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Transgenic Resistance to Insects: Gene Flow 411<br />

particular fi tness components of the weed (Ellstrand, Prentice, and Hancock, 1999).<br />

However, plant breeders have released many cultivars with new genes for resistance to<br />

insect pests, diseases, and environmental stress over many years. Any impacts resulting<br />

from the introgression of such traits into weedy species are equally likely for the products<br />

of plant breeding and genetic modifi cation. The risks are no different and the use of resistance<br />

genes in cultivars from traditional breeding has not enhanced the survival and<br />

spread of weeds in the past. When serious weeds have arisen following hybridization of<br />

crops and wild species, their aggressive nature has arisen from a coupling of morphological<br />

traits conferring weedy attributes and the synchrony of development with the crop<br />

rather than a gain in fi tness from resistance genes (de Wet and Harlan, 1975).<br />

Another problem highlighted by natural hybridization between crops and their wild<br />

relatives is the increased potential of extinction of wild taxa. Some “genetically aggressive”<br />

species, referred to as compilospecies (Harlan and de Wet, 1963), may completely<br />

assimilate another locally rare species through repeated cycles of hybridization and introgression,<br />

causing it to become extinct. The highly successful wheat crop is considered to<br />

have assimilated genes from more than one species of Aegilops (Harlan and de Wet, 1963).<br />

Extinction by hybridization does not depend on relative fi tness, but on patterns of mating<br />

(Ellstrand, Prentice, and Hancock, 1999). The impact of the release of transgenic crops will<br />

be no different than the impact of existing nontransgenic crops. Interspecifi c hybridization<br />

is a common process, but hybrids are rare, and most are sterile, and there is a very low<br />

chance of gene introgression into the wild relatives (Fitter, Perrins, and Williamson, 1990).<br />

Transgenic plants may also become weeds, except in the context of their normal agricultural<br />

environment. Gene escape may occur when a plant species invades a seminatural<br />

habitat or the gene is transferred into the wild relative, and persists in uncultivated land.<br />

Its spread can be checked by methods similar to any other single trait. There are differences<br />

among plant species to disperse from the environment other than the one in which<br />

they are released, and their ability to establish feral populations. Such an event has to be<br />

compared with that of the original plant.<br />

The process of introgression between a transgenic crop modifi ed for better agronomic<br />

characters and a wild relative could potentially lead to increased weediness and adaptation<br />

to the environment (Gueritaine et al., 2002). However, formation of a hybrid and hybrid<br />

progeny could be associated with functional imbalance and low fi tness, which reduces the<br />

risk of gene escape and establishment of the wild species in the fi eld. Inter- and intraspecifi<br />

c gene fl ow between transgenic and nontransgenic plants leads to development of<br />

new genetic materials in cultivated or wild populations. Introgression of novel genes into<br />

wild relatives of crops may have a substantial impact on crop evolution or wild populations<br />

leading either to more desirable cultivars, more aggressive weeds, and/or extinction<br />

of rare and endangered species. A better understanding of gene fl ow is therefore essential<br />

for deployment of transgenic plants. Such information would be helpful in assessing the<br />

ecological risk of transgene escape from cultivated germplasm to closely related wild<br />

relatives. The consequences of gene fl ow from crop to wild relatives include genetic assimilation,<br />

wherein alien genes replace host genes, and demographic swamping, wherein<br />

hybrids are less fertile than their wild parents and as a result the wild populations shrink<br />

(Haygood, Ives, and Andow, 2003). Factors infl uencing gene fl ow include mating system,<br />

mode of pollination, mode of seed dispersal, and the habitat characteristics where the<br />

crops are grown (Messeguer, 2003). Hybridization is perhaps the most serious genetic<br />

threat to endangered species, with extinction often taking place in less than fi ve generations<br />

(Wolf, Takebayashi, and Rieseberg, 2000). Pollen can function as a vehicle to disseminate<br />

introduced, genetically engineered genes throughout a plant population or into a

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