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Review
Mechanism and Active Variety of Allelochemicals
PENG Shao-Lin1, 3*, WEN Jun1, GUO Qin-Feng2
(1. South China Institute of Botany, The Chinese Academy of Sciences, Guangzhou 510650, China;
2. U.S.Geological Survey, 8711 37th St.SE, Jamestown, ND 58401, USA;
3. State Key Laboratory for Biocontrol, School of Life Sciences, Zhongshan University, Guangzhou 510275, China)
Abstract: This article summarizes allelochemicalsactive variety, its potential causes and function
mechanisms. Allelochemicalsactivity varies with temperature, photoperiod, water and soils during natural
processes, with its initial concentration, compound structure and mixed degree during functional processes,
with plant accessions, tissues and maturity within-species, and with research techniques and operation
processes. The prospective developmental aspects of allelopathy studies in the future are discussed.
Future research should focus on: (1) to identify and purify allelochemicals more effectively, especially for
agriculture, (2) the functions of allelopathy at the molecular structure level, (3) using allelopathy to explain
plant species interactions, (4) allelopathy as a driving force of succession, and (5) the significance of
allelopathy in the evolutionary processes.
Key words: allelopathy; active variety; functional mechanism
Allelopathy is commonly defined as any direct or indi- mechanisms and active variety of allelochemicals have also
rect effects (stimulation or inhibitory) by one plant, includ- dramatically increased. The purposes of this paper are (1)
ing microorganisms, on another through production of to review the studies as to how the activity of
chemical compounds that escape into environment (Rice, allelochemicals, varies with natural process, functional
1984). It has long been recognized since Democritus, a Gre- process, excreting ways and techniques and operational
cian scholar, who realized the chemical interactions between processes in investigation; (2) to discuss and categorize
plants since B.C. 3; in China, such a phenomenon was re- the potential causes or mechanisms for these variation;
corded in Qi Min Yao Shu in many agricultural processes. and (3) to identify future research needs and directions.
Since the 1960s, following the rapid advances in chemistry,
plant physiology, biochemistry and ecology, the role of
1 Variation of the AllelochemicalsActivities
allelopathy has increasingly become investigated (Anon, 1.1 Changes in allelochemicalsactivity during natural
2000; Mallik, 2000). Following the establishments of the processes
standard phytotoxic bioassay for allelochemicals and the Studies on the allelochmicalsactivity need to take many
related nomenclature system, related studies then become biotic and abiotic factors into account. Some studies (Xu et
more precise and systemic (Macias et al., 2000a; 2000b). al., 1999; Zhou, 1999; Kong et al., 2002) show that allelopa-
More recently, growing efforts are being made to examine thy is more intensive in the adverse or harsh environment
the role of allelopathy in controlling the spreads of weeds, when water, light or nutrition is limited. Plants often use
pest insects and diseases. allelopathy as a means to increase their competitive ability
Allelochemicals are the small molecular weight com- and therefore survival rates.
pounds excreted from plants during the process of second- 1.1.1 Allelochemiclasactivity varies with temperature
ary metabolism (Rice, 1984). These chemicals usually accu- and photoperiod The intensity of light, photoperiod and
mulated in plants, soils, and other surrounding organisms. temperature all play important roles in the synthesizing of
These compounds also vary in chemical composition, con- allelochemicals. In general, longer photoperiod and higher
centration and localization in plant tissues and from plant- temperature favor the allelochmicalsactivity. For example,
to-plant with changes in both biotic and abiotic conditions Pramanil et al. (2000) found that the autotoxicity of root
(Waller and Einhellig, 1999). As more allelopathic phenom- exudates from Cucumis sativus changed with temperature
ena being discovered and described, studies on the and photoperiod. The rate of root exudation in vegetative
and reproductive stages could be doubled under elevated the same species. Variation in allelochemicalsactivity dur-
temperature and elongated photoperiod. Lobon et al. (2002) ing the excrete process indicates that plant itself is also a
also demonstrated that the allelopathic potential of the exu- key controlling factor. Some studies (Jensen et al., 2001)
dates of Cistus ladanifer was enhanced by high tempera- showed that a certain gene controls the output of
ture and long photoperiod. Chon et al. (2000) found that allelochemicals and different allelochemicals are controlled
prolonging the photoperiod can increase the autotoxicity by different genes and in different time. This proves the
of Medicago sativa L. feasibility of using genes to study the mechanisms of
1.1.2 Allelochemiclasactivity varies with water Water allelopathy.
is the necessary condition of life and changes in water 1.2.1 Allelochemicalsactivity varies with plant acces-
conditions pose significant effects on the allelochemicals sions Different plants, even different accessions of the
activity. In general, the activity will be enhanced when water same plant, will produce different allelochemicals. Wu et al.
is in shortage. For example, some literature (Li et al., 2001) (2000a) analyzed the phenolic acids in root tissues of 58
reported that the concentration of chlorogenic acid in some wheat accessions, and demonstrated that the concentra-
plants may be increased because of the shortage of water. tions of allelochemicals of different accessions were very
Some terpenes, such as -pinene, -pinene, cineole, cam- different. Wu et al. (2000b) evaluated the allelopathy in the
phor and can, increase their volatiles under dry conditions. seedlings of 453 wheat accessions against Lolium rigidum
Dias and Dias (2000) found that prolonging drought can by using the equal-compartment-agar method (ECMA). He
increase the allelopathic activity of Datura stramonium. demonstrated that there was a considerable genetic varia-
1.1.3 Effects of soils on allelochemiclasactivity There tion in allelopathic activity in wheat germplasms. Tang and
is a massive interface between plant roots and soils, where Sun (2002) studied allelopathy of 700 rice accessions against
material exchanges and reciprocal effects between each vegetable and their result demonstrated that allelopathic
other occur. Therefore, soil is a critical factor that affects intensity varied with the accessions and the allelopathy
the allelochemicalsactivity. Rojo et al. (2000) found that among local varieties was stronger than that among culti-
phytoplankton structure and dynamics were related to al- vated varieties. Kamara et al. (2000) investigated the ef-
lelopathy in a semiarid wetland following the increase of fects extracts from levels and much of 14 trees on maize
eutrophication. The study of Hall et al. (1983) showed that germination, growth and yield in the laboratory and field
the input of N, P can reduce the allelopathic inhibitory of experiment, and all data showed great variations in
Solidago pacifica Juz, Festuca ovina L. and Helianthus allelochemicalsactivity.
annuus L. Roth et al. (2000) studied the allelopathy of sor- 1.2.2 Allelochemicalsactivity varies with plant tissues
ghum on wheat under several tillage systems, and found Most tissues of plant, such as leaf, flower, fluid, stem, root
that allelopathy might be reduced by prompt tillage and and seed, even litter, can release a certain amount of
other practices that promote rapid decomposition of sor- allelochemicals into the surrounding environments. These
ghum stover. Kidd and Proctor (2000) examined the growth allelochemicals can be very different as different parts or
response of ecotypes of Holcus lanatus L. in different soil tissues of plants have different physiological functions.
types in Northwest Europe to phenolic acids, and found The extracts from the roots and stems were reported (Mo
that the allelopathic activity varies with soil pH. Souto et and Fan, 2001) that have autotoxicity and inhibit the root-
al. (2000) studied the relationships between phenolics and ing and germination processes of Braguiera gymnorrhiza,
soil microorganisms in spruce forests and found that phe- yet other parts of the plants can stimulate its germination.
nolic compounds can stimulate fungi and cellulose Wu et al. (2001) examined the changes in allelopathic con-
hydrolyzers in the winter, but inhibit cellulose hydrolyzers tent 2,4-dihydroxy-7-methoxy 1,4-benzoxazin 3-one
in summer. Mattner and Parbery (2001) demonstrated that (DIMBOA) in different parts of wheat, and found that
rust may enhance the allelopathy of Lolium perenne L. DIMBOA level in the root tissues is the highest followed
against Trifolium repens L. Blum et al. (2000) found that by the stems. Ben-Hammouda et al. (2002) studied barley
bulk-soil and rhizosphere bacteria can have considerable autotoxicity from the roots, stems and leaves extraction of
influence on the types and concentrations of phytotoxins, barley, and the result showed that the leaves were the most
including phenolic acids on the root surface. important source of allelopathic substances, and the roots
1.2 Within-species variations in allelochmicalsactivity were the last. Ben-Hammouda et al. (2001) also investi-
Allelochemicalsactivity is highly species-specific and gated the phytotoxicity of Hordeum vulgare on Triticum
changes among tissues and also with maturity of plants of durum and T. aestivum, and showed that the allelopathic
PENG Shao-Lin et al.: Mechanism and Active Variety of Allelochemicals
potential increased with physiological maturity, and leaves characterized the different phytotoxic potentials of two
and roots were the most phytotoxic plant pats in H. vulgare closely related dwarf-shrub species, Empetrum nigrum and
plant parts. Huang et al. (2000) studied the changes of total E. hermaphroditum, and found that these negative effects
phenolic content in decomposing Chinese fir, and found were related to the different substitution of a bibenzyl in
the order of total phenolic content in different parts of the two species. Macias et al. (2000c) tested 11 natural and
stump-roots was as follows (from the highest to the lowest): synthetic podolactones and found that their activities re-
root > stump heartwood > stump-roots. quired specific structure. They then proposed a model for
1.2.3 Allelochemicalsactivity varies with plant maturity potential natural herbicide. Goo et al. (2001) extracted the
Plant tissuesmaturity also affects the allelochemicalscon- underground portion of Allium fistulosum and detected
tent and intensity. Some studies (Wang et al., 2001; Hu the allelopathic polysaccharide. He showed that sugar
and Kong, 2002) found that the quantity and content of moiety and/or molecular size were important for the
allelochemicals in soybean stubs were different in different allelochemicalsactivity based on the evidence of their ef-
decomposing time and growth stages. Sharma et al. (2000) fects on the growth of rice seedings.
found that the allelopathic intensity increased by the age 1.3.3 Allelochemicalsactivity varies with mixed degree
of Populus deltoids. Huang et al. (2000) found the total Purity and mixture are very different in their chemical and
phenolic contents in stump-roots of Chinese fir decreased physical properties. Therefore, allelochemicalsactivity may
with increasing age of stump-roots. vary with the mixed degree. Chaves et al. (2001) detected
1.3 Changes in allelochemicalsactivity during func- 11 allelochemicals in the exudates of Cistus ladanifer, and
tional processes studied the effect of each of them and their combinations
Allelochemicals is a kind of small molecular secondary on germination, cotyledon emergence, root length, and
metabolite. The functional processes in allelopathy are cotyledon length of Rumex crispus. He discovered that the
chemical reactions among plants. The rate and intensity of later produced a greater negative effect than when acting
chemical reactions are controlled by many factors, among alone. Saario et al. (2002) extracted allelochemicals, phe-
which the compound structure and initial concentration nolic acid and benzoic acid from air-dried shoots of
are especially important. Therefore, predicting the Vaccinium vitisidaea, and found that both pure acids have
allelochemicalsactivity must take compound structure and strong inhibitory effects on seed germination of Lepidium
detected concentration into account. sativum while the mixed acid has no allelopathy. Dias and
1.3.1 Allelochemicalsactivity varies with its initial con- Moreira (2002) studied the effects of phytotoxic activity of
centration Under certain conditions, the rate of an el- both single and combined application of water soluble and
ementary reaction is positively related to the reactants con- volatile compounds of Cistus ladsnifer on the germination
centration with reactive coefficient as product. Wang et al. and early root growth of subterranean clover. Antagonism
(2001) found that malonic acid and 1, 2-benzennedicarboxylic was found between water solution and volatiles resulting
acid inhibited the day matter accumulation in seeds, plant in a reduction of inhibition or a shift from inhibition to
height, and germination of soybean, and the allelopathic stimulation, but the early root growth was always inhibited
intensity increased as concentration increased. Hu and and only by water solubles. These results showed that the
Kong (2002) found that the allelochemicals, excreted by simultaneous presence of water solubles and volatiles might
root of Arachis hypogaea before four-leaves, stimulated lead to changes of the chemical nature of metabolites. Kong
the seedling growth of rice in low concentration while in- et al. (1998) examined the interactions among allelochemicals
hibited in high concentration. Sinkkonen (2001) developed of Ageretum conyzoides and found synergistic actions
a biological response model for the density-dependent among allelochemicals.
chemical interference based on the plant responses to many 1.4 Possible effects of research techniques and opera-
phytochemicals changes from stimulatory to inhibitory tion processes on allelochemicalsactivity
as the concentration of the phytochemical increases. Many factors can affect the result of extraction and pos-
Mucciarelli et al. (2000) investigated the effects of 3, sible human or operational errors or bias involve extract
4-dihydroxybenzoic acid on Nicotiana tabacum L. selection and extraction conditions during the study. These
Their result showed strong inhibitory at the highest con- factors have not been paid enough attention so far yet
centration but stimulation effects at the lowest they can make comparison among studies very difficultly.
concentration. For this reason, standardization in the extraction and op-
1.3.2 Effects of compound structure Nilsson et al. (2000) erational processes is thus critically needed.
1.4.1 Allelochemicalsactivity varies with extraction CO2 concentration, will increase the secondary metabolites.
The selection of extracts is very important because it di- Optimum Defense, OD hypothesis (Chapin, 1987): Plants
rectly affects the feasibility of the operational process and will produce secondary metabolites only when they attain
the quality and quantity of the products, therefore extrac- more defense profits than growth profits.
tion selection must be taken into account in the study of Resource Availability, RA hypothesis (Coley, 1985):
allelochemicalsactivity.Yanet al. (2000) studied the soy- Resource availability in the environment has been proposed
bean soil extracts using water, phosphoric buffer solution- as the major determinant for both the amount and type of
ether and ethanol on soybean and wheat germination, and plant defense. When resources are limited, plants with in-
showed the allelopathic potentials were very different when herently slow growth are favored over those with fast
extracts were used. Kato-Noguchi (2000; 2001) also dis- growth rates; slow growth rates in turn favor more invest-
covered different allelopathic potentials when n-hexane- ments in defenses.
soluble, acetone-soluble and water-soluble were used for Comparing the four hypotheses above mentioned (Kong
the extracts of Evolvulus alsinoides and Melissa officinalis, et al., 2000) showed that the most remarkable difference
with water extraction having the strongest allelopathic among them is whether plants produce the secondary me-
activity. Burgos and Talbert (2000) studied differential ac- tabolites actively or passively. The first two hypotheses
tivities of 3H-benzoxazolinone (BOA), 2,4-dihydroxy-1,4 (CNB and GOB) consider the metabolites as passive prod-
(2H) benzoxazine-3-one (DIBOA) and crude water extract ucts due to changes in the surrounding environments that
of Secale cereale Elbonin culture dish bioassays using led to substance accumulation in plants. The last two theo-
several vegetable and weed species, and showed that ries (OD and RA), however, consider the metabolites as
DIBOA extract had the strongest allelopathic intensity. voluntary products due to changes in the production cost.
1.4.2 Allelochemicalsactivity varies with extraction Although these four hypotheses cannot explain all the va-
condition The extraction condition can affect the allelo- rieties of allelochemicals, they have valuable implications
pathic potential as well. For example, temperature affects in practical applications.
the activity and saturation of alleochemicals and time af-
2 Mechanisms of AllelochemicalsFunctions
fects the extraction degree of allelochemicals. The compari-
son among allelochemicals obtained from different extract- Presently, studies on the allelopathy mechanisms mostly
ing conditions showed that extracting time and tempera- concentrate on the physiology. Allelochemicals first dam-
ture have important influence on the allelochemicals analy- age the cytolemma, and then send the stress information
sis result (Wang and Zhao, 2001). Prates et al. (2000) evalu- into the cell through the target point on the cytolemma to
ated the effect of cold and hot aqueous extracts of Leucaena affect the adsorption of incretions and ions. The growth of
leucocephala on germination and growth of Zea mays, and plant can be either stimulated or inhibited due to cell divi-
showed that the extracts obtained using cold water had no sion and photosynthesis caused by the adsorption of
phytotoxic effect on the germination and plant growth while incretions, ions and water.
the extraction using hot water caused reduction in root Studies in physiology can be very insightful for allel-
length. opathy research. However, most of these studies have fo-
1.5 The induced mechanism of the allelochemicals cused on a single plant function and cannot explain the
variety mechanisms of allelopathy fully. Other mechanism such as
There are mainly four parts about the induced mecha- genetics, need to incorporated onto more comprehensive
nism of the allelochemicalsvariety in physiology: studies to investigate the similarities and differences in al-
Carbon/Nutrient Balance, CNB hypothesisBryant et lelopathy therefore to form an integrating allelopathy mecha-
al., 1983: Secondary metabolites, such as phenols, terpene nism theory.
and other compounds whose structure is C, H, O) in plants 2.1 The effect on cytolemma
associate with C/N; on the contrary, secondary metabolites, Allelochemicals can affect the structure, function and
such as alkaloid, are negative correlation with C/N. permeability of cytolemma. For example, Pflumacher (2002)
Growth/Differentiation Balance, GDB hypothesis has shown that algae themselves can produce a
(Wareing and Phillips, 1981): Plants give priority to growth cyanobacterial secondary metabolite, microcystin-LR,
in plenteous resource while to differentiation in shortage. which has a strong allelopathy on aquatic macrophytes
Any factors, which affect plant growth more than photo- such as Ceratophyllum demersum and Myriophyllum
synthesis such as undernutrition, low temperature, and high spicatum by changing the pigment patterns. Galindo et al.
PENG Shao-Lin et al.: Mechanism and Active Variety of Allelochemicals
bacteria Escherichia produced colicin to allot their spatial response to multiple environmental factors. Bioscience, 37:
patterns in evolutionary processes. Dobson and Bergstrom 4977.
(2000) studied the evolution of pollen odors and found that Chaves N, Sosa T, Alias J C, Escudero J C. 2001. Identification
allelopathy serves multiple functions in both pollination and effects of interaction phytotoxic compounds from exu-
and defense: by protecting the male gametophyte and in- date of cistus ladanifer leaves. J Chem Ecol, 27: 611621.
creasing its dispersal by animals. These references demon- Chittapur B M, Hunshal C S, Shenoy H. 2001. Allelopathy in
strated that allelopathy and evolution exhibit two-way parasitic weed management: role of catch and trap crops. Al-
functions. However, much remains to be done regarding lelopathy J, 8: 147159.
the production and function of allelopathy during evolu- Chon S U, Coutts J H, Nelson C J. 2000. Effects of light, growth
tion in the context of many other related factors and how media, and seedling orientation on bioassays of alfalfa
allelopathy may affect future evolutional processes. autotoxicity. Agron J, 92: 715720.
Coley P D, Bryant J P, Chapin F S . 1985. Resource availabil-
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