Eve Final Flashcards
Codon bias influenced by
-mutational bias and genetic drift
Natural selection
Gene expression
Translational efficiency
Codon bias
No random usage of synonymous codons to encode a given amino acid
(Different codons that correspond to the same amino acid appear at different frequencies in the genome)
Codon bias example
Mutations from G to A and from C to T are TWICE AS COMMON
(Tends to favor accumulation of codons with A and T bases)
Natural selection as a cause of mutation bias
translation of highly expressed genes is most efficient when their codons correspond to transfer RNA that are common; so selection favors these codons
Can also favor codons that produce messages that are less prone to translation errors
Codon bias driven by selection
tends to be stronger in genes that are highly expressed and in species with very large population sizes such as free living microbes
Codon bias and population size
Species with smaller population sizes drift, overwhelms, whatever selection actions anonymous mutations, and codon bias is very weak or absent
STRONGER IN LARGE POP
Synonymous mutations weakly selected, in LARGER POP…
Selection MORE IMPORTANT; Ns»>1
SMALLER POP MORE INFLUENCED BY DRIFT
Male driven evolution
male germ line mutation rate is higher than the female germ line mutation rate, the Y chromosome is exposed to a more hostile mutation environment so its going to incur more mutations per generation
Y chromosome evolves very very fast
mutation rate. Is 3 x greater
Nonsynonymous sites are more
Clock-like than synonymous sites
dN
Non synonymous site
Expected dN/dS for a “typical” gene
<1
Expected dN/dS for a gene coding for an unconstrained protein
=1
Recurrent directional selection
dN/dS >1
What requires dN/dS»1
Very conservative test for adaptive evolution, as large numbers of selective fixations
Directional selection and evolution rate
Very rapid evolution
What does a smaller dN/dS ratio suggest
rates of protein evolution are slowed down by functional constraint
Recurrent directional selection ratio; dN/dS»_space;1
Very RARE
have to have a lot of amino acid changes going through under selection
Very frequent adaptive evolution has been acting on it
BIG dN/dS tells…
evolving adaptively - potentially its biology is changing
Examples of genes dN/dS >1
- Vertebrate immune system
- Viral coat proteins, other pathogens
- Reproduction related genes
More powerful way of detecting adaptive protein divergence (compared to dN/dS) ?
McDonald-Kreitman (MK) test
McDonald Kreitman test
Under the null hypothesis (neutral evolution), polymorphism and divergence result from genetic drift
Rapidly evolving proteins should also show high levels of protein polymorphism within species
Polymorphic and fixed a.a variants are compared to polymorphic and fixed synonymous variants
Can only detect recurrent adaptive fixations and can only reject if there are sufficient numbers of variants in a gene
A protein under very LOW FUNCTIONAL CONSTRAINT should show a lot of
Protein polymorphism
diverging very quickly between species
Gene under STRONG CONSTRAINT
should show very little polymorphism at amino acid level and show be evolving very slowly at the protein level between species
polymorphism is
positively correlated with recombination
stronger selection/lower recombination
larger hitchhiking effects
hitchhiking effect
as the copies of the beneficial mutation spreads to higher frequency, the neutral variance linked to them hitchhike to higher frequency
mutation becomes fixed, genetic variation is eliminated in the region nearby
Hitchhiking: Regions further from beneficial mutation
Regions of the chromosome further from the beneficial mutation retain variation because recombination joints together chromosomes that carry the beneficial mutation with chromosomes that carry neutral variants
polymorphism remains at regions farther away on the chromosome
this is a telltale sign used by geneticist to find evidence of recent adaptation
background selection
rather than the spread of beneficial mutants having this effect on the genome, its the removal of deleterious mutations that have this effect on the genome
Introduce mutation but selection removes that chromosomal region out
(basically reducing effective population size of chromosomes)
rapid spread of beneficial allele to intermediate frequency
generates long regions of linkage disequilibrium
evolution of human skin pigmentation
indirect result of selection on thermoregulation
- selection for hair loss
- COMPENSATORY selection for skin pigmentation
theory for evolution of lighter skin
selection for vitamin D related traits at high latitudes
more pigment protecting against folate depletion at low latitudes carries a cost and is unnecessary at higher latitudes
light skin alleles evolution
significant amount of lighter pigmentation seems to have evolved independently in groups of European and Asians
eye color: before or after
eye color might have evolved before pigmentation
Species: BSC
groups of actually or potentially interbreeding populations, which are reproductively isolated from other such groups
GENE FLOW (reproductive isolation) is focus
cryptic species
isolated but with little phenotypic differentiation
problems with BSC
asexuals
extinct groups
gene flow between “good” species
breeding experiments impractical
phylogenetic species concept
two populations are species if they are different in MORPHOLOGY, DNA
species are often
reproductively isolated AND phenotypically diverged
speciation is the
evolution of reproductive isolation or the absence of gene flow between two populations is required for them to no longer exchange genes with each other
mechanisms of reproductive isolation
pre zygotic
post zygotic
premating barriers
features that impede transfer of gametes to members of other species
potential mates do not meet
potential mates meet but do not mate
postmating
mating occurs, but zygotes are not formed
postzygotic barriers
hybrids are formed but have reduced fitness
extrinsic
hybrids have low fitness for ENVIRONMENTAL reasons
not well adapted
cant obtain mates
intrinsic
low hybrid fitness is independent of environmental incompatibility
Haldane’s Rule
when hybrid breakdown is differentially expressed in the two sexes, the HETEROgametic sex is usually affected more severely
which appears first; sterility or inviability
hybrid sterility often appears before hybrid inviability
Most common mode of speciation
Allopathic speciation
Allopatric speciation
Two populations have to become geographically separated to eventually evolve these reproductive isolating mechanisms
Might be able to reproduce if they come back together; depends how diverged they are
Allopatric divergence
populations that are more geographically distant from one another have accumulated more genetic differences from each other
More reproductively isolated they are
The more geographically different the population are the less likely they are to meet