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IV. Quantitative and Evolutionary Genetics
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21. Evolution and Speciation
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Table 21.4 Amino Acid Differences (By Percentage) in Cytochrome c Between Different Organisms
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Human being Pig, bovine, sheep Horse Chicken, turkey Snapping turtle Bullfrog Tuna Carp Lamprey Fruit y Screwworm y Silkworm moth Sesame Sun ower Wheat Candida krusei Baker s yeast Neurospora crassa Rhodospirillum rubrum
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10 0
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12 3 0
13 9 11 0
14 9 11 8 0
17 11 13 11 10 0
20 16 18 16 17 14 0
17 11 13 14 13 13 8 0
19 13 15 17 18 20 18 12 0
27 22 22 23 22 20 23 21 27 0
25 20 20 21 22 20 22 20 26 2 0
29 25 27 26 26 27 30 25 30 14 13 0
35 38 39 40 38 41 42 40 44 42 41 39 0
38 40 41 41 39 42 43 41 44 41 40 40 10 0
38 40 41 41 41 43 44 42 46 42 40 40 13 13 0
46 45 46 45 47 46 43 45 50 43 43 43 47 47 45 0
41 41 42 41 44 43 43 42 45 42 42 44 44 43 42 25 0
44 43 43 44 45 45 45 43 47 38 38 44 48 49 48 39 38 0
65 64 64 64 64 65 65 64 66 65 64 65 65 67 66 72 69 69 0
Source: From M. O. Dayhoff, ed., Atlas of Protein Sequence and Structure, National Biomedical Research Foundation, Washington, D.C., 1972. Reprinted with permission.
honor of Linus Pauling, who, along with E. Zuckerkandl, rst proposed the concept of a molecular clock in 1963. If the values of k (such as those in table 21.5) form a normal distribution around 10 9, then 10 9 would be the rate of the molecular evolutionary clock. So far, the data have been too limited to determine the distribution. Although controversy still exists, the neutralists have interpreted the relative constancy of the molecular evolutionary clock as strong evidence in support of the neutral gene hypothesis. A constant rate of molecular evolution over many groups of organisms over many different time intervals implies that the substitution rate is a stochastic or random process rather than a directed or selectional process. This is not to say that no adapted changes occur in proteins or that there are no constraints. In fact, the evidence suggests that three classes of amino acids can be grouped in terms of substitution rate: invariant, moderately variant, and hypervariant. It seems possible that virtually no substitutions of amino acids will occur in and around the active site of the enzyme since any amino acid change in that area might be deleterious or lethal. For ex-
ample, a segment of cytochrome c that runs from amino acids 70 to 80 is invariant in all organisms tested.This area includes a binding site of the protein.
DNA Variation
If the neutralist view of molecular evolution is correct, we should be able to make some predictions about rates of change in DNA. For example, we predict that DNA under greater constraint should amass fewer base changes than DNA under lesser constraint. We could test this by looking at the accumulation of mutations in the three positions of the codon, or we could look at DNA that is not directly translated, such as pseudogenes (see chapter 15) or introns, which are probably under lesser constraint. Let us rst look at the three positions of the codon. A reexamination of the codon dictionary (see table 11.4) shows that the third, or wobble, position of the codon should be under less constraint. Eight amino acids belong to unmixed families; their amino acids are de ned by the rst and second positions coupled with any of the
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