CS 563 September 17, 2018: Instructor: Antonia Tetteh

CS 563
Lecture 2
Instructor: Antonia Tetteh
September 17, 2018
CS 563 Population Genetics, 1

KNUST
Today’s lecture
• Quantifying genetic variation

• Deriving genotypic and allelic frequencies
• Hardy-Weinberg equilibrium
CS 563 Population and 2

Quantitative Genetics, KNUST
Quantifying genetic variation
Variation at a single locus
• If we want to have sufficient description of the genetic
constitution of a population at a single locus, we need to
specify two things:
• 1. What genotypes are present
• 2. How many of each genotype there are in the population
• The genetic constitution of a population, refers to

• 1. The genes it carries,
• 2. The array of gene frequencies, that is, specification of
the alleles present at every locus, and
• 3. The numbers or proportions of the different alleles at
each locus.
CS 563 Population and Quantitative 3

Genetics, KNUST
Let’s take an example from the Australian Aborigine population

having the MN blood group. From this population, 730
individuals were screened for their genotype. This variation is
protein variation of red blood cell antigens using
immunological techniques. The data is presented below:
Blood group No. of Genotype Notation
Individuals frequency
MM 22 22/730 = 0.030 P
MN 216 216/730= 0.296 H
NN 492 492/730= 0.674 Q
Total 730 1.000 P+H+Q=1

• Genotype frequencies give the description of a population at

one time. It is not a description of a breeding group, since
whole genotypes are not transmitted between generations
• Alleles (also genes) are the entities transmitted across

generations. We can describe the genetic constitution of a
population by specifying the allele frequencies

Calculating allele frequencies (diploid individuals )
Blood group No. of No. of alleles
Individuals M N
MM 22 44 0
MN 216 216 216
NN 492 0 984
Total 730 260 1200

Total no. of alleles: 260 + 1,200 = 1,460
Computing allele frequencies:
Frequency of M allele
f(M) = p = 260/1460 = 0.178
f(N) = q = 1200/1460 = 0.822

• We can also compute the allelic frequencies straight from the

genotype frequencies
• Let p=f(M) and q=f(N). Thus, p=f(MM) + ½ f(MN) and

q=f(NN) + ½ f(MN)
From the data

p= 0.030 + ½( 0.296) =0.178
q= 0.674 +½(0.296) =0.822
So p=P + H/2 and

q=Q + H/2
Note that p + q =1
Quantifying genetic variation
Variation at many loci (multilocus measures)
• At more than one locus, two parameters are used to
quantify variation.
• 1) Determine the proportion of polymorphic loci, P
• 2) Determine the heterozygosity of the population
averaged over all loci
• H = mean heterozygosity.
• Heterozygosity is an individual or population-level
parameter
• In a population, a polymorphic gene is one for which the
most common allele has a frequency of less than 0.95
Some authors prefer a more stringent cutoff at 0.99
• Conversely, a monomorphic gene is one that is not
polymorphic, has a frequency of most common allele >0.95;
or frequency of least common allele <0.05

Proportion of polymorphic loci
• The cutoff at 0.95 (sometimes 0.99) in the definition of

polymorphism is arbitrary, but it serves to focus attention on
those genes in which allelic variation is most common
• Suppose we have data for 5 allozyme loci for Zea mays.

Genotype Allozyme Loci
Adh Mdh Pgi-A Pgm Ldh
FF 32 1 50 8 0
FS 16 9 0 24 1
SS 2 40 0 18 49
Total 50 50 50 50 50
f(F) 0.80 0.11 1.00* 0.40 0.01*

f(FS) 0.32 0.18 0.00 0.48 0.02
f(S) 0.20 0.89 0.00 0.60 0.99

For Adh
Genotype No. of individuals F S
FF 32 64
FS 16 16 16
SS 2 0 4
Total 50 80 20
Total no. of alleles = 80 + 20 = 100
f(F) = 80/100 = 0.80
f(S) = 20/100 = 0.20
f(FS) = 16/50 = 0.32

• Proportion of polymorphic loci
The loci that show variation are Adh, Mdh, Pgm (most
common allele has a frequency of less than 0.95)
Both Pgi and Ldh are monomorphic (frequency of least
common allele <0.05)
Polymorphic loci =3; total no. of loci evaluated =5
P = proportion of polymorphic loci = 3/5 = 0.60
• Mean heterozygosity
Take the average of the frequencies of the heterozygous
genotypes across all loci
H = [f(FS)Adh + f(FS)Mdh + f(FS)Pgi + f(FS)Pgm + f(FS)Ldh]/5

= (0.32 + 0.18 + 0.00 + 0.48 + 0.02)/5 = 0.20
From this calculation, the mean frequency of heterozygotes in any
locus in this population is 20%

Heterozygosity
• Heterozygosity provides a great deal of information
about the structure and even history of a population.
• High heterozygosity means lots of genetic variability.
• Low heterozygosity means little genetic variability.
• We often compare the observed level of heterozygosity,
Ho to what we expect under Hardy-Weinberg
equilibrium (HWE).
• If the observed heterozygosity is lower than expected,
we seek to attribute the discrepancy to forces such as
inbreeding. If heterozygosity is higher than expected, we
might suspect an isolate-breaking effect or admixture,
and heterozygosity maintained by a balancing selection
(the mixing of two previously isolated populations).
Heterozygosity
• Several measures of heterozygosity exist. The value of
these measures will range from zero (no heterozygosity)
to nearly 1.0 (for a system with a large number of equally
frequent alleles). Heterozygosity (expected): The
proportion of loci expected to be heterozygous in an
individual
HO (observed heterozygosity) is the observed proportion
of heterozygotes, averaged over loci.
HE (expected heterozygosity) is also known as gene
diversity (= D; preferred, less ambiguous term) and is
calculated as 1.0 minus the sum of the squared gene
frequencies

Heterozygosity
• HE, or Nei’s gene diversity, D. The

simplest way to calculate it for a single
locus is as
HE  1  p 2
i

• On the whole, there is a positive relationship between

amount of polymorphism and degree of heterozygosity
• This relationship is expected because the greater the

proportion of polymorphic genes in a population, the more
genes that are expected to be heterozygous on average
• Vertebrates have the lowest average amount of variation

among the groups
• Plants come next and invertebrates have the highest

Population
• In population genetics, the word population does not usually

refer to an entire species; it refers instead to a group of
organisms of the same species living within a sufficiently
restricted geographical area and any member can
potentially mate with any other member (provided that
they are of the opposite sex)
• In the transmission the genotypes from parents, a new set of

genotypes is constituted in the progeny, from the genes
transmitted in the gametes. The genes carried by the
population thus have continuity from generation to generation,
but the genotypes in which they appear do not.

The ideal population: the Hardy-Weinberg Equilibrium
(HWE)
• Populations are dynamic groups, that change over space and

time. The change may arise from
– Genetic forces
– Ecological forces
– Evolutionary forces
• As population geneticists, we are interested in modeling this

change. To do this, we need a null (basal) population to
compare our empirical and theoretical findings to, i.e., a
reference population where no change occurs.
• We thus define an ideal population, where nothing happens,
as our null/basal population, or reference population and
mating and movement of individuals around is uniform
(panmixia)
(HWE)
The ideal population that we imagine has the following
properties:
• The organisms are diploid
• Reproduction is sexual and gene frequencies are equal
between males and females
• Discrete, nonoverlapping generations
• Random mating with respect to genotype in question
• The population is panmictic (couples form randomly
(panmixia), and their gametes encounter each other
randomly (pangamy))
• Infinite population size
• No migration
• No mutation
• No natural selection
(HWE)
• If we have this ideal population, we expect to see the

following properties:
• 1. The population is stable with respect to gene and

genotype frequencies. There is no tendency for the
population to change from generation to generation.
Mendelian segregation and random mating preserves the
existing genetic variation in the absence of other forces
• 2. In this ideal population, the genotype frequencies in the

progeny produced by random mating among the parents is
determined solely by the gene frequencies of the parents
• An ideal population that has these properties is said to be in
Hardy-Weinberg equilibrium (HWE)

Testing the population properties
• That the population is stable, that is, gene frequencies do

not change generation after generation
Consider a locus A, with two alleles A1 and A2. The frequencies
of the two alleles are given as
f(A1) = p = P+H/2
f(A2) = q = Q+H/2
where the frequencies of the genotypes A1A1, A1A2 and A2A2
are P, H, Q, respectively

1. Let’s see what happens in the next generation. The random

union of gametes is shown in the table below:
Male parent gametes
A1(p) A2(q)
Female A1(p) A1A1(p2) A1A2(pq) progeny
Parent A2(q) A1A2(pq) A2A2(q2) progeny
Gametes
Thus, the genotype frequencies of the progeny are
f(A1A1) = P = p2
f(A1A2) = H = 2pq
f(A2A2) = Q = q2

• 2. The gene frequencies of the progeny can thus be derived
from their genotype frequencies
f(A1) = P + H/2
= p2 + 2pq/2
= p2 + pq
= p(p +q)
=p
and
f(A2) = Q + H/2
= q2 + 2pq/2
= q2 + pq
= q(q +p) Thus the gene frequencies of the
=q progeny are equal to the gene
since p+q=1 frequencies of the parents - nothing
has changed. Population is stable.

• For autosomal loci, it takes one generation of random mating
to get a population in HWE
• Testing for Hardy-Weinberg Equilibrium
Given population genotype frequency data for a gene, we can
establish whether these frequencies are consistent with the
population being at HWE at this locus, i.e., genotype
frequencies in the progeny determined solely by gene
frequencies of the parents
The following data is for genotypes at the MN blood group for

individuals in a human population
Genotype MM MN NN Total
Observed 22 216 492 730
Calculate the observed gene frequencies from the data
CS 563 Population and

24
Hypothesis
• Genotype frequencies have not changed over

one generation of random mating =H0-the null
hypothesis
• If this is true, then we have Hardy-Weinberg
equilibrium
• The alternative hypothesis, HA = genotype
frequencies have changed over one generation
of random mating

Testing for Hardy-Weinberg Equilibrium
p = f(M) = [#MM + ½(#MN)]/Total
= [22 +½(216)]/730
= 0.178
So q = 1 - p = 0.822
• p means that for any zygote, the probability that the egg will
contain allele M is 0.178
• q means that for any zygote, the probability that the egg will
contain allele N is 0.822
2. Given these gene frequencies, calculate the genotype

frequencies expected from these gene frequencies under
HWE
Expected gene frequencies:
f(MM) = p2= (0.178)2 = 0.0317
f(MN) = 2pq = 2(0.178)(0.822) = 0.2926
f(NN) = q2 = (0.822)2 = 0.6757
• 3. Now that we have the expected genotype frequencies after

first generation of random mating, we can calculate the
expected number of individuals per genotype given the
observed total sample size (n=730)
• E(#MM) = p2 × total = (0.0317)(730) = 23.14
E(#MN) =2pq × total = (0.2926)(730) = 213.60
E(#NN) =q2 × total = (0.6757)(730) = 493.26
• 4. Let’s recap at this point:

Genotype Expected # Observed #
MM 23.14 22
MN 213.60 216
NN 493.26 492
• We can compare the observed and expected genotype
numbers using the χ2 goodness-of-fit test.
 ( observed  exp ected ) 2

2 
for all classes exp ected
(22  23.14)2 (216  213.60)2 (492  493.26)2

 
2
   0.086
23.14 213.60 493.26
The number of degrees of freedom for this sample is 1 [df = # of
classes - # of constraints; 3 – 1 (for calculation of q from p) -1
(for data)]
The critical Χ2 at 5% significance level for df=1 is 3.84. The Χ2est of
0.086 is < Χ2 crit of 3.84. so the genotype frequencies have not
changed over one generation of random mating. The
population is in Hardy-Weinberg equilibrium with respect to the
MN blood group
Estimating heterozygote frequencies for dominant alleles
• In situations where one desires to know the numbers of

heterozygous individuals at a particular locus in a population,
by assuming that the population is in Hardy-Weinberg
equilibrium allows the calculation of the frequency of the
heterozygotes
• For example, a dominant allele A1 at a locus A, produces

A1A1, A1A2, and A2A2 where cannot phenotypically distinguish
A1A homozygotes from A1A2 heterozygotes
• The example below illustrates how one can calculate the

heterozygote frequency by assuming HWE

Estimating heterozygote frequencies for dominant alleles
Industrial melanism in the moth

• In a heavily polluted area in Biston betularia
Birmingham, UK, the
melanin (black) Biston
betularia moths account for
87% of the moth population.
The allele leading to
melanism, B, is dominant
over the recessive type, b
• Since the melanic allele is

dominant, the frequency of
the melanic forms is equal
to the sum of the
homozygotes BB and
heterozygote Bb
frequencies
Estimating heterozygote frequencies for dominant
alleles
• f(Black moths) = f(BB) + 1/2f(Bb) = =0.87
• Frequency of non-melanic forms is simply the frequency of
the recessive homozygote
• F(non-melanic moths) = f(bb) = q2 = 1.00 - 0.87 = 0.13
• Thus, q = f(b) = √(0.13) = 0.36
• p = f(B) = 1 – 0.36 = 0.64
• Now,
f(BB) = p2 = (0.64)2 = 0.41
f(Bb) = 2pq = 2(0.64)(0.36) = 0.46
Thus we know that 87% of the moths in Birmingham are black. If
we assume HWE, then this 87% can be partitioned into 41%
homozygotes and 46% heterozygotes
Frequency of heterozygotes
• In a population consisting of an infinite number of individuals
(i.e., a very large population), which is panmictic (marriages
occur randomly), and in the absence of mutation and
selection, the frequency of the genotypes will be the
development of (p+q)2, p and q being the allele frequencies.
• For loci with rare, recessive alleles, most recessive alleles will
be in heterozygotes
• F(Aa) > f(AA)
• Since most human disease alleles are recessive and rare, this
would mean that most disease genes can be found in carriers
who do not show the disease symptoms. Only the individuals
homozygous for the recessive allele would show the disease
• Example: the disease sickle cell anemia in humans has an
incidence of 1 in 1700 among Africans. The genotype
frequency is therefore q2. Frequency of the disease allele,
q = √(q2) = √(1/1700) = 0.024. The nondisease allele, p,
will be 1 - 0.024 = 0.976
Frequency of heterozygotes
• Assuming HWE, the frequency of heterozygotes is

2pq = 2(0.024)(1-0.024) = 0.047
• Or approximately 1 in 21. Thus carriers are common in the
population while affected individuals are rare

Hardy-Weinberg equation
• Hardy and Weinberg in 1908 formulated the
relationship that can be used to predict allele
frequencies when genotype frequencies are
given, or predict genotype frequencies when
allele frequencies are given. The relationship is
known as the Hardy-Weinberg equation
p  2 pq  q  1
2 2
where p and q are allele frequencies of a diploid

locus

Prediction of genotype frequencies
• The genotype frequencies of a population in

HWE can be predicted from two graphs:
• (1) Graphical representation of HWE
• (2) De Finetti diagram

• The figure shows the Graphical representation of
correspondence between Hardy-Weinberg equilibrium
the frequency q, of the a
allele and the genotype Frequency of A allele (p)
frequencies in the case of
two alleles in a panmictic
system. The heterozygotes
are most frequent when
allele frequencies are
equal and 0.5. When an
allele is rare, the
corresponding
homozygote frequency is
rarer because it is a
square of the allele
frequency. In this case all
the subjects who have this
allele are heterozygotes Frequency of a allele (q)

De Finetti diagram
The diagram is named after the
Italian statistician, Bruno (1905-
1988). It is based on an equilateral
triangular coordinate system in
which the frequencies of all
coordinates from any point within
the triangle and drawn to make a
perpendicular intersection to the
sides of the triangle must add up
to 1.0. The three perpendicular
intersections are the genotype
frequencies. The parabola is the
HW expected genotype
frequencies. The dashed lines
represent the frequency of each
genotype from 0 to 1. To use the
diagram, read the distance from
the point in the triangle chosen to
the side of the triangle at the
perpendicular intersection CS 563 Population and
Quantitative Genetics, KNUST 37
Multiple allelic systems
• The Hardy-Weinberg proportions for genes with three or more
alleles are extensions of the two-allele case
• Let’s consider a locus, A, with n alleles
• A1, A2, A3,…., An
having frequencies p1, p2, p3,….,pn
where p1 + p2 +p3 + ….+ pn = 1
We expect the genotype frequencies to be as follows:
pi2 = frequency of AiAi homozygotes

2pipj =frequency of AiAj heterozygotes and
pj2 = frequency of AjAj homozygotes
With more than two alleles at a locus, the genotype frequencies
are determined by the gene frequencies in the same way as
with two alleles

Deviations from Hardy-Weinberg equilibrium
• Deviations in parental Hardy-Weinberg genotype frequencies
from that in progeny in the next generation would not be
realized in the following instances:
1. If the two sexes have different allele frequencies even under
panmitic conditions
2. If a small number of parents mated just by chance, then not all
genotypes would be produced. This is analogous to flipping a
coin only a few times - this will not produce an equal number of
heads or tails
3. Natural selection- this is a biological process that causes
some genotypes in either parental or progeny generations to
be more frequent than others- HW expectations would not be
met
CS 563 Population and
39
• 4. Ploidy differences between males and females as seen in
chromosomal sex determination and haplo-diploid organisms
– In chromosomal sex determination, as in mammals, birds, and
Lepidoptera (butterflies and moths), one sex is determined by
possession of two identical chromosomes (homogametic sex),
while the other sex is determined by possession of two different
chromosomes (heterogametic sex).
– In mammals, females are homogametic (XX) while males are
heterogametic (XY).
– In birds and Lepidopterans, we have the opposite – males are
homogametic (ZZ) and females are heterogametic (ZW).
– In haplo-diploid organisms such as bees and wasps
(Hymenoptera) males are haploid (hemizygous) for all
chromosomes while females are diploid for all chromosomes

Sex-linked loci
• In sex-linked loci, genotype frequencies differ between the

sexes.
• The genotype frequencies for the homogametic sex (XX

females in mammals) are in Hardy-Weinberg proportions just
like autosomal loci and the relationship between gene
frequency and genotype frequency is the same as with
autosomal gene
• Genotype frequencies for the heterogametic sex (XY males in

mammals) are equal to allele frequencies since each
individual carries only one gene instead of two

• In such instances, prediction of genotype frequencies at

one locus under Hardy-Weinberg assumptions requires
keeping track of allele and genotype frequencies in both
sexes in loci and specific chromosomes.

Sex-linked loci
Male gamete
X-bearing sperm Y sperm
A1(p) or A2(q) Y
Female A1(p) A1A1(p2) A1A2(pq) A1Y(p)

gamete A2(q) A1A2(pq) A2A2(q2) A2Y(q)
Attainment of HWE in sex-linked loci requires that gene

frequencies are equal in the two sexes. Remember, for
autosomal loci, it takes one generation of random mating to
reach Hardy-Weinberg equilibrium

Sex-linked loci
• At generation t, let pm and pf denote the frequencies for
X-linked allele A1 in males and females, respectively. Let
pm* and pf* be the frequency in the previous generation,
t-1.
• Since the frequency of A1 in a male gamete in any
generation is equal to its frequency in females in the
previous generation (t-1)
pm = pf*
• Also, since the frequency of A1 alleles in females in
generation t is the average of the allele frequencies of
these alleles in males and females in the previous
generation t-1,
pf = (pm*+ pf*)/2
Sex-linked loci
• The difference in allele frequencies between

males and females in generation t is
pf – pm = (pm* + pf*)/2 – pf*

Sex-linked loci
p p
* *
p *
p f  pm  m f
 f
2 1
pm  p f  2 p f
* * *
p f  pm 
2
pm  p f
* *
p f  pm 
2
( p f  pm )
* *
p f  pm 
2
Sex-linked loci
• Thus, in every generation the difference between male

and female allele frequencies is halved and is in the
opposite direction of the previous generation
• The gene frequency in the population as a whole does
not change but its distribution between the two sexes
oscillates as the population approaches equilibrium

Other factors affecting departure from HWE
5. Mating system- effect of nonrandom mating on genotype frequencies
6. Gene flow
Mating systems
• Mating patterns of organisms do not actually exhibit the random-
mating assumptions, rather, organisms follow systems that create
predictable deviations from HW expected genotype frequencies
a) Assortative mating: this is a term used to describe nonrandom
mating. Two kinds occur:
i) Positive assortative mating
The case where individuals with like genotypes or phenotypes tend
to mate
ii) Negative assortative mating or disassortative mating
The case where individuals with unlike genotypes or phenotypes
tend to mate
•
Mating systems
• Non-random mating has effect on expected genotype
frequencies in a population
• Mating among relatives = consanguineous mating or
biparental mating. This increases the probability that a
progeny is homozygous compared to random mating
because relatives are more likely to share alleles –
identical by descent (idb). This has some similarity to
selfing
• Sexual autogamy or self fertilization = mating where
an individual can mate with itself because it possesses
both sexes. This occurs in plants and nematodes

Mating systems
• Disassortative mating- where individuals with

unlike genotypes have higher probability of
mating
• An example is in mice where the Major
Histocompatibility Loci (MHC) produces proteins
involved in self or nonself recognition response
for avoidance

Other factors creating departure from HWE
Nonrandom mating effects

• Effects of nonrandom mating on genotype
frequencies can be measured by comparing
Hardy-Weinberg expected frequency of
heterozygotes to that observed in a population
• The quantity, fixation index, F, is used to
compare how much heterozygosity is present in
an actual population relative to expected levels
of heterozygosity under random mating

Fixation index
• Fixation index is given by
• Where He is the HW expected frequency of

heterozygotes based on allele frequencies, Ho is
observed frequency of heterozygotes
• F ranges from -1 to +1

Other factors affecting departure from HWE
• 7. When multiple loci control a trait, and there is

linkage disequilibrium
• For a two-locus (A and B) situation, Linkage
Equilibrium (LE), also known as Gametic
Equilibrium is the random association of alleles
at locus A with alleles at locus B
• The two-locus genotypes and their frequencies
at equilibrium are obtained by random union of
gametes, as for the one locus case

Multiple loci and linkage disequilibrium
• When we have random association at a single locus we
can predict the genotype frequencies from a product of
the allele frequencies. This gives the expected genotype
frequencies as shown below:
Locus Allele Frequency

A A1 p1
A2 p2
B B1 q1
B2 q2
The genotype frequencies (HWE) are:

Locus A = p12 : 2p1p2: p22
Locus B = q12 : 2q1q2: q22

Multiple loci and linkage disequilibrium
• The two-locus genotypes and their frequencies at
equilibrium are obtained by random union of gametes, as
for the one locus case
• In this case, there are four possible kinds of gametes (i.e., the
haplotype) per locus. Expected frequencies at equilibrium =
product of the allele frequencies at each locus separately:
Gamete Type Exp. Obs

(Haplotype) Freq. Freq.
• A1B1 p1 q1 P11
• A1B2 p1 q2 P12
• A2B1 p2 q1 P21
• A2B2 p2 q2 P22
Where p1 +p2 = 1
q1 +q2 = 1 Quantitative Genetics, KNUST
Recap
• HW expectations provide a null model , a

prediction based on idealized or simplified
situations where no biological processes are
acting on the population and genotype
frequencies are the result of random mating.
• Actual populations can be compared to the null
model to test the hypothesis about the
evolutionary forces acting on allele and
genotype frequencies


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CS 563

CS 563 Population Genetics, 1

• Quantifying genetic variation

CS 563 Population and 2

• The genetic constitution of a population, refers to

CS 563 Population and Quantitative 3

Let’s take an example from the Australian Aborigine population

Total 730 1.000 P+H+Q=1

• Genotype frequencies give the description of a population at

• Alleles (also genes) are the entities transmitted across

CS 563 Population and 5

Total 730 260 1200

CS 563 Population and 6

• We can also compute the allelic frequencies straight from the

• Let p=f(M) and q=f(N). Thus, p=f(MM) + ½ f(MN) and

From the data

So p=P + H/2 and

CS 563 Population and 8

• The cutoff at 0.95 (sometimes 0.99) in the definition of

CS 563 Population and 9

f(F) 0.80 0.11 1.00* 0.40 0.01*

CS 563 Population and 10

Genotype No. of individuals F S

CS 563 Population and 11

H = [f(FS)Adh + f(FS)Mdh + f(FS)Pgi + f(FS)Pgm + f(FS)Ldh]/5

CS 563 Population and 12

CS 563 Population and 14

• HE, or Nei’s gene diversity, D. The

CS 563 Population and 15

• On the whole, there is a positive relationship between

• This relationship is expected because the greater the

• Vertebrates have the lowest average amount of variation

• Plants come next and invertebrates have the highest

CS 563 Population and 16

• In population genetics, the word population does not usually

• In the transmission the genotypes from parents, a new set of

CS 563 Population and 17

• Populations are dynamic groups, that change over space and

• As population geneticists, we are interested in modeling this

• If we have this ideal population, we expect to see the

• 1. The population is stable with respect to gene and

• 2. In this ideal population, the genotype frequencies in the

CS 563 Population and 20

• That the population is stable, that is, gene frequencies do

CS 563 Population and 21

1. Let’s see what happens in the next generation. The random

CS 563 Population and 22

CS 563 Population and 23

The following data is for genotypes at the MN blood group for

Calculate the observed gene frequencies from the data

CS 563 Population and

• Genotype frequencies have not changed over

CS 563 Population and 25

2. Given these gene frequencies, calculate the genotype

• 3. Now that we have the expected genotype frequencies after

• 4. Let’s recap at this point:

 ( observed  exp ected ) 2

(22  23.14)2 (216  213.60)2 (492  493.26)2

• In situations where one desires to know the numbers of

• For example, a dominant allele A1 at a locus A, produces

• The example below illustrates how one can calculate the

CS 563 Population and 29

Industrial melanism in the moth