Microevolution

Freeman Ch 23

Variation exists among individuals in a population

Some trait differences are heritable

All individuals do not survive & reproduce

Individuals with certain traits survive & produce

relatively more offspring

=
AdaptationsEvolution

= Fitness

descent w modification or gradual change over

Roadmap

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Outline
What is a population?
Genetic variation
Link to evolution
Creating new combinations
Mutation, Sexual reproduction

Agents of microevolution
Natural selection, Genetic drift, Gene flow,
mutation
Calculating allele frequencies in populations
Mendelian vs. population genetics
Hardy-Weinberg principle as a null hypothesis
Variables and conditions of the model

Scales of Evolution

All living organisms:

New species:

Populations:

Billions of years

Millions of years

10s--1000s of years
Microevolution is a
change in allele
frequencies in a
population over
generations

Population = a group of individuals of

the same species that live in the same
area, can interbreed, and vary in the
traits they posses
Natural selection acts on individuals,
but populations are the units that
evolve

Homo sapiens

Evolutionary Genetics
Variation in heritable traits is a
prerequisite for evolution
Genetic variation among individuals is
caused by differences in genes or
other DNA segments
Sexual reproduction can result in genetic
variation by recombining existing alleles
New genes and alleles can arise by
mutation or gene duplication

Fig. 13-8

A1

Parents

A1

A2

A3

Meiosis

Gametes

A1

A2

A3

Random
fertilization
Offspring,
with new
combinations
of alleles

A1

A2

A1

and

A3

Genotypes
Gene pool: all of the alleles for all of
the loci in all of the individuals of a
population
If 2+ alleles for loci,
then individual can be
homozygous or
heterozygous

Fixed if only one

allele for the locus in
the population

Allele Frequencies
Each allele has a frequency in the
population
Fraction of total alleles in a populations gene
pool
Each diploid individual contributes two alleles to
the gene pool for a particular locus

Ex. This individual

contributes one red allele
and one white allele to
the gene pool

 For example, consider a

population of 500 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 in the gene pool
CR (320 2) 160 800
CW (20 2) 160 200

What is the total # of alleles in

the gene pool?

p = dominant allele, q = recessive allele

To calculate the freqquency of each
allele: total # of each allele / total #
alleles in population
p freq CR 800 / (800 200) 0.8
q freq CW 200 / (800 200) 0.2

The sum of alleles is always 1 (or

100%, p + q = 1)
0.8 0.2 1

Agents of Microevolution
Genetic variation in a population must
exist in order for evolution to occur,
BUT variation does NOT guarantee
that the population will evolve
These factors alter allele frequencies
(p and q) and bring about most
evolutionary change

Natural Selection
Genetic drift
Gene flow
Mutation

Natural Selection
Differential reproductive success results
in certain alleles being passed to the
next generation in greater proportions
Can cause adaptive evolution, an
improvement in the match between
organisms and their environment

Tracking Modes of Selection

Use histograms to track allele
frequencies and visualize effects of
selection

Positive vs. Negative

Selection
Positive = the selective pressure that
causes an allele to increase in
frequency until fixation (100%)
Alleles that increase fitness experience +
selection

Negative = the selective pressure that

causes an allele to decrease in
frequency until it is lost (0%)
Alleles that decrease fitness experience selection

Direction of
Evolution

Colored
areas are
experienci
ng
negative
selection

Ex: Smaller size Mean size

has higher fitness has highest
fitness

Mean size
has lowest
fitness

Before selection

Directional
selection
tends to
reduce
genetic
diversity in a
population.

After selection

Number of individuals

(a) Directional selection changes the average value

of a trait.
Normal distribution
Low
fitness

High
fitness

Change in
average
value

Value of a trait

(b) For example, directional selection caused average

body size to increase in a cliff swallow population.

Percentage of birds

Original population
(N = 2880)

Change in
average
value

Body size class

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Survivors
(N = 1027)

Before selection

Number of individuals

(a) Stabilizing selection reduces the amount of variation in

a trait.
Low
fitness

High fitness

Low
fitness

Reduction
in variation

After selection

Value of a trait

20

100
70
50

Mortality

30
20

15

10

Very high
mortality
on extremes

10
7
5
3
2

Birth weight (pounds)

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10

11

Percentage of mortality

Percentage of newborn population

(b) For example, very small and very large babies are the most
likely to die, leaving a narrower distribution of birthweights.

Before selection

Number of individuals

(a) Disruptive selection increases the amount of

variation in a trait.
High
fitness

Low fitness

High
fitness

Increase in
variation
After selection

Value of a trait

(b) For example, only juvenile black-bellied seedcrackers

that had short or extremely long beaks survived long enough
to breed.

Number of individuals

Original population

Highest mortality
at intermediate
range
Survivors
Survivors

Beak length (mm)

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Balancing Selection
Multiple mechanisms
Heterozygote Advantage, Frequency-Dependent
Selection

When multiple alleles are maintained in a

gene pool through natural selection
Diploidy maintains genetic variation

Heterozygote Advantage
When heterozygotes have a higher
fitness than homozygotes
Example: sickle cell anemia also confers
malaria resistance

Frequency Dependent
Selection

Fitness of a phenotype depends on

its frequency relative to other
phenotypes

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Sexual Selection
A special type of natural selection, for
mating success
Cost of reproduction is higher for
females than for males
Eggs larger, more energetically expensive

In most species, males mate with

multiple females
Females become a limited resource over
which males must compete

the struggle between the individuals

of one sex (generally males) for the
possession of the other sex.
Charles Darwin

Tufted Coquettes

Sexual Selection
Traits under sexual selection exhibit adaptions
changes in males (and females) that lead to an
increased ability to secure a mate
Secondary sex characters:
Body size
Color morphs
Ornamental feathers
Weapons (horns, antlers, tusks)
These traits can exhibit sexual dimorphisms
Behaviors:
Fighting, mate choice, dances, vocalizations

Birds of Paradise
Bright plumage and elaborate
behaviors attract females
Sexual selection stronger than
natural selection

(b) Variation in reproductive success is high in males.

Males

Over 90% of the male seals

die without reproducing

Percentage of
males born

(a) Males compete for the opportunity to mate with

females.

A few successful males

father 81100 offspring

Lifetime reproductive success:

Number of offspring weaned

Percentage of
females born

(c) Variation in reproductive success is relatively low in

females.
Females
Only about 60% of the females
die without leaving offspring
The most successful females
produce 10 offspring

Lifetime reproductive success:

Number of offspring weaned

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 Intrasexual selection is direct

competition among individuals of one
sex (often males) for mates of the
opposite sex
Encroaches on energy for reproduction

Intersexual selection (mate choice)

occurs when individuals of one sex
(usually females) are choosy in
selecting their mates
Male showiness can increase a males
chances of attracting a female, while
decreasing his chances of survival

Genetic Drift
Changes in allele frequencies due to
chance or random events, not fitness
of organisms
Significant in small populations, often
reduces genetic variation
Alleles can be eliminated or fixed

Genetic Drift: The Bottleneck

Effect
A sudden reduction in population size
The resulting gene pool is different the
original populations gene pool

Genetic Drift: The Founder

Effect
When a few individuals become isolated
from a larger population
Allele frequencies in the small founder
population do not capture full diversity of
source population
Low genetic variation

Effect of Genetic Drift

(b) Population size = 400

In this population, the allele A1 drifted

to fixation in less than 20 generations

In this population,
the allele A1 was lost

Generation

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Frequency of allele A 1

Frequency of allele A 1

(a) Population size = 4

Generation

Gene Flow
The movement of alleles among
populations
Alleles transferred through the movement
of individuals or gametes
Gene flow tends to reduce variation
among populations over time (i.e.
connectivity)

 Gene flow is an important agent of

evolutionary change in modern human
populations

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Mutations
Permanent changes in DNA
sequence, an important evolutionary
process that adds new alleles
Point = change in single base pair
Chromosome = change in # or
composition of chromosomes
Lateral gene transfer = genes move
from one species to another

Relative fitness

Rate of mutation is slow, but beneficial mutations can spread

due to selection

Relative fitness (of descendant

generations compared with
original population) increased
in sudden bursts over time

Time (generations)

Lateral Gene Transfer can result in adaptive

novelties
(b) Origin of genes for carotenoid synthesis enzymes in
aphids

(a) Redgreen color polymorphism in pea aphids

Gray aphids
Fungus
genes

Lateral gene transfer Duplication

Diversification

Aphid
genes

1 mm

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Red aphids
Deletion

Green aphids

Three color morphs of aphids

are maintained through
frequency dependent
selection

The Hardy-Weinberg Model

A method to calculate genotypic
frequency
A null hypothesis for evolutionary
change

 Genetic variation in a population

must exist in order for evolution to
occur, BUT variation does NOT
guarantee that the population will
evolve
Are the allele frequencies of our
population changing? Why?

Selection
Genetic Drift
Gene flow (migration)
Mutation

Hardy-Weinberg Model
In a population where gametes contribute to
the next generation randomly & Mendelian
inheritance occurs, allele and genotype
frequencies remain constant from generation
to generation
Such a population is in Hardy-Weinberg
Equilibrium (HWE)
HWE is our null hypothesis
We predict there is no change
If our data fit the equilibrium, we can accept the
hypothesis

Hardy Weinberg Model

p2 + 2pq + q2 = 1
Equation built from a modified Punnett
square & gives the frequency of three
genotypes
p2 = frequency of homozygous dominant
2pq = frequency of heterozygote
q2 = frequency of homozygous recessive

Parental allele frequencies:

p = 0.7

q = 0.3

All eggs in gene pool

p = 0.7

All sperm
in gene
pool

p = 0.7

q = 0.3

p2 = 0.49

A1 A1
A1 A2
A2 A2

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p2 = 0.49
2pq = 0.42
q2 = 0.09

pq = 0.21

pq = 0.21

Offspring genotype
frequencies:

q = 0.3

q2 = 0.09

Allele
frequencies
have not
changed

Offspring allele
frequencies:

A1
A2

p = 0.49 +12 (0.42) = 0.70

1

q = 2(0.42) + 0.09 = 0.30

Parental allele
frequencies

Allele
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q p = pq

q q = q2

0.21 + 0.21 = 0.42

Offspring allele
frequencies

Offspring genotype
frequencies

p q = pq

p p = p2

A1

p = 0.7

Allele

A2

q = 0.3

Conditions for HWE

The Hardy-Weinberg theorem describes a
hypothetical population that is not evolving
In real populations, allele and genotype frequencies
do change over time
5 conditions for HWE are rarely met in nature:

These factors
have the most
direct effect
on allele
frequencies

1.Extremely large population sizes

2.Random mating
3.No mutations
4.No natural selection
5.No gene flow/migration

Large Breeding Population

Large populations are less effected by

Random Mating

Mutations

Natural Selection

Gene Flow

Hardy-Weinberg Equilibrium
1. Calculate allele frequencies for a
population
2. Predict expected genotype
frequencies as if the population is in
HWE

3. Calculate expected population

4. Compare predicted and observed
data
If calculated = actual, then population IS NOT
evolving
If calculated actual, then population IS evolving

Practice Case
Consider a population of 100
[diploid] wildflowers where
color gene exhibits
incomplete dominance
2 alleles, 3 phenotypes

Hardy-Weinberg Equilibrium
1. Calculate allele frequencies for
the population

100 wildflowers = 200 alleles for

that locus
Genotype

#
individuals

Red (RR)
Pink (Rw)
White
(ww)
Total

70
20
10

# Alleles
contributed to
gene pool
140
40
20

100

200

Hardy-Weinberg Equilibrium
1. Calculate allele frequencies for
the population

Frequency of R = p = 160/200 =
0.8
Frequency of w = q = 40/200 = 0.2

Genotyp
#
e
individuals
Red (RR)
70
Pink (Rw)
20
White
10
(ww)

#R
140
20
0

#w
0
20
20

Alleles

#R

160

#w

40

Total 200

Hardy-Weinberg Equilibrium
1. Calculate allele frequencies for
the population

Double check that p + q = 1

0.8 + 0.2 = 1

Hardy-Weinberg Equilibrium
2. Predict genotype frequencies if
the population is in HWE

Hardy-Weinberg Equilibrium
2. Predict genotype frequencies if
the population is in HWE

p2 + 2pq + q2 = 1

0.64 + 0.32 + 0.04 = 1

R
(p = 0.8)
R
RR
(p = 0.8) (p2 = 0.64)

w
(q = 0.2)
Rw
(pq =
0.16)
w
Rw
ww
(q = 0.2) (pq = 0.16) (q2 = 0.04)

Hardy-Weinberg Equilibrium
3. Calculate expected population

p2 = 0.64 x 100 individuals = 64 red

2pq = 0.32 x 100 individuals = 32 pink

q2 = 0.04 x 100 individuals = 4 white

Hardy-Weinberg Equilibrium
3. Compare calculated and actual
Calculated
Actual
64 red (p2 = 0.64) 70 red (p2 = 0.70)
32 pink (2pq =
20 pink (2pq =
0.32)
0.20)
4 white (q2 = 0.04)
10 white (q2 =
0.10)
Calculated Actual, so NOT at HWE