Genes Within Populations Chapter 20 1 Genetic Variation and Evolution • Darwin: Evolution is descent with modification. • Evolution: changes through time. 1. Species accumulate difference; 2. Descendants differ from their ancestors; 3. New species arise from existing ones. 2 Natural selection: mechanism of evolutionary change Natural selection: proposed by Darwin as the mechanism of evolution. • individuals have specific inherited characteristics; • they produce more surviving offspring; • the population includes more individuals with these specific characteristics; • the population evolves and is better adapted to its present environment. 3 Darwin’s theory for how long necks evolved in giraffes 4 Natural selection: mechanism of evolutionary change Inheritance of acquired characteristics: Proposed by JeanBaptiste Lamarck. • Individuals passed on physical and behavioral changes to their offspring; • Variation by experience…not genetic; • Darwin’s natural selection: variation a result of preexisting genetic differences. 5 Lamarck’s theory of how giraffes’ long necks evolved. 6 Hardy-Weinberg Principle Godfrey H. Hardy: English mathematician. Wilhelm Weinberg: German physician. Concluded that: The original proportions of the genotypes in a population will remain constant from generation to generation as long as five assumptions are met: 7 Hardy-Weinberg Principle Five assumptions : 1. 2. 3. 4. 5. No mutation takes place No genes are transferred to or from sources Random mating is occurring The population size is very large No selection occurs other 8 Hardy-Weinberg Principle Calculate genotype frequencies with a binomial expansion: (p+q)2 = p2 + 2pq + q2 • • • • p = individuals homozygous for first allele; 2pq = individuals heterozygous for both alleles; q = individuals homozygous for second allele; because there are only two alleles: p plus q must always equal 1 (that is the total population) 9 Hardy-Weinberg Principle Using Hardy-Weinberg equation to predict frequencies in subsequent generations. 10 Agents of Evolutionary Change A population not in Hardy-Weinberg equilibrium indicates that one or more of the five evolutionary agents are operating in a population. • Mutation: A change in a cell’s DNA. – Mutation rates are generally so low they have little effect on Hardy-Weinberg proportions of common alleles. – Ultimate source of genetic variation. 11 Gene flow: A movement of alleles from one population to another. Powerful agent of change; Tends to homogenize allele frequencies. 12 Agents of Evolutionary Change • Nonrandom Mating: mating with specific genotypes: – Shifts genotype frequencies – Assortative Mating: does not change frequency of individual alleles; increases the proportion of homozygous individuals – Disassortative Mating: phenotypically different individuals mate; produce excess of heterozygotes 13 Genetic Drift • Genetic drift: Random fluctuation in allele frequencies over time by chance. • important in small populations – founder effect - few individuals found new population (small allelic pool) – bottleneck effect - drastic reduction in population, and gene pool size 14 Genetic Drift: A bottleneck effect 15 Selection • Artificial selection: a breeder selects for desired characteristics. 16 Selection • Natural selection: environmental conditions determine which individuals in a population produce the most offspring. • 3 conditions for natural selection to occur: – Variation must exist among individuals in a population; – Variation among individuals must result in differences in the number of offspring surviving; – Variation must be genetically inherited. 17 Selection Pocket mice from the Tularosa Basin 18 Selection to match climatic conditions Selection for pesticide resistance 19 Fitness and Its Measurement • Fitness is a combination of: – Survival: how long does an organism live – Mating success: how often it mates – Number of offspring per mating that survive Body size and egg-laying in water striders. 20 Interactions Among Evolutionary Forces • Mutation and genetic drift may counter selection. • The magnitude of drift is inversely related to population size. 21 Interactions Among Evolutionary Forces • Gene flow may promote or constrain evolutionary change: – Spread a beneficial mutation; – Impede adaptation by continual flow of inferior alleles from other populations. • Extent to which gene flow can hinder the effects of natural selection depends on the relative strengths of gene flow: – High in birds & wind-pollinated plants; – Low in sedentary species. 22 Maintenance of Variation • Frequency-dependent selection: depends on how frequently or infrequently a phenotype occurs in a population. – Negative frequency-dependent selection: rare phenotypes are favored by selection. – Positive frequency-dependent selection: common phenotypes are favored; variation is eliminated from the population. • Strength of selection changes through time. 23 Maintenance of Variation Negative frequency dependent selection Positive frequencydependent selection. 24 Maintenance of Variation • Oscillating selection: selection favors one phenotype at one time, and a different phenotype at another time. • Galápagos Islands ground finches – Wet conditions favor big bills (abundant seeds); – Dry conditions favor small bills; Heterozygotes may exhibit greater fitness than homozygotes. 25 Maintenance of Variation • Homozygous dominant phenotype: no anemia; susceptible to malaria. • Heterozygous phenotype: no anemia; less susceptible to malaria Frequency of sickle cell allele. 26 Maintenance of Variation Disruptive selection: acts to eliminate intermediate types. Disruptive selection for large and small beaks in blackbellied seedcracker finch of west Africa. 27 Maintenance of Variation Directional selection: acts to eliminate one extreme from an array of phenotypes Directional selection for negative phototropism in Drosophila. 28 Maintenance of Variation Stabilizing selection: acts to eliminate both extremes. Stabilizing selection for birth weight in humans. 29 Experimental Studies of Natural Selection • In some cases, evolutionary change can occur rapidly; • Evolutionary studies can be devised to test evolutionary hypotheses; • Guppy studies (Poecilia reticulata) in the lab and field – Populations above the waterfalls: low predation – Populations below the waterfalls: high predation 30 Experimental Studies • High predation environment - Males exhibit drab coloration and tend to be relatively small and reproduce at a younger age. • Low predation environment - Males display bright coloration, a larger number of spots, and tend to be more successful at defending territories. 31 The Limits of Selection • Genes have multiple effects: – Pleiotropy: sets limits on how much a phenotype can be altered. • Evolution requires genetic variation – Thoroughbred horse speed – Compound eyes of insects: same genes affect both eyes – Control of ommatidia number in left and right eye 32 Experimental Studies Selection for increased speed in racehorses is no longer effective. 33 Experimental Studies Phenotypic variation in insect ommatidia. 34