Striatal Spiny Projection Neuron (SPN) plasticity rule (Jedrzejewska-Szmek et al 2016)

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1 . Jedrzejewska-Szmek J, Damodaran S, Dorman DB, Blackwell KT (2017) Calcium dynamics predict direction of synaptic plasticity in striatal spiny projection neurons. Eur J Neurosci 45:1044-1056 [PubMed]
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Model Information (Click on a link to find other models with that property)
Model Type: Dendrite;
Brain Region(s)/Organism:
Cell Type(s):
Gap Junctions:
Receptor(s): AMPA; NMDA; GabaA;
Transmitter(s): Glutamate;
Simulation Environment: GENESIS;
Model Concept(s): Calcium dynamics; Synaptic Plasticity;
Implementer(s): Jedrzejewska-Szmek, Joanna ; Damodaran, Sriraman ; Dorman, Daniel B ; Blackwell, Avrama [avrama at];
Search NeuronDB for information about:  GabaA; AMPA; NMDA; Glutamate;
//spines.g for including spines in the MSN model.

if ({exists {getglobal len_neck}})
    include MScell/spineParams.g
include MScell/AddCaSpines.g

function make_spineProto

    float shell_thick, Ca_tau, kB, kE, r

    float surf_head=dia_head*len_head*{PI}
    float surf_neck=len_neck*dia_neck*{PI}

    // ############# electrical compartments
    if (!{exists spine})
        create symcompartment spine

    addfield spine position
		addfield spine pathlen    // path length to soma, = position for prototype .p file
		addfield spine parentprim
		addfield spine parentsec
		addfield spine parenttert

    setfield  spine  \
           Cm     {{spineCM}*surf_neck} \
           Ra     /*{spineRa}*/ { 4.0*len_neck*{neckRA}/(dia_neck*dia_neck*{PI})}     \
           Em     {ELEAK}     \
           initVm {EREST_ACT} \
           Rm     {{spineRM}/surf_neck} \
           inject  0.0         \
           dia     {dia_neck}       \
           len     {len_neck}    \
           position 0.0   

    create symcompartment spine/{spcomp1}
    addfield spine/{spcomp1} position
    addfield spine/{spcomp1} pathlen
		addfield spine/{spcomp1} parentprim
		addfield spine/{spcomp1} parentsec
		addfield spine/{spcomp1} parenttert

    setfield spine/{spcomp1}          \
         Cm     {{spineCM}*surf_head} \
         Ra     {4.0*{spineRA}*len_head/(dia_head*dia_head*{PI})}     \
         Em     {ELEAK}           \
         initVm {EREST_ACT}       \
         Rm     {{spineRM}/surf_head} \
         inject  0.0              \
         dia     {dia_head}         \
         len     {len_head}       \
         position 0.0
    /*combine neck-head of CA1 CA1_spine */
    addmsg spine/{spcomp1} spine RAXIAL Ra Vm 
    addmsg spine spine/{spcomp1} AXIAL Ra Vm

    // ******** make calcium objects **************************
    enable /library
    if ({spinecalcium}==0)
        //This function adds calcium and buffer shells, connects them, and adds pumps
        //The function is in AddCaSpines.g
		neckSlabs={add_difshell_spine {CalciumName} "neck" {dia_neck} {len_neck} {neckSlabs} {neck_thickness_inc}}
		headSlabs={add_difshell_spine {CalciumName} "head" {dia_head} {len_head} {headSlabs} {head_thickness_inc}}
        echo "new slabs, neck=" {neckSlabs} ", head=" {headSlabs}

        //Connect head to neck, calcium and diffusible buffers

		addmsg spine/{spcomp1}/{CalciumName}{headSlabs} spine/{CalciumName}1 DIFF_DOWN prev_C thick
		addmsg spine/{CalciumName}1 spine/{spcomp1}/{CalciumName}{headSlabs} DIFF_UP prev_C thick
		addmsg spine/{spcomp1}/{CalciumName}{headSlabs}{bname2} spine/{CalciumName}1{bname2} DIFF_DOWN prev_free thick
		addmsg spine/{CalciumName}1{bname2} spine/{spcomp1}/{CalciumName}{headSlabs}{bname2} DIFF_UP prev_free thick
 		addmsg spine/{spcomp1}/{CalciumName}{headSlabs}{bname4} spine/{CalciumName}1{bname4} DIFF_DOWN prev_free thick
		addmsg spine/{CalciumName}1{bname4} spine/{spcomp1}/{CalciumName}{headSlabs}{bname4} DIFF_UP prev_free thick
        if ({calciumdye}>0)
            addmsg spine/{spcomp1}/{CalciumName}{headSlabs}{bnamefluor} spine/{CalciumName}1{bnamefluor} DIFF_DOWN prev_free thick
            addmsg spine/{CalciumName}1{bnamefluor} spine/{spcomp1}/{CalciumName}{headSlabs}{bnamefluor} DIFF_UP prev_free thick
            if ({calciumdye} == 4)
                addmsg spine/{spcomp1}/{CalciumName}{headSlabs}{egtaname} spine/{CalciumName}1{egtaname} DIFF_DOWN prev_free thick
                addmsg spine/{CalciumName}1{egtaname} spine/{spcomp1}/{CalciumName}{headSlabs}{egtaname} DIFF_UP prev_free thick

    elif ({spinecalcium}==1)  // Sabatini's model.       Sabatini, 2001,2004
		create Ca_concen  spine/{spcomp1}/{bufferNMDA}  // to create simplified Ca_pool here! 
        kE =86.0                                   // Carter and Sabatini, 2004
        Ca_tau = 25.0e-3                            
        r= (1+kE)/Ca_tau
        if ({calciumdye}==3)
            kB = 220                     // Fluo-4, taken from Yasuda,et,al. 2004,STEK
            Ca_tau = (1+kE+kB)/r         // re-calculate time constant because of application of the new calcium-dye
            kB = 70                      // Fluo-5F
            Ca_tau = (1+kE+kB)/r
        elif ({calciumdye}==1)
            echo "spines.g, ca_concen: need parameter kB for fura, single tau"
        float shell_vol= {PI}*(dia_head*dia_head/4.0)*len_head
        setfield spine/{spcomp1}/{bufferNMDA} \
                                 B          {1.0/(2.0*96494*shell_vol*(1+kE+kB))} \
                                 tau        {Ca_tau}                         \
                                 Ca_base    {Ca_basal}   \
                                 thick      {shell_thick} 

    disable /library
    /**** add NMDA/AMPA channels**************************/

    pushe spine/{spcomp1}

        addSynChannel . {AMPAname} {AMPAgmax} {bufferNMDA}
        addNMDAchannel . {NMDAname} {bufferNMDA} {NMDAgmax} {ghk_yesno}

    /*****************  add L-type, R-type, and T-type Calcium Channels*****************/

    //  addCaChannel {obj} {compt} {Gchan} {CalciumBuffer}
        if ({spinecalcium}==0)
            addCaChannelspines CaL12_channel      .  {gCaL12spine}    {CalciumName}2         // HVA CaL
            addCaChannelspines CaL13_channel      .  {gCaL13spine}    {CalciumName}1      // LVA CaL
            addCaChannelspines CaR_channel        .  {gCaRspine}      {CalciumName}2
            addCaChannelspines CaT_channel        .  {gCaTspine}      {CalciumName}2
        elif ({spinecalcium}==1)
            copy   {bufferNMDA} {bufferNR}
            copy   {bufferNMDA} {bufferLT}
            copy   {bufferNMDA} {bufferAll}

            addmsg {NMDAname} {bufferAll} I_Ca Ik

            if ({gCaL12spine} > {SMALLNUMBER})
                addCaChannelspines CaL12_channel      .  {gCaL12spine}    {bufferLT}         // HVA CaL
                addmsg CaL12_channelGHK {bufferAll} I_Ca Ik
            if ({gCaL13spine} > {SMALLNUMBER})
                addCaChannelspines CaL13_channel      .  {gCaL13spine}    {bufferLT}      // LVA CaL
                addmsg CaL13_channelGHK {bufferAll} I_Ca Ik
            if ({gCaRspine} > {SMALLNUMBER})
                addCaChannelspines CaR_channel        .  {gCaRspine}      {bufferNR}
                addmsg CaR_channelGHK {bufferAll} I_Ca Ik
            if ({gCaTspine} > {SMALLNUMBER})
                addCaChannelspines CaT_channel        .  {gCaTspine}      {bufferLT}
                addmsg CaT_channelGHK {bufferAll} I_Ca Ik


    return {neckSlabs}
//******************done making spines*********************************

//*****************begin function to add spines*********************************

function add_spines_evenly(cellpath,spineproto,a,b,density,bottomslab)
/* "spineproto"   :   spine prototype
** "density" :   1/um,  spine density; The number of spines in one compartment = density * compartment length. 
    str cellpath,compt,spineproto,spine,bottomslab
    int number,i
    float dia,len,a,b,density,position
    float separation,spineloc
    float parentPathlen, spinepath

    if(!{exists /library/{spineproto}})
        echo "The spine protomodel has not been made!"

    str bottomslabName={CalciumName}@{bottomslab}
    int totalspines=0

    foreach compt ({el {cellpath}/#[TYPE={compartment}]}) 
        if (!{{compt}=={{cellpath}@"/axIS"} || {compt}=={{cellpath}@"/ax"}}) 
            dia={getfield {compt} dia}
            position={getfield {compt} position}
            parentPathlen={getfield {compt} pathlen}
            len={getfield {compt} len}
            float xstart={getfield {compt} x0}

            //if the compartment is not a spine ,
            // and its position is between [a,b]
			if ({position>=a} && {position<b} )
		        str parentsec={getfield {compt} parentsec}
                if ({parentsec}=={"/cell/secdend11"})
                    number = density * len * 1e6

                // make sure that each compartment has at least one spine
				    if (number == 0)
					    number = number + 1

					    spine = "spine"@"_"@{i}
                        copy /library/{spineproto} {compt}/{spine}
                        //assign position and pathlen to the spine, this is specific to our one dimensional morphology
                        //The 2nd term of pathlen needs to be  changed if spineloc is changed for real morphology
                        setfield {compt}/{spine} position {spineloc}
                        setfield {compt}/{spine}/{spcomp1} position {spineloc}
                        setfield {compt}/{spine} pathlen {spinepath}
                        setfield {compt}/{spine}/{spcomp1} pathlen {(parentPathlen-len/2)+(spineloc-xstart)}
                        //parents of spine are same as parents of compartment
                        setfield {compt}/{spine} parentprim {getfield {compt} parentprim}
                        setfield {compt}/{spine}/{spcomp1} parentprim {getfield {compt} parentprim}
                        setfield {compt}/{spine} parentsec {getfield {compt} parentsec}
                        setfield {compt}/{spine}/{spcomp1} parentsec {getfield {compt} parentsec}
                        setfield {compt}/{spine} parenttert {getfield {compt} parenttert}
                        setfield {compt}/{spine}/{spcomp1} parenttert {getfield {compt} parenttert}
                        //messages between spineneck and dendrite compartment
                        addmsg {compt}/{spine} {compt} RAXIAL Ra Vm
                        addmsg {compt} {compt}/{spine} AXIAL Ra Vm
                        if ({spinecalcium}==0)
                            //send calcium diffusion message from bottom shell of spine neck to outer shell of compartment
                            addmsg {compt}/{spine}/{bottomslabName} {compt}/{upperShell} DIFF_DOWN prev_C thick
                            addmsg {compt}/{upperShell} {compt}/{spine}/{bottomslabName} DIFF_UP prev_C thick
                            //send buffer diffusion messages from bottom shell of spine neck to outer shell of compartement
                            addmsg {compt}/{spine}/{bottomslabName}{bname2} {compt}/{upperShell}{bname2} DIFF_DOWN prev_free thick
                            addmsg {compt}/{upperShell}{bname2} {compt}/{spine}/{bottomslabName}{bname2} DIFF_UP prev_free thick
                            addmsg {compt}/{spine}/{bottomslabName}{bname4} {compt}/{upperShell}{bname4} DIFF_DOWN prev_free thick
                            addmsg {compt}/{upperShell}{bname4} {compt}/{spine}/{bottomslabName}{bname4} DIFF_UP prev_free thick
                            if ({calciumdye} > 0)
                                addmsg {compt}/{spine}/{bottomslabName}{bnamefluor} {compt}/{upperShell}{bnamefluor}  DIFF_DOWN prev_free thick
                                addmsg {compt}/{upperShell}{bnamefluor} {compt}/{spine}/{bottomslabName}{bnamefluor}  DIFF_UP prev_free thick
                                if ({calciumdye} == 4)
                                    addmsg {compt}/{spine}/{bottomslabName}{egtaname} {compt}/{upperShell}{egtaname}  DIFF_DOWN prev_free thick
                                    addmsg {compt}/{upperShell}{egtaname} {compt}/{spine}/{bottomslabName}{egtaname}  DIFF_UP prev_free thick


                    end //end of for i< number
        //// Loop to set up CONNECTCROSS MESSAGES Between child spines and child dendrites of current compartment
                    int outmsgcount = {getmsg {compt} -outgoing -count}
                    int i
                    for (i = 0; {i} < {outmsgcount}; i = {i} + 1)
                        str msgtype = {getmsg {compt} -outgoing -type {i}}
                        str child =   {getmsg {compt} -outgoing -destination {i}}
                        if ({strcmp {msgtype} "CONNECTHEAD"} == 0) //If the message is CONNECTHEAD We Want to connectcross this child (spine or dend) with every other spine-only child
                            str subcompt
                            foreach subcompt ({el {compt}/#[TYPE={compartment}]}) //This will get list of spine compartments
                                if (!{strcmp {subcompt} {child}}==0)  //If the element we are crossconnecting is not itself
                                    addmsg {child} {subcompt} CONNECTCROSS Ra Vm //This will add messages both ways for every spine-spine and msgs from dend-to-spine but not spine-to-dend
                                    if (!{strncmp {getpath {child} -tail} "spine" 5} ==0) //child is dendrite (not spine), also add message: addmsg {subcompt} {child}
                                        //echo adding connectcross message from {child} to {subcompt}
                                        addmsg {subcompt} {child} CONNECTCROSS Ra Vm
                                    end // if child is dendrite (not spine)
                                end //if subcompt, child not identical
                            end //foreach subcompt (spine)
                        end //if message is connect tail
                    end //for i = 0:outmsgcount
                end //end if parentprim is prim1
            end // end of if position...

        end // end of if ... axIS

    end // end of "foreach" loop

    return totalspines