Hardy Weinberg Equilibrium

Carol E. Lee, University of Wisconsin-Madison

Copyright ©2020; do not upload without permission

Gregor Mendel

Wilhem Weinberg
(1822-1884) (1862 – 1937)
G. H. Hardy
(1877 - 1947)
 Lectures 4-12: Mechanisms of Evolution
(Microevolution)
• Hardy Weinberg Principle (Mendelian Inheritance)
• Genetic Drift
• Mutation
• Sex: Recombination and Random Mating
• Epigenetic Inheritance
• Natural Selection
These are mechanisms acting WITHIN populations,
hence called “population genetics”—EXCEPT for
epigenetic modifications, which act on individuals
in a Lamarckian manner
Recall from Previous Lectures
Darwin’s Observation

Evolution acts through changes in

allele frequency at each
generation

Leads to average change in

characteristic of the population
 Recall from Lecture on History of
Evolutionary Thought
Darwin’s Observation

HOWEVER, Darwin did not

understand how genetic
variation was passed on from
generation to generation
 Gregor Mendel, “Father of Modern
Genetics”
http://www.biography.com/people/gregor-mendel-39282#synopsis

Gregor Mendel

• Mendel presented a mechanism

for how traits got passed on

“Individuals pass alleles on to

their offspring intact”

(the idea of particulate (genes)

inheritance)
(1822-1884)
 Gregor Mendel, “Father of Modern Genetics”
http://www.biography.com/people/gregor-mendel-39282#synopsis

Mendel’s Laws of Inheritance

• Law of Segregation
only one allele passes from each parent on
to an offspring
• Law of Independent
Assortment
– different pairs of alleles are
passed to offspring
independently of each other
 Using 29,000 pea plants, Mendel discovered the 1:3
ratio of phenotypes, due to dominant vs. recessive
alleles

• In cross-pollinating plants with either yellow or green peas,

Mendel found that the first generation (f1) always had yellow
seeds (dominance). However, the following generation (f2)
consistently had a 3:1 ratio of yellow to green.
• Mendel uncovered the underlying
mechanism, that there are dominant and
recessive alleles
 Hardy-Weinberg Principle
Mathematical description of Mendelian inheritance

Jeremy Irons
playing GH Hardy in
the film “The Man
Who Knew Infinity” ↓

Wilhem Weinberg
Godfrey H. Hardy (1862 – 1937)
(1877-1947) German obstetrician-gynecologist
English Mathematician
Testing for Hardy-Weinberg equilibrium can
be used to assess whether a population is
evolving
 The Hardy-Weinberg Principle

• A population that is not evolving shows allele and

genotypic frequencies that are in Hardy Weinberg
equilibrium

• If a population is not in Hardy-Weinberg

equilibrium, it can be concluded that the
population is evolving
Evolutionary Mechanisms
(will put population out of HW Equilibrium):

• Genetic Drift
• Natural Selection
• Mutation
• Migration
*Epigenetic modifications change expression of alleles but not
the frequency of alleles themselves, so they won’t affect the
actual inheritance of alleles
However, if you count the phenotype frequencies, and not the
genotype frequencies , you might see phenotypic frequencies
out of HW Equilibrium due to epigenetic silencing of alleles.
(epigenetic modifications can change phenotype, not genotype)
Requirements of HW Evolution
Violation

Large population size Genetic drift

Random Mating Inbreeding & other

No Mutations Mutations

No Natural Selection Natural Selection

No Migration Migration

An evolving population is one that

violates Hardy-Weinberg Assumptions
 Fig. 23-5a

MAP

CANADA
AREA

ALASKA
Beaufort Sea
•What is a “population?”

NO RRIT
TE
RT OR
HW IE
A group of individuals within

ES S
Porcupine

T
a species that is capable of herd range
interbreeding and producing
fertile offspring

(definition for sexual species)

Fortymile
herd range

YUKON
ALASK A
In the absence of Evolution…

Patterns of inheritance should always be in

“Hardy Weinberg Equilibrium”

Following the transmission rules of Mendel

 Hardy-Weinberg Equilibrium
• According to the Hardy-Weinberg principle,
frequencies of alleles and genotypes in a
population remain constant from generation to
generation

• Also, the genotype frequencies you see in a

population should be the Hardy-Weinberg
expectations, given the allele frequencies
 “Null Model for No
Evolution”
• A population in Hardy-Weinberg Equilibrium
serves as the Null Model (for no evolution) to
test if evolution is happening
Example: Is this population in Hardy
Weinberg Equilibrium?

AA Aa aa
Generation 1 0.25 0.50 0.25
Generation 2 0.20 0.60 0.20
Generation 3 0.10 0.80 0.10
Hardy-Weinberg Theorem
In a non-evolving population,
frequency of alleles and genotypes
remain constant over generations

You should be able to

predict the genotype
frequencies, given the
allele frequencies
 important concepts
• gene: A region of genome sequence (DNA or RNA), that is
the unit of inheritance , the product of which contributes to
phenotype
• locus: Location in a genome (used interchangeably with
“gene,” if the location is at a gene… but, locus can be anywhere,
so meaning is broader than gene)
• loci: Plural of locus
• allele: Variant forms of a gene (e.g. alleles for different eye
colors, BRCA1 breast cancer allele, etc.)
• genotype: The combination of alleles at a locus (gene)
• phenotype: The expression of a trait, as a result of the
genotype and regulation of genes (green eyes, brown hair, body
size, finger length, cystic fibrosis, etc.)
 important concepts
• allele: Variant forms of a gene (e.g. alleles for different eye
colors, BRCA1 breast cancer allele, etc.)

• We are diploid (2 chromosomes), so we have 2 alleles

at a locus (any location in the genome)

• However, there can be many alleles at a locus in a

population.
– For example, you might have inherited a blue eye allele from
your mom and a brown eye allele from your dad… you can’t
have more alleles than that (only 2 chromosomes, one from
each parent)
– BUT, there could be many alleles at this locus in the
population, blue, green, grey, brown, etc.
 A2 A1
Eggs
• Alleles in a A1 A3 A2
population of A1
A4
diploid organisms A2
A3 A1
Sperm
A4

A1
A1
Random Mating (Sex)
A1A1 A1A1
Zygotes
A1A3
A1A1

• Genotypes A2A4

A3A1
 A2 A1
Eggs
A1 A3 A2
A1
A4
So then can we A2
A3 A1
predict the expected Sperm
A4
% of alleles and A1 A1
genotypes in the
population at each
generation? Zygotes
A1A1 A1A1

A1A3
A1A1

A2A4

A3A1
Hardy-Weinberg Theorem
In a non-evolving population,
frequency of alleles and genotypes
remain constant over generations
Fig. 23-6

Alleles in the population

Frequencies of alleles
Gametes produced
p = frequency of
Each egg: Each sperm:
CR allele = 0.8

q = frequency of
CW allele = 0.2 80% 20% 80% 20%
chance chance chance chance

Hardy-Weinberg proportions indicate the

expected allele and genotype frequencies,
given the starting frequencies
• By convention, if there are 2 alleles at a locus, p and
q are used to represent their frequencies

• The frequency of all alleles in a population will add

up to 1

– For example, p + q = 1
If p and q represent the relative frequencies of the
only two possible alleles in a population at a
particular locus, then for a diploid organism (2
chromosomes),
(p + q) 2 = 1

= p2 + 2pq + q2 = 1
– where p2 and q2 represent the frequencies of the
homozygous genotypes and 2pq represents the
frequency of the heterozygous genotype
What about for a triploid organism?
 What about for a triploid organism?
• (p + q)3 = 1

= p3 + 3p2q + 3pq2 + q3 = 1

Potential offspring: ppp, ppq, pqp, qpp,

qqp, pqq, qpq, qqq

How about tetraploid? You work it out.

 Hardy Weinberg Theorem
ALLELES
Probability of A = p p+q=1
Probability of a = q

GENOTYPES
AA: p x p = p2
Aa: p x q + q x p = 2pq
aa: q x q = q2

p2 + 2pq + q2 = 1
 More General HW Equations
• One locus three alleles: (p + q + r)2 = p2 + q2 + r2 + 2pq +2pr +
2qr

• One locus n # alleles: (p1 + p2 + p3 + p4 … …+ pn)2 = p12 + p22 +

p32 + p42… …+ pn2 + 2p1p2 + 2p1p3 + 2p2p3 + 2p1p4 + 2p1p5 + …
… + 2pn-1pn

• For a polyploid (more than two chromosomes):

(p + q)c, where c = number of chromosomes

• If multiple loci (genes) code for a trait, each locus follows the
HW principle independently, and then the alleles at each loci
contribute to and/or interact to influence the trait
 Calculating allele frequencies
We can then define the relative frequencies of A1A1 and A1A2
genotypes as:

Frequency of A1A1 = Number of A1A1 (N11)/Total Number N

Frequency of A1A2 = Number of A1A2 (N12)/Total Number N
Frequency of A2A2 = Number of A2A2 (N22)/Total Number N

Then, the proportion of Allele A1:

If there are two alleles, then the frequency of A2: 1 - p

 ALLELE Frequencies
Frequency of A = p = 0.8
Frequency of a = q = 0.2
p+q=1

Expected GENOTYPE Frequencies

AA: p x p = p2 = 0.8 x 0.8 = 0.64
Aa: p x q + q x p = 2pq
= 2 x (0.8 x 0.2) = 0.32
aa: q x q = q2 = 0.2 x 0.2 = 0.04

p2 + 2pq + q2
Allele frequencies remain the same at
next generation = 0.64 + 0.32 + 0.04 = 1
Expected Allele Frequencies at 2nd Generation
p = AA + Aa/2 = 0.64 + (0.32/2) = 0.8
q = aa + Aa/2 = 0.04 + (0.32/2) = 0.2
 Hardy Weinberg Theorem
ALLELE Frequency
Frequency of A = p = 0.8 p+q=1
Frequency of a = q = 0.2

Expected GENOTYPE Frequency

AA: pxp= p2 = 0.8 x 0.8 = 0.64
Aa: p x q + q x p = 2pq = 2 x (0.8 x 0.2) = 0.32
aa : qxq= q2 = 0.2 x 0.2 = 0.04

p2 + 2pq + q2 = 0.64 + 0.32 + 0.04 = 1

Expected Allele Frequency at 2nd Generation

p = AA + Aa/2 = 0.64 + (0.32/2) = 0.8
q = aa + Aa/2 = 0.04 + (0.32/2) = 0.2
Similar example,
But with different starting allele frequencies

p q
 p2
2pq
q2
 Calculating Allele Frequencies from # of Individuals

• The frequency of an allele in a population can be

calculated from # of individuals:

– For diploid organisms, the total number of alleles at

a locus is the total number of individuals x 2

– The total number of dominant alleles at a locus is 2

alleles for each homozygous dominant individual

– plus 1 allele for each heterozygous individual; the

same logic applies for recessive alleles
Calculating Allele and Genotype Frequencies from
# of Individuals
AA Aa aa
120 60 35 (# of individuals)
#A = (2 x AA) + Aa = 240 + 60 = 300
#a = (2 x aa) + Aa = 70 + 60 = 130
Proportion A = 300/total = 300/430 = 0.70
Proportion a = 130/total = 130/430 = 0.30
A + a = 0.70 + 0.30 = 1
Proportion AA = 120/215 = 0.56
Proportion Aa = 60/215 = 0.28
Proportion aa = 35/215 = 0.16
AA + Aa + aa = 0.56 + 0.28 +0.16 = 1
 Applying the Hardy-Weinberg Principle

• Example: estimate frequency of a disease allele in

a population
• Phenylketonuria (PKU) is a metabolic disorder that
results from homozygosity for a recessive allele

• Individuals that are homozygous for the deleterious

recessive allele cannot break down phenylalanine, results
in build up à mental retardation
• The occurrence of PKU is 1 per 10,000 births
• How many carriers of this disease in the
population?
– Rare deleterious recessives often remain in a
population because they are hidden in the
heterozygous state (the “carriers”)
– Natural selection can only act on the homozygous
individuals where the phenotype is exposed
(individuals who show symptoms of PKU)

– We can assume HW equilibrium if:

• There is no migration from a population with different
allele frequency
• Random mating
• No genetic drift
• Etc
 So, let’s calculate HW frequencies
• The occurrence of PKU is 1 per 10,000 births
(frequency of the disease allele):
q2 = 0.0001
q = sqrt(q2 ) = sqrt(0.0001) = 0.01

• The frequency of wildtype (normal) alleles is:

p = 1 – q = 1 – 0.01 = 0.99

• The frequency of carriers (heterozygotes) of the

deleterious allele is:
2pq = 2 x 0.99 x 0.01 = 0.0198
or approximately 2% of the U.S. population
Conditions for Hardy-Weinberg Equilibrium
• The Hardy-Weinberg theorem describes a
hypothetical population

• The five conditions for nonevolving populations

are rarely met in nature:
– No mutations
– Random mating
– No natural selection
– Extremely large population size
– No gene flow

• So, in real populations, allele and genotype

frequencies do change over time
 DEVIATION
from
Hardy-Weinberg Equilibrium
Indicates that

EVOLUTION
Is happening
 Hardy-Weinberg across a Genome

• In natural populations, some loci might be out of

HW equilibrium, while being in Hardy-Weinberg
equilibrium at other loci

• For example, some loci might be undergoing

natural selection and become out of HW
equilibrium, while the rest of the genome remains
in HW equilibrium
Allele A1 Demo
How can you tell whether a population
is out of HW Equilibrium?
• Perform HW calculations to see if it looks like
the population is out of HW equilibrium

• Then apply statistical tests to see if the

deviation is significantly different from what
you would expect by random chance
Example: Does this population remain in
Hardy Weinberg Equilibrium across
Generations?

AA Aa aa
Generation 1 0.25 0.50 0.25
Generation 2 0.20 0.60 0.20
Generation 3 0.10 0.80 0.10
 AA Aa aa
Generation 1 0.25 0.50 0.25
Generation 2 0.20 0.60 0.20
Generation 3 0.10 0.80 0.10

■ In this case, allele frequencies (of A and a) did not

change.

■ ***However, the population did go out of HW

equilibrium because you can no longer predict
genotypic frequencies from allele frequencies

■ For example, p = 0.5, p2 = 0.25, but in Generation 3,

the observe p2 = 0.10
How can you tell whether a population
is out of HW Equilibrium?

1. When allele frequencies are changing across

generations

2. When you cannot predict genotype

frequencies from allele frequencies (means
there is an excess or deficit of genotypes
than what would be expected given the allele
frequencies)
Example: Does this population remain in
Hardy Weinberg Equilibrium across
Generations?

AA Aa aa
Generation 1 0.25 0.50 0.25
Generation 2 0.24.5 0.50 0.25.5
Generation 3 0.25 0.50.5 0.24.5
 Testing for Deviaton from Hardy-
Weinberg Expectations
• A c2 goodness-of-fit test can be used to determine if a
population is significantly different from the expectations of
Hardy-Weinberg equilibrium.
• If we have a series of genotype counts from a population,
then we can compare these counts to the ones predicted by
the Hardy-Weinberg model.
• O = observed counts, E = expected counts, sum across
genotypes
 Example
• Genotype Count: AA 30 Aa 55 aa 15
• Calculate the c2 value:
Genotype Observed Expected (O-E)2/E
AA 30 33 0.27
Aa 55 49 0.73
aa 15 18 0.50
Total 100 100 1.50
• Since c2 = 1.50 < 3.841 (from Chi-square table, alpha = 0.05),
we conclude that the genotype frequencies in this population
are not significantly different than what would be expected if
the population is in Hardy-Weinberg equilibrium.
 Testing for Deviaton from Hardy-
Weinberg Expectations
• A c2 goodness-of-fit test can be used to determine if a
population is significantly different from the expectations of
Hardy-Weinberg equilibrium.
• If we have a series of genotype counts from a population,
then we can compare these counts to the ones predicted by
the Hardy-Weinberg model.
• O = observed counts, E = expected counts, sum across
genotypes

56
Testing for Deviaton from Hardy-Weinberg Expectations
• O = observed counts, E = expected counts, sum across
genotypes

• We test our c2 value against the Chi-square distribution (sum

of square of a normal distribution), which represents the
theoretical distribution of sample values under HW
equilibrium à Less likely to get
these values by
chance

• And determine how likely it is to get our result simply by

chance (e.g. due to sampling error); i.e., do our Observed
values differ from our Expected values more than what we
would expect by chance (= significantly different)? 57
 Test for Deviation from HW equilibrium
• Genotype Count Generation 4:
AA 65 Aa 31 aa 4
• Calculate the c2 value:
Genotype Observed Expected (O-E)2/E
AA 65 64.8 0.00062
Aa 31 31.4 0.0051
aa 4 3.8 0.0105
Total 100 100 0.016
• Since c2 = 0.016 < 3.841 (from Chi-square table for critical values,
alpha = 0.05), we conclude that the genotype frequencies in this
population are not significantly different than what would be
expected if the population were in Hardy-Weinberg equilibrium. 58
• The chi-squared distribution is used because it is the sum of squared
normal distributions
• Calculate Chi-squared test statistic
• Figure out degrees of freedom
• Select confidence interval (P-value)
• Compare your Chi-squared value to the theoretical
distribution (from the table) and accept or reject the null
hypothesis.
– If the test statistic > than the critical value, the null hypothesis (H0
= there is no difference between the distributions) can be rejected
with the selected level of confidence, and the alternative
hypothesis (H1 = there is a difference between the distributions)
can be accepted.
– If the test statistic < than the critical value, the null hypothesis
cannot be rejected 59
Test for Significance of Deviation from HW Equilibrium

Degrees of Freedom
is n – 1
= 2 alleles (p, q) -1 =
1

60
 Testing for significance
• The results come out not significantly different from HW
equilibrium
• This does not necessarily mean that genetic drift (or
some other evolutionary force) is not happening, but
that we cannot conclude that genetic drift is happening
• Either we do not have enough power (not enough data,
small sample size), or genetic drift is not happening
• Sometimes it is difficult to test whether evolution is
happening, even when it is happening... The signal
needs to be sufficiently large to be sure that you can’t
get the results by chance (like by sampling error)
61
 Test for Deviation from HW equilibrium
• Genotype Count Generation 4 à increase sample size
AA 65000 Aa 31000 aa 4000
• Calculate the c2 value:
Genotype Observed Expected (O-E)2/E
AA 65000 64800 0.617
Aa 31000 31400 5.10
aa 4000 3800 10.32
Total 100,000 100,000 16.04
• Since c2 = 16.04 > 3.841 (from Chi-square table for critical values,
alpha = 0.05), we conclude that the genotype frequencies in this
population ARE significantly different than what would be
expected if the population were in Hardy-Weinberg equilibrium. 62
Test for Significance of Deviation from HW Equilibrium

Degrees of Freedom
is n – 1
= 2 alleles (p, q) -1 =
1

63
• One generation of Random Mating could put a
population back into Hardy Weinberg
Equilibrium
Examples of Deviation from
Hardy-Weinberg Equilibrium
 What would Genetic Drift look
like?
• Most populations are experiencing some level
of genetic drift, unless they are incredibly
large
 Examples of Deviation from
Hardy-Weinberg Equilibrium
AA Aa aa
Generation 1 0.64 0.32 0.04
Generation 2 0.63 0.33 0.04
Generation 3 0.64 0.315 0.045
Generation 4 0.65 0.31 0.04

Is this population in HW equilibrium?

If not, how does it deviate?
What could be the reason?
 Examples of Deviation from
Hardy-Weinberg Equilibrium
AA Aa aa
Generation 1 0.64 0.32 0.04
Generation 2 0.63 0.33 0.04
Generation 3 0.64 0.315 0.045
Generation 4 0.65 0.31 0.04

This looks like a case of Genetic Drift, where allele

frequencies are fluctuating randomly across
generations; should perform a Goodness of Fit test
to determine whether the frequencies deviate
significantly from HW expectations
 Examples of Deviation from
Hardy-Weinberg Equilibrium

AA Aa aa
0.64 0.36 0

Is this population in HW equilibrium?

If not, how does it deviate?
What could be the reason?
 Examples of Deviation from
Hardy-Weinberg Equilibrium

AA Aa aa
0.64 0.36 0

This might be a case of Negative

Selection disfavoring aa
 Examples of Deviation from
Hardy-Weinberg Equilibrium

AA Aa aa
0.25 0.70 0.05

Is this population in HW equilibrium?

If not, how does it deviate?
What could be the reason?
 Examples of Deviation from
Hardy-Weinberg Equilibrium

AA Aa aa
0.25 0.70 0.05

This appears to be a case of Heterozygote

Advantage (or Overdominance)
 Examples of Deviation from
Hardy-Weinberg Equilibrium

AA Aa aa
0.10 0.10 0.80

Is this population in HW equilibrium?

If not, how does it deviate?
What could be the reason?
 Examples of Deviation from
Hardy-Weinberg Equilibrium

AA Aa aa
0.10 0.10 0.80

Selection appears to be favoring aa

 Summary

(1) A nonevolving population is in HW

Equilibrium

(2) Evolution occurs when the requirements for

HW Equilibrium are not met

(3) HW Equilibrium is violated when there is

Genetic Drift, Migration, Mutations, Natural
Selection, and Nonrandom Mating
 Hardy Weinberg Equilibrium

Gregor Mendel

Wilhem Weinberg
(1822-1884) (1862 – 1937)
G. H. Hardy
(1877 - 1947)
 Fig. 23-7-4
80% CR ( p = 0.8) 20% CW (q = 0.2)

Sperm
CR CW
(80%) (20%)
Perform the same
calculations using
(80%)
CR

percentages
Eggs

64% ( p2) 16% ( pq)

CR CR CR CW
16% (qp) 4% (q2)
(20%)
CW

CR CW CW CW

64% CR CR, 32% CR CW, and 4% CW CW

Gametes of this generation:

64% CR + 16% CR = 80% CR = 0.8 = p
4% CW + 16% CW = 20% CW = 0.2 = q
Genotypes in the next generation:

64% CR CR, 32% CR CW, and 4% CW CW plants

Fig. 23-7-1
80% CR (p = 0.8) 20% CW (q = 0.2)

Sperm
CR CW
(80%) (20%)
(80%)
CR
Eggs

64% (p2) 16% (pq)

CR CR CR CW

16% (qp) 4% (q2)

(20%)
CW

CR CW CW CW
Fig. 23-7-2

64% CRCR, 32% CRCW, and 4% CWCW

Gametes of this generation:

64% CR + 16% CR = 80% CR = 0.8 = p

4% CW + 16% CW = 20% CW = 0.2 = q

Fig. 23-7-3

64% CRCR, 32% CRCW, and 4% CWCW

Gametes of this generation:

64% CR + 16% CR = 80% CR = 0.8 = p

4% CW + 16% CW = 20% CW = 0.2 = q

Genotypes in the next generation:

64% CRCR, 32% CRCW, and 4% CWCW plants

1. Nabila is a Saudi Princess who is arranged to marry her
first cousin. Many in her family have died of a rare blood
disease, which sometimes skips generations, and thus
appears to be recessive. Nabila thinks that she is a carrier of
this disease. If her fiancé is also a carrier, what is the
probability that her offspring will have (be afflicted with) the
disease?

(A) 1/4
(B) 1/3
(C) 1/2
(D) 3/4
(E) zero
The following are numbers of pink and white flowers in a population.

Pink White
Generation 1: 901 302
Generation 2: 1204 403
Generation 3: 1510 504

2. Which of the following is most likely to be TRUE?

(A) The heterozygotes are probably pink
(B) The recessive allele here (probably white) is clearly deleterious
(C) Evolution is occurring, as allele frequencies are changing greatly over time
(D) Clearly there is a heterozygote advantage
(E) The frequencies above violate Hardy-Weinberg expectations
The following are numbers of purple and white peas in a population.
(A1A1) (A1A2) (A2A2)
Purple Purple White
Generation 1: 360 480 160
Generation 2: 100 200 200
Generation 3: 0 100 300

3. What are the genotype frequencies at each generation?

(A) Generation 1: 0.30, 0.50, 0.20
Generation 2: 0.20, 0.40, 0.40
Generation 3: 0, 0.333, 0.666
(B) Generation 1: 0.36, 0.48, 0.16
Generation 2: 0.10, 0.20, 0.20
Generation 3: 0, 0.10, 0.30
(C) Generation 1: 0.36, 0.48, 0.16
Generation 2: 0.20, 0.40, 0.40
Generation 3: 0, 0.25, 0.75
(D) Generation 1: 0.36, 0.48, 0.16
Generation 2: 0.36, 0.48, 0.16
Generation 3: 0.36, 0.48, 0.16
4. From the example on the previous slide, what are the
frequencies of alleles at each generation?

(A) Generation1: Dominant allele (A1) = 0.6, Recessive allele (A2) = 0.4
Generation2: Dominant allele = 0.4, Recessive allele = 0.6
Generation3: Dominant allele = 0.125, Recessive allele = 0.875

(B) Generation1: Dominant allele = 0.6, Recessive allele = 0.4

Generation2: Dominant allele = 0.6, Recessive allele = 0.4
Generation3: Dominant allele = 0.6, Recessive allele = 0.4

(C) Generation1: Dominant allele = 0.6, Recessive allele = 0.4

Generation2: Dominant allele = 0.5, Recessive allele = 0.5
Generation3: Dominant allele = 0.25, Recessive allele = 0.75

(D) Generation1: Dominant allele = 0.4, Recessive allele = 0.6

Generation2: Dominant allele = 0.5, Recessive allele = 0.5
Generation3: Dominant allele = 0.25, Recessive allele = 0.75
5. From the example two slides ago, which evolutionary
mechanism might be operating across generations?

(A) Mutation
(B) Selection favoring A1
(C) Heterozygote advantage
(D) Selection favoring A2
(E) Inbreeding
Answers:

1. Parents: Aa x Aa = Offspring: AA (25%), Aa (50%), aa (25%)

Answer = A
2. A
3. C
4. A
5. D