under a skewed offspring distribution among individuals in a - PowerPoint PPT Presentation

Expected time to coalescence and F ST under a skewed offspring distribution among individuals in a population Bjarki Eldon (with John Wakeley) Mathematics and Informatics in Evolution and Phylogeny June 10-12, 2008

High variance in offspring distribution • broadcast spawning and external fertilization • type III survivorship curves Surviv orship urv es • very large population sizes I • low genetic variation I I • large number of singleton genetic variants I I I ❅ P er en t Life Span ❅ ❅ ors ❅ surviv ❅ ❅ ❅ ❅ # ❅ ❅ 0 100

Standard reproduction models have finite variance in offspring number Moran model of overlapping generations: a single randomly chosen individual produces one offspring Coalescent timescale: N 2 generations 2

A modified Moran model A special case of the models of Pitman (1999) and Sagitov (1999) The number U of offspring is a random variable The probability G n,x of an x -merger (2 ≤ x ≤ n ): � u �� N − u � N x n − x � G n,x = P U ( u ) � N � u =2 n  1 − φN − γ / 2 if u = 2    P U ( u ) =   φN − γ / 2 if u = ψN , 0 < ψ < 1 

Population subdivision with migration Conservative migration between finite number D of subpopulations gives convergence to the structured coalescent if Nm is finite as N → ∞ � N γ , N 2 � N γ ≡ min , γ > 0 N γ is the coalescence timescale; m γ N γ < ∞ as N → ∞ m γ is rescaled migration; λ γ = I γ ≥ 2 + φψ 2 I γ ≤ 2 , φ > 0, 0 < ψ < 1 λ γ is the rate of coalescence of two lines

Time to coalescence for two lines sampled from same ( T 0 ) or different ( T 1 ) subpopulations + D − 1 E ( T 0 ) = D < D = E ( T 1 ) λ γ λ γ N γ m γ λ γ = I γ ≥ 2 + φψ 2 I γ ≤ 2 , N γ ≡ min ( N γ , N 2 )

Indicators of population subdivision - F ST and N ST F ST defined in terms of probabilities of identity N ST defined in terms of average numbers of pairwise differences 1 1 F ST = , N ST = 1 + N γ m γ D D 2 1 + N γ m γ ( D − 1) 2 + θ/ 2 D λ γ D − 1 λ γ λ γ D − 1 In a Wright-Fisher population: 1 1 F ST = , N ST = D D 2 D 1 + 4 Nm 1 + 4 Nm ( D − 1) 2 + θ D − 1 D − 1 λ γ = I γ ≥ 2 + φψ 2 I γ ≤ 2 , N γ ≡ min � N γ , N 2 �

SUMMARY (i) multiple mergers coalescent processes may better apply to some marine organisms (ii) coalescent times are shorter than in the standard coalescent (iii) patterns indicating population subdivision can be observed in DNA sequence data even if the usual migration rate Nm is very, very large