NEURON {
POINT_PROCESS GRANULE_Nmda_det_vi
NONSPECIFIC_CURRENT i
RANGE Q10_diff,Q10_channel
RANGE g , ic
RANGE Cdur,Erev,T,Tmax
RANGE Rb, Ru, Rd, Rr, Ro, Rc,rb,gmax,RdRate
RANGE tau_1, tau_rec, tau_facil, U, u0
RANGE PRE
RANGE Used
RANGE MgBlock,v0_block,k_block
RANGE diffuse,Trelease,lamd, Diff, M, Rd, nd, syntype, gmax_factor
}
UNITS {
(nA) = (nanoamp)
(mV) = (millivolt)
(umho) = (micromho)
(mM) = (milli/liter)
(pS) = (picosiemens)
(nS) = (nanosiemens)
(um) = (micrometer)
PI = (pi) (1)
}
PARAMETER {
syntype
gmax_factor = 1
: Parametri Presinaptici
gmax = 18800 (pS) : 7e3 : 4e4
Q10_diff = 1.5
Q10_channel = 2.4
U = 0.42 (1) < 0, 1 >
tau_rec = 8 (ms) < 1e-9, 1e9 >
tau_facil = 5 (ms) < 0, 1e9 >
M = 21.515 : numero di (kilo) molecole in una vescicola
Rd = 1.03 (um)
Diff = 0.223 (um2/ms)
tau_1 = 1 (ms) < 1e-9, 1e9 >
u0 = 0 (1) < 0, 1 >
Tmax = 1 (mM)
: Postsinaptico, Westbrook scheme
Cdur = 0.3 (ms)
Rb = 5 (/ms/mM) : binding
Ru = 0.1 (/ms) : unbinding
RdRate = 12e-3 (/ms) : desensitization
Rr = 9e-3 (/ms) : resensitization
Ro = 3e-2 (/ms) : opening
Rc = 0.966 (/ms) : closing
Erev = -3.7 (mV) : 0 (mV)
v0_block = -20 (mV) : -16 -8.69 (mV) : -18.69 (mV) : -32.7 (mV)
k_block = 13 (mV)
nd = 1
kB = 0.44 (mM)
: Diffusion
diffuse = 1
lamd = 20 (nm)
celsius (degC)
}
ASSIGNED {
v (mV) : postsynaptic voltage
i (nA) : current = g*(v - Erev)
ic (nA) : current = g*(v - Erev)
g (pS) : actual conductance
rb (/ms) : binding
T (mM)
x
Trelease (mM)
tspike[100] (ms) : will be initialized by the pointprocess
PRE[100]
Mres (mM)
MgBlock
numpulses
tzero
gbar_Q10 (mho/cm2)
Q10 (1)
}
STATE {
: Channel states (all fractions)
C0 : unbound
C1 : single bound
C2 : double bound
D : desensitized
O : open
}
INITIAL {
rates(v)
C0 = 1
C1 = 0
C2 = 0
D = 0
O = 0
T = 0
numpulses=0
gbar_Q10 = Q10_diff^((celsius-30)/10)
Q10 = Q10_channel^((celsius-30)/10)
Mres = 1e3 * (1e3 * 1e15 / 6.022e23 * M) : (M) to (mM) so 1e3, 1um^3=1dm^3*1e-15 so 1e15
FROM i=1 TO 100 { PRE[i-1]=0 tspike[i-1]=0 } :PRE_2[500]=0}
tspike[0]=1e12 (ms)
if(tau_1>=tau_rec){
printf("Warning: tau_1 (%g) should never be higher neither equal to tau_rec (%g)!\n",tau_1,tau_rec)
tau_rec=tau_1+1e-5
:printf("tau_rec has been set to %g\n",tau_rec)
}
}
FUNCTION imax(a,b) {
if (a>b) { imax=a }
else { imax=b }
}
FUNCTION diffusione(){
LOCAL DifWave,i,cntc,fi
DifWave=0
cntc=imax(numpulses-100,0)
FROM i=cntc TO numpulses{
fi=fmod(i,100)
tzero=tspike[fi]
if(t>tzero){
DifWave=DifWave+PRE[fi]*Mres*exp(-Rd*Rd/(4*Diff*(t-tzero)))/((4*PI*Diff*(1e-3)*lamd)*(t-tzero))^nd
}
}
diffusione=DifWave
}
BREAKPOINT {
rates(v)
SOLVE kstates METHOD sparse
g = gmax * gbar_Q10 * O * gmax_factor
: E' piu' logico spostare * MgBlock * PRE sul calcolo della corrente!
i = (1e-6) * g * (v - Erev) * MgBlock
ic = i
}
KINETIC kstates {
:if ( diffuse && (t>tspike[0]) ) { Trelease= T + diffusione() } else { Trelease=T }
Trelease = diffusione()
rb = Rb * Trelease :^2 / (Trelease + kB)^2
~ C0 <-> C1 (rb*Q10,Ru*Q10) : (fattore*rb,Ru) qui 2* per descrizione part.identiche
~ C1 <-> C2 (rb*Q10,Ru*Q10) : (rb,fattore*Ru) idem
~ C2 <-> D (RdRate*Q10,Rr*Q10)
~ C2 <-> O (Ro*Q10,Rc*Q10)
CONSERVE C0+C1+C2+D+O = 1
}
PROCEDURE rates(v(mV)) {
: E' necessario includere DEPEND v0_block,k_block per aggiornare le tabelle!
TABLE MgBlock DEPEND v0_block,k_block FROM -120 TO 30 WITH 150
MgBlock = 1 / ( 1 + exp ( - ( v - v0_block ) / k_block ) )
}
NET_RECEIVE(weight, on, nspike, tzero (ms),y, z, u, tsyn (ms)) {LOCAL fi
: *********** ATTENZIONE! ***********
:
: Qualora si vogliano utilizzare impulsi di glutammato saturanti e'
: necessario che il pulse sia piu' corto dell'intera simulazione
: altrimenti la variabile on non torna al suo valore di default.
INITIAL {
y = 0
z = 0
u = u0
tsyn = t
nspike = 1
}
if (flag == 0) {
: Qui faccio rientrare la modulazione presinaptica
nspike = nspike + 1
if (!on) {
tzero = t
on = 1
z = z*exp( - (t - tsyn) / (tau_rec) ) : RESCALED !
z = z + ( y*(exp(-(t - tsyn)/tau_1) - exp(-(t - tsyn)/(tau_rec)))/((tau_1/(tau_rec))-1) ) : RESCALED !
y = y*exp(-(t - tsyn)/tau_1)
x = 1-y-z
if (tau_facil > 0) {
u = u*exp(-(t - tsyn)/tau_facil)
u = u + U * ( 1 - u )
} else { u = U }
y = y + x * u
T=Tmax*y
fi=fmod(numpulses,100)
PRE[fi]=y : PRE[numpulses]=y
:PRE=1 : Istruzione non necessaria ma se ommesso allora le copie dell'oggetto successive alla prima non funzionano!
:}
: all'inizio numpulses=0 !
tspike[fi] = t
numpulses=numpulses+1
tsyn = t
}
net_send(Cdur, nspike)
}
if (flag == nspike) {
tzero = t
T = 0
on = 0
}
}