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Spatial effects in parasite-produced marine diseases

This repository contains the code used to produce the results published in Royal Society Open Science

Abstract

Marine infectious diseases are more prevalent in recent times due to climate change and other anthropogenic pressures, posing a substantial threat to marine ecosystems and the conservation of their biodiversity. An important subset of marine organisms are sessile, for which the most common mechanism for disease transmission is direct contact with waterborne parasites. Only recently, some deterministic compartmental models have been proposed to describe this kind of epidemics, being these models based on non-spatial descriptions where space is homogenised and parasite mobility is not explicitly accounted for. However, in realistic situations, epidemic transmission is conditioned by the spatial distribution of hosts and the parasites mobility patterns. Thus, the interplay between these factors is expected to have a crucial effect in the evolution of the epidemic, so calling for a explicit description of space. In this work we develop a spatially-explicit individual-based model to study disease transmission by waterborne parasites in sessile marine populations. We investigate the impact of spatial disease transmission, performing extensive numerical simulations and analytical approximations. Specifically, the effects of parasite mobility into the epidemic threshold and the temporal evolution of the epidemic are assessed. We show that larger values of pathogen mobility have two main implications: more severe epidemics, as the number of infections increases, and shorter time-scales to extinction. Moreover, an analytical expression for the basic reproduction number of the spatial model is derived as function of the non-spatial counterpart, which characterises a transition between a disease-free and a propagation phase, in which the disease propagates over a large fraction of the system. This allows to determine a phase diagram for the epidemic model as function of the parasite mobility and the basic reproduction number of the non-spatial model.

Requirements

  • Julia v 1.6 or higher with the following libraries

    • StatsBase
    • Plots (for animations, but not necessary to run the simulations)

Tutorial

DISCLAIMER: This is a tutorial to show how to use the library provided in this repository, and can be run in a standard computer. Although all figures in the paper are reproducible using the code provided in the repository (not the tutorial istelf), the computational time needed might be extremly large for some of them. For instance, the phase diagram shown in Figure 4 was obtained after 1-2 weeks of computation in a cluster using distributed computing (each pixel correspond to a simulation averaging over 1000 realisations paralelised with 10 cores. Furthermore, taking advantage of the cluster, several of these pixeles could be computed at the same time, in parallel). Consequently, trying to reproduce this figure in a single machine and without parallel computing could take many months of computation.

#Load IBM library
include("SIRP_IBM_Library.jl")

Auxiliary functions for easy use of library

function init_model(method_hosts, method_parasites, parameters; fraction=0.75)
    
    I = []
    R = []
    
    P = zeros(x, y)
    
    if method_hosts == "full"
        
        S = 1:x*y
        
    elseif method_hosts == "fraction"
        
        S = sample(1:x*y, Int32(x*y*fraction), replace=false)
        
    else
        
        println("This method is not defined")
        
    end
        
    if method_parasites == "center"
        
        P[Int32(floor(x/2)), Int32(floor(y/2))] = P0
            
    elseif method_parasites == "random"
        
        P[rand(1:x*y)] = P0
        
    else
       
        println("This method is not defined")
        
    end

    S0 = length(S)
    P = vec(P)

    grid = construct_squared_grid(x, Int32)

    params = Parameters{Float32}(parameters, zeros(5))

    vars = Variables{Int32}(S, I, R, P)
    
    println("Model initialised with R_0:", R0(params; S0=x*y/S0))
    
    return grid, params, vars
    
end

R0(params; S0=1.0) = (params.rates[2] * params.rates[4]) / (params.rates[1]*(params.rates[3] + params.rates[4])) * (params.rates[end]/(params.rates[3] + params.rates[end])) 

R0_MF(params; S0=1.0) = params.rates[2] * params.rates[4] / (params.rates[1]*(params.rates[3] + params.rates[4]))

Simulate system

Here we show an example on how to simulate the system. The user must choose the number of rows and columns of the grid (x,y), the values for the parameters of the model, the initial conditions, and the maximum time to run, t_end (it is system simulated time, not computation time). Then we run the init_model function to get the initialised variables for grid, params and vars and then we run the IBM(t_end, grid, vars, params) function.

#Choose parameters
x = 100
y = 100

γ = 1.0
λ = 15.0
μ = 1.0
β = 1.0
κ = 2.0

P0 = 50

t_end = 100

parameters = [γ, λ, μ, β, κ]

method_hosts = "full" #full or fraction. if random, the fraction of hosts can be specified (default at 0.75)
method_parasites = "center" #center or random. P0 initialised at center or random cell.

#Initialise the model
grid, params, vars = init_model(method_hosts, method_parasites, parameters; fraction=0.75)

#Simulate
S_f, I_f, R_f, P_f, time = @time IBM(t_end, grid, vars, params);

#We reduce the number of points for plotting
L = length(S_f)

delta = Int(round(L / 1000)) #So we will have 1000 points

S_f_plot = S_f[1:delta:end]
I_f_plot = I_f[1:delta:end]
R_f_plot = R_f[1:delta:end]
P_f_plot = P_f[1:delta:end]

time_plot = time[1:delta:end]

plot(time_plot, [S_f_plot I_f_plot R_f_plot P_f_plot], lw=5, color=["green" "red" "black" "blue"], 
    label=["S" "I" "R" "P"], guidefontsize=20, labelfontsize=20, legendfontsize=20, xlabel="Time", 
    ylabel="Nº of individuals", legend=:right, size=(800, 600), tickfontsize=14)

Model initialised with R_0:5.0

14.590865 seconds (34.70 M allocations: 10.822 GiB, 6.05% gc time, 13.28% compilation time)

example

Animation

"Fast" animation (only hosts are shown)

Here we simulate the system and save an animation showing only the changes in host states, but no parasites are shown

#Choose parameters
x = 30
y = 30

γ = 1.0
λ = 15.0
μ = 1.0
β = 1.0
κ = 1.0

P0 = 100

t_end = 50

parameters = [γ, λ, μ, β, κ]

method_hosts = "full" #full or fraction. if random, the fraction of hosts can be specified (default at 0.75)
method_parasites = "center" #center or random. P0 initialised at center or random cell.

#Initialise the model
grid, params, vars = init_model(method_hosts, method_parasites, parameters; fraction=0.75)

#Simulate the system and save animation
anim = @time IBM_animation_fast(t_end, grid, vars, params; every_frame=100,
    color=cgrad([:black, :red, :green]), size=(800, 600))

println("\nSaving animation...")

name = string("IBM_anim_P_", P0, "_L_", x, "_κ_", params.rates[5], "_.gif")

@time gif(anim, name, fps=100)

Model initialised with R_0:3.75

53.820113 seconds (140.98 M allocations: 10.129 GiB, 7.68% gc time, 7.52% compilation time)

Saving animation...

5.986004 seconds (1.44 M allocations: 79.455 MiB, 9.01% compilation time)

┌ Info: Saved animation to │ fn = /home/alex/Pinna Nobilis/Stochastic models/IBM_anim_P_100_L_30_κ_1.0_.gif └ @ Plots /home/alex/.julia/packages/Plots/Ye1xf/src/animation.jl:114

Animation1

Complete animation

In this animation we show changes in hosts states, parasite spread and population dynamics, but the computational time is larger.

#Choose parameters
x = 20
y = 20

γ = 1.0
λ = 15.0
μ = 1.0
β = 1.0
κ = 1.0

P0 = 100

t_end = 20

parameters = [γ, λ, μ, β, κ]

method_hosts = "full" #full or fraction. if random, the fraction of hosts can be specified (default at 0.75)
method_parasites = "center" #center or random. P0 initialised at center or random cell.

#Initialise the model
grid, params, vars = init_model(method_hosts, method_parasites, parameters; fraction=0.75)

#Simulate the system and save animation
anim = @time IBM_animation(t_end, grid, vars, params; every_frame=100)

println("\nSaving animation...")

name = string("IBM_anim_P_", P0, "_L_", x, "_κ_", params.rates[5], "_.gif")

@time gif(anim, name, fps=100)

Model initialised with R_0:3.75

254.217243 seconds (177.24 M allocations: 29.603 GiB, 61.74% gc time, 0.30% compilation time)

Saving animation...

3.517080 seconds (568 allocations: 43.641 KiB)

┌ Info: Saved animation to 
│   fn = /home/alex/Pinna Nobilis/Stochastic models/IBM_anim_P_100_L_20_κ_1.0_.gif
└ @ Plots /home/alex/.julia/packages/Plots/Ye1xf/src/animation.jl:114

Anim2

Custom host spatial setting

Also a custom spatial setting for the hosts can be achieved, but this requeres further programming knowledge.

In this example we remove the hosts located at the central rows and columns. Although the parasites can still move tho those sites, we observe that the separation is enough to stop the transmission if we are near the critical point, $R_0\sim1$.

(The transmission that occurs among the separated zones of the plot is due to periodic boundary conditions, not long range transmission)

#Choose parameters
x = 30
y = 30

γ = 1.0
λ = 15.0
μ = 1.0
β = 1.0
κ = 1.0

parameters = [γ, λ, μ, β, κ]

P0 = 200

I = []
R = []

P = zeros(x, y)
P[Int32(floor(x/4)), Int32(floor(y/4))] = P0
P = vec(P)

#Custom S0
S0 = zeros(x, y)

forbidden_idx = [15, 16]

for i in 1 : x
    
    for j in 1 : y
        
        if i  forbidden_idx && j  forbidden_idx
            
            S0[i, j] = 1.0
            
        end
        
    end
    
end

S = findall(vec(S0) .> 0)

grid = construct_squared_grid(x, Int32)

params = Parameters{Float32}(parameters, zeros(5))

vars = Variables{Int32}(S, I, R, P);

t_end = 30

anim = @time IBM_animation_fast(t_end, grid, vars, params; every_frame=100, 
    color=cgrad([:black, :blue, :red, :green]), size=(800,600))

println("\nSaving animation...")

name = string("IBM_anim_P_", P0, "_L_", x, "_κ_", params.rates[5], "_obstacle.gif")

@time gif(anim, name, fps = 100)

44.496868 seconds (56.60 M allocations: 4.309 GiB, 52.90% gc time)

Saving animation...

2.660894 seconds (536 allocations: 42.531 KiB)

┌ Info: Saved animation to │ fn = /home/alex/Pinna Nobilis/Stochastic models/IBM_anim_P_200_L_30_κ_1.0_obstacle.gif └ @ Plots /home/alex/.julia/packages/Plots/Ye1xf/src/animation.jl:114

Anim3

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