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Deployment of Insect-Resistant Transgenic Crops for Pest Management 329<br />

Furner, 1992; Assaad, Tucker, and Signer, 1993). Homology-induced silencing plays a<br />

role in regulation of endogenous genes whereby the endogenous repeats are frequently<br />

hypermethylated and packaged into transcriptionally inactive heterochromatin. These<br />

inactivations are also trans acting (Mayer, Heidmann, and Niedenhof, 1993). Common<br />

promoter sequences between two genes occupying nonallelic locations may result in<br />

trans-inactivation of the primary transgene (Matzke, Neuhuber, and Matzke, 1993). A mere<br />

90 bp of homology within a 35S promoter fragment between a second inserted gene and<br />

the hypermethylated copies of a primary transgene with 35S promoter sequences result in<br />

silencing of the inserted second gene (Vaucheret, 1993). Homology-based gene silencing<br />

(co-suppression) has been observed to occur only in a fraction of transformants that carry<br />

a sense copy of the endogenous gene. At the molecular level, co-suppression involves transcriptional<br />

inhibition (Ingelbrecht et al., 1994) and posttranscriptional RNA degradation,<br />

while transcription is unaffected (Mueller et al., 1995). Even single copy transgenes get<br />

inactivated due to DNA methylation (Antequera, Boyes, and Bird, 1990). Condensation of<br />

chromatin structures is mediated by proteins that specifi cally recognize methylated DNA<br />

(Lewis et al., 1992). The methylation pattern of the integration region also has a signifi cant<br />

infl uence on the methylation state of the transgene (Prols and Meyer, 1992).<br />

Performance Limitations<br />

The Bt toxins cannot produce the same dramatic effects on insect mortality as the<br />

synthetic insecticides. The farmers need to be educated about the effi cacy and mode of<br />

action of transgenic crops. The expectations have to be real, and remedial measures should<br />

be taken as the situation warrants. The effects of the transgenic crops on insects will be<br />

relatively slower, but cumulative over time. Transgenic crops may not be able to withstand<br />

the pest density in some seasons. Therefore, careful monitoring of pest populations is an<br />

essential component of pest management involving the transgenic crops. The value of the<br />

transgenic crops can be best realized when deployed as a component of pest management<br />

for sustainable crop production (Sharma and Ortiz, 2000). However, enough information<br />

has not been generated involving transgenic crops in a genuine IPM system to demonstrate<br />

long-term benefi ts of the transgenic crops, especially if environmental and human<br />

health hazards are taken into account. Currently deployed transgenic crops produce only<br />

one Bt toxin protein, while the Bt strains used for commercial formulations produce<br />

several toxins in addition to other factors that increase insect mortality. The current cryIAb<br />

construct employs PEP-carboxylase promoter, which enables expression in green tissue<br />

and, as a result, the expression is greater in young plants. Insects that migrate into the<br />

plant whorl or stem tissue with incomplete chlorophyll formation may escape the toxin<br />

protein. If the toxin is expressed in insuffi cient amounts in such tissues, the insects can<br />

develop mechanisms to withstand low levels of toxins in the transgenic plants. Behavioral<br />

avoidance of the tissue expressing the toxin gene can be another component in insect<br />

resistance to the transgenic plants. Therefore, care should be taken to express the toxins in<br />

suffi cient amounts at the site of damage or feeding by the insects.<br />

Secondary Pest Problems<br />

Most crops are attacked by a large number of insect pests. In the absence of competition<br />

from the major insect pests, the secondary pests tend to assume a major pest status (Hilder<br />

and Boulter, 1999). The Bt toxins may be ineffective against some insect pests, for example,<br />

leaf hoppers, mirid bugs, root feeders, mites, etc. This will offset some of the advantages

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