Synaptic gating at axonal branches, and sharp-wave ripples with replay (Vladimirov et al. 2013)

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Accession:150446
The computational model of in vivo sharp-wave ripples with place cell replay. Excitatory post-synaptic potentials at dendrites gate antidromic spikes arriving from the axonal collateral, and thus determine when the soma and the main axon fire. The model allows synchronous replay of pyramidal cells during sharp-wave ripple event, and the replay is possible in both forward and reverse directions.
Reference:
1 . Vladimirov N, Tu Y, Traub RD (2013) Synaptic gating at axonal branches, and sharp-wave ripples with replay: a simulation study. Eur J Neurosci 38:3435-47 [PubMed]
Model Information (Click on a link to find other models with that property)
Model Type: Neuron or other electrically excitable cell; Axon; Dendrite; Connectionist Network;
Brain Region(s)/Organism:
Cell Type(s): Hippocampus CA1 pyramidal GLU cell; Hippocampus CA1 basket cell;
Channel(s): I Na,t; I A; I K; I K,leak; I K,Ca; I Calcium; I Potassium; I_AHP;
Gap Junctions: Gap junctions;
Receptor(s): GabaA; AMPA;
Gene(s):
Transmitter(s): Gaba; Glutamate;
Simulation Environment: NEURON;
Model Concept(s): Action Potential Initiation; Oscillations; Synchronization; Axonal Action Potentials; Place cell/field; Conduction failure; Brain Rhythms;
Implementer(s): Vladimirov, Nikita ;
Search NeuronDB for information about:  Hippocampus CA1 pyramidal GLU cell; GabaA; AMPA; I Na,t; I A; I K; I K,leak; I K,Ca; I Calcium; I Potassium; I_AHP; Gaba; Glutamate;
begintemplate pyramidalCA1
public init, topol, basic_shape, subsets, geom, biophys, geom_nseg, biophys_inhomo
public synlist, x, y, z, position, x_index, y_index, rotateOY
public connect2targetSoma, connect2targetCollat, connect2targetSomaSpikelet

public soma, basal, shaft, apical, ais, axon, collat, axonal, NoR
public all, level1, level2, level3, level4, level5, level6
public level7, level8, level9, level10, level11
public setcurrentbias_soma, setcurrentbias_AIS, reduceCa, setindexXY
objref synlist

proc init() {
  topol()
  subsets()
  geom()
  geom_nseg()
  biophys()
  reduceCa(0.5)
  double_dend_cond()
  synlist = new List()
  synapses()
  x = y = z = 0 // only change via position
}

create soma, basal[28], shaft, apical[34], ais, axon[3], collat[2], NoR[2]

proc topol() { local i
  for i = 0, 1 connect basal[i](0), soma(0)
  for i = 2, 3 connect basal[i](0), soma(0)
  for i = 4, 5 connect basal[i](0), basal(1)
  for i = 6, 7 connect basal[i](0), basal[1](1)
  for i = 8, 9 connect basal[i](0), basal[2](1)
  for i = 10, 11 connect basal[i](0), basal[3](1)
  for i = 12, 13 connect basal[i](0), basal[4](1)
  for i = 14, 15 connect basal[i](0), basal[5](1)
  for i = 16, 17 connect basal[i](0), basal[6](1)
  for i = 18, 19 connect basal[i](0), basal[7](1)
  for i = 20, 21 connect basal[i](0), basal[8](1)
  for i = 22, 23 connect basal[i](0), basal[9](1)
  for i = 24, 25 connect basal[i](0), basal[10](1)
  for i = 26, 27 connect basal[i](0), basal[11](1)
  connect shaft(0), soma(1)
  for i = 0, 1 connect apical[i](0), shaft(1)
  for i = 2, 3 connect apical[i](0), apical[i-2](1)
  for i = 4, 5 connect apical[i](0), apical[i-4](1)
  for i = 6, 7 connect apical[i](0), apical[4](1)
  for i = 8, 9 connect apical[i](0), apical[5](1)
  for i = 10, 11 connect apical[i](0), apical[6](1)
  for i = 12, 13 connect apical[i](0), apical[7](1)
  for i = 14, 15 connect apical[i](0), apical[8](1)
  for i = 16, 17 connect apical[i](0), apical[9](1)
  for i = 18, 19 connect apical[i](0), apical[11](1)
  for i = 20, 21 connect apical[i](0), apical[13](1)
  for i = 22, 23 connect apical[i](0), apical[15](1)
  for i = 24, 25 connect apical[i](0), apical[17](1)
  for i = 26, 33 connect apical[i](0), apical[i-8](1)
  connect ais(0), soma(0)
  connect axon[0](0), ais(1)
  connect collat[0](0), ais(1)
  connect axon[1](0), axon[0](1)
  connect NoR[0](0), axon[1](1)
  connect axon[2](0), NoR[0](1)
  connect collat[1](0), NoR[0](1)
  connect NoR[1](0), axon[2](1)
  basic_shape()
}
proc basic_shape() {
  soma {pt3dclear() pt3dadd(0, 0, 0, 1) pt3dadd(0, 15, 0, 1)}
  basal {pt3dclear() pt3dadd(0, 0, 0, 1) pt3dadd(-74, -44, 0, 1)}
  basal[1] {pt3dclear() pt3dadd(0, 0, 0, 1) pt3dadd(-14, -59, 0, 1)}
  basal[2] {pt3dclear() pt3dadd(0, 0, 0, 1) pt3dadd(14, -59, 0, 1)}
  basal[3] {pt3dclear() pt3dadd(15, 0, 0, 1) pt3dadd(90, -44, 0, 1)}
  basal[4] {pt3dclear() pt3dadd(-74, -44, 0, 1) pt3dadd(-119, -44, 0, 1)}
  basal[5] {pt3dclear() pt3dadd(-74, -44, 0, 1) pt3dadd(-94, -89, 0, 1)}
  basal[6] {pt3dclear() pt3dadd(-14, -59, 0, 1) pt3dadd(-29, -89, 0, 1)}
  basal[7] {pt3dclear() pt3dadd(-14, -59, 0, 1) pt3dadd(0, -89, 0, 1)}
  basal[8] {pt3dclear() pt3dadd(30, -59, 0, 1) pt3dadd(15, -89, 0, 1)}
  basal[9] {pt3dclear() pt3dadd(30, -59, 0, 1) pt3dadd(45, -89, 0, 1)}
  basal[10] {pt3dclear() pt3dadd(90, -44, 0, 1) pt3dadd(110, -89, 0, 1)}
  basal[11] {pt3dclear() pt3dadd(90, -44, 0, 1) pt3dadd(135, -44, 0, 1)}
  basal[12] {pt3dclear() pt3dadd(-119, -44, 0, 1) pt3dadd(-164, -14, 0, 1)}
  basal[13] {pt3dclear() pt3dadd(-119, -44, 0, 1) pt3dadd(-164, -74, 0, 1)}
  basal[14] {pt3dclear() pt3dadd(-89, -89, 0, 1) pt3dadd(-119, -104, 0, 1)}
  basal[15] {pt3dclear() pt3dadd(-89, -89, 0, 1) pt3dadd(-89, -134, 0, 1)}
  basal[16] {pt3dclear() pt3dadd(-29, -89, 0, 1) pt3dadd(-59, -104, 0, 1)}
  basal[17] {pt3dclear() pt3dadd(-29, -89, 0, 1) pt3dadd(-29, -119, 0, 1)}
  basal[18] {pt3dclear() pt3dadd(0, -89, 0, 1) pt3dadd(-5, -119, 0, 1)}
  basal[19] {pt3dclear() pt3dadd(0, -89, 0, 1) pt3dadd(0, -119, 0, 1)}
  basal[20] {pt3dclear() pt3dadd(15, -89, 0, 1) pt3dadd(15, -119, 0, 1)}
  basal[21] {pt3dclear() pt3dadd(15, -89, 0, 1) pt3dadd(20, -119, 0, 1)}
  basal[22] {pt3dclear() pt3dadd(45, -89, 0, 1) pt3dadd(45, -119, 0, 1)}
  basal[23] {pt3dclear() pt3dadd(45, -89, 0, 1) pt3dadd(75, -104, 0, 1)}
  basal[24] {pt3dclear() pt3dadd(105, -89, 0, 1) pt3dadd(105, -134, 0, 1)}
  basal[25] {pt3dclear() pt3dadd(105, -89, 0, 1) pt3dadd(135, -104, 0, 1)}
  basal[26] {pt3dclear() pt3dadd(135, -44, 0, 1) pt3dadd(180, -74, 0, 1)}
  basal[27] {pt3dclear() pt3dadd(135, -44, 0, 1) pt3dadd(180, -14, 0, 1)}
  shaft {pt3dclear() pt3dadd(15, 0, 0, 1) pt3dadd(15, 30, 0, 1)}
  apical {pt3dclear() pt3dadd(15, 30, 0, 1) pt3dadd(-44, 60, 0, 1)}
  apical[1] {pt3dclear() pt3dadd(15, 30, 0, 1) pt3dadd(75, 60, 0, 1)}
  apical[2] {pt3dclear() pt3dadd(-44, 60, 0, 1) pt3dadd(-74, 90, 0, 1)}
  apical[3] {pt3dclear() pt3dadd(75, 60, 0, 1) pt3dadd(105, 90, 0, 1)}
  apical[4] {pt3dclear() pt3dadd(-44, 60, 0, 1) pt3dadd(-44, 105, 0, 1)}
  apical[5] {pt3dclear() pt3dadd(75, 60, 0, 1) pt3dadd(75, 105, 0, 1)}
  apical[6] {pt3dclear() pt3dadd(-44, 105, 0, 1) pt3dadd(-74, 135, 0, 1)}
  apical[7] {pt3dclear() pt3dadd(-44, 105, 0, 1) pt3dadd(-29, 135, 0, 1)}
  apical[8] {pt3dclear() pt3dadd(75, 105, 0, 1) pt3dadd(60, 135, 0, 1)}
  apical[9] {pt3dclear() pt3dadd(75, 105, 0, 1) pt3dadd(105, 135, 0, 1)}
  apical[10] {pt3dclear() pt3dadd(-74, 135, 0, 1) pt3dadd(-89, 165, 0, 1)}
  apical[11] {pt3dclear() pt3dadd(-74, 135, 0, 1) pt3dadd(-74, 180, 0, 1)}
  apical[12] {pt3dclear() pt3dadd(-29, 135, 0, 1) pt3dadd(0, 150, 0, 1)}
  apical[13] {pt3dclear() pt3dadd(-29, 135, 0, 1) pt3dadd(-29, 180, 0, 1)}
  apical[14] {pt3dclear() pt3dadd(60, 135, 0, 1) pt3dadd(30, 150, 0, 1)}
  apical[15] {pt3dclear() pt3dadd(60, 135, 0, 1) pt3dadd(60, 180, 0, 1)}
  apical[16] {pt3dclear() pt3dadd(105, 135, 0, 1) pt3dadd(120, 165, 0, 1)}
  apical[17] {pt3dclear() pt3dadd(105, 135, 0, 1) pt3dadd(105, 180, 0, 1)}
  apical[18] {pt3dclear() pt3dadd(-74, 180, 0, 1) pt3dadd(-89, 195, 0, 1)}
  apical[19] {pt3dclear() pt3dadd(-74, 180, 0, 1) pt3dadd(-59, 195, 0, 1)}
  apical[20] {pt3dclear() pt3dadd(-29, 180, 0, 1) pt3dadd(-44, 195, 0, 1)}
  apical[21] {pt3dclear() pt3dadd(-29, 180, 0, 1) pt3dadd(-14, 195, 0, 1)}
  apical[22] {pt3dclear() pt3dadd(60, 180, 0, 1) pt3dadd(45, 195, 0, 1)}
  apical[23] {pt3dclear() pt3dadd(60, 180, 0, 1) pt3dadd(75, 195, 0, 1)}
  apical[24] {pt3dclear() pt3dadd(105, 180, 0, 1) pt3dadd(90, 195, 0, 1)}
  apical[25] {pt3dclear() pt3dadd(105, 180, 0, 1) pt3dadd(120, 195, 0, 1)}
  apical[26] {pt3dclear() pt3dadd(-89, 195, 0, 1) pt3dadd(-89, 225, 0, 1)}
  apical[27] {pt3dclear() pt3dadd(-59, 195, 0, 1) pt3dadd(-59, 225, 0, 1)}
  apical[28] {pt3dclear() pt3dadd(-44, 195, 0, 1) pt3dadd(-44, 225, 0, 1)}
  apical[29] {pt3dclear() pt3dadd(-14, 195, 0, 1) pt3dadd(-14, 225, 0, 1)}
  apical[30] {pt3dclear() pt3dadd(45, 195, 0, 1) pt3dadd(45, 225, 0, 1)}
  apical[31] {pt3dclear() pt3dadd(75, 195, 0, 1) pt3dadd(75, 225, 0, 1)}
  apical[32] {pt3dclear() pt3dadd(90, 195, 0, 1) pt3dadd(90, 225, 0, 1)}
  apical[33] {pt3dclear() pt3dadd(120, 195, 0, 1) pt3dadd(120, 225, 0, 1)}
  ais {pt3dclear() pt3dadd(0, 0, 0, 1) pt3dadd(0, -44, 0, 1)}
  axon[0] {pt3dclear() pt3dadd(0, -44, 0, 1) pt3dadd(0, -144, 0, 1)}
  axon[1] {pt3dclear() pt3dadd(0, -144, 0, 1) pt3dadd(0, -244, 0, 1)}
  axon[2] {pt3dclear() pt3dadd(0, -244, 0, 1) pt3dadd(0, -344, 0, 1)}
  collat[0] {pt3dclear() pt3dadd(0, -144, 0, 1) pt3dadd(-200, -144, 0, 1)}
  collat[1] {pt3dclear() pt3dadd(0, -244, 0, 1) pt3dadd(200, -244, 0, 1)}
}

objref all, level1, level2, level3, level4, level5, level6
objref level7, level8, level9, level10, level11, soma_dendrites, dendrites, axonal
proc subsets() { local i
  objref all, level1, level2, level3, level4, level5, level6
  objref level7, level8, level9, level10, level11
  all = new SectionList()
  soma_dendrites = new SectionList()
  dendrites = new SectionList()
  axonal = new SectionList()
    soma { all.append() soma_dendrites.append() }
    for i=0, 27 basal[i] { all.append() soma_dendrites.append() dendrites.append()}
    shaft { all.append() soma_dendrites.append() dendrites.append()}
    for i=0, 33 apical[i] {all.append() soma_dendrites.append() dendrites.append()}
    
	ais { all.append() axonal.append() }
    for i=0, 2 axon[i] { all.append() axonal.append() }
	for i=0, 1 collat[i] { all.append() axonal.append() }
	for i=0, 1 NoR[i] { all.append() axonal.append() }

  level1 = new SectionList()
    for i=12, 27 basal[i] { level1.append() }

  level2 = new SectionList()
    for i=4, 11 basal[i] { level2.append() }

  level3 = new SectionList()
    for i=0, 3 basal[i] { level3.append() }

  level4 = new SectionList()
    soma { level4.append() }

  level5 = new SectionList()
    { shaft level5.append() }

  level6 = new SectionList()
    for i=0, 1 apical[i] { level6.append() }

  level7 = new SectionList()
    for i=2, 5 apical[i] { level7.append() }

  level8 = new SectionList()
    for i=6, 9 apical[i] { level8.append() }

  level9 = new SectionList()
    for i=10, 17 apical[i] { level9.append() }

  level10 = new SectionList()
    for i=18, 25 apical[i] { level10.append() }

  level11 = new SectionList()
    for i=26, 33 apical[i] { level11.append() }

}
proc geom() {
  forsec level1 {  L = 70  diam = 2  }
  forsec level2 {  L = 40  diam = 3.14  }
  forsec level3 {  L = 15  diam = 5  }
  forsec level4 {  L = 25.5  diam = 30  } //soma
  forsec level5 {  L = 50  diam = 10  } //shaft
  forsec level6 {  L = 50  diam = 6.3  }
  forsec level7 {  L = 50  diam = 5  }
  for i=2, 3 apical[i] { L = 70 diam = 2.78 } // oblique, level 7
  forsec level8 {  L = 60  diam = 3.14  }
  forsec level9 {  L = 60  diam = 2.5  }
  apical[10] { L = 50 diam = 1.4 } //oblique, level 9
  apical[12] { L = 50 diam = 1.4 } //oblique, level 9
  apical[14] { L = 50 diam = 1.4 } //oblique, level 9
  apical[16] { L = 50 diam = 1.4 } //oblique, level 9
  forsec level10 {  L = 60  diam = 1.6  }
  forsec level11 {  L = 60  diam = 1.6  }
  ais {  L = 40 diam = 2 }
  axon[0] {  L = 10  diam = 1  }
  axon[1] {  L = 60  diam = 1.4  } //now myelinated
  axon[2] {  L = 60  diam = 1.4  } //now myelinated
  for i=0, 1 NoR[i] { L = 1 diam = 1.1 } // Nodes of Ranvier
  for i=0, 1 collat[i] {  L = 200  diam = 0.5  }
}
external lambda_f
proc geom_nseg() {
  forsec all { nseg = int((L/(0.1*lambda_f(100))+.9)/2)*2 + 1  }
}
// apply the biophysical properties of Traub et al 2005 model to the cell
// $o1 is the cell
proc biophys(){
//general params:
    forsec all { 
	   insert pas 
	   cm = 0.75
	}   
	forsec soma_dendrites {
	    g_pas =   2.E-05
        Ra =   200.
	}
	forsec axonal {
       g_pas =   0.001 
       Ra =   100.
       insert NaFax
	   gbar_NaFax =   0.300 
       insert Kdrax
       gbar_Kdrax =   0.400
    }
// overwrite the general params:
	for i=0,1 NoR[i] { 
		g_pas = 0.4
	}
	for i=1,2 axon[i]{ //myelinated parts of the axon
		cm = 0.02
		gbar_NaFax =   0.010
		gbar_Kdrax =   0.013
	}
	soma { 
       insert NaFsd
	   gbar_NaFsd =   0.070 
       insert Kdrsd
       gbar_Kdrsd =   0.170
	   insert Ca
	   gbar_Ca = 0.001
	   insert Cad
	   phi_Cad = 24
	   beta_Cad = 0.001
	   insert Kahp
	   gbar_Kahp = 0.0008
	   insert Kc
	   gbar_Kc = 0.020
	   insert Ka
	   gbar_Ka = 0.0005
	   }
	forsec level1 { // distal basal dend
	   insert Ca
	   gbar_Ca = 0.001
	   insert Cad
	   phi_Cad = 148
	   beta_Cad = 0.05
	   insert Kahp
	   gbar_Kahp = 0.0008
	   insert Kc
	   gbar_Kc = 0.004
	   }
    forsec level2 { // middle basal dend
	   insert Ca
	   gbar_Ca = 0.001
	   insert Cad
	   phi_Cad = 164
	   beta_Cad = 0.05
	   insert Kahp
	   gbar_Kahp = 0.0008
	   insert Kc
	   gbar_Kc = 0.004
	   insert Ka
	   gbar_Ka = 0.0005
	   }
    forsec level3 { // proximal basal dend
       insert NaFsd
	   gbar_NaFsd =   0.001
       insert Kdrsd
       gbar_Kdrsd =   0.015
	   insert Ca
	   gbar_Ca = 0.001
	   insert Cad
	   phi_Cad = 123
	   beta_Cad = 0.05
	   insert Kahp
	   gbar_Kahp = 0.0008
	   insert Kc
	   gbar_Kc = 0.008
	   insert Ka
	   gbar_Ka = 0.0005
	   }
 // level4 is soma
 // level5 is shaft
    shaft { 
       insert NaFsd
	   gbar_NaFsd =   0.003
       insert Kdrsd
       gbar_Kdrsd =   0.020
	   insert Ca
	   gbar_Ca = 0.001
	   insert Cad
	   phi_Cad = 18
	   beta_Cad = 0.05
	   insert Kahp
	   gbar_Kahp = 0.0008
	   insert Kc
	   gbar_Kc = 0.008
	   insert Ka
	   gbar_Ka = 0.0005
	   }
    forsec level6 {
       insert NaFsd
	   gbar_NaFsd =   0.003
       insert Kdrsd
       gbar_Kdrsd =   0.020
	   insert Ca
	   gbar_Ca = 0.001
	   insert Cad
	   phi_Cad = 29
	   beta_Cad = 0.05
	   insert Kahp
	   gbar_Kahp = 0.0008
	   insert Kc
	   gbar_Kc = 0.008
	   insert Ka
	   gbar_Ka = 0.0005
	   }
	forsec level7 {
	   insert Ca
	   gbar_Ca = 0.002
	   insert Cad
	   phi_Cad = 37
	   beta_Cad = 0.05
	   insert Kahp
	   gbar_Kahp = 0.0008
	   insert Kc
	   gbar_Kc = 0.004
	   }
	forsec level8 {
	   insert Ca
	   gbar_Ca = 0.003
	   insert Cad
	   phi_Cad = 13
	   beta_Cad = 0.05
	   insert Kahp
	   gbar_Kahp = 0.0008
	   insert Kc
	   gbar_Kc = 0.012
	   }
	forsec level9 {
	   insert Ca
	   gbar_Ca = 0.003
	   insert Cad
	   phi_Cad = 16
	   beta_Cad = 0.05
	   insert Kahp
	   gbar_Kahp = 0.0008
	   insert Kc
	   gbar_Kc = 0.012
	   }
	forsec level10 {
	   insert Ca
	   gbar_Ca = 0.001
	   insert Cad
	   phi_Cad = 25
	   beta_Cad = 0.05
	   insert Kahp
	   gbar_Kahp = 0.0008
	   insert Kc
	   gbar_Kc = 0.004
	   }
	forsec level11 {
	   insert Ca
	   gbar_Ca = 0.001
	   insert Cad
	   phi_Cad = 25
	   beta_Cad = 0.05
	   insert Kahp
	   gbar_Kahp = 0.0008
	   insert Kc
	   gbar_Kc = 0.004
	   }
// reversal potentials
	forsec axonal {
       ek =  -90.
       e_pas =  -65.
       ena =   50.
	}
	forsec soma_dendrites {
       ek =  -80.
       e_pas = -65.
       if (ismembrane("NaFsd")) { ena =   50. }
	}
	forall { 
		insert extracellular
		xraxial =  1e+09
		xraxial[1] =  1e+09
		xg = 1e+09
		xg[1] = 1e+09
		xc = 0
		xc[1] = 0
		e_extracellular = 0 
	}
}
proc biophys_inhomo(){}
proc position() { local i
  soma for i = 0, n3d()-1 {
    pt3dchange(i, $1-x+x3d(i), $2-y+y3d(i), $3-z+z3d(i), diam3d(i))
  }
  x = $1  y = $2  z = $3
}
obfunc connect2target() { localobj nc //$o1 target point process, optional $o2 returned NetCon
  soma nc = new NetCon(&v(1), $o1)
  nc.threshold = 10
  if (numarg() == 2) { $o2 = nc } // for backward compatibility
  return nc
}
proc synapses() {}

// adds steady current to soma
// $1 is the current in nA, "-" for hyperpolarizing, "+" for depolarizing
proc setcurrentbias_soma(){ local i, area_soma 
 soma { 
      area_soma = PI*diam*L*1e-8  // unts converted [mcm2->cm2], area_soma is the area of a soma membrane
      if (ismembrane("bias")!=1) {insert bias}
	  amp_bias = -$1*1e-6/area_soma //note conversion [nA->mA] by 1e-6 and change of sign
    }
}

proc setcurrentbias_AIS(){ local i, area_section 
 ais { 
      area_section = PI*diam*L*1e-8  // unts converted [mcm2->cm2], area_section is the area of a section membrane
      if (ismembrane("bias")!=1) {insert bias}
	  amp_bias = -$1*1e-6/area_section //note conversion [nA->mA] by 1e-6 and change of sign
    }
}

proc double_dend_cond() {
   spine_area_multiplier = 2.0
   forsec dendrites {
        if (ismembrane("NaFsd")) { gbar_NaFsd *= spine_area_multiplier }
        if (ismembrane("Kc")) { gbar_Kc *= spine_area_multiplier }
        if (ismembrane("Kdrsd")) { gbar_Kdrsd *= spine_area_multiplier }
        if (ismembrane("Ka")) { gbar_Ka *= spine_area_multiplier }
		if (ismembrane("Kahp")) { gbar_Kahp *= spine_area_multiplier }
        if (ismembrane("Ca")) { gbar_Ca *= spine_area_multiplier }
        if (ismembrane("pas")) { g_pas *= spine_area_multiplier }
        cm = cm * spine_area_multiplier
   }
 }
proc reduceCa() {
	forsec all {
		if (ismembrane("Ca")) { gbar_Ca *= $1 }
	}
}

proc setindexXY() {
	x_index = $1
	y_index = $2
}
obfunc connect2targetSoma() { localobj nc //$o1 target point process, optional $o2 returned NetCon
  soma nc = new NetCon(&v(1), $o1)
  nc.threshold = -10
  if (numarg() == 2) { $o2 = nc } // for backward compatibility
  return nc
}
obfunc connect2targetSomaSpikelet() { localobj nc //$o1 target point process, optional $o2 returned NetCon
  soma nc = new NetCon(&v(1), $o1)
  nc.threshold = -55
  if (numarg() == 2) { $o2 = nc } // for backward compatibility
  return nc
}

obfunc connect2targetCollat() { localobj nc //$o1 target point process, optional $o2 returned NetCon
  collat[0] nc = new NetCon(&v(1), $o1)
  nc.threshold = 0
  if (numarg() == 2) { $o2 = nc } // for backward compatibility
  return nc
}

proc rotateOY(){ local alpha, i, xcenter, zcenter
// rotate cell along axis OY (vertical), argument in radians
	alpha = $1
	soma { xcenter = x3d(0)	zcenter = z3d(0) }
	forsec all {
	for i = 0, n3d()-1 {
		pt3dchange(i, xcenter + (x3d(i) - xcenter)*cos(alpha) - (z3d(i) - zcenter)*sin(alpha), y3d(i), zcenter + (x3d(i) - xcenter)*sin(alpha) + (z3d(i) - zcenter)*cos(alpha), diam3d(i))
    }
	}
}
endtemplate pyramidalCA1

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