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200<br />

Rüdiger Hampp and Andreas Maier<br />

Pioneering work in this field has been carried out by Garbaye (for a review<br />

see Garbaye 1994). Experiments carried out with Picea abies, Pinus nigra,<br />

Pinus sylvestris, Pseudotsuga menziesii, and Quercus robur (Garbaye et al.<br />

1992) indicated that soil bacteria can stimulate the inoculation of roots with<br />

ectomycorrhiza-forming fungi, thereby also reducing the adverse effect of<br />

pathogens. Both effects resulted in a better seedling growth, and thus the term<br />

“helper bacteria” was coined. For more recent work, see Dunstan et al. (1998)<br />

and Poole et al. (2001).<br />

5 Bacteria Associated with Sporocarps and Ectomycorrhiza<br />

Twenty seven bacterial species were isolated from both the sporocarps of Suillus<br />

grevillei and the ECMs of S. grevillei/Larix decidua (Varese et al. 1996). The<br />

genera Pseudomonas, Bacillus, and Streptomyces were predominant. From<br />

sporocarps of white truffles (Tuber sp.), bacterial strains such as Micrococcus,<br />

Moraxella, Staphylococcus and Pseudomonas could be isolated (Citterio et al.<br />

1995). Gram-positive bacteria seldom stimulated in vitro fungal growth.<br />

Among gram-negative bacteria, Pseudomonas strains enhanced growth.<br />

Streptomyces significantly inhibited the fungus. Bacterial supernatants were<br />

not effective.Volatiles enhanced fungal growth to some extent, but not significantly.<br />

Most of the bacteria isolated produced siderophores.<br />

A distinction between the different structures of ECM showed that bacteria<br />

primarily occurred on the <strong>surface</strong> of the mantle and in the interhyphal spaces<br />

(Schelkle et al.1996),but also deep within the mantle (Foster and Marks 1967).<br />

Bacteria of subclasses of proteobacteria (containing <strong>plant</strong>-growth-promoting<br />

rhizobacteria such as Burkholderia, Azospirillum, Acetobacter and Herbaspirillum)<br />

were detected in high numbers on mantle <strong>surface</strong>s (Mogge et al.<br />

2000). The two most common fungi on beech, Lactarius vellereus and Lactarius<br />

subdulcis, were associated with members of the a- and b-subclasses of the<br />

proteobacteria. These bacteria have been shown to be abundant in winter and<br />

early spring (Timonen et al. 1998).<br />

Electron microscopy of ECM with Pinus sylvestris and S. bovinus and Paxillus<br />

involutus (Nurmiaho-Lassila et al. 1997) also revealed bacteria on the<br />

mantle <strong>surface</strong> and at inter- and intracellular locations in the mantle and the<br />

Hartig net (S. bovinus). Fungal strands were colonized only by a few bacteria,<br />

while the outermost external fine hyphae had extensive monolayers of bacteria<br />

attached.<br />

ECM with P. involutus were mostly devoid of bacteria, while the external<br />

mycelium supported bacteria (Nurmiaho-Lassila et al.1997).From their observations,<br />

the authors conclude that single ECM fungi create defined mycorrhizosphere<br />

habitats with distinct populations of bacteria. Knowing that several<br />

different types of ECM can be formed on the same root in close vicinity,a large<br />

local biodiversity of ECM-specific bacterial populations could be postulated.

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