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Genetically Modified Organisms as Invasive Species? 305<br />

of others (Kennedy and Smith 1995). Thus, if impact is measured in terms of<br />

a change in ecosystem function, then no significant impacts have been<br />

detected and, indeed, seem unlikely (Timms-Wilson et al. 2002; Griffiths et al.<br />

2003).<br />

17.5.2 Potential Impacts in Plant Populations<br />

Transgenes which would alter the invasiveness of plants, ultimately causing<br />

the extinction of other species, would be unacceptable. Fitness and invasiveness<br />

are sometimes used interchangeably in the literature, yet this is not<br />

always appropriate. In fact, only when a species is invading a habitat for the<br />

first time is it appropriate to equate fitness and invasiveness – under those circumstances,<br />

population growth rate (fitness) can be used as a measure of how<br />

invasive that genotype is. When considering the invasion of transgenes into<br />

wild populations, enhanced fitness may result in a change in gene frequencies<br />

but not necessarily a change in invasiveness. This latter attribute depends<br />

upon the factors responsible for limiting and regulating the population. For<br />

example, the extent to which the enhanced fecundity of Bt wild sunflowers<br />

(Snow et al. 2003) will enhance invasiveness depends upon the extent to which<br />

wild sunflowers are seed-limited and the herbivores remain susceptible to the<br />

transgene products. Rates of co-evolution may be quite rapid in response to Bt<br />

genes (Shelton et al. 1993; Ferré and van Rie 2002), in which case any increases<br />

in fitness may be transient. If wild sunflower populations are seed-limited and<br />

herbivore populations remain susceptible, then enhanced fitness may translate<br />

into increased abundance. A review of seed sowing experiments aimed at<br />

unravelling the extent to which natural populations are seed-limited (Turnbull<br />

et al. 2000) demonstrated that approximately 50 % of seed augmentation<br />

experiments showed some evidence of seed limitation; this tends to occur in<br />

early successional habitats and with early successional species. In other<br />

words, if succession proceeds, any changes in abundance resulting from<br />

enhanced fecundity may also be transient. The most likely problems to occur<br />

are in disturbed (agricultural) habitats.<br />

17.5.3 Potential Impacts in Animal Populations<br />

Theoretical scenarios have been raised in which the release of transgenic fish<br />

could have very detrimental impacts on wild populations, and one of these is<br />

the Trojan gene hypothesis. This hypothesis illustrates how data on relative<br />

fitness for part of the life cycle could mislead as to the potential long-term<br />

impact of released transgenic salmon. If transgenic escapees have a mating<br />

advantage but viability of offspring is reduced, then models predict that the<br />

transgene will spread through the wild population but that reduced viability

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