% Simplified model of the repressilator

% This function implements the basic model of the repressilator
% All parameter values were taken from 
% Nature. 2000 Jan 20;403(6767):335-8.

% This model was developed by members of the 2003 Synthetic Biology Class
% on Engineered Blinkers

function dydt = repressilator(t, y)

%store the state variables under more meaningful names
mRNA_cI = y(1);
mRNA_lacI = y(2);
mRNA_tetR = y(3);
protein_cI = y(4);
protein_lacI = y(5);
protein_tetR = y(6);

% set the max transcription rate in transcripts per second
k_transcription_cI = 0.5;
k_transcription_lacI = 0.5;
k_transcription_tetR = 0.5;

% set the leakage transcription rate (ie transcription rate if promoter region bound
% by repressor) in transcripts per second
k_transcription_leakage = 5e-4;

% Set the mRNA and protein degradation rates (per second)
mRNA_half_life = 120; % in seconds
k_mRNA_degradation = log(2)/mRNA_half_life;
protein_half_life = 600; % in seconds
k_protein_degradation = log(2)/protein_half_life;

translation_efficiency = 20; % proteins per transcript lifespan
average_mRNA_lifespan = 1/k_mRNA_degradation; % in seconds
k_translation = translation_efficiency/average_mRNA_lifespan; % proteins per transcript per sec

% set the Hill coefficients of the repressors
n_tetR = 2;
n_cI = 2;
n_lacI = 2;

% Set the dissociation constant for the repressors to their target promoters
% in per molecule per second
KM_tetR = 40;
KM_cI = 40;
KM_lacI = 40;

% the differential equations governing the state variables:
% mRNA concentration = transcription given repressor concentration - mRNA degradation + transcription leakage
dydt(1,:) = k_transcription_cI/(1 + (protein_tetR / KM_tetR)^n_tetR) - k_mRNA_degradation*mRNA_cI + k_transcription_leakage;
dydt(2,:) = k_transcription_lacI/(1 + (protein_cI / KM_cI)^n_cI)- k_mRNA_degradation*mRNA_lacI + k_transcription_leakage;
dydt(3,:) = k_transcription_tetR/(1 + (protein_lacI / KM_lacI)^n_lacI) - k_mRNA_degradation*mRNA_tetR + k_transcription_leakage;
% protein concentration = translation - protein degradation
dydt(4,:) = k_translation*mRNA_cI - k_protein_degradation*protein_cI;
dydt(5,:) = k_translation*mRNA_lacI - k_protein_degradation*protein_lacI;
dydt(6,:) = k_translation*mRNA_tetR - k_protein_degradation*protein_tetR;