the evolution of sex: costs and benefits• although sex may have originated from the action of a...
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The Evolution of Sex: Costs and Benefits
Lukas Schärer
Evolutionary Biology
Zoological Institute
University of Basel
11.10.2011 Evolution of Sex and its Consequences HS 11
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• costs of sexual reproduction
• benefits of sexual reproduction• reminder about levels of selection
• sex benefits populations• group selection
• evolvability
• Muller’s ratchet
• mutational deterministic hypothesis
• sex benefits individuals• the red queen
• sex benefits selfish genetic elements
Summary
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The problem
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• twofold cost of sex
Costs of sexual reproduction
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• sex is costly• recombination (disrupts adapted gene
combinations)
• cellular-mechanical costs (time costs of meiosis, syngamy, and karyogamy)
• fertilisation (wasted gametes and predation risks)
• genome dilution (cost of males, twofold cost of sex)
• sexual selection (competition and specialisation)
• parthenogenesis is very effective• if parthenogenetic offspring are equally fit as sexual
offspring the parthenogens replace the sexuals within a few generations
• but that does not appear to happen, sex is widespread throughout animals and plants
• so what are the benefits of sexual reproduction?
fate of a sexual population with 106 individuals into
which one parthenogenetic female is introduced
Costs of sexual reproduction
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grou
pin
divi
dual
gene
from Maynard Smith & Szathmáry 1999
Benefits of sexual reproduction
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• selection can (at least in theory) act on many different levels• e.g., gene, cell, individual, kin group, population, species
• although sex may have originated from the action of a selfish gene, this is highly unlikely to explain sex in extant organisms
• generally selection appears to be primarily on the individual level• why should sex be an exception?
• group selection is only efficient if migration rates are low and if there is frequent extinction of local populations
• migration weakens the difference between groups due to gene flow
• extinction allows long-lived groups to spread
Reminder about levels of selection
8
grou
p
from Maynard Smith & Szathmáry 1999
Benefits of sexual reproduction
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• competition between sexuals and parthenogens is like competition between species (there generally is no gene flow)
• to maintain sexuals imagine a metapopulation structure where• local parthenogens out-compete local sexuals
• local parthenogenetic populations have a higher risk of extinction
• empty patches are generally colonised by sexuals
• but parthenogens may actually be good colonisers
• for this scenario to work we need long-term reasons why sexual populations die out less often than parthenogenetic populations
Sex benefits populations: group selection
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• if environment changes rapidly• e.g., due to coevolutionary arms-races with competitors, predators or parasites
Sex benefits populations: evolvability
from Maynard Smith 1998
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• stochastic loss of the least-mutated (optimal) class• more severe in small populations
• genotype is lost forever unless the back-mutation occurs
Sex benefits populations: Muller’s ratchet
from Maynard Smith 1998
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• can mutations also have an effect in very large populations?
• yes, but only if mutations have worse effects in combination than would be expected based on the sum of their individual effects
• i.e. if they act synergistically
Sex benefits populations: mutational deterministic hypothesis
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• with recombination• individuals on average have 3 mutations
• which are randomly distributed
• truncation selection at ≥5 mutations• kills 15% of the individuals• leads to an average of 2.45 mutations
• individuals acquire new mutations• probability of 0.55 (new distribution)
• recombination redistributes mutations• to the original distribution
• without recombination• truncation selection at ≥5 mutations
• kills 55% of the individuals
• all remaining have 4 mutations
• probability of 0.55 to get a new mutation• leads to the original distribution
Sex benefits populations: mutational deterministic hypothesis
fromMaynard Smith 1998
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• it appears possible that group level selection has been important for the maintenance of sex
• and the phylogenetic distribution of parthenogenesis largely supports this (a famous exception are the bdelloid rotifers)
• however, it requires that the origin of parthenogenesis is a rare event
• and species that are facultative or cyclical parthenogens can not be explained in this way
Sex benefits populations
light and scanning electron micrographs of bdelloid rotifers and their jaws
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grou
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from Maynard Smith & Szathmáry 1999
indi
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Benefits of sexual reproduction
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• sex may be favoured if the environment is highly unpredictable
• asexual reproduction is like buying 100 identical lottery tickets, whereas with sexual reproduction you buy 50 different tickets
• can explain the benefits of sex in rather exceptional circumstances
Sex benefits individuals: the lottery model
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• sex may help to repair DNA damage• DNA damage is not the same as mutation
• mutation can often not be detected by the cell
• repair of single-strand damage often easy (depends on the damage)
• repair of double-strand damage requires diploidy, but not sex
Sex benefits individuals: DNA repair
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• sex may be favoured if the environment changes rapidly• could it change so rapidly as to require different genotypes every generation?
• creation of rare adapted genotypes against coevolving parasites• negative frequency-depedent selection
Sex benefits individuals: the red queen
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• may require mutual coevolution
Sex benefits individuals: the red queen
Potamopyrgus antipodarum
following Lively & Jokela 1995
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grou
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ne
from Maynard Smith & Szathmáry 1999
indi
vidu
al
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Benefits of sexual reproduction
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• a transposon in an asexually reproducing ‘host’ can not transmit to new ‘hosts’
• if it could induce its host to fuse with another host it could infect new genotypes
• as in some plasmids in bacteria
• in this case sex may spread even if it, in the short term, is disadvantageous to the host (Hickey and Rose 1988)
• this mechanism is more likely to explain the origin of sex rather than its maintenance
Sex benefits selfish genes
22
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pge
ne
from Maynard Smith & Szathmáry 1999
indi
vidu
al
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?
!
✓
✓
Benefits of sexual reproduction
![Page 12: The Evolution of Sex: Costs and Benefits• although sex may have originated from the action of a selfish gene, this is highly unlikely to explain sex in extant organisms • generally](https://reader033.vdocuments.us/reader033/viewer/2022041613/5e396fdcaaea4c5a013c1278/html5/thumbnails/12.jpg)
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• costs of sexual reproduction
• benefits of sexual reproduction• reminder about levels of selection
• sex benefits populations• group selection
• evolvability
• Muller’s ratchet
• mutational deterministic hypothesis
• sex benefits individuals• the red queen
• sex benefits selfish genetic elements
Summary
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Reading