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Does electrophoresis randomly sample the genome Since, on the basis of DNA content, the genome of higher organisms has the potential to contain half a million genes, this question may be dif cult to resolve. Since the
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Tamarin: Principles of Genetics, Seventh Edition
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IV. Quantitative and Evolutionary Genetics
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21. Evolution and Speciation
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Table 21.3 Survey of Genic Heterozygosity
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Proportion of Loci Polymorphic per Population 0.28 0.29 0.30 0.20 0.23 0.43 0.25 0.53 0.47 0.86 0.81 0.42 0.61 0.25
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Species Homo sapiens Mus musculus musculus M. m. brevirostris M. m. domesticus Peromyscus polionotus Drosophila pseudoobscura D. persimilis D. obscura D. subobscura D. willistoni D. melanogaster D. simulans Limulus polyphemus
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Number of Populations 1 4 1 2 7 (regions) 10 1 3 (regions) 6 2 21 10 1 1 4
Number of Loci 71 41 40 41 32 24 24 30 31 28 20 19 18 25
Heterozygosity per Locus 0.067 0.091 0.110 0.056 0.057 0.128 0.106 0.108 0.076 0.184 0.175 0.119 0.160 0.061
Standard Error of Heterozygosity 0.018 0.023 0.022 0.014 0.041 0.040 0.030 0.024 0.032 0.039 0.037 0.052 0.024
Source: The Genetic Basis of Evolutionary Change by R. C. Lewontin, (New York: Columbia University Press, 1974). Reprinted with permission of the publisher. Note: See source (Lewontin, 1974) for individual references.
number of polymorphisms that exist in natural populations could be maintained according to these models.
Which Hypothesis Is Correct
Researchers who favor the concept that most electrophoretic alleles are neutral do not deny that selection exists. They do not hold that evolution is non-adaptive, but say merely that most of the molecular variation (electrophoretic) found in nature is not related to tness it is neutral. Thus, the demonstration that selection actually exists, in electrophoretic systems or otherwise, is not proof against the neutralist view. No one denies the explanation for the maintenance of sickle-cell anemia. Selection at several other electrophoretic systems is also known. For example, R. Koehn showed that different alleles of an esterase locus in a freshwater sh in Colorado produced proteins with different enzyme activities at different water temperatures. Koehn then showed that the alleles were distributed as one would predict on the basis of the water temperature. In other words, the distribution of alleles correlated with the distribution of water temperature. The enzyme produced by the ES-1a allele functioned best at warm temperatures, whereas the enzyme produced by the ES-1b allele functioned best at cold temperatures. The cold-adapted enzyme was prevalent in the sh in colder waters (higher latitudes), and the warm-adapted enzyme was prevalent in the sh in warmer waters (lower latitudes; g. 21.8).
Neutral Alleles
The high incidence of polymorphism that electrophoresis reveals may not be important from an evolutionary point of view. If all or most electrophoretic alleles are neutral (i.e., if no allele is more t than its alternative) or only very slightly deleterious, there is virtually no selection at these loci, and the variation observed in the population is merely a chance accumulation of a combination of mutation and genetic drift.This model, proposed by M. Kimura of Japan, is an alternative to the natural selection model.
Motoo Kimura (1924 1994). (Courtesy of
Dr. Motoo Kimura.)
Tamarin: Principles of Genetics, Seventh Edition
IV. Quantitative and Evolutionary Genetics
21. Evolution and Speciation
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Twenty-One
Evolution and Speciation
morphic loci are either being selected or are neutral. For this proof, many loci must be examined independently a very dif cult undertaking or some grand pattern must emerge supporting one hypothesis or the other.
Grand Patterns of Variation
Clinal Selection
Data on the geographic distribution of alleles fail to adequately support either theory. Often, a single allele predominates over the range of a species ( g. 21.9). Changes in allelic frequency from one geographic area to another can often be attributed to clinal selection, selection along a geographic gradient, in which allelic frequencies change as altitude, latitude, or some other geographic attribute changes. Note in gure 21.9 the general increase in the Es-5b frequency from west to east in the southern United States. But, in line with the neutralist view, geographic patterns similar to those in gure 21.9 can also be produced by neutral alleles with a very low level of migration, as little as one individual per one thousand per generation.
Relation of latitude and frequency of the warmadapted esterase allele Es-I a in populations of the sh Catostomus clarki. Note how the frequency of the allele increases as latitude decreases (warmer water). (From Richard
Koehn, Functional and evolutionary dynamics of polymorphic esterases in catostomid shes, Transactions of the American Fisheries Society, 99:223. Copyright 1970 American Fisheries Society, Bethesda, MD.)
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