: $Id: sns.inc,v 1.29 1997/03/25 00:05:44 billl Exp $ COMMENT USAGE: for most receptors ***************************************************************************** NEURON { POINT_PROCESS NAME } PARAMETER { Cdur = 1.08 (ms) : transmitter duration (rising phase) Alpha = 1 (/ms mM) : forward (binding) rate Beta = 0.02 (/ms) : backward (unbinding) rate Erev = -80 (mV) : reversal potential Deadtime = 1 (ms) : mimimum time between release events GMAX = 1 (umho) : reference conductance DELAY = 0 (ms) } INCLUDE "sns.inc" ***************************************************************************** USAGE: for NMDA receptor ***************************************************************************** NEURON{ POINT_PROCESS NMDA RANGE B } PARAMETER { mg = 1. (mM) : external magnesium concentration Cdur = 1. (ms) : transmitter duration (rising phase) Alpha = 4. (/ms mM) : forward (binding) rate Beta = 0.0067 (/ms) : backward (unbinding) rate 1/150 Erev = 0. (mV) : reversal potential Deadtime = 1 (ms) : mimimum time between release events GMAX = 1 (umho) : reference conductance DELAY = 0 : axonal delay } ASSIGNED { B } INCLUDE "sns.inc" : EXTRA BREAKPOINT MUST BE BELOW THE INCLUDE BREAKPOINT { rates(v) g = g * B : but don't really need to readjust conductance i = i * B : i = g*(v - Erev) } PROCEDURE rates(v(mV)) { TABLE B DEPEND mg FROM -100 TO 80 WITH 180 B = 1 / (1 + exp(0.062 (/mV) * -v) * (mg / 3.57 (mM))) } ***************************************************************************** see end for implementation comments ENDCOMMENT INDEPENDENT {t FROM 0 TO 1 WITH 1 (ms)} NEURON { NONSPECIFIC_CURRENT i RANGE R, g RANGE Ron, Roff, synon : accessible for debugging GLOBAL GMAX, DELAY, Cdur, Alpha, Beta, Erev, Deadtime, Rinf, Rtau, q10, exptemp } INCLUDE "snsarr.inc" : array management routines UNITS { (nA) = (nanoamp) (mV) = (millivolt) (umho) = (micromho) (mM) = (milli/liter) } PARAMETER { celsius q10 = 1. exptemp = 37. } ASSIGNED { v (mV) : postsynaptic voltage i (nA) : current = g*(v - Erev) g (umho) : conductance R : fraction of open channels, Ron + Roff Ron : activation state while syn's turned on Roff : activation state for decaying syns Rinf : steady state channels open Rtau (ms) : time constant of channel binding synon : number of syns turned on at a time drive : drive for ODE toward Rinf qq10 : rate speed-up due to Q10 edt : rate factor for Ron edb : decay factor for Roff edc : rate factor for increasing Rcurr dt } INITIAL { : initialize GLOBAL parameters, in case user changes one : this repeats unnecessarily for each instance Rinf = Alpha / (Alpha + Beta) qq10 = q10^((celsius-exptemp)/10.) Rtau = 1 / (Alpha + Beta) / qq10 edt = exp(-dt/Rtau) edb = exp(-Beta * dt) edc = exp(-Cdur/Rtau) drive = Rinf * (1. - edt) : initialize RANGE parameters synon = 0 R = 0 Ron = 0 Roff = 0 : do not initialize QUEU if it hasn't been allocated by init_arrays() if (nsyn > 0) { initq() } else { VERBATIM printf("WARNING nsyn==0 "); ENDVERBATIM } } BREAKPOINT { SOLVE release R = Ron + Roff g = GMAX * R i = g*(v - Erev) } PROCEDURE release() { if (nsyn>0) { : do not try accessing a queue that hasn't been allocated VERBATIM int who; QueU *pqueu; SynS *ppst; pqueu = (QueU *)((unsigned long) queu); while (t >= pqueu[(int)begsyn].time) { /* somebody spiked delay time ago */ ppst = (SynS *)((unsigned long) lpst); who = pqueu[(int)begsyn].index; /* who spiked? */ /* calculate the decay that occurred since last activity ended */ ppst[who].Rcurr *= exptable(-Beta*(t-ppst[who].last)); /* transfer the value from Roff to Ron */ Ron += ppst[who].Rcurr; Roff -= ppst[who].Rcurr; synon += ppst[who].pgm; /* weight syn by percent gmax */ ppst[who].last = t + Cdur; /* time when syn will turn off */ popqh1(Cdur); /* next (also add Cdur to value on the queu) */ } while (t >= pqueu[(int)endsyn].time) { /* somebody needs to be turned off */ ppst = (SynS *)((unsigned long) lpst); /* will do this assign twice in rare cases */ who = pqueu[(int)endsyn].index; /* who spiked? */ /* solve Rcurr forward in time till end of syn activity */ ppst[who].Rcurr = ppst[who].pgm*Rinf*(1.-edc) + ppst[who].Rcurr*edc; Ron -= ppst[who].Rcurr; Roff += ppst[who].Rcurr; /* transfer from on to off */ synon -= ppst[who].pgm; /* remove this percent of gmax */ if (synon<1e-11 && synon>-1e-11) { synon=0.; } if (synon==0. || Ron < 0.) { Ron = 0.; } popqh2(); /* next */ } /* update R */ if (synon > 0) { /* one or more synapses turned on? */ Ron = synon*drive + Ron * edt; /* drive R toward Rinf */ } Roff *= edb; /* Roff always decays toward 0 */ return 0; ENDVERBATIM } } FUNCTION getR(x) { VERBATIM { SynS pst; double rnow; double end, rinf; pst = (PSTCAST[(int)_lx]); end = pst.last; rinf = pst.pgm * Rinf; if (end < 0.) { /* not yet turned on */ rnow = 0.; } else if (t >= end - dt/2) { /* decay */ rnow = pst.Rcurr * exptable(-Beta*(t-end)); } else { /* turning on */ rnow = rinf + (pst.Rcurr - rinf)*exptable((t-(end-Cdur))/(-Rtau)); } if (pst.pgm != 0.) { _lgetR = rnow/pst.pgm; /* scale it to 1 so it looks like a state variable */ } else { _lgetR = 0.; } } ENDVERBATIM } : only gets called for negative numbers FUNCTION exptable(x) { TABLE FROM -10 TO 0 WITH 1000 if ((x > -10) && (x < 0)) { exptable = exp(x) } else { exptable = 0. } }