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Mechanisms and Inheritance of Resistance to Insect Pests 139<br />

Oilseeds<br />

Resistance in soybean to defoliating insects is polygenic. Heritability estimates for resistance<br />

to the cabbage looper, T. ni (Luedders and Dickerson, 1977) and E. varivestis (Sisson<br />

et al., 1976) suggest quantitative inheritance. The F2 plants from a cross between the parents<br />

resistant and susceptible to the soybean looper, P. includens also exhibit partial dominance<br />

or a quantitative inheritance (Kilen, Hatchett, and Hartwig, 1977). Johnson and<br />

Beard (1977) observed that a phytomelanin (achene) layer conferring resistance to sunfl<br />

ower moth, Homoeosoma electellum (Hulst.), in different Helianthus species is inherited as a<br />

dominant trait.<br />

Alfalfa<br />

A single dominant gene in alfalfa, Medicago sativa L., and sweet clover, Melilotus infesta<br />

Guss., controls resistance to the pea aphid, A. pisum (D.V. Glover and Stanford, 1966), and<br />

the sweet clover aphid, Therioaphis riehmi (Bomer) (Manglitz and Gorz, 1968). Resistance to<br />

spotted alfalfa aphid, Therioaphis maculata (Buckton), is controlled by several genes (E. Glover<br />

and Melton, 1966), indicating that resistance is quantitative. Combining ability effects in<br />

alfalfa for resistance to E. fabae are signifi cant and additive in nature (Soper, Mcintosh, and<br />

Elden, 1984; Elden, Elgin, and Soper, 1986).<br />

Potato<br />

Resistance to the green peach potato aphid, Myzus persicae (Sulzer) in cultivated and wild<br />

Solanum species is expressed as a partially dominant trait (Sams, Lauer, and Radcliffe,<br />

1976). One dominant gene controls glandular trichome-mediated resistance to M. persicae<br />

in Solanum tarijense Hawkes and S. berthaultii Hawkes, but in S. phureja ( Juz et Buk.) S.<br />

berthaultii crosses, two genes control the expression of resistance (R.W. Gibson, 1979).<br />

Resistance to M. persicae is mediated by a complex interaction between trichome density<br />

and droplet size of exudates, and is a quantitatively inherited trait (Mehlenbacher,<br />

Plaisted, and Tingey, 1983, 1984). Resistance to potato tuber moth, P. operculella, has been<br />

derived from Solanum sparsipilum (Bitter) Juz. & Buk., and is controlled by a few major<br />

genes (Ortiz et al., 1990).<br />

Grain Legumes<br />

Resistance in Lima bean, Phaseolus lunatus L., to the leafhopper, Empoasca kraemeri Ross &<br />

Moore, is due to the quantitative effect of several genes, and is inherited as a recessive trait<br />

(Lyman and Cardona, 1982). Both additive and dominant gene effects are responsible for<br />

E. kraemeri resistance in cultivars of the common bean, Phaseolus vulgaris L. (Kornegay and<br />

Temple, 1986). Inheritance of hooked trichomes is complex, and is controlled by additive,<br />

dominant, and epistatic gene effects (Park et al., 1994). There is also evidence for transgressive<br />

segregation (levels of resistance greater than that of the resistant parent) in some<br />

progenies from crosses between resistant and susceptible bean cultivars. Resistance in<br />

wild strains of P. vulgaris to Zabrotes subfasciatus (Boheman) is controlled by the toxic seed<br />

protein, arcelin. The presence of arcelin is inherited as a dominant trait (Romero Andreas,<br />

Yandell, and Bliss, 1986; Kornegay, Cardona, and Posso, 1993). Resistance to bean weevil,<br />

Acanthoscelides obtectus (Say), is derived from a wild Phaseolus accession, and is inherited as<br />

a complementary effect of two recessive genes (Kornegay and Cardona, 1991). Resistance

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