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Next: 3 Results Up: Re-entrant activity and its Previous: 1 Introduction

2 The numerical model

The Oxsoft equations (Noble 1990) for a single isopotential cell are in the form of a system of ordinary differential equations:

  equation21

where C is the capacitance of a single cell, V the membrane potential in mV and tex2html_wrap_inline1445 is a vector of the activation and inactivation gating variables and the ionic concentrations that determine the total membrane current tex2html_wrap_inline1447 . The full equations, together with the parameter values, are listed in the Appendix, and can also be obtained in the form of a Pascal program in Oxsoft HEART. This model was incorporated into a partial differential equation model for an excitable medium in the plane (x,y),

  equation37

where D is the diffusion coefficient for V, tex2html_wrap_inline1455 is the Laplacian operator tex2html_wrap_inline1457 and F(t) is a spatially uniform, time dependent forcing that models external electric current applied to the tissue. The diffusion coefficient tex2html_wrap_inline1461 was chosen to give a conduction velocity of tex2html_wrap_inline1463 that corresponds to intercellular (gap junction) conductance of tex2html_wrap_inline1465 for tex2html_wrap_inline1467 long cells (Rice et al. 1995).

Calculations were performed using the explicit Euler method with five-node approximation of the Laplacian on a rectangular grid of 200 tex2html_wrap_inline1469 200 to 300 tex2html_wrap_inline1469 300 nodes with a time and space step of tex2html_wrap_inline1473 and tex2html_wrap_inline1475 (for the majority of experiments) or 0.001 ms and 0.05 mm (i.e. less than the myocyte length) for a few control computations. Steps used in other published ventricular simulations were: 0.025 ms and 0.25 mm in Courtemanche & Winfree (1991) and 0.1 ms and 0.25 mm in Efimov et al. (1995). The boundaries were modelled as impermeable,

  equation55

with the medium large enough so that the exact form of boundary conditions does not influence the spiral wave behaviour.

At the space step of 0.1 mm, the upstroke velocity of the solitary wave was 570 V/s, and the CV 0.38 m/s; and at space step of 0.05 mm, 500 V/s and 0.45 m/s correspondingly. As the space step is reduced, the CV with tex2html_wrap_inline1461 converges to 0.485 m/s; the CV is within 5% of this `true' value at tex2html_wrap_inline1479 i.e. less than the length of a single cell.

The action of external forcing as the additive term F(t) in (2) corresponds to injecting current into each cell and does not correspond to extracellular field stimulation. However, realistic models of external current require consideration of each cell as an extended object described by partial differential equations, or at least integration of transmembrane currents over each cell's surface [14] and so massively increase the computational load. This paper is mostly concerned with the spontaneous evolution of spiral waves, and the qualitative features of resonant drift and so this simplification is acceptable. We measured the external forcing F in V/s in terms of its dimensionality in (2); this can easily be rescaled to amperes per cell by dividing by cell capacitance C. The time dependent F(t) used to produce resonant drift of the spiral wave was a series of rectangular pulses of 2 ms duration, applied at a fixed time after an action potential was detected at the recording site by V increasing through -10 mV.

Spiral waves were initiated in two ways, by a cut wavefront or twin pulse protocol. A plane wave was initiated at one edge of the medium by a 2 ms duration stimulation of a strip 0.65 mm wide, by a current that gave a tex2html_wrap_inline1491 of 50 V/s and the excitation allowed to propagate to the centre of the medium. The wavefront was then cut, and all the variables on one side of the cut reset to their equilibrium values. This numerically convenient but artificial method allows spirals to be initiated in a 2 tex2html_wrap_inline1469 2 cm medium. The twin pulse protocol requires a larger (3 tex2html_wrap_inline1469 3 cm) medium, in which a plane wave is initiated at the lower border by 10 ms stimulation of 50 V/s of a one mm strip, and 180 ms later (after the wavefront has propagated through the medium, establishing a gradient in refractoriness) the second stimulus is applied: a 4 ms stimulation of 40 V/s over the left 3 tex2html_wrap_inline1469 2.5 cm area of the medium. These large areas are necessary for the initiation of reentry in a homogeneous tissue; re-entry persists in smaller media, see figure 1. For anisotropic media we took (2) with the Laplacian replaced by tex2html_wrap_inline1499 or tex2html_wrap_inline1501 with tex2html_wrap_inline1503 , which is equivalent to 3-fold compression in x or y direction. Simulating 1 s of activity in a 2 tex2html_wrap_inline1469 2 cm medium with tex2html_wrap_inline1511 and tex2html_wrap_inline1513 took about 180 hours on a single MIPS R8000 75 MHz central processor unit.


next up previous
Next: 3 Results Up: Re-entrant activity and its Previous: 1 Introduction

Vadim Biktashev
Mon Mar 31 15:56:29 GMT 1997