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Transgenic Resistance to Insects: Nature of Risk and Risk Management 433<br />

Assessment of Risk to Agriculture<br />

The importance of assessing the invasiveness of genetically modifi ed crops has been the<br />

subject of much debate (Fitter, Perrins, and Williamson, 1990). While it is important to be<br />

cautious when predicting weediness based on plant attributes, an assessment of such<br />

characteristics can be made with much greater confi dence when assessing whether wellestablished<br />

crops can invade agricultural or natural ecosystems. Modern crop cultivars no<br />

longer possess the ability to become weeds, as this ability has been severely reduced in the<br />

absence of gene introgression from wild relatives. Common distinctive attributes of weeds,<br />

such as seed dormancy, phenotypic plasticity, indeterminate growth, continuous fl owering<br />

and seed production, and seed dispersal have been bred out of the crop plants over<br />

thousands of generations (Baker, 1965). Such changes appeared early in the domestication<br />

of crop plants as a consequence of repeated sowing and harvesting cycles (Harlan, 1992).<br />

These characters are not candidates for gene transfer back into crops, whether by genetic<br />

modifi cation or traditional breeding, because they would severely reduce the agronomic<br />

performance of a crop for modern farming practices. Furthermore, these attributes do not<br />

arise from a single or few gene transfers. Therefore, genetically modifi ed crops are no<br />

more likely to become weeds outside farming situations than the crop cultivars developed<br />

through traditional breeding in the past.<br />

Selection for Weedy Traits<br />

Weeds generally exhibit a preference for habitats disturbed by human activities, such as<br />

cultivated fi elds, fi eld margins, gardens, roadsides, soil dumps, and waste sites (Harlan<br />

and de Wet, 1965). One feature that all weeds tend to share is high phenotypic plasticity<br />

that allows continuous adaptation to changing environments. It is suspected that genetic<br />

modifi cation for adaptation to drought or heat stress may change some of these characteristics<br />

of the wild and weedy relatives of crops through gene fl ow. Closely related species<br />

can sometimes be highly successful as weeds as they possess different combinations of<br />

weedy characteristics (Williamson, 1993). Whether a genetically modifi ed plant species<br />

becomes a serious weed in a new environment may relate more to its ability to grow in the<br />

new environment coupled with the absence of effective natural enemies such as insect<br />

herbivores and diseases (Williamson, 1994). There has been a close association between<br />

the evolution of weeds and the domestication of crop plants (de Wet and Harlan, 1975;<br />

Harlan, 1992), and some weeds mimic crop characteristics such as growth form and<br />

maturity cycle (Barrett, 1983). Selection away from cultivated attributes and gene introgression<br />

between genetically modifi ed crops and wild relatives may lead to evolution of<br />

weeds with better adaptation to the environment, and this aspect needs to be carefully<br />

assessed for each transgene and crop combination.<br />

Invasiveness<br />

An appropriate measure of invasiveness of a plant is its fi nite rate of increase (Crawley,<br />

1986), the constant at which a population increases over time, assuming a stable age distribution,<br />

and the absence of density-dependent constraints. Risk assessment encompasses<br />

all demographic processes regulating population growth of a species, such as rate of<br />

growth and maturity, fecundity, seed survival, seed germination, and seedling survival to

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