Lecture 16

Population Genetics 2

About the same percentage (2.3%) of

Neanderthal Genome as Europeans but less
fragmented!

Application of our new

knowledge about the
HW-Equilibrium

Important assumptions of Hardy

and Weinberg

infinite population size (insuring randomness)

No mutation
slow
rare
fair meiosis
No selection
between rather then within population
No migration
random mating (inbreeding)

Evolution as change in allele

Frequency over time!
Hardy-Weinberg Null model (no change!)
Change:
Mutation (generating novel alleles but also
changes allele frequencies)
Genetic Drift (change is due to chance)
Natural Selection (change due to heritable
differences in survival and reproduction)
Non-random mating and inbreeding
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Mutations Allele frequency

Changes
u = Mutation rate
Probability of fixation under neutral
evolution is 1/u
Proability a new mutation will arise in the
population is given by population size N
and mutation rate u (in diploid organism u
times 2N)
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Mutation-Drift Equilibrium

*Genetic drift is the change in the frequency of an allele in a

population due to random sampling.
9

Under Mutation-
Drift Equilibrium

Pfixed = u = rate of evolution

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We can use this fixed differences

to estimate mutation rates!

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"Non-Darwinian Evolution" (1969)

Jack L. King and Thomas H. Jukes

the vast majority of molecular differences are

selectively "neutral." That is, the molecular
changes represented by these differences do
not influence the fitness of the individual
organism. As a result, the theory regards these
genomic features as neither subject to, nor
explicable by, natural selection.
Kimura 1986 Neutral theory of molecular
evolution
Is this really Non-Darwinian Evolution?
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The effect of inbreeding on

allele and genotype
frequencies!

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Increase in Homozygotes by
non-random mating (e.g.
inbreeding)
Gametes
From:

males
p

females
p

p*p = p2

p*q

q*p

q*q = q2

Homozygous genotypes dont change but heterozygous genotypes

decline every generation by 50%!
Inbreeding Coefficient = F = 1 (Heterozygosityobserved/Heterozygosityexpected)
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Why is inbreeding bad (negative fitness

effects) or not seen as often?
What are the results of inbreeding in
humans!

15

Hemophilia of Queen Victoria

Only males are affected by this disease. Why?

Recessive Sex (X) linked disease! Hemophilia B (factor IX deficiency) occurs
in around 1 in about 20,00034,000 male births.
The non-sex-linked hemophilia C due to coagulant factor XI deficiency,
which can affect either sex, is more common in Jews of Ashkenazi
(east European) descent.

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17

Definitions/Notes
Fitness of a biological entity = average per capita rate of increase in
numbers
Natural selection can act on any difference in fitness among
phenotypically different classes of biological identities (entities maybe
alleles, genotypes, subgroups of genotypes, populations, species).
Note: If genetic drift lead to the replacement of a neutral allele we have
witnessed a change of allele frequency but natural selection has not
occurred. There is no average difference between the alleles, no bias
towards the increase of one relative to the other.

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Selection of and selection for

selection
- of objects (=of size)
- for properties
(= for red balls)
Woodpecker brains (= of) were selected to
withstand traumas (= for) to be able to
gain access to wood boring insects (fitness
advantage)!

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Adaptations...
are traits that evolved through natural/sexual
selection!

Object = of?

property = for?

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First Question Adaptation or Drift ?

Color pattern of Cepea nemoralis

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Drosselschmiede

22

change in
Allele/Phenotype
frequency

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change in
Allele/Phenoptype
frequency

Selection coefficient (s): Difference between the mean relative fitness of individuals
of a given genotype/phenotype and that of a reference genotype (in our case s = 0.1).
We can say, that genotype A has a selective advantage over genotype B or s = +0.1
for A but -0.1 for B.
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How can we measure the fitness effect of a character?

Manipulating the trait and measure fitness (tail length in
widow birds!

Adaptation not adaptive storytelling! Just because something looks complex

does not mean it evolved by natural selection.

Populus -

http://www.cbs.umn.edu/research/resources/populus

Open Populus Open Model Open Mendelian Genetics Genetic Drift

Populus
You should obtain this
window

Make sure that the following

settings are included:

- Set frequencies collectively

should be CHECKED.
- Permit Selfing should be
UNCHECKED.
- Number of loci = 6.
- Runtime = Other put in 100

Populus activity
This is what you should obtain (or at least a very similar
graph):

You can open multiple graphs by doing this process a second

time. That is:
Open Model Open Mendelian Genetics Genetic Drift