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Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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One-Dimensional <strong>Systems</strong> 59current (nA)I1L (V)I(V)0-1I Na,p (V)-2-60 -40 -20 0 20 40 60membrane potential (mV)aderivative of membrane potential (mV/ms)100V=F(V)50F(V)=-I(V)/C0-50-60 -40 -20 0 20 40 60membrane potential, V (mV)bFigure 3.5: a. I-V relations of the leak current, I L , fast Na + current, I Na , and comb<strong>in</strong>edcurrent I(V ) = I L (V ) + I Na (V ) <strong>in</strong> the I Na,p -model (3.5). Dots denote I 0 (V ) data fromlayer 5 pyramidal cell <strong>in</strong> rat visual cortex. b. The right-hand side of the I Na,p -model(3.5).dynamical systems are usually written <strong>in</strong> the form˙V = F (V ) , V (0) = V 0 ∈ R , (3.6)for example,˙V = −80 − V , V (0) = −20 ,where V is a scalar time-dependent variable denot<strong>in</strong>g the current state of the system,˙V = V t = dV/dt is its derivative with respect to time t, F is a scalar function (itsoutput is one-dimensional) that determ<strong>in</strong>es the evolution of the system, e.g., the righthandside of (3.5) divided by C; see Fig. 3.5b. V 0 ∈ R is an <strong>in</strong>itial condition, and R isthe real l<strong>in</strong>e, i.e., a l<strong>in</strong>e of real numbers (R n would be the n-dimensional real space).In the context of dynamical systems, the real l<strong>in</strong>e R is called phase l<strong>in</strong>e or state l<strong>in</strong>e(phase space or state space for R n ) to stress the fact that each po<strong>in</strong>t <strong>in</strong> R correspondsto a certa<strong>in</strong>, possibly <strong>in</strong>admissible state of the system, and each state of the systemcorresponds to a certa<strong>in</strong> po<strong>in</strong>t <strong>in</strong> R. For example, the state of the Ohmic membrane(3.1) is just its membrane potential V ∈ R. The state of the Hodgk<strong>in</strong>-Huxley model(see Sect. 2.3) is the four-dimensional vector (V, m, n, h) ∈ R 4 . The state of the I Na,p -model (3.5) is aga<strong>in</strong> its membrane potential V ∈ R, because the value m = m ∞ (V ) isunequivocally def<strong>in</strong>ed by V .When all parameters are constant, the dynamical system is called autonomous.When at least one of the parameters is time-dependent, the system is non-autonomous,denoted as ˙V = F (V, t).To solve (3.6) means to f<strong>in</strong>d a function V (t) whose <strong>in</strong>itial value is V (0) = V 0and whose derivative is F (V (t)) at each moment t ≥ 0. For example, the functionV (t) = V 0 + at is an explicit analytical solution to the dynamical system ˙V = a. Theexponentially decay<strong>in</strong>g function V (t) = E L + (V 0 − E L )e −g Lt/C depicted <strong>in</strong> Fig. 3.7,

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