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202<br />

Rüdiger Hampp and Andreas Maier<br />

fluorescent pseudomonad strains (Founoune et al. 2002). After 3 months of<br />

co-culture, the bacterial inoculants disappeared, showing how difficult such<br />

experiments are to interpret. Obviously, the amount of inoculum supplied can<br />

also play an important role (Frey-Klett et al. 1999).<br />

7 Possible Mechanisms of Interaction<br />

As pointed out by Schelkle and Peterson (1996), and in addition to the production<br />

of antibiotics, protective or “helper” effects could be due to competition<br />

for nutrients in the rhizosphere. The formation of siderophores, for<br />

example could be such a mechanism. Siderophores are iron chelators which<br />

make iron available for uptake by the bacteria. As these compounds are<br />

species-specific, Fe-chelates can only be taken up by those bacteria that are<br />

able to produce them. Protective bacteria synthesizing siderophores could<br />

thus out-compete pathogens with regard to Fe (Neidhardt et al. 1990). Similar<br />

mechanisms are known for ECM fungi (Watteau and Berthelin 1990).<br />

Siderophore release from bacteria into the mycorrhizosphere could also<br />

improve absorption of Fe by the mycorrhizal fungus.<br />

Protection from pathogens could, however, also be a mass effect, simply<br />

due to the large number of nonpathogenic bacteria that accumulate in the rhizosphere<br />

due to the high nutrient supply. However, as outlined by Garbaye<br />

(1994), there can be many more mechanisms, such as an improved receptivity<br />

of the root for mycorrhizal infection, a modification of the rhizospheric soil,<br />

improvement of the root-fungus recognition, stimulation of germination of<br />

fungal propagules, as well as an enhancement of fungal growth in the rhizosphere<br />

(see also Brule et al. 2001) which would increase the probability of contact<br />

between fungus and root (compare Dunstan et al. 1998).<br />

In nutrient-poor acidic forest soils modification by micro-organisms<br />

should be an important factor; the C-rich environment provided by the <strong>plant</strong><br />

is attractive for soil micro-organisms, leading to the formation of functionally<br />

compatible microbial communities. These are jointly able to co-mobilize soil<br />

nutrients such as P and N in and around the vegetative mycelium. In addition,<br />

N-fixing bacterial species including Bacillus spp. are possibly present in the<br />

mycorrhizosphere of forest trees (Li et al. 1992) as the vegetative mycelium<br />

represents a niche that is ideally suited for the selection and enrichment of<br />

associative N-fixing bacteria (Sen 2000).<br />

In many of the possible mechanisms, phytohormones such as IAA could<br />

play an important role. A study on the rooting of derooted shoot hypocotyls<br />

of spruce showed that Laccaria bicolor and Pseudomonas fluorescens BBc6<br />

(MHB) both increased the number of roots formed per rooted hypocotyl<br />

(Karabaghli et al. 1998). The same effect was caused by the addition of IAA<br />

alone (control). Both organisms produced IAA in pure culture.

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