COMMENT

na.mod

Sodium channel, Hodgkin-Huxley style kinetics.  

Kinetics were fit to data from Huguenard et al. (1988) and Hamill et
al. (1991)

qi is not well constrained by the data, since there are no points
between -80 and -55.  So this was fixed at 5 while the thi1,thi2,Rg,Rd
were optimized using a simplex least square proc

voltage dependencies are shifted approximately from the best
fit to give higher threshold

Author: Zach Mainen, Salk Institute, 1994, zach@salk.edu

ENDCOMMENT

INDEPENDENT {t FROM 0 TO 1 WITH 1 (ms)}

NEURON {
	SUFFIX na
	USEION na READ ena WRITE ina
	RANGE m, h, gna, gbar
	GLOBAL tha, thi1, thi2, qa, qi, qinf, thinf
	RANGE minf, hinf, mtau, htau
	GLOBAL Ra, Rb, Rd, Rg
	GLOBAL q10, temp, tadj, vmin, vmax, vshift
}

PARAMETER {
	gbar = 1000   	(pS/um2)	: 0.12 mho/cm2
	vshift = -10	(mV)		: voltage shift (affects all)
								
	tha  = -35	(mV)		: v 1/2 for act		(-42)
	qa   = 9	(mV)		: act slope		
	Ra   = 0.182	(/mV/ms)	: open (v)		
	Rb   = 0.124	(/mV/ms)	: close (v)		

	thi1  = -50	(mV)		: v 1/2 for inact 	
	thi2  = -75	(mV)		: v 1/2 for inact 	
	qi   = 5	(mV)	        : inact tau slope
	thinf  = -65	(mV)		: inact inf slope	
	qinf  = 6.2	(mV)		: inact inf slope
	Rg   = 0.0091	(/mV/ms)	: inact (v)	
	Rd   = 0.024	(/mV/ms)	: inact recov (v) 

	temp = 23	(degC)		: original temp 
	q10  = 2.3			: temperature sensitivity

	v 		(mV)
	dt		(ms)
	celsius		(degC)
	vmin = -120	(mV)
	vmax = 100	(mV)
}


UNITS {
	(mA) = (milliamp)
	(mV) = (millivolt)
	(pS) = (picosiemens)
	(um) = (micron)
} 

ASSIGNED {
	ina 		(mA/cm2)
	gna		(pS/um2)
	ena		(mV)
	minf 		hinf
	mtau (ms)	htau (ms)
	tadj
}
 

STATE { m h }

INITIAL { 
	trates(v+vshift)
	m = minf
	h = hinf
}

BREAKPOINT {
        SOLVE states
        gna = tadj*gbar*m*m*m*h
	ina = (1e-4) * gna * (v - ena)
} 

LOCAL mexp, hexp 

PROCEDURE states() {   :Computes state variables m, h, and n 
        trates(v+vshift)      :             at the current v and dt.
        m = m + mexp*(minf-m)
        h = h + hexp*(hinf-h)
        VERBATIM
        return 0;
        ENDVERBATIM
}

PROCEDURE trates(v (mV)) {  
                      
        LOCAL tinc
        TABLE minf, mexp, hinf, hexp
	DEPEND dt, celsius, temp, Ra, Rb, Rd, Rg, tha, thi1, thi2, qa, qi, qinf
	
	FROM vmin TO vmax WITH 199

	rates(v): not consistently executed from here if usetable == 1

        tadj = q10^((celsius - temp)/10(degC))
        tinc = -dt * tadj

        mexp = 1 - exp(tinc/mtau)
        hexp = 1 - exp(tinc/htau)
}


PROCEDURE rates(vm (mV)) {  
        LOCAL  a, b

	a = trap0(vm,tha,Ra,qa)
	b = trap0(-vm,-tha,Rb,qa)
	mtau = 1/(a+b)
	minf = a*mtau

		:"h" inactivation 

	a = trap0(vm,thi1,Rd,qi)
	b = trap0(-vm,-thi2,Rg,qi)
	htau = 1/(a+b)
	hinf = 1/(1+exp((vm-thinf)/qinf))
}


FUNCTION trap0(v (mV), th (mV), a (/mV/ms), q (mV)) (/ms) {
	if (fabs(v/th) > 1e-6) {
	        trap0 = a * (v - th) / (1 - exp(-(v - th)/q))
	} else {
	        trap0 = a * q
 	}
}