Allelopathy Journal 14 (1): 1-12 (2004)

Tables: 2, Figs : 3

0971-4693/94 US $ 5.00
International Allelopathy Foundation 2004

Allelopathic Plants. Ageratum conyzoides L.

CHUIHUA KONG*, FEI HU, XIAOHUA XU1, WENJU LIANG2 and CHAOXIAN ZHANG3

Institute of Tropical & Subtropical Ecology, South China Agricultural University,

Guangzhou, 510642, China
E. Mail: chkong@scau.edu.cn
(Received in revised form : April 5, 2004)

CONTENTS
1.
2.
3.
4.
5.
6.
7.
8.

INTRODUCTION
MORPHOLOGY
ECONOMIC IMPORTANCE
ALLELOPATHIC RESEARCH
ALLELOCHEMICALLS
FUTURE LINES OF RESEARCH
ACKNOWLEDGEMENTS
REFERENCES
ABSTRACT

The genus Ageratum consists of about 40 spp. found in tropical and

subtropical countries, however, only two spp. A. conyzoides L. and A. houstonianum
Mill are well known. The allelopathy of only Ageratum conyzoides L. has been studied
in bioassays, pot cultures and field investigations. Its allelopathic potential varied with
growth stages and environmental conditions. It releases more volatile allelochemicals
(ageratochromene and its derivatives, monoterpenes, sesquiterpenes and flavones)
under adverse conditions. These allelochemicals not only inhibited the germination
and growth of associated plants, but also adversely affected the microbes and insects.
Intercropping of Ageratum conyzoides in citrus orchards effectively suppressed the
weeds and controlled other insect pests, hence, practiced in large areas in South China.
Besides, the mode of release, transformation in soils of major allelcochemicals from A.
conyzoides has been studied.
Key words: Ageratochromene, Ageratum conyzoides, allelochemicals, allelopathy,
chemical interactions, citrus orchard, flavones, pathogens, terpenes, weeds.

1. INTRODUCTION
There are approximately 40 spp. in the genus Ageratum, family Compositae
(Asteraceae) and are native to Central America (28). Only two species, A. conyzoides L.
*

Correspondence author.
State Key Laboratory of Elemental-Organic Chemistry, Nankai University, Tianjin 300071, China; 2 Institute of
Applied Ecology, Chinese Academy of Sciences, Shenyang 110016, China; 3 Institute of Plant Protection, Chinese
Academy of Agricultural Sciences Beijing 100094, China.
1

Kong et al

and A. houstonianum Mill are well known (3,28). Most taxa are found in Mexico, the
Caribbean and Florida (USA). A. conyzoides and A. houstonianum have spread to West
Africa, Southeast Asia, South China, India, Australia and South America (35,47). They are
pioneer plants growing in waste and ruined sites (where sufficient water is available) and
cultivated fields. In Central America A. conyzoides has been bred for many colours of
flowers (47,49). Although it is harmful to crops and invades cultivated fields and interferes
with the natural community compositions, but is a folk medicine in several countries and
also
has
anti-microbial,
insecticidal
and
nematicidal
activities
(34).
A. conyzoides contains many secondary metabolites: flavonoids, chromenes, benzofurans
and terpenoids (34,35,37) and some of them are inhibitory to other organisms. This species
appears to be a valuable agricultural resource (34).

2. MORPHOLOGY AND HABITAT

A. conyzoides is an annual erect, branched herb growing 15 to 100 cm tall. Its
stem is covered with fine white hairs, leaves are opposite, pubescent with long petioles and
include glandular trichomes. It has a shallow tape root system. The inflorescence contains
30 to 50 pink or purple flowers arranged a corymb and are self-incompatible (13). The
fruit is an achene with an aristate pappus and is easily dispersed by wind and animals fir
(Fig.1). Seeds are positively photoblastic and remain viable upto 12 months (35). The
seeds germinate between 20-25C. It prefers a moist, well drained soil but may tolerate dry
conditions (29). The species has great morphological variations and appears highly
adaptable to varying ecological conditions (9). It often becomes dominant and forms a
stand in natural community and is resistant to common insects or diseases (30).

Above-ground parts
Figure 1. Morphology of A. conyzoides

Root

Allelopathic Ageratum conyzoides

3. ECONOMIC IMPORTANCE
The allelopathic strategies have been utilized in South China agriculture, where,
Ageratum conyzoides is intercropped as understory in citrus orchards to suppress the
growth of other weeds. Besides, intercropping of A. conyzoides makes the citrus orchard
environment more favourable for predatory mites (Amblyseius spp.), which are most
effective natural predators of the citrus red mite (Panonychus citri) (30,38,48). The
ageratochromene, demethoxy-ageratochromene, -caryophyllene, -bisabolene and
E--farnesene allelochemicals are the major constituents of the volatiles in the air of
A. conyzoides intercropped in citrus orchard and their amounts ranged from 1 to 2.4
g/m3. These volatile allelochemicals attracted greatly the predatory mite A. newsami but
repelled slightly the P. citri. The population density of predatory mite A. newsami in the
A. conyzoides intercropped citrus orchard could be maintained at a high level through the
release of volatile allelochemicals from plants of A. conyzoides. Hence, the volatile
allelochemicals could regulate the population of the mites in citrus orchard and the
chemical interactions among predator, pest and citrus may be achieved through
intercropping.
The intercropping of A. conyzoides also controls the major weeds (Bidens pilosa,
Digitaria sanguinalis and Cyperus difformis) in citrus orchards. Moreover, in
A. conyzoides intercropped citrus orchard soil, the population of major pathogenic fungi
(Phytophthora citrophthora, Pythium aphanidermatum and Fusarium solani) was also
reduced, as the concentration of three flavones, ageratochromene, and its two dimers
ranged from 11 to 93g/g soil (10,22,23). The presence of these allelochemicals in soils
controlled some weeds and diseases in citrus orchards. Besides, the reversible dynamic
transformations between the ageratochromene and its dimers in the A. conyzoides
intercropped citrus orchard soil may be an important mechanism to maintain an effective
concentration of bioactive allelochemicals for inhibition of weeds and soil pathogenic
fungi (23). Thus, the intercropping A. conyzoides in citrus orchards markedly reduced the
population of major weeds and soil pathogenic fungi (10,22).
These allelochemicals present in the A. conyzoides intercropped citrus orchard
soil could slightly inhibit the growth of citrus seedling at high concentrations (> 300 g.
g-1) in greenhouse, but their allelopathic inhibition did not occur in intercropped citrus
orchard. Further studies revealed that there was a reversibly dynamic transformation
between ageratochromene and its two dimers in the A. conyzoides intercropped citrus
orchard soil, which ageratochromene releasing from ground A. conyzoides plants might be
transformed into its dimers, and the dimers might be remonomerized in the soils.
Ageratochormene in the soil firstly polymerized into dimers after 26 days under high
organic matter and fertility of soil, and then degraded gradually into benzonic acid and its
derivatives, 2methyl-propanoic-acid and acetic acid after 34 days (Fig. 2). However, the
dynamic transformation did not occur in the soil with low organic matter and fertility.
Ageratochromene had no dimerization and degraded directly within 18 days under the low
organic matter and fertility soil. Obviously, the transforming and degrading of
ageratochormene was significantly correlated with the organic matter and fertility in the
soil. The dimerization was not correlated with microorganisms in the soil, but the

Kong et al

MeO
MeO

2 MeO

MeO

O
Ha

Hb

MeO
MeO

Dimer A
COOH

n =0

Hb

MeO
MeO

(CH3O)n

Ha

MeO
MeO

Dimer B

+ CH3CHCOOH + CH3COOH
CH3

Figure 2. Transformation and degrading of ageratochormene in the soil. Ageratochormene

transformation in highly content organic matter and fertility soil was from step 1 to step 3,
but only step 3 occurred in low content organic and fertility soil.

biodegradation of both ageratochromene and its two dimers may have occurred,
particularly in the soil with low organic matter and fertility (23,27).
Generally, allelopathy occurs at seed germination and early growth stages of plant
species, hence, perennial adult citrus trees may resist the influence of allelochemicals
released from understory A. conyzoides. Besides, the large rhizosphere and soil microflora
of the citrus may rapidly biodegrade these allelochemicals and reduce their concentration
to low and non-toxic level. Therefore, A. conyzoides is intercropped in citrus orchards in
>150,000 ha in South China and gave substantial ecological and economic benefits (30). It
is an excellent example of applied aspects of allelopathy in agro-ecosystem.

4. ALLELOPATHIC RESEARCH
Allelopathic effects of extracts, volatiles and residues from A. conyzoides have
been studied (Table 1). A conyzoides releases many kinds of allelochemicals through
leaching, volatilizing and residue decomposion into the environment.
Table 1. Inhibitory allelopathic effects of A conyzoides on plants in bioassays, pot and field
Treatments
Volatile oil

Recipient species

Cucumis sativus
Lolium multiforum
Lysopersicon

Effects on recipient spp.

Lab Bioassays
Fresh weight, root length and shoot
height
Fresh weight, root length, shoot height
and contents of chlorophyll
Fresh weight, root length and shoot
height

Reference
20,24
20,24,25,48
20,24

Allelopathic Ageratum conyzoides

Phaseolus aureus
Raphanus sativus
Triticum aestivum
Aqueous
extracts

Allium sativum
Amaranthus caudatus
Cucumis sativus
Cucumis anguria
Digitaria sanguinalis
Glycine max
Lactuca sativa
Lolium multiforum
Phaseolus aureus
Oryza sativa
Raphanus mungo
Triticum aestivum
Zea mays

Purified
Allelochem
icals

Volatile oil

Aqueous
extracts

Cucumis sativus
Echinochloa crus-gall
Lolium multiforum
Lysopersicon
Phaseolus aureus
Raphanus sativus

Arachis hypogaea

Fresh weight, root length, shoot

and contents of chlorophyll
Fresh weight, root length, shoot
and contents of chlorophyll
Fresh weight, root length and
height
Germination, fresh weight, root
and shoot height

height

20,24,25,48

height

9,20,24,25,48

shoot

24

length

Germination, fresh weight, roots length

and shoot height
Germination
Inhibited germination, root length, and
shoot height
Germination, root length and shoot
height
Germination, root length and shoot
height
Germination, fresh weight, root length
and shoot height
Germination, fresh weight, root length
and shoot height
Germination, roots length and shoot
height
Germination, fresh weight, root length
and shoot height
Germination, fresh weight, root length
and shoot height
Germination, fresh weight, root length
and shoot height

Root length and shoots height of

seedling
Pot culture
Germination, fatty enzyme, soluble
protein, branching, number of pegs and
number of pods, delayed flowering

18
8
29
18
1
18
8
8,44
1
8,20
1,8,44
20,25

20,25,32

27,37

Amaranthus
retroflexus
Brassica campestris
Cucumis sativus
Echinochloa crus-galli
Lolium multiforum

Fresh weight, content of chlorophyll,

POD activity

21,27

Cucumis sativus
Oryza sativa
Zea mays

Growth
Growth
Growth and uptake of P and Zn

1
9,15-17
42

Kong et al

Residues

Arachis hypogaea

Amaranthus
retroflexus
Amarenthus spinosus
Amarenthus viridis,
Biden pilosa,
Carex heterostachya,
Cyperus rotundus,
Cyperus duclouss,
Cuscuta australis,
Cuscuta chinensis,
Digitaria sanguinals,
Polygomun
iapathifolium,
Polygomun
caeepitosum
Oryza sativa

Field studies
Germination, root and shoot length,
number of nodules, chlorophyll content,
number of branches, number of peg and
number of pod
Number of plants in the field
Inhibited growth
Decreased plants population in the field

Growth

5,14
5

1,15,17

4.1. Bioassays and pot studies

In bioassays and pot studies (Table 1), aqueous extracts and the volatiles of
A. conyzoides are allelopathic to crops [rice, wheat, corn, cucumber, radish, tomato, peanut,
soybean and mungbean (8,20,21)] and weeds [Bidens pilosa, Digitaria sanguinalis,
Amaranthus retroflexus, Amaranthus viridis, Cuscuta australis, Cuscuta chinensis,
Cyperus roundus and Cyperus difformis (5)]. A. conyzoides significantly inhibited the
germination and growth and especially the root growth of seedlings of test species.

Radish

Response index (RI)

Mung bean

Ryegrass

0.5
0
A

-0.5
-1

Figure 3. Physiological effects of A conyzoides on test crops A: chlorophyll content, B: Relative

plasma membrane permeability, C: POD activity, D: MDA contents, E: Soluble proteins
contents

Influence of the volatile oil form A. conyzoides on physiological effects in radish,

mungbean and ryegrass were investigated under lab conditions (Fig. 3). The contents of
chlorophyll, soluble proteins and POD activity of test crops were decreased, while their

Allelopathic Ageratum conyzoides

plasma membrane permeability and MDA content were greatly increased (21, 50). Further,
effect of A. conyzoides was also determined on the cytostatic activity in the root
meristematic cells of Allium cepa and a mitodepressive effect was observed (43). This
effect increased with the duration of treatment, till complete inhibition of cell division. In
addition, A. conyzoides significantly inhibited the uptake of both P and Zn in wheat (44).
4.2. Field studies
A. conyzoides often invades the cultivated fields and reduces the crop productivity
(36, 40, 41), through interference (1,14-18,35,38-41). However, it is also beneficial to
some crops in several agro-ecosystems. In south China, A. conyzoides is traditionally used
as green manure in fields, to increase the crop yields and to control the weeds. Its
allelopathic effects depend upon the time of green manuring and its decaying period in
soils (5,27). Such allelopathic effects have been observed upto 30 days after soil
incorporation (5).
4.3. Allelopathy and growing conditions
A. conyzoides growing under favourable weather conditions (sunshine,
temperature and rainfall) is less allelopathic (9,21), but becomes more allelopathic under
adverse conditions. Its allelopathic effects also vary with growth stages, habitats and other
weather factors (8,20,21). Its allelopathic potential is stronger during flowering than other
growth stages. It becomes more allelopathic under deficient nutrient and water or light,
competition with other weeds and infection with Erysiphe cichoracearum and aphids
(Aphiids gossypii) attack. Allelopathy is an adaptation mechanism of plants to strengthen
the ability to compete with neighboring plants (19,21,26). The allelopathic potential of
A. conyzoides was correlated with environmental stress. However, not all stresses
increased its allelopathic potential. Only a few natural factors could induce allelopathy,
anthropogenic factor, such as physical damage and treatment with 2,4-D, did not affect its
allelopathic potential (21).

5. ALLELOCHEMICALS
5.1 Volatile allelochemicals
Ageratochromene and its derivatives, monoterpenes and sesquiterpenes are major
components of the volatiles from A. conyzoides (20-23). These individual constituents and
their mixtures significantly inhibited the germination and growth of various crops and
weeds (Table 2). Noteworthy, Aheratochemene is less in the volatiles of A. conyzoides, but
more in A. houstonianum, but the contents of other volatile allelochemicals are similar in
both spp. (33, 36, 45). However, the allelopathy of A. houstonianum has not received much
attention.
5.2 Allelochemicals in residues and aqueous extracts
Ten flavones including one glycoside, agetochromene, demethoxyageratochromene and stigmast-5,22-diene-3 -ol are major allelochemicals of residues and

Kong et al

Table 2. Bioactivities of allelochemicals from A. conyzoides

Allelochemical

Source

Recipient plant spp.

Effect on Recipient spp.

Reference

Ageratochromene

Leaf,
stem,
soil
flower,
oil

Fresh weight, germination

content of chlorophyll,
POD activity

5,20,21

Demethoxyageratochromene

Bisabolene

Oil

Farnesene

Oil

Stigmastol

Leaf,
stem

Methyl-flavone

Leaf,
stem,
soil
Leaf,
stem,
soil
Leaf,
stem,
soil
Leaf,
stem, oil,
soil
Leaf,
stem,
flower

Growth, fresh weight, root

length, shoot height and
contents of chlorophyll
Growth, fresh weight, root
length, shoot height
Growth, fresh weight, root
length, shoot height
Growth, fresh weight, root
length, shoot height
Growth, fresh weight, root
length, shoot height and
contents of chlorophyll
Germination, growth

20,21

Caryophyllene

Leaf,
stem, oil
flower
Oil

Radish, mungbean,
tomato, wheat,
ryegrass, maize,
peanut, Bidens pilosa,
Digitaria sanguinalis
and Cyperus difformis
Radish, mungbean,
tomato, wheat,
ryegrass, maize, peanut
Radish, mungbean,
tomato
Radish, mungbean,
tomato
Radish, mungbean,
tomato
mungbean, tomato,
wheat, ryegrass, maize,

Germination, growth

23

Germination, growth

23

Germination, growth

20,21

Growth, fresh weight, root

length, shoot height and
contents of chlorophyll,
POD activity MDA
contents, soluble proteins
contents

20,21,27

Hydroxy-flavone
Flavone
glycoside
Caryophyllene
oxide
Volatile oil

Bidens pilosa,
Digitaria sanguinalis
and Cyperus difformis
Bidens pilosa,
Digitaria sanguinalis
and Cyperus difformis
Bidens pilosa,
Digitaria sanguinalis
and Cyperus difformis
Radish, mungbean,
tomato
Radish, mungbean,
tomato, wheat,
ryegrass, maize, peanut

20,21
20,21
20,21
8
23

aqueous extracts from A. conyzoides. Particularly, ageratochromene and its derivative

demethoxy-ageratochromene may be released into the environment through volatilization,
leaching and residue decomposion (8,10,24,32). These allelochemicals inhibited the
growth of crops and weeds (Table 2).
5.3. Allelochemicals interactions
The allelocheimcals from living A. conyzoides are mainly released in air through
volatilization from shoots. Actually, the concentration of volatile released by A. conyzoides

Allelopathic Ageratum conyzoides

is so high that the unpleasant odour can be smelled in the fields. These volatile
allelochemicals may be dissolved in fog, dew or rainfall and comes back to soil in the
vicinity of plants and then directly affect their growth and development (24). Its nonvolatile allelochemicals may be released into the soils by leaching or decomposition of
residues and then inhibits the germination and root growth of neighboring plants (10). A
few volatile allelochemicals including terpenes, may also be released by leaching.
Allelopathic effects of allelochemicals mixtures were more intense than pure
individuals. For example, fenchyl acetate and -bisabolene are not inhibitory individually,
but in mixture with ageratochromene, they become more inhibitory to growth of acceptor
plants (20,25). It indicates allelopathic synergism among allelochemicals of A. conyzoides.
5.4. Biological action of allelochemicals on insects
The allelochemicals of A. conyzoides possess multiple functions, they showed
striking bioactivities on insects. The insecticidal activity of allelochemicals may be one of
the most important biological functions (11,12). Ageratochromene is the first
antiallatotropins isolated from genus Ageratum (3). Both ageratochromene and its
derivatives have antijuveenile hormonal activity. The application of volatile oil of A.
conyzoides on cowpea seed exhibited insecticidal activity against weevil. Significant
oviposition deterrence and complete inhibition of emergenece of adult insects from oiltreated beans were evident at 2.5 to 10l/9.5g beans, with no adverse physiological
effects (6). Assays conducted in India revealed high nymphal mortality of the oil to
Nymphs of Schistocerca gregaria. Many insects such as Mucsa domestica, Chilo partellus,
Sitophlus oryzae, Sitophlus aeamais, Thlaspida japonica, Leptocarsia, Leptocarsia
chinesis, Dysdercus flavidus, Lucilia caesar, Tribolimn confusum, Mythimna separata,
Culex pipiens pallens,and Plutelia xylostella were reported to be inhibited by
allelochemicals of A conyzoides (2,4,11,12,13,31,43).
An incresing number of studies have revealed that the toxicity of
ageratochromene was due to a highly reactive precocene-3,4-epoxide, a metabolite
produced in inscet species from cytochrome P-450. in addition, 3, 4 double bond in
ageratochromene played no significant role in the toxicity but that the oxidative
dealkylation process at C7 position, as a tocopherol-like antioxidants, might be reponsible
for cytotoxicity (4, 6, 7). Both under the experimental and field conditions, populations of
Plutelia xylostella were significantly decreased after ageratochormene was sprayed on
several crops. When ageratochromen was applied on crops, the oviposition deterrent to
imago of Plutelia xylostella occurred firstly, the duration of larvae was prolonged, and
then the imago reproductivity of Plutelia xylostella was reduced. It showed that bilogical
actions of ageratochormene on Plutelia xylostella was not killed directly, but effectively
lowed its populations (11, 12).

6. FUTURE LINES OF RESEARCH

I.

Evaluate the relationships between allelopathic traits of A. conyzoides and its

interference.
II. Study the physiological mechanisms of A. conyzoides allelochemicals on other
organisms.

10

Kong et al

III. Develop and apply allelopathic potentials of A. conyzoides in the agroecosystems.

IV. Modify and synthesize the analogues of allelochemicals and demonstrate their
relationships of structure and function and develop new pesticides.

7. ACKNOWLEDGEMENTS
Authors thank Professor S.S. Narwal, CCS Haryana Agricultural University,
India, for his helpful comment and corrections in the English version. This work was
financially supported by the National Natural Foundation of China (30170182; 39670141)
and the State Science & Tech Program of China (2001BA509B07).

8. REFERENCES
1. Bhatt, B.P., Tomar, J.M.S. and Misra, L.K. (2001). Allelopathic effects of weeds on germination and growth
of legumes and cereal crops of North Eastern Himalayas Allelopathy Journal 8: 225-231.
2. Bouda, A., Tapondjou L.A., Fontem, D. A. and Gumedzoe, M. Y. D. (2001). Effect of essential oils from
leaves of Ageratum conyzoides, Lantana camara and Chromolaena odorata on the mortality of
Sitophilus zaemais (Coleoptera, Curculionidae). Journal of Stored Products Research 37: 103109.
3. Bowers, W.S., Ohta, T., Cleeve, J.S. and Marsella, P. (1976). Discovery of insect anti-juvenile hormones in
plants. Science 193:542-547.
4. Casas, J., Gorchs, G., Sanches-Baeza, F., Teixidor, P. and Messeguer, A. J. (1992). Inhibition of rat live
nucrosomal lipid peroxidation elicited by simple 2,2-dimethylchormenes and chromenes
structurally related to precocenes Journal of Agriculture and Food Chemistry 40:585-590.
5. Chen, J. J., Kong, C. H., Hu, F., Tan, Z. W. and Liang, J. N. (2002). Allelopathy of Ageratum conyzoides
.Allelopathic effects of residues on peanut and related weeds in the field. Acta Ecologica Sinica
22:1196-1201.(In Chinese).
6. Gbolade, A. A., Onayade, O. A. and Ayinde, B. A. (1999) Insecticidal activity of Ageratum conyzoides L.
volatile oil against Callosobruchus maculatus F. in seed treatment and fumigation laboratory tests.
Insect Science and Its Application 19:237-241.
7. Gonzalez, A. G., Zahira, E. A., Grillo. T. A., Luis, J. G., Rivera, A. and Calle J. (1991). Methoxyflavones from
Ageratum conyzoides. Phytochemistry 30: 1269-1271.
8. Hu. F. and Kong, C. H. (1997). Allelopathy of Ageratum conyzoides. I. Allelopathy of Ageratum conyzoides
aqueous extract and isolation and identification of its allelochemicals. Chinese Journal of Applied
Ecology 8:204-308. (In Chinese).
9. Hu. F. and Kong, C. H. (2002). Allelopathy of Ageratum conyzoides. VI. Effects of meteorological conditions
on allelopathy of Ageratum conyzoides. Chinese Journal of Applied Ecology 13:76-80. (In
Chinese).
10. Hu., F., Kong, C. H., Xu, X. H. and Zhou, B. (2002). Inhibitory effect of flavones from Ageratum conyzoides
on the major pathogens in citrus orchard. Chinese Journal of Applied Ecology 13:1166-1168. (In
Chinese).
11. Hunag, S. S., Pan, L. Q. and Kong, C. H. (2000). Allelopathy of Ageratum conyzoides. IV. Control effect of
Ageratochormene on the population of diamondback moth (DBM), Plutella xylostella L. Acta
Ecologica Sinica 20(Supp.): 173-178.(In Chinese).
12. Hunag, S. S., Pan, L. Q., Zeng, L. and Kong, C. H. (2001). Effects of allelochemicals from Ageratum
conyzoides on the field population of diamondback moth (DBM), Plutella xylostella. Acta
Phytophylaciga Sinica 28:357-261. (In Chinese).
13. Jhansi, P. and Ramanujam, C.G.K. (1987). Pollen analysis of extracted and squeezed honey of Hyderabad.
India Geophytology 17:237240.
14. Jha, P. K., Sah, J. P. and Chettri, M. K. (1992). Amaranth productivity under biological stresses. Crop
Research (Hisar) 5: 195-198.

Allelopathic Ageratum conyzoides

11

15. Kalita, D., Choudhury, H. and Dey, S. C. (1999). Assessment of allelopathic potential of some common
upland rice weed species on morpho-phydiological properties of rice (Oryza sativa L.) plant.
Crop Research (Hisar) 17:41-45.
16. Kalita, D., Choudhury, H. and Dey, S. C. (1999). Allelopathic effectiveness of some common rice weed
species on germination, radicle and plumule growth of rice (Oryza sativa L.) seed. Crop Research
(Hisar) 17:183-187.
17. Kalita, D. and Dey, S. C. (1998). Allelopathic effects of shot and root extracts of some weeds on growth of
rice. Geobios 25:269-372.
18. Kato-Noguchi, H. (2001). Assessment of the allelopathic potential of Ageratum conyzoides. Biologia
Plantarum 44: 309-311.
19. Kong, C. H. (1998). Problems needed attention on allelopathy research. Chinese Journal of Applied Ecology
9:332-336. (In Chinese).
20. Kong, C. H., Hu, F., Xu, T. and Lu, Y. H. (1999). Allelopathic potential and chemical constituents of volatile
oil from Ageratum conyzoides. Journal of Chemical Ecology 25: 2347-2356.
21. Kong, C. H., Hu, F. and Xu, X. H. (2002). Allelopathic potential and chemical constituents of volatiles from
Ageratum conyzoides under stress. Journal of Chemical Ecology 28:1185-1194.
22. Kong, C.H., Hunag, S. S. and Hu F. (2001). Allelopathy of Ageratum conyzoides.V. Biological activites of
the volatile oil from ageratum on fungi, insects and plants and its chemical constituents. Acta
Ecologica Sinica 21:874-587. (In Chinese).
23. Kong , C.H., Liang, W.J., Hu, F., Xu, X.H., Wang, P., Jiang, Y., Xing, X.B. (2004). Allelochemicals and their
transformations in the Ageratum conyzoides intercropped citrus orchard soils. Plant Soil (in Press).
24. Kong, C. H., Xu. T. and Hu, F. (1998). Allelopathy of Ageratum conyzoides. II. Releasing mode and activity
of main allelochemicals. Chinese Journal of Applied Ecology 9: 257-260. (In Chinese).
25. Kong, C. H., Xu. T. and Hu, F. (1998). Study on interactions among allelochemicals of Ageratum conyzoides.
Acta Phytoecologica Sinica 22:403-408. (In Chinese).
26. Kong, C. H., Xu. T., Hu, F. and Huang, S. S. (2000). Allelopathy under environmental stress and its induced
mechanism. Acta Ecologica Sinica 20:849-854. (In Chinese).
27. Kong, C. H., Xu, X. H., Chen, J. J., Hu, F. and Tan, Z. W. (2002). Allelopathy of Ageratum conyzoides.
Transformation of main allelochemical in the soil. Acta Ecologica Sinica 22:1189-1195. (In
Chinese).
28. Kossmann, G. and Groth, D. (1993). Plantea Infestantes e Nocivas. BASF Brasileira, So Paulo.
29. Ladeira, A. M., Zaidan, L. B. P. and Figueiredo, Ribeiro, R. D. C. L. (1987). Ageratum conyzoides
L.(Compositae): Germination, flowering and occurrence of phenolic derivatives at different
stages of development. Hoehnea 14:53-62
30. Liang, W. G. and Hunag, M. D. (1994). Influence of citrus orchard ground cover plants on anthropod
communities in China: A review. Agriculture, Ecosystems and Environment 50:29-37.
31. Lu, R. (1982). A study of insect anti-juvenile hormones- chemical composition of Ageratum conyzoides L and
its action to insect. Kunchong Zhishi 19(4):22-25. (In Chinese).
32. Lu, Y. H., Kong, C. H., Dong, C., Huang S. S., and Luo S. M. (2001). Allelopathy of Ageratum conyzoides
L.VII. Syntheses of analogs of ageratochormene and their inhibitory effects on plants and
microorganisms. Acata Ecologica Sinica 21:890-895. (In Chinese).
33. Menut, C., Lamaty, G., Zollo, P. H. Amvam, Kuiate, J. R. and Bessiere, J.M. (1993). Aromatic plants of
tropical central Africa: Part X. Chemical composition of the essential oils of Ageratum
houstonianum Mill. and Ageratum conyzoides L. from Cameron. Flavor and Fragrance Journal
8:1-4.
34. Ming, L. C. (1999). Ageratum conyzoides: A tropical source of medicinal and agricultural products. In:
Perspectives on New Crops and New Uses (Ed., J. Janick). Pp 469-473. Alexandria, VA USA:
ASHS Press.
35. Okunade, A. L. (2002). Ageratum conyzoides L.(Asteraceae). Fitoterapia 73: 1-16.
36. Pari, K., Rao, P. J., Subrahmanyam, B., Rasthogi, J. N. and Devakumar, C. (1998). Benzofuran and other
constituents of the essential oil of Ageratum conyzoides. Phytochemistry 49:1385-1388.
37. Prasad, K. and V.C. Srivastava. (1991). The toxic effect of some weeds on germination and initial growth of
groundnut (Arachis hypogea). Indian Journal of Agricultural Science 61:493494.
38. Pu, T.S., Liao,K. Y. and T. Chang. (1990). Investigations on predations mite resources in Citrus orchards in
Guang Xi and their utilization. Acta Phytophyarica Sinica. 17:355358. (In Chinese).
39. Roder, W., Phengchanh, S. and Keoboulapha, B. (1995). Relationships between soil, fallow period, weeds
and rice yield in slash-and-burn systems of Laos. Plant and Soil 176: 27-36.

12

Kong et al

40. Roder, W., Keoboulapha, B., Phengchanh, S., Prot, J. C. and Matias, D. (1998). Effects of residue
management and fallow length on weeds and rice yields. Weed Research 38: 167-174.
41. Shabana, N., Husain, S. I. and Nisar, S. (1990). Allelopathic effects of some plants on the larval emergence of
Meloidogyne incognita. Journal of Indian Applied and Pure Biology 5:129130.
42. Singh, S. P., Pal, U. R. and Luka, K. (1989). Allelopathic effects of three weeds serious weeds of Nigerian
savanna on germination and seedling vigour of soybean and maize. Journal of Agronomy and
Crop Science 162:236-240.
43. Saenz, M.T, Garcia, M.D. and Gomez, M.A. (1999). Effect of hexanoic and chloroformic extracts of Achillea
ageratum on mitosis in Allium cepa. Cytobios 99: 143-148.
44. Saxena, S, Sharma, K, Kumar, S. Sand, N. K. and Rao, P. B. (2003). Effect of weed extracts on uptake of P
and Zn in wheat varieties. Allelopathy Journal 11 : 201-215.
45. Sharma, K. and Sharma, O.P. (2001). Analysis of precocenes in the essential oil of Ageratum spp. by reversephase high performance liquid chromatography. Phytochemical Analysis 12 : 263-265.
46. Singh,H. P., Daizy, R. B. and Ravinder, K. K. (2002). Allelopathic effect of two volatile monoterpenes
against bill goat weed (Ageratum conyzoides L.). Crop Protection 21: 347-350.
47. Stadler, J., Mungai, G. and Brandl R. (1998). Weed invasion in East Africa: insights from herbarium records.
African Journal of Ecology 36:15-22.
48. Van, M. P. and Lenteren, J.C. (2002). Survey of current crop management practices in a mixed-rice field
landscape, Mekong Delta, Vietnam - potential of habitat manipulation for improved control of
citrus leaf miner and citrus red mite. Agriculture Ecosystems & Environment 88: 35-48.
49. Waterhouse, D.F. (1993). Prospects for biological control of paddy weeds in Southeast Asia and some recent
success in the biological control of aquatic weed. Canberra Extension. Bulletin (Australia). 366.
50. Xu, T., Kong, C. H. and Hu, F. (1999). Allelopathy of Ageratum conyzoides. III. Allelopathic effects of
volatile oil from Ageratum conyzoides on plants under different nutrient levels. Chinese Journal
of Applied Ecology 10:748-750. (In Chinese).