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Chapter 23 The Evolution of Populations PowerPoint Lectures for Biology, Seventh Edition Neil Campbell and Jane Reece Lectures by Chris Romero Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Overview: The Smallest Unit of Evolution • 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 • Natural selection does act on individuals, however, to see the impact of natural selection we need to look at how a population of organisms changes over a long period of time. • It is the population, not its individuals, that evolves. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Concept 23.1: Population genetics provides a foundation for studying evolution • What is microevolution? • Microevolution is change in the genetic makeup of a population from generation to generation Another way of saying it is: It is the evolution of populations. It is defined as a change in the allele frequencies in a population. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Population Genetics • The Origin of Species convinced most biologists that species are the products of evolution, but acceptance of natural selection as the main mechanism of evolution was more difficult. • Darwin could not explain how chance variations arise in a population or how these variations are transmitted from parents to offspring because he had no understanding of Genetics. Gregor Mendel and Charles Darwin were contemporaries but Mendel’s discoveries were unappreciated at the time even though his principles of heredity would have given credibility to natural selection. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The Modern Synthesis • Population genetics is the study of how populations change genetically over time • Population genetics integrates Mendelian genetics with the Darwinian theory of evolution by natural selection • This modern synthesis focuses on populations as units of evolution Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The modern evolutionary synthesis integrated Darwinian selection and Mendelian inheritance • 1. When Mendel’s research was rediscovered in the early twentieth century, many geneticists believed that the laws of inheritance conflicted with Darwin’s theory of natural selection. • Darwin emphasized those characters that vary along a continuum. These characters are influenced by multiple loci while Mendel investigated “either-or” traits. • An important turning point for evolutionary theory was the birth of population genetics, which emphasizes the extensive genetic variation within populations and recognizes the importance of quantitative characters ( those that vary along a continuum). • Advances in population genetics in the 1930s allowed the ideas of Mendelian Genetics and Darwinism to be reconciled and provided a genetic basis for variation and natural selection. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The modern synthesis • The modern synthesis is a comprehensive theory of evolution, the modern synthesis, took form in the early 1940s and it integrated ideas from paleontology, taxonomy, biogeography, and population genetics. • The architects of the modern synthesis included geneticists Theodosius Dobzhansky and Sewall Wright, biogeographer and taxonomist Ernst Mayr, paleontologist George Simpson, and botanist G. L. Stebbins. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings What is emphasized in this theory? • The modern synthesis emphasizes: 1.The importance of populations as the units of evolution. 2. Natural selection as the most important mechanism of evolution. 3.The idea of gradualism to explain how large changes can evolve as an accumulation of small changes over long periods of time. • While many evolutionary biologists are now challenging some of the assumptions of the modern synthesis, it has shaped most of our ideas about how populations evolve. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Gene Pools and Allele Frequencies • A population is a localized group of individuals capable of interbreeding and producing fertile offspring • The gene pool is the total aggregate of genes in a population at any one time • The gene pool consists of all gene loci in all individuals of the population Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings MAP AREA CANADA ALASKA LE 23-3 Beaufort Sea Porcupine herd range Fairbanks Fortymile herd range Whitehorse A population’s gene pool is defined by its allele frequencies • A population is a group of individuals that belong to the same species and live in the same place. • A species is a group of populations whose individuals have the potential to interbreed and produce fertile offspring. • Populations of a species may be isolated from each other (they exchange genetic material rarely), or they may be integraded with low densities in an intermediate region. • Members of a population are far more likely to breed with members of the same population • Gene pool. is the total aggregate of genes in a population at any one time. A gene pool consists of all alleles at all gene loci in all individuals of a population. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The Hardy-Weinberg Theorem • The Hardy-Weinberg theorem describes a population that is not evolving. • The Hardy-Weinberg theorem describes the gene pool of a non-evolving population. • It states that frequencies of alleles and genotypes in a population’s gene pool will remain constant from generation to generation, provided that only Mendelian segregation and recombination of alleles are at work • Mendelian inheritance preserves genetic variation in a population • In other words, the shuffling of alleles after meiosis and random fertilization should have no effect on the overall gene pool of a population. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 23-4 Generation 1 X CRCR genotype Generation 2 Plants mate CWCW genotype All CRCW (all pink flowers) 50% CW gametes 50% CR gametes come together at random Generation 3 25% CRCR 50% CRCW 50% CR gametes 25% CWCW 50% CW gametes come together at random Generation 4 25% CRCR 50% CRCW 25% CWCW Alleles segregate, and subsequent generations also have three types of flowers in the same proportions Preservation of Allele Frequencies • In a given population where gametes contribute to the next generation randomly, allele frequencies will not change Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Hardy-Weinberg Equilibrium • Hardy-Weinberg equilibrium describes a population in which random mating occurs • It describes a population where allele frequencies do not change Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Hardy-Weinberg equilibrium • All we get by meiosis and sexual reproduction is a reshuffling of the genes that are there but no new ones are introduced so the processes of meiosis and random fertilization just maintain the same allele and genotype frequencies that existed in the previous generation. • The population is in a state of equilibrium called HardyWeinberg equilibrium. • Theoretically, the allele frequencies should remain the same forever. • The Hardy-Weinberg theorem states that the processes involved in a Mendelian system have no tendency to alter allele frequencies from one generation to another. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • 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 – And p2 and q2 represent the frequencies of the homozygous genotypes and 2pq represents the frequency of the heterozygous genotype Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings An example: • We can use the Hardy-Weinberg theorem to estimate the percentage of the human population that carries the allele for a particular inherited disease, phenyketonuria (PKU) in this case. • About 1 in 10,000 babies born in the United States is born with PKU, which results in mental retardation and other problems if left untreated. The disease is caused by a recessive allele. • From the epidemiological data, we know that frequency of homozygous recessive individuals (q2 in the Hardy-Weinberg theorem) = 1 in 10,000 or 0.0001. • The frequency of the recessive allele (q) is the square root of 0.0001 = 0.01. • The frequency of the dominant allele (p) is p = 1 - q or 1 - 0.01 = 0.99. • The frequency of carriers (heterozygous individuals) is 2pq = 2 x 0.99 x 0.01 = 0.0198 or about 2%. • Thus, about 2% of the U.S. population carries the PKU allele. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 23-5 Gametes for each generation are drawn at random from the gene pool of the previous generation: 80% CR (p = 0.8) 20% CW (q = 0.2) Sperm CR CW (20%) p2 pq 64% CRCR 16% CRCW (20%) CR (80%) CW Eggs (80%) qp 4% CWCW 16% CRCW q2 Conditions for Hardy-Weinberg Equilibrium • The Hardy-Weinberg theorem describes a hypothetical population • In real populations, allele and genotype frequencies do change over time Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Why the Hardy-Weinberg equilibrium does not represent a real population? • Natural selection is clearly a violation of the conditions necessary for the Hardy-Weinberg equilibrium. • The Hardy-Weinberg equilibrium expects that all individuals in a population have equal ability to survive and produce viable, fertile offspring. However, in a real population there will be variety among individuals, natural selection will lead some individuals to leave more offspring than others. • Natural selection results in some alleles being passed along to the next generation in numbers disproportionate to their frequencies in the present generation. This is because natural selection accumulates and maintains favorable genotypes in a population. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The five conditions for non-evolving populations are rarely met in nature: – Extremely large population size – No gene flow – No mutations – Random mating – No natural selection Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Populations at Hardy-Weinberg equilibrium must satisfy five conditions. • Very large population size. In small populations, chance fluctuations in the gene pool, genetic drift, can cause genotype frequencies to change over time. • No migrations. Gene flow, the transfer of alleles due to the movement of individuals or gametes into or out of our target population can change the proportions of alleles. • No net mutations. If one allele can mutate into another, the gene pool will be altered. • Random mating. If individuals pick mates with certain genotypes, then the mixing of gametes will not be random and the HardyWeinberg equilibrium does not occur. • No natural selection. If there is differential survival or mating success among genotypes, then the frequencies of alleles in the next variation will deviate from the frequencies predicted by the Hardy-Weinberg equation. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Population Genetics and Human Health • We can use the Hardy-Weinberg equation to estimate the percentage of the human population carrying the allele for an inherited disease Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Concept 23.2: Mutation and sexual recombination produce the variation that makes evolution possible • Two processes, mutation and sexual recombination, produce the variation in gene pools that contributes to differences among individuals Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Mutation • Mutations are changes in the nucleotide sequence of DNA • Mutations cause new genes and alleles to arise Animation: Genetic Variation from Sexual Recombination Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings mutations • A mutation is a change in an organism’s DNA. • A new mutation that is transmitted in gametes can immediately change the gene pool of a population by substituting the mutated allele for the older allele. • Each individual has thousands of genes, any one of which could experience a mutation. • Populations are composed of thousands or millions of individuals that may have experienced mutations. • Over the long term, mutation is a very important to evolution because it is the source of genetic variation that serves as the raw material for natural selection. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Point Mutations • A point mutation is a change in one base in a gene • It is usually harmless but may have significant impact on phenotype Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Mutations That Alter Gene Number or Sequence • Chromosomal mutations that delete, disrupt, or rearrange many loci are typically harmful • Gene duplication is nearly always harmful Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Mutation Rates • Mutation rates are low in animals and plants • The average is about one mutation in every 100,000 genes per generation • Mutations are more rapid in microorganisms Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Sexual Recombination • Sexual recombination is far more important than mutation in producing the genetic differences that make adaptation possible Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Concept 23.3: Natural selection, genetic drift, and gene flow can alter a population’s genetic composition • Three major factors alter allele frequencies and bring about most evolutionary change: – Natural selection – Genetic drift – Gene flow Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Natural Selection • Differential success in reproduction results in certain alleles being passed to the next generation in greater proportions Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Causes of Microevolution • What is microevolution? Microevolution is a generation-to-generation change in a population’s allele frequencies • The Hardy-Weinberg theory provides a baseline against which we can compare the allele and genotype frequencies of an evolving population. • The two main causes of microevolution are genetic drift and natural selection Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings What factors can change the allele frequencies in a population? • Genetic drift • Natural selection • Gene flow • Mutation • All represent departures ( the opposite) from the conditions required for the Hardy-Weinberg equilibrium. • Natural selection is the only factor that adapts a population to its environment, and it always favors the favorable traits. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings How can genetic drift affect a population? • The other three ( genetic drift, gene flow and mutations) may effect populations in positive, negative, or neutral ways. • Genetic drift occurs when changes in gene frequencies from one generation to another occur because of chance events that occur when populations are small in size. The smaller the sample, the greater the chance of deviation from an idealized result. • For example, one would not be too surprised if a coin produced seven heads and three tails in ten tosses, but you would be surprised if you saw 700 heads and 300 tails in 1000 tosses—you expect 500 of each. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Genetic Drift • 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 Animation: Causes of Evolutionary Change Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 23-7 CWCW CRCR CRCR CRCW Only 5 of 10 plants leave offspring CRCR CWCW CRCW CWCW CRCR CRCW CRCW CRCR CRCR CRCR CRCW CRCW Generation 1 p (frequency of CR) = 0.7 q (frequency of CW) = 0.3 CWCW CRCR Only 2 of 10 plants leave offspring CRCR CRCR CRCR CRCR CRCR CRCR CRCR CRCR CRCW CRCW Generation 2 p = 0.5 q = 0.5 CRCR CRCR Generation 3 p = 1.0 q = 0.0 Types of Genetic Drift • The bottleneck effect occurs when the numbers of individuals in a larger population are drastically reduced by a disaster. • By chance, some alleles may be overrepresented and others underrepresented among the survivors. Some alleles may be eliminated altogether. • Bottlenecking is an important concept in conservation biology of endangered species. Populations that have suffered bottleneck incidents have lost at least some alleles from the gene pool. This reduces individual variation and adaptability. • The founder effect occurs when a new population is started by only a few individuals that do not represent the gene pool of the larger population. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The Bottleneck Effect • The bottleneck effect is a sudden change in the environment that may drastically reduce the size of a population • The resulting gene pool may no longer be reflective of the original population’s gene pool Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 23-8 Original population Bottlenecking event Surviving population • Understanding the bottleneck effect can increase understanding of how human activity affects other species Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The Founder Effect • The founder effect occurs when a few individuals become isolated from a larger population • It can affect allele frequencies in a population Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Gene Flow (another factor that changes allele frequency in a population) • Gene flow consists of genetic additions or subtractions from a population, resulting from movement of fertile individuals. • Gene flow causes a population to gain or lose alleles • Gene flow tends to reduce differences between populations over time. • Gene flow is genetic exchange due to migration of fertile individuals between populations. • The migration of people throughout the world is transferring alleles between populations that were once isolated, increasing gene flow. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Concept 23.4: Natural selection is the primary mechanism of adaptive evolution (important factor to change allele frequency in a population) • Natural selection accumulates and maintains favorable genotypes in a population Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Natural Selection happens because of genetic variation • Genetic variation occurs within and between populations • The variation among individuals in a population is a combination of heritable and non-heritable traits. • Phenotype, the observable characteristics of an organism ( its appearance), is the cumulative product of an inherited genotype and many environmental influences. • Only the genetic component of variation can affect evolution.This is because only heritable traits pass from generation to generation. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings What is polygenic inheritance? • When two or more genes determine a single trait. • Quantitative characters are those that vary along a continuum within a population. Examples of this are height, color Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Genetic Variation • Genetic variation occurs in individuals in populations of all species • It is not always heritable Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 23-9 Map butterflies that emerge in spring: orange and brown Map butterflies that emerge in late summer: black and white Variation Within a Population • 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 © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Polymorphism • Phenotypic polymorphism describes a population in which two or more distinct morphs for a character are represented in high enough frequencies to be readily noticeable • Genetic polymorphisms are the heritable components of characters that occur along a continuum in a population Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings What is polymorphism? • Polymorphism occurs when two or more discrete characters are present and noticeable in a population. Polymorphism applies only to discrete characters, not quantitative characters, such as human height, which varies among people in a continuum. • Discrete characters, such as flower color, are usually determined by a single locus (gene) with different alleles.( without intermediates) • The contrasting forms are called morphs, as in the redflowered and white-flowered morphs in our wildflower population. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Measuring Genetic Variation • 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 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Variation Between Populations • Most species exhibit geographic variation differences between gene pools of separate populations or population subgroups Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 23-10 1 2.4 3.14 8.11 9.12 10.16 5.18 6 13.17 19 1 2.19 3.8 4.16 9.10 11.12 13.17 15.18 5.14 7.15 XX 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 © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 23-11 Heights of yarrow plants grown in common garden Mean height (cm) 100 50 0 3,000 2,000 1,000 Sierra Nevada Range 0 Seed collection sites Great Basin Plateau What process is responsible for nearly all the genetic variation in a population? The contribution of mutation to variation is very small. In organisms with sexual reproduction, nearly all the variation results from new combinations of alleles by sexual recombination, in other words, most of the genetic differences among individuals are due to gene recombination of the existing alleles from the population gene pool. • Random segregation of homologous chromosomes and random union of gametes creates a unique assortment of alleles in each individual. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Lets answer these questions to review what we studied so far. • 1.What is meant by “the modern synthesis? • 2.What is a population? A gene pool? A species? • 3.What is the Hardy-Weinberg theorem? What does the Hardy-Weinberg model describe? • 4.What are some practical uses of the Hardy-Weinberg equation? Why is it important? The equation allows the calculation of allele frequencies in a gene pool, if the genotype frequencies are known. It is important in the study of evolution because it tells you what happens in non evolving populations and provides a reference point or a base line to compare to natural populations whose gene pools may be changing. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings More questions…. • 5. What conditions a population must meet to maintain a Hardy-Weinberg equilibrium? Must be very large, isolated, no mutations, random mating, and no natural selection • 6. What five factors can cause microevolution change in a population? Microevolution can be caused by genetic drift, gene flow,mutation, non random mating and natural selection. Each of these is a deviation from the Hardy-Weinberg equilibrium. Of these 5 causes for microevolution, only natural selection leads to adaptations. • 7. What is genetic drift? Why is the effect of genetic drift so great in small populations? Changes in the gene pool of a small population due to chance. The larger the population, the less important is the effect of genetic drift. • 8. What are two situations which result in populations small enough for genetic drift to be important?. The bottleneck effect ( size of the population is reduced drastically by a natural disaster which kills non selectively. The surviving population is unlikely to have the same genetic make up as the original one) and the founder effect ( when a few individuals colonize a new habitat) • 9. How does the bottleneck effects the gene pool of a population? It reduces its genetic variability since some alleles may be absent. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 10. What is gene flow? What are the consequences of gene flow on different populations? Gene flow is the migration of fertile individuals (gametes) between populations. A population may gain or lose alleles this way and results in reducing differences between populations. Extensive gene flow can merge two neighboring populations into one. • 11. What is the affect of mutations on populations? A mutation changes one allele for another. Mutations are rare and have little effect on large populations. If a new mutation increases in frequency is only because that individual produced a larger percentage of offsprings than the rest of the population due to natural selection. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings What are the SOURCES OF GENETIC VARIATION ? • Mutation and sexual recombination generate genetic variation • Mutations are changes in the nucleotide sequence of DNA. • New alleles originate only by mutation but mutations of individual genes are rare and random Only mutations in cell lines that produce gametes can be passed along to offspring. Mutations in somatic cells are lost when the individual dies. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings A Closer Look at Natural Selection • From the range of variations available in a population, natural selection increases frequencies of certain genotypes, fitting organisms to their environment over generations Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Evolutionary Fitness • The phrases “struggle for existence” and “survival of the fittest” are commonly used to describe natural selection but can be misleading • Reproductive success is generally more subtle and depends on many factors Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • 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, compared with contributions of alternative genotypes for the same locus Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Directional, Disruptive, and Stabilizing Selection • Selection favors certain genotypes by acting on the phenotypes of certain organisms • Three modes of selection: – Directional – Disruptive – Stabilizing Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • 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 © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Frequency of individuals LE 23-12 Original population Evolved population Directional selection Original population Phenotypes (fur color) Disruptive selection Stabilizing selection The Preservation of Genetic Variation • Various mechanisms help to preserve genetic variation in a population Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Diploidy • Diploidy maintains genetic variation in the form of hidden recessive alleles Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Balancing Selection • Balancing selection occurs when natural selection maintains stable frequencies of two or more phenotypic forms in a population • Balancing selection leads to a state called balanced polymorphism Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 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 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • An example of heterozygous advantage: The sickle-cell allele causes mutations in hemoglobin but also confers malaria resistance It exemplifies the heterozygote advantage Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 23-13 Frequencies of the sickle-cell allele 0–2.5% 2.5–5.0% 5.0–7.5% Distribution of malaria caused by Plasmodium falciparum (a protozoan) 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 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 23-14 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 a new set of images (a new feeding opportunity). Parental population sample 0.6 Phenotypic variation Experimental group sample 0.5 0.4 Frequencyindependent control 0.3 0.2 0 Plain background Patterned background 20 40 60 Generation number 80 100 Neutral Variation • Neutral variation is genetic variation that appears to confer no selective advantage Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Sexual Selection • Sexual selection is natural selection for mating success • It can result in sexual dimorphism, marked differences between the sexes in secondary sexual characteristics Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Intrasexual selection is competition among individuals of one sex for mates of the opposite sex Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Intersexual selection occurs when individuals of one sex (usually females) are choosy in selecting their mates from individuals of the other sex • Selection may depend on the showiness of the male’s appearance Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The Evolutionary Enigma of Sexual Reproduction • Sexual reproduction produces fewer reproductive offspring than asexual reproduction, a so-called “reproductive handicap” Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings LE 23-16 Asexual reproduction Female Sexual reproduction Generation 1 Female Generation 2 Male Generation 3 Generation 4 • Sexual reproduction produces genetic variation that may aid in disease resistance Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 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 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings