Why did this happen?

LECTURE PRESENTATIONS
For CAMPBELL BIOLOGY, NINTH EDITION
Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson

Chapter 23

The Evolution of Populations

Lectures by
Erin Barley
Kathleen Fitzpatrick

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Did this bird evolve in its lifetime?
Evidence of selection by food source.
Microevolution is a change in allele frequencies in a population over generations

Average beak depth (mm)

10

0
1976 1978
(similar to the (after
prior 3 years) drought)
 –Three mechanisms cause
allele frequency change:
• Natural selection
• Genetic drift
• Gene flow
– Only natural selection causes adaptive
evolution

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Concept 23.1: BIG PICTURE: Genetic
variation makes evolution possible

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Nonheritable variation of Nemoria moths: different phenotypes due to
dietary chemicals
Natural selection can only act on variation with a genetic component

(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

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Variation Between Populations
• Most species exhibit geographic variation,
differences between gene pools of separate
populations
• For example, Madeira is home to several
isolated populations of mice
– Chromosomal variation among populations is
due to drift, not natural selection

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Figure 23.4

1 2.4 3.14 5.18 6 7.15

8.11 9.12 10.16 13.17 19 XX

1 2.19 3.8 4.16 5.14 6.7

9.10 11.12 13.17 15.18 XX

 mummichog fish vary in a cold-adaptive allele along a temperature gradient
1.0
Ldh-Bb allele frequency

0.8

0.6

0.4

0.2

0
46 44 42 40 38 36 34 32 30
Latitude (ºN)
Maine Georgia
Cold (6°C) Warm (21ºC)

HOW DID THIS COME TO BE?

Sources of Genetic Variation
• New genes and alleles can arise by mutation or
gene duplication
– The average is about one mutation in every 100,000
genes per generation

– An ancestral odor-detecting gene has been duplicated

many times: humans have 1,000 copies of the gene,
mice have 1,300
– What is the consequence?

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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:
1. Natural selection
2. Genetic drift
3. Gene flow

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Genetic Drift
• Genetic drift describes how allele frequencies
fluctuate unpredictably from one generation to the
next due to chance events
– The smaller a sample, the greater the chance of
deviation from a predicted result
– genetic drift tends to reduce genetic variation through
losses of alleles

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 Animation: Causes of Evolutionary Change
Right-click slide / select “Play”
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Figure 23.9-1

Example of Genetic drift

CRCR CRCR

CRCW

CWCW CRCR

CRCW

CRCR CRCW

CRCR CRCW

Generation 1
p (frequency of CR) = 0.7
q (frequency of CW) = 0.3
Figure 23.9-2

Example of Genetic drift

CRCR
5 CWCW CRCR
CRCR
plants
CRCW
leave
CRCW
off-
spring
CWCW CRCR CRCR CWCW

CRCW CRCW

CRCR CRCW CWCW CRCR

CRCR CRCW CRCW CRCW

Generation 1 Generation 2
p (frequency of CR) = 0.7 p = 0.5
q (frequency of CW) = 0.3 q = 0.5
Example of Genetic drift

CRCR
5 CWCW CRCR
2 CRCR
CRCR
plants plants
CRCW
leave leave
CRCW CRCR CRCR
off- off-
spring spring
CWCW CRCR CRCR CWCW CRCR CRCR

CRCW CRCW CRCR CRCR

CRCR CRCW CWCW CRCR CRCR

CRCR CRCW CRCW CRCW CRCR 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
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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Bottleneck effect

Original
population
Figure 23.10-2

Original Bottlenecking
population event
The bottleneck effect is a sudden reduction in population size due to a
change in the environment
If the population remains small, it may be further affected by genetic
drift

Original Bottlenecking Surviving

population event population
 • How is the bottleneck effect seen in humans
affecting wild organism populations?

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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

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 Gene Flow
• Gene flow consists of the movement of alleles
amongst 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

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 • Gene flow can decrease the fitness of a population
• Consider, for example, the great tit (Parus major)
on the Dutch island of Vlieland
– Mating causes gene flow between the central and
eastern populations
– Immigration from the mainland introduces alleles
that decrease fitness
– Natural selection selects for alleles that increase
fitness
– Birds in the central region with high immigration
have a lower fitness; birds in the east with low
immigration have a higher fitness
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Gene flow and local adaptation

Population in which the

60 surviving females Central
eventually bred population

50 Central NORTH SEA Eastern

Eastern population
Survival rate (%)

Vlieland,
40 the Netherlands
2 km
30

20

10

0
Females born Females born
in central in eastern Parus major
population population
 • Gene flow can also 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

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Bioflix: Selection within hypothetical
beetle population
Come up with a new example of:

• Natural selection
• Gene flow
• Genetic drift
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 (mutations) arise by
chance
– Beneficial alleles are “sorted” and favored by
natural selection

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 • Relative fitness is the contribution an individual
makes to the gene pool of the next generation,
relative to the contributions of other individuals
– Typically individuals are NOT in direct competition

• 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

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Figure 23.13

Frequency of
individuals
Original population

Phenotypes (fur color)

Original Evolved
population population

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

 Bones shown in
green are movable.

Ligament

Striking
adaptations
have arisen by
natural selection
 Evolution over time, process and outcomes

• 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
• Because the environment can change, adaptive
evolution is a continuous process

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 • 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

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Why are males prettier than females?
Why are males prettier than females?
Why are males prettier than females?
Why are males prettier than females?
Why are males prettier than females?
Why are males prettier than females?
outcome is sexual dimorphism
 • 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 male’s chances of attracting a
female, while decreasing his chances of survival

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 • Why do females like him the best?

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Do all animal species
care about looks?
Figure 23.16
EXPERIMENT
Recording of SC Recording of LC
male’s call male’s call

Female gray
tree frog
SC male gray LC male gray
tree frog tree frog
SC sperm  Eggs  LC sperm

Offspring of Offspring of
SC father LC father

Survival and growth of these half-sibling offspring compared

RESULTS

Offspring Performance 1995 1996

Larval survival LC better NSD
Larval growth NSD LC better
Time to metamorphosis LC better LC better
(shorter) (shorter)
NSD = no significant difference; LC better = offspring of LC males superior to
offspring of SC males.
Diploidy
• What’s the benefit?

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Diploidy
• Diploidy maintains genetic variation in the form
of hidden recessive alleles
– by carrying recessive alleles that are hidden
from the effects of selection

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Balancing Selection
• 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

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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
• The sickle-cell allele causes mutations in
hemoglobin but also confers malaria resistance

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Figure 23.17

Key
Frequencies of the
sickle-cell allele
0–2.5%
2.5–5.0%

Distribution of 5.0–7.5%
malaria caused by 7.5–10.0%
Plasmodium falciparum
(a parasitic unicellular eukaryote) 10.0–12.5%
>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
• For example, frequency-dependent selection
selects for approximately equal numbers of
“right-mouthed” and “left-mouthed” scale-eating
fish
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Figure 23.18

“Left-mouthed”
P. microlepis

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

0.5

0
1981 ’82 ’83 ’84 ’85 ’86 ’87 ’88 ’89 ’90
Sample year
Why Natural Selection Cannot Fashion
Perfect Organisms

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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

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Evolutionary compromise…for the frog
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

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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

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The Hardy-Weinberg Principle
• The Hardy-Weinberg principle states that
frequencies of alleles and genotypes in a
population remain constant from generation to
generation, ∴ not evolving
– If a population does not meet the criteria of the
Hardy-Weinberg principle, it can be concluded that
the population is evolving

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 • The five conditions for nonevolving populations
are rarely met in nature, and include:
1. No mutations
2. Random mating
3. No natural selection
4. Extremely large population size
5. No gene flow

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Figure 23.6

CANADA
MAP

ALASKA
AREA

Beaufort Sea

Porcupine
herd range

Porcupine herd

Fortymile
herd range

Fortymile herd
 • 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

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 • 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

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 • 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

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 • 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

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 • 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

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 • 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

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

Sperm
CR (80%) CW (20%)

CR
(80%)
64% (p2) 16% (pq)
Eggs CRCR CRCW

CW 16% (qp) 4% (q2)

(20%) CRCW CWCW

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

Gametes of this generation:

64% CR R
+ 16% C R W = 80% CR = 0.8 = p
(from CRCR plants) (from C C plants)
4% CW W
+ 16% C R W = 20% CW = 0.2 = q
(from CWCW plants) (from C C plants)
Genotypes in the next generation:

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

Applying the Hardy-Weinberg Principle
• 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

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 • 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

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Try this:

A population of mice displays the recessive

trait for loving the banjo. A survey says it is
about 20% of them.
• What % dislike the banjo?
• What is the frequency of the dominant and
recessive allele?
• What % is heterozygous for this trait?
What is the problem with dog breeding?
What is the problem with dog breeding?
• Limit genetic
variability
• Preserve
harmful
mutations
Fitness
decreased
Review quiz
1. What are the three mechanisms for
altering allele frequency?
2. What is the type of selection reflecting the
fact that really small wolves and really big
wolves don’t survive as well as mediums?
3. Draw the curve.
4. In population of fruit flies, 70% of the
gametes contain A1 alleles. If the pop in
in H-W equilibrium, what proportion carry
both A1 and A2? 0.09