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View from the top: hierarchies and reverse hierarchies in the visual system.

We propose that explicit vision advances in reverse hierarchical direction, as shown for perceptual learning. Processing along the feedforward hierarchy of areas, leading to increasingly complex representations, is automatic and implicit, while conscious perception begins at the hierarchy's top, gradually returning downward as needed. Thus, our initial conscious percept--vision at a glance--matches a high-level, generalized, categorical scene interpretation, identifying "forest before trees." For later vision with scrutiny, reverse hierarchy routines focus attention to specific, active, low-level units, incorporating into conscious perception detailed information available there. Reverse Hierarchy Theory dissociates between early explicit perception and implicit low-level vision, explaining a variety of phenomena. Feature search "pop-out" is attributed to high areas, where large receptive fields underlie spread attention detecting categorical differences. Search for conjunctions or fine discriminations depends on reentry to low-level specific receptive fields using serial focused attention, consistent with recently reported primary visual cortex effects.

Humans↗

Receptive fields of disparity-tuned simple cells in macaque V1.

Binocular simple cells in primary visual cortex (V1) are the first cells along the mammalian visual pathway to receive input from both eyes. Two models of how binocular simple cells could extract disparity information have been put forward. The phase-shift model proposes that the receptive fields in the two eyes have different subunit organizations, while the position-shift model proposes that they have different overall locations. In five fixating macaque monkeys, we recorded from 30 disparity-tuned simple cells that showed selectivity to the disparity in a random dot stereogram. High-resolution maps of the left and right eye receptive fields indicated that both phase and position shifts were common. Single cells usually showed a combination of the two, and the optimum disparity was best correlated with the sum of receptive field phase and position shift.

Animals↗

Functional topography of connections linking mirror-symmetric maps in the mouse olfactory bulb.

In rodents, each main olfactory bulb contains two mirror-symmetric glomerular maps, a feature not found in the initial topographic maps of other sensory systems. Targeting tracer injections to identified glomeruli revealed that isofunctional odor columns-translaminar assemblies connected to a given glomerulus-were specifically and reciprocally interconnected through a mutually inhibitory circuit with exquisite topographic specificity. Thus, instead of containing two mirror-symmetric maps, we propose that the olfactory bulb contains a single integrated map in which isofunctional odor columns are connected through an intrabulbar link, analogous to the specific horizontal connections linking iso-orientation columns in primary visual cortex.

Animals↗

End-stopping and the aperture problem: two-dimensional motion signals in macaque V1.

Our perception of fine visual detail relies on small receptive fields at early stages of visual processing. However, small receptive fields tend to confound the orientation and velocity of moving edges, leading to ambiguous or inaccurate motion measurements (the aperture problem). Thus, it is often assumed that neurons in primary visual cortex (V1) carry only ambiguous motion information. Here we show that a subpopulation of V1 neurons is capable of signaling motion direction in a manner that is independent of contour orientation. Specifically, end-stopped V1 neurons obtain accurate motion measurements by responding only to the endpoints of long contours, a strategy which renders them largely immune to the aperture problem. Furthermore, the time course of end-stopping is similar to the time course of motion integration by MT neurons. These results suggest that cortical neurons might represent object motion by responding selectively to two-dimensional discontinuities in the visual scene.

Animals↗

Effects of cholinergic enhancement on visual stimulation, spatial attention, and spatial working memory.

We compared behavioral and neural effects of cholinergic enhancement between spatial attention, spatial working memory (WM), and visual control tasks, using fMRI and the anticholinesterase physostigmine. Physostigmine speeded responses nonselectively but increased accuracy selectively for attention. Physostigmine also decreased activations to visual stimulation across all tasks within primary visual cortex, increased extrastriate occipital cortex activation selectively during maintained attention and WM encoding, and decreased parietal activation selectively during maintained attention. Finally, lateralization of occipital activation as a function of the visual hemifield toward which attention or memory was directed was decreased under physostigmine. In the case of attention, this effect correlated strongly with a decrease in a behavioral measure of selective spatial processing. Our results suggest that, while cholinergic enhancement facilitates visual attention by increasing activity in extrastriate cortex generally, it accomplishes this in a manner that reduces expectation-driven selective biasing of extrastriate cortex.

Acetylcholine↗

Low N-acetyl-aspartate and high choline in the anterior cingulum of recently abstinent methamphetamine-dependent subjects: a preliminary proton MRS study. Magnetic resonance spectroscopy.

Studies based on animal models report that methamphetamine (MA) abuse diminishes dopamine (DA) and serotonin innervation in frontal brain regions. In this in vivo human study, we used proton magnetic resonance spectroscopy (MRS), which yields measures of N-acetyl-aspartate (NAA), a marker of living neurons, to examine frontal brain regions possibly affected by methamphetamine dependence (MD). We tested the hypothesis that MD subjects would exhibit abnormally low levels of NAA, referenced to creatine (Cr), in anterior cingulate gray matter. We further hypothesized that the primary visual cortex, which receives relatively less DA innervation than the frontal brain regions, would show normal NAA/Cr ratios in MD subjects. Subjects included nine MD men (mean+/-standard deviation (S.D.)=32.5+/-6.4 years) and nine age-matched control men (mean+/-S.D.=32.7+/-6.8 years). The MD subjects were MA-free for 4-13 weeks. Proton MRS metabolites were expressed as ratios of creatine; the absolute values of which did not distinguish controls and MD subjects. With regard to metabolite ratios, the MD men had significantly lower NAA/Cr in the cingulum (mean+/-standard error (S.E.): control=1.46+/-0.03; MD=1.30+/-0.03; Mann-Whitney P=0.01) but not in the visual cortex (mean+/-S.E.: control=1.64+/-0.06; MD=1.69+/-11; Mann-Whitney P=0.52) relative to controls. These results provide evidence for NAA/Cr deficit that is selective to the anterior cingulum, at least with respect to visual cortex, in MD subjects. The neuronal compromise that these changes reflect may contribute to the attentional deficits and dampened reward system in MD.

Adult↗

Surface representation in the visual system.

Perception of surface accompanies the impression that a certain area of the visual field is occupied by some quality, such as color, brightness and transparency. This does not mean, however, that information about surface quality must be obtained throughout the area. It has been shown in many situations that our visual system has ability to interpolate information obtained at the border of the surface and to perceive homogeneous surfaces. The most dramatic demonstration of this is the perceptual filling-in at the blind spot. In order to understand the neural representation of surface in the visual system, we conducted a series of experiments using macaque monkeys. First, we examined if neurons in the primary visual cortex (V1) respond when a homogeneous surface is presented on the receptive field. Neurons representing the parafoveal visual field were tested and it was found that about one third of neurons showed significant responses when the cell's receptive field was contained in a homogeneous surface. Then we examined neuron activities in the retinotopic representation of the blind spot in V1. Although there is no retinal input in the blind spot, a homogeneous surface is perceived within the blind spot as a result of filling-in. We tested whether neurons in this region were activated when a homogeneous surface was perceived in the blind spot as a result of filling-in. We found some neurons in V1 were activated by stimuli which lead to the filling-in. These results indicate that when a surface area is perceived, neurons are activated throughout the region in V1 topographically corresponding to the perceived surface and not restricted to the region representing the border of the surface.

Animals↗

Visual extinction and cortical connectivity in human vision.

Visual extinction is a common, poorly understood, consequence of unilateral cerebral damage, where a patient fails to detect one of two simultaneously presented stimuli (the one more contralateral to the lesion), despite the fact that each stimulus is correctly detected when presented in isolation. The phenomenon implies a failure of shifting attention from an attended object to an unattended one. We show here that pair detection is improved in conditions where the two stimuli presented to the two halves of the visual field are proximal, co-oriented and co-axial. It is further shown that stimulus properties producing reduced extinction correlate with the selectivity pattern of spatial lateral interactions observed in the primary visual cortex. We suggest that neuronal activity in early stages of cortical visual processing encodes, using long-range lateral interactions, an image description in which visual objects are already segmented and marked. Segmentation seems to function properly even in the presence of significant destruction of the parietal cortex leading to extinction.

Cerebral Cortex↗

How can squint change the spacing of ocular dominance columns?

The pattern of ocular dominance columns in primary visual cortex of mammals such as cats and macaque monkeys arises during development by the activity-dependent refinement of thalamocortical connections. Manipulating visual experience in kittens by the induction of squint leads to the emergence of ocular dominance columns with a larger size and larger column-to-column spacing than in normally raised animals. The mechanism underlying this phenomenon is presently unknown. Theory suggests that experience cannot influence the spacing of columns if the development proceeds through purely Hebbian mechanisms. Here we study a developmental model in which Hebbian mechanisms are complemented by activity-dependent regulation of the total strength of afferent synapses converging onto a cortical neurone. We show that this model implies an influence of visual experience on the spacing of ocular dominance columns and provides a conceptually simple explanation for the emergence of larger sized columns in squinting animals. Assuming that during development cortical neurones become active in local groups, which we call co-activated cortical domains (CCDs), ocular dominance segregation is controlled by the size of these groups: (1) Size and spacing of ocular dominance columns are proportional to the size sigma of CCDs. (2) There is a critical size sigma* of CCDs such that ocular dominance columns form if sigma sigma*. This critical size of CCDs is determined by the correlation functions of activity patterns in the two eyes and specifies the influence of experience on ocular dominance segregation. We show that sigma* is larger with squint than with normal visual experience. Since experimental evidence indicates that the size of CCDs decreases during development, ocular dominance columns are predicted to form earlier and with a larger spacing in squinters compared to normal animals.

Animals↗

Dynamics of functional connectivity in visual cortical networks: an overview.

The aim of the Royaumont Symposium was to review various dynamic aspects of adaptive changes in functional connectivity, expressed in cortical networks during development, learning, and possibly during recognition and cognitive processing. The link between the various experimental and theoretical models was the comparison of cellular and molecular mechanisms that could be involved in the up- and down-regulation of functional connectivity, over different time scales. These processes have been investigated using several approaches in parallel: 1) at the molecular/subcellular level, to identify postsynaptic receptors (NMDA, mGluR) and second messengers (calcium protein kinases and phosphatases) involved in the induction of synaptic potentiation and depression, and to characterize diffusible factors (NO), released pre- or postsynaptically, involved in the spatial generalization of local changes to neighboring synapses; 2) at the level of integrating networks, to develop electrophysiological (single and multiple recording), pharmacological and optical imaging techniques in order to compare the dynamics of adaptive processes put into play during the natural development of cortical specificity and connectivity, versus those triggered during forced regimes of temporal correlations between pre- and post synaptic activities. Both in vitro and in vivo approaches have been combined in the primary visual cortex of the developing and adult vertebrate (rat, guinea-pig, ferret, cat and monkey). The various forms of 'slow' synaptic plasticity, demonstrated during epigenesis and selective phases of learning in the adult, can be compared with 'fast' forms of functional coupling (or synchronous firing) shown to develop during the time span required for perception and cognitive processing Phenomenology of the dynamics in functional connectivity and their relative dependence on temporal correlation in neuronal activity have been analyzed in each of these situations. Experimental results have been compared at different levels of neuronal integration (synapse, column map and cell assembly) in order to gain a better understanding of functional grouping within cortical networks.

Adaptation, Physiological↗

Neuronal plasticity as an adaptive property of the central nervous system.

This short review presents examples of plasticity in the brains of vertebrates including man. The basic ability of the nervous system to make functionally relevant adaptations to functional challenges of various kinds during development and adulthood is called plasticity. Enucleation of the eyes or lesioning of the lateral geniculate body during development lead to the generation of a new architectonic area within the nonhuman primate and human primary visual cortex. The enucleation of one eye in rats at various postnatal stages causes profound plastic changes in the callosal system of the visual cortex. The central representation of the periphery in the adult cerebral cortex (somatotopy) can also be altered by adaptive processes. Naturally occurring nerve cell death during pre- and early postnatal development can be manipulated by impairing normal development of neuro-transmission. These findings argue for an important role of transmitter receptors in brain plasticity. The number of receptors shows, for most brain regions and receptor types, an overshoot of growth during ontogeny. After lesions have damaged the adult geniculo-cortical and septo-hippocampal systems, receptors can exhibit plastic changes such as upregulation of the number of binding sites (visual cortex) and modifications in the coupling of receptors, transducer proteins (G-proteins) and second messengers (hippocampus).

Aging↗

Ocular dominance development revisited.

New approaches to the study of ocular dominance development, a model system for the development of neural architecture, indicate that eye-specific columns in primary visual cortex emerge substantially before the onset of the critical period, during which neural connections can be altered by visual experience. The timing, speed and specificity of column emergence implicate molecular patterning mechanisms, along with patterns of neural activity, in the generation of this columnar architecture.

Animals↗

Intrinsic processing in the mammalian superior colliculus.

The mammalian superior colliculus receives visual inputs from the retina and primary visual cortex in its superficial layers and sends descending motor commands from its deeper layers. It is now becoming clear that a connection exists between these layers, but the signal transmission through it is not robust. The induction of burst discharges in the deeper layer neurons by direct visual inputs from the superficial layers may lead to 'express' saccadic eye movements with extremely short reaction times in behaving animals.

Action Potentials↗

Mechanisms of stereoscopic vision: the disparity energy model.

The past year has seen significant advances in our understanding of the role played by the primary visual cortex (V1) in stereoscopic vision. Recently, the mechanism by which complex cells in V1 respond to random-dot stereograms has been characterized; it appears that their response properties greatly reduce the complexity of one of the critical links for stereopsis, the correspondence problem.

Animals↗

Neural mechanisms underlying amblyopia.

The nature of the neural basis of amblyopia is a matter of some debate. Recent neurophysiological data show correlates of amblyopia in the spatial properties of neurons in primary visual cortex. These neuronal deficits are probably the initial manifestation of the visual loss, but there are almost certainly additional deficits at higher levels of the visual pathways.

Amblyopia↗

A conscious route to unconscious vision.

Damage to the primary visual cortex can leave subjects with unconscious residual vision, or 'blindsight'. New research suggests that 'top-down' modulation by intact conscious visual processes can improve performance in the impaired visual domain, even though that domain still remains quite inaccessible to consciousness.

Humans↗

Interhemispheric switching mediates perceptual rivalry.

BACKGROUND: Binocular rivalry refers to the alternating perceptual states that occur when the images seen by the two eyes are too different to be fused into a single percept. Logothetis and colleagues have challenged suggestions that this phenomenon occurs early in the visual pathway. They have shown that, in alert monkeys, neurons in the primary visual cortex continue to respond to their preferred stimulus despite the monkey reporting its absence. Moreover, they found that neural activity higher in the visual pathway is highly correlated with the monkey's reported percept. These and other findings suggest that the neural substrate of binocular rivalry must involve high levels, perhaps the same levels involved in reversible figure alternations. RESULTS: We present evidence that activation or disruption of a single hemisphere in human subjects affects the perceptual alternations of binocular rivalry. Unilateral caloric vestibular stimulation changed the ratio of time spent in each competing perceptual state. Transcranial magnetic stimulation applied to one hemisphere disrupted normal perceptual alternations when the stimulation was timed to occur at one phase of the perceptual switch, but not at the other. Furthermore, activation of a single hemisphere by caloric stimulation affected the perceptual alternations of a reversible figure, the Necker cube. CONCLUSIONS: Our findings suggest that interhemispheric switching mediates perceptual rivalry. Thus, competition for awareness in both binocular rivalry and reversible figures occurs between, rather than within, each hemisphere. This interhemispheric switch hypothesis has implications for understanding the neural mechanisms of conscious experience and also has clinical relevance as the rate of both types of perceptual rivalry is slow in bipolar disorder (manic depression).

Adolescent↗

The dynamics of velocity adaptation in human vision.

Since Barlow and Hill's classic study of the adaptation of the rabbit ganglion cell to movement [1], there have been several reports that motion adaptation is accompanied by an exponential reduction in spike rate, and similar estimates of the time course of velocity adaptation have been found across species [2-4]. Psychophysical studies in humans have shown that perceived velocity may reduce exponentially with adaptation [5,6]. It has been suggested that the reduction in firing of single cells may constitute the neural substrate of the reduction in perceived speed in humans [1,5-7]. Although a model of velocity coding in which the firing rate directly encodes speed may have the advantage of simplicity, it is not supported by psychophysical research. Furthermore, psychophysical estimates of the time course of perceived speed adaptation are not entirely consistent with physiological estimates. This discrepancy between psychophysical and physiological estimates may be due to the unrealistic assumption that speed is coded in the gross spike rate of neurons in the primary visual cortex. The psychophysical data on motion processing are, however, generally consistent with a model in which perceived velocity is derived from the ratio of two temporal channels [8-14]. We have examined the time course of speed adaptation and recovery to determine whether the observed rates can be better related to the established physiology if a ratio model of velocity processing is assumed. Our results indicate that such a model describes the data well and can accommodate the observed difference in the time courses of physiological and psychophysical processes.

Adaptation, Physiological↗