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Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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296 Simple Modelslayer 5 neuronsimple modelI=100 pAI=85 pAI=70 pA<strong>in</strong>put+35 mV200 ms-60 mV<strong>in</strong>putresetAHPrecovery, u100500-50restresetAHPI>0I=0I=60 pA-60 -40 -20 0 20membrane potential, v (mV)spike35Figure 8.12: Comparison of <strong>in</strong> vitro record<strong>in</strong>gs of a regular spik<strong>in</strong>g (RS) pyramidalneurons with simulations of the simple model 100 ˙v = 0.7(v + 60)(v + 40) − u + I,˙u = 0.03{−2(v + 60) − u}, if v ≥ +35, then v ← −50, u ← u + 100.responses to positive pulses, sags and rebound spikes to negative pulses (as <strong>in</strong> Fig. 7.48),relatively short latencies of the first spike, and other resonance phenomena. The evenmore extreme example <strong>in</strong> Fig. 7.42 shows a pyramidal neuron execut<strong>in</strong>g a subthresholdoscillation before switch<strong>in</strong>g to a tonic spik<strong>in</strong>g mode.The difference between the types <strong>in</strong> Fig. 8.13 can be expla<strong>in</strong>ed by the sign of theparameter b <strong>in</strong> the simple model (8.5, 8.6), which depends on the relative contributionsof amplify<strong>in</strong>g and resonant slow currents and gat<strong>in</strong>g variables. When b < 0 (or b ≈ 0,e.g., b = 0.5 <strong>in</strong> Fig. 8.14), the neuron is a pure <strong>in</strong>tegrator near saddle-node on <strong>in</strong>variantcircle bifurcation. Greater values of b > 0 put the model near the transition from an<strong>in</strong>tegrator to a resonator the via co-dimension-2 Bogdanov-Takens bifurcation studied<strong>in</strong> Sect. 6.3.3 and 7.2.11.The sequence of bifurcations when b > 0 is depicted <strong>in</strong> Fig. 8.15. Injection ofdepolariz<strong>in</strong>g current below the neuron’s rheobase transforms the rest<strong>in</strong>g state <strong>in</strong>toa stable focus and results <strong>in</strong> damped oscillations of the membrane potential. The

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