Spatial context in the early visual system

Important visual objects in our everyday life, such as fellow people, passing cars or birds perhaps, are not point-like structures but often occupy considerable amounts of the visual field. However, each photoreceptor in our eyes samples just a tiny portion of the visual field and somehow the visual system should integrate these local signals. This process takes place mainly in the visual cortex and, while higher-order visual areas play an important role in perception of extended structures, it is now well established that visual neurons at the first cortical steps of seeing integrate broad spatial context into their responses. The main purpose of this thesis was to provide detailed information concerning the spatial structure of the mechanisms that underlie integration of spatial context in the early visual system. The opening study of this thesis showed that the antagonistic Gaussians structure that has been used for modeling context integration in single visual neurons provides a relatively accurate description of the process also in the human visual system. The first study introduced a novel method for connecting perceptual and neuroimaging measurements and this method was applied in the second study of this thesis. The second study showed that the human visual system integrates spatial context in terms of its visual field size instead of the size of its cortical representation. The third study showed that context is integrated over an unexpectedly large region of the visual field and that spatially distant context may sometimes increase the contrast response of the visual system. The closing study showed that orientation specificity of the integration of spatial context depends on distance both in single neurons in the macaque primary visual cortex and in human perception. The knowledge acquired in this thesis will be generally useful in applications that require understanding of the human visual system.Arkielämän kannalta tärkeät visuaaliset objektit kuten ihmiset, ohikiitävät autot ja kenties kissat, ovat harvoin pistemäisiä, mutta sen sijaan voivat peittää laajankin alueen näkökentästä. Näköaistinsolut prosessoivat kuvainformaatiota erittäin pieneltä näkökentän alueelta ja näköjärjestelmän tulee jollain tavoin yhdistää nämä paikalliset signaalit. Vaikka näköaivokuoren myöhäisten alueiden merkitys spatiaalisesti laajojen objektien havaitsemisessa onkin merkittävä, nykytietämyksen valossa on kiistatonta että myös varhaisten näköaivokuorten hermosolut integroivat spatiaalista kontekstia laajalta näkökentän alueelta. Tässä väitöskirjassa tutkitaan konteksti-integraation taustalla olevien mekanismien spatiaalista rakennetta varhaisessa näköjärjestelmässä. Väitöskirjan ensimmäisessä osatyössä osoitettiin että konteksti-integraatiota yksittäisissä hermosoluissa kuvaavat kahden antagonistisen Gaussilaisen mallit ovat melko hyviä kuvauksia konteksti-integraatiomekanismien spatiaalisesta rakenteesta myös ihmisen näköjärjestelmässä. Ensimmäisessä osatyössä kehitettiin menetelmä joka mahdollistaa havainto- ja aivokuvantamismittausten uudenlaisen yhdistämisen. Tätä menetelmää sovellettiin toisessa osatyössä, jonka päätulos oli konteksti-integraation riippuvuus ärsykkeen koosta näkökentässä sen sijaan että se olisi sidoksissa ärsykkeen edustuksen kokoon aivokuorella. Kolmannessa osatyössä osoitettiin, että kontekstia integroidaan huomattavan laajalta alueelta ja että spatiaalisesti etäinen konteksti saattaa toisinaan vahvistaa näköjärjestelmän kontrastivastetta. Neljäs tutkimus osoitti, että konteksti-integraation valikoivuus orientaatiolle riippuu etäisyydestä niin ihmisen näköhavainnoissa kuin makaki-apinan ensimmäisen näköaivokuoren soluissakin. Tämän väitöskirjan tuloksia voidaan hyödyntää sovelluksissa joissa tarvitaan tietoa ihmisen näköjärjestelmän toiminnasta

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.CELEST, an NSF Science of Learning Center (SBE-0354378); SyNAPSE program of the Defense Advanced Research Project Agency (HR001109-03-0001); Defense Advanced Research Project Agency (HR001-09-C-0011

Feed-Forward Segmentation of Figure-Ground and Assignment of Border-Ownership

Figure-ground is the segmentation of visual information into objects and their surrounding backgrounds. Two main processes herein are boundary assignment and surface segregation, which rely on the integration of global scene information. Recurrent processing either by intrinsic horizontal connections that connect surrounding neurons or by feedback projections from higher visual areas provide such information, and are considered to be the neural substrate for figure-ground segmentation. On the contrary, a role of feedforward projections in figure-ground segmentation is unknown. To have a better understanding of a role of feedforward connections in figure-ground organization, we constructed a feedforward spiking model using a biologically plausible neuron model. By means of surround inhibition our simple 3-layered model performs figure-ground segmentation and one-sided border-ownership coding. We propose that the visual system uses feed forward suppression for figure-ground segmentation and border-ownership assignment

Dynamic and Integrative Properties of the Primary Visual Cortex

The ability to derive meaning from complex, ambiguous sensory input requires the integration of information over both space and time, as well as cognitive mechanisms to dynamically shape that integration. We have studied these processes in the primary visual cortex (V1), where neurons have been proposed to integrate visual inputs along a geometric pattern known as the association field (AF). We first used cortical reorganization as a model to investigate the role that a specific network of V1 connections, the long-range horizontal connections, might play in temporal and spatial integration across the AF. When retinal lesions ablate sensory information from portions of the visual field, V1 undergoes a process of reorganization mediated by compensatory changes in the network of horizontal collaterals. The reorganization accompanies the brain’s amazing ability to perceptually “fill-inâ€, or “seeâ€, the lost visual input. We developed a computational model to simulate cortical reorganization and perceptual fill-in mediated by a plexus of horizontal connections that encode the AF. The model reproduces the major features of the perceptual fill-in reported by human subjects with retinal lesions, and it suggests that V1 neurons, empowered by their horizontal connections, underlie both perceptual fill-in and normal integrative mechanisms that are crucial to our visual perception. These results motivated the second prong of our work, which was to experimentally study the normal integration of information in V1. Since psychophysical and physiological studies suggest that spatial interactions in V1 may be under cognitive control, we investigated the integrative properties of V1 neurons under different cognitive states. We performed extracellular recordings from single V1 neurons in macaques that were trained to perform a delayed-match-to-sample contour detection task. We found that the ability of V1 neurons to summate visual inputs from beyond the classical receptive field (cRF) imbues them with selectivity for complex contour shapes, and that neuronal shape selectivity in V1 changed dynamically according to the shapes monkeys were cued to detect. Over the population, V1 encoded subsets of the AF, predicted by the computational model, that shifted as a function of the monkeys’ expectations. These results support the major conclusions of the theoretical work; even more, they reveal a sophisticated mode of form processing, whereby the selectivity of the whole network in V1 is reshaped by cognitive state

Change blindness: eradication of gestalt strategies

Arrays of eight, texture-defined rectangles were used as stimuli in a one-shot change blindness (CB) task where there was a 50% chance that one rectangle would change orientation between two successive presentations separated by an interval. CB was eliminated by cueing the target rectangle in the first stimulus, reduced by cueing in the interval and unaffected by cueing in the second presentation. This supports the idea that a representation was formed that persisted through the interval before being 'overwritten' by the second presentation (Landman et al, 2003 Vision Research 43149–164]. Another possibility is that participants used some kind of grouping or Gestalt strategy. To test this we changed the spatial position of the rectangles in the second presentation by shifting them along imaginary spokes (by ±1 degree) emanating from the central fixation point. There was no significant difference seen in performance between this and the standard task [F(1,4)=2.565, p=0.185]. This may suggest two things: (i) Gestalt grouping is not used as a strategy in these tasks, and (ii) it gives further weight to the argument that objects may be stored and retrieved from a pre-attentional store during this task

Neural models of inter-cortical networks in the primate visual system for navigation, attention, path perception, and static and kinetic figure-ground perception

Vision provides the primary means by which many animals distinguish foreground objects from their background and coordinate locomotion through complex environments. The present thesis focuses on mechanisms within the visual system that afford figure-ground segregation and self-motion perception. These processes are modeled as emergent outcomes of dynamical interactions among neural populations in several brain areas. This dissertation specifies and simulates how border-ownership signals emerge in cortex, and how the medial superior temporal area (MSTd) represents path of travel and heading, in the presence of independently moving objects (IMOs). Neurons in visual cortex that signal border-ownership, the perception that a border belongs to a figure and not its background, have been identified but the underlying mechanisms have been unclear. A model is presented that demonstrates that inter-areal interactions across model visual areas V1-V2-V4 afford border-ownership signals similar to those reported in electrophysiology for visual displays containing figures defined by luminance contrast. Competition between model neurons with different receptive field sizes is crucial for reconciling the occlusion of one object by another. The model is extended to determine border-ownership when object borders are kinetically-defined, and to detect the location and size of shapes, despite the curvature of their boundary contours. Navigation in the real world requires humans to travel along curved paths. Many perceptual models have been proposed that focus on heading, which specifies the direction of travel along straight paths, but not on path curvature. In primates, MSTd has been implicated in heading perception. A model of V1, medial temporal area (MT), and MSTd is developed herein that demonstrates how MSTd neurons can simultaneously encode path curvature and heading. Human judgments of heading are accurate in rigid environments, but are biased in the presence of IMOs. The model presented here explains the bias through recurrent connectivity in MSTd and avoids the use of differential motion detectors which, although used in existing models to discount the motion of an IMO relative to its background, is not biologically plausible. Reported modulation of the MSTd population due to attention is explained through competitive dynamics between subpopulations responding to bottom-up and top- down signals

Contour integration: Psychophysical, neurophysiological, and computational perspectives

One of the important roles of our visual system is to detect and segregate objects. Neurons in the early visual system extract local image features from the visual scene. To combine these features into separate, global objects, the visual system must perform some kind of grouping operation. One such operation is contour integration. Contours form the outlines of objects, and are the first step in shape perception. We discuss the mechanism of contour integration from psychophysical, neurophysiological, and computational perspectives