Lecture 25: Evolutionary processes

Some Mechanisms of Evolution

1.Mutation
2.Genetic drift
3.Natural selection
4.Sexual selection
5.Gene flow

1. MUTATION
 Ultimate source of all genetic variation
 Change the genetic code  New ‘alleles’ May have phenotypic effects (some won’t
have any effect “silent mutation”)
o With different amino acid sequences of alleles
o Alter gene expression
E.g: population of organism (gene pool) with 3 different allele (3 different version of a gene).
Mutation in this population, will change the genetic code which will generate new alleles with
different amino acid sequences which will alter gene expression which can have a phenotypic
effect (large, beneficial, deleterious …) or no effect on the phenotype.

Example of mutation in Drosophila: eyes change, wild type, wings, color… (important in the
field of scientific researches)

 Most mutations decrease fitness (ability to survive and reproduce) and are considered
deleterious. (since the organism is already adapted to their local environment).
 To study it, they create lines of organisms that are protected from natural selection so
that mutation can accumulate, because natural selection would normally eliminate.
 The control group is maintained in the normal environment where mutations that are
arising would be eliminated by natural selection.
 Then after a certain number of generations, the mutation accumulation line  when
they are tested by in the normal environment to calculate their fitness
As a result The average mutation reduces fitness by about 2%
 The few beneficial mutations that arise are likely to be of small effect (i.e., “hopeful
monsters” are rare)

2. GENETIC DRIFT causes random changes in allele frequency: a demonstration

o In order to measure evolutionary change, calculate allele frequency (proportion of the
Nb of each type of allele
allele present in the population) (x 100 to have them in
total nb of allele∈the population
%) and the total sum of all the allele frequencies is in a population is 1 (or 100 is in %).
o Genetic drift occurs when there is mortality or differential reproduction that is
independent of those alleles. The specific alleles are neutral to each other (no benefit or
detriment).
 o There is random mortality that takes place, so we calculate the new allele frequency, if
they are maintained in the next generation, an evolution has occurred So the genetic
drift has caused the evolution, but it was completely random. (don’t depend on the type
of alleles, it is a random mortality)
o The genetic drift is much stronger in smaller populations (it can be simulated on a
computer by looking at the allele frequencies of different replicate population and look
at the allele frequency changes throughout time) lead to the fixation of particular
alleles (the loss of genetic variation)
o But in larger population, still random allele frequency changes, they will all fix but the
process is taking longer  less powerful genetic drift

Genetic drift – allele frequency change in the lab

 Experiment: 107 populations of Drosophila. melanogaster (fruit fly)
 2 different allele at a locus (do not influence survival/reproductive success, they just
change the eye colour)
 each generation founded by 8 males and 8 females
 two SELECTIVELY NEUTRAL eye colour alleles
 allele frequencies = 0.5 (half of the flies have the bw75 allele and half of them have the
wild type) in all populations in the generation zero
 For the allele bw75, some of the population have a decrease in the frequency of the
allele and some have an increase. In the next generation, they spread out even more…
 Genetic drift changes allele frequency at random, meaning that half of the time it will go
up and half of the time it will go down. Eventually they will all fix for one allele or the
other after many generations (50/50)

Another approach to evolution: “quantitative” genetic variation

1. Traits influenced by multiple genes (polygenic). (e.g. human height)
2. Usually influenced by environmental conditions. (e.g. nutrition)
3. Typically leads to continuous variation in the trait
 Genetic evolution in the context of mean traits values that change through time
(quantitative variations) used for polygenic traits.

Most traits are polygenic  continuous distribution e.g. Swimming speed, cognitive ability
(IQ test)

There are 2 ways of thinking of genetic variation

i. Single genes with multiple alleles Calculate allele frequencies
ii. Traits with many genes involved environmental effect  continuous distribution and
how the properties of these distribution changed through times (the means)

Changes in allele frequency at multiple genes underlie genetic changes in quantitative traits
Continuous traits distribution is just simply generated by multiple loci with multiple alleles
 quantitative and polygenic variations are still underline by genes and alleles (they are just
summing up at multiple alleles and loci)
So, there is TWO complementary ways of looking at evolutionary change
i. Changes in allele frequency
ii. Changes in mean values of quantitative traits (e.g. beak size of a bird) (most of genetic
variation are quantitative)

1. NATURAL SELECTION (from quantitative traits)

 Primary evolutionary force that is driving variation and evolution through time
 Comes from Darwin’s and Wallace’s insight
 Changed the way of how people view the world and themselves
 All organisms have the potential for exponential growth – and yet most populations
remain relative stable in the number of individuals (they don’t grow up
exponentially): this implies high mortality in nature (key point #1)
 Traits vary among individuals (not all individuals in any population/species are
identical) (key point #2)
 Traits influence survival (some of the differences in natural population will influence
survival probability and reproduction success) (key point #3)
 Traits are heritable (those differences are heritable) (key point #4)
So, variations among organisms that has a genetic basis that influences who lives and who dies
and who reproduces and who doesn’t. So, the individuals with genetic variations and traits that
are best for survivals and reproduction, will be most likely to pass those genes and traits to the
next generation  increase frequency of those alleles and traits in the next generation

heritable variations that would influence the ability to survive and reproduce would pass
on to the next generation  next generation different with natural selection

Prediction: Traits that improve survival should increase in frequency (or size if it is a
quantitative traits) through time

 Example for the power of selection: Darwin used domestic breeds

 This power of selection is also seen in modern, controlled studies (here, 35 generations
of drosophila). They select from an original population, flies with a high number of
bristles, so they only let the fly with lots of bristles reproduce Evolution of flies with
lots of bristles. Same for the population that only have few bristles.
But does it happen in NATURE?

Example: In 1976 Daphne Major drought.

 During drought conditions, plants don’t reproduce which then causes limited food
availability for the finches.
 The abundance of the seeds on the ground decrease dramatically, and the average size and
hardness of the remaining seed increased.
 During the drought, because plants weren’t reproducing, finches were feeding on the
remaining seeds on the ground that are small and soft (only a few finches could eat the hard
seeds) leaving at the end of the drought, big/hard seeds.
 capture-measure-mark-resigh  measure beak length and depth
 a lot of mortality took place (90% of all G.Firtis individuals died during the drought) High
mortality (key point #1 from Darwin’s & Wallace’s insight)
 The traits of the individuals mattered for who lived and who died the pinches with larger
beak were more likely to survive (not a random selection since the traits influence survival
key point #3)
 Traits are heritable: relationships between beak size of parents and offspring (quantitative
traits)  big beak parents produce big beak kids, so almost all the variation in beak size is
genetically based (beak size is highly heritable)
 The beak size after the drought increased in the next generation  Evolution happened
 Natural selection occurred favouring birds with deeper beak depths, the characteristic of
the population has changed.
 Selection and evolution are highly variable  beak sizes and shapes are going up and down
after the drought (depending on what they are feeding on)
 Some of the genes underlying this heritable trait variation have been identified and shown
to also change in allele frequency during selection in this population  It has recently been
showed that although quantitative traits that are influenced by many genes, there are a few
genes that are having big contribution so there is a mix of quantitative genetic and classic
Mendelian approaches.
 During drought period, Allelic generations at these beaks evolved
 A beak size locus in Darwin’s finches facilitated character displacement during a drought
(during the drought, natural selection acting not only at the beak size but also on the alleles
at particular genes that are influencing that beak size)

4. SEXUAL SELECTION (not about survival but about mating success)

i. Intra-sexual: e.g., male-male competition (for access to female)

ii. Inter-sexual: e.g., female choice (based on male traits) example: widow birds (long tails
are favored at mating success)  sexual selection in widowbirds favours long tails

5. GENE FLOW (mixing of genes between population, they can’t evolve individually from
each other anymore)

 Movements of genes among populations

 Tends to oppose the divergent natural selection
 E.g., exchanging individuals between the two selection lines of Drosophila would reduce
their divergence