12.07.2015 Views

Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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300 Simple Models(a)simple modellayer 5 neuron (<strong>in</strong> vitro)20 mV20 ms(b)32 nS8 nSg GABAg AMPAresponse<strong>in</strong>put(c)1 seclayer 5 neuron(<strong>in</strong> vitro)simple modelFigure 8.17: (a) Comparison of responses of a rat motor cortex layer 5 pyramidal neuronof RS type and the simple model (8.5, 8.6) to <strong>in</strong> vivo-like stochastic <strong>in</strong>put (8.7) withthe random conductances <strong>in</strong> (b). Part (a) is a magnification of a small region <strong>in</strong> (c).Shown are simulations of 30 ˙v = 3(v+55)(v+42)−u+I(t), ˙u = 0.01{−0.25(v+55)−u},if v ≥ +10, then v ← −40, u ← u + 90. Data was k<strong>in</strong>dly provided by Niraj S. Desaiand Betsy C. Walcott.In vivo-like conditionsIn Fig. 8.17a (dashed curve) we show the response of <strong>in</strong> vitro recorded layer 5 pyramidalneuron of rat motor cortex to fluctuat<strong>in</strong>g <strong>in</strong> vivo-like <strong>in</strong>put. First, random excitatoryand <strong>in</strong>hibitory conductances, g AMPA (t) and g GABA (t) (Fig. 8.17b), were generated us<strong>in</strong>gthe Ornste<strong>in</strong>-Uhlenbeck stochastic process (Uhlenbeck and Ornste<strong>in</strong> 1930), which wasorig<strong>in</strong>ally developed to describe Brownian motion, but can equally well describe <strong>in</strong>vivo-like fluctuat<strong>in</strong>g synaptic conductances produced by random fir<strong>in</strong>gs (Destexhe etal. 2001). Let E AMPA = 0 mV and E GABA = −65 mV denote the reverse potentials ofexcitatory and <strong>in</strong>hibitory synapses, respectively. The correspond<strong>in</strong>g currentI(t) =excitatory <strong>in</strong>put{ }} {g AMPA (t)(E AMPA − V (t)) +<strong>in</strong>hibitory <strong>in</strong>put{ }} {g GABA (t)(E GABA − V (t)) , (8.7)was <strong>in</strong>jected <strong>in</strong>to the neuron us<strong>in</strong>g the dynamic clamp protocol (Sharp et al. 1993),where V (t) denotes the <strong>in</strong>stantaneous membrane potential of the neuron. The sameconductances were <strong>in</strong>jected <strong>in</strong>to the simple model (8.5, 8.6), whose parameters wereadjusted to fit this particular neuron. The superimposed voltage traces, depicted <strong>in</strong>

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