NATURAL SELECTION AND ADAPTATION

STUDY UNIT 2

Biological evolution

- Change in allele frequencies in a population over time

- Allele frequency: Relative frequency of an allele at a particular locus in a population
- How often an allele is observed within a population
- Allele: Version of a gene that controls particular feature in an organism
- Genotypic frequency: How often a genotype is observed within a population
- Phenotypic frequency: How often a phenotype is observed within a population

Hardy Weinberg principle Neutral alleles

• Do not alter tness
- Random mating
• Average reproductive success does not
- No mutations differ between individuals that carry one
- Large population size (genetic drift) neutral allele or the other
- No natural selection • Frequencies may change in a population by
genetic drift
- No immigration (gene ow)

- Allele and genotype frequencies remain constant across generations

- A population in HWE does not evolve
- HW equation: p 2 + 2pq + q 2 = 1

Forces of evolution that lead to speciation

Recombination and effective
- Each force can increase / decrease genetic diversity in a population population size (Ne) increase
genetic diversity in population
- Genetic diversity: Variation seen within and between species on a molecular level
- Raw material for adaptation
- Provides species with the capability for adaptation in the face of evolutionary change to
overcome speci c evolutionary barriers
- Need variation at genetic level in order to survive
- Associated with the number and frequencies of different states

1. Non-random mating
- Assortative mating: preference for similar genotypes / phenotypes
- Disassortative mating: preference for different genotypes / phenotypes

2. Mutation
- Original source of variation within organisms
- Increases genetic diversity in population
- Mutation rates for most organisms are low
- There is no impact of a new mutation from one generation to the next, unless NS is acting

3. Genetic drift
- Random change in the gene pool of a small population that takes place purely by chance
- Alleles xed / lost at random
- Decreases genetic diversity in population
- Always present in a randomly mating population
- But in large populations, its effect on allele frequencies is so small that it becomes impossible to
quantify
- The smaller the population, the more profound the effect of genetic drift

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 Bottleneck effect
- Severe catastrophic event kills off most of the population
- Small number of original population remains
- Several alleles that were in the population are lost
- Genes in the surviving population occur randomly

- Large reduction in genetic diversity

- Very high probability of inbreeding
- Original population does not exist anymore
- Original diversity is lost forever
- New diversity can be introduced through mutation / migration

Founder effect
- Small group of individuals separates from a larger group and expresses genes that were rare
in the original population
- These individuals are the founders of a new population
- Only have some of the original variation
- Rare genes become more common in the next generations

- Large reduction in genetic diversity

- High probability of inbreeding
- Original population still exists
- Diversity is preserved
- Destination population can gain new diversity through mutation / migration

4. Migration
- Type of gene ow when organisms are introduced into a population
- Change of allele frequency or introduction of new alleles into population
- Increases genetic diversity in population

5. Natural selection

- Any consistent difference in tness among different classes of biological entities

- Fitness = number of offspring an individual leaves in the next generation
- Components of tness = survival + reproduction
- Reproductive success includes survival
- Organisms cannot reproduce when they are dead
- ∴ tness ≈ reproductive success

- For NS to take place, the following genetic features are required:

1. Variation
- NS can only act on standing genetic variation
2. Heritability
- Both advantageous and disadvantageous traits / alleles must be heritable
3. Differential reproductive success
- Presence / absence of a trait / allele must have an in uence on reproductive success
- Those with disadvantageous / detrimental alleles must produce less offspring

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 - Evolution by NS:
- Requires a change in a population across generations
- Occurs if there is a:
1. Correlation between individual’s phenotype and tness
2. Correlation between variation in phenotype and parents and offspring
- In sexually reproducing populations entails changes in allele frequencies that underline
phenotypic variation, which affects tness

• Genotypes A and B (in an asexually reproducing species) differ in a characteristic that affects their
tness
• Fitness of A = 3
• Fitness of B = 4
• If these values are constant across generations, B increases in number much faster than A and will
make up the great majority of the population within a few generations
• B is more t because it reproduces faster in less generations
• ∴ frequency of B has increased and frequency of A has declined
• B (which starts at a frequency of 0.5) makes up almost 90% of the population just 7 generations later

• Fitness is more complicated in sexually reproducing organisms

• Sexual selection = natural selection arising through preference by one sex for certain
characteristics in individuals of the other sex
• Males vary in survival, reproduction and reproductive success
• Males compete for females
• Eg Peacock’s tail

- Natural selection can cause extinction of species

- NS selects for the ttest individuals (“survival of the ttest”)
- These individuals survive and reproduce
- Standing variation exists within populations
- Some individuals are selected for and reproduce
- Other individuals are selected against and die
- Over the course of evolution, species currently in the environment (the survivors) are there
through NS and adaptation

Adaptation

- A characteristic that enhances the survival / reproduction of organisms that Peppered moths
• Industrial evolution
bear it, relative to the alternative character states, in a speci c environment • Darker moths survived
- Evolved by natural selection • Alleles passed on
Elephants
• Elephants with tusks hunted
and killed
• Alleles not passed on
• No elephants with tusks
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- Remember that evolution is blind
- There is no forethought, aim or goal in the course of evolution
- It is simply natural selection acting on standing variation in populations
- Standing variation = variation contained in a population in its current state, not the potential
variation that could result from evolutionary forces

Adaptive evolution observed

- Adaptive evolution can be rapid, especially when species are introduced

Soapberry bugs
Beak length has evolved rapidly as an adaptation to the new host plants

• Bugs feed on seeds of plants in soapberry family by

piercing the enveloping seedpod with its slender beak
• In the last 50 years, related species of Asian trees have
been abundantly planted in Texas and Florida
• The bug populations feed mostly on these species
• Asian tree species in Texas have a larger pod than the
original host plants
• Asian tree species in Florida have a smaller pod than the
original host plants
• ∴ soapberry bug populations in Texas have evolved a
longer beak
• ∴ soapberry bug populations in Florida have evolved a
shorter beak

Overexploitation of populations
Commercial overexploitation has severely depleted populations of many species of sh and has resulted in
evolutionary change

• In many species, there has been a trend toward earlier

sexual maturation at a smaller size
• Larger age classes are more subject to predation
• In some species (Atlantic cod), these changes have a
genetic basis
• Fishing for Atlantic cod of the largest size = evolution of
smaller size reaching sexual maturity
• Trophy hunting for bighorn sheep with the largest horns =
evolution of smaller horns
• In both instances, the quality that adds value to the resource
has been diminished by the response to selection

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 Nature of adaptations

Adaptation
- Evolutionary process by which, over the course of generations, organisms are altered to become
improved with respect to features that affect survival or reproduction
- A characteristic of an organism that evolved by natural selection

Preadaptation
- Possession of the necessary properties to permit a shift to a new niche, habitat or function structure
is preadapted for a new function if it can assume that function without evolutionary modi cation
- Feature fortuitously serves a new function

- Eg Parrots
- Strong, sharp beaks for feeding on fruits and seeds
- Introduction of domesticated sheep into New Zealand
- Some attacked by parrots (kea) which pierced the sheep’s skin and fed on their fat
- Kea’s beak useful for this activity

Exaptation
- Evolution of a function of a gene, tissue or structure other than the one it was originally adapted for
- Adaptive feature which co-opted for a new function during evolution
- The adaptive use of a previously nonadaptive trait

- Eg Puf n
- Wings are exaptations for swimming
- Birds “ y” underwater and in air

- An exaptation may be further modi ed by selection ∴ the modi cations are adaptations for the
feature’s new function
- Eg Penguins
- Wings have been modi ed into ippers that enhance swimming but cannot support ight in air

Selection of and selection for

- A feature is only called "adaptive" when we know what it is adaptive for

- How did it increase tness?
- What is its function?
- Two linked traits will both pass through a selective bottleneck when only one of them is selected for

• Balls are selected for the property of small size

• Balls are not selected for their colour
• ∴ there is selection of red balls, which happen to be the
smallest
• Similarly, NS may select for a particular feature
• There may be incidental selection of other features
correlated with that feature
• The function of a feature is associated with the selection of the
feature itself
• Feature causes its bearers to have higher tness
• Feature may have consequences that were not selected for

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 Recognising adaptations

- Not all traits are adaptations

1. Could be a necessary consequence of physics / chemistry

- Red colour of blood is not an adaptation
- The adaptation is the presence of haemoglobin to bind oxygen
- Redness = byproduct of haemoglobin’s structure

2. A non-adaptive trait may be correlated with an adaptive trait

- This occurs when a trait has been co-inherited with a selected trait and becomes adaptive when
the environment changes

3. The trait may have evolved by other mechanisms (random genetic drift) rather than NS

4. The feature may have evolved because it was correlated with another feature that conferred an
adaptive advantage

5. Character state may be the consequence of phylogenetic history

- A hummingbird has four toes
- But all birds have four toes

How to recognise adaptations

Complexity
We assume that if a trait is complex
- Cannot evolve without natural selection in nature, it evolved through NS

Design
- Correlates with the predictions of a model
- Many plants that grow in hot environments have leaves that are divided into lea ets / tear along
fracture lines
- These features conform to a model
- Thin, hot “boundary layer” of air at the surface of a leaf is more readily dissipated by wind
passing over a small surface
- Divided leaf is cooled more effectively

Experiments
- Show that a feature enhances survival, reproduction or performance
- Several oral characters have evolved convergently in the many plant lineages that have shifted
from insect pollination to bird pollination
- Some of these features are advantageous - facilitate bird pollination and discourage bees
- Bees = comb much of the pollen into a mass and feed to their larvae = less effective pollinators

The comparative method

- Compare sets of species to test a hypotheses
- Takes advantage of natural experiments provided by convergent evolution
- If a feature evolves independently in many lineages due to a similar selection pressure, we can
often infer the function of that feature
- Bird vs bat wings
- Wings are adaptive (and convergent) but the bones in them are not

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 Imperfections and constraints

- Darwin noted: ”natural selection will not produce absolute perfection, nor do we always meet, as far
as we can judge, with this high standard in nature”
- Selection can x only those genetic variants with a higher tness
- The best possible variants often fall short of perfection because of various constraints
- Among these constraints are trade-offs Sacri ce for adaption
- Eg With a xed amount of available energy / nutrients, a plant species might evolve higher
seed numbers but only by reducing the size of its seeds

- Lack of suitable genetic variation may explain cases of phylogenetic constraints

- Species retain nonadaptive features / are unable to evolve adaptive traits
- It makes adaptive sense that swans have more neck vertebrae than birds with shorter necks
- But almost all mammals have seven neck vertebrae (including giraffes and whales) despite the
extreme differences in the length of their necks
- Eg Vagus nerve is 15 feet long which is not “intelligently” designed - feature is to complicated for
evolution to x
- Evolution cannot x a “badly designed” feature

NS and the evolution of diversity

- Any characteristic of an organism is likely to be advantageous under some circumstances but not
others
- Optimal feature = character that maximises tness depends on the context in which it functions
- Eg Camou age in animals - colour and patterns that match the background lower the likelihood
that the animal will be detected by predators
- Different variables may be relevant to a particular species and impose NS on many of its features
- NS is the ultimate cause of of divergence among populations and species
- NS is the source of immense diversity of life

- If different closely related species coexist, those individuals that use different resources from the
other species would suffer less competition and have higher tness
- The species will diverge from the others.
- Character displacement: divergence of species as a consequence of their interaction
- Eg Ground nches in the Galápagos Islands
- Finches with larger, deeper bills feed more ef ciently on larger, harder seeds
- Species with different bill depth differ accordingly in diet and the species that coexist on any
island differ, matching the availability of different seeds

Levels of selection

- A hierarchy in which selection can occur among genes, cell types, individual organisms, populations or
species
- Genic selection = NS at the level of the gene
- Illustrated by transposable elements
- Replicate / proliferate within genome, regardless of whether it has a positive or negative effect
on the organism
- Depict sel sh genetic elements

- Some sel sh alleles exhibit segregation distortion and are passed to a heterozygous individual’s
gametes more than 50% of the time

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 - Segregation distortion can result from meiotic drive (in which meiosis does not follow Mendel’s laws)
and from other processes that happen after the gametes are formed
- Eg t locus of the house mouse
- In a male heterozygous for a t allele and for the normal allele T, the t allele kills sperm that
carry the normal allele
- ∴ more than 90% of the male’s sperm carry t
- Embryos that are tt homozygotes die / are sterile
- Despite the disadvantages to the individual, segregation distortion is so great that the
disadvantageous t allele reaches a high frequency in many populations

- Sel sh genetic elements forcefully illustrate the nature of NS

- It is differential reproductive success which need not result in adaptation / improvement
- Selection among individuals is at a “higher level” than selection among genes
- Selection at the gene level may act in opposition to individual selection
- It may be harmful to individual organisms, and might even cause the extinction of
populations or species

Sel sh genes & unsel sh behaviours

- Selection among genes can be based on the effects that change their frequencies across generations
- These effects can enhance the survival of relatives that share the same gene
- ∴ any gene that has successfully increased in frequency is a sel sh gene

- Altruistic traits = bene t other organisms at a detriment to itself

- Best described by kin selection
- An allele for altruistic behaviour can increase in frequency in a population if the bene ciaries of
the behaviour are related to the individual who performs it
- Kin selection = a form of selection in which alleles differ in tness by in uencing the effect of their
bearers on the reproductive success of individuals who carry the same allele by common descent
- Eg Parental care
- Alleles that enhance a parent’s care-giving behaviour have increased in frequency because they
promote the survival of identical copies of those same alleles that the offspring carry

Selection of groups & organisms

- Evolution of traits by individual selection

- By the replacement of less t for more t individuals across generations
- The possibility of future extinction cannot affect the course of evolution

- An altruistic trait cannot evolve if it reduces the tness of an individual that bears it (even if it
bene ts the population / species)
- An altruistic genotype amid sel sh genotypes will produce fewer offspring
- Will decline in frequency
- The sel sh mutant will increase to xation (even if the population of sel sh organisms is at
higher risk of extinction)
- Group selection: differential production / survival of groups that differ in genetic composition
- Enables the evolution of an altruistic trait
- Eg Population made up of sel sh genotypes (high reproductive rates) might have a higher
extinction rate than populations made up of altruistic genotypes (low reproductive rates)

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 - The survival of altruistic groups of individuals causes the species to evolve altruism (even though
individual selection within each group acts in the opposite direction)

- Williams argued that:

- Adaptive traits bene tting populations / species can be explained by individual bene ts or might
not be adaptations at all
- Eg Egg laying
- Females lay fewer eggs in high-density, scarce-food situations, not to ensure suf cient food
supply for the species but because they cannot form as many eggs
- William’s opposition to group selection:
- Individual turnover (birth/death) is much faster than population turnover (formation/extinction)
- Replacement rate of less t (altruistic) by more t (sel sh) individuals is much greater than the
replacement rate less t by more t populations
- ∴ individual selection prevails over group selection

- Majority believe that kin selection (altruism ad cooperation) play a signi cant role in evolution
- Others believe group selection plays a signi cant role in evolution

Species selection

- Selection among groups of organisms, when the groups involved are species and there is a correlation
between some characteristic and the rate of speciation / extinction
- Does not shape adaptations of organisms but it does affect the disparity (diversity of biological
characteristics)
- Consequence = proportion of species that have one character state rather than another changes over
time
- Eg Prevalence of sexual species compared with closely related asexual forms
- Many groups of plants and animals have given rise to asexually reproducing lineages
- Asexual lineages tend to be young (indicated by their close genetic similarity to sexual forms)
- Few asexual forms that arose long ago have persisted ∴ asexual forms have a higher rate of
extinction than sexual populations

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