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Next: Acknowledgments Up: Excitation wave breaking Previous: Periodic wave conduction block

Numerical illustrations

So far we have considered only plane waves. To verify the estimates and analyse the effect of the shear flow on more complicated autowave patterns, we have performed numerical simulations for the FitzHugh-Nagumo system with the flow incorporated, in the following form

  eqnarray229

We shall refer to the space and time units of this equation as s.u. and t.u. respectively. The parameter values were chosen tex2html_wrap_inline904 , a=0.02, tex2html_wrap_inline908 , tex2html_wrap_inline910 , tex2html_wrap_inline912 and D=1. We solved this system with explicit Euler scheme (forward time, centered space), with space step tex2html_wrap_inline916 , in a rectangular medium tex2html_wrap_inline918 with periodic boundary conditions at x=0,L and non-flux boundary conditions at tex2html_wrap_inline922 . The sizes of the medium, L and M, the velocity gradient, tex2html_wrap_inline766 , and the time step, tex2html_wrap_inline930 , were varied in different experiments. The minimum wavelength of a periodic train in quiescent ( tex2html_wrap_inline770 ) medium is 19.0, and the wavelength of the spiral wave is 41.0.

   figure244
Figure 2: Conduction block of a solitary plane wave in the linear shear flow. Velocity gradient tex2html_wrap_inline938 , medium size tex2html_wrap_inline940 , time step tex2html_wrap_inline942 Shown are snapshots of the E field (darker shade corresponds to higher value) with interval tex2html_wrap_inline946 . Panel (d) corresponds to the moment just before the disappearance of the wave: no excitation (black) and only the recovery tail (lighter shade) left in the medium; some excitation only survived near the boundaries.

The evolution of a solitary plane wave is shown in Fig. 2. The horizontally propagating wave was initiated in the quiescent medium and then the flow was switched on, which corresponds to the conditions of the analytical estimation. The activator and inhibitor are E and g and their characteristic times are, respectively, tex2html_wrap_inline952 and tex2html_wrap_inline954 According to (14), this means that tex2html_wrap_inline956 for our choice of parameters. The threshold value of the velocity gradient necessary for breaking a single plane wave was found numerically to be tex2html_wrap_inline958 , the time tex2html_wrap_inline960 and the wave orientation at the moment of the break tex2html_wrap_inline962 , which, to this order of magnitude, is consistent with the analytical estimates of (14), tex2html_wrap_inline964 , tex2html_wrap_inline966 , and tex2html_wrap_inline968 .

   figure297
Figure 3: Breakup of a spiral wave. Velocity gradient tex2html_wrap_inline970 , medium size tex2html_wrap_inline972 , time step tex2html_wrap_inline974 Shown are snapshots of the E field with interval tex2html_wrap_inline978

We have studied what happens to autowave structures at velocity gradients much less than this threshold. According to (15) and (16), we have for this medium tex2html_wrap_inline980 , and at tex2html_wrap_inline970 the wavebreak of the periodic train occurs at tex2html_wrap_inline984 at the angle of tex2html_wrap_inline986 . These predictions, obtained for plane waves, agree, in order of magnitude, with simulations of the evolution of more complicated autowave patterns, the spiral wave and the target pattern.

In the first example, see Fig. 3, we initiated a spiral wave in a quiescent medium ( tex2html_wrap_inline770 ) and then switched on the shear flow. The spiral wave breaks, and the time ( tex2html_wrap_inline990 ) and orientation of the wave ( tex2html_wrap_inline992 ) are in good correspondence with the theory. The correspondence in this case would be better but for the phenomenon of ``plasticity'' of the excitation wave: as the process is non-stationary, the visual break-up happens later than the time at which conditions of stationary propagation are violated. This explanation was confirmed by numerical experiments when the flow was stopped before the wave broke, but after it should have happened accordingly to the analytical estimate; the wave subsequently broke, despite the absence of the flow.

   figure343
Figure 4: Breakup of a target pattern. Velocity gradient tex2html_wrap_inline970 , medium size tex2html_wrap_inline972 , time step tex2html_wrap_inline974 Shown are snapshots of the E field with interval tex2html_wrap_inline978

In the other example (see Fig. 4), we initiated a series of topologically circular waves by periodical stimulation of a point in the medium, with the period equal to the period of the spiral wave. It can be seen that while the first wave propagates without problems, the propagation of the second is suppressed and the third wave is blocked. This block occurs not for the whole wave but at some points only, leading to wavebreaks which curl up into spirals. The first breaks occur to the third wave, at time tex2html_wrap_inline1004 , that is tex2html_wrap_inline1006 after its initiation, and at a propagation angle of about tex2html_wrap_inline1008 , i.e. the same as in the previous case.

These two examples show that, at least for the particular model chosen, the conditions of the wavebreak are almost the same whatever the origin and shape of the pattern is, and the order of magnitude of the velocity gradient necessary for the wavebreak agrees with the estimate (15) obtained for plane periodic waves based on the quasi-stationary arguments.

As we see in Fig. 3, a spiral wave, at time tex2html_wrap_inline878 succumbs to wavebreaks which then develop into new spiral waves. Thus, in a linear shear flow a ``chain reaction'' of spiral wave births and deaths leads to a ``frazzle gas'' of excitation wavelets. The mechanism for generation of this ``frazzle gas'' is different from that described in [9]. As this mechanism requires only a finite deformation of the medium, we may expect that it can play its role not only in constant flows, but in a more wide variety of situations. If oceanic plankton dynamics are considered as an excitable medium [10] then currents can influence their spatial dynamics.

In a quite different physical context, cardiac muscle is excitable and is moving [11]. Condition (14) assumes the finite deformation of the medium to happen on the time scale of the fast excitation processes, about 1 ms in cardiac muscle, which does not seem realistic. Therefore, any normal motion of the heart is highly unlikely to break the excitation wave in it. On the other hand, the condition for breaking of repetitive waves (15,16) only requires finite deformation of the medium during any time period. The critical deformation is the ratio of the spiral wave period and the refractory period, which is known to be quite close to unity in cardiac tissue. Thus, while unlikely to cause the first breaking, the medium motion may play its role (along with other suggested mechanisms, such as heterogeneity and anisotropy of tissue properties, instabilities of plane wave propagation and scroll wave shape etc.) in the development of fibrillation once the reentrant activity has started and before the muscle has lost contractility. Naturally, the motion of the cardiac tissue is far from being constant linear shear flow, and therefore the results can only be considered as hints to what could be expected in more realistic models of reentrant activity in the heart involving mechano-electric feedback.

An ODE system of analogous form to the one considered in this letter, can also describe the action of an electric field on a chemical excitable medium, where the role of the advection velocity is played by the electric field multiplied by the mobility of the reagent. If all the reagents have the same mobility, the corresponding PDE system is formally equivalent to that for reaction-diffusion with convective flow. Examples of the nontrivial influence of the electric field on chemical excitable media are abundant and can be ``soft'', like modification of propagation velocity of waves and drift of patterns [12], or ``hard'', such as direction-dependent propagation block and splitting of excitation waves [13]. There is an interesting relationship between effects of curvature and electric current on excitation wavefront propagation [14]. Though the case of exactly equal mobilities is rather exotic (at minimum, it requires that all essential reagents have charge of the same sign), and the constant linear shear flow cannot correspond to a constant electric field (as the latter should be potential unless in presence of a nonstationary magnetic field), this analogy still might prove to be a powerful agent for mutual fertilisation of these two different problems. While the nontrivial effect of the electric field is due to the species-dependent variation of the advection velocity, the present study illustrates the effect of space-dependent advection velocity. In the context of this analogy, one hypothesis which can be put forward is that the quenching effect of electric field upon repetitive excitation waves might significantly depend upon the period of these waves -- a feature which, as far as we are aware, has not yet been appreciated.


next up previous
Next: Acknowledgments Up: Excitation wave breaking Previous: Periodic wave conduction block

Vadim Biktashev
Wed Aug 12 18:01:29 BST 1998