Firing neocortical layer V pyramidal neuron (Reetz et al. 2014; Stadler et al. 2014)

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Accession:168148
Neocortical Layer V model with firing behaviour adjusted to in vitro observations. The model was used to investigate the effects of IFN and PKC on the excitability of neurons (Stadler et al 2014, Reetz et al. 2014). The model contains new channel simulations for HCN1, HCN2 and the big calcium dependent potassium channel BK.
References:
1 . Stadler K, Bierwirth C, Stoenica L, Battefeld A, Reetz O, Mix E, Schuchmann S, Velmans T, Rosenberger K, Bräuer AU, Lehnardt S, Nitsch R, Budt M, Wolff T, Kole MH, Strauss U (2014) Elevation in type I interferons inhibits HCN1 and slows cortical neuronal oscillations. Cereb Cortex 24:199-210 [PubMed]
2 . Reetz O, Stadler K, Strauss U (2014) Protein kinase C activation mediates interferon-ß-induced neuronal excitability changes in neocortical pyramidal neurons. J Neuroinflammation 11:185 [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: Neocortex;
Cell Type(s): Neocortex L5/6 pyramidal GLU cell;
Channel(s): I Na,p; I Na,t; I L high threshold; I A; I K; I M; I h; I K,Ca; I Sodium; I Calcium; I Mixed; I Potassium; I Q;
Gap Junctions:
Receptor(s):
Gene(s): HCN1; HCN2;
Transmitter(s):
Simulation Environment: NEURON;
Model Concept(s): Activity Patterns; Detailed Neuronal Models; Action Potentials; Signaling pathways;
Implementer(s): Stadler, Konstantin [konstantin.stadler at ntnu.no];
Search NeuronDB for information about:  Neocortex L5/6 pyramidal GLU cell; I Na,p; I Na,t; I L high threshold; I A; I K; I M; I h; I K,Ca; I Sodium; I Calcium; I Mixed; I Potassium; I Q;
//######################################
//  						    
//  LayerVinit.hoc		
//  --------------
//
// Assign general and channel parameters for the Layer V neuron
//  							
//  Author: Konstantin Stadler
//  Version: 20131108
//  							
//######################################
//	
// This file consists of two parts:
//
//  1) Parameter
//     All numerical values necessary for the model
//
//  2) Allocation
//     Conversion of units (as required by the mod files) 
//     Allocating channel properties  


// ************************************************************************** //
// ************************************************************************** //
// 1) Parameter
// ************************************************************************** //
// ************************************************************************** //


  //---------------------------------------------------------------------------
  // General parameters
  //---------------------------------------------------------------------------

  proc init_General() { local RmPoint, dis
		
	celsius    = 32
	Ri         = 68.022
	Cm         = 1.5431
	
    // parameters for the value of Rm along the dendrites
	RmSoma     = 34963
	RmEnd      = 5357
	RmHalfd    = 405.84
	RmSteep    = 50
			
	v_init     = -89    // Junction potential corrected RMP under ZD 7288, Kole 2006 (PMID: 16467515)
		
	spinescale = 2.0    // scale total area to account for spines
						
	forall {
		Ra=Ri
			
		insert pas {
            g_pas = 1/RmSoma
            cm    = Cm
		}
		insert cad
	}	
	
	access dend1[21]        //That's the soma	
	distance()
	
	for i=0,1090 {	    //Rm is distributed sigmoidal (Stuart and Spruston 1998, PMID: 9570781)
	      dend1[i] {  
		  dis     = distance(0)
		  RmPoint = RmEnd+(RmSoma-RmEnd)/(1+exp((dis-RmHalfd)/RmSteep))
		  g_pas   = 1/RmPoint
		}
	}
  }

  // --------------------------------------------------------------------------
  // Specific parameters, all in pS/um2 
  //---------------------------------------------------------------------------
	
  proc init_AxonParameter() {	
	
	g_Nax_Axon    = 3500       
	temp_Nax_Axon = 23  
							
	g_KV_Axon     = 40  
	temp_KV_Axon  = 23

	g_KM_Axon     = 50          
	temp_KM_Axon  = 35          

	vHact_KM_Axon = -27
	vHtau_KM_Axon = -29 
    }

  proc init_SomaParameter() {
		
		g_Na_Soma      = 420
		temp_Na_Soma   = 23
		
		g_Nap_Soma     = 10
		g_KV_Soma      = 20
		temp_KV_Soma   = 23
					
		g_KM_Soma      = 10
		temp_KM_Soma   = 35
				
		vHact_KM_Soma  = vHact_KM_Axon
		vHtau_KM_Soma  = vHact_KM_Axon

		g_KAprox_Soma  = 150
		g_KAdist_Soma  = 0
		temp_KA_Soma   = 23
	
		g_KBK_Soma     = 0.6
		
		g_CaHVA_Soma   = 2
		g_CaLVA_Soma   = 0
				
		temp_Ca_Soma   = 23
		
		g_HCN_Soma     = 0.95
				
		pHCN1_Soma     = 0.67
		Vrev_HCN_Soma  = -45
			
		Vhakt_HCN_Soma = -100
		k_HCN_Soma     = -10
		Vhtau_HCN_Soma = -100
		temp_HCN_Soma  = 24

		a0t_hcn1_Soma  = 0.00102
		a0t_hcn2_Soma  = 0.00022
  }

  proc init_DendParameter() {	
    // capacitance 
	cm_Dend_Dist = Cm*spinescale
	cm_Dend_Prox = Cm    
	
    // Na channel
	g_Na_ApiDendProx = 350
	g_Na_ApiDendDist = 320 
				  
	  temp_Na_Dend=23
	
	g_Na_BasDendStart = 150
	Na_BasRedFaktor = 0.5
				    
    // KV channel
	g_KV_DendStart = g_KV_Soma 
	g_KV_DendEnd = g_KV_Soma/17.2   
	KV_DendDecFac = 80    
	  temp_KV_Dend = 21	

    // Km channel
	g_KM_DistDend = 0 
	  temp_KM_Dend = temp_KM_Soma //=35	
	  vHact_KM_Dend = vHact_KM_Axon
  	  vHtau_KM_Dend = vHact_KM_Axon	
  	  	
    // KA channels - following Acker & Antic 2009
	KA_ProRed = 1/300           // reduction factor for the proximal KA channel type

	KA_BasIncFaktor = 0.7       // increase factor for the KA density of basal dendrites

	g_KA_BasDendStart = 150     // start density basal dendrites
	g_KA_Max = 2000             // maximal possible value
	g_KA_ApiDend = 300          // density for apical dendrites
	temp_KA_Dend = temp_KA_Soma 

    // BK channels			
	g_KBK_Dend = g_KBK_Soma    
	
    // ca channels	
	g_CaHVA_DistDend = 2   
	g_CaLVA_DistDend = 0.5 
	  temp_Ca_Dend=temp_Ca_Soma //=23	
		
    // HCN, distribution based on Kole 2006 
 	g_HCN_DendStart = g_HCN_Soma         // HCN density at the begin of the dendrites
	g_HCN_DendEnd = 40 * g_HCN_DendStart // HCN density at the end of the dendrites
	HCN_DendLambda = 323	             // steepness of the HCN increase, Kole 2006 (PMID: 16467515)
	
	pHCN1_Dend=0.67		                 //share of HCN1 

	Vrev_HCN_Dend = -45                  // reversal potential

	Vhakt_HCN_Dend =  Vhakt_HCN_Soma
	k_HCN_Dend = k_HCN_Soma
	Vhtau_HCN_Dend = -100
	  temp_HCN_Dend=temp_HCN_Soma 
	
	a0t_hcn1_Dend = a0t_hcn1_Soma
  	a0t_hcn2_Dend = a0t_hcn2_Soma
	  
    //!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!
    // the 1st 100 um of the apical trunk and the 1st 20 um of the basals 
    // only if these values differ from the above, overwrites the former
    //!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!
     
    g_KM_ProxDend = 5
    g_CaHVA_ProxDend = g_CaHVA_Soma
    g_CaLVA_ProxDend = 0 
  
  }


// ************************************************************************** //
// ************************************************************************** //
// 2) Allocation
// ************************************************************************** //
// ************************************************************************** //

  //---------------------------------------------------------------------------
  // Axon
  //---------------------------------------------------------------------------

  proc init_Axon() { 
	
   	axon {
	
		insert nax
            gbar_nax  = g_Nax_Axon
            temp_nax  = temp_Nax_Axon
		insert kv
            gbar_kv   = g_KV_Axon
            temp_kv   = temp_KV_Axon
		insert km
            gbar_km   = g_KM_Axon*(1E-4)
            temp_km   = temp_KM_Axon
            vhalfl_km = vHact_KM_Axon
            vhalft_km = vHtau_KM_Axon
	}

	soma {
 		
		insert nax
            gbar_nax  = g_Nax_Axon
            temp_nax  = temp_Nax_Axon
        insert kv
            gbar_kv   = g_KV_Axon
            temp_kv   = temp_KV_Axon
        insert km
            gbar_km   = g_KM_Axon*(1E-4)
            temp_km   = temp_KM_Axon
            vhalfl_km = vHact_KM_Axon
            vhalft_km = vHtau_KM_Axon
  	}

	for i=0,13 {
	   dend1[i]  {
		insert nax	
                gbar_nax = g_Nax_Axon
				temp_nax = temp_Nax_Axon
		insert kv	
                gbar_kv  = g_KV_Axon
				temp_kv  = temp_KV_Axon
	   	insert km 	
                gbar_km = g_KM_Axon*(1E-4)
		  		temp_km = temp_KM_Axon
		  		vhalfl_km = vHact_KM_Axon
		  		vhalft_km = vHtau_KM_Axon
	   }
	}
  }

  //---------------------------------------------------------------------------
  // Soma
  //---------------------------------------------------------------------------

  proc init_Soma() {	
	
	for i=14,27 {
		dend1[i] {
			
            insert na 	
                gbar_na = g_Na_Soma
                temp_na = temp_Na_Soma

            insert nap	
                gbar_nap = g_Nap_Soma * (1E-4)

            insert kv 	
                gbar_kv = g_KV_Soma
                temp_kv = temp_KV_Soma

            insert km 	
                gbar_km   = g_KM_Soma*(1E-4)
                temp_km   = temp_KM_Soma
                vhalfl_km = vHact_KM_Soma
                vhalft_km = vHtau_KM_Soma

            insert kap	
                gkabar_kap = g_KAprox_Soma*(1E-4)
                temp_kap   = temp_KA_Soma

            insert kad    
                gkabar_kad = g_KAdist_Soma*(1E-4)
                temp_kad   = temp_KA_Soma

            insert kBK 	
                gpeak_kBK = g_KBK_Soma*(1E-4)

            insert ca 	
                gbar_ca = g_CaHVA_Soma
                temp_ca = temp_Ca_Soma

            insert it2 	
                gcabar_it2 = g_CaLVA_Soma*(1E-4)

            insert hcn1		
            insert hcn2

		   // HCN Parameter
            gpeak_hcn1 = g_HCN_Soma*(1E-4)*pHCN1_Soma
            gpeak_hcn2 = g_HCN_Soma*(1E-4)*(1-pHCN1_Soma)
				
            Vrev_hcn1  = Vrev_HCN_Soma
            Vrev_hcn2  = Vrev_HCN_Soma
			
            vhakt_hcn1 = Vhakt_HCN_Soma
            vhakt_hcn2 = Vhakt_HCN_Soma
		
            k_hcn1     = k_HCN_Soma
            k_hcn2     = k_HCN_Soma
			
            vhtau_hcn1 = Vhtau_HCN_Soma
            vhtau_hcn2 = Vhtau_HCN_Soma
			
            temp_hcn1  = temp_HCN_Soma
            temp_hcn2  = temp_HCN_Soma
		
            a0t_hcn1   = a0t_hcn1_Soma
            a0t_hcn2   = a0t_hcn2_Soma
		} 
	}
  }

  //---------------------------------------------------------------------------
  // Dendrites 
  //---------------------------------------------------------------------------

  proc init_Dend() {local dis,ratio,lD,NaTemp,KaTemp,HcnTemp,dif,y0_HCN,A_HCN
		
	access dend1[21]	
	distance()		//measuring distance starting at the soma
	
	ld=0
	
   	for i=28,1090 {		//determine maximal length of the dendrite
    	  dend1[i] {		
    	  	dis=distance(.5)
    	  	if (lD < dis) lD = dis
    	  	}
    	 }
		
    // calculate HCN distribution parameter
   		
  	dif    = g_HCN_DendEnd - g_HCN_DendStart
  	A_HCN  = dif / ( (exp(lD/HCN_DendLambda)) - 1)
  	y0_HCN = g_HCN_DendStart - A_HCN

    // All Dendriten 
			
	for i=28,1090 {
	  dend1[i] {
	  	dis=distance(.5)
	  	
	  	g_pas=g_pas*spinescale   
		cm=cm_Dend_Dist	
             
         // all uniform distributions

        insert km   
            gbar_km = g_KM_DistDend*(1E-4) //distal, proximal see below
            temp_km = temp_KM_Axon
            vhalfl_km = vHact_KM_Dend
            vhalft_km = vHtau_KM_Dend	

        insert kBK  
            gpeak_kBK = g_KBK_Dend*(1E-4)*spinescale

        insert ca
            gbar_ca = g_CaHVA_DistDend*spinescale
            temp_ca = temp_Ca_Dend

        insert it2
            gcabar_it2 = g_CaLVA_DistDend*(1E-4)*spinescale
	     
	     //Na channel basal
		     
	     	insert na
	     	  NaTemp = g_Na_BasDendStart - dis * Na_BasRedFaktor
	     	  if (NaTemp < 0) NaTemp=0 
	     	  gbar_na = NaTemp*spinescale
	     	    temp_na = temp_Na_Dend
	     	  // Der Spinescale muss aus den prox. entfernt werden
	     		     
	     //KV channel
	     	
	     	insert kv   
	     	  dif = (g_KV_DendStart-g_KV_DendEnd)
	     	  gbar_kv = g_KV_DendEnd + ( dif * exp (-dis/KV_DendDecFac))	
		  if (gbar_kv < 0) gbar_kv=0
		  gbar_kv = gbar_kv * spinescale
		    temp_kv = temp_KV_Dend	     
		     
	     //KA channels basal
		       
		insert kap
		insert kad
		  KaTemp = g_KA_BasDendStart + dis * KA_BasIncFaktor
		  
		  if (KaTemp>g_KA_Max) {   
			KaTemp = g_KA_Max
		  }		
		  
		  ratio = KA_ProRed * dis   // share distal/proximal KA type
		  if (ratio < 0) ratio = 0
		  if (ratio > 1) ratio = 1	
		
		  gkabar_kad = KaTemp*ratio*(1E-4)*spinescale
		  gkabar_kap = KaTemp*(1-ratio)*(1E-4)*spinescale
							
		    temp_kap = temp_KA_Dend
		    temp_kad = temp_KA_Dend
		
	     //HCN 
		insert hcn1	
		insert hcn2
 			
		HcnTemp = y0_HCN+A_HCN*exp(dis / HCN_DendLambda)
		HcnTemp = HcnTemp*(1E-4)*spinescale
  			
		gpeak_hcn1 = HcnTemp*pHCN1_Dend
		gpeak_hcn2 = HcnTemp*(1-pHCN1_Dend)
  			
		Vrev_hcn1 = Vrev_HCN_Dend
		Vrev_hcn2 = Vrev_HCN_Dend
  			
		vhakt_hcn1 = Vhakt_HCN_Dend
		vhakt_hcn2 = Vhakt_HCN_Dend
  			
		k_hcn1 = k_HCN_Dend
		k_hcn2 = k_HCN_Dend
  			
		vhtau_hcn1 = Vhtau_HCN_Dend
		vhtau_hcn2 = Vhtau_HCN_Dend
		
		temp_hcn1 = temp_HCN_Dend
		temp_hcn2 = temp_HCN_Dend
	  
	  	a0t_hcn1 = a0t_hcn1_Dend
		a0t_hcn2 = a0t_hcn2_Dend
	  }
	}
	
    // apical dendrites, overwrites the values of 'all dendrites' from above
		
	forsec ApikalDend {	
		     
	     dis=distance(.5)
	     
         gbar_na = g_Na_ApiDendDist*spinescale  
	     
		ratio = KA_ProRed * dis   // share distal/proximal KA type
		if (ratio < 0) ratio = 0
		if (ratio > 1) ratio = 1	
  			
		gkabar_kad = g_KA_ApiDend*ratio*(1E-4)*spinescale
		gkabar_kap = g_KA_ApiDend*(1-ratio)*(1E-4)*spinescale
	}  
	
	
    // the 1st 100 um of the apical trunk 
    // overwrites the values from above / removes spinescale
	
	for i=28,32 {
	  dend1[i] {
		g_pas=g_pas/spinescale
		cm=cm_Dend_Prox
						
		
	  	gbar_km = g_KM_ProxDend*(1E-4)
	  	gpeak_kBK = g_KBK_Dend*(1E-4)
	  	gbar_ca = g_CaHVA_ProxDend
	  	gcabar_it2 = g_CaLVA_ProxDend*(1E-4)
	  	
	  	
	  	gbar_na = g_Na_ApiDendProx
	  	gbar_kv = gbar_kv / spinescale
	  	gkabar_kap = gkabar_kap / spinescale
	  	gkabar_kad = gkabar_kad / spinescale
	  	gpeak_hcn1 = gpeak_hcn1 / spinescale
	  	gpeak_hcn2 = gpeak_hcn2 / spinescale
	   }
	}
		
	for i=43,47 {
	  dend1[i] {
		g_pas=g_pas/spinescale
		cm=cm_Dend_Prox
					
	  	gbar_km = g_KM_ProxDend*(1E-4)
	  	gpeak_kBK = g_KBK_Dend*(1E-4)
	  	gbar_ca = g_CaHVA_ProxDend
	  	gcabar_it2 = g_CaLVA_ProxDend*(1E-4)
		
	
	  	gbar_na = g_Na_ApiDendProx
	  	gbar_kv = gbar_kv / spinescale
	  	gkabar_kap = gkabar_kap / spinescale
	  	gkabar_kad = gkabar_kad / spinescale
	  	gpeak_hcn1 = gpeak_hcn1 / spinescale
	  	gpeak_hcn2 = gpeak_hcn2 / spinescale
	  }
	}
		
	for i=71,72 {
	  dend1[i] {
		g_pas=g_pas/spinescale
		cm=cm_Dend_Prox
	
	  	gbar_km = g_KM_ProxDend*(1E-4)
	  	gpeak_kBK = g_KBK_Dend*(1E-4)
	  	gbar_ca = g_CaHVA_ProxDend
	  	gcabar_it2 = g_CaLVA_ProxDend*(1E-4)
		  	
	  	
	  	gbar_na = g_Na_ApiDendProx
	  	gbar_kv = gbar_kv / spinescale
	  	gkabar_kap = gkabar_kap / spinescale
	  	gkabar_kad = gkabar_kad / spinescale
	  	gpeak_hcn1 = gpeak_hcn1 / spinescale
	  	gpeak_hcn2 = gpeak_hcn2 / spinescale
	  }
	}

    // the 1st 20 um of the basals 
    // overwrites the values from above / removes spinescale
		
	for i=777,778 {
	  dend1[i] {
		g_pas=g_pas/spinescale 
		cm=cm_Dend_Prox
	
	  	gbar_km = g_KM_ProxDend*(1E-4)
	  	gpeak_kBK = g_KBK_Dend*(1E-4)
	  	gbar_ca = g_CaHVA_ProxDend
	  	gcabar_it2 = g_CaLVA_ProxDend*(1E-4)
	  	
	  	gbar_na = gbar_na / spinescale
	  	gbar_kv = gbar_kv / spinescale
	  	gkabar_kap = gkabar_kap / spinescale
	  	gkabar_kad = gkabar_kad / spinescale
	  	gpeak_hcn1 = gpeak_hcn1 / spinescale
	  	gpeak_hcn2 = gpeak_hcn2 / spinescale
	  
	  }
	}
		
	for i=808,810 {
	  dend1[i] {
		g_pas=g_pas/spinescale
		cm=cm_Dend_Prox
	
	  	gbar_km = g_KM_ProxDend*(1E-4)
	  	gpeak_kBK = g_KBK_Dend*(1E-4)
	  	gbar_ca = g_CaHVA_ProxDend
	  	gcabar_it2 = g_CaLVA_ProxDend*(1E-4)
	  	
	  	gbar_na = gbar_na / spinescale
	  	gbar_kv = gbar_kv / spinescale
	  	gkabar_kap = gkabar_kap / spinescale
	  	gkabar_kad = gkabar_kad / spinescale
	  	gpeak_hcn1 = gpeak_hcn1 / spinescale
	  	gpeak_hcn2 = gpeak_hcn2 / spinescale
	  
	  }
	}
	
	for i=923,924 {
	  dend1[i] {
		g_pas=g_pas/spinescale
		cm=cm_Dend_Prox
	
	  	gbar_km = g_KM_ProxDend*(1E-4)
	  	gpeak_kBK = g_KBK_Dend*(1E-4)
	  	gbar_ca = g_CaHVA_ProxDend
	  	gcabar_it2 = g_CaLVA_ProxDend*(1E-4)
	  	
	  	gbar_na = gbar_na / spinescale
	  	gbar_kv = gbar_kv / spinescale
	  	gkabar_kap = gkabar_kap / spinescale
	  	gkabar_kad = gkabar_kad / spinescale
	  	gpeak_hcn1 = gpeak_hcn1 / spinescale
	  	gpeak_hcn2 = gpeak_hcn2 / spinescale
	  
	  }
	}
	
	for i=957,959 {
	  dend1[i] {
		g_pas=g_pas/spinescale
		cm=cm_Dend_Prox
		
	  	gbar_km = g_KM_ProxDend*(1E-4)
	  	gpeak_kBK = g_KBK_Dend*(1E-4)
	  	gbar_ca = g_CaHVA_ProxDend
	  	gcabar_it2 = g_CaLVA_ProxDend*(1E-4)
	  	
	  	gbar_na = gbar_na / spinescale
	  	gbar_kv = gbar_kv / spinescale
	  	gkabar_kap = gkabar_kap / spinescale
	  	gkabar_kad = gkabar_kad / spinescale
	  	gpeak_hcn1 = gpeak_hcn1 / spinescale
	  	gpeak_hcn2 = gpeak_hcn2 / spinescale
	  
	  }
	}
	
	for i=1006,1007 {
	  dend1[i] {
		g_pas=g_pas/spinescale
		cm=cm_Dend_Prox
		
  		gbar_km = g_KM_ProxDend*(1E-4)
	  	gpeak_kBK = g_KBK_Dend*(1E-4)
	  	gbar_ca = g_CaHVA_ProxDend
	  	gcabar_it2 = g_CaLVA_ProxDend*(1E-4)
	  	
	  	gbar_na = gbar_na / spinescale
	  	gbar_kv = gbar_kv / spinescale
	  	gkabar_kap = gkabar_kap / spinescale
	  	gkabar_kad = gkabar_kad / spinescale
  		gpeak_hcn1 = gpeak_hcn1 / spinescale
	  	gpeak_hcn2 = gpeak_hcn2 / spinescale
	  
	  }
	}
		
	for i=1055,1058 {
	  dend1[i] {
		g_pas=g_pas/spinescale
		cm=cm_Dend_Prox
		
  		gbar_km = g_KM_ProxDend*(1E-4)
	  	gpeak_kBK = g_KBK_Dend*(1E-4)
	  	gbar_ca = g_CaHVA_ProxDend
	  	gcabar_it2 = g_CaLVA_ProxDend*(1E-4)
	  	
	  	gbar_na = gbar_na / spinescale
  		gbar_kv = gbar_kv / spinescale
  		gkabar_kap = gkabar_kap / spinescale
	  	gkabar_kad = gkabar_kad / spinescale
	  	gpeak_hcn1 = gpeak_hcn1 / spinescale
  		gpeak_hcn2 = gpeak_hcn2 / spinescale
	
	  }
	}
		
	for i=1059,1060 {
	  dend1[i] {
		g_pas=g_pas/spinescale
		cm=cm_Dend_Prox
	
	  	gbar_km = g_KM_ProxDend*(1E-4)
	  	gpeak_kBK = g_KBK_Dend*(1E-4)
  		gbar_ca = g_CaHVA_ProxDend
	  	gcabar_it2 = g_CaLVA_ProxDend*(1E-4)
	  	
	  	gbar_na = gbar_na / spinescale
	  	gbar_kv = gbar_kv / spinescale
	  	gkabar_kap = gkabar_kap / spinescale
	  	gkabar_kad = gkabar_kad / spinescale
  		gpeak_hcn1 = gpeak_hcn1 / spinescale
	  	gpeak_hcn2 = gpeak_hcn2 / spinescale
	
	  }
	}
		
	for i=1067,1069 {
	  dend1[i] {
		g_pas=g_pas/spinescale
		cm=cm_Dend_Prox
		
		gbar_km = g_KM_ProxDend*(1E-4)
		gpeak_kBK = g_KBK_Dend*(1E-4)
	  	gbar_ca = g_CaHVA_ProxDend
	 	gcabar_it2 = g_CaLVA_ProxDend*(1E-4)
	  	
	  	gbar_na = gbar_na / spinescale
	  	gbar_kv = gbar_kv / spinescale
	  	gkabar_kap = gkabar_kap / spinescale
	  	gkabar_kad = gkabar_kad / spinescale
	 	gpeak_hcn1 = gpeak_hcn1 / spinescale
	  	gpeak_hcn2 = gpeak_hcn2 / spinescale
	
	  }
	} 
  }

//---------------------------------------------------------------------------
// reversal and start potential    
//---------------------------------------------------------------------------

  proc init_Volt() {
	
	forall {	
		vshift_na=-10 //provides AP threshold of ~ -60 mV at soma 
		vshift_nax=-10
			
		ena = 55
 		ek = -105
 		e_pas = v_init

		taur_cad = 100  
	}
  }



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