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Interactions Between Microorganisms and Soil Micro- and Mesofauna 257<br />

and cannibalism are certainly widespread in decomposer food-webs (Polis<br />

1991; Gunn and Cherrett 1993), which is supported by soil food web analyses<br />

based on stable isotopes (Ponsard and Arditi 2000; Scheu and Falca 2000).<br />

The dominance of generalist predators and the high incidence of IGP and<br />

cannibalism in soil communities add to the intriguing complexity of belowground<br />

food webs. It is likely that the exceptionally high local diversity<br />

ofsoilorganismsatleastinpartresultsfromthefactthatsoilpredatorsare<br />

so closely intermingled. The very large overlap in prey (among predators)<br />

may prevent competitively superior microbi-detritivores from outcompeting<br />

inferior species, despite close resource overlap among them.<br />

The similarity of trophic positions of species in very different taxonomic<br />

groups within microbi-detritivores and predators supports the assumption<br />

that functional redundancy among soil animals is high (Wardle et al.<br />

1997, 1998; Setälä et al. 1998). High redundancy may help to explain why<br />

decomposer populations are extraordinary stable in time (Bengtsson 1994)<br />

and respond little to external perturbations (Scheu and Schaefer 1998; Joergensen<br />

and Scheu 1999). Decomposer diversity–ecosystem functioning<br />

experiments also support the view of high redundancy in detritus communities<br />

(Mikola et al. 2002; Wardle 2002).<br />

2.2<br />

The Detritus vs. Root Exudate-Based Food Web<br />

Plants supply two quite different energy sources to the soil food web, root<br />

exudates and plant debris (Fig. 3). Plant organic matter fuels the detritus<br />

food web, which acts as two distinct, bacterial and fungal-based compartments<br />

(Wardle and Yeates 1993). Carbon is transferred to successively<br />

higher trophic levels and nutrients bound in microbial biomass are liberated<br />

for plant uptake. Root exudates build the resource for a separate short<br />

and fast energy channel. In the rhizosphere, easily degraded carbon sources<br />

become available to microorganisms leading to a biomass often several<br />

times greater than in corresponding nonassociated soil (Wardle 2002). In<br />

contrast to bacterial and fungal channels, the interactions between plants,<br />

microbes and grazers form a loop (‘microbial loop’; Clarholm 1994). The<br />

dominant grazers, protozoa and nematodes may effectively control microbial<br />

species, but themselves appear to be little controlled by higher order<br />

predators (Scheu and Setälä 2002; Wardle 2002), resulting in carbon to be<br />

processed in the loop and not progress to the remainder of the soil food<br />

web.<br />

Theactivityofmicroorganismsinsoilisusuallylimitedbycarbon,<br />

except in the rhizosphere where plants steadily supply easily available carbonsources.Thisfavorsamicrofloratypicallyconsistingoffast-growing

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