Pyramidal Neuron Deep: attenuation in dendrites (Stuart, Spruston 1998)

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Accession:3677
Stuart, G. and Spruston, N. Determinants of voltage attenuation in neocortical pyramidal neuron dendrites. Journal of Neuroscience 18:3501-3510, 1998.
Reference:
1 . Stuart G, Spruston N (1998) Determinants of voltage attenuation in neocortical pyramidal neuron dendrites. J Neurosci 18:3501-10 [PubMed]
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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:
Cell Type(s): Neocortex L5/6 pyramidal GLU cell;
Channel(s): I h;
Gap Junctions:
Receptor(s):
Gene(s):
Transmitter(s):
Simulation Environment: NEURON;
Model Concept(s): Parameter Fitting;
Implementer(s): Spruston, Nelson [Spruston at Northwestern.edu];
Search NeuronDB for information about:  Neocortex L5/6 pyramidal GLU cell; I h;
proc init_params() {
	strdef somareadfile, dendreadfile
	somareadfile="ri22scs.asc"
	dendreadfile="ri22dcs.asc"
	somav="dend1[7].v(0.5)"    /* somatic recording site */
	dendv="dend1[134].v(0.01)"  /* dendritic recording site */
	/* initialize user-defined membrane parameters */
	celsius = 35
	global_ra=147   /* internal resistivity in ohm-cm */
	Cm=1.0          /* specific membrane capacitance in uF/cm^2 */
	Rm=20805        /* specific membrene resistivity in ohm-cm^2 */
	Vleak=-78 	/* -77 for qscale of 10 */
	Vrest=-78       /* resting potential in mV */
	qblock=1	/* fraction of q current blocked by Cs */
	nakblock=1
	spinescale=2.0  /* scale total area to account for spines */
	areascale=1.088663 /* scale total area by an additional factor */
	target=47.2     /* input resistance target */
	/* attenuation = 0.6251 */
	     /* target=29.0     adjusted for low Rm simulation */
	rmscale = 1241 /* good for Ri18 nusag -> low rm simulation */
  global_ra=200
  areascale=1.228847
  /* parameters for low Ri, nonuniform Rm simulation (init_nurm3) */
  /* short pulse fit not too bad */
	if (nurm==3) {
		global_ra=96
		rmsoma=35000
		rmend=7000
		rmhalfdis=400
		rmsteep=150
		areascale=1.060224
	}
  /* initq1 with init_pass */
	if (initq==1 && nurm<1 && qblock<1) {
		maxq=0.000085
		a0_q=0.0006
		b0_q=a0_q
		zeta_q=5
	}
  /* initq1 with init_nurm3 */
	if (initq==1 && nurm==3 && qblock<1) {
		Vleak=-78
		maxq=0.000068
		a0_q=0.0008
		b0_q=a0_q
		zeta_q=4
	}
  /* initq8 with init_pass */
	if (initq==8 && nurm<1 && qblock<1) {
		qscale=0.01
		maxq=0.0045
		qhalfdis=500
		qsteep=125
		a0_q=0.00022
		b0_q=a0_q
		zeta_q=4
	}
  /* initq8 with init_nurm3 */
	if (initq==8 && nurm==3 && qblock<1) {
		maxq=0.01
		qhalfdis=475
		qsteep=75
		a0_q=0.0002
		b0_q=a0_q
		zeta_q=4
		Vleak=-82
	}
}

proc init() { local dtsav
	finitialize(Vrest)
	fcurrent()
	if (writeout>0) {
		somasimvec.record(&dend1[13].v(0.5))
		dendsimvec.record(&dend1[232].v(0.99))
	}
	if (qblock<1) {
		dtsav=dt
		dt=10
		t=1e6
		printf("initializing")
		for i=0,40 {
			printf(".")
			fadvance()
		}
		printf("\n")
		dt=dtsav
		finitialize()
	}
	t=0
	ss_save()
}

proc advance() {
	fadvance()
	ss_subtract()
}

proc ss_save() {
	Vrestsoma=somaticv()
	Vrestdend=dendriticv()
	ss_subtract()
}

proc ss_subtract() {
	vsoma=somaticv()-Vrestsoma-76
	vdend=dendriticv()-Vrestdend-76
}

proc insert_pass() {
	print "Inserting passive membrane properties\n"
	forall {insert pas Ra=global_ra e_pas=Vleak}
}

proc insertq() {
	printf("Inserting sag conductance in all segments: %d pecent blocked\n",qblock*100)
	forall { insert qq gqbar_qq=0 }
}

proc init_pass() {
	global_ra=$1*100
	dendscale=spinescale*areascale
	somascale=areascale*1.0
	print "Initializing uniform passive membrane properties\n"
	forall { g_pas=1/(Rm/dendscale) cm=Cm*dendscale Ra=global_ra e_pas=Vleak}
	/* the following segments are NOT scaled for spines */
	/* these include the axon, soma, 1st 100 um of apical trunk, */
	/*  and the 1st 20 um of basals */
	/* the axon */
	axon { g_pas=1/(Rm/somascale) cm=Cm*somascale Ra=global_ra e_pas=Vleak}
	soma { g_pas=1/(Rm/somascale) cm=Cm*somascale Ra=global_ra e_pas=Vleak}
	/* the real soma */
	for i=0,9 {
		dend1[i] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	}
	/* the 1st 100 um of the apical trunk */
	for i=10,11 {
		dend1[i] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	}
	for i=35,38 {
		dend1[i] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	}
	dend1[46] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	dend1[49] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	/* the 1st 20 um of the basals */
	dend1[434] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	dend1[435] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	dend1[436] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	dend1[439] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	dend1[453] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	dend1[454] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	dend1[508] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	dend1[509] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	dend1[516] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	dend1[517] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	dend1[543] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	dend1[544] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	dend1[566] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	dend1[567] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
}

proc init_nurm() {
	dendscale=spinescale*areascale
	somascale=areascale*1.0
	/* rmscale is set in init_params */
	/* decrease rmscale to steepen decrease in Rm */
	/* as a function of distance (decreases to zero at 600 or so) */
	dis=0
	access dend1[7]
	area(0.5)
	distance()
	print "Initializing nonuniform passive membrane properties\n"
	forall { g_pas=1/(Rm/dendscale) cm=Cm*dendscale Ra=global_ra e_pas=Vleak}
	for i=8,433 {
		access dend1[i]
		dis=distance(0)
		dend1[i] {g_pas=1/((Rm/dendscale)*(1-dis/rmscale)) }
	}
	/* the following segments are NOT scaled for spines */
	/* these include the axon, soma, 1st 100 um of apical trunk, */
	/*  and the 1st 20 um of basals */
	/* the axon */
	axon { g_pas=1/(Rm/somascale) cm=Cm*somascale }
	soma { g_pas=1/(Rm/somascale) cm=Cm*somascale }
	/* the real soma */
	for i=0,9 {
		dend1[i] {g_pas=1/((Rm/somascale)*(1-dis/rmscale)) cm=Cm*somascale }
	}
	/* the 1st 100 um of the apical trunk */
	for i=10,11 {
		dend1[i] {g_pas=1/((Rm/somascale)*(1-dis/rmscale)) cm=Cm*somascale }
	}
	for i=35,36 {
		dend1[i] {g_pas=1/((Rm/somascale)*(1-dis/rmscale)) cm=Cm*somascale }
	}
	dend1[46] {g_pas=1/((Rm/somascale)*(1-dis/rmscale)) cm=Cm*somascale }
	dend1[49] {g_pas=1/((Rm/somascale)*(1-dis/rmscale)) cm=Cm*somascale }
	/* the 1st 20 um of the basals */
	for i=453,454 {
		dend1[i] {g_pas=1/(Rm/somascale) cm=Cm*somascale }
	}
	for i=434,436 {
		dend1[i] {g_pas=1/(Rm/somascale) cm=Cm*somascale }
	}
	dend1[439] {g_pas=1/(Rm/somascale) cm=Cm*somascale }
	for i=508,509 {
		dend1[i] {g_pas=1/(Rm/somascale) cm=Cm*somascale }
	}
	for i=516,517 {
		dend1[i] {g_pas=1/(Rm/somascale) cm=Cm*somascale }
	}
	for i=543,544 {
		dend1[i] {g_pas=1/(Rm/somascale) cm=Cm*somascale }
	}
	for i=566,567 {
		dend1[i] {g_pas=1/(Rm/somascale) cm=Cm*somascale }
	}
}

proc init_nurm2() {
	dendscale=spinescale*areascale
	somascale=areascale*1.0
	/* nonlinear decrease in Rm from rmsoma to rmend */
	/* rmsoma and rmend are set in init_params */
	maxdis=1113
	dis=0
	access dend1[7]
	area(0.5)
	distance()
	print "Initializing nonuniform passive membrane properties\n"
	forall { g_pas=1/(rmsoma/dendscale) cm=Cm*dendscale Ra=global_ra e_pas=Vleak}
	for i=8,433 {
		access dend1[i]
		dis=distance(0)
		rmpoint=(rmsoma-rmend)*exp(-dis/lambdadis)+rmend
		dend1[i] {g_pas=1/(rmpoint/dendscale) }
	}
	/* the following segments are NOT scaled for spines */
	/* these include the axon, soma, 1st 100 um of apical trunk, */
	/*  and the 1st 20 um of basals */
	/* the reconstructed axon */
	axon { g_pas=1/(rmsoma/somascale) cm=Cm*somascale }
	soma { g_pas=1/(rmsoma/somascale) cm=Cm*somascale }
	for i=0,13 {
		access dend1[i]
		dis=distance(0)
		dend1[i] {g_pas=1/(rmsoma/somascale) cm=Cm*somascale }
	}
	/* the real soma */
	for i=0,9 {
		access dend1[i]
		dis=distance(0)
		rmpoint=(rmsoma-rmend)*(1-(-dis/maxdis))+rmend
		dend1[i] {g_pas=1/(rmpoint/somascale) }
	}
	/* the 1st 100 um of the apical trunk */
	for i=10,11 {
		access dend1[i]
		dis=distance(0)
		rmpoint=(rmsoma-rmend)*exp(-dis/lambdadis)+rmend
		dend1[i] {g_pas=1/(rmpoint/somascale) }
	}
	for i=35,36 {
		access dend1[i]
		dis=distance(0)
		rmpoint=(rmsoma-rmend)*exp(-dis/lambdadis)+rmend
		dend1[i] {g_pas=1/(rmpoint/somascale) }
	}
	for i=46,46 {
		access dend1[i]
		dis=distance(0)
		rmpoint=(rmsoma-rmend)*exp(-dis/lambdadis)+rmend
		dend1[i] {g_pas=1/(rmpoint/somascale) }
	}
	for i=49,49 {
		access dend1[i]
		dis=distance(0)
		rmpoint=(rmsoma-rmend)*exp(-dis/lambdadis)+rmend
		dend1[i] {g_pas=1/(rmpoint/somascale) }
	}
	/* the 1st 20 um of the basals */
	for i=453,454 {
		dend1[i] { g_pas=1/(rmsoma/somascale) }
	}
	for i=434,436 {
		dend1[i] { g_pas=1/(rmsoma/somascale) }
	}
	for i=439,439 {
		dend1[i] { g_pas=1/(rmsoma/somascale) }
	}
	for i=508,509 {
		dend1[i] { g_pas=1/(rmsoma/somascale) }
	}
	for i=516,517 {
		dend1[i] { g_pas=1/(rmsoma/somascale) }
	}
	for i=543,544 {
		dend1[i] { g_pas=1/(rmsoma/somascale) }
	}
	for i=566,567 {
		dend1[i] { g_pas=1/(rmsoma/somascale) }
	}
}

proc init_nurm3() {
	areascale=$1
	dendscale=spinescale*areascale
	somascale=areascale*1.0
	/* sigmoidal decrease in Rm from rmsoma to rmend */
	/* rmsoma and rmend, and rmhalfdis and rmsteep are set in init_params */
	maxdis=1153
	dis=0
	access dend1[7]
	area(0.5)
	distance()
	print "Initializing nonuniform passive membrane properties\n"
	forall { g_pas=1/(rmsoma/dendscale) cm=Cm*dendscale Ra=global_ra e_pas=Vleak}
	for i=8,433 {
		access dend1[i]
		dis=distance(0)
		rmpoint=rmend+(rmsoma-rmend)/(1+exp((dis-rmhalfdis)/rmsteep))
		dend1[i] { g_pas=1/(rmpoint/dendscale) }
	}
	/* the following segments are NOT scaled for spines */
	/* these include the axon, soma, 1st 100 um of apical trunk, */
	/*  and the 1st 20 um of basals */
	/* the reconstructed axon */
	axon { g_pas=1/(Rm/somascale) cm=Cm*somascale }
	soma { g_pas=1/(Rm/somascale) cm=Cm*somascale }
	/* the real soma */
	for i=0,9 {
		access dend1[i]
		dis=distance(0)
		rmpoint=rmend+(rmsoma-rmend)/(1+exp((dis-rmhalfdis)/rmsteep))
		dend1[i] {g_pas=1/(rmpoint/somascale) }
	}
	/* the 1st 100 um of the apical trunk */
	for i=10,11 {
		access dend1[i]
		dis=distance(0)
		rmpoint=rmend+(rmsoma-rmend)/(1+exp((dis-rmhalfdis)/rmsteep))
		dend1[i] {g_pas=1/(rmpoint/somascale) }
	}
	for i=35,35 {
		access dend1[i]
		dis=distance(0)
		rmpoint=rmend+(rmsoma-rmend)/(1+exp((dis-rmhalfdis)/rmsteep))
		dend1[i] {g_pas=1/(rmpoint/somascale) }
	}
	for i=46,46 {
		access dend1[i]
		dis=distance(0)
		rmpoint=rmend+(rmsoma-rmend)/(1+exp((dis-rmhalfdis)/rmsteep))
		dend1[i] {g_pas=1/(rmpoint/somascale) }
	}
	for i=49,49 {
		access dend1[i]
		dis=distance(0)
		rmpoint=rmend+(rmsoma-rmend)/(1+exp((dis-rmhalfdis)/rmsteep))
		dend1[i] {g_pas=1/(rmpoint/somascale) }
	}
	/* the 1st 20 um of the basals */
	for i=434,436 {
		dend1[i] { g_pas=1/(rmsoma/somascale) }
	}
	for i=439,439 {
		dend1[i] { g_pas=1/(rmsoma/somascale) }
	}
	for i=453,454 {
		dend1[i] { g_pas=1/(rmsoma/somascale) }
	}
	for i=508,509 {
		dend1[i] { g_pas=1/(rmsoma/somascale) }
	}
	for i=516,517 {
		dend1[i] { g_pas=1/(rmsoma/somascale) }
	}
	for i=543,544 {
		dend1[i] { g_pas=1/(rmsoma/somascale) }
	}
	for i=566,567 {
		dend1[i] { g_pas=1/(rmsoma/somascale) }
	}
}

proc init_nurm4() {
	Cm=$1
	dendscale=spinescale*areascale
	somascale=areascale*1.0
	/* sigmoidal decrease in Rm from rmsoma to rmend */
	/* rmsoma and rmend, and rmhalfdis and rmsteep are set in init_params */
	/* differs from nurm3 in that function applies to basal dendrites also */
	maxdis=1153
	dis=0
	access dend1[7]
	area(0.5)
	distance()
	print "Initializing nonuniform passive membrane properties with nurm4\n"
	forall {g_pas=1/(rmsoma/dendscale) cm=Cm*dendscale Ra=global_ra e_pas=Vleak}
	if (nuri>0) {
		init_nuri()	/* initialize nonuniform Ri */
	}
	for i=0,599 {
		access dend1[i]
		dis=distance(0)
		rmpoint=rmend+(rmsoma-rmend)/(1+exp((dis-rmhalfdis)/rmsteep))
		dend1[i] {g_pas=1/(rmpoint/dendscale) }
	}
	/* the following segments are NOT scaled for spines */
	/* these include the axon, soma, 1st 100 um of apical trunk, */
	/*  and the 1st 20 um of basals */
	/* the reconstructed axon */
	axon { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	soma { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	/* the real soma */
	for i=0,9 {
		dend1[i] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	}
	/* the 1st 100 um of the apical trunk */
	for i=10,11 {
		dend1[i] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	}
	dend1[35] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	dend1[46] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	dend1[49] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	/* the 1st 20 um of the basals */
	for i=750,755 {
		dend1[i] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	}
	/* the 1st 20 um of the basals */
	dend1[434] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	dend1[435] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	dend1[436] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	dend1[439] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	dend1[453] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	dend1[454] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	dend1[508] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	dend1[509] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	dend1[516] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	dend1[517] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	dend1[543] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	dend1[544] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	dend1[566] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
	dend1[567] { g_pas=g_pas*somascale/dendscale cm=Cm*somascale/dendscale }
}

proc initq1() {
	/* uniform sag in all compartments */
/*	Vleak=-69 */
/*	maxq=0.00015 */
/*	a0_q=0.00057 */
/*	b0_q=a0_q */
	forall { gqbar_q=maxq*dendscale*(1-qblock)}
	printf("Initializing sag conductance in all segments - %d pecent blocked\n",qblock*100)
	/* the axon */
	axon { gqbar_q=0}
	soma { gqbar_q=maxq*somascale*(1-qblock)}
	/* the real soma */
	for i=0,9 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	/* the 1st 100 um of the apical trunk */
	for i=10,11 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	for i=35,36 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	dend1[46] { gqbar_q=maxq*somascale*(1-qblock)}
	dend1[49] { gqbar_q=maxq*somascale*(1-qblock)}
	/* the 1st 20 um of the basals */
	for i=453,454 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	for i=434,436 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	dend1[439] { gqbar_q=maxq*somascale*(1-qblock)}
	for i=508,509 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	for i=516,517 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	for i=543,544 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	for i=566,567 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
}

proc initq2() {
	/* sag scaled by a factor of qscale in nonspiny compartments */
	qscale=0
	Vleak=-70
	maxq=0.00018
	a0_q=0.00057
	b0_q=a0_q
	if (qblock<1) {
		forall { gqbar_q=maxq*dendscale*(1-qblock)}
		printf("Initializing sag conductance in all segments - %d pecent blocked\n",qblock*100)
	}
	/* the axon */
	axon { gqbar_q=0 }
	soma { gqbar_q=0 }
	/* the real soma */
	for i=0,9 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)*qscale}
	}
	/* the 1st 100 um of the apical trunk */
	for i=10,11 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)*qscale}
	}
	for i=35,36 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)*qscale}
	}
	dend1[46] { gqbar_q=maxq*somascale*(1-qblock)*qscale}
	dend1[49] { gqbar_q=maxq*somascale*(1-qblock)*qscale}
	/* the 1st 20 um of the basals */
	for i=453,454 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)*qscale}
	}
	for i=434,436 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)*qscale}
	}
	dend1[439] { gqbar_q=maxq*somascale*(1-qblock)*qscale}
	for i=508,509 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)*qscale}
	}
	for i=516,517 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)*qscale}
	}
	for i=543,544 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)*qscale}
	}
	for i=566,567 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)*qscale}
	}
}

proc initq3() {
	/* sag scaled by a factor of qscale */
	/* in all compartments beyond last oblique */
	qscale=20
	Vleak=-76
	maxq=0.00009
	a0_q=0.00057
	b0_q=a0_q
	forall { gqbar_q=maxq*dendscale*(1-qblock)}
	printf("Initializing sag conductance in all segments\n %d pecent blocked\n",qblock*100)
	/* the axon */
	axon { gqbar_q=0 }
	soma { gqbar_q=0 }
	/* the real soma */
	for i=0,9 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	/* the 1st 100 um of the apical trunk */
	for i=10,11 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	for i=35,36 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	dend1[46] { gqbar_q=maxq*somascale*(1-qblock)}
	dend1[49] { gqbar_q=maxq*somascale*(1-qblock)}
	/* the 1st 20 um of the basals */
	for i=453,454 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	for i=434,436 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	dend1[439] { gqbar_q=maxq*somascale*(1-qblock)}
	for i=508,509 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	for i=516,517 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	for i=543,544 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	for i=566,567 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)}
	}
	/* now scale up sag in all compartments distal to last oblique */
	for i=124,433 {
		dend1[i] { gqbar_q=maxq*somascale*(1-qblock)*qscale}
	}
}

proc initq4() {
	/* scale up sag as a function of distance from the soma */
	/* global linear function of distance doesn't seem to work, */
	/* even for ridiculously steep gradients */
	qscale=0.1
	Vleak=-74
	maxq=0.0000001
	a0_q=0.00041
	b0_q=a0_q
	dis=0
	access dend1[7]
	area(0.5)
	distance()
	printf("Initializing sag conductance in all segments\n %d pecent blocked\n",qblock*100)
	forall { gqbar_q=maxq*dendscale*(1-qblock)}
	for i=0,599 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq+(maxq*dendscale*(1-qblock)*dis/qscale)
	}
	/* the axon */
	axon { gqbar_q=0 }
	soma { gqbar_q=0 }
	/* the real soma */
	for i=0,9 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq+(maxq*somascale*(1-qblock)*dis/qscale)
	}
	/* the 1st 100 um of the apical trunk */
	for i=10,11 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq+(maxq*somascale*(1-qblock)*dis/qscale)
	}
	for i=35,36 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq+(maxq*somascale*(1-qblock)*dis/qscale)
	}
	access dend1[46]
	dis=distance(0)
	gqbar_q=maxq+(maxq*somascale*(1-qblock)*dis/qscale)
	access dend1[49]
	dis=distance(0)
	gqbar_q=maxq+(maxq*somascale*(1-qblock)*dis/qscale)
	/* the 1st 20 um of the basals */
	for i=453,454 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq+(maxq*somascale*(1-qblock)*dis/qscale)
	}
	for i=434,436 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq+(maxq*somascale*(1-qblock)*dis/qscale)
	}
	access dend1[439]
	dis=distance(0)
	gqbar_q=maxq+(maxq*somascale*(1-qblock)*dis/qscale)
	for i=508,509 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq+(maxq*somascale*(1-qblock)*dis/qscale)
	}
	for i=516,517 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq+(maxq*somascale*(1-qblock)*dis/qscale)
	}
	for i=543,544 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq+(maxq*somascale*(1-qblock)*dis/qscale)
	}
	for i=566,567 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq+(maxq*somascale*(1-qblock)*dis/qscale)
	}
}

proc initq5() {
	/* scale up sag as a function of distance from the soma */
	/* try linear gradient restricted to apical dendrites */
	/* q=maxq(qscale+(dis/maxdis)) */
	/* qscale is fraction of maxq at soma, must be between 0 and 1 */
	/* smaller qscale values give steeper gradients */
	/* maxq set in init_params */
	maxdis=1153
	qscale=0
	a0_q=0.00041
	b0_q=a0_q
	dis=0
	access dend1[7]
	area(0.5)
	distance()
	printf("Initializing sag conductance in all segments\n %d pecent blocked\n",qblock*100)
	forall { gqbar_q=maxq*qscale*dendscale*(1-qblock) }
	for i=8,433 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq*(qscale+(dis/maxdis))*dendscale*(1-qblock)
	}
	/* the axon */
	axon { gqbar_q=0 }
	soma { gqbar_q=0 }
	/* the real soma */
	for i=0,9 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq*(qscale+(dis/maxdis))*somascale*(1-qblock)
	}
	/* the 1st 100 um of the apical trunk */
	for i=10,11 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq*(qscale+(dis/maxdis))*somascale*(1-qblock)
	}
	for i=35,36 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq*(qscale+(dis/maxdis))*somascale*(1-qblock)
	}
	access dend1[46]
	dis=distance(0)
	gqbar_q=maxq*(qscale+(dis/maxdis))*somascale*(1-qblock)
	access dend1[49]
	dis=distance(0)
	gqbar_q=maxq*(qscale+(dis/maxdis))*somascale*(1-qblock)
	/* the 1st 20 um of the basals */
	for i=453,454 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq*(qscale+(dis/maxdis))*somascale*(1-qblock)
	}
	for i=434,436 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq*(qscale+(dis/maxdis))*somascale*(1-qblock)
	}
	access dend1[439]
	dis=distance(0)
	gqbar_q=maxq*(qscale+(dis/maxdis))*somascale*(1-qblock)
	for i=508,509 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq*(qscale+(dis/maxdis))*somascale*(1-qblock)
	}
	for i=516,517 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq*(qscale+(dis/maxdis))*somascale*(1-qblock)
	}
	for i=543,544 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq*(qscale+(dis/maxdis))*somascale*(1-qblock)
	}
	for i=566,567 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=maxq*(qscale+(dis/maxdis))*somascale*(1-qblock)
	}
}

proc initq8() {
	/* scale up sag as a sigmoidal function of distance from the soma */
	/* q=minq+(maxq-minq)/(1+exp((-dis-qhalfdis)/qsteep)) */
	/* minq=qscale*maxq */
	/* set maxq, qscale, and qhalfdis and qsteep in init_params */
	maxdis=1153
	minq=qscale*maxq
/*	a0_q=0.00041 */
/*	b0_q=a0_q */
	dis=0
	access dend1[7]
	area(0.5)
	distance()
	printf("Initializing sag conductance in all segments\n %d pecent blocked\n",qblock*100)
/*	forall { gqbar_q=qscale*maxq*dendscale*(1-qblock) } */
/*	this method (next line) makes gqbar_q=0 in the basal dendrites */
	forall { gqbar_q=0 }
	for i=8,433 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=minq+(maxq-minq)/(1+exp(-(dis-qhalfdis)/qsteep))
		gqbar_q=gqbar_q*dendscale*(1-qblock)
	}
	/* the reconstructed axon */
	axon { gqbar_q=0 }
	soma { gqbar_q=0 }
	/* the real soma */
	for i=0,9 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=minq+(maxq-minq)/(1+exp(-(dis-qhalfdis)/qsteep))
		gqbar_q=gqbar_q*somascale*(1-qblock)
	}
	/* the 1st 100 um of the apical trunk */
	for i=10,11 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=minq+(maxq-minq)/(1+exp(-(dis-qhalfdis)/qsteep))
		gqbar_q=gqbar_q*somascale*(1-qblock)
	}
	for i=35,35 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=minq+(maxq-minq)/(1+exp(-(dis-qhalfdis)/qsteep))
		gqbar_q=gqbar_q*somascale*(1-qblock)
	}
	for i=46,46 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=minq+(maxq-minq)/(1+exp(-(dis-qhalfdis)/qsteep))
		gqbar_q=gqbar_q*somascale*(1-qblock)
	}
	for i=49,49 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=minq+(maxq-minq)/(1+exp(-(dis-qhalfdis)/qsteep))
		gqbar_q=gqbar_q*somascale*(1-qblock)
	}
	/* the 1st 20 um of the basals */
	access dend1[439]
	dis=distance(0)
	gqbar_q=minq+(maxq-minq)/(1+exp(-(dis-qhalfdis)/qsteep))
	gqbar_q=gqbar_q*somascale*(1-qblock)
	gqbar_q=0
	for i=434,436 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=minq+(maxq-minq)/(1+exp(-(dis-qhalfdis)/qsteep))
		gqbar_q=gqbar_q*somascale*(1-qblock)
		gqbar_q=0
	}
	for i=453,454 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=minq+(maxq-minq)/(1+exp(-(dis-qhalfdis)/qsteep))
		gqbar_q=gqbar_q*somascale*(1-qblock)
		gqbar_q=0
	}
	for i=508,509 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=minq+(maxq-minq)/(1+exp(-(dis-qhalfdis)/qsteep))
		gqbar_q=gqbar_q*somascale*(1-qblock)
		gqbar_q=0
	}
	for i=516,517 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=minq+(maxq-minq)/(1+exp(-(dis-qhalfdis)/qsteep))
		gqbar_q=gqbar_q*somascale*(1-qblock)
		gqbar_q=0
	}
	for i=543,544 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=minq+(maxq-minq)/(1+exp(-(dis-qhalfdis)/qsteep))
		gqbar_q=gqbar_q*somascale*(1-qblock)
		gqbar_q=0
	}
	for i=566,567 {
		access dend1[i]
		dis=distance(0)
		gqbar_q=minq+(maxq-minq)/(1+exp(-(dis-qhalfdis)/qsteep))
		gqbar_q=gqbar_q*somascale*(1-qblock)
		gqbar_q=0
	}
}

proc initq10() {
	qten_qq=$1*10
	/* scale up sag as a sigmoidal function of distance from the soma */
	/* q=minq+(maxq-minq)/(1+exp((-dis-qhalfdis)/qsteep)) */
	/* minq=qscale*maxq */
	/* set maxq, qscale, and qhalfdis and qsteep in init_params */
	/* differs from initq9 in that function applies to basals as well */
	maxdis=1220
	minq=qscale*maxq
	dis=0
	access dend1[13]
	area(0.5)
	distance()
	printf("Initializing sag conductance in all segments with initq10\n")
	forall { gqbar_qq=0 }
	for i=0,1103 {
		access dend1[i]
		dis=distance(0)
		gqbar_qq=minq+(maxq-minq)/(1+exp(-(dis-qhalfdis)/qsteep))
		gqbar_qq=gqbar_qq*dendscale*(1-qblock)
	}
	/* the reconstructed axon */
	axon { gqbar_qq=0 }
	soma { gqbar_qq=0 }
	for i=0,7 {
		dend1[i] { gqbar_qq=0 }
	}
	/* the real soma */
	for i=0,9 {
		dend1[i] { gqbar_qq=gqbar_qq*somascale/dendscale }
	}
	/* the 1st 100 um of the apical trunk */
	for i=10,11 {
		dend1[i] { gqbar_qq=gqbar_qq*somascale/dendscale }
	}
	for i=35,38 {
		dend1[i] { gqbar_qq=gqbar_qq*somascale/dendscale }
	}
	dend1[46] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	dend1[49] {g_pas=1/(Rm/somascale) cm=Cm*somascale}
	/* the 1st 20 um of the basals */
	dend1[434] { gqbar_qq=gqbar_qq*somascale/dendscale }
	dend1[435] { gqbar_qq=gqbar_qq*somascale/dendscale }
	dend1[436] { gqbar_qq=gqbar_qq*somascale/dendscale }
	dend1[439] { gqbar_qq=gqbar_qq*somascale/dendscale }
	dend1[453] { gqbar_qq=gqbar_qq*somascale/dendscale }
	dend1[454] { gqbar_qq=gqbar_qq*somascale/dendscale }
	dend1[508] { gqbar_qq=gqbar_qq*somascale/dendscale }
	dend1[509] { gqbar_qq=gqbar_qq*somascale/dendscale }
	dend1[516] { gqbar_qq=gqbar_qq*somascale/dendscale }
	dend1[517] { gqbar_qq=gqbar_qq*somascale/dendscale }
	dend1[543] { gqbar_qq=gqbar_qq*somascale/dendscale }
	dend1[544] { gqbar_qq=gqbar_qq*somascale/dendscale }
	dend1[566] { gqbar_qq=gqbar_qq*somascale/dendscale }
	dend1[567] { gqbar_qq=gqbar_qq*somascale/dendscale }
}