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306<br />

R. Hails and T. Timms-Wilson<br />

of offspring will cause local extinction of populations (Muir and Howard<br />

1999). Thus, a transgene invasion could have the ultimately undesirable outcome<br />

of population extinction. The mating advantage of transgenics is based<br />

on the idea that mature transgenic fish would be larger than non-transgenics.<br />

However, faster growth does not necessarily lead to larger size at maturity,<br />

and the genotype-by-environment interactions discussed above should also<br />

be taken into account. If food abundance is low, then the lower survival of the<br />

risk-prone transgenic juveniles may prevent transgene spread in the population.Whether<br />

the Trojan gene is a real threat remains an open question, albeit<br />

a theoretical possibility (Reichhardt 2000).<br />

17.6 Conclusions<br />

Much of the current disquiet about the creation of GMOs is that their introduction<br />

into the natural environment could be irreversible, causing a perturbation<br />

to the ecological community which will propagate and possibly grow<br />

in magnitude as it ripples through the ecosystem. The pathways by which<br />

transgenes could invade ecosystems have been considered for microbes,<br />

plants and fish (Figs. 17.1 to 17.3). In bacteria, if genes are introduced into<br />

chromosomes, then horizontal gene flow will be low (hard to detect in ecological<br />

time) but dispersal of bacteria per se is very high. The traditional view is<br />

that the dispersal of bacteria is unhindered by geographic boundaries – the<br />

environment selects (Finlay 2002; Whitfield 2005). This view of ecological<br />

panmixis is peculiar to the microbiological world, and from this arise two<br />

possible consequences. Firstly, the phenotype of the transgenic bacteria is all<br />

important – it will determine the conditions under which selection will occur.<br />

Secondly, panmixis leads to the functional redundancy discussed above. So,<br />

perhaps counter-intuitively, greater mixing means less impact of altered phenotypes<br />

on ecosystem services.<br />

Studies of natural systems can be used to illustrate the same principles of<br />

horizontal gene flow and selection which we have been discussing with<br />

GMMs, the introduction of Lotus corniculatus to New Zealand being one<br />

example.At first sowing, the seeds needed to be inoculated with a natural isolate,<br />

Mesorhizobium loti, a nitrogen-fixing symbiont which passes ammonium<br />

to the plant, in exchange for nutrients. Subsequent inoculation was not necessary.<br />

However, this did not correlate with the establishment of the original<br />

inoculated bacterium, as this strain could no longer be detected in the field.<br />

The symbiotic genes carried on a conjugative element had been acquired by<br />

indigenous Mesorhizobium species, better adapted to the prevailing soil conditions<br />

but now able to maximise use of a new niche – the roots of L. corniculatus<br />

(Sullivan and Ronson 1998). This dataset serves two purposes: it demonstrates<br />

the resilience of indigenous bacteria to invasion by an alien strain but

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