4 Evolution

Campbell’s Biology
Chapter 22: Overview, Concepts 22.1, 22.2
Chapter 23: Concepts 23.2, 23.3, 23.4
1
Chapter 24: Overview, Concepts 24.1, 24.2
 Hardy-Weinberg equation to test for
evolution in populations
• Population
– Group of
individuals of the
same species that
live in the same
area and
interbreed,
producing fertile
offspring

2
 Hardy-Weinberg equation to test for
evolution in populations
• Populations may be
isolated
geographically
– Not much exchange
of genetic material
• If overlapping
– Still interbreed
within population
more often than
not
3
 Hardy-Weinberg equation to test for
evolution in populations
• GENE POOL
– All copies of every type of allele at every locus in
all members of the population
• If only one allele exists for a particular locus in
a population, that allele is FIXED in the gene
pool
– All individuals homozygous for that allele
• If ≥2 alleles for a particular locus
– Individuals can be heterozygous or homozygous

4
 Hardy-Weinberg equation to test for
evolution in populations
• Each allele has a FREQUENCY in the population
– Proportion
• Example
– Population of 500 wildflowers
– 2 alleles: CR and CW code for flower pigment
– Each allele shows incomplete dominance

CRCR CRCW CWCW

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 Hardy-Weinberg equation to test for
evolution in populations
• 500 wildflowers
– 320 plants with red flowers
– 160 plants with pink flowers
– 20 with white flowers
• Diploid
– 1000 copies of flower colour gene in total

CRCR CRCW CWCW

6
 Hardy-Weinberg equation to test for
evolution in populations
• What is the frequency of the CR allele?
320 red flowers (2 x 320)
+ 160 pink flowers (1 x 160)
= 800
800/1000 = 0.8

CRCR CRCW CWCW

7
 Hardy-Weinberg equation to test for
evolution in populations
• Studying a locus with 2 alleles
– Use p to represent frequency of one allele
– Use q to represent frequency of other allele
• Frequency of CR allele will be p
• p = 0.8 or 80%
• Frequency of CW allele will be q
• q = 0.2 or 20%

CRCR CRCW CWCW

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 Hardy-Weinberg equation to test for
evolution in populations
• p + q = 1 (or 100%)
• Loci with >2 alleles
– Sum of all allele frequencies must still equal 1
• Allele and genotype frequencies used to test
whether evolution is occurring in a population

CRCR CRCW CWCW

9
 Hardy-Weinberg equation to test for
evolution in populations
• To test whether evolution is occurring
– Either due to natural selection or other factors
• Look at genetic makeup of population if NO
EVOLUTION occurring at particular locus
– Compare this with data from real population
• If no differences
– Real population NOT evolving
• If differences
– Real population IS evolving
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 Hardy-Weinberg equation to test for
evolution in populations
• Hardy-Weinberg equilibrium principle
– Frequencies of alleles and genotypes in a
population will remain constant from generation
to generation, provided that only Mendelian
segregation and recombination of alleles are at
work
• A gene pool may be in Hardy-Weinberg
equilibrium if it satisfies the above condition

11
 Hardy-Weinberg equation to test for
evolution in populations
• All alleles for given locus
from all individuals in
population put in one bin
– This bin holds gene pool
• Reproduction occurs by
randomly combining 2
alleles from bin
– Assumptions
• Mating is random
• All male-female matings
equally likely

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 Hardy-Weinberg equation to test for
evolution in populations
Wildflowers
• p = 0.8
• q = 0.2
• 800 CR alleles
• 200 CW alleles
• 1000 alleles total in bin

CRCR CRCW CWCW 13

 Hardy-Weinberg equation to test for
evolution in populations
• Each egg has
– 80% chance of receiving CR
allele
– 20% chance of receiving CW
allele
• Same is true for each
sperm

CRCR CRCW CWCW 14

Hardy-Weinberg equation to
test for evolution in populations

15
Hardy-Weinberg equation to test
for evolution in populations

• Use multiplication rule to

calculate frequencies of 3
possible genotypes
• p2 for CRCR
• 2pq for CRCW
• q2 for CWCW
• Genotype frequencies
MUST ADD UP TO 1
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 Hardy-Weinberg equation to test for
evolution in populations
• H-W equilibrium equation states that at locus with 2
alleles, 3 genotypes appear in following proportions:

p2 + 2pq + q2 = 1

• Sum must equal 1 in ANY population

– In equilibrium or not
• Genotype frequencies must equal above equation for
population to be in H-W equilibrium
– Allele frequencies will not change from generation to
generation
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 Hardy-Weinberg equation to test for
evolution in populations
Conditions for population to be in Hardy-
Weinberg equilibrium
1. No mutations
2. Random mating
3. No natural selection
4. Extremely large population size
5. No gene flow

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 Hardy-Weinberg equation to test for
evolution in populations
• Departure from any of the 5 conditions
results in evolution
– Evolution is common in real populations
• Some may be in H-W equilibrium for certain
loci only
• Use H-W equation to
– Test for evolution in a population
– Estimating % of population carrying allele for an
inherited disease

19
 Hardy-Weinberg equation to test for
evolution in populations
• Example with PKU (phenylketonuria)
– Recessive metabolic disorder
• 1/10 000 Canadian babies has PKU
• How many people are carriers in Canada?
• 1/10 000 is frequency of recessive homozygotes
• q2 = 0.0001
• q = 0.01
• p = 1- q = 0.99
• Carriers are heterozygotes (pq)
• 2pq = 2 (0.99)(0.01) = 0.0198
– 2% of Canadian population

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 Natural selection, genetic drift and
gene flow
• Remember the 5 conditions for Hardy-
Weinberg equilibrium
1. No mutations
2. Random mating
3. No natural selection
4. Extremely large population size
5. No gene flow

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 Natural selection, genetic drift, and
gene flow
• Natural selection, genetic drift, and gene flow
– Mechanisms that alter allele frequencies directly
– Cause most evolutionary change
• Natural selection violates condition 3 for H-W
equilibrium
• Genetic drift (which occurs more commonly in
smaller populations) violates condition 4
• Gene flow violates condition 5
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 Natural selection
Example: fruit fly Drosophila melanogaster
• Has allele for resistance to DDT
Before 1930s
• Fruit flies collected in the wild
– Allele for resistance at 0%
After 1960s
• Fruit flies collected in wild
– Allele for resistance had a frequency of 37%

23
 Genetic drift
• Chance events can cause allele frequencies to
fluctuate unpredictably from one generation
to the next, especially in small populations
• Coin flip probabilities
– The more times you flip the coin, the closer your
outcomes will come to theoretical probabilities
– 10 coin flips versus 1000 coin flips

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 Genetic drift
Allele frequencies affected by chance events
• With wildflowers, it could be a moose stepping on some
plants
• Could also happen during fertilization
– CRCW x CRCW could result in a small number of offspring
– None with CW allele

25
 Genetic drift
• Genetic drift can have a big impact on
populations in 2 main ways

1) Founder effect

2) Bottleneck effect

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 Genetic drift
FOUNDER EFFECT
• A few individuals isolated from a
bigger population
• Small group of individuals may start
their own population
– Gene pool of new population may differ
from that of source population

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 Genetic drift
FOUNDER EFFECT
• A few individuals blown from one island to
another by a big storm could result in the
founder effect
– Storm must indiscriminately transport some
individuals

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 Genetic drift
BOTTLENECK EFFECT
• Size of population drastically reduced
– Natural disaster or human actions
• Surviving population is no longer genetically
representative of original population

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 Genetic drift
BOTTLENECK EFFECT
• By chance
– Some alleles may be overrepresented among
survivors
– Other alleles underrepresented

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 Genetic drift
BOTTLENECK EFFECT

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 Genetic drift
• Is significant in small populations
• Can cause allele frequencies to change AT
RANDOM
• Can lead to loss of genetic diversity within a
population
• Can cause deleterious alleles to become fixed

32
 Gene flow
• Transfer of alleles into or out of population
due to movement of fertile individuals or their
gametes
• Reduces genetic differences between
populations
– Can result in two populations combining into one
• Example
– Pollination

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 Evolution
• Descent with modification
• Living species are descendants of ancestral
species that were different from the present-day
ones
• Also defined more narrowly as the change in the
genetic composition of a population from
generation to generation
• “Nothing in biology makes sense except in the
light of evolution”
– Theodosius Dobzhansky
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 Evolution
• Microevolution
– Evolutionary change below the species level (in a
population)
• Macroevolution
– Evolutionary changes that result in new species
(species-level and above)
• POPULATIONS are the smallest unit of
organism that can evolve
– Individuals DO NOT evolve

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 Natural selection
• There is variety in a population
• Individuals in a population that have traits
better suited to their environment tend to
produce more offspring
• Genetically, selection results in alleles being
passed to the next generation in proportions
that DIFFER from those in present generation

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 Natural selection
• Peppered moth (Biston betularia) in England
during the industrial revolution

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 Evolution
• Darwin was inspired by his travels on the
Beagle as well as the works of
– Linnaeus
– Cuvier
– Malthus
– Hutton
– Lamarck
– Lyell
– Wallace

*You do not have to memorize all these extra names, but know who Darwin is 38
39
 Evolution
• ADAPTATIONS
– Inherited characteristics of organisms that
enhance their survival and reproduction

40
 Evolution
• Darwin’s finches
– Beaks adapted to food source on particular island

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 Evolution
• Observation 1: Members of a population vary
in their inherited characteristics
• Observation 2: All species can produce more
offspring than their environment can support

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 Evolution
• Inference 1: Individuals whose inherited traits
give them a higher probability of surviving
and reproducing in a given environment tend
to leave more offspring than other individuals
• Inference 2: This unequal ability of individuals
to survive and reproduce will lead to
accumulation of favourable traits in the
population over generations

43
 Evolution
• NATURAL SELECTION
– Individuals that have certain heritable traits
survive and (because of those traits) reproduce at
higher rate than others
– Over time, natural selection can increase match
between organisms and environment
– If environment changes
– If individuals move to new environment
• Natural selection may result in adaptation to new
conditions, sometimes giving rise to new species
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 Evolution
• Evidence for evolution by natural selection in
1. Fossils
2. Comparative embryology
3. Evolution of drug-resistant bacteria
4. Molecular evidence
5. Homology
• Similarity in characteristics resulting from a shared
ancestry
6. Artificial selection
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Evolution

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 Evolution
• COMPARATIVE EMBRYOLOGY

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 Evolution
• DRUG-RESISTANT BACTERIA

• MOLECULAR EVIDENCE
– The more similar 2 species are, the higher the % of
DNA they will have in common
– True for proteins as well

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 Evolution
• HOMOLOGY
– Similarity in characteristics resulting from a shared
ancestry

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 Evolution
• ARTIFICIAL SELECTION

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 Evolution
• DIVERGENT EVOLUTION
– Population becomes separated from rest of the
species
– Different selection pressures, evolutionary pattern
– The result of divergent evolution is HOMOLOGY

51
 Evolution
• CONVERGENT EVOLUTION
– Organisms occupy similar environments
– Evolve similarities
• Not from a shared ancestry
– The result of convergent evolution is ANALOGY
– Analogy
• Similarity, but not from common ancestry
– Example
• Insect wings and bird wings arose independently

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 Evolution
• CRYPTIC COLOURATION
– Camouflage

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 Evolution
• APOSEMATIC COLOURATION
– Warning colouration

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 Evolution
• Some species protected by
resemblance to other
species
• BATESIAN MIMICRY
– Palatable or harmless
species mimics unpalatable
or harmful species

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 Evolution
• MÜLLERIAN MIMICRY
– 2 or more unpalatable
species resemble each
other
– Black and yellow or red
stripes characterize
unpalatable animals from
insects to snakes

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 Evolution
• RELATIVE FITNESS
– Contribution an individual makes to the gene pool
of the next generation relative to the
contributions of other individuals
• Natural selection can alter frequency
distribution of heritable traits
1. Directional
2. Disruptive
3. Stabilizing

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 Evolution
• DIRECTIONAL SELECTION occurs when
conditions favour individuals exhibiting one
extreme of a phenotypic range

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 Evolution
• DISRUPTIVE SELECTION occurs when
conditions favour individuals at both extremes
of a phenotypic range

59
 Evolution
• STABILIZING SELECTION acts against both
extreme phenotypes and favours intermediate
variants
– Reduces variation

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 Evolution
• Natural selection refers to selection by
– Environment
– Non-random mating (sexual selection)
• Can result in sexual dimorphism
– Differential fertility
• Early maturation, for example

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Evolution
• SPECIATION
– Process by which one
species splits into two
species
• BIOLOGICAL SPECIES
CONCEPT
– Group of populations
whose members have
the potential to
interbreed in nature and
produce viable, fertile
offspring
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 Evolution
• Formation of a new species hinges on
REPRODUCTIVE ISOLATION
– Existence of biological factors that impede
members of two species from interbreeding and
producing viable, fertile offspring
– Barriers may be PRE-ZYGOTIC or POST-ZYGOTIC
– Pre-zygotic
• Block fertilization
– Post-zygotic
• Contribute to reproductive isolation after hybrid
formed
64
 PRE-ZYGOTIC BARRIERS
• Habitat isolation
– Two species occupy different habitats in
same area
– Encounter one another rarely
– Example
• Two garter snake species
• One lives mainly in water
• Other mainly on land

65
 PRE-ZYGOTIC BARRIERS
• Temporal isolation
– Species breed during different
• Times of day
• Seasons
• Years
– Example
• Ranges of western spotted skunk and
eastern spotted skunk overlap
• Western spotted skunk mates in late summer
• Eastern spotted skunk mates in late winter

66
 PRE-ZYGOTIC BARRIERS
• Behavioural isolation
– Courtship rituals and other behaviours
may be unique to species
– Enable mate recognition
– Example
• Blue-footed boobies mate only after species-
specific courtship display

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 PRE-ZYGOTIC BARRIERS
• Mechanical isolation
– Morphological differences prevent mating
from completing
– Example
• Two snails with shells spiraling in different
directions
• Genital openings cannot align

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 PRE-ZYGOTIC BARRIERS
• Gametic isolation
– Sperm of one species may not be able to
fertilize egg of another species
• Maybe sperm cannot survive in reproductive tract
of female
• Maybe sperm cannot penetrate membrane
surrounding egg
– Example
• Similar species of sea urchin release gametes into
surrounding water
• Sperm and eggs from different species cannot fuse
due to proteins on surface of both not binding well
together
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 POST-ZYGOTIC BARRIERS
• Reduced hybrid viability
– Hybrids may not develop completely, or
are very frail

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 POST-ZYGOTIC BARRIERS
• Reduced hybrid fertility
– Hybrid may survive well, but is sterile and
cannot produce normal gametes
– Example
• Donkeys and horses may mate to produce a
mule
• Mule is sterile

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 POST-ZYGOTIC BARRIERS
• Hybrid breakdown
– First generation hybrid may be viable and
fertile
– When hybrids mate with each other or
with either parent species, next
generation is frail or sterile

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 Evolution
• Speciation may be
– ALLOPATRIC
• Gene flow interrupted
when populations are
isolated geographically
• Whether something is a
barrier or not depends on
the species

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Evolution

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 Evolution
• Speciation may be
– SYMPATRIC
• Occurs in populations that
live in same geographic
area
1. Polyploidy
2. Habitat differentiation
3. Sexual selection

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 Evolution
Sympatric speciation by
• POLYPLOIDY
– Accident during cell
division that results in
extra sets of
chromosomes
– More common in plants
than animals
– Grey tree frog thought
to originate this way
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Evolution
Sympatric speciation by
• HABITAT DIFFERENTIATION
– Genetic factors enable subpopulation to exploit
habitat or resource not used by parent population
– North American apple maggot fly
• Arose from population that fed on hawthorns
• Some colonized apple trees introduced by Europeans
(apple trees matured earlier)
• Now temporal isolation between the two species

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 Evolution
Sympatric speciation by
• SEXUAL SELECTION
– Cichlid fish
• 2 closely related sympatric species differ mainly in colour
of breeding males

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