20/3/2024

Genetics and
Inheritance
Ethan Leo

Patterns of inheritance
Campbell 11th ed. Ch 14, 15
Brooker Genetics 6th ed. Ch 4, 5, 6, 28

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Recap: molecular basis of genetics and

inheritance
• Central dogma
• Meiosis

Mendelian inheritance
• H2 syllabus
• Monohybrid cross, dihybrid cross, test cross
• Idea that there are heritable traits in the form of genes and one character is
controlled by one gene
• Revision reading on IVY

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Extensions of mendelian inheritance

• Multiple alleles
• Incomplete dominance
• Codominance
• Heterozygote advantage – key example: sickle cell disease (Campbell 11th ed
Pg 336)
• Autosomal linkage
• Sex linkage
• Lethal alleles
• Epistasis
• Pleiotropy/polygenic inheritance
All are in H2 syllabus revision reading notes on IVY/Brooker 6e Ch 4, 6, 28
on IVY/Campbell 11th ed. Ch 14-15

Incomplete dominance vs codominance

• Incomplete dominance: phenotype of heterozygotes is the intermediate
between phenotypes of either homozygote
• E.g. incompletely dominant red/white flower alleles: red homozygous flower +
white homozygous flower = pink heterozygous flower
• Codominance: phenotype of heterozygotes is different from phenotypes
of either homozygote, usually the sum of phenotypes of both
homozygotes
• E.g. human ABO blood types – IAIA gives A blood type (A antigens expressed),
IBIB gives B blood type (B antigens expressed), IAIB gives AB blood type (both
A and B antigens expressed)

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Lethal alleles
• Alleles that cause death of the organism at
birth
• Organisms with these genotypes do not
contribute to the phenotypic ratio
• Phenotypic ratio does not add up to 4
• Lethal alleles are most commonly recessive
(hence can be maintained in populations) but
there are some dominant ones
• E.g. Huntington’s disease in humans – able to be
maintained in population due to late onset of
lethal symptoms

epistasis
• Effect of 2 genes interacting with each other
• The alleles of one gene can mask the phenotypic effects of
the alleles of a different gene, hence the epistatic gene
modifies how another gene (hypostatic gene) is expressed
• Best illustrated with an example: Coat colour in
Labrador retrievers
• Gene at B/b locus determines colour (B – black; b – brown)
• Gene at E/e locus determines if colour is deposited (E –
deposited; e – not deposited hence yellow) EE/Ee Black
• Alleles at E/e locus can mask the effect of alleles at B/b BB/Bb
locus hence we say that the ee genotype is epistatic over ee
Yellow
B/b locus ee
• We also say this is recessive epistasis since a recessive bb
Brown
genotype is epistatic over the B/b locus EE/Ee

• Results in 9:3:4 phenotypic ratio (modified 9:3:3:1 B/b locus E/e locus
ratio) Coat color

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epistasis
• Please read the section on epistasis in
the H2 notes on IVY thoroughly for
more examples! (under Genetic Basis
for Variation (II))
• Alternatively read Brooker Genetics
chapter 4 (also on IVY)
• Many different types of epistatic ratios
and biochemical pathways
• Take some time to understand them
properly – notice the similarities and
differences with the standard 9:3:3:1
dihybrid cross table
• Very useful summary from the notes (try
to identify types of epistasis in each case
e.g. recessive/dominant) 

Non-mendelian inheritance
These lie entirely outside the Mendelian framework because these forms
of inheritance do not involve nuclear genes.
• Maternal effect
• Extranuclear inheritance
• Epigenetic inheritance
Brooker 6th ed. Ch 5 on IVY (very detailed)
Campbell 11th ed. Ch 15.5

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Maternal effect
• Genotype of mother directly determines phenotype of
offspring – genotype of father or even offspring itself does not
affect phenotype
• This effect was discovered by performing a reciprocal cross
between two true-breeding snails
• Reciprocal cross: Mate a male homozygous for one allele and female
that is homozygous for another allele, then repeat but swap the sex of
the genotypes
• Result:
• F1 generation of first reciprocal cross had different phenotypes from
second reciprocal cross even though F1 generations of both crosses
had same genotypes
• F2 offspring all have the same phenotype even though they have
different genotypes because they all follow the phenotype of the F1 Dd
mothers.
• The mechanism behind the maternal effect is due to how eggs
are developed during oogenesis.
• Nurse cells that surround the eggs actually transfer gene
products during development. These gene products are coded
for by the mother’s genes, and so the offspring follows the
mother’s genotype.
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Extranuclear inheritance
• Extranuclear/maternal inheritance: inheritance of mitochondrial and
chloroplast DNA which are inherited from the mother
• Because the egg cell provides most of the cytoplasm and organelles to the
zygote following fusion
• Paternal leakage: On occasion the sperm also provides mitochondria
• NOT the same as maternal effect!!
• Maternal effect is caused by nuclear gene products from mother being
transferred to egg, extranuclear inheritance as the name suggests does not
involve nuclear genes at all
• Interesting case study: Chloroplast inheritance in Mirabilis jalapa
• Read more at: https://www.khanacademy.org/science/ap-biology/heredity/non-
mendelian-genetics/a/mitochondrial-and-chloroplast-dna-inheritance

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Epigenetic inheritance
• Epigenetics: the study of heritable phenotype
changes/changes in gene expression that do not
involve alterations in the DNA sequence
• 2 main phenomena:
• Genomic imprinting
• Dosage compensation
• Mechanisms: described in H3 epigenetics notes
on IVY
• Builds on H2 knowledge of organisation and control
of pro & eu genome

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genomic imprinting
• A segment of DNA in each parent is marked (e.g. by DNA methylation) and
this mark is passed on to the offspring
• Offspring only express either the allele from the father or the allele from the
mother (not both)
• Whether it is the father or the mother depends on how the genes are marked (parental
origin-dependent gene expression) and varies from species to species and from
imprinted gene to imprinted gene
• Essentially, for a certain species and certain imprinted gene, the allele from
the mother will always be expressed and the allele from the father will always
be silenced, or vice versa
• Hence knowing
1. whether the organism always expresses the allele from the mother or the
father, and
2. which allele was inherited from mother and which allele from father,
• you can predict the phenotype of the organism.

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genomic imprinting
• Example: In mice, Igf2 (insulin-like growth factor) gene allele from
father is always expressed and allele from mother is always silenced.
• Given that allele Igf2 codes for normal growth factor and allele Igf2-
codes for non-functional growth factor, can you predict the phenotypes
(normal or dwarf) of these heterozygous offspring mice?

Answer in 3…2…1…

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genomic imprinting
• Example: In mice, Igf2 (insulin-like growth factor) gene allele from
father is always expressed and allele from mother is always silenced.
• Given that allele Igf2 codes for normal growth factor and allele Igf2-
codes for non-functional growth factor, can you predict the phenotypes
(normal or dwarf) of these heterozygous offspring mice?

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Dosage compensation
• Ensures that the level of expression in genes on sex chromosomes is similar in both
sexes, even though males and females have different sets of sex chromosomes.
• Mechanisms (different in different species since they have different types of sex
chromosomes)
• X-inactivation: expression of one X chromosome is entirely inactivated (e.g. humans) → mosaic!
formation of Barr body
• Drosophila: Double expression of X-linked genes in males
• C. elegans: Decreased expression of both X chromosomes by half hermaphrodite

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problem solving in genetics

Brooker Genetics 6th ed. Ch 2.5
Genetics Problem Solving Guide 2e

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Genetic crosses
• Usually in genetic cross questions, they are asking for the
probability of an offspring having a certain genotype
• H2 method: draw punnett square and count manually
• But for bio olympiad you need a faster method and that is probability
(Math essentially)
• Math you need:
• Probability
• Combinations (whole topic is permutations and combinations but only
combinations is relevant here)/binomial theorem formula
• (Both are in the H2 math syllabus (first two topics of statistics))
• If you have time, try doing the exercises in the Genetics Problem
Solving Guide (available in RI library)

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Probability same time ! mutually exclusive events that fulfil the question

Multiplication Rule (AND) Addition Rule (OR)

• For independent events (probability of • For mutually exclusive events (events
one happening does not affect that cannot happen at the same time)
probability of another happening) Example: Cross Aa x Aa, what is the
Example: Cross Aa x Aa, what is the probability of offspring being
probability of offspring AA genotype? homozygous?
• Offspring must have A from mother • Offspring can be AA OR aa
AND A from father • First step: probability of AA = ¼, and
Required probability probability of aa = ¼
= probability of inheriting A from mother Required probability
TIMES probability of inheriting A from = probability of offspring being AA
father PLUS probability of offspring being aa
=½x½=¼ =¼+¼=½
Essentially, if you ever forget whether to multiply or add, just think about whether it is an “and” scenario or an “or” scenario

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at least 1 girl no girl

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Example 1
A plant with genotype AaBbCc is crossed with a plant with genotype aaBbCc.
What is the probability of an offspring plant having genotype AaBbcc? Assume
that all 3 gene loci are on different chromosomes.
• Approach: Calculate the probability of getting Aa, then probability of getting
Bb, then probability of getting cc. Combining, since we need Aa AND Bb
AND cc to make a plant with genotype AaBbcc, we multiply the three
probabilities found earlier.
Probability of Aa = ½ 2 ways for offspring to be Bb: B from mother, b
from father OR b from mother, B from father –
Probability of Bb = ½ x ½ x 2 = ½ (why x2?) probability of former PLUS probability of latter.
Probability of cc = ½ x ½ = ¼ Hence (½ x ½) + (½ x ½) or ½ x ½ x 2

Probability of AaBbcc = ½ x ½ x ¼ = 1/16

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Example 1
• The punnett square we avoided drawing:
ABC ABc AbC Abc aBC aBc abC abc

aBC AaBBCC AaBBCc AaBbCC AaBbCc aaBBCC aaBBCc aabBCC aabBCc

aBc AaBBCc AaBBcc AaBbCc AaBbcc aaBBCc aaBBcc aabBCc aabBcc

abC AaBbCC AaBbCc AabbCC AabbCc aaBbCC aaBbCc aabbCC aabbCc

abc AaBbCc AaBbcc AabbCc Aabbcc aaBbCc aaBbcc aabbCc aabbcc

2/32 = 1/16

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combinations
• nCr (n choose r) is the number of ways
to choose n unique items out of total r MATH
unique items
• Useful knowledge if the question
requires an extra layer of analysis, e.g.
instead of asking for the probability of
just one offspring having a certain
genotype, they ask for the probability of
multiple offspring out of a total having
that genotype.
• You can cheese the first part with a
punnett square (the probability of getting
the required genotypes) but for the
second part, you have no choice but to
use math

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Example 2
Two individuals, both heterozygous for eye colour allele (Bb), have 5 children.
What is the probability that 2 of the 5 children have blue eyes (bb)?
• Approach: Find probability that 2 children are bb AND 3 children are not
bb, then multiply this by the number of different ways to choose 2 of the 5
children
• Why multiply by number of ways to choose? Because the required
probability must include the first way to choose 2 out of 5 OR the second way
to choose 2 out of 5 OR the third way… and so on. Hence (¼ x ¼ x ¾ x ¾ x
¾) + (¼ x ¼ x ¾ x ¾ x ¾) +…+ (¼ x ¼ x ¾ x ¾ x ¾), total 10 times.
Probability of bb = ½ x ½ = ¼
Probability of not bb = 1 – ¼ = ¾
Probability of 2 children bb and 3 children not bb = ¼ x ¼ x ¾ x ¾ x ¾ =
27/1024
Number of ways to choose 2 children out of 5 to be bb = 5C2 = 10
Required probability = 10 x 27/1024 = 135/512 = 0.264 (3 s.f.)

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Example 2
Two individuals, both heterozygous for eye colour allele (Bb), have 5
children. What is the probability that 2 of the 5 children have blue eyes
(bb)?
• In case you read Brooker: Binomial theorem
Total number Probability of outcome 2
of events

Number of
outcome 1
Number of Probability of outcome 1
outcome 2

• Notice that the first part is the nCr formula for the total number of
combinations, and the second part is the formula to find the probability
of one of these combinations
• Essentially the same as the stuff on the previous slide so just remember
whichever one you prefer

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Parents (not in diagram): homozygotes for each red/blue chromosome

Gene mapping
• Mendelian genetics assumes the Law of
Independent Assortment: alleles of different genes
are assorted into different gametes independently
of each other
• This does not apply to linked genes i.e. genes on
the same chromosome
• Linked genes tend to be inherited together (hence
inheritance of linked genes are NOT
independent)
• Offspring tend to inherit the parental genotype
unless crossing over occurs between the two genes
to form a recombinant chromosome.
• The closer the 2 genes are on a chromosome, the
less likely crossing over will occur → less likely
offspring will inherit a recombinant genotype
• We can use the frequency of recombinant
genotypes being inherited to measure the distance
between linked genes ⇒ genes for eye colour and wing type are 10.7 map
units/centimorgans away from each other
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Gene mapping (3 genes)

• Same process as for 2 genes
• Cross two strains of flies (parents) that are homozygous at all three genes.
One is homozygous dominant at all three genes, and represents the wild-type,
while the other is homozygous recessive at all 3 genes.
• Perform a test cross between the F1 heterozygote and another homozygous
recessive fly to produce F2 flies, and collect data on the F2 generation.
2. Perform test cross between F1
heterozygous and homozygous
1. Cross two true-breeding strains recessive

3. Collect data for F2

generation

Black body, Grey body, red

purple eye, eyes, long wings
vestigial wing (wild-type)

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Gene mapping (3 genes)

Without calculation, what information can we infer from
this set of data?
Parental genotype (has the most individuals)
Recombinants (single and double crossovers)
Double crossover (has the least individuals)
Double crossovers are the ones with the least individuals
because the probability of two crossing over events occurring
together is lower than just 1 occurring. Identifying the double
crossover classes allow us to identify the order of the three
genes. Look for the allele that is the “odd one out” i.e. not from
the same parent as the other two alleles, by comparing the
double crossover classes with the parental genotypes. In the
parental genotype, we have a grey body, red eye, and long wing.
For the double crossover, we have grey body, purple eyes, and
long wing. Clearly, the only one that’s different is the gene for
eye colour. This means eye colour gene is in the middle.
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Gene mapping (3 genes) – calculating exact

distance between each gene
6.07

Body colour Eye colour Wing type

Step 1: Calculate distance between gene in the middle

(eye colour gene) and gene at the side (in this case body
colour gene was chosen first, so ignore the other one
which is wing type)
• Look for data on recombinants between these two
genes, i.e. grey body, purple eye or black body, red eye
Recombination frequency
= (2+30+28+1) / 1005 = 0.0607 (to 3sf)
Distance between eye colour and body colour genes
= 6.07 map units

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Gene mapping (3 genes) – calculating exact

distance between each gene
6.07 12.3

Body colour Eye colour Wing type

Step 2: Calculate distance between gene in the middle

(eye colour gene) and gene on other side (wing type gene)
• Look for data on recombinants between these two
genes, i.e. red eye, vestigial wings or purple eye, long
wings
Recombination frequency
= (61+2+60+1) / 1005 = 0.123 (to 3sf)
Distance between eye colour and wing type genes
= 12.3 map units

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Gene mapping (3 genes) – calculating exact

distance between each gene
6.07 12.3

Body colour Eye colour Wing type

Step 3: Check by calculating distance between genes on the

sides (wing type and body colour genes)
• Look for data on recombinants between these two genes, i.e.
grey body, vestigial wings or black body, long wings
Recombination frequency
= 61 + 30 + 28 + 60 / 1005 = 0.178 (to 3sf)
Distance between body colour and wing type genes
= 17.8 map units
Distance calculated previously = 6.07 + 12.3 = 18.4 (close to
17.8)

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Gene mapping (3 genes) – calculating exact

distance between each gene
6.07 12.3

Body colour Eye colour Wing type

Q: Why is there a discrepancy between calculations 1+2 and 3?

A: We did not consider double crossovers.
• At a phenotypic level, they don't appear to be recombinant for body
colour and wing type
• But at the gene level, they are recombinant for body colour and
wing type due to crossing over
Calc 3 should be:
Actual recombination frequency
= 61 + 30 + 28 + 60 + 2(2+1) / 1005 = 0.184 (to 3sf)
Note that we have to x2 to the number of double crossover
individuals because 2 crossover events occurred for each individual

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Interference 6.07 12.3

Body colour Eye colour Wing type

• Positive interference: occurrence of a crossover in one region of a

chromosome decreases the probability that a second crossover will occur
nearby.

From our example:

Expected likelihood of a double crossover = 0.061 × 0.123 = 0.0075, or 0.75%
Expected number of offspring due to a double crossover, based on a total of
1005 offspring produced = 1005 × 0.0075 = 7.5
Observed number of double crossovers = 3
Coefficient of coincidence (C) = 3/7.5 = 0.40
Interference = 1 − 0.4 = 0.60

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Pedigree analysis
• A pedigree shows the relationship between family members and
phenotypes of each member
• Used to determine patterns of inheritance

means the individual is dead

means monozygotic twins

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Pedigree analysis

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Str analysis
• STR = short tandem repeat
• At particular loci on chromosomes,
humans have a number of short
repeated sequences (each repeat can be
2-5 nucleotides long)
• The number of repeats at each locus
varies between people but is inherited
• The combination of repeats one has is
unique due to the very low probability
of two people sharing the exact same
numbers at each locus

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Str analysis
• Multiple STR loci shown
• At each locus two numbers shown e.g. Huong
has 3/5 at Locus 1  one chromosome has
allele 3; other chromosome allele 5
• Each person can be uniquely identified by the
combination of alleles they have between all
loci
• We can also compare genotypes between
individuals to determine who is related e.g.
Huong may be related to 735 as they share
one allele at each locus: allele 5 at locus 1,
allele 2 at locus 2, allele 5 at locus 3, allele 3
at locus 4, allele 2 at locus 5, and allele 2 at
locus 6.

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Population genetics
Campbell 11th ed. Ch 23, 52
Brooker Genetics 6th ed. Ch 27

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Population genetics
• Population genetics studies allele frequencies in populations and how
they change
• These changes may arise through natural selection, mutations,
migration, or genetic drift

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Hardy-Weinberg equilibrium
• In the absence of evolutionary influences (i.e. selection, mutation,
genetic drift, migration), allele frequencies remain constant from
generation to generation.
• Such a population is in Hardy-Weinberg equilibrium.

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Hardy-Weinberg equilibrium
• If there are 2 alleles for a trait in a • The Hardy-Weinberg equation allows us to
population, we can represent the calculate the frequency of each genotype. p2 is
frequency of the dominant allele with the frequency of the homozygous dominant
p, and the frequency of the recessive genotype; 2pq is the frequency of the
allele with q. The sum of p and q heterozygous genotype; and q2 the frequency
the homozygous recessive genotype. Adding all
should then be 1. these frequencies should also result in 1.

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example (p+q)2 = p2 + 2pq + q2 = 1

If the proportion of individuals with the recessive phenotype is 16%,
determine the frequency of each allele.
Proportion with recessive phenotype = 0.16
Using Hardy-Weinberg equation,
Proportion with recessive phenotype = q2
q = sqrt(0.16) = 0.4
Since p + q = 1, p = 1 - 0.4 = 0.6
We can also find the proportion of the other genotypes:
Dominant = p2 = 0.6 x 0.6 = 0.36
Heterozygous = 2pq = 2(0.6)(0.4) = 0.48

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Hardy-Weinberg equation for more than 2

alleles
3 alleles: (p+q+r)2 = 1
4 alleles: (p+q+r+s)2 = 1

And so on...

Expand the expressions to get the different combinations of alleles

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Effects of natural selection

What happens if natural selection acts on a population in HW equilibrium?
• Recap: Natural selection occurs when individuals with a particular genotype
have higher fitness than others → more likely to survive → produce more
offspring → pass down alleles to next population → proportion of individuals
with allele increases
• In other words, natural selection is a result of differential reproductive
success (fitness) between genotypes
• The symbol W is used to denote fitness; fitness of each genotype is always
quantified relative to the genotype with the highest fitness, which is assigned a
fitness of 1.0
• e.g. WAA = 1.0; WAa = 0.8; Waa = 0.2

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Effects of natural selection

• Effect 1: Proportion of AA increases since it is selected for
Given
WAA = 1.0 p2 = 0.36 (p+q)2 = p2 + 2pq + q2 = 1
WAa = 0.8 2pq = 0.48
Waa = 0.2 q2 = 0.16
In next generation:
Mean fitness of population = WAA(p2) + WAa(2pq) + Waa(q2)
= (1.0)(0.36) + (0.8)(0.48) + 0.2(0.16) = 0.776
Frequency of AA = WAA(p2) / mean fitness = (1.0)(0.36) / 0.776 = 0.464
Frequency of Aa = WAa(2pq) / mean fitness = (0.8)(0.48) / 0.776 = 0.495
Frequency of aa = Waa(q2) / mean fitness = 0.2(0.16) / 0.776 = 0.0412

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Effects of natural selection

• Effect 2: mean fitness of the population increases.
Given
WAA = 1.0 p2 = 0.36
WAa = 0.8 2pq = 0.48
Waa = 0.2 q2 = 0.16
Gen 1: Mean fitness = 0.776
Gen 2: Mean fitness
= WAA(p2) + WAa(2pq) + Waa(q2)
= 1.0(0.464) + 0.8(0.495) + 0.2(0.0412) why does mean fitness increase?
individuals with AA more likely to pass
= 0.868 > 0.776 on their genes, next gen theres is more
AA in the population -> AA has higher
fitness so mean fitness is higher

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Genetic drift
• Recap: Genetic drift refers to a change in allele frequency
due to chance events or random fluctuations, and not due
to a specific evolutionary pressure.
• In smaller populations, there is a chance that an allele “goes
to fixation” (homozygous in 100% of individuals) or is
eliminated completely, due to random fluctuations.
Formula for probability that a new allele goes to fixation
= 1/(2N), where N = population number
Formula for probability of elimination = 1 - 1/(2N)
• It is easier for an allele to go to fixation in small populations
by chance since the smaller N is, the closer 1/2N is to 1.
This is one of the reasons small populations tend to have
poor genetic diversity.
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Kin selection
• Kin selection is an evolutionary strategy employed by some species like
bees, where an individual favours the reproductive success of their
relatives even at a cost to their own fitness.
• Kin selection occurs between related individuals; if individuals are
unrelated the behaviour is known as just altruism
• Kin selection emphasizes the genetic success of an individual, and not
just reproductive success.

usually employed by the hymenoptera class

(e.g bees -> all worker bees assist their queen
in reproducting

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Kin selection and Hamilton’s rule

• According to Hamilton’s rule, an individual will engage

in kin selection if the genetic contribution its relative
makes to the next generation is greater than the cost to
the individual’s own fitness.
• British biologist JBS Haldane explained this by saying “I
would lay down my life for 2 brothers or 8 cousins.”
• Each sibling shares half my alleles on average.
• If I was supposed to have one child but instead died to protect
my two siblings, it would be worth it for me because my two
siblings would together contribute to the next generation the
same alleles that I would have contributed to my own child.
• Same logic for the cousins
• My fitness takes into account not just my own offspring, but also
the offspring of my relatives.
• This called inclusive fitness.

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1) assign the person of interest a value of 1
** the coefficient of relatedness of a and b is diff frm b and a
2) move up down or side is always 1/2 -> then just multiple HAHA

Kin selection and Hamilton’s rule

A mother antelope and its child are galloping along the plains, when they encounter a group of hungry lions. If the two
antelopes try to escape the lions together, there is a 75% chance that both will be consumed and eaten, and a 25% chance
that both will escape alive. If, however, the mother sacrifices herself to the lions, she may be able to buy her baby
additional time to escape.
What is the minimum chance that the baby can have to escape from the lions following such a sacrifice such that the
mother’s action will be evolutionarily favored?
You may assume that the baby antelope, once escaped, will be guaranteed to survive into a reproducing adult, and that the
mother antelope is at the beginning of reproductive age.

If the action is evolutionary favoured, it means rB>C.

r=0.5 because mother-daughter r/s. Meaning if the daughter lives, the mother will have pass on 0.5 of its alleles.
Let p be the required probability
B refers to the benefit to the relative thus INCREASE in chance of survival of the daughter; B= p – 0.25
C refers to the cost to the individual i.e. how much chance of survival she gives up; C = 0.25
For rB>C, 0.5 (p-0.25) > 0.25
p-0.25>0.5
p>0.75
Min chance is 0.75

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