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Dynamical Systems in Neuroscience:

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One-Dimensional <strong>Systems</strong> 79F(V)10 mV0.5 msExcited0Attractorru<strong>in</strong>smembrane potential (mV)ExcitedV-40 mVslow transitionAttractorru<strong>in</strong>sFigure 3.28: Slow transition through the ghost of the rest<strong>in</strong>g state attractor <strong>in</strong> a corticalpyramidal neuron with I = 30 pA (the same neuron as <strong>in</strong> Fig. 3.15). Even thoughthe rest<strong>in</strong>g state has already disappeared, the function F (V ), and hence the rate ofchange, ˙V , is still small when V ≈ −46 mV.3.3.5 Slow transitionAll physical, chemical, and biological systems near saddle-node bifurcations possesscerta<strong>in</strong> universal features that do not depend on particulars of the systems. Consequently,all neural systems near such a bifurcation share common neuro-computationalproperties, which we will discuss <strong>in</strong> detail <strong>in</strong> Chapter 7. Here we take a look at onesuch property — slow transition through the ru<strong>in</strong>s (or ghost) of the rest state attractor,which is relevant to the dynamics of many neocortical neurons.In Fig. 3.28 we show the function F (V ) of the system (3.5) with I = 30 pA,which is greater than the bifurcation value 16 pA, and the correspond<strong>in</strong>g behaviorof a cortical neuron; compare with Fig. 3.15. The system has only one attractor— the excited state, and any solution start<strong>in</strong>g from an arbitrary <strong>in</strong>itial conditionshould quickly approach this attractor. However, the solutions start<strong>in</strong>g from the <strong>in</strong>itialconditions around −50 mV do not seem to hurry. Instead, they slow down near −46mV and spend a considerable amount of time <strong>in</strong> the voltage range correspond<strong>in</strong>g to therest<strong>in</strong>g state, as if the state were still present. The closer is I to the bifurcation value,the more time the membrane potential spends <strong>in</strong> the neighborhood of the rest<strong>in</strong>g state.Obviously, such a slow transition cannot be expla<strong>in</strong>ed by a slow activation of the <strong>in</strong>wardNa + current, s<strong>in</strong>ce Na + activation <strong>in</strong> a cortical neuron is practically <strong>in</strong>stantaneous.The slow transition occurs because the neuron or the system (3.5) <strong>in</strong> Fig. 3.28 isnear a saddle-node bifurcation. Even though I is greater than the bifurcation value,and the rest state attractor is already annihilated, the function F (V ) is barely abovethe V -axis at the “annihilation site”. In other words, the rest state attractor hasalready been ru<strong>in</strong>ed, but its “ru<strong>in</strong>s” (or its “ghost”) can still be felt because˙V = F (V ) ≈ 0(at attractor ru<strong>in</strong>s, V ≈ −46 mV),as one can see <strong>in</strong> Fig. 3.28. In Chapter 7 we will show how this property expla<strong>in</strong>s the

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