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Role of Microorganisms in Carbon Cycling in Soils 145<br />

Plant debris<br />

1cm 1mm<br />

minerals<br />

5µm<br />

bacteria<br />

50 µm<br />

Fig. 3. Hierarchy of soil aggregates (Oades and Waters 1991, modified). The graphs shows:<br />

the bulk soil (above, left), a macroaggregate consisting of smaller aggregates including<br />

plant debris (above, right), an intermediately sized aggregate composed of microaggregates<br />

(below, right), a microaggegate showing minerals and bacteria associated with plant debris<br />

(below, center)andamicroaggregatewithstabilizedsoilorganicmatter(below, left)<br />

al. 1991; Schulten et al. 1993; Desjardins et al. 1994; Guggenberger et al. 1994)<br />

point out that Corg content increases with diminishing particle size, whereas<br />

the C/N ratio decreases. Jastrow et al. (1996) showed that organic matter<br />

recently introduced into soil is predominantly located in larger aggregates.<br />

Tisdall and Oades (1982) demonstrated that macroaggregates (> 250 µm)<br />

can be destroyed by agricultural practices, whereas microaggregates cannot.<br />

Figure 3 shows the concept of the aggregate hierarchy. (1) Plant residues<br />

are introduced into the soil. This young organic matter comes into contact<br />

with the mineral substances, e.g., through macrofaunal action, and<br />

will be mainly contained in large aggregates several millimeters in size.<br />

(2) During the decomposition of organic matter, the size of the organic<br />

particles decreases, and the stability of the remaining Corg increases. This<br />

Corg fraction is then located in smaller aggregates, which partly compose<br />

larger ones. (3) Finally, we observe small mineral–organic matter–microbe<br />

associations. After the death of the microbes, the mineral–organic matter<br />

complexes, that physically protect the organic carbon against further decomposition,<br />

remain. These associations are called microaggregates and<br />

are highly stable.<br />

Physical separation of particle size fractions revealed that much of the<br />

soil microbial biomass is associated with the smaller fractions (fine silt and<br />

clay); the highly variable amount of microbial biomass within the coarse<br />

fractions strongly depends on the quantity and quality of the macroorganic<br />

matter located there (Jocteur Monrozier et al. 1991). Investigations

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