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CA1 pyramidal neuron: as a 2-layer NN and subthreshold synaptic summation (Poirazi et al 2003)

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Accession:20212
We developed a CA1 pyramidal cell model calibrated with a broad spectrum of in vitro data. Using simultaneous dendritic and somatic recordings, and combining results for two different response measures (peak vs. mean EPSP), two different stimulus formats (single shock vs. 50 Hz trains), and two different spatial integration conditions (within vs. between-branch summation), we found the cell's subthreshold responses to paired inputs are best described as a sum of nonlinear subunit responses, where the subunits correspond to different dendritic branches. In addition to suggesting a new type of experiment and providing testable predictions, our model shows how conclusions regarding synaptic arithmetic can be influenced by an array of seemingly innocuous experimental design choices.
References:
1 . Poirazi P, Brannon T, Mel BW (2003) Arithmetic of subthreshold synaptic summation in a model CA1 pyramidal cell. Neuron 37:977-87 [PubMed]
2 . Poirazi P, Brannon T, Mel BW (2003) Pyramidal neuron as two-layer neural network. Neuron 37:989-99 [PubMed]
3 . Poirazi P, Brannon T, Mel BW (2003ab-sup) Online Supplement: About the Model Neuron 37 Online:1-20
4 . Polsky A, Mel BW, Schiller J (2004) Computational subunits in thin dendrites of pyramidal cells. Nat Neurosci 7:621-7 [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:
Cell Type(s): Hippocampus CA1 pyramidal GLU cell;
Channel(s): I Na,p; I Na,t; I L high threshold; I T low threshold; I A; I K; I M; I h; I K,Ca; I Calcium;
Gap Junctions:
Receptor(s): GabaA; GabaB; NMDA; Glutamate;
Gene(s):
Transmitter(s):
Simulation Environment: NEURON;
Model Concept(s): Action Potential Initiation; Activity Patterns; Dendritic Action Potentials; Active Dendrites; Influence of Dendritic Geometry; Detailed Neuronal Models; Action Potentials; Depression; Delay;
Implementer(s): Poirazi, Panayiota [poirazi at imbb.forth.gr];
Search NeuronDB for information about:  Hippocampus CA1 pyramidal GLU cell; GabaA; GabaB; NMDA; Glutamate; I Na,p; I Na,t; I L high threshold; I T low threshold; I A; I K; I M; I h; I K,Ca; I Calcium;
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CA1_multi
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VClamp.omod *
                            
/* Created by Language version: 4.1  of 8/16/98 */
/* NOT VECTORIZED */
#include <stdio.h>
#include <math.h>
#include "scoplib.h"
#undef PI
 
#include "md1redef.h"
#include "section.h"
#include "md2redef.h"

#if METHOD3
extern int _method3;
#endif

#define exp hoc_Exp
extern double hoc_Exp();
 	/*SUPPRESS 761*/
	/*SUPPRESS 762*/
	/*SUPPRESS 763*/
	/*SUPPRESS 765*/
	 extern double *getarg();
 static double *_p; static Datum *_ppvar;
 
#define delta_t dt
#define erev _p[0]
#define thresh _p[1]
#define d _p[2]
#define s _p[3]
#define tauD _p[4]
#define tauS _p[5]
#define taug _p[6]
#define G _p[7]
#define i _p[8]
#define D _p[9]
#define S _p[10]
#define g _p[11]
#define h _p[12]
#define DD _p[13]
#define DS _p[14]
#define Dg _p[15]
#define Dh _p[16]
#define firing _p[17]
#define _nd_area  *_ppvar[0].pval
#define vpre	*_ppvar[2].pval
#define _p_vpre	_ppvar[2].pval
 
#if MAC
#if !defined(v)
#define v _mlhv
#endif
#if !defined(h)
#define h _mlhh
#endif
#endif
 static int hoc_nrnpointerindex =  2;
 /* external NEURON variables */
 extern double dt;
 extern double t;
 /* declaration of user functions */
 static double _hoc_aux();
 static double _hoc_check();
 static int _mechtype;
extern int nrn_get_mechtype();
 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(_vptr) void* _vptr; { destroy_point_process(_vptr);}
 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));
}
 static _hoc_setdata(_vptr) void* _vptr; { Prop* _prop;
 _prop = ((Point_process*)_vptr)->prop;
 _p = _prop->param; _ppvar = _prop->dparam;
 }
 /* connect user functions to hoc names */
 static IntFunc hoc_intfunc[] = {
 0,0
};
 static struct Member_func {
	char* _name; double (*_member)();} _member_func[] = {
 "loc", _hoc_loc_pnt,
 "has_loc", _hoc_has_loc,
 "get_loc", _hoc_get_loc_pnt,
 "aux", _hoc_aux,
 "check", _hoc_check,
 0, 0
};
 /* declare global and static user variables */
#define eta eta_abbott_nmda
 double eta = 0.33;
#define gamma gamma_abbott_nmda
 double gamma = 0.06;
#define mag mag_abbott_nmda
 double mag = 1;
 /* some parameters have upper and lower limits */
 static HocParmLimits _hoc_parm_limits[] = {
 0,0,0
};
 static HocParmUnits _hoc_parm_units[] = {
 "eta_abbott_nmda", "/mM",
 "mag_abbott_nmda", "mM",
 "gamma_abbott_nmda", "/mV",
 "erev", "mV",
 "thresh", "mV",
 "tauD", "ms",
 "tauS", "ms",
 "taug", "ms",
 "G", "umho",
 "g", "umho",
 "h", "umho",
 "i", "nA",
 "vpre", "mV",
 0,0
};
 static double D0 = 0;
 static double S0 = 0;
 static double g0 = 0;
 static double h0 = 0;
 static double t0 = 0;
 static double v = 0;
 /* connect global user variables to hoc */
 static DoubScal hoc_scdoub[] = {
 "eta_abbott_nmda", &eta,
 "mag_abbott_nmda", &mag,
 "gamma_abbott_nmda", &gamma,
 0,0
};
 static DoubVec hoc_vdoub[] = {
 0,0,0
};
 static double _sav_indep;
 static nrn_alloc(), nrn_init(), nrn_cur(), nrn_state();
 
static int _ode_count(), _ode_map(), _ode_spec(), _ode_matsol();
extern int nrn_cvode_;
 
#define _cvode_ieq _ppvar[3]._i
 /* connect range variables in _p that hoc is supposed to know about */
 static char *_mechanism[] = {
 "abbott_nmda",
 "erev",
 "thresh",
 "d",
 "s",
 "tauD",
 "tauS",
 "taug",
 "G",
 0,
 "i",
 0,
 "D",
 "S",
 "g",
 "h",
 0,
 "vpre",
 0};
 
static nrn_alloc(_prop)
	Prop *_prop;
{
	Prop *prop_ion, *need_memb();
	double *_p; Datum *_ppvar;
  if (nrn_point_prop_) {
	_p = nrn_point_prop_->param;
	_ppvar = nrn_point_prop_->dparam;
 }else{
 	_p = (double *)ecalloc(18, sizeof(double));
 	/*initialize range parameters*/
 	erev = 0;
 	thresh = 0.5;
 	d = 0.4;
 	s = 1;
 	tauD = 300;
 	tauS = 20000;
 	taug = 2;
 	G = 1e-11;
  }
 	_prop->param = _p;
 	_prop->param_size = 18;
  if (!nrn_point_prop_) {
 	_ppvar = (Datum *)ecalloc(4, sizeof(Datum));
  }
 	_prop->dparam = _ppvar;
 	/*connect ionic variables to this model*/
 
}
 static _initlists();
  /* some states have an absolute tolerance */
 static Symbol** _atollist;
 static HocStateTolerance _hoc_state_tol[] = {
 0,0
};
 _abbott_nmda_reg() {
	int _vectorized = 0;
  _initlists();
 	_pointtype = point_register_mech(_mechanism,
	 nrn_alloc,nrn_cur, nrn_state, nrn_init,
	 hoc_nrnpointerindex,
	 _hoc_create_pnt, _hoc_destroy_pnt, _member_func,
	 _vectorized);
 _mechtype = nrn_get_mechtype(_mechanism[0]);
 	hoc_register_cvode(_mechtype, _ode_count, _ode_map, _ode_spec, _ode_matsol);
 	hoc_register_tolerance(_mechtype, _hoc_state_tol, &_atollist);
 	hoc_register_var(hoc_scdoub, hoc_vdoub, hoc_intfunc);
 	ivoc_help("help ?1 abbott_nmda /disks/redondo/brannon/rs/bp/mechanism/abbott_nmda.mod\n");
 hoc_register_limits(_mechtype, _hoc_parm_limits);
 hoc_register_units(_mechtype, _hoc_parm_units);
 }
static int _reset;
static char *modelname = "";

static int error;
static int _ninits = 0;
static int _match_recurse=1;
static _modl_cleanup(){ _match_recurse=1;}
static aux();
static check();
 
static int _ode_spec1(), _ode_matsol1();
 static double *_temp1;
 static int _slist1[3], _dlist1[3];
 static int depression();
 
/*CVODE*/
 static int _ode_spec1 () {_reset=0;
 {
   check (  ) ;
   DD = ( 1.0 / tauD ) * ( 1.0 - D ) ;
   DS = ( 1.0 / tauS ) * ( 1.0 - S ) ;
   Dg = ( 1.0 / taug ) * ( - g ) ;
   }
 return _reset;
}
 static int _ode_matsol1() {
 check (  ) ;
 DD = DD  / (1. - dt*( (( 1.0 / tauD ))*(( ( - 1.0 ) )) )) ;
 DS = DS  / (1. - dt*( (( 1.0 / tauS ))*(( ( - 1.0 ) )) )) ;
 Dg = Dg  / (1. - dt*( (( 1.0 / taug ))*(( - 1.0 )) )) ;
}
 /*END CVODE*/
 
static int depression () {_reset=0;
 {
   check (  ) ;
   DD = ( 1.0 / tauD ) * ( 1.0 - D ) ;
   DS = ( 1.0 / tauS ) * ( 1.0 - S ) ;
   Dg = ( 1.0 / taug ) * ( - g ) ;
   }
 return _reset;}
 
static int  aux (  )  {
   h = 1.0 / ( 1.0 + eta * mag * exp ( - ( gamma * v ) ) ) ;
   i = ( g * h * ( v - erev ) ) ;
    return 0; }
 static double _hoc_aux(_vptr) void* _vptr; {
 double _r;
 	_hoc_setdata(_vptr);
 _r = 1.;
 aux (  ) ;
 return(_r);
}
 
static int  check (  )  {
   if ( firing  && ( vpre < thresh ) ) {
     firing = 0.0 ;
     }
   if ( ( vpre >= thresh )  &&  ! firing ) {
     firing = 1.0 ;
     D = d * D ;
     S = s * S ;
     g = g + G * D * S ;
     }
    return 0; }
 static double _hoc_check(_vptr) void* _vptr; {
 double _r;
 	_hoc_setdata(_vptr);
 _r = 1.;
 check (  ) ;
 return(_r);
}
 
static int _ode_count() { return 3;}
 
static int _ode_spec(_nd, _pp, _ppd) Node* _nd; double* _pp; Datum* _ppd; {
	_p = _pp; _ppvar = _ppd; v = _nd->_v;
  _ode_spec1();
 }
 
static int _ode_map(_ieq, _pv, _pvdot, _pp, _ppd, _atol) int _ieq; double** _pv, **_pvdot, *_pp, *_atol; Datum* _ppd; {
	int _i; _p = _pp; _ppvar = _ppd;
	_cvode_ieq = _ieq;
	for (_i=0; _i < 3; ++_i) {
		_pv[_i] = _pp + _slist1[_i];  _pvdot[_i] = _pp + _dlist1[_i];
		_cvode_abstol(_atollist, _atol, _i);
	}
 }
 
static int _ode_matsol(_nd, _pp, _ppd) Node* _nd; double* _pp; Datum* _ppd; {
	_p = _pp; _ppvar = _ppd; v = _nd->_v;
 _ode_matsol1();
 }

static initmodel() {
  int _i; double _save;_ninits++;
 _save = t;
 t = t0;
{
  D = D0;
  S = S0;
  g = g0;
  h = h0;
 {
   firing = 0.0 ;
   D = 1.0 ;
   S = 1.0 ;
   g = 0.0 ;
   }
  _sav_indep = t; t = _save;

}
}

static nrn_init(_nd, _pp, _ppd) Node *_nd; double *_pp; Datum* _ppd; {
 double _v;
 _p = _pp; _ppvar = _ppd;
 _v = _nd->_v;
 v = _v;
 initmodel();
}

static double _nrn_current(_v) double _v;{double _current=0.;v=_v;{ {
   aux (  ) ;
   }
 _current += i;

} return _current;
}

static nrn_cur(_nd, _pp, _ppd) Node *_nd;double *_pp; Datum* _ppd;{
 double _g, _rhs, _v, *_pdiag;
 _p = _pp; _ppvar = _ppd;
_pdiag = &_nd->_d;
 _v = _nd->_v;
 _g = _nrn_current(_v + .001);
 	{ _rhs = _nrn_current(_v);
 	}
 _g = (_g - _rhs)/.001;
 _g *=  1.e2/(_nd_area);
 _rhs *= 1.e2/(_nd_area);
 _rhs -= _g*_v;
 *_pdiag += _g;
 _nd->_rhs -= _rhs;
 
}

static nrn_state(_nd, _pp, _ppd) Node *_nd; double *_pp; Datum* _ppd; {
 double _break, _save;
 double _v;
 _p = _pp; _ppvar = _ppd;
 _v = _nd->_v;
 _break = t + .5*dt; _save = t; delta_t = dt;
 v=_v;
{
 { {
 for (; t < _break; t += delta_t) {
 error =  euler(_ninits, 3, _slist1, _dlist1, _p, &t, delta_t, depression, &_temp1);
 if(error){fprintf(stderr,"at line 66 in file abbott_nmda.mod:\n   SOLVE depression METHOD euler : so now we're clinical psychiatrists\n"); nrn_complain(_p); abort_run(error);}
 
}}
 t = _save;
  depression();
 }
}
}

static terminal(){}

static _initlists() {
 int _i; static int _first = 1;
  if (!_first) return;
 _slist1[0] = &(D) - _p;  _dlist1[0] = &(DD) - _p;
 _slist1[1] = &(S) - _p;  _dlist1[1] = &(DS) - _p;
 _slist1[2] = &(g) - _p;  _dlist1[2] = &(Dg) - _p;
_first = 0;
}

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