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1/18/2013 1 The Evolution of Populations CHAPT ER 23 2 Microevolution Change in allele frequencies over generations Three mechanisms cause allele frequency change: Natural selection (leads to adaptation) Genetic drift Gene flow 3 In contrast to macroevolution 4 Genetic variation makes evolution possible Some phenotypic variation is environmental and is not heritable 5 Genetic variation within a population Variation can be discrete Mendel’s pea flower color Controlled by one locus Either/or trait Variation and be quantitative Most variation is quantitative Eye color Controlled by two or more genes Continuous variation in trait A locus is fixed if all individuals in a population are homozygous for the same allele 6 Measuring variability Genetic diversity is measured in terms of average heterozygosity The average percentage of loci that are heterozygous 7 Genetic variation between populations 8 A cline is a spatial gradual change in a character 9 Generation of genetic diversity Mutation is the ultimate generation of genetic diversity Point mutation Chromosomal rearrangements Mutation rate is higher in rapidly reproducing organisms and in organisms with RNA genome Sexual reproduction shuffles genetic diversity Crossing over, fertilization, independent sorting of chromosomes 10 Populations share a gene pool 11 Allele frequency In two-allele system, designated as p and q 1 Chromosomal rearrangements Mutation rate is higher in rapidly reproducing organisms and in organisms with RNA genome Sexual reproduction shuffles genetic diversity Crossing over, fertilization, independent sorting of chromosomes 10 Populations share a gene pool 11 Allele frequency In two-allele system, designated as p and q 12 Genotype frequency 13 Hardy Weinberg Equilibrium is used to determine if evolution is occurring For a given p and q, the genotype frequency will be p2 + 2pq + q2 if a population is in Hardy Weinberg equilibrium (i.e., a population is not evolving) p2 and q2 represent the frequencies of the homozygous genotypes and 2pq represents the frequency of the heterozygous genotype Natural populations can evolve at some loci, while being in Hardy-Weinberg equilibrium at other loci 1/18/2013 14 In a study of enzyme variation in a species of grasshopper, you find 15 A1A1, 65 A1A2 and 20 A2A2 in a sample of 100 moths. What is the allele frequency of each allele? What is the genotype frequency of each genotype? What is the expected genotype frequency under Hardy- Weinberg Equilibrium? Is the population in Hardy Weinberg Equilibrium? What can you conclude about this population in terms of evolution? 15 In the pea plant, red individuals may be either homozygous (R1R1) or heterozygous (R1R2), whereas white flowers are homozygous (R2R2). In a sample of plants, there are 35 red flowers and 65 white flowers. Assume that the population is not evolving. What are the allele frequencies? What proportion of red flowers is heterozygous (i.e, what percentage of the red flowers are heterozygous)? 16 A population not in Hardy-Weinberg equilibrium indicates that one or more of the five evolutionary agents are operating in a population 17 Natural selection Natural selection causes adaptive evolution (i.e., the evolution of traits that make organisms better matched to their environment) 18 Genetic drift Random fluctuation in allele frequencies over time by chance Important in small populations Leads to loss of genetic variation, causing alleles to become fixed (even maladaptive alleles) Two types Founder effect Bottleneck effect 19 20 Founder effect 21 Founder effect Few individuals found new population (small allelic pool) 22 Bottleneck effect 2 Bottleneck effect 1/18/2013 19 20 Founder effect 21 Founder effect Few individuals found new population (small allelic pool) 22 Bottleneck effect 23 Bottleneck effect 24 Gene flow A movement of alleles from one population to another Powerful agent of change Tends to homogenize allele frequencies 25 Gene flow can counter adaptation 26 Mutation A random change in a cell’s DNA Mutation rates have little effect on H-W equilibrium Ultimate source of genetic variation 27 Non-random mating Mating with specific genotypes Shifts genotype frequencies but has little change on allele frequencies Assortative mating increases the proportion of homozygous individuals Disassortative mating produces excess of heterozygotes 28 Natural selection Natural selection is not evolution Differential survival and/or reproduction Differences in survival and/or reproduction are not due to chance, but due to a heritable trait that increases fitness 29 Directional selection Selection against one extreme Shifts the mean of a population 30 Negative phototropism in Drosophila 31 Disruptive selection Selection against mean Increases variation 32 Black-bellied seedcracker finch of west Africa 33 Stabilizing selection Selection against extremes Reduces variation but does not change mean 34 Birth weight in humans 35 Sexual selection A type of natural selection driven by interactions between the sexes Leads to sexual dimorphism 36 Intrasexual selection 37 Intersexual selection 38 Male traits may advertise good genes 3 Reduces variation but does not change mean 34 Birth weight in humans 35 Sexual selection A type of natural selection driven by interactions between the sexes Leads to sexual dimorphism 36 Intrasexual selection 37 Intersexual selection 38 Male traits may advertise good genes 39 Maintenance of genetic variation Directional and stabilizing selection reduce genetic variation Balancing selection (heterozygote advantage, oscillating selection and frequency dependent selection) maintain variation 40 Oscillating selection Selection favors one phenotype at one time and a different phenotype at another time Fitness of a phenotype does not depend on its frequency Environmental changes lead to oscillation in selection 41 Oscillating selection Galapagos Islands ground finches Wet conditions favor smaller bills Dry conditions favor larger bills 42 Heterozygote advantage Heterozygotes may exhibit greater fitness than homozygotes Keeps deleterious alleles in a population 43 Malaria Malaria caused by Plasmodium falciparum that infects red blood cells Heterozygote advantage arises from balance of opposing selective factors – anemia and malaria 44 Anemia/malaria 45 Distribution of sickle cell allele and malaria 46 Frequency dependent selection Depends on how frequently a phenotype occurs in a population 47 Why aren’t all organisms perfectly matched with their environment? Must be existing variation Evolutionary constraints Trade-offs (i.e., compromises) occur 1/18/2013 48 4