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    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 23Chapter 23

    The Evolution of Populations

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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 apopulation 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

    Whitehorse

    Fortymile

    herd range

    Figure 23.3

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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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    Preservation of Allele Frequencies

    In a given population where gametes contribute

    to the next generation randomly, allelefrequencies will not change

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    Hardy-WeinbergEquilibrium

    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

    Figure 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)

    SpermCR

    (80%)CW

    (20%)

    p2

    64%

    CRCR16%

    CRCW

    16%

    CRCW4%

    CWCWqp

    CR

    (80%)

    Eggs

    CW

    (20%)

    pq

    If the gametes come together at random, the genotype

    frequencies of this generation are in Hardy-Weinberg equilibrium:

    q2

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

    Gametes of the next generation:

    64% CR from

    CRCRhomozygotes

    16% CR from

    CRCWhomozygotes+ = 80% CR= 0.8 =p

    16% CWfrom

    CRCWheterozygotes+ = 20% CW= 0.2 = q

    With random mating, these gametes will result in the same

    mix of plants in the next generation:

    64% CRCR, 32% CRCWand 4% CWCWplants

    p2

    4% CWfrom

    CWCWhomozygotes

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    Ifp and q represent the relative frequencies of

    the only two possible alleles in a population ata particular locus, then

    p2 + 2pq + q2 = 1

    Andp2 and q2represent the frequencies of the

    homozygous genotypes and 2pq represents

    the frequency of the heterozygous genotype

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    Conditions forHardy-WeinbergEquilibrium

    The Hardy-Weinberg theorem

    Describes a hypothetical population

    In real populations

    Allele and genotype frequencies do changeover 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 andHuman 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 thatmakes 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 adaptiveimpact

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    Mutations That AlterGene NumberorSequence

    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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    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

    Figure 23.7

    CRCR

    CRCW

    CRCR

    CWCW CRCR

    CRCW

    CRCW

    CRCWCRCR

    CRCR

    Only 5 of

    10 plants

    leave

    offspring

    CWCW CRCR

    CRCW

    CRCR CWCW

    CRCW

    CWCW CRCR

    CRCW CRCW

    Only 2 of

    10 plants

    leave

    offspring

    CRCR

    CRCR CRCR

    CRCRCRCR

    CRCR

    CRCR

    CRCR

    CRCRCRCR

    Generation 2

    p = 0.5

    q = 0.5

    Generation 3

    p = 1.0q = 0.0

    Generation 1

    p (frequency ofCR) = 0.7q (frequency ofCW) = 0.3

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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

    Original

    population

    Bottlenecking

    eventSurviving

    population

    Figure 23.8 A

    (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.

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    Understanding the bottleneck effect

    Can increase understanding of how human

    activity affects other species

    Figure 23.8 B

    (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.

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    The FounderEffect

    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

    Figure 23.9 A, B

    (a) Map butterflies that

    emerge in spring:

    orange and brown

    (b) Map butterflies that

    emerge in late summer:

    black and white

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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

    XX1913.1710.169.128.11

    1 2.19 3.8 4.16 5.14 6.7

    XX15.1813.1711.129.10Figure 23.10

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    Some examples of geographic variation occur

    as a cline, which is a graded change in a traitalong a geographic axis

    Figure 23.11

    EXPERIMENT Researchers observed that the average size

    of yarrow plants (Achillea) growing on the slopes of the Sierra

    Nevada mountains gradually decreases with increasing

    elevation. To eliminate the effect of environmental differencesat different elevations, researchers collected seeds

    from various altitudes and planted them in a commongarden. They then measured the heights of the

    resulting plants.

    RESULTS The average plant sizes in the commongarden were inversely correlated with the altitudes at

    which the seeds were collected, although the heightdifferences were less than in the plants natural

    environments.

    CONCLUSION The lesser but still measurable clinal variationin yarrow plants grown at a common elevation demonstrates the

    role of genetic as well as environmental differences.

    Meanheight(cm)

    Atitude(m)

    Heights of yarrow plants grown in common garden

    Seed collection sites

    Sierra NevadaRange

    Great BasinPlateau

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    A Closer Look at Natural Selection

    From the range ofvariations available in a

    population

    Natural selection increases the frequencies of

    certain genotypes, fitting organisms to their

    environment o

    ver generations

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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 StabilizingSelection

    Selection

    Favors certain genotypes by acting on the

    phenotypes of certain organisms

    Three modes of selection are

    Directional

    Disruptive

    Stabilizing

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    The three modes of selection

    Fig 23.12 AC

    (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.

    Phenotypes (fur color)

    Original population

    Originalpopulation

    Evolved

    population

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    The Preservation ofGenetic 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

    Figure 23.13

    Frequencies of thesickle-cell allele

    02.5%

    2.55.0%

    5.07.5%

    7.510.0%

    10.012.5%

    >12.5%

    Distribution ofmalaria caused by

    Plasmodium falciparum

    (a protozoan)

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    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

    Phenotypicd

    iversity

    Figure 23.14

    Parental population sample

    Experimental group sample

    Plain background Patterned background

    On pecking a moth image

    the blue jay receives afood 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).

    0.06

    0.05

    0.04

    0.03

    0.02

    0 20 40 60 80 100Generation number

    Frequency-

    independent control

    N l V i i

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    NeutralVariation

    Neutral variation

    Is genetic variation that appears to confer no

    selective advantage

    S l S l ti

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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 malesappearance

    Figure 23.15

    Th E l ti E i f S l R d ti

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    The Evolutionary Enigma of Sexual Reproduction

    Sexual reproduction

    Produces fewer reproductive offspring than asexual

    reproduction, a so-called reproductive handicap

    Figure 23.16

    Asexual reproduction

    Female

    Sexual reproduction

    Female

    Male

    Generation 1

    Generation 2

    Generation 3

    Generation 4

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    If sexual reproduction is a handicap, why has it

    persisted?

    It produces genetic variation that may aid in

    disease resistance

    Why Natural Selection Cannot Fashion Perfect Organisms

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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