Membrane equation. VCl. dv dt + V = V Na G Na + V K G K + V Cl G Cl. G total. C m. G total = G Na + G K + G Cl
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- Domenic Davidson
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1 Spiking neurons
2
3 Membrane equation V GNa GK GCl Cm VNa VK VCl dv dt + V = V Na G Na + V K G K + V Cl G Cl G total G total = G Na + G K + G Cl = C m G total
4 Membrane with synaptic inputs V Gleak GNa GK GCl Cm Vr VNa VK VCl dv dt + V = V r G leak + V Na G Na + V K G K + V Cl G Cl G total G total = G leak + G Na + G K + G Cl = C m G total
5 Membrane with input current V Gleak Cm Iin(t) Vr dv dt + V = V r + 1 G leak I in (t) G leak = G Na + G K + G Cl = C m G leak
6 Voltage-gated channels
7 Leaky integrate-and-fire neuron
8 Rate coding hypothesis: the signal conveyed by a neuron is in the rate of spiking. Spiking irregularity is largely due to noise and does not convey information.
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10 Fly H1 neuron - constant stimulus (de Ruyter et al., 1997) val histogram describing the probability density, ( ), of finding an interspike
11 Fly H1 neuron - time-varying stimulus (de Ruyter et al., 1997)
12 Spike timing can be very precise in response to time-varying signals Mainen & Sejnowski (1995)
13 Spike timing can be very precise in response to time-varying signals MT neuron response to stochastic moving dot stimuli at different levels of coherence (Newsom lab) Analysis by Bair & Koch (1996)
14
15 Neural encoding and decoding
16 Encoding and decoding are related through the joint distribution over stimulus and response h i 0-20 v av (omm/s) n av n v (omm/s) f g v av (n) n v n av (v) 0 decoding d P(v n) P(n v) e encoding v 20 P(n,v) c n 0 P(v) P(n) a v n 0 b (from Spikes)
17 s(t) sensory system stimulus (t) spike train ŝ(t) estimation algorithm estimate From Spikes, by Rieke, Warland, de Ruyter, & Bialek
18 Reconstruction kernel filter amplitude ( /s) a time (ms) Fly H1 neuron response response b Stimulus reconstruction c ŝ(t) = (t) k(t) ˆk(t) = arg min k(t) D [s(t) (t) k(t)] 2E t velocity ( /s) From Spikes, by Rieke, Warland, de Ruyter, & Bialek time (ms)
19 Strategy for estimating information rate 1. Estimate signal from spikes (t)! ŝ(t) 2. Compute noise in estimate ñ(!) = s(!) ˆ s(!) 3. Compute SNR SNR(!) = h s(!) 2 i h ñ(!) 2 i 4. Calculate lower bound to information rate from SNR R = 1 2 Z d! 2 log 2[1 + SNR(!)] Adapted from Spikes, by Rieke, Warland, de Ruyter, & Bialek
20 Coding by on- and off-cells
21 off cell on cell
22 Neural responses are half-wave rectified (action potentials are positive-only). Signals are thus combined in a push-pull fashion, similar to push-pull amplifiers. From: Neural Engineering, by Eliasmith & Anderson
23 Push-Pull decoding
24 Efficient coding model of retina (Karklin & Simoncelli 2012) Objective function: X I(X; R) j hr j i j a b ON center OFF center c
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