Tables: 2, Figs : 3
0971-4693/94 US $ 5.00
International Allelopathy Foundation 2004
CONTENTS
1.
2.
3.
4.
5.
6.
7.
8.
INTRODUCTION
MORPHOLOGY
ECONOMIC IMPORTANCE
ALLELOPATHIC RESEARCH
ALLELOCHEMICALLS
FUTURE LINES OF RESEARCH
ACKNOWLEDGEMENTS
REFERENCES
ABSTRACT
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).
Above-ground parts
Figure 1. Morphology of A. conyzoides
Root
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
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
Reference
20,24
20,24,25,48
20,24
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
height
20,24,25,48
height
9,20,24,25,48
shoot
24
length
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
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
Radish
Mung bean
Ryegrass
0.5
0
A
-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).
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
Source
Reference
Ageratochromene
Leaf,
stem,
soil
flower,
oil
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
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
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
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).
10
Kong et al
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).
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