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Transgenic Resistance to Insects: Gene Flow 419<br />

for development of resistance in the pest populations also need to be assessed, and strategies<br />

devised to use different genes in different crops in the same environment. Transgenic<br />

crops containing protease inhibitors may pose similar problems, and their use needs to be<br />

carefully planned to avoid the evolution of pest populations capable of withstanding<br />

the transgene. Resistance to abiotic stress factors may present additional challenges, as<br />

this would enable the plants to grow in environments where they were unable to do well<br />

earlier (Fraley, 1992). This confers additional advantage to the transgenic plant, and there<br />

are chances for gene transfer through cross-pollination. The exotic species model can be<br />

used to assess the risk of introducing transgenic plants with resistance to abiotic stress<br />

factors. The risk assessment in such cases requires more information, and should take<br />

into account the nature of competitive advantage conferred by the transgene under<br />

specifi c conditions.<br />

Gene Flow and Enhanced Fitness of Herbivores<br />

Widespread cultivation of transgenic crops with resistance to insect pests and diseases<br />

will impose intense selection pressure on pest populations to adapt to the transgene.<br />

Development of insect pest and disease-resistant cultivars has been one of the primary<br />

objectives of plant breeding for many years (Simmonds et al., 1999). The history of plant<br />

breeding has clearly established that insect pest and pathogen populations can quickly<br />

adapt to crop cultivars with new resistance genes (Bonman, Khush, and Nelson, 1992;<br />

McIntosh and Brown, 1997). However, crop improvement is an ongoing process, and plant<br />

breeders have not stopped breeding for insect pest and disease resistance simply because<br />

the target pest or disease might overcome the resistance. Development of management<br />

strategies to minimize the evolution of insect and pathogen populations that might overcome<br />

the resistance genes has been ongoing for many years. In order to establish an<br />

appropriate resistance management plan, it is important to understand the nature of interactions<br />

between the host crop, the insect pest/pathogen population(s), and the mechanisms<br />

involved. The key to maintaining an effective management plan is regular monitoring of<br />

the response of the insect pest/pathogen populations to the cultivars grown by the farmers.<br />

Although many resistance genes have been identifi ed in crop germplasm, there is no<br />

easy way to predict the quality or durability of these resistance genes. The “breakdown”<br />

of resistance is usually associated with qualitative resistance conferred by major genes<br />

(R genes), where resistance versus susceptibility results from a gene-for-gene interaction<br />

between the R genes in the host and avirulence genes in the pathogen (Flor, 1971). The<br />

resistance conferred by many R genes has not been durable as a consequence of rapid<br />

changes in pathogen populations (Leach et al., 2001). The most widely cited examples of<br />

durable resistance against bacterial or fungal pathogens involve multigenic quantitative<br />

traits (Johnson, 1984; Parlevliet, 2002). However, there are examples where single R genes<br />

have conferred highly durable resistance, for example, the Lr34 gene conferring resistance<br />

to leaf rust in wheat (Kolmer, 1996) and the Xa4 gene conferring bacterial blight resistance<br />

in rice (Bonman, Khush, and Nelson, 1992). The experience gained from plant breeding<br />

over many years will help defi ne the appropriate management approaches for insect pest-<br />

and disease-resistant transgenic crops to prevent or minimize the establishment of insect<br />

pests and pathogen populations that may overcome the resistance mechanism underpinning<br />

the resistance genes. It is not the transgenic versus nontransgenic status of the<br />

crop plants that may result in projected problems, but the way the crops are grown and<br />

managed will have a greater infl uence. The widespread cultivation of genetically modifi ed<br />

crops with insect or disease resistance is no more likely to result in the development of

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