Biology
Sylvia S. Mader Michael Windelspecht
Chapter 16
How Populations
Evolve
Lecture Outline
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Outline
• 16.1 Genes, Populations, and Evolution
• 16.2 Natural Selection
• 16.3 Maintenance of Diversity
16.1 Genes, Populations, and
Evolution
• A
population
is all the members of a single
species occupying a particular area at the same
time.
• Diversity exists among members of a population.
•
Population genetics
is the study of this
diversity in terms of allele differences.
Evaluates the diversity of a population by studying genotype and phenotype frequencies over time
Genes, Populations, and
Evolution
• In the 1930s, population geneticists began to
describe variations in a population in terms of
alleles
•
Microevolution
pertains to evolutionary
changes within a population.
Various alleles at all the gene loci in all individuals make up the gene pool of the population.
The gene pool of a population can be described in terms of:
• Genotype frequencies
• Allele frequencies
The Genetic Basis of Body
Color in the Peppered Moth
5
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d d
D d
D D
Phenotype:
Genotype:
Alleles:
homozygous
homozygous heterozygous
chromosome
Genes, Populations, and
Evolution
• Allele Frequencies
The proportion of each allele within a population’s gene pool.
Frequencies of the dominant and recessive allele must add up to 1.
• This relationship is described by the expression p + q = 1
• p is the frequency of one allele and q is the frequency of the other
Microevolution involves a change in these allele frequencies within populations over time.
• If the gene frequencies do not change over time, microevolution has not occurred.
How Hardy-Weinberg Equilibrium
Is Estimated
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D
d
Equilibrium
genotype frequencies: Gene pool:
Population = 25 moths, 50 alleles
DD Dd Dd Dd
D = 10 d = 40
DD
Dd
Dd Dd Dd Dd
Dd D d
dd
p + q = 1 P2 + 2pq + q2 = 1
p = frequency of D q = frequency of d
p = frequency of D q = frequency of d Frequency of DD Frequency of D
dd dd dd dd
dd dd dd dd
dd dd dd dd
dd dd dd dd 1.00 1.00
p = 10/50 alleles
Frequency of d
q = 40/50 alleles
= 0.20
= 0.80
= 0.04
= 0.32 p2= freq D2
= (0.2)(0.2) Frequency of Dd 2pq = 2(freq D x freq d)
= 2(0.20 x 0.80) Frequency of dd
= (0.08)(0.80) = 0.64 q2 = freq d 2
Allele frequencies:
d d d d d d d d d d d d d d d d d d d d d d d d d d d d d d d d D D D D D d d d
Genes, Populations, and
Evolution
• The
Hardy-Weinberg Equalibrium
states
that:
Allele frequencies in a population will remain
constant assuming:
• No Mutations
• No Gene Flow
• Random Mating
• No Genetic Drift
• No Selection
Hardy-Weinberg Equilibrium
9
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D
d
eggs
0.80 d 0.20 D
0.20
0.80
0.64 dd 0.16 Dd
Offspring 0.04 DD
spe
Genes, Populations, and
Evolution
• Hardy-Weinberg Equilibrium:
Required conditions are rarely (if ever) met
Deviations from a Hardy-Weinberg equilibrium indicate that evolution has taken place
• Analysis of allele changes in populations over time determines the extent to which evolution has occurred.
Mechanisms of Microevolution
12
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d D d D d D d D Dd X Conclusion: Genotype frequencies: Allele frequencies: Allele frequencies: Genotype frequencies: DD Dd dd If... 0.20 D 0.80 d
p2 + 2pq + q2 = 1
DD = 0.04 Dd = 0.32 dd = 0.64
...then we expect
If...
Nonrandom mating
...then we observe
If...
DD Dd Dd DD
or
Selection
...then we observe
Dd DD dd 0.20 D 0.80 d DD
p2 + 2pq + q2 = 1
DD = 0.04 Dd = 0.32 dd = 0.64
dd
p2 + 2pq + q2 = 1
DD Dd
dd
dd = 0.70
Dd = 0.20 DD = 0.10
p2 + 2pq + q2 = 1
0.20 D 0.80 d
0.20 d 0.80 D
dd = 0.04
Dd = 0.32 DD = 0.64
F1 generation F2 generation
X X
No change in allele frequencies No change in genotype frequencies Evolution has not occurred
No change in allele frequencies Genotype frequencies change Evolution has not occurred
Allele frequencies change Genotype frequencies change Evolution has occurred Random mating
Genes, Populations, and
Evolution
• Causes of Microevolution
Genetic mutations
• The raw material for evolutionary change • Provide new alleles
• Some mutations might be more adaptive than others
– Ex: Genetic mutations affecting pigment color in
peppered moths have provided the variation needed for natural selection to occur
Genes, Populations, and
Evolution
• Causes of Microevolution
Gene Flow
(gene migration)
• Movement of alleles between populations when:
– Gametes or seeds (in plants) are carried into another population
– Breeding individuals migrate into or out of population
• Continual gene flow reduces genetic divergence between populations
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Pisum sativum Pisum arvense
gene flow
Genes, Populations, and
Evolution
• Causes of Microevolution
Genetic Drift
• Changes in the allele frequencies of a population due to change rather than selection by the environment
• Does not necessarily lead to adaptation to the environment • Occurs by disproportionate random sampling from population
– Can cause the gene pools of two isolated populations to become dissimilar
– Some alleles are lost and others become fixed (unopposed)
• Likely to occur:
– After a bottleneck
– When severe inbreeding occurs, or – When founders start a new population
• Stronger effect in small populations
Genetic Drift
17
natural disaster kills five green frogs
20% of population 10% of
population
Genes, Populations, and
Evolution
•
Bottleneck Effect
A random event prevents a majority of
individuals from entering the next generation
The next generation is composed of alleles
that just happened to make it
Genes, Populations, and
Evolution
•
Founder Effect
When a new population is started from just a
few individuals
The alleles carried by population founders are
dictated by chance
Formerly rare alleles will either:
• Occur at a higher frequency in the new population, or
• Be absent in new population
Bottleneck and Founder Effects
Change Allele Frequencies
a. b. c. d.
Original population gene pool = 3,800 alleles*
Remnant population gene pool = 90 alleles*
*1 marble = 10 alleles
26%
13% 8%
53%
11%
45% 44%
Genes, Populations, and
Evolution
•
Nonrandom Mating
When individuals do not choose mates
randomly
• Assortative mating:
– Individuals select mates with the same phenotype with respect to a certain characteristic
– Individuals reject mates with differing phenotype
– Increases the frequency of homozygotes for certain loci
16.2 Natural Selection
• Adaptation of a population to the biotic and
abiotic environment
Requires:
• Variation - The members of a population differ from one another
• Inheritance - Many differences are heritable genetic differences
• Differential Adaptiveness - Some differences affect survivability
• Differential Reproduction – Some differences affect the likelihood of successful reproduction
23
Natural Selection
• Results in:
A change in allele frequencies of the gene pool
Improved fitness of the population
24
Natural Selection
• Most traits are
polygenic
- variations in the trait
result in a bell-shaped curve
• Three types of selection occur:
(1) Directional Selection
• The curve shifts in one direction
– Bacteria become resistant to antibiotics
25
Natural Selection
• Three types of selection occur (cont):
(2)
Stabilizing Selection
• The peak of the curve increases and tails decrease • Example - human babies with low or high birth
weight are less likely to survive
(3)
Disruptive Selection
• The curve has two peaks
26
Three Types of Natural Selection
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Nu mbe r of Indiv idua ls Nu mbe r of Indiv idua ls stabilizing selection
a. b. c.
Peak narrows.
disruptive selection
Two peaks result. directional selection
Peak shifts.
27
Human Birth Weight
(stabilizing selection)
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Pe rc ent of B irths i n Popu latio n Pe rc ent In fant Mortality 20 15 10 5 10 20 30 50 70 100 5 7 3 2 .9 1.4 1.8 2.3 2.7
28
Directional Selection
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After More Time After Time Initial Distribution Merychippus b. Equus Hyracotherium a.
Body Size Body Size Body Size
29
Disruptive Selection
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Initial Distribution After Time After More Time Banding Pattern Banding Pattern Banding Pattern Nu mb er of Ind iv idu als Nu mb er of Ind iv idu als Nu mb er of Ind iv idu als b. a.
30
Natural Selection
•
Sexual selection
-
adaptive changes in
males and females lead to an increased
ability to secure a mate.
Males - increased ability to compete with other
males for a mate
31
Natural Selection
• Female Choice
Choice of a mate is a serious consideration because females produce few eggs
• Good genes hypothesis: Females choose mates on the basis of traits that improve the chance of survival.
• Runaway hypothesis: Females choose mates on the basis of traits that improve male appearance.
• Male Competition
Males can father many offspring because they continuously produce sperm in great quantity.
32
Dimorphism
• The drab
females tend to
choose
flamboyant
males as
mates.
33
Sexual Selection:
Competition
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34
Sexual Selection:
Competition
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a: © Y. Arthus-Bertrand/Peter Arnold, Inc.; b: © Neil McIntre/Getty Images
35
Natural Selection
• A study of sexual selection among humans
shows that female choice and male competition
apply to humans too
Women must invest more in having a child than men.
Men need only contribute sperm
• Generally more available for mating than are women.
More available men results in competition
Men also have a choice
36
16.3 Maintenance of Diversity
• Genetic Variability
Populations with limited variation may not be able to adapt to new conditions
Maintenance of variability is advantageous to the population
37
Maintenance of Diversity
• Natural selection causes imperfect adaptations
Depends on evolutionary history
Imperfections are common because of necessary compromises
• The environment plays a role in maintaining
diversity
Disruptive selection due to environmental differences promotes polymorphisms in a population
If a population occupies a wide range, it may have several subpopulations designated as subspecies
38
Subspecies Help Maintain Diversity
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.(E.o. lindheimeri, E.o. quadrivittata): © Zig Leszczynski/Animals Animals/Earth Scenes; (E.o. spiloides): © Joseph Collins/Photo Researchers, Inc.; (E.o. rossalleni): © Dale Jackson/Visuals Unlimited; (E.o. obsoleta): © William Weber/Visuals Unlimited
Pantheropsis obsoleta lindheimeri Pantheropsis obsoleta rossalleni Pantheropsis obsoleta spiloides
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Maintenance of Diversity
• Recessive alleles:
Heterozygotes shelter recessive alleles from selection
Heterozygotes allow even lethal alleles to remain in the population at low frequencies virtually forever
Sometimes recessive alleles confer an advantage to heterozygotes
• The sickle-cell anemia allele is detrimental in homozygote • However, heterozygotes are more likely to survive malaria • The sickle-cell allele occurs at a higher frequency in malaria
40
Maintenance of Diversity
•
Heterozygote Advantage
Assists the maintenance of genetic, and
therefore phenotypic, variations in future
generations.
In sickle cell disease heterozygous individuals
don’t die from sickle-cell disease, and they
41