Cat Area Research Articles

Functional Imaging of Primary Visual Cortex Using Flavoprotein Autofluorescence

Neuronal autofluorescence, which results from the oxidation of flavoproteins in the electron transport chain, has recently been used to map cortical responses to sensory stimuli. This approach could represent a substantial improvement over other optical imaging methods because it is a direct (i.e., nonhemodynamic) measure of neuronal metabolism. However, its application to functional imaging has been limited because strong responses have been reported only in rodents. In this study, we demonstrate that autofluorescence imaging (AFI) can be used to map the functional organization of primary visual cortex in both mouse and cat. In cat area 17, orientation preference maps generated by AFI had the classic pinwheel structure and matched those generated by intrinsic signal imaging in the same imaged field. The spatiotemporal profile of the autofluorescence signal had several advantages over intrinsic signal imaging, including spatially restricted fluorescence throughout its response duration, reduced susceptibility to vascular artifacts, an improved spatial response profile, and a faster time course. These results indicate that AFI is a robust and useful measure of large-scale cortical activity patterns in visual mammals.

Standing Waves and Traveling Waves Distinguish Two Circuits in Visual Cortex

The visual cortex represents stimuli through the activity of neuronal populations. We measured the evolution of this activity in space and time by imaging voltage-sensitive dyes in cat area V1. Contrast-reversing stimuli elicit responses that oscillate at twice the stimulus frequency, indicating that signals originate mostly in complex cells. These responses stand clear of the noise, whose amplitude decreases as 1/frequency, and yield high-resolution maps of orientation preference and retinotopy. We first show how these maps are combined to yield the responses to focal, oriented stimuli. We then study the evolution of the oscillating activity in space and time. In the orientation domain, it is a standing wave. In the spatial domain, it is a traveling wave propagating at 0.2-0.5 m/s. These different dynamics indicate a fundamental distinction in the circuits underlying selectivity for position and orientation, two key stimulus attributes.

Effect of binocular retinal lesions on CRMP2 and CRMP4 but not Dyn I and Syt I expression in adult cat area 17

Removal of retinal input from a restricted region of adult cat visual cortex leads to a substantial reorganization of the retinotopy within the sensory-deprived cortical lesion projection zone (LPZ). Still little is known about the molecular mechanisms underlying this cortical map reorganization. We chose two members of the collapsin response mediator protein (CRMP) family, CRMP2 and CRMP4, because of their involvement in neurite growth, and compared gene and protein expression levels between normal control and reorganizing visual cortex upon induction of central retinal lesions. Parallel analysis of Dynamin I (Dyn I) and Synaptotagmin I (Syt I), two molecules implicated in the exocytosis-endocytosis cycle, was performed because changes in neurotransmitter release have been implicated in cortical plasticity. Western blotting and real-time polymerase chain reaction revealed a clear time-dependent effect of retinal lesioning on CRMP2 and CRMP4 expression, with maximal impact 2 weeks post-lesion. Altered CRMP levels were not a direct consequence of decreased visual activity in the LPZ as complete surgical removal of retinal input to one hemisphere had no effect on CRMP2 or CRMP4 expression. Thus, CRMP expression is correlated to cortical reorganization following partial deafferentation of adult visual cortex. In contrast, Dyn I and Syt I were not influenced and thereby do not promote exocytosis-endocytosis cycle modifications in adult cat cortical plasticity.

The effects of reversible inactivation of postero-temporal visual cortex on neuronal activities in cat's area 17

‘Spontaneous’ and visually evoked action potentials were recorded from single neurons in cytoarchitectonic area 17 (striate cortex, area V1) of anaesthetized and immobilized cats, prior to, during and after brief reversible inactivation of the ipsilateral postero-temporal visual (PTV) cortex (presumed homologue of primate inferotemporal cortex). Inactivation of PTV cortex resulted: 1) in significant changes in the response magnitude (mostly a reduction) to optimal and/or sub-optimal visual stimuli in over 55% of area 17 cells and 2) significant changes (usually a reduction) in the ‘spontaneous’ (background) activity of about two-thirds of the cells in which inactivation of PTV cortex significantly affected the magnitude of responses to optimal stimuli. In over 85% of the significantly affected area 17 cells, rewarming PTV cortex to normal temperature (36 °C) resulted in the recovery of both the magnitude of responses and the background activity to levels not significantly different from pre-inactivation levels. Irrespective of the significance of changes in the magnitude of responses, in a substantial proportion of area 17 cells, inactivation of PTV cortex resulted in changes in some receptive field characteristics. Thus, there were substantial (20% or more) changes in orientation tuning widths (in over a quarter of the sample) and/or direction selectivity indices (in about a third of the sample). Thus, the feedback signals originating from PTV cortex, like signals originating from some other ‘higher-order’ visual cortical areas exert a clear modulatory influence on the responsiveness, background activity and some receptive field properties of neurons in the ipsilateral area 17.

Pattern motion selectivity in population responses of area 18

It is commonly believed that the complexity of visual stimuli represented by individual neurons increases towards higher cortical areas. However, even in early visual areas an individual neuron's response is influenced by stimuli presented outside its classical receptive field. Thus, it has been proven difficult to characterize the coding of complex stimuli at the level of single neurons. We therefore investigated population responses using optical imaging in cat area 18 to complex stimuli, plaids. Plaid stimuli are composed of two superimposed gratings moving in different directions. They may be perceived as either two separate surfaces or as a global pattern moving in intermediate direction to the components' direction of motion. We found that in addition to activity maps representing the individual components' motion, plaid stimuli produced activity distributions matching the predictions from a pattern-motion model in central area 18. Thereby, relative component- and pattern-like modulations followed the degree of psychophysical pattern bias in the stimulus. Thus, our results strongly indicate that area 18 exhibits a substantial response to pattern-motion signals at the population level suggesting the presence of intrinsic or extrinsic mechanisms that allow for integration of motion responses from far outside the classical receptive field.

Neural Basis for Stereopsis from Second-Order Contrast Cues

Humans and animals use visual cues such as brightness and color boundaries to identify objects and navigate through environments. However, even when these cues are not available, we can effortlessly perform these tasks by using second-order cues such as contrast variation (envelope) of patterns on surfaces. Previously, numerous psychophysical studies examined properties of binocular depth processing based on the contrast-envelope cues and suggested the existence of a stereo system that uses these cues. However, its physiological substrate has not been identified yet. Here, we show that a subset of cortical neurons in cat area 18 show binocular interactions for the contrast-envelope stimuli. These neurons are capable of representing a variety of depths in the three-dimensional space based on the information available from contrast cues alone. Furthermore, these neurons show similar disparity-tuning curves for borders defined by both luminance and contrast cues. This cue-invariant tuning is consistent with a linear binocular convergence model for monocular luminance and contrast-envelope processing pathways.

Spatiotemporal Specificity of Neuronal Activity Directs the Modification of Receptive Fields in the Developing Retinotectal System

The precise temporal relation between pre- and postsynaptic activity modulates the strength of synaptic connections. Despite its extensive characterization in vivo and in vitro, the degree to which spike timing-dependent plasticity can shape receptive field properties is unclear. We use in vivo patch-clamp recordings of tectal neurons in developing Xenopus tadpoles to control the precise timing of action potentials with respect to the arrival of a subset of visual inputs evoked by local light stimulation on the retina. The pattern of visual inputs onto a tectal neuron was tracked over time by rapid reverse correlation mapping of receptive fields. Spike timing-dependent potentiation or depression of a subset of synapses reliably shifts the spatial receptive fields toward or away from the trained subregion of visual space, respectively. These results demonstrate that natural patterns of activity evoked by sensory stimuli play an instructive role in the developing nervous system.

The layout of functional maps in area 18 of strabismic cats

Strabismus (or squint) is both a well-established model for developmental plasticity and a frequent clinical symptom. To analyze experience-dependent plasticity of functional maps in the brain we used optical imaging of intrinsic signals to visualize both orientation and ocular dominance domains in cat area 18. In strabismic animals, iso-orientation domains exhibited a pinwheel-like organization, as previously described for area 18 of normally raised animals and for area 17 of both normally raised and strabismic cats. In area 18, mean pinwheel density was similar in the experimental (2.2 pinwheel centers per mm 2 cortical surface) and control animals (2.3/mm 2 in normally raised animals), but significantly lower than in area 17 of both normally raised and strabismic cats (2.7–3.4/mm 2). A comparison of orientation and ocular dominance domains revealed that iso-orientation domains were continuous across the borders of ocular dominance domains and tended to cross these borders at steep angles. Thus, the orientation map does not seem to be modified by experience-dependent changes in afferent activity. Together with our recent observation that strabismus does not enhance the segregation of ocular dominance domains in cat area 18, the present data indicate that the layout of functional maps in area 18 is less susceptible to experience-dependent manipulations than in area 17.

Basal Forebrain Cholinergic Input Is Not Essential for Lesion-Induced Plasticity in Mature Auditory Cortex

The putative role of the basal forebrain cholinergic system in mediating lesion-induced plasticity in topographic cortical representations was investigated. Cholinergic immunolesions were combined with unilateral restricted cochlear lesions in adult cats, demonstrating the consequence of cholinergic depletion on lesion-induced plasticity in primary auditory cortex (AI). Immunolesions almost eliminated the cholinergic input to AI, while cochlear lesions produced broad high-frequency hearing losses. The results demonstrate that the near elimination of cholinergic input does not disrupt reorganization of the tonotopic representation of the lesioned (contralateral) cochlea in AI and does not affect the normal representation of the unlesioned (ipsilateral) cochlea. It is concluded that cholinergic basal forebrain input to AI is not essential for the occurrence of lesion-induced plasticity in AI.

Neuronal Mechanisms for Illusory Brightness Perception in Humans

Biological visual systems are extraordinarily capable of recovering the shape and brightness of objects from sparse and fragmentary information. Using functional magnetic imaging, we show that two associative areas of the dorsal pathway--in the caudal region of the intrapariatal sulcus and in the lateral occipital sulcus--respond specifically to the Craik-O'Brien-Cornsweet illusion generated by high-pass filtered edges. Other visual areas, including primary visual cortex, also respond strongly to the retinotopic location of the edge, but these areas respond equally well to a line of matched contrast and detectability, rather than specifically to the brightness illusion. The reconstruction of surface and/or its brightness seems to be achieved by associative areas from the information about visual features provided by the primary visual cortices, even where there is no physical difference in luminance.

Optical imaging in cat area 18: Strabismus does not enhance the segregation of ocular dominance domains

While early-onset strabismus leads to clearly segregated domains of the left and the right eye in cat primary visual cortex (area 17), far less is known about experience-dependent plasticity of ocular dominance in area 18. We therefore used optical imaging of intrinsic signals to analyze the influence of strabismus on cortical maps in cat area 18. Monocular visual stimulation of the left and right eye with moving square wave gratings of four different orientations induced patchy activity maps. Unlike our previous observations in cat area 17, the monocular activity maps in area 18 of strabismic cats were rather similar so that functional ocular dominance domains were not clearly segregated. Imaging of the 17/18 border region confirmed this observation and revealed a sudden change in the segregation of the left and right eye domains across the border. Our results demonstrate that modified visual input can have different consequences for different visual areas: while the decorrelation of activity between the two eyes (as induced by strabismus) clearly enhances the segregation of ocular dominance domains in cat area 17, area 18 does not show this effect although electrophysiological studies have confirmed that the percentage of binocularly driven neurons is as reduced as in area 17.

The anterior ectosylvian visual area of the ferret: a homologue for an enigmatic visual cortical area of the cat?

The present paper describes the results of electrophysiological mapping experiments focused on the anterior limb of the ectosylvian gyrus of the ferret (Mustela putorius). The aim was to determine if the ferret possessed a homologous cortical area to the anterior ectosylvian visual area (AEV) of the domestic cat, but not clearly delineated in any other mammal studied to date. We were able to gather data on the visuotopic organization of a region that we consider to be a homologue of cat AEV. The visual map in this area showed a distinct visuotopic organization and covered a large extent of the visual field. Within the ferret AEV there were clusters of bimodal recording sites (somato-visual and audio-visual) that were located adjacent to surrounding unimodal cortical areas (such as the second somatosensory area and primary and secondary auditory areas). The ferret AEV, like that of the cat, was topographically isolated from other visual cortical areas by intervening auditory and somatosensory areas. Taken together these features suggest that the region described here as AEV in the ferret is indeed a direct homologue of the previously described cat AEV.

Does Neuronal Synchrony Underlie Visual Feature Grouping?

Previous research suggests that synchronous neural activity underlies perceptual grouping of visual image features. The generality of this mechanism is unclear, however, as previous studies have focused on pairs of neurons with overlapping or collinear receptive fields. By sampling more broadly and employing stimuli that contain partially occluded objects, we have conducted a more incisive test of the binding by synchrony hypothesis in area MT. We find that synchrony in spiking activity shows little dependence on feature grouping, whereas gamma band synchrony in field potentials can be significantly stronger when features are grouped. However, these changes in gamma band synchrony are small relative to the variability of synchrony across recording sites and do not provide a robust population signal for feature grouping. Moreover, these effects are reduced when stimulus differences nearby the receptive fields are eliminated using partial occlusion. Our findings suggest that synchrony does not constitute a general mechanism of visual feature binding.

Contrast response in visual cortex: Quantitative assessment with intrinsic optical signal imaging and neural firing

While previous studies showed that intrinsic optical signals spatially correspond with electrophysiological responses in mammalian visual cortex, the quantitative correspondence of their response strengths is open to question. Measurement of both signals' strength as functions of visual stimulus contrast provides an opportunity for quantitative comparison. Towards that end, the spatial and temporal properties of the optical signal impose important constraints upon quantification of its strength. We used intrinsic optical signal imaging and single unit recording to measure responses to drifting gratings at contrasts ranging from 10–80% in cat area 18. We calculated the average difference images for pairs of oppositely moving, or orthogonally oriented, gratings at each contrast and evaluated three different methods for quantifying optical signal strength. After about 2.5 s, the spatial patterns of optical images and the time course of their strength were contrast-invariant. This “space–time-contrast separability” for optical response implies a spatial uniformity of the optical contrast response functions, provides an objective basis to guide temporal averaging of optical signals, and validates a scalar metric of optical signal strength. Optically measured contrast response functions increase monotonically and saturate at high contrasts, qualitatively resembling those from single units. However, quantitative comparison reveals a nonlinear relationship with neural firing, such that the optical response reaches half of its maximum when the neural response has reached only around 20% of its maximum. This relationship suggests that intrinsic optical signals are relatively more sensitive to weak signals than neural firing.

Role of synaptic and intrinsic membrane properties in short-term receptive field dynamics in cat area 17.

We examined the mechanisms through which the prolonged presentation of either a high-contrast stimulus or an artificial scotoma [equivalent to the stimulation of the receptive field (RF) surround] induces changes in the RF properties of neurons intracellularly recorded in cat primary visual cortex. Discharge and synaptic RFs were quantitatively characterized using bright and dark bars randomly flashed in various positions. Compared with the lack of stimulation (0% contrast for 15-30 s), stimulation with high-contrast sine-wave gratings (15-30 s) was followed by a strong reduction in gain and a weak but significant reduction in width of spike discharge RFs. These reductions were accompanied by a membrane potential hyperpolarization, a decrease of synaptic RF width, and varying changes of synaptic RF gain. Passive hyperpolarization by DC injection also produced significant reduction in the width and gain of discharge RF. Mimicking, in single neurons, high-contrast stimulation with high-intensity current injection also induced a membrane potential hyperpolarization, whose amplitude was correlated with discharge RF gain and width changes. Recovery from adaptation to high-contrast stimulation during the period of gray screen or scotoma presentation was associated with an increase in gain and discharge RF size. Stimulation of the RF surround with an artificial scotoma did not have any additional aftereffects over those of adaptation to a gray screen, indicating that the contraction and expansion of RF gain and size are attributable to intrinsic and synaptic mechanisms underlying adaptation and de-adaptation to strong visual stimuli.

Light-induced Fos expression in phosphate-activated glutaminase- and neurofilament protein-immunoreactive neurons in cat primary visual cortex

Previous double-stainings in the cat visual cortex [E. Van der Gucht, S. Clerens, K. Cromphout, F. Vandesande, L. Arckens, Differential expression of c- fos in subtypes of GABAergic cells following sensory stimulation in the cat primary visual cortex, Eur. J. Neurosci. 16 (2002) 1620–1626] showed that a minority of Fos-immunoreactive nuclei was located in distinct subclasses of inhibitory neurons following sensory stimulation. This report describes double-stainings between Fos and phosphate-activated glutaminase (PAG) or Fos and neurofilament protein (SMI-32) revealing that, following a short-term visual experience, Fos is also expressed in neurochemically distinct subpopulations of non-GABAergic, pyramidal neurons in supra- and infragranular layers of cat area 17.

A quantitative map of the circuit of cat primary visual cortex.

We developed a quantitative description of the circuits formed in cat area 17 by estimating the "weight" of the projections between different neuronal types. To achieve this, we made three-dimensional reconstructions of 39 single neurons and thalamic afferents labeled with horseradish peroxidase during intracellular recordings in vivo. These neurons served as representatives of the different types and provided the morphometrical data about the laminar distribution of the dendritic trees and synaptic boutons and the number of synapses formed by a given type of neuron. Extensive searches of the literature provided the estimates of numbers of the different neuronal types and their distribution across the cortical layers. Applying the simplification that synapses between different cell types are made in proportion to the boutons and dendrites that those cell types contribute to the neuropil in a given layer, we were able to estimate the probable source and number of synapses made between neurons in the six layers. The predicted synaptic maps were quantitatively close to the estimates derived from the experimental electron microscopic studies for the case of the main sources of excitatory and inhibitory input to the spiny stellate cells, which form a major target of layer 4 afferents. The map of the whole cortical circuit shows that there are very few "strong" but many "weak" excitatory projections, each of which may involve only a few percentage of the total complement of excitatory synapses of a single neuron.

Functional segregation of plural regions representing cardinal contours in cat primary visual cortex.

Our previous data based on an imaging study suggested that, in cat area 17, the representations of cardinal orientations overlap less than the representation of their nearby angles. The purpose of this study was to further investigate the underlying single-cell properties. Optical imaging was performed first to map the cortical regions corresponding to the four principal contours, the two cardinals and the two obliques. The cortical region activated by a principal orientation but not by the +10 degrees or -10 degrees neighbouring angles, namely the area with optically relative independent orientation selectivity (RIOS), was mapped together with the regions that overlapped with the +10 degrees and/or -10 degrees neighbouring angles (non-RIOS). Electrode penetrations were targeted to the RIOS and non-RIOS regions in each of the four orientations. A comparison between the RIOS and the non-RIOS regions documented a significantly higher percentage of cells with the orientation preference of the cardinal orientations in the cardinal RIOS region than that seen in the other regions. Additionally, the difference in the tuning width of cells between the RIOS and non-RIOS in the cardinal region was significantly larger than the difference between the RIOS and non-RIOS in the oblique region. The cells in the cardinal RIOS region were tuned more sharply and the cells in cardinal non-RIOS region more broadly than the oblique RIOS and/or the non-RIOS region, which showed no significant difference. These data strongly suggest the existence of functional segregation in the region corresponding to the cardinal contours.

Energy requirements and body surface area of cats and dogs.

.Understanding whether energy requirements and othermetabolic functions vary with BW, metabolic BW, or BSAis crucial when carefully managing patients that vary inbody size (eg, dogs and cats). For example, use of BW oran exponent of BW that is too high will typically under-estimate the energy requirements of smaller animals andoverestimate energy requirements in larger animals.Overfeeding can be especially life-threatening for mal-nourished patients and will promote obesity in well-nour-ished patients. The relationship between size and func-tion may also yield insight into metabolic processes. The idea that heat loss from a warm-bodied animalshould be roughly proportional to BSA was first sug-gested as early as 1839.

Axons in cat visual cortex are topologically self-similar.

The axonal arbors of the different types of neocortical and thalamic neurons appear highly dissimilar when viewed in conventional 2D reconstructions. Nevertheless, we have found that their one-dimensional metrics and topologies are surprisingly similar. To discover this, we analysed the axonal branching pattern of 39 neurons (23 spiny, 13 smooth and three thalamic axons) that were filled intracellularly with horseradish peroxidase (HRP) during in vivo experiments in cat area 17. The axons were completely reconstructed and translated into dendrograms. Topological, fractal and Horton-Strahler analyses indicated that axons of smooth and spiny neurons had similar complexity, length ratios (a measure of the relative increase in the length of collateral segments as the axon branches) and bifurcation ratios (a measure of the relative increase in the number of collateral segments as the axon branches). We show that a simple random branching model (Galton-Watson process) predicts with reasonable accuracy the bifurcation ratio, length ratio and collateral length distribution of the axonal arbors.