: xtrau.mod, based on xtra.mod,v 1.3 2009/02/24 00:52:07 : the "u" stands for "uniform" as in "uniform field" : because this modification was made to accommodate a : uniform extracellular field, : as would occur between two parallel plates 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. xtrau does four 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 extracellular field strength E. This Vector is to be played into the GLOBAL variable E (GLOBAL so only one Vector.play() needs to be executed), which is multiplied by the RANGE variable d ("distance between the local node and the zero potential plane") and then by -1. That 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 (transfer resistance between the local node and the monopolar recording electrode), 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_xtrau 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("xtrau")) { for (x,0) sum += er_xtrau(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. 3. Enables 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. 4. Enables local storage of the distance d of internal nodes from the zero potential plane. 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 xtrau RANGE rx, er, d RANGE x, y, z GLOBAL E POINTER im, ex } PARAMETER { : default transfer resistance between recording electrode and node rx = 1 (megohm) : mV/nA x = 0 (1) : spatial coords y = 0 (1) z = 0 (1) } ASSIGNED { v (millivolts) : is (milliamp) E (volt/m) : field strength d (micron) : distance of node from zero potential plane ex (millivolts) im (milliamp/cm2) er (microvolts) area (micron2) } INITIAL { : ex = is*rx*(1e6) ex = -E*d*(1e-3) er = (10)*rx*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 : } : : PROCEDURE f() { : : 1 mA * 1 megohm is 1000 volts : : but ex is in mV : ex = E*d*(1e-3) : er = (10)*rx*im*area : } : With NEURON 5.5 and later, abandon the BREAKPOINT block and PROCEDURE f(), : and instead use BEFORE BREAKPOINT and AFTER BREAKPOINT BEFORE BREAKPOINT { : before each cy' = f(y,t) setup : ex = is*rx*(1e6) ex = -E*d*(1e-3) } AFTER SOLVE { : after each solution step er = (10)*rx*im*area }