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IMPACT OF PATHOGENS ON PLANT INTERFERENCE AND<br />

ALLELOPATHY<br />

The production of glucosinolates by plants in the Capareles and their subsequent<br />

hydrolysis to toxic isothiocyanates, thiocyanates, and nitriles has been one of the<br />

most intensively studied systems in allelopathy, due partly to the similarity of these<br />

breakdown products to some synthetically produced soil fumigants (and therefore<br />

termed biofumigation; Kirkegaard et al., 2000). Glucosinolates are important in plant<br />

defence against insects, pathogens and nematodes. Jay et al. (1999) showed that<br />

infection of Brassica napus with beet western yellows virus increased glucosinolate<br />

concentration in tissues by 14%. Similarly, Li et al. (1999) found that infection of B.<br />

napus by Sclerotinia sclerotiorum increased glucosinolate content in resistant, but<br />

not in susceptible varieties. Exposure to the pathogenic bacterium, Erwinia carotovora,<br />

triggered the production of glucosinolates in Arabidopsis thaliana (Brader et al.,<br />

2001). The hydrolysis products from these glucosinolates inhibited the growth of E.<br />

carotovora in culture. Furthermore, Tierens et al. (2001) demonstrated that a range<br />

of pathogens were more aggressive in infecting an A. thaliana mutant that did not<br />

produce glucosinolates than the wild type, suggesting the importance of glucosinolates<br />

in protecting against infection. However, the role of glucosinolates in disease resistance<br />

may be species specific, since Andreasson et al. (2001) found that infection of B.<br />

napus by Leptosphaeria maculans had no effect on glucosinolate concentration in the<br />

resistant or susceptible host. Despite the ability for at least some pathogens to increase<br />

glucosinolate production in the Capareles, no one has investigated whether this directly<br />

translates to an increased allelopathic effect against neighbouring plants.<br />

3.2.2. The Effect of Rust on Ryegrass Allelopathy<br />

Perennial ryegrass (Lolium perenne) and white clover (Trifolium repens) are important<br />

components of improved pastures grown in temperate regions worldwide. The ability<br />

of ryegrass to become dominant in pastures has led to numerous investigations that<br />

demonstrate its potential to interfere with companion plants through allelopathy (Naqvi,<br />

1972; Naqvi and Muller, 1975; Newman and Rovira, 1975; Newman and Miller,<br />

1977; Gussin and Lynch, 1980; Buta et al., 1987; Quigley et al., 1990; Sutherland<br />

and Hoglund, 1990; Wardle et al., 1991; Prestidge et al., 1992; Chung and Miller,<br />

1995; Mattner, 1998; Mattner and Parbery, 2001). For example, in a study investigating<br />

the allelopathic ability of nine pasture species, Takahashi et al. (1988) found that<br />

leachate from soil surrounding ryegrass was the most inhibitory to the growth of the<br />

target species, including clover. In a subsequent experiment, they circulated nutrient<br />

solution between the roots of ryegrass and clover to eliminate competition effects.<br />

They found that the growth of clover declined in the system, particularly when the<br />

proportion of ryegrass was high. When they incorporated XAD-4 resin into the system<br />

(which selectively traps organic hydrophobic compounds) clover grew normally.<br />

Moreover, ryegrass became yellow and stunted when grown alone in the system, but<br />

this was prevented by the presence of the resin or clover (Takahashi et al., 1991). In<br />

a further experiment, root exudate from ryegrass not only inhibited the growth of<br />

clover, but also lettuce seedlings. The phytotoxic fraction of the extract contained pmethoxybenzoic,<br />

lauric, myristic, pentadecanoic, palmitoleic, palmitic, oleic and stearic<br />

87

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