The Evolution of Populations: Understanding Genetic Variation and the Factors that Drive Evolution

1. Chapter 23 The Evolution of Populations

2. 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

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

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. 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. 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 preserves genetic variation in a population

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

11. Preservation of Allele Frequencies • In a given population where gametes contribute to the next generation randomly, allele frequencies will not change

12. 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. 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

14. 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 (80%) CW (20%) p2 pq CR (80%) Eggs 64% CRCR 16% CRCW 4% CWCW 16% CRCW CW (20%) qp q2

15. 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. 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. 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. 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. Mutation • Mutations are changes in the nucleotide sequence of DNA • Mutations cause new genes and alleles to arise Animation: Genetic Variation from Sexual Recombination

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. 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. 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. Sexual Recombination • Sexual recombination is far more important than mutation in producing the genetic differences that make adaptation possible

25. 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. Natural Selection • Differential success in reproduction results in certain alleles being passed to the next generation in greater proportions

27. 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

28. LE 23-7 CWCW CRCR CRCR CRCR CRCR CRCW CRCW CRCR CRCR Only 5 of 10 plants leave offspring Only 2 of 10 plants leave offspring CRCR CRCR CRCR CWCW CWCW CRCR CRCW CRCW CRCR CRCR CRCR CWCW CRCR CRCW CRCR CRCR CRCR CRCW CRCW CRCR CRCW Generation 1 p (frequency of CR) = 0.7 q (frequency of CW) = 0.3 Generation 3 p = 1.0 q = 0.0 Generation 2 p = 0.5 q = 0.5

29. 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. Understanding the bottleneck effect can increase understanding of how human activity affects other species

32. 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. 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. Concept 23.4: Natural selection is the primary mechanism of adaptive evolution • Natural selection accumulates and maintains favorable genotypes in a population

35. 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. 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. 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. 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. Variation Between Populations • Most species exhibit geographic variation differences between gene pools of separate populations or population subgroups

41. LE 23-10 1 2.4 3.14 5.18 6 7.15 8.11 9.12 10.16 13.17 19 XX 1 2.19 3.8 4.16 5.14 6.7 9.10 11.12 13.17 15.18 XX

42. 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 100 Mean height (cm) 50 0 3,000 Altitude (m) 2,000 Sierra Nevada Range Great Basin Plateau 1,000 0 Seed collection sites

44. 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. 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. 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. 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. 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 Frequency of individuals Phenotypes (fur color) Original population Evolved population Directional selection Disruptive selection Stabilizing selection

50. The Preservation of Genetic Variation • Various mechanisms help to preserve genetic variation in a population