Compartmentalization of GABAergic inhibition by dendritic spines (Chiu et al. 2013)

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Accession:143604
A spiny dendrite model supports the hypothesis that only inhibitory inputs on spine heads, not shafts, compartmentalizes inhibition of calcium signals to spine heads as seen in paired inhibition with back-propagating action potential experiments on prefrontal cortex layer 2/3 pyramidal neurons in mouse (Chiu et al. 2013).
Reference:
1 . Chiu CQ, Lur G, Morse TM, Carnevale NT, Ellis-Davies GC, Higley MJ (2013) Compartmentalization of GABAergic inhibition by dendritic spines. Science 340:759-62 [PubMed]
Model Information (Click on a link to find other models with that property)
Model Type: Synapse; Dendrite;
Brain Region(s)/Organism: Neocortex;
Cell Type(s): Neocortex L2/3 pyramidal GLU cell;
Channel(s): I Na,t; I L high threshold; I K;
Gap Junctions:
Receptor(s): GabaA;
Gene(s):
Transmitter(s):
Simulation Environment: NEURON;
Model Concept(s): Dendritic Action Potentials; Influence of Dendritic Geometry;
Implementer(s): Carnevale, Ted [Ted.Carnevale at Yale.edu]; Morse, Tom [Tom.Morse at Yale.edu];
Search NeuronDB for information about:  Neocortex L2/3 pyramidal GLU cell; GabaA; I Na,t; I L high threshold; I K;
// param3.hoc
/*
Contains the functions for param3.hoc which calculated these
simulations for changes in the 3 parameters mshift (in ca current),
ECl, gbar_ca

3. My calculation protocol details and
possible additions: For the following please send three traces

I) spine control Ca trace (protocol bAP)
II) spine Ca with inhib (bAPspineinhib)
III) neighboring spine Ca (bAPspineinhib)
II and II are acquired on same simulation run

IV) calculate also dCa, well actually
dCa(inhibited)/dCa(control) = ([Ca2+]peak,inh - B)/([Ca2+]peak,ctrl - B)
calculated from I), II):
dCa_ratio.txt
V) record also the dCa(inhibited), dCa(control) values separately
dCa_inhib.txt
dCa_ctrl.txt

a) mshift -15 mV, 0, 15 mV (code as m15, zero, 15)
b) ECl=-90, -70, -40 mV (code as m90, m70, m40)
c) gbar_ca 0.5x, 1x, 2x (code as 0p5x, 1x, 2x)

These protocols results spine0.dat, spine1.dat,  will fall under a
folder called param3 (parameter studies under Summary email's point
number 3 (the points 1 and 2 were calculated under the 5conditions
folder)):

param3/mshift_m15/[bAP, bAPspineinhib]
param3/mshift_zero/[bAP, bAPspineinhib]
param3/mshift_15/[bAP, bAPspineinhib]
param3/ECl_m90/[bAP, bAPspineinhib]
param3/ECl_m70/[bAP, bAPspineinhib]
param3/ECl_m40/[bAP, bAPspineinhib]
param3/gbar_ca0p5x/[bAP, bAPspineinhib]
param3/gbar_ca1x/[bAP, bAPspineinhib]
param3/gbar_ca2x/[bAP, bAPspineinhib]
The above 9 folders have two protocols each for a total of 18 protocols
in addition each folder
param3/X/
has files
dCa_ratio.txt (uses results in bAP, bAPspineinhib sub folders)
dCa_inhib.txt (ratio numerator, uses result from bAPspineinhib/spine1.txt)
dCa_ctrl.txt (ratio denominator, uses result from bAP/spine1.txt)

-----------
Additional work - translate matlab dCa_ratio.m to dCa_ratio.hoc, which
does the same thing in hoc.
-----------

The calcium and voltage trace files are stored in "proto_dir/file" 
where protocol directory proto_dir can be upto

inhibspine, inhibdend, inhibdend, bAP, bAPinhibspine, bAPinhibdend,
bAP1xinhibdend

and files can be as many as spine0_ca, spine1_ca, spine2_ca, dend_ca, spine0_v,
spine1_v, spine2, dend_v

however here unlike in FiveConditions.hoc, the protocols and files are limited to
X/bAP X/bAPspineinhib
and spine1_ca, spine0_ca where spine1 is the middle spine head, and spine0 is the
proximal spine head (receives a smaller bAP typically because of the slight increase
in bAP amplitude as a function of distance along the dendrite.
*/
strdef param_dir
strdef proto_dir
strdef filename
objref f // temporary file object
f = new File()

objref dend_v_vec, spine0_v_vec, spine1_v_vec, spine2_v_vec
objref dend_cai_vec, spine0_cai_vec, spine1_cai_vec, spine2_cai_vec
// tstop= 1 // test condition
// below are graphs (the 5 conditions turned into 6 with the addition
// of protocol bAP + 1x inhibition in dendrite
num_of_conditions = 18
objref  param3_v[num_of_conditions], param3_ca[num_of_conditions], param3_gates[num_of_conditions]

// add the activation and inactivation curves for the ca chan in spines and dend
objref m_ca_dend_vec, h_ca_dend_vec, m_ca_spine1_vec, h_ca_spine1_vec
objref  m_ca_spine0_vec, h_ca_spine0_vec

for i=0, num_of_conditions-1 {
  param3_v[i] = new Graph(0)
  param3_ca[i] = new Graph(0)
  param3_gates[i] = new Graph(0)
}
proto_num=0  // index into above graphs of protocol results

proc setup_vectors_to_record() {
// record voltages in the dendrite and the Spine[X].head X=0,1,2 (corresponds to paper spines 1 2 3)
  dend_v_vec = new Vector()
  dend_cai_vec= new Vector()

  spine0_v_vec= new Vector()
  spine1_v_vec= new Vector()
  spine2_v_vec= new Vector()
  spine0_cai_vec = new Vector()
  spine1_cai_vec = new Vector()
  spine2_cai_vec = new Vector()

  m_ca_dend_vec = new Vector()
  h_ca_dend_vec = new Vector()
  m_ca_spine1_vec = new Vector()
  h_ca_spine1_vec = new Vector()
  m_ca_spine0_vec = new Vector()
  h_ca_spine0_vec = new Vector()

  t_vec = new Vector()
  t_vec.indgen(0, tstop, dt)

  spinelocation=adjacent_shaft_x_loc // record the dendrite where Spine[1] is attached
  sprint(tmpstr, "dendrite.v(%f)", adjacent_shaft_x_loc) // used to be 180/dendrite.L
  dend_v_vec.label(tmpstr)
  spine0_v_vec.label("Spine[0].head.v(0.5)")
  spine1_v_vec.label("Spine[1].head.v(0.5)")
  spine2_v_vec.label("Spine[2].head.v(0.5)")

  sprint(tmpstr,"dendrite.cai(%g)", spinelocation) // dend. comp. adj. to spine
  dend_cai_vec.label(tmpstr)
  spine0_cai_vec.label("Spine[0].head.cai(0.5)")
  spine1_cai_vec.label("Spine[1].head.cai(0.5)")
  spine2_cai_vec.label("Spine[2].head.cai(0.5)")

  sprint(tmpstr,"dendrite.m_ca(%g)", spinelocation) // dend. comp. adj. to spine
  m_ca_dend_vec.label(tmpstr)
  sprint(tmpstr,"dendrite.h_ca(%g)", spinelocation) // dend. comp. adj. to spine
  h_ca_dend_vec.label(tmpstr)
  m_ca_spine1_vec.label("Spine[1].head.m_ca(0.5)")
  h_ca_spine1_vec.label("Spine[1].head.h_ca(0.5)")
  m_ca_spine0_vec.label("Spine[0].head.m_ca(0.5)")
  h_ca_spine0_vec.label("Spine[0].head.h_ca(0.5)")

  dend_v_vec.record(&dendrite.v(spinelocation), t_vec)
  dend_cai_vec.record(&dendrite.cai(spinelocation), t_vec)

  spine0_v_vec.record(&Spine[0].head.v(0.5), t_vec)
  spine1_v_vec.record(&Spine[1].head.v(0.5), t_vec)
  spine2_v_vec.record(&Spine[2].head.v(0.5), t_vec)

  spine0_cai_vec.record(&Spine[0].head.cai(0.5), t_vec)
  spine1_cai_vec.record(&Spine[1].head.cai(0.5), t_vec)
  spine2_cai_vec.record(&Spine[2].head.cai(0.5), t_vec)

  m_ca_dend_vec.record(&dendrite.m_ca(spinelocation), t_vec)
  h_ca_dend_vec.record(&dendrite.h_ca(spinelocation), t_vec)
  m_ca_spine1_vec.record(&Spine[1].head.m_ca(0.5), t_vec)
  h_ca_spine1_vec.record(&Spine[1].head.h_ca(0.5), t_vec)
  m_ca_spine0_vec.record(&Spine[0].head.m_ca(0.5), t_vec)
  h_ca_spine0_vec.record(&Spine[0].head.h_ca(0.5), t_vec)
}

proc  write_vectors() {
  sprint(tmpstr,"param3/%s/%s/spine0_cai.dat" , param_dir, proto_dir)
  write_vec(tmpstr,t_vec, spine0_cai_vec)
  sprint(tmpstr,"param3/%s/%s/spine1_cai.dat" , param_dir, proto_dir)
  write_vec(tmpstr,t_vec, spine1_cai_vec)
/* JUST THE ABOVE CALCIUM TRACES  
  sprint(tmpstr,"param3/%s/spine2_cai.dat" , proto_dir)
  write_vec(tmpstr,t_vec, spine2_cai_vec)
*/
  sprint(tmpstr,"param3/%s/%s/spine0_v.dat" , param_dir, proto_dir)
  write_vec(tmpstr,t_vec, spine0_v_vec)
  sprint(tmpstr,"param3/%s/%s/spine1_v.dat" , param_dir, proto_dir)
  write_vec(tmpstr,t_vec, spine1_v_vec)
/* AND (ABOVE) THEIR ASSOCIATED VOLTAGE TRACES
  sprint(tmpstr,"param3/%s/spine2_v.dat" , proto_dir)
  write_vec(tmpstr,t_vec, spine2_v_vec)
  
  sprint(tmpstr,"param3/%s/dend_v.dat" , proto_dir)
  write_vec(tmpstr,t_vec,dend_v_vec)
  sprint(tmpstr,"param3/%s/dend_cai.dat" , proto_dir)
  write_vec(tmpstr,t_vec, dend_cai_vec)

  sprint(tmpstr,"param3/%s/m_ca_dend_vec.dat" , proto_dir)
  write_vec(tmpstr,t_vec, m_ca_dend_vec)
  
  sprint(tmpstr,"param3/%s/h_ca_dend_vec.dat" , proto_dir)
  write_vec(tmpstr,t_vec, h_ca_dend_vec)

  sprint(tmpstr,"param3/%s/m_ca_spine1_vec.dat" , proto_dir)
  write_vec(tmpstr,t_vec, m_ca_spine1_vec)

  sprint(tmpstr,"param3/%s/h_ca_spine1_vec.dat" , proto_dir)
  write_vec(tmpstr,t_vec, h_ca_spine1_vec)

  sprint(tmpstr,"param3/%s/m_ca_spine0_vec.dat" , proto_dir)
  write_vec(tmpstr,t_vec, m_ca_spine0_vec)

  sprint(tmpstr,"param3/%s/h_ca_spine0_vec.dat" , proto_dir)
  write_vec(tmpstr,t_vec, h_ca_spine0_vec)
*/
  spine0_cai_vec.line(param3_ca[proto_num],t_vec,3,1)
  spine1_cai_vec.line(param3_ca[proto_num],t_vec,1,1)
/*
  spine2_cai_vec.line(param3_ca[proto_num],t_vec,2,1)
*/
  dend_cai_vec.line(param3_ca[proto_num],t_vec,5,1)
  // graphs
  param3_ca[proto_num].exec_menu("View = plot")
  sprint(tmpstr,"param3 (18 cases) cai under protocol %s %s" , param_dir, proto_dir)
  param3_ca[proto_num].label(.5,.95,tmpstr)
  param3_ca[proto_num].view(0, 0, 500 , 0.0011, 180, 280, 300, 200)
  
  spine0_v_vec.line(param3_v[proto_num],t_vec,3,1)
  spine1_v_vec.line(param3_v[proto_num],t_vec,1,1)
//  spine2_v_vec.line(param3_v[proto_num],t_vec,2,1)
  dend_v_vec.line(param3_v[proto_num],t_vec,5,1)
  param3_v[proto_num].exec_menu("View = plot")

  sprint(tmpstr,"param3 (18 cases) v under protocol %s %s" , param_dir, proto_dir)
  param3_v[proto_num].label(.5,.95,tmpstr)
  param3_v[proto_num].view(114, -70, 6 , 78, 200, 300, 300, 200)

  m_ca_spine0_vec.line(param3_gates[proto_num], t_vec,1, 1)
  h_ca_spine0_vec.line(param3_gates[proto_num], t_vec,2, 1)
  m_ca_spine1_vec.line(param3_gates[proto_num], t_vec,3, 1)
  h_ca_spine1_vec.line(param3_gates[proto_num], t_vec,4, 1)
  m_ca_dend_vec.line(param3_gates[proto_num], t_vec,5, 1)
  h_ca_dend_vec.line(param3_gates[proto_num], t_vec,6, 1)
  param3_gates[proto_num].exec_menu("View = plot")
  sprint(tmpstr,"param3 (18 cases) m_ca, h_ca under protocol %s %s" ,  param_dir, proto_dir)
  param3_gates[proto_num].label(.5,.95,tmpstr)
}

proc run_model() {
  print "model running with protocol ", param_dir,", ", proto_dir
  setup_vectors_to_record()
  init()
  run()
  write_vectors()
}

// Start of top-level script to run model with param3's 18 Conditions

// first switch to semi-high (0.05 is high) neck resistance state:
forsec "neck" {diam=0.07}
forsec "neck" {Ra=200}
// function is being broken up into 1)without and 2)with
// functions so that graphs can be made with the vectors while they are
// available.  These new functions immediately follow below
proc bAP_without_with_spineinhib() {  // run each param3 bAP 
  // case with and without spine inhibition
  // the NC[] index is defined: 0=dend, 1, 2, 3 is Spine[0, 1, 2]
  // spine_choice is 1, 2, or 3 whose default was 2 (middle spine) 
  // 1) bAP                    //////////////////////////////////

  // turn on bAP
  MultIClamp[0].number=1 // turn on current injection that creates bAP
  NC[spine_choice].weight = 0.0 // 0.0004 is 4 nS (0 spine synapse is off)

  proto_dir="bAP"
  run_model()
  proto_num=proto_num+1

  prestimulus_time=99.9
  prestimulus_index=1998 // the 1999th element
  // baseline B
  B=spine1_cai_vec.x[prestimulus_index]
  index_end = 150/dt // might be 3000 (dt=0.05) or 6000 (dt=0.025) to find
                     // an index after the first AP but hopefully before
                     // the second if another AP should occur
  Ca_peak_ctrl = spine1_cai_vec.c(0,index_end).max()
  // leave bAP on

  // 2) bAP + inhib in spine   //////////////////////////////////

  // bAP left on from above, now spine inihib turned back on
  NC[spine_choice].weight = 0.0004 // 4e-4 uS = 0.4 nS

  proto_dir="bAPspineinhib"
  run_model()
  Ca_peak_inhib = spine1_cai_vec.c(0, index_end).max()
  proto_num=proto_num+1
  dCa_ratio=(Ca_peak_inhib-B)/(Ca_peak_ctrl-B)

  sprint(tmpstr,"param3/%s/dCa_inhib.txt",param_dir)
  f.wopen(tmpstr)
  sprint(tmpstr,"%g\n",Ca_peak_inhib-B)
  f.printf(tmpstr)
  f.close()

  sprint(tmpstr,"param3/%s/dCa_ctrl.txt",param_dir)
  f.wopen(tmpstr)
  sprint(tmpstr,"%g\n",Ca_peak_ctrl-B)
  f.printf(tmpstr)
  f.close()

  sprint(tmpstr,"param3/%s/dCa_ratio.txt",param_dir)
  f.wopen(tmpstr)
  sprint(tmpstr,"%g\n",dCa_ratio)
  f.printf(tmpstr)
  f.close()
}
proc bAP_without_spineinhib() {  // run each param3 bAP 
  // case without spine inhibition
  // the NC[] index is defined: 0=dend, 1, 2, 3 is Spine[0, 1, 2]
  // spine_choice is 1, 2, or 3 whose default was 2 (middle spine) 
  // 1) bAP                    //////////////////////////////////

  // turn on bAP
  MultIClamp[0].number=1 // turn on current injection that creates bAP
  NC[spine_choice].weight = 0.0 // 0.0004 is 4 nS (0 spine synapse is off)

  proto_dir="bAP"
  run_model()
  proto_num=proto_num+1

  prestimulus_time=99.9
  prestimulus_index=1998 // the 1999th element
  // baseline B
  B=spine1_cai_vec.x[prestimulus_index]
  index_end = 150/dt  // two limit to first AP hopefully
  Ca_peak_ctrl = spine1_cai_vec.c(0,index_end).max()
  // leave bAP on
} // end of bAP (alone) without inhibition

proc bAP_with_spineinhib() {  // run each param3 bAP 
  // case with spine inhibition
  // the NC[] index is defined: 0=dend, 1, 2, 3 is Spine[0, 1, 2]
  // spine_choice is 1, 2, or 3 whose default was 2 (middle spine) 
  // 2) bAP + inhib in spine   //////////////////////////////////

  // bAP left on from above, now spine inihib turned back on
  NC[spine_choice].weight = 0.0004 // 4e-4 uS = 0.4 nS

  proto_dir="bAPspineinhib"
  run_model()
  index_end = 150/dt // to limit to first AP hopefully
  Ca_peak_inhib = spine1_cai_vec.c(0, index_end).max()
  proto_num=proto_num+1
  dCa_ratio=(Ca_peak_inhib-B)/(Ca_peak_ctrl-B)

  sprint(tmpstr,"param3/%s/dCa_inhib.txt",param_dir)
  f.wopen(tmpstr)
  sprint(tmpstr,"%g\n",Ca_peak_inhib-B)
  f.printf(tmpstr)
  f.close()

  sprint(tmpstr,"param3/%s/dCa_ctrl.txt",param_dir)
  f.wopen(tmpstr)
  sprint(tmpstr,"%g\n",Ca_peak_ctrl-B)
  f.printf(tmpstr)
  f.close()

  sprint(tmpstr,"param3/%s/dCa_ratio.txt",param_dir)
  f.wopen(tmpstr)
  sprint(tmpstr,"%g\n",dCa_ratio)
  f.printf(tmpstr)
  f.close()
}

/* *******************
This is the master controller section of the code where the
parameters are set to desired values and then the model
is run both with and without inhibition by calling the
function bAP_without_with_spineinhib()

During this process the following graphs and tables are generated

Graph panels of Ca traces for:

1) mshift_graph: mshift_m15 mshift_zero mshift_15
2) ECl_graph: ECl_m90 ECl_m70 ECl_m40
3) gbar_ca_graph: gbar_ca0p5x gbar_ca1x gbar_ca2x

Tables of dCa_ratio and also the numer/denominator that determine
dCa_ratio: The following vectors hold those dCa quantities under the
following conditions.

1) mshift_table_vec, mshift_num_vec, mshift_denom_vec: mshift_m15
mshift_zero mshift_15
2) ECl_table_vec, ECl_num_vec, ECl_denom_vec: ECl_m90 ECl_m70 ECl_m40
3) gbar_ca_table_vec, gbar_ca_num_vec, gbar_ca_denom_vec: gbar_ca0p5x
gbar_ca1x gbar_ca2x

At the end of this script the tables are printed at the oc> prompt.
**********************************************
*/
objref mshift_hbox, ECl_hbox, gbar_ca_hbox
objref mshift_graph[3], ECl_graph[3], gbar_ca_graph[3]
objref mshift_table_vec, mshift_table_header_vec
objref mshift_num_vec, mshift_denom_vec
objref ECl_table_vec, ECl_table_header_vec
objref ECl_num_vec, ECl_denom_vec
objref gbar_ca_table_vec,gbar_ca_table_header_vec
objref gbar_ca_num_vec, gbar_ca_denom_vec

mshift_hbox = new HBox()
ECl_hbox = new HBox()
gbar_ca_hbox = new HBox()

for i=0,2 {
  mshift_hbox.intercept(1)
  mshift_graph[i] = new Graph()
  mshift_hbox.intercept(0)
  ECl_hbox.intercept(1)
  ECl_graph[i] = new Graph()
  ECl_hbox.intercept(0)
  gbar_ca_hbox.intercept(1)
  gbar_ca_graph[i] = new Graph()
  gbar_ca_hbox.intercept(0)
}

mshift_table_vec = new Vector()
mshift_table_header_vec = new Vector()
mshift_num_vec = new Vector()
mshift_denom_vec = new Vector()
ECl_table_vec = new Vector()
ECl_table_header_vec = new Vector()
ECl_num_vec = new Vector()
ECl_denom_vec = new Vector()
gbar_ca_table_vec = new Vector()
gbar_ca_table_header_vec = new Vector()
gbar_ca_num_vec = new Vector()
gbar_ca_denom_vec = new Vector()

proto_num = 0 // initialize counter which is subsequently
// incremented in bAP_without_with_spineinhib()

// param3/mshift_m15/[bAP, bAPspineinhib]
param_dir = "mshift_m15"
mshift_ca = -15

bAP_without_spineinhib()

spine0_cai_vec.line(mshift_graph[0],t_vec,1,0)
spine1_cai_vec.line(mshift_graph[0],t_vec,1,0)
mshift_denom_vec.append(Ca_peak_ctrl-B)
mshift_table_header_vec.append(mshift_ca)

bAP_with_spineinhib()

spine0_cai_vec.line(mshift_graph[0],t_vec,3,0)
spine1_cai_vec.line(mshift_graph[0],t_vec,2,0)
mshift_num_vec.append(Ca_peak_inhib-B)
sprint(tmpstr,"cai vs t in two spine heads, mshift=%g, black ctrl, red inhib, blue adj. spine", mshift_ca)
mshift_graph[0].label(0.4,0.9,tmpstr)
mshift_graph[0].exec_menu("View = plot")

// param3/mshift_zero/[bAP, bAPspineinhib]
param_dir = "mshift_zero"
mshift_ca = 0

bAP_without_spineinhib()

spine0_cai_vec.line(mshift_graph[1],t_vec,1,0)
spine1_cai_vec.line(mshift_graph[1],t_vec,1,0)
mshift_denom_vec.append(Ca_peak_ctrl-B)
mshift_table_header_vec.append(mshift_ca)

bAP_with_spineinhib()

spine0_cai_vec.line(mshift_graph[1],t_vec,3,0)
spine1_cai_vec.line(mshift_graph[1],t_vec,2,0)
mshift_num_vec.append(Ca_peak_inhib-B)
sprint(tmpstr,"cai vs t in two spine heads, mshift=%g, black ctrl, red inhib, blue adj. spine", mshift_ca)
mshift_graph[1].label(0.4,0.9,tmpstr)
mshift_graph[1].exec_menu("View = plot")

// param3/mshift_15/[bAP, bAPspineinhib]
param_dir = "mshift_15"
mshift_ca = 15

bAP_without_spineinhib()

spine0_cai_vec.line(mshift_graph[2],t_vec,1,0)
spine1_cai_vec.line(mshift_graph[2],t_vec,1,0)
mshift_denom_vec.append(Ca_peak_ctrl-B)
mshift_table_header_vec.append(mshift_ca)

bAP_with_spineinhib()

spine0_cai_vec.line(mshift_graph[2],t_vec,3,0)
spine1_cai_vec.line(mshift_graph[2],t_vec,2,0)
mshift_num_vec.append(Ca_peak_inhib-B)
sprint(tmpstr,"cai vs t in two spine heads, mshift=%g, black ctrl, red inhib, blue adj. spine", mshift_ca)
mshift_graph[2].label(0.4,0.9,tmpstr)
mshift_graph[2].exec_menu("View = plot")
mshift_hbox.map()
print "mshift tables header"
{mshift_table_header_vec.printf()}
print "dCa_inhib (numerators)"
mshift_num_vec.printf()
print "dCa_ctrl (denominators)"
mshift_denom_vec.printf()
print "dCa_ratio"
mshift_num_vec.c.div(mshift_denom_vec).printf()
print " "
mshift_ca=0 // reset to default for future simulations not related to shifting ca activation

// param3/ECl_m90/[bAP, bAPspineinhib]
param_dir = "ECl_m90"
Exp2Syn[2].e = -90 // middle spines gabaa synapse reversal potential

bAP_without_spineinhib()

spine0_cai_vec.line(ECl_graph[0],t_vec,1,0)
spine1_cai_vec.line(ECl_graph[0],t_vec,1,0)
ECl_denom_vec.append(Ca_peak_ctrl-B)
ECl_table_header_vec.append(Exp2Syn[2].e)

bAP_with_spineinhib()

spine0_cai_vec.line(ECl_graph[0],t_vec,3,0)
spine1_cai_vec.line(ECl_graph[0],t_vec,2,0)
ECl_num_vec.append(Ca_peak_inhib-B)
sprint(tmpstr,"cai vs t in two spine heads, ECl=%g, black ctrl, red inhib, blue adj. spine", Exp2Syn[2].e)
ECl_graph[0].label(0.4,0.9,tmpstr)
ECl_graph[0].exec_menu("View = plot")

// param3/ECl_m70/[bAP, bAPspineinhib]
param_dir = "ECl_m70"
Exp2Syn[2].e = -70 // middle spines gabaa synapse reversal potential

bAP_without_spineinhib()

spine0_cai_vec.line(ECl_graph[1],t_vec,1,0)
spine1_cai_vec.line(ECl_graph[1],t_vec,1,0)
ECl_denom_vec.append(Ca_peak_ctrl-B)
ECl_table_header_vec.append(Exp2Syn[2].e)

bAP_with_spineinhib()

spine0_cai_vec.line(ECl_graph[1],t_vec,3,0)
spine1_cai_vec.line(ECl_graph[1],t_vec,2,0)
ECl_num_vec.append(Ca_peak_inhib-B)
sprint(tmpstr,"cai vs t in two spine heads, ECl=%g, black ctrl, red inhib, blue adj. spine", Exp2Syn[2].e)
ECl_graph[1].label(0.4,0.9,tmpstr)
ECl_graph[1].exec_menu("View = plot")

// param3/ECl_m40/[bAP, bAPspineinhib]
param_dir = "ECl_m40"
Exp2Syn[2].e = -40 // middle spines gabaa synapse reversal potential

bAP_without_spineinhib()

spine0_cai_vec.line(ECl_graph[2],t_vec,1,0)
spine1_cai_vec.line(ECl_graph[2],t_vec,1,0)
ECl_denom_vec.append(Ca_peak_ctrl-B)
ECl_table_header_vec.append(Exp2Syn[2].e)

bAP_with_spineinhib()

spine0_cai_vec.line(ECl_graph[2],t_vec,3,0)
spine1_cai_vec.line(ECl_graph[2],t_vec,2,0)
ECl_num_vec.append(Ca_peak_inhib-B)
sprint(tmpstr,"cai vs t in two spine heads, ECl=%g, black ctrl, red inhib, blue adj. spine", Exp2Syn[2].e)
ECl_graph[2].label(0.4,0.9,tmpstr)
ECl_graph[2].exec_menu("View = plot")
ECl_hbox.map()
print "ECl tables header"
{ECl_table_header_vec.printf()}
print "dCa_inhib (numerators)"
ECl_num_vec.printf()
print "dCa_ctrl (denominators)"
ECl_denom_vec.printf()
print "dCa_ratio"
ECl_num_vec.c.div(ECl_denom_vec).printf()
print " "

Exp2Syn[2].e = -70 // reset to default middle spines gabaa synapse reversal potential

// param3/gbar_ca0p5x/[bAP, bAPspineinhib]
// First store the default vaules:
default_soma_gbar_ca = soma.gbar_ca(0.5)
default_dend_gbar_ca = dendrite.gbar_ca(0.5)

param_dir = "gbar_ca0p5x"
soma for(x,0) gbar_ca(x) = default_soma_gbar_ca * 0.5
dendrite for(x,0) gbar_ca(x) = default_dend_gbar_ca * 0.5
forsec "Spine" gbar_ca(0.5) = default_dend_gbar_ca * 0.5

bAP_without_spineinhib()

spine0_cai_vec.line(gbar_ca_graph[0],t_vec,1,0)
spine1_cai_vec.line(gbar_ca_graph[0],t_vec,1,0)
gbar_ca_denom_vec.append(Ca_peak_ctrl-B)
gbar_ca_table_header_vec.append(0.5)

bAP_with_spineinhib()

spine0_cai_vec.line(gbar_ca_graph[0],t_vec,3,0)
spine1_cai_vec.line(gbar_ca_graph[0],t_vec,2,0)
gbar_ca_num_vec.append(Ca_peak_inhib-B)
sprint(tmpstr,"cai vs t in two spine heads, gbar_ca=%gx, black ctrl, red inhib, blue adj. spine", .5)
gbar_ca_graph[0].label(0.4,0.9,tmpstr)
gbar_ca_graph[0].exec_menu("View = plot")

// param3/gbar_ca1x/[bAP, bAPspineinhib]
param_dir = "gbar_ca1x"
soma for(x,0) gbar_ca(x) = default_soma_gbar_ca
dendrite for(x,0) gbar_ca(x) = default_dend_gbar_ca
forsec "Spine" gbar_ca(0.5) = default_dend_gbar_ca
bAP_without_spineinhib()

spine0_cai_vec.line(gbar_ca_graph[1],t_vec,1,0)
spine1_cai_vec.line(gbar_ca_graph[1],t_vec,1,0)
gbar_ca_denom_vec.append(Ca_peak_ctrl-B)
gbar_ca_table_header_vec.append(1)

bAP_with_spineinhib()

spine0_cai_vec.line(gbar_ca_graph[1],t_vec,3,0)
spine1_cai_vec.line(gbar_ca_graph[1],t_vec,2,0)
gbar_ca_num_vec.append(Ca_peak_inhib-B)
sprint(tmpstr,"cai vs t in two spine heads, gbar_ca=%gx, black ctrl, red inhib, blue adj. spine", 1)
gbar_ca_graph[1].label(0.4,0.9,tmpstr)
gbar_ca_graph[1].exec_menu("View = plot")

// param3/gbar_ca2x/[bAP, bAPspineinhib]
param_dir = "gbar_ca2x"
soma for(x,0) gbar_ca(x) = default_soma_gbar_ca * 2
dendrite for(x,0) gbar_ca(x) = default_dend_gbar_ca * 2
forsec "Spine" gbar_ca(0.5) = default_dend_gbar_ca * 2

bAP_without_spineinhib()

spine0_cai_vec.line(gbar_ca_graph[2],t_vec,1,0)
spine1_cai_vec.line(gbar_ca_graph[2],t_vec,1,0)
gbar_ca_denom_vec.append(Ca_peak_ctrl-B)
gbar_ca_table_header_vec.append(2)

bAP_with_spineinhib()

spine0_cai_vec.line(gbar_ca_graph[2],t_vec,3,0)
spine1_cai_vec.line(gbar_ca_graph[2],t_vec,2,0)
gbar_ca_num_vec.append(Ca_peak_inhib-B)
sprint(tmpstr,"cai vs t in two spine heads, gbar_ca=%gx, black ctrl, red inhib, blue adj. spine", 2)
gbar_ca_graph[2].label(0.4,0.9,tmpstr)
gbar_ca_graph[2].exec_menu("View = plot")
gbar_ca_hbox.map()
print "gbar_ca tables header"
{gbar_ca_table_header_vec.printf()}
print "dCa_inhib (numerators)"
gbar_ca_num_vec.printf()
print "dCa_ctrl (denominators)"
gbar_ca_denom_vec.printf()
print "dCa_ratio"
gbar_ca_num_vec.c.div(gbar_ca_denom_vec).printf()
print " "

// restore to default
soma for(x,0) gbar_ca(x) = default_soma_gbar_ca
dendrite for(x,0) gbar_ca(x) = default_dend_gbar_ca
forsec "Spine" gbar_ca(0.5) = default_dend_gbar_ca

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