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Host population structure and the evolution of parasites. Mike Boots. Our Infectious Diseases. Theory on the evolution of parasites. Evolutionary game theory ‘Adaptive Dynamics’ Can strains invade when rare? Generally a simple haploid genetic assumption Small mutations
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Host population structure and the evolution of parasites Mike Boots
Our Infectious Diseases
Theory on the evolution of parasites • Evolutionary game theory • ‘Adaptive Dynamics’ • Can strains invade when rare? • Generally a simple haploid genetic assumption • Small mutations • Ecological feedbacks
Theory on the evolution of parasites • Infectivity is maximised • Infectious period maximised • Mortality due to infection (virulence) minimised • Recovery rate minimised • Trade-offs related to exploitation of the host explain variation
Virulence as a cost to transmission Transmission Virulence
S I Natural Mortality + Virulence Natural Mortality Transmission Reproduction Lattice Models (Spatial structure within populations) S S S S S I S I I
Simulation results for the evolution of transmission with individuals on a lattice where interactions are all local Mean Transmission TIME Max transmission = 150 No trade-offs between transmission and virulence
I S Global Infection (L) I S Local Infection (1-L) Intermediate Levels of Spatial Structure
5 4 3 2 1 0 0.0 0.2 0.4 0.6 0.8 1.0 Maximum virulence Mean Virulence Linear trade-off with virulence and transmission L (Proportion of global infection)
event conditional prob that I is a neighbour of an S site in an SI pair = # neighbours (fixed) transmission rate Host Parasite models between local and mean-field Pair-wise Approximation: differential equations for pair densities eg, PSI(t) =prob randomly chosen pair is in state SI
Host Parasite models between local and mean-field Pair-wise Approximation: differential equations for pair densities PSI(t) =prob randomly chosen pair is in state SI eg, event =
Host Parasite models between local and mean-field Pair-wise Approximation: differential equations for pair densities PSI(t) =prob randomly chosen pair is in state SI eg, LI event prob that a site is infected (1-LI) = proportion of global infection LI=0 (local), LI=1 (mean-field)
Host Parasite models between local and mean-field • Derive correlation Eqns: for each pair and singleton from states S, I, R and 0 (empty sites). with params 0<LI,Lr<1 for global proportions of reproduction for pathogen and host. • Pair closure: determine qI/SI in terms of qI/S (from Monte Carlo sims). • Analysis:Stability analysis (long term behaviours) • Bifurcation analysis, continuation (limit cycles)
Local density of infecteds Virulence Transmission Background Mortality Global density of susceptibles Invasion Condition > 0 J is a mutant strain I is the resident strain Hat notation denotes quasi steady state
Pairwise Invasion Plots (Linear trade-off between transmission and virulence)
Does the analysis agree with the simulations? • Yes: There is an ES virulence with spatial structure and maximization with global infection • Yes: The ES virulence increases as the proportion of global infection increases • But: The ESS is lost before L=1.0 • Weak selection gradients mean this is not seen when simulation is run for a set time period
Standard assumption of the evolution of virulence theory Transmission Virulence The role of trade-off shape
Simulation Approximation Evolution with a saturating trade-off in a spatial model
S S S S R I R I I The role of recovery: The Spatial Susceptible Infected Removed (SIR) Model
The role of recovery No recovery =0
The role of recovery =0.1 Increased ES virulence Wider region of bistability
The role of recovery =0.2 Bi-stability region reduces
The role of recovery =0.3
The role of recovery =0.4
The role of recovery Max ES virulence increases Recovery rate
Conclusions • Spatial structure crucial to evolutionary outcomes • Bi-stability leading to the possibility of dramatic shifts in virulence • Shapes of trade-offs are important • Approximate analysis is useful in spatial evolutionary models
Collaborators • Akira Sasaki (Kyushu University) • Masashi Kamo (Kyushu: Institute for risk management, Tsukuba) • Steve Webb