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300 P. Lavelle et al.<br />

Fig. 4. Comparison of genetic structures of bacteria based on PCR IGS analysis of 16S and<br />

23S DNA, in soil microsites: Ew– control with no earthworms; Ew + non-ingested soil<br />

treated with earthworms; CAST earthworm casts; BUR burrow wall of earthworm gallery;<br />

GC earthworm gut content<br />

Changes in the activity and composition of microbial communities in<br />

earthworm biogenic structures have been observed by many authors (Parle<br />

1963; Scheu 1987; Tiwari and Mishra 1993; Tiunov and Scheu 1999). Microbial<br />

numbers and biomass increase by a factor of 2.3–4.7 in burrow walls<br />

of Lumbricus terrestris mainly due to changes in the individual bacterial<br />

cell volume (Tiunov and Scheu 1999), with a 3.7–9.1 increase in respiration<br />

rate. These results confirm earlier estimates by Bhatnagar (1975) who<br />

claimedthatupto5–25%oftotalsoilmicroflorawasincludedinthis<br />

microenvironment in temperate pastures of central France. Recent studies<br />

using molecular techniques have shown evidence of the occurrence<br />

of selective microbial communities in Lumbricidae earthworm casts and<br />

burrows (Fig. 4; Kersanté 2003). The bacterial community in the gut contents<br />

differed the most from the original soil, casts and to a lesser extent,<br />

burrow walls, having intermediate communities. This suggests that digestive<br />

processes would select a rather specific microbial community through<br />

the Sleeping Beauty effect, and progressive changes would occur in ageing<br />

structured communities that would slowly converge towards a common<br />

“bulk soil” type of community. An alternative hypothesis, although difficult<br />

to support, could be that earthworms harbour in their guts a specific

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