Linking Visual Cortex to Visual Perception: An Alternative to the Gestalt Bubble

Lehar's lively discussion builds on a critique of neural models of vision that is incorrect in its general and specific claims. He espouses a Gestalt perceptual approach, rather than one consistent with the "objective neurophysiological state of the visual system" (p. 1). Contemporary vision models realize his perceptual goals and also quantitatively explain neurophysiological and anatomical data.

Synchronized Oscillations During Cooperative Feature Lining in a Cortical Model of Visual Perception

A neural network model of synchronized oscillations in visual cortex is presented to account for recent neurophysiological findings that such synchronization may reflect global properties of the stimulus. In these experiments, synchronization of oscillatory firing responses to moving bar stimuli occurred not only for nearby neurons, but also occurred between neurons separated by several cortical columns (several mm of cortex) when these neurons shared some receptive field preferences specific to the stimuli. These results were obtained for single bar stimuli and also across two disconnected, but colinear, bars moving in the same direction. Our model and computer simulations obtain these synchrony results across both single and double bar stimuli using different, but formally related, models of preattentive visual boundary segmentation and attentive visual object recognition, as well as nearest-neighbor and randomly coupled models.

Synchronized Oscillations During Cooperative Feature Linking in a Cortical Model of Visual Perception

A neural network model of synchronized oscillator activity in visual cortex is presented in order to account for recent neurophysiological findings that such synchronization may reflect global properties of the stimulus. In these recent experiments, it was reported that synchronization of oscillatory firing responses to moving bar stimuli occurred not only for nearby neurons, but also occurred between neurons separated by several cortical columns (several mm of cortex) when these neurons shared some receptive field preferences specific to the stimuli. These results were obtained not only for single bar stimuli but also across two disconnected, but colinear, bars moving in the same direction. Our model and computer simulations obtain these synchrony results across both single and double bar stimuli. For the double bar case, synchronous oscillations are induced in the region between the bars, but no oscillations are induced in the regions beyond the stimuli. These results were achieved with cellular units that exhibit limit cycle oscillations for a robust range of input values, but which approach an equilibrium state when undriven. Single and double bar synchronization of these oscillators was achieved by different, but formally related, models of preattentive visual boundary segmentation and attentive visual object recognition, as well as nearest-neighbor and randomly coupled models. In preattentive visual segmentation, synchronous oscillations may reflect the binding of local feature detectors into a globally coherent grouping. In object recognition, synchronous oscillations may occur during an attentive resonant state that triggers new learning. These modelling results support earlier theoretical predictions of synchronous visual cortical oscillations and demonstrate the robustness of the mechanisms capable of generating synchrony.

Temporal Dynamics of Decision-Making during Motion Perception in the Visual Cortex

How does the brain make decisions? Speed and accuracy of perceptual decisions covary with certainty in the input, and correlate with the rate of evidence accumulation in parietal and frontal cortical "decision neurons." A biophysically realistic model of interactions within and between Retina/LGN and cortical areas V1, MT, MST, and LIP, gated by basal ganglia, simulates dynamic properties of decision-making in response to ambiguous visual motion stimuli used by Newsome, Shadlen, and colleagues in their neurophysiological experiments. The model clarifies how brain circuits that solve the aperture problem interact with a recurrent competitive network with self-normalizing choice properties to carry out probablistic decisions in real time. Some scientists claim that perception and decision-making can be described using Bayesian inference or related general statistical ideas, that estimate the optimal interpretation of the stimulus given priors and likelihoods. However, such concepts do not propose the neocortical mechanisms that enable perception, and make decisions. The present model explains behavioral and neurophysiological decision-making data without an appeal to Bayesian concepts and, unlike other existing models of these data, generates perceptual representations and choice dynamics in response to the experimental visual stimuli. Quantitative model simulations include the time course of LIP neuronal dynamics, as well as behavioral accuracy and reaction time properties, during both correct and error trials at different levels of input ambiguity in both fixed duration and reaction time tasks. Model MT/MST interactions compute the global direction of random dot motion stimuli, while model LIP computes the stochastic perceptual decision that leads to a saccadic eye movement.

How Does the Cerebral Cortex Work? Developement, Learning, Attention, and 3D Vision by Laminar Circuits of Visual Cortex

A key goal of behavioral and cognitive neuroscience is to link brain mechanisms to behavioral functions. The present article describes recent progress towards explaining how the visual cortex sees. Visual cortex, like many parts of perceptual and cognitive neocortex, is organized into six main layers of cells, as well as characteristic sub-lamina. Here it is proposed how these layered circuits help to realize the processes of developement, learning, perceptual grouping, attention, and 3D vision through a combination of bottom-up, horizontal, and top-down interactions. A key theme is that the mechanisms which enable developement and learning to occur in a stable way imply properties of adult behavior. These results thus begin to unify three fields: infant cortical developement, adult cortical neurophysiology and anatomy, and adult visual perception. The identified cortical mechanisms promise to generalize to explain how other perceptual and cognitive processes work.

Fast Synchronization of Perpetual Grouping in Laminar Visual Cortical Circuits

Perceptual grouping is well-known to be a fundamental process during visual perception, notably grouping across scenic regions that do not receive contrastive visual inputs. Illusory contours are a classical example of such groupings. Recent psychophysical and neurophysiological evidence have shown that the grouping process can facilitate rapid synchronization of the cells that are bound together by a grouping, even when the grouping must be completed across regions that receive no contrastive inputs. Synchronous grouping can hereby bind together different object parts that may have become desynchronized due to a variety of factors, and can enhance the efficiency of cortical transmission. Neural models of perceptual grouping have clarified how such fast synchronization may occur by using bipole grouping cells, whose predicted properties have been supported by psychophysical, anatomical, and neurophysiological experiments. These models have not, however, incorporated some of the realistic constraints on which groupings in the brain are conditioned, notably the measured spatial extent of long-range interactions in layer 2/3 of a grouping network, and realistic synaptic and axonal signaling delays within and across cells in different cortical layers. This work addresses the question: Can long-range interactions that obey the bipole constraint achieve fast synchronization under realistic anatomical and neurophysiological constraints that initially desynchronize grouping signals? Can the cells that synchronize retain their analog sensitivity to changing input amplitudes? Can the grouping process complete and synchronize illusory contours across gaps in bottom-up inputs? Our simulations show that the answer to these questions is Yes.

Spiking Dynamics during Perceptual Grouping in the Laminar Circuits of Visual Cortex

Grouping of collinear boundary contours is a fundamental process during visual perception. Illusory contour completion vividly illustrates how stable perceptual boundaries interpolate between pairs of contour inducers, but do not extrapolate from a single inducer. Neural models have simulated how perceptual grouping occurs in laminar visual cortical circuits. These models predicted the existence of grouping cells that obey a bipole property whereby grouping can occur inwardly between pairs or greater numbers of similarly oriented and co-axial inducers, but not outwardly from individual inducers. These models have not, however, incorporated spiking dynamics. Perceptual grouping is a challenge for spiking cells because its properties of collinear facilitation and analog sensitivity to inducer configurations occur despite irregularities in spike timing across all the interacting cells. Other models have demonstrated spiking dynamics in laminar neocortical circuits, but not how perceptual grouping occurs. The current model begins to unify these two modeling streams by implementing a laminar cortical network of spiking cells whose intracellular temporal dynamics interact with recurrent intercellular spiking interactions to quantitatively simulate data from neurophysiological experiments about perceptual grouping, the structure of non-classical visual receptive fields, and gamma oscillations.

Linking Attention to Learning, Expectation, Competition, and Consciousness

The concept of attention has been used in many senses, often without clarifying how or why attention works as it does. Attention, like consciousness, is often described in a disembodied way. The present article summarizes neural models and supportive data and how attention is linked to processes of learning, expectation, competition, and consciousness. A key them is that attention modulates cortical self-organization and stability. Perceptual and cognitive neocortex is organized into six main cell layers, with characteristic sub-lamina. Attention is part of unified design of bottom-up, horizontal, and top-down interactions among indentified cells in laminar cortical circuits. Neural models clarify how attention may be allocated during processes of visual perception, learning and search; auditory streaming and speech perception; movement target selection during sensory-motor control; mental imagery and fantasy; and hallucination during mental disorders, among other processes.

The Art of Seeing and Painting

The human urge to represent the three-dimensional world using two-dimensional pictorial representations dates back at least to Paleolithic times. Artists from ancient to modern times have struggled to understand how a few contours or color patches on a flat surface can induce mental representations of a three-dimensional scene. This article summarizes some of the recent breakthroughs in scientifically understanding how the brain sees that shed light on these struggles. These breakthroughs illustrate how various artists have intuitively understand paradoxical properties about how the brain sees, and have used that understanding to create great art. These paradoxical properties arise from how the brain forms the units of conscious visual perception; namely, representations of three-dimensional boundaries and surfaces. Boundaries and surfaces are computed in parallel cortical processing streams that obey computationally complementary properties. These streams interact at multiple levels to overcome their complementary weaknesses and to transform their complementary properties into consistent percepts. The article describes how properties of complementary consistency have guided the creation of many great works of art.

A Solution of the Figure-ground Problem for Biological Vision

A neural network model of 3-D visual perception and figure-ground separation by visual cortex is introduced. The theory provides a unified explanation of how a 2-D image may generate a 3-D percept; how figures pop-out from cluttered backgrounds; how spatially sparse disparity cues can generate continuous surface representations at different perceived depths; how representations of occluded regions can be completed and recognized without usually being seen; how occluded regions can sometimes be seen during percepts of transparency; how high spatial frequency parts of an image may appear closer than low spatial frequency parts; how sharp targets are detected better against a figure and blurred targets are detector better against a background; how low spatial frequency parts of an image may be fused while high spatial frequency parts are rivalrous; how sparse blue cones can generate vivid blue surface percepts; how 3-D neon color spreading, visual phantoms, and tissue contrast percepts are generated; how conjunctions of color-and-depth may rapidly pop-out during visual search. These explanations arise derived from an ecological analysis of how monocularly viewed parts of an image inherit the appropriate depth from contiguous binocularly viewed parts, as during DaVinci stereopsis. The model predicts the functional role and ordering of multiple interactions within and between the two parvocellular processing streams that join LGN to prestriate area V4. Interactions from cells representing larger scales and disparities to cells representing smaller scales and disparities are of particular importance.

Laminar Cortical Dynamics of Visual Form and Motion Interactions During Coherent Object Motion Perception

How do visual form and motion processes cooperate to compute object motion when each process separately is insufficient? A 3D FORMOTION model specifies how 3D boundary representations, which separate figures from backgrounds within cortical area V2, capture motion signals at the appropriate depths in MT; how motion signals in MT disambiguate boundaries in V2 via MT-to-Vl-to-V2 feedback; how sparse feature tracking signals are amplified; and how a spatially anisotropic motion grouping process propagates across perceptual space via MT-MST feedback to integrate feature-tracking and ambiguous motion signals to determine a global object motion percept. Simulated data include: the degree of motion coherence of rotating shapes observed through apertures, the coherent vs. element motion percepts separated in depth during the chopsticks illusion, and the rigid vs. non-rigid appearance of rotating ellipses.

Laminar Cortical Dynamics of Visual Form and Motion Interactions During Coherent Object Motion Perception

How do visual form and motion processes cooperate to compute object motion when each process separately is insufficient? Consider, for example, a deer moving behind a bush. Here the partially occluded fragments of motion signals available to an observer must be coherently grouped into the motion of a single object. A 3D FORMOTION model comprises five important functional interactions involving the brain’s form and motion systems that address such situations. Because the model’s stages are analogous to areas of the primate visual system, we refer to the stages by corresponding anatomical names. In one of these functional interactions, 3D boundary representations, in which figures are separated from their backgrounds, are formed in cortical area V2. These depth-selective V2 boundaries select motion signals at the appropriate depths in MT via V2-to-MT signals. In another, motion signals in MT disambiguate locally incomplete or ambiguous boundary signals in V2 via MT-to-V1-to-V2 feedback. The third functional property concerns resolution of the aperture problem along straight moving contours by propagating the influence of unambiguous motion signals generated at contour terminators or corners. Here, sparse “feature tracking signals” from, e.g., line ends, are amplified to overwhelm numerically superior ambiguous motion signals along line segment interiors. In the fourth, a spatially anisotropic motion grouping process takes place across perceptual space via MT-MST feedback to integrate veridical feature-tracking and ambiguous motion signals to determine a global object motion percept. The fifth property uses the MT-MST feedback loop to convey an attentional priming signal from higher brain areas back to V1 and V2. The model's use of mechanisms such as divisive normalization, endstopping, cross-orientation inhibition, and longrange cooperation is described. Simulated data include: the degree of motion coherence of rotating shapes observed through apertures, the coherent vs. element motion percepts separated in depth during the chopsticks illusion, and the rigid vs. non-rigid appearance of rotating ellipses.

Cortical Dynamics of Visual Motion Perception: Short-Range and Long Range Apparent Motion

This article describes further evidence for a new neural network theory of biological motion perception that is called a Motion Boundary Contour System. This theory clarifies why parallel streams Vl-> V2 and Vl-> MT exist for static form and motion form processing among the areas Vl, V2, and MT of visual cortex. The Motion Boundary Contour System consists of several parallel copies, such that each copy is activated by a different range of receptive field sizes. Each copy is further subdivided into two hierarchically organized subsystems: a Motion Oriented Contrast Filter, or MOC Filter, for preprocessing moving images; and a Cooperative-Competitive Feedback Loop, or CC Loop, for generating emergent boundary segmentations of the filtered signals. The present article uses the MOC Filter to explain a variety of classical and recent data about short-range and long-range apparent motion percepts that have not yet been explained by alternative models. These data include split motion; reverse-contrast gamma motion; delta motion; visual inertia; group motion in response to a reverse-contrast Ternus display at short interstimulus intervals; speed-up of motion velocity as interfiash distance increases or flash duration decreases; dependence of the transition from element motion to group motion on stimulus duration and size; various classical dependencies between flash duration, spatial separation, interstimulus interval, and motion threshold known as Korte's Laws; and dependence of motion strength on stimulus orientation and spatial frequency. These results supplement earlier explanations by the model of apparent motion data that other models have not explained; a recent proposed solution of the global aperture problem, including explanations of motion capture and induced motion; an explanation of how parallel cortical systems for static form perception and motion form perception may develop, including a demonstration that these parallel systems are variations on a common cortical design; an explanation of why the geometries of static form and motion form differ, in particular why opposite orientations differ by 90°, whereas opposite directions differ by 180°, and why a cortical stream Vl -> V2 -> MT is needed; and a summary of how the main properties of other motion perception models can be assimilated into different parts of the Motion Boundary Contour System design.

Cortical Dynamics of Visual Motion Perception: Short-Range and Long-Range Apparent Motion

This article describes further evidence for a new neural network theory of biological motion perception. The theory clarifies why parallel streams Vl --> V2, Vl --> MT, and Vl --> V2 --> MT exist for static form and motion form processing among the areas Vl, V2, and MT of visual cortex. The theory suggests that the static form system (Static BCS) generates emergent boundary segmentations whose outputs are insensitive to direction-ofcontrast and insensitive to direction-of-motion, whereas the motion form system (Motion BCS) generates emergent boundary segmentations whose outputs are insensitive to directionof-contrast but sensitive to direction-of-motion. The theory is used to explain classical and recent data about short-range and long-range apparent motion percepts that have not yet been explained by alternative models. These data include beta motion; split motion; gamma motion and reverse-contrast gamma motion; delta motion; visual inertia; the transition from group motion to element motion in response to a Ternus display as the interstimulus interval (ISI) decreases; group motion in response to a reverse-contrast Ternus display even at short ISIs; speed-up of motion velocity as interflash distance increases or flash duration decreases; dependence of the transition from element motion to group motion on stimulus duration and size; various classical dependencies between flash duration, spatial separation, ISI, and motion threshold known as Korte's Laws; dependence of motion strength on stimulus orientation and spatial frequency; short-range and long-range form-color interactions; and binocular interactions of flashes to different eyes.

A Laminar Cortical Model for 3D Perception of Slanted and Curved Surfaces and of 2D Images: Developement, attention, and Bistability

A model of laminar visual cortical dynamics proposes how 3D boundary and surface representations of slated and curved 3D objects and 2D images arise. The 3D boundary representations emerge from interactions between non-classical horizontal receptive field interactions with intracorticcal and intercortical feedback circuits. Such non-classical interactions contextually disambiguate classical receptive field responses to ambiguous visual cues using cells that are sensitive to angles and disparity gradients with cortical areas V1 and V2. These cells are all variants of bipole grouping cells. Model simulations show how horizontal connections can develop selectively to angles, how slanted surfaces can activate 3D boundary representations that are sensitive to angles and disparity gradients, how 3D filling-in occurs across slanted surfaces, how a 2D Necker cube image can be represented in 3D, and how bistable Necker cuber percepts occur. The model also explains data about slant aftereffects and 3D neon color spreading. It shows how habituative transmitters that help to control developement also help to trigger bistable 3D percepts and slant aftereffects, and how attention can influence which of these percepts is perceived by propogating along some object boundaries.