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81 Cards in this Set

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Linkage disequilibrium
Allele at one locus is non-randomly associated with allele at another locus
Non-random assoc. of alleles at 2 different genes in a population
Linkage equilibrium
Allele at one locus is independent of allele at another locus
Evol at one gene doesn't affect evol at another gene
D
coefficient of linkage disequilibrium
D=0 if
genes are independently sorting, because...
all combinations are equally frequent
When linked genes show LD
chromosome freqs will change from one generation to the next, moving toward linkage equilibrium
Factors that create linkage disquilibrim
1.
2.
3.
4.
5.
1. Selection on multilocus genotypes
2. Physical linkage
3. Genetic drift
4. Population admixture
5. Assortative mating
What reduces LD?
Sexual reproduction. How?
Meiosis, genetic recomination, outbreeding
Usefulness of LD?
1. Detect physical linkage
2. Map genes for trait of interest
3. Infer population history
4. Detect positive selection on genes
LD example:
Ashkenazim with 8-repeat locus more likely to have 84 GG disease mutation
Able to date disease via the rate of recombination (750-2325 years ago)
Selective sweep
(aka hitchhiking) neutral genetic variation linked to a positively selected mutation will also become more prevalent
That haplotype becomes only one in pop, resulting in large reduction in total genetic variation in that chromosome region.
Sexual reproduction reduces LD
via meiosis and genetic recombination
Sexual reproduction is costly because...
1. Cost of finding mates (time and energy)
2. Exposure to disease
3. Possible infertility
4. Two-fold cost of males
Explain the 2-fold cost of males
Parthenogenetic females can make 2x as many offspring
Draw it out.
Why sexual selection better?
1. Genetically unique individuals
2. Recombin. create new multilocus genotypes
3. New combinations may be more fit
4. Breakdown LD
Muller's ratchet
Genetic load (deleterious mutations) increases; decreases fitness
Sexual pop. can purge deleterious mutns. Long term benefit of sex.
What are the short-term benefits of sexual repro?
1. Adaptation in a heterogeneous environment
2. Some multilocus g.t.s favorable in one envt; less favorable in future envt.
3. Quick evolution for orgs locked in evol arms race.
But...
Sometimes sexual repro is favored by competitors (e.g. parasites favoring sexual repro in snails)
How is LD created?
1.
2.
3.
1. Selection. How?
2. Drift. How?
3. Migration.How?
1. can't predict freq of haplotype by multiplying freq of alleles
2. Finite population - fixed for A. Mutation causes A->a. Now always linked to b because occurred on same chromosome.
3. Pops start in LE. Mix pops together and observe LD.
Compare sexual selection to natural selection
Nat Sel: selecting more fit indivs (better at repro)
Sex Sel: selecting indivs better at obtaining mates
Derived character
evolved in response to a selective agent.
"Everything was perfectly designed for its purpose." Viewpoint known as
Panglossian. Held until 1960s when...
biologists began recognizing trade-offs.
Antagonistic pleiotropy
Gene that regulates more than one trait; some may be bad for fitness.
What did the Spandrels of San Marcos analogy predict?
Architectural consequences of the arches coming together were not intentional, just there
Can't assume every feature is an adaptation.
How can we test hypotheses about adaptations?
1. Experimentation
2. Direct observation
3. Comparative method
1. Involves some kind of manipulation
2. Relies on measurement making w/o interference
3. Spp. are not indepentdent data points
Testing an adaptation example: Why do zonosemata flies wave their wings?
To deter jumping spiders by mimicking major predator.
Comparative method accounts for...
phylogeny
Remove effects of relatedness so evaluation is more compatible with statistical methods.
Independent contrast
A comparative method testing mode for adaptation (example- testes size in large bat social group)
1. Compare sister taxa as pairs
2. Any difference btwn species must have been acquired AFTER divergence from a common ancestor
3. Slide all points (now lines) so one point meets origin of graph
4. Treat remaining points like a regular regression
What are limits to adaptation
trade-offs, constraints.
What is the Darwinian Demon?
1. Sexually mature at birth
2. Infinite offspring
3. Reproduces constantly
4. Immortal
Why doesn't the Darwinian Demon exist?
1. Energy-limited
2. Biological processes NOT instantaneous
3. Physiological and evolutionary TRADE-OFFS
4. Ecological context (competition, density-dependence).
Sexual dimorphism
Phenotypic differences
among males and females
of the same species in
secondary sexual traits.
Sexual selection:
Exaggerated male traits may arise because they
increase male mating success. This advantage should
be large enough to compensate for negative
effects on survival or fecundity.
Difference between natural selection and sexual selection
Natural
1. Variation in trait among individuals
2. Variation in trait is heritable
3. Some individuals are more successful at SURVIVING to reproduction than others
4. Reproductive success is non-random
(individuals with favorable form of trait REPRODUCE more)
Sexual:
1.Variation in trait among individuals (expression of trait may be restricted to 1 sex)
2. Variation in trait is heritable
3. Some individuals are more successful at OBTAINING MATINGS than others
4. Reproductive success is non-random (individuals with favorable form of trait MATE more)
Darwin's quote about sexual vs. natural selection
“Sexual selection depends on the success of certain individuals over others OF THE SAME SEX, in relation to the propagation of the species; while natural selection depends on the success OF BOTH SEXES, at all ages, in relation to the general conditions of life.”
What did Trivers 1972 explain?
Exaggerated traits mostly in males because...
Differences in parental investment among males and females.
When does sexual selection occur?
Member of one sex (usually male) compete with each other for access to mates.
If sexual selection occurs in males, what can be expected?
• Males limited by number of females to mate with
• Females limited by quality of mate
• Greater sexual dimorphisms
• More ornamented or brightly colored males.
What are the consequences of sexual selection?
1. Male-male competition
2. Female mate choice.
Examples for each adaptation testing method:
1. Experimental
2. Direct observation
3. Independent contrast
3. Testes size in bats
Forms of male-male competition
1. Form I
2. Form II
3. Form III
1. Combat
2. Sperm competition
3. Infanticide.
Which form of male-male competition may lead to alternative mating strategies?
Form I: Combat.
Example?
Sneaker males
What does sperm competition favor in males?
Mate guarding, large testes, copulatory plug, structures that remove other sperm.
What are the explanations (hypotheses) of female choice?
1. Good genes hypoth
2. Direct benefit
3. Sexy sons hypoth
4. Pre-existing sensory bias hypoth.
Good genes hypothesis says females asses genetic fitness in males...
If trait is costly, indicates male quality, developmental
robustness, resistance to parasites
e.g. long calls in frogs.
Direct benefit hypothesis
If males provide resources, females directly assess quality of resources
May translate into higher repro fitness for both sexes.
Sexy sons
Runaway process; preference has no genetic correlation
Females mate with sexy males because sons will also be sexy. Thus she will have higher repro fitness.
Pre-existing sensory bias
male trait exploits pre-existing sensory bias
e.g. male fiddler crabs build hoods over burrows - look like shelters to females. Males court from within.
Male choice e.g. in pipefish
Females larger and have stripes. Males are the choosy ones.
Types of social interactions
Cooperative
Selfish
Altruistic
Spiteful.
Kin selection - selection applied to the family.
Natural selection favoring the spread of alleles that increase indirect fitness.
indirect fitness
reproduction by relatives made possible by an individual's actions.
Kin selection is...
-Natural selection based on inclusive fitness
-Process by which traits are favored because of their beneficial effects on the fitness of relatives.
Kin selection predicts...
1. that idivs must dispense benefits more often to kin than to non-kin
2. certain behaviors to be more common among relatives: altruism, cooperation
3. selfish and aggressive behaviors to be less common among relatives.
What is Hamilton's rule?
Br - C > 0
B= benefit to recipient
C = cost to actor
r = coefficient of relatedness.
Selection favors altruism if...
it increases indirect fitness
Cost/benefit are measures in units of surviving offspring.
Kin selection requires...
kin recognitiion. Examples?
Spade footed toad morph.
In kin selection, the allele for altruism...
Increases by playing the odds
The more closely the actor is related to the beneficiary, the more likely the beneficiary also carries the altruism allele.
Greenbeard effect
an alternative to kin selection. Does not require discrimination among relatives
Allele causes carrier to grow a green beard, which is recognizable on others. Behave altruistically toward other green beards, and the greenbeard allele will spread.
Slime mold aggregates as greenbeard example
wild-type csA cells are suckers, make up larger portion of stalk in lab (non-reproductive), but...
In soil, wild-type cells are altruists toward each other, so altruistic allele higher in freq than knock-out cell.
Eusociality
-Indivs cooperate in caring for young
-Reproductive division of labor (sterile workers likely)
-Generations overlap.
Why is eusociality common on hymenoptera?
Haplodiploidy
Bros. and sisters only related through their mother.
Haplodiploidy
male haploid (develop from unfertilized eggs); females diploid
So sisters are 3/4 related to e/o, 1/2 to own offspring and 1/4 to brothers.
Kin selection predicts conflict of interest
Sisters more related to each other than brothers
Favors 3:1 sex ratio. Workers destroy male eggs.
Does haplodiploidy explain eusociality?
Problems:
-Multiple matings by queen could make workers more related to offspring than to sisters
-More than one queen may exist in a colony
-many eusocial species aren't haplodiploid (termites, naked mole-rats).
Eusociality evolution
Eusociality evolved in taxa that build nests and take case of young for extended periods
Paper wasps - workers not sterile
-but more successful when co-found nests.
Parent-offspring conflict
Parent related to offspring equally (r=0.5).
-Conflct btwn parent and offspring
-Indiv offspring monopolize parent
-Parent wants to maximize fitness of ALL offspring.
Types of parent-offspring conflict
-Weaning conflict
-Harassment
-Siblicide in boobies
-Mom gets more benefit by reproducing again than producing milk for current offspring
-Bee-eaters harass closely related kin (prevent from nesting) to recruit them as helpers to their parents?
-First hatchling pushes out unhatched egg; older sib kills younger in food shortage (long term)
What is cooperation among unrelated individuals called?
Reciprocal altruism
What does reciprocal altruism require?
1. Repeated interactions among same individuals
2. Many opportunities for altruism to occur
3. Indivs have good memories
4. Altruists interact in symmetrical situations.
Life history is an organism's...
life-time pattern of growth, differentiation, storage and reproduction.
List some important life history traits
Size at birth
Age at maturity
Size at maturity
Number of reproductions
Clutch size
Nat sel can act on any of these traits to improve fitness.
How can LH traits evolve to maximize fitness?
1. Higher survival through ages
2. Higher fecundity at each repro stage
3. Higher fecundity earlier in life
4. Longer repro lifespan
5. Earlier age of first repro.
Basic problem for Darwinian Demon?
Resource allocation in a finite world
-Survival, maintenance
-Growth
-Reproduction.
Why do orgs age and die?
Nat'l selection should oppose senescence and death, but it doesn't.
2 theories:
1. Rate of Living theory
2. Evolutionary theory
Rate of living theory of aging
Aging is caused by the accumulation of irreparable damage to cells and tissues (repl error and metabolic by-products).
What are the predictions in the Rate of Living theory of aging?
1. Aging rate correlated with metabolic rate
2. Species should not be able to evolve longer life spans because already selected for maximal repair
Data does not support the age/metabolic rate correlation.
-Lifetime energy expenditure does vary
-Possible to increase life-span with artificial selection.
Aging due to cumulative effects of cell division?
Cells only capable of a limited number of divisions based on telomere length
Jury still out on this hypothesis
Evolutionary theory of aging
Probability of survival and advantage of maintaining and repairing somatic tissues declines with age. Aging not due to cellular damage so much as failure to repair damage later in life.
The Evol Theory of Again states that
Genes that negatively affect repro or survival at later stages of life are only weakly selected against.
Aging is due to:
1. Deleterious mutations - mutation accumulation
2. Trade-offs between repair and reproduction.
Genes reducing survivorship and reproduction
1. Early in life...
2. Late in life...
1. early in life are strongly selected against
2. late in life are weakly selected against
e.g. DNA mismatch repair enzyme.
Antagonistic Pleiotropy hypothesis
a trade-off and aging relationship
-aging is caused by alleles with pleiotropic effects that are advantageous early in life and deleterious later in life
e.g. D. methuselah homozygotes live 35% longer BUT trade-off in that they have fewer progeny.
Phenotypic plasticity
When selection favors different phenotypes at different times, characters can adjust.