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Genetically Modified Organisms as Invasive Species? 299<br />

17.4 Transgene Spread<br />

Gene flow creates the opportunities for transgene spread but the rate at which<br />

this will occur depends upon the relative fitness of transgenic versus wild<br />

type genotypes.<br />

17.4.1 Transgene Spread in Bacterial Populations<br />

There is only one experiment which has directly compared the fitness of<br />

transgenic and non-transgenic bacteria in ‘semi-field’ conditions, when the<br />

transgene has been located on plasmids. The transgene in this case was a<br />

marker. This was conducted in an ecotron (http://www.cpb.bio.ic.ac.uk/<br />

ecotron/ecotron.html), a facility which houses replicated microcosms containing<br />

simplified ecological communities. The aim of the experiment was to<br />

compare the population dynamics of Pseudomonas fluorescens SBW25R (the<br />

control) with the same bacterium carrying a gene cassette which had been<br />

introduced into the genome in three different ways: directly inserted into the<br />

bacterial chromosome (SBW25R::KX), as a similar insertion but including a<br />

lysogenic phage (SBW25::KX-F101), and inserted into a conjugative plasmid<br />

(SBW25R pQBR11::KX). Following seed dressing, all bacterial strains successfully<br />

colonized the phytosphere of chickweed (Stellaria media) and<br />

became established. However, densities of the introduced strain in the lysogenic-phage<br />

treatment were consistently lower than in the other treatments,<br />

at times approaching the limits of detection. The lower density of this strain<br />

was attributed to cell deaths caused by lysis following phage induction,<br />

which was expected in the phytosphere (Ashelford et al. 1999). In the plasmid<br />

treatment, the densities of P. fluorescens SBW25R (pQBR11::KX) on day<br />

53 were significantly lower on roots and leaves than was the case for the<br />

plasmid-free control, with plasmid carriage resulting in a marked reduction<br />

in colonizing fitness. However, from day 53 onwards, in each of the four<br />

chambers the population density of the plasmid-carrying strain increased<br />

significantly on roots (25-fold) and leaves (13-fold) until, by day 95, these<br />

densities either matched or slightly exceeded those of the control. These<br />

results can be explained if carriage of the plasmid were typically associated<br />

with a cost to the host (as has frequently been demonstrated; Lilley and Bailey<br />

1997) but that, periodically, an unidentified plasmid gene (or genes)<br />

improves the fitness of the bacteria, perhaps by conferring the ability to<br />

utilise or tolerate certain substrates exuded by the plants as they mature.<br />

Variation in the fitness of bacteria carrying plasmids is correlated with plant<br />

growth stage in the field.<br />

Because of the rates of horizontal gene flow expected, there has yet to be a<br />

field release of bacteria containing transgenic plasmids. However, there have

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