Optimal information processing in neural populations. Nigel Stocks School of Engineering, University of Warwick, UK. Collaborators: A. Nikitin R. P. Morse M. D. McDonnell - School of Engineering, University of Warwick, UK - Aston University, Birmingham UK - University of South Australia, Australia. 1 Sensory signal transduction Cochlea Inner Hair Cell Data from H1 neuron of fly (taken from ‘Spikes’, Rieke et al) 2 HAIR CELLS (mechanical transducers) K+ Stereocilia (“hairs”) Ca+ 5 µm 3 Electron microscope image courtesy of Dr. David Furness, Keele University. Noise sources K+ Brownian motion of stereocilia, 2-3nm rms (at threshold displacements of 0.3nm can be detected). Thermal noise of ion channels. Stochastic nature of synaptic transmission. SNR ~ 0-10dB Ca+ 5 µm 4 Electron microscope images courtesy of Dr. David Furness, Keele University. SNR ~ 0dB SNR ~ 70dB To understand performance need to consider collective behaviour of neurons. How do we maximise information flow through a population of noisy neurons? 5 Array of N McCulloch-Pitts Neurons Random Gaussian signal 2 Y = Number of units ‘on’ (discrete output) Each unit has the transfer function given by 1 if yi = 0 if S (x t) + ηi > θ i S (x t) + ηi < θ i 6 N=1 (single neuron) Threshold=0 Threshold=2.83 σ = noise power signal power 7 Maximising Information for identical thresholds 8 Array of N McCulloch-Pitts Neurons (comparators) Random Gaussian signal Y = Number of units ‘on’ (discrete output) Each unit has the transfer function given by 1 if yi = 0 if S (x t) + ηi > θ i S (x t) + ηi < θ i 9 N>1 and {ϑi } = 0 Suprathreshold Stochastic Resonance (SSR) Information, I (bits), Threshold=0 Threshold=0 N. G. Stocks, Phys. Rev. Lett 84, 2310 (2000) 10 10 comparators without noise Input signal y1 y2 y3 y4 y5 y6 y7 y8 y9 y10 Output response 11 10 comparators with noise Input signal y1 y2 y3 y4 y5 y6 y7 y8 y9 y10 Output response 12 SSR in Real Neurons 13 Experiments with additive noise Suprathreshold (+6 dB) signal. 1 second duration. Same waveform for each presentation. Stimulating electrodes Isolated sciatic nerve Current source Noise waveform, that differed for each presentation 14 Information transmission with additive noise Noise 1 Response 1 virtual fibre 1 Noise 2 Response 2 virtual fibre 2 Noise 3 Response 3 virtual fibre 3 Noise 4 Response 4 virtual fibre 4 Responses from 1 sciatic nerve fibre15 Experimental Results SSR Classical SR 0.3 Signal -6dB Signal 6dB 0.5 Mutual information (bits) M u tu al in fo rm atio n (b its) 0.6 0.4 0.3 1 fibre 2 fibres 4 fibres 8 fibres 16 fibres 20 fibres 0.2 0.1 0.0 -50 -40 -30 -20 -10 0 Noise level (dB re base level) 10 20 1 fibre 2 fibres 4 fibres 8 fibres 16 fibres 20 fibres 0.2 0.1 0.0 -50 -40 -30 -20 -10 0 10 20 Noise level (dB re base level) 16 Period histograms. Comparison of single fibre acoustic data to LIF model Model Squirrel Monkey Anderson D.J, J. Acoustic Soc. Of Am. 54, 361 (1973) 17 Information curves Membrane noise Signal 28dB 18 Information curves Membrane noise ‘natural’ noise Signal 28dB 19 1 Signal 6dB 0.5 1 neurone 4 neurones 8 neurones 16 neurones 0.8 0.4 I (bits) 0.6 0.3 0.4 1 fibre 2 fibres 4 fibres 8 fibres 16 fibres 20 fibres 0.2 0.1 0.2 0.0 -50 -40 -30 -20 -10 0 10 0 -60 20 -40 -20 0 20 40 σ (dB) Noise level (dB re base level) 1 1 neurone 4 neurones 8 neurones 16 neurones 0.8 0.6 I (bits) Mutual information (bits) 0.6 0.4 0.2 20 0 -60 -40 -20 0 20 40 ERROR: invalidrestore OFFENDING COMMAND: restore STACK: -savelevel-savelevel-dictionary-