BIO 111 - Chapter 20

Because learning changes everything.
Chapter 20
Genes Within
Populations
BIOLOGY
Thirteenth Edition
Raven, Johnson, Mason, Losos,
Duncan
© 2023 McGraw Hill, LLC. All rights reserved. Authorized only for instructor use in the classroom.
No reproduction or further distribution permitted without the prior written consent of McGraw Hill, LLC.
Lecture Outline
20.1 Genetic Variation and Evolution

20.2 Changes in Allele Frequency
20.3 Five Agents of Evolutionary Change
20.4 Quantifying Natural Selection
20.5 Reproductive Strategies
20.6 Natural Selection’s Role in Maintaining
Variation
20.7 Selection Acting on Traits Affected by
Multiple Genes
20.8 Experimental Studies of Natural
Selection
20.9 Interactions Among Evolutionary
Forces
20.10 The Limits of Selection
tamoncity/Shutterstock
© McGraw Hill, LLC 2

Genetic Variation and Evolution
Genetic variation
• Differences in alleles of genes found within individuals in a
population.
• Natural populations contain much variation.
Evolution
• How an entity changes through time.
• Development of modern concept traced to Darwin.
• “Descent with modification”.

Evolution
“Through time, species accumulate differences; as a result,

descendants differ from their ancestors. In this way, new
species arise from existing ones.”
• Charles Darwin.

Many processes lead to evolutionary change
• Darwin was not the first to propose a theory of evolution
• Unlike his predecessors, however, Darwin proposed

natural selection as the mechanism of evolution
• Natural selection can lead to change in allele frequencies

– frequencies of alleles of a gene from generation to
generation

Population genetics
• Study of properties of genes in a population
• Evolution results in a change in the genetic composition of

a population
• Natural populations contain substantial genetic variation
• Genetic variation is required for evolution to occur

Figure 20.1
Rosita So Image/Getty Images

Figure 20.2
Diverse Images/Universal Images Group/Getty Images

Genetic variation
Genetic variation in populations now measured using

increasingly sophisticated tools
Examples of variation due to genetic differences
• Human blood groups
SNPs
• Used to assess patterns in over 300 species
• >100,000 human genomes partially or wholly sequenced
• Extensive genetic variation has been documented
• African genomes have the most variation

Changes in Allele Frequency
• Genetic variation was a puzzle to Darwin and his

contemporaries
• Mendel’s work largely unknown
• Scientists thought selection should favor an optimal form;

eliminate variation
• Blending inheritance widely accepted – offspring expected

to be phenotypically intermediate relative to parents

Hardy–Weinberg principle 1
Predicts genotype frequencies

Hardy–Weinberg equilibrium
• Proportions of genotypes do not change in a population as
long as:
1. No mutation takes place
2. No genes are transferred to or from other sources (no immigration

or emigration)
3. Mating is random
4. The population size is very large
5. No selection occurs

Hardy–Weinberg principle 2
Can be written as an equation

Used to calculate allele frequencies
Frequency of first allele is p, second allele is q
p + q = 1 in a population
• p = B for black coat color.
• Black cat is BB or Bb.
• q = b for white coat color.

• White cats are bb.
p 2  2 pq  q 2  1
BB  Bb  bb  1
Figure 20.3 (1)
Access the text alternative for slide images.

Figure 20.3 (2)

Making Hardy–Weinberg predictions 1
• If all 5 assumptions for Hardy-Weinberg equilibrium are

true, allele and genotype frequencies do not change from
one generation to the next
• In reality, most populations will not meet all 5 assumptions
• Primary use of the equation is to determine whether

evolutionary processes are operating in a population and
hypothesize what the processes are

Making Hardy–Weinberg predictions 2
What makes populations vary from Hardy-Weinberg

equilibrium?
• Natural selection might favor homozygotes over

heterozygotes.
• Individuals may choose to mate with genetically similar

individuals.
• Influx of individuals from other populations
• Mutations occurring

Five Agents of Evolutionary Change

Agents of evolutionary change 1
Mutation
• Rates generally low
• Other evolutionary processes usually more important in
changing allele frequency
• Ultimate source of genetic variation
• Makes evolution possible
Gene flow
• Movement of alleles from one population to another
• Animal physically moves into new population
• Drifting of gametes or immature stages of plants or animals into
an area
• Pollen and seeds can travel long distances

Agents of evolutionary change 2
Nonrandom mating
• Assortative mating
• Phenotypically similar individuals mate
• Increases proportion of homozygous individuals
• Disassortative mating
• Phenotypically different individuals mate
• Produces excess of heterozygotes

Genetic drift
Genetic drift
• In small populations, allele frequency may change by
chance alone.
• Population must be large to be in H-W equilibrium.
• Magnitude of genetic drift is inversely related to population

size.
• Can lead to the loss of alleles in isolated populations and
uncommon alleles are more vulnerable.
• Founder effect
• Bottleneck effect

Founder effect
• One or a few individuals disperse and become the

founders of a new, isolated population
• Some alleles are lost, and others change in frequency
• Organisms on islands
• Self-pollinating plants
• Amish populations

Bottleneck effect
• Drastic reduction in population size due to drought,

disease, other natural forces
• Survivors may constitute a random genetic sample of the

original population
• Results in loss of genetic variability

Figure 20.5

Figure 20.6
Northern Elephant Seal

• Bottleneck case study
• Nearly hunted to extinction in 19th century
• As a result, species has lost almost all of its genetic variation
• Population now numbers in tens of thousands

Selection favors some genotypes over others
• Some individuals leave behind more progeny than others,

and the rate at which they do so is affected by phenotype
and behavior
• Artificial selection – breeder selects desired characteristics
• Natural selection – environmental conditions determine

which individuals produce the most offspring

Evolution by natural selection 1
These conditions must be met for evolution by natural

selection to occur
1. Variation must exist among individuals in a population
2. Variation among individuals must result in differences in

the number of offspring surviving in the next generation
3. Variation must have a genetic basis

Evolution by natural selection 2
Natural selection and evolution are not the same

• Natural selection is a process.
• Only one of several processes that can result in evolution.
• Evolution is the historical record, or outcome, of change

through time.
Result of evolution driven by natural selection is that
populations become better adapted to their environment

Selection to avoid predators
Pocket mice come in different colors

• Population living on rocks favor dark color.
• Populations living on sand favor light color.

Selection to match climatic conditions
• Enzyme allele frequencies often vary with latitude
• Allele frequencies of enzyme lactate dehydrogenase in

fish vary geographically
• Enzymes formed by these alleles function differently at

different temperatures

Selection for pesticide and microbial resistance
Widespread use of insecticides has led to the rapid evolution

of resistance in more than 500 pest species
Housefly has pesticide resistance alleles:

• pen gene decreases insecticide uptake.
• kdr and dld-r genes decrease target sites for insecticide.
Evolution of resistance to antibiotics in many disease-

causing pathogens
• With each new drug, resistant organisms emerge;
dangerous for human health.

Figure 20.8

Quantifying Natural Selection
Fitness
• Individuals with one phenotype leave more surviving

offspring in the next generation than individuals with an
alternative phenotype.
• Relative concept; the most fit phenotype is simply the one

that produces, on average, the greatest number of
offspring.

Measuring fitness
Most fit phenotype is assigned fitness value of 1

Population of toads: green and brown
• Green toads have 4 offspring.
• Brown toads leave 2.5 offspring.
• Green fitness 4/4=1.
• Brown fitness 2.5/4=0.625.

Many components of fitness
Fitness has many components

• Survival
• Sexual selection – some individuals more successful at
attracting mates
• Number of offspring per mating
• Traits favored for one component may be a disadvantage
for others
Selection favors phenotypes with the greatest fitness
• Phenotype with greater fitness usually increases in
frequency.
Figure 20.9
• Larger female water striders lay more eggs per day

• But large females survive for a shorter period of time
• As a result, intermediate-sized females produce the most
offspring over the course of their entire lives and thus have the
highest fitness

Reproductive Strategies
• Males and females usually differ in how they attempt to

maximize fitness
• Females evaluate a male’s quality and then decide

whether to mate
• Peahens prefer to mate with peacocks with more eyespots

on their tailfeathers

Figure 20.10
(a) Carole_R/Flickr Flash/Getty Images; (b) Bruce Beehler/Science Source

Parental Investment 1
Refers to the energy and time each sex invests in producing

and rearing offspring
Females have a higher parental investment
Sexes face very different selective pressures
Males best increase their fitness by mating with as many

females as possible
Females are limited by number of eggs that can be

produced, so a female should be choosy in picking the male
that can provide greatest benefit

Parental Investment 2
However in cases with biparental care, mate choice should

be more equal
In some cases male investment exceeds that of females
• Male Mormon crickets transfer a protein-containing packet
to females during mating.
• Males are the choosy sex.
• Males that choose larger females leave more offspring.

Figure 20.11

Sexual selection
• Competition for mates

• Intrasexual selection – competitive interactions between
members of one sex
• Intersexual selection – mate choice
• Secondary sexual characteristics – antlers and horns
used to combat other males; long tail feathers and bright
plumage used to “persuade” members of opposite sex
• Sexual dimorphism – Differences between sexes (males
larger than females)
• Sperm competition – selects for features that increase
probability that a male’s sperm will fertilize the eggs
Intrasexual selection
• Individuals of one sex (usually males) compete with each

other for the opportunity to mate
• A few successful males may engage in an inordinate

number of mating, whereas most males don’t mate at all
• Elephant seal males control territories on breeding

beaches; a few dominant males do most of the breeding

Figure 20.12
©Cathy & Gordon ILLG

Intersexual selection
Active choice of a mate

Direct benefits of mate choice
• Females can benefit from choosing the male that can provide
the best care.
• Males may provide no care but maintain territories that provide
food and refuge.
Indirect benefits of mate choice
• One possible indirect benefit is higher-quality offspring.
• Choose the male that is healthiest or oldest.
• Handicap hypothesis – only genetically superior mates survive
with a handicap such as a long tail that is a hinderance in flying
– hypothesis still being debated.

Sensory Exploitation 1
Some courtship displays appear to have evolved from a

predisposition in females to respond to certain stimuli
Sensory exploitation – evolution in males of a signal that

exploits preexisting biases
Túngara frog
• Males have a short burst of sound at the end of their calls.
• Females of this species and other related species are

particularly attracted to this call.

Figure 20.13
W. Perry Conway/Getty Images

Sensory Exploitation 2
Opportunity for sensory exploitation may be widespread
• Experiment to see if birds had latent preferences for

particular stimuli in the zebra finch.
• Long, differently colored feathers were glued vertically to

the tops of the heads of male birds, like nothing found in
nature.
• Females were strongly attracted to males with white

crests, but not red or green ones.
• If a mutation arose that caused a male to have a white

crest on his head it would give a great advantage.

Figure 20.14

Natural Selection’s Role in Maintaining Variation
Frequency-dependent selection
• Fitness of a phenotype depends on its frequency within
the population.
• Negative frequency-dependent selection
• Rare phenotypes favored by selection
• Rare forms may not be in “search image”; preyed upon less
frequently
• Positive frequency-dependent selection

• Favors common form
• Tends to eliminate variation
• “Oddballs” stand out

Figure 20.15
• Negative frequency-dependent selection

• In water boatman, fish eat the most common color type
more than they would by chance alone

Positive frequency-dependent selection

Oscillating selection
Selection favors one phenotype at one time and another

phenotype at another time
Effect will be to maintain genetic variation in the population
Medium ground finch of Galápagos Islands
• Birds with big bills favored during drought.
• Birds with smaller bills favored in wet conditions.

Heterozygote advantage
Heterozygotes are favored over homozygotes

Works to maintain both alleles in the population
Sickle cell anemia
• Hereditary disease affecting hemoglobin.
• Causes severe anemia.
• Homozygotes for sickle cell allele usually die before
reproducing (without medical treatment).

Sickle cell allele
• Why is the sickle cell

allele not eliminated?
• Leading cause of
death in central Africa
is malaria
• Heterozygotes for
sickle cell allele do
not suffer anemia and
are much less
susceptible to malaria

Selection Acting on Traits Affected by Multiple Genes
Many traits affected by more than one gene
Selection operates on all the genes for the trait
Changes the population depending on which genotypes are

favored
Types of selection
• Disruptive
• Directional
• Stabilizing

Figure 20.18

Disruptive selection
Acts to eliminate intermediate types

Different beak sizes of African black-bellied seedcracker
finch
• Available seeds fall into 2 categories.
• Favors bill sizes for one or the other.
• Birds with intermediate-sized beaks are at a disadvantage
with both seed types – they are unable to open large
seeds and too clumsy to efficiently process small seeds.

Figure 20.19

Directional selection
Acts to eliminate one extreme

Often occurs in nature when the environment changes
In Drosophila, eliminated flies that moved toward the light
• Now fewer in resulting population have that behavior.

Figure 20.20

Stabilizing selection
• Acts to eliminate both extremes
• Makes intermediate more common by eliminating

extremes
• In humans, infants with intermediate weight at birth have

the highest survival rate

Figure 20.21

Experimental Studies of Natural Selection
To study evolution, biologists have traditionally investigated

what has happened in the past
• Fossils or DNA evidence
Laboratory studies on fruit flies common for nearly 100 years
Only recently started with evolutionary lab and field

experiments

Natural selection in guppies 1
Guppy coloration
• Found in small streams in northeastern South America and
Trinidad.
• Some are capable of colonizing portions of streams above
waterfalls.
• Different dispersal methods
• Other species not able to make it upstream
• Dispersal barriers create 2 different environments.

• Predators rare above waterfall

Pike cichlid (predator) rare above waterfall

• Killifish rarely eats guppies.
• Guppy males larger and gaudier.
Predator common below waterfall

• Individuals that are more drab and reproduce earlier.

Figure 20.22

Guppy lab study

• Other explanations are possible for field results.
• 10 large pools
• Added pike cichlids to 4, killifish to 4, and 2 left as controls.
• 14 months and 10 guppy generations later.
• Guppies in killifish and control pool – large and colorful.
• Guppies in pike cichlid pools – smaller and drab.
Field experiments confirmed that variations between populations in

nature were the result of genetic differences
Scientists can test hypotheses about evolution in nature

Figure 20.23

Interactions Among Evolutionary Forces 1
Mutations and genetic drift may counter selection

• In nature, mutation rates are rarely high enough to counter
selection.
• Selection is nonrandom but genetic drift is random.
• Drift may decrease an allele favored by selection.
• Selection usually overwhelms drift except in small populations.

Interactions Among Evolutionary Forces 2
Gene flow can be

• Constructive
• Spread beneficial mutation to other populations.
• Constraining
• Can impede adaptation by continual flow of inferior alleles from
other populations.

Slender bent grass at copper mines
• High concentration of metal ions in the soil surrounding

abandoned mines
• Alleles of certain genes confer the ability to grow on soils
high in heavy metals, but resistant individuals grow slower
on uncontaminated sites
• Expect resistance allele at 100% at mine site, 0%
elsewhere
• Instead, resistance allele occurs at intermediate levels in
many areas (pollen is carried by wind)
• Gene flow between sites high enough to counteract
natural selection

Figure 20.24

Limits of Selection
Multiple phenotypic effects of alleles

• Same gene affects size of hen’s comb and rate at which
she lays – selection for hens that lay many eggs but have a
small comb would be difficult.
Lack of genetic variation
• Gene pool of thoroughbred horses limited and performance
times have not improved for more than 50 years.
• Phenotypic variation may not have genetic basis.
• Interactions between genes – epistasis.
• Selective advantage of an allele at one gene may vary from
one genotype to another.

Selection for increased speed in racehorses is no longer
effective

Differences in the number of ommatidia in fly eyes does
not have a genetic basis

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Because learning changes everything.

20.1 Genetic Variation and Evolution

© McGraw Hill, LLC 2

© McGraw Hill, LLC 3

“Through time, species accumulate differences; as a result,

© McGraw Hill, LLC 4

• Darwin was not the first to propose a theory of evolution

• Unlike his predecessors, however, Darwin proposed

• Natural selection can lead to change in allele frequencies

© McGraw Hill, LLC 5

• Study of properties of genes in a population

• Evolution results in a change in the genetic composition of

• Natural populations contain substantial genetic variation

• Genetic variation is required for evolution to occur

© McGraw Hill, LLC 6

Rosita So Image/Getty Images

© McGraw Hill, LLC 7

Diverse Images/Universal Images Group/Getty Images

© McGraw Hill, LLC 8

Genetic variation in populations now measured using

© McGraw Hill, LLC 9

• Genetic variation was a puzzle to Darwin and his

• Mendel’s work largely unknown

• Scientists thought selection should favor an optimal form;

• Blending inheritance widely accepted – offspring expected

© McGraw Hill, LLC 10

Predicts genotype frequencies

2. No genes are transferred to or from other sources (no immigration

4. The population size is very large

© McGraw Hill, LLC 11

Can be written as an equation

• q = b for white coat color.

Access the text alternative for slide images.

© McGraw Hill, LLC 13

Access the text alternative for slide images.

© McGraw Hill, LLC 14

• If all 5 assumptions for Hardy-Weinberg equilibrium are

• In reality, most populations will not meet all 5 assumptions

• Primary use of the equation is to determine whether

© McGraw Hill, LLC 15

What makes populations vary from Hardy-Weinberg

• Natural selection might favor homozygotes over

• Individuals may choose to mate with genetically similar

• Influx of individuals from other populations

© McGraw Hill, LLC 16

Access the text alternative for slide images.

© McGraw Hill, LLC 17

© McGraw Hill, LLC 18

• Increases proportion of homozygous individuals

• Produces excess of heterozygotes

© McGraw Hill, LLC 19

• Magnitude of genetic drift is inversely related to population

© McGraw Hill, LLC 20

• One or a few individuals disperse and become the

• Some alleles are lost, and others change in frequency

© McGraw Hill, LLC 21

• Drastic reduction in population size due to drought,

• Survivors may constitute a random genetic sample of the

• Results in loss of genetic variability

© McGraw Hill, LLC 22

Access the text alternative for slide images.

© McGraw Hill, LLC 23