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82<br />

SCOTT W. MATTNER<br />

(Putnam, 1985). Initially, the reason why plants devote resources to the production of<br />

these compounds was not understood as they were regarded as functionless waste<br />

products (Mothes, 1955). It is now increasingly accepted, however, that these<br />

compounds function as defensive agents against pathogens, insects and neighbouring<br />

plants (allelopathy). There is an enormous diversity of allelochemicals produced by<br />

plants (Bansal, 1994), with classification based on chemical structure (Whittaker and<br />

Feeney, 1971; Mandava, 1985; Putnam 1985) or on origin and chemical properties<br />

(Rice, 1984). For example, Rice (1984) recognised 14 main categories of<br />

allelochemicals. Of these groups, however, the phenolics are considered by many as<br />

the most important (Putnam and Duke, 1979; Mandava, 1985; Inderjit, 1996).<br />

Putative allelochemicals have been isolated from a variety of different plant organs,<br />

including shoots, roots, flowers, rhizomes, fruit and seed (Rice, 1984). They are<br />

stored in cell vacuoles so as not to interfere with the donor plant itself (Chou, 1989).<br />

Furthermore, secondary metabolites may be bound to sugars as glycosides or occur as<br />

polymers and crystals rendering them ineffective against the donor (Whittaker and<br />

Feeney, 1971). The release of allelochemicals into the environment may occur through<br />

volatilisation, root exudation, leaching or plant residue decomposition (Rice, 1984).<br />

<strong>Allelochemicals</strong> induce a wide range of symptoms in receiver plants, ranging<br />

from sudden wilting and death (e.g. tomato (Lycopersicon esculentum) grown in the<br />

vicinity of black walnut (Juglans nigra), Hale and Orcutt, 1987), to the more common<br />

subtle changes in growth. Determination of this reaction depends on the mode of<br />

action, the concentration, and the susceptibility of the receiver plant to the<br />

allelochemical. Allelopathic effects may be direct, such as affecting plant metabolism<br />

and growth, or indirect, such as altering of soil properties and nutrient status (Inderjit<br />

and Weiner, 2001). Rizvi et al. (1992) explained 12 plant functions that allelochemicals<br />

may affect, including membrane permeability, stomata function and photosynthesis,<br />

and cytology and ultrastructure.<br />

3. THE INFLUENCE OF PATHOGENS ON INTERFERENCE<br />

AND ALLELOPATHY<br />

Natural plant populations increase in production and number of individuals until<br />

constrained by environmental limitations (Burdon, 1987). The constraint of plant<br />

growth by the environment is often mediated through plant interference. Therefore,<br />

the ability of plants to interfere with their neighbours is important in determining<br />

their abundance in a community. Plant pathogens generally reduce the development,<br />

production and longevity of their hosts. In plant communities, this ‘burden of a<br />

parasite’ may result in a partly unoccupied niche that resistant plants re-inhabit.<br />

Overall, pathogens play a significant role in the ecology of plant communities by<br />

maintaining (Peters and Shaw, 1996) or reducing (Burdon, 1991) species diversity,<br />

driving succession (Van der Putten et al., 1993), ensuring plants do not establish<br />

under their parents (Augsberger, 1984) and help determine the long-term composition<br />

of plant communities (Dobson and Crawley, 1994).

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