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27 Root exudation and rhizosphere biology 413<br />

Plant autoinhibition mediated by secondary metabolites has been the<br />

subject of much research (Singh et al. 1999). However, (±)-catechin is one<br />

of the first secondary metabolites determined to act as both an allelochemical<br />

and an autoinhibitor. Autoinhibition of seedling establishment may<br />

influence plant populations in several ways. First, autoinhibition may be<br />

a mechanism through which adults avoid establishment of intraspecific<br />

competitors in dense populations (Schenk et al. 1999). Second, seedlings<br />

that produce autoinhibitors may reduce intraspecific competition by preventing<br />

the establishment of their siblings and unrelated seedlings (Dyer<br />

2004). Third, autoinhibitors that delay germination, rather than killing<br />

seeds or seedlings, may postpone germination in areas where intraspecific<br />

competition from adults would prevent seedling survival (Picman<br />

and Picman 1984). Determining the specific effects of autoinhibition on<br />

C. maculosa populations will require further research to determine whether<br />

(±)-catechin reduces C. maculosa seedling root elongation or delays germination<br />

under field conditions.<br />

Autoinhibition by C. maculosa is probably most important in wellestablished<br />

stands, where soil (±)-catechin is likely to be concentrated<br />

and widespread. (±)-Catechin concentrations lower than 0.5 mg ml −1 have<br />

little effect on C. maculosa root elongation, and no effect on germination<br />

(Perry et al. 2005b). However, much lower (±)-catechin concentrations have<br />

large effects on other, more (±)-catechin-sensitive species (Bais et al. 2003;<br />

Weir et al. 2003; Perry et al. 2005a). Thus, C. maculosa seedlings should<br />

have an advantage over more (±)-catechin-sensitive species in soils with<br />

moderate (±)-catechin concentrations, while very high (±)-catechin concentrations<br />

may prevent establishment of C. maculosa seedlings as well<br />

as other species. In well-established C. maculosa populations with high<br />

soil (±)-catechin concentrations, autoinhibition, water stress (Jacobs and<br />

Sheley 1998), and seed predation (Muller-Schärer and Schroeder 1993)<br />

probably operate together to limit C. maculosa seedling establishment.<br />

27.5<br />

(±)-Catechin Effects on Soil Communities<br />

In addition to the inhibitory effects of (±)-catechin on plants, (+)-catechin<br />

and (–)-catechin both reduce survival of some soil organisms. However,<br />

the two enantiomers appear to affect different organisms. Bais et al. (2002)<br />

found that (+)-catechin, but not (–)-catechin, inhibited growth of several<br />

soil bacteria. Six bacterial strains were exposed to each of the enantiomers<br />

or the racemic mixture, applied to filter discs. Xanthomonas campestris<br />

ssp. vesicatoria, Erwinia amylovora, Pseudomonas fluorescens,andE. carotovora<br />

were each susceptible to 100 µg ml −1 (+)-catechin or to 200 µg ml −1

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