treatments for broad groups of pests and commodities were
seen as a means of enhancing trade. Area-wide pest management was valued
for preharvest pest control and improvement of quarantine security for
export products. However, some treatments such as irradiation were not
accepted by all countries and this slowed their adoption. Follett and Neven
(2005) concluded that efforts for standardization of phytosanitary measures
and research would improve information exchanges and market access nego-
tiations.
Target pests
A pest of regulatory concern that could become established in an area where
it is not found is a quarantine pest risk, and requires quarantine action.
Mango fruit pests include internal pulp feeders (i.e. fruit y immatures), seed
and fruit pulp pests (i.e. mango weevils and fruit caterpillars) and external
pests (i.e. scales, mealybugs, thrips and mites) (see Pea et al., Chapter 10,
this volume). External pests pose detection risks as surface hitchhikers that
Table 15.4. Examples of summary information for exporting Australian mangoes to
the EU and New Zealand (Source: adapted from AQIS, 2008b).
Documentation
Required for
EU New Zealand
Import Permit No No
Phytosanitary Certicate Yes Yes
a
Additional Declaration No Yes
b
Post Entry Quarantine No No
EX188 No No
EX46 No No
Radiation Statement No No
a
Treatment details, including date of treatment, are to be endorsed on the Phytosanitary
Certicate in the treatment section. The treatment is to be shown as: irradiation at minimum
250 gray.
b
The mangoes in this consignment have been treated in accordance with Appendix 12 of the
Bilateral Quarantine Arrangement between New Zealand Ministry of Farming and AQIS.
Postharvest Technology 559
can be detected visually by inspectors. Such pests need to be controlled in the
eld and removed before the fruits are exported. Internal pests, such as wee-
vils, fruit y immatures or larvae of Lepidoptera, pose additional risks
because of difculties of detection and their potential to damage fruit esh
and/or mango seed. Immatures of mango seed weevil (Sternochetus mangiferae
(F.)) occur in mango seed (but not esh) in most of Africa, Asia, Australia, the
Pacic and the Carribbean (Waite, 2002), and are difcult to kill in situ with-
out damaging the market quality of the treated mangoes. Orchard-control
measures and surveying are discussed by Hansen (1991, 1993), Waite (2002)
and Wittenberg (2007). The mango pulp weevil (Sternochetus gravis (F.), syn.
Sternochetus frigidus (F.)) occurs in India, Bangladesh, part of the island of
Palawan in the Philippines and a few other regions in South-east Asia (Waite,
2002; Astridge and Baron, 2007c; Catindig and Kong, 2007; Walker, 2007a). It
causes severe damage to the fruit pulp only (de Jesus et al., 2007).
Follett and Gabbard (2000) concluded that mango seed weevil does not
seriously affect mango yield or marketability. Nevertheless, the seed weevil
is a major quarantine concern for countries which have not recorded it or
claim area freedom, that is Middle Eastern countries and China (Waite, 2002).
Seed and pulp-attacking Lepidoptera pests are quarantine risks in some
countries (Waite, 2002; Walker, 2007b; Yarrow and Chandler, 2007). Entry of
mangoes from countries having mango weevil and other Sternochetus spe-
cies may be restricted or prohibited into countries free of these pests. Exten-
sive surveying, sampling, implementation of eld control measures and/or
area-freedom certication and maintenance may be necessary for approval
of market access (Johnson and Heather, 1995; Waite, 2002). Disinfestation
treatments that ensure weevils are not able to reproduce may be acceptable
when dosages for mortality damage fruit excessively.
Fruit y disinfestation
Tephritidae are the most important mango pests and occur wherever man-
goes are grown (Waite, 2002). Eggs are oviposited below the peel. The wound
provides an opening for microorganisms and scars the peel. Larvae feed and
tunnel throughout the pulp. Fruit ies infest tropical and temperate fruits. It
is the risk to temperate climate fruits and commodities produced in y-free
areas that has prompted the development of quarantine restrictions and
treatments for fruit y hosts.
At present, quarantine treatments against fruit ies are not required for
fruit entering the European Union (EU), despite the large production of tem-
perate fruit in fruit-y-free regions. Fly infestation has not been perceived as
a threat because winter temperatures throughout much of the region effec-
tively prevent establishment of the ies, despite geographical continuity
with the distribution range of the Mediterranean fruit y (Ceratitis capitata
(Wiedemann)). Canada does not require fruit y disinfestation of tropical
produce for the same reason. Exotic fruit y pests could become established
in southern Florida, Texas and California because of their subtropical climate.
The USA requires that mangoes be disinfested by vapour heat, irradiation,
hot water or hot air.
G.I. Johnson and P.J. Hofman 560
In Queensland, Australia, y larvae infestation of mangoes in the mar-
keting chain is rare, despite the widespread occurrence of endemic species of
fruit ies (Bactrocera spp.). Preharvest control measures, and grading out of
coloured fruit at the packhouse, effectively eliminate infestation of most com-
mercial consignments; however, mangoes consigned to the Australian states
in temperate regions free of the ies must be disinfested against fruit ies to
help ensure area freedom of temperate-fruit-production areas (RSPM, 2004;
Jessup et al., 2007). When effective eld control and grade-out of ripening
fruit is in place, the mandatory disinfestation of mature-green mangoes
entering the exclusion zone is probably unnecessary.
Potentially acceptable quarantine treatments that disinfest mangoes
include vapour heat, hot air, hot water immersion, irradiation, quick-freezing,
combination treatments and some miscellaneous treatments (Taylor et al.,
2002; Ducamp Collin et al., 2007). The major constraints in the development
of treatments have been the susceptibility of mangoes to heat, cold and irra-
diation damage and O
2
depletion, and the extensive research and negotiation
required to obtain market access approvals to high-end markets (i.e. the USA,
the EU and Japan). Treatments need to be veried as non-damaging to a
range of cultivars by fruit size by environments likely to be encountered
(Jacobi and Gowanlock, 1995; ISPM, 2007b). Treatments that cannot be used
because they lower fruit quality at dosages that kill pests are methyl bromide
fumigation (Spalding et al., 1977) and cold temperature storage (Kane and
Marcellin, 1978).
Vapour heat
Vapour heat treatment (VHT) involves heating air that is nearly saturated
with moisture, and passing the air stream across the fruit (Jacobi et al., 2001b).
When the temperature of the mango fruit is at or below the dew point of air,
condensation occurs on the fruit surface and rapidly heats the fruit by con-
ductive energy transfer. The core of the fruit next to the seed is heated to
c. 45C for the required time before cooling. Fruit have to be sorted for size
before treatment because of different rates of attaining the required core
temperature.
Vapour heat is used worldwide to disinfest mangoes of fruit ies. Jacobi
et al. (2001b) list the VHT protocols approved for importation of mangoes
into Japan from the Philippines, Taiwan, Thailand, Australia and Mexico.
Conditions range from 4347C pulp core temperature for 10 min to 6 h;
however, the most common treatment conditions are 4647C for 1030 min.
Melon fruit y (Bactrocera cucurbitae Coquillett) immatures in mangoes from
Okinawa were killed at 44 0.3C core temperature for 3 h (Sunagawa et al.,
1987). Taiwanese mangoes infested with melon y can be disinfested with
vapour heat at 47.5C until the centre pulp is >46.5C for 45 min (Kuo et al.,
1987).
A VHT schedule was approved against Queensland fruit y (Bactrocera
tryoni), in Kensington Pride, R2E2, Keitt, Palmer and Kent from Aus-
tralia for the Japanese market (AQIS, 2008b), which consists of a core tem-
perature of 47C for 15 min. The United States Department of Agriculture,
Postharvest Technology 561
Animal and Plant Health Inspection Service Plant Protection and Quarantine
(USDA-APHIS PPQ) approved VHT as a quarantine treatment for Mexican
fruit y (Anastrepha ludens (Loew)) and other Anastrepha species in Manila,
and for mangoes from Taiwan infested with oriental fruit y (Anonymous,
1994a). Generic guidelines for use of VHT in treating commodities for the
USA market are provided by the USDA-APHIS PPQ manual on vapour heat.
Mangoes from Taiwan imported into Australia must be treated until the pulp
temperature has been 46.5C for 30 min (AQIS, 2008a).
Hot air
Hot or forced hot air systems also heat the air to 4050C, but at a lower RH.
Relative humidity usually remains >50%, depending on ambient RH, but is
never high enough to produce condensation. Heat is transferred to the fruit
by convection, with no condensation of water on the skin (Gaffney and Arm-
strong, 1990; Jacobi et al., 2001b). Relative humidity should be high enough to
prevent fruit desiccation during treatment. Transfer of heat from the air to
the skin is slow compared with VHT. Mangan and Ingle (1992) reported that
a mean centre pulp temperature of >47C killed all stages of West Indian fruit
y, Anastrepha obliqua (Macquart), in Mexican mangoes, and Sharp (1992)
found a centre pulp temperature of >46C killed all stages of Caribbean fruit
y, Anastrepha alletis (Loew), in Florida-grown mangoes.
Hot water
Provided that fruit are not damaged, hot water immersion is environmen-
tally safe and efcient for killing mango pests. Use of hot water to kill fruit
y eggs and larvae intensied in the USA when the Environmental Pro-
tection Agency (EPA) removed ethylene dibromide from the market as a
chemical fumigant because of health concerns (Anonymous, 1983). Sharp and
Spalding (1984) showed that mangoes could be disinfested of Caribbean fruit
y using hot water. The work led to more studies in Haiti and a disinfestation
method for West Indian fruit y (Sharp et al., 1988), as well as Mediterranean
fruit y and other Anastrepha spp. in Texas and Mexico (Sharp et al., 1989a, b),
Puerto Rico (Segarra-Carmona et al., 1990) and Peru (Sharp and Picho-Martinez,
1990). Nascimento et al. (1992) developed a hot water treatment for fruit ies
in mangoes in Brazil. Hot-water-treated mangoes may be imported into the
USA from Mexico, Central America, South America and the West Indies
(Anonymous, 1994a). Typical treatments include 46.1C for 65 min for smaller
fruit to 90 min for larger fruit (Jacobi et al., 2001b). Large commercial hot-
water-treatment facilities have been constructed, certied by the USDA-
APHIS PPQ, and used in Mexico, Central and South America, and the West
Indies. Generic guidelines for the use of hot water are provided by the USDA-
APHIS PPQ manual for hot water treatment.
In Australia, Smith (1992) showed that immersing ve Australian mango
cultivars in 48C water for 30 min killed eggs and larvae of Bactrocera aquilo-
nis; however, Kensington Pride is more sensitive to hot water than to vapour
heat, so the latter has been adopted for disinfestation of mangoes in Australia
(Jacobi et al., 1994). Grov et al. (1997) found that treatment of several cultivars
G.I. Johnson and P.J. Hofman 562
in hot water at 46.1C for 90 min followed by refrigeration for 24 h did not
damage fruit, although some cultivars showed severe lenticel damage.
Refrigeration of Tommy Atkins fruit immediately after treatment resulted
in scald development. Weevils in Alphonso mangoes from India were not
killed when infested mangoes were immersed in water at 4852C for up to
90 min and 5470C for up to 5 min (Shukla and Tandon, 1985).
Compared with hot air treatments, hot water treatments can damage the
skin, partly because of rapid heat transfer from the water to the skin com-
pared with from the skin to the centre of the fruit. Damage includes skin scald-
ing, lenticel damage, cavities, white starchy areas in the esh and delayed
ripening (Jacobi et al., 2001b). Several factors inuence damage severity after
heat treatment, for example cultivar, temperature and duration (Jacobi et al.,
2001b). Immature fruit have low heat tolerance, and small fruit are damaged
by heat more readily than large fruit. Conditioning treatments (i.e. 37C core
temperature, for at least 12 h in air) can reduce injury, and preharvest condi-
tions, especially rainfall before harvest, can increase skin damage (Esguerra
and Lizada, 1990; Esguerra et al., 1990; Jacobi and Wong, 1992; Jacobi et al.,
1994, 1995; Jacobi and Giles 1997). Better understanding of these inuences
could increase the commercial potential for hot water disinfestation.
Hot water dips could pose human health risks. Sivapalasingam et al.
(2003) reported that an outbreak of Salmonella enterica that infected 72 patients
from 13 USA states may have been due to contamination of hot-water-dipped
mangoes from a single farm in Brazil. No outbreaks were reported among
consumers in the EU of mangoes from the same farm, and the EU does not
require hot water disinfestation.
Irradiation
Irradiation involves rays (at <1000 Gy), X-rays, electrons and microwaves
(Thomas, 1986; Velasco and Medina, 2004; Follett et al., 2007; Moreno et al.,
2007). A 2005 FAO/International Atomic Energy Agency (IAEA) report indi-
cated that >20 irradiation facilities have been planned, constructed or reno-
vated in ten countries, some of which are mango exporters (Eustice, 2007).
Radiation treatments have been developed for fruit ies in mangoes from
Florida, Mexico, India and Australia. Von Windeguth (1986) treated mangoes
with 76 Gy and disinfested them of Caribbean fruit y eggs and larvae. Third
instar Mediterranean fruit y larvae in Mexican mangoes irradiated with 250
Gy did not emerge from pupae, and 60 Gy applied to third instar Mexican
fruit y, and West Indian fruit y in Mexican mangoes prevented adult emer-
gence (Bustos et al., 1992). Bustos et al. (2004) recommended a generic dose of
150 Gy for control of Mexican fruit y (A. ludens), the West Indian fruit y (A.
obliqua), the sapote fruit y (Anastrepha serpentina) and the Mediterranean
fruit y (C. capitata) in mango. Kensington Pride mangoes infested with
eggs and larvae of Queensland fruit y and Bactrocera jarvisi (Tryon) are
disinfested with 74101 Gy (Heather et al., 1991).
International guidelines for the use of irradiation as a phytosanitary
measure are available (ISPM, 2003), and recently a fast track process has been
proposed as an Annex to ISPM 28 (ISPM, 2008), which endorses irradiation
Postharvest Technology 563
at 70 Gy as a generic treatment to control Anastrepha spp. in fruit and vegeta-
bles by extrapolating work on mango by Bustos et al. (2004). Heather (2004)
provides generic guidelines for the development of irradiation protocols for
disinfestation. Fruits are never exposed to radioactive materials (Anony-
mous, 1986) and most modern treatment units use an electron beam process
rather than a radioactive source for irradiation.
Irradiation can be used for controlling seed weevil and lepidopterous
pests in fruit. Seo et al. (1974) reported that 206 and 329 Gy killed mango
weevil in Hawaiian mango. Thomas (1975) showed that 500 Gy killed all
mango weevil larvae and pupae and 750 Gy prevented adults from emerging
from mangoes in Africa. A dose of 500 Gy, however, did not disinfest
Alphonso mangoes of seed weevil (Shukla and Tandon, 1985). Indian man-
goes from approved packhouses must be irradiated with a minimum of 400
Gy at an approved and certied irradiation treatment facility using Cobalt-60
(APEDA, 2007). A quarantine treatment of 300 Gy has been approved to ster-
ilize mango seed weevil in mangoes exported from Hawaii to USA mainland
markets (Follett, 2004). Follett and Lower (2000) demonstrated control of
Cryptophlebia illepida (Butler), Cryptophlebia ombrodelta (Lower) and Cryp-
tophlebia illepida (Lepidoptera: Tortricidae), and an irradiation quarantine dose
of 250 Gy has been approved for Hawaiian mangoes. The treatment also
controls fruit ies (Follett, 2004).
USA regulations covering irradation are described in the Code of Federal
Regulations GPO Access (2008), revised annually (Wall, 2008), and this sum-
marizes approved treatments for a range of pests (EPA, 2002) (Table 15.5),
Table 15.5. Minimum absorbed dose of gamma irradiation required by USDA for specic
pests (Source: adapted from EPA, 2002).
Scientic name Common name Minimum absorbed dose (Gy)
Anastrepha ludens Mexican fruit y 70
Anastrepha obliqua West Indian fruit y 100
Anastrepha serpentina Sapote fruit y 100
Anastrepha suspensa Caribbean fruit y 70
Bactrocera cucurbitae Melon fruit y 150
Bactrocera dorsalis Oriental fruit y 150
Bactrocera jarvisi Jarvis fruit y 100
Bactrocera tryoni Queensland fruit y 100
Brevipalpus chilensis False red spider mite 300
Ceratitis capitata Mediterranean fruit y 150
Cryptophlebia illepida Koa seed worm 250
Grapholita molesta Oriental fruit moth 200
Sternochetus mangiferae Mango seed weevil 300
All other fruit ies of the family Tephritidae which are not
listed above
150
Plant pests of the class Insecta not listed above, except
pupae and adults of the order Lepidoptera
400
G.I. Johnson and P.J. Hofman 564
many of which can infest mangoes. The USDA-APHIS PPQ manual on irradia-
tion provides generic guidelines. Irradiation was approved for the USA market
as a phytosanitary treatment for all fresh fruits and vegetables from all coun-
tries in 2002. Effects of -irradiation on mango fruit quality and disease control
have been reported (Mitchell et al., 1992; Moreno et al., 2006; Reyes and Cisneros-
Zevallos, 2007; Wall, 2008). Only marginal disease control was obtained with
Kensington Pride at the highest non-deleterious doses for mature-green fruit
(300 Gy), with additive effects of disease control treatments and irradiation on
disease reduction (Johnson et al., 1990a). Disease control may be more effective
in cultivars with greater tolerance of irradiation (van der Linde and Thord-Gray,
1986; Johnson et al., 1990a). Other types of irradiation have been evaluated
for mango disinfestation but none has been adequately suitable.
Quick freezing
Quick freezing of mango, lowering the temperature to 17C and holding at
6C or below for 48 h is used to disinfest mangoes for processing (Anony-
mous, 1994a; PPQ, 2007). The process is not approved for importing man-
goes with seeds from most of the West Indies, French Guiana, all countries
outside of North, Central and South America, Oceania, Hawaii, South-east
Asia, the Philippines and the Republic of South Africa into continental USA
because mango weevil could be present (Anonymous, 1994a; PPQ, 2007).
Fumigation
Fumigation is an ideal methodology for ensuring effective control when the
fumigant is effective and safe to use. Until 1994, New Zealand required fumi-
gation of mangoes from Australia, the Cook Islands and the Philippines
using 33, 29 or 22 g/m
3
ethylene dibromide at 1015, 15.519.5, or 20C and
above, respectively, at normal atmosphere pressure (NAP) to disinfest man-
goes of fruit ies before entry. As part of the international phase-out of ozone-
depleting substances, the process was banned in 1994 (Anonymous, 1992;
N.W. Heather, personal communication, Brisbane, 1994) and most applica-
tions as a fruit fumigant have ceased worldwide. Methyl bromide was phased
out completely in the USA in 2005, but some emergency uses for quarantine
applications may be permitted, e.g. to destroy a serious quarantine pest in an
imported consignment or to meet ofcial requirements of an importing coun-
try (EPA, 2008). Mangoes imported into Australia from countries where fruit
ies occur must be fumigated with 1635 g/m
3
ethylene dibromide for 2 h at
2126C or above (Anonymous, 1985, 1988).
Phosphine is widely used as a fumigant of durable produce (grains and
tobacco). It provides effective control of fruit y larvae and other pests in
temperate fruits under experimental conditions (Horn and Horn, 2004).
However, phosphine when mixed with water is highly explosive and the
vapour is toxic to humans, so prospects for utilization are not strong.
Miscellaneous treatments
CHEMICAL TREATMENTS. Postharvest chemical treatments using dimethoate are
effective against Queensland fruit y with Kensington Pride (Swaine et al.,
Postharvest Technology 565
1984). The treatment is required for Australian-grown mangoes entering all
Australian states except Queensland and New South Wales, but is under review.
The USA and the EU do not allow the use of chemicals to disinfest mangoes.
NATURAL PRODUCTS. The short shelf life of mango and the high level of insect
mortality required obviates the use of natural products for disinfestation.
Suhaila and Halim (1994) reported the potential of low toxicity, insecticidal
compounds from edible plants that may be effective for topical application to
harvested fruit. Extracts of black pepper (Piper nigrum) were particularly active
in laboratory tests against vinegar y (Drosophila melanogaster (Meigen)).
ATMOSPHERES. CA and MA regimes could have potential for disinfesting man-
goes, but there has been less interest in the technology because heat treatments
and irradiation are faster (Ke and Kader, 1992; Yahia and Tiznado-Hernandez,
1993; Yahia and Vazquez-Moreno, 1993; Yahia, 1994; Len et al., 2000). Treat-
ments are limited to regimes which do not adversely affect ripe fruit quality.
Len et al. (2000) found that CA of 1% O
2
and 30 or 50% CO
2
disinfested
Manila mangoes of A. obliqua, but damage (as spongy tissue) was unaccept-
ably high.
Shrink-wrapping has been ineffective as a quarantine treatment to disin-
fest mangoes of fruit y immatures. Gould and Sharp (1990) reported that
the time needed to disinfest Florida-grown mangoes infested with Caribbean
fruit y eggs and larvae exceeded the shelf life of wrapped mangoes.
COMBINATION TREATMENTS. Serial applications of two or more treatments, which
alone do not achieve quarantine security, have been used to disinfest mangoes.
Seo et al. (1972) reported that eggs and larvae of Mediterranean fruit y, orien-
tal fruit y and melon y were killed in mangoes immersed in water at 46.3C
for 120 min and then fumigated with ethylene dibromide. Lin et al. (1976)
reported that all oriental fruit y and melon y larvae in Taiwan-grown man-
goes were killed when fruit were immersed in 4850C water for 120 min,
hydrocooled, dried and cooled, and then fumigated with ethylene dibromide.
Controlled Atmosphere/Temperature Treatment System (or CATTS)
technology applies a short heat treatment in a low O
2
/high CO
2
environ-
ment, and controls quarantine insect pests while maintaining commodity
quality (Mitcham, 2007; Neven, 2008). Trials using CATTS with mangoes have
been conducted in Australia with promising results. Varith et al. (2007) evalu-
ated a microwave-vapour heat treatment (MW-VHT) disinfestation technol-
ogy for mangoes: the microwave component for pre-heating and the VHT
component for the holding process. Temperatures of 4655C and holding
times of 220 min effectively disinfested fruit of oriental fruit y eggs without
effects on physico-chemical parameters, compared to untreated fruit. There
was less heat damage compared with conventional VHT only fruit. MW-VHT
shortened the process time by 90% compared with the conventional VHT.
PACKAGING. Some markets, for example Japan and the USA, require that fruit
must be packed into insect-proof packages following disinfestation to preclude
G.I. Johnson and P.J. Hofman 566
reinfestation during transportation or storage. The disinfestation facility
feeds fruit into an insect-proof area within which waxing (optional), grading
and packing occur.
15.7 Preparing Fruit for Market
Surface coatings
Surface coatings are used to improve fruit appearance and to alter gas per-
meability to reduce moisture loss or retard ripening. Commercial use of sur-
face coatings on mango fruit needs to be considered carefully because of the
ne balance between benecial and undesirable effects on fruit quality. Neg-
ative effects of coatings include reduction in chlorophyll loss (Fonseca et al.,
2004a), anaerobic conditions and off-avours (Amarante and Banks, 2001)
and skin damage, possibly due to cytotoxic reactions with other components
in the coating formulation (Bower et al., 2003). Generally, coatings have less
effect on delaying ripening during cold storage, compared with extending
the shelf life at typical ripening temperatures (Amarante and Banks, 2001).
Less signicant effects are observed in more mature and in ripening fruit.
Coatings often delay skin colour change rather than softening, which increases
the risk of soft, green fruit with less consumer appeal.
Coatings are generally emulsions of synthetic (e.g. polyethylene) or nat-
ural (e.g. polysaccharides, carnauba, beeswax, etc.) origin. Surface coatings
containing waxes, oils (e.g. carnauba, beeswax, etc.) and resins (e.g. shellac)
have a greater effect on limiting water loss then reducing O
2
and CO
2
perme-
ability, compared with those containing polysaccharides, (e.g. those based on
cellulose) (Amarante and Banks, 2001). Formulations based on shellac result
in a shinier appearance than those based on carnauba wax and polysaccharide-
based waxes (Baldwin et al., 1999; Hoa and Ducamp, 2008).
Factors other than coating formulation can affect fruit gas permeability,
i.e. cultivar, variations in skin permeability between fruit, inconsistency in
coating thickness during application, interference from water during appli-
cation causing coating cracking and coating thickness and evenness-of-
spread over the fruit surface. The effect of coating on fruit quality can vary
with holding conditions because of larger temperature effects on respiration
rate than on coating permeability.
Shorter and Joyce (1994) found commercially formulated Avocado and
Passionfruit Wax, a polyethylene and shellac emulsion, and Technimul 9122
Wax, a polyethylene-based emulsion, were acceptable with Kensington Pride
mango, while Peach Wax, a polyethylene-based emulsion and starch solution
was unacceptable. With Peach Wax, deleterious modied atmosphere effects
on colour development, softening and avour were obtained (El Ghaouth
et al., 1992b; Shorter and Joyce, 1994). Coating Tommy Atkins mango with a
carnauba-based coating and BeeCoat (based on beeswax) reduced water loss,
shrinkage, chlorophyll breakdown, CI and decay after cold storage, and Bee-
Coat also reduced red lenticel discoloration (Feygenberg et al., 2005). With
Postharvest Technology 567
Tommy Atkins, polysaccharide and carnauba-based coatings modied the
atmosphere within the fruit and reduced decay, but only the polysaccharide-
based coating delayed ripening (Baldwin et al., 1999). The carnauba-based
coating signicantly reduced water loss compared with the polysaccharide-
based coating treatments; carnauba-based coatings result in lower water per-
meability and higher O
2
/CO
2
permeability.
Coatings may reduce surface defects. Excessive water loss is associated
with increased skin CI in avocado and mango, and carnauba-based coatings
reduce CI in cold-stored mangoes (Bower et al., 2003). In this study, the carnauba-
based coating contained numerous holes, which allowed respiration gas
exchange (thereby preventing anaerobic respiration), while still providing
efcient control of water loss. Surface coatings may also reduce sapburn, skin
browning and lenticel damage (Shorter and Joyce, 1994), but incorporating
these potential benets into commercial systems may be difcult.
Waxes should be applied by roller brushes in a specically designed wax
applicator or by very light hand application. Dipping fruit in a wax emulsion
is not recommended. A uniform ow of fruit through the wax applicator
must be maintained to prevent uneven wax application. Fruit should be dry
before entering the wax applicator, otherwise foaming of water-emulsion
waxes may occur. Brushes on the wax applicator need to be completely satu-
rated with the wax mixture before any fruit passes over them. Complete cov-
erage of the entire fruit surface is essential. Patchy application can be caused
by insufcient wax, too few brushes following application (minimum of six
brushes required), poor and/or inadequate drying facilities, and overload-
ing of the unit. Brushes should be kept soft with regular washing with hot
water.
Packaging
Packaging provides conveniently sized carriage units for product, protects
individual fruit from contact rub and compression damage, and excludes
dirt, pests and contaminants. McGregor (1987) and Hilton (1994) discussed
key aspects of packaging for tropical produce. Packaging is also a marketing
tool. Design and colours of symbols and text on carton exteriors portray a
marketing image. Manufacturers of consumer products exploit packaging to
great advantage (along with advertising) to increase both rst-time and
repeat sales. Marketing and design professionals may be involved in the
development and customer evaluation of product packaging. Cultural pref-
erences need to considered, e.g. use or avoidance of red for some Asian mar-
kets. Packaging is the external face of brand loyalty. Consistent product
performance and quality is the core.
Some constraints to packaging may be specied by market regulations,
including carton dimensions and labelling requirements. Country of origin,
cultivar, grower, packing shed, market agent, count (number per carton and
weight range) and class may be required. The word mangoes should be
clearly visible (Anonymous, 1993). The information appears on the narrow
G.I. Johnson and P.J. Hofman 568
sides of the cartons. Storage and product use information can also be printed
on the cartons. Many QA systems require adequate labelling linked to appro-
priate record keeping for plate-to-farm traceability. Clear labelling facilitates
correct delivery, allows immediate buyer recognition of product prole and
ensures maintenance of accurate sales records. An exporting country may
nd it of value to identify individual packers by barcoding or numbers
stamped on cartons, so that sources of faulty packaging can be traced. Some
countries also use date codes which enable exporters to determine the fresh-
ness of the produce at the point of export and evaluate an importers capacity
to achieve adequate turnover of the fruit without prolonged storage. It also
provides invaluable feedback on the efciency of the total distribution chain.
Cartons used for export should be clean, strong, unbroken and new. The
water absorption capacity of the material should be evaluated as excess
absorption will lead to collapse on the pallet. The cartons strength will
depend on the starch used by the manufacturer, the outer liner and the direc-
tion and numbers of uting in the carton (Anonymous, 1994b). There is
increasing pressure in the EU for recyclable packing material. Cartons that
are recyclable should be marked with the appropriate international symbol.
Returnable plastic crates are increasingly being used for domestic trade, but
the return cost would make this less protable for international trade.
Inspection
In some countries, independent inspectors check the fruit prior to palletizing
to ensure that the relevant marketing, residue and phytosanitary standards
have been met. Fruit for Japan is disinfested under the supervision of a Japa-
nese inspector. Further inspections are usually made at the port of exit. Some
exporting countries require a declaration by the grower to ensure that fruit
will comply with the standards specied by importing countries.
Palletizing
Handling mangoes on pallets allows convenient movement of large volumes
of fruit. McGregor (1987) described critical features and arrangements for
loading. The disadvantages of pallets for export are the cost, lower numbers
of cartons per sea container and loss. Some domestic markets have pallet
share systems. Relevant markets and transporters should be consulted con-
cerning required pallet dimensions and appropriate access for fork-lift sys-
tems. The correctly sized pallet, for example as designated by the ISO, which
is designed to t snugly into a standard sea container, should also be used for
the local market.
Precision stacking with each box tted exactly on top of the one below
minimizes risk of damage. Collapsed or lopsided pallet stacks have usually
been due to careless stacking and/or loose placement in the shipping con-
tainer. Pallet slats should not block ventilation holes in the cartons. Cartons
Postharvest Technology 569
should be register-stacked so that ventilation is continuous. Link sheets, which
bind the cartons together at intervals, should also be designed to ensure con-
tinuous ventilation through the pallet. In the cold room, pallets should not be
stacked against a wall or placed directly against each other (Boelema, 1987).
Precooling
Precooling removes eld heat from the product and lowers the temperature
to that required for ripening, transportation or storage. Precooling also
reduces the cooling demand on any in-transit cooling system. Precooling
concepts and systems are described by Thompson et al. (2002). Forced-air
cooling systems efciently and rapidly remove eld heat, and are preferred
for bringing fruit to storage temperature. High RH systems are preferred as
they reduce fruit water loss. Hydrocooling can increase the risk of infection
by wound pathogens (i.e. Rhizopus spp.) and are less effective with large fruit.
Kitinoja and Kader (2003) describe low-cost cooling facilities for use in devel-
oping countries.
Ethylene and ripening
Induction of ripening is routinely employed with mangoes. There are effec-
tive low technology methods involving calcium carbide (releases acetylene
which mimics ethylene) or the leaves of particular trees (Lizada, 1994). More
sophisticated systems include generation of ethylene from ethanol using
catalytic conversion, pure ethylene gas, or a mixture of ethylene and an inert
gas (CO
2
or N
2
) to reduce the risk of explosion with 330% ethylene in air
(Reid, 2002). A number of automatic ethylene control systems are available
(PDS, 2008) to maintain ethylene concentrations within required limits.
Climacteric fruit have differing sensitivities to ethylene. Kensington Pride
mango is sensitive to concentrations as low as 0.01 l/l (OHare et al., 1994).
Ripening is enhanced with concentrations up to 510 l/l, with very little ben-
et at >50100 l/l (Nguyen, 2003). There is more yellow colour on the ripe
fruit when ripened at 20C with 10 l/l ethylene for 3 days compared with no
ethylene, resulting in a more attractive appearance. Also, diseases are gener-
ally less in these fruit (Table 15.6), presumably because fruit ripen more quickly
with less time for disease development. Good ethylene treatment can improve
presentation appearance and increase saleable life (dened as the days from
when the fruit reach at least 60% yellow skin colour to when the fruit had lost
saleability because of disease) (Ledger et al., 2002a).
Ethylene can also reduce quality if not used appropriately. Ripening
Kensington Pride fruit at <18C with ethylene can result in soft fruit with
less yellow skin colour, most likely because ethylene stimulated softening to
a greater extent than chlorophyll loss (Nguyen, 2003). Ripe fruit disease can
also be greater. These effects can be aggravated with concentrations above
100 l/l (Fig. 15.5). Kensington Pride fruit must be cooled to <24C before
G.I. Johnson and P.J. Hofman 570
the start of ethylene treatment; otherwise, skin spotting can develop (Ledger,
2003a). Ripening at 1822C is recommended for maximum yellow skin colour,
less disease and higher avour volatiles (Hofman, 1997; Lalel et al., 2004).
The relatively high respiration rate of ripening mangoes can result in
CO
2
accumulation in the ripening room, particularly if the room is full and
there is poor ventilation. Carbon dioxide concentrations up to 5.3% have
Table 15.6. Days for fruit to reach the eating soft stage (days to ripe at 20C),
percentage weight loss/day and percentage of fruit surface area affected by stem
rots in Kensington Pride mango fruit treated with 25 l/l 1-methylcyclopropene
(1-MCP) for 14 h at 20C followed by exposure to 100 l/l ethylene for 24 h at 20C.
Fruit were then ripened at 20C. Means followed by the same letter in each column
are not signicantly different (P >0.05) (Source: Hofman et al., 2001).
Treatment Days to ripe Weight loss (%)/day Stem rots (%)
Untreated 13.6
b
0.3
a
9.6
b
Ethylene 7.9
a
0.4
b
1.3
a
1-MCP 18.7
c
0.3
a
18
c
1-MCP + ethylene 18.2
c
0.3
a
25.8
d
Ethylene concentration (l/l)
0 10 100 1000 0 10 100 1000 0 10 100 1000
G
r
e
e
n

c
o
l
o
u
r

(
%
)
0
1
2
5
10
20
30
40
50
60
70
24 h
72 h
Treatment at 15C Treatment at 20C
Treatment at 25C
Fig. 15.5. Effect of ethylene concentration and time in ethylene and ripening
temperature on the percentage skin surface area with green colour of Kensington
Pride mangoes at eating soft; least signicant difference = 5.16 (P <0.05) (Source:
Nguyen et al., 2002). Note the increased green colour on the skin of ripe fruit with
lower ripening temperature and higher ethylene concentrations and duration.
Postharvest Technology 571
been recorded in ripening rooms (Ledger, 2007), which can cause more green
colour and a dull appearance on the ripe fruit (Nguyen, 2003). Ripening room
CO
2
concentrations should be maintained at <1% with adequate ventilation
to minimize fruit quality loss (Kernot et al., 1999; Ledger, 2007).
Accidental exposure of mangoes to ethylene and its analogues from adja-
cent ripening rooms, exhaust fumes from internal compression engines or
wound ethylene produced from damaged/ripening fruit can cause prema-
ture ripening. Various systems can remove unwanted ethylene, for example
oxidizing mechanisms such as potassium permanganate either in sachets or
in ethylene scrubbing units in storage rooms, catalytic oxidizers or ozone-based
systems (Reid, 2002). Smartfresh (active ingredient 1-methylcyclopropene;
1-MCP) is a relatively new approach for preventing undesirable ethylene
effects. 1-MCP is a structural analogue of ethylene and irreversibly binds to
the ethylene receptors in the plant, thus preventing ethylene-initiated ripen-
ing. Ripening re-commences as additional ethylene-receptor sites are pro-
duced in the fruit (Blankenship and Dole, 2003). Generally, Smartfresh
treatment is applied in well-sealed cold rooms or plastic tents as soon as pos-
sible after packing. 1-MCP concentrations of 250200,000 l/l for 12 h are
optimum for delaying ripening (Jiang and Joyce, 2000; Hofman et al., 2001;
Adkins et al., 2002; Penchaiya et al., 2006), although most reports state 250
1000 l/l. 1-MCP treatment completely negated any effect of subsequent eth-
ylene on ripening, and can almost double the days to eating soft compared
with ethylene-treated fruit ripened at 20C (Table 15.6) (Adkins et al., 2002).
However, the 1-MCP effects were less in more mature fruits (Alves et al.,
2004), and ethylene exposure before 1-MCP will negate any 1-MCP benet
(Adkins et al., 2002). Any benecial effects of 1-MCP also appear to be less
with longer-term storage (Hofman, unpublished results). 1-MCP treatment
can cause more disease on ripe fruit, because the longer days to ripen allows
more disease development (Hofman et al., 2001). Sourcing fruit from well-
managed orchards can help minimize this effect (Adkins et al., 2005).
For some domestic markets, on-farm treatment of mangoes with ethyl-
ene is used to ensure that fruit have more attractive colour when they are
displayed at the wholesale market 4872 h after dispatch from the farm. This
practice improves returns as fruit can be delivered to retail outlets ready-to-
eat. Ethylene induction of ripening is undesirable for more distant markets
because fruit arrive at the market too ripe for sale, with greater risk of bruis-
ing and disease.
15.8 Pre- and Post-shipping Storage
Cool storage
Cool storage is important when delivery time from harvest to the consumer
exceeds the typical ripening time (510 days). The ideal storage temperature
is dictated by the risk of CI, fruit ripening and disease development during stor-
age, and storage time. CI is rst noted as greying of the skin, which intensies
G.I. Johnson and P.J. Hofman 572
with lower temperatures and longer duration (Phakawatmongkol et al., 2004;
Suresh et al., 2004). In more severe cases esh discoloration and abnormal
ripening can occur. CI development can occur at regimes of 312C (Sadasi-
vam el al., 1971; Thomas and Oke, 1983; Chaplin et al., 1986 a, b, 1991a, b;
Smillie et al., 1987; Thomas and Joshi, 1988; Medlicott et al., 1990b). Longer
storage times require greater care with temperature selection, the quality of
the fruit being stored and conditions before and after harvest. Storage should
be for the minimum period necessary. The following factors affect the opti-
mum storage temperatures and durations:
Genetic differences cultivars differ in chilling sensitivity (Phakawat-
mongkol et al., 2004).
Maturity less mature fruit ripen more slowly at a given temperature,
and are more prone to CI and other storage-related disorders (Medlicott,
1985; Medlicott et al., 1987, 1990 a, b; Oosthuyse, 1993). Such fruit may
not soften at all when exposed to temperatures that are suitable for stor-
age of more mature fruit. In South Africa, adequately mature fruit can be
stored at 810C for 2128 days (Oosthuyse, 1994). Placement in cold
storage without delay and post-storage exposure to temperatures that
promote ripening (e.g. 20C) are important preconditions for success.
Duration of storage Kensington Pride mangoes can be stored at 10 C
for 3 weeks or at 7C for 2 weeks, after which skin colour development
can be affected (McLaughlan and Wells, 1994). Generally, the shorter the
storage time, the greater the tolerance to storage temperatures outside
the 1012C range.
Delays between harvest and cold storage, and ripeness stage the longer
the delay between harvest and cold storage, the greater the risk of ripen-
ing during storage. This applies particularly for more mature fruit. If
prolonged, a delay may render refrigerated storage ineffective in pre-
venting fruit from becoming soft during transit, despite the apparent ab-
sence of softening on dispatch (Oosthuyse, 1994). Fruit should be picked,
packed and placed in cold storage within 24 h. For fruit that have rip-
ened, storage temperatures of less than 8C can be used for up to 21 days
without deterioration in quality during storage; however, the fruit will
deteriorate rapidly after removal from storage (Van Straten and Oost-
huyse, 1994). Some cultivars may be more sensitive to ripe storage, since
Kensington Pride fruit at the mid-climacteric stage will start to lose
appearance after 3 days at 10C because of increased disease and mild CI
(H. Nguyen et al., 2004).
Disease load and fruit tolerance of disease certain mango cultivars are
very tolerant of postharvest pathogens (e.g. see Hassan, 2007). The con-
ditions under which mangoes are grown may be unfavourable for infec-
tion. In these situations, storage temperatures can be higher to reduce the
risk of CI.
Development of CI in mango and other fruits is closely associated with
antioxidant activity (Arafat, 2005; Kondo et al., 2005). Mango fruit held at 6C
for 1020 days had lower antioxidant activity in the skin compared with fruit
Postharvest Technology 573
stored at 12C. Application of several jasmonate derivatives before storage
reduced CI at 67C (Gonzlez-Aguilar et al., 2000; Kondo et al., 2005), pos-
sibly through an antioxidant mechanism. Other chemical treatments can
also reduce CI. Polyamines occur naturally in fruit and decrease during
storage under chill-inducing conditions, and application before storage can
reduce CI (Nair et al., 2003). Salicylic acid appears to be involved in cell
wall stability. Application of methyl salicylate, which breaks down to sali-
cylic acid, signicantly reduced CI in Zill mangoes stored at 7C (Han et al.,
2006). 2,4-D can also reduce mango CI, possibly through interaction with
natural plant hormones and antioxidant levels in the fruit (Wang et al.,
2008). Some of these treatments could have commercial application, but
may have residue implications.
Decay is a major limitation to storage life. The incidence of postharvest
decay on fruit that ripen after refrigerated storage is positively related to the
duration of storage and the extent of ripening during storage (Oosthuyse,
1991, 1992, 1994). Disease development after post-storage exposure to ripen-
ing temperatures can be reduced by minimizing the shipping period and by
storing fruit at temperatures that inhibit softening and ground skin colour
development. If CI occurs, disease develops earlier and will be more exten-
sive (Oosthuyse, 1990).
Controlled and modied atmosphere storage
Decreasing the O
2
and/or increasing the CO
2
concentration can have several
advantages with respect to storage (i.e. reduced ethylene production, better
avour retention, slowing softening and green skin colour loss and reduced
CI) (Thompson, 1998; Yahia, 2006). However, if the O
2
concentration is too
low (dependent on cultivar, storage temperature, fruit maturity and ripeness
stage) anaerobic respiration will commence, with associated production of
ethanol and acetaldehydes, leading to off-avours and physiological disor-
ders (Bender et al., 2000). Atmosphere modication generally has less benet
for tropical fruit compared with temperate fruit, but does have commercial
potential for sea freight to distant markets. Atmosphere control can be active
or passive, or combinations of the two. Surface coatings (see Surface coatings
section under 15.7 Preparing Fruit for Market, this chapter) also provide
modied atmosphere inside the fruit.
With CA systems, O
2
and CO
2
concentrations are actively monitored and
controlled by injecting N
2
and CO
2
, or bleeding air into the container as
required. In more passive systems, such as the MaXtrend

system (Maxtend,
2008) fruit respiration directly lowers O
2
concentrations, and its concentra-
tion is monitored and manipulated by venting as required. In some cases, the
container is ushed with N
2
at the start of storage to rapidly establish the
desired atmospheres. Excess CO
2
is absorbed with hydrated lime. In MA sys-
tems, atmospheres are modied by placing a semi-permeable membrane
around the fruit (usually plastic lm), and relying on fruit respiration to
modify the atmosphere.
G.I. Johnson and P.J. Hofman 574
McLauchlan and Barker (1994) suggested 4% CO
2
and 24% O
2
for CA
storage of Kensington Pride mangoes at 13C, and recommended further
research on atmospheres <2% O
2
and >10% CO
2
. Oxygen had the biggest
effect on retarding skin colour and softening, with signicant retardation
when decreasing from 4 to 2%. Subsequent research suggested that concen-
trations of 1.52% may be more effective in retarding softening, although
these concentrations may increase the risk of off-avours. Tommy Atkins
and Haden can tolerate 23% O
2
for 23 weeks at 12C, but lower concentra-
tions were not tested (Bender et al., 2000). In Kensington Pride there was
little additional capacity for CO
2
concentrations between 6 and 10% to retard
softening or loss of green colour (McLauchlan and Barker, 1994), although
there may be some benet for storage of 12 weeks at concentrations >10%
(Bender et al., 2000). For Delta R2E2 mangoes 3% O
2
and 6% CO
2
have been
recommended (Lalel and Singh, 2006); however, this is a rm-eshed culti-
var, which can perhaps tolerate higher O
2
concentrations to improve vola-
tiles, compared with the softer Kensington Pride. Longer storage times with
CA could cause higher disease levels (Johnson et al., 1990b), higher acidity in
the esh at eating soft (McLauchlan and Barker, 1994; Bender et al., 2000) and
slower loss of green colour compared with non-stored fruit (Bender et al.,
2000). For cultivars that normally have higher acidity, CA-stored fruit may
need to be ripened for several more days to lower acidity.
Cold storage and CA can reduce volatiles production following ripening
at room temperature. As the skin CI severity increases with decreasing stor-
age temperature, total volatiles production appears to decrease (Singh et al.,
2004). In Kensington Pride and R2E2, CA storage signicantly reduces
total concentrations of aroma volatile compounds compared with air-stored
fruit, irrespective of storage period between 2438 days (Singh et al., 2004;
Lalel and Singh, 2006). Decreasing the O
2
concentration from 3 to 1% at 6 or
8% CO
2
or increasing CO
2
concentration from 6 to 8% signicantly increased
most of the monoterpenes, including terpinolene. Cold-stored fruit are
known to have less aroma than those ripened without storage.
Fruit can tolerate short periods with <1% O
2
or >20% CO
2
(Yahia, 2006).
This has been utilized for insect disinfestation (see Disinfestation section
under 15.6 Packhouse Measures, this chapter). Mango can tolerate low O
2

concentrations for 5 days at 20C (Yahia, 1994). These short-term CA treat-
ments may improve storage life or reduce CI during subsequent cold storage
without atmosphere modication; this has been noted with avocado (Truter
and Eksteen, 1987; Pesis et al., 1994). Preliminary investigations suggested
little benet of 2060% CO
2
for 18 days before cold storage of Kensington
Pride (Meiburg et al., 1998).
The optimum conditions for storage of each product to provide maximum
storage life without quality loss must be determined taking into account
cultivar, season and growing conditions. Measurement of chlorophyll uo-
rescence has been used to monitor product performance under CA, with ad-
justment of gas conditions to achieve the optimal storage-life/quality balance.
Changes in chlorophyll characteristics and therefore chlorophyll uores-
cence under CA occur before CI symptoms are obvious (DeEll and Toivonen,
Postharvest Technology 575
2003a). Thus monitoring changes in chlorophyll uorescence characteristics
can provide advance warning of the potential for CI, and allow adjustment
of storage conditions to minimize its development (DeEll and Toivonen,
2003b). This concept has now been marketed as HarvestWatch (Harvest-
Watch, 2008), and some preliminary success has been obtained with apples
(Stephens and Tanner, 2005; DeLong et al., 2007).
Modied atmosphere packaging (MAP) generally cannot reliably achieve
the low O
2
concentrations required to signicantly delay softening without
damaging the fruit, but MAP can still have benecial effects relative to the
costs of CA (Pesis et al., 2000; Rosa et al., 2001; Singh et al., 2001; Castro et al.,
2005; Yahia, 2006). However, MAP can reduce quality if the cultivar/holding
temperature/lm permeability/storage time combination is not optimal
(Sornsrivichai et al., 1989), resulting in anaerobic conditions and off-avours.
Excess moisture retention inside the bags can increase disease problems
(Joyce and Patterson, 1994). Special lms have been developed with higher
water vapour transmission rates (Pesis et al., 2000) or moisture absorption
materials can be included. Ethylene absorption sachets can reduce chloro-
phyll loss and red discoloration around the lenticels (Rosa et al., 2001).
MAP reduces weight loss (Singh and Janes, 2001; Bower et al., 2003),
which maintains saleable weight, but may also reduce CI. There may be a
direct relation between these, since weight loss can contribute to CI in avo-
cado, and the reduction in CI obtained in mango through the use of wax
coatings has been attributed to the same mechanism (Bower et al., 2003). Pesis
et al. (2000) considered that lenticel discoloration is a symptom of mild CI,
and noted that MAP reduced the red coloration around the lenticels in the
blushed area and the green coloration around the lenticels in the green area
of Keitt mangoes. Less lenticel spotting occurs in Kensington Pride man-
goes stored under MAP (Yuen et al., 1993). It is not clear whether the reduc-
tion in lenticel damage was due to CO
2
/O
2
or humidity modication.
Cultivar, lm type, number and mass of fruit per package, temperature,
RH, time of storage, maturity of the fruit and production conditions are
important for developing MAP systems (Brecht et al., 2003; Yahia, 2006).
Important challenges are the differential effects of temperature on fruit respi-
ration and lm permeability, resulting in differing gas concentrations around
the fruit as temperature uctuates. Success with MAP depends on a consistent
or at least predictable cold chain removal of the plastic lm before signicant
temperature uctuations are likely to occur and using MAP lms that are
unlikely to cause anaerobic conditions within the temperature range experi-
enced in the cold chain. Brecht et al. (2003) suggested an approach to designing
exible CA/MA systems to account for variations in the cold chain.
15.9 Transport
Transportation of tropical fruit and vegetables has been reviewed by McGregor
(1987) and Thompson (2002). For local markets (<3 h access), transportation
of fruit in non-refrigerated carriers is feasible, particularly if the fruit has
G.I. Johnson and P.J. Hofman 576
been precooled and transported at night with few stops. Fruit must be shel-
tered from direct sun and rain. For sea export, fruit must be cooled to the
required vessel carrying temperature (or within 2C thereof) and the cold chain
must be maintained until the fruit is displayed for purchase.
Some retailers prefer fruit that are ready to eat within 12 days of receipt. In
these situations, the ideal scenario is to ripen the product as close as possible to
the retail outlet to minimize physical damage to the soft fruit. However, where
end-market location and transport arrangements allow delivery to market
within 3 days, ripening on-farm has advantages by reducing costs for growers,
and extending ownership of the product. Transport time is a major consideration
for determining optimum systems (Ledger, 2003b) (Table 15.7).
When the export dispatch facility is >1 h away from the packhouse, the
following road transport recommendations apply:
The refrigerated truck should be clean and in good mechanical condi-
tion. The insulation and oor should be in a sound condition, the door
seals must be intact and the doors must close very tightly.
Table 15.7. Ripening and transport recommendations for Kensington Pride mango within
Australia, to cater for the ripe-for-tonight programme of major retail chains (Source: Ledger,
2003b).
System Aim Recommended handling conditions
System 1 ripen
at market
To deliver uniformly backward
fruit to the market destination
and then use ethylene to ripen
fruit ready for retail sale.
Temperature is managed
through the chain to prevent
mixed ripening and to avoid
temperatures >22C. This is
the preferred system for
Northern Territory and northern
Western Australia growers send-
ing >2000 km to market
Precool fruit to transport temperature
within 1215 h of packing
Transport at 1216C for trips of
12 days and 12C for longer trips
Ripen at the market using 10 ppm
ethylene for 23 days at 1820C
Continue to hold fruit at 1820C
until ready for sale
Store at 1012C to slow ripening
for a maximum of 3 days
System 2 ripen
on farm
To deliver fruit to the market
destination ready for retail
sale within 12 days. Fruit are
ripened evenly using ethylene
to colour stage 3 (3050%
yellow) before transport and
temperature is managed
through the chain to avoid high
temperatures >22C. Ripening
on farm is not recommended
for transport times >4 days
Precool to 1820C within 1215 h
of packing
Ripen using 10 ppm ethylene for
23 days at 1820C
Hold at 1820C until colour stage
3 (3050% yellow)
Transport at 1216C for trips of
12 days and 12C for 34 day
trips
Hold at market at 1820C until
ready for sale
Store at 1012C to slow ripening
for a maximum of 3 days
Postharvest Technology 577
The refrigeration equipment must be correctly set on air delivery and
must be calibrated for each journey. Equipment needs to function reli-
ably and receive regular servicing. Air should be delivered at the set
point and uctuations should not exceed 0.5C from set point. Refriger-
ated vehicles should be tted with temperature loggers monitoring the
delivery air, and with a digital display on the outside of the box. Refrig-
erated vehicles are not usually designed for, or capable of, lowering fruit
temperatures so the fruit must be at the relevant shipping temperature
when loading.
Because of the shorter time involved, air-transported fruit may have less
stringent temperature requirements than sea-export fruit. Airlines carry-
ing cargo may need to be consulted concerning the normal hold tem-
peratures in their aircraft.
Sea-export fruit should be held under refrigeration until loading. Sea
transport can be in refrigerated vessels, with entire refrigerated decks
lled with pallets, or in sea containers, each of which is linked to a cen-
tral ducted refrigeration system in refrigerated container vessels. Alter-
natively, integral containers with their own individual cooling systems
or integral CA containers may be used.
Close temperature monitoring on the vessels is essential. By monitoring
delivery air temperatures (DAT) and return air temperatures (RAT), it is pos-
sible to assess whether fruit is heating up due to respiration or inadequate
precooling, and to take necessary steps (Anonymous, 1989; Eksteen, 1990).
While most refrigerated container vessels monitor individual container air
temperatures, including DAT and RAT, it is sometimes advisable to include
additional temperature loggers which can measure air and fruit pulp tem-
peratures for an entire journey.
The sea-freight component is generally the most time-consuming part of
the whole eld-to-supermarket voyage (for example see Table 15.8). Essen-
tial activities before and after transport can be signicant, for a relatively
perishable product like mango. Minimizing time delays in each component
of the distribution chain is important. To reduce product deterioration,
Table 15.8. Typical packing and shipping schedules for mangoes consigned by sea
to the EU from South Africa.
Operation Days required
Picking and packing 1
Precooling and accumulation of load 4
Transport to port 2
Port handling and accumulation of load 3
Voyage time 17
Discharge handling 1
Transport and distribution 2
Total 30
G.I. Johnson and P.J. Hofman 578
producers and marketers should encourage training in perishable product
handling and QA systems for personnel from trucking, sea-freight and air-
freight companies who are responsible for loading, unloading and maintain-
ing storage facilities.
Co-shipment or storage with fruit or owers that produce high levels of
ethylene can cause unanticipated triggering of mango ripening. Co-shipment
with papaya (Carica papaya) increases mango ripening (OHare et al., 1994).
Conversely, co-shipment of carambolas (Averrhoa carambola) with mangoes
caused ripening of the carambolas. The development of specialized packaging
materials to eliminate extraneous ethylene may reduce the risk of unwanted
ripening, although mixed transport should be avoided.
15.10 Marketing
Modern supermarket chains require large quantities of uniform produce that
can be purchased on contract for delivery at a particular time to stores across
a city or country. This allows the supermarket chain to promote the product
at a special price. Mangoes are generally priced per fruit rather than by
weight, although this is changing. Barcoding and/or Price Lookup Codes
(PLU) on the labels of individual fruit for electronic checkout processing
improves monitoring of purchase habits and stock control. The International
Federation of Produce Standards (IFPS) (2008) provides a forum for stan-
dardization of produce labelling and the PLUs are applicable internationally.
Proctor and Cropley (1994) cautioned the need to ensure that label adhesives
comply with food additive restrictions in the EU.
Networks and cooperatives
Marketing cooperatives or networks can assist individual producers to obtain
critical mass in an industry, and full buyer expectations of large supply and
seasonal spread of production (Glogoski, 1995; Grifn, 1995; Higginbottom,
1995; M.C. Nguyen et al., 2004).
Promotion and consumer education
Mangoes are increasingly popular among afuent consumers in the EU,
North America and northern Asia. In the tropics, they are reminders of a
non-urban living, which has become less common because of rapid indus-
trialization and migration to the cities. Whether for domestic use or export,
mangoes must compete in the fresh market with other equally attractive,
nutritious, aromatic and tasty fruit. Mangoes must also increasingly compete
with the snack food, beverage and entertainment industries.
Consumer education can encourage consumption and sales. Customers
can be educated how to select and store mangoes and how to use both the
Postharvest Technology 579
fresh and the processed products in a variety of ways, thereby increasing
total demand. Production of mango slices in take-away packs can tap domes-
tic and export markets for ready-to-eat, healthy products and circumvent
some disinfestation requirements (see Raymundo et al., Chapter 17, this vol-
ume). Siriphanich (1994) has reviewed minimal processing of tropical fruit
and noted the advantages of gaining market access and reducing transporta-
tion costs.
15.11 Conclusions
Mango production has been based almost entirely on Mangifera indica, albeit
a variable meld of thousands of cultivars which may be derived from inter-
specic hybrids of a few closely related species (Kostermans and Bompard,
1993). Given its perishable nature, capitalizing more on the diversity of exist-
ing germplasm to develop cultivars with superior storage traits linked to
customer appeal could deliver major benets.
Future improvements in postharvest technology and quarantine treat-
ment will come from renement of preharvest management, for example
reducing disease inoculum and increasing fruit resistance to disease, reduc-
ing harvest costs and fruit damage, improving postharvest treatments and
systems, and supply chain approaches to enhance fruit longevity and quality
and reduce the risks of product damage. Improvements will also accrue from
the provision of user-friendly information for supply chain personnel, but
only if the information is utilized and implemented. Increases in throughput
via the automation of harvesting and treatment systems for fruit will increase
as production and marketing costs escalate. Labour saving and work ef-
ciency will also become more critical. Innovative transport arrangements
may become necessary as regional development places greater pressures on
transport systems. International, collaborative joint-marketing ventures will
ensure year-round supplies of uniform quality fruit, and per capita consump-
tion of mangoes will increase (Johnson, 1995).
Acknowledgements
The authors acknowledge the contributions of the co-authors of Johnson et al.
(1997), which this book chapter supercedes, and Leanne Taylor and Roberto
Marques for assistance with references. The authors also thank the Depart-
ment of Primary Industries and Fisheries for research programme support.
References
Abu-Goukh, A.B.A. and Mohamed, H.I. (2004) Effect of harvesting method on quality
and shelf life of mango fruits. Tropical Science 44, 7376.
G.I. Johnson and P.J. Hofman 580
Adkins, M., Hofman, P.J. and Stubbings, B. (2002) Manipulating the Ripening of Tropical
and Subtropical Fruits with 1-MCP (Ethylbloc
R
). Final Report for Project FR00029.
Horticulture Australia Ltd, Sydney.
Adkins, M.F., Hofman, P.J., Stubbings, B.A. and Macnish, A.J. (2005) Manipulating avo-
cado fruit ripening with 1-methylcyclopropene. Postharvest Biology and Technol-
ogy 35, 3342.
Agricultural and Processed Food Products Export Development Authority (APEDA)
(2007) Ministry of Commerce and Industry, Government of India. Available at:
http://www.apeda.com/apedawebsite/Announcements/GUIDELINES MANGOES
TO USA.pdf (accessed 11 October 2008).
Akem, C.N. (2006) Mango anthracnose disease: present status and future research pri-
orities. Plant Pathology 5, 266273.
Alves, R.E., Filgueiras, H.A.C., Almeida, A.S., Pereira, M.E.C., Cocozza, F.M. and Jorge,
J.T. (2004) Postharvest ripening of Tommy Atkins mangoes on two maturation
stages treated with 1-MCP. Acta Horticulturae 645, 627632.
Amarante, C. and Banks, N.H. (2001) Postharvest physiology and quality of coated fruits
and vegetables. Horticultural Reviews 26, 161238.
Amesbury, K., Kernot, I. and Rudge, T. (2002) Tableland Marketing Mango Grading
Guide. Available at: http://www.mangoes.net.au/images/pdf/Mango_Grade_Chart.
pdf (accessed 10 March 2008).
Anesiadis, T., Karaoglanidis, G.S. and Tzavella-Klonari, K. (2003) Protective, curative
and eradicant activity of the strobilurin fungicide azoxystrobin against Cercospora
beticola and Erysiphe betae. Journal of Phytopathology 15, 647651.
Anonymous (1983) Ethylene dibromide: intent to cancel registrations of pesticide prod-
ucts containing ethylene dibromide; determination concluding the rebuttable pre-
sumption against registration; availability of position documents. Federal Register
48, 4623446248.
Anonymous (1985) Treatments for fresh fruit and vegetables. In: Plant Quarantine Treat-
ment Schedule. Australian Quarantine and Inspection Service, Canberra, pp. 2123.
Anonymous (1986) Irradiation in the production, processing, and handling of food.
Food and Drug Administration. Federal Register 51, 1337613399.
Anonymous (1988) Conditions for importation of fruit and vegetables. In: Plant Quaran-
tine Manual, section E. Australian Quarantine Inspection Service, Canberra, pp. 159.
Anonymous (1989) Guide to Food Transport. Fruit and Vegetables. Mercantila, Copen-
hagen, pp. 162163.
Anonymous (1992) New Zealand Ministry of Agriculture and Fisheries Treatment Re-
quirements, NASS: 152.02, Border Protection Specications: Clearance of Fresh
Produce, Section 2.0, TB. Wellington, North Island, New Zealand.
Anonymous (1993) International Standardisation of Fruit and Vegetables: Mangoes. Or-
ganisation for Economic Cooperation and Development. Available at http://ingen-
taconnect.com/content/oecd/16080149/1993/00001993/00000001/5193033e
Anonymous (1994a) Plant Pest and Quarantine Treatment Manual. United States De-
partment of Agriculture (USDA), Animal and Plant Health Inspection Service,
Hyattsville, Maryland, USA.
Anonymous (1994b) Mango Farmer/Packhouse/Exporter Guidelines. South African
Mango Growers Association, Tzaneen, South Africa.
Arafat, L.A.E.T. (2005) Chilling injury in mangoes. PhD thesis, Wageningen University,
Wageningen, the Netherlands.
Arpaia, M.L. (1994) Preharvest factors inuencing postharvest quality of tropical and
subtropical fruit. HortScience 29, 982985.
Askar, A. and Treptow, H. (1993) Quality Assurance in Tropical Fruit Processing. Springer-
Verlag, Berlin.
Postharvest Technology 581
ASEAN_GAP (2007) Association of South-east Asian Nations (ASEAN) Good Agricul-
tural Practice. Standards and Manuals. Available at: http://www.aphnet.org/gap/
ASEANgap.html (accessed 31 October 2007).
Astridge, D. and Baron, Z. (2007a) Fruit spotting bug in mangoes. In: Delivering Mango
Research. Proceedings of the Sixth Australian Mango Conference. Gold Coast,
Queensland, Australia, pp. 1516.
Astridge, D. and Baron, Z. (2007b) IPM in mangoes. In: Delivering Mango Research.
Proceedings of the Sixth Australian Mango Conference. Gold Coast, Queensland,
Australia, pp. 1415.
Astridge, D. and Baron, Z. (2007c) Mango pulp weevil in mangoes of the Philippines.
(Abstract). In: Delivering Mango Research. Proceedings of the Sixth Australian
Mango Conference. Gold Coast, Queensland, Australia, p. 17.
Australian Quarantine and Inspection Service (AQIS) (2008a) AQIS Import Conditions
Database. Search mango. Available at: http://www.aqis.gov.au/icon32/asp/ex_
querycontent.asp (accessed 3 April 2008).
Australian Quarantine and Inspection Service (AQIS) (2008b) Phyto search. Available at:
http://www.aqis.gov.au/phyto/asp/ex_search.asp (accessed 7 April 2008).
Bagshaw, B. and Brown, B. (1989) Disorders. In: Ridgeway, R. (ed.) Mango Pests and
Disorders. Queensland Department of Primary Industries, Brisbane, Queensland,
Australia, pp. 2228.
Baker, I.W. (1986) Mango maturity investigations. In: Proceedings of the First Australian
Mango Research Workshop. Commonwealth Scientic and Industrial Research Or-
ganization (CSIRO), Melbourne, pp. 271273.
Baldwin, E.A., Burns, J.K., Kazokas, W., Brecht, J.K., Hagenmaier, R.D., Bender, R.J. and
Pesis, E. (1999) Effect of two edible coatings with different permeability character-
istics on mango (Mangifera indica L.) ripening during storage. Postharvest Biology
and Technology 17, 215226.
Bally, I.S.E. (2007) The effect of preharvest nutrition and crop load on fruit quality and
postharvest disease in mango (Mangifera indica L.). PhD thesis, University of
Queensland, Brisbane, Queensland, Australia.
Bally, I.S.E., OHare, T.J. and Holmes, R.J. (1997) Detrimental effects of detergent in the
development of mango skin browning. Acta Horticulturae 455, 612621.
Bandyopadhyay, C., Gholap, A.S. and Mamdapur, V.R. (1985) Characterisation of alke-
nyl resorcinol in mango (Mangifera indica L.) latex. Journal of Agricultural and
Food Chemistry 33, 377379.
Biosecurity New Zealand (BANZ) (2008) Import Health Standards. Available at: http://
www.biosecurity.govt.nz/commercial-imports/import-health-standards/search (ac-
cessed 7 April 2008).
Barba, R.C. (1974) Induction of owering of the mango by chemical spray. In: Proceed-
ings of the Fifth Crop Science Society of the Philippines Meeting. pp. 154160.
Bartley, J.P. and Schwede, A. (1987) Volatile avour components in the headspace of the
Australian Bowen mango. Journal of Food Science 52, 353360.
Bender, R.J., Brecht, J.K., Sargent, S.A. and Huber, D.J. (2000) Mango tolerance to re-
duced oxygen levels in controlled atmosphere storage. Journal of the American
Society for Horticultural Science 125, 707713.
Bezuidenhout, J.L.J., Robbertse, P.J. and Kaiser, C. (2005) Anatomical investigation of lenticel
development and subsequent discolouration of Tommy Atkins and Keitt mango
(Mangifera indica L.) fruit. Journal of Horticultural Science and Biotechnology 80, 1822.
Blankenship, S.M. and Dole, J.M. (2003) 1-Methylcyclopropene: a review. Postharvest
Biology and Technology 28, 125.
Boelema, T. (1987) Long-distance transport of avocados. South African Avocado Grow-
ers Association Yearbook 10, 153156.
G.I. Johnson and P.J. Hofman 582
Bondad, N.D. and Linsagan, E. (1979) Flowering in mango induced with potassium ni-
trate. HortScience 14, 527528.
Bower, J.P., Dennison, M.T. and Fowler, K. (2003) Avocado and mango cold storage
damage as related to water loss control. Acta Horticulturae 628, 401406.
Brady, C. (1994) Biochemical and molecular approaches to fruit ripening and senes-
cence. In: Champ, B.R., Highley, E. and Johnson, G.I. (eds) Postharvest Handling of
Tropical Fruits. Australian Centre for International Agricultural Research (ACIAR)
Proceedings 50. ACIAR, Canberra, pp. 205217.
Brecht, J.K., Chau, K.V., Fonseca, S.C., Oliveira, F.A.R., Silva, F.M., Nunes, M.C.N. and
Bender, R.J. (2003) Maintaining optimal atmosphere conditions for fruits and veg-
etables throughout the postharvest handling chain. Postharvest Biology and Tech-
nology 27, 87101.
Brown, B.I., Wells, I.A. and Murray, C.F. (1986) Factors affecting the incidence and se-
verity of mango sapburn and its control. ASEAN Food Joumal 2, 127132.
Buchanan, A. (1994) Tropical fruits: the social political and economic issues. In: Champ,
B.R., Highley, E. and Johnson, G.I. (eds) Postharvest Handling of Tropical Fruits.
Australian Centre for International Agricultural Research (ACIAR) Proceedings 50.
ACIAR, Canberra, pp. 1826.
Bunt, C. and Piccone, M. (1994) Quality assurance: a total approach. In: Champ, B.R.,
Highley, E. and Johnson, G.I. (eds) Postharvest Handling of Tropical Fruits. Austra-
lian Centre for International Agricultural Research (ACIAR) Proceedings 50. ACIAR,
Canberra, pp. 5664.
Burdon, J.N. and Moore, K.G. (1991) The soft-nose disorder of mango fruit. Planter 67,
477481.
Bustos, M.E., Enkerlin, H.W., Toledo, A.J., Reyes, F.J. and Casimiro, G.A. (1992) Irradia-
tion of mangoes as a quarantine treatment. In: Use of Irradiation as a Quarantine
Treatment of Food and Agricultural Commodities. Proceedings of the Final Research
Coordination Meeting, International Atomic Energy Agency (IAEA), Vienna,
Austria.
Bustos, M.E., Enkerlin, W., Reyes, J. and Toledo, J. (2004) Irradiation of mangoes as a
postharvest quarantine treatment for fruit ies (Diptera: Tephritidae). Journal of Eco-
nomic Entomology 97, 286292.
Castro, J.V., Pfaffenbach, L.B., Carvalho, C.R.L. and Rossetto, C.J. (2005) Effects of lm
packaging and cold storage on postharvest quality of Tommy Atkins mangoes.
Acta Horticulturae 682, 16831687.
Catindig, J.L.A. and Kong, L.H. (2007) Asia Pacic Alien Species Database. Mango Pulp
Weevil. Available at: http://apasd-niaes.dc.affrc.go.jp/ (accessed 31 October
2007).
Chalutz, E., Droby, S., Wilson, C.L. and Wisniewski, M.E. (1992) DV-induced resistance
to postharvest diseases of citrus fruit. Journal of Phytochemistry and Photobiology
15, 367374.
Chaplin, G.R., Nuevo, P.A., Graham, D. and Cole, S.P. (1986a) Chilling responses of
Kensington mango fruit stored under variable low temperature regimes. ASEAN
Food Journal 2, 133138.
Chaplin, G.R., Graham, D. and Cole, S.P. (1986b) Reduction of chilling injury in mango
fruit by storage in polyethylene bags. ASEAN Food Journal 2, 139142.
Chaplin, G.R., Cole, S.P., Landragan, M., Nuevo, P.A., Lam, P.F. and Graham, D. (1991a)
Chilling injury and storage of mango (Mangifera indica L.) fruit under low tempera-
tures. Acta Horticulturae 291, 461471.
Chaplin, G.R., McBride, R.L., Abdullah, A. and Nuevo, P.A. (1991b) Sensory and physi-
cochemical quality of Kensington mangoes after storage at low temperature. ASE-
AN Food Journal 6, 109113.
Postharvest Technology 583
Chin, D., Brown, H. and Thistleton, B. (2007) Mango pulp weevil in mangoes of the
Philippines. In: Delivering Mango Research. Proceedings of the Sixth Australian
Mango Conference. Gold Coast, Queensland, Australia, pp. 1823.
Chitarra, A.B., Chitarra, M.I.F. and Evangelista, R.M. (2001) Biochemical changes in
mango fruits, Tommy Atkins treated with calcium chloride preharvest and stored
under refrigeration. Acta Horticulturae 553, 7981.
Cipollini, M.L. and Stiles, E.W. (1992) Relative risks of microbial rot for eshy fruits:
signicance with respect to dispersal and selection for secondary defence. Advanc-
es in Ecological Research 23, 3591.
Coates, L.M. and Johnson, G.I. (1993) Effective disease control in heat-disinfested fruit.
Postharvest News and Information 4, 35N40N.
Coates, L.M., Johnson, G.I. and Cooke, A.W. (1993) Postharvest disease control in man-
goes using high humidity hot air and fungicide treatments. Annals of Applied Biol-
ogy 123, 441448.
Coates, L.M., Davis, R.D. and Cooke, A.W. (1995) Postharvest disease control in mango
potential options for reducing chemical use. In: Proceedings of Mango 2000 Pro-
duction Workshop, Townsville.
CODEX (2008a) CODEX Alimentarius Commission. Available at: http://www.codexali-
mentarius.net/web/index_en.jsp (accessed 21 March 2008).
CODEX (2008b) Codex Standards List, 2008. CODEX Alimentarius Current Ofcial
Standards. Available at: http://www.codexalimentarius.net/web/standard_list.
do?lang=en (accessed 21 March 2008).
CODEX-MRL (2008) Limits for Prochloraz Assorted Tropical and Sub-tropical Fruits
Inedible Peel. Database Listings of Maximum Residue Limits; Extraneous Maximum
Residue Limits. Ofcial Standards Pesticide Residues in Food Searchable Data-
base. World Health Organization (WHO)/Food and Agriculture Organization (FAO)
Food Standards CODEX Alimentarius. Available at: http://www.codexalimentarius.
net/mrls/pestdes/jsp/pest_q-e.jsp (accessed 4 April 2008).
Coetzer, L.A., Robbertse, P.J. and De Wet, E. (1991) Die invloed van boortoedienings op
vrugproduksie en verkoelde opberging van mangos. South African Mango Growers
Association Yearbook 11, 2931.
Cojocaru, M., Droby, S., Glotter, E., Goldman, H., Gottlieb, H., Jacoby, B. and Prusky,
D. (1985) 5-(12-heptadecerlyl)-resorcinol, the major component of the antifungal
agent from the peel of mango fruit. Phytochemistry 25, 10931095.
Cond, B.D. and Pitkethly, R.N. (2007) Mango scab in Australia. In: Delivering Mango
Research. Proceedings of the Sixth Australian Mango Conference. Gold Coast,
Queensland, Australia, pp. 810.
Cooke, A.W. and Johnson, G.I. (1994) Mango postharvest disease control: effect of rain
at harvest, fungicide treatments and fruit brushing on fruit appearance. In: Champ,
B.R., Highley, E. and Johnson, G.I. (eds) Postharvest Handling of Tropical Fruits.
Australian Centre for International Agricultural Research (ACIAR) Proceedings 50.
ACIAR, Canberra, pp. 448449.
Crous, P.W., Slippers, B., Wingeld, M.J., Rheeder, J., Marasas, W.F.O., Philips, A.J.L.,
Alves, A., Burgess, T., Barber, P. and Groenewald, J.Z. (2006) Phylogenetic lineages
in the Botryosphaeriaceae. Studies in Mycology 55, 235253.
Cunningham, I.C. (1986) Mango insect pests. In: Proceedings of the First Australian
Mango Research Workshop. Commonwealth Scientic and Industrial Research
Organization (CSIRO), Melbourne, pp. 221224.
Cunningham, I.C. (1991a) Management of mango insect pests. Acta Horticulturae 291,
379388.
Cunningham, I.C. (1991b) Common mango scales in Queensland. Acta Horticulturae
291, 409412.
G.I. Johnson and P.J. Hofman 584
Cunningham, I.C. (1991c) Mango seed weevil in Queensland. Acta Horticulturae 291,
413417.
Dann, E., Zainuri, Hassan, K., Irving, D.E., Cooke, A.W. and Coates, L. (2005) Self-defence
for mangoes. Mango Matters Autumn/Winter issue, 3536.
Dann, E., Coates, L., Cooke, A.W., Smith, L. and Dean, J. (2007) Novel methods for
integrated control of mango postharvest diseases. In: Delivering Mango Research.
Proceedings of the Sixth Australian Mango Conference. Gold Coast, Queensland,
Australia, pp. 57.
Davenport, T.L. and Nez-Elisea, R. (1997) Reproductive physiology. In: Litz, R.E. (ed.) The
Mango: Botany, Production and Uses. CAB International, Wallingford, UK, pp. 69146.
DeEll, J.R. and Toivonen, P.M.A. (eds) (2003a) Practical Applications of Chlorophyll Fluo-
rescence in Plant Biology. Kluwer Academic Publishers, Dordrecht, the Nether-
lands, 280 pp.
DeEll, J.R. and Toivonen, P.M.A. (2003b) Use of chlorophyll uorescence in postharvest
quality assessments of fruits and vegetables. In: DeEll, J.R. and Toivonen, P.M.A.
(eds) Practical Applications of Chlorophyll Fluorescence in Plant Biology. Kluwer
Academic Publishers, Dordrecht, the Netherlands, pp. 203242.
de Jesus, L.R.A., Calumpang, S.M.F., Medina, J.R. and Ohsawa, K. (2007) Floral volatiles
of Mangifera indica L. (cv. Carabao) attractive to Sternochetus frigidus (Fabr.)
(Coleoptera: Curculionidae). The Philippine Agricultural Scientist 86, 282289.
DeLong, J.M., Prange, R.K. and Harrison, P.A. (2007) Chlorophyll uorescence-
based low-O
2
CA storage of organic Cortland and Delicious apples. Acta Horti-
culturae 737, 3137.
Diedhiou, P.M., Mbaye, N., Drame, A. and Samb, P.I. (2007) Alteration of postharvest
diseases of mango (Mangifera indica) through production practices and climatic
factors. African Journal of Biotechnology 6, 10871094.
Dodd, J.C., Estrada, A.B., Matcham, J., Jeffries, P. and Jeger, M.J. (1991a) The effect of
climatic factors on Colletotrichum gloeosporioides, the causal agent of mango an-
thracnose, in the Philippines. Plant Pathology 40, 568575.
Dodd, J.C., Bugante, R., Koomen, I., Jefries, P. and Jeger, M.J. (1991b) Pre- and post-harvest
control of mango anthracnose in the Philippines. Plant Pathology 40, 576583.
Dodd, J.C., Estrada, A. and Jeger, M.J. (1992) Epidemiology of Colletotrichum gloeospo-
rioides in the tropics. In: Bailey, J.A. and Jeger, M.J. (eds) Colletotrichum: Biology,
Pathology and Control. CAB International, Wallingford, UK, pp. 308325.
Dodd, J.C., Prusky, D. and Jeffries, P. (1997) Fruit diseases. In: Litz, R.E. (ed.) The Mango:
Botany, Production and Uses. CAB International, Wallingford, UK, pp. 257280.
Droby, S., Prusky, D., Jacoby, B. and Goldman, A. (1986) The presence of an antifungal
compound and its relation to the latency of Alternaria alternata in unripe peels of
mango fruits. Physiological Plant Pathology 29, 173183.
Droby, S., Prusky, D., Jacoby, B. and Goldman, A. (1987) Induction of antifungal resor-
cinols in esh of unripe mango fruits and its relation to latent infection by Alter-
naria alternata. Physiological and Molecular Plant Pathology 30, 285292.
Ducamp Collin, M.-N., Arnaud, C., Kagy, V. and Didier, C. (2007) Fruit ies: disinfesta-
tion, techniques used, possible application to mango. Fruits 62, 223236.
du Plooy, G.W., van der Merwe, C. and Korsten, L. (2004) Differences in the surface
structures of three mango cultivars and the effect of kaolin on these structures.
South African Mango Growers Association Yearbook 24, 2936.
du Plooy, G.W., van der Merwe, C.F. and Korsten, L. (2006) Lenticel discolouration in
mango (Mangifera indica L.) fruit a cytological study of mesophyll cells from af-
fected tissue. Journal of Horticultural Science and Biotechnology 81, 869873.
Eckert, J.W. (1983) Control of postharvest diseases with antimicrobial agents. In: Diseases
of Fruits and Vegetables. North Atlantic Treaty Organization (NATO) Advisory
Postharvest Technology 585
Study Institute Series, Series A Life Sciences. Available at http://www.fao.org/
agris/search/display.do?f=./1984/v1002/US8268297.xml; US8268297, New York,
pp. 265285.
Eksteen, G.J. (1990) Temperature changes during sea shipment of mangoes in ducted
containers. South African Mango Growers Association Yearbook 10, 2627.
El Ghaouth, A. and Wilson, C.L. (1995) Biologically based technologies for the control
of postharvest diseases. Postharvest News Information 6, 5N11N.
El Ghaouth, A., Arul, J., Asselin, A. and Benhamou, N. (1992a) Antifungal activity of
chitosan on postharvest pathogens: induction of morphological and cytological
alterations in Rhizopus stolonifer. Mycological Research 96, 769779.
El Ghaouth, A.E., Ponnampalam, R., Castaigne, F. and Arul, J. (1992b) Chitosan coating
to extend the storage life of tomatoes. HortScience 27, 10161018.
Environmental Protection Agency (EPA) (2002) Irradiation Phytosanitary Treatment of
Imported Fruits and Vegetables Federal Register (Volume 67, Number 205). Rules
and Regulations: 6501665029. Available at: http://www.wais.access.gpo.gov
(accessed 23 October 2008).
Environmental Protection Agency (EPA) (2008) The Phaseout of Methyl Bromide. Avail-
able at: http://www.epa.gov/ozone/mbr/ (accessed 3 April 2008).
Erwin, D.C. (1973) Systemic fungicides: disease control, translocation and mode of
action. Annual Review of Phytopathology 11, 389400.
Esguera, E.B. and Lizada, M.C.C. (1990) The postharvest behaviour and quality of Cara-
bao mangoes subjected to vapour heat treatment. ASEAN Food Journal 5, 612.
Esguerra, E.B., Brena, S.R., Reyes, M.U. and Lizada, M.C.C. (1990) Physiological break-
down in vapor heat-treated Carabao mango. Acta Horticulturae 269, 425434.
Estrada, A.B., Dodd, J.C. and Jeffries, P. (1993) Effect of environment on the in vitro
growth and development of Colletotrichum gloeosporioides isolates from the Phil-
ippines. Acta Horticulturae 341, 360370.
EurepGAP (2007) What is EurepGAP? Available at: http://www.eurepgap.org/Languages/
English/about.html (accessed 31 October 2007).
Eustice, R.F. (2007) Recent Developments in Food Irradiation: a Global Review. Min-
nesota Beef Council. Available at: http://www.mnbeef.org/New%20Site/HTML/irra
diationSub/2RecentDevelopments.htm (accessed 31 October 2007).
Everett, K.R., Owen, S.G. and Cutting, J.G.M. (2005) Testing efcacy of fungicides
against postharvest pathogens of avocado (Persea Americana cv. Hass). New Zea-
land Journal of Plant Protection 58, 8995.
Fallik, E. and Grinberg, S. (1992) Hinokitiol: a natural substance that controls postharvest dis-
eases in eggplant and pepper fruits. Postharvest Biology and Technology 2, 137144.
FAOSTAT (2008) FAOSTAT On-line. Rome. United Nations Food and Agriculture Orga-
nization. Available at: http://faostat.fao.org/default.aspx?lang=en (accessed 21
March 2008).
Feygenberg, O., Hershkovitz, V., Ben-Arie, R., Jacob, S., Pesis, E. and Nikitenko, T. (2005)
Postharvest use of organic coating for maintaining bio-organic avocado and mango
quality. Acta Horticulturae 682, 10571061.
FFTC-GAP (2007) Good Agricultural Practice in Asia and Oceana. Available at http://agnet.org/
library/ubk/54/
Fitzell, R.D. and Peak, C.M. (1984) The epidemiology of anthracnose disease in mango:
inoculum sources, spore production and dispersal. Annals of Applied Biology 104,
5359.
Fitzell, R.D., Peak, C.M. and Darnell, R.E. (1984) A model for estimating infection levels
of anthracnose disease of mango. Annals of Applied Biology 104, 451458.
Follett, P.A. (2004) Irradiation quarantine treatments for mango seed weevil and Cryp-
tophlebia spp. In: Irradiation as a Phytosanitary Treatment of Food and Agricultural
G.I. Johnson and P.J. Hofman 586
Commodities. IAEA-TECDOC-1427. International Atomic Energy Agency (IAEA),
Vienna, Austria, pp. 918.
Follett, P.A. and Gabbard, Z. (2000) Effect of mango weevil (Coleoptera: Curculionidae)
damage on mango seed viability in Hawaii. Journal of Economic Entomology 93,
12371240.
Follett, P.A. and Lower, R.A. (2000) Irradiation to ensure quarantine security for Cryp-
tophlebia spp. (Lepidoptera: Tortricidae) in Sapindacious fruits from Hawaii. Jour-
nal of Economic Entomology 93, 18481854.
Follett, P.A. and Neven, L.G. (2005) Current trends in quarantine entomology. Annual
Review of Entomology 51, 359385.
Follett, P.A., Yang, M., Lu, K. and Chen, T. (2007) Irradiation for postharvest control of
quarantine pests. Formosan Entomology 27, 115.
Fonseca, M.J.O., Cecon, P.R., Salomo, L.C.C. and Pushmann, R. (2004a) Pulp and skin
pigments in mango Haden treated with fungicides and wax. Acta Horticulturae
645, 633638.
Fonseca, M.J.O., Salomo, L.C.C., Cecon, P.R. and Puschmann, R. (2004b) Fungicides
and wax in postharvest preservation of mango Haden. Acta Horticulturae 645,
557563.
Food and Fertilizer Technology Centre (FFTC) (2007) Good Agricultural Practice in Asia
and Oceana. Available at: http://www.agnet.org/library/vbk/54/ (accessed 7 Octo-
ber 2007).
Gaffney, J.J. and Armstrong, J.W. (1990) High-temperature forced-air research facility for
heating fruits for insect quarantine treatments. Journal of Economic Entomology 83,
19591964.
Good Agricultural Practice (GAP) Small Producers in Export Horticulture: a Guide to
Best Practice. Natural Resources Institute. Available at htto://nri.org/NRET/
GLOBALGAP (2007) Global GAP. Available at: htttp://www.globalgap.org/cms/front_
content.php?idart=3&idcat=9&lang=1 (accessed 31 October 2007).
Glogoski, A. (1995) Group marketing redressing the balance of power. In: Smarter
Marketing, Better Prots. Proceedings of the Second Annual Queensland Fruit and
Vegetable Growers Conference, Brisbane, Queensland, pp. 16.
Goguey, T. (1993) Study of the effects of three ower-inducing substances on Ken and
Zill mango. Acta Horticulturae 341, 216224.
Gonzlez-Aguilar, G.A., Fortiz, J., Cruz, R., Baez, R. and Wang, C.Y. (2000) Methyl jas-
monate reduces chilling injury and maintains postharvest quality of mango fruit.
Journal of Agricultural and Food Chemistry 48, 515519.
Good Agricultural Practice (GAP) (2003) Small Producers in Export Horticulture. A
Guide to Best Practice. Agricultural and Environmental Practices: Good Agricul-
tural Practice. Natural Resources Institute. Available at: http://www.nri.org/NRET/
SPCDR/Chapter4/agriculture-4-4.htm (accessed 7 October 2007).
Gould, W.P. and Sharp, J.L. (1990) Caribbean fruit y (Diptera: Tephritidae) mortality
induced by shrink-wrapping infested mangoes. Journal of Economic Entomology
83, 23242326.
Grifn, D. (1995) Grower networking to achieve international competitiveness. In:
Smarter Marketing, Better Prots. Proceedings of the Second Annual Queensland
Fruit and Vegetable Growers Conference, Brisbane, Queensland, pp. 716.
Grov, T., Steyn, W.P. and de Beer, M.S. (1997) Determining suitable non-damaging
treatments for post-harvest disinfestation of fruit y in mango. South African Mango
Growers Association Yearbook 17, 119125.
GPOAccess (2008) Code of Federal Regulations. Available at: http://gpoaccess.gov/cfr/
index.html (accessed 10 March 2008).
Postharvest Technology 587
Gullino, M.L. and Kuijpers, L.A. (1994) Social and political implications of managing
plant diseases with restricted fungicides in Europe. Annual Review of Phytopathol-
ogy 32, 559579.
Hall, A., Bockett, G., Taylor, S., Sivamohan, M.K.V. and Clark, N. (2001) Why research
partnerships really matter: innovation theory, institutional arrangements and im-
plications for developing new technology for the poor. World Development 29,
783797.
Han, J., Tian, S., Meng, X. and Ding, Z. (2006) Response of physiologic metabolism and
cell structures in mango fruit to exogenous methyl salicylate under low-temperature
stress. Physiologia Plantarum 128, 125133.
Hansen, J.D. (1991) Mango weevil biology, eld control and postharvest technology: a
review. International Journal of Tropical Agriculture 9, 234244.
Hansen, J.D. (1993) Dynamics and control of the mango seed weevil. Acta Horticultu-
rae 341, 415420.
Hardy, S., Fallow, K. and Barkley, P. (2004) Using Copper Sprays to Control Diseases in
Citrus. Citrus Fact Sheet, New South Wales Department of Primary Industries. Avail-
able at: http://www.dpi.nsw.gov.au/__data/assets/pdf_le/0003/138171/copper-
spray-citrus.pdf (accessed 3 January 2008).
HarvestWatch (2008) HarvestWatch Monitoring Produce Health. Availablet at:
http://www.intbox.com/technology.asp?page=2582 (accessed 28 February 2008).
Harvey, E.C. (1987) Maturity indices for quality control and harvest maturity. In: Prins-
ley, R.T. and Tucker, G. (eds) Mangoes a Review. The Commonwealth Secretariat,
London, pp. 3955.
Hassan, M.K. (2007) Constitutive alk(en)ylresorcinols and resistance to postharvest dis-
ease in mango (Mangifera indica L.). PhD thesis, University of Queensland, Bris-
bane, Queensland.
Hassan, M.K., Dann, E.K., Irving, D.E. and Coates, L.M. (2007) Concentrations of constitu-
tive alk(en)ylresorcinols in peel of commercial mango varieties and resistance to post-
harvest anthracnose. Physiological and Molecular Plant Pathology 17, 158165.
Heather, N.W. (1994) Quarantine disinfestation of tropical fruits: non-chemical options.
In: Champ, B.R., Highley, E. and Johnson, G.I. (eds) Postharvest Handling of Tropi-
cal Fruits. Australian Centre for International Agricultural Research (ACIAR) Pro-
ceedings 50. ACIAR, Canberra, pp. 272279.
Heather, N. (2004) Generalized quarantine disinfestation research protocol. In: Irradiation
as a Phytosanitary Treatment of Food and Agricultural Commodities. IAEA-TECDOC-
1427. International Atomic Energy Agency (IAEA), Vienna, Austria, pp. 171178.
Heather, N.W., Corcoran, R.J. and Banos, C. (1991) Disinfestation of mangoes with
gamma irradiation against two Australian fruit ies (Diptera: Tephritidae). Journal of
Economic Entomology 84, 13041307.
Hegnauer, R. (1994) Phytochemistry and chemotaxonomy of the Anacardiaceae with
special emphasis on Mangifera. In: Kostermans, A.J.G.H. and Bompard, J.M. (1993)
The Mangoes, their Botany, Nomenclature, Horticulture and Utilisation. Academic
Press, London, pp. 1116.
Hewett, E. (2006) An overview of preharvest factors inuencing postharvest quality of
horticultural products. International Journal of Postharvest Technology and Innovation
1, 415.
Higginbottom, D. (1995) Group marketing the future. In: Smarter Marketing, Better
Prots. Proceedings of the Second Annual Queensland Fruit and Vegetable Grow-
ers Conference, Brisbane, Queensland, pp. 1722.
Hilton, D.J. (1994) Impact and vibration damage to fruit during handling and transporta-
tion. In: Champ, B.R., Highley, E. and Johnson, G.I. (eds) Postharvest Handling of
G.I. Johnson and P.J. Hofman 588
Tropical Fruits. Australian Centre for International Agricultural Research (ACIAR)
Proceedings 50. ACIAR, Canberra, pp. 116126.
Hoa, T.T. and Ducamp, M.-N. (2008) Effects of different coatings on biochemical changes of
Cat Hoa Loc mangoes in storage. Postharvest Biology and Technology 48, 150152.
Hofman, P.J. (1997) What causes green, ripe mangoes. Mango Care 20, 1315.
Hofman, P.J. (1998) Production factors inuence fruit quality and response to postharvest
treatments. In: Coates, L.M., Hofman, P.J. and Johnson, G.I. (eds) Disease Control
and Storage Life Extension in Fruit. Australian Centre for International Agricultural
Research (ACIAR), Canberra, pp. 618.
Hofman, P.J. and Ledger, S.N. (2006) Using a supply chain approach to guide R&D. Acta
Horticulturae 699, 219226.
Hofman, P.J. and Mullen, C. (2005) The Role of Rootstocks and Nutrition in the Quality
of Hass Avocado. Final Report of Project AV00013. Horticulture Australia Ltd,
Sydney.
Hofman, P.J. and Smith, L.G. (1994) Preharvest effects on postharvest quality of sub-
tropical and tropical fruit. In: Champ, B.R., Highley, E. and Johnson, G.I. (eds)
Postharvest Handling of Tropical Fruits. Australian Centre for International Agricul-
tural Research (ACIAR) Proceedings 50. ACIAR, Canberra, pp. 261268.
Hofman, P.J., Smith, L.G. and Meiburg, G.F. (1995) Effect of position on the tree and
time of owering on fruit quality in mango. In: Australasian Postharvest Horticul-
ture Conference: Science and Technology for the Fresh Food Revolution, Mel-
bourne p. 38.
Hofman, P.J., Smith, L.G., Joyce, D.C., Johnson, G.I. and Meiburg, G.F. (1997a) Bagging
of mango (Mangifera indica cv. Keitt) fruit inuences fruit quality and mineral com-
position. Postharvest Biology and Technology 12, 8391.
Hofman, P.J., Smith, L.G., Meiburg, G.F. and Giles, J.E. (1997b) Production locality affects
mango fruit quality. Australian Journal of Experimental Agriculture 37, 801808.
Hofman, P.J., Smith, L.G., Meiburg, G.F., Holmes, R. and Barker, J.A. (1998) Mango fruit
quality is affected by production conditions. In: Coates, L.M., Hofman, P.J. and
Johnson, G.I. (eds) Disease Control and Storage Life Extension in Fruit. Australian
Centre for International Agricultural Research (ACIAR) Proceedings 81. ACIAR,
Canberra, pp. 3139.
Hofman, P.J., Jobin-Dcor, M., Meiburg, G.F., Macnish, A.J. and Joyce, D.C. (2001) Rip-
ening and quality responses of avocado, custard apple, mango and papaya fruit to
1-methylcyclopropene. Australian Journal of Experimental Agriculture 41, 567572.
Holmes, R. (2003) Checklist for sapburn. Mango Care 39, 1011.
Holmes, R. and Bally, I. (1994) Harvest aid update. Mango Care 11, 1011.
Holmes, R.J., Ledger, S.N. and Macleod, W.N.B. (1993) Handling systems to reduce
sapburn. Acta Horticulturae 341, 528532.
Horn, F. and Horn, P. (2004) Fresh fruit fumigation with phosphine as alternative for
methyl bromide. Paper 58. In: Proceedings 2004 of the Annual International Re-
search Conference on Methyl Bromide Alternatives and Emissions Reductions.
Available at: http://mbao.org/2004/Proceedings04/mbrpro04.html (accessed 1 May
2008).
International Federation for Produce Standards (IFPS) (2008) Available at: http://www.
plucodes.com/ (accessed 7 April 2008).
International Plant Protection Convention (IPPC) (2008) International Standards for Phy-
tosanitary Measures (ISPMs). International Phytosanitary Portal. Available at: https://
www.ippc.int/IPP/En/default.jsp (accessed 2 April 2008).
International Standardisation Organisation (ISO) (2008) Available at: http://www.iso.org/
iso/home.htm (accessed 2 April 2008).
Postharvest Technology 589
International Standards for Phytosanitary Measures (ISPM) (2003) Guidelines for the use
of irradiation as a phytosanitary measure. In: International Standards for Phytosani-
tary Measures Secretariat of the International Plant Protection Convention, 2006.
ISPM No. 18. Secretariat of the International Plant Protection Convention, Food
and Agriculture Organization of the United Nations, Rome, Italy, pp. 209220.
International Standards for Phytosanitary Measures (ISPM) (2007a) International Stan-
dards for Phytosanitary Measures 1 to 29. Available at: https://www.ippc.int
(accessed 31 October 2007).
International Standards for Phytosanitary Measures (ISPM) (2007b) Phytosanitary Treat-
ments for Regulated Pests. ISPM No. 28. Available at: https://www.ippc.int
(accessed 31 October 2007).
International Standards for Phytosanitary Measures (ISPM) (2008) Irradiation Treatment
for Anastrepha ludens; A. serpentina and Others. Annex to ISPM No. 28. Available
at: https://www.ippc.int (accessed 3 April 2008).
Jacobi, K.K. and Giles, J.E. (1997) Quality of Kensington mango (Mangifera indica
Linn.) fruit following combined vapour heat disinfestation and hot water disease
control treatments. Postharvest Biology and Technology 12, 285292.
Jacobi, K.K. and Gowanlock, D. (1995) Ultrastructural studies of Kensington mango
(Mangifera indica Linn.) heat injuries. HortScience 30, 102103.
Jacobi, K.K. and Wong, I.S. (1992) Quality of Kensington mango (Mangifera indica
Linn.) following hot water and vapour heat treatments. Postharvest Biology and
Technology 1, 349359.
Jacobi, K.K., Coates, L.M. and Wong, L.S. (1994) Heat disinfestation of mangoes: effect
on fruit quality and disease control. In: Champ, B.R., Highley, E. and Johnson, G.I.
(eds) Postharvest Handling of Tropical Fruits. Australian Centre for International
Agricultural Research (ACIAR) Proceedings 50. ACIAR, Canberra, pp. 280287.
Jacobi, K.K., Wong, L.S. and Giles, J.E. (1995) Effect of fruit maturity on quality and
physiology of high-humidity hot air-treated Kensington mango (Mangifera indica
Linn.). Postharvest Biology and Technology 5, 149159.
Jacobi, K.K., MacRae, E.A. and Hetherington, S.E. (2001a) Loss of heat tolerance in
Kensington mango fruit following heat treatments. Postharvest Biology and Tech-
nology 21, 321330.
Jacobi, K.K., MacRae, E.A. and Hetherington, S.E. (2001b) Postharvest heat disinfesta-
tion treatments of mango fruit. Scientia Horticulturae 89, 171193.
Jeffries, P. and Koomen, I. (1992) Strategies and prospects for biological control of dis-
eases caused by Colletotrichum. In: Bailey, J.A. and Jeger, M.J. (eds) Colletotri-
chum: Biology, Pathology and Control. CAB International, WalIingford, UK,
pp. 337357.
Jessup, A.J., Dominiak, B., Woods, B., de Lima, C.P.F., Tomkins, A. and Smallridge, C.J.
(2007) Area-wide management of fruit ies in Australia. In: Vreysen, M.J.B., Robin-
son, A.S. and Hendrichs, A. (eds) Area-wide Control of Insect Pests from Research
to Field Implementation. Springer, Dordrecht, the Netherlands, pp. 685697.
Jiang, Y. and Joyce, D.C. (2000) Effects of 1-methylcyclopropene alone and in combina-
tion with polyethylene bags on the postharvest life of mango fruit. Annals of
Applied Biology 137, 321327.
Joel, D.M. (1978) The secretory ducts of mango fruits: a defence system effective against
the Mediterranean fruit y. Israel Journal of Botany 1, 4445.
Joel, D.M. (1980) Resin ducts in the mango fruit: a defence system. Journal of Experi-
mental Botany 31, 17071718.
John, K.S., Bhat, S.G. and Rao, U.J.S.P. (2002) Involvement of peroxidase and polyphe-
nol oxidase in mango sap-injury. Journal of Food Biochemistry 26, 403414.
G.I. Johnson and P.J. Hofman 590
Johnson, G.I. (1994) Mango, stem end rot, Part III. In: Ploetz, R.C., Zentmyer, G.A.,
Nishijima, W.T., Rohrbach, K.G. and Ohr, H.D. (eds) Compendium of Tropical Fruit
Diseases. American Phytopathological Society (APS) Press, St Paul, Minnesota,
pp. 3941.
Johnson, G.I. (1995) Global outlook. In: Mango 2000 Marketing Seminar Handbook.
Department of Primary Industry Brisbane, Queensland, pp. 2130.
Johnson, G.I. (1997) Mango disease losses: balancing the economy and ecology. Acta
Horticulturae 455, 575586.
Johnson, G.I. and Coates, L.M. (1991) Quality assurance in tropical fruit future disease
control prospects. In: Sustainable Management of Pests, Diseases and Weeds, Aus-
tralian Horticulture Clean and Green in the 90s, Proceedings of the First National
Conference, Australian Society of Horticultural Science. Macquarie University, Syd-
ney, pp. 7176.
Johnson, G.I. and Coates, L.M. (1993) Postharvest diseases of mango. Postharvest News
and Information 4, 27N34N.
Johnson, G.I. and Heather, N.W. (1995) Disease and pest control in tropical fruit. In:
Champ, B.R. and Highley, E. (eds) Postharvest Technology for Agricultural Products
in Vietnam. Australian Centre for International Agricultural Research (ACIAR) Pro-
ceedings 60. ACIAR, Canberra, pp. 100126.
Johnson, G.I. and Hofman, P.J. (eds) (2004) Agri-product Supply-Chain Management in
Developing Countries. Australian Centre for International Agricultural Research
(ACIAR) Proceedings 119. ACIAR, Canberra.
Johnson, G.I. and Sangchote, S. (1994) Control of postharvest diseases of tropical fruits:
challenges for the 21st century. In: Champ, B.R., Highley, E. and Johnson, G.I. (eds)
Postharvest Handling of Tropical Fruits. Australian Centre for International Agricul-
tural Research (ACIAR) Proceedings 50. ACIAR, Canberra, pp. 140161.
Johnson, G.I., Muirhead, I.F., Mayers, P. and Cooke, T. (1989a) Diseases. In: Ridgeway,
R. (ed.) Mango Pests and Disorders. Department of Primary Industries, Brisbane,
Queensland, pp. 19.
Johnson, G.I., Muirhead, I.F. and Rappel, L.M. (1989b) Mango post-harvest disease con-
trol: a review of research in Australia, Malaysia and Thailand. ASEAN Food Journal
4, 139141.
Johnson, G.I., Boag, T.S., Cooke, A.W., Izard, M., Panitz, M. and Sangchote, S. (1990a)
Interaction of postharvest disease control treatments and gamma radiation on man-
goes. Annals of Applied Biology 116, 245257.
Johnson, G.I., Sangchote, S. and Cooke, A.W. (1990b) Control of stem-end rot (Dothi-
orella dominicana) and other postharvest diseases of mangoes (cv. Kensington
Pride) during short- and long-term storage. Tropical Agriculture 67, 183187.
Johnson, G.I., Mead, A.J, Cooke, A.W. and Dean, J.R. (1991) Mango stem end rot patho-
gens infection levels between owering and harvest. Annals of Applied Biology
119, 465473.
Johnson, G.I., Mead, A.J., Cooke, A.W. and Dean, J.R. (1992) Mango stem end rot patho-
gens fruit infection by endophytic colonisation of the inorescence and pedicel.
Annals of Applied Biology 120, 225234.
Johnson, G.I., Cooke, A.W. and Mead, A.J. (1993) Infection and quiescence of mango
stem end rot pathogens. Acta Horticulturae 341, 329336.
Johnson, G.I., Cooke, A.W., Mayers, P.E., Bagshaw, J.S., Ledger, S.N., Yuen, C.M.C., Tan,
S.C. and Joyce, D.C. (1996) Mangoes. In: Coates, L., Cooke, T., Persley, D., Beattie,
B. and Wade, N. (eds) Postharvest Diseases of Horticultural Produce. Vol. 2: Tropi-
cal Fruit. Queensland Department of Primary Industries, Brisbane, Queensland,
pp. 4662.
Postharvest Technology 591
Johnson, G.I., Sharp, J.L., Milne, D.L. and Oosthuyse, S.A. (1997) Postharvest technol-
ogy and quarantine treatments. In: Litz, R.E. (ed.) The Mango: Botany, Production
and Uses, CAB International, Wallingford, UK, pp. 447508.
Johnson, G.I., Highley, E. and Joyce, D.C. (eds) (1998) Disease Resistance in Fruit. Aus-
tralian Centre for International Agricultural Research (ACIAR) Proceedings 80.
ACIAR, Canberra.
Johnson, G.I., Weinberger, K. and Wu, M.H. (2008) The Vegetable Industry in Tropical
Asia: an Overview of Production and Trade, with a Focus on Thailand, Indonesia,
the Philippines, Vietnam and India [CD-ROM]. Shanhua, Taiwan: AVRDC - The
World Vegetable Center. 56 pp. (Exploration series; no. 1.)
Joshi, G.D. and Roy, S.K. (1985) Spongy tissue in mango a physiological disorder. Indian
Horticulture 29, 2122.
Joyce, D.C. and Patterson, B. (1994) Postharvest water relations in horticultural crops:
principles and problems. In: Champ, B.R., Highley, E. and Johnson, G.I. (eds) Post-
harvest Handling of Tropical Fruits. Australian Centre for International Agricultural
Research (ACIAR) Proceedings 50. ACIAR, Canberra, pp. 228238.
Joyce, D.C., Hockings, P.D., Mazucco, R.A., Shorter, A.J. and Brereton, I.M. (1993) Heat
treatment injury of mango fruit revealed by non-destructive magnetic resonance
imaging. Postharvest Biology and Technology 3, 305311.
Joyce, D.C., Macnish, A.J., Hofman, P.J., Simons, D.H. and Reid, M.S. (1999) Use of
1-methylcyclopropene to modulate banana ripening. In: Kanellis, A.K., Chang, C.,
Klee, H., Bleecker, A.B., Pech, J.C. and Grierson, D. (eds) Biology and Biotechnol-
ogy of the Plant Hormone Ethylene II. Kluwer Academic Publishers, Dordrecht, the
Netherlands, pp. 189190.
Joyce, D.C., Shorter, A.J. and Hockings, P.D. (2001) Mango fruit calcium levels and the
effect of postharvest calcium inltration at different maturities. Scientia Horticultu-
rae 91, 8199.
Kane, O. and Marcellin, P. (1978) Incidence of ripening and chilling the mitrochondria
from mango fruits. Plant Physiology 61, 634638.
Karunanayake, K.O.L.C. (2007) Natural defence mechanisms in mango fruit and their
potential in management of postharvest diseases. PhD thesis, University of Peride-
nya, Peridenya, Sri Lanka.
Katrodia, J.S. (1989) Spongy tissue in mango causes and control measures. Acta Hor-
ticulturae 231, 814826.
Katrodia, J.S. and Sheth, I.K. (1989) Spongy tissue development in mango fruit of cultivar Al-
phonso in relation to temperature and its control. Acta Horticulturae 231, 827834.
Ke, D. and Kader, A.A. (1992) Potential of controlled atmospheres for postharvest insect
disinfestation of fruits and vegetables. Postharvest News and Information 3, 31N
37N.
Keil, H., Wasserman, D. and Dawson, E.R. (1946) Mango dermatitis and its relationship
to poison-ivy hypersensitivity. Annals of Allergy 4, 268281.
Kernot, I.I., Meurant, N., Holmes, R., MacLeod, N., Fullelove, G. and Bally, I. (eds)
(1999) Mango Information Kit. Queensland Department of Primary Industries, Bris-
bane, Queensland.
Kitagawa, H., Matsui, T., Kawada, K. and Agravante, J.U. (1992) Bagging of fruit on the
tree to control disease. Acta Horticulturae 321, 871875.
Kitinoja, L. and Kader, A.A. (2003) Small-scale Postharvest Handling Practices a Man-
ual for Horticultural Crops. University of California, Davis, California.
Kobiler, I., Shalom, Y., Roth, I., Akerman, M., Vinokur, Y., Fuchs, Y. and Prusky, D. (2001)
Effect of 2,4-dichlorophenoxyacetic acid on the incidence of side and stem end rots
in mango fruits. Postharvest Biology and Technology 23, 2332.
G.I. Johnson and P.J. Hofman 592
Kondo, S., Kittikorn, M. and Kanlayanarat, S. (2005) Preharvest antioxidant activities of
tropical fruit and the effect of low temperature storage on antioxidants and jas-
monates. Postharvest Biology and Technology 36, 309318.
Koomen, I., Dodd, J.C., Jeger, M.J. and Jeffries, P. (1990) Postharvest biocontrol of an-
thracnose disease of mangoes. Journal of the Science of Food and Agriculture 50,
137138.
Korsten, L. (2006) Advances in control of postharvest diseases in tropical fresh produce.
International Journal of Postharvest Technology and Innovation 1, 4861.
Korsten, L., Van Harmelen, M.W.S., Heitmann, A., De Villiers, E.E. and De Jager, E.S.
(1991) Biological control of postharvest mango fruit diseases. South African Mango
Growers Association Yearbook 11, 6567.
Korsten, L., Lonsdale, J.H., De Villiers, E.E. and De Jager, E.S. (1992) Preharvest biologi-
cal control of mango diseases. South African Mango Growers Association Yearbook
12, 7278.
Korsten, L., De Villiers, E.E. and Lonsdale, J.H. (1993) Biological control of mango post-
harvest disease in the packhouse. South African Mango Growers Association Year-
book 13, 117121.
Korsten, L., De Villiers, E.E., Wehner, F.C. and Kotze, J.M. (1994) A review of biological
control of postharvest diseases of subtropical fruits. In: Champ, B.R., Highley, E.
and Johnson, G.I. (eds) Postharvest Handling of Tropical Fruits. Australian Centre for
International Agricultural Research (ACIAR) Proceedings 50. ACIAR, Canberra,
pp. 172185.
Kostermans, A.J.G.H. and Bompard, J.M. (1993) The Mangoes, their Botany, Nomencla-
ture, Horticulture and Utilisation. Academic Press, London.
Kramer, A. (1973) Fruits and vegetables. In: Kramer, A. and Twigg, B.A. (eds) Quality
Control for the Food Industry. Vol. 2. AVI, Westport, Connecticut, USA, pp. 157
228.
Kuo, L.S., Su, C.Y., Hseu, C.Y., Chao, Y.F., Chen, H.Y., Liao, J.Y. and Chu, C.F. (1987)
Vapour Heat Treatment for Elimination of Dacus dorsalis and Dacus cucurbitae In-
festing Mango Fruits. Taiwan Bureau of Commodity Inspection and Quarantine,
Ministry of Economic Affairs, Taipei, Taiwan.
Lahav, E. and Whiley, A.W. (2002) Irrigation and mineral nutrition. In: Whiley, A.W.,
Schaffer, B. and Wolstenholme, B.N. (eds) The Avocado: Botany, Production and
Uses. CAB International, Wallingford, UK, pp. 259297.
Lalel, H.J.D. and Singh, Z. (2006) Controlled atmosphere storage of Delta R2E2 mango
fruit affects production of aroma volatile compounds. Journal of Horticultural Sci-
ence and Biotechnology 81, 449457.
Lalel, H.J.D., Singh, Z. and Tan, S. (2003a) Glycosidically-bound aroma volatile com-
pounds in the skin and pulp of Kensington Pride mango fruit at different stages of
maturity. Postharvest Biology and Technology 29, 205218.
Lalel, H.J.D., Singh, Z. and Tan, S.C. (2003b) Maturity stage at harvest affects fruit ripen-
ing, quality and biosynthesis of aroma volatile compounds in Kensington Pride
mango. Journal of Horticultural Science and Biotechnology 78, 225233.
Lalel, H.J.D., Singh, Z. and Tan, S.C. (2004) Ripening temperatures inuence biosynthe-
sis of aroma volatile compounds in Kensington Pride mango fruit. Journal of Hor-
ticultural Science and Biotechnology 79, 146157.
Ledger, S.N. (1986) Mango packaging. In: Proceedings of the First Australian Mango
Research Workshop. Commonwealth Scientic and Industrial Research Organiza-
tion (CSIRO), Melbourne, pp. 314319.
Ledger, S.N. (1991a) Impact and pressure resistance of mangoes postharvest handling
of mangoes. Acta Horticulturae 291, 479488.
Postharvest Technology 593
Ledger, S.N. (1991b) Post-harvest handling of mangoes. Acta Horticulturae 291,
479488.
Ledger, S.N. (2003a) Dont gas hot fruit. Mango Care 39, 23.
Ledger, S.N. (2003b) Two handling systems. Mango Care 36, 1314.
Ledger, S.N. (2004) Control of Postharvest Diseases in Mango. Department of Primary
Industries and Fisheries (DPI&F) note. Department of Primary Industries and Fisher-
ies, Brisbane, Australia.
Ledger, S.N. (2007) Managing mango ripening and storage rooms. In: Delivering Mango
Research. The Sixth Amistar Australian Mango Conference. Gold Coast, Queensland,
pp. 2629. Available at: http://www.mangoes.net.au/downloads/File/PDF/industry_
conference/mango_conference_07_proceedings.pdf (accessed 4 April 2008).
Ledger, S.N. and Bagshaw, J. (1994) Stimulating change in horticulture quality manage-
ment. Australasian Postharvest Conference Proceedings. University of Queensland
Gatton College, Lawes, Queensland, pp. 3540.
Ledger, S.N. and Premier, R. (2006) Interpretive Guide for Association of South-east
Asian Nations (ASEAN) Good Agricultural Practice (GAP) Produce Quality Module
Good Agricultural Practices for Production of Fresh Fruit and Vegetables in ASEAN
Countries, Quality Assurance Systems for ASEAN Fruit and Vegetables Project. ASE-
AN-Australia Development Cooperation Project. Available at: http://www.aphnet.
org/gap/ASEANgap.html (accessed 1 May 2008).
Ledger, S.N., Campbell, T.P., Holmes, R.J. and Kernot, I.I. (2002a) The saleable life index.
Mango Care 36, 23.
Ledger, S.N., Holmes, R. and Kernot, I. (2002b) Better mangoes special edition. Mango
Care 36, 114.
Len, D.M., Ortega, D.A., Cabrera, H., de la Cruz, J., Parkin, K.L. and Garcia, H.S.
(2000) Postharvest disinfestation of mango (Mangifera indica cv. Manila) with con-
trolled atmospheres. Journal of Applied Horticulture 2, 7175.
Lim, T.K. and Kuppelweiser, W. (1993) Mango sapburn amelioration in the Northern
Territory. Acta Horticulturae 341, 518527.
Lin, T.H., Tseng, F.C., Chang, C.R. and Wang, I.Y. (1976) Multiple treatment for disinfest-
ing oriental fruit y in mangoes. Plant Protection Bulletin 18, 231241.
Lizada, M.C.C. (1994) Fruit handling systems in developing countries. In: Champ, B.R.,
Highley, E. and Johnson, G.I. (eds) Postharvest Handling of Tropical Fruits. Austra-
lian Centre for International Agricultural Research (ACIAR) Proceedings 50. ACIAR,
Canberra, pp. 109115.
Lonsdale, J.H. (1992) In search of an effective postharvest treatment for the control of
postharvest diseases of mangoes. South African Mango Growers Association Year-
book 12, 3236.
Lonsdale, J.H. (1993) Strategies for the control of postharvest diseases of mango. South
African Mango Growers Association Yearbook 13, 109116.
Loveys, B.R., Robinson, S.P., Brophy, J.J. and Chacko, E.K. (1992) Mango sapburn: com-
ponents of fruit sap and their role in causing skin damage. Australian Journal of
Plant Physiology 19, 449457.
Lunn, D. (2004) Prochloraz. Food and Agriculture Organization (FAO) Pesticide Man-
agement Pesticide Specications and Quality Control Standards. Available at:
http://www.fao.org/ag/AGP/AGPP/Pesticid/JMPR/Download/2004_eva/Prochloraz.
pdf (accessed 28 October 2007).
MacLeod, A.J. and Snyder, C.H. (1985) Volatile components of two cultivars of mango
from Florida. Journal of Agriculture and Food Chemistry 33, 380384.
MacLeod, A.J., MacLeod, G. and Snyder, C.H. (1988) Volatile aroma constituents of
mango (cv. Kensington). Phytochemistry 27, 21892193.
G.I. Johnson and P.J. Hofman 594
Mahayothee, B., Mhlbauer, W., Neidhart, S., Leitenberger, M. and Carle, R. (2004)
Non-destructive determination of maturity of Thai mangoes by near-infrared spec-
troscopy. Acta Horticulturae 645, 581588.
Mahendra, M.S., Suryadi, M. and Padmono, N. (2002) Current status of the rural econ-
omy and measures for revitalising it and increasing farmers income in Indonesia:
development of substantial Gedong Gincu mango plantations in West Java. In:
The Ninth Japanese International Research Centre for Agricultural Sciences Inter-
national Symposium 2002 Value-Addition to Agricultural Products. Tsukuba, Ja-
pan, pp. 4952.
Mangan, R.L. and Ingle, S.J. (1992) Forced hot-air quarantine treatment for mangoes
infested with West Indian fruit y (Diptera: Tephritidae). Journal of Economic Ento-
mology 85, 18591864.
Maqbool, M., Malik, A.U. and Jabbar, A. (2007) Sap dynamics and its management in
commercial mango cultivars of Pakistan. Pakistan Journal of Botany 39, 15651574.
Marigheto, N., Venturi, L. and Hills, B. (2008) Two-dimensional NMR relaxation studies
of apple quality. Postharvest Biology and Technology 48, 331340.
Maxtend (2008) Maxim freshness global service. Available at: www.maxtend.com.au
(accessed 7 April 2008).
McGregor, B.M. (1987) Tropical Products Transport Handbook. United States Department
of Agriculture (USDA), Agricultural Handbook No. 668. USDA, Washington, DC.
McKenzie, C.B. (1994) The background skin colour of exported mango fruit in relation
to tree nitrogen status. South African Mango Growers Association Yearbook 14,
2628.
McLauchlan, R.L. and Barker, L.R. (1994) Controlled atmospheres for Kensington man-
go storage: classical atmospheres. In: Johnson, G.I. and Highley, E. (eds) Develop-
ment of Postharvest Handling Technology for Tropical Tree Fruits. Australian Centre
for International Agricultural Research (ACIAR) Proceedings 58. ACIAR, Canberra,
pp. 2529.
McLauchlan, R.L. and Wells, L.A. (1994) Storage and ripening temperatures for
Kensington mangoes. In: Johnson, G.I. and Highley, E. (eds) Development of
Postharvest Handling Technology for Tropical Tree Fruits. Australian Centre for
International Agricultural Research (ACIAR) Proceedings 58. ACIAR, Canberra,
pp. 2529.
McMaugh, T. (2005) Guidelines for the surveillance for plant pests in Asia and the Pa-
cic. Australian Centre for International Agricultural Research (ACIAR) Mongraph
MN119 [English], MN119a [Indonesian]. ACIAR, Canberra. Available at: http://
www.aciar.gov.au/publication/MN119+%5BEnglish%5D%2C+MN119a+%5BInd
onesian%5D (accessed 1 May 2008.).
Mead, A.J. and Winston, E.C. (1991) Description of the disorder stem-end cavity, pos-
sible causes and suggestions for reducing incidence in packing sheds. Acta Horti-
culturae 291, 265271.
Medlicott, A.P. (1985) Mango fruit ripening and the effects on maturity, temperature and
gases. PhD thesis, Wolverhampton Polytechnic, Wolverhampton, UK.
Medlicott, A.P., Reynolds, S.B., New, S.W. and Thompson, A.K. (1987) Harvest maturity
effects on mango fruit ripening. Tropical Agriculture 65, 153157.
Medlicott, A.P., NDiaye, M. and Sigrist, J.M.M. (1990a) Harvest maturity and concen-
tration and exposure time to acetylene inuence initiation of ripening in mangos.
Journal of the American Society of Horticultural Science 115, 426430.
Medlicott, A.P., Sigrist, J.M.M. and Sy, O. (1990b) Ripening of mangoes following low
temperature storage. Journal of the American Society of Horticultural Science 115,
430434.
Postharvest Technology 595
Meiburg, G.F., Hofman, P.J., Smith, L.G., Cooke, A.W. and Barker, J.A. (1998) Quality of
Kensington mangoes after short duration exposure to high carbon dioxide concen-
trations. In: Coates, L.M., Hofman, P.J. and Johnson, G.I. (eds) Disease Control in
Storage Life Extension in Fruit. Australian Centre for International Agricultural Re-
search (ACIAR) Proceedings No. 81. ACIAR, Canberra, pp. 5560.
Milne, D.L. (1994) Postharvest handling of avocado, mango and lychee for export from
South Africa. In: Champ, B.R., Highley, E. and ]ohnson, G.I. (eds) Postharvest Han-
dling of Tropical Fruits. Australian Centre for International Agricultural Research
(ACIAR) Proceedings 50. ACIAR, Canberra, pp. 7389.
Minnis, D. (1993) Horticultural marketing whats really involved. In: Australasian Post-
harvest Conference Proceedings, Department of Primary Industries, Gatton, Queen-
sland, Australia, pp. 203206.
Minnis, D.E. (1994) Prospects for marketing tropical fruits in Asia. In: Champ, B.R.,
Highley, E. and Johnson, G.I. (eds) Postharvest Handling of Tropical Fruits. Austra-
lian Centre for International Agricultural Research (ACIAR) Proceedings 50. ACIAR,
Canberra, pp. 5664.
Mitcham, E.J. (2007) Heat with controlled atmospheres. In: Tang, J., Mitcham, E., Wang,
S. and Lurie, S. (eds) Heat Treatments for Postharvest Pest Control: Theory and
Practice. CAB International, Wallingford, UK, pp. 251268.
Mitchell, G.E., McLauchlan, L., Isaacs, A.R., Williams, D.J. and Nottingham, S.M. (1992)
Effect of low dose irradiation on composition of tropical fruits and vegetables. Jour-
nal of Food Composition and Analysis 5, 291311.
Mizrach, A. (2008) Ultrasonic technology for quality evaluation of fresh fruit and vege-
tables in pre- and postharvest processes. Postharvest Biology and Technology 48,
315330.
Moreno, M., Castell-Perez, M.E., Gomes, C., Da Silva, P.F. and Moreira, R.G. (2006) Effects
of electron beam irradiation on physical, textural, and microstructural properties of
Tommy Atkins mangoes (Mangifera indica L.). Journal of Food Science 71, E80E86.
Moreno, M., Castell-Perez, M.E., Gomes, C., Da Silva, P.F., Kim, J. and Moreira, R.G.
(2007) Irradiation for postharvest quarantine insects. Formosan Entomology 27,
115.
Muirhead, I.F. (1976) Postharvest control of mango anthracnose with benomyl and hot
water. Australian Journal of Experimental Agriculture and Animal Husbandry 16,
600603.
Muirhead, I.F. and Grattidge, R. (1986) Postharvest diseases of mango the Queensland
experience. In: Proceedings of the First Australian Mango Workshop. Common-
wealth Scientic and Industrial Research Organization (CSIRO), Melbourne,
pp. 248252.
Muller, A.T. and Burt, J.R. (1989) Postharvest storage control of mango stem end rot with
fungicidal dips. Australian Journal of Experimental Agriculture 29, 125127.
Murray, E. and George, E. (1994) Fruit packing house operations to improve returns. In:
Champ, B.C., Highley, E. and Johnson, G.I. (eds) Postharvest Handling of Tropical
Fruits. Australian Centre for International Agricultural Research (ACIAR) Proceed-
ings 50. ACIAR, Canberra, pp. 105108.
Nair, S.S., Singh, Z. and Tan, S. (2003) Chilling injury development in Kensington Pride
mango fruit in relation to free polyamines. In: Australian Postharvest Horticulture
Conference. Brisbane, Queensland, pp. 278279.
Nascimento, A.S., Malavasi, A., Morgante, J.S. and Duarte, A.L.A. (1992) Hot-water im-
mersion treatment for mangoes infested with Anastrepha fraterculus, A. obliqua,
and Ceratitis capitata (Diptera: Tephritidae) in Brazil. Journal of Economic Entomol-
ogy 85, 456460.
G.I. Johnson and P.J. Hofman 596
Natural Resources Institute (NRI) (2008a) Small Producers in Export Horticulture: a
Guide to Best Practice. Elements for Successful Business. Natural Resources Institute
UK: the University of Greenwich. Available at: http://www.nri.org/projects/NRET/
SPCDR/Chapter1/elements-1-5-1.htm (accessed 28 March 2008).
Natural Resources Institute (NRI) (2008b) Smallholders in Export Horticulture: a Guide
to Best Practice. Natural Resources Institute UK: the University of Greenwich. Avail-
able at: http://www.nri.org/projects/NRET/SPCDR/index.htm (accessed 1 April
2008).
Neven, L.G. (2008) Organic quarantine treatments for tree fruits. HortScience 43, 2226.
Nguyen, H.X. (2003) Pre-harvest and postharvest factors affecting skin colour and other
quality attributes of Kensington Pride mango (Mangifera indica Linn.). PhD thesis,
University of Queensland, Brisbane, Queensland.
Nguyen, H., Hofman, P.J., Smith, L.G., Stubbings, B., Adkins, M. and McConchie, R.
(2002) Effect of ethylene and ripening temperatures on the skin colour and esh
characteristics of ripe Kensington Pride mango fruit. Acta Horticulturae 575, 635
642.
Nguyen, H., Hofman, P., Holmes, R., Bally, I., Stubbings, B. and McConchie, R. (2004)
Effects of nitrogen on the skin colour and other quality attributes of ripe Kensington
Pride mango Mangifera indica L. fruit. Journal of Horticultural Science and Bio-
technology 79, 204210.
Nguyen, M.C., Wei, W., Vo, T.T., Rankin, M. and Russell, I. (2004) Getting farmers to
work together: the experiences of mango growers in the Mekong Delta region of
Vietnam. In: Johnson, G.I. and Hofman, P.J. (eds) Agriproduct Supply-Chain Man-
agement in Developing Countries. Australian Centre for International Agricultural
Research (ACIAR) Proceedings 119. ACIAR, Canberra, pp. 107111.
OHare, T.J. (1994) The susceptibility of Thai and Australian mango cultivars to sap
injury and possible means of control. In: Johnson, G.I. and Highley, E. (eds) Devel-
opment of Postharvest Handling Technology for Tropical Tree Fruits. Australian
Centre for International Agricultural Research (ACIAR) Proceedings 58. ACIAR,
Canberra, pp. 2124.
OHare, T.J. and Prasad, A. (1993) The alleviation of sap-induced skin injury by calcium
hydroxide. Acta Horticulturae 321, 372381.
OHare, T.J., McLaughlan, R.L. and Turner, C.D. (1994) Effect of co-shipment of exotic
tropical fruit on existing fruit transported in bulk shipment. Proceedings of the Aus-
tralasian Postharvest Conference. University of Queensland, Gatton, Queensland,
pp. 245250.
OHare, T.J., Bally, I.S.E., Dahler, J.M., Saks, Y. and Underhill, S.J.R. (1999) Characterisa-
tion and induction of etch browning in the skin of mango fruit. Postharvest Biol-
ogy and Technology 16, 269277.
Oka, K., Saito, F., Yasuhara, T. and Sugimoto, A. (2004) A study of cross-reactions be-
tween mango contact allergens and urushiol. Contact Dermatitis 51, 292296.
Oosthuyse, S.A. (1990) Effect of lowering the temperature of the cold storage regime of
export mangoes after four weeks of storage. South African Mango Growers Asso-
ciation Yearbook 10, 3856.
Oosthuyse, S.A. (1991) Correlating pre-storage ripeness of mangoes with the proportion
of good quality fruit present after four weeks of cold-storage. South African Mango
Growers Association Yearbook 11, 4953.
Oosthuyse, S.A. (1992) Long-term refrigerated storage of mangoes: what are the issues?
South African Mango Growers Association Yearbook 12, 6165.
Oosthuyse, S.A. (1993) Disorders of breless mangos growth in South Africa for export.
South African Mango Growers Association Yearbook 13, 8088.
Postharvest Technology 597
Oosthuyse, S.A. (1994) Quality of mature Zill mangoes after long-term refrigerated
storage as determined by pre-storage ripeness and cold-storage regime. South Afri-
can Mango Growers Association Yearbook 14, 3742.
Oosthuyse, S.A. (1995) The South African industry experience. South African Mango
Growers Association Yearbook 15, 111.
Oosthuyse, S.A. (1998) Effect of environmental conditions at harvest on the incidence of
lenticel damage in mango. South African Mango Growers Association Yearbook
18, 1517.
Oosthuyse, S.A. (1999) Effects of each of the stages in the pack-line on the incidence of
lenticel damage in Keitt and Tommy Atkins mango. South African Mango Grow-
ers Association Yearbook 19, 3739.
Opara, L.U. and Nguyen, H.X. (1999) Postharvest handling and environmental factors
inuencing mango fruit quality a review. Journal of South Pacic Agriculture 6,
119.
Pacic Data Systems (PDS) (2008) Ethylene/Ripening Gas Controlling System. Pacic
Data Systems. Available at: http://www.pacdatasys.com.au/dataloggers/Systems_
Overview.htm (accessed 7 April 2008).
Peacock, B.E. (1986) Postharvest handling of mangoes. In: Proceedings of the First
Australian Mango Research Workshop. Commonwealth Scientic and Industrial
Research Organization (CSIRO), Melbourne, pp. 295313.
Peak, E.M., Fitzell, R.D., Hannah, R.S. and Batten, D.J. (1986) Development of a micro-
processor-based data recording system for predicting plant disease based on studies
on mango anthracnose. Computers and Electronics in Agriculture 1, 251262.
Pea, J. (2004) Integrated pest management and monitoring techniques for mango pests.
Acta Horticulturae 645, 151161.
Penchaiya, P., Jansasithorn, R. and Kanlavanarat, S. (2006) Effect of 1-MCP on physio-
logical changes in mango Nam Dok Mai. Acta Horticulturae 712, 717721.
Pesis, E., Ampunpong, C., Shusiri, B. and Hewett, E.W. (1994) Enhancement of ethylene
and CO
2
production in apple fruit following short-term exposure to high CO
2
. Post-
harvest Biology and Technology 4, 309317.
Pesis, E., Aharoni, D., Aharon, Z., Ben-Arie, R., Aharoni, N. and Fuchs, Y. (2000) Modi-
ed atmosphere and modied humidity packaging alleviates chilling injury symp-
toms in mango fruit. Postharvest Biology and Technology 19, 93101.
Phakawatmongkol, W., Ketsa, S. and van Doorn, W.G. (2004) Variation in fruit chilling
injury among mango cultivars. Postharvest Biology and Technology 32, 115118.
Pieiro, M., Berania, L. and Ros, D. (2004) Improving the Quality and Safety of Fresh
Fruits and Vegetables: a Practical Approach. Manual for Trainers. Food and Agricul-
ture Organization (FAO), Rome. Available at: http://www.fao.org/docrep/007/
y5488e/y5488e00.htm (accessed 1 May 2008).
Plan, M.R.R., Joyce, D.C., Ogle, H.J. and Johnson, G.I. (2002) Mango stem-end rot
(Botryosphaeria dothidea) disease control by partial-pressure inltration of fungi-
cides. Australian Journal of Experimental Agriculture 42, 625629.
Ploetz, R.C. (ed.) (2003) Diseases of Tropical Fruit Crops. CAB International, Wallingford,
UK.
Ploetz, R.C. (2004) The major diseases of mango: strategies and potential for sustainable
management. Acta Horticulturae 645, 137150.
Ploetz, R.C. (2007) Diseases of tropical perennial crops: challenging problems in di-
verse environments. Plant Disease 91, 644663.
Ploetz, R.C. and Prakash, O. (1997) Foliar, oral and soilborne diseases. In: Litz, R.E.
(ed.) The Mango: Botany, Production and Uses. CAB International, Wallingford,
UK, pp. 281325.
G.I. Johnson and P.J. Hofman 598
Pofey, M., Owens, G., Kulkani, V., Smith, E.S.C. and Conde, B. (1999) Mango Management
Flowering to Market. Agnote, 301 No. D9, December 1999. Agdex No: 234/11.
Plant Protection and Quarantine (PPQ) (2007) 5. Treatment Manual Treatment Schedules
T100 Schedules for Fruit, Nuts, and Vegetables. United States Department of Ag-
riculture (USDA) Animal and Plant Health Inspection Service (APHIS). Available at:
http://www.aphis.usda.gov/import_export/plants/manuals/ports/downloads/
treatment_pdf/05_02_t100schedules.pdf (accessed 31 October 2007).
Proctor, F.J. and Cropley, J.P. (1994) Trends and changes in the European market for
tropical fruits and their impact on technology requirements. In: Champ, B.R., High-
ley, E. and Johnson, G.I. (eds) Postharvest Handling of Tropical Fruits. Australian
Centre for International Agricultural Research (ACIAR) Proceedings 50. ACIAR,
Canberra, pp. 6572.
Prusky, D. and Keen, N.T. (1993) Involvement of preformed antifungal compounds in
the resistance of subtropical fruits to fungal decay. Plant Disease 77, 114119.
Prusky, D. and Plumbley, R.A. (1992) Quiescent infections of Colletotrichum in tropical
and subtropical fruit. In: Bailey, J.A. and Jeger, M.J. (eds) Colletotrichum: Biology,
Pathology and Control. CAB International, Wallingford, UK, pp. 289307.
Prusky, D., Fuchs, Y. and Zauberman, G. (1980) A method for preharvest assessment of
latent infections in fruit. Annals of Applied Biology 98, 7985.
Prusky, D., Keen, N.T., Sims, J.J. and Midland, S.L. (1982) Possible involvement of an
antifungal compound in latency of Colletotrichum gloeosporioides in unripe avo-
cado fruits. Phyiopathology 72, 15781582.
Prusky, D., Plumbley, R.A. and Kobiler, I. (1992) Modulation of natural resistance of
avocado fruit by CO
2
treatments. Physiological and Molecular Plant Pathology 39,
325334.
Prusky, D., Gat, Z. and Burd, P. (1993a) Effect of relative humidity during growth on the
incidence of quiescent infections of Altemaria alternata. Plant Disease 77, 249252.
Prusky, D., Kobiler, I., Zauberman, G. and Fuchs, Y. (1993b) Preharvest conditions and
postharvest treatments affecting incidence of decay in mango fruits during storage.
Acta Horticulturae 341, 307320.
Prusky, D., Plumbley, R.A., Kobiler, I., Zauberman, G. and Fuchs, Y. (1993c) The effect
of elevated CO
2
levels on the symptom expression of Colletotrichum gloeospori-
oides on avocado fruits. Plant Pathology 42, 900904.
Prusky, D., Fuchs, Y., Kobiler, I., Roth, I., Weksler, A., Shalom, Y., Fallik, E., Zauberman,
G., Pesis, E., Akerman, M., Ykutiely, O., Weissblum, A., Regev, R. and Artes, L.
(1999) Effect of hot water brushing, prochloraz treatment and waxing on the inci-
dence of black spot decay caused by Alternaria alternata in mango fruits. Posthar-
vest Biology and Technology 15, 165174.
Prusky, D., Kobiler, I., Akerman, M. and Miyara, I. (2006) Effect of acidic solutions and
acidic prochloraz on the control of postharvest decay caused by Alternaria alternata
in mango and persimmon fruit. Postharvest Biology and Technology 42, 134141.
Regional Standards for Phytosanitary Measures (RSPM) (2004) Guidelines for the Estab-
lishment, Maintenance and Verication of Fruit Fly Pest Free Areas in North America.
RSPM No. 17. North American Plant Protection Organisation (NAPPO). Available at:
http://www.nappo.org/Standards/REVIEW/RSPMNo17-e.pdf (accessed 1 May 2008).
Reid, M.S. (2002) Ethylene in postharvest technology. In: Kader, A.A. (ed.) Postharvest
Technology of Horticultural Crops, 3rd edn. University of California, Oakland,
California, pp. 149162.
Reuveni, M. (2000) Efcacy of trioxystrobin (Flint), a new strobilurin fungicide, in con-
trolling powdery mildews on apple, mango and nectarine, and rust on prune trees.
Crop Protection 19, 335341.
Postharvest Technology 599
Reuveni, M., Harpaz, M. and Reuveni, R. (1998) Integrated control of powdery mildew
in eld-grown mango trees by foliar sprays of mono-potassium phosphate fertiliser,
sterol inhibitor fungicides and the stroilurin Kresoxym-methyl. European Journal of
Plant Pathology 104, 853860.
Reyes, L.F. and Cisneros-Zevallos, L. (2007) Electron-beam ionizing radiation stress effects
on mango fruit (Mangifera indica L.) antioxidant constituents before and during post-
harvest storage. Journal of Agricultural and Food Chemistry 55, 61326139.
Reyes, M.U., Paull, R.E., Gautz, L.D., Armstrong, J.W. and Follett, P.A. (2000) Non-
destructive inspection of mango fruit (Mangifera indica L.) with soft X-ray imaging.
Acta Horticulturae 509, 787792.
Robinson, S.P., Loveys, B.R. and Chacko, E.K. (1993) Polyphenol oxidase enzymes in
the sap and skin of mango fruit. Australian Journal of Plant Physiology 20, 99107.
Rolle, R.S. (2006) Improving postharvest management and marketing in the Asia-Pacic
region: issues and challenges. In: Postharvest Management of Fruit and Vegetables
in the Asia-Pacic Region Reports of the Asian Productivity Organisation (APO)
Seminar on Reduction of Postharvest Losses of Fruit and Vegetables held in India;
and Marketing and Food Safety: Challenges in Postharvest Management of Ag-
ricultural/Horticultural Products in Islamic Republic of Iran. Asian Productivity
Organisation, pp. 2331. Available at: http://www.apo-tokyo.org/00e-books/AG-18_
PostHarvest.htm (accessed 1 May 2008).
Rosa, B.A., Aharoni, D., Feygenberg, O., Aharoni, N., Keynan, A. and Pesis, E. (2001)
Effect of modied atmosphere packaging on mango ripening. Acta Horticulturae
553, 607609.
Sadasivam, R., Muthuswamy, S., Sundaraj, J.S. and Vasudevan, V. (1971) Note on chill-
ing injury in mango (Mangifera indica L.) fruits in refrigerated storage. Indian Jour-
nal of Agricultural Science 41, 715716.
Saranwong, S., Sornsrivichai, J. and Kawano, S. (2004) Prediction of ripe-stage eating
quality of mango fruit from its harvest quality measured non-destructively by near
infrared spectroscopy. Postharvest Biology and Technology 31, 137145.
Schoorl, D. and Holt, J.E. (1982) Planning for change in horticulture a case study in
packing shed design. Agricultural Systems 8, 7786.
Schoorl, D. and Holt, J.E. (1985) Design of a packing shed for the ripening, packing and
handling of tomatoes. Queensland Agricultural Journal 111, 185190.
Segarra-Carmona, A.E., Franqui, R.A., Ramirez-Ramos, L.V., Santiago, L.R. and Torres-
Rivera, C.N. (1990) Hot-water dip treatments to destroy Anastrepha obliqua larvae
(Diptera: Tephritidae) in mangoes from Puerto Rico. Journal of Agriculture of the
University of Puerto Rico 74, 441447.
Self, G., de Assis, J.S. and Caron, V.C. (2006) Effects of postharvest handling on lenticel
spotting of Tommy Atkins mangoes from Northeast Brazil. Acta Horticulturae 712,
543550.
Seo, S.T., Chambers, D.L., Akamine, E.K., Komura, M. and Lee, C.Y.L. (1972) Hot water-
ethylene dibromide fumigation-refrigeration treatment for mangoes infested by orien-
tal and Mediterranean fruit ies. Journal of Economic Entomology 65, 13721374.
Seo, S.T., Kobayashi, R.M., Chambers, D.L., Steiner, L.F., Lee, C.Y.L. and Komura, M.
(1974) Mango weevil: cobalt 60 gamma-irradiation of packaged mangoes. Journal
of Economic Entomology 67, 504505.
Sepiah, M. (1986) Effectiveness of hot water, hot benomyl and cooling on postharvest
diseases of mango. ASEAN Food Journal 2, 117120.
Seymour, G.B., Ndiaye, M., Wainwright, H. and Tucker, G.A. (1990) Effects of cultivar
and harvest maturity on ripening of mangoes during storage. Journal of Horticul-
tural Science 65, 479483.
G.I. Johnson and P.J. Hofman 600
Sharp, J.L. (1992) Hot-air quarantine treatment for mango infested with Caribbean fruit
y (Diptera: Tephritidae). Journal of Economic Entomology 85, 23022304.
Sharp, J.L. and Heather, N.W. (2002) Quarantine treatments for pests of tropical fruits.
In: Pea, J.E., Sharp, J.L. and Mysoki, M. (eds) Tropical Fruit Pests and Pollinators
Biology, Economic Importance, Natural Enemies and Control. CAB International,
Wallingford, UK, pp. 391406.
Sharp, J.L. and Picho-Martinez, H. (1990) Hot-water quarantine treatment to control
fruit ies in mangoes imported into the United States from Peru. Journal of Eco-
nomic Entomology 83, 19401943.
Sharp, J.L. and Spalding, D.H. (1984) Hot water as a quarantine treatment for Florida
mangos infested with Caribbean fruit y. Proceedings of the Florida State Horticul-
tural Society 97, 355357.
Sharp, J.L., Ouye, M.T., Thalman, R., Hart, W., Ingle, S. and Chew, V. (1988) Submersion
of Francis mango in hot water as a quarantine treatment for the West Indian fruit
y and the Caribbean fruit y (Diptera: Tephritidae). Journal of Economic Entomology
81, 14311436.
Sharp, J.L., Ouye, M.T., Ingle, S.J. and Hart, W.G. (1989a) Hot-water quarantine treat-
ment for mangoes from Mexico infested with Mexican fruit y and West Indian fruit
y (Diptera: Tephritidae). Journal of Economic Entomology 82, 16571662.
Sharp, J.L., Ouye, M.T., Ingle, S.J., Hart, W.G., Enkerlin, W.R.H., Hilario Celedonio, H.,
Jorge Toledo, A., Stevens, L., Quintero, E., Reyes, J.F. and Schwarz, A. (1989b) Hot-
water quarantine treatment for mangoes from the state of Chiapas, Mexico, infested
with Mediterranean fruit y and Anastrepha serpentina (Wiedemann) (Diptera:
Tephritidae). Journal of Economic Entomology 82, 16631666.
Shorter, A. and Joyce, D.C. (1994) Effect of surface coatings, on sap burn of Kensington
Pride mango fruit. Tropical Agriculture 71, 243246.
Shukla, R.P. and Tandon, P.L. (1985) Bio-ecology and management of the mango weevil,
Sternocbetus mangiferae (Fabricius) (Coleoptera: Curculionidae). International
Journal of Tropical Agriculture 3, 293303.
Simmons, S.L. (1998) The effects of preharvest factors on postharvest quality of Kensing-
ton Pride mango (Mangifera indica L.). PhD thesis, University of Queensland, Bris-
bane, Queensland.
Simmons, S.L., Hofman, P.J. and Hetherington, S.E. (1995) The effects of water stress on
mango fruit quality. Proceedings of the Mango 2000 Production workshop,
191197.
Simmons, S.L., Hofman, P.J., Whiley, A.W. and Hetherington, S.E. (1998) Effects of
leaf:fruit ratios on fruit growth, mineral concentration and quality of mango
(Mangifera indica L. cv. Kensington Pride). Journal of Horticultural Science and
Biotechnology 73, 367374.
Singh, R.N., Singh, G., Mishra, J.S. and Rao, O. (1987) Studies on the effect of pre- and
post-harvest treatment of calcium nitrate and calcium chloride on the storage life of
Amrapali mango. Progressive Horticulture 19, 19.
Singh, Z. and Janes, J. (2001) Effects of postharvest application of ethephon on fruit rip-
ening, quality and shelf life of mango under modied atmosphere packaging. Acta
Horticulturae 553, 599602.
Singh, Z., Janes, J. and Tan, S.C. (2000) Effects of different surfactants on calcium uptake
and its effects on fruit ripening, quality and postharvest storage of mango under
modied atmosphere packaging. Acta Horticulturae 509, 413417.
Singh, Z., Janes, J. and Suresh, N. (2001) Packaging materials affect physiological weight
loss, fruit colour and quality of mango during storage. Acta Horticulturae 553, 603
604.
Postharvest Technology 601
Singh, Z., Lalel, H.J.D. and Suresh, N. (2004) A review of mango fruit aroma volatile
compounds state of the art research. Acta Horticulturae 645, 519527.
Siriphanich, J. (1994) Minimal processing of tropical fruits. In: Champ, B.R., Highley, E.
and Johnson, G.I. (eds) Postharvest Handling of Tropical Fruits. Australian Centre
for International Agricultural Research (ACIAR) Proceedings 50. ACIAR, Canberra,
pp. 127137.
Sivapalasingam, S., Barrett, E., Kimura, A., Van Duyne, S., De Witt, W., Ying, M., Frisch,
A., Phan, Q., Gould, E., Shillam, P., Reddy, V., Cooper, T., Hoekstra, M., Higgins, C.,
Sanders, J.P., Tauxe, R.V. and Slutsker, L. (2003) A multistate outbreak of serotype
newport infection linked to mango consumption: impact of water-dip disinfestation
technology. Clinical Infectious Diseases 37, 15851590.
Slippers, B., Johnson, G.I., Crous, P.W., Coutinho, T.A., Wingeld, B.D. and Wingeld,
M.J. (2005) Phylogenetic and morphological re-evaluation of the Botryosphaeria
species causing diseases of Mangifera indica. Mycologia 97, 102113.
Smillie, R.M., Hetherington, S.E., Nott, R., Chaplin, G.R. and Wade, N.L. (1987) Appli-
cations of chlorophyll uorescence to the postharvest physiology and storage of
mango and banana fruit and the chilling tolerance of mango cultivars. ASEAN Food
Journal 3, 5559.
Smith, E.S.C. (1992) Fruit y disinfestation in mangoes by hot water dipping. In: Ooi,
P.A.C., Lim, G.S. and Teng, P.S. (eds) Proceedings of the Third International Confer-
ence on Plant Protection in the Tropics. Malaysian Plant Protection Society No. 6.
Malaysian Plant Protection Society, Kuala Lumpur, pp. 188195.
Smith, T.E., Hofman, P.J., Stephenson, R.A., Asher, C.J. and Hetherington, S.E. (1997)
Improving boron nutrition improves Hass avocado fruit size and quality. In: Cutting,
J. (ed.) Conference 97: Searching for Quality. Joint Conference of the Australian Avo-
cado Growers Federation and the New Zealand Avocado Growers Association.
Rotorua, New Zealand, pp. 131137.
Snowdon, A. (1990) A Colour Atlas of Post-harvest Disease and Disorders of Fruits and
Vegetables, Vol. 1. Fruits and General Introduction. Wolfe Scientic, London.
Snowdon, A. (1994) Diagnosing the causes of outturn problems in imported tropical
fruits. In: Champ, B.R., Highley, E. and Johnson, G.I. (eds) Postharvest Handling of
Tropical Fruits. Australian Centre for International Agricultural Research (ACIAR)
Proceedings 50. ACIAR, Canberra, pp. 94101.
Sonneveld, C. (2006) Measures to assure better food safety, marketing, and consumer
satisfaction in fruit and vegetables. In: Postharvest Management of Fruit and Vege-
tables in the Asia-Pacic Region Reports of the Asian Productivity Organisation
(APO) Seminar on Reduction of Postharvest Losses of Fruit and Vegetables held in
India; and Marketing and Food Safety: Challenges in Postharvest Management of
Agricultural/Horticultural Products in Islamic Republic of Iran. Asian Productivity
Organisation, pp. 8499. Available at: http://www.apo-tokyo.org/00e-books/AG-
18_PostHarvest.htm (accessed 1 May 2008).
Sornsrivichai, J., Anusadorn, P., Oogaki, C. and Gemma, H. (1989) Storage life and qual-
ity of mango (Mangifera indica L. cv. Keaw Sawoey) fruits stored in seal-packaging
by plastic lms and under low pressure at different temperatures. Japanese Journal
of Tropical Agriculture 33, 617.
Spalding, D.H., Benschoter, C.A., von Windeguth, D.L., King, J.R., Reeder, W.F. and
Burditt, A.K., Jr (1977) Methyl bromide and phosphine fumigation injury to avo-
cados and mangos. Proceedings of the Florida State Horticultural Society 90,
268270.
Stephens, B.E. and Tanner, D.J. (2005) The harvest watch system measuring fruits
healthy glow. Acta Horticulturae 687, 363364.
G.I. Johnson and P.J. Hofman 602
Stovolt, G.E. and Dirou, G.E. (2004) Blight Disease in Mangoes. Agnote, DPI-489 First
Edition 2004. Agdex 217/10. New South Wales Department of Primary Industries,
New South Wales.
Subedi, P.P., Walsh, K.B. and Owens, G. (2007) Prediction of mango eating quality at
harvest using short-wave near infrared spectrometry. Postharvest Biology and Tech-
nology 43, 326334.
Subramanayam, H., Krishnamurthy, S., Subhadra, N.V. and Dalal, V. (1980) Studies on
internal breakdown, a physiological ripening disorder in Alphonso mangoes
(Mangifera indica L.). Tropical Science 13, 203209.
Suhaila, M. and Halim, M.T. (1994) Anti-fruit-y activity of extracts of black pepper and
other edible plants. In: Champ, B.R., Highley, E. and Johnson, G.I. (eds) Postharvest
Handling of Tropical Fruits. Australian Centre for International Agricultural Re-
search (ACIAR) Proceedings 50. ACIAR, Canberra, pp. 371372.
Sunagawa, K., Kume, K. and Lwaizumi, R. (1987) The effectiveness of vapour heat treat-
ment against the melon y, Dacus cucurbitae Coquillett, in mango and fruit toler-
ance to the treatment. Research Bulletin of Plant Protection Japan 23, 1320.
Sunarharum, W.B. (2007) Identication of key volatiles responsible for the unique aromas
and avours of different mango varieties. MSc thesis, University of Queensland, Bris-
bane, Queensland.
Sundravadana, S., Alice, D., Kuttalam, S. and Samiyappan, R. (2006) Control of mango
anthracnose by azoxystrobin. Tunisian Journal of Plant Protection 1, 109114.
Sundravadana, S., Alice, D., Kuttalam, S. and Samiyappan, R. (2007) Efcacy of azox-
ystrobin on Colletotrichum gloeosporiodes Penz. growth and on controlling mango
anthracnose. Journal of Agricultural and Biological Science 2, 1015.
Suresh, N., Singh, Z. and Tan, S.C. (2004) Chilling injury in relation to ethylene biosyn-
thesis in Kensington Pride mango fruit. Journal of Horticultural Science and Bio-
technology 79, 8290.
Suslow, T. (2000) Postharvest Handling for Organic Crops. University of California Pub-
lication 7254. University of California, Davis, California.
Swaine, G., Melksham, K.J. and Corcoran, R.J. (1984) Dimethoate dipping of Kensing-
ton mango against Queensland fruit y. Australian Journal of Experimental Agricul-
ture and Animal Husbandry 14, 620623.
Taylor, R.W.D., Devereau, A.D. and Golob, P. (2002) Remedial treatments in pest man-
agement. In: Golob, P., Farrell, G. and Orchard, J.E. (eds) Crop Post-harvest: Sci-
ence and Technology, Vol. 1: Principles and Practice. Blackwell Science, Oxford,
UK, pp. 321359.
Terry, L.A. and Joyce, D.C. (2004) Elicitors of induced disease resistance in postharvest
horticultural crops: a brief review. Postharvest Biology and Technology 32, 113.
Thomas, A.C. (1975) The Application of Ionizing Radiation to the Shelf Life Extension of
Mangoes in South Africa. Republic of South Africa Atomic Energy Board, Pelinada-
ba, Pretoria, PEL-244, South Africa.
Thomas, P. (1986) Radiation preservation of foods of plant origin. III. Tropical fruits: banana,
mangoes, and papayas. Critical Reviews in Food Science and Nutrition 23, 147206.
Thomas, P. and Joshi, M.R. (1988) Reduction in chilling injury in ripe Alphonso mango
fruit in cold storage by temperature conditioning. International Journal of Food Sci-
ence and Technology 23, 447455.
Thomas, P. and Oke, M.S. (1983) Improvement in quality and storage of Alphonso
mangoes by cold adaptation. Scientia Horticulturae 19, 257262.
Thomas, P., Kannan, A., Degwekar, V.H. and Ramamurthy, M.S. (1995) Non-destructive
detection of seed weevil-infested mango fruits by X-ray imaging. Postharvest Biol-
ogy and Technology 5, 161165.
Postharvest Technology 603
Thompson, A.K. (1998) Controlled Atmosphere Storage of Fruit and Vegetables. CAB
International, Wallingford, UK.
Thompson, J.F. (2002) Tranportation. In: Kader, A.A. (ed.) Postharvest Technology of
Horticultural Crops, 3rd edn. University of California, Oakland, California, pp. 259
269.
Thompson, J.F., Mitchell, F.G. and Kasmire, R.F. (2002) Cooling horticultural commodi-
ties. In: Kader, A.A. (ed.) Postharvest Technology of Horticultural Crops, 3rd edn.
University of California, Oakland, California, pp. 97112.
Timmer, L.W. and Zitko, S.E. (1996) Evaluation of copper fungicides and rates of metallic
copper for control of Melanose on grapefruit in Florida. Plant Disease 80, 166169.
Torres, J.D., Talens, P., Carot, J.M., Chiralt, A. and Escrichea, I. (2007) Volatile prole of
mango (Mangifera indica L.) as affected by osmotic dehydration. Food Chemistry
101, 219228.
Truter, A.B. and Eksteen, G.J. (1987) Controlled and modied atmospheres to extend
storage life of avocados. South African Avocado Growers Association Yearbook 10,
151153.
Underhill, S. and Dahler, J. (1995) Classifying skin browning. Mango Care 14, 67.
Van der Linde, H.J. and Thord-Gray, R.S. (1986) Irradiation as a post harvest treatment
for mangoes. South African Mango Growers Association Yearbook 6, 2426.
Van Straten, B. and Oosthuyse, S.A. (1994) Die effek van koelopberging by 4 of 8C op
die tempo van kwalititeit verslegting van ryp Sensation mango vrugte. South Afri-
can Mango Growers Association Yearbook 14, 3436.
Varith, J., Sirikajornjaru, W. and Kiatsiriroat, T. (2007) Microwave-vapor heat disinfesta-
tion on oriental fruit y eggs in mangoes. Journal of Food Processing and Preserva-
tion 31, 253269.
Velasco, L.R.I. and Medina, C. (2004) Soft X-ray imaging for non-destructive detection
of mango pulp weevil (Sternochetus frigidus Fabr.) infestation in fresh mature green
Carabao mango (Mangifera indica L.) fruits. The Philippine Agricultural Scientist
87, 160164.
Vijaysegaran, S. (1994) Preharvest fruit y control: strategies for the tropics. In: Champ,
B.R., Highley, E. and Johnson, G.I. (eds) Postharvest Handling of Tropical Fruits.
Australian Centre for International Agricultural Research (ACIAR) Proceedings 50.
ACIAR, Canberra, pp. 288303.
Vinning, G.S. and Young, J. (2006) Linking production and marketing of fruit and vegeta-
bles for better farm incomes in the Asia-Pacic region. In: Postharvest Management
of Fruit and Vegetables in the Asia-Pacic Region Reports of the Asian Productivity
Organisation (APO) Seminar on Reduction of Postharvest Losses of Fruit and Veg-
etables held in India; and Marketing and Food Safety: Challenges in Postharvest
Management of Agricultural/Horticultural Products in Islamic Republic of Iran. Asian
Productivity Organisation, pp. 5369. Available at: http://www.apo-tokyo.org/00e-
books/AG-18_PostHarvest.htm (accessed 1 May 2008).
von Windeguth, D.L. (1986) Gamma irradiation as a quarantine treatment for Caribbean
fruit y infested mangos. Proceedings of the Florida State Horticultural Society 99,
131134.
Wagner, L.J. and Flores, H.E. (1994) Effect of taxol and related compounds on growth of
plant pathogenic fungi. Phytopathology 84, 11731178.
Wainwright, H. and Burbage, M.B. (1989) Physiological disorders in mango (Mangifera
indica L.) fruit. Journal of Horticultural Science 64, 125135.
Waite, G. (2002) Pests and pollinators of mango. In: Pea, J.E., Sharp, J.L. and Mysoki,
M. (eds) Fruit Pests and Pollinators. Biology, Economic Importance, Natural Enemies
and Control. CAB International, Wallingford, UK, pp. 103130.
G.I. Johnson and P.J. Hofman 604
Walker, K. (2007a) Mango Pulp Weevil (Sternochetus frigidus) Pest and Diseases Image
Library. Available at: http://www.padil.gov.au (accessed 2 April 2008).
Walker, K. (2007b) Red Banded Mango Caterpillar (Deanolis sublimbalis). Pest and Diseases
Image Library. Available at: http://www.padil.gov.au (accessed 31 October 2007).
Wall, M. (2008) Quality of postharvest horticultural crops after irradiation treatment.
Stewart Postharvest Review 2008, 2, Article 1.
Wang, B., Wang, J., Liang, H., Yi, J., Zhang, J., Lin, L., Wu, Y., Feng, X., Cao, J. and Jiang,
W. (2008) Reduced chilling injury in mango fruit by 2,4-dichlorophenoxyacetic acid
and the antioxidant response. Postharvest Biology and Technology 48, 172181.
Watada, A.E., Herner, R.C., Kader, A.A. and Romani, R.J. (1984) Terminology for the de-
scription of developmental stages of horticultural crops. HortScience 19, 2021.
Wells, I. and Littlemore, J. (1989) Dipping mangoes, watch the Benlate concentration.
Queensland Fruit and Vegetable News 60, 18.
Whiley, A.W. (1999) Investigation into the effect of calcium on the fruit disorder stem
end cavity in the mango cultivar Kensington Pride. In: Whiley, A.W. (ed.) Flower-
ing Behaviour and Subsequent Productivity in Mango. Final Report for Project no.
9012. Australian Centre for International Agricultural Research (ACIAR), Canberra.
Whiley, A.W. and Hofman, P.J. (2007) Development of Best Practice Pre- and Post-
harvest Protocols for Production of Calypso mango. Final Report for Project
FR02049. Horticulture Australia Ltd, Sydney.
Wikipedia (2007) Good Agricultural Practice. Available at: http://en.wikipedia.org/wiki/
Good_Agricultural_Practices (accessed 31 October 2007).
Willingham, S.L., Coates, L.M., Cooke, A.W., Dean, J.R. and Beasley, D.R. (1999) Field
management of mango postharvest pathogens. In: Asia-Pacic Plant Pathology for
the New Millenium. 12th Biennial Australasian Plant Pathology Conference. Can-
berra, p. 138.
Wills, R.B.H., Yuen, M.C.C., Sabari, Laksmi, L.D.S. and Suyanti (1988) Effect of calcium
inltration on delayed ripening of three mango cultivars in Indonesia. ASEAN Food
Journal 4, 6768.
Wilson, C.L. and Pusey, P.L. (1985) Potential for biological control of postharvest plant
diseases. Plant Disease 69, 375378.
Wilson, C.L., El Ghaouth, A., Chalutz, E., Droby, S., Stevens, C., Lu, J.Y., Khan, V. and
Arul, J. (1994) Potential of induced resistance to control postharvest diseases of
fruits and vegetables. Plant Disease 78, 837844.
Winston, E. (1986) Observations of internal mango esh breakdown: need for standard-
ization of terminology. In: Proceedings of the First Australian Mango Research
Workshop. Commonwealth Scientic and Industrial Research Organization
(CSIRO), Melbourne, pp. 7782.
Wittenberg, L. (2007) Mapping mango seed weevil. In: Delivering Mango Research. The
Sixth Amistar Australian Mango Conference. Gold Coast, Queensland, pp. 3133.
Wolstenholme, B.N. (2004) Nitrogen the manipulator element: managing inputs and
outputs in different enviroments. South African Avocado Growers Association
Yearbook 27, 6278.
World Trade Organisation (WTO) (2008) World Trade Organisation Technical Barriers to
Trade. Available at: http://www.wto.org/english/tratop_e/tbt_e/tbt_e.htm (accessed
4 April 2008).
Wright, M.A. and Stringer, A. (1973) The toxicity of thiabendazole, benomyl, methyl
benzimidazol-2-yl carbamate and thiophanate-methyl to the earthworm, Lumbri-
cus terrestris. Pesticide Science 4, 431432.
Yahia, E.M. (1994) The potential use of insecticidal atmospheres for mango, avocado,
and papaya fruits. In: Champ, B.R., Highley, E. and Johnson, G.I. (eds) Postharvest
Postharvest Technology 605
Handling of Tropical Fruits. Australian Centre for International Agricultural Re-
search (ACIAR) Proceedings No. 50. ACIAR, Canberra, pp. 373374.
Yahia, E.M. (2006) Modied and controlled atmospheres for tropical fruits. Stewart Post-
harvest Review 2006(5), article 6.
Yahia, E.M. and Tiznando-Hernandez, M. (1993) Tolerance and responses of harvested
mango to insecticidal low oxygen atmospheres. HortScience 28, 10311033.
Yahia, E.M. and Vazquez-Moreno, L. (1993) Responses of mango to insecticidal oxygen
and carbon dioxide atmospheres. Food Science and Technology 26, 4248.
Yarrow, R. and Chandler, L. (2007) Informal testing of the RBMC pheromone over early
mango fruiting season at Lockerbie Station via Bamaga, Cape York Peninsula. In:
Delivering Mango Research. The Sixth Amistar Australian Mango Conference. Gold
Coast, Queensland, pp. 2425.
Yashoda, H.M., Prabha, T.N. and Tharanathan, R.N. (2006) Mango ripening: changes in
cell wall constituents in relation to textural softening. Journal of the Science of Food
and Agriculture 86, 713721.
Yogaratnam, N. and Johnson, D.S. (1982) The application of foliar sprays containing ni-
trogen, magnesium, zinc and boron to apple trees. Journal of Horticultural Science
57, 159164.
Young, T.W. (1957) Soft-nose, a physiological disorder in mango fruits. Proceedings of
the Florida State Horticulture Society 70, 280283.
Young, T.W. and Miner, J.T (1961) Relationship of nitrogen and calcium to soft-nose
disorder in mango fruit. Proceedings of the American Society of Horticultural Sci-
ence 58, 201208.
Yuen, C.M.C., Tan, S.C., Joyce, D. and Chettri, P. (1993) Effect of postharvest calcium
and polymeric lms on ripening and peel injury in Kensington Pride mango. ASE-
AN Food Journal 8, 110113.
Zainuri (2006) Defence mechanisms and induced resistance in Kensington Pride man-
go. PhD thesis, University of Queensland, Brisbane, Queensland.
Zainuri, Wearing, A.H., Coates, L. and Terry, L. (2001) Effects of phosphonate and salicylic
acid treatments on anthracnose disease development and ripening of Kensington
Pride mango fruit. Australian Journal of Experimental Agriculture 41, 805813.
Zainuri, Irving, D.E., Dann, E.K., Coates, L.M. and Wearing, A.H. (2003) Activating man-
go fruit defence to anthracnose disease. In: Australian Postharvest Horticulture
Conference. Brisbane, Queensland, pp. 149150.
Zeng, K. and Waibo, J. (2005) Induced resistance to anthracnose (Colletotrichum gloeo-
sporioides) in mango by postharvest treatment with salicylic acid. Journal of China
Agricultural University 10, 3640.
Zeng, K.F., Cao, J.K. and Jiang, W. (2006) Enhancing disease resistance in harvested
mango (Mangifera indica L.) fruit by salicylic acid. Journal of the Science of Food
and Agriculture 85, 694698.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
606 (ed. R.E. Litz)
16 World Mango Trade and the Economics
of Mango Production
E.A. Evans and O.J. Mendoza
University of Florida, Florida, USA
16.1 Introduction 606
16.2 Recent Trends in World and USA Mango Production, Trade and Consumption 607
World situation 607
USA mango production, imports and consumption 611
16.3 Sample Costs and Returns Associated with the Establishment and Production of
Mango Orchards 613
General approach to estimating cost of production of orchard crops 614
Main assumptions 614
Discussion of establishment phase budget 617
Discussion of production phase budget 621
Protability analysis 624
16.4 Conclusions 626
16.1 Introduction
Worldwide mango production occurs in over 90 countries. Although only a
relatively small proportion of total mango production enters international
trade (<4%), the volume traded has increased substantially since the late
1990s. Among the factors responsible for the increased mango production,
trade and consumption are lower prices, year-round availability, fewer
horticultural trade barriers, changes in food consumption preferences,
longer shelf life for perishables and consumer interest in healthier foods.
Although not a major mango producer, the USA has developed most of the
popular cultivars traded on the international market, and is the largest
single-country mango importer. The costs and returns and general practices
of establishing and maintaining orchards vary considerably from coun-
try to country and within each country (different regions and production
systems).
World Mango Trade and Economics 607
16.2 Recent Trends in World and USA Mango Production, Trade and
Consumption
World situation
Asia accounts for approximately 77% of global mango production, and the
Americas and Africa account for approximately 13% and 9%, respectively
(FAOSTAT, 2007). In 2005, world production of mango was estimated to have
reached 28.51 million t, an increase from the 27.82 million t recorded in the
previous year. Between 1996 and 2005, production grew at an average annual
rate of 2.6%. Table 16.1 shows the worlds top ten mango-producing coun-
tries, which account for about 85% of the worlds production.
India is the largest producer, accounting for 38.58% of global production
from 2003 to 2005. During that period, the Indian mango crop averaged
10.79 million t, followed by China and Thailand at 3.61 million t (12.90%) and
1.73 million t (6.20%), respectively. Other leading mango-producing coun-
tries and their respective shares of world production during the 20032005
period include Mexico (5.50%), Indonesia (5.29%), Pakistan (4.48%), Brazil
(4.30%), the Philippines (3.53%), Nigeria (2.61%) and Egypt (1.28%).
Although currently only 3.3% of the world production of mango is traded
globally, this represents a noticeable increase over the quantities traded since
the late 1980s. In terms of distribution, Mexico, Brazil, Peru, Ecuador and
Haiti supply the majority of North Americas imports. India and Pakistan are
the predominant suppliers to the West Asian market. South-east Asian coun-
tries get most of their supplies from the Philippines and Thailand. European
Union (EU) buyers source mango mainly from South America and Asia.
In 2005, global exports of mango reached 912,853 t, a slight decrease of
0.73% compared with the previous year, and were valued at US$543,100,000
(FAOSTAT, 2007). Table 16.2 shows the top ten major mango-exporting coun-
tries. India is the largest producer but only recently has overtaken Mexico as
the number one exporter of the fruit. For the 20032005 period, Mexico and
India dominated the export trade with shares of 22.64% and 20.25%, respec-
tively, followed by Brazil (13.18%) and Pakistan (6.94%). Other major export-
ers include the Netherlands (major re-exporter), Peru, Ecuador, the Philippines,
Thailand and China.
World imports of mango increased from 397,623 t in 1996 to 826,584 t in
2005. The USA is the number one importer of mango. During the 20032005
period, the USA imported 271,848 t, or approximately a third of the total
mango imports (Table 16.3).
The Netherlands imported 88,300 t of mangoes (10.62%), although most of it
is redistributed throughout the EU. Other prominent importing countries that
are also major redistributors are the United Arab Emirates (6.82%) and Saudi
Arabia (5.32%). Most of these imports are redistributed to other countries within
the Middle East. Although China (4.91%) appears as a major importer, the quan-
tities imported have been declining. For example, China imported 57,000 t in
2004 and only 19,000 t in 2005. This could be due to increases in domestic pro-
duction in response to an increase in domestic demand driven by rising per
E
.
A
.

E
v
a
n
s

a
n
d

O
.
J
.

M
e
n
d
o
z
a
6
0
8
Table 16.1. Worlds ten major mango producers, 19962005 (1000 t) (Source: FAOSTAT, 2007).
Countries 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 20032005 (%)
India 11,000 11,000 10,230 9,780 10,500 10,060 10,640 10,780 10,800 10,800 38.58
China 2,074 2,410 2,562 3,127 3,211 3,273 3,513 3,571 3,582 3,673 12.90
Thailand 1,181 1,198 1,088 1,462 1,633 1,700 1,700 1,700 1,700 1,800 6.20
Mexico 1,189 1,500 1,474 1,508 1,559 1,577 1,523 1,362 1,573 1,679 5.50
Indonesia 783 1,088 600 827 876 923 1,403 1,526 1,438 1,478 5.29
Pakistan 908 914 917 916 938 990 1,037 1,035 1,056 1,674 4.48
Brazil 593 508 469 456 538 782 842 1,254 1,358 1,000 4.30
Philippines 898 1,005 945 866 848 882 956 1,006 968 985 3.53
Nigeria 656 689 731 729 730 730 730 730 730 730 2.61
Egypt 203 231 223 287 299 325 287 319 375 380 1.28
Others 3,248 3,230 3,347 3,656 3,597 3,731 4,001 4,327 4,242 4,308 15.34
World total 22,733 23,773 22,584 23,615 24,730 24,973 26,634 27,609 27,822 28,508 100.00
W
o
r
l
d

M
a
n
g
o

T
r
a
d
e

a
n
d

E
c
o
n
o
m
i
c
s
6
0
9
Table 16.2. Worlds ten major mango-exporting countries, 19962005 (1000 t) (Source: FAOSTAT, 2007).
Countries 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 20032005 (%)
Mexico 148 187 209 204 207 195 195 216 213 195 22.64
India 27 45 47 38 39 46 42 179 156 223 20.25
Brazil 24 23 39 54 67 94 104 138 111 114 13.18
Pakistan 18 25 39 41 48 52 48 60 82 49 6.94
Netherlands 21 25 17 37 34 43 33 58 51 69 6.42
Peru 11 6 11 20 21 27 35 40 60 58 5.71
Ecuador 0 2 7 0 26 34 30 38 41 40 4.31
Philippines 40 45 53 35 40 39 36 38 36 25 3.61
Thailand 8 9 10 10 9 11 9 8 33 2 1.55
China 12 7 9 10 5 5 15 22 10 4 1.31
Others 80 104 87 103 132 121 127 126 127 135 14.08
World total 391 478 529 552 628 666 673 923 920 913 100.00
E
.
A
.

E
v
a
n
s

a
n
d

O
.
J
.

M
e
n
d
o
z
a
6
1
0
Table 16.3. Worlds top ten major mango-importing countries, 19962005 (1000 t) (Source: FAOSTAT, 2007).
Countries 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 20032005 (%)
USA 171 187 197 219 235 238 263 278 276 261 32.70
Netherlands 25 34 35 63 62 70 71 91 76 98 10.62
United Arab Emirates 28 37 48 48 42 46 52 62 58 51 6.82
Saudi Arabia 10 16 14 9 28 36 35 40 42 51 5.32
China 36 40 47 33 33 34 38 47 57 19 4.91
Bangladesh 5 9 0 11 21 21 14 43 37 36 4.63
UK 16 18 18 23 22 27 24 32 37 47 4.63
Germany 13 17 17 24 23 25 28 32 33 37 4.11
France 18 23 22 31 26 26 27 32 35 35 4.09
Malaysia 14 6 21 1 20 27 31 26 45 19 3.59
Others 61 68 66 84 114 106 101 142 148 173 18.58
World total 398 454 486 545 628 656 684 825 843 827 100.00
World Mango Trade and Economics 611
capita income. Other noticeable importers include Bangladesh and the UK
(4.63% each), Germany (4.11%), France (4.09%) and Malaysia (3.59%).
The most popular export mango cultivars continue to be Kent, Tommy
Atkins, Haden and Keitt. These cultivars have fruit with a red blush and are
less brous, rmer and more suited for long-distance transportation. The green
cultivars are only now being widely accepted in the international market and
include cultivars such as Ataulfo and Amelie. Other cultivars which are gain-
ing signicance in international trade include: Alphonso, Dudhpeda, Kesar,
Sindhu, Pairi, Desi, Chausa, Langra and Katchamita. Most of these newer
cultivars on the international scene are coming from India and Pakistan.
Since the late 1990s, per capita consumption has increased noticeably in
the USA, Japan and China, mainly due to higher income levels, improved
advertising and lower mango prices. On the international scene, prices for
most mango varieties have declined considerably over the decade, dropping
about 50%. For example, the average price for mango in the European mar-
ket was US$12/kg in 1996, compared to just over US$6/kg today. The reduc-
tion in price is partly due to the increased availability of tropical fruits. There
is consensus, however, that prices have stabilized but could increase with
proper promotional efforts.
Although the quantity of processed mango fruit that is traded internation-
ally is small compared with the fresh fruit trade (<7%) there is evidence to
suggest that it is increasing. Such products include mango juice, pickled man-
goes, mango chutney, mango pulp and paste, mango pure, dried fruit, mango
slices in brine and mango our. India is the main exporter of processed mango
followed by Pakistan, Brazil and Zimbabwe. Major importers include the
United Arab Emirates, Saudi Arabia, Kuwait, USA, UK and Canada.
USA mango production, imports and consumption
The USA is not a major mango producer even though most of the commer-
cially traded varieties have been developed in Florida. USA production,
mainly in Florida, remains fairly stable at a little under 3000 t/year.
The USA is currently the worlds leading importer of fresh mangoes,
accounting for 32.70% of the total imports during the 20032005 period
(FAOSTAT, 2007). Figure 16.1 shows the trend of mango imports into the
USA between 1997 and 2006. Overall, the graph indicates a steady increase
in the volume of mango imports. Between 1997 and 2006, imports increased
from 187,193 t to 298,088 t, an average annual growth rate of 5.46%. Mango
imports were valued at about US$233,100,000 in 2006 (USDA, Foreign Agri-
cultural Service, 2007).
The main sources of USA imports of mango are Mexico, Peru, Ecuador
and Brazil. Mexico is the main supplier of mango to the USA (60.78% share
in 2006) (Fig. 16.2). In recent years, Brazil, Peru and Ecuador have become
signicant exporters to the USA, competing with Mexico at the beginning
and the end of the season. The USA exports very few of its mango imports,
mainly to Canada and the UK.
E.A. Evans and O.J. Mendoza 612
USA consumption of mango has increased steadily, from a per capita
level of 0.5 kg in 1996 to 1 kg in 2005 (USDA, Economic Research Service,
2007). The growth in USA consumption of mango is driven by many factors,
such as year-round availability, lower prices, consumer preferences and more
0
50,000
100,000
150,000
200,000
250,000
300,000
350,000
1997 1998 1999 2000 2001 2002 2003 2004 2005 2006
Year
U
S
A

t
o
t
a
l

i
m
p
o
r
t
s

(
t
)
Fig. 16.1. USA total imports of mango (t), 19972006 (Source: USDA Foreign
Agricultural Service, 2007).
0
20,000
40,000
60,000
80,000
100,000
120,000
140,000
160,000
180,000
200,000
U
S
A

t
o
t
a
l

i
m
p
o
r
t
s

(
t
)
1997 1998 1999 2000 2001 2002 2003 2004 2005 2006
Year
Mexico Peru Ecuador Brazil
Fig. 16.2. USA total imports of mango (t) by country, 19972006 (Source: USDA
Foreign Agricultural Service, 2007).
World Mango Trade and Economics 613
disposable income. However, mango consumption levels are still considered
relatively low when compared to other fruits, for example bananas (11 kg)
and oranges (5 kg).
USA prices for mango vary widely by cultivar and season, mainly due to
the fact that the commodity demand is price inelastic (sensitive to variations
in quantities available, a 1% increase in the quantity tends to lead to a >1%
fall in the price). In general, mango prices have been steadily declining over
the past decade. Table 16.4 shows the average cost, insurance and freight
(CIF) prices for mango imports into the USA from main supply sources dur-
ing the 19982006 period. Although prices have decreased noticeably from
the 1998 level, they do appear to have stabilized in the last couple of years.
16.3 Sample Costs and Returns Associated with the Establishment and
Production of Mango Orchards
In this section, we have estimated the per hectare costs to establish an 18 ha
commercial mango orchard in Florida USA, as well as the annual estimated
costs and net returns per hectare after establishment. We have not provided
too much detail on production practices (since such information is readily
available elsewhere), but rather focus on the methodology of estimating the
costs and returns of establishment and production. Although somewhat hypo-
thetical, the data presented are based on a combination of information
obtained from the growers, economic engineering using recommended prac-
tices, and discussions with industry experts. We begin by briey describing
the general methodology, followed by a listing of some of the major assumptions
used in the analysis. Estimates of the establishment costs are then discussed,
and the chapter closes with a discussion of protability of a mango operation.
Table 16.4. Average cost, insurance and freight (CIF) prices for selected varieties from
main suppliers to the USA, 19982006 (US$/kg) (Source: USDA Agricultural Marketing
Service, 2007).
Year
Country of origin
Year average Brazil Ecuador Haiti Peru Mexico
1998 3.43 3.21 n/a
a
3.61 2.09 3.09
1999 2.13 1.67 2.24 1.89 1.78 1.94
2000 2.09 1.69 2.05 1.65 1.72 1.84
2001 1.74 1.65 2.24 1.85 1.69 1.83
2002 1.67 1.47 2.13 1.61 1.61 1.70
2003 1.72 1.28 1.96 1.45 1.45 1.57
2004 1.65 1.83 1.98 1.43 1.43 1.66
2005 1.67 1.94 2.11 1.58 1.67 1.79
2006 1.65 1.67 2.11 1.39 1.72 1.71
a
n/a, not available.
E.A. Evans and O.J. Mendoza 614
General approach to estimating cost of production of orchard crops
The general approach for estimating the cost of production of perennial crops
(orchards and vineyards), which usually take more than a year to begin pro-
duction, is to develop two separate budgets: one for the establishment
phase and the other for the production phase. The establishment phase bud-
get reects the sum total of all expenses (expressed on a yearly and per unit
basis) that are incurred over the years to bring an orchard into meaningful
(mature trees) production. Once this amount is determined, it is treated as if
it were an expense incurred in the purchase of a capital item, for example
machinery to be used in the production phase of the orchard. As in the case
of a capital item, an equal annual amount (amortized amount) is charged to
the production operation as an expense spread over the estimated future life
of the orchard. In other words, the amortized amount is included as a non-
cash overhead expenditure in the production phase budget when determin-
ing net returns from the enterprise. The production phase budget estimates
the costs and returns on an annual per unit basis associated with the mainte-
nance of the orchard after it has been established. Although the procedure
seems daunting, it is made much easier by using spreadsheet software with
built-in formulas for calculating amortization.
Main assumptions
The following assumptions were used to estimate production costs and
returns for 18 ha operations.
Land
The hypothetical farm consists of 20 ha. A mango orchard is being estab-
lished on 18 ha, with roads, irrigation system and farmstead occupying 2 ha.
The 18 ha orchard is considered large enough to use machinery and equip-
ment efciently. The orchard is farmed by the owner with the help of hired
part-time labour.
Site preparation
It is assumed that the land is relatively clear, with no costs being included for
major land preparation such as timber clearing, rock removal or land level-
ling. However, if these operations are required, they should be included. The
main operations considered here are associated with fencing and road con-
struction for travelling and harvesting. These operations are usually done 1
year prior to planting, but costs are shown in the rst year.
Planting, training and pruning
Trees are planted on 7.5 m 7.5 m spacing, or 175 trees/ha. The life of the
orchard in this study is projected to be 25 years. The variety considered in
this study is Tommy Atkins. Pruning and training begin in the third year,
and labour time required for pruning increases in the successive years.
World Mango Trade and Economics 615
Hedging and topping operations are carried out immediately after fruit har-
vesting.
Fertilization
During the rst 3 years, an N-P-K fertilizer (6% nitrogen) is spread by hand
six times each year. After year 3, the frequency of application decreases to
four times each year. Table 16.5 shows the annual fertilizer rates assumed. In
addition, the trees are given annual nutritional sprays of copper, zinc, man-
ganese and boron. Iron is applied in chelated form as a soil drench two to
three times each year.
Irrigation
Total irrigation costs include the cost of pumping water and irrigation labour.
Water for our irrigation system is supplied from a well; therefore, the cost of
the water is zero. Irrigation costs for individual orchards vary, depending on
the amount of water pumped, pumping system, energy source and irrigation
district.
Weed control management
Weeds in the tree rows are controlled with applied pre- and post-emergent
(residual) contact herbicides such as Roundup

. With ve applications each


year, spot sprays are applied at the rate of approximately 5 l/ha. However, as
the trees grow larger and shade reduces weed growth, the number of appli-
cations is reduced to approximately three per annum.
Harvest
Mango fruits are best harvested using clippers and hand-carried harvesting
bags (about US$0.11/kg). With large trees, fruits are harvested using picking
poles, with or without attached clippers, which are equipped with bags into
which the fruit falls. After harvesting, the fruits are usually piled under a tree
on the ground. The fruit is then loaded onto trucks in the eld and hauled to
packing houses for US$0.11/kg.
Yields and returns
A major assumption is that the orchard requires approximately 7 years to
reach maturity; hence the rst 6 years are considered the establishment phase.
Even though trees require about 7 years before reaching maturity, they will
start having saleable fruits from about the third year. Table 16.6 shows the
typical yield assumptions used in the analysis. Annual gross eld yields at
maturity are assumed to be 22,000 kg/ha.
Table 16.5. Annual fertilizer rates @ 6% N.
Year 1 2 3 4 5 6 7+
Rate (kg/ha) 236 472 707 786 1100 1257 1415
E.A. Evans and O.J. Mendoza 616
Labour
Hourly wages for workers are US$10.00 for skilled workers and US$7.00 for
eld workers. Adding 34% for the employers share of federal and state pay-
roll taxes, insurance and other possible benets, yields a labour rate of US$13/h
for skilled labour and US$9/h for eld labour.
Cash overhead
Cash overhead consists of numerous cash expenses paid during each year
that are assigned to the entire farm, not to a particular operation. These costs
include property taxes, ofce expenses, capital interest, liability and property
insurance, equipment repairs, sanitation services and crop insurance.
PROPERTY TAXES. In the USA, most counties charge a base property tax rate of
approximately 1% on the assessed value of the property.
INTEREST ON OPERATING CAPITAL. Interest on operating capital is based on cash
operating costs and is calculated at a nominal rate of 5%/year. A nominal
interest rate is the going market cost of borrowed funds.
INSURANCE. Insurance for farm investment differs depending on assets included
and amount of coverage. Property insurance provides coverage for property
loss and the standard practice in the USA according to the American Society
of Agricultural Engineers (ASAE) is to charge about 0.67% of the average
value of the assets over their useful life. Liability insurance covers accidents
on the farm (US$580/year).
OFFICE EXPENSES. Ofce and business operating expenses are estimated at
US$355/ha, and include ofce supplies, telephone, bookkeeping, account-
ing, legal fees, road maintenance and miscellaneous administrative charges.
SANITATION SERVICES. Sanitation services offer portable toilets for orchards and
cost the farm US$1900/year (US$106/ha). This cost includes delivery and
servicing of a single toilet and washing unit.
CROP INSURANCE. Multi-peril crop insurance is purchased at a cost of US$445/ha.
Non-cash overhead
Non-cash overhead is calculated as the capital recovery cost for equipment
and other farm investments. Capital recovery cost is the annual depreciation
Table 16.6. Yield assumptions.
Year 1 2 3 4 5 6 7+
Yield (kg/ha) 0 0 1,700 4,500 9,000 15,500 22,000
World Mango Trade and Economics 617
and interest costs for a capital investment. It is the amount of money required
each year to be set aside so that a capital item can be fully replaced at the end
of its useful life. It can be viewed as the value of the portion of the capital
item that was utilized in the production process during that year. The for-
mula for the calculation of the annual capital recovery costs is the PMT Excel
formula:
Capital Recovery = PMT (I, N, (PP SV)) + (I)*(SV)
Where:
PMT = the formula that calculates the payment for a loan based on con-
stant payments and a constant interest rate.
I = the interest rate used to represent the cost of borrowed capital (in this
case, 5%).
N = the number of years.
PP = the purchase price for the capital item.
SV = an estimate of the remaining value of an investment at the end of its
useful life. In this case, the percentage of remaining value is calculated
from equations developed by the ASAE based on equipment type and
years of life.
Farm equipment on a mango orchard in the region is purchased either
new or used. The study shows the current purchase price for new equip-
ment. The new purchase price is adjusted to 60% to indicate a mix of new and
used equipment.
The orchard is irrigated using a sprinkler irrigation system. The life of
the irrigation system is estimated at 20 years. The irrigation system is installed
before the orchard is planted and includes costs of a pumping system, instal-
lation labour and design and materials.
Although it is assumed that the grower owns the land, the going rental
rate in South Florida of approximately US$2500/ha is used to reect the
opportunity (economic) cost of land.
Discussion of establishment phase budget
Table 16.7 shows the layout of the establishment phase budget. The budget
reects the annual per hectare costs incurred in establishing a mango orchard.
Here it is assumed that a mango orchard reaches maturity in the seventh
year. As such, the budget reects all associated cash and non-cash expenses
incurred over 6 years.
The information required for each year is similar except for the rst 2 or
3 years. With reference to Table 16.7, year 1 includes pre-planting costs (site
preparation and orchard layout), which in this case amounts to US$5000/ha.
Most of the pre-planting costs most likely would occur in the year before, but
the practice is to include these costs in year 1. Planting costs include new
trees, digging holes for trees, setting trees, and wrapping and staking trees.
Both pre-planting and planting costs appear only in year 1. Total per hectare
E
.
A
.

E
v
a
n
s

a
n
d

O
.
J
.

M
e
n
d
o
z
a
6
1
8
Table 16.7. Sample costs to establish a mango orchard in south Florida (source: compiled by the authors).
Operation Unit
Year (US$/ha)
1st 2nd 3rd 4th 5th 6th
Pre-planting costs
Site preparation 4,500
Orchard layout 500
Total pre-planting costs 5,000
Planting costs
Mango trees (175 trees/ha) US$25.00 4,375
Digging, planting, wrapping and staking US$10.00 1,750
Total planting costs 6,125
Replanting costs
Replaced trees 5% 5% 225
Digging, planting, wrapping and staking
replaced trees
90
Total replanting costs 315
Cultural costs
Orchard pruning 0 0 0 260 260 260
Herbicide 100 100 100 100 100 100
Fungicide 0 0 0 1,360 1,360 1,360
Fertilizer 80 160 242 270 430 600
Insecticide 200 200 200 200 200 200
Mowing 390 390 240 180 180 180
Irrigate and walk lines 90 90 90 90 67 67
Pest control advisor 0 0 150 150 150 150
Pollination 0 0 0 0 450 450
Rodent control (squirrels) 60 60 60 60 60 60
Total cultural costs 920 1,000 1,082 2,670 3,257 3,427
Harvesting and marketing costs
Picking US$0.11/kg US$0.11 0 0 187 495 990 1,705
W
o
r
l
d

M
a
n
g
o

T
r
a
d
e

a
n
d

E
c
o
n
o
m
i
c
s
6
1
9
Packing and shipping US$0.11/kg US$0.11 0 0 187 495 990 1,705
Sales charge @ 10% of FOB
a
price 10% 0 0 153 405 810 1,395
Total harvesting, marketing and inspection
costs
0 0 527 1,395 2,790 4,805
Interest on operating capital @ 5% 5% 602 66 54 134 163 171
Total operating costs 12,647 1,381 1,663 4,199 6,210 8,403
Cash overhead costs
Liability insurance 30 30 30 30 30 30
Crop insurance 0 0 445 445 445 445
Sanitation fee 0 0 106 106 106 106
Ofce expenses 355 355 355 355 355 355
Property taxes 150 150 150 150 150 150
Property insurance 250 250 250 250 250 250
Investment repairs 185 190 210 210 210 210
Interest on operating capital (cash overhead) 5% 49 49 77 77 77 77
Total cash overhead costs 1,019 1,024 1,623 1,623 1,623 1,623
Total cash costs 13,666 2,405 3,286 5,822 7,833 10,027
Income from production 0 0 1,530 4,050 8,100 13,950
Net cash costs for the year 13,666 2,405 1,756 1,772 267 3,923
Accumulated net cash costs 13,666 16,070 17,827 19,598 19,332 15,408
Non-cash overhead
Land rent 2,500 2,500 2,500 2,500 2,500 2,500
Machinery and equipment 405 405 405 405 405 405
Building 185 185 185 185 185 185
Tools (shovels, picking bags, saws, etc.) 20 20 20 20 20 20
Shop tools 42 42 42 42 42 42
Irrigation system 380 380 380 380 380 380
Drippers 30 100 0 0 0 0
Sprinklers 0 0 30 30 30 30
Total non-cash overhead costs 3,562 3,632 3,562 3,562 3,562 3,562
Total net cost for the year 17,228 6,037 5,318 5,334 3,295 361
a
FOB, freight on board.
E.A. Evans and O.J. Mendoza 620
pre-planting and planting costs are estimated at US$11,125 (US$5000 +
US$6125). In year 1, it is also assumed that there are no replanting costs asso-
ciated with replacement trees. However, provisions are made to accommo-
date this cost in subsequent years. Here we assume that 5% of the trees planted
in year 1 will be replaced in year 2. Cultural costs are then estimated, includ-
ing standard operations such as pruning, fertilizing and mowing. In our
example, the total cultural costs in year 1 are estimated at US$920/ha. Next
are harvesting and marketing costs. This subheading includes the costs of
such operations as picking, packing and shipping. Other charges include
sales charges (if these operations were commissioned) and inspection and
assessment fees. In both years 1 and 2, it is not expected that trees will have
saleable crops and, in fact, it is recommended that efforts should be made to
deower such trees to prevent fruiting and to foster better tree growth. Con-
sequently, there are no marketing costs in those years. To the sum for pre-
planting, planting, replanting, cultural, and harvesting and marketing costs
is added an interest charge of 5%. This reects the economic (opportunity
cost) cost of money that was either borrowed or owner-nanced, to cover these
expenses. Thus the total operating (variable) costs in year 1 is US$12,647/ha.
Overhead costs are xed costs (in the sense that they must be paid irre-
spective of the level of production) that relate to the entire farm. For the pur-
pose of budgeting they are allocated on a per hectare basis. There are two
subcategories of overhead costs: cash overhead and non-cash overhead.
Cash overhead costs must be paid with cash during the year. They include
liability and crop insurance, taxes and ofce expenses. Interest is charged on
cash overhead costs. Total cash overhead costs plus total operating costs
equals total cash costs. Total cash costs represent the total cash expended in
any given year for producing and marketing a crop. In this example total
cash costs for year 1 is US$13,666 (US$12,647 + US$1019)/ha. Non-cash over-
head costs reect capital recovery associated with asset ownership is added
later and discussed below.
The next major category is income from production. Mango trees will
start supporting economic crops from about the third year, even though the
trees are not yet fully mature. The income from the sales of these crops is
deducted (credited) from the total cash costs for that year. In years 1 and 2, there
is no income. However, in year 3 there is income. Based on our analysis, the
income (US$1530) is then deducted from total cash costs for that year (US$3286)
to give the net cash costs for the year (US$1756). The negative net cash costs
recorded for year 5 (US$267) implies that, for that year, revenues from
the sales of fruit exceeded total cash expenditures for that year (US$7833
US$8100).
The accumulated net cash costs category is self-explanatory; it keeps a
running tally of cash costs incurred during the establishment years. Thus,
based on our analysis, the accumulated net cash costs in year 6 amounts to
US$15,408/ha, reecting the sum of the net cash costs for years 1 through to
6, i.e. US$13,666 + US$2405 + US$1756 + US$1772 + (US$267) + (US$3923).
This amount does not include capital recovery costs for equipment and
machinery that a grower owns and that is used in establishing an orchard.
World Mango Trade and Economics 621
To complete the establishment phase budget, it is necessary to include
the non-cash overhead charges. As previously dened, these are not cash
payments. They include an estimate of the amount of money that should be
set aside to help with replacing an investment item at the end of its useful
life. They usually reect depreciation and interest charges associated with a
capital item (i.e. the contribution of a capital item to the production process
for a given year). If some of these services were commissioned they would be
included in cash charges. The same is true for land rent. Here it is included
among the non-cash overhead charges because we assumed that the grower
already owns the land and therefore is not paying cash for the land. Since the
land could have been sold and the proceeds placed in a bank to earn interest,
the rent charged against production represents, in a sense, lost interest
from not selling the land and placing the proceeds in a money-bearing bank
account. In other words, it is included as payment to the owner for using his
land in this manner. The sum of non-cash overhead and net cash cost equals
the total net cost for that year. Table 16.7 shows that, in year 1, the total net
cost is US$17,228/ha, out of which US$13,666 and US$3562 are the net cash
costs and the non-cash overhead costs, respectively.
The nal category is total accumulated net costs, which keeps a tally of
the total cost incurred since the start of the establishment phase. Based on the
information provided in Table 16.7, the data reveal that, at the end of the
6 years, the investor would have invested US$36,850/ha to establish the
orchard. Of this amount, the cash costs amount to US$15,408 and non-cash
costs due to ownership of xed assets were estimated at US$21,442. Thus,
the total cost to establish the 18 ha mango orchard is US$663,300 (18
US$36,850) of which the cash amount (ignoring xed costs) would be US$277,344
(18 US$15,408).
The full establishment costs of US$663,300 (US$36,850/ha) is then
amortized at 5% real rate of interest for the estimated life of the orchard (25
years) to give an annual cost for capital recovery (interest and depreciation)
of US$2615/ha. This amount will be charged as part of the non-cash over-
head charges (Table 16.8) in computing the cost to produce a hectare of mango
during the production phase (after the orchard has been established).
Discussion of production phase budget
Table 16.8 provides the estimates of the annual growing costs (after establish-
ment) for mango in south Florida. The budget is similar to the one used in the
establishment phase except that there are no provisions for pre-planting,
planting and replanting expenses, and the estimates are for a typical year.
Also of importance, and shown in italics in Table 16.8, is the amortized estab-
lishment cost (US$2615/ha) obtained from Table 16.7. This reects the annual
amount that must be recovered for the investment made in establishing an
orchard.
Based on Table 16.8, it can be seen that the total annual cash cost to pro-
duce a hectare of mango, assuming a yield of 22,000 kg/ha (Table 16.6) is
E
.
A
.

E
v
a
n
s

a
n
d

O
.
J
.

M
e
n
d
o
z
a
6
2
2
Table 16.8. Sample cost per hectare to produce mango in south Florida.
Operation Unit
Costs (US$/ha)
Operation
time (h/ha)
Labour
cost
Fuel, lube
and repairs
Material
cost Custom/rent
Total
cost
Cultural costs (materials, labour, fuel,
lube and repairs)
Orchard pruning ( 1) 15 135 25 0 150 310
Herbicide ( 4) 8 72 37 40 0 149
Fungicide ( 12) 30 390 36 940 0 1,366
Fertilizer ( 4) 12 156 24 792 0 972
Insecticide ( 1) 4 36 25 40 0 101
Mowing ( 6) 12 156 25 0 0 181
Irrigate and walk lines ( 1) 3 27 0 40 0 67
Pest control advisor ( 1) 0 0 0 0 150 150
Pollination ( 1) 0 0 0 0 450 450
Rodent control (squirrels) ( 1) 0 0 0 0 60 60
Machinery repair 5 65 54 20 0 139
Total cultural costs 89 1,037 226 1,872 810 3,945
Harvesting and marketing costs
Picking US$0.11/kg US$0.11 0 0 0 0 2,420 2,420
Packing and shipping US$0.11/kg US$0.11 0 0 0 0 2,420 2,420
Sales charge @ 10% of FOB price 10% 0 0 0 0 1,100 1,980
Total harvesting and marketing costs 0 0 0 0 5,940 6,820
Interest on operating capital @ 5% 5% 197
Total operating costs/hectare 1,037 226 1,872 6,750 10,962
Cash overhead costs
Liability insurance 30
Crop insurance 445
Leaf analysis 12
W
o
r
l
d

M
a
n
g
o

T
r
a
d
e

a
n
d

E
c
o
n
o
m
i
c
s
6
2
3
Soil analysis 12
Sanitation fee 106
Ofce expenses 355
Property taxes 150
Property insurance 250
Investment repairs 207
Interest on operating capital (cash overhead) 5% 78
Total cash overhead costs 1,645
Total cash costs/hectare 12,607
Cost per
producing
hectare
Annual
cost: capital
recovery/ha Total cost
Non-cash overhead
Land rent 2,500 2,500 2,500
Machinery and equipment 3,648 405 405
Building 2,964 185 185
Tools (shovels, picking bags, saws, etc.) 217 20 20
Shop tools 464 42 42
Irrigation system 4,817 380 380
Sprinklers 20 20 20
Amortized establishment cost 2,615 2,615
Total non-cash overhead costs 14,630 6,167 6,167
Total cost/hectare 18,774
E.A. Evans and O.J. Mendoza 624
US$12,607 or about US$0.57/kg. Of this total, cultural costs accounts for
US$3945 (31%), harvesting and marketing costs amount to US$6820 (54%)
and cash overhead costs US$1645 (13%). When non-cash overhead costs are
added in, the economic cost to produce a hectare of mango in South Florida
is estimated at US$18,774, or about US$0.85/kg.
Protability analysis
Table 16.9 utilizes the information presented in Tables 16.6 and 16.8 along
with information obtained elsewhere to analyse the protability of mango
production in South Florida based on 2006 data. Based on an average freight
on board (FOB) price of US$0.90/kg and marketable yield of 22,000 kg/
ha (actual yield would be slightly more), the gross returns are calculated as
US$19,800/ha. Subtracting total operating costs of US$10,962 (cultural, and
harvesting and marketing costs) from this amount gives a gross margin of
US$8838/ha. In other words, based on these assumptions, the enterprise is
protable in the short run since gross margin is positive. This implies that the
returns from crop sales are more than sufcient to cover the variable costs
incurred in the production and marketing of the crop and there are still funds
left over to go towards paying overhead costs.
To be viable in the long run, the amount remaining should be able to
cover overhead (cash and non-cash) and provide a reasonable return to the
operator. Subtracting cash overhead costs (US$1645) and non-cash overhead
costs (US$6167) from gross margin (US$8838) gives net returns of US$1026/
ha. Viewed slightly differently, from total revenue of US$19,800 obtained from
selling a hectare of the crop, after paying all costs (variable and xed) the
amount of US$1026/ha remain as payment to management. This results in a
return of approximately 7% on cash and non-cash investment.
1
Given that prices and yield can vary, Table 16.10 shows a sensitivity anal-
ysis for changes in price and yield variables. The sensitivity analysis is cal-
culated on net returns but could be done on gross margin. As seen in Table
16.10, a 10% price increase combined with a 10% yield increase would result
in over a 335% net return per hectare increase (from US$1026 to US$4460/
ha). A 10% price increase would have a greater impact on net return than
would a 10% yield increase. In the case of the former, the net returns would
increase from US$1026/ha to US$2984/ha (191%). In the case of the latter, net
returns would increase to US$2324 (127%). This underscores the importance
of doing whatever is necessary to improve crop prices such as improving the
quality of the product and engaging in direct marketing where possible.
Assuming marketable yields of 22,000 kg/ha, the break-even price, or
the price at which net returns (economic prots) would be zero, is computed
as US$0.85/kg. In other words, at this price the grower is just able to cover
both variable and xed costs. Any FOB price above this would result in pos-
itive net return and vice versa. Likewise, if prices were to remain at US$0.90/
kg, growers would have to ensure that they obtain yields in excess of about
20,000kg/ha (break-even volume) to generate positive returns.
World Mango Trade and Economics 625
Table 16.9. Costs and returns to produce mango in south Florida, 2007.
Unit Quantity/ha
Price or
cost per
unit (US$)
Value or
cost/ha
(US$)
Gross returns kg 22,000 0.90 19,800
Total gross returns for mangoes 19,800
Operating costs
Pruning Tree 175 1.77 310
Herbicide Application 4 37.25 149
Fungicide Application 12 113.83 1,366
Insecticide Application 1 101.00 101
Fertilizer Application 4 243.00 972
Mow middles Times 6 30.17 181
Pest control advisor ha 1 150.00 150
Irrigation ha 1 67.00 67
Pollination ha 1 450.00 450
Rodent control ha 1 60.00 60
Machinery repair ha 1 139.00 139
Interest on operating capital @ 5% 5% 197
Harvesting and marketing costs
Picking US$0.11/kg kg 22,000 0.11 2,420
Packing and shipping US$0.11/kg kg 22,000 0.11 2,420
Sales charge @ 10% of FOB price kg 22,000 0.09 1,980
Total operating costs/hectare 10,962
Gross Margin 8,838
Cash overhead costs
Liability insurance 30
Crop insurance 445
Leaf analysis 12
Soil analysis 12
Sanitation fee 106
Ofce expenses 355
Property taxes 150
Property insurance 250
Investment repairs 207
Interest on operating capital
(cash overhead) @ 5%
5% 78
Total cash overhead costs 1,645
Total cash costs/hectare 12,607
Non-cash overhead
Land rent 2,500
Machinery and equipment 405
Building 185
Tools (shovels, picking bags,
saws, etc.)
20
Shop tools 42
(Continued)
E.A. Evans and O.J. Mendoza 626
16.4 Conclusions
The major trends and developments occurring within the world and USA
mango market have been discussed and the methodology used to compute
the costs of establishing and maintaining an orchard has been demon-
strated. Several trends were highlighted, including the increasing levels of
production, trade and consumption of mangoes and the declining or stag-
nating prices. Although the case for establishing an orchard was hypotheti-
cal, the statistics are based on information obtained from growers, economic
engineering using recommended practices, and discussions with industry
experts.
Acknowledgements
The authors are indebted to many farmers and agricultural researchers who
provided us with innumerable advice and relevant comments about this
study. We wish to thank those who kindly offered suggestions for improve-
ment, including Dr Jonathan Crane, Dr Carlos Balerdi, Scott Smith, Luis D.
Roman, Sikavas Nalampang, Denys Benjamin and Valderez Gonzalez. A spe-
cial thanks to Carol Fountain for editing the manuscript.
Table 16.9. (Continued)
Unit Quantity/ha
Price or
cost per
unit (US$)
Value or
cost/ha
(US$)
Irrigation system 380
Sprinklers 20
Amortized establishment cost 2,615
Total non-cash overhead costs 6,167
Total cost/hectare 18,774
Net returns above total costs 1,026
Table 16.10. Sensitivity analysis.
Yield (kg/ha)
FOB price (US$/kg)
0.72 0.81 0.90 0.99 1.08
19,800 3833 2052 272 1508 3288
22,000 2891 933 1026 2984 4943
24,200 1949 187 2324 4460 6597
World Mango Trade and Economics 627
Note
1
This is obtained by adding the interests charged on operating and cash overhead costs
to the net returns to give an adjusted net income. Next, subtract these interests from the
total costs per hectare to give an adjusted total cost per hectare. Finally, express the
adjusted net income as a percentage of the adjusted total cost per hectare.
References
FAOSTAT (2007) Food and Agriculture Organization of the United Nations database.
Available at: http://faostat.fao.org/ (accessed 30 November 2007).
United States Department of Agriculture (USDA) Agricultural Marketing Service (2007)
USDA, Agricultural Marketing Service: Fruit and Vegetable Market News Website.
Available at: http://www.marketnews.usda.gov/portal/fv (accessed 31 March 2008).
United States Department of Agriculture (USDA) Economic Research Service (2007)
Fruit and Tree Nut Yearbook, 2005. Available at: http://www.ers.usda.gov/Data/
FoodConsumption/FoodAvailSpreadsheets.htm (accessed 31 March 2008).
United States Department of Agriculture (USDA) Foreign Agricultural Service (2007)
USDA Foreign Agricultural Service: Market and Trade Data. Available at: http://
www.fas.usda.gov/markettradedata.asp (accessed 3 November 2007).
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
628 (ed. R.E. Litz)
17 Fruit Processing
L.C. Raymundo, M.T. Ombico and T.M. de Villa
University of the Philippines Los Baos, Laguna, the Philippines
17.1 Introduction 628
17.2 Dehydrated Mango Products 630
Dried mango 630
Mango fruit bar 631
Mango fruit roll 632
Vacuum-puffed dried mango 632
17.3 Spray-dried Mango Powders 634
Spray-dried mango fruit powder and instant mango juice 634
Spray-dried green mango powder 635
Spray-dried instant green mango shake 636
17.4 Capital Investment Costs 637
17.5 Raw Material Requirements of the Mango Processing Plant 638
17.6 Conclusion 639
17.1 Introduction
Processing is a value-adding step that lls the gap between farm production
and marketing. The objective of processing is to convert highly perishable
but prime quality farm commodities to more stable forms. When the process
is accomplished with the least alteration in the nutritional value and aes-
thetic properties of the food, high consumer acceptance is assured. Com-
pletely new product lines can likewise be created out of fresh raw materials
through processing. In other instances, the raw material may undergo such
extensive physical alteration during processing that the product is used dif-
ferently by consumers. Culinary experts devise new uses for these products
to t the changing lifestyles of present-day consumers. The availability in the
market, for example, of pre-mixed condiments and various meat powders
used for avouring foods such as instant noodles and similar convenience
Fruit Processing 629
foods, has simplied the life of working women who do not have the time
or the skill to prepare food the way their mothers and grandmothers did
at home. The products are affordable and easy to prepare; hence, consumer
acceptance is high. In addition, these foods are found everywhere, from
neighbourhood stores in the suburbs and countryside to supermarkets in
urban centres. The product and process research and development work of
food scientists and food technologists, the business acumen of entrepreneurs
and the marketing expertise of product distributors ensure the availability of
the food products.
The global competitiveness of agricultural produce of a country can be con-
siderably enhanced by utilizing appropriate technologies to produce high-
quality processed foods. Research focusing particularly along the areas of
processing equipment, product design and packaging has made the latest
techniques available for the manufacture of new generations of food prod-
ucts. Tropical fruits have had an excellent record of breaking into the world
market because of their exotic avours (Plate 83). When processed into pure,
juice, nectar or simply dried fruit slices, they can be shipped long distances
with only minimum changes in quality.
The popularity of fruits may be attributed to consumer perception of
their health benets. In the case of fruit juices, many consumers are now
looking for healthy alternatives to the traditional carbonated beverages. Fruits
are rich in dietary bre as well as phytonutrients, especially antioxidants,
and have no cholesterol. The availability in the market of natural fruit juices
derived from fresh fruits is a welcome alternative to synthetic juices that are
being passed off as fruit juices but are, in reality, sugar-based, fruit-avoured
beverages prepared from articial ingredients.
Mango fruits are usually eaten fresh as dessert or as relish depending on
their stage of maturity when picked. In the Philippines and elsewhere, fresh
mangoes are available throughout the year because the ower-induction
technology rst described by Barba (1974) allows growers to produce fruit
out-of-season. The off-season fruits, however, are still expensive. Most mango
growers have not put the technology into practice due to the high cost of
additional farm inputs necessary for its successful implementation during
the rainy season.
At the peak of the harvest season, on the other hand, oversupply of
fresh mangoes depresses the market price to the detriment of the growers.
Moreover, high temperatures combined with high relative humidity (RH)
and intense sunlight during the harvest season accelerate the metabolic
processes associated with ripening in fresh mangoes, rendering them sus-
ceptible to microbial attack, particularly by Colletotrichum gloeosporioides
Penz., the cause of anthracnose. The physico-chemical changes that occur
during ripening also lead to fruit deterioration and loss of quality. Thus,
fresh mangoes are processed to facilitate better distribution and to stabilize
prices.
Mango processing was previously reviewed by Nanjundaswamy (1997),
who described the status of processing technologies and products in India.
This review discusses current trends in mango processing.
L.C. Raymundo et al. 630
17.2 Dehydrated Mango Products
Dehydration works on the principle that by lowering the water (H
2
O) con-
tent of foods below a certain threshold level, growth of many spoilage micro-
organisms is prevented, thus preserving the food. As more and more H
2
O is
removed from the material, its solids content becomes concentrated, further
making the food less suitable for microbial growth.
The choice of the most appropriate drying system is determined largely
by the cost-effectiveness of the process. Sun drying is the most inexpensive
method for drying foods. The products, however, are rather susceptible to
contamination by dirt, insects, rodents, faecal matter and microorganisms.
The process also requires several days to dry each load or batch of raw mate-
rials since it relies on the availability of sunlight. Temperature control is vir-
tually impossible. Other systems have been designed that are more hygienic
and equally cost-effective compared to sun drying. The use of solar dryers
practically eliminates most of the inherent defects of sun drying. However,
the reliance of solar dryers on sunlight as the source of energy for evaporat-
ing H
2
O from the material being dried makes the system commercially unre-
liable. Solar panel collector-equipped dryers with provision to store energy
from sunlight can operate continuously and efciently.
A mechanical dryer such as the convection oven provides the processor with
control over the system that is essential for a successful drying operation. It elim-
inates most of the problems mentioned above. Although the initial investment
cost is high due to the acquisition of the dryer, in the long run it is more eco-
nomical than sun drying because the drying time per load is much shorter.
Oil-, steam-, liquid propane gas- or electric-powered air heaters are the
alternative sources of energy for the dryers. Dryers have also utilized a wood-
red furnace that heats the air entering the drying chamber. Its main advan-
tage is that trimmings from raw materials, packing materials and other trash
can be burned in the furnace, keeping the compound clean.
Dried mango
Dehydrated or dried mango slices are among the rst products manufac-
tured commercially from ripe mango fruits that caught the attention of con-
sumers in the international market for processed tropical fruits (Plates 83 and
84). The product was developed in Cebu, the Philippines, from where it
spread to the neighbouring islands of Panay and Negros. It is now produced
in many regions of the Philippine archipelago where mango is abundant. In
addition, it is a popular product in Thailand and elsewhere in South and
South-east Asia.
In the Philippines, the Carabao mango is the preferred variety for dehy-
dration or drying. The fruit should be at the rm-ripe stage. When over-ripe
fruit is used as raw material, a dark-coloured product will invariably result.
Although the dried pieces from over-ripe mangoes have a more distinct ripe
mango avour that attracts customers, the shelf life is considerably shorter.
Fruit Processing 631
The fruit is washed thoroughly. The cheeks are sliced with a sharp
stainless-steel knife. Each slice is then cut into two or three pieces length-
wise. The esh is scooped from the skin with a stainless-steel scoop or ladle.
These operations are done manually; however, a simple and novel peeling
and slicing machine for ripe mangoes has been developed and patented in
Australia (as cited by Nanjundaswamy, 1997).
Sugar syrup is prepared by adding 175 kg sugar, 1.7 kg citric acid and
0.85 kg sodium metabisulte in 175 l H
2
O to make a 45 Brix sugar concen-
tration and heating to 90C to dissolve the metabisulte. The prepared
mango slices (1 t) are added to the syrup. The preparation is heated to 90C
and then allowed to stand for at least 6 h. Subsequently, the sugar concen-
tration of the syrup is adjusted by dissolving more sugar until the concen-
tration reaches 50 Brix using a hand refractometer. After 6 h, the mango
pieces are again removed from the syrup and the sugar concentration is
adjusted to 60 Brix using a hand refractometer for the plumping stage.
The syrup is reheated to 90C; the mango slices are added to the syrup and
soaked for a further 6 h.
The nal step in the process involves the removal of mango pieces from
the syrup. They are lightly rinsed with H
2
O to remove the excess sugar that
may crystallize on the surface of the mango during drying. The slices are
then loaded in drying trays and dehydrated at 4050C in a convection dryer.
Drying time usually requires 18 h. The dried mango pieces are unloaded
from the dryer and reconditioned at room temperature overnight. Each
piece is coated with confectioners sugar. The product is then packed in
aluminium-foil-laminated pouches and sealed. Recent improvements in
dryer design can reduce drying time to 8 h. As a result, up to three batches of
prepared mango slices can be processed daily instead of only two batches
every 36 h. The process for the production of dried mangoes has been
described by Raymundo et al. (1999).
Mango fruit bar
The product is also commonly referred to as mango leather (Plate 83). The
processing of mango fruit bar has also been described previously by Amor-
iggi (1992) and Raymundo et al. (1999, unpublished work, 2006). Pure pro-
cessed from ripe Carabao mango is the preferred raw material for the
manufacture of mango fruit bar in the Philippines, although Pico is also
suitable because of the orange colour of the pure.
The total solids content of 1 t of mango pure is adjusted to 25 Brix with
the use of a hand refractometer by adding sugar to the pure. The amount of
sugar required depends on the initial total solids content of the mango pure.
Then 2 kg each of citric acid and sodium metabisulte are blended into the
pure. Juice of calamansi, also known as the Philippine lemon ( Citrofor-
tunella microcarpa Wij. (Bunge); Citrus mitis Blanco) may be used instead of
citric acid at the rate of c.20 kg per batch, although the resulting cost of the
product will be slightly higher. There is no real difference in the appearance
L.C. Raymundo et al. 632
and avour of the nished product. Citrus juice is generally utilized if the
client prefers an all-natural product.
The prepared pure is heated for 2 min at 80C with constant stirring to
avoid scorching. The hot mixture is carefully transferred to stainless-steel
drying trays. The trays are loaded into the dryer and dried for 14 h at 55C.
At the end of the drying operation, the dried sheets of mango should be pli-
able and should not stick to the ngers when touched.
The sheets of mango are removed from the trays. Six layers of the dried
mango leather are stacked on top of each other. The edges are trimmed and
bars measuring 2 4 cm are cut from the stack. Each bar is coated with con-
fectioners sugar, wrapped in cellophane then packed in labelled plastic bags.
Mango fruit roll
The processing of mango fruit roll has been described previously (UPLB,
1996; Raymundo et al., 1999, unpublished work, 2006). The product and pro-
cess are similar to mango fruit bar, and they only differ in the presentation.
The total solids content of 1 t of mango pure is adjusted to 25 Brix using a
hand refractometer by adding sugar to the pure. The amount of sugar
needed depends on the initial total solids content of the mango pure. Then
2 kg each of citric acid and sodium metabisulte are blended into the pure.
As with mango fruit bar, citric acid may be replaced by calamansi juice at the
rate of c.20 kg per batch without affecting the overall sensory quality of
the fruit roll, if the client species an all-natural product.
The prepared pure is then heated for 2 min at 80C with constant stir-
ring to avoid scorching. The hot mixture is carefully transferred to stainless-
steel drying trays. The trays are loaded into a convection dryer and dried for
1216 h at 55C.
The dried sheets of mango are removed from the trays. Confectioners
sugar is sprinkled over a stainless-steel-topped table to avoid sticking of
the sheets on subsequent rolling. Each sheet is rolled manually into 1 cm
diameter pieces. The ends are trimmed; and each roll is sliced into 5 cm long
pieces. The pieces are coated with confectioners sugar and wrapped with
cellophane. The rolls are then packed in appropriate containers.
Vacuum-puffed dried mango
The use of vacuum for pufng and drying mango and similar food materials
is not as widespread as explosion pufng (Eskew et al., 1963), high tempera-
ture short time (HTST) pneumatic drying and centrifugal uidized bed (CFB)
drying (Brown et al., 1972), because the vacuum-puff dryer is still expensive.
With the increasing consumer demand for high-quality processed foods and
their willingness to pay a higher price for such products, vacuum-puff dehy-
dration could become an economically viable investment opportunity for
entrepreneurs, particularly in the mango processing industry.
Fruit Processing 633
Vacuum-puffed mango pieces will readily rehydrate due to the porosity
created during the pufng and drying stages of the process. The signicantly
shorter drying time (Candelaria and Raymundo, 1994b) also makes possible
the drying of four loads of prepared mango slices in 24 h.
The ripe mangoes are washed thoroughly in chlorinated H
2
O, then sliced
either mechanically or with a stainless-steel knife. The esh is scooped from
the skin with a sharp stainless-steel ladle (Candelaria, 1993; Candelaria and
Raymundo, 1994a). The fruit pieces are then heated to 90C in 30 Brix syrup
containing 1% sodium metabisulte, and steeped in the syrup for 46 h. The
mango slices are removed from the syrup, rinsed briey in H
2
O, arranged in
stainless-steel trays and loaded into the vacuum oven.
The mango pieces thus prepared are initially heated at a positive pres-
sure of 4050 kPa until the maximum tissue temperature of 100C is reached,
usually within 8 min. The pressure is released and the hot mango pieces are
dried at 70 to 80 kPa vacuum at a temperature of 4550C. Total dehydration
time under vacuum is 6 h. The above pressure-temperature combinations pro-
vide the most desirable puff and rehydration characteristics (Candelaria and
Raymundo, 1994b).
With the present technology, vacuum-puffed dried mango from a 1 t
batch of prepared mango slices is more expensive to produce than convec-
tion oven-dried mango (Table 17.1). The production cost needs to be reduced
for the product to be market competitive. Research that focuses on devising
a system to assure a continuous supply of mango fruits is required. The plant
must operate on a year-round basis in order to optimize the use of its equip-
ment and facilities.
The facilities can be used for the production of other vacuum-puffed
fruits (i.e. bananas and muskmelon) as well as vegetables (i.e. white potato
and maize kernels) (Candelaria, 1993; Candelaria and Raymundo, 1994b)
among others, which can be used as raw materials in the manufacture of
instant foods.
Table 17.1. A comparison of the protability of different mango product lines.
Product Volume (kg)
Cost of goods
sold (US$)
Gross prot
(US$)
Net prot before
tax (US$)
Mango powder
a
124,800 251,460 966,540 439,983
Instant mango juice
a
202,890 467,355 1,560,645 965,088
Green mango powder
a
141,221 245,226 1,166,984 600,216
Instant green mango shake
a
124,800 209,468 1,308,532 473,975
Vacuum-puffed dried mango
b
124,800 1,141,807 730,193 246,588
Dried mango
c
67,392 398,492 140,644 84,386
Mango fruit bar
c
115,200 921,600 461,816 277,090
Mango fruit roll
c
115,200 921,600 461,816 277,090
a
US$10/kg powder.
b
US$15/kg.
c
US$8/kg.
L.C. Raymundo et al. 634
17.3 Spray-dried Mango Powders
Spray-drying is a process in which a liquid feed is nely dispersed or atomized
to form droplets, which are eventually sprayed into a heated air chamber. The
process facilitates the rapid evaporation of H
2
O from the feed droplets, thereby
forming the powder particles. The product obtained using the technology is a
free-owing powder that may or may not be instantly soluble in H
2
O, depend-
ing on the formulation of the liquid feed that has been used. By modifying the
formulation and adjusting the process parameters, a plain powder is obtained
that can be dry-mixed with sugar, modied starches or similar components.
The powder is used for avouring confectioneries and pharmaceutical prepa-
rations as well as in the manufacture of baby foods and tropical fruit drinks
fortied with nutrients to replace those portions lost during processing.
Tropical fruit juice powders that are rapidly soluble in H
2
O are produced
directly by spray-drying fruit juices and pures. By dry-mixing spray-dried
plain fruit powder with sweeteners, a ready-to-drink juice is made. The latter
method has the added advantage in that it allows formulation of exclusive
blends of fruit drinks. As a result, consumers have a wider range of products
from which to choose that will suit their individual preferences.
By converting fruit pulps into powder or by instantizing them, their shelf
life is prolonged. Consequently, this value-adding step simplies exportation
since many of the restrictions normally imposed on fresh produce by import-
ing countries are offset by the process. Transportation cost is also reduced by
at least 85%, which reects the amount of H
2
O removed from the fresh juice.
Instant juices are more convenient for consumers since they can be reconsti-
tuted easily. In the pharmaceutical and cosmetics industries, there is a large
demand for natural tropical fruit powders as avouring and colouring agents
to add to their usual line of fruit-avoured products. Powders are also more
convenient to handle during the manufacturing process.
Spray-drying is by far the most cost-effective method for transforming fruit
pulps into powder. Fruit pulp is very heat-sensitive and requires special treat-
ments to produce competitively priced powders of superior quality. Further-
more, because of the rapid drying cycle and simplicity of operation, continuous
production is achieved which contributes to the low operational cost. The short
holding time of the powder inside the drying chamber reduces the risks of pow-
der burn. Human contact with the liquid feed and powder is also minimized as
a result of the short holding time. The processes are, therefore, very hygienic and
the product is ready for packaging as it leaves the dryer. Unlike other drying sys-
tems, including convection oven-drying and drum-drying, there is no need to
purchase a hammer mill for grinding the dry akes into a powder. The main
drawback of spray-drying, however, is the relatively high initial investment cost.
Spray-dried mango fruit powder and instant mango juice
Khalid and Sial (1974), Anonymous (1998) and Diaz (2000) have discussed
the recovery of fruit powder and production of instant mango juice powder
Fruit Processing 635
using the technology. Both products use mango pure as the raw material.
They differ only in the composition of the liquid feed. The liquid feed is
mango pure with the total solids adjusted to the right consistency, thereby
allowing the pure to be discharged through a high-speed nozzle in the form
of ne droplets into the drying chamber that quickly dries to a yellow free-
owing powder. The patent application for the manufacture of spray-dried
mango powders is still pending at the Philippine Patent Ofce. The process
parameters used, therefore, cannot be discussed in detail in this chapter. The
parameters used are the same as reported previously, namely, the inlet tem-
perature, the feed rate, air speed and the outlet temperature (Welti and Lau-
ente, 1983; Liang and King, 1991; Anonymous, 1998). The powder of instant
mango juice is comparatively dense so that it gets wet easily, enabling it to
sink immediately during reconstitution. The plain mango fruit powder, on the
other hand, does not disperse as readily; it has a tendency to clump on the H
2
O
surface. However, it will dissolve quickly in hot H
2
O or with the use of an
ordinary household blender.
A modest capacity spray-drying plant equipped with a NIRO SD 12.5R
model spray-dryer made in Denmark, valued at approximately US$207,860,
with an evaporative capacity of 25 kg H
2
O/h will require an initial invest-
ment of about US$313,860 to make it operational. In addition, if fresh man-
goes are be used as raw materials, a pure processing facility will cost around
US$98,000. The gure represents expenses for buying and installation of
equipment, and does not include the costs of the building, land and ancillary
expenses such as environmental pollution control system and spray-dryer
accessories.
At the evaporative capacity rate of 25 kg H
2
O/h, the facility can produce
124.8 t/year of plain mango powder and 202.8 t/year of instant mango juice
at a total cost of US$251,460 and US$467,355, respectively (Table 17.1). The
net prot for plain mango powder is US$439,983 and US$965,088 for instant
mango juice before taxes.
Spray-dried green mango powder
The pure of green mangoes can be converted to a powder (UPLB, 2005) just
like the pure of ripe mangoes by spray-drying (Plate 85a). The spray-dried
powder can be mixed with other condiments and used as a souring agent for
exotic or native dishes, or as the raw material in the manufacture of instant
green mango shake (see below). The powder may be dry-mixed with sugar,
powdered honey, caramel powder or powdered sugar syrup to instantize it.
During this process, the mixture can be fortied with vitamins (e.g. ascorbic
acid) and other nutrients.
The estimated cost of goods sold using a commercial spray-dryer with
an evaporative capacity of 25 kg H
2
O/h is US$245,226/year (Table 17.1). At
the rated capacity of the spray-drying plant of 141,221 kg/year, the net prot
before taxes for green mango fruit powder is US$600,216 at a selling price of
US$10/kg of powder.
L.C. Raymundo et al. 636
Spray-dried instant green mango shake
Green mango shake on cracked ice is a very popular thirst quencher in South-
east Asia and elsewhere (Plate 85b). Its origin can be traced back to the coun-
tryside where nely diced fresh green mango pieces are mixed with H
2
O,
sugar and ice to make a cheap, wholesome summer drink during the mango
season. The practice has since been modied and upgraded to cater to upscale
domestic markets and abroad.
The beverage is an excellent source of vitamin C. When the green mango
pure is spray-dried, a free-owing white to greenish-white powder is pro-
duced that will dissolve instantly even in cold H
2
O. The powdering process
offers unprecedented convenience to the consumer, especially when the pow-
der is packed in sachets. Since no articial or synthetic colours and avour-
ing agents are included in the liquid-feed formulation, the natural taste and
aroma of green mango is retained in the powder.
Using a commercial spray-dryer with an evaporative capacity of 25 kg
H
2
O/h, the estimated cost of goods sold is US$209,468/year. At the rated
capacity of spray-drying plant of 124,800 kg, the net prot before taxes for
instant green mango shake is US$473,975/year at a selling price of US$10/kg
of powder (Table 17.1).
The technology for the production of instant green mango shake and
green mango powder was developed at UPLB (2005). If the proper feed for-
mulation and process parameters are applied, spray-drying is an efcient
and hygienic method for producing cheap but high-quality mango fruit
powder and instant mango juice. Table 17.2 demonstrates that the spray-
dried mango powders have much lower microbial load, i.e. 2.810.4 10
2

colony forming units (cfu)/g, compared to the industry standard of 1 10
4
to
5 10
5
cfu/g total plate count. Except for mango powder, the mould and
yeast counts are within the limit of 1 10
2
cfu/g. Locally rened sugar and
imported modied starch, on the other hand, have much higher mould
and yeast counts than the Heinz standard for powdered products (Shapton
Table 17.2. Microbial load of raw materials and spray-dried mango fruit powder and instant
mango juice (cfu/g).
Material Total plate count Mould count Yeast count
Mango powder 2.8 10
2
2 10
2
<10
Instant mango juice 3.6 10
2
0.4 10
2
<10
Green mango powder 10.4 10
2
0.95 10
2
<10
Instant green mango shake 9.8 10
2
0.85 10
2
1 10
2
Mango pure 0.5 10
2
0.2 10
2
<10
Green mango pure 0.8 10
2
0.1 10
2
<10
Sugar, rened 11 10
2
43 10
2
20 10
2
Modied starch 63 10
2
7.5 10
2
5 10
2
H.J. Heinz standard (Shapton and
Shapton, 1991)
1 10
4
to 5 10
5
1 10
2
1 10
2
Fruit Processing 637
and Shapton, 1991) at 11 10
2
and 63 10
2
cfu/g total plate count, respec-
tively. These materials are essential ingredients of the liquid-feed formula-
tion used for the production of the spray-dried powders.
The formulation and processing of numerous mango products popu-
lar in South Asian countries were reported previously (Nanjundaswamy,
1997).
17.4 Capital Investment Costs
An initial investment of approximately US$100,000 is needed for the estab-
lishment of a mango dehydration plant. The facility can also be used for dry-
ing other farm commodities such as vegetables, spices and other high-value
crops. The cost of acquiring the land, building construction and ancillary
costs such as environmental pollution abatement as well as quality assurance
laboratory facilities are not included in the estimate.
A commercial dryer such as that manufactured by Tsung Hsing Food
Machinery Co. Ltd of Taiwan with a loading capacity of 500600 kg prepared
mango slices costs c.US$11,000. At the rated capacity of the plant of 1 t per
batch, 67,392 kg of dried mango pieces can be produced annually. The cost of
goods sold is US$398,492 with a net prot before taxes of US$84,386/year. If
the facility is used for the manufacture of mango fruit bar and mango fruit
roll, the annual production is 115,200 kg of either fruit bar or fruit roll with a
net prot before taxes of US$277,090/year (Table 17.1).
The total cost of equipment for the vacuum-pufng plant with a loading
capacity of 1 t is approximately US$122,000, broken down as US$100,000 for
fabrication of the dryer and US$22,000 for other plant equipment. The total
capitalization, which includes the cost of raw materials, salaries and equip-
ment per year, is US$648,240 excluding interest on capital.
Spray-dryers come in various capacities ranging from NIRO SD Micro
Spray-dryer ideal for powdering small quantities of raw materials for research
and development to units that produce powders at evaporative rates in excess
of 500 kg H
2
O/h. The investment cost for equipment increases with the capac-
ity of the dryer (Table 17.3). For a small-scale operation producing powders or
ours from commercial pure, the initial investment is US$182,000 for a 5 kg
H
2
O evaporative capacity dryer while a plant equipped with a 12 kg evapora-
tive capacity dryer requires US$262,000 for equipment. On the other hand, an
investment of US$313,860 is needed to equip a medium-scale drying plant
with a 25 kg H
2
O/h evaporative capacity such as NIRO SD 12.5R.
The estimates do not include land, building and construction costs as
well as the cost of pollution statement facilities. The cost of land is dependent
on where the plant will be located while building and construction costs are
determined by the capacity of the spray-dryer unit to be acquired.
Total capitalization is US$317,200, US$535,947 and US$806,265 for the
5 kg, 12 kg and 25 kg H
2
O/h evaporative capacity units, respectively. The
estimate includes the cost of raw materials, salaries and equipment per annum.
Interest on capital is not included.
L.C. Raymundo et al. 638
1.5 Raw Material Requirements of the Mango Processing Plant
The amount of mango pure, mango slices and fresh mango required
monthly and annually to run the plant continuously varies with the prod-
uct line (Table 17.4). In general, more raw materials are needed to process
dried mango, mango fruit roll and mango fruit bar compared to the spray-
dried mango powders. As a result, the net income is lower than that derived
from the production and sale of spray-dried mango powders (Tables 17.1
and 17.4).
Table 17.3. Basic equipment for the spray-drying plant.
Equipment and accessories
Evaporative rate
5 kg H
2
O/h 12 kg H
2
O/h 25 kg H
2
O/h
NIRO Production Minor 110,000
NIRO SD 6.3R 156,000
NIRO SD 12.5R 207,860
Spray-dryer accessories 30,000 25,000 25,000
Homogenizer/emulsier 20,000 45,000 45,000
Packaging equipment 5,200 5,200 5,200
Refrigerated rooms (modular, 1 t capacity,
@ US$2,500/unit)
5,000 10,000 10,000
Walk-in freezers (modular 1 t capacity
@ US$4,000/unit)
4,000 8,000 8,000
Stainless steel kettles (500 l capacity,
with stirrer @ US$2,500/unit),
5,000 10,000 10,000
Stainless steel work tables 2,800 2,800 2,800
Total 182,000 262,000 313,860
Table 17.4. Estimated volume of raw material needed for the operation of a mango
spray-drying and a mango dehydration plant.
Product line
Mango pure/slices Fresh mango
kg/month kg/year kg/month kg/year
Mango powder 21,000 252,000 42,000 504,000
Instant mango juice 19,500 234,000 39,000 468,000
Green mango powder 9,100 109,200 18,200 218,400
Instant green mango shake 7,800 93,600 15,600 187,200
Vacuum-puffed dried mango 104,000 1,248,000 208,000 2,496,000
Dried mango 34,670 416,040 69,340 832,080
Mango fruit roll 24,000 288,000 48,000 576,000
Mango fruit bar 24,000 288,000 48,000 576,000
Fruit Processing 639
The spray-drying facility requires 187.2218.4 t/year of green mangoes
and 468504 t/year of ripe mango fruits at the evaporative capacity of 25 kg
H
2
O/h to produce the volume of powders in Table 17.1. On the other hand,
576 t/year of ripe fruits are required to produce mango fruit roll or fruit bar.
For dried mango, 832 t/year of fresh fruits are needed to produce 67.4 t/year
of the product while to produce 124.8 t/year of vacuum-puffed dried mango,
2496 t/year of fresh fruits are necessary.
In terms of area, c.3848 ha can supply the green mango requirement of
the spray-drying facility. Approximately 100 ha are needed to produce 124.8
t of ripe mango powder or 202.9 t of instant mango juice from fresh ripe
mango fruits. The fresh fruit requirement for the manufacture of fruit roll
and fruit bar can be supplied by 115 ha of mangoes. The 832 t and 2496 t of
fresh fruit are equivalent to the annual harvest from 166 ha and 500 ha of
mangoes to produce 67.4 t and 124.8 t of dried mango and vacuum-puffed
dried mango, respectively. The estimates are based on a yield of 5 t from 100
trees/ha as well as the biennial fruiting habit of mango.
17.6 Conclusion
Processing of mango is a protable business venture once economies of scale are
attained, i.e. when the processor has the proper proportions of raw materials,
labour and machinery to meet a given market demand. Fresh mangoes are now
available year-round. But the supply is still insufcient to satisfy the demand by
the fresh fruit market and the mango-processing sector. It is, therefore, essential
that a farming system for mango be designed in order to minimize the cost of
production of off-season fruits and ensure the sustainable operation of mango
processing facilities. Since raw material sourcing is the primary cause of the dif-
culties encountered by processors of dried mangoes, mango in syrup and
similar products, growers need more assistance for them to adopt the tech-
nology for off-season mango ower induction. The problem is not as serious
for product lines utilizing mango pure as starting material since the pure
can be processed during the peak of the harvest season and held in cold stor-
age for later use. Once the system is in place, the processed mango industry
is expected to develop further and become a major revenue generator.
In the near future, with the advances in the eld of genetic engineering,
it may be possible to eliminate the biennial fruiting habit of many current
mango cultivars or, at the very least, minimize its inuence on the perfor-
mance of the crop. Species of Mangifera and some non-cultivated, wild types
of mango can be the source of desirable traits that may be incorporated in the
next generation of mango cultivars, such as their innate ability to bear fruits
during the rainy season (see Bompard, Chapter 2, this volume).
The technologies for processing mangoes are readily available. Others
are being developed in research laboratories to cope with the changing needs
of consumers. The main problem is to ensure a continuous supply of high-
quality fresh mango fruit in order to produce the prime quality commodities
that consumers expect from the industry.
L.C. Raymundo et al. 640
References
Amoriggi, G. (1992) The marvelous mango bar. Ceres 24, 2528.
Anonymous (1998) Instruction Manual for Spray Drying Plant. NIRO A/S Soeborg, Denmark.
Barba, R.C. (1974) Induction of owering of the mango by chemical spray. Proceedings
of the Crop Science Society of the Philippines 5, 154160.
Brown, G.E., Farkas, D.F. and De Marchena, E.S. (1972) Centrifugal uidized bed.
Blanches, dryers and puff piece-form foods. Food Technology 26, 2330.
Candelaria, N.M. (1993) Dehydration of vacuum-puffed fruits and vegetables. PhD dis-
sertation, University of the Philippines Los Baos, Laguna, the Philippines.
Candelaria, N.M. and Raymundo, L.C. (1994a) Comparative drying and reconstitution,
characteristics of some fruits and vegetables. Philippine Agriculturist 77, 321326.
Candelaria, N.M. and Raymundo, L.C. (1994b) Vacuum pufng and dehydration of fruits
and vegetables. Philippine Agriculturist 77, 251260.
Diaz, J.V. (2000) Development of spray-dried instant mango (Mangifera indica L.) juice pow-
der. MSc thesis, University of the Philippines Los Baos, Laguna, the Philippines.
Eskew, R.K., Cording, J., Jr and Sullivan, J.F. (1963) A gun for explosive, pufng of dehy-
drated fruits and vegetables. Food Engineering 35, 9196.
Khalid, M.A. and Sial, M.B. (1974) Spray-drying of mango juice powder. Mesopotamia
Journal of Agriculture 9, 4756.
Liang, B. and King, L.J. (1991) Factors inuencing ow patterns, temperature and conse-
quence drying rates in spray-drying. Drying Technology 9, 127.
Nanjundaswamy, A.M. (1997) Processing. In: Litz, R.E. (ed.) The Mango: Botany, Pro-
duction and Uses. CABI International, Wallingford, UK, pp. 507542.
Raymundo, L.C., Ombico, M.T. and De Villa, T.M. (1999) Establishment of integrated
mango processing plant. In: Namuco, L.O. and Andam, C.J. (eds) Mango Production
Manual. Philippine Council for Agriculture, Forestry and Natural Resources Research
and Development (PCARRD), Los Baos, Laguna, the Philippines, pp. 97119.
Raymundo, L.C., Ombico, M.T., De Villa, T.M. and Jaen, R.M. (2006) Processes of Fruits,
Nuts, Vegetables and Other Tropical Foods. Fruit and Vegetable Laboratory, Food
Science Cluster, University of the Philippines Los Baos, Laguna, the Philippines.
(Unpublished)
Shapton, D.A. and Shapton, N.F. (1991) Principles and Practices for the Safe Processing
of Foods. Butterworth-Heinemann Ltd, Oxford, UK.
University of the Philippines Los Baos (UPLB) (1996) Standardization of Processing
Method and Evaluation of Different Packaging Material and Storage Conditions for
Mango Roll. Annual Report, Institute of Food Science and Technology, College of
Agriculture, University of the Philippines Los Baos, Laguna, the Philippines.
University of the Philippines Los Baos (UPLB) (2005) Spray-dried Instant Juice and
Powder from Green Mango. Annual Report, Food Science Cluster, College of Agri-
culture, University of the Philippines Los Baos, Laguna, the Philippines.
Welti, J.S. and Lauente, B. (1983) Spray-drying of comminuted orange products. I. In-
uence of air and temperature and feed rate on product quality. Revista de Agro-
nomica y Technologia de Alementos 23, 97106.
CAB International 2009. The Mango, 2nd Edition: Botany, Production and Uses
(ed. R.E. Litz) 641
18 Biotechnology
R.E. Litz,
1
M.A. Gmez-Lim
2
and U. Lavi
3
1
University of Florida, Florida, USA
2
CINVESTAV, Irapuato, Mexico
3
Agricultural Research Organization, Bet Dagan, Israel
18.1 Introduction 641
18.2 Cell and tissue culture 643
Organogenesis 643
Somatic embryogenesis 643
Protoplast isolation and culture 650
Potential for other regeneration pathways 651
18.3 Molecular Breeding and Genetics 651
Marker assisted selection (MAS) 651
Gene cloning 651
Genomics 655
18.4 Genetic Engineering 656
In vitro induced mutations 656
Genetic transformation 659
18.5 In vitro conservation 660
Medium-term storage 660
Long-term storage 661
18.6 Conclusions 662
18.1 Introduction
Improvement of monoembryonic mangoes by selection of superior seedling
trees resulting from open pollinations dates from approximately 500 years
ago (the late 1500s), when the Mogul emperor Akbar established the Lakh
Bagh, a garden of seedling mango trees near Dabangha in Bihar (see Mukher-
jee and Litz, Chapter 1, this volume). Only a few years before this time, the
Portuguese had introduced grafting techniques into India, and the superior
selection mango trees within the Lakh Bagh were vegetatively propagated
and named. Among these early mango selections were Alphonso, Dashe-
hari, Langra, etc. All subsequently named mango selections have been
R.E. Litz et al. 642
derived by identifying seedling trees within populations either from within
uncontrolled open pollinations or, less commonly, among controlled polli-
nated progenies.
Mango cultivar improvement programmes currently exist in several
countries (see Iyer and Schnell, Chapter 4, this volume), and they address
signicant production problems that have a genetic basis. Classical breeding
of mango has obvious limitations, which include, the long juvenile period of
mango trees (7 or more years), the low frequency of fruit set following con-
trolled pollination, the period required for seedling trees to be evaluated for
fruit production, tree architecture, and the cost of maintaining large popu-
lations of seedling trees in order to observe segregation of important hor-
ticultural traits. There is no single ideotype for mango; however, the most
important attributes for scion cultivars must include: compact size, resistance
to anthracnose and other limiting diseases, fruit production (which would
include annual bearing and factors that affect fruit quality, i.e. shape, colour,
avour and size). In many traditional mango-producing countries, consumer
preference for mangoes has been quite conservative, and has been resistant
to the introduction of new cultivars. Although there is a demand for improved
clonal rootstocks, there are few breeding programmes that address this need.
For mango rootstocks, the most important traits include tolerance of abiotic
stress, polyembryony and the ability to limit vegetative growth of the scion
(i.e. dwarng).
Biotechnology refers to the application of molecular biology and somatic
cell genetics to the improvement of plants. Biotechnology can resolve some
of the most serious production problems of important mango cultivars and
improve breeding methodologies. Genomic studies will ultimately associate
genes with specic functions, and this will impact genetic engineering and
molecular breeding of mango. Marker assisted selection (MAS) would facili-
tate the screening of seedling populations for important horticultural traits.
Genetic engineering would permit the targeting and alteration of specic
horticultural traits in existing cultivars, without altering the integrity of clones.
Mango improvement by modern genetics will be freed from the constraints
of the lengthy juvenile period of the species and the additional years required
for tree evaluation. Moreover, the efcient management of mango plant
genetic resources should be greatly facilitated within the next decade by
advances that have been made in molecular biology, cell culture and cryo-
preservation.
Mango biotechnology has been variously reviewed since 2002 (Litz and
Gomez-Lim, 2002, 2005; Litz, 2003, 2004, 2008; Gomez-Lim and Litz, 2007;
Krishna and Singh, 2007). In this chapter, we have addressed the current sta-
tus of mango genomics, gene cloning and cell culture. Within the discussion
of cell culture, the following areas of research are addressed: (i) the utiliza-
tion and potential of cell culture for improving existing mango cultivars by in
vitro-induced mutation followed by selection and genetic transformation; and
(ii) advances in medium- and long-term conservation of genetic resources.
The model species Arabidopsis thaliana has provided a useful tool for studying
the regulation of gene expression of important plant processes, for example
Biotechnology 643
embryo and organ development, including shoots, leaves, roots, owers and
fruit. Genomic analysis and the identication of horticulturally important
genes that control these and other processes will ultimately enable us to under-
stand and control many aspects of mango fruit production (i.e. disease and
pest control, fruit quality, tree architecture, etc.). The development of molecu-
lar approaches will also facilitate mango breeding through the identication
of DNA markers for important traits.
This chapter is a discussion of the current state of mango cell culture,
gene cloning and the manipulation of cell cultures to address plant breeding
objectives by genetic engineering.
18.2 Cell and Tissue Culture
Organogenesis
The earliest report of morphogenesis from mango cell cultures involved
the differentiation of adventitious roots from callus that was initiated from
mango cotyledons on Murashige and Skoog (1962) semi-solid induction
medium (MS) that was supplemented with kinetin and naphthaleneacetic
acid (NAA) (Rao et al., 1982). Of greater signicance was the later report that
callus initiated from leaves from mature mango trees also had morphogenic
potential. Raghuvanshi and Srivastava (1995) induced caulogenic cultures
from fully expanded, young leaves of monoembryonic Amrapali. Disin-
fested leaf pieces were initially explanted into liquid MS medium supple-
mented with the antioxidant 0.05% polyvinylpyrolidone with 2 h subcultures
for up to 24 h on a rotary shaker at 75 rpm. Leaf pieces were then transferred
onto MS semi-solid medium supplemented with 13.0 M kinetin and 1.1 M
indole acetic acid (IAA), the optimum growth regulator formulation for mul-
tiple shoot development from leaf callus. Subculture of individual shoots
onto semi-solid MS medium supplemented with 9.8 M indole butyric acid
(IBA) stimulated root induction and development. Cultures were main-
tained at 25C with a 16 h photoperiod provided by cool white uorescent
lights (5070 mol/m
2
/s).
The major advantage of the organogenic pathway for regenerating mango
is that cultures can be induced whenever there is a leaf ush; however, genetic
manipulation studies require much higher rates of regeneration than has
been hitherto reported.
Somatic embryogenesis
Genetic engineering of mango has been based upon the efcient recovery of
somatic embryos from embryogenic cultures. Efcient regeneration of woody
plants from cell cultures derived from mature phase materials is often difcult
to achieve, and the optimum procedure generally must be determined empir-
ically. Mangifera indica consists of two ecogeographic races: a monoembryonic
R.E. Litz et al. 644
group that probably originated in north-eastern India and northern Myan-
mar and a polyembryonic group that originated in South-east Asia. The pres-
ence of nucellar embryos in seeds of polyembryonic mangoes demonstrates
that cells of the nucellus of the species have morphogenic potential (see
Mukherjee and Litz, Chapter 1, this volume). Sturrock (1968) reported that
polyembryony in mango appeared to be inherited as a recessive trait; how-
ever, Aron et al. (1998) have demonstrated that polyembryony is under the
control of a single dominant gene. Somatic embryogenesis, which is the equiv-
alent morphogenic response in vitro, is a dominant trait in several other spe-
cies, for example lucerne (Reisch and Bingham, 1980) and orchardgrass
(Gavin et al., 1989), although in red clover it apparently is conferred by a reces-
sive gene (Broda, 1984). The presence of morphogenically competent cells in
the nucellus is critical for the induction of embryogenic mango cultures of
both monoembryonic and polyembryonic mango cultivars.
Induction of embryogenic mango cultures from the excised nucellus of
immature mango seeds of polyembryonic and monoembryonic cultivars was
rst described by Litz et al. (1982) and Litz (1984), respectively. The current
standard protocol for induction and maintenance of embryogenic mango
cultures and for development of somatic embryos is derived from DeWald
et al. (1989a, b) and Litz et al. (1993). Table 18.1 provides citations for all in
vitro studies that have involved somatic embryogenesis of mango. The efforts
in this eld have expanded considerably since the late 1990s.
It is impossible to predict the embryogenic response of the nucellus of
different mango cultivars, irrespective of the seed type (i.e. polyembryonic or
monoembryonic). Early reports suggested that the nucellus of polyembry-
onic mangoes responded in vitro more readily than the nucellus of monoem-
bryonic mangoes (Litz, 1986); however, this assumption no longer is considered
to be valid. Litz et al. (1997) reported that the induction of embryogenic com-
petence in the explanted nucellus of monoembryonic Tommy Atkins can be
inhibited by the ethylene antagonist aminoethoxyvinylglycine (AVG) and by
dicyclohexylammonium sulfate (DCHA), an inhibitor of spermidine synthe-
sis. In contrast, induction of embryogenic competence of the explanted nucel-
lus of polyembryonic Tuehau is unaffected by either AVG or DCHA. Litz
and Schaffer (1987) demonstrated that somatic embryogenesis in mango is
partially mediated by spermidine. The biosynthesis of ethylene and/or the
sensitivity of nucellar tissue to ethylene may be important factors for the
induction of embryogenic cultures from this tissue.
Induction
The induction of embryogenic competence is associated with the develop-
mental stage of the nucellus at the time of explanting, and also can be af-
fected by the physiological status of the tree (Litz, 1987). Fruits that are
approximately 3040 days after pollination contain seeds in which the nucel-
lus is at the ideal stage for explanting: relatively thick and intact and easily
removed. The embryo (mass) in fruit and developing seed of this age is usu-
ally no more than half the length of the immature seed. Mango fruit of the appro-
priate stage of development are surface-disinfested with 2030% (v/v) domestic
Biotechnology 645
bleach containing Tween 20 for 30 min. The sterilant is rinsed from the fruit
with three changes of sterile deionized or distilled water, and each fruit is
bisected along its longitudinal axis without damaging the immature seed
under axenic conditions. The immature seed is removed from the bisected
fruit, which is also bisected carefully along its longitudinal axis. Manza-
nilla Ramirez et al. (2000) obtained optimum induction with polyembryonic
Ataulfo, monoembryonic Tommy Atkins and monoembryonic Haden
nucellar explants when the embryo (mass) to immature seed ratio was 1:3.
The zygotic embryo (of a monoembryonic cultivar) or polyembryonic mass
(of a polyembryonic cultivar) is excised and discarded. The nucellus can be
carefully peeled from the interior of the seedcoat using a sterile, at spatula.
Following the transfer of the nucellus onto induction medium in sterile Petri
dishes, the cultures are incubated in darkness at 25C. Thereafter, it is essen-
tial to subculture the nucellar explants onto fresh induction medium at daily
intervals until the oxidation of the explant ceases; oxidation is associated
with darkening of the medium around the explanted nucellus.
A summary of those mango cultivars that have been successfully estab-
lished as embryogenic cultures is provided in Table 18.2. The efciency of
Table 18.1. Summary of published reports on somatic embryogenesis of mango
from nucellar explants.
Country Reference
Colombia Flrez-Ramos et al. (2007)
India Ara et al. (1998, 2000a, b)
Chaturvedi et al. (2004)
Deore et al. (2000)
Jana et al. (1994)
Krishna and Singh (2007)
Laxmi et al. (1999)
Singh et al. (2001, 2002)
Sulekha and Rajmohan (2003)
Thomas (1999)
Mexico Manzanilla Ramirez et al. (2000)
Rivera Domnguez et al. (2004)
Philippines Patena et al. (2002)
USA DeWald et al. (1989a, b)
Lad et al. (1997)
Litz (1984)
Litz and Gomez-Lim (2005)
Litz and Lavi (1997)
Litz and Schaffer (1987)
Litz and Vijayakumar (1988)
Litz et al. (1982, 1983, 1984, 1995, 1997, 1998)
Mathews et al. (1992)
Monsalud et al. (1995)
Pliego Alfaro et al. (1996b)
R.E. Litz et al. 646
induction is cultivar-dependent. For example, Litz et al. (1998) compared the
induction of four mango cultivars, polyembryonic Hindi, monoembry-
onic Lippens, polyembryonic Nam Doc Mai and monoembryonic Tommy
Atkins, and observed that Hindi has the highest embryogenic response,
followed by Lippens, Tommy Atkins and Nam Doc Mai in descending
order. Manzanilla Ramirez et al. (2000) compared the induction responses of
three mango cultivars and observed that polyembryonic Ataulfo was more
embryogenic than either monoembryonic Tommy Atkins or monoembry-
onic Haden in descending order. Optimization of conditions for induction
of embryogenic mango cultures using polyembryonic James Saigon and
polyembryonic Parris as models was described by DeWald et al. (1989a).
The current induction procedure has been only slightly modied since then,
and utilizes a basal medium consisting of B5 (Gamborg et al., 1968) major salts
without ammonium sulfate ((NH
4
)
2
SO
4
), MS minor salts and organic compo-
nents, 60 g/l sucrose, 400 mg/l glutamine, 2.44.8 M 2,4-dichlorophenoxy-
acetic acid (2,4-D) and 2.0 g/l gellan gum. Patena et al. (2002) modied the
standard induction medium by supplementing it with 100 mg/l coconut
water (CW) to control oxidation of the explants.
The auxin 2,4-D has a temporal effect on induction of embryogenic
competence of explanted nucellus of polyembryonic Carabao (Lad et al.,
1997). Induction requires a minimum of 714 days exposure to 2,4-D and a
maximum exposure period of 56 days; acquisition of embryogenic compe-
tence is optimum after approximately 28 days exposure to 2,4-D (Plate 86).
Nurse cultures, which consist of highly embryogenic mango cultures (e.g.
polyembryonic Hindi) have been effective for stimulating induction of
Table 18.2. Induction of somatic embryogenesis from nucellar cultures of mango
(Source: Jana et al., 1994; Litz and Lavi, 1997; Manzanilla Ramirez et al., 2000;
Ara et al., 2000b; Chaturvedi et al., 2004).
Cultivar Seed type Cultivar Seed type Cultivar Seed type
Alphonso Mono Gedong Poly Mulgoa Mono
Ambalavi Poly Golek Poly Mundan Mono
Amrapali Mono Heart Poly Nam Doc Mai Poly
Arumanis Poly Hindi Poly Neelum Mono
Ataulfo Poly Honc Cambodiana Poly Ono Poly
Baneshan Mono Irwin Mono Parris Poly
Brander Poly James Saigon Poly Peach Poly
Brooks Mono Keitt Mono Philippine Poly
Cambodiana Poly Kensington Pride Poly Sabre Poly
Carabao Poly Kur Poly Simmonds Poly
Chausa Mono Langra Benarsi Mono Tommy Atkins Mono
Chino Poly Lippens Mono Tuehau Poly
Dashehari Mono Madu Poly Turpentine Poly
Everbearing Mono Manzano Poly White Langra Mono
Florigon Poly Mikongenesis Poly
Biotechnology 647
embryogenic competence from the nucellus of cultivars that are normally
difcult to induce, for example polyembryonic Nam Doc Mai (Litz et al.,
1998). The nurse culture procedure involves explanting the nucellus onto
sterile lter paper which has been moistened with induction medium and
which overlays the highly embryogenic mango culture (polyembryonic
Hindi) on semi-sterile induction medium. It is not clear if a nucellar callus
is initiated from the explant prior to acquisition of embryogenic competence;
however, somatic embryos can develop directly from the nucellus without
an intermediate callus (Litz, 1987). Embryogenic nucellar cultures are recog-
nizable ap-proximately 30 days after explanting; they are completely orga-
nized, and consist of proembryonal somatic embryos, embryogenic cells, cell
aggregates and proembryonic masses (PEMs) (Litz et al., 1993, 1995; Litz and
Lavi, 1997).
Maintenance
Embryogenic mango cultures are friable and cream to light brown in
colour, although the cultures rapidly darken on semi-solid medium, and
must therefore be subcultured at 34 week intervals. PEMs develop from
globular somatic embryos in the presence of the primary induction agent,
2,4-D. The PEMs originate as globular somatic embryos, but their devel-
opment as individual somatic embryos is arrested in the presence of 2,4-D.
The PEMs increase in diameter with cells of the protoderm dividing rap-
idly; secondary globular somatic embryos develop from these proliferating
embryogenic cells. This highly repetitive pattern of somatic embryogen-
esis in the presence of 2,4-D is the basis for maintenance of embryogenic
cultures.
Embryogenic mango cultures can be maintained as proliferating PEMs
either on semi-solid or in liquid induction medium. Maintenance of embryo-
genic cultures of many cultivars is optimal in liquid induction medium sup-
plemented with 4.8 M 2,4-D (Litz et al., 1984; DeWald et al., 1989a) (Plate 87);
however, rapid proliferation of embryogenic suspension cultures is cultivar-
dependent (Litz et al., 1993). Embryogenic suspension cultures are initiated
by inoculating approximately 400 mg of PEMs into sterile 80 ml maintenance
medium in 250 ml Erlenmeyer asks (or 200 mg of PEMs into 40 ml mainte-
nance medium in 125 ml Erlenmeyer asks). The asks are maintained on a
rotary shaker at 100125 rpm in semi-darkness at 25C with regular transfers
of PEMs into fresh medium at 1014 day intervals. Regular subculturing is
essential to prevent loss of morphogenic potential and darkening of the tis-
sue. The typical embryogenic suspension culture consists of PEMs, embryo-
genic cells and multicellular complexes.
Maturation
Development of somatic embryos from embryogenic cultures maintained on
semi-solid maintenance medium occurs sporadically and without synchroni-
zation, due to the lack of direct contact of parts of a culture with medium
containing 2,4-D. Exposure to 2,4-D is necessary for embryogenic culture
proliferation, while at the same time, somatic embryo development is
R.E. Litz et al. 648
inhibited. In order to stimulate somatic embryo development, embryogenic
cultures must be transferred from maintenance medium to medium without
2,4-D in order to initiate efcient somatic embryo development. For embryo-
genic suspension cultures, the PEMs are decanted through sterile ltration
fabric with a 1000 m opening size. The larger fraction (>1000 m diameter)
is re-inoculated into liquid maintenance medium for continued proliferation
and the smaller fraction is transferred either into liquid medium or onto
semi-solid medium without 2,4-D in order to arrest repetitive somatic
embryogenesis and to initiate somatic embryo development (Plate 88).
The plant growth media and conditions for stimulating somatic embryo
development and maturation are based upon the protocol described by DeWald
et al. (1989b) with minor alterations. Initially, a medium consisting of B5 major
salts, MS minor salts and organic components, 60 g/l sucrose and 400 mg/l
glutamine with or without 2.0 g/l gellan gum is utilized. Different mango
cultivars require different periods for cotyledon differentiation following
subculture to maturation medium (Litz et al., 1993). Either 4.65 M kinetin or
4.44 M benzyladenine (BA) can stimulate the development of cotyledons
and the apical meristem and reduce the maturation period. The cultures are
incubated in darkness at 25C.
When growth of embryogenic cultures of highly responsive cultivars is
optimized as suspension cultures, the early cotyledonary somatic embryos
that develop in liquid medium are hyperhydric. Mathews et al. (1992) and
Monsalud et al. (1995) demonstrated that hyperhydric somatic embryos can-
not develop to maturity, and become necrotic. Hyperhydricity of mango
somatic embryos can be reversed either by partially desiccating heart stage
embryos (23 mm length) under high relative humidity (RH) for 24 h or by
plating them on maturation medium solidied with 6.0 g/l gellan gum
(Monsalud et al., 1995). Reversion of hyperhydricity can result in precocious
germination of mango somatic embryos, although this can be prevented if
500 M abscisic acid (ABA) is included in the modied maturation medium.
Embryogenic cultures of different mango cultivars vary in their response
to subculture from liquid maintenance medium onto maturation medium,
and this must be determined empirically for each cultivar. For example, poly-
embryonic Hindi somatic embryo development (cotyledon differentiation)
from embryogenic suspension cultures follows a step-wise procedure. The
<1000 m diameter fraction from liquid maintenance is subcultured into
liquid maturation medium until the early heart stage of somatic embryo
development is apparent. Subsequently, the heart stage somatic embryos are
subcultured onto semi-solid maturation medium. In contrast, the <1000 m
fraction of monoembryonic Keitt and polyembryonic Carabao embryogenic
suspension cultures can be transferred directly from liquid maintenance
medium onto semi-solid maturation medium.
Mango zygotic embryos require approximately 45 months in order to
develop to maturity in vivo, and mature embryos can be >68 cm long (Plate
89). Consequently, the plant growth media that have been formulated to
stimulate growth and development of mango somatic embryos from the
heart stage to maturity reect the differing requirements of these enlarging
Biotechnology 649
somatic embryos. The medium that has been utilized for development of
mango somatic embryos to maturity consists of B5 major salts, MS minor
salts and organic components, 400 mg/l glutamine, 20% (v/v) lter-sterilized
CW, 40 g/l sucrose and 2.0 g/l gellan gum (DeWald et al., 1989b). As the
somatic embryos enlarge, the sucrose concentration of maturation medium is
gradually reduced to 10 g/l.
During somatic embryo development certain developmental anomalies
become apparent, of which the most frequently observed are polycotyly, fas-
ciation, absence of bipolarity, secondary somatic embryogenesis from the
hypocotyl and precocious germination. Polycotyly and fasciation do not
affect subsequent plant development; however, failure to address problems
associated with absence of bipolarity, secondary embryogenesis and preco-
cious germination can seriously impact the recovery of plants. Control of
precocious germination of developing embryos is occasionally necessary,
since immature somatic embryos that germinate before they are physiologi-
cally mature cannot survive. Some of these developmental anomalies can be
eliminated by maintaining relatively high sucrose concentrations and/or by
incorporating 100 M ABA in the maturation medium (Monsalud et al., 1995;
Pliego-Alfaro et al., 1996b). The cultures are maintained in darkness at 25C
during somatic embryo maturation.
Germination
When somatic embryos begin to germinate, they are nally transferred to
light conditions. The radicle elongates, followed by growth of the taproot.
The shoot apical meristem remains quiescent for approximately 2 weeks
after germination, at which time the shoot elongates. Although many mango
somatic embryos germinate under these conditions, their survival or conver-
sion ex vitro is low, primarily due to apical shoot necrosis, a physiological
disorder that is associated with calcium ion (Ca
++
) deciency. Different strat-
egies have been attempted to improve the conversion rate (i.e. survival of
somatic embryo-derived plants):
The period for embryogenic cultures in/on maintenance medium should 1.
be minimal (Litz and Lavi, 1997).
Ara 2. et al. (1998) rescued in vitro microshoots obtained from germinated
somatic embryos by pulsing them for 24 h with 24.6 M IBA in liquid medi-
um followed by transfer to auxin-free medium in darkness for root develop-
ment.
Somatic embryo shoots have also been rescued by micrografting the 3.
shoots on decapitated in vitro-germinated seedling rootstocks (Plate 90).
Enhanced recovery of mango plantlets can occur following the induction 4.
of photoautotropism by transfer of small plantlets onto minimal plant growth
medium, containing <5% sucrose and 1% (w/v) activated charcoal (Litz et al.,
1993). A lter-sterilized air mixture consisting of 20,000 ppm carbon dioxide
(CO
2
) in a nitrogen gas carrier is introduced into the growing containers, and
the plantlets are exposed to a 16 h photoperiod at 180 mol/s/m
2
provided
by cool white uorescent tubes.
R.E. Litz et al. 650
Ara 5. et al. (1999) reported improved conversion following the encapsulation
of early heart stage somatic embryos in calcium alginate-containing modied
standard mango medium with half-strength major salts and supplemented
with 2.9 M gibberellic acid (GA
3
). The germination of encapsulated somatic
embryos was almost 75% greater than non-encapsulated somatic embryos.
Protoplast isolation and culture
The isolation, culture and regeneration of plantlets from protoplasts isolated
from embryogenic suspension cultures of monoembryonic Amrapali has been
described (Ara et al., 2000a). One gram of embryogenic culture was transferred
from a 34-week-old suspension into 10 ml lter-sterilized growth medium
consisting of B5 major salts, MS minor salts and organic components, supple-
mented with 0.3 M sucrose, 0.4 M mannitol, 0.1 M sorbitol, 2.74 mM glutamine,
1.0% cellulase, 1.0% hemicellulase and 0.5% pectinase (Sigma) with gentle shak-
ing in darkness at 25C for 24 h. The digestion mixture was then passed through
a sieve (50 m) in order to remove debris, and then centrifuged for 5 min at
100 g. The supernatant was discarded, and cell debris was resuspended and
precipitated by centrifugation three times at 3 min for each centrifugation cycle.
The pellet was nally resuspended in 1 ml medium, and layered on 3 ml sucrose
solution (25% w/v) and centrifuged at 100 g for 7 min. Protoplasts were removed
and cultured in maintenance medium, but modied to contain 0.18 M sucrose,
2.74 mM glutamine and 4.5 M 2,4-D. Somatic embryos developed from PEMs
following subculture onto somatic embryo development medium (without
2,4-D), and plantlets were recovered using standard procedures (see Somatic
embryogenesis section in 18.2 Cell and tissue culture, this chapter).
Protoplast technology has not been utilized at this time for mango
improvement, and the likelihood of its exploitation is difcult to predict. The
sexual compatibilities of Mangifera spp. with the common mango are unknown.
Many newly described Mangifera species tolerate stressful environmental
conditions and have pest and disease resistance (see Bompard, Chapter 2,
this volume); however, it is uncertain if they are isolated genetically from
mango. Somatic hybridization might enable the genetic recombination of the
common mango with some of the Mangifera species as a means for develop-
ing new rootstocks. This approach, however, would almost certainly have
little utility for scion development, since the common mango is a tetraploid,
and the ploidy level of many of the Mangifera spp. is unknown. Somatic
hybridization would result in hexaploids or octoploids, and introgression of
useful traits into the common mango would be very difcult.
Potential for other regeneration pathways
The recovery of embryogenic haploid cultures from microspores has been
described for several perennial fruit tree species, including Annona squamosa
(Nair et al., 1983), Citrus aurantifolia (Chaturvedi and Sharma, 1985), Citrus
Biotechnology 651
microcarpa (Chen et al., 1980), Dimocarpus longan (Yang and Wei, 1984) and
Litchi chinensis (Fu and Tang, 1983). Mangifera indica is thought to be an auto-
tetraploid (2n = 4x = 40), and recovery of diploid (n = 2x = 20) plants would
reveal the nature of the ancestral species and simplify genomic analysis.
Chromosome doubling would restore autotetraploidy, and somatic hybrid-
ization of diploid mangoes with wild Mangifera species would allow interest-
ing genetic recombination.
18.3 Molecular Breeding and Genetics
The identication and use of molecular markers for distinguishing mango
from other Mangifera spp. and for identifying different mango cultivars has
been discussed by Bompard, Chapter 2, this volume and by Iyer and Schnell,
Chapter 4, this volume.
Marker assisted selection (MAS)
MAS offers great potential for improvement of quantitative traits in crop
plants. There are clear advantages for the use of molecular markers in plant
breeding, such as a decreased number of breeding generations, the availabil-
ity of a uniform method for scoring, no need to use phenotypic scoring until
the end and, nally, the possibility for obtaining information on the percent-
age of genome contributed by each parent in the offspring. Although molecu-
lar markers have been used for taxonomic purposes with mango (Schnell et al.,
1995; Lpez-Valenzuela et al., 1997; Eiadthong et al., 2000; Ravishankar et al.,
2000; etc.) mango has not been the subject of MAS. It is notable that although
mango has 40 chromosomes, it has a comparatively small haploid genome size
(0.91 pg), which is only three times as large as the recently sequenced genome
of A. thaliana (the plant with the smallest genome size known), about half that
of tomato and comparable to that of rice (Arumuganathan and Earle, 1991).
Clearly, the comparatively small mango genome would facilitate the identi-
cation of molecular markers and the creation of a genetic map.
Gene cloning
Genetic transformation of mango cultures involves the transfer of genes to
manipulate specic processes. Genetic manipulation of any crop requires
that relevant genes must be available. Mango has been the subject of molecu-
lar research for several years, and several genes have been identied.
Early studies showed that changes in mRNA and protein content occur
during fruit ripening (Lpez-Gmez and Gmez-Lim, 1992; Chaimanee et al.,
1999). The level of a number of mRNAs (as assayed by in vitro translation)
changes throughout the ripening process. This method for detecting changes
in mRNAs has low sensitivity and, therefore, only the most abundant proteins
R.E. Litz et al. 652
can be detected. For that reason, the molecular analysis of fruit ripening
requires construction of a gene library. In mango, cDNA libraries have been
constructed, mainly from ripe fruit, and screened using several approaches.
The mRNA for virtually all the ripening-related genes isolated so far have
been shown to be absent or at a low level in immature fruit, increase during
ripening and decline as ripening progresses (Giovannoni, 2001). Unlike other
fruits, none of the identied genes in mango code for enzymes involved in
the ripening process itself (see below).
Mango fruit are highly perishable commodities due to over-ripening,
which is mainly caused by the sharp increase in ethylene production, which
occurs simultaneously with the climacteric peak (Tucker and Grierson, 1987;
see Brecht and Yahia, Chapter 14, this volume). Mangoes have poor storage
quality and storage in controlled or modied atmospheres has been associ-
ated with physiological disorders (Chaplin, 1989). Genetic manipulation of
mango fruit ripening represents an attractive alternative to extend storage
life and, therefore, the isolation of mango genes coding for enzymes involved
in ethylene biosynthesis has been a target for research.
The two key enzymes in the ethylene biosynthetic pathway are those
catalysing the conversion of S-adenosylmethionine (SAM) to 1-aminocyclo-
propane-1-carboxylic acid (ACC) and ACC to ethylene, i.e. ACC synthase
and ACC oxidase or ethylene forming enzyme (EFE), respectively and cDNA
clones coding for mango ACC synthase and ACC oxidase have been identi-
ed (Gmez-Lim, 1993). Their expression during ripening was studied in
pulp and peel in northern blot analysis-type experiments. The ACC synthase
message is undetectable in unripe fruit and starts to appear in turning fruit,
reaching a maximum in ripe fruit (Gmez-Lim, 1993). This pattern of expres-
sion is similar in the peel and in the pulp; however, the message appears in
the pulp before the peel. The ACC oxidase message shows similar kinetics in
both types of tissue, but the message is clearly detectable before any ACC
synthase message becomes detectable (Gmez-Lim, 1993). These results sug-
gest that ACC oxidase is expressed before ACC synthase and that ripening
starts on the inside of mango fruit and proceeds outwards. Ethylene-treated
mango fruits show a different pattern of expression, with ACC oxidase and
ACC synthase appearing initially in the peel.
If ethylene is being actively produced, the gas must be clearly perceived
by plant cells and therefore ethylene sensitivity is important for the ripening
process. Major advances in understanding ethylene signal transduction have
come from a molecular genetic approach using ethylene responsive mutants
of A. thaliana and tomato. Several genes coding for ethylene receptor homo-
logues have been isolated from various plants (Johnson and Ecker, 1998).
Genetic manipulation of the ethylene receptor represents an interesting alter-
native to control ethylene production and delay ripening, particularly in
those fruits where other alternatives, such as storage in controlled atmo-
spheres, have not been effective (Wilkinson et al., 1997). A cDNA coding for a
mango ethylene receptor has been isolated (Gutirrez-Martinez et al., 2001).
The message seems to be present at low levels in unripe fruit and to increase
Biotechnology 653
as the fruit ripens. Mechanical wounding appears to up-regulate the expres-
sion of the receptor.
Several studies have convincingly shown that the prole of released
volatiles changes as ripening proceeds (Olle et al., 1998; Sakho et al., 1998;
Ansari et al., 1999; Saby John et al., 1999; Andrade et al., 2000; Bender et al.,
2000). It is likely that many of these compounds, most of which are terpe-
noids, are determinants of avour and aroma and many of them originated
from the metabolism of fatty acids via the -oxidation pathway. It is known
that several fatty acids, particularly linoleic and oleic acids, decrease in con-
centration during ripening. In this sense, a cDNA for thiolase, an enzyme
from the -oxidation pathway, was identied as having a ripening-specic
pattern and to be up-regulated during ripening (Bojrquez and Gmez-Lim,
1995). A cDNA for acyl CoA oxidase, the key enzyme in the -oxidation path-
way, has isolated from mango fruit and shown to behave similarly to thiolase
(A. Nila-Mendez and M. A. Gmez-Lim, Irapuato, unpublished data). These
enzymes might be involved in metabolism of fatty acids to produce volatile
compounds. The contents of sugars and organic acids can inuence the a-
vour properties of mango (Malundo et al., 2001).
The manipulation of fruit aroma and avour is a long established research
goal and, accordingly, the isolation of genes coding for enzymes involved in
biosynthesis of these compounds has been targeted. The gene coding for
alcohol acyl transferase, an enzyme presumably involved in the synthesis of
compounds implicated in fruit avour, has been identied in mango (GB:
AX025510; patent WO 0032789-A 36).
Alternate oxidase is involved in the cyanide-resistant respiratory path-
way. It has been studied mainly in thermogenic species, and its activity is
correlated with heat production, necessary to volatilize foul-smelling com-
pounds to attract insect pollinators. There is a signicant participation of this
pathway in the climacteric of many fruit. A cDNA coding for mango alter-
nate oxidase has been isolated and the message was detected by Northern
blot analysis in unripe fruit and shown to increase substantially in ripe fruit
(Cruz-Hernandez and Gmez-Lim, 1995). These results were correlated with
similar increases in enzyme activity and protein accumulation. The tempera-
ture in ripe monoembryonic Alfonso fruit is up to 10C higher than in
unripe fruit and this has been attributed to the activity of alternate oxidase
(Kumar et al., 1990). This extra heat might also serve to volatilize aroma-
giving compounds. The results with alternate oxidase were conrmed by
Considine et al. (2001), who isolated several members of the multigene family
of mango alternate oxidase and showed that they were expressed differen-
tially during ripening. They also identied a gene coding for an uncoupling
protein whose mRNA peaked at the turning stage whereas the protein peaked
at the ripe stage (Considine et al., 2001). They suggested a role for alternate
oxidase and the uncoupling protein in post-climacteric senescence. Because
both mRNA and protein for alternate oxidase and the uncoupling protein
increased in a similar pattern, they hypothesized that their expression is con-
trolled simultaneously.
R.E. Litz et al. 654
Ripening of fruit involves a number of metabolic reactions, including
synthesis and turnover of the plant cell wall. In mango, fruit ripening is char-
acterized by a gradual softening, which is caused by progressive depolymer-
ization of pectic and hemicellulosic polysaccharides with signicant loss of
galactose, arabinose and mannose residues at the ripe stage (Yashoda et al.,
2005). This depolymerization is caused by the activity of different hydrolytic
enzymes that are secreted as ripening proceeds (i.e. polygalacturonase and
-galactosidase), which are present in different isoforms in mango pulp
(Prasanna et al., 2005, 2006). Cell wall-metabolizing enzymes from mango, i.e.
exopolygalacturonase (Chaimanee et al., 2000), endo-polygalacturonase (Sun-
tornwat et al., 2000), -galactosidase (S. Parra-Arenas and M. A. Gmez-Lim,
Irapuato, unpublished data) and pectate lyase (Chourasia et al., 2006), have
been cloned and correlate closely with fruit ripening. Furthermore, an ethyl-
ene-responsive, ripening-related expansin has also been identied as being
closely correlated with softening (Sane et al., 2005).
A cDNA clone coding for a homologue of the YPT/Rab class of small
Guanosine-5-triphosphate (GTP)-binding proteins has been identied from
mango and is induced during ripening (Zainal et al., 1996). These proteins
appear to control the secretion of other proteins as well as the fusion of mem-
branes in animal cells (Fischer von Mollard et al., 1994). Some of these small
GTP-binding proteins have homologues in plants, albeit their role has not
been well dened (Staehelin and Moore, 1995). It is tempting to speculate
that, based on the pattern of expression, these small GTP-binding proteins
facilitate secretion of various hydrolytic enzymes during fruit ripening.
Saiprasad et al. (2004) isolated ve ripening-related cDNAs by reverse
transcription-polymerase chain reaction (RT-PCR). They showed similarity
to PRl-1 protein, a transcription initiation factor, a CCR-4 protein and to an
18S ribosomal RNA gene and a 23S ribosomal RNA gene. Likewise, Lycett
et al. (1997) identied by differential screening six clones up-regulated dur-
ing ripening. Unfortunately, none of these clones code for proteins directly
associated with ripening. Only two of them showed homology to a plastid
chromatin and Ypt/Rab11 class of small GTPase, respectively. I A tomato
rab11-like cDNA was used for tomato transformation in antisense (Lu et al.,
2001). The fruits changed colour as expected but failed to soften normally.
This was accompanied by reduced levels of two cell wall hydrolases, pectin-
esterase and polygalacturonase. There were other phenotypic effects in the
plants, including determinate growth, reduced apical dominance, branched
inorescences, abnormal oral structure and ectopic shoots on the leaves. In
some plants, ethylene production was reduced. These data suggest an
additional role for the gene in other ripening processes.
Vasanthaiah et al. (2006) reported the characterization of 27 differentially
expressed genes during mango internal breakdown and found catalase,
superoxide dismutase, ubiquitin, alcohol dehydrogenase, coproporphyrino-
gen oxidase and keratin-associated protein to be up-regulated; whereas,
several ribosomal genes, cysthathionine gamma synthase and fructose bispho-
sphate aldolase were down-regulated. The increased expression of catalase,
coproporhyrinogen III oxidase and keratin genes during internal breakdown
Biotechnology 655
was interpreted as a sign of oxidative stress, which they hypothesized as one
of the probable causes for this disorder.
In addition to these genes, several mango sequences have been reported
in the GenBank. These include a genomic sequence for the large subunit of
ribulose 1,5-bisphosphate carboxylase (U39269), cDNAs for two unidentied
clones (AF370123 and AF061639), a partial cDNA for LFY (AY189684), a cDNA
for xyloglucan endo-transglycosylase (GenBank accession AY600965) and a
putative Pto-like serine/threonine kinase gene (AY693369). The function of
these sequences in mango development remains to be determined.
There have been several reports on therapeutic properties of extracts
from mango leaves or seeds, including anti-inammatory effects (Beltrn
et al., 2004; Garrido et al., 2004; Leiro et al., 2004), anthelminthic and antial-
lergic properties (Garca et al., 2003a), anti-diarrhoeal properties (Sairam
et al., 2003) and even an enhancement of the humoral immune response in
mouse (Garcia et al., 2003b). It is likely that the constituents responsible for
these therapeutic effects will be the subject of intense scrutiny and, if the
right genetic elements are identied, the genetic manipulation of the biosyn-
thetic route may become a reality.
Genomics
The human genome project has been the catalyst for the development of sev-
eral high-throughput technologies that have made it possible to map and
sequence complex genomes. Many bacterial genomes and the genomes of
Sacchromyces cerevisiae, Caenorhabditis elegans, Drosophila melanogaster, A. thali-
ana, Oriza sativa and Populus trichocarpa have been fully sequenced. In addition,
the National Center for Biotechnology Information Entrez Genome Projects
web site reports that sequencing of several more plant genomes is in prog-
ress. Nevertheless, the completion of the entire genomic sequence of a par-
ticular organism represents the end of the structural genomics segment of the
project. It is clear, therefore, that the identication of every gene within the
genomes of model organisms is only the initial step to understand what these
genes do and how they interact to make up a living organism. Understand-
ing the functions of the 20,00050,000 genes comprising plant (and animal)
genomes and the variations within a population and roles in normal devel-
opment will represent a possibly greater task than the mapping and sequenc-
ing efforts currently underway.
Understanding the function of genes and other parts of the genome is
known as functional genomics, and research has involved model organisms
such as A. thaliana and rice. Model organisms offer a cost-effective way to
follow the inheritance of genes through many generations in a relatively
short time. Functional genomics is characterized by high throughput or
large-scale experimental methodologies combined with statistical and com-
putational analysis of the results. The fundamental strategy in a functional
genomics approach is to expand the scope of biological investigation from
studying single genes or proteins to studying all genes or proteins at once in
R.E. Litz et al. 656
a systematic fashion. Computational biology will perform a critical and
expanding role in this area. Whereas structural genomics has been character-
ized by data management, functional genomics will be characterized by min-
ing the data sets for particularly valuable information. Functional genomics
promises to rapidly narrow the gap between sequence and function and to
yield new insights into the behaviour of biological systems. The essential
requirement for the implementation of functional genomics is the availability
of identied sequences, which can be subsequently used in studies such as
high-density arrays. Currently, there are very few sequenced genes from
mango, and these are mainly fruit-specic, and an effort to increase this num-
ber is needed. Considering the cDNA libraries prepared from ripe fruit in
various laboratories around the world, an effort to determine expressed
sequence tags (ESTs) would be worthwhile. At the same time, more cDNA
libraries from tissues other than fruit are needed. At present, the technology
for successful application of functional genomics to mango is well devel-
oped, but the raw material (i.e. identied genes) is lacking. Nevertheless,
over 1000 ESTs from ripe mango pulp have been sequenced and are in the
process of being analysed by microarrays using RNA probes from different
ripening stages and from different mango cultivars (G. Ramirez Zavala and
M.A. Gmez-Lim, Irapuato, unpublished results).
On the other hand, proteomics, the large-scale study of protein structure
and function, is still to be applied to mango. Proteomics is often considered
the next step after genomics but in fact it is more challenging, because a pro-
teome differs from cell to cell and constantly changes through its biochemical
interactions with the genome and the environment. The Human Genome
Project has revealed that there are far fewer protein-coding genes in the
human genome than proteins in the human proteome (20,00025,000 genes
versus >500,000 proteins). The protein diversity is thought to be due to alter-
native splicing and post-translational modication of proteins and protein
degradation (Reddy, 2007). Clearly, this discrepancy implies that protein
diversity cannot be fully characterized by gene expression analysis alone.
Thus proteomics may be more useful for characterizing cells and tissues.
18.4 Genetic Engineering
In vitro induced mutations
There is signicant consumer resistance to replacement of local mango cultivars
by newer selections in many traditional mango-producing countries of South
and South-east Asia. Serious production and postharvest problems that have
a genetic cause (e.g. alternate and irregular bearing, susceptibility to anthra-
cnose and mango malformation, tree shape and size, etc.) cannot be addressed
in the long term using applied physiology. For example, paclobutrazol has
been applied to alternate-bearing mango cultivars to promote owering in
the off-years; however, this can result in severe decline in trees that have been
treated for successive years. As a result, conventional mango breeding in
Biotechnology 657
India has focused upon the development of mango cultivars that are largely
indistinguishable from traditional selections with respect to fruit size, appear-
ance, taste, avour and overall quality.
Despite its impact on crop breeding (Maluszynski, 2001) and particularly
on banana improvement (Novak et al., 1990), mutation breeding has not been
successfully exploited for mango cultivar improvement by production of use-
ful off-types of existing selections. There is some anecdotal evidence that
somatic mutations can occur naturally in mango on the basis of variation that
occurs within seed-propagated polyembryonic cultivars. There are reported
to be marked phenotypic differences within polyembryonic Kensington Pride
trees in Australia, and among polyembryonic cultivars of South-east Asia
(e.g. Arumanis, Golek, etc.). Different phenotypes of polyembryonic Aru-
manis, for example, have been characterized as Arumanis-1, Arumanis-2,
etc. Gan et al. (1981) and Litz et al. (1993) described isozyme variation within
populations of South-east Asian polyembryonic mangoes.
The most important production and postharvest problem of mango in
the humid tropics and subtropics is anthracnose, caused by Colletotrichum
gloeosporiodes (Penz.) Penz. and Sacc. In Penz. The current strategies for con-
trol of this disease involve the use of moderately resistant cultivars (i.e.
monoembryonic Calypso, monoembryonic Keitt and monoembryonic
Tommy Atkins, etc.) and at least weekly applications of fungicides (i.e.
benomyl or maneb or mancozeb) from the time of owering until harvesting
(Dodd et al., 1997). This can result in as many as 25 spray applications in
a season, and is considered an increasingly unsustainable agricultural
practice.
The effect of -irradiation on embryogenic cultures of polyembryonic
Hindi, monoembryonic Keitt and monoembryonic Tommy Atkins was
reported by Litz (2001) as part of the Food and Agriculture Organization
(FAO)/International Atomic Energy Agency (IAEA) Co-ordinated Research
Project entitled Improvement of Tropical and Subtropical Fruit Trees through
Induced Mutations and Biotechnology. Embryogenic mango cultures on semi-
solid maintenance medium were exposed to 0200 Gy irradiation provided
by
60
Co. The lethal dose for 50% mortality (LD
50
) for each cultivar was deter-
mined: it was approximately 125 Gy for Keitt and approximately 100 Gy for
Tommy Atkins. The LD
50
of embryogenic Hindi cultures could not be
established within this dosage range, possibly due to its high rate of prolif-
eration relative to Keitt and Tommy Atkins. This study was conrmed by
Manzanilla Ramirez et al. (2000). The main objective of these studies has been
to develop appropriate technologies for utilizing induced mutations in vitro
for recovery of useful off-types of existing cultivars.
Culture ltrates produced by pathogenic fungi and bacteria can be uti-
lized not only to select for resistance to the pathogen in vitro (Hammerschlag,
1992), but also to induce the host resistance response. In order for in vitro
selection to be used effectively, essential prerequisites include: (i) a highly
morphogenic suspension culture; and (ii) evidence that the culture ltrate or
phytotoxin can produce symptoms of the disease on plant organs as well as
with cells in suspension cultures. Litz et al. (1991) rst reported that the C.
R.E. Litz et al. 658
gloeosporiodes culture ltrate was effective as a selective agent in mango sus-
pension cultures growing in maintenance medium formulation. Somatic
embryos developed from embryogenic cells and PEMs that had survived
exposure to C. gloeosporiodes culture ltrate, and regenerants appeared to
show resistance to inoculation with the pathogen. Jayasankar et al. (1999)
characterized the in vitro effects of the puried C. gloeosporiodes phytotoxin,
colletotrichin (Gohbara et al., 1977, 1978), and crude C. gloeosporiodes culture
ltrate on the mortality and growth of polyembryonic Hindi and polyem-
bryonic Carabao embryogenic cultures. In this study, the LD
50
values for the
effects of C. gloeosporiodes culture ltrate and colletotrichin on embryogenic
cultures and the growth curves of challenged cultures were established.
Later, embryogenic cultures of the same cultivars were either exposed con-
tinuously for four cycles of challenge/selection/regrowth or were challenged
for one, two, three and four complete cycles with colletotrichin and the par-
tially puried culture ltrate of C. gloeosporiodes (Jayasankar and Litz, 1998).
At the end of each cycle, surviving PEMs were physically removed from the
selection medium, cloned and then either rechallenged or subcultured onto
somatic embryo maturation medium.
In order to induce the expression of anti-fungal genes in vitro, at least
three successive challenges with either crude ltrate or colletotrichin were
necessary. This was determined by co-culturing the challenged material with
a virulent strain of C. gloeosporiodes. Maintenance medium was inoculated
with challenged and selected embryogenic cultures at opposite sides of Petri
dishes. After 3 weeks, a virulent strain of the pathogen was inoculated in the
centre of each Petri dish. Co-culture of the pathogen with resistant cultures
resulted in the suppression of mycelium growth; the anti-fungal nature of the
PEMs increased with each cycle of challenge and selection, and was maxi-
mum after three cycles. There was enhanced production of extracellular anti-
fungal proteins chitinase and -1,3-glucanase in selected cultures. An
additional chitinase isozyme at 45 kDa was observed with anti-fungal Hindi
cultures and at 25 kDa with anti-fungal Carabao cultures, with respect to
the controls. The anti-fungal nature of selected, resistant lines in suspension
cultures and in somatic embryos was persistent for more than 2 years follow-
ing selection. Various random amplication of polymorphic DNA (RAPD)
markers were associated with selected cultures that were strongly anti-fun-
gal (Jayasankar et al., 1998). RAPD markers of the unchallenged controls and
of leaves from the parent trees were identical, indicating that exposure to
either colletotrichin or culture ltrate is essential for expression of anti-fungal
genes. The RAPD results also demonstrated that embryogenic cultures are
quite stable genetically, and the expression of anti-fungal genes is not the
result of somaclonal variation. Furthermore, it seems highly probable that
the phytotoxins themselves are highly mutagenic.
In vitro-induced mutation followed by selection could be a highly
efcient method for addressing specic breeding problems of mango assum-
ing that effective selection agents are available. Unfortunately, at this time,
there are relatively few such selection agents that can be utilized in this
manner.
Biotechnology 659
Genetic transformation
Genetic transformation is currently the only practical solution for improving
existing elite selections of perennial species for specic horticultural traits
and for investigating gene function by interference RNA. Transformation of
mango has been reviewed most recently by Litz and Gomez-Lim (2002, 2005)
and Gomez-Lim and Litz (2007).
General protocols
Mathews et al. (1992, 1993) rst reported the genetic transformation of mango
using embryogenic cultures of polyembryonic Hindi and of a monoembry-
onic Keitt zygotic embryo-derived embryogenic line, respectively. These two
studies utilized two different disarmed, engineered strains of Agrobacterium
tumefaciens: (i) strain C58C1 containing the plasmid pGV 3850::1103 with the
selectable marker gene for neophosphate transferase (NPTII) which confers
resistance to the antibiotic kanamycin, both of which were driven by the
CaMV constitutive 35S promoter (Mathews et al., 1993); and (ii) strain A208
containing the plasmid pTiT37-SE::pMON9749, a co-integrate vector, with
genes for NPTII and the scorable marker -glucuronidase (gus or uidA) with
the 35S promoter (Mathews et al., 1992). A report by Cruz Hernandez et al.
(1997) utilized A. tumefaciens strain LBA4404 containing NPTII, -glucuronidase
(GUS) and genes that mediate a horticulturally useful trait in binary plasmid
pBI121 with the CaMV 35S promoter. Mathews and Litz (1990) earlier had
demonstrated that 12.5 g/ml kanamycin sulfate is toxic to embryogenic
suspension cultures; whereas, much higher levels (200 g/ml kanamycin)
are toxic to embryogenic cultures that are grown on semi-solid medium.
These genetic transformation reports have followed a similar two-step
selection (Mathews et al., 1992; Cruz Hernandez et al., 1997). Embryogenic
suspension cultures in their logarithmic phase of growth are separated by
passing them through sterile ltration fabric (1000 m pore size), and the
large fraction (>1000 m) is abraded with a sterile brush on sterile lter
paper. The abraded PEMs are then incubated with acetosyringone-activated
A. tumefaciens for 3 days in liquid maintenance medium, with subculture
into fresh medium at 24 h intervals. The PEMs are then transferred onto
semi-solid maintenance medium supplemented with 200 mg/l kanamycin
sulfate and 500 mg/l cefotaxime. After 10 months on this selection medium,
the PEMs are transferred to semi-solid maintenance medium containing
400 mg/l kanamycin sulfate. Proliferating cultures are subcultured in liq-
uid maintenance medium containing 100 mg/l kanamycin sulfate, and
somatic embryo development is initiated by subculture onto semi-solid
maturation medium. Mathews et al. (1993) regenerated transgenic mango
plants derived from a Keitt zygotic embryo embryogenic culture and
which had been transformed with pGV 3850::1103 containing the selectable
marker gene nptII. Genetic transformation was conrmed by: (i) growth in
selection medium containing inhibitory levels of kanamycin sulfate; (ii)
positive histochemical reaction for GUS with X-GLUC (Jefferson, 1987); and
(iii) Southern hybridization.
R.E. Litz et al. 660
Transient gene expression in embryogenic polyembryonic Kensington
Pride and polyembryonic Carabao cultures has been described using a
biolistic approach using two vectors: (i) pBI426 with GUS-NPTII under the
control of a double CaMV 35S promoter; and (ii) pBINgfp-Ser, which con-
tains NPTII and the green uorescent protein gene (gfp) (Cruz Hernandez
et al., 2000).
Transformation with genes that mediate horticulturally signicant traits
Loss of mango fruit due to spoilage in storage and en route to markets
accounts for a signicant proportion of total production in many developing
countries that have poorly developed infrastructure (i.e. cold storage facili-
ties, poor roads, unreliable transportation, etc.) (see Brecht and Yahia, Chap-
ter 14, this volume) Mango has become an important export commodity for
several developing countries. Extended shelf life and absence of physiologi-
cal disorders that cause internal breakdown of fruit (e.g. soft nose and jelly
seed) of the most important export cultivars (e.g. monoembryonic Tommy
Atkins) are potentially very important, therefore, for the valuable export
trade and for domestic markets.
The mango is a climacteric fruit, and ethylene therefore is a critical regu-
lator of the biochemical processes that occur during ripening. Certain rate-
limiting genes that mediate ethylene production in mango have been cloned.
Cruz Hernandez et al. (1997) described the genetic transformation of embryo-
genic polyembryonic Hindi mango cultures with mango ACC oxidase, ACC
synthase and ACC alternative oxidase cloned in the antisense orientation
and under the control of the CaMV 35S constitutive promoter in the pBI121
binary vector in A. tumefaciens strain LBA4404. Embryogenic cultures were
transformed by the two-step procedure described above. Although the phe-
notype of the transformed lines was not reported, the genetic transforma-
tions were conrmed in each case by the XGLUC reaction for GUS, growth in
the presence of inhibitory levels of kanamycin sulfate, Southern blot hybrid-
ization and NPTII amplication by PCR. Successful regeneration of plants
and inhibition of ethylene production by mature mango fruit could possibly
resolve the production problem of premature ripening (jelly seed) and post-
harvest loss due to spoilage.
18.5 In vitro Conservation
Medium-term storage
Mango embryos cannot tolerate desiccation during maturation, and devel-
opment is not arrested; this is typical of recalcitrant seeds or embryos. With-
out a period of developmental arrest, mango embryos develop to maturity
and germinate in a continuous sequence. Mango seeds (and embryos) cannot
survive for more than 3 or 4 weeks under minimal in vitro growth conditions
(Parisot, 1988), and would perish after several days in a conventional seed
bank.
Biotechnology 661
Monsalud et al. (1995) demonstrated that 45 mm long somatic embryos
representing the late heart stage can be partially desiccated, and stored dry in
Petri dishes for more than 30 days without any loss of viability. On the other
hand, larger somatic embryos cannot survive partial desiccation. This study
has interesting implications for future studies that could focus on the concept
of an articial seed (i.e. somatic embryo) genebank for vegetatively propa-
gated tropical fruit trees.
Mango somatic embryos have been manipulated in order to induce
developmental arrest. Abscisic acid (ABA) is associated with the initiation of
developmental arrest and acquisition of desiccation tolerance of orthodox
type seeds and embryos. Abscisic acid causes developmental arrest in vitro at
relatively low concentrations for somatic embryos of the orthodox type (Bew-
ley and Black, 1985). Pliego Alfaro et al. (1996a, b) were able to arrest the
development of late heart stage polyembryonic Hindi somatic and nucellar
embryos with ABA at 100 M and higher concentrations in the maturation
medium. This strategy arrests growth and development for several months
so long as mango embryos are on ABA-containing maturation medium. ABA
also has a persistent residual effect, and mango somatic embryo develop-
ment was inhibited for approximately 1 month after their transfer onto matu-
ration medium without ABA. Increased osmolarity of the maturation medium
also inhibited somatic embryo development; however, there was no residual
effect following transfer of somatic embryos onto maturation medium with-
out osmoticum.
Long-term storage
Embryogenic mango cultures cannot be stored indenitely, and lose their
regeneration potential over time. The initiation of embryogenic cultures is
dependent on owering and fruit set, which is strictly related to environmen-
tal stimuli, and normally occurs one time each year or on alternative years with
alternate-bearing selections. Long-term storage of embryogenic mango cul-
tures is essential for genetic manipulation studies and will be increasingly
important for the management of genetic resources. Embryogenic mango cul-
tures have been cryopreserved by different procedures (Wu et al., 2003; Rajani
Nadgauda and Pamela Moon, Homestead, Florida, USA, personal communi-
cation). Wu et al. (2003) compared three cryopreservation protocols for embryo-
genic cultures derived from monoembryonic Zihua zygotic embryos:
encapsulation-dehydration, pregrowth-dehydration and vitrication. Encap-
sulation-dehydration was unsuccessful, and only limited survival (8.3%) was
obtained following desiccation of PEMs for 1 h to 58.5% moisture content prior
to freezing in liquid nitrogen. Vitrication, involving treatment of PEMs with
plant vitrication solution 2 (Sakai et al., 1991) for 20 min prior to freezing in
liquid nitrogen, was successful (94.3%). Embryogenic Hindi cultures have
also been introduced into cryogenic storage (Rajani Nadgauda and Pamela
Moon, Homestead, Florida, USA, personal communication) and somatic
embryos have been recovered from these cultures. Two procedures were
R.E. Litz et al. 662
successful: (i) stepwise cooling in which cryoprotected (5% DMSO and 5%
glycerol) embryogenic cultures were cooled in Mr Frosty containers at the
rate of 1C/min from room temperature (25C) to 75C followed by rapid
cooling to 196C; (ii) rapid cooling (vitrication). After cryovials were removed
from liquid nitrogen and rapidly warmed, cultures were thoroughly washed
with maintenance medium and plated on semi-solid maintenance medium.
Somatic embryo development was initiated by subculturing the PEMs on
somatic embryo maturation medium.
18.6 Conclusions
Biotechnology tools have great potential for mango, including advanced
micropropagation procedures, conservation and cultivar improvement. There
are also several strategies to genetically manipulate the crop for biotechno-
logical purposes. Genes in the antisense or sense orientation or RNAi (Small,
2007) can be utilized to inhibit specic genes. The increasing availability of
identied genes should facilitate a better understanding and genetic manip-
ulation of specic developmental processes.
Considering the large number of mango microsatellite sequences identi-
ed and reported in the GenBank, parameters such as heterozygosity, aver-
age gene diversity, frequency of outcrossing, cultivar relationships and a
mango genetic map should be determined in the near future. Correlations
between morphological and DNA markers together with a linkage map
should eventually enable MAS for mango improvement.
Most of the research quoted here has been carried out in a few centres;
however, it is now widely acknowledged that biotechnology is mainstream
research. It is hoped that more research centres will be involved.
References
Andrade, E.H.A., Maia, J.G.S. and Zoghbi, M. das G.B. (2000) Aroma volatile constitu-
ents of Brazilian varieties of mango fruit. Journal of Food Composition and Analysis
13, 2733.
Ansari, S.H., Ali, M., Velasco-Negueruela, A. and Prez-Alonso, M.J. (1999) Volatile
constituents of the fruits of three mango cultivars, Mangifera indica L. Journal of
Essential Oil Research 11, 6568.
Ara, H., Jaiswal, U. and Jaiswal, V.S. (1998) Rooting of microshoots of Mangifera indica
L. cv. Amrapali. Current Science 74, 240242.
Ara, H., Jaiswal, U. and Jaiswal, V.S. (1999) Germination and plantlets regeneration from
encapsulated somatic embryos of mango (Mangifera indica L.). Plant Cell Reports
19, 166170.
Ara, H., Jaiswal, U. and Jaiswal, V.S. (2000a) Plant regeneration from protoplasts of
mango (Mangifera indica L.) through somatic embryogenesis. Plant Cell Reports 19,
622627.
Ara, H., Jaiswal, U. and Jaiswal, V.S. (2000b) Somatic embryogenesis and plantlet regen-
eration in Amrapali and Chausa cultivars of mango (Mangifera indica L.). Current
Science 78, 164169.
Biotechnology 663
Aron, Y., Czosnek, H., Gazit, S. and Degani, C. (1998) Polyembryony in mango (Mangifera
indica L.) is controlled by a single dominant gene. HortScience 33, 12411242.
Arumuganathan, K. and Earle, E.D. (1991) Nuclear DNA content of some important
plant species. Plant Molecular Biology Reporter 9, 208218.
Beltrn, A.E., Alvarez, Y., Xavier, F.E., Hernanz, R., Rodriguez, J., Nunez, A.J., Alonso,
M.J. and Salaices, M. (2004) Vascular effects of the Mangifera indica L. extract
(Vimang). European Journal of Pharmacology 499, 297305.
Bender, R.J., Brecht, J.K., Baldwin, E.A. and Malundo, T.M.M. (2000) Aroma volatiles of
mature-green and tree-ripe Tommy Atkins mangoes after controlled atmosphere
vs. air storage. HortScience 35, 684686.
Bewley, J.D. and Black, M. (1985) Seeds: Physiology of Development and Germination.
Plenum, New York.
Bojrquez, G. and Gmez-Lim, M.A. (1995) Peroxisomal thiolase mRNA is induced
during mango fruit ripening. Plant Molecular Biology 28, 811820.
Broda, Z. (1984) Wegatatvymna popagacja koniczyny czerwonej (Trifolium pratense L.)
proprzez kultury tkankowe ze szczegolnym uwzglednieniem genetycznego uwar-
unkowania zdolnosci do regeneracji z kalusa. Rocsniki Akademii Rolniczej w Poz-
naniu Rozprawy naukowe 140, 541.
Chaimanee, P., Suntornwat, O., Lerrtwikool, N. and Bungaruang, L. (1999) Changes in
gene expression during mango fruit ripening, Journal of Biochemistry Molecular
Biology and Biophysics 3, 7579.
Chaimanee, P., Lertwikoon, N., Bungaruang, L. and Suntornwat, O. (2000) Exo-polygal-
actanase from ripe mango (Mangifera indica Linn. Cv Nam doc Mai). Acta Horticul-
turae 509, 171176.
Chaplin, G.R. (1989) Advances in postharvest physiology of mango. Acta Horticulturae
231, 639648.
Chaturvedi, H.C. and Sharma, A.K. (1985) Production of androgenic plants of Citrus
aurantifolia. Journal of Plant Physiology 119, 473477.
Chaturvedi, H.C., Agnihotri, S., Sharma, M., Sharma, A.K., Jain, M., Gupta, P., Chour-
asia, A. and Kidwai, N.R. (2004) Induced nucellar embryogenesis in vitro for clonal
multiplication of Mangifera indica L. var. Ambalavi: a dwarng rootstock. Indian
Journal of Biotechnology 3, 22228.
Chen, Z., Wang, M. and Liao, H. (1980) The induction of Citrus pollen plants in articial
media. Acta Genetica Sinica 7, 189191 (in Chinese).
Chourasia, A., Sane, V.A. and Nath, P. (2006) Differential expression of pectate lyase
during ethylene-induced postharvest softening of mango (Mangifera indica var.
Dashehari). Physiologia Plantarum 128, 546555.
Considine, M.J., Daley, D.O. and Whelan, J. (2001) The expression of alternative oxi-
dase and uncoupling protein during fruit ripening in mango. Plant Physiology 126,
16191629.
Cruz-Hernandez, A. and Gmez-Lim, M.A. (1995) Alternative oxidase from mango
(Mangifera indica, L.) is differentially regulated during fruit ripening. Planta 197,
569576.
Cruz-Hernandez, A., Gomez-Lim, M.A. and Litz, R.E. (1997) Transformation of mango
somatic embryos. Acta Horticulturae 455, 292298.
Cruz-Hernandez, A., Town, L., Cavallaro, A. and Botella, J.lR. (2000) Transient and sta-
ble transformation in mango by particle bombardment. Acta Horticulturae 509,
237242.
Deore, A.C., Chavan, S.S., Deshpande, R.S. and Dhonushke, B.L. (2000) Factors affecting
in vitro somatic embryogenesis from nucellar tissue of Alphonso mango. Annals of
Plant Physiology, 14, 182186.
R.E. Litz et al. 664
DeWald, S.G., Litz, R.E. and Moore, G.A. (1989a) Optimizing somatic embryo pro-
duction in mango. Journal of the American Society for Horticultural Science 114,
712716.
DeWald, S.G., Litz, R.E. and Moore, G.A. (1989b) Maturation and germination of man-
go somatic embryos. Journal of the American Society for Horticultural Science 114,
837841.
Dodd, J.C., Prusky, D. and Jeffries, P. (1997) Fruit diseases. In: Litz, R.E. (ed.) The Mango:
Botany, Production and Uses. CAB International, Wallingford, UK, pp. 257280.
Eiadthong, W., Yonemori, K., Kanzaki, S., Sugiura, A., Utsunomiya, N. and Subhadrab-
andhu, S. (2000) Amplied fragment length polymorphism analysis for studying
genetic relationships among Mangifera species in Thailand. Journal of the American
Society for Horticultural Science 125, 160164.
Fischer von Mollard, G., Stahl, B., Li, C., Sdhof, T.C. and John, R. (1994) Rab proteins
in regulated exocytosis. Trends in Biochemical Sciences 19, 164168.
Flrez-Ramos, C.P., Buitrago, M.E., Lentini, Z. and Cock, J. (2007) Somatic embryogen-
esis and plantlet regeneration of mango (Mangifera indica L.). Acta Horticulturae
738, 443445.
Fu, L. and Tang, D. (1983) Induction of pollen plants of litchi tree (Litchi chinensis Sonn.)
Acta Genetica Sinica 10, 369374. (in Chinese with English abstract)
Gamborg, O.L., Miller, R.A. and Ojima, K. (1968) Plant cell cultures: I. Nutrient require-
ments of suspension cultures of soybean root cells. Experimental Cell Research 50,
151158.
Gan, Y.Y., Zaini, S. and Idris, A. (1981) Genetic variation in the grafted vegetatively
propagated mango (Mangifera indica). Pertanika 4, 5362.
Garca, D., Escalante, M., Delgado, R., Ubeira, F.M. and Leiro, J. (2003a) Anthelminthic
and antiallergic activities of Mangifera indica L. stem bark components vimang and
mangiferin. Phytotherapy Research 17, 12031208.
Garca, D., Leiro, J., Delgado, R., Sanmartin, M.L. and Ubeira, F.M. (2003b) Mangifera
indica L. extract (vimang) and mangiferin modulate mouse humoral immune re-
sponses. Phytotherapy Research 17, 11821187.
Garrido, G., Gonzalez, D., Lemus, Y., Garcia, D., Lodeiro, L., Quintero, G., Delporte,
C., Nuez-Selles, A.J. and Delgado, R. (2004) In vivo and in vitro anti-inammatory
activity of Mangifera indica L. extract (VIMANG). Pharmacology Research 50,
143149.
Gavin, A.L., Conger, B.V. and Trigiano, R.N. (1989) Sexual transmission of somatic
embryogenesis in Dactylis glomerata. Plant Breeding 103, 251254.
Giovannoni, J. (2001) Molecular biology of fruit maturation and ripening. Annual
Review of Plant Physiology and Plant Molecular Biology 52, 725749.
Gohbara, M., Kosuge, Y., Suzuki, A. and Tamura, S. (1977) Colletotrichin monohydrate
methanol solvate. Acta Crystalogia 33, 12761278.
Gohbara, M., Kosuge, Y., Yamasaki, Y., Suzuki, A. and Tamura, S. (1978) Isolation, struc-
tures and biological activities of colletotrichins, phytotoxic substances from Col-
letotrichium nicotianae. Agricultural and Biological Chemistry 42, 10371043.
Gmez-Lim, M.A. (1993) Mango fruit ripening: physiology and molecular biology. Acta
Horticulturae 341, 484499.
Gmez-Lim, M.A. and Litz, R.E. (2007) Mango. In: Pua, E.C. and Davey, M.R. (eds)
Biotechnology in Agriculture and Forestry. Vol. 60. Springer, Heidelberg,
pp. 5171.
Gutirrez-Martinez, P., Lpez-Gmez, R. and Gmez-Lim, M.A. (2001) Identication of
an etr1-homologue from mango expressing during fruit ripening and wounding.
Journal of Plant Physiology 158, 101108.
Biotechnology 665
Hammerschlag, F.A. (1992) Somaclonal variation. In: Hammerschlag, F.A. and Litz, R.E.
(eds) Biotechnology of Perennial Fruit Crops. CAB International, Wallingford, UK,
pp. 3555.
Jana, M.M., Nadgauda, R.S., Rajmohan, K. and Mascarenhas, A.F. (1994) Rapid somatic
embryogenesis from the nucelli of monoembryonic mango varieties. In Vitro Cel-
lular and Developmental Biology 30, 5557.
Jayasankar, S. and Litz, R.E. (1998) Characterization of embryogenic mango cultures
selected for resistance to Colletotrichum gloeosporioides culture ltrate and phyto-
toxin. Theoretical and Applied Genetics 96, 823831.
Jayasankar, S., Litz, R.E., Schnell, R.J. and Cruz-Hernandez, A. (1998) Embryogenic
mango cultures selected for resistance to Colletotrichum gloeosporioides culture
ltrate show variation in random amplied polymorphic DNA (RAPD) markers. In
Vitro Cellular and Developmental Biology 34, 112116.
Jayasankar, S., Litz, R.E., Gray, D.J. and Moon, P.A. (1999) Responses of mango cultures
and seedling bioassays to a partially puried phytotoxin produced by a mango leaf
isolate of Colletotrichum gloeosporioides Penz. In Vitro Cellular and Developmen-
tal Biology 35, 475479.
Jefferson, R.A. (1987) Assaying chimeric genes in plants: the GUS gene fusion system.
Plant Molecular Biology Reporter 5, 387405.
Johnson, P.R. and Ecker, J.R. (1998) The ethylene gas signal transduction pathway: a
molecular perspective. Annual Review of Genetics 32, 227254.
Krishna, H. and Singh, S.K. (2007) Biotechnological advances in mango (Mangifera in-
dica L.) and their future implication in crop improvementreview. Biotechnology
Advances 25, 223243.
Kumar, S., Patil, B.C. and Sinha, K. (1990) Cyanide resistant respiration is involved in
temperature rise in ripening mangoes. Biochemistry and Biophysics Research Com-
mmunications 168, 818822.
Lad, B.L., Jayasankar, S., Pliego-Alfaro, F.A., Moon, P.A. and Litz, R.E. (1997) Temporal
effect of 2,4-D on induction of embryogenic nucellar cultures, culture maintenance
and maturation of Carabao mango somatic embryos. In Vitro Cellular and Devel-
opmental Biology 33, 253257.
Laxmi, D.V., Sharma, H.C., Kirti, P.B. and Mohan, M.L. (1999) Somatic embryogenesis
in mango (Mangifera indica L.). Current Science 77, 13551358.
Leiro, J., Garcia, D., Arranz, J.A., Delgado, R., Sanmartin, M.L. and Orallo, F. (2004) An
Anacardiaceae preparation reduces the expression of inammation-related genes
in murine macrophages. International Immunopharmacology 4, 9911003.
Litz, R.E. (1984) In vitro somatic embryogenesis from nucellar callus of monoembryonic
mango. HortScience 19, 715717.
Litz, R.E. (1986) The mango. In: Evans, D.A., Sharp, W.R. and Ammirato, P.V. (eds) Hand-
book of Plant Cell Culture. Vol. IV Techniques and Applications. MacMillan Pub-
lishing Co., New York, pp. 612625.
Litz, R.E. (1987) Application of tissue culture to tropical fruit trees. In: Green, C.E., Som-
ers, D.A., Hackett, W.P. and Biesboer, D.D. (eds) Plant Tissue and Cell Culture. Alan
R. Liss, New York, pp. 407418.
Litz, R.E. (2001) Recovery of mango plants with anthracnose resistance following
mutation induction and selection in vitro with the phytotoxin(s) produced by
Colletotrichum gloeosporioides Penz. In: Improvement of Tropical and Subtrop-
ical Fruit Trees through Induced Mutations and Biotechnology. Report of the
First Co-ordinated Research Project. Joint Food and Agriculture Organization
(FAO)/International Atomic Energy Agency (IAEA) Division, Vienna, Austria,
pp. 7378.
R.E. Litz et al. 666
Litz, R.E. (2003) In vitro regeneration and transformation of mango. In: Jaiwal, P.K. and
Singh, R.P. (eds) Plant Genetic Engineering Vol. 6: Improvement of Fruit Crops. Sci
Tech Publishing, Houston, pp. 2340.
Litz, R.E. (2004) Biotechnology and Mango Improvment. Acta Horticulturae 645,
8592.
Litz, R.E. (2008) Mango. In: Kole, C. and Hall, T.C. (eds) Compendium of Transgenic
Crop Plants: Transgenic Tropical and Subtropical Fruits and Nuts. Blackwell Pub-
lishing, Oxford, UK, pp. 163174.
Litz, R.E. and Gomez-Lim, M.A. (2002) Genetic transformation of mango. In: Khacha-
tourians, G., McHughern, A., Scorza, R., Nip, W.K. and Hui, Y.H. (eds) Transgenic
Plants and Crops. Marcel Dekker, New York, pp. 421436.
Litz, R.E. and Gomez-Lim, M.A. (2005) Mangifera indica Mango. In: Litz, R.E. (ed)
Biotechnology of Fruit and Nut Crops. CAB International, Wallingford, UK,
pp. 4061.
Litz, R.E. and Lavi, U. (1997) Biotechnology. In: Litz, R.E. (ed.) The Mango: Botany, Pro-
duction and Uses. CAB International, Wallingford, UK, pp. 401423.
Litz, R.E. and Schaffer, B. (1987) Polyamines in adventitious and somatic embryogenesis
in mango (Mangifera indica L.). Journal of Plant Physiology 128, 51258.
Litz, R.E. and Vijayakumar, N. (1988) In vitro somatic embryogenesis from the nucellus
of Mangifera indica L. Acta Horticulturae 231, 473475.
Litz, R.E., Knight, R.L. and Gazit, S. (1983) Somatic embryos from cultured ovules of
polyembryonic Mangifera indica L. Plant Cell Reports 1, 264266.
Litz, R.E., Knight, R.J., Jr and Gazit, S. (1984) In vitro somatic embryogenesis from
Mangifera indica L. callus. Scientia Horticulturae 22, 233240.
Litz, R.E., Mathews, V.H., Hendrix, R.C. and Yurgalevitch, C. (1991) Mango somatic cell
genetics. Acta Horticulturae 291, 133140.
Litz, R.E., Mathews, V.H., Moon, P.A., Pleigo-Alfaro, F., Yurgalevitch, C. and DeWald,
S.G. (1993) Somatic embryogenesis of mango (Mangifera indica L.). In: Reden-
baugh, K. (ed.) SynSeeds: Application of Synthetic Seeds to Crop Improvement.
CRC Press, Boca Raton, Florida, pp. 409425.
Litz, R.E., Moon, P.A., Monsalud, M.J., Jayasankar, S. and Mathews, H. (1995) Somatic
embryogenesis in Mangifera indica L. (mango). In: Jain, S.M., Gupta, P.K. and New-
ton, R.J. (eds) Somatic Embryogenesis in Woody Plants Vol. 2 Angiosperms. Kluwer
Academic Publishers, Dordrecht, the Netherlands, pp. 341356.
Litz, R.E., Yurgalevitch, C. and Hendrix, R.C. (1997) Effects of 1-aminocyclopropane-1-
carboxylic acid, aminoethoxyvinylglycine, methylglyoxal-bis-(guanylhydrazone)
and dicyclohexylammonium sulfate on induction of embryogenic competence of
mango nucellar explants. Plant Cell Tissue and Organ Culture 51, 171176.
Litz, R.E., Hendrix, R.C., Moon, P.A. and Chavez, V.M. (1998) Induction of embryogenic
mango cultures as affected by genotype, explanting, 2,4-D and embryogenic nurse
culture. Plant Cell Tissue and Organ Culture 53, 1318.
Lpez-Gmez, R. and Gmez-Lim, M.A. (1992) Changes in mRNA and protein synthe-
sis during ripening in mango fruit. Journal of Plant Physiology 141, 8287.
Lpez-Valenzuela, J.A., Martnez, O. and Paredes-Lpez, O. (1997) Geographic differen-
tiation and embryo type identication in Mangifera indica cultivars using random
amplied polymorphic DNA markers. HortScience 32, 105108.
Lu, C., Zainal, Z., Tucker, G.A. and Lycett, G.W. (2001) Developmental abnormalities
and reduced fruit softening in tomato plants expressing an antisense Rab11 GTPase
gene. The Plant Cell 13, 18191833.
Lycett, G.W., Zainal, Z.B., Los, M., Findlay C.L. and Tucker, G.A. (1997) Novel ripening-
specic cDNA clones from mango fruit. Acta Horticulturae 455, 277286.
Biotechnology 667
Malundo, T.M.M., Shewfelt, R.L., Ware, G.O. and Baldwin, E.A. (2001) Sugars and acids
inuence avor properties of mango (Mangifera indica L.). Journal of the American
Society for Horticultural Science 126, 115121.
Maluszynski, M. (2001) Ofcially released mutant varieties the FAO/IAEA database.
Plant Cell Tissue and Organ Culture 65, 175177.
Manzanilla Ramirez, M.A., Robles Gonzalez, M.M. and Medina Urrutia, V.M. (2000)
Avances sobre embriogenesis somatica en mango Ataulfo y Haden. In: Proceedings
Simposio Mango Control de Floracion y Mejoramiento Genetico. Instituto National
de Investigaciones Forestales y Agropecuarias, Apatzingan, Mexico, pp. 4756.
Mathews, H., Litz, R.E., Wilde, D.H., Merkel, S. and Wetzstein, H.Y. (1992) Stable inte-
gration and expression of -glucuronidase and NPT II genes in mango somatic
embryos. In Vitro Cellular and Developmental Biology 28, 172178.
Mathews, H., Litz, R.E., Wilde, H.D. and Wetzstein, H.Y. (1993) Genetic transformation
of mango. Acta Horticulturae 341, 9397.
Mathews, V.H. and Litz, R.E. (1990) Kanamycin sensitivity of mango somatic embryos.
HortScience 25, 965966.
Monsalud, M.J., Mathews, H., Litz, R.E. and Gray, D.J. (1995) Control of hyperhydricity
of mango somatic embryos. Plant Cell Tissue and Organ Culture 42, 195206.
Murashige, T. and Skoog, F. (1962) A revised medium for growth and bioassays with
tobacco tissue cultures. Physiologia Plantarum 15, 473497.
Nair, S., Gupta, P.K. and Mascarenhas, A.F. (1983) Haploid plants from in vitro anther
culture of Annona squamosa L. Plant Cell Reports 2, 198200.
Novak, F.J., Afza, R., van Duren, M. and Omar, M.S. (1990) Mutation induction by
gamma irradiation of in vitro cultured shoot-tips of banana and plantain (Musa
cvs.). Tropical Agriculture 67, 2128.
Olle, D., Baumes, R.L., Bayonove, C.L., Lozano, Y.F., Sznaper, C. and Brillouet, J.M.
(1998) Comparison of free and glycosidically linked volatile components from
polyembryonic and monoembryonic mango (Mangifera indica L.) cultivars. Journal
of Agricultural and Food Chemistry 46, 10941100.
Parisot, E. (1988) Etude de la croissance rhythmique chez de jeunes manguiers (Mangifera
indica L.). Premiere partie: description, germination et conservation de graines
polyembryonnees de manguier. Fruits 43, 97105.
Patena, L.F., Carlos-Refuerzo, L.R. and Barba, R.C. (2002) Somatic embryogenesis and
plantlet regeneration in mango (Mangifera indica L.). Vitro Cellular and Develop-
mental Biology 38, 173177.
Pliego-Alfaro, F., Monsalud, M.J., Litz, R.E., Gray, D.J. and Moon, P.A. (1996a) Effect of
abscisic acid, osmolarity and partial desiccation on the development of recalcitrant
mango (Mangifera indica L.) somatic embryos. Plant Cell Tissue and Organ Culture
44, 6370.
Pliego-Alfaro, F., Litz, R.E., Moon, P.A. and Gray, D.J. (1996b) Effect of abscisic acid, os-
molarity and temperature on in vitro development of recalcitrant mango (Mangifera
indica L.) nucellar embryos. Plant Cell Tissue and Organ Culture 44, 5361.
Prasanna, V., Prabha, T.N. and Tharanathan, R.N. (2005) Multiple forms of beta-galactosidase
from mango (Mangifera indica L. Alphonso) fruit pulp. Journal of the Science of
Food and Agriculture 85, 797803.
Prasanna, V., Prabha, T.N. and Tharanathan, R.N. (2006) Multiple forms of polygalactu-
ronase from mango (Mangifera indica L. cv Alphonso) fruit. Food Chemistry 95,
3036.
Raghuvanshi, S.S. and Srivastava, A. (1995) Plant regeneration of Mangifera indica using
liquid shaker to reduce phenolic oxidation. Plant Cell Tissue and Organ Culture 41,
8385.
R.E. Litz et al. 668
Rao, A.N., Sin, Y.M., Kathagoda, N. and Hutchinson, J.F. (1982) Cotyledon tissue culture
of some tropical fruits. In: Rao, A.N. (ed.) Tissue Culture of Economically Important
Plants. COSTED, Singapore, pp. 124137.
Ravishankar, K.V., Anand, L. and Dinesh, M.R. (2000) Assessment of genetic relatedness
among mango cultivars of India using RAPD markers. The Journal of Horticultural
Science and Biotechnology 75, 198201.
Reddy, A.S. (2007) Alternative splicing of pre-messenger RNAs in plants in the genomic
era. Annual Review of Plant Biology 58, 267294.
Reisch, B. and Bingham, E.T. (1980) The genetic control of bud formation from callus
cultures of diploid alfalfa. Plant Science Letters 20, 7177.
Rivera Domnguez, M., Manzanilla Ramirez, M.A., Robles Gonzlez, M. and Gomez-
Lim, M.A. (2004) Induction of somatic embryogenesis and plant regeneration of
Ataulfo mango (Mangifera indica L.). Plant Cell Tissue and Organ Culture 79,
101104.
Saby John, K., Mohan Rao, L.J., Bhat, S.G. and Prasada Rao, U.J.S. (1999) Characteriza-
tion of aroma components of sap from different Indian mango varieties. Phytochem-
istry 52, 891894.
Saiprasad, G.V.S., Anand, L., Ravishankar, K.V., Mythili, J.V., Nagesh, M. and Joshi, R.
(2004) Isolation and characterization of mRNAs differentially expressed during rip-
ening of mango fruits. Indian Journal of Biotechnology 3, 533537.
Sairam, K., Hemalatha, S., Kumar, A., Srinivasan, T., Ganesh, J., Shankar, M. and Venka-
taraman, S. (2003) Evaluation of anti-diarrhoeal activity in seed extracts of Mangifera
indica. Journal of Ethnopharmacology 84, 1115.
Sakai, A., Kobayashi, S. and Oyama, I. (1991) Cryopreservation of nucellar cells of naval
range (Citrus sinensis Osb. var brasiliensis Tanaka) by vitrication. Plant Cell Reports
9, 3033.
Sakho, M., Chassagne, D., Jaus, A., Chiarazzo, E. and Crouzet, J. (1998) Enzymatic mac-
eration: effects on volatile components of mango pulp. Journal of Food Science 63,
975978.
Sane, V.A., Chourasia, A. and Nath, P. (2005) Softening in mango (Mangifera indica cv.
Dashehari) is correlated with the expression of an early ethylene responsive,
ripening related expansin gene, MiExpA1. Postharvest Biology and Technology 38,
223230.
Schnell, R.A., Ronning, C.M. and Knight, RJ., Jr (1995) Identication of cultivars and
validation of genetic relationships in Mangifera indica L. using RAPD markers. The-
oretical and Applied Genetics 90, 269274.
Singh, S.K., Sharma, H.C., Goswami, A.M. and Singh, S.P. (2001) Abscisic acid aided
normal somatic embryogenesis from nucellar tissue in mango cv. Amprapali. In:
Rai, M., Kumar, S., Singh, R. and Prasad, V.S.R.K. (eds) Recent Trends in Horticul-
tural Research, pp. 161164.
Singh, S.K., Sharma, H.C. and Singh, S.P. (2002) Abscisic acid aided normal somatic
embryogenesis from nucellar tissue in mango cv. Amprapali. In: Kapoor, A.C. (ed.)
Sustainability of Hill Agriculture: Emerging Trends and Possible Solutions. Indian
Agricultural Research Institute, Delhi, pp. 295299.
Small, I. (2007) RNAi for revealing and engineering plant gene functions. Current Opin-
ion in Biotechnology 18, 148153.
Staehelin, L.A. and Moore, I. (1995) The plant Golgi apparatus: structure, functional
organization, and trafcking mechanisms. Annual Review of Plant Physiology and
Plant Molecular Biology 46, 261288.
Sturrock, D. (1968) Genetics of mango polyembryony. Proceedings of the Florida State
Horticultural Society 81, 311314.
Biotechnology 669
Sulekha, G.R. and Rajmohan, K. (2003) Relative response of varieties and explants in the
induction of somatic embryogenesis in mango (Mangifera indica L.). South Indian
Horticulture 52, 512.
Suntornwat, N., Lertwikoon, L., Bungaruang, P., Chaimanell, J., Speirs, J. and Subhadra-
bandhu, S. (2000) Cloning and characterization of putative, endopolygalacturonase
cDNA from ripening mango (Mangifera indica Linn cv. Nam Dok Mai). Acta Horti-
culturae 509, 153158.
Thomas, P. (1999) Somatic embryogenesis and plantlet regeneration from nucellar tissue
of monoembryonic mango. Journal of Horticultural Science and Biotechnology 74,
135139.
Tucker, G.A. and Grierson, D. (1987) Fruit ripening. In: Stumpf, P.K. and Conn, E.E. (eds)
The Biochemistry of Plants Vol. 12. Academic Press, New York, pp. 265313.
Vasanthaiah, H.K.N., Ravishankar, K.V., Shivashankara, K.S., Anand, L., Narayanas-
wamy, P., Mukunda, G. and Prasad, T.G. (2006) Cloning and characterization of
differentially expressed genes of internal breakdown in mango fruit (Mangifera in-
dica). Journal of Plant Physiology 163, 671679.
Wilkinson, J.Q., Lanahan, M.B., Yen, H.C., Giovannoni, J.J. and Klee, H.J. (1997) An
ethylene-inducible component of signal transduction encoded by never-ripe. Science
270, 18071809.
Wu, Y.-J., Huang, X.-L., Xiao, J.-N., Li, X.-J., Zhou, M.-D. and Engelmann, F. (2003)
Cryopreservation of mango (Mangifera indica L.) embryogenic cultures. Cryo-Letters
24, 303314.
Yang, Y.Q. and Wei, W.X. (1984) Induction of anther-derived plantlet in longan. Acta
Genetica Sinica 11, 288293. (in Chinese with English abstract)
Yashoda, H.M., Prabha, T.N. and Tharanathan, R.N. (2005) Mango ripening-chemical
and structural characterization of pectic and hemicellulosic polysaccharides. Car-
bohydrate Research 340, 13351342.
Zainal, Z., Tucker, G.A. and Lycett, G.W. (1996) A rab11-like gene is developmentally
regulated in ripening mango (Mangifera indica L.) fruit. Biochemica et Biophysica
Acta 1314, 187190.

You might also like