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purkinje_bignet21.f
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purkinje_bignet21.f
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c 26 March 2007, network version of /purkinje_parallel/purkinje.f, to run on multiple nodes
! 21 March 2007, version of purkinje.f to run on a node.
c 13 March 2007, begin construction of Purkinje cell model, starting with bask.f
c Sources include Llinas & Sugimori; Roth & Hausser for cell structure and passive
c parameters; de Schutter & Bower & Miyasho et al. for many of the active properties;
c Roth & Hausser for anomalous rectifier; Martina et al for data on delayed rectifier.
c 1 soma cylinder (level 1), 24 smooth dendritic compartments, #2 - 25.
c compartments 2 & 3 are dendritic shaft (level 2), 4 - 25 rest of smooth dendrites (level 3.
c There are 44 "canonical treelets" for the spiny dendrites; 2 treelets attach to
c each of the level 3 smooth dendritic compartments.
c A spiny treelet has 12 compartments: 4 along the main root, then 4 in each of 2 symmetrical
c branches. The spiny dendrites are compartments 26 - 553.
c Axon has 6 10-micron compartments, #554 - 559, level 0.
c Active conductances: gnaf (m cubed h) - axon will use shifted version.
c gnap (m cubed)
c gcaP (m) P channels
c gcaT (m h) T channels
c gcaR (m h) R channels
c gar (m) anomolous rectifier
c gKDR (m fourth) delayed rectifier, leaving out inactivation
c gKM (m) M current, also called persistent K
c gKA (m fourth x h) A current
c gKD (m h) D current
c gKC - try my original structure
c gKAHP - try my original structure.
c 12 Active conductances in all
PROGRAM PURKINJE_NET
PARAMETER (num_purk = 1000, np = 1000, ! i.e. = num_purk
x numnodes = 50)
integer, parameter :: cellspernode = 20
integer, parameter :: totaxgj = 2500
c integer, parameter :: num_axgjcompallow = 1
integer, parameter :: num_axgjcompallow = 3
double precision, parameter :: gapcon = 6.0d-3
c double precision, parameter :: gapcon = 5.5d-3
c double precision, parameter :: gapcon = 0.0d-3
double precision, parameter :: noisepe = 0.8d0/75.d0 ! 0.8 ms ~= 1333 dt
double precision gj_axon554 (cellspernode), gj_axon554_global(np) ! these used by mpi
double precision gj_axon555 (cellspernode), gj_axon555_global(np) ! these used by mpi
double precision gj_axon556 (cellspernode), gj_axon556_global(np) ! these used by mpi
double precision gj_axon557 (cellspernode), gj_axon557_global(np) ! these used by mpi
double precision gj_axon558 (cellspernode), gj_axon558_global(np) ! these used by mpi
double precision gj_axon559 (cellspernode), gj_axon559_global(np) ! these used by mpi
double precision soma_local (cellspernode), soma_global(np) ! these used by mpi
integer na, nb, display, ectr /0/
integer gjtable (totaxgj,4)
c integer table_axgjcompallow (num_axgjcompallow) /558/
integer table_axgjcompallow (num_axgjcompallow)
c x /555,556,557,558,559/ ! IF THIS IS ALTERED, MPI CODE MUST BE ALTERED AS WELL.
x /554,555,556 / ! IF THIS IS ALTERED, MPI CODE MUST BE ALTERED AS WELL.
INTEGER J1, I, J, K, L, O, ISEED, K1, thisno
REAL*8 TIMTOT, Z, Z1, Z2, Z3, curr(559,np), c(559), DT
REAL*8 TIMER, gettime, time1, time2, time
real*8 ranvec(np), seed /137.d0/
c CINV is 1/C, i.e. inverse capacitance
real*8 v(559,np), chi(559,np), cinv(559), gL(559),
x gNaF(559), gNaP(559), gCaP(559), gCaT(559), gCaR(559),
x gar(559), gKDR(559), gKM(559), gKA(559), gKD(559),
x gKC(559), gKAHP(559)
real*8 jacob(559,559), betchi(559), gam(0:559,0:559)
real*8
x mnaf(559,np), hnaf(559,np),
x mnap(559,np),
x mcap(559,np),
x mcat(559,np), hcat(559,np),
x mcar(559,np), hcar(559,np),
x mar (559,np),
x mkdr(559,np),
x mkm(559,np),
x mka(559,np), hka(559,np),
x mkd(559,np), hkd(559,np),
x mkc(559,np),
x mkahp(559,np)
real*8 gampa(559,np),gnmda(559,np),ggaba_a(559,np)
real*8 kdr_shift, cafor(59)
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_nap(0:640),betam_nap(0:640),dalpham_nap(0:640),
X dbetam_nap(0:640),
X alpham_cap(0:640),betam_cap(0:640),dalpham_cap(0:640),
X dbetam_cap(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_car(0:640),betam_car(0:640),dalpham_car(0:640),
X dbetam_car(0:640),
X alphah_car(0:640),betah_car(0:640),dalphah_car(0:640),
X dbetah_car(0:640),
X alpham_ar(0:640), betam_ar(0:640), dalpham_ar(0:640),
X dbetam_ar(0:640),
X alpham_kdr(0:640),betam_kdr(0:640),dalpham_kdr(0:640),
X dbetam_kdr(0:640),
X alpham_km(0:640), betam_km(0:640), dalpham_km(0:640),
X dbetam_km(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_kd(0:640), betam_kd(0:640),dalpham_kd(0:640) ,
X dbetam_kd(0:640),
X alphah_kd(0:640), betah_kd(0:640), dalphah_kd(0:640),
X dbetah_kd(0:640),
X alpham_kc(0:640), betam_kc(0:640), dalpham_kc(0:640),
X dbetam_kc(0:640)
real*8 vL,vk,vna,var,vca,vgaba_a
INTEGER NEIGH(559,6), NNUM(559)
c START EXECUTION PHASE
include 'mpif.h'
call mpi_init (info)
call mpi_comm_rank(mpi_comm_world, thisno, info)
call mpi_comm_size(mpi_comm_world, nodes , info)
time1 = gettime()
time1 = gettime()
na = thisno * cellspernode + 1
nb = na + cellspernode - 1 ! define which cells to be integrated on this node
c kdr_shift = 7.d0
kdr_shift = 10.d0
CALL PURKINJE_SETUP
X (alpham_naf, betam_naf, dalpham_naf, dbetam_naf,
X alphah_naf, betah_naf, dalphah_naf, dbetah_naf,
X alpham_nap, betam_nap, dalpham_nap, dbetam_nap,
X alpham_cap, betam_cap, dalpham_cap, dbetam_cap,
X alpham_cat, betam_cat, dalpham_cat, dbetam_cat,
X alphah_cat, betah_cat, dalphah_cat, dbetah_cat,
X alpham_car, betam_car, dalpham_car, dbetam_car,
X alphah_car, betah_car, dalphah_car, dbetah_car,
X alpham_ar , betam_ar , dalpham_ar , dbetam_ar,
X kdr_shift,
X alpham_kdr, betam_kdr, dalpham_kdr, dbetam_kdr,
X alpham_km , betam_km , dalpham_km , dbetam_km ,
X alpham_ka , betam_ka , dalpham_ka , dbetam_ka ,
X alphah_ka , betah_ka , dalphah_ka , dbetah_ka ,
X alpham_kd , betam_kd , dalpham_kd , dbetam_kd ,
X alphah_kd , betah_kd , dalphah_kd , dbetah_kd ,
X alpham_kc , betam_kc , dalpham_kc , dbetam_kc)
CALL PURKINJE_MAJ (GL,GAM,
X gNaF, gNaP, gCaP, gCaT, gCaR, gar, gKDR,
X gKM, gKA, gKD, gKC, gKAHP,
X CAFOR,JACOB,C,BETCHI,NEIGH,NNUM,thisno)
do i = 1, 559
cinv(i) = 1.d0 / c(i)
end do
c gnaf = 0.d0
c gnap = 0.d0
c gkdr = 0.d0
c gka = 0.d0
gkd = 0.d0
c gkm = 0.d0
gkc = 0.d0
gkahp = 0.d0
gcat = 0.d0
gcaP = 0.d0
gcaR = 0.d0
c gar = 0.d0
c Below introduces somatic shunt
c gL(1) = 100.d0 * gL(1)
gL(1) = 5.d0 * gL(1)
c below 4 statements disconnect axon from soma.
c gam(1,554) = 0.d0
c gam(554,1) = 0.d0
c jacob(1,554) = 0.d0
c jacob(554,1) = 0.d0
MG = 1.0d0
C IN MILLIMOLAR
VL = -80.d0
VK = - 85.d0
VNA = 45.d0
VCA = 135.d0
VAR = -30.d0
VGABA_A = -75.d0
DT = .00060d0
c DT = .00200d0
TIMTOT = 175.00d0
c TIMTOT = 8.d0 * dt
c timtot = 0.d0
c ? initialize membrane state variables?
c v = VL
v = -55.d0
k1 = idnint (4.d0 * (v(1,na) + 120.d0))
hnaf = alphah_naf(k1)/(alphah_naf(k1)+betah_naf(k1))
hka = alphah_ka(k1)/(alphah_ka(k1)+betah_ka(k1))
hkd = alphah_kd(k1)/(alphah_kd(k1)+betah_kd(k1))
hcat=alphah_cat(k1)/(alphah_cat(k1)+betah_cat(k1))
hcar=alphah_car(k1)/(alphah_car(k1)+betah_car(k1))
call purkinje_gapbld (thisno, np, totaxgj, gjtable,
X table_axgjcompallow, num_axgjcompallow, 1)
c Define tonic currents
call durand (seed, np, ranvec)
do L = 1, np
curr(1,L) = 0.35d0 + 0.10d0 * ranvec(L) ! for soma
end do
c Hyperpolarize one or more cells, to try to unmask spikelets
curr(1,21) = -0.25d0
curr(1,22) = -0.25d0
curr(1,23) = -0.25d0
curr(1,24) = -0.25d0
curr(1,25) = -0.25d0
curr(1,26) = -0.25d0
curr(1,27) = -0.25d0
curr(1,28) = -0.25d0
do L = 1, np
do i = 554, 559
curr(i,L) = 0.04d0
end do
end do
O = 0
TIME = 0.d0
TIMER = 0.d0 ! for periodic spikes
2000 TIME = TIME + DT
O = O + 1
IF (TIME .GT. TIMTOT) GO TO 2001
c if (mod(O,8000).eq.0) timer = timer + 4.8d0
c curr(1) = 1.50d0
c curr(1) = 0.75d0
c IF (MOD(O,100).EQ. 0 ) THEN
c IF ( i.eq.i ) THEN
c WRITE (6,904) TIME, v(1), v(14), chi(14),
c X V(25), v(554), v(557), v(558), v(559), curr(559)
c X ,curr(14)
c ENDIF
904 FORMAT(2X,F8.3,2X,8f8.2,f8.4,f9.4)
c Set up currents for ectopic spikes. Note the "quantization" of time introduced this way
if (mod(O,1333).eq.0) then
call durand (seed,np,ranvec)
do L = na, nb
if ((ranvec(L).gt.0.d0).and.(ranvec(L).le.noisepe)) then
curr(559,L) = 0.45d0
ectr = ectr + 1
else
curr(559,L) = 0.d0
endif
end do
end if
CALL PURKINJE_INT_NET (V,CHI,CINV,
X mnaf,hnaf,mnap,mcap,mcat,hcat,mcar,hcar,mar,
x mkdr,mkm,mka,hka,mkd,hkd,mkc,mkahp,
x dt,neigh,nnum,jacob,mg,
x vL,vk,vna,var,vca,vgaba_a,betchi,gam,gL,
x gnaf,gnap,gcaP,gcaT,gCaR,gar,
x gKDR,gKM,gKA,gKD,gKC,gKAHP,
x gampa,gnmda,ggaba_a,
x O,time,
X alpham_naf, betam_naf, dalpham_naf, dbetam_naf,
X alphah_naf, betah_naf, dalphah_naf, dbetah_naf,
X alpham_nap, betam_nap, dalpham_nap, dbetam_nap,
X alpham_cap, betam_cap, dalpham_cap, dbetam_cap,
X alpham_cat, betam_cat, dalpham_cat, dbetam_cat,
X alphah_cat, betah_cat, dalphah_cat, dbetah_cat,
X alpham_car, betam_car, dalpham_car, dbetam_car,
X alphah_car, betah_car, dalphah_car, dbetah_car,
X alpham_ar , betam_ar , dalpham_ar , dbetam_ar,
X alpham_kdr, betam_kdr, dalpham_kdr, dbetam_kdr,
X alpham_km , betam_km , dalpham_km , dbetam_km ,
X alpham_ka , betam_ka , dalpham_ka , dbetam_ka ,
X alphah_ka , betah_ka , dalphah_ka , dbetah_ka ,
X alpham_kd , betam_kd , dalpham_kd , dbetam_kd ,
X alphah_kd , betah_kd , dalphah_kd , dbetah_kd ,
X alpham_kc , betam_kc , dalpham_kc , dbetam_kc ,
X cafor,curr,
X np, initialize, na, nb, thisno, ! np = num_purk, but use shorter name here
X gapcon, totaxgj, gjtable)
c data broadcasting, followed by reading appropriate axonal voltages into locally known voltage array
if (mod(O,5).eq.0) then
! CODE BELOW SHARES ALL AXONAL COMPARTMENTS EXCEPT HILLOCK, AND SOMA.
k = 554
do L = na, nb
gj_axon554 (L - na + 1) = v(k,L)
end do
call mpi_allgather (gj_axon554,
x cellspernode, mpi_double_precision,
x gj_axon554_global, cellspernode, mpi_double_precision,
x mpi_comm_world, info)
do L = 1, np
v(k,L) = gj_axon554_global(L)
end do
c k = table_axgjcompallow(1)
k = 555
do L = na, nb
gj_axon555 (L - na + 1) = v(k,L)
end do
call mpi_allgather (gj_axon555,
x cellspernode, mpi_double_precision,
x gj_axon555_global, cellspernode, mpi_double_precision,
x mpi_comm_world, info)
do L = 1, np
v(k,L) = gj_axon555_global(L)
end do
c k = table_axgjcompallow(2)
k = 556
do L = na, nb
gj_axon556 (L - na + 1) = v(k,L)
end do
call mpi_allgather (gj_axon556,
x cellspernode, mpi_double_precision,
x gj_axon556_global, cellspernode, mpi_double_precision,
x mpi_comm_world, info)
do L = 1, np
v(k,L) = gj_axon556_global(L)
end do
c k = table_axgjcompallow(3)
k = 557
do L = na, nb
gj_axon557 (L - na + 1) = v(k,L)
end do
call mpi_allgather (gj_axon557,
x cellspernode, mpi_double_precision,
x gj_axon557_global, cellspernode, mpi_double_precision,
x mpi_comm_world, info)
do L = 1, np
v(k,L) = gj_axon557_global(L)
end do
c k = table_axgjcompallow(4)
k = 558
do L = na, nb
gj_axon558 (L - na + 1) = v(k,L)
end do
call mpi_allgather (gj_axon558,
x cellspernode, mpi_double_precision,
x gj_axon558_global, cellspernode, mpi_double_precision,
x mpi_comm_world, info)
do L = 1, np
v(k,L) = gj_axon558_global(L)
end do
c k = table_axgjcompallow(5)
k = 559
do L = na, nb
gj_axon559 (L - na + 1) = v(k,L)
end do
call mpi_allgather (gj_axon559,
x cellspernode, mpi_double_precision,
x gj_axon559_global, cellspernode, mpi_double_precision,
x mpi_comm_world, info)
do L = 1, np
v(k,L) = gj_axon559_global(L)
end do
do L = na, nb
soma_local (L - na + 1) = v(1,L)
end do
call mpi_allgather (soma_local,
x cellspernode, mpi_double_precision,
x soma_global, cellspernode, mpi_double_precision,
x mpi_comm_world, info)
do L = 1, np
v(1,L) = soma_global(L)
end do
endif
GO TO 2000
2001 CONTINUE
if (thisno.eq.0) then
write(6,897) ectr
897 format(i8,' axonal pulses on node 0')
time2 = gettime()
write(6,309) time2 - time1
309 format(' Elapsed time = ',f8.0,' secs')
endif
1000 CONTINUE
call mpi_finalize(info)
END
C SETS UP TABLES FOR RATE FUNCTIONS
SUBROUTINE PURKINJE_SETUP
X (alpham_naf, betam_naf, dalpham_naf, dbetam_naf,
X alphah_naf, betah_naf, dalphah_naf, dbetah_naf,
X alpham_nap, betam_nap, dalpham_nap, dbetam_nap,
X alpham_cap, betam_cap, dalpham_cap, dbetam_cap,
X alpham_cat, betam_cat, dalpham_cat, dbetam_cat,
X alphah_cat, betah_cat, dalphah_cat, dbetah_cat,
X alpham_car, betam_car, dalpham_car, dbetam_car,
X alphah_car, betah_car, dalphah_car, dbetah_car,
X alpham_ar , betam_ar , dalpham_ar , dbetam_ar,
X kdr_shift,
X alpham_kdr, betam_kdr, dalpham_kdr, dbetam_kdr,
X alpham_km , betam_km , dalpham_km , dbetam_km ,
X alpham_ka , betam_ka , dalpham_ka , dbetam_ka ,
X alphah_ka , betah_ka , dalphah_ka , dbetah_ka ,
X alpham_kd , betam_kd , dalpham_kd , dbetam_kd ,
X alphah_kd , betah_kd , dalphah_kd , dbetah_kd ,
X alpham_kc , betam_kc , dalpham_kc , dbetam_kc)
INTEGER I,J,K
real*8 minf, hinf, taum, tauh, V, Z,
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_nap(0:640),betam_nap(0:640),dalpham_nap(0:640),
X dbetam_nap(0:640),
X alpham_caP(0:640),betam_caP(0:640),dalpham_caP(0:640),
X dbetam_caP(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_car(0:640),betam_car(0:640),dalpham_car(0:640),
X dbetam_car(0:640),
X alphah_car(0:640),betah_car(0:640),dalphah_car(0:640),
X dbetah_car(0:640),
X alpham_ar(0:640), betam_ar(0:640), dalpham_ar(0:640),
X dbetam_ar(0:640),
X alpham_kdr(0:640),betam_kdr(0:640),dalpham_kdr(0:640),
X dbetam_kdr(0:640),
X alpham_km(0:640), betam_km(0:640), dalpham_km(0:640),
X dbetam_km(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_kd(0:640), betam_kd(0:640),dalpham_kd(0:640) ,
X dbetam_kd(0:640),
X alphah_kd(0:640), betah_kd(0:640), dalphah_kd(0:640),
X dbetah_kd(0:640),
X alpham_kc(0:640), betam_kc(0:640), dalpham_kc(0:640),
X dbetam_kc(0:640)
real*8 kdr_shift
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 gNaF
c T. Miyasho et al. 2001
alpham_naf(i) = 35.d0 / (0.d0 + dexp((v + 5.d0)/(-10.d0)))
betam_naf(i) = 7.d0 / (0.d0 + dexp((v+65.d0)/20.d0))
c T. Miyasho et al. 2001
alphah_naf(i) = 0.225d0 / (1.d0 + dexp((v+80.d0)/10.d0))
betah_naf(i) = 7.5d0 / (0.d0 + dexp((v-3.d0)/(-18.d0)))
c T. Miyasho et al. 2001
alpham_nap(i) = 200.d0 / (1.d0 + dexp((v-18.d0)/(-16.d0)))
betam_nap(i) = 25.d0 / (1.d0 + dexp((v+58.d0)/8.d0))
c P channels, T. Miyasho et al. 2001
alpham_cap(i) = 8.5d0 / (1.d0 + dexp((v-8.d0)/(-12.5d0)))
betam_cap(i) = 35.d0 / (1.d0 + dexp((v+74.d0)/14.5d0))
c T-current, from T. Miyasho et al., 2001
alpham_cat(i) = 2.6d0 / (1.d0 + dexp((v+21.d0)/(-8.d0)))
betam_cat(i) = 0.18d0 / (1.d0 + dexp((v+40.d0)/4.d0))
alphah_cat(i) = 0.0025d0 / (1.d0 + dexp((v+40.d0)/8.d0))
betah_cat(i) = 0.19d0 / (1.d0 + dexp((v+50.d0)/(-10.d0)))
c R-current, from T. Miyasho et al., 2001
alpham_car(i) = 2.6d0 / (1.d0 + dexp((v+7.d0)/(-8.d0)))
betam_car(i) = 0.18d0/ (1.d0 + dexp((v+26.d0)/4.d0))
alphah_car(i) = 0.0025/ (1.d0 + dexp((v+32.d0)/8.d0))
betah_car(i) = 0.19d0/ (1.d0 + dexp((v+42.d0)/(-10.d0)))
c h-current (anomalous rectifier), Roth and Hausser
alpham_ar(i) = 0.00063d0 * dexp (-0.063d0 * (v + 73.2d0))
betam_ar(i) = 0.00063d0 * dexp(0.079d0 * (v +73.2d0))
c delayed rectifier, non-inactivating. Start with mine (e.g. bask.f)
c Perhaps modify after comparison with Martina et al., 2003
v = v + kdr_shift
c minf = 1.d0/(1.d0+dexp((-v-27.d0)/11.5d0))
minf = 1.d0/(1.d0+dexp((-v-20.d0)/11.5d0)) ! elevate threshold
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 - for bask.f
v = v - kdr_shift
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 A current: T. Miyasho et al., 2001
alpham_ka(i) = 1.4d0 / (1.d0 + dexp((v+27.d0)/(-12.d0)))
betam_ka(i) = 0.49d0/ (1.d0 + dexp((v+30.d0)/4.d0))
alphah_ka(i) = 0.0175d0/(1.d0 + dexp((v+50.d0)/8.d0))
betah_ka(i) = 1.3 / (1.d0 + dexp((v+13.d0)/(-10.d0)))
c D current: T. Miyasho et al., 2001
alpham_kd(i) = 8.5d0 / (1.d0 + dexp((v+17.d0)/(-12.5d0)))
betam_kd(i) = 35.0d0/ (1.d0 + dexp((v+99.d0)/14.5d0))
alphah_kd(i) = 0.0015d0/(1.d0 + dexp((v+89.d0)/8.d0))
betah_kd(i) = 0.0055d0 / (1.d0 + dexp((v+83.d0)/(-8.d0)))
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)
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_nap(i) = (alpham_nap(i+1)-alpham_nap(i))/.25d0
dbetam_nap(i) = (betam_nap(i+1)-betam_nap(i))/.25d0
dalpham_cap(i) = (alpham_cap(i+1)-alpham_cap(i))/.25d0
dbetam_cap(i) = (betam_cap(i+1)-betam_cap(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_car(i) = (alpham_car(i+1)-alpham_car(i))/.25d0
dbetam_car(i) = (betam_car(i+1)-betam_car(i))/.25d0
dalphah_car(i) = (alphah_car(i+1)-alphah_car(i))/.25d0
dbetah_car(i) = (betah_car(i+1)-betah_car(i))/.25d0
dalpham_ar(i) = (alpham_ar(i+1)-alpham_ar(i))/.25d0
dbetam_ar(i) = (betam_ar(i+1)-betam_ar(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_km(i) = (alpham_km(i+1)-alpham_km(i))/.25d0
dbetam_km(i) = (betam_km(i+1)-betam_km(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_kd(i) = (alpham_kd(i+1)-alpham_kd(i))/.25d0
dbetam_kd(i) = (betam_kd(i+1)-betam_kd(i))/.25d0
dalphah_kd(i) = (alphah_kd(i+1)-alphah_kd(i))/.25d0
dbetah_kd(i) = (betah_kd(i+1)-betah_kd(i))/.25d0
dalpham_kc(i) = (alpham_kc(i+1)-alpham_kc(i))/.25d0
dbetam_kc(i) = (betam_kc(i+1)-betam_kc(i))/.25d0
2 CONTINUE
END
SUBROUTINE PURKINJE_MAJ (GL,GAM,
X gNaF, gNaP, gCaP, gCaT, gCaR, gar, gKDR,
X gKM, gKA, gKD, gKC, gKAHP,
X CAFOR,JACOB,C,BETCHI,NEIGH,NNUM,thisno)
c Conductances: see main program.
c Note VAR = equil. potential for anomalous rectifier.
c 1 soma cylinder (level 1), 24 smooth dendritic compartments, #2 - 25.
c compartments 2 & 3 are dendritic shaft (level 2), 4 - 25 rest of smooth dendrites (level 3.
c There are 44 "canonical treelets" for the spiny dendrites; 2 treelets attach to
c each of the level 3 smooth dendritic compartments (22 of them).
c A spiny treelet has 12 compartments: 4 along the main root, then 4 in each of 2 symmetrical
c branches. The spiny dendrites are compartments 26 - 553, level 4.
c Axon has 6 5-micron compartments, #554 - 559, level 0.
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
REAL*8 C(559),GL(559),GAM(0:559,0:559)
REAL*8 gnaf(559), gnap(559), gcap(559), gcat(559), gcar(559)
REAL*8 gar(559), gKDR(559), gKM(559), gKA(559), gKD(559),
X gKC(559), gKAHP(559)
REAL*8 JACOB(559,559),RI_SD,RI_AXON,RM_SD,RM_AXON,CDENS
INTEGER LEVEL(559), thisno
REAL*8 GNAF_DENS(0:4), GNAP_DENS(0:4)
REAL*8 GCAP_DENS(0:4), GCAT_DENS(0:4), GCAR_DENS(0:4)
REAL*8 GAR_DENS(0:4)
REAL*8 GKDR_DENS(0:4), GKM_DENS(0:4), GKA_DENS(0:4)
REAL*8 GKD_DENS(0:4), GKC_DENS(0:4), GKAHP_DENS(0:4)
REAL*8 RES, RINPUT, z1, z2, z, somaar
REAL*8 RSOMA, PI, BETCHI(559), CAFOR(559)
REAL*8 RAD(559), LEN(559), GAM1, GAM2, ELEN(559)
REAL*8 RIN, D(559), AREA(559), RI
INTEGER NEIGH(559,6), NNUM(559)
C FOR ESTABLISHING TOPOLOGY OF COMPARTMENTS
RI_SD = 115.d0 ! Roth & Hausser
c RM_SD = 125000.d0 ! Roth & Hausser
c RM_SD = 100000.d0 ! close to Roth & Hausser
RM_SD = 50000.d0 ! close to Miyasho et al.
RI_AXON = 100.d0
c RM_AXON = 125000.d0 ! NOTE same as for SD
RM_AXON = 2000.d0
CDENS = 0.8d0 ! Roth & Hausser
PI = 3.14159d0
c See T. Miyasho et al., Brain Research 2001, for initial densities, but
c there will be changes. e.g. Ca spikes have voltage-dependent fast-repol.
c so I will make dendritic gKDR larger and gKC smaller.
c ALSO, since there is now axon with voltage-shifted gNaF activation, I will
c try much smaller gNaF somatic density (Miyasho et al. have 10,000 ohm-cm-sqd)
c gNaF
gnaf_dens(0) = 3500.d0
gnaf_dens(1) = 5000.d0
c gnaf_dens(1) = 7000.d0
gnaf_dens(2) = 10.d0
gnaf_dens(3) = 0.d0
gnaf_dens(4) = 0.d0
c gNaP
gnap_dens(0) = 0.1d0
gnap_dens(1) = 5.0d0
gnap_dens(2) = 1.0d0
gnap_dens(3) = 0.0d0
gnap_dens(4) = 0.d0
c gCaP
gcaP_dens(0) = 0.0d0
gcaP_dens(1) = 0.0d0
gcaP_dens(2) = 0.0d0
gcaP_dens(3) = 5.0d0
gcaP_dens(4) = 5.0d0
c gCaT
gcaT_dens(0) = 0.0d0
gcaT_dens(1) = 0.0d0
gcaT_dens(2) = 0.5d0
gcaT_dens(3) = 1.5d0
gcaT_dens(4) = 1.5d0
c gCaR
gcaR_dens(0) = 0.0d0
gcaR_dens(1) = 0.0d0
gcaR_dens(2) = 0.0d0
gcaR_dens(3) = 8.0d0
gcaR_dens(4) = 8.0d0
c gar - Miyasho only have Ih at soma, but many princ cells have dendritic Ih
gar_dens(0) = 0.0d0
gar_dens(1) = 0.005d0
gar_dens(2) = 0.005d0
gar_dens(3) = 0.005d0
gar_dens(4) = 0.005d0
c gKDR
c gKDR_dens(0) = 3500.0d0
gKDR_dens(0) = 1000.0d0
c gKDR_dens(1) = 7000.0d0
gKDR_dens(1) = 1000.0d0
gKDR_dens(2) = 0.5d0
gKDR_dens(3) = 0.5d0
gKDR_dens(4) = 0.5d0
c gKM
gKM_dens(0) = 1.00d0
gKM_dens(1) = 1.00d0
gKM_dens(2) = 1.00d0
gKM_dens(3) = 0.04d0
gKM_dens(4) = 0.04d0
c gKA
gKA_dens(0) = 1.0d0
gKA_dens(1) = 15.0d0
gKA_dens(2) = 80.0d0
gKA_dens(3) = 80.0d0
gKA_dens(4) = 80.0d0
c gKD
gKD_dens(0) = 0.0d0
gKD_dens(1) = 0.0d0
gKD_dens(2) = 80.0d0
gKD_dens(3) = 80.0d0
c gKD_dens(3) = 200.0d0
gKD_dens(4) = 80.0d0
c gKD_dens(4) = 200.0d0
c gKC
gKC_dens(0) = 0.0d0
gKC_dens(1) = 0.0d0
gKC_dens(2) = 25.0d0
gKC_dens(3) = 25.0d0
gKC_dens(4) = 25.0d0
c gKAHP
gkahp_dens(0) = 0.d0
gkahp_dens(1) = 0.d0
gkahp_dens(2) = 0.d0
gkahp_dens(3) = 1.60d0
gkahp_dens(4) = 1.60d0
if (thisno.eq.0) then
WRITE (6,9988)
9988 FORMAT(2X,'I',4X,'NADENS',' CADENS(P)',' KDRDEN',' KAHPDE',
X ' KCDENS',' KADENS')
DO 9989, I = 0, 4
WRITE (6,9990) I, gnaf_dens(i), gcaR_dens(i), gkdr_dens(i),
X gkahp_dens(i), gkc_dens(i), gka_dens(i)
9990 FORMAT(2X,I2,2X,F7.2,1X,F6.2,1X,F7.2,1X,F6.2,1X,F6.2,1X,F6.2)
9989 CONTINUE
endif
level(1) = 1
level(2) = 2
level(3) = 2
do i = 4, 25
level(i) = 3
end do
do i = 26, 553
level(i) = 4
end do
do i = 554, 559
level(i) = 0
end do
c connectivity of axon
nnum(554) = 2
nnum(555) = 2
nnum(556) = 2
nnum(557) = 2
nnum(558) = 2
nnum(559) = 1
neigh(554,1) = 1
neigh(554,2) = 555
neigh(555,1) = 554
neigh(555,2) = 556
neigh(556,1) = 555
neigh(556,2) = 557
neigh(557,1) = 556
neigh(557,2) = 558
neigh(558,1) = 557
neigh(558,2) = 559
neigh(559,1) = 558
c connectivity of SD part
nnum(1) = 2 ! SOMA
neigh(1,1) = 554
neigh(1,2) = 2
c Now proximal smooth dendrite: no spiny treelets attached
nnum(2) = 2
neigh(2,1) = 1
neigh(2,2) = 3
nnum(3) = 3
neigh(3,1) = 2
neigh(3,2) = 4
neigh(3,3) = 5
c Now distal smooth dendrite (level 3): 2 spiny treelets to each
nnum(4) = 6
neigh(4,1) = 3
neigh(4,2) = 5
neigh(4,3) = 12
neigh(4,4) = 18
neigh(4,5) = 26 ! base of spiny treelet
neigh(4,6) = 38 ! base of spiny treelet
nnum(5) = 5
neigh(5,1) = 3
neigh(5,2) = 4
neigh(5,3) = 6
neigh(5,4) = 50 ! base of spiny treelet
neigh(5,5) = 62 ! base of spiny treelet
nnum(6) = 4
neigh(6,1) = 5
neigh(6,2) = 7
neigh(6,3) = 74
neigh(6,4) = 86
nnum(7) = 4
neigh(7,1) = 6
neigh(7,2) = 8
neigh(7,3) = 98
neigh(7,4) = 110
nnum(8) = 4
neigh(8,1) = 7
neigh(8,2) = 9
neigh(8,3) = 122
neigh(8,4) = 134
nnum(9) = 4
neigh(9,1) = 8
neigh(9,2) = 10
neigh(9,3) = 146
neigh(9,4) = 158
nnum(10) = 4
neigh(10,1) = 9
neigh(10,2) = 11
neigh(10,3) = 170
neigh(10,4) = 182
nnum(11) = 3
neigh(11,1) = 10
neigh(11,2) = 194
neigh(11,3) = 206
nnum(12) = 5
neigh(12,1) = 4
neigh(12,2) = 18
neigh(12,3) = 13
neigh(12,4) = 218
neigh(12,5) = 230
nnum(13) = 4
neigh(13,1) = 12
neigh(13,2) = 14
neigh(13,3) = 242
neigh(13,4) = 254
nnum(14) = 4
neigh(14,1) = 13
neigh(14,2) = 15
neigh(14,3) = 266
neigh(14,4) = 278
nnum(15) = 4
neigh(15,1) = 14
neigh(15,2) = 16
neigh(15,3) = 290
neigh(15,4) = 302
nnum(16) = 4
neigh(16,1) = 15
neigh(16,2) = 17
neigh(16,3) = 314
neigh(16,4) = 326
nnum(17) = 3
neigh(17,1) = 16
neigh(17,2) = 338
neigh(17,3) = 350
nnum(18) = 5
neigh(18,1) = 4
neigh(18,2) = 12
neigh(18,3) = 19
neigh(18,4) = 362
neigh(18,5) = 374
nnum(19) = 4
neigh(19,1) = 18
neigh(19,2) = 20
neigh(19,3) = 386
neigh(19,4) = 398
nnum(20) = 4
neigh(20,1) = 19
neigh(20,2) = 21
neigh(20,3) = 410
neigh(20,4) = 422
nnum(21) = 4
neigh(21,1) = 20
neigh(21,2) = 22
neigh(21,3) = 434
neigh(21,4) = 446
nnum(22) = 4
neigh(22,1) = 21
neigh(22,2) = 23
neigh(22,3) = 458
neigh(22,4) = 470
nnum(23) = 4
neigh(23,1) = 22
neigh(23,2) = 24
neigh(23,3) = 482
neigh(23,4) = 494
nnum(24) = 4
neigh(24,1) = 23
neigh(24,2) = 25
neigh(24,3) = 506
neigh(24,4) = 518
nnum(25) = 3
neigh(25,1) = 24
neigh(25,2) = 530
neigh(25,3) = 542
c Attachments of bases of treelets: one side to a smooth compartment, the other to next more distal part of treelet
do i = 26, 530, 24
nnum(i) = 2
neigh(i,1) = (i - 26)/24 + 4
neigh(i,2) = i + 1
end do
do i = 38, 542, 24
nnum(i) = 2
neigh(i,1) = (i - 38)/24 + 4
neigh(i,2) = i + 1
end do
c Now connect up the rest of the treelets
do i = 27, 543, 12
nnum(i) = 2
neigh(i,1) = i-1
neigh(i,2) = i+1
end do
do i = 28, 544, 12
nnum(i) = 2
neigh(i,1) = i-1
neigh(i,2) = i+1
end do
do i = 29, 545, 12
nnum(i) = 3
neigh(i,1) = i-1
neigh(i,2) = i+1
neigh(i,3) = i+5