Chapter 23: The Evolution of Populations This chapter begins with the idea that we focused on as we closed the

last chapter: Individuals do not evolve! Populations evolve. The Overview looks at the work of Peter and Rosemary Grant with Galpagos finches to illustrate this point, and the rest of the chapter examines the change in populations over time. As in the last chapter, first read each concept to get the big picture and then go back to work on the details presented by our questions. Dont lose sight of the conceptual understanding by getting lost in the details!

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 1. What is microevolution? Microevolution is a change in allele frequencies in a population over generations 2. What are the three main mechanisms that can cause changes in allele frequency? Natural selection, genetic drift (chance events that alter allele frequency); and gene flow (transfer of alleles between populations). 3. Which is the only mechanism that is adaptive, or improves the match between organisms and their environment? Natural selection

Concept 23.1 Mutation and sexual reproduction produce the genetic variation that makes evolution possible
4. Because Darwin did not know about the work of Gregor Mendel, he could not explain how organisms pass heritable traits to their offspring. In looking at genetic variation, what are discrete characters, and what are quantitative characters?

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 5. Using the techniques of molecular biology, what are the two ways of measuring genetic variation in a population? 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 6. Geographic variation may be shown in a graded manner along a geographic axis known as a cline. What external factors might produce a cline? Why does the existence of a cline suggest natural selection? Most species exhibit geographic variation, differences between gene pools of separate populations or population subgroups Some examples of geographic variation occur as a cline, which is a graded change in a trait along a geographic axis

7. What is the ultimate source of new alleles? 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 8. Mutations are any change in the nucleotide sequence of an organisms DNA. These mutations provide the raw material from which new traits may arise and be selected. What occurs in a point mutation? A point mutation is a change in one base in a gene The effects of point mutations can vary: Mutations in noncoding regions of DNA are often harmless Mutations in a gene might not affect protein production because of redundancy in the genetic code The effects of point mutations can vary: Mutations that result in a change in protein production are often harmful Mutations that result in a change in protein production can sometimes increase the fit between organism and environment 9. What is translocation? An aberration in chromosome structure resulting from attachment of a chromosomal fragment to a non-homologous chromosome. How could it be beneficial? During protein synthesis the third stage in the elongation cycle when the RNA carrying the growing polypeptide moves from the A-site to the P-site on the ribosome. The transport of organic nutrients in the phloem of vascular plants. 10. How does gene duplication occur? How might it play a role in evolution? Chromosomal mutations that delete, disrupt, or rearrange many loci are typically harmful Duplication of large chromosome segments is usually harmful Duplication of small pieces of DNA is sometimes less harmful and increases the genome size Duplicated genes can take on new functions by further mutation 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 11. Much of the genetic variation that makes evolution possible comes through sexual reproduction. What are the three mechanisms by which sexual reproduction shuffles existing alleles? 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

Concept 23.2 The Hardy-Weinberg equation can be used to test whether a population is evolving
The first step in testing whether evolution is occurring in a population is to clarify what we mean by a population

12. What is a population? A population is a localized group of individuals capable of interbreeding and producing fertile offspring 13. What is a gene pool? A gene pool consists of all the alleles for all loci in a population 14. The greater the number of fixed alleles, the lower the species diversity. What does it mean to say that an allele is fixed? A locus is fixed if all individuals in a population are homozygous for the same allele 15. The Hardy-Weinberg principle is used to describe a population that is not evolving. What does this principle state? 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 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

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. If the frequency of alleles in a population remains constant, the population is at Hardy- Weinberg equilibrium. There are five conditions for Hardy-Weinberg equilibrium. It is very important for you to know these conditions, so enter them neatly into the box below. CONDITIONS FOR HARDY-WEINBERG EQUILIBRIUM 1. 2. 3. 4. 5. No mutations Random mating No natural selection Extremely large population size No gene flow

Natural populations can evolve at some loci, while being in HardyWeinberg equilibrium at other loci 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 -p2 (=AA) and -q2 (=aa) representing the frequencies of the homozygous genotypes; and -2pq represents the frequency of the heterozygous genotype (=Aa) Let p = the frequency of A, the dominant allele, and Let q = the frequency of a, the recessive allele

-p + q = 1 -p = 1 q -q = 1 p 17. Suppose in a plant population that red flowers (R ) are dominant to white flowers (r ). In a population of 500 individuals, 25% show the recessive phenotype. How many individuals would you expect to be homozygous dominant and heterozygous for the trait? Solution Let p = the frequency of (R ), the dominant allele, and Let q = the frequency of (r ), the recessive allele 1. q2 (= rr) representing the frequencies of the homozygous recessive = 25% = 0.25. Since q2 = 0.25, q = 0.5 2. p + q = 1, so p = 0.5 3. Homozygous dominant individuals are RR or p2 = 0.25, and they will represent (0.25)(500) = 125 individuals 4. The heterozygous individuals are calculated from 2pq = 2 (0.5)(0.5) = 0.5, and in a population of 500 individuals it will be (0.5)(500) = 250 individuals. 18. In a population of plants 64% exhibit the dominant flower color (red) and 36% of the plants have white flowers. What is the frequency of the dominant allele? (There are a couple of twists in this problem.) Solution This problem requires you to recognize that individuals with the dominant trait can be either homozygous or heterozygous. Therefore, you cannot simply take the square root of 0.64 to get p. For problems of this type you must begin with the homozygous recessive group Let p = the frequency of (R ), the dominant allele, and Let q = the frequency of (r ), the recessive allele 1. q2 = the frequency of the homozygous recessive = 36% = 0.36. Since q2 = 0.36, q = 0.6 2. p + q = 1, so p = 0.4 3. Notice this problem asks for the frequency of the dominant allele (p), not the frequency of the homozygous dominant individuals (p2). Thus, you are done; the frequency of the dominant allele = 40%.

Concept 23.3: Natural selection, genetic drift, and gene flow can alter allele frequencies in a population
19. First, lets try to summarize the BIG IDEA from this section. Scan through the entire concept to pull out this information. Three major factors alter allelic frequency and bring about evolutionary change. List each factor and provide an explanation. Factor Natural selection Genetic drift Explanation Differential success in reproduction results in certain alleles being passed to the next generation in greater proportions 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

The Founder Effect

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

The Bottleneck Effect

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 populations 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

Effects of Genetic Drift: A Summary

1.Genetic drift is significant in small populations 2.Genetic drift causes allele frequencies to change at

Gene flow

random 3.Genetic drift can lead to a loss of genetic variation within populations 4.Genetic drift can cause harmful alleles to become fixed 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 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 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

20. Which of the factors above results in a random, non-adaptive change in allelic frequencies? Genetic drift describes how allele frequencies fluctuate unpredictably from one generation to the next 21. Which of the factors above tends to reduce the genetic differences between populations and make populations more similar? Gene flow consists of the movement of alleles among populations

22. Which of the factors above results in individuals that are better suited to their environment? Natural Selection: Differential success in reproduction results in certain alleles being passed to the next generation in greater proportions 23. Explain what happens to each of these examples of genetic drift: Founder Effect: 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 Bottleneck Effect: 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 populations 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

Concept 23.4: Natural selection is the only mechanism that consistently causes adaptive evolution
24. In evolutionary terms fitness refers only to the ability to leave offspring and contribute to the gene pool of the next generation. It may have nothing to do with being big, or strong, or aggressive. Define relative fitness.

Relative Fitness

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 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 25. What is the relative fitness of a sterile mule? 0.0 Relative fitness is the contribution an individual makes to the gene pool of the next generation, relative to the contributions of other individuals 26. Figure 23.13 is important because it helps explain the three modes of selection. Label each type of selection, and fill n the chart to explain what is occurring.

Type of Selection

How It Works Stabilizing selection favors intermediate variants and acts against extreme phenotypes Directional selection favors individuals at one end of the phenotypic range

Stabilizing Directional

Disruptive

Disruptive selection favors individuals at both extremes of the phenotypic range

27. What is often the result of sexual selection? The Key Role of Natural Selection in Adaptive Evolution Sexual selection is natural selection for mating success 28. What is the difference between intrasexual selection and intrasexual selection? 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 Give an example of each type of selection. Male showiness due to mate choice can increase a males chances of attracting a female, while decreasing his chances of survival 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 29. Explain two ways in which genetic variation is preserved in a population. 1. Diploidy maintains genetic variation in the form of hidden recessive alleles 2. Balancing selection occurs when natural selection maintains stable frequencies of two or more phenotypic forms in a population 30. Discuss what is meant by heterozygous advantage, and use sicklecell anemia as an example. 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

31. Give four reasons why natural selection cannot produce perfect organisms. 1.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

Testing Your Knowledge: Self-Quiz Answers

Now you should be ready to test your knowledge. Place your answers here: 1.________ 2.________ 3.________ 4._________ 5. _________