//**********************       MOSSY CELL         ****************************************

// Mossy Cell template

// extracted from
// Dentate gyrus network model 
// Santhakumar V, Aradi I, Soltesz I (2005) J Neurophysiol 93:437-53 
// https://senselab.med.yale.edu/ModelDB/showModel.cshtml?model=51781&file=\dentategyrusnet2005\DG500_M7.hoc

// ModelDB file along with publication:
// Yim MY, Hanuschkin A, Wolfart J (2015) Hippocampus 25:297-308.
// http://onlinelibrary.wiley.com/doi/10.1002/hipo.22373/abstract

// modified by
// Abraham Nunes / 2022
// Man Yi Yim / 2015
// Alexander Hanuschkin / 2011

begintemplate MossyCell

ndend1=4
ndend2=4
ndend3=4
ndend4=4

public  pre_list, connect_pre, subsets, is_art, is_connected
public vbc2gc, vmc2gc, vhc2gc, vgc2bc, vbc2bc, vmc2bc, vhc2bc, vgc2mc, vbc2mc, vmc2mc, vhc2mc, vgc2hc, vmc2hc
public soma, mcdend1, mcdend2, mcdend3, mcdend4
create soma, mcdend1[ndend1], mcdend2[ndend2], mcdend3[ndend3], mcdend4[ndend4]
public all, adend, bdend, cdend, ddend
objref syn, pre_list, fl

//to include steady state current injection
nst=1
objectvar stim[nst]
double stimdur[nst], stimdel[nst], stimamp[nst]
public stim, stimdur, stimamp, stimdel


objref syn
proc init() {
	pre_list = new List()
	subsets()
	temp()
	synapse()
}

objref all, pdend, ddend

proc subsets() { local i
	objref all, pdend, ddend
	all = new SectionList()
		soma all.append()
		for i=0, 3 mcdend1 [i] all.append()
		for i=0, 3 mcdend2 [i] all.append()
		for i=0, 3 mcdend3 [i] all.append()
		for i=0, 3 mcdend4 [i] all.append()

	pdend  = new SectionList()
		mcdend1 [0] pdend.append()
		mcdend2 [0] pdend.append()
		mcdend3 [0] pdend.append()
		mcdend4 [0] pdend.append()

	ddend  = new SectionList()
		for i=1, 3 mcdend1 [i] ddend.append()
		for i=1, 3 mcdend2 [i] ddend.append()
		for i=1, 3 mcdend3 [i] ddend.append()
		for i=1, 3 mcdend4 [i] ddend.append()
	
}


proc temp() {

	soma {nseg=1 L=20 diam=20} 
		
	mcdend1 [0] {nseg=1 L=50 diam=5.78}
	mcdend1 [1] {nseg=1 L=50 diam=4}
	mcdend1 [2] {nseg=1 L=50 diam=2.5}
 	mcdend1 [3] {nseg=1 L=50 diam=1}

	mcdend2 [0] {nseg=1 L=50 diam=5.78}
	mcdend2 [1] {nseg=1 L=50 diam=4}
	mcdend2 [2] {nseg=1 L=50 diam=2.5}
	mcdend2 [3] {nseg=1 L=50 diam=1}
 		 
	mcdend3 [0] {nseg=1 L=50 diam=5.78}
	mcdend3 [1] {nseg=1 L=50 diam=4}
	mcdend3 [2] {nseg=1 L=50 diam=2.5}
	mcdend3 [3] {nseg=1 L=50 diam=1} 
	
	mcdend4 [0] {nseg=1 L=50 diam=5.78}
	mcdend4 [1] {nseg=1 L=50 diam=4}
	mcdend4 [2] {nseg=1 L=50 diam=2.5}
	mcdend4 [3] {nseg=1 L=50 diam=1} 	

    
	forall {
		insert ccanl
            catau_ccanl = 10
            caiinf_ccanl = 5.e-6
		insert ka
            gkabar_ka=0.00001
		insert nca  
            gncabar_nca=0.00008
		insert lca 
            glcabar_lca=0.0006
		insert sk
            gskbar_sk=0.016
		insert bk
            gkbar_bk=0.0165
		insert ih 
            ghyfbar_ih=0.000005
            ghysbar_ih=0.000005
	    insert ichan2
	        vshiftma_ichan2 = 43                           // value Yim et al. 2015
            vshiftmb_ichan2 = 15                           // value Yim et al. 2015
            vshiftha_ichan2 = 65                           // value Yim et al. 2015
            vshifthb_ichan2 = 12.5                         // value Yim et al. 2015
            vshiftnfa_ichan2 = 18                          // value Yim et al. 2015
            vshiftnfb_ichan2 = 43                          // value Yim et al. 2015
            vshiftnsa_ichan2 = 30                          // value Yim et al. 2015
            vshiftnsb_ichan2 = 55                          // value Yim et al. 2015
	}

	soma {
		insert bk
            gkbar_bk=0.030179002060815					// fitted to Howard et al. 2007
		insert ichan2  
			gnatbar_ichan2=0.054131228197338  			// fitted to Howard et al. 2007
			gkfbar_ichan2=0.043070370589102				// fitted to Howard et al. 2007
			gksbar_ichan2=0.010074989693078  			// fitted to Howard et al. 2007
			gl_ichan2 = 0.000006990410948				// fitted to Howard et al. 2007
			vshiftma_ichan2 = 46.10613308007081     	// fitted to Howard et al. 2007                       
            vshiftmb_ichan2 = 21.587317954511636     	// fitted to Howard et al. 2007                      
            vshiftha_ichan2 = 156.60600048559073       	// fitted to Howard et al. 2007                      
            vshifthb_ichan2 = 22.12030473272491         // fitted to Howard et al. 2007                 
            vshiftnfa_ichan2 = 30.701653087965823       // fitted to Howard et al. 2007                   
            vshiftnfb_ichan2 = 52.57215810598858        // fitted to Howard et al. 2007                   
            vshiftnsa_ichan2 = 24.097976599105685       // fitted to Howard et al. 2007                
            vshiftnsb_ichan2 = 66.46342312305076        // fitted to Howard et al. 2007                 
		insert ka
            gkabar_ka=0.000015360279879					// fitted to Howard et al. 2007
		insert lca 
            glcabar_lca=0.00076494065621				// fitted to Howard et al. 2007
		insert nca  
            gncabar_nca=0.000046076963887				// fitted to Howard et al. 2007
		insert sk
            gskbar_sk=0.021562445425285					// fitted to Howard et al. 2007
		insert ih 
            ghyfbar_ih=0.000009836462022				// fitted to Howard et al. 2007
            ghysbar_ih=0.000008197063742				// fitted to Howard et al. 2007
		
		cm=0.6								
	} 

	
	forsec pdend {
		insert ichan2
			gnatbar_ichan2=0.12  
			gkfbar_ichan2=0.0005
			gl_ichan2 = 0.000044
		cm=2.4
	}
		
	forsec ddend {
		insert ichan2
			gnatbar_ichan2=0.0
			gkfbar_ichan2=0.00
			gl_ichan2 = 0.000044
		cm=2.4
	}
		
	connect mcdend1[0](0), soma(1)
	connect mcdend2[0](0), soma(1)
	connect mcdend3[0](0), soma(0)
	connect mcdend4[0](0), soma(0)
	
	for i=1,3 {connect mcdend1[i](0), mcdend1[i-1](1)}
	for i=1,3 {connect mcdend2[i](0), mcdend2[i-1](1)}
	for i=1,3 {connect mcdend3[i](0), mcdend3[i-1](1)}
	for i=1,3 {connect mcdend4[i](0), mcdend4[i-1](1)}

	forall {Ra=100}
	forall {
        		ena 		= 	50		        // ena was unified from enat=55 (BC, HIPP, MC) and enat=45 (GC) in Santhakumar et al. (2005) <ah>
                ek			=	-90		        // simplified ekf=eks=ek=esk; note the eK was erroneously reported as -105mV in the Yim et al. 2015 <ah>
				ehyf		=	-40
				ehys		=	-40
				el_ichan2	= 	-59
                cao_ccanl	=	2 
			}

} // end proc temp()


//Defining synapses on to Mossy Cells
	objref syn  
	proc synapse() {

	mcdend1 [3] syn = new Exp2Syn(0.7)	//PP(AMPA) syn to dist dend similar to PP to GC
	syn.tau1 = 1.5	syn.tau2 = 5.5	syn.e = 0
	pre_list.append(syn)

	mcdend2 [3] syn = new Exp2Syn(0.7)	//PP(AMPA) syn to dist dend similar to PP to GC
	syn.tau1 = 1.5	syn.tau2 = 5.5	syn.e = 0
	pre_list.append(syn)

	mcdend3 [3] syn = new Exp2Syn(0.7)	//PP(AMPA) syn to dist dend similar to PP to GC
	syn.tau1 = 1.5	syn.tau2 = 5.5	syn.e = 0
	pre_list.append(syn)

	mcdend4 [3] syn = new Exp2Syn(0.7)	//PP(AMPA) syn to dist dend similar to PP to GC
	syn.tau1 = 1.5	syn.tau2 = 5.5	syn.e = 0
	pre_list.append(syn)

	mcdend1 [0] syn = new Exp2Syn(0.5)	//GC(AMPA) syn to prox dend similar to GC>CA3 Jonas '93
	syn.tau1 = 0.5	syn.tau2 = 6.2	syn.e = 0
	pre_list.append(syn)

	mcdend2 [0] syn = new Exp2Syn(0.5)	//GC(AMPA) syn to prox dend similar to GC>CA3 Jonas '93
	syn.tau1 = 0.5	syn.tau2 = 6.2	syn.e = 0
	pre_list.append(syn)

	mcdend3 [0] syn = new Exp2Syn(0.5)	//GC(AMPA) syn to prox dend similar to GC>CA3 Jonas '93
	syn.tau1 = 0.5	syn.tau2 = 6.2	syn.e = 0
	pre_list.append(syn)

	mcdend4 [0] syn = new Exp2Syn(0.5)	//GC(AMPA) syn to prox dend similar to GC>CA3 Jonas '93
	syn.tau1 = 0.5	syn.tau2 = 6.2	syn.e = 0
	pre_list.append(syn)

	mcdend1 [0] syn = new Exp2Syn(0.5)	//MC(AMPA) syn to prox dend similar to CA#>CA3 Aaron
	syn.tau1 = 0.45 	syn.tau2 =2.2	syn.e = 0
	pre_list.append(syn)

	mcdend2 [0] syn = new Exp2Syn(0.5)	//MC(AMPA) syn to prox dend similar to CA#>CA3 Aaron
	syn.tau1 = 0.45	syn.tau2 = 2.2		syn.e = 0
	pre_list.append(syn)

	mcdend3 [0] syn = new Exp2Syn(0.5)	//MC(AMPA) syn to prox dend similar to CA#>CA3 Aaron
	syn.tau1 = 0.45	syn.tau2 = 2.2	syn.e = 0
	pre_list.append(syn)

	mcdend4 [0] syn = new Exp2Syn(0.5)	//MC(AMPA) syn to prox dend similar to CA#>CA3 Aaron
	syn.tau1 = 0.45	syn.tau2 = 2.2	syn.e = 0
	pre_list.append(syn)

	soma syn = new Exp2Syn(0.5)	//BC(GABA) syn to prox dend based on BC>CA3 Bartos PNAS (mice)
	syn.tau1 = 0.3	syn.tau2 = 3.3	syn.e = -70
	pre_list.append(syn)

	mcdend1 [2] syn = new Exp2Syn(0.5)	//HIPP(GABA) syn to prox dend based on Hilar>GC Harney&Jones
	syn.tau1 = .5	syn.tau2 = 6		syn.e = -70
	pre_list.append(syn)

	mcdend2 [2] syn = new Exp2Syn(0.5)	//HIPP(GABA) syn to prox dend based on Hilar>GC Harney&Jones
	syn.tau1 = .5	syn.tau2 = 6		syn.e = -70
	pre_list.append(syn)

	mcdend3 [2] syn = new Exp2Syn(0.5)	//HIPP(GABA) syn to prox dend based on Hilar>GC Harney&Jones
	syn.tau1 = .5	syn.tau2 = 6		syn.e = -70
	pre_list.append(syn)

	mcdend4 [2] syn = new Exp2Syn(0.5)	//HIPP(GABA) syn to prox dend based on Hilar>GC Harney&Jones
	syn.tau1 = .5	syn.tau2 = 6	syn.e =-70
	pre_list.append(syn)

	

// Total of 17 synapses 	0-3 PP; 	4-7 GC; 	8-11 MC; 	12 BC; 	13-16 HIPP 
	} // end proc synapse

	proc connect_pre() {  
	soma $o2 = new NetCon (&v(1), $o1)
	}

func is_art()  { return 0 }

endtemplate MossyCell