MATERNAL EFFECTS

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.