Dynamic cortical interlaminar interactions (Carracedo et al. 2013)

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Accession:150806
"... Here we demonstrate the mechanism underlying a purely neocortical delta rhythm generator and show a remarkable laminar, cell subtype and local subcircuit delineation between delta and nested theta rhythms. We show that spike timing during delta-nested theta rhythms controls an iterative, reciprocal interaction between deep and superficial cortical layers resembling the unsupervised learning processes proposed for laminar neural networks by Hinton and colleagues ... and mimicking the alternating cortical dynamics of sensory and memory processing during wakefulness."
Reference:
1 . Carracedo LM, Kjeldsen H, Cunnington L, Jenkins A, Schofield I, Cunningham MO, Davies CH, Traub RD, Whittington MA (2013) A neocortical delta rhythm facilitates reciprocal interlaminar interactions via nested theta rhythms. J Neurosci 33:10750-61 [PubMed]
Citations  Citation Browser
Model Information (Click on a link to find other models with that property)
Model Type: Realistic Network;
Brain Region(s)/Organism:
Cell Type(s): Neocortex L5/6 pyramidal GLU cell; Neocortex L2/3 pyramidal GLU cell; Neocortex V1 L5B pyramidal pyramidal tract GLU cell; Neocortex fast spiking (FS) interneuron; Neocortex spiking regular (RS) neuron; Neocortex spiking low threshold (LTS) neuron; Neocortex deep neurogliaform interneuron; Neocortex superficial neurogliaform interneuron;
Channel(s): I Na,p; I Na,t; I L high threshold; I A; I K; I M; I h; I K,Ca; I Calcium; I A, slow;
Gap Junctions: Gap junctions;
Receptor(s): GabaA; GabaB; AMPA; NMDA;
Gene(s):
Transmitter(s):
Simulation Environment: FORTRAN;
Model Concept(s): Activity Patterns; Bursting; Oscillations; Sleep;
Implementer(s): Traub, Roger D [rtraub at us.ibm.com];
Search NeuronDB for information about:  Neocortex L5/6 pyramidal GLU cell; Neocortex L2/3 pyramidal GLU cell; Neocortex V1 L5B pyramidal pyramidal tract GLU cell; GabaA; GabaB; AMPA; NMDA; I Na,p; I Na,t; I L high threshold; I A; I K; I M; I h; I K,Ca; I Calcium; I A, slow;
/
CarracedoEtAl2013
readme.txt
dexptablebig_setup.f *
dexptablesmall_setup.f *
fnmda.f *
groucho_gapbld.f *
groucho_gapbld_mix.f *
integrate_deepaxaxx.f *
integrate_deepbaskx.f *
integrate_deepLTSx.f *
integrate_deepng.f *
integrate_nontuftRSXXB.f *
integrate_nrtxB.f *
integrate_spinstelldiegoxB.f *
integrate_supaxaxx.f *
integrate_supbaskx.f *
integrate_supLTSX.f *
integrate_supng.f *
integrate_suppyrFRBxPB.f *
integrate_suppyrRS.f *
integrate_suppyrRSXPB.f *
integrate_tcrxB.f *
integrate_tuftIBVx3B.f *
integrate_tuftRSXXB.f *
makefile *
otis_table_setup.f *
spikewaveS5.f *
synaptic_compmap_construct.f *
synaptic_map_construct.f *
                            
! Integration program for superior & deep basket & axo-axonic cells
! From baskn.f in supergj.f

       SUBROUTINE integrate_supbaskx (O, time, numcell, V, curr,
     &  initialize, firstcell, lastcell,
     &  gAMPA, gNMDA, gGABA_A, Mg, gapcon, totaxgj, gjtable, dt,
     &  chi,mnaf,mnap,
     &  hnaf,mkdr,mka,
     &  hka,mk2,hk2,
     &  mkm,mkc,mkahp,
     &  mcat,hcat,mcal,
     &  mar)

           SAVE

       integer, parameter:: numcomp = 59  ! should be compat. with calling prog

       integer numcell, totaxgj, gjtable(totaxgj,4)
       integer initialize, firstcell, lastcell
       INTEGER J1, I, J, K, L, L1, O, K1
       REAL*8  Z, Z1, Z2, curr(numcomp,numcell), c(numcomp)
       REAL*8 dt, time, Mg, gapcon
c Usual dt in this program .002 ms

c CINV is 1/C, i.e. inverse capacitance

       real*8 v(numcomp,numcell), chi(numcomp,numcell), cinv(numcomp),
     x mnaf(numcomp,numcell),hnaf(numcomp,numcell), 
     x mkdr(numcomp,numcell),
     x mka(numcomp,numcell),hka(numcomp,numcell),mk2(numcomp,numcell),
     x hk2(numcomp,numcell),mkm(numcomp,numcell),
     x mkc(numcomp,numcell),mkahp(numcomp,numcell),
     x mcat(numcomp,numcell),hcat(numcomp,numcell),
     x mcal(numcomp,numcell),mar(numcomp,numcell),
     x jacob(numcomp,numcomp),betchi(numcomp),
     x gam(0:numcomp,0:numcomp),gL(numcomp),gnaf(numcomp),
     x gnap(numcomp),gkdr(numcomp),gka(numcomp),
     x gk2(numcomp),gkm(numcomp),gkc(numcomp),gkahp(numcomp),
     x gcat(numcomp),gcaL(numcomp),gar(numcomp),
     x cafor(numcomp), ggaba_a(numcomp,numcell),
     x gampa(numcomp,numcell),gnmda(numcomp,numcell)
       real*8
     X alpham_naf(0:640),betam_naf(0:640),dalpham_naf(0:640),
     X   dbetam_naf(0:640),
     X alphah_naf(0:640),betah_naf(0:640),dalphah_naf(0:640),
     X   dbetah_naf(0:640),
     X alpham_kdr(0:640),betam_kdr(0:640),dalpham_kdr(0:640),
     X   dbetam_kdr(0:640),
     X alpham_ka(0:640), betam_ka(0:640),dalpham_ka(0:640) ,
     X   dbetam_ka(0:640),
     X alphah_ka(0:640), betah_ka(0:640), dalphah_ka(0:640),
     X   dbetah_ka(0:640),
     X alpham_k2(0:640), betam_k2(0:640), dalpham_k2(0:640),
     X   dbetam_k2(0:640),
     X alphah_k2(0:640), betah_k2(0:640), dalphah_k2(0:640),
     X   dbetah_k2(0:640),
     X alpham_km(0:640), betam_km(0:640), dalpham_km(0:640),
     X   dbetam_km(0:640),
     X alpham_kc(0:640), betam_kc(0:640), dalpham_kc(0:640),
     X   dbetam_kc(0:640),
     X alpham_cat(0:640),betam_cat(0:640),dalpham_cat(0:640),
     X   dbetam_cat(0:640),
     X alphah_cat(0:640),betah_cat(0:640),dalphah_cat(0:640),
     X   dbetah_cat(0:640),
     X alpham_caL(0:640),betam_caL(0:640),dalpham_caL(0:640),
     X   dbetam_caL(0:640),
     X alpham_ar(0:640), betam_ar(0:640), dalpham_ar(0:640),
     X   dbetam_ar(0:640)
       real*8 vL,vk,vna,var,vca,vgaba_a

        INTEGER NEIGH(numcomp,5), NNUM(numcomp)
        real*8 fastna_shift

c the f's are the functions giving 1st derivatives for evolution of
c the differential equations for the voltages (v), calcium (chi), and
c other state variables.
       real*8 fv(numcomp), fchi(numcomp),fmnaf(numcomp),
     x fhnaf(numcomp),fmkdr(numcomp),
     x fmka(numcomp),fhka(numcomp),fmk2(numcomp),fhk2(numcomp),
     x fmkm(numcomp),fmkc(numcomp),fmkahp(numcomp),
     x fmcat(numcomp),fhcat(numcomp),fmcal(numcomp),fmar(numcomp)

c below are for calculating the partial derivatives
       real*8 dfv_dv(numcomp,numcomp), dfv_dchi(numcomp), 
     x  dfv_dmnaf(numcomp),
     x  dfv_dhnaf(numcomp),dfv_dmkdr(numcomp),
     x  dfv_dmka(numcomp),dfv_dhka(numcomp),
     x  dfv_dmk2(numcomp),dfv_dhk2(numcomp),
     x  dfv_dmkm(numcomp),dfv_dmkc(numcomp),
     x  dfv_dmkahp(numcomp),dfv_dmcat(numcomp),
     x  dfv_dhcat(numcomp),dfv_dmcal(numcomp),
     x  dfv_dmar(numcomp)

        real*8 dfchi_dv(numcomp), dfchi_dchi(numcomp),
     x dfmnaf_dmnaf(numcomp), dfmnaf_dv(numcomp),dfhnaf_dhnaf(numcomp),
     x dfhnaf_dv(numcomp),dfmkdr_dmkdr(numcomp),dfmkdr_dv(numcomp),
     x dfmka_dmka(numcomp),dfmka_dv(numcomp),
     x dfhka_dhka(numcomp),dfhka_dv(numcomp),
     x dfmk2_dmk2(numcomp),dfmk2_dv(numcomp),
     x dfhk2_dhk2(numcomp),dfhk2_dv(numcomp),
     x dfmkm_dmkm(numcomp),dfmkm_dv(numcomp),
     x dfmkc_dmkc(numcomp),dfmkc_dv(numcomp),
     x dfmcat_dmcat(numcomp),dfmcat_dv(numcomp),dfhcat_dhcat(numcomp),
     x dfhcat_dv(numcomp),dfmcal_dmcal(numcomp),dfmcal_dv(numcomp),
     x dfmar_dmar(numcomp),dfmar_dv(numcomp),dfmkahp_dchi(numcomp),
     x dfmkahp_dmkahp(numcomp), dt2

         INTEGER  K0
       REAL*8 OPEN(numcomp),gamma(numcomp),gamma_prime(numcomp)
c gamma is function of chi used in calculating KC conductance
       REAL*8 alpham_ahp(numcomp), alpham_ahp_prime(numcomp)
       REAL*8 gna_tot(numcomp),gk_tot(numcomp),gca_tot(numcomp)
       REAL*8 gca_high(numcomp), gar_tot(numcomp)
c this will be gCa conductance corresponding to high-thresh channels
       REAL*8 A, BB1, BB2

c do initialization on 1st time step
c      if (O.eq.1) then
       if (initialize.eq.0) then

c Program fnmda assumes A, BB1, BB2 defined in calling program
c as follows:
         A = DEXP(-2.847d0)
         BB1 = DEXP(-.693d0)
         BB2 = DEXP(-3.101d0)

       CALL  SUPBASK_SETUP
     X   (alpham_naf, betam_naf, dalpham_naf, dbetam_naf,
     X    alphah_naf, betah_naf, dalphah_naf, dbetah_naf,
     X    alpham_kdr, betam_kdr, dalpham_kdr, dbetam_kdr,
     X    alpham_ka , betam_ka , dalpham_ka , dbetam_ka ,
     X    alphah_ka , betah_ka , dalphah_ka , dbetah_ka ,
     X    alpham_k2 , betam_k2 , dalpham_k2 , dbetam_k2 ,
     X    alphah_k2 , betah_k2 , dalphah_k2 , dbetah_k2 ,
     X    alpham_km , betam_km , dalpham_km , dbetam_km ,
     X    alpham_kc , betam_kc , dalpham_kc , dbetam_kc ,
     X    alpham_cat, betam_cat, dalpham_cat, dbetam_cat,
     X    alphah_cat, betah_cat, dalphah_cat, dbetah_cat,
     X    alpham_caL, betam_caL, dalpham_caL, dbetam_caL,
     X    alpham_ar , betam_ar , dalpham_ar , dbetam_ar)

        CALL SUPBASKMAJ (GL,GAM,GKDR,GKA,GKC,GKAHP,GK2,GKM,
     X              GCAT,GCAL,GNAF,GNAP,GAR,
     X    CAFOR,JACOB,C,BETCHI,NEIGH,NNUM)

          do i = 1, 59
             cinv(i) = 1.d0 / c(i)
          end do

C  IN MILLIMOLAR

        VL = -65.d0
        VK =  -100.d0
        VNA = 50.d0
        VCA = 125.d0
        VAR = -40.d0
        VGABA_A = -75.d0


c ? initialize membrane state variables?
        do L = 1, numcell
        do i = 1, numcomp
          v(i,L) = VL
	  chi(i,L) = 0.d0
	mnaf(i,L) = 0.d0
	mkdr(i,L) = 0.d0
	mk2(i,L) = 0.d0
	mkm(i,L) = 0.d0
	mkc(i,L) = 0.d0
	mkahp(i,L) = 0.d0
	mcat(i,L) = 0.d0
	mcal(i,L) = 0.d0
	mar(i,L) = 0.d0

        k1 = idnint (4.d0 * (vL + 120.d0))

      hnaf(i,L) = alphah_naf(k1)/(alphah_naf(k1)+betah_naf(k1))
      hka(i,L) = alphah_ka(k1)/(alphah_ka(k1)+betah_ka(k1))
      hk2(i,L) = alphah_k2(k1)/(alphah_k2(k1)+betah_k2(k1))
      hcat(i,L)=alphah_cat(k1)/(alphah_cat(k1)+betah_cat(k1))
         end do
         end do


          do i = 1, numcomp
                  gnap(i) = 0.d0
                  gk2(i) = 0.d0
                  gkm(i) = 0.d0
                  gkahp(i) = 0.d0
                  gcat(i) = 0.d0
                  gar(i) = 0.d0
		  open(i) = 0.d0
          end do
             goto 1000
c End initialization
             endif

c          do L = 1, numcell
           do L = firstcell, lastcell


       DO I = 1, numcomp
          FV(I) = -GL(I) * (V(I,L) - VL) * cinv(i)
c         DO 302, J = 1, NNUM(I)
          DO J = 1, NNUM(I)
             K = NEIGH(I,J)
302     FV(I) = FV(I) + GAM(I,K) * (V(K,L) - V(I,L)) * cinv(i)
          END DO
        END DO
301    CONTINUE


        CALL FNMDA (V, OPEN, numcell, numcomp, MG, L, 
     &    A, BB1, BB2)

      DO i = 1, numcomp
421    FV(I) = FV(I) + ( CURR(I,L)
     X   - (gampa(I,L) + open(i) * gnmda(I,L))*V(I,L)
     X   - ggaba_a(I,L)*(V(I,L)-Vgaba_a) ) * cinv(i)
      END DO
c above assumes equil. potential for AMPA & NMDA = 0 mV

       do m = 1, totaxgj
        if (gjtable(m,1).eq.L) then
         L1 = gjtable(m,3)
         igap1 = gjtable(m,2)
         igap2 = gjtable(m,4)
 	fv(igap1) = fv(igap1) + gapcon *
     &   (v(igap2,L1) - v(igap1,L)) * cinv(igap1)
        else if (gjtable(m,3).eq.L) then
         L1 = gjtable(m,1)
         igap1 = gjtable(m,4)
         igap2 = gjtable(m,2)
 	fv(igap1) = fv(igap1) + gapcon *
     &   (v(igap2,L1) - v(igap1,L)) * cinv(igap1)
        endif
       end do ! do m

c      do i = 1, ngap_FS(L) ! obsolete gj code
c      L1 = list_gap_FS(L,i)
c       fv(dendsite) = fv(dendsite) + gapconid_FS *
c    &   (vdgap_global_FS(L1) - v(dendsite,L)) * cinv(dendsite)
c      end do  ! obsolete gj code

       do i = 1, numcomp
        gamma(i) = dmin1 (1.d0, .004d0 * chi(i,L))
        if (chi(i,L).le.250.d0) then
          gamma_prime(i) = .004d0
        else
          gamma_prime(i) = 0.d0
        endif
       end do

c     DO 88, I = 1, numcomp
      DO I = 1, numcomp
       gna_tot(i) = gnaf(i) * (mnaf(i,L)**3) * hnaf(i,L) +
     x     gnap(i) * (mnaf(i,L)**3)
       gk_tot(i) = gkdr(i) * (mkdr(i,L)**4) +
     x             gka(i)  * (mka(i,L)**4) * hka(i,L) +
     x             gk2(i)  * mk2(i,L) * hk2(i,L) +
     x             gkm(i)  * mkm(i,L) +
     x             gkc(i)  * mkc(i,L) * gamma(i) +
     x             gkahp(i)* mkahp(i,L)
       gca_tot(i) = gcat(i) * (mcat(i,L)**2) * hcat(i,L) +
     x              gcaL(i) * (mcaL(i,L)**2)
       gca_high(i) =
     x              gcaL(i) * (mcaL(i,L)**2)
       gar_tot(i) = gar(i) * mar(i,L)


88     FV(I) = FV(I) - ( gna_tot(i) * (v(i,L) - vna)
     X  + gk_tot(i) * (v(i,L) - vK)
     X  + gca_tot(i) * (v(i,L) - vCa)
     X  + gar_tot(i) * (v(i,L) - var) ) * cinv(i)
       END DO

         do i = 1, numcomp
         do j = 1, numcomp
          if (i.ne.j) then
            dfv_dv(i,j) = jacob(i,j)
          else
            dfv_dv(i,j) = jacob(i,i) - cinv(i) *
     X  (gna_tot(i) + gk_tot(i) + gca_tot(i) + gar_tot(i)
     X   + ggaba_a(i,L) + gampa(i,L)
     X   + open(i) * gnmda(I,L) )
          endif
         end do
         end do

          do i = 1, numcomp
        dfv_dchi(i)  = - cinv(i) * gkc(i) * mkc(i,L) *
     x                     gamma_prime(i) * (v(i,L)-vK)
        dfv_dmnaf(i) = -3.d0 * cinv(i) * (mnaf(i,L)**2) *
     X    (gnaf(i) * hnaf(i,L) + gnap(i)) * (v(i,L) - vna)
        dfv_dhnaf(i) = - cinv(i) * gnaf(i) * (mnaf(i,L)**3) *
     X                    (v(i,L) - vna)
        dfv_dmkdr(i) = -4.d0 * cinv(i)*gkdr(i) * (mkdr(i,L)**3)
     X                   * (v(i,L) - vK)
        dfv_dmka(i)  = -4.d0 * cinv(i)*gka(i) * (mka(i,L)**3) *
     X                   hka(i,L) * (v(i,L) - vK)
        dfv_dhka(i)  = - cinv(i) * gka(i) * (mka(i,L)**4) *
     X                    (v(i,L) - vK)
       dfv_dmk2(i)  = - cinv(i)*gk2(i) * hk2(i,L) * (v(i,L)-vK)
       dfv_dhk2(i)  = - cinv(i)*gk2(i) * mk2(i,L) * (v(i,L)-vK)
       dfv_dmkm(i)  = - cinv(i)*gkm(i) * (v(i,L) - vK)
       dfv_dmkc(i)  = - cinv(i)*gkc(i) * gamma(i) * (v(i,L)-vK)
       dfv_dmkahp(i)= - cinv(i)*gkahp(i) * (v(i,L) - vK)
       dfv_dmcat(i)  = -2.d0 * cinv(i) * gcat(i) * mcat(i,L) *
     X                    hcat(i,L) * (v(i,L) - vCa)
        dfv_dhcat(i) = - cinv(i) * gcat(i) * (mcat(i,L)**2) *
     X                  (v(i,L) - vCa)
        dfv_dmcal(i) = -2.d0 * cinv(i) * gcal(i) * mcal(i,L) *
     X                      (v(i,L) - vCa)
        dfv_dmar(i) = - cinv(i) * gar(i) * (v(i,L) - var)
          end do

         do i = 1, numcomp
          fchi(i) = - cafor(i) * gca_high(i) * (v(i,L) - vca)
     x       - betchi(i) * chi(i,L)
          dfchi_dv(i) = - cafor(i) * gca_high(i)
          dfchi_dchi(i) = - betchi(i)
         end do

       do i = 1, numcomp
        alpham_ahp(i) = dmin1(0.2d-4 * chi(i,L),0.01d0)
        if (chi(i,L).le.500.d0) then
          alpham_ahp_prime(i) = 0.2d-4
        else
          alpham_ahp_prime(i) = 0.d0
        endif
       end do

       do i = 1, numcomp
        fmkahp(i) = alpham_ahp(i) * (1.d0 - mkahp(i,L))
     x                  -.001d0 * mkahp(i,L)
        dfmkahp_dmkahp(i) = - alpham_ahp(i) - .001d0
        dfmkahp_dchi(i) = alpham_ahp_prime(i) *
     x                     (1.d0 - mkahp(i,L))
       end do

          do i = 1, numcomp


       K1 = IDNINT ( 4.d0 * (V(I,L) + 120.d0) )
       IF (K1.GT.640) K1 = 640
       IF (K1.LT.  0) K1 =   0

             fastNa_shift = -2.5d0
       K0 = IDNINT ( 4.d0 * (V(I,L)+  fastNa_shift+ 120.d0) )
       IF (K0.GT.640) K0 = 640
       IF (K0.LT.  0) K0 =   0


        fmnaf(i) = alpham_naf(k0) * (1.d0 - mnaf(i,L)) -
     X              betam_naf(k0) * mnaf(i,L)
        fhnaf(i) = alphah_naf(k1) * (1.d0 - hnaf(i,L)) -
     X              betah_naf(k1) * hnaf(i,L)
        fmkdr(i) = alpham_kdr(k1) * (1.d0 - mkdr(i,L)) -
     X              betam_kdr(k1) * mkdr(i,L)
        fmka(i)  = alpham_ka (k1) * (1.d0 - mka(i,L)) -
     X              betam_ka (k1) * mka(i,L)
        fhka(i)  = alphah_ka (k1) * (1.d0 - hka(i,L)) -
     X              betah_ka (k1) * hka(i,L)
        fmk2(i)  = alpham_k2 (k1) * (1.d0 - mk2(i,L)) -
     X              betam_k2 (k1) * mk2(i,L)
        fhk2(i)  = alphah_k2 (k1) * (1.d0 - hk2(i,L)) -
     X              betah_k2 (k1) * hk2(i,L)
        fmkm(i)  = alpham_km (k1) * (1.d0 - mkm(i,L)) -
     X              betam_km (k1) * mkm(i,L)
        fmkc(i)  = alpham_kc (k1) * (1.d0 - mkc(i,L)) -
     X              betam_kc (k1) * mkc(i,L)
        fmcat(i) = alpham_cat(k1) * (1.d0 - mcat(i,L)) -
     X              betam_cat(k1) * mcat(i,L)
        fhcat(i) = alphah_cat(k1) * (1.d0 - hcat(i,L)) -
     X              betah_cat(k1) * hcat(i,L)
        fmcaL(i) = alpham_caL(k1) * (1.d0 - mcaL(i,L)) -
     X              betam_caL(k1) * mcaL(i,L)
        fmar(i)  = alpham_ar (k1) * (1.d0 - mar(i,L)) -
     X              betam_ar (k1) * mar(i,L)

       dfmnaf_dv(i) = dalpham_naf(k0) * (1.d0 - mnaf(i,L)) -
     X                  dbetam_naf(k0) * mnaf(i,L)
       dfhnaf_dv(i) = dalphah_naf(k1) * (1.d0 - hnaf(i,L)) -
     X                  dbetah_naf(k1) * hnaf(i,L)
       dfmkdr_dv(i) = dalpham_kdr(k1) * (1.d0 - mkdr(i,L)) -
     X                  dbetam_kdr(k1) * mkdr(i,L)
       dfmka_dv(i)  = dalpham_ka(k1) * (1.d0 - mka(i,L)) -
     X                  dbetam_ka(k1) * mka(i,L)
       dfhka_dv(i)  = dalphah_ka(k1) * (1.d0 - hka(i,L)) -
     X                  dbetah_ka(k1) * hka(i,L)
       dfmk2_dv(i)  = dalpham_k2(k1) * (1.d0 - mk2(i,L)) -
     X                  dbetam_k2(k1) * mk2(i,L)
       dfhk2_dv(i)  = dalphah_k2(k1) * (1.d0 - hk2(i,L)) -
     X                  dbetah_k2(k1) * hk2(i,L)
       dfmkm_dv(i)  = dalpham_km(k1) * (1.d0 - mkm(i,L)) -
     X                  dbetam_km(k1) * mkm(i,L)
       dfmkc_dv(i)  = dalpham_kc(k1) * (1.d0 - mkc(i,L)) -
     X                  dbetam_kc(k1) * mkc(i,L)
       dfmcat_dv(i) = dalpham_cat(k1) * (1.d0 - mcat(i,L)) -
     X                  dbetam_cat(k1) * mcat(i,L)
       dfhcat_dv(i) = dalphah_cat(k1) * (1.d0 - hcat(i,L)) -
     X                  dbetah_cat(k1) * hcat(i,L)
       dfmcaL_dv(i) = dalpham_caL(k1) * (1.d0 - mcaL(i,L)) -
     X                  dbetam_caL(k1) * mcaL(i,L)
       dfmar_dv(i)  = dalpham_ar(k1) * (1.d0 - mar(i,L)) -
     X                  dbetam_ar(k1) * mar(i,L)

       dfmnaf_dmnaf(i) =  - alpham_naf(k0) - betam_naf(k0)
       dfhnaf_dhnaf(i) =  - alphah_naf(k1) - betah_naf(k1)
       dfmkdr_dmkdr(i) =  - alpham_kdr(k1) - betam_kdr(k1)
       dfmka_dmka(i)  =   - alpham_ka (k1) - betam_ka (k1)
       dfhka_dhka(i)  =   - alphah_ka (k1) - betah_ka (k1)
       dfmk2_dmk2(i)  =   - alpham_k2 (k1) - betam_k2 (k1)
       dfhk2_dhk2(i)  =   - alphah_k2 (k1) - betah_k2 (k1)
       dfmkm_dmkm(i)  =   - alpham_km (k1) - betam_km (k1)
       dfmkc_dmkc(i)  =   - alpham_kc (k1) - betam_kc (k1)
       dfmcat_dmcat(i) =  - alpham_cat(k1) - betam_cat(k1)
       dfhcat_dhcat(i) =  - alphah_cat(k1) - betah_cat(k1)
       dfmcaL_dmcaL(i) =  - alpham_caL(k1) - betam_caL(k1)
       dfmar_dmar(i)  =   - alpham_ar (k1) - betam_ar (k1)

          end do

       dt2 = 0.5d0 * dt * dt

        do i = 1, numcomp
          v(i,L) = v(i,L) + dt * fv(i)
           do j = 1, numcomp
        v(i,L) = v(i,L) + dt2 * dfv_dv(i,j) * fv(j)
           end do
        v(i,L) = v(i,L) + dt2 * ( dfv_dchi(i) * fchi(i)
     X          + dfv_dmnaf(i) * fmnaf(i)
     X          + dfv_dhnaf(i) * fhnaf(i)
     X          + dfv_dmkdr(i) * fmkdr(i)
     X          + dfv_dmka(i)  * fmka(i)
     X          + dfv_dhka(i)  * fhka(i)
     X          + dfv_dmk2(i)  * fmk2(i)
     X          + dfv_dhk2(i)  * fhk2(i)
     X          + dfv_dmkm(i)  * fmkm(i)
     X          + dfv_dmkc(i)  * fmkc(i)
     X          + dfv_dmkahp(i)* fmkahp(i)
     X          + dfv_dmcat(i)  * fmcat(i)
     X          + dfv_dhcat(i) * fhcat(i)
     X          + dfv_dmcaL(i) * fmcaL(i)
     X          + dfv_dmar(i)  * fmar(i) )

        chi(i,L) = chi(i,L) + dt * fchi(i) + dt2 *
     X   (dfchi_dchi(i) * fchi(i) + dfchi_dv(i) * fv(i))
        mnaf(i,L) = mnaf(i,L) + dt * fmnaf(i) + dt2 *
     X   (dfmnaf_dmnaf(i) * fmnaf(i) + dfmnaf_dv(i)*fv(i))
        hnaf(i,L) = hnaf(i,L) + dt * fhnaf(i) + dt2 *
     X   (dfhnaf_dhnaf(i) * fhnaf(i) + dfhnaf_dv(i)*fv(i))
        mkdr(i,L) = mkdr(i,L) + dt * fmkdr(i) + dt2 *
     X   (dfmkdr_dmkdr(i) * fmkdr(i) + dfmkdr_dv(i)*fv(i))
        mka(i,L) =  mka(i,L) + dt * fmka(i) + dt2 *
     X   (dfmka_dmka(i) * fmka(i) + dfmka_dv(i) * fv(i))
        hka(i,L) =  hka(i,L) + dt * fhka(i) + dt2 *
     X   (dfhka_dhka(i) * fhka(i) + dfhka_dv(i) * fv(i))
        mk2(i,L) =  mk2(i,L) + dt * fmk2(i) + dt2 *
     X   (dfmk2_dmk2(i) * fmk2(i) + dfmk2_dv(i) * fv(i))
        hk2(i,L) =  hk2(i,L) + dt * fhk2(i) + dt2 *
     X   (dfhk2_dhk2(i) * fhk2(i) + dfhk2_dv(i) * fv(i))
        mkm(i,L) =  mkm(i,L) + dt * fmkm(i) + dt2 *
     X   (dfmkm_dmkm(i) * fmkm(i) + dfmkm_dv(i) * fv(i))
        mkc(i,L) =  mkc(i,L) + dt * fmkc(i) + dt2 *
     X   (dfmkc_dmkc(i) * fmkc(i) + dfmkc_dv(i) * fv(i))
        mkahp(i,L) = mkahp(i,L) + dt * fmkahp(i) + dt2 *
     X (dfmkahp_dmkahp(i)*fmkahp(i) + dfmkahp_dchi(i)*fchi(i))
        mcat(i,L) =  mcat(i,L) + dt * fmcat(i) + dt2 *
     X   (dfmcat_dmcat(i) * fmcat(i) + dfmcat_dv(i) * fv(i))
        hcat(i,L) =  hcat(i,L) + dt * fhcat(i) + dt2 *
     X   (dfhcat_dhcat(i) * fhcat(i) + dfhcat_dv(i) * fv(i))
        mcaL(i,L) =  mcaL(i,L) + dt * fmcaL(i) + dt2 *
     X   (dfmcaL_dmcaL(i) * fmcaL(i) + dfmcaL_dv(i) * fv(i))
        mar(i,L) =   mar(i,L) + dt * fmar(i) + dt2 *
     X   (dfmar_dmar(i) * fmar(i) + dfmar_dv(i) * fv(i))
         end do


              end do



2001          CONTINUE


1000    CONTINUE
        END



C  SETS UP TABLES FOR RATE FUNCTIONS
       SUBROUTINE SUPBASK_SETUP
     X   (alpham_naf, betam_naf, dalpham_naf, dbetam_naf,
     X    alphah_naf, betah_naf, dalphah_naf, dbetah_naf,
     X    alpham_kdr, betam_kdr, dalpham_kdr, dbetam_kdr,
     X    alpham_ka , betam_ka , dalpham_ka , dbetam_ka ,
     X    alphah_ka , betah_ka , dalphah_ka , dbetah_ka ,
     X    alpham_k2 , betam_k2 , dalpham_k2 , dbetam_k2 ,
     X    alphah_k2 , betah_k2 , dalphah_k2 , dbetah_k2 ,
     X    alpham_km , betam_km , dalpham_km , dbetam_km ,
     X    alpham_kc , betam_kc , dalpham_kc , dbetam_kc ,
     X    alpham_cat, betam_cat, dalpham_cat, dbetam_cat,
     X    alphah_cat, betah_cat, dalphah_cat, dbetah_cat,
     X    alpham_caL, betam_caL, dalpham_caL, dbetam_caL,
     X    alpham_ar , betam_ar , dalpham_ar , dbetam_ar)

      INTEGER I,J,K

      real*8 minf, hinf, taum, tauh, V, Z, shift_hnaf,
     X  shift_mkdr,
     X alpham_naf(0:640),betam_naf(0:640),dalpham_naf(0:640),
     X   dbetam_naf(0:640),
     X alphah_naf(0:640),betah_naf(0:640),dalphah_naf(0:640),
     X   dbetah_naf(0:640),
     X alpham_kdr(0:640),betam_kdr(0:640),dalpham_kdr(0:640),
     X   dbetam_kdr(0:640),
     X alpham_ka(0:640), betam_ka(0:640),dalpham_ka(0:640) ,
     X   dbetam_ka(0:640),
     X alphah_ka(0:640), betah_ka(0:640), dalphah_ka(0:640),
     X   dbetah_ka(0:640),
     X alpham_k2(0:640), betam_k2(0:640), dalpham_k2(0:640),
     X   dbetam_k2(0:640),
     X alphah_k2(0:640), betah_k2(0:640), dalphah_k2(0:640),
     X   dbetah_k2(0:640),
     X alpham_km(0:640), betam_km(0:640), dalpham_km(0:640),
     X   dbetam_km(0:640),
     X alpham_kc(0:640), betam_kc(0:640), dalpham_kc(0:640),
     X   dbetam_kc(0:640),
     X alpham_cat(0:640),betam_cat(0:640),dalpham_cat(0:640),
     X   dbetam_cat(0:640),
     X alphah_cat(0:640),betah_cat(0:640),dalphah_cat(0:640),
     X   dbetah_cat(0:640),
     X alpham_caL(0:640),betam_caL(0:640),dalpham_caL(0:640),
     X   dbetam_caL(0:640),
     X alpham_ar(0:640), betam_ar(0:640), dalpham_ar(0:640),
     X   dbetam_ar(0:640)
C FOR VOLTAGE, RANGE IS -120 TO +40 MV (absol.), 0.25 MV RESOLUTION


       DO 1, I = 0, 640
          V = dble (I)
          V = (V / 4.d0) - 120.d0

c gNa
           minf = 1.d0/(1.d0 + dexp((-V-38.d0)/10.d0))
           if (v.le.-30.d0) then
            taum = .0125d0 + .1525d0*dexp((v+30.d0)/10.d0)
           else
            taum = .02d0 + .145d0*dexp((-v-30.d0)/10.d0)
           endif
c from interneuron data, Martina & Jonas 1997, tau x 0.5
           alpham_naf(i) = minf / taum
           betam_naf(i) = 1.d0/taum - alpham_naf(i)

            shift_hnaf =  0.d0
        hinf = 1.d0/(1.d0 +
     x     dexp((v + shift_hnaf + 58.3d0)/6.7d0))
        tauh = 0.225d0 + 1.125d0/(1.d0+dexp((v+37.d0)/15.d0))
c from interneuron data, Martina & Jonas 1997, tau x 0.5
            alphah_naf(i) = hinf / tauh
            betah_naf(i) = 1.d0/tauh - alphah_naf(i)

          shift_mkdr = 0.d0
c delayed rectifier, non-inactivating
       minf = 1.d0/(1.d0+dexp((-v-shift_mkdr-27.d0)/11.5d0))
            if (v.le.-10.d0) then
             taum = .25d0 + 4.35d0*dexp((v+10.d0)/10.d0)
            else
             taum = .25d0 + 4.35d0*dexp((-v-10.d0)/10.d0)
            endif
              alpham_kdr(i) = minf / taum
              betam_kdr(i) = 1.d0 /taum - alpham_kdr(i)
c from Martina, Schultz et al., 1998

c A current: Huguenard & McCormick 1992, J Neurophysiol (TCR)
            minf = 1.d0/(1.d0 + dexp((-v-60.d0)/8.5d0))
            hinf = 1.d0/(1.d0 + dexp((v+78.d0)/6.d0))
        taum = .185d0 + .5d0/(dexp((v+35.8d0)/19.7d0) +
     x                            dexp((-v-79.7d0)/12.7d0))
        if (v.le.-63.d0) then
         tauh = .5d0/(dexp((v+46.d0)/5.d0) +
     x                  dexp((-v-238.d0)/37.5d0))
        else
         tauh = 9.5d0
        endif
           alpham_ka(i) = minf/taum
           betam_ka(i) = 1.d0 / taum - alpham_ka(i)
           alphah_ka(i) = hinf / tauh
           betah_ka(i) = 1.d0 / tauh - alphah_ka(i)

c h-current (anomalous rectifier), Huguenard & McCormick, 1992
           minf = 1.d0/(1.d0 + dexp((v+75.d0)/5.5d0))
           taum = 1.d0/(dexp(-14.6d0 -0.086d0*v) +
     x                   dexp(-1.87 + 0.07d0*v))
           alpham_ar(i) = minf / taum
           betam_ar(i) = 1.d0 / taum - alpham_ar(i)

c K2 K-current, McCormick & Huguenard
             minf = 1.d0/(1.d0 + dexp((-v-10.d0)/17.d0))
             hinf = 1.d0/(1.d0 + dexp((v+58.d0)/10.6d0))
            taum = 4.95d0 + 0.5d0/(dexp((v-81.d0)/25.6d0) +
     x                  dexp((-v-132.d0)/18.d0))
            tauh = 60.d0 + 0.5d0/(dexp((v-1.33d0)/200.d0) +
     x                  dexp((-v-130.d0)/7.1d0))
             alpham_k2(i) = minf / taum
             betam_k2(i) = 1.d0/taum - alpham_k2(i)
             alphah_k2(i) = hinf / tauh
             betah_k2(i) = 1.d0 / tauh - alphah_k2(i)

c voltage part of C-current, using 1994 kinetics, shift 60 mV
              if (v.le.-10.d0) then
       alpham_kc(i) = (2.d0/37.95d0)*dexp((v+50.d0)/11.d0 -
     x                                     (v+53.5)/27.d0)
       betam_kc(i) = 2.d0*dexp((-v-53.5d0)/27.d0)-alpham_kc(i)
               else
       alpham_kc(i) = 2.d0*dexp((-v-53.5d0)/27.d0)
       betam_kc(i) = 0.d0
               endif
c Speed-up of C kinetics here.
          alpham_kc(i) = 2.d0 * alpham_kc(i)
           betam_kc(i) = 2.d0 *  betam_kc(i)

c high-threshold gCa, from 1994, with 60 mV shift & no inactivn.
            alpham_cal(i) = 1.6d0/(1.d0+dexp(-.072d0*(v-5.d0)))
            betam_cal(i) = 0.1d0 * ((v+8.9d0)/5.d0) /
     x          (dexp((v+8.9d0)/5.d0) - 1.d0)

c M-current, from plast.f, with 60 mV shift
        alpham_km(i) = .02d0/(1.d0+dexp((-v-20.d0)/5.d0))
        betam_km(i) = .01d0 * dexp((-v-43.d0)/18.d0)

c T-current, from Destexhe et al., 1996, pg. 170
         minf = 1.d0/(1.d0 + dexp((-v-52.d0)/7.4d0))
         hinf = 1.d0/(1.d0 + dexp((v+80.d0)/5.d0))
         taum = 1.d0 + .33d0/(dexp((v+27.d0)/10.d0) +
     x                  dexp((-v-102.d0)/15.d0))
         tauh = 28.3d0 +.33d0/(dexp((v+48.d0)/4.d0) +
     x                     dexp((-v-407.d0)/50.d0))
              alpham_cat(i) = minf / taum
              betam_cat(i) = 1.d0/taum - alpham_cat(i)
              alphah_cat(i) = hinf / tauh
              betah_cat(i) = 1.d0 / tauh - alphah_cat(i)

1        CONTINUE

         do 2, i = 0, 639

      dalpham_naf(i) = (alpham_naf(i+1)-alpham_naf(i))/.25d0
      dbetam_naf(i) = (betam_naf(i+1)-betam_naf(i))/.25d0
      dalphah_naf(i) = (alphah_naf(i+1)-alphah_naf(i))/.25d0
      dbetah_naf(i) = (betah_naf(i+1)-betah_naf(i))/.25d0
      dalpham_kdr(i) = (alpham_kdr(i+1)-alpham_kdr(i))/.25d0
      dbetam_kdr(i) = (betam_kdr(i+1)-betam_kdr(i))/.25d0
      dalpham_ka(i) = (alpham_ka(i+1)-alpham_ka(i))/.25d0
      dbetam_ka(i) = (betam_ka(i+1)-betam_ka(i))/.25d0
      dalphah_ka(i) = (alphah_ka(i+1)-alphah_ka(i))/.25d0
      dbetah_ka(i) = (betah_ka(i+1)-betah_ka(i))/.25d0
      dalpham_k2(i) = (alpham_k2(i+1)-alpham_k2(i))/.25d0
      dbetam_k2(i) = (betam_k2(i+1)-betam_k2(i))/.25d0
      dalphah_k2(i) = (alphah_k2(i+1)-alphah_k2(i))/.25d0
      dbetah_k2(i) = (betah_k2(i+1)-betah_k2(i))/.25d0
      dalpham_km(i) = (alpham_km(i+1)-alpham_km(i))/.25d0
      dbetam_km(i) = (betam_km(i+1)-betam_km(i))/.25d0
      dalpham_kc(i) = (alpham_kc(i+1)-alpham_kc(i))/.25d0
      dbetam_kc(i) = (betam_kc(i+1)-betam_kc(i))/.25d0
      dalpham_cat(i) = (alpham_cat(i+1)-alpham_cat(i))/.25d0
      dbetam_cat(i) = (betam_cat(i+1)-betam_cat(i))/.25d0
      dalphah_cat(i) = (alphah_cat(i+1)-alphah_cat(i))/.25d0
      dbetah_cat(i) = (betah_cat(i+1)-betah_cat(i))/.25d0
      dalpham_caL(i) = (alpham_cal(i+1)-alpham_cal(i))/.25d0
      dbetam_caL(i) = (betam_cal(i+1)-betam_cal(i))/.25d0
      dalpham_ar(i) = (alpham_ar(i+1)-alpham_ar(i))/.25d0
      dbetam_ar(i) = (betam_ar(i+1)-betam_ar(i))/.25d0
2      CONTINUE

         do i = 640, 640
      dalpham_naf(i) =  dalpham_naf(i-1)
      dbetam_naf(i) =  dbetam_naf(i-1)
      dalphah_naf(i) = dalphah_naf(i-1)
      dbetah_naf(i) = dbetah_naf(i-1)
      dalpham_kdr(i) =  dalpham_kdr(i-1)
      dbetam_kdr(i) =  dbetam_kdr(i-1)
      dalpham_ka(i) =  dalpham_ka(i-1)
      dbetam_ka(i) =  dbetam_ka(i-1)
      dalphah_ka(i) =  dalphah_ka(i-1)
      dbetah_ka(i) =  dbetah_ka(i-1)
      dalpham_k2(i) =  dalpham_k2(i-1)
      dbetam_k2(i) =  dbetam_k2(i-1)
      dalphah_k2(i) =  dalphah_k2(i-1)
      dbetah_k2(i) =  dbetah_k2(i-1)
      dalpham_km(i) =  dalpham_km(i-1)
      dbetam_km(i) =  dbetam_km(i-1)
      dalpham_kc(i) =  dalpham_kc(i-1)
      dbetam_kc(i) =  dbetam_kc(i-1)
      dalpham_cat(i) =  dalpham_cat(i-1)
      dbetam_cat(i) =  dbetam_cat(i-1)
      dalphah_cat(i) =  dalphah_cat(i-1)
      dbetah_cat(i) =  dbetah_cat(i-1)
      dalpham_caL(i) =  dalpham_caL(i-1)
      dbetam_caL(i) =  dbetam_caL(i-1)
      dalpham_ar(i) =  dalpham_ar(i-1)
      dbetam_ar(i) =  dbetam_ar(i-1)
       end do   

       END

        SUBROUTINE SUPBASKMAJ
C BRANCHED ACTIVE DENDRITES
     X             (GL,GAM,GKDR,GKA,GKC,GKAHP,GK2,GKM,
     X              GCAT,GCAL,GNAF,GNAP,GAR,
     X    CAFOR,JACOB,C,BETCHI,NEIGH,NNUM)
c Conductances: leak gL, coupling g, delayed rectifier gKDR, A gKA,
c C gKC, AHP gKAHP, K2 gK2, M gKM, low thresh Ca gCAT, high thresh
c gCAL, fast Na gNAF, persistent Na gNAP, h or anom. rectif. gAR.
c Note VAR = equil. potential for anomalous rectifier.
c Soma = comp. 1; 4 dendrites each with 13 compartments, 6-comp. axon
c Drop "glc"-like terms, just using "gl"-like
c CAFOR corresponds to "phi" in Traub et al., 1994
c Consistent set of units: nF, mV, ms, nA, microS

        INTEGER, PARAMETER:: numcomp = 59
        REAL*8 C(numcomp),GL(numcomp),GAM(0:numcomp,0:numcomp)
        REAL*8 GNAF(numcomp),GCAT(numcomp)
        REAL*8 GKDR(numcomp),GKA(numcomp),GKC(numcomp)
        REAL*8 GKAHP(numcomp),GCAL(numcomp),GAR(numcomp)
        REAL*8 GK2(numcomp),GKM(numcomp),GNAP(numcomp)
        REAL*8 JACOB(numcomp,numcomp)
        REAL*8 RI_SD,RI_AXON,RM_SD,RM_AXON,CDENS
        INTEGER LEVEL(numcomp)
        REAL*8 GNAF_DENS(0:9), GCAT_DENS(0:9), GKDR_DENS(0:9)
        REAL*8 GKA_DENS(0:9), GKC_DENS(0:9), GKAHP_DENS(0:9)
        REAL*8 GCAL_DENS(0:9), GK2_DENS(0:9), GKM_DENS(0:9)
        REAL*8 GNAP_DENS(0:9), GAR_DENS(0:9)
        REAL*8 RES, RINPUTi, ELEN(numcomp)
        REAL*8 RSOMA, PI, BETCHI(numcomp), CAFOR(numcomp)
        REAL*8 RAD(numcomp), LEN(numcomp), GAM1, GAM2
        REAL*8 RIN, D(numcomp), AREA(numcomp), RI, Z
        INTEGER NEIGH(numcomp,5), NNUM(numcomp), i, j, k, it
C FOR ESTABLISHING TOPOLOGY OF COMPARTMENTS

        RI_SD = 200.d0
c       RM_SD = 50000.d0
        RM_SD = 25000.d0
        RI_AXON = 100.d0
        RM_AXON = 1000.d0
        CDENS = 1.d0

        PI = 3.14159d0

        gnaf_dens(0) = 400.d0
        gnaf_dens(1) =  60.d0
        gnaf_dens(2) =  60.d0
        gnaf_dens(3) =  60.d0
        do i = 4, 9
c         gnaf_dens(i) = 60.d0
          gnaf_dens(i) = 10.d0
        end do

        gkdr_dens(0) = 400.d0
        gkdr_dens(1) = 100.d0
        gkdr_dens(2) = 100.d0
        gkdr_dens(3) = 100.d0
        do i = 4, 9
         gkdr_dens(i) = 10.d0
c        gkdr_dens(i) = 60.d0
        end do

        gnap_dens(0) = 0.d0
        do i = 1, 9
          gnap_dens(i) = 0.01d0 * gnaf_dens(i)
        end do

        gcat_dens(0) = 0.d0
        do i = 1, 3
          gcat_dens(i) = 0.05d0
        end do
        do i = 4, 9
          gcat_dens(i) = 2.d0
        end do

        gcal_dens(0) = 0.d0
        do i = 1, 3
c         gcal_dens(i) = 0.5d0
          gcal_dens(i) = 0.1d0
        end do
        do i = 4, 9
c         gcal_dens(i) = 0.5d0
          gcal_dens(i) = 0.2d0
        end do

        gka_dens(0) = 1.d0
        gka_dens(1) =  1.d0
        gka_dens(2) =  1.d0
        gka_dens(3) =  1.d0
        do i = 4, 9
         gka_dens(i) = 1.0d0
        end do

        gkc_dens(0) = 0.d0
        do i = 1, 9
c        gkc_dens(i) = 10.00d0
         gkc_dens(i) = 25.00d0
        end do

        gkm_dens(0) = 0.d0
        do i = 1, 9
         gkm_dens(i) = 0.50d0
        end do

        gk2_dens(0) = .5d0
        do i = 1, 9
         gk2_dens(i) = 0.50d0
        end do

        gkahp_dens(0) = 0.d0
        do i = 1, 9
         gkahp_dens(i) = 0.10d0
        end do

        gar_dens(0) = 0.d0
        do i = 1, 9
         gar_dens(i) = 0.025d0
        end do

c       WRITE   (6,9988)
9988    FORMAT(2X,'I',4X,'NADENS',' CADENS(L)',' KDRDEN',' KAHPDE',
     X     ' KCDENS',' KADENS')
c       DO 9989, I = 0, 9
        DO I = 0, 9
c         WRITE (6,9990) I, gnaf_dens(i), gcaL_dens(i), gkdr_dens(i),
c    X  gkahp_dens(i), gkc_dens(i), gka_dens(i)
9990    FORMAT(2X,I2,2X,F6.2,1X,F6.2,1X,F6.2,1X,F6.2,1X,F6.2,1X,F6.2)
        END DO
9989    CONTINUE


        level(1) = 1
        do i = 2, 41, 13
         level(i) = 2
        end do
        do i = 3, 42, 13
           level(i) = 3
           level(i+1) = 3
        end do
        do i = 5, 44, 13
           level(i) = 4
           level(i+1) = 4
           level(i+2) = 4
        end do
        do i = 8, 47, 13
           level(i) = 5
           level(i+1) = 5
           level(i+2) = 5
        end do
        do i = 11, 50, 13
           level(i) = 6
           level(i+1) = 7
           level(i+2) = 8
           level(i+3) = 9
        end do

        do i = 54, 59
         level(i) = 0
        end do

c connectivity of axon
        nnum(54) = 2
        nnum(55) = 3
        nnum(56) = 3
        nnum(58) = 3
        nnum(57) = 1
        nnum(59) = 1
         neigh(54,1) =  1
         neigh(54,2) = 55
         neigh(55,1) = 54
         neigh(55,2) = 56
         neigh(55,3) = 58
         neigh(56,1) = 55
         neigh(56,2) = 57
         neigh(56,3) = 58
         neigh(58,1) = 55
         neigh(58,2) = 56
         neigh(58,3) = 59
         neigh(57,1) = 56
         neigh(59,1) = 58

c connectivity of SD part
          nnum(1) = 5
          neigh(1,1) = 54
          neigh(1,2) =  2
          neigh(1,3) = 15
          neigh(1,4) = 28
          neigh(1,5) = 41

          do i = 2, 41, 13
           nnum(i) = 3
           neigh(i,1) = 1
           neigh(i,2) = i + 1
           neigh(i,3) = i + 2
          end do

          do i = 3, 42, 13
           nnum(i) = 4
           neigh(i,1) = i - 1
           neigh(i,2) = i + 1
           neigh(i,3) = i + 2
           neigh(i,4) = i + 3
          end do

          do i = 4, 43, 13
           nnum(i) = 3
           neigh(i,1) = i - 2
           neigh(i,2) = i - 1
           neigh(i,3) = i + 3
          end do

          do i = 5, 44, 13
           nnum(i) = 3
           neigh(i,1) = i - 2
           neigh(i,2) = i + 1
           neigh(i,3) = i + 3
          end do

          do i = 6, 45, 13
           nnum(i) = 3
            neigh(i,1) = i - 3
            neigh(i,2) = i - 1
            neigh(i,3) = i + 3
          end do

          do i = 7, 46, 13
           nnum(i) = 2
           neigh(i,1) = i - 3
           neigh(i,2) = i + 3
          end do

          do i = 8, 47, 13
           nnum(i) = 2
           neigh(i,1) = i - 3
           neigh(i,2) = i + 3
          end do

          do i = 9, 48, 13
           nnum(i) = 1
           neigh(i,1) = i - 3
          end do

          do i = 10, 49, 13
           nnum(i) = 1
           neigh(i,1) = i - 3
          end do

          do i = 11, 50, 13
           nnum(i) = 2
           neigh(i,1) = i - 3
           neigh(i,2) = i + 1
          end do

          do i = 12, 51, 13
           nnum(i) = 2
           neigh(i,1) = i - 1
           neigh(i,2) = i + 1
          end do

          do i = 13, 52, 13
           nnum(i) = 2
           neigh(i,1) = i - 1
           neigh(i,2) = i + 1
          end do

          do i = 14, 53, 13
           nnum(i) = 1
           neigh(i,1) = i - 1
          end do

c        DO 332, I = 1, 59
         DO I = 1, numcomp
c          WRITE(6,3330) I, NEIGH(I,1),NEIGH(I,2),NEIGH(I,3),NEIGH(I,4),
c    X NEIGH(I,5)
3330     FORMAT(2X,I5,I5,I5,I5,I5,I5)
         END DO
332      CONTINUE
c         DO 858, I = 1, 59
          DO I = 1, 59
c          DO 858, J = 1, NNUM(I)
           DO J = 1, NNUM(I)
            K = NEIGH(I,J)
            IT = 0
c           DO 859, L = 1, NNUM(K)
            DO L = 1, NNUM(K)
             IF (NEIGH(K,L).EQ.I) IT = 1
            END DO
859         CONTINUE
             IF (IT.EQ.0) THEN
c             WRITE(6,8591) I, K
8591          FORMAT(' ASYMMETRY IN NEIGH MATRIX ',I4,I4)
             ENDIF
           END DO
           END DO
858       CONTINUE

c length and radius of axonal compartments
          do i = 54, 59
            len(i) = 50.d0
          end do
c         rad(54) = 0.80d0
c         rad(55) = 0.7d0
          rad(54) = 0.70d0
          rad(55) = 0.6d0
          do i = 56, 59
           rad(i) = 0.5d0
          end do

c  length and radius of SD compartments
          len(1) = 20.d0
          rad(1) = 7.5d0

          do i = 2, 53
           len(i) = 40.d0
          end do

          rad(2) =   1.06d0
          rad(3) =   rad(2) / 1.59d0
          rad(4) =   rad(2) / 1.59d0
          rad(5) =   rad(2) / 2.53d0
          rad(6) =   rad(2) / 2.53d0
          rad(7) =   rad(2) / 1.59d0
          rad(8) =   rad(2) / 2.53d0
          rad(9) =   rad(2) / 2.53d0
          rad(10) =  rad(2) / 1.59d0
          rad(11) =  rad(2) / 2.53d0
          rad(12) =  rad(2) / 2.53d0
          rad(13) =  rad(2) / 2.53d0
          rad(14) =  rad(2) / 2.53d0

          do i = 15, 53
           rad(i) = rad(i-13)
          end do

c       WRITE(6,919)
919     FORMAT('COMPART.',' LEVEL ',' RADIUS ',' LENGTH(MU)')
c       DO 920, I = 1, 59
c920      WRITE(6,921) I, LEVEL(I), RAD(I), LEN(I)
921     FORMAT(I3,5X,I2,3X,F6.2,1X,F6.1,2X,F4.3)

c       DO 120, I = 1, 59
        DO I = 1, numcomp
          AREA(I) = 2.d0 * PI * RAD(I) * LEN(I)
C NO CORRECTION FOR CONTRIBUTION OF SPINES TO AREA
          K = LEVEL(I)
          C(I) = CDENS * AREA(I) * (1.D-8)

           if (k.ge.1) then
          GL(I) = (1.D-2) * AREA(I) / RM_SD
           else
          GL(I) = (1.D-2) * AREA(I) / RM_AXON
           endif

          GNAF(I) = GNAF_DENS(K) * AREA(I) * (1.D-5)
          GNAP(I) = GNAP_DENS(K) * AREA(I) * (1.D-5)
          GCAT(I) = GCAT_DENS(K) * AREA(I) * (1.D-5)
          GKDR(I) = GKDR_DENS(K) * AREA(I) * (1.D-5)
          GKA(I) = GKA_DENS(K) * AREA(I) * (1.D-5)
          GKC(I) = GKC_DENS(K) * AREA(I) * (1.D-5)
          GKAHP(I) = GKAHP_DENS(K) * AREA(I) * (1.D-5)
          GCAL(I) = GCAL_DENS(K) * AREA(I) * (1.D-5)
          GK2(I) = GK2_DENS(K) * AREA(I) * (1.D-5)
          GKM(I) = GKM_DENS(K) * AREA(I) * (1.D-5)
          GAR(I) = GAR_DENS(K) * AREA(I) * (1.D-5)
c above conductances should be in microS
         END DO
120           continue

         Z = 0.d0
c        DO 1019, I = 2, 53
         DO I = 2, 53
           Z = Z + AREA(I)
         END DO
1019     CONTINUE
c        WRITE(6,1020) Z
1020     FORMAT(2X,' TOTAL DENDRITIC AREA ',F7.0)

c       DO 140, I = 1, 59
        DO I = 1, numcomp
c       DO 140, K = 1, NNUM(I)
        DO K = 1, NNUM(I)
         J = NEIGH(I,K)
           if (level(i).eq.0) then
               RI = RI_AXON
           else
               RI = RI_SD
           endif
         GAM1 =100.d0 * PI * RAD(I) * RAD(I) / ( RI * LEN(I) )

           if (level(j).eq.0) then
               RI = RI_AXON
           else
               RI = RI_SD
           endif
         GAM2 =100.d0 * PI * RAD(J) * RAD(J) / ( RI * LEN(J) )

         GAM(I,J) = 2.d0/( (1.d0/GAM1) + (1.d0/GAM2) )
         END DO
         END DO
140     CONTINUE
c gam computed in microS

c       DO 299, I = 1, 59
        DO I = 1, numcomp
299       BETCHI(I) = .05d0
        END DO
        BETCHI( 1) =  .02d0

c       DO 300, I = 1, 59
        DO I = 1, numcomp
c300     D(I) = 2.D-4
300     D(I) = 1.D-4
        END DO
c       DO 301, I = 1, 59
        DO I = 1, numcomp
c        IF (LEVEL(I).EQ.1) D(I) = 5.D-3
         IF (LEVEL(I).EQ.1) D(I) = 2.D-4
        END DO
301     CONTINUE
C  NOTE NOTE NOTE  (DIFFERENT FROM SWONG)


c      DO 160, I = 1, 59
       DO I = 1, numcomp
160     CAFOR(I) = 5200.d0 / (AREA(I) * D(I))
       END DO
C     NOTE CORRECTION

c       do 200, i = 1, 59
        do i = 1, numcomp
200     C(I) = 1000.d0 * C(I)
        end do
C     TO GO FROM MICROF TO NF.

c     DO 909, I = 1, 59
      DO I = 1, numcomp
       JACOB(I,I) = - GL(I)
c     DO 909, J = 1, NNUM(I)
      DO J = 1, NNUM(I)
         K = NEIGH(I,J)
         IF (I.EQ.K) THEN
c            WRITE(6,510) I
510          FORMAT(' UNEXPECTED SYMMETRY IN NEIGH ',I4)
         ENDIF
         JACOB(I,K) = GAM(I,K)
         JACOB(I,I) = JACOB(I,I) - GAM(I,K)
       END DO
       END DO
909   CONTINUE

c 15 Jan. 2001: make correction for c(i)
          do i = 1, numcomp
          do j = 1, numcomp
             jacob(i,j) = jacob(i,j) / c(i)
          end do
          end do

c      DO 500, I = 1, 59
       DO I = 1, numcomp
c       WRITE (6,501) I,C(I)
501     FORMAT(1X,I2,' C(I) = ',F7.4)
        END DO
500     CONTINUE
        END