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Genetically Modified Organisms as Invasive Species? 301<br />

In the first group, crop–wild relative hybrids containing the transgene were<br />

grown under experimental conditions often close to the conditions found in<br />

cultivation and, most particularly, with the experimental plants being<br />

released from plant competition. The densities of those natural enemies<br />

which were transgene targets were then manipulated, the results illustrating<br />

that, under appropriate conditions, those hybrids containing the transgenes<br />

have greater fitness. For example, F1 hybrids between B. napus and B. rapa<br />

containing Bt transgenes were found to have a fecundity advantage under<br />

high insect pressure (Vacher et al. 2004). These studies do little more than<br />

illustrate that the transgene will behave as expected in the hybrid as well as<br />

the crop plant, conferring a selective advantage when those plant populations<br />

are affected by the natural enemy (herbivore or pathogen) targeted by the<br />

transgene.<br />

In the second group, far fewer studies have addressed the same questions<br />

under natural field conditions, without manipulating natural enemy pressure.<br />

One exception to this involved a Bt gene backcrossed into wild sunflower populations.<br />

The transgenic backcrossed line had significantly higher fecundity,<br />

compared to the backcrossed control line (Snow et al. 2003). All else being<br />

equal, this would lead to enhanced fitness of plants carrying this transgene.<br />

The difference between these two classes of study is crucial in determining<br />

the frequency of those conditions under which the transgene would be<br />

expected to confer a selective advantage and, therefore, the rate at which it<br />

would spread. The difficulties associated with estimating the relative fitness of<br />

transgenic hybrids may in part explain why the first class of studies is so<br />

much more common than the second. Herbivores and pathogens occur sporadically<br />

in space and time in natural communities: over many years and at<br />

numerous sites, pathogen-resistance genes may provide little advantage until<br />

that one year when a new, virulent pathogen sweeps through an area. Thus,<br />

experiments conducted over a limited number of years and sites run the risk<br />

of being unable to detect any fitness differences. The temptation is then to<br />

manipulate natural enemy pressure to demonstrate the obvious – as long as<br />

some significant result is obtained. The key question is ‘what role do natural<br />

enemies play in regulating existing plant populations?’ So, the most informative<br />

studies will not necessarily involve transgenic plants at all but, rather,<br />

underpinning ecological processes in natural communities.<br />

Another parameter in determining the relative selective advantage of<br />

transgenic plants is the cost of carrying the transgene in the absence of the<br />

target. Costs would contribute to the rate at which transgene frequency may<br />

decline when the population is not under that specific selection pressure.<br />

Again, studies of natural herbivore and pathogen resistance in non-transgenic<br />

plants can be highly informative. A recent review revealed that, if<br />

genetic background is controlled, then 82 % of studies demonstrated fitness<br />

costs associated with carrying herbivore-resistance genes (Strauss et al. 2002),<br />

either as direct costs, such as a trade-off in resources allocated to defence or

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