Fast Spiking Basket cells (Tzilivaki et al 2019)

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Accession:237595
"Interneurons are critical for the proper functioning of neural circuits. While often morphologically complex, dendritic integration and its role in neuronal output have been ignored for decades, treating interneurons as linear point neurons. Exciting new findings suggest that interneuron dendrites support complex, nonlinear computations: sublinear integration of EPSPs in the cerebellum, coupled to supralinear calcium accumulations and supralinear voltage integration in the hippocampus. These findings challenge the point neuron dogma and call for a new theory of interneuron arithmetic. Using detailed, biophysically constrained models, we predict that dendrites of FS basket cells in both hippocampus and mPFC come in two flavors: supralinear, supporting local sodium spikes within large-volume branches and sublinear, in small-volume branches. Synaptic activation of varying sets of these dendrites leads to somatic firing variability that cannot be explained by the point neuron reduction. Instead, a 2-stage Artificial Neural Network (ANN), with both sub- and supralinear hidden nodes, captures most of the variance. We propose that FS basket cells have substantially expanded computational capabilities sub-served by their non-linear dendrites and act as a 2-layer ANN."
Reference:
1 . Tzilivaki A, Kastellakis G, Poirazi P (2019) Challenging the point neuron dogma: FS basket cells as 2-stage nonlinear integrators Nature Communications 10(1):3664 [PubMed]
Model Information (Click on a link to find other models with that property)
Model Type: Neuron or other electrically excitable cell;
Brain Region(s)/Organism: Hippocampus; Prefrontal cortex (PFC);
Cell Type(s): Hippocampus CA3 interneuron basket GABA cell; Neocortex layer 5 interneuron;
Channel(s):
Gap Junctions:
Receptor(s):
Gene(s):
Transmitter(s):
Simulation Environment: NEURON; MATLAB; Python;
Model Concept(s): Active Dendrites; Detailed Neuronal Models;
Implementer(s): Tzilivaki, Alexandra [alexandra.tzilivaki at charite.de]; Kastellakis, George [gkastel at gmail.com];
Search NeuronDB for information about:  Hippocampus CA3 interneuron basket GABA cell;
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TzilivakiEtal_FSBCs_model
Multicompartmental_Biophysical_models
mechanism
x86_64
.libs
ampa.mod *
ampain.mod *
cadyn.mod *
cadynin.mod *
cal.mod *
calc.mod *
calcb.mod *
can.mod *
cancr.mod *
canin.mod *
car.mod *
cat.mod *
catcb.mod *
cpampain.mod *
gabaa.mod *
gabaain.mod *
gabab.mod *
h.mod *
hcb.mod *
hin.mod *
ican.mod *
iccb.mod *
iccr.mod *
icin.mod *
iks.mod *
ikscb.mod *
ikscr.mod *
iksin.mod *
kadist.mod *
kadistcr.mod *
kadistin.mod *
kaprox.mod *
kaproxcb.mod *
kaproxin.mod *
kca.mod *
kcain.mod *
kct.mod *
kctin.mod *
kdr.mod *
kdrcb.mod *
kdrcr.mod *
kdrin.mod *
naf.mod *
nafcb.mod *
nafcr.mod *
nafin.mod *
nafx.mod *
nap.mod *
netstim.mod *
NMDA.mod *
NMDAIN.mod *
sinclamp.mod *
vecstim.mod *
ampa.c
ampa.lo
ampain.c
ampain.lo
cadyn.c
cadyn.lo
cadynin.c
cadynin.lo
cal.c
cal.lo
calc.c
calc.lo
calcb.c
calcb.lo
can.c
can.lo
cancr.c
cancr.lo
canin.c
canin.lo
car.c
car.lo
cat.c
cat.lo
catcb.c
catcb.lo
cpampain.c
cpampain.lo
gabaa.c
gabaa.lo
gabaain.c
gabaain.lo
gabab.c
gabab.lo
h.c
h.lo
hcb.c
hcb.lo
hin.c
hin.lo
ican.c
ican.lo
iccb.c
iccb.lo
iccr.c
iccr.lo
icin.c
icin.lo
iks.c
iks.lo
ikscb.c
ikscb.lo
ikscr.c
ikscr.lo
iksin.c
iksin.lo
kadist.c
kadist.lo
kadistcr.c
kadistcr.lo
kadistin.c
kadistin.lo
kaprox.c
kaprox.lo
kaproxcb.c
kaproxcb.lo
kaproxin.c
kaproxin.lo
kca.c
kca.lo
kcain.c
kcain.lo
kct.c
kct.lo
kctin.c
kctin.lo
kdr.c
kdr.lo
kdrcb.c
kdrcb.lo
kdrcr.c
kdrcr.lo
kdrin.c
kdrin.lo
libnrnmech.la *
mod_func.c
mod_func.lo *
naf.c
naf.lo
nafcb.c
nafcb.lo
nafcr.c
nafcr.lo
nafin.c
nafin.lo
nafx.c
nafx.lo
nap.c
nap.lo
netstim.c
netstim.lo
NMDA.c
NMDA.lo
NMDAIN.c
NMDAIN.lo
sinclamp.c
sinclamp.lo
special
vecstim.c
vecstim.lo
                            
/* Created by Language version: 6.2.0 */
/* VECTORIZED */
#define NRN_VECTORIZED 1
#include <stdio.h>
#include <stdlib.h>
#include <math.h>
#include "scoplib_ansi.h"
#undef PI
#define nil 0
#include "md1redef.h"
#include "section.h"
#include "nrniv_mf.h"
#include "md2redef.h"
 
#if METHOD3
extern int _method3;
#endif

#if !NRNGPU
#undef exp
#define exp hoc_Exp
extern double hoc_Exp(double);
#endif
 
#define nrn_init _nrn_init__NMDAIN
#define _nrn_initial _nrn_initial__NMDAIN
#define nrn_cur _nrn_cur__NMDAIN
#define _nrn_current _nrn_current__NMDAIN
#define nrn_jacob _nrn_jacob__NMDAIN
#define nrn_state _nrn_state__NMDAIN
#define _net_receive _net_receive__NMDAIN 
#define release release__NMDAIN 
 
#define _threadargscomma_ _p, _ppvar, _thread, _nt,
#define _threadargsprotocomma_ double* _p, Datum* _ppvar, Datum* _thread, _NrnThread* _nt,
#define _threadargs_ _p, _ppvar, _thread, _nt
#define _threadargsproto_ double* _p, Datum* _ppvar, Datum* _thread, _NrnThread* _nt
 	/*SUPPRESS 761*/
	/*SUPPRESS 762*/
	/*SUPPRESS 763*/
	/*SUPPRESS 765*/
	 extern double *getarg();
 /* Thread safe. No static _p or _ppvar. */
 
#define t _nt->_t
#define dt _nt->_dt
#define Alpha _p[0]
#define Beta _p[1]
#define e _p[2]
#define iNMDA _p[3]
#define g _p[4]
#define gmax _p[5]
#define ica _p[6]
#define Ron _p[7]
#define Roff _p[8]
#define Rinf _p[9]
#define Rtau _p[10]
#define synon _p[11]
#define B _p[12]
#define DRon _p[13]
#define DRoff _p[14]
#define v _p[15]
#define _g _p[16]
#define _tsav _p[17]
#define _nd_area  *_ppvar[0]._pval
#define _ion_ica	*_ppvar[2]._pval
#define _ion_dicadv	*_ppvar[3]._pval
 
#if MAC
#if !defined(v)
#define v _mlhv
#endif
#if !defined(h)
#define h _mlhh
#endif
#endif
 
#if defined(__cplusplus)
extern "C" {
#endif
 static int hoc_nrnpointerindex =  -1;
 static Datum* _extcall_thread;
 static Prop* _extcall_prop;
 /* external NEURON variables */
 /* declaration of user functions */
 static double _hoc_mgblock();
 static int _mechtype;
extern void _nrn_cacheloop_reg(int, int);
extern void hoc_register_prop_size(int, int, int);
extern void hoc_register_limits(int, HocParmLimits*);
extern void hoc_register_units(int, HocParmUnits*);
extern void nrn_promote(Prop*, int, int);
extern Memb_func* memb_func;
 extern Prop* nrn_point_prop_;
 static int _pointtype;
 static void* _hoc_create_pnt(_ho) Object* _ho; { void* create_point_process();
 return create_point_process(_pointtype, _ho);
}
 static void _hoc_destroy_pnt();
 static double _hoc_loc_pnt(_vptr) void* _vptr; {double loc_point_process();
 return loc_point_process(_pointtype, _vptr);
}
 static double _hoc_has_loc(_vptr) void* _vptr; {double has_loc_point();
 return has_loc_point(_vptr);
}
 static double _hoc_get_loc_pnt(_vptr)void* _vptr; {
 double get_loc_point_process(); return (get_loc_point_process(_vptr));
}
 extern void _nrn_setdata_reg(int, void(*)(Prop*));
 static void _setdata(Prop* _prop) {
 _extcall_prop = _prop;
 }
 static void _hoc_setdata(void* _vptr) { Prop* _prop;
 _prop = ((Point_process*)_vptr)->_prop;
   _setdata(_prop);
 }
 /* connect user functions to hoc names */
 static VoidFunc hoc_intfunc[] = {
 0,0
};
 static Member_func _member_func[] = {
 "loc", _hoc_loc_pnt,
 "has_loc", _hoc_has_loc,
 "get_loc", _hoc_get_loc_pnt,
 "mgblock", _hoc_mgblock,
 0, 0
};
#define _f_mgblock _f_mgblock_NMDAIN
#define mgblock mgblock_NMDAIN
 extern double _f_mgblock( _threadargsprotocomma_ double );
 extern double mgblock( _threadargsprotocomma_ double );
 
static void _check_mgblock(double*, Datum*, Datum*, _NrnThread*); 
static void _check_table_thread(double* _p, Datum* _ppvar, Datum* _thread, _NrnThread* _nt, int _type) {
   _check_mgblock(_p, _ppvar, _thread, _nt);
 }
 /* declare global and static user variables */
#define Cmax Cmax_NMDAIN
 double Cmax = 1;
#define Cdur Cdur_NMDAIN
 double Cdur = 1;
#define mg mg_NMDAIN
 double mg = 1;
#define usetable usetable_NMDAIN
 double usetable = 1;
 /* some parameters have upper and lower limits */
 static HocParmLimits _hoc_parm_limits[] = {
 "usetable_NMDAIN", 0, 1,
 0,0,0
};
 static HocParmUnits _hoc_parm_units[] = {
 "Cmax_NMDAIN", "mM",
 "Cdur_NMDAIN", "ms",
 "mg_NMDAIN", "mM",
 "Alpha", "/ms",
 "Beta", "/ms",
 "e", "mV",
 "iNMDA", "nA",
 "g", "umho",
 0,0
};
 static double Roff0 = 0;
 static double Ron0 = 0;
 static double delta_t = 0.01;
 /* connect global user variables to hoc */
 static DoubScal hoc_scdoub[] = {
 "Cmax_NMDAIN", &Cmax_NMDAIN,
 "Cdur_NMDAIN", &Cdur_NMDAIN,
 "mg_NMDAIN", &mg_NMDAIN,
 "usetable_NMDAIN", &usetable_NMDAIN,
 0,0
};
 static DoubVec hoc_vdoub[] = {
 0,0,0
};
 static double _sav_indep;
 static void nrn_alloc(Prop*);
static void  nrn_init(_NrnThread*, _Memb_list*, int);
static void nrn_state(_NrnThread*, _Memb_list*, int);
 static void nrn_cur(_NrnThread*, _Memb_list*, int);
static void  nrn_jacob(_NrnThread*, _Memb_list*, int);
 static void _hoc_destroy_pnt(_vptr) void* _vptr; {
   destroy_point_process(_vptr);
}
 
static int _ode_count(int);
static void _ode_map(int, double**, double**, double*, Datum*, double*, int);
static void _ode_spec(_NrnThread*, _Memb_list*, int);
static void _ode_matsol(_NrnThread*, _Memb_list*, int);
 
#define _cvode_ieq _ppvar[5]._i
 static void _ode_matsol_instance1(_threadargsproto_);
 /* connect range variables in _p that hoc is supposed to know about */
 static const char *_mechanism[] = {
 "6.2.0",
"NMDAIN",
 "Alpha",
 "Beta",
 "e",
 0,
 "iNMDA",
 "g",
 "gmax",
 "ica",
 0,
 "Ron",
 "Roff",
 0,
 0};
 static Symbol* _ca_sym;
 
extern Prop* need_memb(Symbol*);

static void nrn_alloc(Prop* _prop) {
	Prop *prop_ion;
	double *_p; Datum *_ppvar;
  if (nrn_point_prop_) {
	_prop->_alloc_seq = nrn_point_prop_->_alloc_seq;
	_p = nrn_point_prop_->param;
	_ppvar = nrn_point_prop_->dparam;
 }else{
 	_p = nrn_prop_data_alloc(_mechtype, 18, _prop);
 	/*initialize range parameters*/
 	Alpha = 4;
 	Beta = 0.02;
 	e = 0;
  }
 	_prop->param = _p;
 	_prop->param_size = 18;
  if (!nrn_point_prop_) {
 	_ppvar = nrn_prop_datum_alloc(_mechtype, 6, _prop);
  }
 	_prop->dparam = _ppvar;
 	/*connect ionic variables to this model*/
 prop_ion = need_memb(_ca_sym);
 	_ppvar[2]._pval = &prop_ion->param[3]; /* ica */
 	_ppvar[3]._pval = &prop_ion->param[4]; /* _ion_dicadv */
 
}
 static void _initlists();
  /* some states have an absolute tolerance */
 static Symbol** _atollist;
 static HocStateTolerance _hoc_state_tol[] = {
 0,0
};
 
#define _tqitem &(_ppvar[4]._pvoid)
 static void _net_receive(Point_process*, double*, double);
 static void _update_ion_pointer(Datum*);
 extern Symbol* hoc_lookup(const char*);
extern void _nrn_thread_reg(int, int, void(*)(Datum*));
extern void _nrn_thread_table_reg(int, void(*)(double*, Datum*, Datum*, _NrnThread*, int));
extern void hoc_register_tolerance(int, HocStateTolerance*, Symbol***);
extern void _cvode_abstol( Symbol**, double*, int);

 void _NMDAIN_reg() {
	int _vectorized = 1;
  _initlists();
 	ion_reg("ca", -10000.);
 	_ca_sym = hoc_lookup("ca_ion");
 	_pointtype = point_register_mech(_mechanism,
	 nrn_alloc,nrn_cur, nrn_jacob, nrn_state, nrn_init,
	 hoc_nrnpointerindex, 1,
	 _hoc_create_pnt, _hoc_destroy_pnt, _member_func);
 _mechtype = nrn_get_mechtype(_mechanism[1]);
     _nrn_setdata_reg(_mechtype, _setdata);
     _nrn_thread_reg(_mechtype, 2, _update_ion_pointer);
     _nrn_thread_table_reg(_mechtype, _check_table_thread);
  hoc_register_prop_size(_mechtype, 18, 6);
  hoc_register_dparam_semantics(_mechtype, 0, "area");
  hoc_register_dparam_semantics(_mechtype, 1, "pntproc");
  hoc_register_dparam_semantics(_mechtype, 2, "ca_ion");
  hoc_register_dparam_semantics(_mechtype, 3, "ca_ion");
  hoc_register_dparam_semantics(_mechtype, 4, "netsend");
  hoc_register_dparam_semantics(_mechtype, 5, "cvodeieq");
 	hoc_register_cvode(_mechtype, _ode_count, _ode_map, _ode_spec, _ode_matsol);
 	hoc_register_tolerance(_mechtype, _hoc_state_tol, &_atollist);
 pnt_receive[_mechtype] = _net_receive;
 pnt_receive_size[_mechtype] = 5;
 	hoc_register_var(hoc_scdoub, hoc_vdoub, hoc_intfunc);
 	ivoc_help("help ?1 NMDAIN /home/cluster/aleka/MainPath/Desktop/FSBC_model/Multicompartmental_Biophysical_models/mechanism/x86_64/NMDAIN.mod\n");
 hoc_register_limits(_mechtype, _hoc_parm_limits);
 hoc_register_units(_mechtype, _hoc_parm_units);
 }
 static double *_t_mgblock;
static int _reset;
static char *modelname = "simple NMDA receptors";

static int error;
static int _ninits = 0;
static int _match_recurse=1;
static void _modl_cleanup(){ _match_recurse=1;}
 
static int _ode_spec1(_threadargsproto_);
/*static int _ode_matsol1(_threadargsproto_);*/
 static double _n_mgblock(_threadargsprotocomma_ double _lv);
 static int _slist1[2], _dlist1[2];
 static int release(_threadargsproto_);
 
/*CVODE*/
 static int _ode_spec1 (double* _p, Datum* _ppvar, Datum* _thread, _NrnThread* _nt) {int _reset = 0; {
   DRon = ( synon * Rinf - Ron ) / Rtau ;
   DRoff = - Beta * Roff ;
   }
 return _reset;
}
 static int _ode_matsol1 (double* _p, Datum* _ppvar, Datum* _thread, _NrnThread* _nt) {
 DRon = DRon  / (1. - dt*( ( ( ( - 1.0 ) ) ) / Rtau )) ;
 DRoff = DRoff  / (1. - dt*( ( - Beta )*( 1.0 ) )) ;
 return 0;
}
 /*END CVODE*/
 static int release (double* _p, Datum* _ppvar, Datum* _thread, _NrnThread* _nt) { {
    Ron = Ron + (1. - exp(dt*(( ( ( - 1.0 ) ) ) / Rtau)))*(- ( ( ( ( synon )*( Rinf ) ) ) / Rtau ) / ( ( ( ( - 1.0 ) ) ) / Rtau ) - Ron) ;
    Roff = Roff + (1. - exp(dt*(( - Beta )*( 1.0 ))))*(- ( 0.0 ) / ( ( - Beta )*( 1.0 ) ) - Roff) ;
   }
  return 0;
}
 static double _mfac_mgblock, _tmin_mgblock;
  static void _check_mgblock(double* _p, Datum* _ppvar, Datum* _thread, _NrnThread* _nt) {
  static int _maktable=1; int _i, _j, _ix = 0;
  double _xi, _tmax;
  static double _sav_mg;
  if (!usetable) {return;}
  if (_sav_mg != mg) { _maktable = 1;}
  if (_maktable) { double _x, _dx; _maktable=0;
   _tmin_mgblock =  - 140.0 ;
   _tmax =  80.0 ;
   _dx = (_tmax - _tmin_mgblock)/1000.; _mfac_mgblock = 1./_dx;
   for (_i=0, _x=_tmin_mgblock; _i < 1001; _x += _dx, _i++) {
    _t_mgblock[_i] = _f_mgblock(_p, _ppvar, _thread, _nt, _x);
   }
   _sav_mg = mg;
  }
 }

 double mgblock(double* _p, Datum* _ppvar, Datum* _thread, _NrnThread* _nt, double _lv) { 
#if 0
_check_mgblock(_p, _ppvar, _thread, _nt);
#endif
 return _n_mgblock(_p, _ppvar, _thread, _nt, _lv);
 }

 static double _n_mgblock(double* _p, Datum* _ppvar, Datum* _thread, _NrnThread* _nt, double _lv){ int _i, _j;
 double _xi, _theta;
 if (!usetable) {
 return _f_mgblock(_p, _ppvar, _thread, _nt, _lv); 
}
 _xi = _mfac_mgblock * (_lv - _tmin_mgblock);
 if (isnan(_xi)) {
  return _xi; }
 if (_xi <= 0.) {
 return _t_mgblock[0];
 }
 if (_xi >= 1000.) {
 return _t_mgblock[1000];
 }
 _i = (int) _xi;
 return _t_mgblock[_i] + (_xi - (double)_i)*(_t_mgblock[_i+1] - _t_mgblock[_i]);
 }

 
double _f_mgblock ( _threadargsprotocomma_ double _lv ) {
   double _lmgblock;
 _lmgblock = 1.0 / ( 1.0 + exp ( 0.072 * - _lv ) * ( mg / 3.57 ) ) ;
   
return _lmgblock;
 }
 
static double _hoc_mgblock(void* _vptr) {
 double _r;
   double* _p; Datum* _ppvar; Datum* _thread; _NrnThread* _nt;
   _p = ((Point_process*)_vptr)->_prop->param;
  _ppvar = ((Point_process*)_vptr)->_prop->dparam;
  _thread = _extcall_thread;
  _nt = (_NrnThread*)((Point_process*)_vptr)->_vnt;
 
#if 1
 _check_mgblock(_p, _ppvar, _thread, _nt);
#endif
 _r =  mgblock ( _p, _ppvar, _thread, _nt, *getarg(1) );
 return(_r);
}
 
static void _net_receive (_pnt, _args, _lflag) Point_process* _pnt; double* _args; double _lflag; 
{  double* _p; Datum* _ppvar; Datum* _thread; _NrnThread* _nt;
   _thread = (Datum*)0; _nt = (_NrnThread*)_pnt->_vnt;   _p = _pnt->_prop->param; _ppvar = _pnt->_prop->dparam;
  if (_tsav > t){ extern char* hoc_object_name(); hoc_execerror(hoc_object_name(_pnt->ob), ":Event arrived out of order. Must call ParallelContext.set_maxstep AFTER assigning minimum NetCon.delay");}
 _tsav = t;   if (_lflag == 1. ) {*(_tqitem) = 0;}
 {
   if ( _lflag  == 0.0 ) {
     _args[2] = _args[2] + 1.0 ;
     if (  ! _args[1] ) {
       _args[3] = _args[3] * exp ( - Beta * ( t - _args[4] ) ) ;
       _args[4] = t ;
       _args[1] = 1.0 ;
       synon = synon + _args[0] ;
           if (nrn_netrec_state_adjust && !cvode_active_){
    /* discon state adjustment for cnexp case (rate uses no local variable) */
    double __state = Ron;
    double __primary = (Ron + _args[3] ) - __state;
     __primary += ( 1. - exp( 0.5*dt*( ( ( ( - 1.0 ) ) ) / Rtau ) ) )*( - ( ( ( ( synon )*( Rinf ) ) ) / Rtau ) / ( ( ( ( - 1.0 ) ) ) / Rtau ) - __primary );
    Ron += __primary;
  } else {
 Ron = Ron + _args[3]  ;
         }
     if (nrn_netrec_state_adjust && !cvode_active_){
    /* discon state adjustment for cnexp case (rate uses no local variable) */
    double __state = Roff;
    double __primary = (Roff - _args[3] ) - __state;
     __primary += ( 1. - exp( 0.5*dt*( ( - Beta )*( 1.0 ) ) ) )*( - ( 0.0 ) / ( ( - Beta )*( 1.0 ) ) - __primary );
    Roff += __primary;
  } else {
 Roff = Roff - _args[3]  ;
         }
 }
     net_send ( _tqitem, _args, _pnt, t +  Cdur , _args[2] ) ;
     }
   if ( _lflag  == _args[2] ) {
     _args[3] = _args[0] * Rinf + ( _args[3] - _args[0] * Rinf ) * exp ( - ( t - _args[4] ) / Rtau ) ;
     _args[4] = t ;
     synon = synon - _args[0] ;
         if (nrn_netrec_state_adjust && !cvode_active_){
    /* discon state adjustment for cnexp case (rate uses no local variable) */
    double __state = Ron;
    double __primary = (Ron - _args[3] ) - __state;
     __primary += ( 1. - exp( 0.5*dt*( ( ( ( - 1.0 ) ) ) / Rtau ) ) )*( - ( ( ( ( synon )*( Rinf ) ) ) / Rtau ) / ( ( ( ( - 1.0 ) ) ) / Rtau ) - __primary );
    Ron += __primary;
  } else {
 Ron = Ron - _args[3]  ;
       }
     if (nrn_netrec_state_adjust && !cvode_active_){
    /* discon state adjustment for cnexp case (rate uses no local variable) */
    double __state = Roff;
    double __primary = (Roff + _args[3] ) - __state;
     __primary += ( 1. - exp( 0.5*dt*( ( - Beta )*( 1.0 ) ) ) )*( - ( 0.0 ) / ( ( - Beta )*( 1.0 ) ) - __primary );
    Roff += __primary;
  } else {
 Roff = Roff + _args[3]  ;
       }
 _args[1] = 0.0 ;
     }
   gmax = _args[0] ;
   } }
 
static int _ode_count(int _type){ return 2;}
 
static void _ode_spec(_NrnThread* _nt, _Memb_list* _ml, int _type) {
   double* _p; Datum* _ppvar; Datum* _thread;
   Node* _nd; double _v; int _iml, _cntml;
  _cntml = _ml->_nodecount;
  _thread = _ml->_thread;
  for (_iml = 0; _iml < _cntml; ++_iml) {
    _p = _ml->_data[_iml]; _ppvar = _ml->_pdata[_iml];
    _nd = _ml->_nodelist[_iml];
    v = NODEV(_nd);
     _ode_spec1 (_p, _ppvar, _thread, _nt);
  }}
 
static void _ode_map(int _ieq, double** _pv, double** _pvdot, double* _pp, Datum* _ppd, double* _atol, int _type) { 
	double* _p; Datum* _ppvar;
 	int _i; _p = _pp; _ppvar = _ppd;
	_cvode_ieq = _ieq;
	for (_i=0; _i < 2; ++_i) {
		_pv[_i] = _pp + _slist1[_i];  _pvdot[_i] = _pp + _dlist1[_i];
		_cvode_abstol(_atollist, _atol, _i);
	}
 }
 
static void _ode_matsol_instance1(_threadargsproto_) {
 _ode_matsol1 (_p, _ppvar, _thread, _nt);
 }
 
static void _ode_matsol(_NrnThread* _nt, _Memb_list* _ml, int _type) {
   double* _p; Datum* _ppvar; Datum* _thread;
   Node* _nd; double _v; int _iml, _cntml;
  _cntml = _ml->_nodecount;
  _thread = _ml->_thread;
  for (_iml = 0; _iml < _cntml; ++_iml) {
    _p = _ml->_data[_iml]; _ppvar = _ml->_pdata[_iml];
    _nd = _ml->_nodelist[_iml];
    v = NODEV(_nd);
 _ode_matsol_instance1(_threadargs_);
 }}
 extern void nrn_update_ion_pointer(Symbol*, Datum*, int, int);
 static void _update_ion_pointer(Datum* _ppvar) {
   nrn_update_ion_pointer(_ca_sym, _ppvar, 2, 3);
   nrn_update_ion_pointer(_ca_sym, _ppvar, 3, 4);
 }

static void initmodel(double* _p, Datum* _ppvar, Datum* _thread, _NrnThread* _nt) {
  int _i; double _save;{
  Roff = Roff0;
  Ron = Ron0;
 {
   Rinf = Cmax * Alpha / ( Cmax * Alpha + Beta ) ;
   Rtau = 1.0 / ( Cmax * Alpha + Beta ) ;
   synon = 0.0 ;
   }
 
}
}

static void nrn_init(_NrnThread* _nt, _Memb_list* _ml, int _type){
double* _p; Datum* _ppvar; Datum* _thread;
Node *_nd; double _v; int* _ni; int _iml, _cntml;
#if CACHEVEC
    _ni = _ml->_nodeindices;
#endif
_cntml = _ml->_nodecount;
_thread = _ml->_thread;
for (_iml = 0; _iml < _cntml; ++_iml) {
 _p = _ml->_data[_iml]; _ppvar = _ml->_pdata[_iml];

#if 0
 _check_mgblock(_p, _ppvar, _thread, _nt);
#endif
 _tsav = -1e20;
#if CACHEVEC
  if (use_cachevec) {
    _v = VEC_V(_ni[_iml]);
  }else
#endif
  {
    _nd = _ml->_nodelist[_iml];
    _v = NODEV(_nd);
  }
 v = _v;
 initmodel(_p, _ppvar, _thread, _nt);
 }
}

static double _nrn_current(double* _p, Datum* _ppvar, Datum* _thread, _NrnThread* _nt, double _v){double _current=0.;v=_v;{ {
   B = mgblock ( _threadargscomma_ v ) ;
   g = ( Ron + Roff ) * 1.0 * B ;
   iNMDA = g * ( v - e ) ;
   ica = 7.0 * iNMDA / 10.0 ;
   iNMDA = 3.0 * iNMDA / 10.0 ;
   }
 _current += ica;
 _current += iNMDA;

} return _current;
}

static void nrn_cur(_NrnThread* _nt, _Memb_list* _ml, int _type) {
double* _p; Datum* _ppvar; Datum* _thread;
Node *_nd; int* _ni; double _rhs, _v; int _iml, _cntml;
#if CACHEVEC
    _ni = _ml->_nodeindices;
#endif
_cntml = _ml->_nodecount;
_thread = _ml->_thread;
for (_iml = 0; _iml < _cntml; ++_iml) {
 _p = _ml->_data[_iml]; _ppvar = _ml->_pdata[_iml];
#if CACHEVEC
  if (use_cachevec) {
    _v = VEC_V(_ni[_iml]);
  }else
#endif
  {
    _nd = _ml->_nodelist[_iml];
    _v = NODEV(_nd);
  }
 _g = _nrn_current(_p, _ppvar, _thread, _nt, _v + .001);
 	{ double _dica;
  _dica = ica;
 _rhs = _nrn_current(_p, _ppvar, _thread, _nt, _v);
  _ion_dicadv += (_dica - ica)/.001 * 1.e2/ (_nd_area);
 	}
 _g = (_g - _rhs)/.001;
  _ion_ica += ica * 1.e2/ (_nd_area);
 _g *=  1.e2/(_nd_area);
 _rhs *= 1.e2/(_nd_area);
#if CACHEVEC
  if (use_cachevec) {
	VEC_RHS(_ni[_iml]) -= _rhs;
  }else
#endif
  {
	NODERHS(_nd) -= _rhs;
  }
 
}
 
}

static void nrn_jacob(_NrnThread* _nt, _Memb_list* _ml, int _type) {
double* _p; Datum* _ppvar; Datum* _thread;
Node *_nd; int* _ni; int _iml, _cntml;
#if CACHEVEC
    _ni = _ml->_nodeindices;
#endif
_cntml = _ml->_nodecount;
_thread = _ml->_thread;
for (_iml = 0; _iml < _cntml; ++_iml) {
 _p = _ml->_data[_iml];
#if CACHEVEC
  if (use_cachevec) {
	VEC_D(_ni[_iml]) += _g;
  }else
#endif
  {
     _nd = _ml->_nodelist[_iml];
	NODED(_nd) += _g;
  }
 
}
 
}

static void nrn_state(_NrnThread* _nt, _Memb_list* _ml, int _type) {
double* _p; Datum* _ppvar; Datum* _thread;
Node *_nd; double _v = 0.0; int* _ni; int _iml, _cntml;
#if CACHEVEC
    _ni = _ml->_nodeindices;
#endif
_cntml = _ml->_nodecount;
_thread = _ml->_thread;
for (_iml = 0; _iml < _cntml; ++_iml) {
 _p = _ml->_data[_iml]; _ppvar = _ml->_pdata[_iml];
 _nd = _ml->_nodelist[_iml];
#if CACHEVEC
  if (use_cachevec) {
    _v = VEC_V(_ni[_iml]);
  }else
#endif
  {
    _nd = _ml->_nodelist[_iml];
    _v = NODEV(_nd);
  }
 v=_v;
{
 {   release(_p, _ppvar, _thread, _nt);
  } }}

}

static void terminal(){}

static void _initlists(){
 double _x; double* _p = &_x;
 int _i; static int _first = 1;
  if (!_first) return;
 _slist1[0] = &(Ron) - _p;  _dlist1[0] = &(DRon) - _p;
 _slist1[1] = &(Roff) - _p;  _dlist1[1] = &(DRoff) - _p;
   _t_mgblock = makevector(1001*sizeof(double));
_first = 0;
}

#if defined(__cplusplus)
} /* extern "C" */
#endif

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