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