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IV. Quantitative and Evolutionary Genetics
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21. Evolution and Speciation
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STUDY OBJECTIVE 1: To analyze the mechanisms of evo-
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lution and speciation
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The theory of evolution by natural selection was put forward by Charles Darwin, who recognized the natural variation among individuals within a population of similar organisms. He noted also that offspring are overproduced in nature, and this overproduction inevitably leads to competition for scarce resources. Darwin assumed that, when competition occurs, the most t will survive; through time, then, a population will become better adapted to its environment through the process of natural selection. Applying the algebra of population genetics to this theory leads to the modern concept of evolution, neo-Darwinism. Cladogenic speciation occurs when reproductive isolating mechanisms arise, usually after gene ow in a population is blocked. Different populations of a species can then evolve independently. When individuals from the isolates can no longer interbreed, speciation has taken place. If the isolates then come in contact again, they will remain as separate species. Speciation may occur gradually or in a punctuated manner; it can be by allopatric, parapatric, or sympatric mechanisms. STUDY OBJECTIVE 2: To investigate the mechanisms of
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amount of heterozygosity occurred in natural populations. Attempts to explain this variation have led to two major competing theories: (1) variation is maintained selectively and (2) variation is not under selective pressure, but is instead neutral. Two areas of evidence support the neutralist view. First, the molecular evolutionary clock (the per-year, per-amino acid, substitution rate) appears to be fairly constant at 10 9. This constancy implies that the majority of amino acid changes are the result of stochastic processes. Second, there have been greater numbers of nucleotide substitutions in DNA under lesser constraint than in DNA under greater constraint. For example, the third, or wobble, position of the codon has accumulated more mutations than the other two positions. We conclude that natural selection creates adapted organisms, but the majority of base and amino acid changes may be neutral. STUDY OBJECTIVE 3: To discuss sociobiology, the evolu-
tion of social behavior
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the maintenance of genetic variation in natural populations, both selective and neutral 596 603
Evolution depends on variation. In 1966, Lewontin and Hubby, using electrophoresis, showed that a tremendous
Sociobiology is another term for evolutionary behavioral ecology. It attempts to provide evolutionary explanations for social behaviors. Apparent altruistic behavior can be explained either as kin selection or as sel shness. Sterile insect castes have come about because of the unusual haplodiploid sex-determining mechanism in the eusocial hymenoptera. There is much controversy about and little information for applying sociobiological principles to human behavior.
S O L V E D
PROBLEM 1: What are the roles of reproductive isolating
P R O B L E M S
If weak reproductive isolating mechanisms have evolved, natural selection usually favors strengthening them by selecting against hybrids and against any mating behavior that leads to the formation of hybrids.
PROBLEM 2: What is our modern evolutionary concept
mechanisms in the process of evolution Answer: Reproductive isolating mechanisms prevent individuals in two populations from mating with each other or producing viable offspring. These mechanisms can be prezygotic or postzygotic. They usually evolve while populations are isolated from each other, either physically or during parapatric or sympatric speciation. For example, if a species is split by a new river, the populations on either side of the river can evolve in isolation from each other. Reproductive isolating mechanisms usually evolve irrespective of the other facets of evolution taking place. Thus, if, after time, the two populations come into contact (the river dries up), reproductive isolating mechanisms may have evolved to prevent mating.
of altruism Answer: An altruistic act is one in which an individual risks the loss of tness in order to bene t another individual. Human beings value these sel ess acts; however, they are not favored in natural animal populations, except under very speci c circumstances, because altruistic acts should be selected against. In other words, all other things being equal, an individual that did not do altruistic acts would have a higher tness than one that did
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