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Tamarin: Principles of Genetics, Seventh Edition
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II. Mendelism and the Chromosomal Theory
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5. Sex Determination, Sex Linkage, and Pedigree Analysis
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e ended chapter 3 with a discussion of the chromosomal theory of heredity, stated lucidly in 1903 by Walter Sutton, that genes are located on chromosomes. In 1910, T. H. Morgan, a 1933 Nobel laureate, published a paper on the inheritance of white eyes in fruit ies. The mode of inheritance for this trait, discussed later in this chapter, led inevitably to the conclusion that the locus for this gene is on a chromosome that determines the sex of the ies: when a white-eyed male was mated with a red-eyed female, half of the F2 sons were white-eyed and half were red-eyed; all F2 daughters were red-eyed. Not only was this the rst evidence that localized a particular gene to a particular chromosome, but this study also laid the foundation for our understanding of the genetic control of sex determination.
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sex chromosome, as in some grasshoppers and beetles; females are usually XX and males X0. And in the compound chromosome case, several X and Y chromosomes combine to determine sex, as in bedbugs and some beetles. We need to emphasize that the chromosomes themselves do not determine sex, but the genes they carry do. In general, the genotype determines the type of gonad, which then determines the phenotype of the organism through male or female hormonal production.
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The XY System
The XY situation occurs in human beings, in which females have forty-six chromosomes arranged in twentythree homologous, homomorphic pairs. Males, with the same number of chromosomes, have twenty-two homomorphic pairs and one heteromorphic pair, the XY pair ( g. 5.1). During meiosis, females produce gametes that contain only the X chromosome, whereas males produce two kinds of gametes, X- and Y-bearing ( g. 5.2). For this reason, females are referred to as homogametic and males as heterogametic. As you can see from gure 5.2, in people, fertilization has an equal chance of producing either male or female offspring. In Drosophila, the system is the same, but the Y chromosome is almost 20% larger than the X chromosome ( g. 5.3). Since both human and Drosophila females normally have two X chromosomes, and males have an X and a Y chromosome, it seems impossible to know whether maleness is determined by the presence of a Y chromosome or the absence of a second X chromosome. One way to resolve this problem would be to isolate individuals with odd numbers of chromosomes. In chapter 8, we examine the causes and outcomes of anomalous chromosome numbers. Here, we consider two facts from that chapter. First, in rare instances, individuals form, although they are not necessarily viable, with extra sets of chromosomes. These individuals are referred to as polyploids (triploids with 3n, tetraploids with 4n, etc.). Second, also infrequently, individuals form that have more or fewer than the normal number of any one chromosome. These aneuploids usually come about when a pair of chromosomes fails to separate properly during meiosis, an occurrence called nondisjunction. The existence of polyploid and aneuploid individuals makes it possible to test whether the Y chromosome is male determining. For example, a person or a fruit y that has all the proper nonsex chromosomes, or autosomes (forty-four in human beings, six in Drosophila), but only a single X without a Y would answer our question. If the Y were absolutely male determining, then this X0 individual should be female. However, if the sex-determining mechanism is a result of the number of X chromosomes, this individual should be a male. As it turns out, an X0 individual is a Drosophila male and a human female.
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