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Residual processing of chromatic signals in the absence of a geniculostriate projection.

We have investigated the residual processing of chromatic signals in a subject with unilateral damage to the primary visual cortex using psychophysical, pupillometric and functional magnetic resonance imaging (fMRI) methods. Of particular interest was to establish the correlation between the subject's ability to make use of chromatic signals in the blind hemifield to discriminate between different coloured targets, the corresponding residual pupil colour responses and the level and location of cortical activation generated by the same stimuli as revealed by fMRI. The results obtained using the three different experimental approaches are consistent and suggest that retrograde degeneration of thalamic and retinal chromatic processing mechanisms caused by damage to the primary visual cortex in man does not abolish completely the ability to process chromatic signals particularly when large, long-wavelength stimuli are employed.

Color Perception↗

Improvement of temporal resolution in fMRI using slice phase encode reordered 3D EPI.

A magnetic resonance imaging (MRI) pulse sequence was developed for acquiring high temporal resolution 3D functional magnetic resonance images (fMRI). The technique uses a 3D-acquisition scheme that increases the SNR per unit time. Compared to the previously developed methods, the proposed one does not sacrifice the spatial resolution or require a reduction in the number of acquired slices. The method was tested on six control subjects using a well-known visual activation task. Statistical analysis of the acquired data showed significant activation in the primary visual cortex of each subject. The proposed acquisition method makes use of the periodicity of the applied stimulus. A given kz plane is scanned sequentially using a modified 3D EPI pulse sequence while the stimulus cycle (SC) is applied. The same SC is repeated for each acquisition of a new kz plane sequence. After data acquisition, temporal reordering is done on the data to obtain the proper 3D image sequence. Since the familiarization of the subject with the activation task is an important aspect that might affect the performance of the method, habituation of the primary visual cortex was tested by applying a similar visual stimulus and acquiring data using standard 2D Echo Planar Imaging (EPI). No habituation was observed within a duration of 7 min 12 sec.

Adult↗

A network that uses few active neurones to code visual input predicts the diverse shapes of cortical receptive fields.

Computational models of primary visual cortex have demonstrated that principles of efficient coding and neuronal sparseness can explain the emergence of neurones with localised oriented receptive fields. Yet, existing models have failed to predict the diverse shapes of receptive fields that occur in nature. The existing models used a particular "soft" form of sparseness that limits average neuronal activity. Here we study models of efficient coding in a broader context by comparing soft and "bard" forms of neuronal sparseness. As a result of our analyses, we propose a novel network model for visual cortex. The model forms efficient visual representations in which the number of active neurones, rather than mean neuronal activity, is limited. This form of hard sparseness also economises cortical resources like synaptic memory and metabolic energy. Furthermore, our model accurately predicts the distribution of receptive field shapes found in the primary visual cortex of cat and monkey.

Animals↗

Ocular dominance predicts neither strength nor class of disparity selectivity with random-dot stimuli in primate V1.

We address two unresolved issues concerning the coding of binocular disparity in primary visual cortex. Experimental studies and theoretical models have suggested a relationship between a cell's ocular dominance, assessed with monocular stimuli, and its tuning to binocular disparity. First, the disparity energy model of disparity selectivity suggests that there should be a correlation between ocular dominance and the strength of disparity tuning. Second, several studies have reported a relationship between ocular dominance and the shape of the disparity tuning curve, with cells dominated by one eye more likely to have disparity tuning of the tuned-inhibitory type. We investigated both of these relationships in single neurons recorded from the primary visual cortex of awake fixating macaques, using dynamic random-dot patterns as a stimulus. To classify disparity tuning curves quantitatively, we develop a new measure of symmetry, which can be applied to any function. We find no evidence for any correlation between ocular dominance and the nature of disparity tuning. This places constraints on the circuitry underlying disparity tuning.

Algorithms↗

Contrast dependence of perceptual grouping in brain-damaged patients with visual extinction.

Extinction is manifested in conditions of bilateral simultaneous stimulation, as a failure to detect the stimulus contra-lateral to the side of a cerebral lesion, while the same stimulus is correctly detected there when presented in isolation. The phenomenon is usually interpreted in terms of impaired mobilization of attention from an attended to an unattended object. We have recently shown, using pairs of Gabor patches as stimuli, that pair detection is maximally improved in conditions where the two stimuli presented simultaneously to the two halves of the visual field are co-oriented and co-axial and their location is not too eccentric. Here we add new information by showing that contrast isotropy of the stimulus pair is important in producing this orientation-similarity gain. The further advantage of co-oriented co-linear stimuli over co-oriented parallel (vertical) stimuli was shown exclusively with iso-contrast stimulus pairs, and was significantly enhanced when the contrast level of the stimulus pair was low. Stimulus properties producing reduced extinction seem to correlate with the selectivity pattern and contrast dependence of (a) spatial lateral facilitation observed in psychophysical studies with normal observers, and (b) long-range interactions observed in the primary visual cortex. Thus, two remote visual stimuli seem to be processed as a single object when the corresponding neuronal activities are linked via long-range lateral interactions. The present demonstration of contrast dependency in such processing, strengthens our previous conjecture that even in the presence of significant, extinction producing, parietal damage, the primary visual cortex preserves the capacity to encode, using long-range lateral interactions, an image description in which visual objects are already segregated from background.

Aged↗

Functional magnetic resonance imaging mirrors recovery of visual perception after repetitive tachistoscopic stimulation in patients with partial cortical blindness.

We investigated three patients with partial cortical blindness after brain injury by means of functional magnetic resonance imaging (fMRI) before and after the application of a daily visual stimulation-therapy over a period of 6 months. Before therapy, fMRI data showed a severely reduced blood-oxygen-level-dependent (BOLD) signal in primary visual cortex when compared to healthy volunteers. Following several months of rehabilitative therapy a neuropsychological improvement of visual functions was accompanied by an increase in BOLD signal of residual perilesional regions whereas fMRI data of the control group remained unchanged. A high capacity of functional recovery and synaptic plasticity of surviving perilesional neuronal structures of primary visual cortex followed by an increased input into post-connected visual areas can be discussed as a basis for the reoccurrence of visual functions.

Adult↗

Human face perception traced by magneto- and electro-encephalography.

The temporal and spatial processing of face perception in normal subjects was traced by magnetoencephalography (MEG) and electroencephalography (EEG). We used 5 different visual stimuli: (1) face with opened eyes, (2) face with closed eyes, (3) eyes, (4) scrambled face, and (5) hand, and they were shown in random order. Subjects were asked to count the number of hand stimuli. To analyze the complicated brain responses to visual stimuli, we used brain electric source analysis (BESA) as the spatio-temporal multiple source model. In MEG recording, the 1M and 2M components were identified in all subjects. The 1M component was recorded to all kinds of stimuli. The 2M component was clearly identified only to face stimulation in all subjects, but to eyes stimulation in only 3 subjects with a small amplitude. The 2M component was not identified to scrambled face nor hand stimulation. The 2M component was recorded from the right hemisphere in all subjects, but in only 5 of 10 subjects from the left hemisphere. The mean peak latencies of the 1M and 2M components were approximately 132 and 179 ms, respectively. The interpeak latency between 1M and 2M was approximately 47 ms on average but the interindividual difference was large. There was no significant difference of the 2M latency between face with opened eyes and face with closed eyes. The 1M component was generated in the primary visual cortex in the bilateral hemispheres, and the 2M component was generated in the inferior temporal cortex, around the fusiform gyrus. In the EEG recording, face-specific components, positive at the vertex, P200 (Cz), and the negative at the temporal areas, N190 (T5') and N190 (T6'), were clearly recorded. The EEG results were fundamentally compatible with the MEG results. The amplitude of the component recorded from the right hemisphere was significantly larger than that from the left hemisphere. These findings suggest that the fusiform gyrus is considered to play an important role in face perception in humans, and that the right hemisphere is more dominant. Face perception takes place approximately 47 ms after the primary response to visual stimulation in the primary visual cortex, but the period of information transfer to the fusiform gyrus is variable among subjects. Detailed temporal and spatial analyses of the processing of face perception can be achieved with MEG.

Adult↗

Enhanced responsiveness of human extravisual areas to photic stimulation in patients with severely reduced vision.

Lesions in the primary visual cortex induce severe loss of visual perception. Depending on the size of the lesion, the visual field might be affected by small scotomas, hemianopia, or complete loss of vision (cortical blindness). In many cases, the whole visual field of the patient is affected by the lesion, but diffuse light-dark discrimination remains (residual rudimentary vision, RRV). In other cases, a sparing of a few degrees can be found (severely reduced vision, SRV). In a follow-up study, we mapped visually induced cerebral activation of three subjects with SRV using functional magnetic resonance imaging. We were especially interested in the visual areas that would be activated if subjects could perceive the stimulus consciously although information flow from V1 to higher visual areas was strongly reduced or virtually absent. Because subjects were only able to discriminate strong light from darkness, we used goggles flashing intense red light at a frequency of 3 Hz for full visual field stimulation. Besides reduced activation in V1, we found activation in the parietal cortex, the frontal eye fields (FEF), and the supplementary eye fields (SEF). In all patients, FEF activation was pronounced in the right hemisphere. These patterns were never seen in healthy volunteers. In a patient who recovered completely, we observed that extrastriate activation disappeared in parallel with the visual field restitution. This result suggests that damage to the primary visual cortex changes the responsiveness of parietal and extravisual frontal areas in patients with SRV. This unexpected result might be explained by increased stimulus-related activation of attention-related networks.

Adult↗

Parietal neglect and visual awareness.

The last decade has seen a resurgence of interest in the neural correlates of conscious vision, with most discussion focused on the 'blindsight' that can follow damage to primary visual cortex, in the occipital lobe. We suggest that new insights into the neural basis of visual awareness may be gleaned from a different neuropsychological phenomenon, namely visual 'neglect' after injury to regions in the parietal lobe. Neglect provides several revealing contrasts with occipital blindsight. Here we summarise four key findings. First, unlike the deficits caused by damage to primary visual cortex, the loss of awareness in parietal neglect is characteristically not strictly retinotopic. Second, visual segmentation processes are preserved in neglect, and can influence what will reach the patient's awareness. Third, extensive unconscious processing takes place for those stimuli on the neglected side which escape awareness, including some degree of object identification. Finally, parietal damage affects initial stages of motor planning as well as perception. These findings are consistent with recent data on single-cell activity in the monkey brain. They also suggest why areas in the inferior parietal lobe may play a prominent role in visual awareness.

Animals↗

Cortical maps of separable tuning properties predict population responses to complex visual stimuli.

In the earliest cortical stages of visual processing, a scene is represented in different functional domains selective for specific features. Maps of orientation and spatial frequency preference have been described in the primary visual cortex using simple sinusoidal grating stimuli. However, recent imaging experiments suggest that the maps of these two spatial parameters are not sufficient to describe patterns of activity in different orientation domains generated in response to complex, moving stimuli. A model of cortical organization is presented in which cortical temporal frequency tuning is superimposed on the maps of orientation and spatial frequency tuning. The maps of these three tuning properties are sufficient to describe the activity in orientation domains that have been measured in response to drifting complex images. The model also makes specific predictions about how moving images are represented in different spatial frequency domains. These results suggest that the tangential organization of primary visual cortex can be described by a set of maps of separable neuronal receptive field features including maps of orientation, spatial frequency, and temporal frequency tuning properties.

Animals↗

Meridional anisotropy in visual processing: implications for the neural site of the oblique effect.

The contention is examined that the oblique effect, i.e., the well-known performance deficit in detecting orientation difference in oblique lines as compared to vertical and horizontal ones, has its origin in a relative deficiency of neurons with obliquely-oriented receptive fields in the primary visual cortex. Psychophysical observations demonstrate a prominent oblique effect also in visual tasks involving widely-separated elements and other stimuli that would elicit little or no response in oriented neurons in the visual cortex. Conversely, some tasks, e.g. position discrimination, exhibit no oblique effect even with short, high-contrast lines. When the comparison with the reference can be accomplished during a single brief exposure rather than sequential ones, thresholds for orientation differences between adjacent contours in oblique meridians are also elevated compared to those in the vertical and horizontal, but to a lesser extent. In one particular texture discrimination task some but not all observers have a conspicuous oblique effect. The discrimination only of the direction of streaming random dots, not of their speed, is poorer for motions in oblique meridians. The findings imply that the neural locus for the oblique effect is more central than the primary visual cortex.

Contrast Sensitivity↗

Cortical organization in shrews: evidence from five species.

Cortical organization was examined in five shrew species. In three species, Blarina brevicauda, Cryptotis parva, and Sorex palustris, microelectrode recordings were made in cortex to determine the organization of sensory areas. Cortical recordings were then related to flattened sections of cortex processed for cytochrome oxidase or myelin to reveal architectural borders. An additional two species (Sorex cinereus and Sorex longirostris) with visible cortical subdivisions based on histology alone were analyzed without electrophysiological mapping. A single basic plan of cortical organization was found in shrews, consisting of a few clearly defined sensory areas located caudally in cortex. Two somatosensory areas contained complete representations of the contralateral body, corresponding to primary somatosensory cortex (S1) and secondary somatosensory cortex (S2). A small primary visual cortex (V1) was located closely adjacent to S1, whereas auditory cortex (A1) was located in extreme caudolateral cortex, partially encircled by S2. Areas did not overlap and had sharp, histochemically apparent and electrophysiologically defined borders. The adjacency of these areas suggests a complete absence of intervening higher level or association areas. Based on a previous study of corticospinal connections, a presumptive primary motor cortex (M1) was identified directly rostral to S1. Apparently, in shrews, the solution to having extremely little neocortex is to have only a few small cortical subdivisions. However, the small areas remain discrete, well organized, and functional. This cortical organization in shrews is likely a derived condition, because a wide range of extant mammals have a greater number of cortical subdivisions.

Animals↗

Rapid regulation of brain-derived neurotrophic factor mRNA within eye-specific circuits during ocular dominance column formation.

The neurotrophin brain-derived neurotrophic factor (BDNF) has emerged as a candidate retrograde signaling molecule for geniculocortical axons during the formation of ocular dominance columns. Here we examined whether neuronal activity can regulate BDNF mRNA in eye-specific circuits in the developing cat visual system. Dark-rearing throughout the critical period for ocular dominance column formation decreases levels of BDNF mRNA within primary visual cortex, whereas short-term (2 d) binocular blockade of retinal activity with tetrodotoxin (TTX) downregulates BDNF mRNA within the lateral geniculate nucleus (LGN) and visual cortical areas. Brief (6 hr to 2 d) monocular TTX blockade during the critical period and also in adulthood causes downregulation in appropriate eye-specific laminae in the LGN and ocular dominance columns within primary visual cortex. Monocular TTX blockade at postnatal day 23 also downregulates BDNF mRNA in a periodic fashion, consistent with recent observations that ocular dominance columns can be detected at these early ages by physiological methods. In contrast, 10 d monocular TTX during the critical period does not cause a lasting decrease in BDNF mRNA expression in columns pertaining to the treated eye, consistent with the nearly complete shift in physiological response properties of cortical neurons in favor of the unmanipulated eye known to result from long-term monocular deprivation. These observations demonstrate that BDNF mRNA levels can provide an accurate "molecular readout" of the activity levels of cortical neurons and are consistent with a highly local action of BDNF in strengthening and maintaining active synapses during ocular dominance column formation.

Aging↗

Increased glutamate, GABA and glutamine in lateral geniculate nucleus but not in medial geniculate nucleus caused by visual attention to novelty.

This study is concerned with cortico-thalamic neural mechanisms underlying attentional phenomena. Previous results from this laboratory demonstrated that the visual sector of the GABAergic thalamic reticular nucleus is selectively c-fos activated in rats that are naturally paying attention to features of a novel-complex environment, and that this activation is dependent on top-down glutamatergic inputs from the primary visual cortex. By contrast, the acoustic sector of the thalamic reticular nucleus is not activated despite noise generated by exploration and c-fos activation of brainstem acoustic centers (e.g. dorsal cochlear nucleus, inferior colliculus). A prediction of these results is that the levels of the neurotransmitters glutamate and GABA, and the glutamate-related amino acid glutamine, will be increased in the lateral geniculate nucleus (LGN), but not in the medial geniculate nucleus (MGN) of rats that explore a novel-complex environment in comparison to levels of these amino acids in control rats. By means of neurochemical analysis of these amino acids (HPLC) the results of this study confirmed this prediction. The results are consistent with the previously proposed 'focal attention' hypothesis postulating that a focus of attention in the primary visual cortex generates top-down center-surround facilitatory-inhibitory effects on geniculocortical transmission via corticoreticulogeniculate pathways. The results also supports the notion that a main function of corticothalamic pathways to relay thalamic nuclei is attention-dependent modulation of thalamocortical transmission.

Animals↗

Developmental changes in the relationship between type 2 synapses and spiny neurons in the monkey visual cortex.

This study continues an exploration of synaptic development in the primary visual cortex of the monkey (Macaca nemestrina). In a prior study (Mates and Lund, '83a), we observed that type 2 synapses on the cell bodies of spiny stellate neurons of lamina 4C appeared not only to increase in number during early postnatal development but also subsequently decreased during maturation. Using quantitative, stereological electron microscopic methods, we examined the maturation of this synapse population from embryonic day 159 to adult, on spiny stellate neurons of 4C alpha and beta and, for comparison, on pyramidal neurons in upper and lower lamina 6. Tissue was also taken for comparison from two animals reared to 8 weeks of age with binocular eyelid closure from birth. We confirmed that a marked increase and subsequent decrease occurred in this somal type 2 synapse population on both neuron populations. However, due to the infrequency of the smooth dendritic neurons (approximately 5% of the neuron population) giving rise to the type 2 contacts, and due to expansion of the neuropil during maturation increasing intercell distances against constant volume of the type 2 axon arbors, it is concluded that the decrease in type 2 somal synapses may represent a redistribution to dendrites rather than loss from the neuropil. Cells of lamina 4C beta (receiving input from the parvocellular lateral geniculate nucleus-LGN) show a slower initial accumulation of type 2 contacts compared to neurons of lamina 4C alpha (receiving input from magnocellular LGN), or to pyramidal neurons of lamina 6.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Claustral afferents to the rat's visual cortex.

Horseradish peroxidase (HRP) was injected electrophoretically into the rat's primary visual cortex (V1), and three other retinotopically organized cortical areas, anterior medial visual area (AM), posterior medial visual area (PM) and anterior lateral area (AL). While the HRP injections into V1 labelled many neurons in the ipsilateral claustrum, the injections in any of the three other visual areas labelled very few claustral neurons.

Afferent Pathways↗

Gene expression changes and molecular pathways mediating activity-dependent plasticity in visual cortex.

Two key models for examining activity-dependent development of primary visual cortex (V1) involve either reduction of activity in both eyes via dark-rearing (DR) or imbalance of activity between the two eyes via monocular deprivation (MD). Combining DNA microarray analysis with computational approaches, RT-PCR, immunohistochemistry and physiological imaging, we find that DR leads to (i) upregulation of genes subserving synaptic transmission and electrical activity, consistent with a coordinated response of cortical neurons to reduction of visual drive, and (ii) downregulation of parvalbumin expression, implicating parvalbumin-expressing interneurons as underlying the delay in cortical maturation after DR. MD partially activates homeostatic mechanisms but differentially upregulates molecular pathways related to growth factors and neuronal degeneration, consistent with reorganization of connections after MD. Expression of a binding protein of insulin-like growth factor-1 (IGF1) is highly upregulated after MD, and exogenous application of IGF1 prevents the physiological effects of MD on ocular dominance plasticity examined in vivo.

Animals↗

Relationship between contrast adaptation and orientation tuning in V1 and V2 of cat visual cortex.

Previous studies investigating the response properties of neurons in the primary visual cortex of cats and primates have shown that prolonged exposure to optimally oriented, high-contrast gratings leads to a reduction in responsiveness to subsequently presented test stimuli. We recorded from 119 neurons in cat V1 and V2 and found that in a high proportion of cells contrast adaptation also occurs for gratings oriented orthogonal to a neuron's preferred orientation, even though this stimulus did not elicit significant increases in spiking activity. Approximately 20% of neurons adapted equally to all orientations tested and a further 46% showed at least some adaptation to orthogonally oriented gratings, whereas 20% of neurons did not adapt to orthogonal gratings. The magnitude of contrast adaptation was positively correlated with adapting contrast, but was not related to the spiking activity of the cells. Highly direction selective neurons produced stronger adaptation to orthogonally oriented gratings than other neurons. Orientation-related adaptation was correlated with the rate of change of orientation tuning in consecutive cells along electrode penetrations that traveled parallel to the cortical layers. Nonoriented adaptation was most common in areas where orientation preference changed rapidly, whereas orientation-selective adaptation was most common in areas where orientation preference changed slowly. A minority of neurons did not show contrast adaptation (14%). No major differences were found between units in different cortical layers, V1 and V2, or between complex and simple cells. The relevance of these findings to the current understanding of adaptation within the context of orientation column architecture is discussed.

Action Potentials↗