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Druckmann, S.

Publications and source records attributed to Druckmann, S..

27 records · Page 2Linked to original sources

Towards a More General Understanding of the Algorithmic Utility of Recurrent Connections

Lateral and recurrent connections are ubiquitous in biological neural circuits. Yet while the strong computational abilities of feedforward networks have been extensively studied, our understanding of the role and advantages of recurrent computations that might explain their prevalence remains an important open challenge. Foundational studies by Minsky and Roelfsema argued that computations that require propagation of global information for local computation to take place would likely particularly benefit from the sequential, parallel nature of processing in recurrent networks. Such "tag propagation" algorithms perform repeated, local propagation of information and were originally introduced in the context of detecting connectedness, a task that is challenging for feedforward networks. Here, we advance the understanding of the utility of lateral and recurrent computation by first performing a large-scale empirical study of neural architectures for the computation of connectedness to explore feedforward solutions more fully and establish robustly the importance of recurrent architectures. In addition, we highlight a tradeoff between computation time and performance and demonstrate hybrid feedforward/recurrent models that perform well even in the presence of varying computational time limitations. We then generalize tag propagation architectures to multiple, interacting propagating tags, and demonstrate that these are efficient computational substrates for more general computations of connectedness by introducing and solving an abstracted biologically inspired decision-making task. Our work thus clarifies and expands the set of computational tasks that can be solved efficiently by recurrent computation, yielding hypotheses for structure in population activity that may be present in such tasks. Author SummaryIn striking contrast to the majority of current-day artificial neural network research which primarily uses feedforward architectures, biological brains make extensive use of lateral and recurrent connections, raising the possibility that this difference makes a fundamental contribution to the gap in computational power between real neural circuits and artificial neural networks. Thus, despite the challenge of making effective comparisons between different network architectures, developing a more detailed understanding of the computational role played by such connections is a pressing need. Here, we leverage the computational capabilities of large-scale machine learning to robustly explore how differences in architectures affect a networks ability to learn tasks that require propagation of global information. We first focus on the task of determining whether two pixels are connected in an image which has an elegant and efficient recurrent solution: propagate a connected label or tag along paths. Inspired by this solution, we show that it can be generalized in many ways, including propagating multiple tags at once and changing the computation performed on the result of the propagation. Strikingly, this simple expansion of the tag propagation network is sufficient to solve a crucial abstraction to temporal connectedness at the core of many decision-making problems, which we illustrate for a an abstracted competitive foraging task Our results shed light on the set of computational tasks that can be solved efficiently by recurrent computation and how these solutions may relate to the structure of neural activity.

neuroscience↗

Transforming representations of movement from body- to world-centric space

When an animal moves through the world, its brain receives a stream of information about the bodys translational movement. These incoming movement signals, relayed from sensory organs or as copies of motor commands, are referenced relative to the body. Ultimately, such body-centric movement signals must be transformed into world-centric coordinates for navigation1. Here we show that this computation occurs in the fan-shaped body in the Drosophila brain. We identify two cell types in the fan-shaped body, PFNd and PFNv2,3, that conjunctively encode translational velocity signals and heading signals in walking flies. Specifically, PFNd and PFNv neurons form a Cartesian representation of body-centric translational velocity - acquired from premotor brain regions4,5 - that is layered onto a world-centric heading representation inherited from upstream compass neurons6-8. Then, we demonstrate that the next network layer, comprising h{Delta}B neurons, is wired so as to transform the representation of translational velocity from body-centric to world-centric coordinates. We show that this transformation is predicted by a computational model derived directly from electron microscopy connectomic data9. The model illustrates the key role of a specific network motif, whereby the PFN neurons that synapse onto the same h{Delta}B neuron have heading-tuning differences that offset the differences in their preferred body-centric directions of movement. By integrating a world-centric representation of travel velocity over time, it should be possible for the brain to form a working memory of the path traveled through the environment10-12.

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Object recognition under occlusion revisited: elucidating algorithmic advantages of recurrent computation

Despite the ubiquity of recurrent connections in the brain, their role in visual processing is less understood than that of feedforward connections. Occluded object recognition, an ethologically critical cognitive capacity, is thought to rely on recurrent processing of visual information, but it remains unclear whether and how recurrent processing improves recognition of occluded objects. Using convolutional models of the visual system, we demonstrate how a distinct form of computation arises in recurrent-but not feedforward- networks that leverages information about the occluder to "explain-away" the occlusion-- i.e., recognition of the occluder provides an account for missing or altered features, potentially rescuing recognition of occluded objects. This occurs without any constraint placed on the computation and is observed both across a systematic architecture sweep of convolutional models and in a model explicitly constructed to approximate the primate visual system. Building on these computational results, we conduct a behavioral experiment to study explaining-away in humans, and find evidence consistent with explaining-away. Finally, to demonstrate a specific mechanism for explaining-away, we develop an experimentally inspired recurrent model that recovers fine-grained features of occluded stimuli by explaining-away. Recurrent connections capability to explain away may extend to more general cases where undoing context-dependent changes in representations could benefit perception.

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Decoding spoken English phonemes from intracortical electrode arrays in dorsal precentral gyrus

ObjectiveTo evaluate the potential of intracortical electrode array signals for brain-computer interfaces (BCIs) to restore lost speech, we measured the performance of classifiers trained to discriminate a comprehensive basis set for speech: 39 English phonemes. We classified neural correlates of spoken-out-loud words in the "hand knob" area of precentral gyrus, which we view as a step towards the eventual goal of decoding attempted speech from ventral speech areas in patients who are unable to speak. ApproachNeural and audio data were recorded while two BrainGate2 pilot clinical trial participants, each with two chronically-implanted 96-electrode arrays, spoke 420 different words that broadly sampled English phonemes. Phoneme onsets were identified from audio recordings, and their identities were then classified from neural features consisting of each electrodes binned action potential counts or high-frequency local field potential power. We also examined two potential confounds specific to decoding overt speech: acoustic contamination of neural signals and systematic differences in labeling different phonemes onset times. Main resultsA linear decoder achieved up to 29.3% classification accuracy (chance = 6%) across 39 phonemes, while a recurrent neural network classifier achieved 33.9% accuracy. Parameter sweeps indicated that performance did not saturate when adding more electrodes or more training data, and that accuracy improved when utilizing time-varying structure in the data. Microphonic contamination and phoneme onset differences modestly increased decoding accuracy, but could be mitigated by acoustic artifact subtraction and using a neural speech onset marker, respectively. SignificanceThe ability to decode a comprehensive set of phonemes using intracortical electrode array signals from a nontraditional speech area suggests that placing electrode arrays in ventral speech areas is a promising direction for speech BCIs.

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A comparison of neuronal population dynamics measured with calcium imaging and electrophysiology

Calcium imaging with fluorescent protein sensors is widely used to record activity in neuronal populations. The transform between neural activity and calcium-related fluorescence involves nonlinearities and a low-pass filter, but the effects of the transformation on analyses of neural populations are not well understood. We compared neuronal spikes and fluorescence in matched neural populations in behaving mice. We report multiple discrepancies between analyses performed on the two types of data, which were only partially resolved by spike inference algorithms applied to fluorescence. To model the relation between spiking and fluorescence we simultaneously recorded spikes and fluorescence from individual neurons. Using these recordings we developed a model transforming spike trains to synthetic-imaging data. The model recapitulated the differences in analyses. Our analysis highlights challenges in relating electrophysiology and imaging data, and suggests forward modeling as an effective way to understand differences between these data.

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Targeted photostimulation uncovers circuit motifs supporting short-term memory

Short-term memory is associated with persistent neural activity without sustained input, arising from the interactions between neurons with short time constants1,2. A variety of neural circuit motifs could account for measured neural activity3-7. A mechanistic understanding of the neural circuits supporting short-term memory requires probing network connectivity between functionally characterized neurons8. We performed targeted photostimulation of small (< 10) groups of neurons, while imaging the response of hundreds of other neurons9,10, in anterior-lateral motor cortex (ALM) of mice performing a delayed response task11. Mice were instructed with brief auditory stimuli to make directional movements (lick left or lick right), but only after a three second delay epoch. ALM contains neurons with delay epoch activity that is selective for left or right choices. Targeted photostimulation of groups of neurons during the delay epoch allowed us to observe the functional organization of population activity and recurrent interactions underlying short-term memory. These experiments revealed strong coupling between neurons sharing similar selectivity. Brief photostimulation of functionally related neurons produced changes in activity in sparse subpopulations of nearby neurons that persisted for several seconds following stimulus offset, far outlasting the duration of the perturbation. Photostimulation produced behavioral biases that were predictable based on the selectivity of the perturbed neuronal population. These results suggest that ALM contains multiple intercalated modules, consisting of recurrently coupled neurons, that can independently maintain persistent activity.

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An orderly single-trial organization of population dynamics in premotor cortex predicts behavioral variability

Animals are not simple input-output machines. Their responses to even very similar stimuli are variable. A key, long-standing question in neuroscience is understanding the neural correlates of such behavioral variability. To reveal these correlates, behavior and neural population must be related to one another on single trials. Such analysis is challenging due to the dynamical nature of brain function (e.g. decision making), neuronal heterogeneity and signal to noise difficulties. By analyzing population recordings from mouse frontal cortex in perceptual decision-making tasks, we show that an analysis approach tailored to the coarse grain features of the dynamics was able to reveal previously unrecognized structure in the organization of population activity. This structure was similar on error and correct trials, suggesting what may be the underlying circuit mechanisms, was able to predict multiple aspects of behavioral variability and revealed long time-scale modulation of population activity.

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Kilohertz frame-rate two-photon tomography

Point-scanning two-photon microscopy enables high-resolution imaging within scattering specimens such as the mammalian brain, but sequential acquisition of voxels fundamentally limits imaging speed. We developed a two-photon imaging technique that scans lines of excitation across a focal plane at multiple angles and uses prior information to recover high-resolution images at over 1.4 billion voxels per second. Using a structural image as a prior for recording neural activity, we imaged visually-evoked and spontaneous glutamate release across hundreds of dendritic spines in mice at depths over 250 {micro}m and frame-rates over 1 kHz. Dendritic glutamate transients in anaesthetized mice are synchronized within spatially-contiguous domains spanning tens of microns at frequencies ranging from 1-100 Hz. We demonstrate high-speed recording of acetylcholine and calcium sensors, 3D single-particle tracking, and imaging in densely-labeled cortex. Our method surpasses limits on the speed of raster-scanned imaging imposed by fluorescence lifetime.

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Multiplicative Updates for Optimization Problems with Dynamics

We consider the problem of optimizing general convex objective functions with nonnegativity constraints. Using the Karush-Kuhn-Tucker (KKT) conditions for the nonnegativity constraints we will derive fast multiplicative update rules for several problems of interest in signal processing, including non-negative deconvolution, point-process smoothing, ML estimation for Poisson Observations, nonnegative least squares and nonnegative matrix factorization (NMF). Our algorithm can also account for temporal and spatial structure and regularization. We will analyze the performance of our algorithm on simultaneously recorded neuronal calcium imaging and electrophysiology data.

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