Neocort. pyramidal cells subthreshold somatic voltage controls spike propagation (Munro Kopell 2012)

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Accession:136309
There is suggestive evidence that pyramidal cell axons in neocortex may be coupled by gap junctions into an ``axonal plexus" capable of generating Very Fast Oscillations (VFOs) with frequencies exceeding 80 Hz. It is not obvious, however, how a pyramidal cell in such a network could control its output when action potentials are free to propagate from the axons of other pyramidal cells into its own axon. We address this problem by means of simulations based on 3D reconstructions of pyramidal cells from rat somatosensory cortex. We show that somatic depolarization enables propagation via gap junctions into the initial segment and main axon, while somatic hyperpolarization disables it. We show further that somatic voltage cannot effectively control action potential propagation through gap junctions on minor collaterals; action potentials may therefore propagate freely from such collaterals regardless of somatic voltage. In previous work, VFOs are all but abolished during the hyperpolarization phase of slow-oscillations induced by anesthesia in vivo. This finding constrains the density of gap junctions on collaterals in our model and suggests that axonal sprouting due to cortical lesions may result in abnormally high gap junction density on collaterals, leading in turn to excessive VFO activity and hence to epilepsy via kindling.
Reference:
1 . Munro E, Kopell N (2012) Subthreshold somatic voltage in neocortical pyramidal cells can control whether spikes propagate from the axonal plexus to axon terminals: a model study. J Neurophysiol 107:2833-52 [PubMed]
Model Information (Click on a link to find other models with that property)
Model Type: Realistic Network; Neuron or other electrically excitable cell; Axon;
Brain Region(s)/Organism: Neocortex;
Cell Type(s): Neocortex L5/6 pyramidal GLU cell; Neocortex U1 L2/6 pyramidal intratelencephalic GLU cell;
Channel(s): I Na,t; I K; I Sodium; I Potassium;
Gap Junctions: Gap junctions;
Receptor(s):
Gene(s):
Transmitter(s):
Simulation Environment: NEURON; MATLAB;
Model Concept(s): Oscillations; Detailed Neuronal Models; Axonal Action Potentials; Epilepsy;
Implementer(s): Munro, Erin [ecmun at math.bu.edu];
Search NeuronDB for information about:  Neocortex L5/6 pyramidal GLU cell; Neocortex U1 L2/6 pyramidal intratelencephalic GLU cell; I Na,t; I K; I Sodium; I Potassium;
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Munro_Kopell_corticalcontrol
NEURON
cell_geoms
run_files
readme.txt
extrema.mod
gap.mod
k2_Traub.mod *
ka_Traub.mod *
kdr_Jonas.mod
kdr_Traub.mod
kdr_Yu.mod
naf_Jonas.mod
naf_Traub.mod
naf_Yu.mod
pas_basket.mod
pas_chand.mod
030625DS2.hoc
axon_templates.hoc
C040896A-P3.hoc
C040896A-P3axgeom.hoc
C230797B-P4.hoc
C270999B-P2axgeom.hoc
C280199C-P1.hoc
C290500C-P1axgeom.hoc
cell_templates.hoc
expcell_templates.hoc
gap_junction.hoc
gj_propagation_test.ses
junction_stats.hoc
kinetics.hoc
kinetics_wholecell.hoc
mosinit.hoc
propagation_test.ses
PropagationSearch.hoc
simulation_base.hoc
vs-arg_cutoff.hoc
vs-arg_cutoff_Jonas.hoc
vs-expcell_gj.hoc
vs-expcell_gj_gL.hoc
vs-expcell_gj_Jonas.hoc
vs-expcell_gjCC.hoc
vs-expcell_gjISgNa.hoc
vs-expcell_gjsISgNa.hoc
vs-generic_cutoff.hoc
vs-generic_cutoff_Jonas.hoc
                            
load_file("kinetics.hoc")

proc choose_stim_site(){local i,avg_diam,totalL,curL,arcl,l1,l2,arcd localobj cell,rvp,sec_list,sn
    // choose the "stimulation site" of the crux of the axon for the cell
    // which is always 1/4 of the electrotonic length away from the crux
    cell = $o1
    kinetics = $2

    rvp = new RangeVarPlot("v")
    cell.axon[cell.junction_site]{ rvp.begin(0) }
    cell.axon[cell.extstim_site]{ rvp.end(0.5) }
    sn = new String()
    sprint(sn.s,".*axon[%d]",cell.extstim_site) // cell.junction_site)
    sec_list = new SectionList()
    rvp.list(sec_list)
    avg_diam = 0
    totalL = 0
    forsec sec_list{
        if(issection(sn.s)){ // in extstim_site, only go as far as stim
            for(i=1;i<n3d() && arc3d(i)<0.5*L;i=i+1){
                arcd = (diam3d(i-1)+diam3d(i))/2
                arcl = arc3d(i)-arc3d(i-1)
                avg_diam = avg_diam + arcd*arcl
            }
            //arcl = arc3d(i)-arc3d(i-1)
            //l1 = (0.5*L-arc3d(i-1))/2
            //l2 = arcl-l1
            //avg_diam = avg_diam + ((diam3d(i-1)*l1+diam3d(i)*l2)/arcl)*(l1*2)
            //totalL = totalL + l1*2
            totalL = totalL + L/2
        }else{ // between extstim and junction_site, use full length
            for(i=1;i<n3d();i=i+1){
                arcd = (diam3d(i-1)+diam3d(i))/2
                arcl = arc3d(i)-arc3d(i-1)
                avg_diam = avg_diam + arcd*arcl
            }
            totalL = totalL + L
        }
    }
    avg_diam = avg_diam/totalL
    print "avg_diam=", avg_diam
    if(kinetics==1){
        cell.stim_length = elength(avg_diam)/8
    }else{
        cell.stim_length = elength(avg_diam)/3
    }
    print "elength=", elength(avg_diam)
    if(cell.stim_length>totalL-cell.stim_length){
        print "totalL = ", totalL, ", stim_length = ", cell.stim_length, ", extstim_length = ", totalL-cell.stim_length
        return
    }
    curL = 0
    forsec sec_list{
        if(cell.stim_loc == 0 && curL+L>cell.stim_length){
            cell.stim_loc = (cell.stim_length-curL)/L
            cell.stim_sr = new SectionRef()
        }
        if(cell.extstimrec_loc == 0 && curL+L>totalL-cell.stim_length){
            cell.extstimrec_loc = (totalL-cell.stim_length-curL)/L
            cell.extstim_sr = new SectionRef()
        }
        curL = curL + L
    }
}

begintemplate SimpleAxon
public axon,extstim_site,stim_sr,stim_loc,junction_site,prop_site,IS,se,nsection,stim_length,extstimrec_loc,extstim_sr
external insert_Traub,insert_Jonas,insert_Yu,insert_conglomerate,insert_modTraub,tstop,choose_stim_site

create axon[31]
objref this,se,stim_sr,extstim_sr

proc init(){
    xopen("C270999B-P2axgeom.hoc")
    nsection = 31
    extstim_site = 8
    junction_site = 5
    prop_site = 10
    IS = 4
    
    forall {
        nseg = int(L/$2) + 1
    }
    
    kinetics = $1
    if(kinetics==0){
        insert_Traub()
    }else if(kinetics==1){
        insert_Jonas()
    }else if(kinetics==2){
        insert_Yu()
    }else if(kinetics==3){
        insert_conglomerate()
    }else{
        insert_modTraub(this,IS)
    }
    forall{
        insert extr
    }
    stim_length = 0
    stim_loc = 0
    extstimrec_loc = 0
  choose_stim_site(this,kinetics)
  print "in SimpleAxon init: axon[junction_site].diam=", axon[junction_site].diam

  axon[IS]{
    se = new SEClamp(0)
    se.dur1 = tstop
    se.amp1 = axon[extstim_site].e_pas
    somaL = 0.01
    somar = 16.136
    rho_soma = Ra*somaL/(PI*somar^2)*10^(-2)
    rho_IS = Ra*L/(PI*(diam/2)^2)*10^(-2)
    se.rs = (rho_soma+rho_IS)/2
  }
}

endtemplate SimpleAxon


begintemplate TypicalAxon
public axon,extstim_site,stim_sr,stim_loc,junction_site,prop_site,IS,se,nsection,stim_length,extstim_sr,extstimrec_loc
external insert_Traub,insert_Jonas,insert_Yu,insert_conglomerate,insert_modTraub,tstop,choose_stim_site

create axon[65]
objref this,se,stim_sr,extstim_sr

proc init(){
    xopen("C040896A-P3axgeom.hoc")
    nsection = 65
    extstim_site = 43
    //stim_site = 42
    junction_site = 40 
    //axon[junction_site]{ distance() }
    //axon[stim_site]{
    //stim_loc = (35-distance(0))/L
    //}
    prop_site = 2
    IS = 0
    
    forall {
        nseg = int(L/$2) + 1
    }
    
    kinetics = $1
    if(kinetics==0){
        insert_Traub()
    }else if(kinetics==1){
        insert_Jonas()
    }else if(kinetics==2){
        insert_Yu()
    }else if(kinetics==3){
        insert_conglomerate()
    }else{
        insert_modTraub(this,IS)
    }
    forall{
        insert extr
    }
    stim_length = 0
    stim_loc = 0
    extstimrec_loc = 0
    choose_stim_site(this,kinetics)
    print "axon[junction_site]=", axon[junction_site].diam
    
    axon[IS]{
        se = new SEClamp(0)
        se.dur1 = tstop
        se.amp1 = axon[extstim_site].e_pas
        somaL = 0.01
        somar = 23.13
        rho_soma = Ra*somaL/(PI*somar^2)*10^(-2)
        rho_IS = Ra*L/(PI*(diam/2)^2)*10^(-2)
        se.rs = (rho_soma+rho_IS)/2
    }
}

endtemplate TypicalAxon


begintemplate ComplexAxon
public axon,extstim_site,stim_loc,prop_site,junction_site,IS,se,nsection,stim_sr,stim_length,extstim_sr,extstimrec_loc
external insert_Traub,insert_Jonas,insert_Yu,insert_conglomerate,insert_modTraub,tstop,choose_stim_site

create axon[75]
objref this,se,stim_sr,extstim_sr

proc init(){local stim_dist,avg_diam,totalL localobj seclist
    xopen("C290500C-P1axgeom.hoc")
    nsection = 75
    extstim_site = 12
    prop_site = 39
    IS = 0
    
    forall {
        nseg = int(L/$2) + 1
    }
    
    kinetics = $1
    if(kinetics==0){
        insert_Traub()
    }else if(kinetics==1){
        insert_Jonas()
    }else if(kinetics==2){
        insert_Yu()
    }else if(kinetics==3){
        insert_conglomerate()
    }else{
        insert_modTraub(this,IS)
    }
    forall{
        insert extr
    }
    junction_site = 10
    stim_length = 0
    stim_loc = 0
    extstimrec_loc = 0
    choose_stim_site(this,kinetics)
  //stim_site = 10
  //stim_loc = elength(axon[stim_site].diam)/(4*axon[stim_site].L)

  axon[IS]{
    se = new SEClamp(0)
    se.dur1 = tstop
    se.amp1 = axon[extstim_site].e_pas
    somaL = 5.3779
    somar = 2.49
    rho_soma = Ra*somaL/(PI*somar^2)*10^(-2)
    rho_IS = Ra*L/(PI*(diam/2)^2)*10^(-2)
    se.rs = (rho_soma+rho_IS)/2
}

}

endtemplate ComplexAxon

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