COMMENT
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
kca.mod
Calcium-dependent potassium channel
Based on
Pennefather (1990) -- sympathetic ganglion cells
taken from
Reuveni et al (1993) -- neocortical cells
Author: Zach Mainen, Salk Institute, 1995, zach@salk.edu
20150512 NTC
Cleaned code up a bit.
Made threadsafe.
See na.mod in ModelDB entry 2488.
Special comment:
This mechanism borrowed heavily from na.mod in ModelDB entry 2488.
That code, which was intended to be used only at 37 deg C
(see comments from 20120514 in that file),
calculated ionic conductance as the product
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"
This deviates from the standard HH formula
g = gbar*product_of_gating_variables
and has the unfortunate consequence of not only making the
effective channel density differ from the nominal (i.e. user-assigned)
channel density, but it would also make the effective channel density
depend on temperature!
Sooner or later this is guaranteed to confound studies of the effects
of temperature on model operation.
It would also be a debugging nightmare, not least because
the ModelView tool--so handy for discovering the properties of a
model and verifying a close match between the computational model
and the conceptual model--would report the NOMINAL channel density,
not the effective channel density.
To eliminate this problem, every statement of the form
g = tadj*gbar*gating variables
has been replaced by
g = gbar*gating variables
Furthermore, the numerical value assigned to gbar,
whether by an assignment statement in the PARAMETER block
or by a hoc or Python statement,
must now use the actual conductance density.
For this particular mechanism, tadj at 37 deg C is 3.20936
so the actual conductance density is 3.20936 times
the nominal conductance density.
ENDCOMMENT
NEURON {
THREADSAFE
SUFFIX kca
USEION k READ ek WRITE ik
USEION ca READ cai
RANGE n, gk, gbar
RANGE ninf, ntau
GLOBAL Ra, Rb, caix
GLOBAL q10, temp, tadj, vmin, vmax
}
UNITS {
(mA) = (milliamp)
(mV) = (millivolt)
(pS) = (picosiemens)
(um) = (micron)
(mM) = (milli/liter)
}
PARAMETER {
gbar = 10 (pS/um2) : 0.001 mho/cm2
: v (mV)
: cai (mM)
caix = 1
Ra = 0.01 (/ms) : max act rate
Rb = 0.02 (/ms) : max deact rate
: dt (ms)
: celsius (degC)
temp = 23 (degC) : original temp
q10 = 2.3 : temperature sensitivity
vmin = -120 (mV)
vmax = 100 (mV)
}
ASSIGNED {
v (mV)
celsius (degC)
cai (mM)
a (/ms)
b (/ms)
ik (mA/cm2)
gk (pS/um2)
ek (mV)
ninf
ntau (ms)
tadj
}
STATE { n }
INITIAL {
: since tadj is a per-thread GLOBAL
: all threads must calculate its value at initialization
tadj = q10^((celsius - temp)/(10 (degC)))
rates(cai)
n = ninf
}
BREAKPOINT {
SOLVE states METHOD cnexp
: gk = tadj*gbar*n
gk = gbar*n
ik = (1e-4) * gk * (v - ek)
}
: LOCAL nexp
DERIVATIVE states { :Computes state variable n
rates(cai) : at the current v
n' = (ninf-n)/ntau
:n = n + nexp*(ninf-n)
}
UNITSOFF
PROCEDURE rates(cai(mM)) {
a = Ra * cai^caix
b = Rb
: tadj = q10^((celsius - temp)/10)
tadj = q10^((celsius - temp)/(10 (degC)))
ntau = 1/tadj/(a+b)
ninf = a/(a+b)
: ntau = 1/(a+b)
: ninf = a*ntau
: tinc = -dt * tadj
: nexp = 1 - exp(tinc/ntau)
}
UNITSON