1. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings PowerPoint Lectures for Biology, Seventh Edition Neil Campbell and Jane Reece Lectures by Chris Romero Chapter 23 The Evolution of Populations

2. 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

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4. Concept 23.1: Population genetics provides a foundation for studying evolution Microevolution is change in the genetic makeup of a population from generation to generation

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6. 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

7. 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

8. LE 23-3 MAP AREA CANADA ALASKA Beaufort Sea Porcupine herd range NORTHWEST TERRITORIES Fairbanks Fortymile herd range Whitehorse ALASKA YUKON

9. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The Hardy-Weinberg Theorem The Hardy-Weinberg theorem describes a population that is not evolving It states that frequencies of alleles and genotypes in a population’s gene pool remain constant from generation to generation, provided that only Mendelian segregation and recombination of alleles are at work Mendelian inheritance p reserves genetic variation in a population

10. LE 23-4 Generation 3 25% C R C R Generation 4 50% C R C W 25% C W C W 50% C W gametes 50% C R come together at random 25% C R C R 50% C R C W 25% C W C W Alleles segregate, and subsequent generations also have three types of flowers in the same proportions gametes Generation 2 Generation 1 CRCRCRCR CWCWCWCW genotype Plants mate All C R C W (all pink flowers) 50% C R 50% C W gametes come together at random X

11. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Preservation of Allele Frequencies In a given population where gametes contribute to the next generation randomly, allele frequencies will not change

12. 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

13. 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 – p 2 + 2pq + q 2 = 1 – And p 2 and q 2 represent the frequencies of the homozygous genotypes and 2pq represents the frequency of the heterozygous genotype

14. LE 23-5 Gametes for each generation are drawn at random from the gene pool of the previous generation: 80% C R (p = 0.8) 20% C W (q = 0.2) Sperm C R (80%) C W (20%) pqp2p2 16% C R C W 64% C R Eggs C W (20%) C R (80%) 16% C R C W qp 4% C W q2q2

15. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Conditions for Hardy-Weinberg Equilibrium The Hardy-Weinberg theorem describes a hypothetical population In real populations, allele and genotype frequencies do change over time

16. 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

17. 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

18. 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

19. 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 Animation: Genetic Variation from Sexual Recombination

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21. 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

22. 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

23. 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

24. 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

25. 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

26. 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

27. 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 Animation: Causes of Evolutionary Change

28. LE 23-7 CRCRCRCR CRCRCRCR CWCWCWCW CRCRCRCR CRCWCRCW CRCRCRCR CRCWCRCW CWCWCWCW CWCWCWCW CRCWCRCW CRCWCRCW CRCRCRCR CRCWCRCW CRCWCRCW CRCRCRCR CRCRCRCR CRCWCRCW CWCWCWCW CRCWCRCW CRCRCRCR Only 5 of 10 plants leave offspring Only 2 of 10 plants leave offspring CRCRCRCR CRCRCRCR CRCRCRCR CRCRCRCR CRCRCRCR CRCRCRCR CRCRCRCR CRCRCRCR CRCRCRCR CRCRCRCR Generation 2 p = 0.5 q = 0.5 Generation 3 p = 1.0 q = 0.0 Generation 1 p (frequency of C R ) = 0.7 q (frequency of C W ) = 0.3

29. 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

30. LE 23-8 Original population Bottlenecking event Surviving population

31. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Understanding the bottleneck effect can increase understanding of how human activity affects other species

32. 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

33. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Gene Flow Gene flow consists of genetic additions or subtractions from a population, resulting from movement of fertile individuals or gametes Gene flow causes a population to gain or lose alleles It tends to reduce differences between populations over time

34. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Concept 23.4: Natural selection is the primary mechanism of adaptive evolution Natural selection accumulates and maintains favorable genotypes in a population

35. 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

36. LE 23-9 Map butterflies that emerge in spring: orange and brown Map butterflies that emerge in late summer: black and white

37. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Variation Within a Population Both discrete and quantitative characters contribute to variation within a population Discrete characters can be classified on an either- or basis Quantitative characters vary along a continuum within a population

38. 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

39. 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

40. 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

41. LE 23-10 12.43.145.1867.15 8.119.1210.1613.1719XX 1 9.1011.1213.1715.18XX 2.193.84.165.146.7

42. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Some examples of geographic variation occur as a cline, which is a graded change in a trait along a geographic axis

43. LE 23-11 Heights of yarrow plants grown in common garden Sierra Nevada Range Great Basin Plateau Seed collection sites 100 50 0 3,000 2,000 1,000 0 Mean height (cm) Altitude (m)

44. 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

45. 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

46. 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

47. 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

48. 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

49. LE 23-12 Original population Evolved population Phenotypes (fur color) Original population Directional selectionDisruptive selectionStabilizing selection Frequency of individuals

50. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The Preservation of Genetic Variation Various mechanisms help to preserve genetic variation in a population

51. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Diploidy Diploidy maintains genetic variation in the form of hidden recessive alleles

52. 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

53. 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

54. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The sickle-cell allele causes mutations in hemoglobin but also confers malaria resistance It exemplifies the heterozygote advantage

55. LE 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% Distribution of malaria caused by Plasmodium falciparum (a protozoan)

56. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Frequency-Dependent Selection In frequency-dependent selection, the fitness of any morph declines if it becomes too common in the population

57. LE 23-14 Parental population sample Experimental group sample 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). Plain backgroundPatterned background 0.6 0.5 0.4 0.3 0.2 020406080100 Generation number Frequency- independent control Phenotypic variation

58. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Neutral Variation Neutral variation is genetic variation that appears to confer no selective advantage

59. 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

60. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Intrasexual selection is competition among individuals of one sex for mates of the opposite sex

61. 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

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63. The Evolutionary Enigma of Sexual Reproduction Sexual reproduction produces fewer reproductive offspring than asexual reproduction, a so-called “reproductive handicap”

64. LE 23-16 Asexual reproduction Female Generation 1 Generation 2 Generation 3 Generation 4 Sexual reproduction Female Male

65. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Sexual reproduction produces genetic variation that may aid in disease resistance

66. 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