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412 L.G. Perry et al.<br />

27.4<br />

(±)-Catechin and C. maculosa Autoinhibition<br />

In addition to reducing the success of C. maculosa’s interspecific competitors,<br />

(±)-catechin appears to reduce C. maculosa seedling recruitment<br />

in well-established populations. In C. maculosa stands in North America,<br />

adult plants are sometimes widely spaced, with the interspaces between<br />

plants unoccupied, occupied by C. maculosa seedlings that fail to establish,<br />

or occupied by other species. Perry et al. (2005b) examined whether this<br />

pattern might be accounted for by (±)-catechin inhibition of C. maculosa<br />

seedling establishment (i.e., autoinhibition).<br />

To test whether organic compounds in the soil in C. maculosa stands<br />

limited C. maculosa seedling establishment, Perry et al. (2005b) added activated<br />

carbon, which adsorbs organic compounds (Mahall and Callaway<br />

1992), to the soil around adult C. maculosa plants in a well-established<br />

population in the field. Activated carbon addition increased C. maculosa<br />

seedling densities by more than 30%, indicating a strong negative effect of<br />

organic compounds in the soil around C. maculosa adults on C. maculosa<br />

seedling establishment. Seedling densities were still visibly greater in the<br />

soil with activated carbon than in the soil without activated carbon in some<br />

plots 1 year after the start of the experiment (L. Perry, personal observation),<br />

indicating that the effects of organic compounds in the soil around<br />

C. maculosa adults on C. maculosa seedling establishment are persistent.<br />

Measurements of soil (±)-catechin around adult C. maculosa at the field<br />

site indicated that (±)-catechin concentrations in the site were extremely<br />

high (mean 1.55 mg g −1 dry soil). C. maculosa seedling densities at the site<br />

were very low (mean 28 seedlings per square meter). To test whether the<br />

high soil (±)-catechin concentrations could account in part for the low<br />

C. maculosa seedling establishment, Perry et al. (2005b) examined the response<br />

of C. maculosa seedlings to (±)-catechin in laboratory experiments.<br />

Treatment with 1.0 mg ml −1 of (±)-catechin reduced C. maculosa seedling<br />

root lengths by 50%, demonstrating that C. maculosa soil (±)-catechin<br />

concentrations are sufficient to substantially reduce C. maculosa seedling<br />

growth. Higher (±)-catechin concentrations (2 and 4 mg ml −1 ) reduced<br />

C. maculosa root elongation by as much as 75%. Reduced root elongation<br />

did not result in seedling mortality under laboratory conditions, but would<br />

be expected to reduce survival under field conditions with limited water. In<br />

addition, when (±)-catechin was maintained in solution in 10% methanol<br />

for 2 days after treatment, treatment with 1.0 mg ml −1 of (±)-catechin reduced<br />

C. maculosa germination by more than 70%. Tetrazolium analyses of<br />

seed viability indicated that (±)-catechin treatment did not reduce seed survival,<br />

suggesting that (±)-catechin dissolved in methanol may have delayed<br />

germination of C. maculosa seeds.

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