The Evolution of Populations: Powerpoint Lectures For

Copyright 2005 Pearson Education, Inc.
publishing as Benjamin Cummings

PowerPoint Lectures for
Biology, Seventh Edition
Neil Campbell and J ane Reece
Lectures by Chris Romero
Chapter 23
The Evolution of Populations
Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings
The Smallest Unit of Evolution
One common misconception about evolution is
that individual organisms evolve, in the
Darwinian sense, during their lifetimes
Natural selection acts on individuals, but
populations evolve
However, the evolutionary impact of natural
selection is only apparent in tracking how a
population of organisms changes over time.

Genetic variations in populations
Contribute to evolution
Figure 23.1
How do we study evolution?
Population genetics provides a foundation for
studying evolution
Microevolution
Is change in the genetic makeup of a
population from generation to generation
Figure 23.2
The Modern Synthesis
Population genetics--Is the study of how populations
change genetically over time
Reconciled Darwins and Mendels ideas and
Integrates Mendelian genetics with the Darwinian
theory of evolution by natural selection
What was missing in Darwins explanation was an
understanding of inheritance that could explain how chance
variations arise in a population while also accounting for the
precise transmission of these variations from parents to
offspring.
Although Gregor Mendel and Charles Darwin were
contemporaries, Mendels discoveries were unappreciated at
the time, even though his principles of heredity would have
given credibility to natural selection.
Gene Pools and Allele Frequencies
A population
Is a localized group of individuals that are capable of interbreeding and producing fertile
offspring
One definition of a species (among others) is a group of populations whose individuals have
the potential to interbreed and produce fertile offspring in a nature.
MAP
AREA

Fairbanks

Whitehorse
Fortymile
herd range
Figure 23.3
Populations of a
species may be
isolated from each
other, such that they
exchange genetic
material rarely, or
they may intergrade
with low densities in
an intermediate
region.
The gene pool
Is the total aggregate of genes in a population
at any one time
Consists of all gene loci in all individuals of the
population
Each locus is represented twice in the genome
of a diploid individual.
Individuals can be homozygous or
heterozygous for these homologous loci.
The Hardy-Weinberg Theorem
The Hardy-Weinberg theorem
Describes a population that is not evolving
States that the frequencies of alleles and
genotypes in a populations gene pool remain
constant from generation to generation
provided that only Mendelian segregation and
recombination of alleles are at work
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
Figure 23.4
Generation
1
C
R
C
R

genotype
C
W
C
W

genotype
Plants mate
All C
R
C
W

(all pink flowers)
50% C
R

gametes
50% C
W

gametes
Come together at random
Generation
2
Generation
3
Generation
4
25% C
R
C
R
50% C
R
C
W
25% C
W
C
W

50% C
R

gametes
50% C
W

gametes
Come together at random
25% C
R
C
R
50% C
R
C
W
25% C
W
C
W

Alleles segregate, and subsequent
generations also have three types
of flowers in the same proportions
A population in Hardy-Weinberg equilibrium

Figure 23.5
Gametes for each generation are drawn at random from
the gene pool of the previous generation:
80% C
R
(p = 0.8) 20% C
W
(q = 0.2)
Sperm
C
R

(80%)
C
W

(20%)
p
2

64%
C
R
C
R

16%
C
R
C
W

16%
C
R
C
W

4%
C
W
C
W

qp
C
R

(
8
0
%
)

E
g
g
s

C
W

(
2
0
%
)

pq
If the gametes come together at random, the genotype
frequencies of this generation are in Hardy-Weinberg equilibrium:
q
2

64% C
R
C
R
, 32% C
R
C
W
, and 4% C
W
C
W

Gametes of the next generation:
64% C
R
from
C
R
C
R
homozygotes
16% C
R
from
C
R
C
W
homozygotes
+ =
80% C
R
= 0.8 = p
16% C
W
from
C
R
C
W
heterozygotes
+ =
20% C
W
= 0.2 = q
With random mating, these gametes will result in the same
mix of plants in the next generation:
64% C
R
C
R
, 32% C
R
C
W
and 4% C
W
C
W
plants
p
2
4% C
W
from
C
W
C
W
homozygotes
If p and q represent the relative frequencies of
the only two possible alleles in a population at
a particular locus, then
p
2
+ 2pq + q
2
= 1
And p
2
and q
2
represent the frequencies of the
homozygous genotypes and 2pq represents
the frequency of the heterozygous genotype
Conditions for Hardy-Weinberg Equilibrium
The Hardy-Weinberg theorem
Describes a hypothetical population
In real populations
Allele and genotype frequencies do change
over time
Evolution does occur in relation to the
environment

The 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, phenyketonuria (PKU) in this case.
About 1 in 10,000 babies born in the United States
is born with PKU, which results in mental
retardation and other problems if left untreated.
The disease is caused by a recessive allele.
How is evolution possible?
Mutation and sexual recombination produce the
variation that makes evolution possible
Produce the variation in gene pools that
contributes to differences among individuals
Natural selection, genetic drift, and gene flow can
also alter a populations genetic composition

Mutation
Mutations
Are changes in the nucleotide sequence of DNA
Cause new genes and alleles to arise
Figure 23.6
Point Mutations
A point mutation
Is a change in one base in a gene
Can have a significant impact on phenotype
Is usually harmless, but may have an adaptive
impact
A deletion occurs when a chromosome fragment lacking
a centromere is lost during cell division.
This chromosome will be missing certain genes.
A duplication occurs when a fragment becomes attached
as an extra segment to a sister chromatid.
Copyright 2002 Pearson Education, Inc., publishing as Benjamin Cummings
Fig. 15.13a & b
Mutations That Alter Gene Number or Sequence
An inversion occurs when a chromosomal fragment
reattaches to the original chromosome but in the reverse
orientation.
In translocation, a chromosomal fragment joins a
nonhomologous chromosome.
Some translocations are reciprocal, others are not.
Fig. 15.13c & d
Mutations That Alter Gene Number or Sequence
Mutation Rates
Mutation rates
Tend to be low in animals and plants
Average about one mutation in every 100,000
genes per generation
Are more rapid in microorganisms
Over the long term, mutation is a very
important to evolution because it is the original
source of genetic variation that serves as the
raw material for natural selection.
Sexual Recombination
In sexually reproducing populations
sexual recombination is far more important
than mutation in producing the genetic
differences that make adaptation possible
Genes can not be definitively determined or
selected for by natural means but can be doen
via artificial selection and biotechnology
Natural Selection
Differential success in reproduction
Results in certain alleles being passed to the
next generation in greater proportions
Survival of the fittest, the most adaptive and
the luckiest.
Able to make more babies that survive than
the others
Accumulates and maintains favorable
genotypes in a population
Genetic Drift
Statistically, the smaller a sample
The greater the chance of deviation from a predicted result
Describes how allele frequencies can fluctuate unpredictably from
one generation to the next
Tends to reduce genetic variation
Figure 23.7
C
R
C
R

C
R
C
W

C
R
C
R

C
W
C
W
C
R
C
R

C
R
C
W

C
R
C
W

C
R
C
W
C
R
C
R

C
R
C
R

Only 5 of
10 plants
leave
offspring
C
W
C
W
C
R
C
R

C
R
C
W

C
R
C
R

C
W
C
W

C
R
C
W

C
W
C
W
C
R
C
R

C
R
C
W
C
R
C
W

Only 2 of
10 plants
leave
offspring
C
R
C
R

C
R
C
R
C
R
C
R

C
R
C
R
C
R
C
R

C
R
C
R

C
R
C
R

C
R
C
R

C
R
C
R
C
R
C
R

Generation 2
p = 0.5
q = 0.5
Generation 3
p = 1.0
q = 0.0
Generation 1
p (frequency of C
R
) = 0.7
q (frequency of C
W
) = 0.3
The Bottleneck Effect
In the bottleneck effect
A sudden change in the environment may
drastically reduce the size of a population
The gene pool may no longer be reflective of
the original populations gene pool
Original
population
Bottlenecking
event
Surviving
population
Figure 23.8 A
(a) Shaking just a few marbles through the
narrow neck of a bottle is analogous to a
drastic reduction in the size of a population
after some environmental disaster. By chance,
blue marbles are over-represented in the new
population and gold marbles are absent.
Human impacts on bottlenecked species
Understanding the bottleneck effect
Can increase understanding of how human
activity affects other species
Figure 23.8 B
(b)
Similarly, bottlenecking a population
of organisms tends to reduce genetic
variation, as in these northern
elephant seals in California that were
once hunted nearly to extinction.
The Founder Effect
The founder effect
Occurs when a few individuals become
isolated from a larger population due to
catastrophes or choice (early America)
Can affect allele frequencies in a population
Founder effects have been demonstrated in
human populations that started from a small
group of colonists and explorers.
Gene Flow
Gene flow
Causes a population to gain or lose alleles
Results from the movement of fertile
individuals or gametes
Tends to reduce differences between
populations over time
Genetic Variation
Genetic variation
Occurs in individuals in populations of all
species
Is not always heritable
Figure 23.9 A, B
(a) Map butterflies that
emerge in spring:
orange and brown
(b) Map butterflies that
emerge in late summer:
black and white
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 (red or white)
Quantitative characters
Vary along a continuum within a population (height,
length, etc.)
Quantitative variation is usually due to polygenic
inheritance in which the additive effects of two or more
genes influence a single phenotypic character.

Polymorphism

Phenotypic polymorphism
Describes a population in which two or more
distinct morphs for a character are each
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 the number of polymorphisms in a population
by determining the amount of heterozygosity at the
gene level and the molecular level
Gene diversity- Average heterozygosity
Measures the average percent of gene loci that are
heterozygous.
In the fruit fly (Drosophila), about 86% of their
13,000 gene loci are homozygous (fixed).
About 14% (1,800 genes) are heterozygous.
Nucleotide diversity measures the level of difference in
nucleotide sequences (base pair differences) among
individuals in a population.
In fruit flies, about 1% of the bases are different
between two individuals.
Two individuals would differ at 1.8 million of the 180
million nucleotides in the fruit fly genome.
Humans have relatively little genetic variation.
Gene diversity is about 14% in humans.
Nucleotide diversity is only 0.1%.
You and your neighbor have the same nucleotide
at 999 out of every 1,000 nucleotide sites in your
DNA.
Variation Between Populations
Most species exhibit geographic variation
Differences between gene pools of separate
populations or population subgroups
1 2.4 3.14 5.18 6 7.15
XX 19 13.17 10.16 9.12 8.11
1 2.19 3.8 4.16 5.14 6.7
XX 15.18 13.17 11.12 9.10
Figure 23.10
Some examples of geographic variation occur
as a cline, which is a graded change in a trait
along a geographic axis
Figure 23.11
EXPERIMENT Researchers observed that the average size
of yarrow plants (Achillea) growing on the slopes of the Sierra
Nevada mountains gradually decreases with increasing
elevation. To eliminate the effect of environmental differences
at different elevations, researchers collected seeds
from various altitudes and planted them in a common
garden. They then measured the heights of the
resulting plants.
RESULTS The average plant sizes in the common
garden were inversely correlated with the altitudes at
which the seeds were collected, although the height
differences were less than in the plants natural
environments.
CONCLUSION The lesser but still measurable clinal variation
in yarrow plants grown at a common elevation demonstrates the
role of genetic as well as environmental differences.
M
e
a
n

h
e
i
g
h
t

(
c
m
)

A
t
i
t
u
d
e

(
m
)

Heights of yarrow plants grown in common garden
Seed collection sites
Sierra Nevada
Range
Great Basin
Plateau
A Closer Look at Natural Selection
From the range of variations available in a
population
Natural selection increases the 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
Can be misleading
Reproductive success
Is generally more subtle and depends on many
factors
Types of FITNESS
Darwinian 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 as compared to the 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 are
Directional
Disruptive
Stabilizing
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
The three modes of selection
Fig 23.12 AC
(a) Directional selection shifts the overall
makeup of the population by favoring
variants at one extreme of the
distribution. In this case, darker mice are
favored because they live among dark
rocks and a darker fur color conceals them
from predators.
(b) Disruptive selection favors variants
at both ends of the distribution. These
mice have colonized a patchy habitat
made up of light and dark rocks, with the
result that mice of an intermediate color are
at a disadvantage.
(c) Stabilizing selection removes
extreme variants from the population
and preserves intermediate types. If
the environment consists of rocks of
an intermediate color, both light and
dark mice will be selected against.
Phenotypes (fur color)
Original population
Original
population
Evolved
population
The Preservation of Genetic Variation
Balancing selection
Occurs when natural selection maintains stable
frequencies of two or more phenotypic forms in a
population
Leads to a state called balanced polymorphism
Heterozygote advantage
Frequency dependent selection
Diploidy
Maintains genetic variation in the form of hidden
recessive alleles
Heterozygote Advantage
Some individuals who are heterozygous at a
particular locus
Have greater fitness than homozygotes --
greater survivorship and reproductive success
than homozygotes.
Natural selection
Will tend to maintain two or more alleles at that
locus
GENERALLY referred to as the HYBRID VIGOR

EXAMPLE of the heterozygote advantage

The sickle-cell allele
Causes mutations in hemoglobin but also
confers malaria resistance
Figure 23.13
Frequencies of the
sickle-cell allele
02.5%
2.55.0%
5.07.5%
7.510.0%
10.012.5%
>12.5%
Distribution of
malaria caused by
Plasmodium falciparum
(a protozoan)
Frequency-Dependent Selection
The fitness of any morph declines if it becomes too common
in the population
P
h
e
n
o
t
y
p
i
c

d
i
v
e
r
s
i
t
y

Figure 23.14
Parental population sample
Experimental group sample
Plain background Patterned background
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
to a new set of images (a
new feeding opportunity).
0.06
0.05
0.04
0.03
0.02
0 20 40 60 80 100
Generation number
Frequency-
independent control
Blue jays selected the more obvious moths which led to their
eventual modification of their shape and color so as NOT to be
eaten
The relationships between parasites and their hosts
often demonstrate this type of selection.
Hosts often vary in their defense against parasites and
parasites in their ability to infect hosts.
Those parasites that are capable of infecting the most
common host type will increase in abundance.
The rarer host types will increase as the genetic
frequencies in the parasite population shifts.
These shifts in genetic frequencies among hosts and
among parasites maintain variation in both populations.

Frequency-Dependent Selection
Host-parasite between clones of aquatic snails
and a parasitic worm.

Fig. 23.11
In these snails which reproduce asexually, the most common snail
clones suffer the higher infection rates than the least
common clone, suggesting
frequency-
dependent
selection.
Neutral Variation
Neutral variation
Is genetic variation that appears to confer no
selective advantage
For example, the diversity of human
fingerprints seems to confer no selective
advantage to some individuals over others.
Much of the protein and DNA variation
detectable by methods like electrophoresis
may be neutral in their adaptive qualities.

Sex is an evolutionary enigma.
It is far inferior to asexual reproduction as
measured by reproductive output.
If a population consisted of half sexual females
and half asexual females, the asexual condition
would increase.
All offspring of asexual females would be
reproductive daughters.
Only half of the offspring of sexual females would be
daughters; the other half would be necessary males.
Natural selection maintains sexual reproduction-WHY?
The Evolutionary Enigma of Sexual Reproduction
Sexual reproduction
Produces fewer reproductive offspring than asexual
reproduction, a so-called reproductive handicap
Figure 23.16
Asexual reproduction
Female
Sexual reproduction
Female
Male
Generation 1
Generation 2
Generation 3
Generation 4
Sexual Selection
Is natural selection for
mating success
Can result in sexual
dimorphism, marked
differences between
the sexes in
secondary sexual
characteristics- MALE
and Female!
However, some species
have very little physical
differences- which is boy
and which is girl cat?
Intrasexual selection
Is a direct 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
May depend on the showiness of the males
appearance
Figure 23.15
In most organisms,
it is FEMALE
choice that rules!
Why and HOW women choose.???
The underlying bases of female choice are probably
not aesthetic.
Current research is investigating the hypothesis that
females use these sexual advertisements to measure
the general health of a male.
Individuals with infections or other problems are likely
to have a relatively dull, disheveled plumage.
These individuals are unlikely to win many females.
For the female that chooses a healthy mate, even if his
inclination is just a prewired response to visual signals,
the benefit is a greater probability of having healthy
offspring.

Why have sex?
If sexual reproduction is a handicap, why has it
persisted?
It produces genetic variation that may aid in
disease resistance
Sex provides a mechanism for changing the
distribution of alleles and varying them among
offspring- due to recombination and
independent assortment
HOW to ensure genetic diversity and LESS chance of
INCEST/ sharing similar genetic stock?
MARRY outside of
your:
Community
City
County
Country
Continent
Culture
PLANET?

Jocey is Hispanic while Franky is Chinese
Why Natural Selection Cannot Fashion Perfect Organisms
Evolution is limited by historical constraints
Adaptations are often compromises
Chance and natural selection interact and may
not be adaptive all the time!
Selection can only edit existing variations

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