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Mechanisms of Evolution Microevolution Population Genetics Key Concepts • 23.1: Population genetics provides a foundation for studying evolution • 23.2: Mutation and sexual recombination produce the variation that makes evolution possible • 23.3: Natural selection, genetic drift, and gene flow can alter a population’s genetic composition • 23.4: Natural selection is the primary mechanism of adaptive evolution • Concept 23.4: Natural selection is the primary mechanism of adaptive evolution • Natural selection – Accumulates and maintains favorable genotypes in a population – Is the only deviation from the Hardy-Weinberg that leads to adaptation (of the population to the environment) 1 A Closer Look at Natural Selection • From the range of variations available in a population – Natural selection increases the frequencies of certain genotypes, fitting organisms to their environment over generations Evolutionary Fitness • The phrases “struggle for existence” and “survival of the fittest” – Are commonly used to describe natural selection – Can be misleading • Reproductive success – Is generally more subtle and depends on many factors • Fitness – Is the contribution an individual makes to the gene pool of the next generation, relative to the contributions of other individuals • Relative fitness – Is the contribution of a genotype to the next generation as compared to the contributions of alternative genotypes for the same locus 2 3 Modes of Selection • Selection – Favors certain genotypes by acting on the phenotypes of certain organisms • Three modes of selection are – Directional – Disruptive – Stabilizing • Directional selection – Favors individuals at one end of the phenotypic range • Disruptive selection – Favors individuals at both extremes of the phenotypic range • Stabilizing selection Frequency of individuals – Favors intermediate variants and acts against extreme phenotypes Original population Evolved population (a) Directional selection shifts the overall makeup of the population by favoring variants at one extreme of the distribution. In this case, darker mice are favored because they live among dark rocks and a darker fur color conceals them from predators. Original population Phenotypes (fur color) (b) Disruptive selection favors variants at both ends of the distribution. These mice have colonized a patchy habitat made up of light and dark rocks, with the result that mice of an intermediate color are at a disadvantage. (c) Stabilizing selection removes extreme variants from the population and preserves intermediate types. If the environment consists of rocks of an intermediate color, both light and dark mice will be selected against. Fig 23.12 A–C 3 The Preservation of Genetic Variation • Various mechanisms help to preserve genetic variation in a population Diploidy • Diploidy – Maintains genetic variation in the form of hidden recessive alleles Balancing Selection • Balancing selection – Occurs when natural selection maintains stable frequencies of two or more phenotypic forms in a population – Leads to a state called balanced polymorphism 4 Heterozygote Advantage • Some individuals who are heterozygous at a particular locus – Have greater fitness than homozygotes • Natural selection – Will tend to maintain two or more alleles at that locus • The sickle-cell allele – Causes mutations in hemoglobin but also confers malaria resistance – Exemplifies the heterozygote advantage Distribution of malaria caused by Plasmodium falciparum (a protozoan) Figure 23.13 Frequencies of the sickle-cell allele 0–2.5% 2.5–5.0% 5.0–7.5% 7.5–10.0% 10.0–12.5% >12.5% Frequency-Dependent Selection • In frequency-dependent selection – The fitness of any morph declines if it becomes too common in the population 5 Example of frequencydependent selection On pecking a moth image the blue jay receives a food reward. If the bird does not detect a moth on either screen, it pecks the green circle to continue to a new set of images (a new feeding opportunity). Parental population sample Phenotypic diversity Experimental group sample 0.06 0.05 0.04 Frequencyindependent control 0.03 0.02 0 Plain background Patterned background 20 60 40 80 Generation number 100 Figure 23.14 Neutral Variation • Is genetic variation that appears to confer no selective advantage Sexual Selection • Sexual selection – Is natural selection for mating success – Can result in sexual dimorphism, marked differences between the sexes in secondary sexual characteristics 6 Intrasexual selection • Is a direct competition among individuals of one sex for mates of the opposite sex Intersexual selection • Occurs when individuals of one sex (usually females) are choosy in selecting their mates from individuals of the other sex • May depend on the showiness of the male’s appearance Figure 23.15 The Evolutionary Enigma of Sexual Reproduction • Sexual reproduction – Produces fewer reproductive offspring than asexual reproduction, a so-called reproductive handicap Sexual reproduction Asexual reproduction Generation 1 Female Female Generation 2 Male Generation 3 Generation 4 Figure 23.16 7 • If sexual reproduction is a handicap, why has it persisted? – It produces genetic variation that may aid in disease resistance Why Natural Selection Cannot Fashion Perfect Organisms • Evolution is limited by historical constraints • Adaptations are often compromises • Chance and natural selection interact • Selection can only edit existing variations 8