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184<br />

Michael Kaldorf et al.<br />

1995). Similarly, the effects of transgenic potatoes producing the lectin GNA<br />

on nontarget soil organisms could not be attributed to the formation of the<br />

lectin GNA itself (Griffiths et al. 2000). So far, only one GMP producing an<br />

antibacterial substance, namely T4 lysozyme-producing potatoes with<br />

enhanced resistance to Erwinia carotovora, has been investigated in detail for<br />

nontarget effects on soil bacteria. Most soil bacteria were lysozyme-sensitive<br />

when tested in laboratory experiments with pure cultures (de Vries et al.<br />

1999). In addition, increased killing of Bacillus subtilis was observed on the<br />

root <strong>surface</strong> of T4 lysozyme-producing potatoes from the field, and this effect<br />

was ascribed directly to the lysozyme release by the roots (Ahrenholtz et al.<br />

2000). Nevertheless, the production of lysozyme had only a minor influence<br />

on the bacterial phyllo- and rhizosphere communities (Heuer and Smalla<br />

1999; Heuer et al. 2002), which was considered negligible relative to natural<br />

factors by the authors. Further studies with the same system which focused on<br />

potentially beneficial <strong>plant</strong>-associated microbes like auxin-producing bacteria<br />

or bacteria antagonistic to the pathogenic E. carotovora did not reveal correlations<br />

between the transgenic character of <strong>plant</strong>s and the pheno- or genotypic<br />

features of bacterial isolates (Lottmann and Berg 2001). Thus, up to now,<br />

there is no direct evidence from field experiments that the primary product of<br />

transgene expression is responsible for significant changes in the soil microbial<br />

community in any GMP. Instead, secondary effects of GMP generation,<br />

like somaclonal variation and changes in general <strong>plant</strong> metabolism induced<br />

by the transgene insertion or expression, may contribute to a major part of<br />

the effects described above.<br />

3 Impact of Genetically Modified Plants on Symbiotic<br />

Interactions<br />

The question whether the genetical transformation of <strong>plant</strong>s might reduce<br />

their ability to form mutualistic symbioses with microorganisms has been<br />

addressed in a surprisingly small number of studies.<br />

Biological nitrogen fixation accounts for about 65 % of the nitrogen utilized<br />

in agriculture worldwide (Vance and Graham 1995). The ability to<br />

reduce atmospheric nitrogen to ammonia (nitrogen fixation) is restricted to<br />

prokaryotes. Beside free-living and <strong>plant</strong>-associated bacteria, members of the<br />

Rhizobiaceae living symbiotically in the typical root nodules of legumes such<br />

as alfalfa, clover, pea, and soybean are the agriculturally most important<br />

group of nitrogen fixing organisms. The symbiotic interaction between rhizobia<br />

and legumes requires a sequential signal exchange between both partners,<br />

and therefore, exhibits a high degree of host specificity (Bothe 1993). Transgenic<br />

<strong>plant</strong>s have been used as a tool to investigate the host recognition of rhizobia<br />

(Diaz et al. 1989, 2000). For example, the transfer of lectin genes between<br />

different legumes has been shown as a possible way to modify host specificity

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