Evolutionsbiologie
ETH Zürich
ETH Zürich
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Cartes-fiches
Bottleneck effect
Population has undergone a temporary restriction that has severely reduces its genetic diversity; Epidemic, natural disaster like fire or flood
Founder effect
Populations are small when they hav just been founded; Allele frequency in new population are likely, by chance, to be different than before => founder effect; direct result of sampling error
Chance, Determinism and Evolution
Mutational input is random; Mutation is random with respect to effects on fitness, no drive to mutate in advantageous directions; Evolution by selection is non-random, some gentic variants reproduce at higher rates than others; Evolution by genetic drift is random, chance events result in changes in allele frequency; Power of drift is inversely proportional to the power of selection
Effect of drift alone on diversity
All alleles would drift to either fixation or loss; Drift will reduce genetic diversity until only one allele is left
Effect of population size on rate of change by drift
Evolution by genetic drift occurs more rapidly in small populations; but it can also change frequencies in large populations over long periods of time; on allele will drift to fixation and all others are lost
Loss of heterozygosity
Only surviving oryx deer population in zoos and natural reserves, 50 breeding individuals; diploid organism, 2 alleles per gene; Heterozygosity in the absence of mutation and selection: Hnew = Hold * (1 - 1/2N) = Hold * 0.99
Probability of fixation by genetic drift
Probability that a given allele will drift to fixation is equal to its fequency in the population; Probability of fixation: P(fix) = x/2N for diploids, x = number of allele copies, N = number of diploid individuals
Genetic drift in experimental fruit fly populations
107 populations with each 8 males and females; All populations initially heterozygous for eye-color gene brown, genotype bw/bw75; Drifted towards extreme allele frequencies, equally in both directions; Heterozygosity decreased over time
Effective population size NE
Are especially sensitive to unequal sex ratios in populations; the higher inequality the lower NE; The lower NE the higher the effect of genetic drift; NE = (4 Nmales*Nfemales)/(Nmales*Nfemales)
Non-random mating (NRM)
Causes deviations from HW equilibrium; Inbreeding: disproportionate mating among relatives, reduces heterozygote frequency relative to Hardy-Weinberg; Assorttative mating: positive with like and like mating or negative with opposite attraction; NRM doesn't cause evolution but influences it => Inbreeding doesn't alter allele frequency but genotype distribution; Coefficient of inbreeding F
General analysis of inbreeding
Heterozygosity H in an inbred population is always less than ina random mating population, inbreeding coefficient is non-zero => Hinbred = Hrandom*(1 - F)
Inbreeding can depress average fitness
Inbreeding depression (sigma): reduced fitness in individuals due to kin matings => sigma = 1 - omegaselfed / omegaoutcrossed; results from: reduction in the frequency of superior-fitness heterozygotes, increase in frequency of homozygotes for deleterious alleles; Inbreeding depression in humans, higher mortality; Selection of inbreeding avoidance: mate choice, migration, self-incompatibility
Conservation biology example: greater prairie chicken
Declining population size, declining male numbers => declining fitness; Population crashed (bottleneck), ineffectiveness of selection in small populations, increased inbreeding and increased chance evolution 8drift), reduced heterozygosity, loss of variants, accumulation of deleterious alleles, reduced fitness; Migration increases H and fitness, populations size grows, selection gets more effective in cleansing deleterious alleles => conservation could stop extinction vortex
Two locus version HWE
Evolution at one locus cann affect evolution at another locus
Linkage
Two loci are on the same nonrecombining stretch of a chromosome; 1. Loci are close enough together on the same chromosome that they remain together after meiotic crossing 2. In nonrecombining organisms, the loci always stay together on the same chromosome
Linkage and Selection
If two loci are linked, selection on one locus can affect fate of the other; Multilocus genotype on one chromosome is called a haplotype
In linkage equilibrium (LE)
Genotype at one locus is evolutionary independent of genoytpes at another locus; Genes are on seperate chromosomes or far apart on the same; Haploptype frequencies calculated directly from allele frequencies
In linkage disequilibrium (LD)
Non-random association between genotype at one locus and its genotype at the other locus; haplotype frequencies can't be calculated directly from allele frequencies (Violates condition of Hardy-Weinberg)
A population is in LE when:
1. The frequency of B on chromosomes carrying allele A equals the frequency of B on chromosomes with a 2. Frequency of any haplotype can be calculated by multiplying the frequencies of the constituent alleles; gXY = f(XY) = f(X)*f(Y) 3. Coefficient of linkage disequilibrium D is 0, D = gAB*gab - gAb*gaB, g = genoytpe frequency
Evolutiionary forces increasing LD
Under Hardy-Weinberg and linkage equilibrium, haplotype frequencies don't change over time; Deviations from HWE suggest one of three mechanisms at work: 1. Selection on multi-locus genotypes 2. Genetic drift 3. Population admixture (Migration)
Disequilibirum due to selection on haplotypes
A non-random subset of haplotypes survives
Sexual recombination reduces LD
Curves showing the decline in LD, according to D' = D(1 - r) with different values of r; r reflects likelihood of uncoupling by recombination, rmax = 0.5 for fully oncoupled genes on different chromosomes; HW assumes rate of decline proportional to r
Coefficient of linkage disequilibrium D
Maximum value is 0.25, minimum is -0.25; Ex: f(A) = f(a) = f(B) = f(b) = 0.5, a) gAB = gab = 0.5, gaB = gAb = 0 => D = 0.25, b) gAB = gab = 0, gaB = gAb = 0.5 => D = -0.25
Special form of LD: Hitchhiking
Selection on one locus causes changes of frrequencies of alleles at a linked locus that is not under selection; When selection is positive, it's called hitchhiking; PROBLEM, when not considering linkages, locus wrongly accused of being under selection
Hitchhiking of b with A
a) Increase in A leads to increase in b because A is together with more b than B, and the alleles tend to stay together (LD) b) Increase of A has no effect on B or b because the two loci are uncoupled (LE)
Why have sex?
Life originated without sex (cell doubling in bacteria); Downside of sex: Find mates, choose/compete for mates, risk of diseases, offspring has to be fertile, one offspring is only half the bill
Paradox of sex
Sex diulutes genes by half; Imaginary null population with sex vs. no sex; doesn't affect 1. number of offspring per mother 2. survival or reproductive successs of offspring; Population founded of two sexual individuals and one asexual; Every generation, each female produces 4 more offspring => asexuals increase with every generation
Male prental care explaining sex?
Male care offspring increase female fitness, Ex: humans, birds; BUT in most species, males give away their genes and run...
Sex helps fitness in long run
Testing wether the second assumption of Maynard-Smith ist true: survival and reproductive success of offspring; Red and black markers to distinguish between sexual and artificially asexual beetles; In the oresence of insecticide, sexual individuals survive better the more of it there is
Evolutionary reasons for sex
Linkage disequilibrium happens; Sex can have no effect, and hence no benefit in a population at linkage equilibrium, sex happens; Why is sex beneficial: 1. Those that posit creation of linkage disequilibrium by drift, sex allows selection to purge unfavorable multi-locus allele combinations that might otherwise drift to fixation 2. Those that posit creation of linkage disequilibrium by selection, sex allows selection to promote new favorable multi-locus allele combinations that would otherwise not be created
Muller's Ratchet, drift model for sex
In small populations: asexual populations accumulate more and more deleterious mutations by drift, the best can be lost by drift, forward mutations to deleterious mutations greater than backward mutations, increased genetic load, decreased average fitness, sex breaks the ratchet, allows resetting to zero mutation genotypes; Ratchet stronger in small populations, when N>1000, drift is too weak Ex: mutation accumulation in populations with strong bottleneck
Benefits of sex manifest only in the long term stopping Muller's ratchet, in short term asexual females outcompete sexual ones
Rapid adaption to environments, selection model for sex
Sex mode is more likely to create new variants being more fit to novel environments, Ex: more parasites => more male snails and thus more sexual reproduction
First sex
Volvox; Individuals retain ability to become female, male or asexual until late development
Identifying Adaptations
How to know whether a trait is adaptive? Adaptation are traits enhancing fitness; Test the effect on fitness by removing a trait
Testing a Mutualism Hypothesis
Hypothesis: oxpecker-host relationship is mutualistic; Birds thought to reduce tick load and gain easy meal; BUT birds were not interested in tick load relative to blood/earwax; Shooing birds consistently reduced number and severity of open skin wounds on host => Original hypothesis unlikely
Adaptation Caveats
Many trait variations are likely to not reflect adaptations; Not all phenotypic traits are adaptations; Adaptations do not reflect optimal engineering design
Testing adaptation hypotheses
Experiments in which study groups differ by only one experimentally sontrolled variable; Observe; Perform comparative studies to test correlations among variables that suggest causal relationships
Designing a good experiment
1. Ask an interesting question 2. Generate a list of most plausible answers 3. Design experimental treatments that distinguish between all plausible hypotheses 4. Include control groups 5. All treatment groups to same conditions (except for tested variable) 6. Randomize 7. Replicate sufficiently
Why do Zonosemata flies wave marked wings?
Behavior similar to fly's major predator; 3 hypothesis: 1. no mimicry (null) 2. mimicry to scare away other predators 3. mimicry to scare away major predator
Different bat species have different sized testes. Why?
Sperm competition hypthesis: multiple matings selection for larger ejaculates => increase chance of fertilization; Larger groups allow for more multiple matings; Testes size correlate with group size; Correlation analysis supports hypothesis; BUT if species with similar traits are also more closely related than species with different traits, we've got non-independence of our data points => Patterns may be shaped by phylogenetic relatedness rather than selection
If correlation holds up after adjustment for phylogenetic relationships => Correlation reflects causation; Here, bat testes size correlates with group size after correction for phylogenetic relatedness