7, 8, 9, 10, 11, 12 Flashcards
3 types of selection
positive (directional) selection
negative (purifying) selection
selection to maintain variation (balancing selection)
describe the distribution of polygenic traits
- continuous distribution
- typical of size/number traits
- example of quantitative inheritance
draw and describe 3 different modes of selection on quantitative traits
stabilising selection favours average traits
directional selection favours one extreme
disruptive selection favours both extremes
what may be the effects of disruptive selection?
- trait divergence
- speciation, if trait divergence causes a reduction in gene flow
how can we study adaptation?
- test for correlation of alleles or traits with environment over space and time and analyse genomic diversity (genes targeted by selection ought to show distinctive patterns)
- experimental manipulations in field/lab
describe the struggle to determine the agents of selection
- research through today shows thousands of measurements of selection
- demonstrates fitness differences & evolutionary change in traits
and yet: - many fewer convincing cases document the mechanisms (agents) of selection in natural populations
- linking evolution to ecology is difficult
describe the peppered moth (biston betularia) and industrial melanism
- light and dark forms of species that rest on trees
- in the UK, before 1850, dark moths were rare
- dark form caused by a single dominant allele of Cortex gene
- industrial pollution blackened tree trunks near cities and resulted in an increase in the dark form
- dark variant replaced light form in polluted areas, light form predominated in rural unpolluted areas
- mechanism of selection due to predation by birds
- differences in moth crypsis (camouflage) depend on trunk coloration
what happened to the darker melanic variant of the peppered moth after the introduction of the UK ‘clean air act’ in 1956?
there was a decline,
what does the lag in evolutionary response to changes in air pollution levels reflect for peppered moths?
the time required for forests to return to a more natural (unpolluted) state as well as a low initial frequency of the recessive allele for a typical coloration
describe the evolution of heavy metal tolerance in plants
- mine waste is heavily polluted with heavy chemicals such as lead, copper, nickel, cadmium
- heavy metal tolerant genotypes can occur at very low frequency in nearby uncontaminated pastures
- tolerant genotypes invade mine tailings from nearby pastures
- gene flow between pasture and mine is restricted because of flowering time differences between them
- alleles enabling tolerance maintained on mines but not pastures
define a selective sweep
when selection causes a new mutation to increase in frequency so quickly that nearby alleles ‘hitchhike’ and also increase in frequency
describe DNA Variation at glucose-6-phosphate dehydrogenase deficiency in humans
recent natural selection has caused
- low diversity
- high frequency of derived (new) allele
- G6PD gene shows evidence of the recent, rapid spread of resistance allele (consistent with hypothesis of selection for malaria resistance)
describe the long term experimental evolution study of adaptation by E.Coli
- 36 years if propagating 12 population flasks
- > 75000 generations of evolution
- propagated in minimal glucose/citrate medium
results of long term experimental evolution study of adaptation by E.Coli
- all populations rapidly increased in fitness
- similar adaptations across populations (eg larger cell sizes, higher max growth rates on glucose)
- parallel mutations in same genes
- some unique adaptations and distinct genetic changes
- only one strain evolved ability to grow on citrate
define a population
a group of individuals of a single species occupying a given area at the same time
define migration
the movement of individuals from one population to another
define gene flow
the movement of alleles from one population to another
how can we measure gene flow?
use experimental approaches
use neutral genetic markers:
- polymorphic genetic variants that aren’t direct targets of selection
experiment to answer the question: how much gene flow occurs between geographically separated populations?
- establish two populations, fixed for alternative alleles, separated by a given distance
- score FS heterozygotes in offspring
- frequency of heterozygotes = an estimate of gene flow
formally define genetic drift
stochastic changes in allele frequency due to random variation in fecundity and mortality
what does random mean in evolution?
stochastic (unpredictable or random) evolutionary forces:
- mutation
- recombination
- genetic drift
deterministic (predictable or non-random) evolutionary forces:
- natural selection
define population bottlenecks
- a single sharp reduction in abundance, usually followed by a rebound
- causes a loss of diversity
define a founder event
- colonisation by a few individuals that start a new population
- colonising group contains only limited diversity compared to the source population
why is genetic drift more pronounced in small populations?
- more drastic fluctuations in each generation
- more rapid loss of genetic diversity (i.e. faster time to allele fixation or loss)
- less consistency across replicate populations
as distance increases, gene flow
decreases
define isolation by distance
accumulation of local genetic variation due to geographically limited dispersal
phenotypic variation may be:
- adaptive (‘local adaptation’
- due to genetic drift
- phenotypic plasticity
how do we test for local adaptation and plasticity?
reciprocal transplant studies
Genomic analyses
define phenotypic plasticity
the ability of a genotype to modify its phenotype in response to a particular environment
draw 3 graphs for no plasticity, plasticity, and highly variable plasticity
how does phenotypic plasticity occur?
through modifications to development, growth, and/or behaviours under genetic control
in what organisms is phenotypic plasticity common?
in sedentary organisms like plants and corals, but also in animal behaviour
phenotypic plasticity often is an adaptation to…
unpredictable environments
does all phenotypic plasticity result from adaptation?
no
describe reciprocal transplant studies
Growth of equivalent genotypes in contrasting environments and comparisons of their relative
performance
why are reciprocal transplant studies useful?
- Can separate phenotypic variation into genetic and environmental components
- Enables measurement of selection against non‐local genotypes
- Can provide evidence for/against local adaptation
Clausen‐Keck‐Hiesey Transplant Conclusions
- Differences between populations due to BOTH plasticity and genetics
- Evidence for widespread local adaptation
– Local populations had highest fitness
How do we test for plasticity and adaptation in species that we can’t manipulate experimentally?
genomic studies
Tradeoffs Associated With Skin Pigmentation
High UV radiation:
– Degrades folate, critical in highly dividing tissues (e.g. embryos, testes)
– May have selected for increased pigmentation
- Strong purifying selection on MC1R in equatorial
regions
Low UV radiation:
– Reduced vitamin D synthesis
- VitD critical for bone development, immunity, etc.
– May have selected for reduced pigmentation
was there a history of local adaptation in skin pigmentation?
Numerous genes known to affect skin pigmentation
* These genes show higher between‐population differentiation than most others
–> Evidence supporting a history of local adaptation
* Pigmentation genes show evidence for positive selection in regions with distinctive skin
colouration
what was the confusion about the link between micro and macroevolution?
can processes of microevolution lead to macroevolution?
taxonomic (morphological) species concept
based primarily on distinct measurable differences
biological species concept
based on inter-fertility among individuals
why is it so hard to define a species?
concepts vary among groups of organisms and among scientists. There is no universal species concept.
- geographic isolation alone is NOT sufficient
- isolation does NOT have to be absolute (what cutoff?)
Darwin’s definition of a species
groups of organisms that are sufficiently similar in phenotype
Ernst Meyer’s view on distinguishing species
reproductive isolation as key to distinguishing species
what species does the BSC not apply to?
does not apply well for bacteria, asexuals, highly self-fertilising species…or fossils
allopatric speciation
often called geographic speciation
- due to involvement of geographical isolation
- much more common and easier to evolve
stages where reproductive isolation can occur
pre-zygotic:
- finding a compatible mate and mating
- fertilisation
post-zygotic:
- development and growth of zygote (F1)
- adult survival & reproduction
- growth, survival, reproduction of offspring (F2)
pre-zygotic barriers
prevent mating or fertilisation so no zygote is formed:
- geographical, ecological
- temporal, behavioural (mate recognition)
- mechanical (genital stricture compatibility)
- cellular (sperm-egg compatibility)
use Rhagoletis pomonella (Apple Maggot Flies) as an example of pre-zygotic isolation
- host shift after arrival of domesticated applies in 1800s
- differences in timing of host planting fruiting (apple vs hawthorn)
- different timing of fly mating on preferred host plant)
- reduces fly gene flow by 94% in sympatry (same region)
describe pre-zygotic isolation in abalone
binding of sperm lysin protein to egg vitelline envelope receptor (VERL) required for fertilization (molecular lock and key)
Lysin/VERL interaction has coevolved
– Different evolutionary changes in different species
– Causes reproductive isolation due to fertilization
incompatibility
post-zygotic barriers
prevent proper functioning of zygotes
once they are formed
* Caused by combinations of genes with low fitness in the
hybrid
* Arise as an indirect byproduct of evolution acting
separately in different populations (cannot be directly
favored by natural selection)
intrinsic post-zygotic barriers vs extrinsic post-zygotic barriers
Intrinsic Post‐zygotic Barriers:
* Inviability, sterility, or abnormal development of hybrids
Extrinsic Post‐zygotic Barriers:
* Ecological mismatch of hybrid phenotype to environment
example of intrinsic post-zygotic isolation
Mule is a sterile hybrid cross of:
* Male donkey (62 chromosomes)
* Female horse (64 chromosomes)
Hinny is a sterile hybrid of:
* Male horse (64 chromosomes)
* Female donkey (62 chromosomes)
relation between genetic distance and post-zygotic isolation in fruit flies
- The more that fly pairs are genetically differentiated,
the more likely they are to be reproductively isolated
example of extrinsic post-zygotic isolation
aposematic helicons butterflies
Hybrids have aberrant colour patterns
* Higher predation risk
* Lower mating success
local adaptation by different populations can lead to
reproductive isolation and speciation
distinct evolutionary responses happen due to
different selective pressures