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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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enzyme occurs in several different allelic forms that differ by single amino acids. Thus, both forms (A and B) will dehydrogenate glucose-6-phosphate both are fully functional enzymes but because they differ by an amino acid, they can be distinguished by their rate of migration in an electrical eld (one form moves faster than another). This electrical separation, termed electrophoresis, is carried out by placing samples of the enzymes in a supporting gel, usually starch, polyacrylamide, agarose, or cellulose acetate ( g. 5.8 and box 5.2). After the electric current is applied for several hours, the enzymes move in the gel as bands, revealing the distance each enzyme traveled. Since blood serum is a conglomerate of proteins from many cells, the serum of a female heterozygote ( g. 5.8, lane 3) has both A and B forms ( bands), whereas any single cell (lanes 4 10) has only one or the other. Since the gene for glucose-6-phosphate dehydrogenase is carried on the X chromosome, this electrophoretic display indicates that only one X is active in any particular cell. Another aspect of the glucose-6-phosphate dehydrogenase system provides further proof of the Lyon hypothesis. If a cell has both alleles functioning, both A and B proteins should be present. Since the functioning glucose-6-phosphate dehydrogenase enzyme is a dimer (made up of two protein subunits), 50% of the enzymes should be heterodimers (AB). These would form a third, intermediate band between the A form (AA dimer) and the B form (BB dimer; g. 5.9). The lack of heterodimers in the blood of heterozygotes is further proof that both G-6-PD alleles are not active within the same cells. That is, in any one cell, only AA or BB dimers can form, because no single cell has both the A and B forms.
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The Lyon hypothesis has been demonstrated with many X-linked loci, but the most striking examples are those for color phenotypes in some mammals. For example, the tortoiseshell pattern of cats is due to the inactivation of X chromosomes ( g. 5.10). Tortoiseshell cats are normally females heterozygous for the yellow and black alleles of the X-linked color locus. They exhibit patches of these two colors, indicating that at a certain stage in development, one or the other of the X chromosomes was inactivated and all of the ensuing daughter cells in that line kept the same X chromosome inactive. The result is patches of coat color. The X chromosome is inactivated starting at a point called the X inactivation center (XIC). That region contains a gene called XIST (for X inactive-speci c transcripts, referring to the transcriptional activity of this gene in the inactivated X chromosome). The XIST gene has been putatively identi ed as the gene that initiates the inactivation of the X chromosome. This gene is known to be active only in the inactive X chromosome in a normal XX female. Another aspect of Lyonization is that several other loci are known to be active on the inactivated X chromosome; they are active in both X chromosomes, even though one is heterochromatic (inactivated). Although several of these loci are in the pseudoautosomal region of the short arm of the X chromosome, several other of the thirty or more genes known to be active are on other places on the mammalian X chromosome. Active genes on the inactive X include the gene for the enzyme steroid sulphatase; the red-cell antigen Xga; MIC2; a ZFY-like gene termed ZFX; the gene for Kallmann syndrome; and several others.
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A form
AA homodimer AB heterodimer
B form
BB homodimer
Electrophoretic gel stained for glucose-6-phosphate dehydrogenase. Lanes 1 3 contain blood from an AA homozygote, a BB homozygote, and an AB heterozygote, respectively. Lanes 4 10 contain homogenates of individual cells of an AB heterozygote.
Electrophoretic gel stained for glucose-6-phosphate dehydrogenase. Lanes 1 and 2 contain blood serum from AA and BB women, respectively, and lane 3 contains serum from an AB heterozygote. Lane 4 shows the pattern expected if both the A and B alleles were active within the same cell.
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