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Chapter 23
The Evolution of Populations
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
: Population genetics:
foundation for studying
evolution
Microevolution is change in the genetic makeup of a population from
generation to generation
Population genetics is the study of how populations change genetically over time
Population genetics integrates Mendelian genetics with the Darwinian theory of
evolution by natural selection
This modern synthesis focuses on populations as units of evolution
Gene Pools and Allele
Frequencies
•
A population is a localized group of individuals capable of interbreeding and
producing fertile offspring
The gene pool is the total aggregate of genes in a population at any one time
The gene pool consists of all gene loci in all individuals of the population
The Hardy-Weinberg Theorem
The Hardy-Weinberg theorem describes a population that is not
evolving
It states that frequencies of alleles and genotypes in a population’s
gene pool remain constant from generation to generation, provided
that only Mendelian segregation and recombination of alleles are at
work
Mendelian inheritance preserves genetic variation in a
It describes a population where allele frequencies do
not change
populationHardy-Weinberg equilibrium describes a
population in which random mating occurs
Preservation of Allele Frequencies
In a given population where gametes contribute to the next generation
randomly, allele frequencies will not change.
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
And p2 and q2 represent the frequencies of the homozygous genotypes and 2pq
represents the frequency of the heterozygous genotype
LE 23-5
Gametes for each generation are
drawn at random from the gene pool
of the previous generation:
80% CR (p = 0.8)
20% CW (q = 0.2)
Sperm
CR
CW
(20%)
p2
pq
64%
CRCR
16%
CRCW
(20%)
CR
(80%)
CW
Eggs
(80%)
qp
4%
CWCW
16%
CRCW
q2
Conditions for Hardy-Weinberg
Equilibrium
The Hardy-Weinberg theorem describes a hypothetical population
In real populations, allele and genotype frequencies do change over timeThe
five conditions for non-evolving populations are rarely met in nature:
Extremely large population size
No gene flow
No mutations
Random mating
No natural selection
Population Genetics and Human
Health
We can use the Hardy-Weinberg equation to estimate the percentage of the
human population carrying the allele for an inherited disease
Mutation
Mutation and sexual recombination produce the variation that makes
evolution possible
Two processes, mutation and sexual recombination, produce the variation
in gene pools that contributes to differences among individuals
Mutations are changes in the nucleotide sequence of DNA
Mutations cause new genes and alleles to arise
Animation: Genetic Variation from Sexual Recombination
Mutations
A point mutation is a change in one base in a gene
It is usually harmless but may have significant impact on phenotype
Chromosomal mutations that delete, disrupt, or rearrange many loci are
typically harmful
Gene duplication is nearly always harmful
Mutation rates are low in animals and plants
The average is about one mutation in every 100,000 genes per generation
Mutations are more rapid in microorganisms
Sexual Recombination
Sexual recombination is far more important than mutation in producing
the genetic differences that make adaptation possible
Natural selection, genetic drift, and gene flow can alter
a population’s genetic composition
Three major factors alter allele frequencies and bring about most
evolutionary change:
Natural selection
Genetic drift
Gene flow
Differential success in reproduction results in certain alleles being passed to the
next generation in greater proportions
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
Animation: Causes of Evolutionary Change
LE 23-7
CWCW
CRCR
CRCR
CRCW
Only 5 of
10 plants
leave
offspring
CRCR
CWCW
CRCW
CWCW
CRCR
CRCW
CRCW
CRCR
CRCR
CRCR
CRCW
CRCW
Generation 1
p (frequency of CR) = 0.7
q (frequency of CW) = 0.3
CWCW
CRCR
Only 2 of
10 plants
leave
offspring
CRCR
CRCR
CRCR
CRCR
CRCR
CRCR
CRCR
CRCR
CRCW
CRCW
Generation 2
p = 0.5
q = 0.5
CRCR
CRCR
Generation 3
p = 1.0
q = 0.0
The Bottleneck Effect
The bottleneck effect is a sudden change in the environment that may
drastically reduce the size of a population
The resulting gene pool may no longer be reflective of the original
population’s gene pool
Understanding the bottleneck effect can increase understanding of how
human activity affects other species
LE 23-8
Original
population
Bottlenecking
event
Surviving
population
The Founder Effect/Gene Flow
The founder effect occurs when a few individuals become isolated from a
larger population
It can affect allele frequencies in a population
Gene flow consists of genetic additions or subtractions from a population,
resulting from movement of fertile individuals or gametes
Gene flow causes a population to gain or lose alleles
It tends to reduce differences between populations over time
Natural selection is the primary
mechanism of adaptive evolution
Natural selection accumulates and maintains favorable genotypes in a
population
Genetic Variation
Genetic variation occurs in individuals in populations of all species
It is not always heritable
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
LE 23-9
Map butterflies that
emerge in spring:
orange and brown
Map butterflies that
emerge in late summer:
black and white
Polymorphism
Phenotypic polymorphism describes a population in which two or more
distinct morphs for a character are represented in high enough frequencies
to be readily noticeable
Genetic polymorphisms are the heritable components of characters that
occur along a continuum in a population
Measuring Genetic Variation
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.
Most species exhibit geographic variation differences between gene pools
of separate populations or population subgroups
LE 23-10
1
2.4
3.14
8.11
9.12
10.16
5.18
6
13.17
19
1
2.19
3.8
4.16
9.10
11.12
13.17
15.18
5.14
7.15
XX
6.7
XX
•
Some examples of geographic variation occur as a cline, which is a graded
change in a trait along a geographic axis
Heights of yarrow plants grown in
common garden
Seed collection sites
A Closer Look at Natural
Selection
From the range of variations available in a population, natural selection
increases frequencies of certain genotypes, fitting organisms to their
environment over generations
Evolutionary Fitness
The phrases “struggle for existence” and “survival of the fittest” are
commonly used to describe natural selection but can be misleading
Reproductive success is generally more subtle and depends on many
factors
Fitness is the contribution an individual makes to the gene pool of the next
generation, relative to the contributions of other individuals
Relative fitness is the contribution of a genotype to the next generation,
compared with contributions of alternative genotypes for the same locus
Directional, Disruptive, and
Stabilizing Selection
Selection favors certain genotypes by acting on the phenotypes of certain
organisms
Three modes of selection:
Directional=Directional selection favors individuals at one end of the
phenotypic range
Disruptive-Disruptive selection favors individuals at both extremes of the
phenotypic range
Stabilizing-Stabilizing selection favors intermediate variants and acts against
extreme phenotypes
Frequency of
individuals
LE 23-12
Original
population
Evolved
population
Directional selection
Original population
Phenotypes (fur color)
Disruptive selection
Stabilizing selection
The Preservation of Genetic Variation
Various mechanisms help to preserve genetic variation in a population
Diploidy maintains genetic variation in the form of hidden recessive alleles
Balancing selection occurs when natural selection maintains stable
frequencies of two or more phenotypic forms in a population
Balancing selection leads to a state called balanced polymorphism
Some individuals who are heterozygous at a particular locus have greater
fitness than homozygotes=HETEROZYGUS ADVANTAGE-SICKLE
CELL AMEMIA
Natural selection will tend to maintain two or more alleles at that locus
LE 23-13
Frequencies of the
sickle-cell allele
0–2.5%
2.5–5.0%
5.0–7.5%
Distribution of
malaria caused by
Plasmodium falciparum
(a protozoan)
7.5–10.0%
10.0–12.5%
>12.5%
Selection
In frequency-dependent selection, the fitness of any morph declines if it
becomes too common in the population
Neutral variation is genetic variation that appears to confer no selective
advantage
Sexual selection is natural selection for mating success
It can result in sexual dimorphism, marked differences between the sexes in
secondary sexual characteristics
LE 23-14
On pecking a moth
image the blue jay
receives a food reward.
If the bird does not
detect a moth on
either screen, it pecks
the green circle to
continue a new set
of images (a new
feeding opportunity).
Parental population sample
0.6
Phenotypic
variation
Experimental group sample
0.5
0.4
Frequencyindependent control
0.3
0.2
0
Plain background
Patterned background
20
40
60
Generation number
80
100
Intrasexual selection is competition among individuals of one sex for
mates of the opposite sex
Intersexual selection occurs when individuals of one sex (usually females)
are choosy in selecting their mates from individuals of the other sex
Selection may depend on the showiness of the male’s appearance
The Evolutionary Enigma of Sexual
Reproduction
Sexual reproduction produces fewer reproductive offspring than asexual
reproduction, a so-called “reproductive handicap”
Sexual reproduction produces genetic variation that
may aid in disease resistance
LE 23-16
Asexual reproduction
Female
Sexual reproduction
Generation 1
Female
Generation 2
Male
Generation 3
Generation 4
Why Natural Selection Cannot
Fashion Perfect Organisms
Evolution is limited by historical constraints
Adaptations are often compromises
Chance and natural selection interact
Selection can only edit existing variations