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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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The McGraw Hill Companies, 2001
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lectrophoresis, a technique for separating relatively similar types of molecules (for example, proteins and nucleic acids), has opened up new and exciting areas of research in population, biochemical, and molecular genetics. It has allowed us to see variations in large numbers of loci, previously dif cult or impossible to sample. In biochemical genetics, electrophoretic techniques can be used to study enzyme pathways. In molecular genetics,
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Electrophoresis
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electrophoresis is used to sequence nucleotides (see chapter 13) and to assign various loci to particular chro-
mosomes. In population genetics (see chapter 21), electrophoresis has made it possible to estimate the amount of variability that occurs in natural populations. Here we discuss protein electrophoresis, a process that entails placing a sample often blood serum or a cell homogenate at the top of a gel prepared from a suitable substrate (e.g., hydrolyzed starch, polyacrylamide, or cellulose acetate) and a buffer. An electrical current is
Vertical starch gel apparatus. Current ows from the upper buffer chamber to the lower one by way of the paper wicks and the starch gel. Cooling water ows around the system.
O 1 J Q 2 M Q 3 M M 4 J M 5 J L 6 H J 7 H M 8 G M 9 G J 10 J J
(R. P. Canham, Serum protein variations and selection in uctuating populations of cricetid rodents, Ph.D. thesis, University of Alberta, 1969. Reproduced by permission.)
Figure 2 Ten samples of deer mouse (Peromyscus maniculatus) blood studied for general protein. Al is albumin and Tf is transferrin, the two most abundant proteins in mammalian blood. The six Tf allozymes are labeled G, H, J, L, M, and Q. (R. P. Canham, Serum protein variations and
selection in uctuating populations of cricetid rodents, Ph.D. thesis, University of Alberta, 1969. Reproduced by permission.)
Tamarin: Principles of Genetics, Seventh Edition
II. Mendelism and the Chromosomal Theory
5. Sex Determination, Sex Linkage, and Pedigree Analysis
The McGraw Hill Companies, 2001
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passed through the gel to cause charged molecules to move ( g. 1), and the gel is then treated with a dye that stains the protein. In the simplest case, if a protein is homogeneous (usually the product of a homozygote), it forms a single band on the gel. If it is heterogeneous (usually the product of a heterozygote), it forms two bands. This is because the two allelic protein products differ by an amino acid; they have different electrical charges and therefore travel through the gel at different rates (see g. 5.8). The term allozyme refers to different electrophoretic forms of an enzyme controlled by alleles at the same locus. Figure 2 shows samples of mouse blood serum that have been stained for protein. Most of the staining reveals albumins and -globulins (transferrin). Because they are present in very small concentrations, many enzymes present in the serum are not visible, but a stain that is specific for a particular enzyme can make that enzyme visible on the gel.
For example, lactate dehydrogenase (LDH) can be located because it catalyzes this reaction:
LDH lactic acid NAD pyruvic acid NADH
Thus, we can stain speci cally for the lactate dehydrogenase enzyme by adding the substrates of the enzyme (lactic acid and nicotinamide adenine dinucleotide, NAD ) and a suitable stain speci c for a product of the enzyme reaction (pyruvic acid or nicotinamide adenine dinucleotide, reduced form, NADH). That is, if lactic acid and NAD are poured on the gel, only lactate dehydrogenase converts them to pyruvic acid and NADH. We can then test for the presence of NADH by having it reduce the dye, nitro blue tetrazolium, to the blue precipitate, formazan, an electron carrier. We then add all the preceding reagents and look for blue bands on the gel ( g. 3). In addition to its uses in population genetics and chromosome mapping, electrophoresis has been ex-
tremely useful in determining the structure of many proteins and for studying developmental pathways. As we can see from the lactate dehydrogenase gel in gure 3, ve bands can occur. In some tissues of a homozygote, these bands occur roughly in a ratio of 1:4:6:4:1. This pattern can come about if the enzyme is a tetramer whose four subunits are random mixtures of two gene products (from the A and B loci). Thus we would get AAAA (1/16) AAAB (4/16) AABB (6/16) ABBB (4/16) BBBB (1/16) (Note that the ratio 1:4:6:4:1 is the expansion of [A + B]4, and the relative intensity of each band the number of protein doses is calculated from the rule of unordered events described in chapter 4.)
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