CAMPBELL

BIOLOGY TENTH
EDITION

Reece • Urry • Cain • Wasserman • Minorsky • Jackson

23
The Evolution
of Populations

© 2014 Pearson Education, Inc.

The Smallest Unit of Evolution

▪ A common misconception is that organisms evolve

during their lifetimes
▪ Natural selection acts on individuals, but only
populations evolve
▪ Consider, for example, a population of medium
ground finches on Daphne Major Island
▪ During a drought, large-beaked birds were more
likely to crack large seeds and survive
▪ The finch population evolved by natural selection

© 2014 Pearson Education, Inc.

 ▪ Microevolution is a change in allele frequencies
in a population over generations
▪ Three mechanisms cause allele frequency change
▪ Natural selection
▪ Genetic drift
▪ Gene flow
▪ Only natural selection causes adaptive evolution

© 2014 Pearson Education, Inc.

 Genetic variation makes evolution possible.
CONCEPT 23.1

© 2014 Pearson Education, Inc.

Concept 23.1: Genetic variation makes
evolution possible
▪ Variation in heritable traits is a prerequisite for
evolution
▪ Mendel’s work on pea plants provided evidence of
discrete heritable units (genes)

© 2014 Pearson Education, Inc.

Genetic Variation

▪ Genetic variation among individuals is caused by

differences in genes or other DNA segments
▪ Phenotype is the product of inherited genotype
and environmental influences
▪ Natural selection can only act on variation with a
genetic component

© 2014 Pearson Education, Inc.

 ▪ Some phenotypic differences are determined by a
single gene and can be classified on an either-or
basis
▪ Other phenotypic differences are determined by
the influence of two or more genes and vary along
a continuum within a population

© 2014 Pearson Education, Inc.

Figure 23.3

Hair color in horses-determined by multiple genes 🡺 wide variety

© 2014 Pearson Education, Inc.
Genetic Variation from Sexual Recombination

© 2014 Pearson Education, Inc.

 ▪ Genetic variation can be measured as gene
variability or nucleotide variability
▪ For gene variability, 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
▪ Nucleotide variation rarely results in phenotypic
variation

© 2014 Pearson Education, Inc.

 ▪ Some phenotypic variation does not result from
genetic differences among individuals, but rather
from environmental influences
▪ Only genetically determined variation can have
evolutionary consequences

© 2014 Pearson Education, Inc.

Figure 23.5

Nonheritable Variation
(a) (b)

Oak flower diet Oak leaf diet

© 2014 Pearson Education, Inc.

Sources of Genetic Variation

▪ New genes and alleles can arise by mutation or

gene duplication
▪ Sexual reproduction can result in genetic variation
by recombining existing alleles

© 2014 Pearson Education, Inc.

 The Hardy-Weinberg equation can be used to test whether a
population is evolving.
CONCEPT 23.2

© 2014 Pearson Education, Inc.

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

© 2014 Pearson Education, Inc.

Gene Pools and Allele Frequencies

▪ A population is a localized group of individuals

capable of interbreeding and producing fertile
offspring
▪ A gene pool consists of all the alleles for all loci in
a population
▪ A locus is fixed if all individuals in a population are
homozygous for the same allele

© 2014 Pearson Education, Inc.

 Figure 23.6 One species, two populations

CANAD
MAP

ALASK
ARE
A

A
A
Porcupine
herd range

Fortymile
herd range
Porcupine Fortymile
herd herd

© 2014 Pearson Education, Inc.

Figure 23.UN01

CR CR

Incomplete Dominance:
If there are two or more alleles
CW CW
for a locus, diploid individuals
may be either homozygous or
heterozygous
CR CW

© 2014 Pearson Education, Inc.

 ▪ The frequency of an allele in a population can be
calculated
▪ For diploid organisms, the total number of alleles at
a locus is the total number of individuals times 2
▪ The total number of dominant alleles at a locus is
two alleles for each homozygous dominant
individual plus one allele for each heterozygous
individual; the same logic applies for recessive
alleles

© 2014 Pearson Education, Inc.

 ▪ By convention, if there are two alleles at a locus, p
and q are used to represent their frequencies
▪ The frequency of all alleles in a population will add
up to 1
▪ For example, p + q = 1

© 2014 Pearson Education, Inc.

 ▪ For example, consider a population of wildflowers
that is incompletely dominant for color
▪ 320 red flowers (CRCR)
▪ 160 pink flowers (CRCW)
▪ 20 white flowers (CWCW)
▪ Calculate the number of copies of each allele
▪ CR = (320 × 2) + 160 = 800
▪ CW = (20 × 2) + 160 = 200

© 2014 Pearson Education, Inc.

 ▪ To calculate the frequency of each allele
▪ p = freq CR = 800 / (800 + 200) = 0.8
▪ q = freq CW = 200 / (800 + 200) = 0.2
▪ The sum of alleles is always 1
▪ 0.8 + 0.2 = 1

© 2014 Pearson Education, Inc.

The Hardy-Weinberg Equation

▪ The Hardy-Weinberg equation describes the

genetic makeup we expect for a population that is
not evolving at a particular locus
▪ If the observed genetic makeup of the population
differs from expectations under Hardy-Weinberg, it
suggests that the population may be evolving

© 2014 Pearson Education, Inc.

Hardy-Weinberg Equilibrium

▪ In a population where gametes contribute to the

next generation randomly and Mendelian
inheritance occurs, allele and genotype
frequencies remain constant from generation to
generation
▪ Such a population is in Hardy-Weinberg
equilibrium

© 2014 Pearson Education, Inc.

 Frequencies of alleles
p = frequency of CR allele =
0.8
q = frequency of CW allele =
0.2

Figure 23.7 Alleles in the population

Selecting alleles
at random from
a gene pool

Gametes produced
Each egg: Each
sperm:

80% 20% 80% 20%

chance chance chance chance
© 2014 Pearson Education, Inc.
 ▪ Hardy-Weinberg equilibrium describes the
constant frequency of alleles in such a gene pool
▪ Consider, for example, the same population of 500
wildflowers and 1,000 alleles where
▪ p = freq CR = 0.8
▪ q = freq CW = 0.2

© 2014 Pearson Education, Inc.

 ▪ The frequency of genotypes can be calculated
▪ CRCR = p2 = (0.8)2 = 0.64
▪ CRCW = 2pq = 2(0.8)(0.2) = 0.32
▪ CWCW = q2 = (0.2)2 = 0.04
▪ The frequency of genotypes can be confirmed
using a Punnett square

© 2014 Pearson Education, Inc.

Figure 23.8
80% CR (p = 0.8) 20% CW (q = 0.2)

Sperm
CR p = 0.8 CW q = 0.2

CR
p = 0.8
0.64 (p2) 0.16 (pq)
Eggs CRCR CRCW

CW 0.16 (qp) 0.04 (q2)

q = 0.2 CRCW CWCW

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

Gametes of this generation:

64% CR R
+ 16% C R W = 80% CR = 0.8 =
(from CRCR plants) (from C C plants) p
4% CW W
W W + 16% C R W = 20% CW = 0.2 =
(from C C plants) (from C C plants) q
With random mating, these gametes will result in the same
mix of genotypes in the next generation:

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

© 2014 Pearson Education, Inc.
 ▪ 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
▪ where p2 and q2 represent the frequencies of the
homozygous genotypes and 2pq represents the
frequency of the heterozygous genotype

How to Solve Hardy Weinberg Problems

© 2014 Pearson Education, Inc.

Conditions for Hardy-Weinberg Equilibrium

▪ The Hardy-Weinberg theorem describes a

hypothetical population that is not evolving
▪ In real populations, allele and genotype
frequencies do change over time

© 2014 Pearson Education, Inc.

 ▪ The five conditions for non-evolving populations
are rarely met in nature
1. No mutations
2. Random mating
3. No natural selection
4. Extremely large population size
5. No gene flow
*Natural populations can evolve at some loci, while
being in Hardy-Weinberg equilibrium at other loci*

© 2014 Pearson Education, Inc.

Applying the Hardy-Weinberg Equation
▪ We can assume the locus that causes
phenylketonuria (PKU) is in Hardy-Weinberg
equilibrium given that
1. The PKU gene mutation rate is low
2. Mate selection is random with respect to whether
or not an individual is a carrier for the PKU allele
3. Natural selection can only act on rare
homozygous individuals who do not follow dietary
restrictions
4. The population is large
5. Migration has no effect as many other populations
have similar allele frequencies
© 2014 Pearson Education, Inc.
 ▪ The occurrence of PKU is 1 per 10,000 births
▪ q2 = 0.0001
▪ q = 0.01
▪ The frequency of normal alleles is
▪ p = 1 − q = 1 − 0.01 = 0.99
▪ The frequency of carriers is
▪ 2pq = 2 × 0.99 × 0.01 = 0.0198
▪ or approximately 2% of the U.S. population

© 2014 Pearson Education, Inc.

 Natural selection, genetic drift and gene flow can alter allele
frequencies in a population.
CONCEPT 23.3

© 2014 Pearson Education, Inc.

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

© 2014 Pearson Education, Inc.

Natural Selection

▪ Differential success in reproduction results in

certain alleles being passed to the next generation
in greater proportions
▪ For example, an allele that confers resistance to
DDT in fruit flies increased in frequency after DDT
was used widely in agriculture

© 2014 Pearson Education, Inc.

 ▪ Natural selection can cause adaptive evolution,
an improvement in the match between organisms
and their environment

© 2014 Pearson Education, Inc.

Natural Selection

© 2014 Pearson Education, Inc.

Genetic Drift

▪ The smaller a sample, the greater the chance of

random 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

© 2014 Pearson Education, Inc.

Figure 23.9–3

5 plants 2 plants
leave leave
offspring offspring
CRCR CRCW CRCR
CWCW
CRCR CRCW CRCR
CRCR
R R
CRCR
C C R R
C C
R R
CRCW
C C CRCR
CWCW
W
C C W CRCR CRCR
R
C C W W
C C W CRCR
CRCR
R W
C C
CRCW CRCR
R R
C C CRCR
R W R W
C C C C CRCR

Generation 1 Generation 2 Generation 3

p (frequency of CR) = 0.7 p = 0.5 p = 1.0
q (frequency of CW) = 0.3 q = 0.5 q = 0.0

© 2014 Pearson Education, Inc.

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

© 2014 Pearson Education, Inc.

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 population’s gene pool
▪ If the population remains small, it may be further
affected by genetic drift

© 2014 Pearson Education, Inc.

Figure 23.10–3

Original Bottlenecking Surviving

population event population

© 2014 Pearson Education, Inc.

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

© 2014 Pearson Education, Inc.

 Case Study: Impact of Genetic Drift on the
Greater Prairie Chicken
▪ Loss of prairie habitat caused a severe reduction
in the population of greater prairie chickens in
Illinois
▪ The surviving birds had low levels of genetic
variation, and only 50% of their eggs hatched

© 2014 Pearson Education, Inc.

Figure 23.11
Pre-bottleneck Post-bottlenec
(Illinois, 1820) k
(Illinois, 1993)

Greater prairie
chicken Range
of greater
prairie
(a) chicken

Number Percenta
Location Population
of alleles ge
size
per of eggs
locus hatched
Illinois
1930–1960s 1,000–25,000 5.2 93
1993 <50 3.7 <50

Kansas, 1998 750,000 5.8 99

(no
bottleneck)
Nebraska, 1998 75,000– 5.8 96
(no 200,000
bottleneck)
(b)
© 2014 Pearson Education, Inc.
 ▪ Researchers used DNA from museum specimens
to compare genetic variation in the population
before and after the bottleneck
▪ The results showed a loss of alleles at several loci
▪ Researchers introduced greater prairie chickens
from populations in other states and were
successful in introducing new alleles and
increasing the egg hatch rate to 90%

© 2014 Pearson Education, Inc.

Effects of Genetic Drift: A Summary

1. Genetic drift is significant in small populations

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

© 2014 Pearson Education, Inc.

Gene Flow

▪ 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 variation among
populations over time

© 2014 Pearson Education, Inc.

 ▪ Gene flow can decrease the fitness of a
population

▪ Consider, for example, the great tit (Parus

major) on the Dutch island of Vlieland
(Netherlands)
▪ Mating causes gene flow between the
central and eastern populations

▪ Immigration from the mainland introduces

alleles that decrease fitness on the island

▪ Natural selection removes alleles that

decrease fitness

▪ Birds born in the central region with high

immigration have a lower fitness; birds born
in the east with low immigration have a
higher fitness

© 2014 Pearson Education, Inc.

Figure 23.12

Central
population

NORTH SEA Eastern

Vlieland, population
N the Netherlands
Parus major 2 km

50 Population in which
the surviving
females
40
Survival rate (%)

eventually
Centralbred
Eastern
30

20

10

0
Females born in Females born in
central population eastern
population
© 2014 Pearson Education, Inc.
 ▪ Gene flow can increase the fitness of a population
▪ Consider, for example, the spread of alleles for
resistance to insecticides
▪ 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
▪ Gene flow is an important agent of evolutionary
change in modern human populations

© 2014 Pearson Education, Inc.

Mechanisms of Evolution: genetic drift

▪ [Link]
Y

© 2014 Pearson Education, Inc.

 Natural selection is the only mechanism that consistently causes
adaptive evolution.
CONCEPT 23.4

© 2014 Pearson Education, Inc.

Concept 23.4: Natural selection is the only
mechanism that consistently causes adaptive
evolution
▪ Evolution by natural selection involves both
chance and “sorting”
▪ New genetic variations arise by chance
▪ Beneficial alleles are “sorted” and favored by
natural selection
▪ Only natural selection consistently increases the
frequencies of alleles that provide reproductive
advantage

© 2014 Pearson Education, Inc.

Natural Selection: A Closer Look

▪ Natural selection brings about adaptive evolution

by acting on an organism’s phenotype

© 2014 Pearson Education, Inc.

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

© 2014 Pearson Education, Inc.

 ▪ 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 individuals

© 2014 Pearson Education, Inc.

Directional, Disruptive, and Stabilizing Selection

▪ There are three modes of selection

▪ Directional selection favors individuals at one
extreme 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

© 2014 Pearson Education, Inc.

Figure 23.13

Original
population

Frequency of
individuals
Original Evolved Phenotypes (fur color)
population population

(a) Directional selection (b) Disruptive selection (c) Stabilizing selection

© 2014 Pearson Education, Inc.

The Key Role of Natural Selection in Adaptive
Evolution
▪ Striking adaptations have
arisen by natural selection
▪ For example, certain
octopuses can change color
rapidly for camouflage
▪ For example, the jaws of
snakes allow them to
swallow prey larger than
their heads

© 2014 Pearson Education, Inc.

 ▪ Natural selection increases the frequencies of
alleles that enhance survival and reproduction
▪ Adaptive evolution occurs as the match between a
species and its environment increases
▪ Because the environment can change, adaptive
evolution is a continuous process
▪ Genetic drift and gene flow do not consistently
lead to adaptive evolution as they can increase or
decrease the match between an organism and its
environment

© 2014 Pearson Education, Inc.

Sexual Selection

▪ Sexual selection is natural selection for mating

success
▪ It can result in sexual dimorphism, marked
differences between the sexes in secondary
sexual characteristics

© 2014 Pearson Education, Inc.

 ▪ Intrasexual selection is
direct 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 Intrasexual Competition Red Deer
▪ Male showiness due to mate
choice can increase a male’s
chances of attracting a
female, while decreasing his
chances of survival

© 2014 Pearson Education, Inc.

Intersexual- Bird of Paradise
 ▪ How do female preferences evolve?
▪ The “good genes” hypothesis suggests that if a
trait is related to male genetic quality or health,
both the male trait and female preference for that
trait should increase in frequency

© 2014 Pearson Education, Inc.

Balancing Selection

▪ Diploidy maintains genetic variation in the form of

recessive alleles hidden from selection in
heterozygotes
▪ Balancing selection occurs when natural
selection maintains stable frequencies of two or
more phenotypic forms in a population
▪ Balancing selection includes
▪ Heterozygote advantage
▪ Frequency-dependent selection

© 2014 Pearson Education, Inc.

Heterozygote Advantage

▪ 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
▪ Heterozygote advantage can result from stabilizing
or directional selection

© 2014 Pearson Education, Inc.

 ▪ A mutation in an allele that codes for part of the
hemoglobin protein causes sickle-cell disease, but
also confers malaria resistance
▪ In regions where the malaria parasite is common,
selection favors individuals heterozygous for the
sickle-cell allele

© 2014 Pearson Education, Inc.

Frequency-Dependent Selection

▪ In frequency-dependent selection, the fitness of

a phenotype declines if it becomes too common in
the population
▪ Selection favors whichever phenotype is less
common in a population
▪ For example, frequency-dependent selection
results in approximately equal numbers of
“right-mouthed” and “left-mouthed” scale-eating
fish

© 2014 Pearson Education, Inc.

Figure 23.18

“Left-mouthed”
P. microlepis

1.0
“Right-mouthe
d”
P. microlepis
“left-mouthed”
Frequency of

individuals

0.5

0
1981 ’82 ’83 ’84 ’85 ’86 ’87 ’88 ’89 ’90
Sample year
© 2014 Pearson Education, Inc.
Why Natural Selection Cannot Fashion 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

© 2014 Pearson Education, Inc.