Visual resolution and experience: acuity deficits in cats following early selective visual deprivation

Cats reared during the first 5 months of life in environments that contain contours of a single orientation show a diminished ability to resolve gratings of the orthogonal orientation in later life. It is argued that these perceptual deficits result from changes in the organization of the visual cortex induced by the selected early visual input.     

Ocular dominance column development: analysis and simulation

The visual cortex of many adult mammals has patches of cells that receive inputs driven by the right eye alternating with patches that receive inputs driven by the left eye. These ocular dominance patches (or "columns") form during early life as a consequence of competition between the activity patterns of the two eyes. A mathematical model of several biological mechanisms that can account for this development is presented. Analysis of this model reveals the conditions under which ocular dominance segregation will occur and determines the resulting patch width. Simulations of the model also exhibit other phenomena associated with early visual development, such as topographic refinement of cortical receptive fields, the confinement of input cell connections to patches, monocular deprivation plasticity including a critical period, and the effect of artificially induced strabismus. The model can be used to predict the results of proposed experiments and to discriminate among various mechanisms of plasticity.     

Bottom-up dependent gating of frontal signals in early visual cortex

The frontal eye field (FEF) is one of several cortical regions thought to modulate sensory inputs. Moreover, several hypotheses suggest that the FEF can only modulate early visual areas in the presence of a visual stimulus. To test for bottom-up gating of frontal signals, we microstimulated subregions in the FEF of two monkeys and measured the effects throughout the brain with functional magnetic resonance imaging. The activity of higher-order visual areas was strongly modulated by FEF stimulation, independent of visual stimulation. In contrast, FEF stimulation induced a topographically specific pattern of enhancement and suppression in early visual areas, but only in the presence of a visual stimulus. Modulation strength depended on stimulus contrast and on the presence of distractors. We conclude that bottom-up activation is needed to enable top-down modulation of early visual cortex and that stimulus saliency determines the strength of this modulation.     

Functional organization of primate visual cortex revealed by high resolution optical imaging

A high spatial resolution optical imaging system was developed to visualize cerebral cortical activity in vivo. This method is based on activity-dependent intrinsic signals and does not use voltage-sensitive dyes. Images of the living monkey striate (VI) and extrastriate (V2) visual cortex, taken during visual stimulation, were analyzed to yield maps of the distribution of cells with various functional properties. The cytochrome oxidase--rich blobs of V1 and the stripes of V2 were imaged in the living brain. In V2, no ocular dominance organization was seen, while regions of poor orientation tuning colocalized to every other cytochrome oxidase stripe. The orientation tuning of other regions of V2 appeared organized as modules that are larger and more uniform than those in V1.     

Role of subplate neurons in functional maturation of visual cortical columns

The subplate forms a transient circuit required for development of connections between the thalamus and the cerebral cortex. When subplate neurons are ablated, ocular dominance columns do not form in the visual cortex despite the robust presence of thalamic axons in layer 4. We show that subplate ablation also prevents formation of orientation columns. Visual responses are weak and poorly tuned to orientation. Furthermore, thalamocortical synaptic transmission fails to strengthen, whereas intracortical synapses are unaffected. Thus, subplate circuits are essential not only for the anatomical segregation of thalamic inputs but also for key steps in synaptic remodeling and maturation needed to establish the functional architecture of visual cortex.     

Reward timing in the primary visual cortex

We discovered that when adult rats experience an association between visual stimuli and subsequent rewards, the responses of a substantial fraction of neurons in the primary visual cortex evolve from those that relate solely to the physical attributes of the stimuli to those that accurately predict the timing of reward. In addition to revealing a remarkable type of response plasticity in adult V1, these data demonstrate that reward-timing activity-a "higher" brain function-can occur very early in sensory-processing paths. These findings challenge the traditional interpretation of activity in the primary visual cortex.     

Temporal specificity in the cortical plasticity of visual space representation

The circuitry and function of mammalian visual cortex are shaped by patterns of visual stimuli, a plasticity likely mediated by synaptic modifications. In the adult cat, asynchronous visual stimuli in two adjacent retinal regions controlled the relative spike timing of two groups of cortical neurons with high precision. This asynchronous pairing induced rapid modifications of intracortical connections and shifts in receptive fields. These changes depended on the temporal order and interval between visual stimuli in a manner consistent with spike timing-dependent synaptic plasticity. Parallel to the cortical modifications found in the cat, such asynchronous visual stimuli also induced shifts in human spatial perception.     

Neural mechanisms of visual attention: how top-down feedback highlights relevant locations

Attention helps us process potentially important objects by selectively increasing the activity of sensory neurons that represent the relevant locations and features of our environment. This selection process requires top-down feedback about what is important in our environment. We investigated how parietal cortical output influences neural activity in early sensory areas. Neural recordings were made simultaneously from the posterior parietal cortex and an earlier area in the visual pathway, the medial temporal area, of macaques performing a visual matching task. When the monkey selectively attended to a location, the timing of activities in the two regions became synchronized, with the parietal cortex leading the medial temporal area. Parietal neurons may thus selectively increase activity in earlier sensory areas to enable focused spatial attention.     

A map of visual space induced in primary auditory cortex

Maps of sensory surfaces are a fundamental feature of sensory cortical areas of the brain. The relative roles of afferents and targets in forming neocortical maps in higher mammals can be examined in ferrets in which retinal inputs are directed into the auditory pathway. In these animals, the primary auditory cortex contains a systematic representation of the retina (and of visual space) rather than a representation of the cochlea (and of sound frequency). A representation of a two-dimensional sensory epithelium, the retina, in cortex that normally represents a one-dimensional epithelium, the cochlea, suggests that the same cortical area can support different types of maps. Topography in the visual map arises both from thalamocortical projections that are characteristic of the auditory pathway and from patterns of retinal activity that provide the input to the map.     

Neuroscience. Noise makes sense in neuronal computing

The united efforts of assemblies of neurons in the brain's primary visual cortex translate incoming visual signals into action potentials. These action potentials encode, for example, the contrast and orientation of different parts of the image. Some neurons are sensitive to one particular orientation, other are sensitive to other orientations, but all neurons respond equally well to the image contrast. In a Perspective, Volgushev and Eysel explain the finding (Anderson et al.) that neurons are able to maintain this sensitivity to the orientation of a stimulus regardless of the contrast by adding noise to the membrane potential, such that action potentials can also be generated in response to weak signals at low contrast.     

The role of area 17 in visual imagery: convergent evidence from PET and rTMS

Visual imagery is used in a wide range of mental activities, ranging from memory to reasoning, and also plays a role in perception proper. The contribution of early visual cortex, specifically Area 17, to visual mental imagery was examined by the use of two convergent techniques. In one, subjects closed their eyes during positron emission tomography (PET) while they visualized and compared properties (for example, relative length) of sets of stripes. The results showed that when people perform this task, Area 17 is activated. In the other, repetitive transcranial magnetic stimulation (rTMS) was applied to medial occipital cortex before presentation of the same task. Performance was impaired after rTMS compared with a sham control condition; similar results were obtained when the subjects performed the task by actually looking at the stimuli. In sum, the PET results showed that when patterns of stripes are visualized, Area 17 is activated, and the rTMS results showed that such activation underlies information processing.     

Single-neuron activity and tissue oxygenation in the cerebral cortex

Blood oxygen level-dependent functional magnetic resonance imaging uses alterations in brain hemodynamics to infer changes in neural activity. Are these hemodynamic changes regulated at a spatial scale capable of resolving functional columns within the cerebral cortex? To address this question, we made simultaneous measurements of tissue oxygenation and single-cell neural activity within the visual cortex. Results showed that increases in neuronal spike rate were accompanied by immediate decreases in tissue oxygenation. We used this decrease in tissue oxygenation to predict the orientation selectivity and ocular dominance of neighboring neurons. Our results establish a coupling between neural activity and oxidative metabolism and suggest that high-resolution functional magnetic resonance imaging may be used to localize neural activity at a columnar level.     

Feedback connections to the lateral geniculate nucleus and cortical response properties

The cerebral cortex receives sensory input from the periphery by means of thalamic relay nuclei, but the flow of information goes both ways. Each cortical area sends a reciprocal projection back to the thalamus. In the visual system, the synaptic relations that govern the influence of thalamic afferents on orientation selectivity in the cortex have been studied extensively. It now appears that the connectivity of the corticofugal feedback pathway is also fundamentally linked to the orientation preference of the cortical cells involved.     

Paths of information flow through visual cortex.

The main route of information flow in the cerebral cortex is from the middle layers of cortex to upper and lower layers. However, upper layers of the cat primary visual cortex can be directly driven by inputs from secondary visual cortex when activity in middle layers is disrupted. Upper-layer activity can be driven either by middle layers or by direct corticocortical inputs. One consequence of this result is that areas of cortex thought to be carrying out low-order analysis may be able to extract considerable information from higher order areas.     

Interhemispheric effects of cortical lesions on brain biochemistry

Unilateral lesions in the visual and somesthetic cortex of rat brain cause a slight but significant increase inc the cortical cholinesterase activity in the contralateral hemisphere. There is some indication of strain differences in this effect. No change in cholinesterase activity is found in the subcortical brain. These findings may be helpful in understanding mechanisms of interdependence among brain areas.     

Neuronal correlates of eye movements in the visual cortex of the cat

About 10 percent of the cells in the visual cortex of awake cats do not respond to stationary parallel stripes in any orientation or to stripes moving across the visual field in any direction at a moderate speed (up to 132 degrees per second), but these cells are either excited or inhibited during saccadic eye movements when the animal faces a patterned visual environment. Of nineteen such cells tested in total darkness, seven discharged in association with eye movements. For saccade-related discharges, the latency during retinal stimulation is typically shorter than the latencey in total darkness.     

G protein-coupled receptor-dependent development of human frontal cortex

The mammalian cerebral cortex is characterized by complex patterns of anatomical and functional areas that differ markedly between species, but the molecular basis for this functional subdivision is largely unknown. Here, we show that mutations in GPR56, which encodes an orphan G protein-coupled receptor (GPCR) with a large extracellular domain, cause a human brain cortical malformation called bilateral frontoparietal polymicrogyria (BFPP). BFPP is characterized by disorganized cortical lamination that is most severe in frontal cortex. Our data suggest that GPCR signaling plays an essential role in regional development of human cerebral cortex.     

Electrophysiological evidence that abnormal early visual experience can modify the human brain

Visual resolution in humans is nearly equal for vertically and horizontally oriented detail, but for some subjects there is a substantial difference in resolving power for these orientations. Although subjects who exhibit this difference invariably have ocular astigmatism, optical explanations of the effect can be ruled out. Direct evidence has been found for an electrophysiological correlate to the psychophysical finding. Subjects who have reduced resolution for a pattern of a particular orientation also show a decreased evoked potential response elicited by a target of the same orientation. The results are consistent with the hypothesis that a deficiency of specific features in the early visual input can alter the organization of the visual pathways.     

Neuronal activity in the lateral intraparietal area and spatial attention

Although the parietal cortex has been implicated in the neural processes underlying visual attention, the nature of its contribution is not well understood. We tracked attention in the monkey and correlated the activity of neurons in the lateral intraparietal area (LIP) with the monkey's attentional performance. The ensemble activity in LIP across the entire visual field describes the spatial and temporal dynamics of a monkey's attention. Activity subtending a single location in the visual field describes the attentional priority at that area but does not predict that the monkey will actually attend to or make an eye movement to that location.     

The contribution of noise to contrast invariance of orientation tuning in cat visual cortex

Feedforward models of visual cortex appear to be inconsistent with a well-known property of cortical cells: contrast invariance of orientation tuning. The models' fixed threshold broadens orientation tuning as contrast increases, whereas in real cells tuning width is invariant with contrast. We have compared the orientation tuning of spike and membrane potential responses in single cells. Both are contrast invariant, yet a threshold-linear relation applied to the membrane potential accurately predicts the orientation tuning of spike responses. The key to this apparent paradox lies in the noisiness of the membrane potential. Responses that are subthreshold on average are still capable of generating spikes on individual trials. Unlike the iceberg effect, contrast invariance remains intact even as threshold narrows orientation selectivity. Noise may, by extension, smooth the average relation between membrane potential and spike rate throughout the brain.