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Allelochemicals Biologica... - Name

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92<br />

SCOTT W. MATTNER<br />

from three ryegrass cultivars infected with three different strains of endophyte, all<br />

inhibited the growth of clover, by up to 27% compared with extracts from endophytefree<br />

ryegrass. Both ryegrass cultivar and endophyte strain influenced the degree that<br />

ryegrass extracts inhibited clover, but this did not relate to the type of alkaloids produced<br />

by the different endophyte strains. These studies suggest that both environmental<br />

and genetic influences may moderate the triggers for enhanced allelopathy by endophyte<br />

infected grasses, and this may explain the discrepancy in results between individual<br />

studies. Nonetheless, the majority of evidence suggests that endophyte infection has<br />

the capacity to increase ryegrass allelopathy, which is a further added benefit conferred<br />

by this mutualist to its host. Although the endophyte is a non-pathogenic organism,<br />

its ability to stimulate plant allelopathy adds further weight to the hypothesis that<br />

infection increases the allelopahtic ability of host plants.<br />

3.2.4. Other Systems<br />

The ability of rusts to stimulate allelopathy in their hosts may not be limited to ryegrass,<br />

as the rusts Puccinia hordei and Uromyces troflii-repentis increased the suppression<br />

of white clover by barley grass (Hordeum leporinum) and subterranean clover (Trifolium<br />

subterraneum), respectively (Mattner, 1998). Yet, the effect was not universal since<br />

Puccinia coronata in wild oat (Avena fatua), Puccinia graminis in cocksfoot (Dactylis<br />

glomerata) and Puccinia recondita in soft brome (Bromus mollis) all failed to increase<br />

their host’s allelopathic ability.<br />

Kong et al. (2002) studied the allelopathic potential of goatweed (Ageratum<br />

conyzoides) under different environmental stresses, including infection by powdery<br />

mildew (Erysiphe cichoracearum). Infection stimulated the production of 17 of the<br />

24 volatile chemicals produced by goatweed that they investigated, with total volatile<br />

production increasing by 50%. Exposure to the volatiles released by infected goatweed<br />

stimulated the growth of peanut (Arachis hypogaea), redroot amaranth (Amaranthus<br />

retroflexus), Italian ryegrass (Lolium multiflorum) and cucumber (Cucumis sativus)<br />

compared with volatiles from healthy goatweed. In contrast, volatiles from infected<br />

plants inhibited the growth of three fungal pathogens (Rhizoctonia solani, Botrytis<br />

cinerea, and Sclerotinia sclerotiorum). For this reason, they postulated that the<br />

allelochemicals stimulated by fungal infection in goatweed are more important in the<br />

defence of the plant against infection, rather than against competition from<br />

neighbouring plants. Nonetheless, this study currently provides the clearest<br />

demonstration of the ability of pathogens to influence the allelopathic ability of plants.<br />

4. IMPLICATIONS FOR PATHOGEN<br />

AGGRESSIVENESS AND EVOLUTION<br />

Plant/pathogen interactions in natural populations are often explained in terms of coevolution,<br />

which is ‘the joint evolution of two (or more) taxa that have close ecological<br />

relationships but do not exchange genes, and in which reciprocal selective pressures<br />

operate to make the evolution of either taxon partially dependent upon the evolution

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