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Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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Synchronization (see www.izhikevich.com) 469(a)synchronization<strong>in</strong>-phase anti-phase out-of-phase(b)phase-lock<strong>in</strong>g1:2 2:13:2Figure 10.12: Examples of fundamental types of synchronization of spik<strong>in</strong>g activity toperiodic pulsed <strong>in</strong>puts (synchronization is 1:1 phase-lock<strong>in</strong>g).T-T sunstablestabilityregionPRC00stablephaseshiftstimulus phase,stabilityregionTFigure 10.13: Fixed po<strong>in</strong>ts of the Po<strong>in</strong>carephase map (10.1.5).marked by the bold curves <strong>in</strong> Fig. 10.13.Now consider the Class 1 and Class 2 I Na + I K -oscillators shown <strong>in</strong> Fig. 10.6. ThePRC <strong>in</strong> Class 1 regime is mostly positive, imply<strong>in</strong>g that such an oscillator can easilysynchronize with faster <strong>in</strong>puts (T − T s > 0) but cannot synchronize with slower <strong>in</strong>puts.Indeed, the oscillator can only advance its phase to catch up faster pulse tra<strong>in</strong>s, but itcannot delay the phase to wait for the slower <strong>in</strong>put. Synchronization with the <strong>in</strong>puthav<strong>in</strong>g T s ≈ T is only marg<strong>in</strong>al. In contrast, Class 2 I Na + I K -oscillator does not havethis problem because its PRC has well-def<strong>in</strong>ed positive and negative regions.10.1.8 Phase lock<strong>in</strong>gp:q-phase-lock<strong>in</strong>g occurs when the oscillator fires p spikes for every q <strong>in</strong>put pulses,such as the 3:2-phase-lock<strong>in</strong>g <strong>in</strong> Fig. 10.12b or 2:2 phase-lock<strong>in</strong>g <strong>in</strong> Fig. 10.14, whichtypically occurs when pT ≈ qT s . The <strong>in</strong>tegers p and q need not be relatively prime <strong>in</strong>the case of pulsed-coupled oscillators. Synchronization, i.e., 1:1 phase-lock<strong>in</strong>g, as wellas p:1 phase-lock<strong>in</strong>g corresponds to a fixed po<strong>in</strong>t of the Po<strong>in</strong>care phase map (10.3) withp fired spikes per one <strong>in</strong>put pulse. Indeed, the map tells only the phase of the oscillatorat each pulse, but does not tell the number of oscillations made between the pulses.Each p:q-locked solution corresponds to a stable periodic orbit of the Po<strong>in</strong>carephase map with the period q (so that ϑ n = ϑ n+q for any n). Such orbits <strong>in</strong> maps (10.4)

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