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

monooxygenase, lipid transfer protein, heat shock protein, DNA-J protein,<br />

and blue copper-binding protein (Bais et al. 2003). Phenylpropanoid and<br />

terpenoid phytoalexin pathway genes, a number of which produce enzymes<br />

that can act as antioxidants (Sticher et al. 1997), were also induced in the<br />

roots 1 h after catechin treatment. However, the most interesting change<br />

in gene expression that they observed was the upregulation of ten genes<br />

in the first 10 min after catechin treatment. These included genes associated<br />

with calcium signaling and oxidative stress, as well as four unknown<br />

genes lacking homology with genes from other organisms. Identification<br />

of the function of these unknown genes could provide insights into the<br />

factors that cause plants to be susceptible to catechin, and into potential<br />

mechanisms of resistance.<br />

27.3.4<br />

(±)-Catechin Is Present at Phytotoxic<br />

Concentrations in C. maculosa Soils<br />

While laboratory experiments demonstrated that (±)-catechin is a potent<br />

phytotoxin with strong effects on plant biochemistry and gene expression,<br />

field observations were required to gauge the importance of (±)-catechin<br />

in C. maculosa competitive interactions under natural conditions. In particular,<br />

for (±)-catechin to influence plant interactions, it must be present<br />

at phytotoxic concentrations in C. maculosa field soils. Several studies<br />

have reported exceptionally high soil (±)-catechin concentrations in North<br />

American C. maculosa soils (Bais et al. 2002, 2003; Perry et al. 2005b), indicating<br />

that soil (±)-catechin is present in sufficient quantities in C. maculosa<br />

soil to inhibit plant neighbors. Bais et al. (2003) reported a mean soil<br />

(±)-catechin concentration of 2.24±0.20 mg g −1 and Perry et al. (2005b)<br />

reported a mean soil (±)-catechin concentration of 1. 55 ± 1.27 mg g −1 dry<br />

soil. In addition, tests of the soil extracts from one site confirmed that the<br />

(±)-catechin in C. maculosa soils has similar phytotoxicity to (±)-catechin<br />

from commercial sources (Perry et al. 2005b). Bais et al. (2002) found that<br />

(±)-catechin concentrations declined with distance from the C. maculosa<br />

taproot, and with increasing soil depth. However, Perry et al. (2005b) found<br />

that soil (±)-catechin concentrations did not change with distance from the<br />

C. maculosa taproot but remained high as far as 25 cm from the taproot,<br />

indicating that high soil (±)-catechin concentrations may be ubiquitous in<br />

at least some well-established C. maculosa populations. The differences in<br />

soil (±)-catechin concentrations among the studies are not surprising, since<br />

the studies were conducted in different locations and at different times. The<br />

effects of soil characteristics, climate, and season on (±)-catechin secretion,<br />

stability, and soil absorption are not yet understood.

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