Lecture 21 Neural Modeling II Martin Giese Aim of this Class Account for experimentally observed effects in motion perception with the simple neuronal model presented in Lecture 18: • • • • • • Dependence on stimulus strength Stochastic nature of perception Psychometric function Bistable motion perception Hysteresis Perceptual switching Model Neuron Model Neuron Population of real neurons Similar tuning properties: – – – position direction speed Visual Field Simplified Neuron Model Model for single local motion detector (consisting of many neurons with similar tuning) “Time constant” du (t ) u (t ) s (t ) h dt Rate of change of average membrane potential Resting level parameter Stimulus intensity (input signal) Average membrane potential (over all neurons) Phase Diagram du dt du (t ) u (t ) s h dt attractor (u stays constant) u* u u increases “Phase diagram”: u decreases u* du dt u Attractor Follow the arrows in the phase diagram: u(t) changes always in the direction of the attractor u* u(t) u(t) u* u(t) u* u* t t t u* u attractor du dt With Noise State fluctuates near attractor u(t) Probability u* u t u* u attractor du dt With Noise (for Experts) Exact mathematical description: Stochastic Process (Ornstein-Uhlenbeck) u(t) u* t du (t ) u (t ) s h (t ) dt white noise Single Neuron Model Activation and Perception Assumption: Motion is seen only when u(t) is larger than a threshold value T. u T u T u(t) t No stimulus seen u(t) t Stimulus t seen No Stimulus s = 0 du dt du (t ) u (t ) 0 h dt motion seen attractor u* -h “Phase diagram”: u T (threshold) u* du dt u Large Stimulus s > 0 du dt du (t ) u (t ) s h dt motion seen attractor u* T “Phase diagram”: u s-h u* du dt u Psychometric Function Stimulus: -h motion seen T u P no u u u * Psychometric function: P(seen) 1 P weak u u u* 0 P strong u u * u s Two Neuron Model Inhibition Motivation Bistable perception by lateral inhibition between motion detectors ! Inhibition Two Uncoupled Neurons Neuron 1: Neuron 2: du1 (t ) u1 (t ) s1 (t ) h dt du 2 (t ) u 2 (t ) s2 (t ) h dt No inhibition. 2D Phase Diagram u Attractor: pair of activations [u1*, u2*] u2 attractor u* u* du dt u * du dt u u1 Synaptic Coupling Neuron for “horizontal” Neuron for “vertical” Inhibition Coupling through nonlinear threshold function ! (Katz & Miledi, 1967) Two Coupled Neurons Inhibition of neuron 1 by neuron 2 du1 (t ) u1 (t ) s1 (t ) h w1 f (u2 (t )) Neuron 1: dt du2 (t ) Neuron 2: u2 (t ) s2 (t ) h w2 f (u1 (t )) dt f(u) Inhibition of neuron 2 by neuron 1 u 2D Phase Diagram Two attractors ! 2 Attractors (for Experts) Linear system theory predicts for purely linear interaction: • not more than one isolated attractor • if multiple attractors, they are not asymptotically stable (in all directions in phase space) vertical motion seen 2 Attractors horizontal motion not seen Two attractors ! vertical motion not seen horizontal motion seen Bistability basin of attraction for vertical perceptual ambiguity basin of attraction for horizontal Perceptual Switching Fluctuations can push state in the other basin of attraction perceptual switch Influence of the Aspect Ratio Change of aspect ratio: • • different motion detectors activated change of the mutual inhibition Stimulus Inhibition Relative Stability horizontal and vertical equally stable horizontal more stable than vertical Hysteresis State can not follow change of attractor immediately tendency to stay in the same basin of attraction hysteresis Switching Probability Medium aspect ratio switch possible Extreme aspect ratio switch very unlikely Programming NeuDyn.m Simulates neural model with 1or 2 neurons and returns the time series of activation. ut = NeuDyn(tsim, u0, S, tau, Q, W); NeuDyn.m Simulates neural model with 1or 2 neurons and returns the time series of activation. simulation time steps time constant strength of fluctuations ut = NeuDyn(tsim, u0, S, tau, Q, W); NeuDyn.m Simulates neural model with 1or 2 neurons and returns the time series of activation. ut = NeuDyn(tsim, u0, S, tau, Q, W); initial activation stimulus strength (number / vector) NeuDyn.m Simulates neural model with 1or 2 neurons and returns the time series of activation. ut = NeuDyn(tsim, u0, S, tau, Q, W); simulated activity time series (vector / matrix) interaction (vector / matrix) NeuDyn.m Simulates neural model with 1or 2 neurons and returns the time series of activation. simulation time steps time constant strength of fluctuations ut = NeuDyn(tsim, u0, S, tau, Q, W); initial activation stimulus strength simulated activity interaction time series Literature Additional readings (for interested people): Hock, H.S., Kelso, J.A.S., Schöner, G. (1993). Bistability and hysteresis in the organization of apparent motion patterns. Journal of Experimental Psychology: Human Perception and Performance, 19(1), 63-80. Giese, M.A. (1999). Dynamic Neural Field Theory of Motion Perception, Kluwer, Dordrecht, NL. Wilson, H.R. (1999). Spikes, Decisions, and Actions. Oxford University Press, Oxford, UK.