Rhesus Monkey Layer 3 Pyramidal Neurons: V1 vs PFC (Amatrudo, Weaver et al. 2012)

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Accession:144553
Whole-cell patch-clamp recordings and high-resolution 3D morphometric analyses of layer 3 pyramidal neurons in in vitro slices of monkey primary visual cortex (V1) and dorsolateral granular prefrontal cortex (dlPFC) revealed that neurons in these two brain areas possess highly distinctive structural and functional properties. ... Three-dimensional reconstructions of V1 and dlPFC neurons were incorporated into computational models containing Hodgkin-Huxley and AMPA- and GABAA-receptor gated channels. Morphology alone largely accounted for observed passive physiological properties, but led to AP firing rates that differed more than observed empirically, and to synaptic responses that opposed empirical results. Accordingly, modeling predicts that active channel conductances differ between V1 and dlPFC neurons. The unique features of V1 and dlPFC neurons are likely fundamental determinants of area-specific network behavior. The compact electrotonic arbor and increased excitability of V1 neurons support the rapid signal integration required for early processing of visual information. The greater connectivity and dendritic complexity of dlPFC neurons likely support higher level cognitive functions including working memory and planning.
Reference:
1 . Amatrudo JM, Weaver CM, Crimins JL, Hof PR, Rosene DL, Luebke JI (2012) Influence of highly distinctive structural properties on the excitability of pyramidal neurons in monkey visual and prefrontal cortices. J Neurosci 32:13644-60 [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; Prefrontal cortex (PFC);
Cell Type(s): Neocortex L2/3 pyramidal GLU cell;
Channel(s): I N; I K;
Gap Junctions:
Receptor(s):
Gene(s):
Transmitter(s):
Simulation Environment: NEURON;
Model Concept(s): Activity Patterns; Influence of Dendritic Geometry; Detailed Neuronal Models; Electrotonus; Conductance distributions; Vision;
Implementer(s): Weaver, Christina [christina.weaver at fandm.edu];
Search NeuronDB for information about:  Neocortex L2/3 pyramidal GLU cell; I N; I K;
/
V1_PFC_ModelDB
README
kvz_nature.mod *
naz_nature.mod *
vsource.mod *
actionPotentialPlayer.hoc *
add_axon.hoc
analyticFunctions.hoc *
analyze_EPSC.m
aux_procs.hoc
batchrun.hoc
custominit.hoc
define_PFC.hoc
electro_procs.hoc *
figOptions.hoc
fixnseg.hoc *
init_model.hoc
init_PFC.hoc
Jul16IR3f_fromSWCthenManual_Nov22-11.hoc
load_scripts.hoc *
main_fig10_pfc.hoc
main_fig10_v1baseline.hoc
main_fig10_v1tuned.hoc
main_fig9_pfcElec.hoc
main_fig9_v1Elec.hoc
main_PFC-ApBas_fig11epsc.hoc
main_PFC-ApBas_fig12ipsc.hoc
main_V1-ApBas_fig11epsc.hoc
main_V1-ApBas_fig12ipsc.hoc
May3IR2t_ImportFromSWCthenManual_Aug19-11.hoc
measureMeanAtten.hoc
mosinit.hoc
PFC-V1_AddSynapses.hoc
plot_seClamp_i.ses
plot_seClamp_IPSC.ses
read_EPSCsims_mdb.m
read_IPSCsims_mdb.m
readcell.hoc
readNRNbin_Vclamp.m
rigPFCmod.ses
synTweak.hoc
vsrc.ses
                            
/******  These lines are copied from ~/research/Neuron/shared/hoc/syn_params.hoc  *******************/

//define these now, redefine when used
objref stimEsyndend[10], connEsyndend[10], Esyndend[10]
objref stimIsyndend[10], connIsyndend[10], Isyndend[10]
objref stimEsynsoma[1], connEsynsoma[1], Esynsoma[1]
objref stimIsynsoma[1], connIsynsoma[1], Isynsoma[1]
objref stimEsynprox[1], connEsynprox[1], Esynprox[1]
objref stimIsynprox[1], connIsynprox[1], Isynprox[1]
objref stimEsyndist[1], connEsyndist[1], Esyndist[1]
objref stimIsyndist[1], connIsyndist[1], Isyndist[1]

EXTHZ = 1 //Hz //10 //
INTHZ = 5.5 //Hz //65 //
NOISE = 1

ISI_EXC = 1000/INTHZ
ISI_INH = 1000/INTHZ
START_EXC = 50
START_INH = 50

//densities of AMPA, GABA are per 100um^2, 
GABAaDENDDEN=1//10 //density of GABAa at dendrites 
AMPADENDDEN=1//60 //density of AMPA at dendrites 
TOTAL_GABAa_g=491.4 //nS
TOTAL_AMPA_g =5898 //nS 


SEEDNUM = 1
SEED = 6514302

NSEG= 27
NSEG_SPINE=3 

OFFSET=50 //start 
ONSET=50  //end of applying events
NOISE=1
CURRINJ=0.03 // current injected
DT = 0.025
TSTOP = 1000 //
PLOTSMS=4


// These parameter values are modified from Traub, Contreras, et al. 2005:  
// Single-column thalamocortical network model exhibiting gamma oscillations,
// sleep spindles and epileptogenic bursts.  J Neurophysiol 93(4):  2194-2232,
// (2005).  PMID:  15525801.  See Appendix B, starting on page 2224.
//
// These are from superficial pyramid parameters reported in that reference.
// Note some alternative conductances reported for different connection types,
// e.g. pyramid->pyramid AMPA connections vs. stellate->pyramid connections.
// See Traub et al. to see which connections use which conductance values.


//
// values for PFC model from Amatrudo, Weaver et al (2012)
//
  AMPA_tau1 = 0.15  //(ms), time constant for AMPA
  AMPA_tau2 = 4     //(ms), 

  AMPA_Esyn = 0  //reversal potential 
  AMPA_g=0.00054 // change in conductance

//
// revised AMPA_g for V1 model (low gAMPA) from Amatrudo, Weaver et al (2012)
//
//  AMPA_g=0.00017 // (uS) 1.0nS // change in conductance;  other connections used 0.25 nS, 0.5 nS


  GABAa_g=0.001  // 1.0nS // change in conductance; other connections used 0.05 nS, 0.1 nS
  GABAa_Esyn = -81 //reversal potential for superficial pyramids; other neuron types in the model used -75 mV
  GABAa_tau1 = 6.0 //  
  GABAa_tau2 = 6.0 //  

/**** update, Jan 23 2012:  Jennie found that the amplitude of PSCs for V1 cells are about HALF what they are for PFC.  

Mean amplitude: 12.5 pA PFC;  6 pA V1
Rise times were approximately:  PFC:  1.6 ms;  1.1 ms for V1
Half width: PFC  7 ms;  5 ms for V1
Decay Time:  PFC 7 ms; 5 ms for V1.

So if we use the SAME synaptic parameters for both cells, the model and data differ substantially.

****/


START_EXC = 100

/*********************************************

    input	$o1	SectionList where synapses are needed
		$o2	vector of synapse locations
		$3	starting time of excitation
		$4	timing between subsequent PSPs
*********************************************/
func AddSynapses() { local cnt, nSyn, DELAY, ST_EXC, stopTime

    nSyn = $o2.size()
    ST_EXC = $3
    DELAY  = $4

    objref stimEsyndend[nSyn], connEsyndend[nSyn], Esyndend[nSyn]

    cnt = 0
    forsec $o1 {
	Esyndend[cnt] = new Exp2Syn($o2.x[cnt])
	Esyndend[cnt].tau1=AMPA_tau1  //--- ms decay time constatn
	Esyndend[cnt].tau2=AMPA_tau2  //--- ms decay time constatn
	Esyndend[cnt].e=AMPA_Esyn //-- mV reversal potential

	stimEsyndend[cnt]= new NetStim($o2.x[cnt])
	stimEsyndend[cnt].interval=  1e9
	stimEsyndend[cnt].start= $3 + cnt*DELAY
	stimEsyndend[cnt].noise=0
	stimEsyndend[cnt].number=1

        connEsyndend[cnt] = new NetCon(stimEsyndend[cnt],Esyndend[cnt],-55,0,AMPA_g)

        cnt += 1
    }
    stopTime = START_EXC+(nSyn+2)*DELAY
    printf("Added %d synapses, final time %d\n",nSyn,START_EXC + nSyn*DELAY)

    return stopTime
}

objref sref

/*********************************************

    input	$1	total number of excitatory apical synapses
    		$2	total number of excitatory basal synapses
		$3	starting time of excitation
		$4	timing between subsequent PSPs
		$o5	SectionList to which sections are added
		$o6	vector of locations

*********************************************/
func AddExcSynapses_byNumber() { local cnt, nSyn, DELAY, ST_EXC, stopTime, SAspacing, totSA, SAcnt, numSec, cntSec, xloc

    nASyn   = $1
    nBSyn   = $2
    nSyn = nASyn + nBSyn

    ST_EXC = $3
    DELAY  = $4

    objref stimEsyndend[nSyn], connEsyndend[nSyn], Esyndend[nSyn]
    soma { distance(0) }
    $o5 = new SectionList()
    $o6 = new Vector()

    // apical synapses first
    if( nASyn > 0 ) {

        totSA = 0
        forsec apical {
	    for(x) {
	        totSA += area(x)
            }
        }

        SAspacing = totSA / nASyn
        printf("Spacing for apical EXC synapses:  every %g = %g / %d microns ^ 2\n",SAspacing,totSA,nASyn)
        SAcnt = 0
        cnt = 0

        /***  no excitatory synapses added to soma ***/
        numSec = 0
        forsec apical { numSec += 1 }

        cntSec = 0
        forsec apical {
            cntSec += 1
	    for(x) {
	        SAcnt += area(x)
	        while( SAcnt >= SAspacing ) {
		    // add synapse here

   		    sref = new SectionRef()
		    $o5.append()
		    $o6.append(x)

		    //printf("SynLoc[%d] %s(%g) Dist %g\tCutoff %g > %g, total %g out of %g\n",cnt,secname(),x,distance(x), SAcnt,SAspacing,SAcnt+cnt*SAspacing,totSA)
		    Esyndend[cnt] = new Exp2Syn(x)
		    Esyndend[cnt].tau1=AMPA_tau1  //--- ms decay time constatn
		    Esyndend[cnt].tau2=AMPA_tau2  //--- ms decay time constatn
		    Esyndend[cnt].e=AMPA_Esyn //-- mV reversal potential

		    stimEsyndend[cnt]= new NetStim(x)
		    stimEsyndend[cnt].interval=  1e9
		    stimEsyndend[cnt].start= $3 + cnt*DELAY
		    stimEsyndend[cnt].noise=0
		    stimEsyndend[cnt].number=1

	            connEsyndend[cnt] = new NetCon(stimEsyndend[cnt],Esyndend[cnt],-55,0,AMPA_g)

	            cnt += 1

		    SAcnt -= SAspacing
	        }
    	    }
            // if it's the end of the SectionList and we still have a synapse to add, put it here.
	    if( cnt < nASyn && cntSec == numSec ) {
	        //printf("---Added %d synapses, need a total of %d\n",cnt,nSyn)
	        while( cnt < nASyn ) {
	            xloc = 1

 		    // add synapse here

		    sref = new SectionRef()
		    $o5.append()
		    $o6.append(xloc)

		    //printf("\t\tSynLoc[%d] %s(%g) Dist %g\n",cnt,secname(),xloc,distance(xloc))
		    Esyndend[cnt] = new Exp2Syn(xloc)
		    Esyndend[cnt].tau1=AMPA_tau1  //--- ms decay time constatn
		    Esyndend[cnt].tau2=AMPA_tau2  //--- ms decay time constatn
		    Esyndend[cnt].e=AMPA_Esyn //-- mV reversal potential

		    stimEsyndend[cnt]= new NetStim(xloc)
		    stimEsyndend[cnt].interval=  1e9
		    stimEsyndend[cnt].start= $3 + cnt*DELAY
		    stimEsyndend[cnt].noise=0
		    stimEsyndend[cnt].number=1

	            connEsyndend[cnt] = new NetCon(stimEsyndend[cnt],Esyndend[cnt],-55,0,AMPA_g)

	            cnt += 1
	        }
            }
        }
        printf("Finished with apical dendrites, added %d synapses, %g SA till next synapse\n",cnt,SAcnt)

    } else { printf("No apical synapses added.\n") }

    // now basal synapses
    if( nBSyn > 0 ) {

        totSA = 0
        forsec basal {
	    for(x) {
	        totSA += area(x)
            }
        }

        SAspacing = totSA / nBSyn
        printf("Spacing for basal EXC synapses:  every %g = %g / %d microns ^ 2\n",SAspacing,totSA,nBSyn)
        SAcnt = 0
        // note: we do NOT reinitialize 'cnt' here.

        numSec = 0
        forsec basal { numSec += 1 }

        cntSec = 0
        forsec basal {
            cntSec += 1
	    for(x) {
	        SAcnt += area(x)
	        while( SAcnt >= SAspacing ) {
		    // add synapse here

		    sref = new SectionRef()
		    $o5.append()
		    $o6.append(x)

		    //printf("SynLoc[%d] %s(%g) Dist %g\tCutoff %g > %g, total %g out of %g\n",cnt,secname(),x,distance(x), SAcnt,SAspacing,SAcnt+cnt*SAspacing,totSA)
		    Esyndend[cnt] = new Exp2Syn(x)
		    Esyndend[cnt].tau1=AMPA_tau1  //--- ms decay time constatn
		    Esyndend[cnt].tau2=AMPA_tau2  //--- ms decay time constatn
		    Esyndend[cnt].e=AMPA_Esyn //-- mV reversal potential

		    stimEsyndend[cnt]= new NetStim(x)
		    stimEsyndend[cnt].interval=  1e9
		    stimEsyndend[cnt].start= $3 + cnt*DELAY
		    stimEsyndend[cnt].noise=0
		    stimEsyndend[cnt].number=1

	            connEsyndend[cnt] = new NetCon(stimEsyndend[cnt],Esyndend[cnt],-55,0,AMPA_g)

	            cnt += 1

		    SAcnt -= SAspacing
	        }
    	    }
            // if it's the end of the SectionList and we still have a synapse to add, put it here.
	    if( cnt < nSyn && cntSec == numSec ) {
	        //printf("---Added %d synapses, need a total of %d\n",cnt,nSyn)
	        while( cnt < nSyn ) {
	            xloc = 1

		    // add synapse here

		    sref = new SectionRef()
		    $o5.append()
		    $o6.append(xloc)

		    //printf("\t\tSynLoc[%d] %s(%g) Dist %g\n",cnt,secname(),xloc,distance(xloc))
		    Esyndend[cnt] = new Exp2Syn(xloc)
		    Esyndend[cnt].tau1=AMPA_tau1  //--- ms decay time constatn
		    Esyndend[cnt].tau2=AMPA_tau2  //--- ms decay time constatn
		    Esyndend[cnt].e=AMPA_Esyn //-- mV reversal potential

		    stimEsyndend[cnt]= new NetStim(xloc)
		    stimEsyndend[cnt].interval=  1e9
		    stimEsyndend[cnt].start= $3 + cnt*DELAY
		    stimEsyndend[cnt].noise=0
		    stimEsyndend[cnt].number=1

	            connEsyndend[cnt] = new NetCon(stimEsyndend[cnt],Esyndend[cnt],-55,0,AMPA_g)

	            cnt += 1
	        }
            }
        }
        printf("Finished with basal dendrites, added %d synapses, %g SA till next synapse\n",cnt,SAcnt)

    } else {  printf("No basal synapses added.\n")  }


    printf("Added %d excitatory synapses, final time %d\n",nSyn,START_EXC + nSyn*DELAY)

    return nSyn
}



/*********************************************

    input	$1	total number of inhibitory synapses
		$2	maximal distance from soma for dendritic inhib synapses
		$3	starting time of excitation
		$4	timing between subsequent PSPs
		$o5	SectionList to which sections are added
		$o6	vector of locations
		$7	count of sections added

*********************************************/
func AddInhSynapses() { local cnt, nSyn, DELAY, ST_EXC, stopTime, SAspacing, totSA, SAcnt, numSec, cntSec, xloc

    nSyn   = $1
    ST_EXC = $3
    DELAY  = $4

    totSA = countSA($2)
    SAspacing = totSA / nSyn
    printf("Spacing for INH synapses:  every %g = %g / %d microns ^ 2\n",SAspacing,totSA,nSyn)

    objref stimIsyndend[nSyn], connIsyndend[nSyn], Isyndend[nSyn]

    soma { distance(0) }

    SAcnt = 0
    cnt = 0
    $o5 = new SectionList()
    $o6 = new Vector()

    forsec "soma" { 
	for(x) {
	    SAcnt += area(x)
	    while( SAcnt >= SAspacing ) {
		sref = new SectionRef()
		$o5.append()
		$o6.append(x)

		// add synapse here
		//printf("SynLoc %s(%g)\tCutoff %g > %g, total %g out of %g\n",secname(),x,SAcnt,SAspacing,SAcnt+cnt*SAspacing,totSA)
		Isyndend[cnt] = new Exp2Syn(x)  	// PICK UP HERE!!!! 
print "\tAdded Exp2Syn"
		Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
print "\tset tau1"
		Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
print "\tset tau2"
		Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential
print "\tset Esyn"

		stimIsyndend[cnt]= new NetStim(x)
print "\tAdded NetStim"
		stimIsyndend[cnt].interval=  1e9
		stimIsyndend[cnt].start= $3 + cnt*DELAY
		stimIsyndend[cnt].noise=0
		stimIsyndend[cnt].number=1
print "\tset NetStim params"

	        connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)
print "\tAdded NetCon"

        	cnt += 1

		SAcnt -= SAspacing

	    }

	}
    }
    printf("Finished with soma, added %d synapses, %g SA till next synapse\n",cnt,SAcnt)

    numSec = 0
    forsec dendritic { numSec += 1 }

    cntSec = 0
    forsec dendritic {
        cntSec += 1
	for(x) {
	    if( distance(x) > $2 ) continue

	    SAcnt += area(x)
	    while( SAcnt >= SAspacing ) {
		// add synapse here

		sref = new SectionRef()
		$o5.append()
		$o6.append(x)

		//printf("SynLoc[%d] %s(%g) Dist %g\tCutoff %g > %g, total %g out of %g\n",cnt,secname(),x,distance(x), SAcnt,SAspacing,SAcnt+cnt*SAspacing,totSA)
		Isyndend[cnt] = new Exp2Syn(x)
		Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		stimIsyndend[cnt]= new NetStim(x)
		stimIsyndend[cnt].interval=  1e9
		stimIsyndend[cnt].start= $3 + cnt*DELAY
		stimIsyndend[cnt].noise=0
		stimIsyndend[cnt].number=1

	        connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	        cnt += 1

		SAcnt -= SAspacing
	    }
    	}
        // if it's the end of the SectionList and we still have a synapse to add, put it here.
	if( cnt < nSyn && cntSec == numSec ) {
	    //printf("---Added %d synapses, need a total of %d\n",cnt,nSyn)
	    while( cnt < nSyn ) {
	        xloc = 1

		// add synapse here

		sref = new SectionRef()
		$o5.append()
		$o6.append(xloc)

		//printf("\t\tSynLoc[%d] %s(%g) Dist %g\n",cnt,secname(),xloc,distance(xloc))
		Isyndend[cnt] = new Exp2Syn(xloc)
		Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		stimIsyndend[cnt]= new NetStim(xloc)
		stimIsyndend[cnt].interval=  1e9
		stimIsyndend[cnt].start= $3 + cnt*DELAY
		stimIsyndend[cnt].noise=0
		stimIsyndend[cnt].number=1

	        connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	        cnt += 1
	    }
        }
    }
    printf("Finished with dendrites, added %d synapses, %g SA till next synapse\n",cnt,SAcnt)

    stopTime = START_EXC+(nSyn+2)*DELAY
    printf("Added %d inhibitory synapses, final time %d\n",nSyn,START_EXC + nSyn*DELAY)

    $7 = cnt

    return stopTime
}





/*********************************************

    input	$1	total number of inhibitory synapses
		$2	maximal distance from soma for dendritic inhib synapses
		$3	starting time of excitation
		$4	timing between subsequent PSPs
		$o5	SectionList to which sections are added
		$o6	vector of locations
		$7	count of sections added
		$8	proportion of all synapses to distribute proximally

    Distribute a proportion of the total number of inhibitory synapse close to 
    the soma.  Then, distribute the rest uniformly across the whole soma and 
    dendrites.

*********************************************/
func AddInhSynapses_ProxAndUni() { local cnt, nSyn, DELAY, ST_EXC, stopTime, SAspacing, totSA, prxSA, SAcnt, numSec, cntSec, xloc, nClose, nFar

    nSyn   = $1
    ST_EXC = $3
    DELAY  = $4
    nClose = int(nSyn * $8 + 0.5 )
    nFar   = nSyn - nClose 


    objref stimIsyndend[nSyn], connIsyndend[nSyn], Isyndend[nSyn]
    $o5 = new SectionList()
    $o6 = new Vector()

    soma { distance(0) }

    numSec = 0
    forsec dendritic { numSec += 1 }

    /**** STEP 1:  distribute a proportion the synapses on the soma & proximal dendrites.  ****/

    prxSA = countSA($2)
    SAspacing = prxSA / nClose
    printf("Spacing for proximal INH synapses:  every %g = %g / %d microns ^ 2\n",SAspacing,prxSA,nClose)

    SAcnt = 0
    cnt = 0
    forsec "soma" { 
	for(x) {
	    SAcnt += area(x)
	    while( SAcnt >= SAspacing ) {
		sref = new SectionRef()
		$o5.append()
		$o6.append(x)

		// add synapse here
		//printf("SOMA:  SynLoc %s(%g)\tCutoff %g > %g, total %g out of %g\n",secname(),x,SAcnt,SAspacing,SAcnt+cnt*SAspacing,totSA)
		Isyndend[cnt] = new Exp2Syn(x)  	
		Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		stimIsyndend[cnt]= new NetStim(x)
		stimIsyndend[cnt].interval=  1e9
		stimIsyndend[cnt].start= $3 + cnt*DELAY
		stimIsyndend[cnt].noise=0
		stimIsyndend[cnt].number=1

	        connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

        	cnt += 1

		SAcnt -= SAspacing

	    }

	}
    }
    printf("Finished with soma, added %d synapses, %g SA till next synapse\n",cnt,SAcnt)

    cntSec = 0
    forsec dendritic {
        cntSec += 1
	for(x) {
	    if( distance(x) > $2 ) continue

	    SAcnt += area(x)
	    while( SAcnt >= SAspacing ) {
		// add synapse here

		sref = new SectionRef()
		$o5.append()
		$o6.append(x)

		//printf("PROX:  SynLoc[%d] %s(%g) Dist %g\tCutoff %g > %g, total %g out of %g\n",cnt,secname(),x,distance(x), SAcnt,SAspacing,SAcnt+cnt*SAspacing,totSA)
		Isyndend[cnt] = new Exp2Syn(x)
		Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		stimIsyndend[cnt]= new NetStim(x)
		stimIsyndend[cnt].interval=  1e9
		stimIsyndend[cnt].start= $3 + cnt*DELAY
		stimIsyndend[cnt].noise=0
		stimIsyndend[cnt].number=1

	        connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	        cnt += 1

		SAcnt -= SAspacing
	    }
    	}
    }
    printf("Finished with proximal dendrites, added %d synapses, %g SA till next synapse\n",cnt,SAcnt)


    /**** STEP 2:  distribute remaining synapses througout medial and distal dendrites.  ****/

    if( nFar > 0 ) {

        totSA = countSA(1e9)
        SAspacing = (totSA - prxSA ) / nFar
        printf("Spacing for distal INH synapses:  every %g = %g / %d microns ^ 2\n",SAspacing,(totSA-prxSA),nFar)

        SAcnt = 0
        cntSec = 0
        forsec dendritic {
            cntSec += 1
	    for(x) {
	        if( distance(x) <= $2 ) continue

   	        SAcnt += area(x)
	        while( SAcnt >= SAspacing ) {
		    // add synapse here

 		    sref = new SectionRef()
		    $o5.append()
		    $o6.append(x)

		    //printf("DISTAL:  SynLoc[%d] %s(%g) Dist %g\tCutoff %g > %g, total %g out of %g\n",cnt,secname(),x,distance(x), SAcnt,SAspacing,SAcnt+cnt*SAspacing,(totSA-prxSA))
		    Isyndend[cnt] = new Exp2Syn(x)
		    Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		    Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		    Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		    stimIsyndend[cnt]= new NetStim(x)
		    stimIsyndend[cnt].interval=  1e9
		    stimIsyndend[cnt].start= $3 + cnt*DELAY
		    stimIsyndend[cnt].noise=0
		    stimIsyndend[cnt].number=1

	            connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	            cnt += 1

		    SAcnt -= SAspacing
	        }
    	    }
            // if it's the end of the SectionList and we still have a synapse to add, put it here.
	    if( cnt < nSyn && cntSec == numSec ) {
	        //printf("---Added %d synapses, need a total of %d\n",cnt,nSyn)
	        while( cnt < nSyn ) {
	            xloc = 1

		    // add synapse here

		    sref = new SectionRef()
		    $o5.append()
		    $o6.append(xloc)

		    //printf("\t\tSynLoc[%d] END %s(%g) Dist %g\n",cnt,secname(),xloc,distance(xloc))
		    Isyndend[cnt] = new Exp2Syn(xloc)
		    Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		    Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		    Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		    stimIsyndend[cnt]= new NetStim(xloc)
		    stimIsyndend[cnt].interval=  1e9
		    stimIsyndend[cnt].start= $3 + cnt*DELAY
		    stimIsyndend[cnt].noise=0
		    stimIsyndend[cnt].number=1

	            connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	            cnt += 1
	        }
            }
        }
        printf("Finished with dendrites, added %d synapses, %g SA till next synapse\n",cnt,SAcnt)    
    }


    stopTime = START_EXC+(nSyn+2)*DELAY
    printf("Added %d inhibitory synapses, final time %d\n",nSyn,START_EXC + nSyn*DELAY)

    $7 = cnt

    return stopTime
}









/*********************************************

    input	$1	base density of inhibitory synapses
		$2	maximal distance from soma for dendritic inhib synapses
		$3	starting time of excitation
		$4	timing between subsequent PSPs
		$o5	SectionList to which sections are added
		$o6	vector of locations
		$7	count of sections added
		$8	inhib synapse density on proximal spines
		$9	inhib synapse density on distal spines

    Specify a baseline density of inhibitory synapse throughout dendrites, and 
    extra density terms to represent inhib synapses that would have been on 
    dendritic spines.  Distribute these uniformly in their respective regions.

*********************************************/
func AddInhSynapses_ProxDist_byDensity() { local cnt, nSyn, DELAY, ST_EXC, stopTime, Lspacing, totSA, prxSA, Lcnt, numSec, cntSec, xloc, nClose, nFar, baseDens, prxSpDens, dstSpDens, totL, prxL, dstL, lastX

    baseDens = $1
    ST_EXC = $3
    DELAY  = $4

    prxL = find_totLen($2)
    totL = find_totLen(1e9)
    dstL = totL - prxL
    prxSpDens = $8
    dstSpDens = $9

    nClose = int(prxL * (baseDens + prxSpDens) + 0.5 )
    nFar   = int(dstL * (baseDens + dstSpDens) + 0.5 )
    nSyn = nClose + nFar
    printf("Inh syn density:  %g base/t%g prox Spines/t%g dist Spines\n",baseDens,prxSpDens,dstSpDens)
    printf("Want a total of %d synapses: %d close and %d far from soma.\n",nSyn,nClose,nFar)



    objref stimIsyndend[nSyn], connIsyndend[nSyn], Isyndend[nSyn]
    $o5 = new SectionList()
    $o6 = new Vector()

    soma { distance(0) }

    numSec = 0
    forsec dendritic { numSec += 1 }

    /**** STEP 1:  distribute a proportion the synapses on the soma & proximal dendrites.  ****/

    Lspacing = prxL / nClose
    printf("Spacing for proximal INH synapses:  every %g = %g / %d microns of length\n",Lspacing,prxL,nClose)

    Lcnt = 0
    cnt = 0
    forsec "soma" { 
	for(x) {
	    if( x==0 || x == 1 ) continue

	    Lcnt += L / nseg
	    while( Lcnt >= Lspacing ) {
		sref = new SectionRef()
		$o5.append()
		$o6.append(x)

		// add synapse here
		//printf("SOMA:  SynLoc %s(%g)\tCutoff %g > %g, total %g out of %g\n",secname(),x,Lcnt,Lspacing,Lcnt+cnt*Lspacing,totL)
		Isyndend[cnt] = new Exp2Syn(x)  	
		Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		stimIsyndend[cnt]= new NetStim(x)
		stimIsyndend[cnt].interval=  1e9
		stimIsyndend[cnt].start= $3 + cnt*DELAY
		stimIsyndend[cnt].noise=0
		stimIsyndend[cnt].number=1

	        connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

        	cnt += 1

		Lcnt -= Lspacing

	    }

	}
    }
    printf("Finished with soma, added %d synapses, %g L till next synapse\n",cnt,Lcnt)

    cntSec = 0
    forsec dendritic {
        cntSec += 1
	for(x) {
	    if( distance(x) > $2 ) { continue }
	    //if( distance(x) > $2 ) { printf("---- Skip %s\t",secname() ) continue }
	    if( x==0 || x==1 ) continue

	    Lcnt += L / nseg
	    //printf("\n\t+++\t%s(%g) L add %g -> Lcnt %g\n",secname(),x,L/nseg,Lcnt)

	    while( Lcnt >= Lspacing ) {

                if( cnt > nClose ) { printf("\ttrying to add synapse %d\n",cnt) }
		// add synapse here

		sref = new SectionRef()
		$o5.append()
		$o6.append(x)

		//printf("PROX:  SynLoc[%d] %s(%g) Dist %g\tCutoff %g > %g, total %g out of %g\n",cnt,secname(),x,distance(x), Lcnt,Lspacing,Lcnt+cnt*Lspacing,prxL)
		Isyndend[cnt] = new Exp2Syn(x)
		Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential
 
		stimIsyndend[cnt]= new NetStim(x)
		stimIsyndend[cnt].interval=  1e9
		stimIsyndend[cnt].start= $3 + cnt*DELAY
		stimIsyndend[cnt].noise=0
		stimIsyndend[cnt].number=1

	        connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	        cnt += 1

		Lcnt -= Lspacing
	    } 
    	}
    }
    printf("Finished with proximal dendrites, added %d synapses, %g length till next synapse\n",cnt,Lcnt)


    /**** STEP 2:  distribute remaining synapses througout medial and distal dendrites.  ****/

    //Lcnt = 0

    if( nFar > 0 ) {	// continue here.

        Lspacing = dstL / nFar
        printf("Spacing for distal INH synapses:  every %g = %g / %d microns of length\n",Lspacing,dstL,nFar)

        //Lcnt = 0
        cntSec = 0
        forsec dendritic {
            cntSec += 1
	    for(x) {
	        if( x==0 || x==1 ) continue
	        if( distance(x) <= $2 ) continue

   	        Lcnt += L / nseg
	        while( Lcnt >= Lspacing && cnt < nSyn ) {
		    // add synapse here

 		    sref = new SectionRef()
		    $o5.append()
		    $o6.append(x)

		    //printf("DISTAL:  SynLoc[%d] %s(%g) Dist %g\tCutoff %g > %g, total %g out of %g\n",cnt,secname(),x,distance(x), Lcnt,Lspacing,Lcnt+cnt*Lspacing,dstL)
		    Isyndend[cnt] = new Exp2Syn(x)
		    Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		    Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		    Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		    stimIsyndend[cnt]= new NetStim(x)
		    stimIsyndend[cnt].interval=  1e9
		    stimIsyndend[cnt].start= $3 + cnt*DELAY
		    stimIsyndend[cnt].noise=0
		    stimIsyndend[cnt].number=1

	            connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	            cnt += 1

		    Lcnt -= Lspacing
	        }
    	    }
            // if it's the end of the SectionList and we still have a synapse to add, put it here.
	    if( cnt < nSyn && cntSec == numSec ) {
	        //printf("---Added %d synapses, need a total of %d\n",cnt,nSyn)
	        while( cnt < nSyn ) {
	            xloc = 1

		    // add synapse here

		    sref = new SectionRef()
		    $o5.append()
		    $o6.append(xloc)

		    //printf("\t\tSynLoc[%d] END %s(%g) Dist %g\n",cnt,secname(),xloc,distance(xloc))
		    Isyndend[cnt] = new Exp2Syn(xloc)
		    Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		    Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		    Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		    stimIsyndend[cnt]= new NetStim(xloc)
		    stimIsyndend[cnt].interval=  1e9
		    stimIsyndend[cnt].start= $3 + cnt*DELAY
		    stimIsyndend[cnt].noise=0
		    stimIsyndend[cnt].number=1

	            connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	            cnt += 1
	        }
            }
        }
        printf("Finished with dendrites, added %d synapses, %g length till next synapse\n",cnt,Lcnt)    
    }


    stopTime = START_EXC+(nSyn+2)*DELAY
    printf("Added %d inhibitory synapses, final time %d\n",nSyn,START_EXC + nSyn*DELAY)

    $7 = cnt

    return nSyn
    //return stopTime
}







/*********************************************

    input	$1	base density of inhibitory synapses
		$2	maximal distance from soma for dendritic inhib synapses
		$3	starting time of excitation
		$4	timing between subsequent PSPs
		$o5	SectionList to which sections are added
		$o6	vector of locations
		$7	count of sections added
		$8	inhib synapse density on proximal spines
		$9	inhib synapse density on distal spines
		$10	if 0, do only apical; if 1, do only basal; else do both.

    Specify a baseline density of inhibitory synapse throughout dendrites, and 
    extra density terms to represent inhib synapses that would have been on 
    dendritic spines.  Distribute these uniformly in their respective regions.

*********************************************/
func AddInhSynapses_ProxDist_byDensityAB() { local cnt, nSyn, DELAY, ST_EXC, stopTime, Lspacing, totSA, prxSA, Lcnt, numSec, cntSec, xloc, nClose, nFar, baseDens, prxSpDens, dstSpDens, totL, prxL, dstL, lastX

    baseDens = $1
    ST_EXC = $3
    DELAY  = $4

    if( $10 == 0 ) {
        prxL = find_totApicLen($2)
        totL = find_totApicLen(1e9)
    } else {
        if( $10 == 1 ) {
            prxL = find_totBasLen($2)
            totL = find_totBasLen(1e9)
        } else {
            prxL = find_totLen($2)
            totL = find_totLen(1e9)
        }
    }
    dstL = totL - prxL

    prxSpDens = $8
    dstSpDens = $9

    nClose = int(prxL * (baseDens + prxSpDens) + 0.5 )
    nFar   = int(dstL * (baseDens + dstSpDens) + 0.5 )
    nSyn = nClose + nFar
    printf("Inh syn density:  %g base/t%g prox Spines/t%g dist Spines\n",baseDens,prxSpDens,dstSpDens)
    printf("Want a total of %d synapses: %d close and %d far from soma.\n",nSyn,nClose,nFar)



    objref stimIsyndend[nSyn], connIsyndend[nSyn], Isyndend[nSyn]
    $o5 = new SectionList()
    $o6 = new Vector()

    soma { distance(0) }

    numSec = 0
    if( $10 == 0 ) { forsec apical { numSec += 1 }
    } else {
        if( $10 ==1 ) {  forsec basal { numSec += 1 }
        } else {
            forsec dendritic { numSec += 1 }
        }
    }

    /**** STEP 1:  distribute a proportion the synapses on the soma & proximal dendrites.  ****/

    Lspacing = prxL / nClose
    printf("Spacing for proximal INH synapses:  every %g = %g / %d microns of length\n",Lspacing,prxL,nClose)

    Lcnt = 0
    cnt = 0

    /****  this section is the same regardless of whether we consider apical or basal only, or all dendrites ****/
    forsec "soma" { 
	for(x) {
	    if( x==0 || x == 1 ) continue

	    Lcnt += L / nseg
	    while( Lcnt >= Lspacing ) {
		sref = new SectionRef()
		$o5.append()
		$o6.append(x)

		// add synapse here
		//printf("SOMA:  SynLoc %s(%g)\tCutoff %g > %g, total %g out of %g\n",secname(),x,Lcnt,Lspacing,Lcnt+cnt*Lspacing,totL)
		Isyndend[cnt] = new Exp2Syn(x)  	
		Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		stimIsyndend[cnt]= new NetStim(x)
		stimIsyndend[cnt].interval=  1e9
		stimIsyndend[cnt].start= $3 + cnt*DELAY
		stimIsyndend[cnt].noise=0
		stimIsyndend[cnt].number=1

	        connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

        	cnt += 1

		Lcnt -= Lspacing

	    }

	}
    }
    printf("Finished with soma, added %d synapses, %g L till next synapse\n",cnt,Lcnt)

    cntSec = 0


    if( $10 != 1 ) {
        forsec apical {
            cntSec += 1
	    for(x) {
	        if( distance(x) > $2 ) { continue }
  	        if( x==0 || x==1 ) continue

	        Lcnt += L / nseg

	        while( Lcnt >= Lspacing ) {

                    if( cnt > nClose ) { printf("\ttrying to add synapse %d\n",cnt) }
		    // add synapse here

	  	    sref = new SectionRef()
		    $o5.append()
		    $o6.append(x)

	  	    Isyndend[cnt] = new Exp2Syn(x)
		    Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		    Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		    Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential
 
		    stimIsyndend[cnt]= new NetStim(x)
		    stimIsyndend[cnt].interval=  1e9
		    stimIsyndend[cnt].start= $3 + cnt*DELAY
		    stimIsyndend[cnt].noise=0
		    stimIsyndend[cnt].number=1

	            connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	            cnt += 1

	  	    Lcnt -= Lspacing
	        } 
    	    }
        }
    } 

    if( $10 != 0 ) {
        forsec basal {
            cntSec += 1
	    for(x) {
	        if( distance(x) > $2 ) { continue }
  	        if( x==0 || x==1 ) continue

	        Lcnt += L / nseg

	        while( Lcnt >= Lspacing ) {

                    if( cnt > nClose ) { printf("\ttrying to add synapse %d\n",cnt) }
		    // add synapse here

	  	    sref = new SectionRef()
		    $o5.append()
		    $o6.append(x)

	  	    Isyndend[cnt] = new Exp2Syn(x)
		    Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		    Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		    Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential
 
		    stimIsyndend[cnt]= new NetStim(x)
		    stimIsyndend[cnt].interval=  1e9
		    stimIsyndend[cnt].start= $3 + cnt*DELAY
		    stimIsyndend[cnt].noise=0
		    stimIsyndend[cnt].number=1

	            connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	            cnt += 1

	  	    Lcnt -= Lspacing
	        } 
    	    }
        }
    } 

    
    printf("Finished with proximal dendrites, added %d synapses, %g length till next synapse\n",cnt,Lcnt)


    /**** STEP 2:  distribute remaining synapses througout medial and distal dendrites.  ****/

    if( nFar > 0 ) {	// continue here.

        Lspacing = dstL / nFar
        printf("Spacing for distal INH synapses:  every %g = %g / %d microns of length\n",Lspacing,dstL,nFar)

        cntSec = 0

	if( $10 == 0  ) {
            forsec apical {
                cntSec += 1
	        for(x) {
	            if( x==0 || x==1 ) continue
	            if( distance(x) <= $2 ) continue

     	            Lcnt += L / nseg
	            while( Lcnt >= Lspacing && cnt < nSyn ) {
		        // add synapse here

 		        sref = new SectionRef()
		        $o5.append()
		        $o6.append(x)

		        Isyndend[cnt] = new Exp2Syn(x)
		        Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		        Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		        Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		        stimIsyndend[cnt]= new NetStim(x)
		        stimIsyndend[cnt].interval=  1e9
		        stimIsyndend[cnt].start= $3 + cnt*DELAY
		        stimIsyndend[cnt].noise=0
		        stimIsyndend[cnt].number=1

	                connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	                cnt += 1

		        Lcnt -= Lspacing
	            }
    	        }
                // if it's the end of the SectionList and we still have a synapse to add, put it here.
	        if( cnt < nSyn && cntSec == numSec ) {
	            while( cnt < nSyn ) {
	                xloc = 1

		        // add synapse here

		        sref = new SectionRef()
		        $o5.append()
		        $o6.append(xloc)

		        Isyndend[cnt] = new Exp2Syn(xloc)
		        Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		        Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		        Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		        stimIsyndend[cnt]= new NetStim(xloc)
		        stimIsyndend[cnt].interval=  1e9
		        stimIsyndend[cnt].start= $3 + cnt*DELAY
		        stimIsyndend[cnt].noise=0
		        stimIsyndend[cnt].number=1

	                connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	                cnt += 1
	            }
                }
            }
	}
	if( $10 == 1 ) {
            forsec basal {
                cntSec += 1
	        for(x) {
	            if( x==0 || x==1 ) continue
	            if( distance(x) <= $2 ) continue

     	            Lcnt += L / nseg
	            while( Lcnt >= Lspacing && cnt < nSyn ) {
		        // add synapse here

 		        sref = new SectionRef()
		        $o5.append()
		        $o6.append(x)

		        Isyndend[cnt] = new Exp2Syn(x)
		        Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		        Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		        Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		        stimIsyndend[cnt]= new NetStim(x)
		        stimIsyndend[cnt].interval=  1e9
		        stimIsyndend[cnt].start= $3 + cnt*DELAY
		        stimIsyndend[cnt].noise=0
		        stimIsyndend[cnt].number=1

	                connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	                cnt += 1

		        Lcnt -= Lspacing
	            }
    	        }
                // if it's the end of the SectionList and we still have a synapse to add, put it here.
	        if( cnt < nSyn && cntSec == numSec ) {
	            while( cnt < nSyn ) {
	                xloc = 1

		        // add synapse here

		        sref = new SectionRef()
		        $o5.append()
		        $o6.append(xloc)

		        Isyndend[cnt] = new Exp2Syn(xloc)
		        Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		        Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		        Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		        stimIsyndend[cnt]= new NetStim(xloc)
		        stimIsyndend[cnt].interval=  1e9
		        stimIsyndend[cnt].start= $3 + cnt*DELAY
		        stimIsyndend[cnt].noise=0
		        stimIsyndend[cnt].number=1

	                connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	                cnt += 1
	            }
                }
            }
	}
	if( ($10 != 0 ) && ($10 != 1) ) {
            forsec dendritic {
                cntSec += 1
	        for(x) {
	            if( x==0 || x==1 ) continue
	            if( distance(x) <= $2 ) continue

     	            Lcnt += L / nseg
	            while( Lcnt >= Lspacing && cnt < nSyn ) {
		        // add synapse here

 		        sref = new SectionRef()
		        $o5.append()
		        $o6.append(x)

		        Isyndend[cnt] = new Exp2Syn(x)
		        Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		        Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		        Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		        stimIsyndend[cnt]= new NetStim(x)
		        stimIsyndend[cnt].interval=  1e9
		        stimIsyndend[cnt].start= $3 + cnt*DELAY
		        stimIsyndend[cnt].noise=0
		        stimIsyndend[cnt].number=1

	                connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	                cnt += 1

		        Lcnt -= Lspacing
	            }
    	        }
                // if it's the end of the SectionList and we still have a synapse to add, put it here.
	        if( cnt < nSyn && cntSec == numSec ) {
	            while( cnt < nSyn ) {
	                xloc = 1

		        // add synapse here

		        sref = new SectionRef()
		        $o5.append()
		        $o6.append(xloc)

		        Isyndend[cnt] = new Exp2Syn(xloc)
		        Isyndend[cnt].tau1=GABAa_tau1  //--- ms decay time constatn
		        Isyndend[cnt].tau2=GABAa_tau2  //--- ms decay time constatn
		        Isyndend[cnt].e=GABAa_Esyn //-- mV reversal potential

		        stimIsyndend[cnt]= new NetStim(xloc)
		        stimIsyndend[cnt].interval=  1e9
		        stimIsyndend[cnt].start= $3 + cnt*DELAY
		        stimIsyndend[cnt].noise=0
		        stimIsyndend[cnt].number=1

	                connIsyndend[cnt] = new NetCon(stimIsyndend[cnt],Isyndend[cnt],-55,0,GABAa_g)

	                cnt += 1
	            }
                }
            }
	}

        printf("Finished with dendrites, added %d synapses, %g length till next synapse\n",cnt,Lcnt)    
    }


    stopTime = START_EXC+(nSyn+2)*DELAY
    printf("Added %d inhibitory synapses, final time %d\n",nSyn,START_EXC + nSyn*DELAY)

    $7 = cnt

    return nSyn
}





proc SynapsesOff() { local i

    for i = 0, $1-1 {
        stimEsyndend[i].start = 1e9
    }
}

proc SynapseOn() {
    stimEsyndend[$1].start = $2
}



/********
	input   $1	value of gAMPA
		$2	number of synapses
********/
proc adjust_gAMPA() {  local i

    for i = 0, $2-1 {
	connEsyndend[i].weight = $1
    }
}


/********
	input   $1	value of gAMPA
		$2	number of synapses
********/
proc adjust_gGABA() {  local i

    for i = 0, $2-1 {
	connIsyndend[i].weight = $1
    }
}



/********
	input   $1	value of tau1 for Exp2Syn
		$2	number of synapses
********/
proc adjust_tau1() {  local i

    for i = 0, $2-1 {
	Esyndend[i].tau1 = $1
    }
}



/********
	input   $1	value of tau1 for Exp2Syn
		$2	number of synapses
********/
proc adjust_tau2() {  local i

    for i = 0, $2-1 {
	Esyndend[i].tau2 = $1
    }
}




/********
	input   $1	value of tau1 for Exp2Syn
		$2	number of synapses
********/
proc adjust_Itau1() {  local i

    for i = 0, $2-1 {
	Isyndend[i].tau1 = $1
    }
}



/********
	input   $1	value of tau1 for Exp2Syn
		$2	number of synapses
********/
proc adjust_Itau2() {  local i

    for i = 0, $2-1 {
	Isyndend[i].tau2 = $1
    }
}


/******************
	input 	$1 	max distance to count, or -1 if full dendrites only should be counted
******************/
func countSA() { local sa

    sa = 0

    if( $1 >= 0 ) { forsec "soma" for(x) sa += area(x) }

    forsec dendritic {
	for(x) {
            if( $1< 0 ||  distance(x) <= $1 ) {
                //printf("%s\t%g\t%g\t%g\n",secname(), x, distance(x), area(x) )
		sa += area(x)
	    }
        }
    }

    return sa
}



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