Post-synaptic facilitation and network dynamics underlying stimulus-specific combination sensitivity.
Paper
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The authors' code
MATLAB · 298 lines · 9.5 KB · MIT
- function results = Figure_2()
- P = get_common_params();
- cases(1) = struct('title','(A) AB-0 ms delay','from1',100,'to1',150,'from2',150,'to2',200,'reverse_order',false);
- cases(2) = struct('title','(B) AB-early B','from1',100,'to1',150,'from2',120,'to2',170,'reverse_order',false);
- cases(3) = struct('title','(C) AB-50 ms delay','from1',100,'to1',150,'from2',200,'to2',250,'reverse_order',false);
- cases(4) = struct('title','(D) BA-0 ms delay','from1',100,'to1',150,'from2',150,'to2',200,'reverse_order',true);
- figure('Color','w');
- tiledlayout(2,2,'Padding','compact','TileSpacing','compact');
- results = struct([]);
- for k = 1:4
- [t, sol] = run_case(P, cases(k));
- nexttile;
- plot_case(t, sol, cases(k).title);
- results(k).t = t;
- results(k).sol = sol;
- results(k).case = cases(k);
- end
- end
- function P = get_common_params()
- P.Iapp1 = 200;
- P.Iapp2 = 200;
- P.gCaTINT1 = 0.5;
- P.gCaTX2 = 0.5;
- P.gCaTCSN = 0.8;
- P.gSKX2 = 2;
- P.gSKCSN = 2;
- P.gGAi1csn = 10;
- P.gAMx2csn = 1.2;
- P.gAMCNCN = 5;
- P.gNaINT = 800;
- P.gKINT = 1200;
- P.gHINT = 4;
- P.CINT = 20;
- P.krINT = 0.01;
- P.gNaX = 450;
- P.gKX = 60;
- P.gHX = 6;
- P.CX = 20;
- P.krX = 0.3;
- P.taur0 = 5;
- P.taur1 = 15;
- P.thetarrT = 68;
- P.sigmarrT = 2;
- P.thetarT = -67;
- P.sigmarT = 2;
- P.thetaaT = -85;
- P.sigmaaT = -8;
- P.thetab = 0.4;
- P.sigmab = -0.1;
- P.phirT = 0.2;
- P.VL = -70;
- P.VK = -90;
- P.VNa = 70;
- P.VH = -30;
- P.Tmax = 1;
- P.VT = 2;
- P.Kps = 5;
- P.gL = 2;
- P.gCa = 19;
- P.arGA = 5;
- P.adGA = 0.18;
- P.VGA = -100;
- P.arAM = 1.1;
- P.adAM = 0.19;
- P.VAM = 0;
- P.taunbar = 10;
- P.tauh = 1;
- P.Ca_ex = 2.5;
- P.RTF = 26.7;
- P.thetam = -35;
- P.thetan = -30;
- P.thetas = -20;
- P.sigmam = -5;
- P.sigman = -5;
- P.sigmas = -0.05;
- P.f = 0.1;
- P.eps = 0.0015;
- P.kCa = 0.3;
- P.bCa = 0.1;
- P.ks = 0.5;
- P.taurs = 1500;
- P.thetarf = -105;
- P.thetars = -105;
- P.sigmarf = 5;
- P.sigmars = 25;
- P.prf = 100;
- P.tspan = 0:0.01:900;
- P.INTinit = [-67.6; 0.5e-03; 0.996; 0.1355; 0.1; 5.5e-04; 0.034];
- P.Xinit = [-75; 0.01; 0.01; 0.01; 0.01; 0.01; 0.01];
- P.init_cond = [P.INTinit; P.Xinit; P.Xinit; 0.01; 0.01; 0.01];
- end
- function [t, sol] = run_case(P, caseDef)
- P.from1 = caseDef.from1;
- P.to1 = caseDef.to1;
- P.from2 = caseDef.from2;
- P.to2 = caseDef.to2;
- P.reverse_order = caseDef.reverse_order;
- [t, sol] = ode113(@(t,sol) SimpleFTN_local(t, sol, P), P.tspan, P.init_cond);
- end
- function plot_case(t, sol, titleStr)
- vINT1 = sol(:,1);
- vX2 = sol(:,8);
- vCSN = sol(:,15);
- step = 150;
- i = 1;
- hold on
- plot(t, vINT1-step*i, 'Color', [0.46,0.01,0.65], 'LineWidth', 1);
- text(60, -step*i, 'A');
- i = i + 1;
- plot(t, vX2-step*i, 'Color', [0.61,0.62,0.67], 'LineWidth', 1);
- text(60, -step*i, 'B');
- i = i + 1;
- plot(t, vCSN-step*i, 'Color', [0.04,0.17,0.55], 'LineWidth', 1);
- text(60, -step*i, 'CSN');
- i = i + 1;
- yMin = min([vINT1; vX2; vCSN]) - (i * step) - 100;
- yMax = max([vINT1; vX2; vCSN]) + 100;
- xlim([50 300]);
- ylim([yMin yMax]);
- scaleLengthX = 10;
- scaleLengthY = 50;
- x_start = 270;
- y_start = yMin + 200;
- line([x_start, x_start + scaleLengthX], [y_start, y_start], 'Color', 'k', 'LineWidth', 2);
- line([x_start, x_start], [y_start, y_start + scaleLengthY], 'Color', 'k', 'LineWidth', 2);
- text(x_start + scaleLengthX/2, y_start - 30, '10 ms', 'HorizontalAlignment', 'center', 'FontSize', 9);
- text(x_start - 5, y_start + scaleLengthY/2, '50 mV', 'HorizontalAlignment', 'right', 'FontSize', 9);
- title(titleStr, 'FontWeight', 'normal');
- set(gca, 'YTick', [], 'XTick', [], 'XColor', 'none', 'YColor', 'none');
- box off
- hold off
- end
- function output = SimpleFTN_local(t, sol, P)
- VINT1 = sol(1); nINT1 = sol(2); hINT1 = sol(3); rTINT1 = sol(4); CaiINT1 = sol(5); rfINT1 = sol(6); rsINT1 = sol(7);
- VX2 = sol(8); nX2 = sol(9); hX2 = sol(10); rTX2 = sol(11); CaiX2 = sol(12); rfX2 = sol(13); rsX2 = sol(14);
- VCSN = sol(15); nCSN = sol(16); hCSN = sol(17); rTCSN = sol(18); CaiCSN = sol(19); rfCSN = sol(20); rsCSN = sol(21);
- sGAi1csn = sol(22);
- sAMx2csn = sol(23);
- sAMCNCN = sol(24);
- sinfINT1 = 1/(1 + exp((VINT1-P.thetas)/P.sigmas));
- iCaLINT1 = P.gCa*(sinfINT1^2)*VINT1*(P.Ca_ex/(1-exp((2*VINT1)/P.RTF)));
- aTinfINT1 = 1/(1 + exp((VINT1-P.thetaaT)/P.sigmaaT));
- bTinfINT1 = 1/(1 + exp((rTINT1-P.thetab)/P.sigmab)) - 1/(1 + exp(-P.thetab/P.sigmab));
- rTinfINT1 = 1/(1 + exp((VINT1-P.thetarT)/P.sigmarT));
- taurTINT1 = P.taur0 + P.taur1/(1 + exp((VINT1-P.thetarrT)/P.sigmarrT));
- iCaTINT1 = P.gCaTINT1*(aTinfINT1^3)*(bTinfINT1^2)*VINT1*(P.Ca_ex/(1-exp((2*VINT1)/P.RTF)));
- iLINT1 = P.gL*(VINT1-P.VL);
- ninfINT1 = 1/(1 + exp((VINT1-P.thetan)/P.sigman));
- taunINT1 = P.taunbar/cosh((VINT1-P.thetan)/(2*P.sigman));
- iKINT1 = P.gKINT*(nINT1^4)*(VINT1-P.VK);
- minfINT1 = 1/(1 + exp((VINT1-P.thetam)/P.sigmam));
- alphahINT1 = 0.128*exp(-(VINT1+50)/18);
- betahINT1 = 4/(1 + exp(-(VINT1+27)/5));
- hinfINT1 = alphahINT1/(alphahINT1 + betahINT1);
- iNaINT1 = P.gNaINT*(minfINT1^3)*hINT1*(VINT1-P.VNa);
- rfinfINT1 = 1/(1 + exp((VINT1-P.thetarf)/P.sigmarf));
- taurfINT1 = P.prf/(-7.4*(VINT1+70)/(exp(-(VINT1+70)/0.8)-1) + 65*exp(-(VINT1+56)/23));
- rsinfINT1 = 1/(1 + exp((VINT1-P.thetars)/P.sigmars));
- iHINT1 = P.gHINT*(P.krINT*rfINT1 + (1-P.krINT)*rsINT1)*(VINT1-P.VH);
- TINT1 = P.Tmax/(1 + exp(-(VINT1-P.VT)/P.Kps));
- minfX2 = 1/(1 + exp((VX2-P.thetam)/P.sigmam));
- ninfX2 = 1/(1 + exp((VX2-P.thetan)/P.sigman));
- tauNX2 = P.taunbar./cosh((VX2-P.thetan)/(2*P.sigman));
- alphaHX2 = 0.128*exp(-(VX2+50)/18);
- betaHX2 = 4/(1 + exp(-(VX2+27)/5));
- hinfX2 = alphaHX2/(alphaHX2 + betaHX2);
- iNaX2 = P.gNaX*(minfX2^3)*hX2*(VX2-P.VNa);
- iKX2 = P.gKX*(nX2^4)*(VX2-P.VK);
- sinfX2 = 1/(1 + exp((VX2-P.thetas)/P.sigmas));
- iCaLX2 = P.gCa*(sinfX2^2)*VX2*(P.Ca_ex/(1-exp((2*VX2)/P.RTF)));
- aTinfX2 = 1/(1 + exp((VX2-P.thetaaT)/P.sigmaaT));
- bTinfX2 = 1/(1 + exp((rTX2-P.thetab)/P.sigmab)) - 1/(1 + exp(-P.thetab/P.sigmab));
- rTinfX2 = 1/(1 + exp((VX2-P.thetarT)/P.sigmarT));
- taurTX2 = P.taur0 + P.taur1/(1 + exp((VX2-P.thetarrT)/P.sigmarrT));
- iCaTX2 = P.gCaTX2*(aTinfX2^3)*(bTinfX2^2)*VX2*(P.Ca_ex/(1-exp((2*VX2)/P.RTF)));
- kinfX2 = (CaiX2^2)/(CaiX2^2 + (P.ks^2));
- iSKX2 = P.gSKX2*kinfX2*(VX2-P.VK);
- rinffX2 = 1/(1 + exp((VX2-P.thetarf)/P.sigmarf));
- rinfsX2 = 1/(1 + exp((VX2-P.thetars)/P.sigmars));
- taurfX2 = P.prf./(-7.4*(VX2+70)./(exp(-(VX2+70)/0.8)-1) + 65*exp(-(VX2+56)/23));
- iHX2 = P.gHX*(P.krX*rfX2 + (1-P.krX)*rsX2)*(VX2-P.VH);
- iLX2 = P.gL*(VX2-P.VL);
- TX2 = P.Tmax/(1 + exp(-(VX2-P.VT)/P.Kps));
- minfCSN = 1/(1 + exp((VCSN-P.thetam)/P.sigmam));
- ninfCSN = 1/(1 + exp((VCSN-P.thetan)/P.sigman));
- tauNCSN = P.taunbar./cosh((VCSN-P.thetan)/(2*P.sigman));
- alphaHCSN = 0.128*exp(-(VCSN+50)/18);
- betaHCSN = 4/(1 + exp(-(VCSN+27)/5));
- hinfCSN = alphaHCSN/(alphaHCSN + betaHCSN);
- iNaCSN = P.gNaX*(minfCSN^3)*hCSN*(VCSN-P.VNa);
- iKCSN = P.gKX*(nCSN^4)*(VCSN-P.VK);
- sinfCSN = 1/(1 + exp((VCSN-P.thetas)/P.sigmas));
- iCaLCSN = P.gCa*(sinfCSN^2)*VCSN*(P.Ca_ex/(1-exp((2*VCSN)/P.RTF)));
- aTinfCSN = 1/(1 + exp((VCSN-P.thetaaT)/P.sigmaaT));
- bTinfCSN = 1/(1 + exp((rTCSN-P.thetab)/P.sigmab)) - 1/(1 + exp(-P.thetab/P.sigmab));
- rTinfCSN = 1/(1 + exp((VCSN-P.thetarT)/P.sigmarT));
- taurTCSN = P.taur0 + P.taur1/(1 + exp((VCSN-P.thetarrT)/P.sigmarrT));
- iCaTCSN = P.gCaTCSN*(aTinfCSN^3)*(bTinfCSN^3)*VCSN*(P.Ca_ex/(1-exp((2*VCSN)/P.RTF)));
- kinfCSN = (CaiCSN^2)/(CaiCSN^2 + (P.ks^2));
- iSKCSN = P.gSKCSN*kinfCSN*(VCSN-P.VK);
- rinffCSN = 1/(1 + exp((VCSN-P.thetarf)/P.sigmarf));
- rinfsCSN = 1/(1 + exp((VCSN-P.thetars)/P.sigmars));
- taurfCSN = P.prf./(-7.4*(VCSN+70)./(exp(-(VCSN+70)/0.8)-1) + 65*exp(-(VCSN+56)/23));
- iHCSN = P.gHX*(P.krX*rfCSN + (1-P.krX)*rsCSN)*(VCSN-P.VH);
- iLCSN = P.gL*(VCSN-P.VL);
- TCSN = P.Tmax/(1 + exp(-(VCSN-P.VT)/P.Kps));
- iGAi1csn = P.gGAi1csn*sGAi1csn*(VCSN-P.VGA);
- iAMx2csn = P.gAMx2csn*sAMx2csn*(VCSN-P.VAM);
- iAMCNCN = P.gAMCNCN*sAMCNCN*(VCSN-P.VAM);
- if ~P.reverse_order
- if (t > P.from1 && t < P.to1)
- curr1 = P.Iapp1;
- else
- curr1 = 0;
- end
- if (t > P.from2 && t < P.to2)
- curr2 = P.Iapp2;
- else
- curr2 = 0;
- end
- else
- if (t > P.from1 && t < P.to1)
- curr2 = P.Iapp2;
- else
- curr2 = 0;
- end
- if (t > P.from2 && t < P.to2)
- curr1 = P.Iapp1;
- else
- curr1 = 0;
- end
- end
- output = zeros(24,1);
- output(1) = (-iNaINT1-iKINT1-iCaLINT1-iCaTINT1-iHINT1-iLINT1+curr1)/P.CINT;
- output(2) = (ninfINT1-nINT1)/taunINT1;
- output(3) = (hinfINT1-hINT1)/P.tauh;
- output(4) = P.phirT*(rTinfINT1-rTINT1)/taurTINT1;
- output(5) = -P.f*(P.eps*(iCaLINT1+iCaTINT1) + P.kCa*(CaiINT1-P.bCa));
- output(6) = (rfinfINT1-rfINT1)/taurfINT1;
- output(7) = (rsinfINT1-rsINT1)/P.taurs;
- output(8) = (-iNaX2-iKX2-iCaLX2-iCaTX2-iSKX2-iHX2-iLX2+curr2)/P.CX;
- output(9) = (ninfX2-nX2)/tauNX2;
- output(10) = (hinfX2-hX2)/P.tauh;
- output(11) = P.phirT*(rTinfX2-rTX2)/taurTX2;
- output(12) = -P.f*(P.eps*(iCaLX2+iCaTX2) + P.kCa*(CaiX2-P.bCa));
- output(13) = (rinffX2-rfX2)/taurfX2;
- output(14) = (rinfsX2-rsX2)/P.taurs;
- output(15) = (-iNaCSN-iKCSN-iCaLCSN-iCaTCSN-iSKCSN-iHCSN-iLCSN-iGAi1csn-iAMx2csn-iAMCNCN)/P.CX;
- output(16) = (ninfCSN-nCSN)/tauNCSN;
- output(17) = (hinfCSN-hCSN)/P.tauh;
- output(18) = P.phirT*(rTinfCSN-rTCSN)/taurTCSN;
- output(19) = -P.f*(P.eps*(iCaLCSN+iCaTCSN) + P.kCa*(CaiCSN-P.bCa));
- output(20) = (rinffCSN-rfCSN)/taurfCSN;
- output(21) = (rinfsCSN-rsCSN)/P.taurs;
- output(22) = P.arGA*TINT1*(1-sGAi1csn) - P.adGA*sGAi1csn;
- output(23) = P.arAM*TX2*(1-sAMx2csn) - P.adAM*sAMx2csn;
- output(24) = P.arAM*TCSN*(1-sAMCNCN) - P.adAM*sAMCNCN;
- end
Figure_2.m at commit 873b631, under MIT · at the source
Overview
Abstract
Combination-sensitive neurons (CSNs) transform specific combinations of sensory features into selective neural responses, supporting the temporal organization of complex signals. We developed a biophysically realistic computational model, constrained by intrinsic and synaptic properties, to examine how temporally ordered stimulus pairs can be associated across delays of hundreds of milliseconds. The model shows that upstream inhibitory-delay circuits can transiently preserve information about the first stimulus while progressively shaping the excitability of an output neuron. Together with post-inhibitory rebound and facilitation, these dynamics create a delay-dependent primed state that funnels separated inputs into a narrow coincidence-detection window. This mechanism enables sharply timed, selective responses to stimulus pairs without requiring sustained spiking in the output neuron. Our results provide a circuit-level link between coincidence detection and longer-timescale temporal integration, offering a general framework for temporal coding in sensory systems.
Reproduced under the paper's license (CC BY), from the paper cited above.
Repository
Its files are read in the Code ↔ Paper reader above.
daoulab/combination-sensitivity-model
873b631e0e6cc7156ea34a2f84c353e46747ad94, 13 April 2026Availability: 1 check, the latest on 28 September 2026: the link answers
- 28 September 2026: the link answers
15 files
- Figure_2.m, MATLAB, 298 lines
- Figure_3.m, MATLAB, 402 lines
- Figure_4.m, MATLAB, 427 lines
- Figure_5.m, MATLAB, 338 lines
- Figure_6.m, MATLAB, 342 lines
- Figure_7.m, MATLAB, 164 lines
- Figure_8.m, MATLAB, 469 lines
- Figure_S1.m, MATLAB, 125 lines
- Figure_S2.m, MATLAB, 285 lines
- Figure_S3.m, MATLAB, 295 lines
- Figure_S4.m, MATLAB, 397 lines
- Figure_S5.m, MATLAB, 340 lines
- Figure_S6.m, MATLAB, 357 lines
- LICENSE, License, 21 lines
- README.md, Text, 107 lines
The paper's code and data availability statement is in the Data section.
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Data and code availability
• Data: This study did not generate new unique datasets. All results reported in this paper were produced from computational simulations and can be reproduced from the code provided below. • Code: MATLAB source codes that generate every figure and supplementary figure in the paper is available on GitHub at https://
Reproduced under the paper's license (CC BY), from the paper cited above.
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Version 1, 28 September 2026: the first record
Recorded: type, language, journal, volume, issue, pages, dates, 2 authors, 3 keywords, 2 funders, 92 references.
Cite
This paper
Merabi, Z., & Daou, A. (2026). Post-synaptic facilitation and network dynamics underlying stimulus-specific combination sensitivity. iScience, 29(6), 116125. https://
BibTeX
@article{merabi2026post,
author = {Merabi, Zeina and Daou, Arij},
title = {{Post-synaptic facilitation and network dynamics underlying stimulus-specific combination sensitivity}},
journal = {iScience},
year = {2026},
month = may,
volume = {29},
number = {6},
pages = {116125},
publisher = {Elsevier},
issn = {2589-0042},
doi = {10.1016/
url = {https://
pmid = {42256266},
pmcid = {PMC13235528}
}
RIS
TY - JOUR
AU - Merabi, Zeina
AU - Daou, Arij
TI - Post-synaptic facilitation and network dynamics underlying stimulus-specific combination sensitivity
T2 - iScience
J2 - iScience
PY - 2026
DA - 2026/
VL - 29
IS - 6
SP - 116125
SN - 2589-0042
PB - Elsevier
DO - 10.1016/
UR - https://
LA - en
ER -
CSL-JSON
{
"id": "10.1016/
"type": "article-journal",
"title": "Post-synaptic facilitation and network dynamics underlying stimulus-specific combination sensitivity",
"container-title": "iScience",
"author": [
{
"family": "Merabi",
"given": "Zeina"
},
{
"family": "Daou",
"given": "Arij"
}
],
"container-title-short":
"volume": "29",
"issue": "6",
"page": "116125",
"DOI": "10.1016/
"PMID": "42256266",
"PMCID": "PMC13235528",
"ISSN": "2589-0042",
"publisher": "Elsevier",
"URL": "https://
"language": "en",
"issued": {
"date-parts": [
[
2026,
5,
27
]
]
}
}
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