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Physico-Chemical and Molecular Markers for Resistance to Insect Pests 185<br />

against the target insect pest(s). In addition, crops frequently suffer from a number of<br />

primary herbivores. This suggests that single and multiple transgenes will need to be<br />

combined in the same variety with other sources, mechanisms, and targets of insect pest<br />

resistance in order to generate highly effective and sustainable seed-based technologies.<br />

From an evolutionary point of view, the development of plants with several genes for<br />

insect resistance will be expected to decrease the probability of insect pests to overcome<br />

newly deployed seed-based resistance technologies and thereby increase the effectiveness<br />

and prolong the life of such new varieties (Hadi, McMullen, and Finer, 1996; Karim,<br />

Riazuddin, and Dean, 1999), for example, the activity of Bt genes in transgenic plants is<br />

enhanced by the serine protease inhibitors (MacIntosh et al., 1990; Zhao et al., 1997) and<br />

tannic acid (Gibson et al., 1995). However, this may have some metabolic cost to the plant<br />

in some cases, and different resistance gene products may also have deleterious or<br />

nullifying interactions.<br />

Combining transgene- and QTL-mediated resistance can be used as a viable strategy for<br />

insect control. A QTL conditioning maize earworm resistance in soybean PI 229358 and<br />

the cry1Ac transgene from the recurrent parent Jack-Bt has been pyramided into BC 2F 3<br />

plants by MAS (Walker et al., 2002). Segregating individuals were genotyped through SSR<br />

markers linked to an antibiosis/antixenosis QTL on LG M, and tested for the presence of<br />

cry1Ac. MAS was used during and after the two backcrosses to develop a series of BC 2F 3<br />

plants with or without the cry1Ac transgene and the QTL conditioning insect resistance.<br />

The BC 2F 3 plants homozygous for parental alleles at markers on LG M, and which either<br />

had or lacked Cry1Ac, were assigned to one of four possible genotype classes. These plants<br />

were used in no-choice detached leaf feeding bioassays with corn earworm, H. zea and<br />

soybean looper, P. includens. Fewer larvae of either species survived on leaves expressing<br />

the Cry1Ac protein. Though not as great as the effect of Cry1Ac, the PI 229358-derived LG<br />

M QTL also had a detrimental effect on larval weights of both species, and on defoliation<br />

by maize earworm, but did not reduce defoliation by the soybean looper. Weights of soybean<br />

looper larvae fed on foliage from transgenic plants with the PI-derived QTL were<br />

lower than those of larvae fed transgenic tissue with the corresponding Jack chromosomal<br />

segment (Walker et al., 2002). Therefore, combining transgene- and QTL-mediated resistance<br />

to lepidopteran insects may be a viable strategy for insect control.<br />

Marker-Assisted versus Phenotypic Selection<br />

Expression of physicochemical markers is infl uenced by the environment, and this makes<br />

them less reliable than molecular markers. A number of methods have been used for mapping<br />

QTLs associated with the traits of interest (Knot and Hailey, 1992; Karp et al., 1997).<br />

Among the molecular markers, RFLPs are the most reliable (Caetano-Anolles, Bassam, and<br />

Gresshoff, 1991; J.J. Lin et al., 1996), and can be converted into SCARs by sequencing the<br />

two ends of the DNA clones and designing oligonucleotide primers based on end sequences<br />

(Williams et al., 1991). Molecular markers and QTLs associated with insect resistance can<br />

be used to transfer the resistance genes from sources of resistance into high-yielding cultivars.<br />

It is important that the marker cosegregates with the gene, and is closely linked<br />

(1 cM or less) with the trait of interest. There are several advantages of using molecular<br />

markers to breed for insect resistance. Some types of molecular markers behave in a<br />

codominant manner to detect heterozygotes in segregating populations. Morphological

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