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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


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Overview: The Smallest Unit of Evolution One common misconception about evolution is that individual organisms evolve, in the Darwinian sense, during their lifetimes Natural selection acts on individuals, but populations evolve

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Genetic variations in populations


Contribute to evolution

Figure 23.1
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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

Figure 23.2
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The Modern Synthesis Population genetics


Is the study of how populations change genetically over time Reconciled Darwins and Mendels ideas

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The modern synthesis


Integrates Mendelian genetics with the Darwinian theory of evolution by natural selection Focuses on populations as units of evolution

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Gene Pools and Allele Frequencies A population


Is a localized group of individuals that are capable of interbreeding and producing fertile offspring
MAP AREA

Fairbanks

Fortymile herd range

Whitehorse

Figure 23.3
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The gene pool


Is the total aggregate of genes in a population at any one time Consists of all gene loci in all individuals of the population

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The Hardy-Weinberg Theorem The Hardy-Weinberg theorem


Describes a population that is not evolving States that the frequencies of alleles and genotypes in a populations gene pool remain constant from generation to generation provided that only Mendelian segregation and recombination of alleles are at work

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Mendelian inheritance
Preserves genetic variation in a population
Generation 1 CWCW CRCR genotype genotype Plants mate

Generation 2 All CRCW (all pink flowers)

50% CW gametes Come together at random

50% CR gametes

Generation 3 25% CRCR

50% CRCW

25% CWCW

50% CR gametes

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

Figure 23.4
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Preservation of Allele Frequencies In a given population where gametes contribute to the next generation randomly, allele frequencies will not change

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Hardy-Weinberg Equilibrium Hardy-Weinberg equilibrium


Describes a population in which random mating occurs Describes a population where allele frequencies do not change

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A population in Hardy-Weinberg equilibrium


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
CR (80%)

pq

Eggs

p2 64% CRCR CW (20%)

16% CRCW

qp

16% CRCW

4% CWCW

q2 If the gametes come together at random, the genotype frequencies of this generation are in Hardy-Weinberg equilibrium: 64% CRCR, 32% CRCW, and 4% CWCW

Gametes of the next generation: 16% CR from 64% CR from + RCW homozygotes RCR homozygotes C C 4% CW from CWCW homozygotes

= =

80% CR = 0.8 = p 20% CW = 0.2 = q

16% CW from CRCW heterozygotes

With random mating, these gametes will result in the same mix of plants in the next generation:

Figure 23.5

64% CRCR, 32% CRCW and 4% CWCW plants

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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

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Conditions for Hardy-Weinberg Equilibrium The Hardy-Weinberg theorem


Describes a hypothetical population

In real populations
Allele and genotype frequencies do change over time

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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

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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

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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

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Mutation Mutations
Are changes in the nucleotide sequence of DNA

Cause new genes and alleles to arise

Figure 23.6
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Point Mutations A point mutation


Is a change in one base in a gene Can have a significant impact on phenotype Is usually harmless, but may have an adaptive impact

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Mutations That Alter Gene Number or Sequence Chromosomal mutations that affect many loci
Are almost certain to be harmful May be neutral and even beneficial

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Gene duplication
Duplicates chromosome segments

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Mutation Rates Mutation rates


Tend to be low in animals and plants Average about one mutation in every 100,000 genes per generation Are more rapid in microorganisms

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Sexual Recombination In sexually reproducing populations, sexual recombination


Is far more important than mutation in producing the genetic differences that make adaptation possible

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Concept 23.3: Natural selection, genetic drift, and gene flow can alter a populations genetic composition Three major factors alter allele frequencies and bring about most evolutionary change
Natural selection Genetic drift Gene flow

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Natural Selection Differential success in reproduction


Results in certain alleles being passed to the next generation in greater proportions

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Genetic Drift Statistically, the smaller a sample


The greater the chance of deviation from a predicted result

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Genetic drift
Describes how allele frequencies can fluctuate unpredictably from one generation to the next Tends to reduce genetic variation
CRCR CRCW CRCR Only 5 of 10 plants leave offspring CWCW CRCW CRCR Only 2 of 10 plants leave offspring CRCR CRCR CRCR

CWCW CRCW CRCR CRCR

CRCR

CRCR CRCW

CWCW

CRCR CRCR

CRCR

CRCR CRCR

CRCW CRCW CRCW

CWCW

CRCR CRCW CRCR

CRCR

Generation 1 p (frequency of CR) = 0.7 q (frequency of CW) = 0.3

Generation 2 p = 0.5 q = 0.5

Generation 3 p = 1.0 q = 0.0

Figure 23.7
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The Bottleneck Effect In the bottleneck effect


A sudden change in the environment may drastically reduce the size of a population The gene pool may no longer be reflective of the original populations gene pool

(a) Shaking just a few marbles through the narrow neck of a bottle is analogous to a drastic reduction in the size of a population after some environmental disaster. By chance, blue marbles are over-represented in the new population and gold marbles are absent.

Original population

Bottlenecking event

Surviving population

Figure 23.8 A
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Understanding the bottleneck effect


Can increase understanding of how human activity affects other species

(b) Similarly, bottlenecking a population of organisms tends to reduce genetic variation, as in these northern elephant seals in California that were once hunted nearly to extinction.

Figure 23.8 B

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The Founder Effect The founder effect


Occurs when a few individuals become isolated from a larger population Can affect allele frequencies in a population

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Gene Flow Gene flow


Causes a population to gain or lose alleles Results from the movement of fertile individuals or gametes Tends to reduce differences between populations over time

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Concept 23.4: Natural selection is the primary mechanism of adaptive evolution Natural selection
Accumulates and maintains favorable genotypes in a population

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Genetic Variation Genetic variation


Occurs in individuals in populations of all species Is not always heritable

(a) Map butterflies that emerge in spring: orange and brown

(b) Map butterflies that emerge in late summer: black and white

Figure 23.9 A, B
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Variation Within a Population Both discrete and quantitative characters


Contribute to variation within a population

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Discrete characters
Can be classified on an either-or basis

Quantitative characters
Vary along a continuum within a population

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Polymorphism Phenotypic polymorphism


Describes a population in which two or more distinct morphs for a character are each 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
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Measuring Genetic Variation Population geneticists


Measure the number of polymorphisms in a population by determining the amount of heterozygosity at the gene level and the molecular level

Average heterozygosity
Measures the average percent of loci that are heterozygous in a population
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Variation Between Populations Most species exhibit geographic variation


Differences between gene pools of separate populations or population subgroups
1 2.4 3.14 5.18 6 7.15

8.11

9.12

10.16

13.17

19

XX

2.19

3.8

4.16

5.14

6.7

Figure 23.10

9.10

11.12

13.17

15.18

XX

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Some examples of geographic variation occur as a cline, which is a graded change in a trait along a geographic axis
Heights of yarrow plants grown in common garden EXPERIMENT Researchers observed that the average size Mean height (cm) Atitude (m) of yarrow plants (Achillea) growing on the slopes of the Sierra Nevada mountains gradually decreases with increasing elevation. To eliminate the effect of environmental differences at different elevations, researchers collected seeds from various altitudes and planted them in a common garden. They then measured the heights of the resulting plants.

RESULTS The average plant sizes in the common garden were inversely correlated with the altitudes at which the seeds were collected, although the height differences were less than in the plants natural environments.

Sierra Nevada Range

Great Basin Plateau

CONCLUSION The lesser but still measurable clinal variation in yarrow plants grown at a common elevation demonstrates the role of genetic as well as environmental differences.

Seed collection sites

Figure 23.11
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A Closer Look at Natural Selection From the range of variations available in a population
Natural selection increases the frequencies of certain genotypes, fitting organisms to their environment over generations

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Evolutionary Fitness The phrases struggle for existence and survival of the fittest
Are commonly used to describe natural selection Can be misleading

Reproductive success
Is generally more subtle and depends on many factors

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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 as compared to the contributions of alternative genotypes for the same locus

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Directional, Disruptive, and Stabilizing Selection Selection


Favors certain genotypes by acting on the phenotypes of certain organisms

Three modes of selection are


Directional Disruptive Stabilizing

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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
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The three modes of selection


Original population

Original population

Evolved population

Phenotypes (fur color)

(a) Directional selection shifts the overall makeup of the population by favoring variants at one extreme of the distribution. In this case, darker mice are favored because they live among dark rocks and a darker fur color conceals them from predators.

(b) Disruptive selection favors variants at both ends of the distribution. These mice have colonized a patchy habitat made up of light and dark rocks, with the result that mice of an intermediate color are at a disadvantage.

(c) Stabilizing selection removes extreme variants from the population and preserves intermediate types. If the environment consists of rocks of an intermediate color, both light and dark mice will be selected against.

Fig 23.12 AC
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The Preservation of Genetic Variation Various mechanisms help to preserve genetic variation in a population

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Diploidy Diploidy
Maintains genetic variation in the form of hidden recessive alleles

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alancing Selection Balancing selection


Occurs when natural selection maintains stable frequencies of two or more phenotypic forms in a population Leads to a state called balanced polymorphism

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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

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The sickle-cell allele


Causes mutations in hemoglobin but also confers malaria resistance Exemplifies the heterozygote advantage

Distribution of malaria caused by Plasmodium falciparum (a protozoan)

Figure 23.13
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Frequencies of the sickle-cell allele 02.5% 2.55.0% 5.07.5% 7.510.0% 10.012.5% >12.5%

Frequency-Dependent Selection In frequency-dependent selection


The fitness of any morph declines if it becomes too common in the population

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An example of frequency-dependent selection


Parental population 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 to a new set of images (a new feeding opportunity).

Experimental group sample


Phenotypic diversity

0.06 0.05 0.04 0.03 0.02 0 20 Frequencyindependent control

Plain background

Patterned background

60 40 80 Generation number

100

Figure 23.14
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Neutral Variation Neutral variation


Is genetic variation that appears to confer no selective advantage

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Sexual Selection Sexual selection


Is natural selection for mating success Can result in sexual dimorphism, marked differences between the sexes in secondary sexual characteristics

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Intrasexual selection
Is a direct competition among individuals of one sex for mates of the opposite sex

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Intersexual selection
Occurs when individuals of one sex (usually females) are choosy in selecting their mates from individuals of the other sex May depend on the showiness of the males appearance

Figure 23.15
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The Evolutionary Enigma of Sexual Reproduction Sexual reproduction


Produces fewer reproductive offspring than asexual reproduction, a so-called reproductive handicap
Asexual reproduction Female Generation 1 Female Sexual reproduction

Generation 2

Male

Generation 3

Generation 4

Figure 23.16
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If sexual reproduction is a handicap, why has it persisted?


It produces genetic variation that may aid in disease resistance

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Why Natural Selection Cannot Fashion Perfect Organisms

Evolution is limited by historical constraints Adaptations are often compromises

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Chance and natural selection interact Selection can only edit existing variations

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