GENETIC DRIFT

I. GENETIC DRIFT

 random changes (chance) in the frequencies of 2 or more alleles or genotypes from one
generation to the next
 caused by chance events of survival, reproduction, and meiosis
 based on luck, not fitness
o other factors that affect genetic drift: age of populations; no. of breeding males and
females

II. FUNDAMENTAL FEATURES OF GENETIC DRIFT

1. Genetic drift is unbiased.

 allele frequency is as likely to go up as to
go down
⤷ natural selection: favor one allele
over another, genetic drift: does not

2. Random fluctuations in allele frequency are

larger in smaller population.
 outcomes = more predictable (averaging
over a  number of random events)
 reason why genetic drift is stronger in
small populations

3. Genetic drift cause genetic variation to be

lost.
 an allele frequency that fluctuates
randomly up and down will
eventually reach either p/q = 0 or 1
4. Genetic drift causes populations that are
initially identical to become different.
 different populations, different
genotypic variance

5. An allele can become fixed without the benefit of

natural selection.
 allele 1: fixed, allele 2: lost
 if natural selection is not acting
 GENETIC DRIFT

III. HOW STRONG IS GENETIC DRIFT?

• Genetic drift is stronger in smaller populations than in larger populations

• Pelagibacter ubique
− most numerous organisms on earth
− population size of 1028
• How do we measure the strength of genetic drift?
 Through this formula: 1/(Ne)
• Effective Population Size (Ne) – the number of individuals in a population who
contribute offspring to the next generation (introduced by Wright)
 in evolution, population size means effective population size
 effective population size is usually smaller than the real census population
size (not everyone breeds and leaves offspring)

EXAMPLE: 100 adult zebras:

Nm = # males 60 females

Nf = # females 12 males with harems

28 male bachelors
4𝑁𝑁𝑚𝑚 𝑁𝑁𝑓𝑓
𝑁𝑁𝑒𝑒 =
𝑁𝑁𝑚𝑚 + 𝑁𝑁𝑓𝑓
REMEMBER:
4 × 12 × 60
𝑁𝑁𝑒𝑒 = = 40  value = strong drift
12 + 60
∴ 20 Nm and 20 Nf in an ideal population  value = weak drift
mating randomly

III. WHY DO WE CARE ABOUT EFFECTIVE POPULATION SIZE?

• Ne: actual unit of evolution (rather than the census size N)

 only the alleles that actually get passed onto the next generation count
 evolutionary dead ends - individuals that do not mate or have offspring
• Ne = 1 or extremely low: population is completely inbred (even if the census size is large)

IV. CONSEQUENCES OF GENETIC DRIFT

• If population size is reduced:

 at the allelic level
- random fixation of alleles (loss of alleles)
 at the genotypic level
- loss of heterozygosity (due to fewer alleles)
• Probability of loss of alleles is greater in smaller populations
 example:
o there are 50 different alleles in population  a new population is
founded by only 10 individuals  new population will be unable to
capture all 50 alleles, and many of the alleles will be lost
 GENETIC DRIFT

V. 2 MAIN TYPES OF GENETIC DRIFT

1. Bottleneck Effect
 occurs when a natural disaster, or any
similar event, randomly kills a large
portion of the population
 leaving survivors whose allele
frequencies are different from the
previous population

2. Founder Effect
 involves small groups of
individuals separating from a
larger population
 forming a small and new
colonies
 invade new isolated
habitat

VI. A HUMAN FOUNDER EFFECT EXAMPLE

• The Amish Community in northern USA,

 was founded by a small number of colonists which possessed the gene for
polydactyly (extra toes or fingers)
− population  in size but remained genetically isolated
− polydactyly more frequent in the Amish community than in other
communities

VII. RANDOM FIXATION OF ALLELES

• Fixation: When an allele

frequency becomes 100%
− other alleles are lost by
chance
• Fluctuations are much larger
in smaller populations
 GENETIC DRIFT

• Some effects of genetic drift (fixation of alleles) are best understood by looking backward
in time
− done through Genetic Genealogy, the use of DNA and historical records to infer
the relationship between individuals
o tracing family history using gene tree
− all copies of a gene in any population, species, or group of species trace back
to a single ancestral gene copy at some point in the past

VIII. GENETIC DRIFT VS NATURAL SELECTION

• population size: relative importance of selection and drift

o  Ne: caused by natural selection
o  Ne: caused by genetic drift
 many deleterious mutations (fixed in humans and other species)

IX. GENETIC DRIFT OFTEN LEADS TO INBREEDING

• mating among genetic relatives due to small

population size
− results to loss of some alleles
− alleles at a locus - most likely
homozygous
• Wright developed a method
− calculate to measure the chance of
being a homozygote upon inbreeding
− Fx: coefficient of inbreeding
− FA: coefficient for the common ancestor
𝒏𝒏+𝟏𝟏
• 𝑭𝑭𝒙𝒙 = 𝜮𝜮 ��𝟏𝟏�𝟐𝟐� (𝟏𝟏 + 𝑭𝑭𝑨𝑨 )�
 GENETIC DRIFT

X. CONSEQUENCES OF INBREEDING

 small population size  genetic drift = loss of alleles,  in heterozygosity,  exposure

of deleterious alleles
o deleterious recessive alleles: exposed as homozygous
 less masked in the heterozygous state

XI. DRIFT AND GENETIC VARIATION WITHIN SPECIES

• Polymorphism - typically high in the regions between genes and within introns
• Exons (coding regions) - less variable
 synonymous mutation - change in 3rd base of the codon
 nonsynonymous mutation - change in 1st or 2nd base of the codon
 neutral mutations - common in non-coding regions
 deleterious mutations - common in coding regions
 removed through purifying selection or negative selection
 some retain due to drift

XII. EXAMPLES OF INBREEDING

• Agriculture
− Holstein cows
 neurological disorders
 autoimmune diseases
 fertility problems
• Royal Families
− America
 Porphyria - accumulation of porphyrin precursors, causes insanity
 dominant, more intense in homozygous form
 George III, loss of American Colony
− Spain
 Hapsburg jaw - facial deformity (elongated jaw)
 kings and queens of Spanish Habsburg dynasty
 cognitively disabled + impotent
− Egypt
 Tutankhamun (King Tut) - best known pharaoh
 clubfoot, a cleft palate, and a severe bone disease
 parents: brother + sister
• Genetic Immigration (Migration)
− Diana + Charles (7th cousin)
− Kate & Megan (neither aristocrats nor royals)