Eggs and embryos are expected to be influenced by the maternal environment in
which they develop. Even those removed from the body of the mother at an early stage receive cytoplasm and nutrients in the eggs from the mother, and special influences on gene action may have already taken effect. Certain potentialities of the egg are known to be determined before fertilization, and, in some cases, these have been influenced by the surrounding maternal environment. Such predetermination by genes of the mother, rather than those of the progeny, is called maternal effect. Existence of a maternal effect is commonly substantiated or disproved by reciprocal crosses. If a maternal effect is involved, results from reciprocal crosses will be different from each other, with genes of the mother being expressed.
Maternal Effect in Snail Shell Coiling
One of the earliest and best-known examples of a maternal effect is that of the direction of coiling in shells of the snail Limnaea peregra. Some strains of this species have dextral shells, which coil to the right; other have sinistral shells, which coil to the left. This characteristic is determined by the genotype of the mother (not her phenotype) rather than by the genes of the developing snail. Allele s+ for right-handed coiling is dominant over allele s for coiling to the left. When crosses (Fig. 20.13) were made between females coiled to the right and males coiled left, the F1 snails were all coiled to the right. The usual 3:1 ratio was not obtained in the F2 because the phenotype of ss was not expressed. Instead, the pattern determined by the mother`s (P) genes (s+s+) was expressed in the F1, and the F1 mother`s genotype (s+s) was expressed in the F 2. When ss individuals were inbred, only progeny that coiled to the left were produced. When the s+s+ or s+s snails were inbred, however, they produced off-spring that all coiled to the right. From the reciprocal cross between left-coiling females and right-coiling males (Fig. 20.14), all F1 progeny were coiled to the left. The F2 all coiled to the right; but, when each F2 snail was inbred, those with the genotype ss produced progeny that coiled to the left. Further investigation of coiling in snails has shown that the spindle formed in the metaphase of the first cleavage division influences the direction of coiling. The spindle of potential “dextral” snails is tipped to the right, but that of “sinistral” snails is tipped to the left. This difference in the arrangement of the spindle is controlled by the genes of the mother. They determine the orientation of the spindle, which in turn influences further cell division and results in the adult pattern of coiling. The actual phenotypic characteristic, therefore, is influenced directly by the mother, with no immediate relation to the genes in the egg, sperm, or progeny. However, most other snail traits do not show the maternal-effect pattern. The striping color pattern, for example, is also determined in the early embryo, but it is controlled directly by chromosomal genes of both parents. In this example, comparable color patterns are obtained from the results of reciprocal crosses.
Maternal Effect in Drosophila
At the university of Texas abnormal growth in the head region of Drosophila melanogaster appeared sporadically in a sample from a wild population collected at Acahuizotla, Mexico. At the University of Utah, these flies were inbred and selected for the abnormal head growths over a period of several years. The proportion of flies expressing the trait, named “tumorous head” (tu-h; Fig. 20.15), was increased to about 76 percent at 22oC when the flies were raised on a cornmeal and molasses medium. When reciprocal crosses were conducted, a maternal effect was indicated. Tu-h females mated separately with three wild-type males and males from 11 laboratory stocks produced from 14 to 52 percent (average of 30 percent) of abnormal flies in the first generation. From the reciprocal cross between tu-h males and the same three wild-type and the same 11 laboratory stock females, … percent (average of less than 1 percent) of tumor-head flies were obtained. Further studies demonstrated the maternal effect. Genes of the mother were … an influence in the direction of abnormal … on the heads of adult progeny during the first … of development. Two major genes were found to control the tumorous head trait: (1) a sex linked gene at 64,5 map units on the X chromosome controlling the maternal effect and (2) a structural gene at 58 map units on the third chromosome controlling the tumorous head phenotype.