Adaptations Flashcards
What does the evolution of senescence involve?
random effects (drift and mutation) and trade-offs
What is extrinsic damage?
(bad luck)
death/injury/illness from causes for which your genes can’t help much (ie. accidents, overwhelming predators or diseases, weather, etc.)
selection cannot help with this
What is intrinsic damage?
(your genes’ fault)
death/injury/illness from developmental or survival issues that genes could have solved (ie. cancer, heart attacks)
selection can help with this – but didn’t
What is the senescence theory?
if extrinsic damage tended to kill your ancestors when they were old, then intrinsic damage will have evolved to kill you when you are old also, even if you are now protected from extrinsic damage
bad luck tripped up your ancestors, then their own genes evolved to kick them when they were down
What is the mutation accumulation theory to why we have genes that make us senesce?
(random effects)
deleterious mutations that act in old age passively accumulated because selection against them was weak, because frequent extrinsic damage meant that individuals were unlikely to be alive then anyway
involves only random effects (mutation and drift), and failure of selection to act
What is the antagonistic pleiotropy theory to why we have genes that make us senesce?
(trade-offs)
deleterious mutations that act in old age were selected for because of their beneficial effects earlier in life
What are somatic mutations?
mutation may have an effect on the body, but won’t get passed to offspring because it’s somatic – not in the germ-line that would make gametes
What are germ-line mutations?
mutation that happens in the germ-line likely won’t have any effect on that body, but it could get passed to offspring where it would act
What is pleiotropy?
gene has more than one effect
Why might pleiotropic genes be selected for?
high rates of extrinsic damage
- early selection is much stronger than late selection, as old-age individuals are so likely to die anyway
advantage of early reproduction
- better to reproduce early than late (assuming one’s body can be ready early) because early reproducing genes get a shorter generation time and therefore multiply faster
- gaining early babies more than compensates for losing late ones
- reproducing earlier (having a shorter generation time) is advantageous – genotype that starts reproduction earlier will outcompete a slower one, because it will multiply faster if all else is equal
What is a counter-adaptation?
adaptation that counters change to organism’s environment
What is another term for coevolution?
reciprocal adaptation
What is entangled fates?
association that persists through offspring generation after generation that selects for cooperation (inhibits cheating) between lineages – within species, or between species
What are the two consequences of vertical transmission?
- co-speciation (parallel phylogenies)
- evolution of cooperation because of entangled fates
What are transposable elements?
sequences of DNA that can move within and between genomes
persist in genome merely because they have succeeded in replicating themselves – not necessarily intrinsically good or bad
Are transposable elements selfish?
yes – do not normally perform useful functions for the organism
What deleterious effects can transposable elements have?
- metabolic cost of maintaining extra DNA
- introduce mutations when they move
- counter-adaptations to suppress their activity
- can disrupt gene expression
What is isogamy?
same-sized gametes – ie. single-celled alga
What is anisogamy?
some gametes are really big (eggs), some are really small (sperm) – ie. animals, land plants
differences in gametes size means:
- eggs (and consequences of making them) are costly
- sperm are cheap
What is the operational sex ratio?
ratio of sexually competing males that are ready to mate to sexually competing females that are ready to mate
Is variance in reproductive greater for males or females? Why?
males
- females are more or less assured of some offspring (if they are healthy)
- some males get a lot of offspring, others don’t – males are competing
(roles are reversed in some species – if males invest more in caring for offspring, they are the choosier)
What is intrasexual selection?
males compete amongst each other, and females mate with winner
What is intersexual selection?
females choose among males according to their qualities
Female Intersexual Selection
-
What are the costs of sexual reproduction?
- cost of producing males – by dividing resources between daughters and sons, sexual females should be quickly outcompeted by asexual variants (all else being equal)
- destruction of favourable gene combinations
- cost of finding mate
- risk of not finding mate
- risk of disease transmission during matching
What is Fisher’s sex ratio?
total reproductive value of males in a population is exactly equal to total value of all females, because each sex must supply half the ancestry of all future generations of the species
sex ratio will so adjust itself, under the influence of natural selection, that the total parental expenditure incurred in respect of children of each sex, shall be equal
Evolution of Sex Ratios
Which sex has the advantage?
the rarer sex typically contributes more to next generation, selecting for mutations that equalize sex ratio
Evolution of Sex Ratios
What type of ratio would reduce the cost of sex?
female-biased sex ratios – populations with these ratios would be able to grow faster
When do trait values show a continuous distribution?
when many genetic loci contribute equally to a trait
What is the breeding value of a genotype?
tells you how much that individual contributes to the trait mean of its offspring, when assessed against a variety of genetic backgrounds
measures degree to which individual’s phenotype can be expected to be transmitted to that individual’s offspring
NOT a direct measure of the phenotype of an individual
Why does breeding value differ from direct measure of the phenotype?
because it is a function of additive effects of alleles
What are additive effects?
determines how much the mean of a trait changes given that an individual carries a particular allele
What does the additive effect of an allele depend on?
- h (dominance coefficient)
- frequencies of alleles at that locus
- s (selection coefficient) – matter if the trait is under selection
What are additive effects of alleles (A)?
average effect of all alleles that contribute to a phenotype
What are dominance effects (D)?
effects of dominance interactions among alleles at each locus
What are epistasis effects (I)?
effect of interactions among alleles at different loci – reflects reshuffling of alleles over a set of loci
Why does the breeding value differ from the direct measure of the phenotype?
when two individuals produce offspring, they can only contribute 1 allele
- that allele in the offspring may find itself paired in a new dominance relationship
- across loci, that allele will find itself in new epistatic relationships
Why don’t the other components of phenotypic variation matter (much) for QT evolution?
- non-additive effects (D, I) are disrupted every generation, and therefore do not have a predictable effect on offspring
- environmental effects (E) are (by definition) not genetically based
- these three types of variation affect phenotypic trait values (P), but are not predictably passed onto offspring
What is additive genetic variance (VA)?
variance in breeding values within a population
What does evolution by natural selection require?
- variation in a trait
- variation in fitness associated with different phenotypes
- that the trait is heritable
What is narrow sense heritability?
describes proportion of phenotypic variance due to additive genetic variance among individuals, OR extent to which we expect trait variation in to be passed from parents to offspring
What is broad sense heritability?
describes proportion of phenotypic variance due to total genetic variance among individuals
Are heritability (h^2) estimates for populations or individuals
populations – measure of the properties of a trait in a particular population
Does heritability indicate the degree to which a trait is genetically based?
no
rather, it measures proportion of phenotypic variance that is the result of (additive) genetic factors – ie. trait may be under genetic control, but if there is no additive genetic variance associated with the trait, h2 will be 0
Estimating Heritability in Nature Using Parent-offspring Correlation
h^2 = slope of line describing relationship between parent trait and offspring trait values
Estimating Heritability: Parent-offspring Regression
What is the midparent value?
average of phenotypic value of parents
Estimating Heritability: Parent-offspring Regression
What is the midoffspring value?
mean phenotypic value of offspring
Estimating Heritability in Nature Using Other Forms of Relatedness
heritability can also be estimated using other measures of relatedness, and then associating relatedness with trait variation
information about relatedness can come from pedigrees, or from analysis of genomic markers segregating in populations (ie. SNPs)
What would you need in order to predict how much the mean trait value would shift under directional selection?
- starting mean trait value
- heritability of the trait
- strength of selection
Is the selection differential (S) larger or smaller with stronger selection?
larger
What are the possible reasons for limits in long-term selection experiments?
- biological/physiological limits (ie. below a certain level of seed oil, plants are not viable)
- pleiotropy: genes that contribute to traits under selection may have other roles, with negative effects on survival or reproduction
- linkage: genes that contribute to traits may be tightly linked to loci with negative effects on fitness (recombination between these loci would solve this problem)
- lack of variation: if all loci that contribute to a trait go to fixation (= no additive variance) – this is less common than we might expect
What are the possible sources of more extreme traits compared to the original population?
- new environments change the selective value of traits so that traits that were previously neutral (or even deleterious) become advantageous
- new combinations of alleles that were not previously found together or expressed are favoured
- new ‘beneficial’ (desirable) mutations that arise by chance, and then are favoured by selective breeding (longer term)
- in some cases, new environments can reveal different phenotypes and make them available for selection
Building evidence for adaptation involves providing evidence of evolution by natural selection by….
- establishing an association between trait values and fitness values
- establishing that there is a genetic basis for the trait / knowing the genetic basis of the trait (which genes and alleles contribute to the phenotype)
- understanding the mechanism of selection (demonstrating with realistic tests that exposure to a specific setting / ecological context leads to evolution in the trait values)
What are the two possible sources of adaptive alleles (new adaptations)?
- new mutations arising in species or populations as they adapt
- standing variation – alleles already present in species (as neutral or deleterious alleles), which become favoured in new selective context
(traits with complex genetic underpinnings (ie. quantitative traits) also are adaptive)
What is likely to happen to novel mutations that are neutral?
likely to be lost due to drift (even beneficial mutations may be lost)
What type of dynamic would affect novel mutations?
context – if novel environment is so harsh that it requires truly novel traits, and individuals can’t escape the conditions, mutation is unlikely to solve the challenge because relevant mutations are unlikely to appear at just the right moment
What populations risk extinction when the environment changes?
populations that are small and have low additive genetic variation
large additive genetic variation and/or large population can allow evolution to rescue declining populations
What is QTL mapping?
(mapping of quantitative traits) scores set of F2 individuals for random genetic markers, and builds linkage maps
- variation in traits of these same F2 is then statistically associated with the markers
- loci explain small/large proportions of phenotypic variance
How can we uncover the genetic basis of adaptive traits?
- QTL mapping
- simple plots that show distribution of phenotypic trait scores in F2 population (as well as parental and F1 values), give some hint about genetic basis and degree of dominance of traits
Are mutations of very large effect common?
relatively uncommon, but many effect sizes are also not super small
Small effect loci/mutations have lower potential to…
`yield large, rapid changes in trait means in response to selection
Large effect loci/mutations have potential to…
be highly disadvantageous, if they take you away from the optimum that selection is favouring
but these mutations are quickly removed by selection, and therefore we should still see evidence of large effect beneficial mutations
Do large populations change more slowly or quickly than smaller populations due to drift?
slowly
When can gene flow act?
if two populations differ in frequency or presence/absence of different alleles, then gene flow can change their allele frequencies
What type of populations share more gene flow?
nearby populations