Neural Coding and Decoding Albert E. Parker Center for Computational Biology Department of Mathematical Sciences Montana State University Collaborators: Alexander Dimitrov John P. Miller Zane Aldworth Tomas Gedeon Brendan Mumey Problem: How does neural ensemble activity represent information about sensory stimuli? Our Approach: • Build a model using Information Theory My Research: Probability Theory • Use the model to determine a coding scheme My Research: Numerical Optimization Techniques Neural Coding and Decoding. Goal: Determine a coding scheme: How does neural ensemble activity represent information about sensory stimuli? Demands: • An animal needs to recognize the same object on repeated exposures. Coding has to be deterministic at this level. • The code must deal with uncertainties introduced by the environment and neural architecture. Coding is by necessity stochastic at this finer scale. Major Problem: The search for a coding scheme requires large amounts of data How to determine a coding scheme? Idea: Model a part of a neural system as a communication channel using Information Theory. This model enables us to: • Meet the demands of a coding scheme: o Define a coding scheme as a relation between stimulus and neural response classes. o Construct a coding scheme that is stochastic on the finer scale yet almost deterministic on the classes. • Deal with the major problem: o Use whatever quantity of data is available to construct coarse but optimally informative approximations of the coding scheme. o Refine the coding scheme as more data becomes available. • Investigate the cricket cercal sensory system. Stimulus and Response Classes X stimulus sequences response sequences Y stimulus/response sequence pairs distinguishable classes of stimulus/response pairs Information Theoretic Quantities A quantizer or encoder, Q, relates the environmental stimulus, X, to the neural response, Y, through a process called quantization. In general, Q is a stochastic map Q( y| x ): X Y The Reproduction space Y is a quantization of X. This can be repeated: Let Yf be a reproduction of Y. So there is a quantizer q ( y f | y ): Y Yf Use Mutual information to measure the degree of dependence between X and Yf. q ( y f , y ) p( x , y ) y I ( X , Yf ) q ( y f , y ) p( x , y ) log p ( x ) p ( y ) q ( y , y ) y,y f y f Use Conditional Entropy to measure the self-information of Yf given Y H (Yf |Y ) p( y ) q ( y y,y f f | y ) log(q ( y f | y )) The Model Problem: To determine a coding scheme between X and Y requires large amounts of data Idea: Determine the coding scheme between X and Yf, a squashing (reproduction) of Y, such that: Yf preserves as much information (mutual information) with X as possible and the self-information (entropy) of Yf |Y is maximized. That is, we are searching for an optimal mapping (quantizer): q * ( y f | y) : Y Y f that satisfies these conditions. Justification: Jayne's maximum entropy principle, which states that of all the quantizers that satisfy a given set of constraints, choose the one that maximizes the entropy. Equivalent Optimization Problems Maximum entropy: maximize F(q(yf|y)) = H(Yf|Y) constrained by I(X;Yf ) Io Io determines the informativeness of the reproduction. Deterministic annealing (Rose, ’98): maximize F(q(yf|y)) = H(Yf|Y) + I(X,Yf ). Small favor maximum entropy, large - maximum I(X,Yf ). Simplex Algorithm: maximize I(X,Yf ) over vertices of constraint space Implicit solution: qy f | y e q DI q p y y , y e yf q DI q p y ? ? Modeling the cricket cercal sensory system as a communication channel Nervous system Signal Communication channel Wind Stimulus and Neural Response in the cricket cercal system Neural Responses (over a 30 minute recording) caused by white noise wind stimulus. X Time in ms. A t T=0, the first spike occurs Y Some of the air current stimuli preceding one of the neural responses Neural Responses (these are all doublets) for a 12 ms window T, ms Quantization: A quantizer is any map f: Y -> Yf from Y to a reproduction space Yf with finitely many elements. Quantizers can be deterministic y f y or qy f | y probabilistic refined Y Yf Y Applying the algorithm to cricket sensory data. Yf 1 2 1 Yf 2 3 Y Conclusions We • model a part of the neural system as a communication channel. • define a coding scheme through relations between classes of stimulus/response pairs. - Coding is probabilistic on the individual elements of X and Y. - Coding is almost deterministic on the stimulus/response classes. To recover such a coding scheme, we • propose a new method to quantify neural spike trains. - Quantize the response patterns to a small finite space (Yf). - Use information theoretic measures to determine optimal quantizer for a fixed reproduction size. - Refine the coding scheme by increasing the reproduction size. • present preliminary results with cricket sensory data.