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Genetically Modified Organisms as Invasive Species? 297<br />

Transgenes placed on the chromosome will have much lower rates of escape<br />

than those placed on plasmids within the bacteria, even though gene flow<br />

between chromosomes of unrelated bacteria does occur in evolutionary time,<br />

via conjugation or, more commonly, transduction.<br />

17.3.1.4 Evidence for Gene Transfer from GMMs<br />

Most, if not all studies of the field and microcosm release of GMMs have<br />

included a component to specifically study the possibility of gene transfer to<br />

or from the transgenic bacteria, allowing us to test the prediction posed<br />

above. To date, no investigation has revealed an outcome which had not been<br />

predicted and no evidence has been provided demonstrating the invasion of<br />

transgenes from genetically stable constructs. Therefore, from existing evidence,<br />

it is apparent that the likelihood of gene transfer directly equates to the<br />

method of genetic construction. When introduced genes were located on<br />

mobile genetic elements such as conjugative plasmids, transfer has been<br />

observed in laboratory investigations. However, as this outcome was entirely<br />

predictable, field releases have involved bacteria genetically modified to carry<br />

transgenes on their chromosomes. Insertions have been mediated by the use<br />

of transposons, disarmed transposons or by site-directed homologous<br />

recombination. The order in which these three approaches are listed represents<br />

their relative genetic stability and, therefore, the likelihood of being<br />

transferred (Bailey et al. 1995; Troxler et al. 1997). In all the reports of field<br />

testing of bacteria modified to carry transgenes on their chromosomes, none<br />

have found either the transfer or loss of the transgenes. Even in the most<br />

appropriately designed laboratory investigations, transfer frequencies were<br />

negligible or effectively zero (Bailey et al. 1995; Troxler et al. 1997).<br />

17.3.2 Gene Escape in Plant Communities<br />

Gene escape from a plant species involves two steps: hybridisation with a wild<br />

relative, and the survival and reproduction of resulting hybrids. Successful<br />

hybridisation between crop plants and wild relatives was earlier thought to be<br />

infrequent (Ellstrand et al. 1999). This is probably because gene flow was not<br />

perceived to be of significant concern – people had simply not looked.A comprehensive<br />

review of the world’s most important food crops has found that<br />

most spontaneously hybridise somewhere in their range, although rates vary<br />

considerably (Ellstrand 2003a). Oilseed rape can hybridise with close relatives<br />

(reviewed in Hails and Morley 2005), principally wild turnip, Brassica rapa,<br />

with national estimates of 32,000±26,000 hybrids being formed annually in<br />

the UK (Wilkinson et al. 2003). Although this represents a hybridisation rate<br />

of only 0.019 (i.e. 19 hybrids for every 1,000 B. rapa), the sheer number of

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