% Rubinstein model of Ranvier Node's action potentials as in
% Rubinstein JT (1995) Biophys J 68: 779-785.
% Mino H, Rubinstein JT, White JA (2002) Ann Biomed Eng 30: 578-587.
% Bruce IC (2007) Ann Biomed Eng 35: 315-318;
% Voltage is shifted so that resting voltage = 0 mV (the reversal of the leak)
% This script simulates 1000 sweeps per stimulus and calculates
% firing efficiency, mean firing time and firing time variance
% 100 (nsim) simultaneous sweeps, 10 times
% Lines 44, 60, 61, 112 and 113 are intended to make a voltage trace
% It is advisable to reduce nsim before uncommenting them
%
% Coupled activation particles, 8-state Na channel
% Diffusion Approximation with steady state values of variables in the random term
nsim=100;
gNa=0.02569*265; %mS/cm2
ENa=144; %mV
Cm=0.0000714*265; Rm=1953.49/265;
NNa=1000;
Tstop=1; dt=0.001; %ms
points = round(Tstop/dt);
Idel=0; Idur=0.1; %ms, ms
threshold=80;
NaNs=0;
currents=5:0.1:6.5;
Eff=[];
meanFT=[];
varFT=[];
NaNs=[];
tic()
for curr=1:length(currents)
Iamp=ones(1,nsim)*currents(curr);
firetimes=[];
NNaNs=0;
for nn=1:10
p=1;
% Uncomment this only with a small nsim
% vrec = zeros(points,nsim);
v = 0*ones(1,nsim);
am=1.872*(v-25.41)./(1-exp((25.41-v)/6.06));
bm=3.973*(21-v)./(1-exp((v-21)/9.41));
ah=-0.549*(27.74+v)./(1-exp((v+27.74)/9.06));
bh=(22.57)./(1+exp((56-v)/12.5));
M=am./bm;
H=ah./bh;
statesum=(1+H).*(1+M).^3;
s=[ones(1,nsim);3*M;3*M.^2;M.^3;H;3*M.*H;3*M.^(2).*H;M.^(3).*H]./(ones(8,1)*statesum);
firetime=zeros(1,nsim);
firing=zeros(1,nsim);
for t = dt:dt:Tstop
%vrec(p,:) = v; %uncomment with small nsim
%p=p+1;
if any(~firing&v>=threshold)
ind=find(v>=threshold&~firing);
firetime(ind)=t;
firing(ind)=1;
end
am=1.872*(v-25.41)./(1-exp((25.41-v)/6.06));
bm=3.973*(21-v)./(1-exp((v-21)/9.41));
ah=-0.549*(27.74+v)./(1-exp((v+27.74)/9.06));
bh=(22.57)./(1+exp((56-v)/12.5));
Iapp=Iamp*(t>Idel&t<(Idel+Idur));
Imemb=-Iapp+gNa.*s(8,:).*(v-ENa)+v/Rm;
% deterministic transition matrix
transition=[3*am.*s(1,:)-(2*am+bm+ah).*s(2,:)+2*bm.*s(3,:)+bh.*s(6,:);
2*am.*s(2,:)-(2*bm+am+ah).*s(3,:)+3*bm.*s(4,:)+bh.*s(7,:);
am.*s(3,:)-(3*bm+ah).*s(4,:)+bh.*s(8,:);
-(3*am+bh).*s(5,:)+bm.*s(6,:)+ah.*s(1,:);
3*am.*s(5,:)-(2*am+bm+bh).*s(6,:)+2*bm.*s(7,:)+ah.*s(2,:);
2*am.*s(6,:)-(2*bm+am+bh).*s(7,:)+3*bm.*s(8,:)+ah.*s(3,:);
am.*s(7,:)-(3*bm+bh).*s(8,:)+ah.*s(4,:)];
%The stochastic terms (R) are only function of state variables
R=randn(10,nsim).*sqrt((dt/NNa)*[6*am.*bh.*bm.^3;
12*am.^(2).*bh.*bm.^2;
6*am.^(3).*bh.*bm;
2*ah.*bh.*bm.^3;
6*ah.*bh.*am.*bm.^2;
6*ah.*bh.*am.^(2).*bm;
2*ah.*bh.*am.^3;
6*am.*ah.*bm.^3;
12*am.^(2).*ah.*bm.^2;
6*am.^(3).*ah.*bm]./(ones(10,1)*((ah+bh).*(am+bm).^3)));
%Stochastic terms that correspond to state variables 2-8
Wt=[-R(1,:)+R(2,:)+R(5,:);
-R(2,:)+R(3,:)+R(6,:);
-R(3,:)+R(7,:);
R(8,:)-R(4,:);
-R(8,:)+R(9,:)-R(5,:);
-R(9,:)+R(10,:)-R(6,:);
-R(10,:)-R(7,:)];
s(2:8,:)=s(2:8,:)+dt*transition+Wt;
s(1,:)=ones(1,nsim)-sum(s(2:8,:),1);
v=v-dt*Imemb/Cm;
end
fprintf('round %d Iamp %g\n',nn,currents(curr))
% clf %uncomment with small nsim
% plot(dt:dt:Tstop,vrec)
firetimes=[firetimes firetime];
NNaNs=NNaNs + sum(1*(isinf(v)|isnan(v)));
end
Eff=[Eff;length(find(firetimes~=0))];
meanFT=[meanFT;mean(firetimes(firetimes~=0))];
if length(find(firetimes~=0))>1
varFT=[varFT;var(firetimes(firetimes~=0))];
else
varFT=[varFT;0];
end
NaNs=[NaNs;NNaNs];
end
realt=toc();
fprintf('time = %g\n',realt)
csvwrite(sprintf('EffN-Rb8DAss-N%g-dt%g-%gs.csv',NNa,dt,realt),[currents' Eff meanFT varFT NaNs])
figure(1);
clf
subplot(3,1,1)
plot(currents,Eff)
subplot(3,1,2)
plot(currents,meanFT)
subplot(3,1,3)
plot(currents,varFT)