Dendritica (Vetter et al 2001)

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Accession:7907
Dendritica is a collection of programs for relating dendritic geometry and signal propagation. The programs are based on those used for the simulations described in: Vetter, P., Roth, A. & Hausser, M. (2001) For reprint requests and additional information please contact Dr. M. Hausser, email address: m.hausser@ucl.ac.uk
Reference:
1 . Vetter P, Roth A, Häusser M (2001) Propagation of action potentials in dendrites depends on dendritic morphology. J Neurophysiol 85:926-37 [PubMed]
Model Information (Click on a link to find other models with that property)
Model Type: Dendrite;
Brain Region(s)/Organism:
Cell Type(s): Hippocampus CA1 pyramidal cell; Hippocampus CA3 pyramidal cell; Neocortex U1 L2/6 pyramidal intratelencephalic cell; Cerebellum Purkinje cell;
Channel(s): I Na,t; I L high threshold; I p,q; I K; I M; I K,Ca;
Gap Junctions:
Receptor(s):
Gene(s):
Transmitter(s):
Simulation Environment: NEURON;
Model Concept(s): Action Potential Initiation; Bursting; Active Dendrites; Influence of Dendritic Geometry; Detailed Neuronal Models; Axonal Action Potentials; Action Potentials;
Implementer(s): Hausser, M [M.Hausser at ucl.ac.uk];
Search NeuronDB for information about:  Hippocampus CA1 pyramidal cell; Hippocampus CA3 pyramidal cell; Cerebellum Purkinje cell; Neocortex U1 L2/6 pyramidal intratelencephalic cell; I Na,t; I L high threshold; I p,q; I K; I M; I K,Ca;
Files displayed below are from the implementation
/* Relating dendritic geometry and signal propagation   */
/* Philipp Vetter, Arnd Roth and Michael Hausser        */
/* settings.hoc Version 1.0                  11.10.1999 */

/**************************************************************
  ActiveModels
***************************************************************/

act0	   = "act0"
act1	   = "act1"
act2	   = "act2"
act3	   = "act3"
act4	   = "act4"
act5	   = "act5"
act6	   = "act6"
act7	   = "act7"
act8	   = "act8"
act9	   = "act9"
act10	   = "act10"


proc get_settings() {

	get_standard()		// initialize standard settings  (T, activespine ...) 
	active_get(ActiveModel)			/* get simulation values          */
  	cell_name(cell)				/* set cell-related strdefs       */
	activecell = cellname
	print "loading ", cellname  /* up to here, only flags and strdefs are set */

		}

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

proc active_get() {	/* get the simulation values for ActiveModel $s1 */

			if (strcmp($s1,act0) 	   == 0) act0_set()
			if (strcmp($s1,act1) 	   == 0) act1_set()
			if (strcmp($s1,act2) 	   == 0) act2_set()
			if (strcmp($s1,act3) 	   == 0) act3_set()
			if (strcmp($s1,act4) 	   == 0) act4_set()
			if (strcmp($s1,act5) 	   == 0) act5_set()
			if (strcmp($s1,act6) 	   == 0) act6_set()
			if (strcmp($s1,act7) 	   == 0) act7_set()
			if (strcmp($s1,act8) 	   == 0) act8_set()
			if (strcmp($s1,act9) 	   == 0) act9_set()
			if (strcmp($s1,act10) 	   == 0) act10_set()


		  }

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

proc activemodel_set() { /* this puts the active models in the graphics menu */
			xmenu("Conductances")
				xradiobutton("Standard ","actname_set(0)")
				xradiobutton("Standard (A. Destexhe)","actname_set(1)")
				xradiobutton("high Rax","actname_set(2)")
				xradiobutton("low Rax","actname_set(3)")
				xradiobutton("high Cm","actname_set(4)")
				xradiobutton("low Cm","actname_set(5)")
				xradiobutton("high Rax (A. Destexhe)","actname_set(6)")
				xradiobutton("Migliore both","actname_set(7)")
				xradiobutton("high Rax (A. Destexhe)","actname_set(8)")
				xradiobutton("high Rax (A. Destexhe)","actname_set(9)")
				xradiobutton("high Rax (A. Destexhe)","actname_set(10)")
			xmenu()

			}

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

proc actname_set() { local n
		    n = $1
		    if (n==0) actname = "standard"
		    if (n==1) actname = "standard (A. Destexhe)"
		    if (n==2) actname = "high Rax"
		    if (n==3) actname = "low Rax"
		    if (n==4) actname = "high Cm"
		    if (n==5) actname = "low Cm"
		    if (n==6) actname = "high Rax (A. Destexhe)"
		    if (n==7) actname = "Migliore both abs"
		    if (n==8) actname = "Migliore both rel"
			
 		    sprint(ActiveModel,"act%d",n)
		    sprint(cellnameact,"%s using %s conductances",activecell,actname)
		    if (numarg()<2) print cellnameact
		    doEvents()
 		}

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

proc act0_set() {		/* Zach Nature standard (kvz|naz) */

Rax		= 150		/* passive membrane properties */
Rm         	= 12000
C_m        	= 1
G_kv		= 30		/* active membrane properties */
G_kvnode	= 500
G_na		= 35
G_nanode	= 35000
		 }

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

proc act1_set() {		/* Destexhe standard (kvd|nad) */

Rax		= 150		
Rm         	= 12000
C_m        	= 1.0
G_kv		= 40		
G_kvnode	= 1200
G_na		= 50
G_nanode	= 40000
		 }

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

proc act2_set() {  		/* Zach high Rax (kvz|naz)   */

Rax		= 200		
Rm         	= 12000
C_m        	= 1.0
G_kv		= 20		
G_kvnode	= 100
G_na		= 28
G_nanode	= 30000
		 }

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

proc act3_set() {  		/* Zach low Rax (kvz|naz)   */

Rax		= 70		
Rm         	= 12000
C_m        	= 1.0
G_kv		= 35		
G_kvnode	= 500
G_na		= 32
G_nanode	= 32000
		 }

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

proc act4_set() {  		/* Zach high C_m  (kvz|naz)   */

Rax		= 150		
Rm         	= 8500
C_m        	= 1.5
G_kv		= 35		
G_kvnode	= 35
G_na		= 42
G_nanode	= 35000
		 }

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

proc act5_set() {  		/* Zach low C_m (kvz|naz)   */

Rax		= 150		
Rm         	= 18500
C_m        	= 0.65
G_kv		= 50		
G_kvnode	= 85
G_na		= 24
G_nanode	= 24000
		 }

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

proc act6_set() {		/* Destexhe (kvd|nad) high Rax */

Rax		= 200		
Rm         	= 12000
C_m        	= 1.0
G_kv		= 40		
G_kvnode	= 2000
G_na		= 50
G_nanode	= 40000
		 }


proc act7_set() {		/* Migliore standard (kvz|naz) */

Rax		= 150		/* passive membrane properties */
Rm         	= 12000
C_m        	= 1
G_kv		= 0 // 30		/* active membrane properties */
G_kvnode	= 500
G_na		= 380
G_nanode	= 35000

Kad_current     = 1
Gbar_kad 	= 0.048  		/* Ka distal settings (Migliore) */
Kad_rel 	= 0
Kad_slope 	= 1

Kap_current     = 1
Gbar_kap 	= 0.048            	/* Ka proximal settings (Migliore) */
Kap_rel 	= 0
Kap_slope 	= 1

		 }


proc act8_set() {		/* Migliore standard (kvz|naz) relative */

Rax		= 150		/* passive membrane properties */
Rm         	= 12000
C_m        	= 1
G_kv		= 0 // 30		/* active membrane properties */
G_kvnode	= 500
G_na		= 380
G_nanode	= 35000

Kad_current     = 1
Gbar_kad 	= 0.048  		/* Ka distal settings (Migliore) */
Kad_rel 	= 1
Kad_slope 	= 1

Kap_current     = 1
Gbar_kap 	= 0.048            	/* Ka proximal settings (Migliore) */
Kap_rel 	= 1
Kap_slope 	= 1

		 }


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

proc get_standard()	{ /* standard settings, implemented substitutively, whenever a neuron is loaded */
		Kap_current 	= 0		/* substitutively no Kap_current */
		Kad_current 	= 0		/* substitutively no Kad_current */
		Iq_current 	= 0		/* substitutively no Iq_current */
		Ca_current	= 0		/* substitutively no Ca_current */
		KCa_current	= 0		/* substitutively no KCa_current */
		Km_current	= 0		/* substitutively no Km_current */
		nonuniform_Rm	= 0		/* substitutively no nonuniform Rm */
		electrotonicL	= 0		/* substitutively physical lengths */
		activespine	= 1		/* model spines with active membrane */
		currentdist	= 1		/* status for which distlist to use*/
		currentcell	= 0		/* number of currently active cell */
		simMode 	= 0		/* 0 IClamp 1 Vclamp simulation  */
		celsius    	= 37		/* general settings */
		originx         = 0.5           /* where to add axon */
		Ek         	= -90
		Ena        	= 60
		cells 		= 1
		swc		= 0		/* flag is turned on for Duke Southampton files */
		synthetic       = 0             /* synthetic neurons need longer simulation time */

		St_del         	= 0		/* stimulation (duration,delay,amplitude) */
		St_amp		= 1		
		St_dur		= 9999

		Sim_durI	= 0		/* simulation (duration & dt) */
		Sim_dtI		= 0.25		/* run for Sim_durI with Sim_dtI, 
						   then Sim_dur with Sim_dt */
		Sim_dur		= 15		
		Sim_dt		= 0.025

		Rec_del		= Sim_durI	/* recording options (for display) */
		Rec_dur		= Sim_dur
		Rec_dt		= Sim_dt		

		/* passive membrane properties, which aren't being changed in general */
		Rm_node 	= 50
		Cm_myelin  	= 0.04

		G_ca		= .3		/* 13.8.98 */
		G_kca		= .1
		G_km		= 3
		Rm_end		= Rm_half = Rm_steep = 0
		G_qq		= 0.02		/* Iq settings */
		Qq_end 		= 20
		Qq_steep	= 439
		Qq_half		= 50


		Gbar_kad = 0.048  		/* Ka distal settings (Migliore) */
		Kad_rel = 0
		Kad_slope = 1

		Gbar_kap = 0.048            /* Ka proximal settings (Migliore) */
		Kap_rel = 0
		Kap_slope = 1
			}



		diam_int       	= 0.01   // diameter interval 
				    //(consider tapering effect when looking at impedance matching)
		Rest_dur 	= 100	/* simulation settings for achieving rest */
		purkinjestep 	= 0.25
		figcurrent 	= -1		/* settings for fig() */
		fsize 		= 1
		movie           = 0


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

proc passive_ref() {  	/* reference values for passive membrane properties
			   Stuart & Spruston J. Neurosci (1998) */
				  
			n = numarg()
				
			if (n < 2) {	/* control values */
					Rin             = 32.1
                        		Tau             = 12.4
	                        	Slope           = -0.0014029
        	                	Offset          = 0.95564     	}

			if (n == 2) {	/* CsCl data */
					Rin             = 51.7
                        		Tau             = 20.9
	                        	Slope           = -0.00088573  
        	                	Offset          = 0.9856      	}
	           }

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

amp_max  	= 46.8		/* values from Stuart, Schiller & Sakmann J Physiol (1997) */
amp_bas  	= 50		/* amp_max + amp_bas = AP amplitude */
amp_lmd  	= 155		
vmax_lmd 	= 246
vmax_amp 	= 697
plat_s   	= 500		/* slope peak latency */
olat_s   	= 1200		/* slope onset latency */
half_s		= 0.002		/* slope halfwidth */
half_b          = 0.46 		/* baseline halfwidth */




// need to set these values in order to set flags

asections_max   = -9999
adarea_max      = -9999
adarea_maxdist  = -9999
diam_mean	= -9999
deq_max		= -9999
deq_maxdist	= -9999
sections_mean	= -9999
taper				= -9999
branchdensityII			= -9999
branchdensity			= -9999
branchdensityII_noend		= -9999
branchdensity_noend		= -9999
mean_stem_dendrite_diam		= -9999
darea_max			= -9999
ataper				= -9999
abranchdensityII		= -9999
abranchdensity			= -9999
abranchdensityII_noend		= -9999
abranchdensity_noend		= -9999
amean_stem_dendrite_diam	= -9999
sections_max			= -9999
sections_maxdist		= -9999
taper_mean			= -9999

powers = 0
geometric = 0
equiv = 0
forward = 0
Yval = 3014
Xval1= 1000
Xval2= 1000
Xval3= 1000
Axon = 0
actname_set(0,"don't")

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

proc rec_set() {	/* set the recording vectors during simulation 	*/
			/* $1 		rec_del delay			*/
			/* $2 		rec_dur duration		*/
			/* $3 		rec_dt timestep			*/

			n = numarg()
			if (n>0)  rec_del = $1 else rec_del = Rec_del
			if (n>1)  rec_dur = $2 else rec_dur = Rec_dur
			if (n>2)  rec_dt  = $3 else rec_dt  = Rec_dt

			time.indgen(rec_del,rec_del+rec_dur,rec_dt)

			vsoma.record(&Soma.sec.v(0.5),time)
			vnode.record(&node[0].v(0.5),time)
			vdend.record(&Distal.sec.v(1),time)
			voltrec.record(&Vclamp.sec.v(Vclampx),time)
			synapserec.record(&Synapse.sec.v(Synapsex),time)
			
		}

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

proc st_set()   {       /* set Iclamp parameters  				*/
			/* $1 		st_amp amplitude			*/
			/* $2 		st_del delay				*/
			/* $3 		st_dur duration				*/

			n = numarg()
			if (n>0)  st_amp = $1 else st_amp = St_amp
			if (n>1)  st_del = $2 else st_del = St_del
			if (n>2)  st_dur = $3 else st_dur = St_dur
			
			st.dur = st_dur
			st.del = st_del
			st.amp = st_amp 
			vclamp.dur1 = st_dur
			synapse.onset = 2
                }                       

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

proc set_synapse() {      local i 
			/* set Synapse parameters				*/
			/* $1 		synapse_gmax			*/
			/* $2 		synapse_tau0				*/
			/* $3 		synapse_tau1				*/
			/* $4 		synapse_onset				*/
			/* $5 		synapse_e				*/


			n = numarg()
			if (n>0)  synapse_gmax  = $1 else synapse_gmax = 0.1
			if (n>1)  synapse_tau0 = $2 else synapse_tau0 = 0.2
			if (n>2)  synapse_tau1 = $3 else synapse_tau1 = 3.0
			if (n>3)  synapse_onset = $4 else synapse_onset = 2
			if (n>4)  synapse_e = $5 else synapse_e = 0

			synapse.gmax   = synapse_gmax
			synapse.tau0   = synapse_tau0	
			synapse.tau1   = synapse_tau1
			synapse.onset = synapse_onset
			synapse.e    = synapse_e 

		}

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

proc sim_set()   {       local i 
			/* set simulation  parameters  				*/
			/* $1 		sim_dur					*/
			/* $2 		sim_dt					*/
			/* $3 		sim_durI ( this is before sim_dur)	*/
			/* $4 		sim_dtI 				*/

			n = numarg()
			if (n>0)  sim_dur 	= $1 else sim_dur 	= Sim_dur
			if (n>1)  sim_dt 	= $2 else sim_dt	= Sim_dt
			if (n>2)  sim_durI 	= $3 else sim_durI	= Sim_durI
			if (n>3)  sim_dtI 	= $4 else sim_dtI	= Sim_dtI
			
			rec_set(sim_durI,sim_dur,sim_dt)
		}

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

proc spine_set()	{ /* sets spines active if $1 = activespine == 1 */

	if (numarg()) activespine = $1 

	
	forall                { spine_pk  =1 }
        forsec dendritesI     {	spine_pk  = spinescale }
	forsec dendritesII    {	spine_pk  = spinescaleII }
	forsec dendritesIII   { dendarea = 0			// spine corrections Rapp
				for(x) dendarea+= area(x)
				spinarea        = 10*L
				spine_pk   = (spinarea+dendarea)/dendarea
                               }

	forall for(x) if (!activespine) { active_pk = 1/spine_pk(x)}  else { active_pk(x)=1 }

			}

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

proc active_set()  {	/* set active model parameters  */
			/* $1 		gbar_na		*/
			/* $2 		gbar_kv		*/
			/* $3 		gbar_na (node) 	*/
			/* $4 		g_qq	 	*/
			/* $5 		qq_end	 	*/
			/* $6 		qq_steep 	*/
			/* $7 		qq_half	 	*/

		n = numarg()
		if (n>0)  g_na     = $1 else g_na     = G_na
		if (n>1)  g_kv     = $2 else g_kv     = G_kv
		if (n>2)  g_nanode = $3 else g_nanode = G_nanode
		if (n>3)  g_kvnode = $4 else g_kvnode = G_kvnode
		if (n>4)  g_qq      = $5 else g_qq		= G_qq
		if (n>5)  qq_end    = $6 else qq_end		= Qq_end 
		if (n>6)  qq_steep  = $7 else qq_steep 	= Qq_steep
		if (n>7)  qq_half   = $8 else qq_half		= Qq_half

		if (Iq_current)	forsec all \
		gbar_qq=qq_end+(g_qq-qq_end)/(1+exp((distance(0)-qq_half)/qq_steep))
		
		forsec all   for(x)   { gbar_kv = g_kv*spine_pk(x)*active_pk(x)  
			  	     gbar_na = g_na*spine_pk(x)*active_pk(x)
				     if (Iq_current) gbar_qq *spine_pk(x)*active_pk(x) }

		forsec "myelin"         gbar_na = g_na
  
		forsec "node"         {	gbar_na = g_nanode
		               		gbar_kv = g_kvnode}

		forsec "hill"         {	gbar_na = g_nanode
		               		gbar_kv = g_kvnode }

		forsec "iseg"         {	gbar_na = g_nanode
		               		gbar_kv = g_kvnode }
	

	   }

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

proc passive_set() { /* set passive properties	*/
		     /* $1 	rm		*/
		     /* $2 	rax		*/
		     /* $3 	c_m 		*/
		     /* $4 	cm_myelin	*/
		     /* $5 	qq_end	 	*/
		     /* $6 	qq_steep 	*/
		     /* $7 	qq_half	 	*/

	n = numarg()
	if (n>0)  rm        = $1      else rm     	= Rm
	if (n>1)  rax       = abs($2) else rax    	= Rax
	if (n>2)  c_m       = $3      else c_m    	= C_m
	if (n>3)  cm_myelin = $4      else cm_myelin	= Cm_myelin
	if (n>4)  rm_node   = $5      else rm_node	= Rm_node
	if (n>5)  rm_end    = $6      else rm_end	= Rm_end 
	if (n>6)  rm_steep  = $7      else rm_steep	= Rm_steep
	if (n>7)  rm_half   = $8      else rm_half	= Rm_half

	if (!nonuniform_Rm) rm_end = rm	 /* only nonuniform distr if nonuniform_Rm==1 */ 
	origin.sec distance(0,originx)
        forall {if (rm-rm_end==0) g_pas =1/rm else \
		          g_pas = 1/(rm+(rm-rm_end)/(1+exp((distance(0)-rm_half)/rm_steep)))
               cm    = c_m
               Ra    = rax
                      }
	forsec "myelin"  	cm    = cm_myelin	
        forsec "node"    	g_pas = 1/rm_node	
        forsec "iseg"    	g_pas = 1/rm	/* make sure iseg & hill have uniform rm */	
        forsec "hill"    	g_pas = 1/rm	

        forsec dendrites     {	cm     *= spine_pk		// spine corrections
                               	g_pas  *= spine_pk }

		}



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

proc ca_set()  {/* set Calcium channel parameters  */
		/* $1 		gbar_ca		*/

		n = numarg()
		if (n>0)  g_ca     = $1 else g_ca     = G_ca
		if (Ca_current) {

		forsec all           gbar_ca	= g_ca*spine_pk*active_pk  

		   }}



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

proc kca_set()  {	/* set KCalcium channel parameters  */
			/* $1 		gbar_kca		*/

		n = numarg()
		if (n>0)  g_kca     = $1 else g_kca     = G_kca
		if (KCa_current){
		forsec all for(x) gbar_kca = g_kca*spine_pk(x)*active_pk(x)  /* stdard spinecorrection */
		   }}

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

proc set_vclamp() { local n
		n = numarg()
		if (n>0) forsec $s1 Vclamp = new SectionRef()
		if (n>1) Vclampx = $2
		Vclamp.sec vclamp.loc(Vclampx)
		vclamp.rs = 0.001  // 1 kOhm series resistance
		vclamp.dur1 = rec_dur
		}

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

proc km_set()  {	/* set km_channel parameters  */
			/* $1 		gbar_km		*/

		n = numarg()
		if (n>0)  g_km     = $1 else g_km     = G_km

		if (Km_current) {

		forsec all     	gbar_ca = g_km*spine_pk(x)*active_pk(x)  /* standard spinecorrection */
		   }}

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

proc set_waveform() { local n
	      n = numarg()

	      if (n==1)	waveform = $o1.c
	      if (n==0) { sprint(cellvar,"../data/%s/simvclamp_p18",ActiveModel)
			  fileob.ropen(cellvar)
    			  waveform.vread(fileob) }
	      if (n==2) { fileob.ropen($s1)
    			  waveform.vread(fileob) }

	      if (n==3) { fileob.chooser("", "Browse for waveforms", "*","Type file name","Cancel")
    			while (fileob.chooser()) {
			fileob.getname(str1)
			fileob.ropen(str1)  
			waveform.vread(fileob) } }

			}

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

proc reset() {  /* set the whole environment to its standard settings */


		       threshold_dur = 20   // sim_dur when calculating threshold
	if (synthetic) threshold_dur = 50   // synthetic geometries can be very long
	if (forward)   threshold_dur = 40   // forward propagation may take longer 10/3/99

	simMode=0  
	st_set()
	set_vclamp()
	if (numarg()!=1) sim_set()
	rec_set()
	spine_set()
	active_set()
	set_synapse()
	passive_set()
	ca_set()
	kca_set()
	km_set()
	lkad_set()
	lkap_set()
	     }

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

proc distal_vclamp() { local n
		/* VClamps p18 somatic AP waveform at section $s1 */
		/* waveform scaled by $2                          */
		/* if numarg() > 2 specify Waveform file   */
		/* if numarg() > 3 plot results   */
		/* e.g. distal_vclamp("dend[1]",0.5,"coolwave",1)   */

		n = numarg()
		if (n>1) scaling = $2 else scaling = 1

		set_dendrite($s1)
	      	set_vclamp($s1)
		passive_set(rm,rax,c_m,cm_myelin,rm_node,rm_end,rm_steep,rm_half)

		active_set(g_na,g_kv,g_nanode,g_kvnode)
		st_set(st_amp,st_del,st_dur)
		rinput_calc()

		simMode = 1 /* voltage clamping */
		if (n>2) { fileob.ropen($s3)
    	    		   histx.vread(fileob) 
			   sim(scaling,histx)		
			   sim(scaling,histx) } \
		else     { sim(scaling)
			   sim(scaling) }
		sim_calc()
		if (n>3) sim_plot()
		}

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

proc set_dendrite() {
		/* e.g. set_dendrite("dend[1]") will set dend[1] as the dendrite to be plotted */

		sprint(str1,"%s Distal = new SectionRef()",$s1)
		execute(str1)
		}

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

proc save_dendrite() { 	fileob.wopen($s1)
			vdend.vwrite(fileob)
			}

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





proc lkad_set()  {	/* set active model parameters  */
			/* $1 		gkabar_kad	*/
			/* $2 		kad_slope	*/
			/* $3 		kad_rel	        */

	n = numarg()
	if (n>0)  gbar_kad   = $1 else gbar_kad     = Gbar_kad
	if (n>1)  kad_slope  = $2 else kad_slope    = Kad_slope
	if (n>2)  kad_rel    = $3 else kad_rel	    = Kad_rel

	insert_channels()

forsec dendrites  if(ismembrane("kad")) for(x) gkabar_kad(x) = gbar_kad*factor_kad(x)*spine_pk*active_pk 

	   }


proc lkap_set()  {	/* set active model parameters  */
			/* $1 		gkabar_kap	*/
			/* $2 		kap_slope	*/
			/* $3 		kap_rel	        */

	n = numarg()
	if (n>0)  gbar_kap   = $1 else gbar_kap     = Gbar_kap
	if (n>1)  kap_slope  = $2 else kap_slope    = Kap_slope
	if (n>2)  kap_rel    = $3 else kap_rel	    = Kap_rel
	insert_channels()
forsec dendrites  if(ismembrane("kap")) for(x) gkabar_kap(x) = gbar_kap*factor_kap(x)*spine_pk*active_pk 

	   }










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