23 - Evolutionofpopulations Text

Chapter 23
The Evolution of Populations
PowerPoint Lectures for

Biology, Seventh Edition
Neil Campbell and Jane Reece
Lectures by Chris Romero

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings
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
Genetic variations in populations

Contribute to evolution
Figure 23.1
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
The Modern Synthesis

Population genetics
Is the study of how populations change
genetically over time
Reconciled Darwins and Mendels ideas
The modern synthesis

Integrates Mendelian genetics with the
Darwinian theory of evolution by natural
selection
Focuses on populations as units of evolution
Gene Pools and Allele Frequencies

A population
MAP
AREA
CANADA
ALASKA
Is a localized group of individuals that are capable of

interbreeding and producing fertile offspring
Beaufort Sea
Porcupine
herd range
N
TE OR
RR TH
IT W E
O S
RI T
ES
Fortymile
herd range
ALASKA
YUKON
Fairbanks
Whitehorse
Figure 23.3
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
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
Mendelian inheritance
Preserves genetic variation in a population
Generation
1
CWCW
CRCR
genotype
genotype
Plants mate
Generation
2
All CRCW
(all pink flowers)
50% CR
gametes
50% CW
gametes
Come together at random
Generation
3
25% CRCR
50% CRCW
50% CR
gametes
25% CWCW
50% CW
gametes
Come together at random
Generation
4
25% CRCR
Figure 23.4
50% CRCW
25% CWCW
Alleles segregate, and subsequent

generations also have three types
of flowers in the same proportions
Preservation of Allele Frequencies

In a given population where gametes contribute
to the next generation randomly, allele
frequencies will not change
Hardy-Weinberg Equilibrium
Hardy-Weinberg equilibrium
Describes a population in which random
mating occurs
Describes a population where allele
frequencies do not change
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%)
pq
CR
(80%)
p2
64%
CRCR
CW
(20%)
Eggs
p2
16%
CRCW
16%
CRCW
qp
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
+
CRCW homozygotes
CRCR homozygotes
4% CW from
CW CW homozygotes
16% CW from
CRCW heterozygotes
80% CR = 0.8 = p
20% CW = 0.2 = q
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
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
Conditions for Hardy-Weinberg Equilibrium

The Hardy-Weinberg theorem
Describes a hypothetical population
In real populations
Allele and genotype frequencies do change
over time
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
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
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
Mutation
Mutations
Are changes in the nucleotide sequence of DNA
Cause new genes and alleles to arise
Figure 23.6
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
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
Gene duplication
Duplicates chromosome segments
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
Sexual Recombination
In sexually reproducing populations, sexual
recombination
Is far more important than mutation in
producing the genetic differences that make
adaptation possible
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
Natural Selection
Differential success in reproduction
Results in certain alleles being passed to the
next generation in greater proportions
Genetic Drift
Statistically, the smaller a sample
The greater the chance of deviation from a
predicted result
Genetic drift
Describes how allele frequencies can fluctuate
unpredictably from one generation to the next
Tends to reduce genetic variation
CWCW
CRCR
CRCR
Only 5 of
10 plants
leave
offspring
CRCW
CWCW
CRCR
CRCR
CRCW
CWCW
CRCR
CRCW
CRCW
CRCR
CWCW
CRCW
CRCR
CRCR
CRCW
Generation 1
p (frequency of CR) = 0.7
q (frequency of CW) = 0.3
CRCR
CRCR
CRCR
CRCR
CRCR
CRCR
CRCR
CRCR
CRCW
CRCW
Generation 2
p = 0.5
q = 0.5
Figure 23.7
Only 2 of
10 plants
leave
offspring
CRCR
CRCR
Generation 3
p = 1.0
q = 0.0
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
Figure 23.8 A
Bottlenecking
event
Surviving
population
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
The Founder Effect

The founder effect
Occurs when a few individuals become
isolated from a larger population
Can affect allele frequencies in a population
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
Concept 23.4: Natural selection is the primary

mechanism of adaptive evolution
Natural selection
Accumulates and maintains favorable
genotypes in a population
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
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
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
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
Variation Between Populations

Most species exhibit geographic variation
Differences between gene pools of separate
populations or population subgroups
Figure 23.10
2.4
3.14
5.18
8.11
9.12
10.16
13.17
2.19
3.8
4.16
9.10
11.12
13.17
15.18
7.15
19
XX
5.14
6.7
XX
Some examples of geographic variation occur

as a cline, which is a graded change in a trait
along a geographic axis
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 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.
Mean height (cm)
Heights of yarrow plants grown in common garden
Atitude (m)
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.
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.
Figure 23.11
Sierra Nevada
Range
Great Basin
Plateau
Seed collection sites
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
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
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
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
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
Frequency of individuals
The three modes of selection
Original
population
Original population
Phenotypes (fur color)
Evolved
population
(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.
Fig 23.12 AC
(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.
The Preservation of Genetic Variation

Various mechanisms help to preserve genetic
variation in a population
Diploidy
Diploidy
Maintains genetic variation in the form of
hidden recessive alleles
Balancing 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
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
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
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
An example of frequency-dependent selection

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).
Parental population sample
Experimental group sample

Phenotypic diversity
0.06
0.05
0.04
Frequencyindependent control
0.03
0.02
0
Plain background
Patterned background
Figure 23.14
20
60
40
80
Generation number
100
Neutral Variation
Neutral variation
Is genetic variation that appears to confer no
selective advantage
Sexual Selection
Sexual selection
Is natural selection for mating success
Can result in sexual dimorphism, marked
differences between the sexes in secondary
sexual characteristics
Intrasexual selection
Is a direct competition among individuals of
one sex for mates of the opposite sex
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
The Evolutionary Enigma of Sexual Reproduction

Sexual reproduction
Produces fewer reproductive offspring than asexual
reproduction, a so-called reproductive handicap
Sexual reproduction
Asexual reproduction
Generation 1
Female
Female
Generation 2
Male
Generation 3
Generation 4
Figure 23.16
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
Evolution is limited by historical constraints

Adaptations are often compromises
Chance and natural selection interact

Selection can only edit existing variations

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Chapter 23

The Evolution of Populations

PowerPoint Lectures for

Lectures by Chris Romero

Overview: The Smallest Unit of Evolution

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Genetic variations in populations

Concept 23.1: Population genetics provides a

The Modern Synthesis

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

The modern synthesis

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Gene Pools and Allele Frequencies

Is a localized group of individuals that are capable of

The gene pool

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

The Hardy-Weinberg Theorem

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Come together at random

Come together at random

Alleles segregate, and subsequent

Preservation of Allele Frequencies

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

A population in Hardy-Weinberg equilibrium

Gametes of the next generation:

With random mating, these gametes will result in the same

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

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

If p and q represent the relative frequencies of

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Conditions for Hardy-Weinberg Equilibrium

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

The five conditions for non-evolving

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Population Genetics and Human Health

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Concept 23.2: Mutation and sexual

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Cause new genes and alleles to arise

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Mutations That Alter Gene Number or Sequence

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Concept 23.3: Natural selection, genetic drift,

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

The Bottleneck Effect

(a) Shaking just a few marbles through the

Understanding the bottleneck effect

(b) Similarly, bottlenecking a population

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

The Founder Effect

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings