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Quasi-Stationary Approximation

The system (15) contains a singular small parameter, the ratio of the characteristic times of the intracellular conductivity tex2html_wrap_inline965 , tex2html_wrap_inline971 , and of the membrane excitability, tex2html_wrap_inline973 . Biophysical data [Weidmann, 1952] give the intracellular resistivity of the order of 250 Ohm tex2html_wrap_inline975 cm, which for the cell size of 10-80  tex2html_wrap_inline977 m gives tex2html_wrap_inline965 of the order of several tex2html_wrap_inline977 S, so tex2html_wrap_inline983 , which is much less than tex2html_wrap_inline985 . Numerical calculations of the simplified two-compartment model (15) would require time steps not greater than tex2html_wrap_inline971 , while if we can get rid of the small parameter, times steps of around tex2html_wrap_inline973 would be adequate.

We exclude the small parameter tex2html_wrap_inline965 by ``quasi-stationary'' arguments. Rewrite the first two equations of (15) as

   eqnarray169

where

displaymath993

and

displaymath995

Now, in (17) the term tex2html_wrap_inline997 dominates over tex2html_wrap_inline999 , and omitting the nonlinear term tex2html_wrap_inline999 , (17) becomes a linear equation. If the characteristic time of tex2html_wrap_inline1003 change is bigger than tex2html_wrap_inline971 , then this equation describes the fast approach of tex2html_wrap_inline1007 to its quasi-stationary value,

displaymath1009

and we can substitute this value into (16), which yields

  equation192

Equations (18) and the last pair of (15) form a closed system, which depends only on the ratio tex2html_wrap_inline1011 . If we assume that the duration of pulses tex2html_wrap_inline1003 is shorter than the characteristic time tex2html_wrap_inline1015 of the slow variables then tex2html_wrap_inline1017 . This final simplification gives:

  eqnarray205

This model is almost as simple as the original (1) ordinary differential equation (e.g., it has three equations more than the 17 variable Noble et al. [1990] ordinary differential system we use for ventricular excitation), but describes the effect of external current. This has been obtained from (15) assuming the characteristic time of the external curent pulses, tex2html_wrap_inline1019 , is

displaymath1021

In practice tex2html_wrap_inline1019 , tex2html_wrap_inline973 and tex2html_wrap_inline1027 are all of the order of 1 ms.

The general model (14) can be simplified by quasi-stationary arguments to

  equation224

with separate ordinary differential equations for v at each point of the membrane. If the external field tex2html_wrap_inline905 is fixed in direction and varies only in magnitude, then the surface integral in (20) can be reduced, in a Lebesgue style, to an ordinary integral

  equation233

where the kernel K(s) is determined by the cell geometry, the conductivities, and the direction of the external field, and because of electroneutrality of the cell

displaymath1035

The simple model (19) corresponds to evaluation of the integral in (21) at two points tex2html_wrap_inline1037 .

Below we explore numerically some properties of the model (19) and its generalisation to spatially extended media. To validate the two-point approximation of the integral (21), we compare the two-point results with those of a five-point evaluation of (21), in the form

  eqnarray242


next up previous
Next: Action onto a Up: A model for the Previous: Simplified Two-Compartment Model

Vadim Biktashev
Fri Mar 28 21:26:28 GMT 1997