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Decoding cortical neuronal signals: network models, information estimation and spatial tuning.

We have studied the encoding of spatial pattern information by complex cells in the primary visual cortex of awake monkeys. Three models for the conditional probabilities of different stimuli, given the neuronal response, were fit and compared using cross-validation. For our data, a feed-forward neural network proved to be the best of these models. The information carried by a cell about a stimulus set can be calculated from the estimated conditional probabilities. We performed a spatial spectroscopy of the encoding, examining how the transmitted information varies with both the average coarseness of the stimulus set and the coarseness differences within it. We find that each neuron encodes information about many features at multiple scales. Our data do not appear to allow a characterization of these variations in terms of the detection of simple single features such as oriented bars.

Animals↗

A model for the neuronal implementation of selective visual attention based on temporal correlation among neurons.

We propose a model for the neuronal implementation of selective visual attention based on temporal correlation among groups of neurons. Neurons in primary visual cortex respond to visual stimuli with a Poisson distributed spike train with an appropriate, stimulus-dependent mean firing rate. The spike trains of neurons whose receptive fields do not overlap with the "focus of attention" are distributed according to homogeneous (time-independent) Poisson process with no correlation between action potentials of different neurons. In contrast, spike trains of neurons with receptive fields within the focus of attention are distributed according to non-homogeneous (time-dependent) Poisson processes. Since the short-term average spike rates of all neurons with receptive fields in the focus of attention covary, correlations between these spike trains are introduced which are detected by inhibitory interneurons in V4. These cells, modeled as modified integrate-and-fire neurons, function as coincidence detectors and suppress the response of V4 cells associated with non-attended visual stimuli. The model reproduces quantitatively experimental data obtained in cortical area V4 of monkey by Moran and Desimone (1985).

Animals↗

Neurophysiological correlates of perceptual learning in the human brain.

Rapid learning processes are crucial for human object recognition. We report here on the alterations in neurophysiological activity in the human brain induced by repeated presentation of visual stimuli. In psychophysical experiments the percentage of correct responses increased significantly within less than 30 minutes in untrained observers. This stimulus-specific improvement was not carried over to differently oriented stimuli. Similar learning effects were observed in component latencies of evoked potential field distributions. The occurrence of specific potential field configurations reflected perceptual learning. A spatio-temporal activation pattern with steep gradients over the primary visual cortex appeared to be correlated with plasticity in the human visual system.

Adult↗

The effects of physostigmine on the response characteristics of the cat visual evoked potential.

Steady-state pattern visual evoked potentials were recorded from the surface of the cat primary visual cortex before and after the intravenous administration of physostigmine, an agent that blocks the enzyme responsible for the breakdown of synaptically released acetylcholine. Under pentobarbital anesthesia, physostigmine increased the amplitude and changed the phase of the second response harmonic of the visual evoked potential, whereas the amplitude and phase of the fourth harmonic were not affected. These effects persisted for 15 to 45 minutes and were blocked by prior treatment with scopolamine or atropine. In addition, scopolamine or atropine administered 5 to 10 minutes after physostigmine returned the visual evoked potential to the baseline state. In comparison, when nitrous oxide was used, physostigmine caused a marked reduction in visual evoked potential amplitude, an effect that was reversed by subsequent atropine. These results indicate that the cholinergic system influences the visual evoked potential via a muscarinic pathway and that this influence is strongly affected by the anesthetic regimen used.

Acetylcholine↗

Decreases in peripheral-type benzodiazepine receptors in postmortem brains of chronic schizophrenics.

We measured the peripheral-type benzodiazepine receptors (PBRs), a marker of gliosis, in 26 brain areas (cerebral cortex, thalamus and extrapyramidal system) of the postmortem brains of 13 chronic schizophrenics and 10 controls, using [3H] PK 11195 as a ligand for the receptor assay. The specific [3H] PK 11195 binding was significantly decreased in three brain areas (superior parietal cortex, primary visual area and putamen) of schizophrenics, although there were no changes in the binding in the other brain areas. Scatchard analysis revealed that there were decreases in both the Bmax and Kd of [3H] PK 11195 binding in the brain areas. These results were almost in accordance with a number of neuropathological studies reporting that there was no change or reduction in glial cells in the brain regions of schizophrenics and suggested that the decreased density of PBRs in the brain may be involved in the pathophysiology of schizophrenia, associated with reduced production of neurosteroids coupled to PBRs.

Adult↗

Tuning of striate neurons to cross-shaped figures in conditions of local blockade of intracortical inhibition.

The recently observed selective sensitivity to cross-shaped and angular figures was studied in 85 primary visual cortex (field 17) neurons in cats before and after local blockade of GABA(A)ergic inhibition by microiontophoretic application of the GABA antagonist bicuculline. Two opposite effects were seen: half of the neurons studied showed decreases or complete loss of sensitivity to crosses, and a third of the cells showed increases or the appearance of sensitivity to crosses. These data provide evidence for significant roles for intracortical inhibition in providing sensitivity to crosses and intersecting lines in two types of visual cortex neurons, the effects on these two types of neuron being opposite.

Action Potentials↗

Visual corticopontine projections in the guinea pig: an autoradiographic study.

The goal of this study is to characterize the anatomical organization of the visual cortical output to the basal pontine nuclei in the guinea pig. Data from the literature show that guinea pigs exhibit different optokinetic oculomotor behaviors with respect to rats and rabbits. Namely, they present a fast rise in eye movement velocity at stimulus onset and a better performance in monocular horizontal stimulation. Possible differences in the visual corticopontocerebellar pathway might explain these peculiarities. The pontine projections from the primary visual cortex were studied with the method of the anterograde axonal transport of [3H]leucine. The terminal labeling forms prominent patches, ipsilaterally to the cortical injection, throughout the rostrocaudal extent of the pontine nuclei, predominantly in the dorsolateral region. At the intermediate rostrocaudal level, some foci of labeling are found ventrolaterally as well. Sparse fields are present also in the medial pontine nuclei and in the nucleus reticularis tegmenti pontis, but only when the injection site extends to secondary visual areas, either lateral or medial. The present description of the corticopontine projections in guinea pigs is in substantial agreement with the projections previously described in rats, with a few differences, namely: (1) the recipient area extends more caudally; (2) secondary visual areas project to the nucleus reticularis tegmenti pontis.

Animals↗

Characteristics of surround inhibition in cat area 17.

The effects of stimuli falling outside the 'classical receptive field' and their influence on the orientation selectivity of cells in the cat primary visual cortex are still matters of debate. Here we examine the variety of effects of such peripheral stimuli on responses to stimuli limited to the receptive field. We first determined the extent of the classical receptive field by increasing the diameter of a circular patch of drifting grating until the response saturated or reached a maximum, and by decreasing the diameter of a circular mask in the middle of an extended grating, centred on the receptive field, until the cell just began to respond. These two estimates always agreed closely. We then presented an optimum grating of medium-to-high contrast filling the classical receptive field while stimulating the surround with a drifting grating that had the same parameters as the central stimulus but was varied in orientation. For all but five neurons (of 37 tested), surround stimulation produced clear suppression over some range of orientations, while none showed explicit facilitation under these conditions. For 11 cells (34% of those showing suppression), the magnitude of suppression did not vary consistently with the orientation of the surround stimulus. In the majority of cells, suppression was weakest for a surround grating oriented orthogonal to the cell's optimum. Nine of these cells (28%) exhibited maximum inhibition at the optimum orientation for the receptive field itself, but for 12 cells (38%) there was apparent 'release' from inhibition for surround gratings at or near the cell's optimum orientation and direction, leaving inhibition either maximal at angles flanking the optimum (9 cells) or broadly distributed over the rest of the orientation range (3 cells). This implies the existence of a subliminal facilitatory mechanism, tightly tuned at or near the cell's optimum orientation, extending outside the classical receptive field. For just two cells of 13 tested the preferred orientation for a central grating was clearly shifted towards the orientation of a surrounding grating tilted away from the cell's optimum. The contrast gain for central stimulation at the optimal orientation was measured with and without a surround pattern. For nine of 25 cells tested, surround stimulation at the cell's optimum orientation facilitated the response to a central grating of low contrast (< or =0.1) but inhibited that to a higher-contrast central stimulus: the contrast-response gain is reduced but the threshold contrast is actually decreased by surround stimulation. Hence the receptive field is effectively larger for low-contrast than for high-contrast stimuli. Inhibition from the periphery is usually greatest at or around the cell's optimum, while suppression within the receptive field has been shown to be largely non-selective for orientation. Inhibition by orientations flanking the optimum could serve to sharpen orientation selectivity in the presence of contextual stimuli and to enhance orientational contrast; and it may play a part in orientation contrast illusions.

Animals↗

Self-organizing maps for visual feature representation based on natural binocular stimuli.

We model the stimulus-induced development of the topography of the primary visual cortex. The analysis uses a self-organizing Kohonen model based on high-dimensional coding. It allows us to obtain an arbitrary number of feature maps by defining different operators. Using natural binocular stimuli, we concentrate on discussing the orientation, ocular dominance, and disparity maps. We obtain orientation and ocular dominance maps that agree with essential aspects of biological findings. In contrast to orientation and ocular dominance, not much is known about the cortical representation of disparity. As a result of numerical simulations, we predict substructures of orientation and ocular dominance maps that correspond to disparity maps. In regions of constant orientation, we find a wide range of horizontal disparities to be represented. This points to geometrical relations between orientation, ocular dominance, and disparity maps that might be tested in experiments.

Algorithms↗

Cross-modal plasticity after monocular enucleation of the adult rabbit.

The mature brain undergoes compensatory reorganization of the primary visual cortex (V1) in response to retinal lesions. This study demonstrates that V1 also supports cross-modal reorganization by observing an increase in tactile responses in V1 after monocular enucleation of the adult rabbit. The proportion of tactile-responsive V1 neurons increased from 0% to 31%, in an area of cortex equivalent to 40 degrees of visual space. Retrograde fiber-tracing analysis suggests that intracortical connections from association areas may underlie these novel responses. Cortical plasticity of this kind may be involved in recovery from sensory system damage and could provide an enhanced sense of touch to the blind.

Action Potentials↗

The attentional repulsion effect in perception and action.

The attentional repulsion effect refers to the perceived displacement of a Vernier stimulus in a direction that is opposite to a brief peripheral cue. The twofold purpose of the present study was to: (1) replicate the perceptual effect using a Vernier discrimination task, and (2) determine whether the effect would also affect action using a guided localization task. A perceptual attentional repulsion effect was found in experiment 1 and a similar effect was found in experiment 2, with a computer mouse localization task, and in experiment 3, with a guided limb localization task (in both cases pointing responses were biased in the direction opposite to that of the cue). These findings suggest that the attentional repulsion effect occurs early in visual processing, probably affecting the receptive fields of the position-coding units in primary visual cortex before "object-perception" and "object-action" information is segregated into separate pathways.

Analysis of Variance↗

Neurons in the posteromedial lateral suprasylvian area of the cat are sensitive to binocular positional depth cues.

Single units in the posteromedial lateral suprasylvian area of the cat are known to be very sensitive to movement. A proportion of these cells can encode movement in depth, but it is unclear whether posteromedial lateral suprasylvian cells only rely upon motion cues to evaluate stimulus depth or whether they can also code for spatial cues. The present study aims at assessing the sensitivity to spatial disparity of binocular cells, in the postero-medial lateral suprasylvian area, in order to determine whether these units are tuned to positional depth cues. A total of 126 single cells located in the posteromedial lateral suprasylvian area of anesthetized, paralyzed cats were examined. As recordings were performed in the central visual field representation, receptive fields were small. A third of the receptive fields were surrounded by an inhibitory region and almost three-quarters of the cells were direction-selective. Most cells (110/114) were binocular, and a large proportion of single neurons responded to stimuli appearing on the fixation plane by increasing (tuned excitatory cells, 43%) or decreasing (tuned inhibitory cells, 14%) their response rate. A smaller proportion of cells increased their firing rate in response to crossed (near cells, 10%) or uncrossed (far cells, 6%) spatial disparities, hence demonstrating respective preference for stimuli presumably appearing in front of or behind the fixation plane. As compared to primary visual cortex, the proportion of disparity-sensitive cells in posteromedial lateral suprasylvian area is similar, but selectivity is significantly coarser. As the posteromedial lateral suprasylvian area can code for both spatial and temporal aspects of stimuli, this area might be involved in the spatiotemporal integration of depth cues, a process that may also participate in the control of accommodation and vergence.

Animals↗

Column spacing in normal and visually deprived monkeys.

A recent model for the development of the pattern of eye-dominance domains in primary visual cortex predicts that stimulus conditions during early visual life determine the spacing (or periodicity) of ocular dominance columns (ODC). The model predicts that normal binocular visual experience consists of highly correlated binocular stimulation and leads to relatively narrow ODC spacing, while abnormal binocular visual stimulation attendant with strabismus consists of non-correlated, incoherent, and asynchronous stimulation and leads to wider than normal ODC spacing. Evidence in support of the model has been presented for strabismus in the kitten. We tested the predictions of the model in normal monkeys and others subjected to various forms of abnormal visual experience during infancy. We identified and measured the inter-column spacing (or periodicity) in the V1 cortex of 19 adult monkeys (M. mulatta) using the cytochrome-oxidase (CO) histochemical method. There were no significant differences in the V1 inter-column spacing between normal adult monkeys (n=5) and other adult monkeys having had monocular-form deprivation (n=5), experimental anisometropia (n=5), or experimental strabismus (n=4) early in life. The quality of early binocular visual experience is not a significant determinant of the inter-column spacing in primate V1 cortex. Therefore, the model predicting an increase in the ODC periodicity with strabismus is not supported.

Analysis of Variance↗

Binocular interactions and spatial disparity sensitivity in the superior colliculus of the Siamese cat.

In Siamese cats, a genetically determined massive misrouting of retinal ganglion cells toward the contralateral hemisphere, as well as an accompanying strabismus, is believed to underlie the extreme paucity of binocular cells in the primary visual cortex. However, binocular cells have been shown to be present in more important numbers at the collicular level. The present study aims at investigating binocular interactions and sensitivity to spatial disparity in the superior colliculus of the Siamese cat. The activity of single units was recorded in the superficial layers of paralyzed and anesthetized Siamese cats. Although most collicular cells were monocularly driven, a significant proportion could be driven through both eyes (34/216 or 16%). Upon isolation of a binocular cell, the receptive fields were separated, then simultaneously stimulated with two light bars. A temporal delay was introduced between the arrival of the bars in the receptive fields to generate spatial disparities (-3 degrees to +3 degrees, in 0.5 degrees or 1 degree steps). Results showed that some binocular cells presented disparity tuning profiles similar to the tuned excitatory (12/34), tuned inhibitory (2/34), near (2/34) and far (3/34) cells found at various cortical levels in the normal cat. These interactions might allow for coarse binocular fusion as well as play a role in the initiation of vergence and the fixation of the eyes upon the appropriate plane of vision.

Animals↗

Neuronal responses from beyond the classic receptive field in V1 of alert monkeys.

Responses of primary visual cortex (V1) neurons to stimuli inside the classic receptive field (CRF) can be modulated by stimuli outside the CRF. We recently reported that responses of most V1 neurons to a line in the CRF center are inhibited by large surround-stimuli and that this modulation is stimulus selective. Here we report that a significant proportion of V1 neurons in alert monkeys respond directly to stimuli outside the CRF with very long latency and much reduced selectivity. When surround stimuli are presented alone, three response patterns can be distinguished in 153 single- or multiunits tested: (1) 31.4% have no significant response; (2) 50.3% show excitatory responses that are significantly higher than spontaneous activity. The average latency of these responses is about 145 ms, 2-3 times longer than center responses; (3) 18.3% show suppressed spontaneous activity after stimulus onset. The direct surround responses are found to be only weakly selective for the orientation of contextual lines, and not selective for other contextual patterns tested. While the outburst of responses to stimuli within the CRF is not affected by reducing stimulus duration from 500 ms to 50 ms, late excitatory surround responses are virtually eliminated. We propose that the late excitatory surround responses to extra-CRF stimulation alone are the reflection of feedback from higher cortical areas and may contribute to reduced contextual inhibition of cells in V1. This could play a role in figure-ground segregation.

Animals↗

Lewy body and Alzheimer pathology in a family with the amyloid-beta precursor protein APP717 gene mutation.

Mutations in the amyloid precursor protein (APP) gene cause one form of early onset familial Alzheimer's disease (AD). One such family has been studied genetically and neuropathologically and represents the basis of the present report. Four siblings with the APP717 Val to Ile mutation, aged 59, 65, 61 and 64 years, apolipoprotein E (APOE) genotyped 2,4 (first three) and 2,3 respectively, had severe AD, Braak stage VI with frequent neurofibrillary tangles in the primary visual cortex, Brodmann area 17. The first one also met McKeith criteria for the limbic stage of dementia with Lewy bodies but did not have substantia nigra Lewy bodies. The second two met McKeith criteria for the neocortical stage of dementia with Lewy bodies and both had substantia nigra Lewy bodies. The fourth had AD but no Lewy bodies. A cousin without the APP717 mutation who was APOE 3, 4, developed dementia at age 60 and died at age 75. She had severe cerebrovascular atherosclerosis, less severe AD, Braak stage V, with sparing of area 17. She also had Lewy bodies in the substantia nigra and in the cortex and met McKeith criteria for neocortical stage of dementia with Lewy bodies. Extrapyramidal features were present in all five. Lewy bodies have been described in 53% of reported autopsies on individuals with the APP717 Val to Ile mutation coincident with dementia and AD neuropathologic changes. These observations suggest an association between the chromosome 21 APP mutation and Lewy body formation, possibly mediated by other environmental or genetic factors.

Aged↗

Fatigue in patients with lupus is not associated with disturbances in cerebral blood flow as detected by SPECT.

OBJECTIVES: Fatigue is a common complaint in patients with systemic lupus erythematosus (SLE). We investigated whether focal or general disturbances of cerebral blood flow (CBF), as assessed by SPECT, were associated with the presence of fatigue in an unselected group of SLE patients. METHODS: Fifty-six patients were included. Mean age was 47.5 years (+/-12.7), mean disease duration 14.7 years (+/-8.9), and disease activity measured by SLE disease activity index (SLEDAI) was 5.7 (+/-5.4). Fatigue was assessed by the Fatigue Severity Scale (FSS) and CBF by Tc-99m-hexamethyl propylamine oxime (HMPAO)-SPECT. The images were read and processed quantitatively by a computer program using the primary visual cortex as reference region and > 15% CBF deviation as definition of abnormality. RESULTS: The mean FSS score was 4.6 (+/-1.8). SPECT revealed focal CBF disturbances in 17 patients (30.4 %). Generalized symmetrical CBF reductions were present in 32 patients (57.1 %). There were no significant associations between CBF disturbances in any region of the brain and the degree of fatigue. CONCLUSIONS: Fatigue in SLE patients is not related to focal or general CBF disturbances. Therefore, factors that do not influence blood flow seem responsible for the fatigue phenomenon.

Adult↗

Chronically implanted epidural electrodes in Göttinger minipigs allow function tests of epiretinal implants.

BACKGROUND: To test the function of implantable devices for electrical stimulation of the retina, long-term registration tests of cortical-evoked potentials are required. Skin electrodes are not appropriate to provide representative recordings, due to the voluminous pneumatic frontal sinus of minipigs. Therefore, epidural electrodes were permanently implanted in minipigs and tested with visual and electrical retinal stimulation. The present study describes long-term recordings of electrically evoked cortical potentials in minipigs. METHODS: Three-channel silver-silver chloride electrodes were fixed on the epidura dorsal to the primary visual cortex of one hemisphere in Göttinger minipigs. Repeated light stimulation was performed and platinum-polyimide film multielectrode arrays were implanted on the retina for electrical stimulation and were later removed. Cortical potentials were recorded after stimulation with short biphasic charge-balanced currents. RESULTS: For up to 18 months, the implanted epidural electrodes allowed recording of visual cortex potentials evoked by visual or electrical retina stimulation. Small changes of response amplitudes in subsequent experiments indicated a stable location and recording properties of the cortex electrodes. Visual stimulation often yielded stronger responses for the contralateral eye. Superthreshold electrical retina stimulation evoked cortical responses with less delay compared to visual stimulation. We found threshold currents of 50 microA for charge-balanced biphasic trains of current impulses. Postoperative examination showed an attached and unchanged retina. CONCLUSIONS: The minipig model is appropriate for the implantation of epiretinal stimulation electrodes and for the long-term tests of retinal implants by recording of cortical responses with chronically implanted epidural electrodes.

Animals↗