Evolutionary Biology 196.
217
Dr Gayle Ferguson (lectures 12 & 13)
Genetic drift: evolution at random
Topics covered
The concept of genetic drift: random change in allele frequencies over time
Relevance of the Hardy-Weinberg Principle to finite populations undergoing
microevolutionary processes
Genetic drift as a sampling error
The effect of population size on probability of allele fixation by drift
The effect of initial allele frequency on the probability of fixation by drift
Coalescence in the history of gene lineages e.g. Mitochondrial Eve
Effective population size
Bottlenecks
Founder effects
Examples of genetic drift operating in natural populations
The neutral theory of molecular evolution
At the end of these lectures you should:
Understand the process of genetic drift: namely, that the frequencies of neutral alleles
fluctuate at random over time as a consequence of sampling error at each generation,
and that fluctuation inevitably leads to a loss of genetic variation (in small or isolated
populations) and the random loss of one (or more) allele(s) and the random fixation of
another;
Understand how different alleles are fixed by chance in different populations;
Understand the relationship between effective population size and the time to fixation
by genetic drift;
Explain the neutral theory of molecular evolution in terms of functional constraints
and synonymous and non-synonymous mutations;
Explain the basis of the molecular clock;
Understand how, given an estimate of the mutation rate, the level of sequence
variation provides a basis for estimating the historical effective size (Ne) of a
population (e.g. the human population, by this calculation, has descended from an
African population of about 10,000 breeding members, from which colonists migrated
into Europe and Asia perhaps only 40,000 years ago).
A question to contemplate
From Futuyma: Some evolutionary biologists have argued that the neutral theory
should be taken as the null hypothesis to explain genetic variation within species or
populations and genetic differences among them. In this view, adaptation and natural
selection should be the preferred explanation only if genetic drift cannot explain the
data. Others might argue that since there is so much evidence that natural selection
has shaped species characteristics, selection should be the explanation of choice and
that the burden of proof should fall on advocates of the neutral theory. Why might
one of these points of view be more convincing than the other?
How might you go about determining which of drift or selection is responsible for the
evolution of a trait, say for example the evolutionary loss of useless organs such as
eyes in many cave-dwelling animals?
Write your own summary of these lectures here taking no more than two paragraphs.
References
Futuyama (2009). Evolution. MacMillan (Chapter 10)
Templeton et al (2001). Disrupting evolutionary processes: The effect of habitat
fragmentation on collared lizards in the Missouri Ozarks. PNAS 98(10): 5426-5432
Grant PR (2002). Founder effects and Silvereyes. PNAS 99(12): 7818-7820
Evolutionary Biology 196.217
Dr Gayle Ferguson (lectures 17 & 18)
Natural selection and adaptation
Topics covered
What is an adaptation?
Historical perspective: William Paley and the argument from design
selection for vs selection of
Definitions and components of fitness
An introduction to the levels of selection
How do we determine if a character is adaptive?
The confounding effects of physics and chemistry, phylogenetic history,
developmental constraint, genetic drift, hitch-hiking and pleiotropic effects
Pre-adaptations and exaptations
Experimental investigation of adaptation by natural selection: the comparative method
An example: the relationship between testes size and monogamy/polygamy in monkeys
The importance of independent phylogenetic contrasts
Examples of natural selection in action: (1) the power of experiments in bacteria
(different mutational effects on fitness; hitch-hiking; (2) inversion polymorphism in
Drosophila pseudoobscura (parallel trajectories); (3) sexual selection (long-tailed
Widowbirds and Guppies - survival and reproductive fitness are not always coupled);
When selection operates on more than one level: (1) genic selection vs individual
selection (segregation distortion in Drosophila); (2) individual selection vs group
selection (Flour beetle)
At the end of this lecture you should:
Know that a feature is an adaptation for a particular function if it has evolved by
natural selection for that function by enhancing the relative rate of success (i.e.
fitness) of biological entities with that feature;
Know that natural selection is a consistent difference in fitness among
phenotypically different biological entities, and is the antithesis of chance;
Understand that natural selection can occur at different levels and that sometimes
multi-level selection may cause conflict;
Understand why genic and individual selections are likely to be the most important
levels at which natural selection acts;
Understand that not all features are adaptations and be aware of the various
confounding factors that give rise to the appearance of adaptation;
Be able to explain how the comparative method can be used to test whether or not a
feature may be an adaptation for a particular function, and discuss experimental
design with reference to illustrative examples;
Be able to describe and give examples of different kinds of natural selection in action,
e.g. sexual selection, individual selection and group selection.
A question to contemplate
How might you test whether or not a population is better adapted after a certain
evolutionary change than before?
Write your own summary of these lectures here taking no more than two paragraphs.
References
Futuyama (2009). Evolution MacMillan (Chapter 11)
Ganetzky (2000). Tracking down a cheating gene. American Scientist 88(2): 128
[Link]
Evolutionary Biology 196.217
Dr Gayle Ferguson (lectures 19-21)
The Genetical Theory of Natural Selection
Topics covered
The modern evolutionary synthesis
Population biology/genetics: definitions and approach
The roles of the 5 microevolutionary processes in bringing about changes in the
gene pool (i.e. allele frequencies) of a population
The relationship between genotype, phenotype, and fitness with respect to
natural selection
Fitness: definitions and mathematical representation of this
The components of fitness in sexual species
Selection coefficient: definitions and mathematical representation of this
Modes of selection (acting on discrete, single-locus, two allele polymorphisms):
homozygote advantage (directional selection), heterozygote advantage (overdominance), heterozygote disadvantage (under-dominance)
Modes of selection (acting on continuously-varying, quantitative traits):
directional selection, stabilising (normalising) selection, diversifying
(destabilising, disruptive) selection
Directional selection: examples (e.g. peppered moth and others); relationship
between dominance/recessiveness and directional selection.
Adaptive landscapes: stable and unstable equilibrium points in populations
undergoing selection; stable and unstable polymorphisms
Balancing selection: mechanisms by which stable polymorphisms are
maintained; examples
Multiple stable equilibria: mechanisms by which history and chance may play a
role in determining the outcome of evolutionary change; examples
Crossing valleys on adaptive landscapes: the interaction of selection and genetic
drift
Molecular signatures of natural selection: neutral theory and the ratio of
synonymous to non-synonymous mutations; long haplotypes as a consequence of
selection and linkage disequilibrium; selective sweeps; molecular signatures of
directional selection vs purifying selection vs balancing selection; examples, e.g.
the lactase persistence in human populations
Even at a single locus, the diverse genetic effects of natural selection cannot be
summarized by the slogan survival of the fittest. Selection may indeed fix the fittest
genotype, or it may maintain a population in a state of stable polymorphism, in which
inferior genotypes may persist. (Futumya)
At the end of these lectures you should:
Understand the theory behind the statement above; understand how the five
microevolutionary processes contribute to change in genetic composition of
populations over time; understand the concept and components of fitness; be able to
describe the different types of selection that may act upon discrete and continuous
polymorphic traits, illustrate these with examples, and (where appropriate) express
these in terms of fitness, selection coefficients, adaptive landscapes, and stable
equilibria; be able to describe how stable polymorphisms may be maintained in a
population; be able to describe situations where the final equilibrium state to which
selection brings a population depends on its initial genetic constitution, and give
examples; understand the relationship between population size, strength of selection,
and the likely outcome of evolution (i.e. selection vs genetic drift); be able to explain
how variation in DNA sequences can provide evidence of natural selection (and link
this to predictions of the neutral theory covered in earlier lectures); be able to
explain how selection can also be inferred from DNA sequence comparisons among
different species (i.e. the incidence of non-synonymous vs synonymous nucleotide
substitutions)
A question to contemplate:
Considering the principles of mutation, genetic drift, and natural selection discussed
in these and previous lectures, do you expect adaptive evolution to occur more rapidly
in small or in large populations? Why? Answer the same question with respect to
non-adaptive (neutral) evolution.
Write your own summary of the lecture here taking no more than a paragraph.
Reference:
Futuyma: Chapter 12