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what does absolute time estimation allow for
to connect biological and geological history
phylogenies with branch lengths proportion to time provide
more information about evolutionary history than unrooted trees with branch lengths in units of subsitution/sites
problems with fossil record
biased towards organisms that fossilize well
geological processes destroy them
gaps and missing data
molecular clock
mutations accumulate in proteins in a linear relationship to species divergence time
zuckerkandl and pauling (molec clock)
rate of evolution is approximately constant over time in all lineages
changes within a protein are clock like and sequence divergence measures elapsed time
original molec clock hypothesis
for one protein evolutionary rate is constant across lineages
does not claim different proteins change at the same rate
does not claim evolution is gradual at the phenotypic level
molecular clock is predicted by
neutral theory
in small populations there are — number of new mutants but — probability of fixation
lower higherin
in large populations there are — number of new mutants but — probability of fixation
higher , lower
neutral theory
evolutionary rate is independet of population size and there is a constant neutral rate of evolution
relative rate test evaluates
whether molecular clock holds for a given taxa and sequence
is there a global molecular clock
no, rates differ among mammalian orders and within them, humans are slowest primates. evertyhing has to accomodate rate variation
why is the clock not constant (selectionist explanation)
rate reflects the strength and direction of selection which varies with the enviornment and life hsitory
why is the clock not constant (neutralist explanation)
rate reflects the mutation rate which varies for mechanical reasons
generation time, efficinecy of dna repair, and metabolic rate
strict global clock model
one rate parameter for the whole tree
substution ratei s constant over time
(not true known from relative rate test)
relaxed clocks
every branch has its own rate drawn from distirbution
uncorrelated relaxed clock
each barnch draws indepdendelty, usually from log normalau
autocorrelated relaxed clock
substitution rate evovle gradually over time. closely related lienages have similar rates.
autocorrelated relaxed clock says rate is —- uncorrelated says its —-
heritated, not heritable
where are clocks the hardest
extremes
over short intervals mutations are counted not substitutions
over long intervals you undercount because of saturation
clock calibration
a fossil fixes the age of one node it sites on
every other node is dated by propagating outwards
propagation needs an assumption about how the rate behaves between calibrated node and everywhere else