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Transcript
Evolution of mutation rate 
evolution of sex
The nature and dynamics of
beneficial variation
Why does
mutation rate
evolve?
Mutation supply rate
Mutator tradeoff?
Why sex?
• In with the good?
• Out with the bad?
Why not sex?
The twofold cost of sex
In with the good
• Variation among siblings = “tangled bank”
• Creates novel/adaptive genotypes more
rapidly (Fisher-Muller)
• Red Queen hypothesis
Recombination
No recombination
Related advantages of
recombination
• Sex generates novel combinations of alleles, some of which may
have greater fitness.
• The fate of a beneficial mutation with sex and recombination
depends less on the genetic background on which it arises (less
interference among loci).
• Also known as the The Vicar of Bray Hypothesis after:
an English cleric noted for an ability to change his religion whenever
a new monarch ascended the throne [to emphasize] that there may
be great advantages of easily and gracefully adapting to changed
circumstances. Bell 1986.
• Probability of fixation of beneficial mutations is higher in sexuals with
frequent recombination than asexuals with no recombination. The
effect substantially favors sexual individuals ONLY if beneficial
mutations occur frequently throughout the genome.
The most curious part of the thing was, that the trees and the other things round them never changed
their places at all: however fast they went, they never seemed to pass anything. "I wonder if all the
things move along with us?" thought poor puzzled Alice. And the Queen seemed to guess her thoughts,
for she cried, "Faster! Don't try to talk!“
Not that Alice had any idea of doing THAT. She felt as if she would never be able to talk again, she was
getting so much out of breath, and still the Queen cried "Faster! Faster!" and dragged her along.
"Are we nearly there?" Alice managed to pant out at last.
"Nearly there!" the Queen repeated. "Why, we passed it ten minutes ago! Faster!" And they ran on for a
time in silence, with the wind whistling in Alice's ears, and almost blowing her hair off her head . . . until
suddenly, just as Alice was getting quite exhausted, they stopped, and she found herself sitting on the
ground, breathless and giddy.
The Queen propped her up against a tree, and said kindly, "You may rest a little now."
Alice looked round her in great surprise. "Why, I do believe we've been under this tree the whole time!
Everything's just as it was!“
"Of course it is," said the Queen, "what would you have it?"
"Well, in OUR country," said Alice, still panting a little, "you'd generally get to somewhere else -- if you
ran very fast for a long time, as we've been doing."
"A slow sort of country!' said the Queen. "Now, HERE, you see, it takes all the running YOU can do, to
keep in the same place. If you want to get somewhere else, you must run at least twice as fast as that!"
Lewis Carroll
Alice Through the Looking Glass
1865
Out with the bad
• Allows reconstruction of mutation-free
individuals
• Removes deleterious mutations
• See Wiki: Kondrashov’s hatchet
Epistasis and Kondrashov’s hatchet
Out with
the bad:
Muller’s
ratchet
Recombination
No recombination
Prokaryotic recombination 
eukaryotic sex
But some of the key variables can
be manipulated in prokaryotes
In with the good
Fisher-Muller: increase rate of
adaptation?
Greater adaptation in
rec+ lines
rec+ - rec-
spoT mutations
fix more quickly
in rec+ lines
spoT mutations
become
deleterious in reclines owing to
interference
among beneficial
mutations in other
competing clones
Fitness is affected
by the presence of
other competing
phage.
Open symbols: low
MOI
Closed symbols:
high MOI