COMMENT
This mechanism is intended to be used in conjunction
with the extracellular mechanism. Pointers specified
at the hoc level must be used to connect the
extracellular mechanism's e_extracellular and i_membrane
to this mechanism's ex and im, respectively.
xtra does three useful things:
1. Serves as a target for Vector.play() to facilitate
extracellular stimulation. Assumes that one has initialized
a Vector to hold the time sequence of the stimulus current.
This Vector is to be played into the GLOBAL variable is
(GLOBAL so only one Vector.play() needs to be executed),
which is multiplied by the RANGE variable rx ("transfer
resistance between the stimulus electrode and the local
node"). This product, called ex in this mechanism, is the
extracellular potential at the local node, i.e. is used to
drive local e_extracellular.
2. Reports the contribution of local i_membrane to the
total signal that would be picked up by an extracellular
recording electrode. This is computed as the product of rx,
i_membrane (called im in this mechanism), and the surface area
of the local segment, and is reported as er. The total
extracellularly recorded potential is the sum of all er_xtra
over all segments in all sections, and is to be computed at
the hoc level, e.g. with code like
func fieldrec() { local sum
sum = 0
forall {
if (ismembrane("xtra")) {
for (x) sum += er_xtra(x)
}
}
return sum
}
Bipolar recording, i.e. recording the difference in potential
between two extracellular electrodes, can be achieved with no
change to either this NMODL code or fieldrec(); the values of
rx will reflect the difference between the potentials at the
recording electrodes caused by the local membrane current, so
some rx will be negative and others positive. The same rx
can be used for bipolar stimulation.
Multiple monopolar or bipolar extracellular recording and
stimulation can be accommodated by changing this mod file to
include additional rx, er, and is, and changing fieldrec()
to a proc.
3. Allows local storage of xyz coordinates interpolated from
the pt3d data. These coordinates are used by hoc code that
computes the transfer resistance that couples the membrane
to extracellular stimulating and recording electrodes.
Prior to NEURON 5.5, the SOLVE statement in the BREAKPOINT block
used METHOD cvode_t so that the adaptive integrators wouldn't miss
the stimulus. Otherwise, the BREAKPOINT block would have been called
_after_ the integration step, rather than from within cvodes/ida,
causing this mechanism to fail to deliver a stimulus current
when the adaptive integrator is used.
With NEURON 5.5 and later, this mechanism abandons the BREAKPOINT
block and uses the two new blocks BEFORE BREAKPOINT and
AFTER BREAKPOINT, like this--
BEFORE BREAKPOINT { : before each cy' = f(y,t) setup
ex = is*rx*(1e6)
}
AFTER SOLVE { : after each solution step
er = (10)*rx*im*area
}
This ensures that the stimulus potential is computed prior to the
solution step, and that the recorded potential is computed after.
ENDCOMMENT
NEURON {
SUFFIX xtra
RANGE rx, er, coeff, recx
RANGE x, y, z
GLOBAL is
POINTER im, ex
}
PARAMETER {
: default transfer resistance between stim electrodes and axon
coeff=1
recx=1 :ext rex given eld pos
rx = 1 (megohm) : mV/nA
x = 0 (1) : spatial coords
y = 0 (1)
z = 0 (1)
}
ASSIGNED {
v (millivolts)
is (milliamp)
ex (millivolts)
im (milliamp/cm2)
er (microvolts)
area (micron2)
}
INITIAL {
ex = is*rx*coeff*(1e6)
er = (10)*recx*im*area
: this demonstrates that area is known
:UNITSOFF
:printf("area = %f\n", area)
:UNITSON
}
: Use BREAKPOINT for NEURON 5.4 and earlier
: BREAKPOINT {
: SOLVE f METHOD cvode_t
: }
: With NEURON 5.5 and later, abandon the BREAKPOINT block
: and instead use BEFORE BREAKPOINT and AFTER BREAKPOINT
BREAKPOINT { : before each cy' = f(y,t) setup
: ex = is*rx*(1e6)
:er = (10)*rx*im*area
:print "ex", ex
:print "er", er
}
PROCEDURE f() {
: 1 mA * 1 megohm is 1000 volts
: but ex is in mV
ex = is*rx*(1e6)
er = (10)*recx*im*area
}
BEFORE BREAKPOINT { : before each cy' = f(y,t) setup
ex = is*rx*coeff*(1e6)
}
AFTER SOLVE { : after each solution step
er = (10)*recx*im*area
}