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IMPACT OF PATHOGENS ON PLANT INTERFERENCE AND<br />

ALLELOPATHY<br />

81<br />

1988; Grace and Tilman, 1990; Casper and Jackson, 1997) and allelopathy (Putnam<br />

and Duke, 1978; Rice, 1984; Inderjit et al., 1995; Anaya, 1999) already occur in the<br />

literature.<br />

The view of competition in the present chapter is ‘the interaction between<br />

individuals brought about by the shared requirements for a resource in limited supply,<br />

and leading to a reduction in the survivorship, growth and/or reproduction of the<br />

individuals concerned’ (Begon et al., 1986). Plants mostly compete for the resources<br />

of light, water and nutrients (Donald, 1963), and less frequently for carbon dioxide,<br />

oxygen and space (Trenbath, 1974). In the exact sense of the definition, plants do not<br />

compete so long as these resources are in excess of the needs of both. When plants do<br />

compete, however, the outcome always reduces the growth of at least one of the<br />

competitors. The outcome of competition between two plants depends on the ability<br />

of each species to secure and utilise resources, i.e. their competitiveness. A highly<br />

competitive plant may be one that has a high rate of uptake of a particular resource or<br />

a low requirement for that resource (Grace and Tilman, 1990). Plants may differ in<br />

their ability to compete for individual resources, while environmental differences may<br />

also influence their overall competitive ability. Differences in competitive ability, in<br />

turn, help to structure the composition of mixed plant communities.<br />

2.2. Allelopathy<br />

Molisch first advanced the term allelopathy in 1937. It derives from the Greek words<br />

‘allelon’, meaning of each other, and ‘pathos’, meaning to suffer (Mandava, 1985).<br />

Despite the origin of its root words, Molisch used the term to refer to the chemical<br />

interactions between all plants (higher plants and microorganisms), including<br />

stimulatory as well as inhibitory effects. Some authors have considered that the concept<br />

covers only inhibitory effects (Rice, 1974; Putnam, 1985; Boyette and Abbas, 1995),<br />

yet others exclude microorganisms from the definition (Putnam and Duke, 1978;<br />

Putnam, 1985; Pratley, 1996). Many inhibitory chemicals produced by plants, however,<br />

stimulate growth at low concentrations (Liu and Lovett, 1993; Pratley, 1996), and<br />

microorganisms can mediate allelopathy (Rice, 1992; Bremner and McCarty, 1993).<br />

For these reasons, the definition of allelopathy used in this chapter closely follows<br />

that of Molisch’s, ‘the beneficial and detrimental chemical interaction among [plant]<br />

organisms including microorganisms’ (Rice, 1984). Although some authors have<br />

confused allelopathy with competition, the distinguishing feature is that allelopathy<br />

involves an addition of a chemical to the environment, whereas competition involves<br />

the shared utilisation of some limited factor required for growth (Muller, 1969; Rice<br />

1984).<br />

Any chemical produced by a plant (donor) that stimulates or inhibits the growth<br />

of a neighbour (receiver or receptor) is broadly termed an allelochemical. Typically,<br />

allelochemicals are secondary metabolites (Whittaker and Feeney 1971; Rice, 1984;<br />

Rizvi et al., 1992), produced as by-products of the acetate and shikimic acid pathways.<br />

They may also form as degradation products from the action of microbial enzymes

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