A Computational Analysis of the Function of Three Inhibitory Cell Types in Contextual Visual Processing

Most cortical inhibitory cell types exclusively express one of three genes, parvalbumin, somatostatin and 5HT3a. The visual responses of cortical neurons are affected not only by local cues, but also by visual context. As the inhibitory neuron types have distinctive synaptic sources and targets over different spatial extents and from different areas, we conjecture that they possess distinct roles in contextual processing. We use modeling to relate structural information to function in primary visual cortex (V1) of the mouse, and investigate their role in contextual visual processing. Our findings are threefold. First, the inhibition mediated by parvalbumin positive (PV) cells mediates local processing and could underlie their role in boundary detection. Second, the inhibition mediated by somatostatin-positive (SST) cells facilitates longer range spatial competition among receptive fields. Third, non-specific top-down modulation to interneurons expressing vasoactive intestinal polypeptide (VIP), a subclass of 5HT3a neurons, can selectively enhance V1 responses.Comment: 39 pages, 5 figures, 4 supplemental figures, 2 table

Solving constraint-satisfaction problems with distributed neocortical-like neuronal networks

Finding actions that satisfy the constraints imposed by both external inputs and internal representations is central to decision making. We demonstrate that some important classes of constraint satisfaction problems (CSPs) can be solved by networks composed of homogeneous cooperative-competitive modules that have connectivity similar to motifs observed in the superficial layers of neocortex. The winner-take-all modules are sparsely coupled by programming neurons that embed the constraints onto the otherwise homogeneous modular computational substrate. We show rules that embed any instance of the CSPs planar four-color graph coloring, maximum independent set, and Sudoku on this substrate, and provide mathematical proofs that guarantee these graph coloring problems will convergence to a solution. The network is composed of non-saturating linear threshold neurons. Their lack of right saturation allows the overall network to explore the problem space driven through the unstable dynamics generated by recurrent excitation. The direction of exploration is steered by the constraint neurons. While many problems can be solved using only linear inhibitory constraints, network performance on hard problems benefits significantly when these negative constraints are implemented by non-linear multiplicative inhibition. Overall, our results demonstrate the importance of instability rather than stability in network computation, and also offer insight into the computational role of dual inhibitory mechanisms in neural circuits.Comment: Accepted manuscript, in press, Neural Computation (2018

Context-Sensitive Binding by the Laminar Circuits of V1 and V2: A Unified Model of Perceptual Grouping, Attention, and Orientation Contrast

A detailed neural model is presented of how the laminar circuits of visual cortical areas V1 and V2 implement context-sensitive binding processes such as perceptual grouping and attention. The model proposes how specific laminar circuits allow the responses of visual cortical neurons to be determined not only by the stimuli within their classical receptive fields, but also to be strongly influenced by stimuli in the extra-classical surround. This context-sensitive visual processing can greatly enhance the analysis of visual scenes, especially those containing targets that are low contrast, partially occluded, or crowded by distractors. We show how interactions of feedforward, feedback and horizontal circuitry can implement several types of contextual processing simultaneously, using shared laminar circuits. In particular, we present computer simulations which suggest how top-down attention and preattentive perceptual grouping, two processes that are fundamental for visual binding, can interact, with attentional enhancement selectively propagating along groupings of both real and illusory contours, thereby showing how attention can selectively enhance object representations. These simulations also illustrate how attention may have a stronger facilitatory effect on low contrast than on high contrast stimuli, and how pop-out from orientation contrast may occur. The specific functional roles which the model proposes for the cortical layers allow several testable neurophysiological predictions to be made. The results presented here simulate only the boundary grouping system of adult cortical architecture. However we also discuss how this model contributes to a larger neural theory of vision which suggests how intracortical and intercortical feedback help to stabilize development and learning within these cortical circuits. Although feedback plays a key role, fast feedforward processing is possible in response to unambiguous information. Model circuits are capable of synchronizing quickly, but context-sensitive persistence of previous events can influence how synchrony develops. Although these results focus on how the interblob cortical processing stream controls boundary grouping and attention, related modeling of the blob cortical processing stream suggests how visible surfaces are formed, and modeling of the motion stream suggests how transient responses to scenic changes can control long-range apparent motion and also attract spatial attention.Defense Advanced Research Projects agency and the Office of Naval Research (N00014-95-1-0409); National Science Foundation (IRI 94-01659, IRI 97-20333); ONR (N00014-92-J-1309, N00014-95-1-0657

Towards a Theory of the Laminar Architecture of Cerebral Cortex: Computational Clues from the Visual System

One of the most exciting and open research frontiers in neuroscience is that of seeking to understand the functional roles of the layers of cerebral cortex. New experimental techniques for probing the laminar circuitry of cortex have recently been developed, opening up novel opportunities for investigating ho1v its six-layered architecture contributes to perception and cognition. The task of trying to interpret this complex structure can be facilitated by theoretical analyses of the types of computations that cortex is carrying out, and of how these might be implemented in specific cortical circuits. We have recently developed a detailed neural model of how the parvocellular stream of the visual cortex utilizes its feedforward, feedback, and horizontal interactions for purposes of visual filtering, attention, and perceptual grouping. This model, called LAMINART, shows how these perceptual processes relate to the mechanisms which ensure stable development of cortical circuits in the infant, and to the continued stability of learning in the adult. The present article reviews this laminar theory of visual cortex, considers how it may be generalized towards a more comprehensive theory that encompasses other cortical areas and cognitive processes, and shows how its laminar framework generates a variety of testable predictions.Defense Advanced Research Projects Agency and the Office of Naval Research (N00014-95-0409); National Science Foundation (IRI 94-01659); Office of Naval Research (N00014-92-1-1309, N00014-95-1-0657

A roadmap to integrate astrocytes into Systems Neuroscience.

Systems neuroscience is still mainly a neuronal field, despite the plethora of evidence supporting the fact that astrocytes modulate local neural circuits, networks, and complex behaviors. In this article, we sought to identify which types of studies are necessary to establish whether astrocytes, beyond their well-documented homeostatic and metabolic functions, perform computations implementing mathematical algorithms that sub-serve coding and higher-brain functions. First, we reviewed Systems-like studies that include astrocytes in order to identify computational operations that these cells may perform, using Ca2+ transients as their encoding language. The analysis suggests that astrocytes may carry out canonical computations in a time scale of subseconds to seconds in sensory processing, neuromodulation, brain state, memory formation, fear, and complex homeostatic reflexes. Next, we propose a list of actions to gain insight into the outstanding question of which variables are encoded by such computations. The application of statistical analyses based on machine learning, such as dimensionality reduction and decoding in the context of complex behaviors, combined with connectomics of astrocyte-neuronal circuits, is, in our view, fundamental undertakings. We also discuss technical and analytical approaches to study neuronal and astrocytic populations simultaneously, and the inclusion of astrocytes in advanced modeling of neural circuits, as well as in theories currently under exploration such as predictive coding and energy-efficient coding. Clarifying the relationship between astrocytic Ca2+ and brain coding may represent a leap forward toward novel approaches in the study of astrocytes in health and disease

Multiscale sampling model for motion integration

Biologically plausible strategies for visual scene integration across spatial and temporal domains continues to be a challenging topic. The fundamental question we address is whether classical problems in motion integration, such as the aperture problem, can be solved in a model that samples the visual scene at multiple spatial and temporal scales in parallel. We hypothesize that fast interareal connections that allow feedback of information between cortical layers are the key processes that disambiguate motion direction. We developed a neural model showing how the aperture problem can be solved using different spatial sampling scales between LGN, V1 layer 4, V1 layer 6, and area MT. Our results suggest that multiscale sampling, rather than feedback explicitly, is the key process that gives rise to end-stopped cells in V1 and enables area MT to solve the aperture problem without the need for calculating intersecting constraints or crafting intricate patterns of spatiotemporal receptive fields. Furthermore, the model explains why end-stopped cells no longer emerge in the absence of V1 layer 6 activity (Bolz & Gilbert, 1986), why V1 layer 4 cells are significantly more end-stopped than V1 layer 6 cells (Pack, Livingstone, Duffy, & Born, 2003), and how it is possible to have a solution to the aperture problem in area MT with no solution in V1 in the presence of driving feedback. In summary, while much research in the field focuses on how a laminar architecture can give rise to complicated spatiotemporal receptive fields to solve problems in the motion domain, we show that one can reframe motion integration as an emergent property of multiscale sampling achieved concurrently within lamina and across multiple visual areas.This work was supported in part by CELEST, a National Science Foundation Science of Learning Center; NSF SBE-0354378 and OMA-0835976; ONR (N00014-11-1-0535); and AFOSR (FA9550-12-1-0436). (CELEST, a National Science Foundation Science of Learning Center; SBE-0354378 - NSF; OMA-0835976 - NSF; N00014-11-1-0535 - ONR; FA9550-12-1-0436 - AFOSR)Published versio