Showing papers on "Receptive field published in 1987"

Visual Processing in Monkey Extrastriate Cortex

Abstract: The neuronal processes that lead to visual perception have attracted intense interest since Kuffier's studies of receptive field organization in cat retinal ganglion cells over three decades ago (Kuffier 1953). A variety of ana­ tomical and physiological approaches have been employed to analyze the organization of thc visual pathway between the retina and striate cortex (VI ) and the transformations of visual information that occur at each stage (see Hubel & Wiesel 1977, Stone 1 983, Shapley & Lennie 1985). The growth in understanding of the retinostriate pathway has been accompanied by increasing interest in visual processing in the expanse of extrastriate cortex beyond V I . Studies of extrastriate cortex in many spec­ ies showed that it comprises a mosaic of visual areas that can be dis­ tinguished by several anatomical and physiological criteria (reviewed by Kaas 1 978, Zeki 1 978, Cowey 1979, Van Essen 1 979, 1985, Wagor et al 1980, Tusa et al 1 98 1) . The literature in this field is large, and we do not attempt to review all relevant studies. Rather, we concern ourselves with three recent devel­ opments that have yielded insight into information processing and flow within extrastriate cortex. The first of these is the convergence of ana tom-

The two-dimensional spatial structure of simple receptive fields in cat striate cortex

Abstract: 1. A reverse correlation (6, 8, 25, 35) method is developed that allows quantitative determination of visual receptive-field structure in two spatial dimensions. This method is applied to simple cells in the cat striate cortex. 2. It is demonstrated that the reverse correlation method yields results with several desirable properties, including convergence and reproducibility independent of modest changes in stimulus parameters. 3. In contrast to results obtained with moving stimuli, we find that the bright and dark excitatory subregions in simple receptive fields do not overlap to any great extent. This difference in results may be attributed to confounding the independent variables space and time when using moving stimuli. 4. All simple receptive fields have subregions that vary smoothly in all directions in space. There are no sharp transitions either between excitatory subregions or between subregions and the area surrounding the receptive field. 5. Simple receptive fields vary both in the number of subregions observed, in the elongation of each subregion, and in the overall elongation of the field. In contrast with results obtained using moving stimuli, we find that subregions within a given receptive field need not be the same length. 6. The hypothesis that simple receptive fields can be modeled as either even symmetric or odd symmetric about a central axis is evaluated. This hypothesis is found to be false in general. Most simple receptive fields are neither even symmetric nor odd symmetric. 7. The hypothesis that simple receptive fields can be modeled as the product of a width response profile and an orthogonal length response profile (Cartesian separability) is evaluated. This hypothesis is found to be true for only approximately 50% of the cells in our sample.

Visual properties of neurons in area V4 of the macaque : sensitivity to stimulus form

Abstract: Area V4, a visuotopically organized area in prestriate cortex of the macaque, is the major source of visual input to the inferior temporal cortex, known to be crucial for object recognition. To exa...

Dynamic receptive field plasticity in rat spinal cord dorsal horn following C-primary afferent input

Abstract: The central terminals of cutaneous primary afferent neurons are spatially ordered in the dorsal horn in a highly organized fashion such that a point-to-point map represents the body surface. This afferent terminal somatotopic map correlates with the map of the receptive fields of the cells on which they terminate. The location, size and modality of the cutaneous receptive fields of dorsal horn neurons necessarily depend upon the anatomical presence of afferent nerve fibres which deliver information from the periphery, directly or indirectly, to the cells. However the receptive field size and modality of a cell do not depend only on anatomical connections. Excitatory and inhibitory interneurons, descending influences and facilitations or depressions of synaptic contacts can alter receptive field properties. Here we show that prolonged and substantial cutaneous receptive field changes can be produced by brief inputs from peripheral unmyelinated afferent fibres.

Receptive field properties of neurons in area V3 of macaque monkey extrastriate cortex.

Abstract: Receptive field properties of 147 neurons histologically verified to be located in area V3 were investigated during semichronic recording from paralyzed anesthetized macaque monkeys Quantitative analyses were made of neuron selectivities for direction, orientation, speed, binocular disparity, and color The majority of neurons in V3 (76%) were strongly orientation selective; 40% demonstrated strong direction selectivity Most cells were tuned for stimulus speed and almost half showed optimum responses at 16 degrees/s The distribution of optimum speeds ranged primarily from 4 to 32 degrees/s Several cells in V3 displayed multi-peaked orientation- and/or direction-tuning curves These cells had two or more narrowly tuned peaks that were not co-axial In some ways, they resemble higher-order hypercomplex cells of cat area 19 and may subserve a higher level of form or motion analysis than is seen at antecedent visual areas Roughly half (45%) of the cells were selective for binocular disparity Approximately half of these were tuned excitatory in that they showed weak responses when tested through either eye alone, but showed strong binocular facilitation centered on the fixation plane The other disparity-selective cells were tuned inhibitory or asymmetric in their responses in front and behind the fixation plane Contrary to previous reports, approximately 20% of the neurons in V3 were color selective in terms of showing a severalfold greater response to the best monochromatic wavelength compared with the worst Color-tuning curves of the subset of color selective cells had, on average, a full bandwidth at half maximum response of 80-100 nm A comparison of the receptive field properties of neurons in V3 to those in other areas of visual cortex suggests that V3, like MT, is well suited for the analysis of several aspects of stimulus motion V3 may also be involved in some aspects of form analysis, particularly at low contrast levels Comparison with area VP, a thin strip of cortex anterior to ventral V2, which was previously considered part of V3, indicates that direction selectivity is much more prevalent in V3 than in VP Conversely, color-selective cells are the majority in VP but a minority in V3 This suggests that visual information is processed differently in the upper and lower visual fields

Spatiotemporal convergence and divergence in the rat S1 "barrel" cortex.

Abstract: The size and response magnitude of receptive fields were evaluated for cells in the rat cortical barrel-field by using standard vibrissal deflections of 1.14 degrees. Such stimuli fell within the plateau region of stimulus-response curves. The response of all neurones to all vibrissae within and surrounding centre-receptive fields were analysed for probability and latency of response. It was found that cells in supragranular layers had small centre-receptive fields (average 1.6 vibrissae) with small excitatory surrounds (1.5 vibrissae) while cells in the granular layers had small, powerful centre-receptive fields (1.4) with moderately large excitatory surrounds (2.6). Neurones in infragranular layers possessed large but weak centre-receptive fields (2.6) with large surrounds (3.5). Sixty-four neurones in layer IV were studied, the precise locations of which were identified by using dye lesioning and cytochrome oxidase staining. There were no differences in receptive field size for cells within septa and barrel hollows, but the latter were twice as likely to produce two or more spikes per stimulus from the principal vibrissa (65% against 33%). Histological analysis showed that the principal vibrissa was synonymous with the appropriate vibrissa for the barrel on 86% (55 of 64) of occasions. A quantitative analysis of convergent input to three neighbouring barrels (E1, E2, and D1) showed considerable graded overlap of receptive field surrounds, although facial hair adjacent to the mystacial pad only influenced cells on the edge of the barrel-field. Individual vibrissae exhibited significant divergent input to adjacent inappropriate barrels, being preferentially directed to distant septal rather than barrel hollow cells. An analysis of latencies showed that 40% of barrel hollow cells and 48% of barrel septal cells responded at short (less than 10 ms) latencies to their appropriate vibrissa. In contrast, responses to inappropriate vibrissae were overwhelmingly of long latency (10-greater than 30 ms), only 2% of inappropriate responses from barrel hollow cells and 13% from septal cells being of short latency. These results suggest that direct inputs largely project to appropriate barrels. The possibility that divergent inputs are generated by intracortical mechanisms is discussed.

Endstopped neurons in the visual cortex as a substrate for calculating curvature.

Abstract: Neurons in the visual cortex typically respond selectively to the orientation, and velocity and direction of movement, of moving-bar stimuli. These responses are generally thought to provide information about the orientation and position of lines and edges in the visual field. Some cells are also endstopped, that is selective for bars of specific lengths. Hubel and Wiesel first observed that endstopped hypercomplex cells could respond to curved stimuli and suggested they might be involved in detection of curvature, but the exact relationship between endstopping and curvature has never been determined. We present here a mathematical model relating endstopping to curvature in which the difference in response of two simple cells gives rise to endstopping and varies in proportion to curvature. We also provide physiological evidence that endstopped cells in area 17 of the cat visual cortex are selective for curvature, whereas non-endstopped cells are not, and that some are selective for the sign of curvature. The prevailing view of edge and curve determination is that orientations are selected locally by the class of simple cortical cells and then integrated to form global curves. We have developed a computational theory of orientation selection which shows that measurements of orientation obtained by simple cells are not sufficient because there will be strong, incorrect responses from cells whose receptive fields (RFs) span distinct curves (Fig. 1). If estimates of curvature are available, however, these inappropriate responses can be eliminated. Curvature provides the key to structuring the network that underlies our theory and distinguishes it from previous lateral inhibition schemes.

Sensorimotor integration in the primate superior colliculus. II: Coordinates of auditory signals

Abstract: Based on the findings of the preceding paper, it is known that auditory and visual signals have been translated into common coordinates at the level of the superior colliculus (SC) and share a motor circuit involved in the generation of saccadic eye movements It is not known, however, whether the translation of sensory signals into motor coordinates occurs prior to or within the SC Nor is it known in what coordinates auditory signals observed in the SC are encoded The present experiment tested two alternative hypotheses concerning the frame of reference of auditory signals found in the deeper layers of the SC The hypothesis that auditory signals are encoded in head coordinates predicts that, with the head stationary, the response of auditory neurons will not be affected by variations in eye position but will be determined by the location of the sound source The hypothesis that auditory responses encode the trajectory of the eye movement required to look to the target (motor error) predicts that the response of auditory cells will depend on both the position of the sound source and the position of the eyes in the orbit Extracellular single-unit recordings were obtained from neurons in the SC while monkeys made delayed saccades to auditory or visual targets in a darkened room The coordinates of auditory signals were studied by plotting auditory receptive fields while the animal fixated one of three targets placed 24 degrees apart along the horizontal plane For 99 of 121 SC cells, the spatial location of the auditory receptive field was significantly altered by the position of the eyes in the orbit In contrast, the responses of five sound-sensitive cells isolated in the inferior colliculus were not affected by variations in eye position The possibility that systematic variations in the position of the pinnae associated with different fixation positions could account for these findings was controlled for by plotting auditory receptive fields while the pinnae were mechanically restrained Under these conditions, the position of the eyes in the orbit still had a significant effect on the responsiveness of collicular neurons to auditory stimuli The average magnitude of the shift of the auditory receptive field with changes in eye position (129 degrees) did not correspond to the magnitude of the shift in eye position (24 degrees) Alternative explanations for this finding were considered One possibility is that, within the SC, there is a gradual transition from auditory signals in head coordinates to signals in motor error coordinates(ABSTRACT TRUNCATED AT 400 WORDS)

Functional properties of parietal visual neurons: Radial organization of directionalities within the visual field

Abstract: Parietal visual neurons (PVNs) were studied in waking monkeys as they executed a simple fixation-detection task. Test visual stimuli of varied direction, speed, and extent were presented during the fixation period; these stimuli did not control behavior. Most PVNs subtend large, bilateral receptive fields and are exquisitely sensitive to stimulus motion and direction but insensitive to stimulus speed. The directional preferences of PVNs along meridians are opponently organized, with the preferred directions pointing either inward toward or outward away from the fixation point. Evidence presented in the preceding paper (Motter et al., 1987) indicates that opponent directionality along a single meridian is produced by a feed-forward inhibition of 20 degrees-30 degrees spatial extent. The observations fit a double-Gaussian model of superimposed but unequal excitatory and inhibitory receptive fields: When the former is larger, inward directionality results; when smaller, outward directionality results. We examine here the distribution of the meridional directional preferences in the visual field. Tests showed that opponent organization is not produced by differences in local directional properties in different parts of the receptive field. The distribution of response intensities from one meridian to another is adequately described by a sine wave function. These data indicate a best radial direction for each neuron with a broad distribution of response intensities over successive meridians. Thus, any single PVN, with rare exceptions, cannot signal radial stimulus direction precisely. We then determined how accurately the population response predicted radial stimulus direction by the application of a linear vector summation model. The resulting population vector varied from stimulus direction by an average of 9 degrees. Whether or not the perception of the direction of motion depends upon a population vector remains uncertain. PVNs are especially sensitive to object movement in the visual surround, particularly in the periphery of the visual field. This, combined with their large receptive fields and their wide but flat sensitivity to stimulus speed, makes them especially sensitive to optic flow. This is discussed in relation to the role of the parietal visual system in the visual guidance of projected movements of the arm and hand, in the guidance of locomotion, and in evoking the illusion of vection.

Local precision of visuotopic organization in the middle temporal area (MT) of the macaque.

Abstract: The representation of the visual field in the middle temporal area (MT) was examined by recording from single neurons in anesthetized, immobilized macaques. Measurements of receptive field size, variability of receptive field position (scatter) and magnification factor were obtained within the representation of the central 25°. Over at least short distances (less than 3 mm), the visual field representation in MT is surprisingly orderly. Receptive field size increases as a linear function of eccentricity and is about ten times larger than in V1 at all eccentricities. Scatter in receptive field position at any point in the visual field representation is equal to about one-third of the receptive field size at that location, the same relationship that has been found in V1. Magnification factor in MT is only about onefifth that reported in V1 within the central 5° but appears to decline somewhat less steeply than in V1 with increasing eccentricity. Because the smaller magnification factor in MT relative to V1 is complemented by larger receptive field size and scatter, the point-image size (the diameter of the region of cortex activated by a single point in the visual field) is roughly comparable in the two areas. On the basis of these results, as well as on our previous finding that 180° of axis of stimulus motion in MT are represented in about the same amount of tissue as 180° of stimulus orientation in V1, we suggest that a stimulus at one point in the visual field activates at least as many functional “modules” in MT as in V1.

Corticofugal feedback influences the generation of length tuning in the visual pathway.

Abstract: In the feline visual system, neurons exhibiting sensitivity to the length of a moving contour were first observed in the cortex and described as 'hypercomplex cells'. In these cells an increase in stimulus length beyond an optimal value leads to a rapid decline in response. This decline has been attributed to an intracortical inhibitory input which may be driven by layer VI cells with very long receptive fields. It is now clear, however, that cells in the dorsal lateral geniculate nucleus (dLGN), exhibit a degree of length tuning similar to that of cortical 'hypercomplex cells', suggesting that this response property could be generated subcortically. Alternatively, as the dLGN receives a massive corticofugal projection from layer VI cells in the visual cortex, it is possible that this input has a function in generating length tuning in the dLGN. We have investigated this issue by comparing the length tuning of dLGN cells with and without corticofugal feedback. The data show that corticofugal feedback makes a highly significant contribution to the length tuning of dLGN cells. This raises the possibility that length tuning is an emergent property of the geniculo-cortical loop.

Physiological evidence for serial processing in somatosensory cortex.

Abstract: Removal of the representation of a specific body part in the postcentral cortex of the macaque resulted in the somatic deactivation of the corresponding body part in the second somatosensory area. In contrast, removal of the entire second somatosensory area had no grossly detectable effect on the somatic responsivity of neurons in the postcentral cortex. This direct electrophysiological evidence for serial cortical processing in somesthesia is similar to that found earlier for vision and, taken together with recent anatomical evidence, suggests that there is a common cortical plan for the processing of sensory information in the various sensory modalities.

Effects of early unilateral blur on the macaque's visual system. II. Anatomical observations.

Abstract: We studied the properties of visual cortical and lateral geniculate neurons in 5 macaque monkeys raised with the vision of one eye blurred by daily instillation of atropine. This rearing reduced the degree of binocular interaction in striate cortical neurons and caused a modest shift in eye dominance away from the atropine-treated eye. It also led to a difference in the spatial properties of neurons driven by the 2 eyes: neurons driven by the treated eye tended to have lower optimal spatial frequencies, poorer spatial resolution, and lower contrast sensitivity than neurons driven by the untreated eye. Some of the few binocularly driven neurons had receptive fields with sharply different spatial properties in the 2 eyes, with the treated eye’s receptive field always having poorer spatial resolution. In striate cortex, the effects on neuronal spatial properties were less marked in layer 4 than in more superficial or deeper layers; there was no difference in the spatial properties of lateral geniculate neurons driven by the 2 eyes. A small sample of extrastriate cortical neurons from a single animal showed effects similar to those seen in striate cortex. The striate cortical changes varied consistently from animal to animal: The less affected animals had no discernible eye dominance shift and relatively small differences in spatial properties between the eyes; the more affected animals had substantial eye dominance shifts and larger interocular spatial differences. These variations were also reflected in, and consistent with, behavioral and anatomical measurements performed in the same monkeys. Depriving an animal of vision in one eye early in life profoundly alters the function of its central visual pathways. Following even

Two classes of single-input X-cells in cat lateral geniculate nucleus. II: Retinal inputs and the generation of receptive-field properties

Abstract: The retinal inputs to cells in the cat's lateral geniculate nucleus (LGN) were directly recorded to study the basis for the properties of two classes of LGN X-cells: Xs (single) and XL (lagged). Th...

Nonlinear directionally selective subunits in complex cells of cat striate cortex.

Abstract: 1. We have analyzed receptive fields (RFs) of directionally selective (DS) complex cells in the striate cortex of the cat. We determined the extent to which the DS of a complex cell depends on spat...

Plasticity in the organization of adult cerebral cortical maps: a computer simulation based on neuronal group selection.

Abstract: Recent experimental evidence from the somatosensory, auditory, and visual systems documents the existence of functional plasticity in topographic map organization in adult animals. This evidence suggests that an ongoing competitive organizing process controls the locations of map borders and the receptive field properties of neurons. A computer model based on the process of neuronal group selection has been constructed that accounts for reported results on map plasticity in somatosensory cortex. The simulations construct a network of locally connected excitatory and inhibitory cells that receives topographic projections from 2 receptor sheets corresponding to the glabrous and dorsal surfaces of the hand (a typical simulation involves approximately 1500 cells, 70,000 intrinsic and 100,000 extrinsic connections). Both intrinsic and extrinsic connections undergo activity-dependent modifications according to a synaptic rule based on heterosynaptic interactions. Repeated stimulation of the receptor sheet resulted in the formation of neuronal groups-local sets of strongly interconnected neurons in the network. Cells in most groups were found to have similar receptive fields: they were exclusively glabrous or dorsal despite equal numbers of anatomical connections from both surfaces. The sharpness of map borders was due to the sharpness of the underlying group structure; shifts in the locations of these borders resulted from competition between groups. Following perturbations of the input, the network underwent changes similar to those observed experimentally in monkey somatosensory cortex. Repeated local tapping on the receptor sheet resulted in a large increase in the magnification factor of the stimulated region. Transection of the connections from a glabrous region resulted in the organization of a new representation of corresponding dorsal region. The detailed simulations provide several insights into the mechanisms of such changes, as well as a series of predictions about cortical behavior for further experimental test.

Two classes of single-input X-cells in cat lateral geniculate nucleus. I. Receptive-field properties and classification of cells.

Abstract: Cells in the cat's dorsal lateral geniculate nucleus (LGN) were studied by presentation of visual stimuli and also by simultaneous recording of their ganglion cell inputs in the retina. This paper ...

Spatial properties of neurons in the monkey striate cortex.

Abstract: Contrast sensitivity as a function of spatial frequency was determined for 138 neurons in the foveal region of primate striate cortex. The accuracy of three models in describing these functions was assessed by the method of least squares. Models based on difference-of-Gaussians (DOG) functions where shown to be superior to those based on the Gabor function or the second differential of a Gaussian. In the most general case of the DOG models, each subregion of a simple cell's receptive field was constructed from a single DOG function. All the models are compatible with the classical observation that the receptive fields of simple cells are made up of spatially discrete 'on' and 'off' regions. Although the DOG-based models have more free parameters, they can account better for the variety of shapes of spatial contrast sensitivity functions observed in cortical cells and, unlike other models, they provide a detailed description of the organization of subregions of the receptive field that is consistent with the physiological constraints imposed by earlier stages in the visual pathway. Despite the fact that the DOG-based models have spatially discrete components, the resulting amplitude spectra in the frequency domain describe complex cells just as well as simple cells. The superiority of the DOG-based models as a primary spatial filter is discussed in relation to popular models of visual processing that use the Gabor function or the second differential of a Gaussian.

Spatial response properties of acoustically responsive neurons in the superior colliculus of the ferret: a map of auditory space.

Abstract: Extracellular single-unit recordings were made from auditory neurons in the superior colliculus of ferrets anesthetized with either a neuroleptic or a combination of barbiturate with paralysis. The...

Common and differential effects of attentive fixation on the excitability of parietal and prestriate (V4) cortical visual neurons in the macaque monkey

Abstract: The excitability of cortical neurons of prestriate area V4 and area PG of the inferior parietal lobule were examined using the method of single-neuron analysis in awake macaque monkeys. Levels of excitability were measured as the intensity of response to optimal visual stimuli placed in the most responsive region of the cell's receptive field. Physically and retinotopically identical stimuli were delivered during eye movement pauses under 3 conditions: during a no-task state in which the animal was awake and alert, but not receiving or expecting rewards or working in any task; between trials of the task state, the intertrial interval, while the animal awaited the appearance of a fixation target; and during the foreperiod of the task state, as the animal attentively fixated a small target light, waiting to detect its dimming in order to receive liquid reward. Experiments were carried out in 6 hemispheres of 4 monkeys; both V4 and PG were examined through the same chamber placements in 2 hemispheres. A total of 478 neurons in V4 and PG were identified as visual; quantitative studies were done on 146 in V4 and 54 in PG. We found in these experiments a common effect, a 3– 4-fold facilitation of the responses of both V4 and PG visual neurons during the task state as compared to in the no-task state, and a differential effect, in that V4 neurons showed a similar 3-4-fold facilitation of responses to stimuli presented during the intertrial interval, whereas PG neuronal responses during this interval were similar to those evoked in the no-task state. We describe the functional properties of V4 neurons studied in the waking state. The findings are discussed in relation to the positions of these 2 areas in the occipitoparietal and occipitotemporal transcortical visual systems and to their respective roles in visuospatial perception and pattern recognition. They are also discussed with regard to the candidate neural mechanisms through which the changes in cortical neuronal excitability might be mediated.

Alpha Ganglion Cells in Mammalian Retinae

Abstract: Retinae from species of six orders of mammals (table 1) were processed by an on-the-slide neurofibrillar staining method to establish whether alpha-type ganglion cells are generally present in placental mammals. Alpha cells of the domestic cat, where they were first defined as a type, are used as a standard of reference. Alpha cells were found in all the twenty species examined; characteristically they have the largest somata and large dendritic fields with a typical branching pattern. In keeping with the common morphology there are inner and outer stratifying subpopulations and therefore a presumptive 'on-centre' and 'off-centre' responsiveness to light. Depending on the species, alpha cells form between 1 and 4% of the ganglion-cell population and their dendritic fields cover the retina three to four times. The morphology of alpha ganglion cells, and many of their quantitative features, are conserved in mammals coming from different habitats and having a wide variety of behaviours. Because it is known different habitats and having a wide variety of behaviours. Because it is known from the cat that alpha ganglion cells have brisk-transient or Y receptive fields it is possible that all placental mammals possess this physiological system.

Functional properties of parietal visual neurons: mechanisms of directionality along a single axis

Abstract: The directional properties of parietal visual neurons (PVNs) were examined using the method of single-neuron analysis in waking monkeys. PVN properties were determined with passive visual stimuli as the animal executed a simple detection task. Parietal area PG was studied in 10 hemispheres of 6 male Macaca mulatta. Each class of parietal neurons was identified in PG: the fixation, projection, visual, and oculomotor neurons; 613 PVNs were identified, 323 were studied quantitatively, and 188 were studied with one or more of the protocols described. The receptive fields of PVNs are commonly large and bilateral, and at the limit some may fill the visual field; for many, the central zone of the visual field is spared when the fields are determined by stimuli that enter from the periphery and transit meridians. The receptive fields vary with the behavioral state, the angle of gaze, and the parameters of the stimuli used to determine them. PVNs are sensitive to stimulus movement but relatively insensitive to stimulus speed; many respond over a speed range of 5 degrees-500 degrees/sec. Stimulus-response relations may be incremental or decremental with increasing speed or show maxima or minima in the midrange of speed, but the response variation over the full range is rarely greater than 2:1. The directional preferences of PVNs with bilateral receptive fields are opponently organized; the preferred directions point either inward toward or outward away from the central line of gaze along the 4 meridians tested, which were equally spaced in the circular dimension of the visual field. The mechanism of the axis directionality of PVNs was studied using conditioning-test paradigms. They revealed a feed-forward inhibition preceding a stimulus, an effect that extends from the leading edge of the stimulus for 10 degrees-20 degrees in front of the moving stimulus and lasts for several hundred milliseconds. A double-Gaussian model of superimposed but unequal excitatory and inhibitory effects suffices to explain the present observations. It places demand upon the projection of functional properties from the contralateral hemisphere or from the ipsilateral prestriate areas that project upon PG over multistaged pathways and minimal demands upon intracortical processing mechanisms.

Characterization of neuronal responses to noxious visceral and somatic stimuli in the medial lumbosacral spinal cord of the rat

Abstract: The cutaneous receptive fields, long ascending projections, and responses to colorectal distension (20-100 mmHg) and tail movement of 252 neurons in spinal segments L6-S1 were characterized in pentobarbital- or halothane-N2O anesthetized, physiologically intact male rats. Seventeen additional neurons were studied in spinalized rats. Neurons studied were located within 0.5 mm of the midline at depths 0.2-1.4 mm from the spinal cord dorsum and included the area immediately dorsal and lateral to the central canal. Colorectal distension and/or antidromic invasion from the contralateral ventral quadrant of the cervical spinal cord were used as search stimuli. One hundred seventeen neurons responded to noxious colorectal distension; many had long ascending projections and convergent somatic input from deep joint receptors, ipsilateral perianal/scrotal cutaneous receptive fields, or both. Stimulus-response functions (SRFs) of 45 neurons to graded colorectal distension were linear, allowing extrapolation of threshold distending pressures to neuronal response. Neurons responsive to colorectal distension were subdivided into four classes based on their initial response colorectal distension (75-80 mmHg, 20 s). Short-latency abrupt (SL-A) neurons were excited at short latency by colorectal distension; activity abruptly returned to base line following termination of distension. Most SL-A neurons had long ascending projections, convergent somatic receptive fields, and 4/6 tested were excited by bradykinin administered intraarterially. The threshold distending pressure, estimated from the SRFs of 19 SL-A neurons, extrapolated to 2.7 mmHg. Short-latency sustained (SL-S) neurons were also excited at short latency by colorectal distension, but responses were sustained for 4-120 s following termination of distension. Most SL-S neurons had long ascending projections, convergent somatic receptive fields, and 18/20 tested were excited by intraarterial bradykinin. The threshold distending pressure, estimated from the SRFs of 20 SL-A neurons, extrapolated to 17.0 mmHg. Long-latency (LL) neurons were excited by colorectal distension at long latency following the onset of distension. No LL neurons had demonstrable long ascending projections, and few had convergent excitatory somatic fields. Three of five LL neurons were excited by intraarterial bradykinin. The threshold distending pressure, estimated from the SRFs of six LL neurons, extrapolated to 9.8 mmHg. Inhibited (INHIB) neurons were spontaneously active and were inhibited by colorectal distension.(ABSTRACT TRUNCATED AT 400 WORDS)

Centrifugal organization of direction preferences in the cat's lateral suprasylvian visual cortex and its relation to flow field processing

Abstract: The cerebral cortex of the cat contains between 1 and 2 dozen representations of the visual field with different functional specializations. Six visual field maps lie along both banks of the suprasylvian sulcus, lateral and anterior to the visual areas in the occipital cortex. We have studied single-unit receptive field properties and their global organization across the visual field in 2 of these lateral suprasylvian areas, PMLS (essentially the Clare-Bishop area) and PLLS. Most neurons in PMLS and PLLS display selectivity for the direction of a light stimulus moving across their receptive fields with various degrees of directional tuning. We have used light spots of different size and velocity projected on a tangent screen in order to determine the direction preference of cells in these 2 areas. A strong tendency was found for neurons to respond best to centrifugal directions, i.e., to movement away from the area centralis. Thus, for these cells direction preference depends on the location of their receptive fields within the visual field. Velocity preference and binocular interaction in these neurons is also globally organized: Velocity preference increases with eccentricity, binocular synergism is strongest in the center of the visual field. Cluster analysis of recording tracks with respect to “radial” and “circular” cell categories reveals a grouping of cells with like properties in the lateral suprasylvian cortex. These new categories are formed by combining “centrifugal” and “centripetal” cells on the one hand and cells with direction preferences orthogonal to these on the other. The radial or centrifugal organization of direction preferences in conjunction with the global arrangement of velocity preference and binocular interaction suggests that PMLS and PLLS are involved in the processing of expanding visual flow fields of motion. Such flow fields are commonly encountered when a visual object moves towards an observer or during forward locomotion.

Abnormal single-unit activity recorded in the somatosensory thalamus of a quadriplegic patient with central pain.

Abstract: We have performed single unit analysis of the activity of cells located in the ventral nuclear group of thalamus in a patient with dysesthetic pain below the level of a clinically complete traumatic spinal cord transection at C5. Cells located in the parasagittal plane 14 mm lateral to the midline responded to tactile stimulation in small facial and intraoral receptive fields, which were characteristic of patients without somatosensory abnormality [30]. In this patient the 16 mm lateral parasagittal plane contained cells with receptive fields located on the occiput and neck instead of the upper extremity as would normally be expected. Cells with receptive fields on the neck and occiput had not previously been observed in recordings from single units (n = 531) responding to somatosensory stimulation [30]. Thus, on the basis of their location in a region of thalamus which normally represents parts of the body below the level of the spinal cord transection and their unusual receptive fields adjacent to these same parts of the body, we propose that the cells in the 16 mm lateral plane have lost their normal afferent input. Analysis of the autopower spectra of spike trains indicates that cells in the 16 mm lateral plane exhibited a higher mean firing rate and greater tendency to fire in bursts than cells in the 14 mm lateral plane (P

Connections between pyramidal neurons in layer 5 of cat visual cortex (area 17).

Abstract: The structural features of two physiologically-characterised pyramidal neurons (PC1 and PC2) closely situated in layer 5b in the visual cortex (area 17) of a single cat were studied using a combination of electrophysiological and anatomical techniques. Both PC1 and PC2 had exceptionally large somata (30–40 μm in diameter). On the basis of this and other morphological features cell PC1 was classified as a Meynert cell. PC1 possessed a very large (2.75° × 4.50°) binocularly driven standard complex receptive field. PC2 was also binocularly driven with a small, B-type receptive field. Both cells had the same preference for the direction and orientation of visual stimuli. PC1 and PC2 could be antidromically activated from stimulating electrodes positioned above the dorsal lateral geniculate nucleus with a response latency indicating that these cells probably innervated the visual tectum or pretectum. In addition to corticoefferent axons, the two neurons possessed extensive intracortical axon arbors that ramified extensively in layers 5 and 6 of the medial and lateral banks of the lateral gyrus in area 17. Axon collaterals from both PC1 and PC2 also innervated a small common target region in area 18. A total of 313 boutons from the axonal arbors of PC1 and PC2 were examined in the electron microscope. All of the identified synaptic junctions were found to establish Gray type 1 asymmetrical contacts. The combined ultrastructural data for both neurons indicated that 80% of boutons were onto dendritic spine heads, with 14%, 6%, and 1% onto small-, medium-, and large-calibre dendritic shafts, respectively. The spectrum of postsynaptic targets showed little variation with respect to lamina, distance from somata, or cortical area. Other large pyramidal neurons in layer 5 and spiny neurons in layer 6 were identified as receiving synaptic input from either PC1 or PC2. Using a computer graphics system, rotations of the bouton distributions revealed the existence of a clustered innervation of layers 5 and 6 in areas 17 and 18 derived from the two identified neurons. The bouton distributions strongly resembled the tangential pattern described previously for the functional slab-like organisation of the cortex. The results provide a morphological basis for the clustered intrinsic connectivity of pyramidal cells in layers 5 and 6 of the cat visual cortex. Furthermore, the results indicate the widespread excitatory influence of large pyramidal neurons on other cells projecting subcortically to sites dealing with visually guided behavior.

Mapping of retinal and geniculate neurons onto striate cortex of macaque

Abstract: A unity ratio between geniculate and ganglion cells can be shown in the macaque visual system. Comparison of the densities (cells/deg2) in the dorsal lateral geniculate nucleus (dLGN) of parvocellular (P) and magnocellular (M) cells, respectively, representing color-opponent and broad-band ganglion cells, with cortical magnification (mm2/deg2) gives the number of afferents per square millimeter in striate cortex (V1). For P cells, this afferent density rises only slightly with eccentricity, indicating that V1 magnification is approximately proportional to the density of P cells. The density of cytochrome oxidase puffs in V1 also rises only slightly with eccentricity. As a result, the number of P-cell afferents per puff-centered module is remarkably constant throughout V1. Our findings thus support a novel hypothesis of peripheral scaling, in which V1 cortical magnification is based on the mapping of just 1 class of afferent onto V1 modules. This "P-cell module" in V1 may be composed of submodules corresponding anatomically to the honeycomb cell in layer 4A of V1 and physiologically to a minimal complete set of color-opponent ganglion cells. In contrast, the afferent density of M cells rises steeply with eccentricity, so that the reciprocal of their afferent density, the cortical "domain" of M cells, declines with eccentricity. This decline is similar to that of point-image area in V1. As a result, the number of M cells per point-image area is nearly constant. This quantity is analogous to the receptive-field coverage factor in the retina, which for M cells is fairly constant and greater than unity at all eccentricities. The results show fundamental differences between the neural maps of these 2 major cell types, differences that are likely to have psychophysical consequences.

Ectosylvian visual area of the cat: location, retinotopic organization, and connections.

Abstract: We have mapped out the ectosylvian visual area (EVA) of the cat in a series of single- and multiunit recording studies. EVA occupies 10-20 mm2 of cortex at the posterior end of the horizontal limb of the anterior ectosylvian sulcus. EVA borders on somatosensory cortex anteriorly, auditory cortex posteriorly, and nonresponsive cortex laterally. EVA exhibits limited retinotopic organization, as indicated by the fact that receptive fields shift gradually with tangential travel of the microelectrode through cortex. However, a point-to-point representation of the complete visual hemifield is not present. We have characterized the afferent and efferent connections of EVA by placing retrograde and anterograde tracer deposits in EVA and in other cortical visual areas. The strongest transcortical fiber projection to EVA arises in the lateral suprasylvian visual areas. Area 20, the granular insula, and perirhinal cortex provide additional sparse afferents. The projection from lateral suprasylvian cortex to EVA arises predominantly in layer 3 and terminates in layer 4. EVA projects reciprocally to all cortical areas from which it receives input. The projection from EVA to the lateral suprasylvian areas arises predominantly in layers 5 and 6 and terminates in layer 1. EVA is linked reciprocally to a thalamic zone encompassing the lateromedial-suprageniculate complex and the adjacent medial subdivision of the latero-posterior nucleus. We conclude that EVA is an exclusively visual area confined to the anterior ectosylvian sulcus and bounded by nonvisual cortex. EVA is distinguished from other visual areas by its physical isolation from those areas, by its lack of consistent global retinotopic organization, and by its placement at the end of a chain of areas through which information flows outward from the primary visual cortex.