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Snail Coiling
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Snails are coiled either to the right (dextrally) or to the left (sinistrally) as determined by holding the snail with the apex up and looking at the opening. The snail is dextrally coiled if the opening comes from the right-hand side and sinistrally coiled if it comes from the left-hand side ( g. 17.1).The inheritance pattern of the coiling is at rst perplexing.
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Figure 17.1 Inheritance of coiling in the pond snail, Limnaea peregra. Reciprocal crosses (D, dominant dextral; d, recessive sinistral coiling) are shown (DD mated with dd in each case). The F1 individuals in both crosses have the Dd genotype but re ect the mother s genotype in respect to coiling; DD mothers produce dextrally coiled offspring, whereas dd mothers produce sinistrally coiled offspring. The F2 individuals in both cases are identical because the genotypes of the F1 mothers are identical (Dd ). The coiling of a snail s shell is determined by its mother s genotype, not phenotype.
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In the left half of gure 17.1, a dextral snail provides the eggs, and a sinistral snail provides the sperm. The offspring are all dextral; presumably, therefore, dextral coiling is dominant. When the F1 are self-fertilized (snails are hermaphroditic), all the offspring are dextrally coiled. The result is unexpected. Nevertheless, when the F2 are self-fertilized, one-fourth produce only sinistral offspring, and three-fourths produce only dextral offspring. If selffertilization is continued through ensuing generations, this 3:1 phenotypic ratio will be revealed as a Mendelian 1:2:1 genotypic ratio, thereby reaf rming the notion of a single locus with two alleles, and dextral dominant. However, something interfered with the expected phenotypic pattern. When the reciprocal cross is made ( g. 17.1, right), the F1 have the same genotype as just described but are coiled sinistrally, as is the female parent. From here on, the results are exactly the same for both crosses. In both cases, the F1 are phenotypically similar to the female parent even though the offspring in both crosses have the same genotype (Dd). The explanation is that the genotype of the maternal parent determines the phenotype of the offspring, with dextral dominant. Thus, the DD mother in gure 17.1 produces F1 progeny that are dextral with a Dd genotype, and the dd mother produces progeny that are sinistral with the same Dd genotype. Why does this pattern occur
A process of spiral cleavage takes place in the zygotes of mollusks and some other invertebrates.The spindle at mitosis is tipped in relation to the axis of the egg. If the spindle is tipped one way, a snail will be coiled sinistrally; if it is tipped the other way, the snail will be coiled dextrally. The direction of tipping is determined by the maternal cytoplasm, which is under the control of the maternal genotype. Obviously, maternal control affects only one generation in each generation, the coiling is dependent on the maternal genotype.
Moth Pigmentation
There are other examples of maternal effects in which the cytoplasm of the mother, under the control of chromosomal genes, controls the phenotype of her offspring. In the our moth, Ephestia k hniella, kynurenin, which is a precursor for pigment, accumulates in the eggs. The recessive allele, a, when homozygous, results in a lack of kynurenin. Reciprocal crosses give different results for larvae and adults. When a nonpigmented female is crossed with a pigmented male, the results are strictly Mendelian; but when the mother is pigmented (a a), all the larvae are pigmented regardless of genotype ( g. 17.2).The initial larval pigmentation comes from residual kynurenin in the eggs, which is then diluted out so that an adult s pigmentation conforms to its own genotype.
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