//######################################
//
// 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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