Dynamics of Perceptual Bistability J Rinzel, NYU w/ N Rubin, A Shpiro, R Curtu, R Moreno • Alternations in perception of ambiguous stimulus – irregular… • Oscillator models – mutual inhibition, switches due to adaptation -- noise gives randomness to period • Attractor models – noise driven, no alternation w/o noise • Constraints from data: <T>, CV Which model is favored? Mutual inhibition with slow adaptation alternating dominance and suppression IV: Levelt, 1968 Oscillator Models for Directly Competing Populations Two mutually inhibitory populations, corresponding to each percept. Firing rate model: r1(t), r2(t) Slow negative feedback: adaptation or synaptic depression. f r1r11 r2r2 f Slow adaptation, a1(t) u τ dr1/dt = -r1 + f(-βr2 - g a1+ I1) τa da1/dt = -a1 + r1 No recurrent excitation …half-center oscillator τ dr2/dt = -r2 + f(-βr1 - g a2+ I2) τa da2/dt = -a2 + r2 τa >> τ , f(u)=1/(1+exp[(θ-u)/k]) w/ N Rubin, A Shpiro, R Curtu Shpiro et al, J Neurophys 2007 Wilson 2003; Laing and Chow 2003 Alternating firing rates Adaptation slowly grows/decays WTA or ATT regime IV adaptation LC model Analysis of Dynamics Fast-Slow dissection: r1 , r2 fast variables a1 , a2 slow variables a1, a2 frozen r1- nullcline r2- nullcline r1 = f(-βr2 - g a1+ I1) r2 = f(-βr1 - g a2+ I2) r2 r1 r1-r2 phase plane, slowly drifting nullclines r1- nullcline r2- nullcline r2 At a switch: • saddle-node in fast dynamics. • dominant r is high while system rides near “threshold” of suppressed populn’s nullcline ESCAPE. Switching occurs when a1-a2 traj reaches a curve of SNs (knees) r1 β =0.9, I1=I2=1.4 a2 a1 Switching due to adaptation: release or escape mechanism f input net input net input Small I, “release” Large I, “escape” Dominant, a ↑ I–ga θ I+αa–ga θ Suppressed, a ↓ I–ga-β Recurrent excitation, secures “escape” Noise leads to random dominance durations and eliminates WTA behavior. τ dri/dt = -ri+ f(-βrj - g ai+ Ii + ni) τa dai/dt = -ai + ri Added to stimulus I1,2 s.d., σ = 0.03, τn = 10 Model with synaptic depression Noise-Driven Attractor Models w/ R Moreno, N Rubin J Neurophys, 2007 No oscillations if noise is absent. Kramers 1940 LP-IV in an attractor model Compare dynamical skeletons: “oscillator” and attractor-based models activity WTA rA=rB OSC gA= gB Observed variability and mean duration constrain the model. With noise 1 sec < mean T < 10 sec 0.4 < CV < 0.6 Noise-free Difficult to arrange low CV and high <T> in OSC regime. Favored: noise-driven attractor with weak adaptation – but not far from oscillator regime. With noise Best fit distribution depends on parameter values. Noise dominated Adaptation dominated I1 , I2 = 0.6 SUMMARY Swartz Foundation and NIH. w/ N Rubin, A Shpiro, R Curtu, R Moreno Experimentally: • Monotonic <T> vs I • <T> and CV as constraints • No correlation between successive cycles Models, one framework – vary params. Mutual, direct inhibition; w/o recurrent excitation Non-monotonic dominance duration vs I1, I2 • Attractor regime, noise dominated • Better match w/ data. • Balance between noise level and adaptation strength. • Oscillator regime, adaptation dominated • Relatively smaller CV. • Relatively greater correlation between successive cycles. • Moreno model (J Neurophys 2007): local inhibition, strong recurrent excitation: monotonic <T> vs I.
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