Reentrant Connections Research Articles

Punching shear tests in flat slabs supported on re-entrant corner columns

The punching strength and deformation capacity of slab-column connections is governing for design at ultimate limit state of flat slabs. This phenomenon has been investigated since decades with experimental programmes in an effort to derive empirical or mechanical design approaches. Such efforts, due to the complexity of the phenomena, have mostly been addressed to axisymmetric inner slab-column connections, which correspond to the simplest academic case and are not directly covering a large variety of practical situations. Although some tests are also available for other configurations, such as edge or corner connections, no experimental evidence is found on relevant cases for practice such as re-entrant corner columns. The lack of experimental data in this latter case raises questions on the design of such connections, as extrapolating results from other cases is not a simple task due to the complexity of the phenomena implied (such as redistributions in the shear field and moment transfer between slab and column). In order to advance the knowledge on the response of re-entrant connections and to provide a sound design approach for them, this paper presents the results of an investigation specifically addressed at this case. To that aim, seven specimens were tested under different conditions, varying the load eccentricity, flexural reinforcement and shear reinforcement arrangement at the edges. The tests were instrumented with refined measurements, allowing to understand in a detailed manner the response of the slabs during testing in the shear-critical region. The experimental results are later compared to current design codes, leading to a number of conclusions with practical relevance. Finally, on the basis of the knowledge acquired from the tests, a mechanical approach is proposed based on the Critical Shear Crack Theory to assess the punching response. This approach is shown to lead to accurate and reliable predictions of the punching strength, improving those of current design codes.

Increasing associative plasticity in temporo-occipital back-projections improves visual perception of emotions

The posterior superior temporal sulcus (pSTS) is a critical node in a network specialized for perceiving emotional facial expressions that is reciprocally connected with early visual cortices (V1/V2). Current models of perceptual decision-making increasingly assign relevance to recursive processing for visual recognition. However, it is unknown whether inducing plasticity into reentrant connections from pSTS to V1/V2 impacts emotion perception. Using a combination of electrophysiological and neurostimulation methods, we demonstrate that strengthening the connectivity from pSTS to V1/V2 selectively increases the ability to perceive facial expressions associated with emotions. This behavior is associated with increased electrophysiological activity in both these brain regions, particularly in V1/V2, and depends on specific temporal parameters of stimulation that follow Hebbian principles. Therefore, we provide evidence that pSTS-to-V1/V2 back-projections are instrumental to perception of emotion from facial stimuli and functionally malleable via manipulation of associative plasticity.

A blinking focal pattern of re-entrant activity in the avian tectum

Re-entrant connections are inherent to nervous system organization; however, a comprehensive understanding of their operation is still lacking. In birds, topographically organized re-entrant signals, carried by axons from the nucleus-isthmi-parvocellularis (Ipc), are distinctly recorded as bursting discharges across the optic tectum (TeO). Here, we used up to 48 microelectrodes regularly spaced on the superficial tectal layers of anesthetized pigeons to characterize the spatial-temporal pattern of this axonal re-entrant activity in response to different visual stimulation. We found that a brief luminous spot triggered repetitive waves of bursting discharges that, appearing from initial sources, propagated horizontally to areas representing up to 28° of visual space, widely exceeding the area activated by the retinal fibers. In response to visual motion, successive burst waves started along and around the stimulated tectal path, tracking the stimulus in discontinuous steps. When two stimuli were presented, the burst-wave sources alternated between the activated tectal loci, as if only one source could be active at any given time. Because these re-entrant signals boost the retinal input to higher visual areas, their peculiar dynamics mimic a blinking "spotlight," similar to the internal searching mechanism classically used to explain spatial attention. Tectal re-entry from Ipc is thus highly structured and intrinsically discontinuous, and higher tectofugal areas, which lack retinotopic organization, will thus receive incoming visual activity in a sequential and piecemeal fashion. We anticipate that analogous re-entrant patterns, perhaps hidden in less bi-dimensionally organized topographies, may organize the flow of neural activity in other parts of the brain as well.

Intracerebral electrical stimulation to understand the neural basis of human face identity recognition.

Recognizing people's identity by their faces is a key function in the human species, supported by regions of the ventral occipito-temporal cortex (VOTC). In the last decade, there have been several reports of perceptual face distortion during direct electrical stimulation (DES) with subdural electrodes positioned over a well-known face-selective VOTC region of the right lateral middle fusiform gyrus (LatMidFG; i.e., the "Fusiform Face Area", FFA). However, transient impairments of face identity recognition (FIR) have been extremely rare and only behaviorally quantified during DES with intracerebral (i.e., depth) electrodes in stereo-electroencephalography (SEEG). The three detailed cases reported so far, summarized here, were specifically impaired at FIR during DES inside different anatomical VOTC regions of the right hemisphere: the inferior occipital gyrus (IOG) and the LatMidFG, as well as a region that lies at the heart of a large magnetic susceptibility artifact in functional magnetic resonance imaging (fMRI): the anterior fusiform gyrus (AntFG). In the first two regions, the eloquent electrode contacts were systematically associated with the highest face-selective and (unfamiliar) face individuation responses as measured with intracerebral electrophysiology. Stimulation in the right AntFG did not lead to perceptual changes but also caused an inability to remember having been presented face pictures, as if the episode was never recorded in memory. These observations support the view of an extensive network of face-selective VOTC regions subtending human FIR, with at least three critical nodes in the right hemisphere associated with differential intrinsic and extrinsic patterns of reentrant connectivity.

Empowering Reentrant Projections from V5 to V1 Boosts Sensitivity to Motion

Evidence from macaques [1] and humans [2, 3] has shown that back projections from extrastriate areas to the primary visual area (V1) determine whether visual awareness will arise. For example, reentrant projections from the visual motion area (V5) to V1 are considered to be critical for awareness of motion [2, 3]. If these projections are also instrumental to functional processing of moving stimuli [4-8], then increasing synaptic efficacy in V5-V1 connections should induce functionally relevant short-term plastic changes, resulting in enhanced perception of visual motion. Using transcranial magnetic stimulation (TMS), we applied a novel cortico-cortical paired associative stimulation (ccPAS) protocol to transiently enhance visual motion sensitivity and demonstrate both the functional relevance of V5-V1 reentrant projections to motion perception and their plasticity. Specifically, we found that ccPAS aimed at strengthening reentrant connectivity from V5 to V1 (but not in the opposite direction) enhanced the human ability to perceive coherent visual motion. This perceptual enhancement followed the temporal profile of Hebbian plasticity [9-18] and was observed only when an optimal timing of 20ms between TMS pulses [2, 3, 5, 6] was used, not when TMS pulses were delivered synchronously. Thus, plastic change is critically dependent on both the direction and timing of connectivity; if either of these requirements was not met, perceptual enhancement did not take place. We therefore provide novel causal evidence that V5-V1 back projections, instrumental to motion perception, are functionally malleable. These findings have implications for theoretical models of visual awareness and for the rehabilitation of visual deficits.

Misbinding of Color and Motion in Human Visual Cortex

A fundamental challenge for the visual system is to integrate visual features into a coherent scene, known as the binding problem. The neural mechanisms of feature binding are hard to identify because of difficulties in separating active feature binding from feature co-occurrence. In previous studies on feature binding, visual features were superimposed and presented simultaneously. Neurons throughout the visual cortex are known to code multiple features. Therefore, the observed binding effects could be due to the physical co-occurrence of features and the sensory representation of feature pairings. It is uncertain whether the mechanisms responsible for perceptual binding were actually recruited. To address this issue, we performed psychophysical and fMRI experiments to investigate the neural mechanisms of a steady-state misbinding of color and motion, because feature misbinding is probably the most striking evidence for the active existence of the binding mechanisms. We found that adapting to the color-motion misbinding generated the color-contingent motion aftereffect, as well as the color-contingent motion adaptation effect in visual cortex. Notably, V2 exhibited the strongest adaptation effect, which significantly correlated with the aftereffect across subjects. Furthermore, effective connectivity analysis using dynamic causal modeling showed that the misbinding was closely associated with enhanced feedback from V4 and V5 to V2. These findings provide strong evidence for active feature binding in early visual cortex and suggest a critical role of reentrant connections from specialized intermediate areas to early visual cortex in this process.

TMS to the “occipital face area” affects recognition but not categorization of faces

The human cortical system for face perception is comprised of a network of connected regions including the middle fusiform gyrus (“fusiform face area” or FFA), the inferior occipital cortex (“occipital face area” or OFA), and the superior temporal sulcus. The traditional hierarchical feedforward model of visual processing suggests information flows from early visual cortex to the OFA for initial face feature analysis to higher order regions including the FFA for identity recognition. However, patient data suggest an alternative model. Patients with acquired prosopagnosia, an inability to visually recognize faces, have been documented with lesions to the OFA but who nevertheless show face-selective activation in the FFA. Moreover, their ability to categorize faces remains intact. This suggests that the FFA is not solely responsible for face recognition and the network is not strictly hierarchical, but may be organized in a reverse hierarchical fashion. We used transcranial magnetic stimulation (TMS) to temporarily disrupt processing in the OFA in neurologically-intact individuals and found participants’ ability to categorize intact versus scrambled faces was unaffected, however face identity discrimination was significantly impaired. This suggests that face categorization but not recognition can occur without the “earlier” OFA being online and indicates that “lower level” face category processing may be assumed by other intact face network regions such as the FFA. These results are consistent with the patient data and support a non-hierarchical, global-to-local model with re-entrant connections between the OFA and other face processing areas.

Tagging cortical networks in emotion: A topographical analysis

Viewing emotional pictures is associated with heightened perception and attention, indexed by a relative increase in visual cortical activity. Visual cortical modulation by emotion is hypothesized to reflect re-entrant connectivity originating in higher-order cortical and/or limbic structures. The present study used dense-array electroencephalography and individual brain anatomy to investigate functional coupling between the visual cortex and other cortical areas during affective picture viewing. Participants viewed pleasant, neutral, and unpleasant pictures that flickered at a rate of 10 Hz to evoke steady-state visual evoked potentials (ssVEPs) in the EEG. The spectral power of ssVEPs was quantified using Fourier transform, and cortical sources were estimated using beamformer spatial filters based on individual structural magnetic resonance images. In addition to lower-tier visual cortex, a network of occipito-temporal and parietal (bilateral precuneus, inferior parietal lobules) structures showed enhanced ssVEP power when participants viewed emotional (either pleasant or unpleasant), compared to neutral pictures. Functional coupling during emotional processing was enhanced between the bilateral occipital poles and a network of temporal (left middle/inferior temporal gyrus), parietal (bilateral parietal lobules), and frontal (left middle/inferior frontal gyrus) structures. These results converge with findings from hemodynamic analyses of emotional picture viewing and suggest that viewing emotionally engaging stimuli is associated with the formation of functional links between visual cortex and the cortical regions underlying attention modulation and preparation for action.

VISUAL FEATURE BINDING WITHIN THE SELECTIVE TUNING ATTENTION FRAMEWORK

We present a biologically plausible computational model for solving the visual feature binding problem, based on recent results regarding the time course and processing sequence in the primate visual system. The feature binding problem appears due to the distributed nature of visual processing in the primate brain, and the gradual loss of spatial information along the processing hierarchy. This paper puts forward the proposal that by using multiple passes of the visual processing hierarchy, both bottom-up and top-down, and using task information to tune the processing prior to each pass, we can explain the different recognition behaviors that primate vision exhibits. To accomplish this, four different kinds of binding processes are introduced and are tied directly to specific recognition tasks and their time course. The model relies on the reentrant connections so ubiquitous in the primate brain to recover spatial information, and thus allow features represented in different parts of the brain to be integrated in a unitary conscious percept. We show how different tasks and stimuli have different binding requirements, and present a unified framework within the Selective Tuning model of visual attention.

Visual binding through reentrant connectivity and dynamic synchronization in a brain-based device.

Effective visual object recognition requires mechanisms to bind object features (e.g. color, shape and motion) while distinguishing distinct objects. Synchronously active neuronal circuits among reentrantly connected cortical areas may provide a basis for visual binding. To assess the potential of this mechanism, we have constructed a mobile brain-based device, Darwin VIII, which is guided by simulated analogues of cortical and sub-cortical areas required for visual processing, decision-making, reward and motor responses. These simulated areas are reentrantly connected and each area contains neuronal units representing both the mean activity level and the relative timing of the activity of groups of neurons. Darwin VIII learns to discriminate among multiple objects with shared visual features and associates 'target' objects with innately preferred auditory cues. We observed the co-activation of globally distributed neuronal circuits that corresponded to distinct objects in Darwin VIII's visual field. These circuits, which are constrained by a reentrant neuroanatomy and modulated by behavior and synaptic plasticity, are necessary for successful discrimination. By situating Darwin VIII in a rich real-world environment involving continual changes in the size and location of visual stimuli due to self-generated movement, and by recording its behavioral and neuronal responses in detail, we were able to show that reentrant connectivity and dynamic synchronization provide an effective mechanism for binding the features of visual objects.

Active Sensing of Visual and Tactile Stimuli by Brain-based Devices

We describe the construction and performance of `brain-based devices? (BBDs), physical devices whose behaviour is controlled by simulated nervous systems modelled on vertebrate neuroanatomy and neurophysiology, that carry out perceptual categorization and selective conditioning to visual and textural stimuli. BBDs take input from the environment through on-board sensors including cameras, microphones and artificial whiskers, and take action based on experiential learning. BBDs have a large-scale neural simulation, a phenotype, a body plan, and the means to learn through autonomous exploration. Key neural mechanisms in the present BBDs include synaptic plasticity, reward or value systems, reentrant connectivity, the dynamic synchronization of neuronal activity, and neuronal units with spatiotemporal response properties. With our BBDs, as with animals, it is the interaction of these neural mechanisms with the sensorimotor correlations generated by active sensing and self motion that is responsible for adaptive behaviour. BBDs permit analysis of activity at all levels of the nervous system during behaviour, and as such they provide a rich source of heuristics for generating hypotheses regarding brain function. Moreover, by taking inspiration from systems neuroscience, BBDs provide a novel architecture for the design of neuromorphic systems.

The neuroanatomy of phenomenal vision: a psychological perspective.

Somewhere in the visual system, phenomenal vision--the seeing of colors, brightness, depths, shades, and motion--is generated not only from the distribution of light on the retina, but also when the eyes are closed, in dreams, hallucinations, phosphenes, and (possibly) imagery. Whether these different forms of phenomenal vision share a common substrate although their origins are different (optical, mechanical, electrical, endogenous) is discussed in the light of evidence from neuropsychological and functional imaging studies. Whereas extrastriate visual cortical areas appear to be involved in all types of phenomenal vision that have been studied, the necessity of a contribution from primary visual cortex is demonstrated by the loss of conscious vision that follows its destruction. If both extrastriate and primary cortical activation are needed, the latter may not just provide an indispensable input, but may also need to receive the output of the extrastriate processing via reentrant connections.

Solutions to the binding problem: progress through controversy and convergence.

Solutions to the binding problem: progress through controversy and convergence.

The temporal correlation hypothesis of visual feature integration: still alive and well.

The temporal correlation hypothesis of visual feature integration: still alive and well.

602 A new approach to assessing drowsiness and attention through continuous EEG (C-EEG) and neural network methods

It has recently been suggested that reentrant connections are essential in systems that process complex information [A. Damasio, H. Damasio, Cortical systems for the retrieval of concrete knowledge: the convergence zone framework, in: C. Koch, J.L. Davis (Eds.), Large Scale Neuronal Theories of the Brain, The MIT Press, Cambridge, 1995, pp. 61–74; G. Edelman, The Remembered Present, Basic Books, New York, 1989; M.I. Posner, M. Rothbart, Constructing neuronal theories of mind, in: C. Koch, J.L. Davis (Eds.), Large Scale Neuronal Theories of the Brain, The MIT Press, Cambridge, 1995, pp. 183–199; C. von der Malsburg, W. Schneider, A neuronal cocktail party processor, Biol. Cybem., 54 (1986) 29–40]. Reentry is not feedback, but parallel signalling in the time domain between spatially distributed maps, similar to a process of correlation between distributed systems. Accordingly, it was expected that during spontaneous reversals of the Necker cube, complex patterns of correlations between distributed systems would be present in the cortex. The present study included EEG (n=4) and MEG recordings (n=5). Two experimental questions were posed: (1) Can distributed cortical patterns present during perceptual reversals be classified differently using a generalised regression neural network (GRNN) compared to processing of a two-dimensional figure? (2) Does correlated cortical activity increase significantly during perception of a Necker cube reversal? One-second duration single trials of EEG and MEG data were analysed using the GRNN. Electrode/sensor pairings based on cortico–cortical connections were selected to assess correlated activity in each condition. The GRNN significantly classified single trials recorded during Necker cube reversals as different from single trials recorded during perception of a two-dimensional figure for both EEG and MEG. In addition, correlated cortical activity increased significantly in the Necker cube reversal condition for EEG and MEG compared to the perception of a non-reversing stimulus. Coherent MEG activity observed over occipital, parietal and temporal regions is believed to represent neural systems related to the perception of Necker cube reversals.

Neural network classifications and correlation analysis of EEG and MEG activity accompanying spontaneous reversals of the Necker cube

It has recently been suggested that reentrant connections are essential in systems that process complex information [A. Damasio, H. Damasio, Cortical systems for the retrieval of concrete knowledge: the convergence zone framework, in: C. Koch, J.L. Davis (Eds.), Large Scale Neuronal Theories of the Brain, The MIT Press, Cambridge, 1995, pp. 61–74; G. Edelman, The Remembered Present, Basic Books, New York, 1989; M.I. Posner, M. Rothbart, Constructing neuronal theories of mind, in: C. Koch, J.L. Davis (Eds.), Large Scale Neuronal Theories of the Brain, The MIT Press, Cambridge, 1995, pp. 183–199; C. von der Malsburg, W. Schneider, A neuronal cocktail party processor, Biol. Cybem., 54 (1986) 29–40]. Reentry is not feedback, but parallel signalling in the time domain between spatially distributed maps, similar to a process of correlation between distributed systems. Accordingly, it was expected that during spontaneous reversals of the Necker cube, complex patterns of correlations between distributed systems would be present in the cortex. The present study included EEG ( n=4) and MEG recordings ( n=5). Two experimental questions were posed: (1) Can distributed cortical patterns present during perceptual reversals be classified differently using a generalised regression neural network (GRNN) compared to processing of a two-dimensional figure? (2) Does correlated cortical activity increase significantly during perception of a Necker cube reversal? One-second duration single trials of EEG and MEG data were analysed using the GRNN. Electrode/sensor pairings based on cortico–cortical connections were selected to assess correlated activity in each condition. The GRNN significantly classified single trials recorded during Necker cube reversals as different from single trials recorded during perception of a two-dimensional figure for both EEG and MEG. In addition, correlated cortical activity increased significantly in the Necker cube reversal condition for EEG and MEG compared to the perception of a non-reversing stimulus. Coherent MEG activity observed over occipital, parietal and temporal regions is believed to represent neural systems related to the perception of Necker cube reversals.

Nested reentrant and recurrent computation in early vision: a Bayesian neuromorphic model applied to hyperacuity

Hyperacuity is demonstrated in a neuromorphic model of the early visual system. The model incorporates Bayesian principles which are embodied in the dynamics of reentrant and recurrent feedback processes. Each retinotopically mapped area in the model represents a transformation of data from the visual field. Sensory information propagates in a bottom-up direction from one area to the next, while information based on Bayesian priors propagates in a top-down direction through reentrant connections. The 'bottom-up' and 'top-down' information maintain a separate existence in distinct layers of the model, but they interact through local connections within each area. Transformations between one area and the next are defined by the reentrant synaptic connections between areas, while local prior probability maps are defined by local recurrent connections within layers. The representation of hyperacuity is accomplished using a model of functional multiplicity: the large ratio of neurons in striate cortex compared with the number of afferent fibers projecting from the lateral geniculate nucleus. High functional multiplicity, in conjunction with hierarchical reentrant processing, allows the model to represent a fine-grained restoration of the line structure of visual input.

If I Only Had a Brain

If I Only Had a Brain Steven Shaviro Burroughs writes: “in this life we have to take things as we find them as the torso murderer said when he discovered his victim was a quadruple amputee.” Good advice for the anatomically deranged, like Cliff Steele. He’s a character in the DC/Vertigo comic book DOOM PATROL; I refer in particular to the issues written by Grant Morrison and illustrated by Richard Case, between 1989 and 1992 (#s 19–63). Cliff has a problem with his body, you see. It happened like this. He used to be a daredevil racing car driver; he had a horrible wreck. Nearly all of him was burned to ashes, but they snatched his brain from the flames. And then they implanted that brain in a new prosthetic body, all shiny metal, ultra high tech, a veritable fighting machine. Now Cliff is the muscle of the DOOM PATROL, a brain turned into brawn. They expect him to be a macho bruiser, when actually he’s quite sensitive underneath. And to add insult to injury, they call him Robotman—a name he violently hates. What would that do for your sense of self-esteem? The life of a superhero these days! Cliff thinks of himself as just a regular guy; Robocop fantasies are the last thing on his mind. But with a metallic casing like this, he can’t exactly blend into the crowd. It’s what Baudrillard calls hyper-visibility, the postmodern condition par excellence. No chance of chilling out with a secret identity, like old Clark Kent used to do. All this metal is a clunky encumbrance, no matter how great its tensile strength. You know you’re in bad shape when you bang your head against a wall, and you still don’t feel a thing. At this point, Cliff doesn’t even really know what his body can do. How good is all this cyber-tech stuff anyway? How accurate and detailed is sensory input? How fast is motor response? What unaccustomed relays and connections now trigger the pain and pleasure centers in Cliff’s brain? Will he ever be able to taste and smell? Can he ever have sex again? What about getting drunk or stoned? “The only good thing about having a human brain in a robot body,” Cliff remarks sardonically at one point, “is that it’s easier to control brain chemistry.” Just the touch of a button, and anxiety is dissipated, alertness is heightened, or memory is enhanced. But alas, this techno-manipulation seems to work only for utilitarian ends, and not for hedonistic ones. “Our machines are disturbingly lively,” Donna Haraway writes, “and we ourselves frighteningly inert.” It might not be so bad, if only you could get used to the situation. After all, Descartes argued long ago that the body is a machine. It shouldn’t matter all that much whether metal or flesh is the medium. In either case, it’s simply a matter of mastering the electrochemical interface: regularizing chains of association, facilitating neural feedback patterns, reinforcing the appropriate re-entrant connections. In short, a question of recognition and memory, of cultivating habits over the course of time. The problem is that Cliff’s mechanical body never stays the same. He’s continually being sent back to the shop for upgrades and repairs. Transistors burn out; programming errors and faulty couplings throw him off stride. He gets into fights, and enemies regularly mangle his metal to bits. As if that weren’t bad enough, Doc Magnus (who built and programmed his body in the first place) and Niles Caulder (the Chief of the DOOM PATROL) tend to use Cliff as a pawn in their ongoing professional rivalry. Neither of them is content to let well enough alone; they are both all too eager to retool him in order to try out their latest cybernetic design ideas. And let’s not even think about those insectoid aliens who at one point fit Cliff out in a new metal carapace with six legs. Life in a robot body, even if you’re strong, is just one humiliation after another. The persistence of memory in the brain...

Modeling perceptual grouping and figure-ground segregation by means of active reentrant connections.

The segmentation of visual scenes is a fundamental process of early vision, but the underlying neural mechanisms are still largely unknown. Theoretical considerations as well as neurophysiological findings point to the importance in such processes of temporal correlations in neuronal activity. In a previous model, we showed that reentrant signaling among rhythmically active neuronal groups can correlate responses along spatially extended contours. We now have modified and extended this model to address the problems of perceptual grouping and figure-ground segregation in vision. A novel feature is that the efficacy of the connections is allowed to change on a fast time scale. This results in active reentrant connections that amplify the correlations among neuronal groups. The responses of the model are able to link the elements corresponding to a coherent figure and to segregate them from the background or from another figure in a way that is consistent with the so-called Gestalt laws.

Integration of distributed cortical systems by reentry: a computer simulation of interactive functionally segregated visual areas

A computer model based on visual cortex has been constructed to analyze how the operations of multiple, functionally segregated cortical areas can be coordinated and integrated to yield a unified perceptual response. We propose that cortical integration arises through the process of reentry--the ongoing, parallel, recursive signaling between separate maps along ordered anatomical connections. To test the efficacy of this reentrant cortical integration (RCI) model, we have carried out detailed computer simulations of 3 interconnected cortical areas in the striate and extrastriate cortex of the macaque. The simulated networks contained a total of over 222,000 units and 8.5 million connections. The 3 modeled areas, called VOR, VOC, and VMO, incorporate major anatomical and physiological properties of cortical areas V1, V3, and V5 but are vastly simplified compared with monkey visual cortex. Simulated area VOR contains both orientation and directionally selective units; simulated area VMO discriminates the direction of motion of arbitrarily oriented objects; and simulated area VOC responds to both luminance and occlusion boundaries in the stimulus. Area VOC is able to respond to illusory contours (Kanizsa, 1979) by means of the same neural architecture used for the discrimination of occlusion boundaries. This architecture also generates responses to structure-from-motion by virtue of reentrant connections from VMO to VOC. The responses of the simulated networks to these illusions are consistent with the perceptual responses of humans and other species presented with these stimuli. The networks also respond in a consistent manner to a novel illusion that combines illusory contours and structure-from-motion. The response synthesized to this combined illusion provides a strong argument supporting the need for a recursive reentrant process in the cortex. Functional integration of the simulated areas in the RCI model were found to depend upon the combined action of 3 reentrant processes: (1) conflicting responses among segregated areas are competitively eliminated, (2) outputs of each area are used by other areas in their own operations, and (3) outputs of an area are "reentered" back to itself (through lower areas) and can thus be used iteratively to synthesize responses to complex or illusory stimuli. Transection of the reentrant connections selectively abolished these integrative processes and led to failure of figural synthesis. The proposed model of reentry suggests a basis for understanding how multiple visual areas as well as other cortical areas may be integrated within a distributed system.