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 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 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
cad.mod *
cagk.mod *
cal.mod *
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car.mod *
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gabaa.mod *
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glutamate.mod *
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hha_old.mod *
hha2.mod *
kadist.mod *
kaprox.mod *
kca.mod *
km.mod *
nap.mod *
nmda.mod *
somacar.mod *
mosinit.hoc *
mosinit.hoc.old *
                            
TITLE K-A channel from Klee Ficker and Heinemann
: modified by Brannon and Yiota Poirazi (poirazi@LNC.usc.edu)
: to account for Hoffman et al 1997 proximal region kinetics
: used only in soma and sections located < 100 microns from the soma

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

}

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

PARAMETER {                       :parameters that can be entered when function is called in cell-setup
        dt              (ms)
	v               (mV)
        ek = -77        (mV)      :K reversal potential  (reset in cell-setup.hoc)
	celsius = 24	(degC)
       	gkabar = 0      (mho/cm2) :initialized conductance
        vhalfn = 11     (mV)      :activation half-potential
        vhalfl = -56    (mV) 	  :inactivation half-potential
        a0n = 0.05      (/ms)     :parameters used
        zetan = -1.5    (1)       :in calculation of
        zetal = 3       (1)       :steady state values
        gmn = 0.55      (1)       :and time constants
        gml = 1         (1)
	lmin = 2        (mS)
	nmin = 0.1      (mS)
	pw = -1         (1)
	tq = -40
	qq = 5
	q10 = 5                   :temperature sensitivity
}


NEURON {
	SUFFIX kap
	USEION k READ ek WRITE ik
        RANGE gkabar,gka
        GLOBAL ninf,linf,taul,taun,lmin
}

STATE {          :the unknown parameters to be solved in the DEs 
	n l
}
 
ASSIGNED {       :parameters needed to solve DE
	ik (mA/cm2)
        ninf
        linf      
        taul
        taun
        gka
}

INITIAL {		:initialize the following parameter using rates()
	rates(v)
	n = ninf
	l = linf
	gka = gkabar*n*l
	ik = gka*(v-ek)
}

BREAKPOINT {
	SOLVE states 
	gka = gkabar*n*l
	ik = gka*(v-ek)
}


FUNCTION alpn(v(mV)) { LOCAL zeta 
  zeta = zetan+pw/(1+exp((v-tq)/qq))
  alpn = exp(1.e-3*zeta*(v-vhalfn)*9.648e4/(8.315*(273.16+celsius))) 
}

FUNCTION betn(v(mV)) { LOCAL zeta
  zeta = zetan+pw/(1+exp((v-tq)/qq))
  betn = exp(1.e-3*zeta*gmn*(v-vhalfn)*9.648e4/(8.315*(273.16+celsius))) 
}

FUNCTION alpl(v(mV)) {
  alpl = exp(1.e-3*zetal*(v-vhalfl)*9.648e4/(8.315*(273.16+celsius))) 
}

FUNCTION betl(v(mV)) {
  betl = exp(1.e-3*zetal*gml*(v-vhalfl)*9.648e4/(8.315*(273.16+celsius))) 
}

LOCAL facn,facl
:if state_borgka is called from hoc, garbage or segmentation violation will
:result because range variables won't have correct pointer.  This is because
: only BREAKPOINT sets up the correct pointers to range variables.
PROCEDURE states() {     : exact when v held constant; integrates over dt step
        rates(v)
        n = n + facn*(ninf - n)
        l = l + facl*(linf - l)
        VERBATIM
        return 0;
        ENDVERBATIM
}

PROCEDURE rates(v (mV)) {                  :callable from hoc
        LOCAL a,qt
        qt = q10^((celsius-24)/10)         : temprature adjastment factor
        a = alpn(v)
        ninf = 1/(1 + a)                   : activation variable steady state value
        taun = betn(v)/(qt*a0n*(1+a))      : activation variable time constant
	if (taun<nmin) {taun=nmin}         : time constant not allowed to be less than nmin
        facn = (1 - exp(-dt/taun))
        a = alpl(v)
        linf = 1/(1+ a)                    : inactivation variable steady state value
	taul = 0.26*(v+50)                 : inactivation variable time constant
	if (taul<lmin) {taul=lmin}         : time constant not allowed to be less than lmin
        facl = (1 - exp(-dt/taul))
}

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