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3 Methaogenic Microbial Communities Asociated with Aquatic Plants 43<br />

Another implication of the observations concerns the structure of the<br />

methanogenic microbial community on the roots, which seem to be very simple,<br />

consisting only of H 2-producing fermenting and H 2-consuming methanogenic<br />

microorganisms. However, experiments with excised rice roots have<br />

demonstrated a more complex community of fermenting bacteria including<br />

vigorous fermentative production of acetate, propionate and butyrate (Conrad<br />

and Klose 1999, 2000). Interestingly, a significant percentage (up to 60 %)<br />

of these fatty acids was produced by reduction of CO 2. The stable isotope signature<br />

of the produced acetate was consistent with the production by CO 2<br />

reduction (Conrad et al. 2002). Acetate production from CO 2 indicates that<br />

homoactogenic bacteria were active, a likely conclusion, since homoacetogenic<br />

Sporomusa are members of the rice root microflora (Rosencrantz et al.<br />

1999). Homoactogens have also been found on the roots of sea grass (Küsel et<br />

al. 1999, 2001). Approximately 30 % of the root epidermal cells of sea grass<br />

were colonized with microorganisms that hybridized with an archaeal probe<br />

suggesting the presence of methanogens (Küsel et al. 1999). Presently, little is<br />

known about the fate of the produced fatty acids. Propionate and butyrate can<br />

potentially be further converted to acetate, CO 2 and H 2 by syntrophic bacteria,<br />

which are present in the anoxic rice soil, followed by H 2/CO 2-dependent<br />

methanogenesis (Krylova et al. 1997). Syntrophic oxidation of acetate, however,<br />

is unlikely since [2- 14 C]acetate was hardly turned over in root preparations<br />

(Lehmann-Richter et al. 1999). The most likely fate of the acetate produced<br />

by the root microflora is its escape into the bulk soil where it is<br />

methanogenically decomposed (Fig. 5). Alternatively, acetate may be a substrate<br />

for anaerobic bacteria using nitrate, ferric iron or sulfate as electron<br />

Fig. 5. Conceptual model of the<br />

localization of methanogenic<br />

archaea (MA), homoacetogenic<br />

bacteria (HAB), methane-oxidizing<br />

bacteria (MOB) and aerobic<br />

bacteria (AB) in vicinity of rice<br />

roots and to each other, and the<br />

flow of organic carbon. The insertion<br />

of lateral roots (and root<br />

tips) are the most likely sites<br />

where O 2 and organic substrates<br />

(e.g., sugars) are released into the<br />

soil

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