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Lecture 9: Population genetics, first-passage problems. Outline: population genetics Moran model fluctuations ~ 1/ N but not ignorable effect of mutations effect of selection neurons: integrate-and-fire models interspike interval distribution no leak with leaky cell membrane
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Lecture 9: Population genetics, first-passage problems • Outline: • population genetics • Moran model • fluctuations ~ 1/N but not ignorable • effect of mutations • effect of selection • neurons: integrate-and-fire models • interspike interval distribution • no leak • with leaky cell membrane • evolution • traffic
Population genetics: Moran model 2 alleles, N haploid organisms
Population genetics: Moran model 2 alleles, N haploid organisms choose 2 individual at random: 1 dies, the other reproduces
Population genetics: Moran model 2 alleles, N haploid organisms choose 2 individual at random: 1 dies, the other reproduces If there are n1 organisms of type 1 before this step, then afterwards there are
Population genetics: Moran model 2 alleles, N haploid organisms choose 2 individual at random: 1 dies, the other reproduces If there are n1 organisms of type 1 before this step, then afterwards there are n1 + 1 with probability x(1 – x) x = n1/N
Population genetics: Moran model 2 alleles, N haploid organisms choose 2 individual at random: 1 dies, the other reproduces If there are n1 organisms of type 1 before this step, then afterwards there are n1 + 1 with probability x(1 – x) x = n1/N n1 – 1 with probability x(1 – x)
Population genetics: Moran model 2 alleles, N haploid organisms choose 2 individual at random: 1 dies, the other reproduces If there are n1 organisms of type 1 before this step, then afterwards there are n1 + 1 with probability x(1 – x) x = n1/N n1 – 1 with probability x(1 – x) n1 with probability x2 + (1 – x)2.
Population genetics: Moran model 2 alleles, N haploid organisms choose 2 individual at random: 1 dies, the other reproduces If there are n1 organisms of type 1 before this step, then afterwards there are n1 + 1 with probability x(1 – x) x = n1/N n1 – 1 with probability x(1 – x) n1 with probability x2 + (1 – x)2. So
Population genetics: Moran model 2 alleles, N haploid organisms choose 2 individual at random: 1 dies, the other reproduces If there are n1 organisms of type 1 before this step, then afterwards there are n1 + 1 with probability x(1 – x) x = n1/N n1 – 1 with probability x(1 – x) n1 with probability x2 + (1 – x)2. So
Population genetics: Moran model 2 alleles, N haploid organisms choose 2 individual at random: 1 dies, the other reproduces If there are n1 organisms of type 1 before this step, then afterwards there are n1 + 1 with probability x(1 – x) x = n1/N n1 – 1 with probability x(1 – x) n1 with probability x2 + (1 – x)2. So or
Population genetics: Moran model 2 alleles, N haploid organisms choose 2 individual at random: 1 dies, the other reproduces If there are n1 organisms of type 1 before this step, then afterwards there are n1 + 1 with probability x(1 – x) x = n1/N n1 – 1 with probability x(1 – x) n1 with probability x2 + (1 – x)2. So or
continuum limit: FP equation (N steps/generation)
continuum limit: FP equation (N steps/generation)
continuum limit: FP equation (N steps/generation) boundary conditions: P(0,t) = P(1,t) = 0
continuum limit: FP equation (N steps/generation) boundary conditions: P(0,t) = P(1,t) = 0 (once an allele dies out, it can not come back)
continuum limit: FP equation (N steps/generation) boundary conditions: P(0,t) = P(1,t) = 0 (once an allele dies out, it can not come back) stochastic differential equation:
continuum limit: FP equation (N steps/generation) boundary conditions: P(0,t) = P(1,t) = 0 (once an allele dies out, it can not come back) stochastic differential equation: notice
heterozygocity Eventually P(x,t) gets concentrated at one boundary,
heterozygocity Eventually P(x,t) gets concentrated at one boundary, i.e., one allele completely dominates the other.
heterozygocity Eventually P(x,t) gets concentrated at one boundary, i.e., one allele completely dominates the other. But it is random which one.
heterozygocity Eventually P(x,t) gets concentrated at one boundary, i.e., one allele completely dominates the other. But it is random which one. Measure this by the heterozygocity
heterozygocity Eventually P(x,t) gets concentrated at one boundary, i.e., one allele completely dominates the other. But it is random which one. Measure this by the heterozygocity use Ito’s lemma:
heterozygocity Eventually P(x,t) gets concentrated at one boundary, i.e., one allele completely dominates the other. But it is random which one. Measure this by the heterozygocity use Ito’s lemma:
heterozygocity Eventually P(x,t) gets concentrated at one boundary, i.e., one allele completely dominates the other. But it is random which one. Measure this by the heterozygocity use Ito’s lemma:
heterozygocity Eventually P(x,t) gets concentrated at one boundary, i.e., one allele completely dominates the other. But it is random which one. Measure this by the heterozygocity use Ito’s lemma:
heterozygocity Eventually P(x,t) gets concentrated at one boundary, i.e., one allele completely dominates the other. But it is random which one. Measure this by the heterozygocity use Ito’s lemma: i.e., diversity dies out in about N generations
fluctuations of x So mean-square fluctuations of x grow initially linearly in t and then saturate
with mutation: Mutation induces a drift term in the FP and sd equation
with mutation: Mutation induces a drift term in the FP and sd equation
with mutation: Mutation induces a drift term in the FP and sd equation
with mutation: Mutation induces a drift term in the FP and sd equation stationary solution:
with mutation: Mutation induces a drift term in the FP and sd equation stationary solution:
fluctuations Use Ito’s lemma on F(x) = x2:
fluctuations Use Ito’s lemma on F(x) = x2:
fluctuations Use Ito’s lemma on F(x) = x2: at steady state:
fluctuations Use Ito’s lemma on F(x) = x2: at steady state:
fluctuations Use Ito’s lemma on F(x) = x2: at steady state:
fluctuations Use Ito’s lemma on F(x) = x2: at steady state: mean square fluctuations:
heterozygocity: small noise (large population):
heterozygocity: small noise (large population):
heterozygocity: small noise (large population): large noise (small population):
heterozygocity: small noise (large population): large noise (small population):
heterozygocity: small noise (large population): large noise (small population): usually one allele dominates, rare transitions