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Development of Resistance to Transgenic Plants 383<br />

of this gene confer resistance in strains from Hawaii, Pennsylvania, and Philippines. The<br />

biphasic nature of the lepidopteran genetic linkage map has been used to detect this gene<br />

in diamondback moth, with 207 amplifi ed fragment length polymorphism (AFLP) DNA<br />

markers. An AFLP marker has been cloned and sequenced for the chromosome containing<br />

the Bt resistance gene. Resistance in O. nubilalis to Bt appears to be inherited as an incompletely<br />

dominant autosomal gene, indicating that if the fi eld resistance turns out to be<br />

dominant, then the high dose/refuge strategy may be ineffective for management of resistance<br />

to this pest.<br />

Strategies for Resistance Management<br />

To preserve the value of Bt endotoxins in crop protection, it will be necessary to implement<br />

resistance management measures from the very beginning. Individual farmers have<br />

limited incentive to adopt resistance management technologies for Bt endotoxins, and the<br />

greatest incentive lies with the Bt industry (Kennedy and Whalon, 1995). However, the<br />

implementation of a coordinated, industry-wide, Bt resistance management effort is likely<br />

to be constrained by competition among different segments of the Bt industry interested<br />

in different technologies (sprays versus transgenic plants), and among producers of Bt<br />

products using the same technology. Successful implementation of resistance management<br />

for Bt endotoxins will require that the Bt industry prepackage resistance management<br />

techno logies, and that these prepackaged resistance management strategies do not<br />

add signifi cantly to the costs or complexity of pest control by the end user. In view of<br />

criticism of the Bt transgenic crops for their ability to enhance development of resistance,<br />

prevention of development of resistance to Bt is the best policy. The Bt resistance management<br />

tactics have been incorporated in the regulation of Bt transgenic crops in most countries<br />

(Haliday, 2001). Transgenic crops must produce high doses of toxin and there should<br />

be nontransgenic crops or other alternative host crops as refuge for the production of<br />

susceptible insect populations.<br />

To increase the effectiveness of the transgenic plants, it is important to implement the<br />

resistance management strategies from the very beginning. A number of conceptual strategies<br />

have been developed for resistance management (McGaughey and Whalon, 1992;<br />

Tabashnik, 1994; Gould, 1998). A high level of migration or a sensible reduction of the fi tness<br />

associated with the change in the genome may lead to a long-lasting effi cacy of the transgenic<br />

crops (Arpaia, Chiriatti, and Giorio, 1998). Most of the strategies for resistance management<br />

are based on mixtures of toxins to be deployed for insect control, tissue-specifi c<br />

production, and induced toxin production. Two or more insecticidal proteins or different<br />

genes can also be introduced into the same plant to slow down the rate of development of<br />

resistance in the target insects. Plants can also be engineered so that the toxin is produced<br />

only in the tissues where the insect feeds. In induced toxin production, the plants can be<br />

engineered in such a way that the toxin is produced when the insect starts feeding.<br />

The strategies for resistance management would depend on the number and nature of<br />

gene action, insect behavior, and insect-genotype-environment interaction. Spring movement<br />

of emerging adults onto wild hosts may delay the development of resistance if the<br />

movement of adults is far enough from the fi eld in which the pupae over-wintered. Increase<br />

in the summer migration and the distance moved will also delay resistance development.<br />

It has been suggested that Bt sunfl ower might not lead to the development of Bt-resistant

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