Chapter 23 (or 7): Population Genetics:

The Evolution of Populations

Oakton Community College - BIO 122

Dr. Paul Gulezian

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
Microevolution is a change in allele
frequencies in a population over generations
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Concept 23.1: Mutation and sexual reproduction

produce the genetic variation that makes evolution
possible
Two processes, mutation and sexual
reproduction, produce the variation in gene
pools that contributes to differences among
individuals

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Genetic Variation
Variation in individual genotype leads to
variation in individual phenotype
Not all phenotypic variation is heritable
Natural selection can only act on variation with
a genetic component for evolution to proceed

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Fig. 23-2

Non-Heritable Variation
(a)

(b)

Fig. 23-2a

(a)

Fig. 23-2b

(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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

 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
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Variation Between
Populations
Most species exhibit geographic variation,
differences between gene pools of separate
populations or population subgroups

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Fig. 23-3

2.4

8.11

9.12

3.14

5.18

10.16 13.17

7.15

19

XX

2.19

3.8

4.16 5.14

9.10 11.12 13.17 15.18

6.7
XX

 Some examples of geographic variation occur

as a cline, which is a graded change in a trait
along a geographic axis

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Fig. 23-4

0.8
0.6
0.4

LdhB b allele frequency

1.0

0.2
0

46

44

Maine
Cold (6C)

42

40

38
36
Latitude (N)

34

32

30

Georgia
Warm (21C)

Mutatio
n
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.

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Point
Mutations
A point mutation is a change in one base in a
gene

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 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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

 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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Mutations That Alter Gene Number or

Sequence
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
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Mutation
Rates
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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Sexual
Reproduction
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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Porcupine
herd range

T
ES S
HW RIE
RT ITO
NO RR
TE

Beaufort Sea

MAP
AREA

Fortymile
herd range

Fortymile herd

CANADA

ALASKA

Porcupine herd

ALASKA
Y U K ON

Fig. 23-5

Porcupine
herd range

T
ES S
HW RIE
RT ITO
NO RR
TE

Beaufort Sea

MAP
AREA

Fortymile
herd range
ALASKA
YUKON

CANADA

ALASKA

Fig. 23-5a

 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 x 2
The total number of dominant alleles at a locus
is 2 alleles for each homozygous dominant
individual plus 1 allele for each heterozygous
individual; the same logic applies for recessive
alleles
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

 By convention, if there are 2 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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

The Hardy-Weinberg
Principle
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
Thus, the H-W principle is a null model.

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Hardy-Weinberg
Equilibrium
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
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Fig. 23-6

Alleles in the population

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

Gametes produced
Each egg:

Each sperm:

80%
20%
chance chance

80%
20%
chance chance

 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
where p2 and q2 represent the frequencies of
the homozygous genotypes and 2pq
represents the frequency of the heterozygous
genotype
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

80% CR ( p = 0.8)

20% CW (q = 0.2)

CW
(20%)

CR
(80%)

Sperm
CR
(80%)

64% ( p2)
CRCR

CW
(20%)

Eggs

Fig. 23-7-1

16% (qp)
CRCW

16% ( pq)
CRCW
4% (q2)
CWCW

Fig. 23-7-2

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

Gametes of this generation:

64% CR + 16% CR= 80% CR = 0.8 = p
4% CW + 16% CW= 20% CW = 0.2 = q

Fig. 23-7-3

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

Gametes of this generation:

64% CR + 16% CR= 80% CR = 0.8 = p
4% CW + 16% CW= 20% CW = 0.2 = q
Genotypes in the next generation:

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

20% CW(q = 0.2)

80% CR(p = 0.8)

CW
(20%)

64% ( p2)
CRCR

16% (pq)
CRCW

CR
(80%)
C
(20%)

Eggs

Sperm
C
(80%)

Fig. 23-7-4

16% (qp)
CRCW

4% (q2)
CWCW

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

Gametes of this generation:

64% CR+ 16% CR= 80% CR = 0.8 = p
4% CW + 16% CW = 20% CW = 0.2 = q
Genotypes in the next generation:

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

Conditions for Hardy-Weinberg

Equilibrium
The Hardy-Weinberg theorem describes a
hypothetical population
In real populations, allele and genotype
frequencies do change over time

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

 The five conditions for nonevolving populations

are rarely met in nature:
No mutations
Random mating
No natural selection
Extremely large population size
No gene flow

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

 Natural populations can evolve at some loci,

while being in Hardy-Weinberg equilibrium at
other loci

Applying the Hardy-Weinberg Principle

We can assume the locus that causes
phenylketonuria (PKU) is in Hardy-Weinberg
equilibrium given that:
The PKU gene mutation rate is low
Mate selection is random with respect to
whether or not an individual is a carrier for the
PKU allele
Natural selection can only act on rare
homozygous individuals who do not follow
dietary restrictions
The population is large
Migration has no effect as many other
populations have similar allele frequencies

What is the frequency of carriers of the PKU allele?

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 x 0.99 x 0.01 = 0.0198
or approximately 2% of the U.S. population
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Natural
Selection
Differential success in reproduction results in
certain alleles being passed to the next
generation in greater proportions

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Genetic
Drift
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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Fig. 23-8-1

CRCR

CRCR
CRCW

CRCR

CWCW
CRCW
CRCR
CRCR

CRCW
CRCW

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

Fig. 23-8-2

CRCR

CRCR

CWCW

CRCW

CRCW

CRCR

CWCW

CWCW

CRCR
CRCW

CRCW
CRCR
CRCR

CRCR

CRCW
CRCW

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

CWCW
CRCW

CRCR
CRCW

Generation 2
p = 0.5
q = 0.5

Fig. 23-8-3

CRCR

CRCR

CWCW

CRCW

CRCR

CRCW

CRCR

CRCW
CRCW

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

CWCW
CRCW

CRCR
CRCR

CWCW

CRCR

CRCW

CRCR

CRCR

CRCW

CRCR

CWCW

CRCR

CRCR
CRCR

CRCR
CRCR

CRCR
CRCW

Generation 2
p = 0.5
q = 0.5

CRCR

CRCR

Generation 3
p = 1.0
q = 0.0

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

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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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Fig. 23-9

Original
population

Bottlenecking
event

Surviving
population

 Understanding the bottleneck effect can

increase understanding of how human activity
affects other species

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Fig. 23-10
Pre-bottleneck Post-bottleneck
(Illinois, 1820) (Illinois, 1993)

Range
of greater
prairie
chicken

(a)

Location

Population
size

Percentage
Number
of alleles of eggs
per locus hatched

Illinois
1,00025,000

5.2

93

<50

3.7

<50

Kansas, 1998
(no bottleneck)

750,000

5.8

99

Nebraska, 1998
(no bottleneck)

75,000
200,000

5.8

96

Minnesota, 1998
(no bottleneck)

4,000

5.3

85

19301960s
1993

(b)

Fig. 23-10a

Pre-bottleneck
(Illinois, 1820)

(a)

Range
of greater
prairie
chicken

Post-bottleneck
(Illinois, 1993)

Fig. 23-10b

Location

Population
size

Number
Percentage
of alleles of eggs
per locus hatched

Illinois
19301960s
1993

1,00025,000
<50

5.2

93

3.7

<50

Kansas, 1998
(no bottleneck)

750,000

5.8

99

Nebraska, 1998
(no bottleneck)

75,000
200,000

5.8

96

Minnesota, 1998
(no bottleneck)

4,000

5.3

85

(b)

 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 population in other states and
were successful in introducing new alleles and
increasing the egg hatch rate to 90%
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Effects of Genetic Drift: A

Summary
1. Genetic drift is significant in small populations
2. Genetic drift causes 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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

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 differences between
populations over time
Gene flow is more likely than mutation to alter
allele frequencies directly
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Fig. 23-11

 Gene flow can decrease the fitness of a

population, particularly if it is adapted to
specific environmental conditions
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
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Fig. 23-12

Index of copper tolerance

70
60

MINE
SOIL

NONMINE
SOIL

NONMINE
SOIL

50

Prevailing wind direction

40
30
20
10
0

20

20

20

40

60

80

Distance from mine edge (meters)

100

120

140

160

Fig. 23-12a

Index of copper tolerance

70
60

MINE
SOIL

NONMINE
SOIL

50

NONMINE
SOIL
Prevailing wind direction

40
30
20
10
0

20

20

0
100
20
40
60
80
Distance from mine edge (meters)

120

140

160

Fig. 23-12b

 Gene flow can also 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
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Concept 23.4: Natural selection is the only

mechanism that consistently causes adaptive
evolution
Only natural selection consistently results in
adaptive evolution

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A Closer Look at Natural

Selection
Natural selection brings about adaptive evolution
by acting on an organisms phenotype

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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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

 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

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Directional, Disruptive, and Stabilizing

Selection
Three modes of selection:
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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Frequency of individuals

Fig. 23-13

Original
Evolved
population population

(a) Directional selection

Original population

Phenotypes (fur color)

(b) Disruptive selection

(c) Stabilizing selection

Frequency of individuals

Fig. 23-13a

Original population

Phenotypes (fur color)

Original population
Evolved population

(a) Directional selection

Frequency of individuals

Fig. 23-13b

Original population

Phenotypes (fur color)

Evolved population

(b) Disruptive selection

Frequency of individuals

Fig. 23-13c

Original population

Phenotypes (fur color)

Evolved population

(c) Stabilizing selection

What kind of selection does this

situation describe?

What kind of selection does this

situation describe?

Stabilizing selection.intermediates are favored.

The Key Role of Natural Selection in Adaptive

Evolution
Natural selection increases the frequencies of
alleles that enhance survival and reproduction
Adaptive evolution occurs as the match
between an organism and its environment
increases

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Fig. 23-14

(a) Color-changing ability in cuttlefish

Movable bones

(b) Movable jaw

bones in
snakes

Fig. 23-14a

(a) Color-changing ability in cuttlefish

Fig. 23-14b

Movable bones

(b) Movable jaw

bones in
snakes

 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

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

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

Fig. 23-15

 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
Male showiness due to mate choice can
increase a males chances of attracting a
female, while decreasing his chances of
survival (an evolutionary tradeoff)
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Sexual Selection: Female Mate Choice

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

Fig. 23-16

EXPERIMENT

Female gray
tree frog
SC male gray
tree frog

LC male gray
tree frog

SC sperm
sperm

Eggs

LC

Offspring of Offspring of
SC father
LC father
Fitness of these half-sibling offspring compared

RESULTS

Fitness Measure

1995

1996

Larvalgrowth

NSD

LCbetter

Larvalsurvival

LCbetter

NSD

Timetometamorphosis

LCbetter
(shorter)

LCbetter
(shorter)

NSD=nosignificantdifference;LCbetter=offspringofLCmales
superiortooffspringofSCmales.

Fig. 23-16a

EXPERIMENT

Female gray
tree frog
LC male gray
tree frog

SC male gray
tree frog

SC sperm Eggs LC sperm

Offspring of Offspring of
LC father
SC father
Fitness of these half-sibling offspring compared

Fig. 23-16b

RESULTS

Fitness Measure

1995

1996

Larvalgrowth

NSD

LCbetter

Larvalsurvival

LCbetter

NSD

Timetometamorphosis

LCbetter
(shorter)

LCbetter
(shorter)

NSD=nosignificantdifference;LCbetter=offspringofLCmales
superiortooffspringofSCmales.

The Preservation of Genetic

Variation
Various mechanisms help to preserve genetic
variation in a population
Diploidy
Balancing Selection
Heterozygote Advantage
Frequency-dependent Selection

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Diploid
y
Diploidy maintains genetic variation in the form
of hidden recessive alleles that selection does
not act upon because they are not manifest in
the phenotype.

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Balancing
Selection
Balancing selection occurs when natural
selection maintains stable frequencies of two or
more phenotypic forms in a population
Heterozygote Advantage
Frequency-dependent selection

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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
Example: The sickle-cell allele causes
mutations in hemoglobin (bad) but also confers
malaria resistance (good)
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Fig. 23-17

Frequencies of the
sickle-cell allele
02.5%

Distribution of
malaria caused by
Plasmodium falciparum
(a parasitic unicellular eukaryote)

2.55.0%
5.07.5%
7.510.0%
10.012.5%
>12.5%

Frequency-Dependent Selection
In frequency-dependent selection, the fitness
of a phenotype declines if it becomes too
common in the population
Selection can favor whichever phenotype is
less common in a population
Example: prey-switching behavior in many
predators leads to multiple forms of prey
coexisting in populations
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Fig. 23-18

Right-mouthed

Frequency of
left-mouthed individuals

1.0

Left-mouthed

0.5

1981 82 83 84 85 86 87 88 89 90
Sample year

Fig. 23-18a

Right-mouthed

Left-mouthed

Fig. 23-18b

Frequency of
left-mouthed individuals

1.0

0.5

1981 8
2

8
3

8 8 8 8
6year
7
4 Sample
5

8
8

8
9

9
0

Neutral
Variation
Neutral variation is genetic variation that
appears to confer no selective advantage or
disadvantage
For example,
Variation in noncoding regions of DNA
(although many of these regions are now
known to be swtiches that control the
expression of other genes)
Variation in proteins that have little effect on
protein function or reproductive fitness
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Why Natural Selection Cannot Fashion

Perfect Organisms
1. Selection can act only on existing variations, it is
not the source of new alleles (mutations are)
2. Evolution is limited by historical constraints
(developmental patterns, for example)
3. Adaptations are often compromises between
different organismal needs (mate attraction,
predator avoidance)
4. Chance events, natural selection, and the
environment interact and conidtions constantly
change so perfection is a moving target
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

You should now be able

to:
1. Explain why the majority of point mutations
are harmless
2. Explain how sexual recombination generates
genetic variability
3. Define the terms population, species, gene
pool, relative fitness, and neutral variation
4. List the five conditions of Hardy-Weinberg
equilibrium
Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

5. Apply the Hardy-Weinberg equation to a

population genetics problem
6. Explain why natural selection is the only
mechanism that consistently produces
adaptive change
7. Explain the role of population size in genetic
drift

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings

8. Distinguish among the following sets of terms:

directional, disruptive, and stabilizing
selection; intrasexual and intersexual
selection
9. List four reasons why natural selection cannot
produce perfect organisms

Copyright 2008 Pearson Education Inc., publishing as Pearson Benjamin Cummings