A special Gauss – Markov neuronal model

posed for time-homogeneous diffusion processes and mentioned in the previous subsection, in a previous paper cf. Di Nardo et al., 1998b it has been proved that the FPT density of a Gauss – Markov process can be obtained by solving the simple, non-singular, Volterra second kind inte- gral Eq. 5, with St C 1 [t , and c [St, t y, t]= St − mt 2 − St − mt 2 h 1 th 2 t − h 2 th 1 t h 1 th 2 t − h 2 th 1 t − y − mt 2 h 2 th 1 t − h 2 th 1 t h 1 th 2 t − h 2 th 1 t × f[St, t y, t]. By making use of this result, in Di Nardo et al. 1998b an efficient numerical procedure based on a repeated Simpson’s rule has been proposed to evaluate FPT densities of Gauss – Markov processes. In the following, we present a special non-sta- tionary Gauss – Markov neuronal model and make use of such numerical procedure to analyze the corresponding firing pdf’s.

3. A special Gauss – Markov neuronal model

In this section, we consider the neuronal model {Xt, t I}, with I [0, + , characterized by the mean mt = r + ljt, 12 where j t = Á Ã Í Ã Ä aq q − a e − tq − e − ta if a q te − tq if a = q, and by the covariance cs, t = s 2 q 2 e − t − sq , 13 with l \ 0, r R, , q \ 0, a \ 0 and s \ 0. As is easily seen, the above conditions characterizing Gauss – Markov processes are satisfied by Eqs. 12 and 13. Indeed, in this case we have h 1 t = s 2 q e tq 2 and h 2 t = e − tq . Hence, here we are defining a Gauss – Markov neuronal model. Re- calling Eq. 11, the coefficients A 1 x, t and A 2 x, t for the underlying process are, respectively, given by A 1 x, t = − 1 q x − r + le − ta , A 2 x, t = s 2 . 14 Hence, the infinitesimal moments of the OU neuronal model turn out to be a special case of model expressed by Eq. 14. Indeed, for l = 0 Eq. 14 yields A 1 x, t = − x − rq; moreover, when l \ 0 and a ¡0 the drift A 1 x, t goes to − x − rq. Let us consider the deterministic model sug- gested by Eqs. 1 and 14 in the absence of randomness, i.e. with s 2 = 0 described by dxt dt = − 1 q x − r + le − ta , xt = x . It is not hard to see that xt = x e − t − t q + r [1 − e − t − t q ] + l [jt − jt e − t − t q ]. 15 Recalling Eqs. 12 and 13, we note that, similarly to the OU model, again the conditional mean Mt t , given by the first of Eq. 10, identifies with Eq. 15. Furthermore, if t = 0 and x = r , from Eq. 15 one has xt = r + ljt, that coincides with the mean mt given by Eq. 12. We note that relations Eq. 14 can be a poste- riori interpreted in the following way. The neu- ron’s membrane potential is not only subject to the usual spontaneous exponential decay and to endogenous random components, but it also expe- riences an external input whose magnitude, how- ever, exponentially damps with the time-constant a . Hence, the effect of such an input depends on the two parameters a and l. In other words, such Fig. 3. Firing pdf’s for the neuronal model of Section 3, for t = 0, x = r = − 70, s 2 = 2, S = − 60, q = 5, l = 14 and a = 1, 2, 3, 4. Decreasing modes refer to increasing values of a. parameters mimic the effect of an external neu- ron’s input whose initial strength l exponentially damps with the time-constant a. When x 5 r , which is the interesting case in neurobiology, from Eq. 15 it follows that xt is initially increasing, to decrease monotonically as t “ towards the resting potential r, after reaching a maximum. Note the significant diversity of behavior of xt for l \ 0 and for l = 0, the latter representing the deterministic version of the OU model. Indeed, if l = 0, xt monotonically tends to the resting potential r for all x r .

4. Numerical results