:Interneuron Cells to Pyramidal Cells GABA with local Ca2+ pool and read public soma Ca2+ pool
NEURON {
POINT_PROCESS interD2pyrVDA_STFD
USEION ca READ eca,ica
NONSPECIFIC_CURRENT igaba
RANGE initW
RANGE Cdur_gaba, AlphaTmax_gaba, Beta_gaba, Erev_gaba, gbar_gaba, W, on_gaba, g_gaba
RANGE eca, tauCa, Icatotal
RANGE ICag, P0g, fCag
RANGE Cainf, pooldiam, z
RANGE lambda1, lambda2, threshold1, threshold2
RANGE fmax, fmin, Wmax, Wmin, maxChange, normW, scaleW, srcid, destid
RANGE pregid,postgid, thr_rp
RANGE F, f, tauF, D1, d1, tauD1, D2, d2, tauD2
RANGE facfactor
}
UNITS {
(mV) = (millivolt)
(nA) = (nanoamp)
(uS) = (microsiemens)
FARADAY = 96485 (coul)
pi = 3.141592 (1)
}
PARAMETER {
srcid = -1 (1)
destid = -1 (1)
Cdur_gaba = 0.7254 (ms)
AlphaTmax_gaba = 7.2609(/ms)
Beta_gaba = 0.2667 (/ms)
Erev_gaba = -75 (mV)
gbar_gaba = 0.6e-3 (uS)
Cainf = 50e-6 (mM)
pooldiam = 1.8172 (micrometer)
z = 2
k = 0.01
tauCa = 50 (ms)
P0g = .01
fCag = .024
lambda1 = 2 : 1 : 1.5 : 2.0
lambda2 = 0.01
threshold1 = 0.47 : 0.45 : 0.4 : 0.43 : 0.8 : 1.0 : 1.1 : 0.55 (uM)
threshold2 = 0.52 : 0.5 : 0.45 : 0.48 : 1.2 : 1.3 : 1.2 : 0.70 (uM)
:GABA Weight
initW = 4.5 : : 3 : 2.5 : 1 : 5 : 6.25 : 5
fmax = 4 : 3.5 : 3 :4
fmin = .8
GAPstart1 = 96000
GAPstop1 = 196000
thr_rp = 1 : .7
facfactor = 1
: the (1) is needed for the range limits to be effective
f = 0 (1) < 0, 1e9 > : 1.3 (1) < 0, 1e9 > : facilitation
tauF = 20 (ms) < 1e-9, 1e9 >
d1 = 0.95 (1) < 0, 1 > : fast depression
tauD1 = 40 (ms) < 1e-9, 1e9 >
d2 = 0.9 (1) < 0, 1 > : slow depression
tauD2 = 70 (ms) < 1e-9, 1e9 >
DAstart1 = 39500
DAstop1 = 40000
DAstart2 = 35900
DAstop2 = 36000
DA_t1 = 0.8
DA_t2 = 1.3 : 1.2
DA_S = 1.5 : 1.8
Beta1 = 0.001 (/ms) : 1/decay time for neuromodulators
Beta2 = 0.0001 (/ms)
}
ASSIGNED {
v (mV)
eca (mV)
ica (nA)
igaba (nA)
g_gaba (uS)
on_gaba
W
t0 (ms)
ICan (mA)
ICag (mA)
Afactor (mM/ms/nA)
Icatotal (mA)
dW_gaba
Wmax
Wmin
maxChange
normW
scaleW
pregid
postgid
rp
tsyn
fa
F
D1
D2
}
STATE { r_nmda r_gaba capoolcon }
INITIAL {
on_gaba = 0
r_gaba = 0
W = initW
t0 = -1
Wmax = fmax*initW
Wmin = fmin*initW
maxChange = (Wmax-Wmin)/10
dW_gaba = 0
capoolcon = Cainf
Afactor = 1/(z*FARADAY*4/3*pi*(pooldiam/2)^3)*(1e6)
fa =0
F = 1
D1 = 1
D2 = 1
}
BREAKPOINT {
SOLVE release METHOD cnexp
}
DERIVATIVE release {
if (t0>0) {
if (rp < thr_rp) {
if (t-t0 < Cdur_gaba) {
on_gaba = 1
} else {
on_gaba = 0
}
} else {
on_gaba = 0
}
}
if (t0>0) {
if (rp < thr_rp) {
if (t-t0 < Cdur_gaba) {
on_gaba = 1
} else {
on_gaba = 0
}
} else {
on_gaba = 0
}
}
r_gaba' = AlphaTmax_gaba*on_gaba*(1-r_gaba)-Beta_gaba*r_gaba
dW_gaba = eta(capoolcon)*(lambda1*omega(capoolcon, threshold1, threshold2)-lambda2*GAP1(GAPstart1, GAPstop1)*W)*dt
: Limit for extreme large weight changes
if (fabs(dW_gaba) > maxChange) {
if (dW_gaba < 0) {
dW_gaba = -1*maxChange
} else {
dW_gaba = maxChange
}
}
:Normalize the weight change
normW = (W-Wmin)/(Wmax-Wmin)
if (dW_gaba < 0) {
scaleW = sqrt(fabs(normW))
} else {
scaleW = sqrt(fabs(1.0-normW))
}
W = W + dW_gaba*scaleW
:Weight value limits
if (W > Wmax) {
W = Wmax
} else if (W < Wmin) {
W = Wmin
}
g_gaba = gbar_gaba*r_gaba*facfactor*DA1(DAstart1,DAstop1)*DA2(DAstart2,DAstop2) : Dopamine effect on GABA
igaba = W*g_gaba*(v - Erev_gaba)
ICag = P0g*g_gaba*(v - eca)
Icatotal = ICag + k*ica*4*pi*((15/2)^2)*(0.01) : icag+k*ica*Area of soma*unit change
capoolcon'= -fCag*Afactor*Icatotal + (Cainf-capoolcon)/tauCa
}
NET_RECEIVE(dummy_weight) {
t0 = t
rp = unirand()
:F = 1 + (F-1)* exp(-(t - tsyn)/tauF)
D1 = 1 - (1-D1)*exp(-(t - tsyn)/tauD1)
D2 = 1 - (1-D2)*exp(-(t - tsyn)/tauD2)
:printf("%g\t%g\t%g\t%g\t%g\t%g\n", t, t-tsyn, F, D1, D2, facfactor)
:printf("%g\t%g\t%g\t%g\n", F, D1, D2, facfactor)
tsyn = t
facfactor = F * D1 * D2
::F = F+f :F * f
if (F > 3) {
F=3 }
if (facfactor < 0.7) {
facfactor=0.7
}
D1 = D1 * d1
D2 = D2 * d2
:printf("\t%g\t%g\t%g\n", F, D1, D2)
}
:::::::::::: FUNCTIONs and PROCEDUREs ::::::::::::
FUNCTION eta(Cani (mM)) {
LOCAL taulearn, P1, P2, P4, Cacon
P1 = 0.1
P2 = P1*1e-4
P4 = 1
Cacon = Cani*1e3
taulearn = P1/(P2+Cacon*Cacon*Cacon)+P4
eta = 1/taulearn*0.001
}
FUNCTION omega(Cani (mM), threshold1 (uM), threshold2 (uM)) {
LOCAL r, mid, Cacon
Cacon = Cani*1e3
r = (threshold2-threshold1)/2
mid = (threshold1+threshold2)/2
if (Cacon <= threshold1) { omega = 0}
else if (Cacon >= threshold2) { omega = 1/(1+50*exp(-50*(Cacon-threshold2)))}
else {omega = -sqrt(r*r-(Cacon-mid)*(Cacon-mid))}
}
FUNCTION DA1(DAstart1 (ms), DAstop1 (ms)) {
LOCAL DAtemp1, DAtemp2, DAtemp3, DAtemp4, DAtemp5, DAtemp6, DAtemp7, DAtemp8, DAtemp9, DAtemp10, DAtemp11, DAtemp12, DAtemp13, DAtemp14, DAtemp15, DAtemp16, DAtemp17, DAtemp18,s
DAtemp1 = DAstart1+4000
DAtemp2 = DAtemp1+4000
DAtemp3 = DAtemp2+4000
DAtemp4 = DAtemp3+4000
DAtemp5 = DAtemp4+4000
DAtemp6 = DAtemp5+4000
DAtemp7 = DAtemp6+4000
DAtemp8 = DAtemp7+4000
DAtemp9 = DAtemp8+4000
DAtemp10 = DAtemp9+4000
DAtemp11 = DAtemp10+4000
DAtemp12 = DAtemp11+4000
DAtemp13 = DAtemp12+4000
DAtemp14 = DAtemp13+4000
DAtemp15 = DAtemp14 + 4000 + 100000 : 100sec Gap
DAtemp16 = DAtemp15 + 4000
DAtemp17 = DAtemp16 + 4000
DAtemp18 = DAtemp17 + 4000
if (t <= DAstart1) { DA1 = 1.0}
else if (t >= DAstart1 && t <= DAstop1) {DA1 = DA_t1} : 2nd tone in conditioning
else if (t > DAstop1 && t < DAtemp1) {DA1 = 1.0 + (DA_t1-1)*exp(-Beta1*(t-DAstop1))} : Basal level
else if (t >= DAtemp1 && t <= DAtemp1+500) {DA1=DA_t1} : 3rd tone
else if (t > DAtemp1+500 && t < DAtemp2) {DA1 = 1.0 + (DA_t1-1)*exp(-Beta1*(t-(DAtemp1+500)))} : Basal level
else if (t >= DAtemp2 && t <= DAtemp2+500) {DA1=DA_t1} : 4th tone
else if (t > DAtemp2+500 && t < DAtemp3) {DA1 = 1.0 + (DA_t1-1)*exp(-Beta1*(t-(DAtemp2+500)))} : Basal level
else if (t >= DAtemp3 && t <= DAtemp3+500) {DA1=DA_t1} : 5th tone
else if (t > DAtemp3+500 && t < DAtemp4) {DA1 = 1.0 + (DA_t1-1)*exp(-Beta1*(t-(DAtemp3+500)))} : Basal level
else if (t >= DAtemp4 && t <= DAtemp4+500) {DA1=DA_t1} : 6th tone
else if (t > DAtemp4+500 && t < DAtemp5) {DA1 = 1.0 + (DA_t1-1)*exp(-Beta1*(t-(DAtemp4+500)))} : Basal level
else if (t >= DAtemp5 && t <= DAtemp5+500) {DA1=DA_t1} : 7th tone
else if (t > DAtemp5+500 && t < DAtemp6) {DA1 = 1.0 + (DA_t1-1)*exp(-Beta1*(t-(DAtemp5+500)))} : Basal level
else if (t >= DAtemp6 && t <= DAtemp6+500) {DA1=DA_t1} : 8th tone
else if (t > DAtemp6+500 && t < DAtemp7) {DA1 = 1.0 + (DA_t1-1)*exp(-Beta1*(t-(DAtemp6+500)))} : Basal level
else if (t >= DAtemp7 && t <= DAtemp7+500) {DA1=DA_t1} : 9th tone
else if (t > DAtemp7+500 && t < DAtemp8) {DA1 = 1.0 + (DA_t1-1)*exp(-Beta1*(t-(DAtemp7+500)))} : Basal level
else if (t >= DAtemp8 && t <= DAtemp8+500) {DA1=DA_t1} : 10th tone
else if (t > DAtemp8+500 && t < DAtemp9) {DA1 = 1.0 + (DA_t1-1)*exp(-Beta1*(t-(DAtemp8+500)))} : Basal level
else if (t >= DAtemp9 && t <= DAtemp9+500) {DA1=DA_t2} : 11th tone - Second Step
else if (t > DAtemp9+500 && t < DAtemp10) {DA1 = 1.0 + (DA_t2-1)*exp(-Beta2*(t-(DAtemp9+500)))} : Basal level
else if (t >= DAtemp10 && t <= DAtemp10+500) {DA1=DA_t2} : 12th tone
else if (t > DAtemp10+500 && t < DAtemp11) {DA1 = 1.0 + (DA_t2-1)*exp(-Beta2*(t-(DAtemp10+500)))} : Basal level
else if (t >= DAtemp11 && t <= DAtemp11+500) {DA1=DA_t2} : 13th tone
else if (t > DAtemp11+500 && t < DAtemp12) {DA1 = 1.0 + (DA_t2-1)*exp(-Beta2*(t-(DAtemp11+500)))} : Basal level
else if (t >= DAtemp12 && t <= DAtemp12+500) {DA1=DA_t2} : 14th tone
else if (t > DAtemp12+500 && t < DAtemp13) {DA1 = 1.0 + (DA_t2-1)*exp(-Beta2*(t-(DAtemp12+500)))} : Basal level
else if (t >= DAtemp13 && t <= DAtemp13+500) {DA1=DA_t2} : 15th tone
else if (t > DAtemp13+500 && t < DAtemp14) {DA1 = 1.0 + (DA_t2-1)*exp(-Beta2*(t-(DAtemp13+500)))} : Basal level
else if (t >= DAtemp14 && t <= DAtemp14+500) {DA1=DA_t2} : 16th tone
else if (t > DAtemp14+500 && t < DAtemp15) {DA1 = 1.0 + (DA_t2-1)*exp(-Beta2*(t-(DAtemp14+500)))} : Basal level
else if (t >= DAtemp15 && t <= DAtemp15+500) {DA1=DA_t2} : 1st tone in Extinction
else if (t > DAtemp15+500 && t < DAtemp16) {DA1 = 1.0 + (DA_t2-1)*exp(-Beta2*(t-(DAtemp15+500)))} : Basal level
else if (t >= DAtemp16 && t <= DAtemp16+500) {DA1=DA_t2} : 2nd tone
else if (t > DAtemp16+500 && t < DAtemp17) {DA1 = 1.0 + (DA_t2-1)*exp(-Beta2*(t-(DAtemp16+500)))} : Basal level
else if (t >= DAtemp17 && t <= DAtemp17+500) {DA1=DA_t2} : 3rd tone
else if (t > DAtemp17+500 && t < DAtemp18) {DA1 = 1.0 + (DA_t2-1)*exp(-Beta2*(t-(DAtemp17+500)))} : Basal level
else if (t >= DAtemp18 && t <= DAtemp18+500) {DA1=1.0} : 4th tone
else { DA1 = 1.0}
}
FUNCTION DA2(DAstart2 (ms), DAstop2 (ms)) {
LOCAL DA2temp1, DA2temp2, DA2temp3, DA2temp4, DA2temp5, DA2temp6, DA2temp7, DA2temp8, DA2temp9, DA2temp10, DA2temp11, DA2temp12, DA2temp13, DA2temp14, DA2temp15, DA2temp16,s
DA2temp1 = DAstart2 + 4000
DA2temp2 = DA2temp1 + 4000
DA2temp3 = DA2temp2 + 4000
DA2temp4 = DA2temp3 + 4000
DA2temp5 = DA2temp4 + 4000
DA2temp6 = DA2temp5 + 4000
DA2temp7 = DA2temp6 + 4000
DA2temp8 = DA2temp7 + 4000
DA2temp9 = DA2temp8 + 4000
DA2temp10 = DA2temp9 + 4000
DA2temp11 = DA2temp10 + 4000
DA2temp12 = DA2temp11 + 4000
DA2temp13 = DA2temp12 + 4000
DA2temp14 = DA2temp13 + 4000
DA2temp15 = DA2temp14 + 4000
if (t <= DAstart2) { DA2 = 1.0}
else if (t >= DAstart2 && t <= DAstop2) {DA2 = DA_S } : 1st shock
else if (t > DAstop2 && t < DA2temp1) {DA2 = 1.0 + (DA_S-1)*exp(-Beta2*(t-(DAstop2+500)))}
else if (t >= DA2temp1 && t <= DA2temp1+100) {DA2=DA_S} : 2nd shock
else if (t > DA2temp1+100 && t < DA2temp2) {DA2 = 1.0 + (DA_S-1)*exp(-Beta2*(t-(DA2temp1+100)))}
else if (t >= DA2temp2 && t <= DA2temp2+100) {DA2=DA_S} : 3rd shock
else if (t > DA2temp2+100 && t < DA2temp3) {DA2 = 1.0 + (DA_S-1)*exp(-Beta2*(t-(DA2temp2+100)))}
else if (t >= DA2temp3 && t <= DA2temp3+100) {DA2=DA_S} : 4th shock
else if (t > DA2temp3+100 && t < DA2temp4) {DA2 = 1.0 + (DA_S-1)*exp(-Beta2*(t-(DA2temp3+100)))}
else if (t >= DA2temp4 && t <= DA2temp4+100) {DA2=DA_S} : 5th shock
else if (t > DA2temp4+100 && t < DA2temp5) {DA2 = 1.0 + (DA_S-1)*exp(-Beta2*(t-(DA2temp4+100)))}
else if (t >= DA2temp5 && t <= DA2temp5+100) {DA2=DA_S} : 6th shock
else if (t > DA2temp5+100 && t < DA2temp6) {DA2 = 1.0 + (DA_S-1)*exp(-Beta2*(t-(DA2temp5+100)))}
else if (t >= DA2temp6 && t <= DA2temp6+100) {DA2=DA_S} : 7th shock
else if (t > DA2temp6+100 && t < DA2temp7) {DA2 = 1.0 + (DA_S-1)*exp(-Beta2*(t-(DA2temp6+100)))}
else if (t >= DA2temp7 && t <= DA2temp7+100) {DA2=DA_S} : 8th shock
else if (t > DA2temp7+100 && t < DA2temp8) {DA2 = 1.0 + (DA_S-1)*exp(-Beta2*(t-(DA2temp7+100)))}
else if (t >= DA2temp8 && t <= DA2temp8+100) {DA2=DA_S } : 9th shock
else if (t > DA2temp8+100 && t < DA2temp9) {DA2 = 1.0 + (DA_S-1)*exp(-Beta2*(t-(DA2temp8+100)))}
else if (t >= DA2temp9 && t <= DA2temp9+100) {DA2=DA_S } : 10th shock
else if (t > DA2temp9+100 && t < DA2temp10) {DA2 = 1.0 + (DA_S-1)*exp(-Beta2*(t-(DA2temp9+100)))}
else if (t >= DA2temp10 && t <= DA2temp10+100) {DA2=DA_S} : 11th shock
else if (t > DA2temp10+100 && t < DA2temp11) {DA2 = 1.0 + (DA_S-1)*exp(-Beta2*(t-(DA2temp10+100)))}
else if (t >= DA2temp11 && t <= DA2temp11+100) {DA2=DA_S } : 12th shock
else if (t > DA2temp11+100 && t < DA2temp12) {DA2 = 1.0 + (DA_S-1)*exp(-Beta2*(t-(DA2temp11+100)))}
else if (t >= DA2temp12 && t <= DA2temp12+100) {DA2=DA_S} : 13th shock
else if (t > DA2temp12+100 && t < DA2temp13) {DA2 = 1.0 + (DA_S-1)*exp(-Beta2*(t-(DA2temp12+100)))}
else if (t >= DA2temp13 && t <= DA2temp13+100) {DA2=DA_S } : 14th shock
else if (t > DA2temp13+100 && t < DA2temp14) {DA2 = 1.0 + (DA_S-1)*exp(-Beta2*(t-(DA2temp13+100)))}
else if (t >= DA2temp14 && t <= DA2temp14+100) {DA2=DA_S} : 15th shock
else if (t > DA2temp14+100 && t < DA2temp15) {DA2 = 1.0 + (DA_S-1)*exp(-Beta2*(t-(DA2temp14+100)))}
else if (t >= DA2temp15 && t <= DA2temp15+100) {DA2=DA_S} : 16th shock
else { DA2 = 1.0}
}
FUNCTION GAP1(GAPstart1 (ms), GAPstop1 (ms)) {
LOCAL s
if (t <= GAPstart1) { GAP1 = 1}
else if (t >= GAPstart1 && t <= GAPstop1) {GAP1 = 1} : During the Gap, apply lamda2*2
else { GAP1 = 1}
}
FUNCTION unirand() { : uniform random numbers between 0 and 1
unirand = scop_random()
}