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2004 All Elo Path Yj
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CONTENTS
1. INTRODUCTION
2. MORPHOLOGY
3. ECONOMIC IMPORTANCE
4. ALLELOPATHIC RESEARCH
5. ALLELOCHEMICALLS
6. FUTURE LINES OF RESEARCH
7. ACKNOWLEDGEMENTS
8. 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.
1. INTRODUCTION
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).
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, β -caryophyllene, α -bisabolene and
E-β-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 μ
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 93μg/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 μg.
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, 2–methyl-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
4 Kong et al
MeO
O MeO
MeO 1 MeO O
MeO MeO
O 2 MeO Ha Ha Hb
Hb +
MeO MeO
O MeO
3 O
Dimer A Dimer B
CH3
n =0 3
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
Lab Bioassays
Volatile oil Cucumis sativus Fresh weight, root length and shoot 20,24
height
Lolium multiforum Fresh weight, root length, shoot height 20,24,25,48
and contents of chlorophyll
Lysopersicon Fresh weight, root length and shoot 20,24
height
Allelopathic Ageratum conyzoides 5
Pot culture
Volatile oil Arachis hypogaea Germination, fatty enzyme, soluble 27,37
protein, branching, number of pegs and
number of pods, delayed flowering
Amaranthus Fresh weight, content of chlorophyll, 21,27
retroflexus POD activity
Brassica campestris
Cucumis sativus
Echinochloa crus-galli
Lolium multiforum
Aqueous Cucumis sativus Growth 1
extracts Oryza sativa Growth 9,15-17
Zea mays Growth and uptake of P and Zn 42
6 Kong et al
Field studies
Residues Arachis hypogaea Germination, root and shoot length, 5
number of nodules, chlorophyll content,
number of branches, number of peg and
number of pod
Amaranthus Number of plants in the field 5,14
retroflexus Inhibited growth
Amarenthus spinosus Decreased plants population in the field 5
Amarenthus viridis,
Biden pilosa,
Carex heterostachya,
Cyperus rotundus,
Cyperus duclouss,
Cuscuta australis,
Cuscuta chinensis,
Digitaria sanguinals,
Polygomun
iapathifolium,
Polygomun
caeepitosum
Oryza sativa Growth 1,15,17
0.5
0
A B C D
-0.5
-1
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).
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 non-
volatile 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 γ-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.
7. ACKNOWLEDGEMENTS
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