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
mechanism
not-currently-used
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VClamp.omod *
                            
TITLE HH channel (activity dependant attenuation)
: Mel-modified Hodgkin - Huxley conductances (after Ojvind et al.)
: Brannon-added attenuation 
: Poirazi-modified Kdr and Na threshold/time constants to make it more stable





NEURON {
	SUFFIX hha
	USEION na READ ena WRITE ina
	USEION k READ ek WRITE ik
	NONSPECIFIC_CURRENT il
	RANGE gnabar, gkbar, gl, el
	RANGE ar2, vhalfs
	RANGE inf, fac, tau
	RANGE taus
	RANGE W
	GLOBAL taumin
}

UNITS {
	(mA) = (milliamp)
	(mV) = (millivolt)
}

INDEPENDENT {t FROM 0 TO 1 WITH 1 (ms)}

PARAMETER {
        a0r=0.0003    (ms)
        b0r=0.0003    (ms)

        zetar=12    (1)
	zetas=12   (1)
	gms=0.2    (1)
        gmr=0.2   (1)

	a0s=0.0003 (ms)
	vvs  = 2 (mV)
        vhalfr=-60   (mV)

	v (mV)
	W = 0.016 (/mV)    : this 1/61.5 mV. see the paper

	celsius = 34	(degC)
	dt (ms)
:	gnabar=.20 (mho/cm2)
:	gkbar=.12 (mho/cm2)
:	gl=.0001 (mho/cm2)
        gnabar=0 (mho/cm2)
	gkbar=0 (mho/cm2)
	gl=0 (mho/cm2)
	ena = 60 (mV)
	ek = -77 (mV)
	el = -70.0 (mV)	: steady state at v = -65 mV
	ar2 = 1.0

	b=0

	taumin=3

}

STATE {
	m h n l s
}

ASSIGNED {
	ina (mA/cm2)
	ik (mA/cm2)
	il (mA/cm2)
	inf[5]
	fac[5]
	tau[5]
}




BREAKPOINT {
	SOLVE states
:	ina = gnabar*m*m*h*s*(v - ena)
	ina = gnabar*m*m*h*s*(v - ena)
	ik = gkbar*n*n*l*(v - ek)
	il = gl*(v - el)
}

INITIAL {
	states()
	s=1
        l=1
        h=1
:	ina = gnabar*m*m*h*s*(v - ena)
	ina = gnabar*m*m*h*s*(v - ena)
	ik = gkbar*n*n*l*(v - ek)
	il = gl*(v - el)
}

PROCEDURE calcg() {
	mhn(v*1(/mV))
	m = m + fac[0]*(inf[0] - m)
	h = h + fac[1]*(inf[1] - h)
	n = n + fac[2]*(inf[2] - n)
        l = l + fac[3]*(inf[3] - l)
	s = s + fac[4]*(inf[4] - s)
}	

PROCEDURE states() {	: exact when v held constant
	calcg()
	VERBATIM
	return 0;
	ENDVERBATIM
}




FUNCTION varss(v, i) {
	LOCAL max, min,vhalf,smooth
	if (i==0) {
		varss = 1 / (1 + exp((v + 40)/(-3))) :Na activation
:		varss = 1 / (1 + exp((v + 39)/(-2.7))) :Na activation
	}
	else if (i==1) {
:		varss = 1 / (1 + exp((v + 45)/(3))) :Na inactivation
		varss = 1 / (1 + exp((v + 45)/(2.5))) :Na inactivation
	}
	else if (i==2) {	
:		varss = 1 / (1 + exp((v + 42)/(-2))) :K activation
		varss = 1 / (1 + exp((v + 40)/(-2))) :K activation
         }
	else if (i==3) {	
		varss = (1 + 1.6/(1+exp((v + 65)/12)))/2.52

	} else {
                :"s" activation system - Migliore 96 model
		max=1
		min=0
		smooth=2
		vhalf=-60
		varss =     alpv(v,vhalfr)
       }
}

FUNCTION alpr(v(mV)) { 
  alpr = exp(1.e-3*zetar*(v-vhalfr)*9.648e4/(8.315*(273.16+celsius))) 
}


FUNCTION alpv(v(mV),vh) {
         alpv = (1+ar2*exp((v-vh)/vvs))/(1+exp((v-vh)/vvs))

}

FUNCTION betr(v(mV)) {
  betr = exp(1.e-3*zetar*gmr*(v-vhalfr)*9.648e4/(8.315*(273.16+celsius))) }


FUNCTION vartau(v, i) {
	LOCAL alpha, beta, sum,tmp

	if (i==0) {
		vartau = 0.05  :Na activation tau
:                if ( v < -51) {
:                    vartau = (0.4/25.0)*(v +75)+0.1
:                } else if (v < 0) {
:                    vartau = (-0.3/50.0)*(v +50)+0.5
:                } else {
:                  vartau = 0.2
:                } 
	}
	else if (i==1) {
              vartau = 0.5 :Na inactivation tau
:	        vartau = (-7.0/50.0)*(v +75)+8
:                if (v > -25) {
:                  vartau = 1
:                } 
        }
	else if (i==2) {
 		vartau = 1.5     :K activation
        }
	else if (i==3) {
                vartau = 350   :K inactivation
      	} else {
	        tmp = betr(v)/(a0r+b0r*alpr(v))
	        if (tmp<taumin) {tmp=taumin}
	VERBATIM
/*	printf("%g %g\n", _lv, _ltmp);*/
	ENDVERBATIM
		vartau=tmp
       }
}	


PROCEDURE mhn(v) {LOCAL a, b :rest = -70
:	TABLE inf, fac DEPEND dt, celsius FROM -100 TO 100 WITH 200
	FROM i=0 TO 4 {
		tau[i] = vartau(v,i)
		inf[i] = varss(v,i)
		fac[i] = (1 - exp(-dt/tau[i]))
	}
:	printf("v: %g inf[3]: %g\n", v, inf[3])
}
















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