Survey
* Your assessment is very important for improving the work of artificial intelligence, which forms the content of this project
* Your assessment is very important for improving the work of artificial intelligence, which forms the content of this project
Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings One misconception is that organisms evolve, in the Darwinian sense, during their lifetimes Natural selection acts on individuals, but only populations evolve Genetic variations in populations contribute to evolution Microevolution is a change in allele frequencies in a population over generations Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-1 Two processes, mutation and sexual reproduction, produce the variation in gene pools that contributes to differences among individuals Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Variation in individual genotype leads to variation in individual phenotype Not all phenotypic variation is heritable Natural selection can only act on variation with a genetic component Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-2 (a) (b) Fig. 23-2a (a) Fig. 23-2b (b) Population geneticists measure polymorphisms in a population by determining the amount of heterozygosity at the gene and molecular levels Average heterozygosity measures the average percent of loci that are heterozygous in a population Nucleotide variability is measured by comparing the DNA sequences of pairs of individuals Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Most species exhibit geographic variation, differences between gene pools of separate populations or population subgroups Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-3 1 2.4 8.11 9.12 3.14 5.18 10.16 13.17 6 7.15 19 XX 1 2.19 3.8 4.16 5.14 9.10 11.12 13.17 15.18 6.7 XX • Some examples of geographic variation occur as a cline, which is a graded change in a trait along a geographic axis Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-4 1.0 0.8 0.6 0.4 0.2 0 46 Maine Cold (6°C) 44 42 40 38 36 Latitude (°N) 34 32 30 Georgia Warm (21°C) Mutations are changes in the nucleotide sequence of DNA Mutations cause new genes and alleles to arise Only mutations in cells that produce gametes can be passed to offspring Animation: Genetic Variation from Sexual Recombination Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Mutation rates are low in animals and plants The average is about one mutation in every 100,000 genes per generation Mutations rates are often lower in prokaryotes and higher in viruses Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Both discrete and quantitative characters contribute to variation within a population Discrete characters can be classified on an eitheror basis Quantitative characters vary along a continuum within a population Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Sexual reproduction can shuffle existing alleles into new combinations In organisms that reproduce sexually, recombination of alleles is more important than mutation in producing the genetic differences that make adaptation possible Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings The first step in testing whether evolution is occurring in a population is to clarify what we mean by a population Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings • A population is a localized group of individuals capable of interbreeding and producing fertile offspring A gene pool consists of all the alleles for all loci in a population A locus is fixed if all individuals in a population are homozygous for the same allele Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-5 Porcupine herd MAP AREA Beaufort Sea Porcupine herd range Fortymile herd range Fortymile herd Fig. 23-5a MAP AREA Beaufort Sea Porcupine herd range Fortymile herd range The frequency of an allele in a population can be calculated For diploid organisms, the total number of alleles at a locus is the total number of individuals x 2 The total number of dominant alleles at a locus is 2 alleles for each homozygous dominant individual plus 1 allele for each heterozygous individual; the same logic applies for recessive alleles Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings By convention, if there are 2 alleles at a locus, p and q are used to represent their frequencies The frequency of all alleles in a population will add up to 1 For example, p + q = 1 Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings The Hardy-Weinberg principle describes a population that is not evolving If a population does not meet the criteria of the Hardy-Weinberg principle, it can be concluded that the population is evolving Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings The Hardy-Weinberg principle states that frequencies of alleles and genotypes in a population remain constant from generation to generation In a given population where gametes contribute to the next generation randomly, allele frequencies will not change Mendelian inheritance preserves genetic variation in a population Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-6 Alleles in the population Frequencies of alleles p = frequency of CR allele = 0.8 q = frequency of CW allele = 0.2 Gametes produced Each egg: Each sperm: 80% 20% chance chance 80% 20% chance chance Hardy-Weinberg equilibrium describes the constant frequency of alleles in such a gene pool If p and q represent the relative frequencies of the only two possible alleles in a population at a particular locus, then p2 + 2pq + q2 = 1 where p2 and q2 represent the frequencies of the homozygous genotypes and 2pq represents the frequency of the heterozygous genotype Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-7-1 80% CR (p = 0.8) 20% CW (q = 0.2) Sperm CR (80%) CW (20%) 64% (p2) CRCR 16% (pq) CRCW 16% (qp) CRCW 4% (q2) CW CW Fig. 23-7-2 64% CRCR, 32% CRCW, and 4% CWCW Gametes of this generation: 64% CR + 16% CR = 80% CR = 0.8 = p 4% CW + 16% CW = 20% CW = 0.2 = q Fig. 23-7-3 64% CRCR, 32% CRCW, and 4% CWCW Gametes of this generation: 64% CR + 16% CR = 80% CR = 0.8 = p 4% CW + 16% CW = 20% CW = 0.2 = q Genotypes in the next generation: 64% CRCR, 32% CRCW, and 4% CWCW plants Fig. 23-7-4 20% CW (q = 0.2) 80% CR ( p = 0.8) Sperm (80%) CW (20%) 64% ( p2) CR CR 16% ( pq) CR CW CR 16% (qp) CR CW 4% (q2) CW CW 64% CR CR, 32% CR CW, and 4% CW CW Gametes of this generation: 64% CR + 16% CR = 80% CR = 0.8 = p 4% CW = 20% CW = 0.2 = q + 16% CW Genotypes in the next generation: 64% CR CR, 32% CR CW, and 4% CW CW plants The Hardy-Weinberg theorem describes a hypothetical population In real populations, allele and genotype frequencies do change over time Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings The five conditions for nonevolving populations are rarely met in nature: No mutations Random mating No natural selection Extremely large population size No gene flow Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings We can assume the locus that causes phenylketonuria (PKU) is in Hardy-Weinberg equilibrium given that: The PKU gene mutation rate is low Mate selection is random with respect to whether or not an individual is a carrier for the PKU allele Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Natural selection can only act on rare homozygous individuals who do not follow dietary restrictions The population is large Migration has no effect as many other populations have similar allele frequencies Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings The occurrence of PKU is 1 per 10,000 births q2 = 0.0001 q = 0.01 The frequency of normal alleles is p = 1 – q = 1 – 0.01 = 0.99 The frequency of carriers is 2pq = 2 x 0.99 x 0.01 = 0.0198 or approximately 2% of the U.S. population Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Three major factors alter allele frequencies and bring about most evolutionary change: Natural selection Genetic drift Gene flow Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Differential success in reproduction results in certain alleles being passed to the next generation in greater proportions Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings The smaller a sample, the greater the chance of deviation from a predicted result Genetic drift describes how allele frequencies fluctuate unpredictably from one generation to the next Genetic drift tends to reduce genetic variation through losses of alleles Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-8-1 CR CR CR CR CR CW CR CR CW CW CR CW CR CR CR CR CR CW CR CW Generation 1 p (frequency of CR) = 0.7 q (frequency of CW ) = 0.3 Fig. 23-8-2 CR CR CR CR CW CW CR CW CR CW CR CR CW CW CW CW CR CR CR CW CR CW CR CR CR CR CR CR CR CW CR CW Generation 1 p (frequency of CR) = 0.7 q (frequency of CW ) = 0.3 CW CW CR CW CR CR CR CW Generation 2 p = 0.5 q = 0.5 Fig. 23-8-3 CR CR CR CR CW CW CR CW CR CW CR CR CW CW CR CR CR CW CR CR CR CW CR CW Generation 1 p (frequency of CR) = 0.7 q (frequency of CW ) = 0.3 CW CW CR CW CR CR CR CR CR CR CW CW CR CR CR CW CR CR CR CR CR CR CR CR CR CR CR CR CR CR CR CW Generation 2 p = 0.5 q = 0.5 CR CR CR CR Generation 3 p = 1.0 q = 0.0 The founder effect occurs when a few individuals become isolated from a larger population Allele frequencies in the small founder population can be different from those in the larger parent population Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings The bottleneck effect is a sudden reduction in population size due to a change in the environment The resulting gene pool may no longer be reflective of the original population’s gene pool If the population remains small, it may be further affected by genetic drift Understanding the bottleneck effect can increase understanding of how human activity affects other species Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-9 Original population Bottlenecking event Surviving population Loss of prairie habitat caused a severe reduction in the population of greater prairie chickens in Illinois The surviving birds had low levels of genetic variation, and only 50% of their eggs hatched Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-10a Pre-bottleneck (Illinois, 1820) (a) Range of greater prairie chicken Post-bottleneck (Illinois, 1993) Fig. 23-10b Location Population size Number Percentage of alleles of eggs per locus hatched Illinois 1,000–25,000 5.2 93 <50 3.7 <50 Kansas, 1998 (no bottleneck) 750,000 5.8 99 Nebraska, 1998 (no bottleneck) 75,000– 200,000 5.8 96 Minnesota, 1998 (no bottleneck) 4,000 5.3 85 1930–1960s 1993 (b) Genetic drift is significant in small populations 2. Genetic drift causes allele frequencies to change at random 3. Genetic drift can lead to a loss of genetic variation within populations 4. Genetic drift can cause harmful alleles to become fixed 1. Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Gene flow consists of the movement of alleles among populations Alleles can be transferred through the movement of fertile individuals or gametes (for example, pollen) Gene flow tends to reduce differences between populations over time Gene flow is more likely than mutation to alter allele frequencies directly Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Gene flow can decrease the fitness of a population In bent grass, alleles for copper tolerance are beneficial in populations near copper mines, but harmful to populations in other soils Windblown pollen moves these alleles between populations The movement of unfavorable alleles into a population results in a decrease in fit between organism and environment Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Gene flow can increase the fitness of a population Insecticides have been used to target mosquitoes that carry West Nile virus and malaria Alleles have evolved in some populations that confer insecticide resistance to these mosquitoes The flow of insecticide resistance alleles into a population can cause an increase in fitness Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Only natural selection consistently results in adaptive evolution Natural selection brings about adaptive evolution by acting on an organism’s phenotype The phrases “struggle for existence” and “survival of the fittest” are misleading as they imply direct competition among individuals Reproductive success is generally more subtle and depends on many factors Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings The phrases “struggle for existence” and “survival of the fittest” are misleading as they imply direct competition among individuals Reproductive success is generally more subtle and depends on many factors Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Relative fitness is the contribution an individual makes to the gene pool of the next generation, relative to the contributions of other individuals Selection favors certain genotypes by acting on the phenotypes of certain organisms Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Three modes of selection: Directional selection favors individuals at one end of the phenotypic range Disruptive selection favors individuals at both extremes of the phenotypic range Stabilizing selection favors intermediate variants and acts against extreme phenotypes Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-13 Original population Original Evolved population population (a) Directional selection Phenotypes (fur color) (b) Disruptive selection (c) Stabilizing selection Fig. 23-13a Original population Phenotypes (fur color) Original population Evolved population (a) Directional selection Fig. 23-13b Original population Phenotypes (fur color) Evolved population (b) Disruptive selection Fig. 23-13c Original population Phenotypes (fur color) Evolved population (c) Stabilizing selection Natural selection increases the frequencies of alleles that enhance survival and reproduction Adaptive evolution occurs as the match between an organism and its environment increases Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-14a (a) Color-changing ability in cuttlefish Fig. 23-14b Movable bones (b) Movable jaw bones in snakes Because the environment can change, adaptive evolution is a continuous process Genetic drift and gene flow do not consistently lead to adaptive evolution as they can increase or decrease the match between an organism and its environment Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Sexual selection is natural selection for mating success It can result in sexual dimorphism, marked differences between the sexes in secondary sexual characteristics Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-15 Intrasexual selection is competition among individuals of one sex (often males) for mates of the opposite sex Intersexual selection, often called mate choice, occurs when individuals of one sex (usually females) are choosy in selecting their mates Male showiness due to mate choice can increase a male’s chances of attracting a female, while decreasing his chances of survival Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings How do female preferences evolve? The good genes hypothesis suggests that if a trait is related to male health, both the male trait and female preference for that trait should be selected for Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-16 EXPERIMENT Female gray tree frog SC male gray tree frog LC male gray tree frog SC sperm Eggs LC sperm Offspring of Offspring of SC father LC father Fitness of these half-sibling offspring compared RESULTS Fitness Measure 1995 1996 Larval growth NSD LC better Larval survival LC better NSD Time to metamorphosis LC better (shorter) LC better (shorter) NSD = no significant difference; LC better = offspring of LC males superior to offspring of SC males. Diploidy maintains genetic variation in the form of hidden recessive alleles Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Balancing selection occurs when natural selection maintains stable frequencies of two or more phenotypic forms in a population Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Heterozygote Advantage • Heterozygote advantage occurs when heterozygotes have a higher fitness than do both 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 Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-17 Frequencies of the sickle-cell allele 0–2.5% Distribution of malaria caused by Plasmodium falciparum (a parasitic unicellular eukaryote) 2.5–5.0% 5.0–7.5% 7.5–10.0% 10.0–12.5% >12.5% Neutral variation is genetic variation that appears to confer no selective advantage or disadvantage For example, Variation in noncoding regions of DNA Variation in proteins that have little effect on protein function or reproductive fitness Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Selection can act only on existing variations 2. Evolution is limited by historical constraints 3. Adaptations are often compromises 4. Chance, natural selection, and the environment interact 1. Copyright © 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings Fig. 23-19