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156 Biotechnological Approaches for Pest Management and Ecological Sustainability<br />

may be valuable tools for introgression of genes of interest from the wild relatives into the<br />

cultigen. The F 6-8 progenies of crosses involving a resistance source from the wild relatives<br />

and the cultivated types can also be used as RILs for mapping insect resistance (provided<br />

the population has been advanced through the generations in a correct manner).<br />

Linkage between a resistance gene and a linked marker may vary greatly. The two may<br />

be completely linked, where no crossing over occurs between the resistance gene and the<br />

marker during meiosis, and the gene and marker are always transferred together from one<br />

generation to another. The resistance gene and a molecular marker at times are incompletely<br />

linked, and crossing over may occur between the gene and the marker during<br />

meiosis, thus breaking the linkage between the marker allele and the resistance allele of<br />

the parents. Estimates of the recombination between the resistance gene and the linked<br />

marker are measured as the recombination frequency (RF), which is measured among<br />

backcross progenies of segregating F 2s, F 2:3 families, or RILs by matching the phenotype<br />

and genotype of each progeny (Lander et al., 1987). Mapping simultaneously estimates all<br />

recombination frequencies for markers segregating as dominant, recessive, and codominant<br />

traits in the mapping population. The linkage between QTL and marker loci is based<br />

on the distribution patterns for the resistance characters linked to insect resistance genes<br />

and the molecular marker at each locus (Lincoln, Daly, and Lander, 1993).<br />

To estimate recombination frequency between a resistance gene and a molecular marker,<br />

researchers often analyze between 100 and 500 F 2:3 progenies or RILs derived from crosses<br />

of parents with known resistance or susceptibility. DNA is collected from the resistant and<br />

susceptible parents, as well as from each F 2 plant or several plants in F 6 RILs. Different DNA<br />

markers from a variety of chromosome locations are screened to identify those detecting<br />

polymorphisms between the parents. If the parent polymorphisms are apparent in the<br />

bulked segregant DNA samples, the marker is referred to as a putatively linked marker,<br />

and DNA of all F 2 plants or RILs families is also evaluated. At this point, QTL mapping<br />

software packages such as Mapmaker, Joinmap, QTL cartographer, etc., are used to correlate<br />

the phenotype and genotype of the plants in a mapping population to develop the<br />

genetic linkage map. Linkage mapping software packages contain mapping functions such<br />

as the Haldane’s (1919) and Kosambi’s (1944) functions to correct any under-estimation.<br />

Markers associated with insect resistance can be physico-chemical factors of the plants<br />

(plant hairs or trichomes, fl ower and seed color, leaf size and shape, thickness of the cell<br />

wall, protein, amino acid, sugar, and fatty acid profi les, secondary metabolites such as<br />

alkaloids, terpenoids, fl avonoids, isozymes, and protease inhibitors) or DNA based [RAPDs<br />

(random-amplifi ed polymorphic DNA), RFLPs (restriction fragment length polymorphisms),<br />

AFLPs (amplifi ed fragment length polymorphic DNA), SCARs (sequence characterized<br />

amplifi ed regions), STS (sequence tagged sites), SSRs (simple sequence repeats),<br />

ALPs (amplicon length polymorphisms), and DArTs (diversity array technology)] (Crouch<br />

et al., 2005).<br />

Physico-chemical Markers Associated with Resistance to Insects<br />

Morphological Markers<br />

Plants have acquired several physicochemical characteristics that contribute to insect resistance<br />

in different crops (Panda and Khush, 1995; Smith, 2005) (Table 6.1). The role of various<br />

physico-chemical components in host plant resistance to insects is discussed below.

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