The Hamiltonian Brain: Efficient Probabilistic Inference with Excitatory-Inhibitory Neural Circuit Dynamics

Probabilistic inference offers a principled framework for understanding both behaviour and cortical computation. However, two basic and ubiquitous properties of cortical responses seem difficult to reconcile with probabilistic inference: neural activity displays prominent oscillations in response to constant input, and large transient changes in response to stimulus onset. Indeed, cortical models of probabilistic inference have typically either concentrated on tuning curve or receptive field properties and remained agnostic as to the underlying circuit dynamics, or had simplistic dynamics that gave neither oscillations nor transients. Here we show that these dynamical behaviours may in fact be understood as hallmarks of the specific representation and algorithm that the cortex employs to perform probabilistic inference. We demonstrate that a particular family of probabilistic inference algorithms, Hamiltonian Monte Carlo (HMC), naturally maps onto the dynamics of excitatory-inhibitory neural networks. Specifically, we constructed a model of an excitatory-inhibitory circuit in primary visual cortex that performed HMC inference, and thus inherently gave rise to oscillations and transients. These oscillations were not mere epiphenomena but served an important functional role: speeding up inference by rapidly spanning a large volume of state space. Inference thus became an order of magnitude more efficient than in a non-oscillatory variant of the model. In addition, the network matched two specific properties of observed neural dynamics that would otherwise be difficult to account for using probabilistic inference. First, the frequency of oscillations as well as the magnitude of transients increased with the contrast of the image stimulus. Second, excitation and inhibition were balanced, and inhibition lagged excitation. These results suggest a new functional role for the separation of cortical populations into excitatory and inhibitory neurons, and for the neural oscillations that emerge in such excitatory-inhibitory networks: enhancing the efficiency of cortical computations.

[1]  Pei-Ji Liang,et al.  Temporal and Spatial Properties of the Retinal Ganglion Cells' Response to Natural Stimuli Described by Treves-Rolls Sparsity , 2009, 2009 3rd International Conference on Bioinformatics and Biomedical Engineering.

[2]  Charles J. Wilson,et al.  Spontaneous subthreshold membrane potential fluctuations and action potential variability of rat corticostriatal and striatal neurons in vivo. , 1997, Journal of neurophysiology.

[3]  Terrence J. Sejnowski,et al.  Assignment of Multiplicative Mixtures in Natural Images , 2004, NIPS.

[4]  Peter Dayan,et al.  Doubly Distributional Population Codes: Simultaneous Representation of Uncertainty and Multiplicity , 2003, Neural Computation.

[5]  Adam M. Packer,et al.  Inferring neural population dynamics from multiple partial recordings of the same neural circuit , 2013, NIPS.

[6]  David C. Knill,et al.  Surface orientation from texture: ideal observers, generic observers and the information content of texture cues , 1998, Vision Research.

[7]  W. Singer,et al.  Modulation of Neuronal Interactions Through Neuronal Synchronization , 2007, Science.

[8]  A. Pouget,et al.  Marginalization in Neural Circuits with Divisive Normalization , 2011, The Journal of Neuroscience.

[9]  Peter Dayan,et al.  Statistical Models of Linear and Nonlinear Contextual Interactions in Early Visual Processing , 2009, NIPS.

[10]  Aapo Hyvärinen,et al.  Statistical Models of Natural Images and Cortical Visual Representation , 2010, Top. Cogn. Sci..

[11]  S. Duane,et al.  Hybrid Monte Carlo , 1987 .

[12]  Peter Dayan,et al.  Cortical Surround Interactions and Perceptual Salience via Natural Scene Statistics , 2012, PLoS Comput. Biol..

[13]  Peter Dayan,et al.  Computational Differences between Asymmetrical and Symmetrical Networks , 1998, NIPS.

[14]  K. Harris,et al.  Gating of Sensory Input by Spontaneous Cortical Activity , 2013, The Journal of Neuroscience.

[15]  T. Branco,et al.  The probability of neurotransmitter release: variability and feedback control at single synapses , 2009, Nature Reviews Neuroscience.

[16]  A. Pouget,et al.  Reading population codes: a neural implementation of ideal observers , 1999, Nature Neuroscience.

[17]  Andrew Gelman,et al.  Handbook of Markov Chain Monte Carlo , 2011 .

[18]  M. Ernst,et al.  Humans integrate visual and haptic information in a statistically optimal fashion , 2002, Nature.

[19]  J. Tenenbaum,et al.  Special issue on “Probabilistic models of cognition , 2022 .

[20]  Tirin Moore,et al.  Rapid enhancement of visual cortical response discriminability by microstimulation of the frontal eye field , 2007, Proceedings of the National Academy of Sciences.

[21]  Tiago Branco,et al.  Dendritic nonlinearities are tuned for efficient spike-based computations in cortical circuits , 2015, eLife.

[22]  K. Svoboda,et al.  Neural Activity in Barrel Cortex Underlying Vibrissa-Based Object Localization in Mice , 2010, Neuron.

[23]  I. Nelken,et al.  Processing of Sounds by Population Spikes in a Model of Primary Auditory Cortex , 2007, Front. Neurosci..

[24]  Wei Ji Ma,et al.  Bayesian inference with probabilistic population codes , 2006, Nature Neuroscience.

[25]  Michael S. Lewicki,et al.  Emergence of complex cell properties by learning to generalize in natural scenes , 2009, Nature.

[26]  P. Fries,et al.  Robust Gamma Coherence between Macaque V1 and V2 by Dynamic Frequency Matching , 2013, Neuron.

[27]  B. McNaughton,et al.  Paradoxical Effects of External Modulation of Inhibitory Interneurons , 1997, The Journal of Neuroscience.

[28]  Wolf Singer,et al.  Neuronal Synchrony: A Versatile Code for the Definition of Relations? , 1999, Neuron.

[29]  P. Lennie,et al.  Information Conveyed by Onset Transients in Responses of Striate Cortical Neurons , 2001, The Journal of Neuroscience.

[30]  P. Fries Neuronal gamma-band synchronization as a fundamental process in cortical computation. , 2009, Annual review of neuroscience.

[31]  A. Borst Seeing smells: imaging olfactory learning in bees , 1999, Nature Neuroscience.

[32]  Shawn D. Burton,et al.  NeuroElectro: a window to the world's neuron electrophysiology data , 2014, Front. Neuroinform..

[33]  Jason Bell,et al.  Contour inflections are adaptable features. , 2014, Journal of vision.

[34]  P. Berkes,et al.  Statistically Optimal Perception and Learning: from Behavior to Neural Representations , 2022 .

[35]  József Fiser,et al.  Spontaneous Cortical Activity Reveals Hallmarks of an Optimal Internal Model of the Environment , 2011, Science.

[36]  Sen Song,et al.  Highly Nonrandom Features of Synaptic Connectivity in Local Cortical Circuits , 2005, PLoS biology.

[37]  Daniel B. Rubin,et al.  The Stabilized Supralinear Network: A Unifying Circuit Motif Underlying Multi-Input Integration in Sensory Cortex , 2015, Neuron.

[38]  Aapo Hyvärinen,et al.  Interpreting Neural Response Variability as Monte Carlo Sampling of the Posterior , 2002, NIPS.

[39]  A. Pouget,et al.  Probabilistic brains: knowns and unknowns , 2013, Nature Neuroscience.

[40]  Peter Dayan,et al.  Optimal Recall from Bounded Metaplastic Synapses: Predicting Functional Adaptations in Hippocampal Area CA3 , 2014, PLoS Comput. Biol..

[41]  Edgar Bermudez Contreras,et al.  Formation and Reverberation of Sequential Neural Activity Patterns Evoked by Sensory Stimulation Are Enhanced during Cortical Desynchronization , 2013, Neuron.

[42]  J. Cowan,et al.  Excitatory and inhibitory interactions in localized populations of model neurons. , 1972, Biophysical journal.

[43]  R. J. van Beers,et al.  Integration of proprioceptive and visual position-information: An experimentally supported model. , 1999, Journal of neurophysiology.

[44]  Radford M. Neal Sampling from multimodal distributions using tempered transitions , 1996, Stat. Comput..

[45]  Michael I. Jordan,et al.  An internal model for sensorimotor integration. , 1995, Science.

[46]  A. Pérez-Villalba Rhythms of the Brain, G. Buzsáki. Oxford University Press, Madison Avenue, New York (2006), Price: GB £42.00, p. 448, ISBN: 0-19-530106-4 , 2008 .

[47]  Guillaume Hennequin,et al.  Fast Sampling-Based Inference in Balanced Neuronal Networks , 2014, NIPS.

[48]  József Fiser,et al.  Neural Variability and Sampling-Based Probabilistic Representations in the Visual Cortex , 2016, Neuron.

[49]  Andrew M. Clark,et al.  Stimulus onset quenches neural variability: a widespread cortical phenomenon , 2010, Nature Neuroscience.

[50]  W. Gerstner,et al.  Spike-Timing-Dependent Plasticity: A Comprehensive Overview , 2012, Front. Syn. Neurosci..

[51]  David J. Field,et al.  Emergence of simple-cell receptive field properties by learning a sparse code for natural images , 1996, Nature.

[52]  R. C. Gerkin,et al.  Brain-wide analysis of electrophysiological diversity yields novel categorization of mammalian neuron types. , 2015, Journal of neurophysiology.

[53]  D. Kullmann,et al.  Plasticity of Inhibition , 2012, Neuron.

[54]  J. Rinn,et al.  DeCoN: Genome-wide Analysis of In Vivo Transcriptional Dynamics during Pyramidal Neuron Fate Selection in Neocortex , 2015, Neuron.

[55]  David M. Sobel,et al.  A theory of causal learning in children: causal maps and Bayes nets. , 2004, Psychological review.

[56]  Xiaoqin Wang,et al.  Sustained firing in auditory cortex evoked by preferred stimuli , 2005, Nature.

[57]  E. Basar,et al.  A review of brain oscillations in cognitive disorders and the role of neurotransmitters , 2008, Brain Research.

[58]  W. Wildman,et al.  Theoretical Neuroscience , 2014 .

[59]  P. Dayan,et al.  Perceptual organization in the tilt illusion. , 2009, Journal of vision.

[60]  Alexandre Pouget,et al.  Probabilistic Interpretation of Population Codes , 1996, Neural Computation.

[61]  Radford M. Neal MCMC Using Hamiltonian Dynamics , 2011, 1206.1901.

[62]  P. Lennie,et al.  Rapid adaptation in visual cortex to the structure of images. , 1999, Science.

[63]  W. Gerstner,et al.  Optimal Control of Transient Dynamics in Balanced Networks Supports Generation of Complex Movements , 2014, Neuron.

[64]  Martin J. Wainwright,et al.  Scale Mixtures of Gaussians and the Statistics of Natural Images , 1999, NIPS.

[65]  J. Maunsell,et al.  Differences in Gamma Frequencies across Visual Cortex Restrict Their Possible Use in Computation , 2010, Neuron.

[66]  Brendon O. Watson,et al.  Brain rhythms and neural syntax: implications for efficient coding of cognitive content and neuropsychiatric disease. , 2012, Dialogues in clinical neuroscience.

[67]  Konrad Paul Kording,et al.  Bayesian integration in sensorimotor learning , 2004, Nature.

[68]  G. Buzsáki,et al.  Preconfigured, skewed distribution of firing rates in the hippocampus and entorhinal cortex. , 2013, Cell reports.

[69]  Edoardo Milotti,et al.  1/f noise: a pedagogical review , 2002, physics/0204033.

[70]  P. Dayan,et al.  Synapses with short-term plasticity are optimal estimators of presynaptic membrane potentials , 2010, Nature Neuroscience.

[71]  R. Reid,et al.  Specificity of monosynaptic connections from thalamus to visual cortex , 1995, Nature.

[72]  J. Tenenbaum,et al.  Theory-based Bayesian models of inductive learning and reasoning , 2006, Trends in Cognitive Sciences.

[73]  Alex M Thomson,et al.  Binomial parameters differ across neocortical layers and with different classes of connections in adult rat and cat neocortex , 2007, Proceedings of the National Academy of Sciences.

[74]  John P. Cunningham,et al.  Empirical models of spiking in neural populations , 2011, NIPS.

[75]  Timothy D. Hanks,et al.  Probabilistic Population Codes for Bayesian Decision Making , 2008, Neuron.

[76]  Wolfgang Maass,et al.  Neural Dynamics as Sampling: A Model for Stochastic Computation in Recurrent Networks of Spiking Neurons , 2011, PLoS Comput. Biol..

[77]  Richard E. Turner,et al.  A Structured Model of Video Reproduces Primary Visual Cortical Organisation , 2009, PLoS Comput. Biol..

[78]  A. Watson A formula for human retinal ganglion cell receptive field density as a function of visual field location. , 2014, Journal of vision.

[79]  R. Tweedie,et al.  Exponential convergence of Langevin distributions and their discrete approximations , 1996 .

[80]  Eero P. Simoncelli,et al.  Natural signal statistics and sensory gain control , 2001, Nature Neuroscience.

[81]  R. Jacobs,et al.  Optimal integration of texture and motion cues to depth , 1999, Vision Research.

[82]  D. Rubin,et al.  Maximum likelihood from incomplete data via the EM - algorithm plus discussions on the paper , 1977 .

[83]  Michael Okun,et al.  Instantaneous correlation of excitation and inhibition during ongoing and sensory-evoked activities , 2008, Nature Neuroscience.