COMMENT
ca.mod
Uses fixed eca instead of GHK eqn
HVA Ca current
Based on Reuveni, Friedman, Amitai and Gutnick (1993)
J. Neurosci. 13:4609-4621.
Author: Zach Mainen, Salk Institute, 1994, zach@salk.edu
26 Ago 2002 Modification of original channel to allow
variable time step and to correct an initialization error.
Done by Michael Hines(michael.hines@yale.e) and
Ruggero Scorcioni(rscorcio@gmu.edu) at EU Advance Course
in Computational Neuroscience. Obidos, Portugal
20110202 made threadsafe by Ted Carnevale
20120514 fixed singularity in PROCEDURE rates
Special comment:
This mechanism was designed to be run at a single operating
temperature--37 deg C--which can be specified by the hoc
assignment statement
celsius = 37
This mechanism is not intended to be used at other temperatures,
or to investigate the effects of temperature changes.
Zach Mainen created this particular model by adapting conductances
from lower temperature to run at higher temperature, and found it
necessary to reduce the temperature sensitivity of spike amplitude
and time course. He accomplished this by increasing the net ionic
conductance through the heuristic of changing the standard HH
formula
g = gbar*product_of_gating_variables
to
g = tadj*gbar*product_of_gating_variables
where
tadj = q10^((celsius - temp)/10)
temp is the "reference temperature" (at which the gating variable
time constants were originally determined)
celsius is the "operating temperature"
Users should note that this is equivalent to changing the channel
density from gbar at the "reference temperature" temp (the
temperature at which the at which the gating variable time
constants were originally determined) to tadj*gbar at the
"operating temperature" celsius.
ENDCOMMENT
NEURON {
THREADSAFE
SUFFIX ca
USEION ca READ eca WRITE ica
RANGE m, h, gca, gbar
RANGE minf, hinf, mtau, htau
GLOBAL q10, temp, tadj, vmin, vmax, vshift
}
UNITS {
(mA) = (milliamp)
(mV) = (millivolt)
(pS) = (picosiemens)
(um) = (micron)
(mM) = (milli/liter)
FARADAY = (faraday) (coulomb)
R = (k-mole) (joule/degC)
PI = (pi) (1)
}
PARAMETER {
gbar = 0.1 (pS/um2) : 0.12 mho/cm2
vshift = 0 (mV) : voltage shift (affects all)
cao = 2.5 (mM) : external ca concentration
cai (mM)
temp = 23 (degC) : original temp
q10 = 2.3 : temperature sensitivity
vmin = -120 (mV)
vmax = 100 (mV)
}
ASSIGNED {
v (mV)
celsius (degC)
ica (mA/cm2)
gca (pS/um2)
eca (mV)
minf hinf
mtau (ms) htau (ms)
tadj
}
STATE { m h }
INITIAL {
tadj = q10^((celsius - temp)/(10 (degC))) : make all threads calculate tadj at initialization
trates(v+vshift)
m = minf
h = hinf
}
BREAKPOINT {
SOLVE states METHOD cnexp
gca = tadj*gbar*m*m*h
ica = (1e-4) * gca * (v - eca)
}
: LOCAL mexp, hexp
:PROCEDURE states() {
: trates(v+vshift)
: m = m + mexp*(minf-m)
: h = h + hexp*(hinf-h)
: VERBATIM
: return 0;
: ENDVERBATIM
:}
DERIVATIVE states {
trates(v+vshift)
m' = (minf-m)/mtau
h' = (hinf-h)/htau
}
PROCEDURE trates(v (mV)) {
TABLE minf, hinf, mtau, htau
DEPEND celsius, temp
FROM vmin TO vmax WITH 199
rates(v): not consistently executed from here if usetable == 1
: tinc = -dt * tadj
: mexp = 1 - exp(tinc/mtau)
: hexp = 1 - exp(tinc/htau)
}
UNITSOFF
PROCEDURE rates(vm (mV)) {
LOCAL a, b
tadj = q10^((celsius - temp)/(10 (degC)))
: a = 0.055*(-27 - vm)/(exp((-27-vm)/3.8) - 1)
: singular at vm = -27
: a = 0.055*3.8*(-27 - vm)/3.8)/(exp((-27-vm)/3.8) - 1)
: let z = (-27 - vm)/3.8
: a = 0.055*3.8*z/(exp(z) - 1)
a = 0.209*efun(-(27+vm)/3.8)
b = 0.94*exp((-75-vm)/17)
mtau = 1/tadj/(a+b)
minf = a/(a+b)
:"h" inactivation
a = 0.000457*exp((-13-vm)/50)
b = 0.0065/(exp((-vm-15)/28) + 1)
htau = 1/tadj/(a+b)
hinf = a/(a+b)
}
UNITSON
FUNCTION efun(z) {
if (fabs(z) < 1e-4) {
efun = 1 - z/2
}else{
efun = z/(exp(z) - 1)
}
}