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Showing papers on "Receptive field published in 1984"


Journal ArticleDOI
TL;DR: The results suggest that a system involved in the processing of color information, especially color-spatial interactions, runs parallel to and separate from the orientation-specific system.
Abstract: Staining for the mitochondrial enzyme cytochrome oxidase reveals an array of dense regions (blobs) in the primate primary visual cortex. They are most obvious in the upper layers, 2 and 3, but can also be seen in layers 4B, 5, and 6, in register with the blobs in layers 2 and 3. We compared cells inside and outside blobs in macaque and squirrel monkeys, looking at their physiological responses and anatomical connections. Cells within blobs did not show orientation selectivity, whereas cells between blobs were highly orientation selective. Receptive fields of blob cells had circular symmetry and were of three main types, Broad-Band Center-Surround, Red-Green Double-Opponent, and Yellow-Blue Double-Opponent. Double-Opponent cells responded poorly or not at all to white light in any form, or to diffuse light at any wavelength. In contrast to blob cells, none of the cells recorded in layer 4C beta were Double-Opponent: like the majority of cells in the parvocellular geniculate layers, they were either Broad-Band or Color-Opponent Center-Surround, e.g., red-on-center green-off-surround. To our surprise cells in layer 4C alpha were orientation selective. In tangential penetrations throughout layers 2 and 3, optium orientation, when plotted against electrode position, formed long, regular, usually linear sequences, which were interrupted but not perturbed by the blobs. Staining area 18 for cytochrome oxidase reveals a series of alternating wide and narrow dense stripes, separated by paler interstripes. After small injections of horseradish peroxidase into area 18, we saw a precise set of connections from the blobs in area 17 to thin stripes in area 18, and from the interblob regions in area 17 to interstripes in area 18. Specific reciprocal connections also ran from thin stripes to blobs and from interstripes to interblobs. We have not yet determined the area 17 connections to thick stripes in area 18. In addition, within area 18 there are stripe-to-stripe and interstripe-to-interstripe intrinsic connections. These results suggest that a system involved in the processing of color information, especially color-spatial interactions, runs parallel to and separate from the orientation-specific system. Color, encoded in three coordinates by the major blob cell types, red-green, yellow-blue, and black-white, can be transformed into the three coordinates, red, green, and blue, of the Retinex algorithm of Land.

1,546 citations


Journal ArticleDOI
TL;DR: The first systematic survey of the responses of IT neurons to both simple stimuli and highly complex stimuli indicates that there may be specialized mechanisms for the analysis of faces in IT cortex.
Abstract: Previous studies have reported that some neurons in the inferior temporal (IT) cortex respond selectively to highly specific complex objects. In the present study, we conducted the first systematic survey of the responses of IT neurons to both simple stimuli, such as edges and bars, and highly complex stimuli, such as models of flowers, snakes, hands, and faces. If a neuron responded to any of these stimuli, we attempted to isolate the critical stimulus features underlying the response. We found that many of the responsive neurons responded well to virtually every stimulus tested. The remaining, stimulus-selective cells were often selective along the dimensions of shape, color, or texture of a stimulus, and this selectivity was maintained throughout a large receptive field. Although most IT neurons do not appear to be “detectors” for complex objects, we did find a separate population of cells that responded selectively to faces. The responses of these cells were dependent on the configuration of specific face features, and their selectivity was maintained over changes in stimulus size and position. A particularly high incidence of such cells was found deep in the superior temporal sulcus. These results indicate that there may be specialized mechanisms for the analysis of faces in IT cortex.

1,387 citations


Journal ArticleDOI
TL;DR: The discharges of single neurones in the parvocellular and magnocellular laminae of the macaque's lateral geniculate nucleus were recorded with glass‐insulated tungsten micro‐electrodes and linearity of spatial summation was examined.
Abstract: The discharges of single neurones in the parvocellular and magnocellular laminae of the macaque's lateral geniculate nucleus (l.g.n.) were recorded with glass-insulated tungsten micro-electrodes. Linearity of spatial summation was examined using the test devised by Hochstein & Shapley (1976). 2 of 272 parvocellular units and 6 of 105 magnocellular units showed clearly non-linear spatial summation. A quantitative index of non-linearity did not suggest the existence of a distinct 'non-linear' class of magnocellular unit. Spatial contrast sensitivity to moving gratings was measured by a tracking procedure in which contrast was adjusted to elicit a reliable modulation of discharge. With the exception of cells that were driven by blue-sensitive cones, measurements of contrast sensitivity did not reveal distinct subgroups of parvocellular units. All had low sensitivity, and those with receptive fields in the fovea could resolve spatial frequencies of up to 40 cycles deg-1. Magnocellular units had substantially higher sensitivity, but poorer spatial resolution. The higher sensitivities of magnocellular units led to their giving saturated responses to stimuli of high contrast. Responses of parvocellular units were rarely saturated by any stimulus. At any one eccentricity the receptive fields of parvocellular units had smaller centres than did those of magnocellular units. Receptive fields of magnocellular units driven by the ipsilateral eye had larger receptive fields than did those driven by the contralateral eye. Parvocellular units were most sensitive to stimuli modulated at temporal frequencies close to 10 Hz; magnocellular units to stimuli modulated at frequencies nearer 20 Hz. The loss of sensitivity as temporal frequency fell below optimum was more marked in magnocellular than parvocellular units. Changes in temporal frequency altered the shapes of the spatial contrast sensitivity curves of both parvocellular and magnocellular units. These changes could be explained by supposing that centre and surround have different temporal properties, and that the surround is relatively less sensitive to higher temporal frequencies.

1,037 citations


Journal ArticleDOI
TL;DR: The topographic organization of striate cortex in the macaque was studied using physiological recording techniques as mentioned in this paper, which facilitated the quantitative analysis of various features of the visual representation, including individual variability in these features as well as in the overall size of the cortex.

924 citations


Journal ArticleDOI
TL;DR: The neuronal circuitry and structure‐function relationships of single neurones in the striate visual cortex of the cat using a combination of electrophysiological and anatomical techniques were studied and two striking correlations were found between dendritic morphology and receptive field type.
Abstract: We have studied the neuronal circuitry and structure-function relationships of single neurones in the striate visual cortex of the cat using a combination of electrophysiological and anatomical techniques. Glass micropipettes filled with horseradish peroxidase were used to record extracellularly from single neurones. After studying the receptive field properties, the afferent inputs of the neurones were studied by determining their latency of response to electrical stimulation at different positions along the optic pathway. Some cells were thus classified as receiving a mono- or polysynaptic input from afferents of the lateral geniculate nucleus (l.g.n.), via X- or Y-like retinal ganglion cells. Two striking correlations were found between dendritic morphology and receptive field type. All spiny stellate cells, and all star pyramidal cells in layer 4A, had receptive fields with spatially separate on and off subfields (S-type receptive fields). All the identified afferent input to these, the major cell types in layer 4, was monosynaptic from X- or Y-like afferents. Neurones receiving monosynaptic X- or Y-like input were not strictly segregated in layer 4 and the lower portion of layer 3. Nevertheless the X- and Y-like l.g.n. fibres did not converge on any of the single neurones so far studied. Monosynaptic input from the l.g.n. afferents was not restricted to cells lying within layers 4 and 6, the main termination zones of the l.g.n. afferents, but was also received by cells lying in layers 3 and 5. The projection pattern of cells receiving monosynaptic input differed widely, depending on the laminar location of the cell soma. This suggests the presence of a number of divergent paths within the striate cortex. Cells receiving indirect input from the l.g.n. afferents were located mainly within layers 2, 3 and 5. Most pyramidal cells in layer 3 had axons projecting out of the striate cortex, while many axons of the layer 5 pyramids did not. The layer 5 cells showed the most morphological variation of any layer, were the most difficult to activate by electrical stimulation, and contained some cells which responded with the longest latencies of any cells in the striate cortex. This suggests that they were several synapses distant from the l.g.n. input. The majority of cells in layers 2, 3, 4 and 6 had the same basic S-type receptive field structure. Only layer 5 contained a majority of cells with spatially overlapping on and off subfields (C- and B-type receptive fields).(ABSTRACT TRUNCATED AT 400 WORDS)

520 citations


Journal ArticleDOI
24 May 1984-Nature
TL;DR: It is found that the auditory receptive fields shifted with changes in eye position, allowing the auditory and visual maps to remain in register, as well as other data suggesting that the primate SC is organized in motor, not sensory, coordinates.
Abstract: The process by which sensory signals are transformed into commands for the control of movement is poorly understood. A potential site for such a transformation is the superior colliculus (SC), which receives auditory, visual and somatosensory inputs1–3 and contains neurones that discharge before saccadic eye movements4–6. Along the primary sensory pathways, signals coding the spatial location of auditory, visual and somatosensory targets are based on distinctly different coordinate systems, and it is not known whether each type of sensory input uses a separate motor pathway or if they are converted into a common coordinate system in order to share a single pre-motor circuit. Sensory neurones in the SC have spatially restricted receptive fields (RFs) and are organized into maps across the collicular surface7–9. Acute experiments have shown a rough correspondence between the spatial positions of RFs of neurones encountered along a single dorsal–ventral penetration of the colliculus, regardless of the modality of the effective stimulus10–14, suggesting that auditory, visual and somatosensory maps might be in register. However, in these conditions the head-centred auditory system and the retinotopic visual system are aligned because the eyes are in the primary orbital position15. Moreover, other data have suggested16–18 that the primate SC is organized in motor, not sensory, coordinates, although in the cat, eye position was found to have no effect on auditory receptive fields19. We therefore sought here to determine what happens to the registration of the auditory and visual maps in the alert, behaving animal. Monkeys, with heads fixed, were trained to make delayed saccadic eye movements to auditory or visual targets from one of three initial fixation points while the activity of single neurones was recorded extracellularly. We found that the auditory receptive fields shifted with changes in eye position, allowing the auditory and visual maps to remain in register.

351 citations


Journal ArticleDOI
TL;DR: The intermediate and deep layers of the cat's superior colliculus are examined for evidence of a neural representation of auditory space and the responses of single units to sounds presented in a free field support the following generalizations.
Abstract: We have examined the intermediate and deep layers of the cat's superior colliculus for evidence of a neural representation of auditory space. We measured the responses of single units to sounds presented in a free field. The results support the following generalizations. Most auditory units in the superior colliculus have sharply delimited receptive fields which form two discrete classes distinguished by their locations and sizes. The remaining units respond to sounds presented at any location. Each auditory unit responds maximally to sounds at a particular horizontal and vertical location within its receptive field, the unit's “best area.” The best areas and receptive field borders of a unit are resistant to changes in the intensity of stimulus. The locations of best areas shift systematically as a function of unit position to form a continuous map of auditory space. The horizontal dimension of space is mapped rostrocaudally, and the vertical dimension is mapped mediolaterally. This map corresponds in orientation with the map of visual space. These data permit us to infer the distribution of unit activity elicited by a sound at any given location. Regardless of its location, a sound activates a substantial portion of the superior colliculus. Indeed, sounds at some locations activate nearly all of the auditory units. However the activated portion of the colliculus contains a restricted region of units which are excited to near their maximum firing rates. The position of this focus of greatest activity varies systematically according to the location of the sound source, thus mapping the location of the sound in space.

350 citations


Journal ArticleDOI
TL;DR: A differential effect of BMI is suggested, suggesting that GABA controls receptive-field size in rapidly adapting regions, and indicates that neurons in rapid adapting regions differ pharmacologically from those in other submodality regions.
Abstract: Extracellular recordings of 209 neurons were obtained with carbon fiber-containing multibarrel micropipettes. The cells were isolated in the primary somatosensory cortex of cats anesthetized with barbiturate and classified according to the nature of their response to natural stimuli, the nature of the surrounding multiunit responses to the same stimuli, the response to thalamic stimulation, and their depth in the cortex. To study factors controlling the excitability of somatosensory neurons, their receptive fields were examined in the presence of iontophoretically administered gamma-aminobutyric acid (GABA), glutamate, and bicuculline methiodide (BMI). Even when the neurons were depolarized to perithreshold levels with glutamate, or when local inhibitory influences mediated by GABA were antagonized by BMI, the apparent specificity for one class of afferent input was maintained. Neurons responding to stimulation of either cutaneous or deep receptors maintained their modality specificity, and neurons in cutaneous rapidly adapting regions never took on slowly adapting properties. When ejected at currents that did not elicit action potentials, glutamate lowered the threshold for activation by cutaneous stimuli but did not enlarge the receptive field. With larger ejecting currents, the neurons developed an on-going discharge, but even at these higher doses, glutamate did not produce an increase in the receptive-field size. Some neurons in regions of cortex exhibiting slowly adapting multiunit responses were relatively insensitive to glutamate. These cells required four to five times more glutamate to evoke discharges than did most neurons. Other cells, previously unresponsive to somatic stimuli, could be shown to possess distinct cutaneous receptive fields when either glutamate or BMI was ejected in their vicinity. Iontophoretically administered BMI altered the firing pattern of somatosensory neurons, causing them to discharge in bursts of 3-15 impulses. BMI enlarged the receptive-field size of neurons in regions displaying rapidly adapting multiunit background discharges but not in those regions with slowly adapting multiunit discharges. This differential effect of BMI, suggesting that GABA controls receptive-field size in rapidly adapting regions, also indicates that neurons in rapidly adapting regions differ pharmacologically from those in other submodality regions. In all cortical regions, BMI blocked the poststimulus inhibitory period that normally followed thalamic stimulation.(ABSTRACT TRUNCATED AT 400 WORDS)

349 citations


Journal ArticleDOI
TL;DR: Numerous highly angled electrode penetrations through the opercular region of macaque striate cortex reveal that layers 4A, 4C alpha, and 4C beta--the primary input sublaminae for axons from the lateral geniculate nucleus (LGN)--are retinotopically organized on a fine scale and populated mostly by monocularly driven cells having small receptive fields and lacking orientation selectivity.
Abstract: Numerous highly angled electrode penetrations through the opercular region of macaque striate cortex reveal that layers 4A, 4C alpha, and 4C beta--the primary input sublaminae for axons from the lateral geniculate nucleus (LGN)--are retinotopically organized on a fine scale and populated mostly by monocularly driven cells having small receptive fields and lacking orientation selectivity. Layer 4B, which does not receive a direct thalamic input, contains orientationally selective cells, and many of these are also direction selective. To a significant degree the response properties of cells in layers 4C alpha and 4C beta reflect the response properties of their respective afferent inputs, from the magno- and parvocellular laminae of the LGN. Accordingly, cells in layer 4C alpha have lower contrast thresholds and larger minimum response fields than do the cells in layer 4C beta. In contrast to this clear-cut separation, the cells of layer 4A (whose major source of direct LGN input arises from the parvocellular layers) exhibit both high and low contrast thresholds. With regard to the precision of retinotopic mapping that is seen in lamina 4C, it is noteworthy that there is substantial overlap among the minimum response fields of neighboring neurons. Due to a larger mean receptive field size, this overlap is greater in layer 4C alpha than it is in 4C beta. In either sublamina, however, the minimum cortical distance that separates different and nonoverlapping parts of the visual field corresponds closely--within a factor of 2--to the known arborizational spreads of single geniculate afferents.

318 citations


Journal ArticleDOI
01 Sep 1984-Brain
TL;DR: It was concluded that a train of impulses in a single tactile unit may produce within the brain of the subject a construct which specifies with great accuracy the skin area of the unit's terminals as well as a tactile subquality which is related to unit properties.
Abstract: Tungsten microelectrodes were inserted percutaneously into the median nerve of alert human subjects for recording and stimulation of single nerve fibres. Impulses from mechanoreceptive units in the glabrous skin of the hand were recorded and single afferents were characterized with respect to unit type (FA I, FA II, SA I, and SA II), as well as size and shape of receptive field, and force threshold. The electrode was then reconnected to an electrical stimulator and short pulse trains (0.25 to 0.5 s, 20 to 100 Hz) were delivered at successively increasing current intensity, while the subject was asked to report any sensation that he noticed in the hand. The first sensation was always that of a localized skin deformation within a small area, typically 2 to 3 mm in diameter, often coinciding with the receptive field of the recorded unit. Spatial matching was also found in many cases for the size, shape and orientation of the perceptive and receptive fields, strongly suggesting that the sensation was accounted for by the recorded unit that had been selectively activated by the current pulses. There were clear differences between group data associated with the four types of units with regard to the quality of the percepts. Vibratory sensation was reported with all FA II units and was common with FA I units, whereas a sustained indentation was often associated with SA I units. Indirect evidence suggested that activation of SA II units usually did not elicit a sensation. It was confirmed that a single impulse in a single FA I unit may elicit a sensory response in the attending subject, whereas a much larger input was required from SA I units, which are also less sensitive to mechanical stimuli. This was one of several findings supporting the impression that differential receptive properties, even within a group of afferents, were associated with different sensory responses. It was concluded that a train of impulses in a single tactile unit may produce within the brain of the subject a construct which specifies with great accuracy the skin area of the unit's terminals as well as a tactile subquality which is related to unit properties.

280 citations


Journal ArticleDOI
TL;DR: It is shown that the vast majority of primate striate cells respond to pure color stimuli, in addition to responding to luminance-varying stimuli, and in general, simple cells are color-selective whereas complex cells response to multiple color regions.

Journal ArticleDOI
TL;DR: It is suggested that one effect of the layer 6 to layer 4 projection could be to contribute to inhibitory features of the receptive fields of layer 4 cells, which are thought to mediate inhibition within the cortex.
Abstract: Although cells in layer 4 of cat striate cortex represent the first stage in the cortical processing of visual information, they have considerably more complicated receptive field properties than the afferents to the layer from the lateral geniculate nucleus. In considering how these properties are generated, we have focused on the intrinsic cortical circuitry, and particularly on the projection to layer 4 from layer 6. Layer 6 pyramidal cells were injected with horseradish peroxidase and examined at the light and electron microscopic level. The labeled axon terminals were found to form asymmetric synapses and to show a strong preference for contacting dendritic shafts. Serial reconstruction of dendrites postsynaptic to labeled layer 6 cell axon terminals showed that a large proportion of the postsynaptic dendrites belonged to smooth and sparsely spiny stellate cells, suggesting a selective innervation of these cell types. In contrast, the geniculate projection to layer 4 made synapses primarily with dendritic spines and, as a result, the large majority of terminals ended on spiny cells. Since smooth and sparsely spiny stellate cells are thought to mediate inhibition within the cortex, we suggest that one effect of the layer 6 to layer 4 projection could be to contribute to inhibitory features of the receptive fields of layer 4 cells.

Journal ArticleDOI
TL;DR: Response properties of single neurons in the middle temporal visual area of anesthetized owl monkeys were determined and quantified for flashed and moving bars of light under computer control for position, orientation, direction of movement, and speed.
Abstract: Response properties of single neurons in the middle temporal visual area (MT) of anesthetized owl monkeys were determined and quantified for flashed and moving bars of light under computer control for position, orientation, direction of movement, and speed. Receptive-field sizes, ranging from 4 to 25 degrees in width, were considerably larger than receptive fields with corresponding eccentricities in the striate cortex. Neurons were highly binocular with most cells equally or nearly equally activated by either eye. Neurons varied in selectivity for axis and direction of moving bars. Some neurons demonstrated little or no selectivity, others were bidirectional on a single axis, while the largest group was highly selective for direction with little or no response to bar movement opposite to the preferred direction. Over 70% of neurons were classified as highly selective and 90% showed some preference for direction and/or axis of stimulus movement. Neurons typically responded to bar movement only over a restricted range of velocities. The majority of neurons responded best to a particular velocity within the 5-60 degrees/s range, with marked attenuation of the response for velocities greater or less than the preferred. Some neurons failed to show significant response attenuation even at the lowest tested velocity, while other neurons preferred velocities of 100 degrees/s or more and failed to attenuate to the highest velocities. Response magnitude varied with stimulus dimensions. Increasing the length of the moving bar typically increased the magnitude of the response slightly until the stimulus exceeded the receptive-field borders. Other neurons responded less to increases in bar length within the excitatory receptive field. Neurons preferred narrow bars less than 1 degree in width, and marked reductions in responses characteristically occurred with wider stimuli. Moving patterns of randomly placed small dots were often as effective as or more effective than single bars in activating neurons. Selectivity for direction of movement remained for the dot pattern. for the dot pattern. Poststimulus time (PST) histograms of responses to bars flashed at a series of 21 different positions across the receptive field, in the "response-plane" format, indicated a spatially and temporally homogeneous receptive-field structure for nearly all neurons. Cells characteristically showed transient excitation at both stimulus onset and offset for all effective stimulus locations. Some cells responded mainly at bright stimulus onset or offset.

Journal ArticleDOI
TL;DR: The characteristics of 222 single neurons in and adjacent to the subnucleus interpolaris were examined electrophysiologically in adult cats anesthetized with chloralose to determine if the functionally identified neurons projected directly to the contralateral posterior thalamus, ipsilateral cerebellum, or cervical spinal cord.
Abstract: In view of continuing uncertainties concerning the organization, afferent inputs, and projection sites of neurons in the subnucleus interpolaris of the trigeminal (V) spinal tract nucleus, the characteristics of 222 single neurons in and adjacent to the subnucleus were examined electrophysiologically in adult cats anesthetized with chloralose. Neurons were tested for orthodromic responsiveness to a variety of stimuli that included nonnoxious tactile stimuli, noxious mechanical and radiant-heat stimuli, and graded electrical stimulation of the skin, mucosa, tooth pulp, and masseter nerve. Antidromic activation techniques were also used to determine if the functionally identified neurons projected directly to the contralateral posterior thalamus, ipsilateral cerebellum, or cervical spinal cord. In addition, the periaqueductal gray matter (PAG) was stimulated to test for conditioning influences from the PAG on orthodromic responses to noxious and nonnoxious oral-facial stimuli. Interpolaris neurons were somatotopically arranged in subnucleus interpolaris in a pattern conforming in general to the medially facing, inverted-head representation characteristic of other parts of the V brain stem sensory nuclear complex. On the basis of their responsiveness to cutaneous stimuli, the neurons could be functionally classified as either cutaneous nociceptive or low-threshold mechanoreceptive (LTM) neurons. The LTM neurons constituted the major neuron type, accounting for over 75% of our neuron sample. Most of them had a localized mechanoreceptive field of less than 100 mm2 in area that was restricted to one V division, and they had skin-evoked response latencies indicative of afferent input predominantly from A-beta cutaneous afferents. A population of nociceptive neurons was also encountered in the lateral, marginal region of interpolaris and at its medial or ventral border with the reticular formation. These neurons were of two types: nociceptive-specific (NS) neurons, which did not respond to nonnoxious stimuli but which required noxious stimuli for their activation; and wide dynamic range ( WDR ) neurons, which responded to both noxious and nonnoxious stimuli applied to the facial skin. Most had an ipsilateral receptive field that was greater than 100 mm2 in area and that often involved two or three V divisions. Their properties generally conformed to those previously described for nociceptive neurons in the medullary dorsal horn (V subnucleus caudalis) and spinal cord dorsal horn. Interpolaris neurons of all classes (LTM, WDR , and NS) were found to have direct axonal projections to the thalamus, cerebellum, and spinal cord.(ABSTRACT TRUNCATED AT 400 WORDS)

Journal ArticleDOI
TL;DR: Each of the three identified interneurones examined in this study was found to have a directional response that matched the response predicted on the basis of its anatomy, which shows a strong correlation between anatomical overlap of primary afferent and interneURone and the existence of a synaptic connexion.
Abstract: The structural relationship between the afferent projection and the dendrites of the interneurones was examined in the cercal-to-giant interneurone system of the cricket using intracellular recording and dye injection techniques. The physiology of the sensory neurones beneath the cercal filiform hairs was investigated by placing a recording pipette over the end of a cut hair and using movements of the pipette to characterize the directionality of the receptor. Most of the filiform receptors could be classified as belonging to one of four major types. Each type is sensitive to a different wind direction and is confined to particular regions of the cercus. The location of the terminal arborizations of each type of sensory cell was revealed by staining with cobalt chloride. Single cells were stained reliably by placing a dye-filled pipette over a cut hair. Each physiological receptor type arborizes in a different region of the central nervous system. Therefore the neuropile is functionally divided according to wind direction. The dendrites of three identified interneurones were examined in the context of this afferent projection. It was found that each of these neurones has dendrites in regions of neuropile corresponding to different wind directions. By searching for unitary synaptic potentials in identified interneurones, it was possible to show a strong correlation between anatomical overlap of primary afferent and interneurone and the existence of a synaptic connexion. Further, when there was no overlap, no synaptic potentials were seen. Therefore the over-all excitatory receptive field of an interneurone could be predicted by examining its dendritic structure. Each of the three identified interneurones examined in this study was found to have a directional response that matched the response predicted on the basis of its anatomy.

Journal ArticleDOI
TL;DR: It is hypothesized that the disruptions, regions of isorepresentation, and regions of gradual change result from the thickening, splitting, and folding of a two‐dimensional representation of the skin surface to occupy a three‐dimensional volume.
Abstract: Multiunit microelectrode mapping techniques were used to investigate the organization of the somatosensory thalamus in squirrel monkeys. Receptive fields and response characteristics were determined for closely spaced recording sites along arrays of electrode penetrations that passed through the ventral thalamus dorsoventrally, rostrocaudally, or lateromedially. The results were related to thalamic architecture and led to the following conclusions: (1) A large, single, systematic representation of the body surface occupied most or all of the ventroposterior nucleus, VP. The nucleus was further defined by a distinct cytoarchitectonic appearance, produced by densely packed, deeply stained neurons. (2) Recording sequences in VP were characterized by (a) abrupt shifts in receptive field locations over short recording distances indicating that the electrode had crossed discontinuities or folds in the representation, (b) long sequences of overlapping receptive fields indicating regions of continuous representation and the maintenance of adjacency in the map, and (c) similar receptive field locations for sites along the trajectory of a penetration indicating regions of isorepresentation. Major somatotopic discontinuities were associated with crossing narrow cell-poor laminae that partially divided VP into subnuclei related to the hand, foot, trunk, and tail in lateral VP and the face in medial VP. Somatotopic discontinuities occurred for electrode penetrations in all three planes, but discontinuities were greater and more frequent for lateromedial electrode penetrations. Lines of isorepresentation and gradual change were most extensive in the rostrocaudal and dorsoventral planes. We hypothesize that the disruptions, regions of isorepresentation, and regions of gradual change result from the thickening, splitting, and folding of a two-dimensional representation of the skin surface to occupy a three-dimensional volume. (3) The magnifications of various skin surfaces in VP were variable so that some skin surfaces, especially the tips of the digits, occupied relatively large portions of the nucleus, while other skin surfaces such as the trunk activated little tissue. It appeared that regions of isorepresentation varied in extent according to magnification factor and position in the map. (4) Within VP, neurons could be classified as slowly adapting or rapidly adapting to maintained skin indentation. Each type of neuron formed small groups or clusters in the nucleus so that several successive recording sites typically encountered one type before a sequence of the other type was observed.(ABSTRACT TRUNCATED AT 400 WORDS)

Journal ArticleDOI
TL;DR: In alert monkeys, single-unit responses to visual stimuli were recorded in the central thalamic region where eye movement-related activity has been observed and the occurrence of these patterns of firing was related to stimulus fixation.
Abstract: In alert monkeys, single-unit responses to visual stimuli were recorded in the central thalamic region where eye movement-related activity has been observed (33). Usually, the stimuli were 1 degree annulus patterns of dim light presented at unpredictable locations on a tangent screen. The animals were trained on two tasks: one in which they delivered the stimulus themselves by pressing a panel that they had to release immediately when the stimulus shape changed to a square, and another one in which the stimulus was turned on by the experimenter and the monkeys were rewarded for fixating this target for a predetermined length of time. In both tasks, continuous stimulus fixation was required. Receptive fields were tested with and without a fixation point. Retinal coordinates of stimuli were obtained by subtracting eye-position coordinates from stimulus coordinates in space, the monkey's head being fixed. Unit responses in the cases where targeting occurred or did not occur were analyzed separately. Transient responses were observed in 63 units and sustained responses in 44 units. Among the 63 units responding transiently, 42 did so irrespective of targeting. Their receptive fields were very large, generally including the fovea, and predominantly contralateral when the fields were asymmetric. The responses of the other 21 units depended on the occurrence of targeting. They were called visually triggered eye movement-related responses (VTEM). VTEM units were further subdivided in 9 units active only with targeting and 12 units showing the classical phenomenon of "response enhancement" under this condition. VTEM units were contrasted to six units that were both passively visually responsive and bursting with saccades, either spontaneous or visually triggered. The latencies of passive visual and VTEM responses to stimulus onset were comprised between 77 and 135 ms in 80% of the units. VTEM units also fired prior to retargeting saccades. Presaccadic units active with spontaneous saccades also discharged with visually elicited saccades. The earliest sign of activation after stimulus onset eliciting a saccade appeared between 80 and 100 ms, that is, in the same range of latencies as passive visual and VTEM units. Sustained visual responses consisted of activation in 18 units and inactivation in 26 units. The occurrence of these patterns of firing was related to stimulus fixation. In the majority of cases, the changes in discharge frequency started before fixation was achieved by a targeting saccade. They terminated before fixation was broken by a saccade away from the stimulus.(ABSTRACT TRUNCATED AT 400 WORDS)

Journal ArticleDOI
TL;DR: Receptive field maps together with electrolytic marking lesions at recording and stimulation sites showed that tonic-on pretectal cells are retinotopically organized and are aggregated in a strip running from the dorso-medial tip of the pretectum to the ventro-lateral boundary, suggesting that they mediate constriction in the light reflex of the rat's pupil.

Journal ArticleDOI
TL;DR: A positive correlation was found between the diameter of the receptive-field center and the dendritic field in Y, X, and W-cells and among all three classes of ganglion cells.
Abstract: The action spike activities of single ganglion cells were recorded from the nasal retina of the intact eye of anesthetized and immobilized cats. Each ganglion cell was identified as a Y-, X-, or W-...

Journal ArticleDOI
TL;DR: A new view of receptive field organization in the X-cell is led to: the center mechanism is spatially homogeneous in its dynamics whereas the surround mechanism are spatially inhomogeneous.

Journal ArticleDOI
TL;DR: Spinal neurons antidromically activated from either the lateral reticular nucleus (LRN) or immediately adjacent areas were identified in the rat lumbar spinal cord and could be subdivided into three main groups according to their location.
Abstract: Spinal neurons antidromically activated from either the lateral reticular nucleus (LRN) or immediately adjacent areas were identified in the rat lumbar spinal cord. In agreement with previous anatomical work (60), these neurons were widely distributed in both the dorsal and ventral horns of the spinal cord and could be subdivided into three main groups according to their location: a) deep ventromedial (DVM) cells, which project more substantially to the LRN than to other supraspinal targets; b) cells of the median portion of the neck of the dorsal horn (mNDH), which project exclusively to the LRN; c) cells lying in other parts of the dorsal horn (superficial layers, nucleus proprius, reticular extension of the neck), by their location, they are indistinguishable from cells projecting to other supraspinal targets. The probability is high that the DVM and mNDH cells contribute exclusively, or at least preferentially, to the lateral component of the spinoreticular tract (lSRT), defined as the direct spinal pathway to the LRN. Although electrophysiological properties of cells were clearly related to their spinal location, several subpopulations could be recognized in each of the three main groups. The majority of DVM neurons were in lamina VII, with some in laminae VI, VIII, and X. With the exception of a few lamina X cells, the DVM neurons had high conduction velocities. Four subpopulations of these neurons were recognized. a) Innocuous proprioceptive cells responded to small changes in joint position, some showing convergence of nonnoxious cutaneous inputs. b) High-threshold cells (approximately 50% of DVM cells). Seventy-five percent of these cells were excited from bilateral receptive fields (mostly symmetric) with noxious cutaneous pinching that extended to subcutaneous tissues. Their evoked responses had long-lasting postdischarges that continued up to several minutes after cessation of the stimulus. c) Inhibited cells had no demonstrable excitatory receptive fields and a high ongoing activity that was tonically depressed by pressure or pinch; poststimulus effects of long duration were observed. d) Cells with no resting discharge and demonstrable excitatory peripheral receptive fields. mNDH cells had recording sites at the medial border of the internal portion of the reticular area of the neck of the dorsal horn.(ABSTRACT TRUNCATED AT 400 WORDS)

Journal ArticleDOI
TL;DR: Preliminary results suggest that the DLPN could supply the smooth-pursuit system with signals concerning the direction and velocity of target image motion on the retina.
Abstract: The visual properties of 77 dorsolateral pontine nucleus (DLPN) cells were studied in two alert monkeys. In 41 cells, presentation of a moving random dot background pattern, while the monkeys fixated a stationary spot, elicited modulations in discharge rate that were related either to (i) the velocity of background motion in a specific direction or to (ii) only the direction of background movement. Thirty-six DLPN cells exhibited responses to small, 0.6–1.7 deg, visual stimuli. Nine such cells exhibited non-direction selective receptive fields that were eccentric from the fovea. During fixation of a stationary bluish spot, the visual responses of 27 DLPN cells to movement of a small, white “test” spot were characterized by two components: (1) as the test spot crossed the fovea in a specific direction, transient velocity-related increases in discharge rate occurred and (2) a maintained, smaller increase in activity was observed for the duration of test spot movement in the preferred direction. This DLPN activity associated with small visual stimuli was also observed during smooth-pursuit eye movements when, due to imperfect tracking, retinal image motion of the target produced slip in the same direction. These preliminary results suggest that the DLPN could supply the smooth-pursuit system with signals concerning the direction and velocity of target image motion on the retina.

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TL;DR: Heterotopic callosal connections may be one factor responsible for binocular vision and also may provide the basis for large, nonoriented receptive fields of units in layer V of rodent visual cortex.

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TL;DR: Motor neurons of T2 to T4 spinal segments were antidromically activated from the medullary reticular formation and the contralateral region in or near the ventral posterior lateral nucleus of the thalamus and support Ruch's convergence projection theory for referred pain.

Journal ArticleDOI
TL;DR: The organization of somatosensory projections to the dysgranular areas of somatic sensory cortex was mapped in albino rats and receptive fields that activate layer IV granule cells in these Dysgranular zones were cutaneous and deep, roughly somatotopic, larger, and required stronger stimulation than the cutaneous light touch RFs of the adjacent granule cell zones.

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TL;DR: The findings are the first to describe the functional effects of dopamine antagonists on single cells in the mammalian retina and on ganglion cell activity in the vertebrate retina.
Abstract: The effects of dopamine antagonists on the extracellularly recorded activity of ON- and OFF-center brisk ganglion cells and ON-OFF directionally selective ganglion cells in the rabbit retina were investigated. Haloperidol, fluphenazine, and cis-flupenthixol, infused in the arterial system supplying the eye, produced similar effects. In general, these drugs reduced the antagonistic surround responses of brisk ganglion cells, reduced the sustained excitation of the center response of ON-center brisk-sustained cells, reduced the leading edge response of ON-OFF directionally selective cells to moving light stimuli along with any sustained excitation to stationary light stimuli, and affected the spontaneous activity of the cells. These drug effects appear to be due to a blockade of D-1 (adenylate cyclase- linked) receptors and not to D-2 receptors. Neither S-sulpiride nor metoclopramide, two specific D-2 antagonists, had much effect. The findings are the first to describe the functional effects of dopamine antagonists on single cells in the mammalian retina and on ganglion cell activity in the vertebrate retina.

Journal ArticleDOI
TL;DR: The proportion of neurones showing accelerated firing during different parts of the step cycle fluctuated more for antidromically identified pyramidal tract neurones (p.t.n.s) than for non‐p.n.'s and when the neurones were subdivided according to the movement evoked by threshold electrical stimulation through the micro‐electrode.
Abstract: Discharge patterns of motor cortical neurones in cats walking steadily on a moving belt have been compared with other functional characteristics of the neurones. In forelimb motor cortex rhythmic discharges occurred in cells with peripheral receptive fields in all parts of the contralateral forelimb and also in cells with no discernible receptive field. Cells discharging at similar times during the step cycle often had very different receptive fields and cells with similar receptive fields (including neighbouring cells) could discharge at similar or at quite different times. In cells with a cutaneous receptive field including the forefoot the discharges during locomotion remained rhythmic (and their phasing relative to the step cycle was unchanged) when the response to mechanical stimulation in the receptive field was temporarily much reduced or abolished by local anaesthesia of the skin. The proportion of neurones showing accelerated firing during different parts of the step cycle fluctuated more for antidromically identified pyramidal tract neurones (p.t.n.s) than for non-p.t.n.s and was highest during the second half of stance in the contralateral forelimb and lowest during swing. When the neurones were subdivided according to the movement evoked by threshold electrical stimulation through the micro-electrode, p.t.n.s and non-p.t.n.s recorded by electrodes evoking elbow flexion showed a wide variety of discharge patterns. For p.t.n.s the discharge rate reached an average of 69 impulses/s during late stance and declined to an average of 26 impulses/s during swing.

Journal ArticleDOI
TL;DR: Investigation of how lateral geniculate neurons and retinal ganglion cells are affected when a drug is applied to the retina that inactivates on-bipolar cells in the mudpuppy found that the surrounds of many off-center cells could still be excited by light stimuli; thus, despite the probable inactivation of on- bipolar cells, not all on- responses in the LGN were abolished.
Abstract: In the cat, lateral geniculate neurons and retinal ganglion cells can almost all be categorized as on-center (excited by light stimuli) or off-center (excited by dark stimuli). We have investigated how these cells are affected when a drug is applied to the retina that, in the mudpuppy, inactivates on-bipolar cells (Slaughter, M. M., and R. F. Miller (1981) Science 211: 182-185). This drug, D,L-2-amino-4-phosphonobutyric acid (APB), is an analogue of glutamate. After injecting APB into the vitreous of the eye, we could record in the optic tract from fibers of off-center retinal ganglion cells that appeared normal in their responses, but we could find no fibers from on-center cells in the injected eye. In the lateral geniculate nucleus (LGN), the outcome was similar. Of a sample of 144 cells studied quantitatively, 13 had extremely poor responses and could not be characterized as on-or off-center. The remaining 131 cells had off-centers and appeared largely normal. Their receptive field organization was unaltered: the field center was excited by dark (off) stimuli and inhibited by light, whereas the surround was inhibited by off stimulation. Of particular interest was the fact that the surrounds of many off-center cells could still be excited by light stimuli; thus, despite the probable inactivation of on-bipolar cells, not all on-responses in the LGN were abolished. As a group, off-center cells were somewhat less responsive then in control experiments. Except for this change, APB appeared to affect only on-center cells.(ABSTRACT TRUNCATED AT 250 WORDS)

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TL;DR: The response of monkey lateral geniculate nucleus cells to flashing spots, annuli, and drifting sine‐wave gratings were recorded with tungsten micro‐electrodes to support the view that spatial and temporal tuning are dynamic properties of some l.g.n. neurones by virtue of descending input from the visual cortex.
Abstract: The response of monkey lateral geniculate nucleus (l.g.n.) cells to flashing spots, annuli, and drifting sine-wave gratings were recorded with tungsten micro-electrodes. These stimuli were presented (a) monocularly, through an aperture in the centre of a radial grating, or (b) dichoptically, in which the spots or drifting gratings were presented to the dominant eye's receptive field, while the centre of the radial grating was positioned on the corresponding retinal location of the other eye. Movement of the radial grating produced changes in the l.g.n. cell responses evoked by the spots and sine-wave gratings. These changes were reversed by cryogenic blockade of the striate cortex. Therefore, radial grating movement altered the responses of l.g.n. cells by activating the corticogeniculate (c.g.) pathway. In about half of all cells, radial grating-induced alterations of centre, or surround, or both responses to spots and annuli were produced. By adopting a simple spatial filtering model of the centre and surround mechanisms, it was possible to predict how these alterations in centre/surround balance would affect the cell's responses to sine-wave gratings. Alterations were observed in the peak and band width of the spatial and/or temporal tuning curves. The radial gratings did not alter the spatial summation properties of cells. Minor alterations in the spectral neutral points of chromatically opponent neurones were occasionally found. These results are interpreted as support for the view that spatial and temporal tuning are dynamic properties of some l.g.n. neurones by virtue of descending input from the visual cortex.

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TL;DR: Single unit electrical activity has been recorded from dorsal horn neurones in the lumbar spinal cord of adult rats which had been treated at birth with either capsaicin or with the solvent-vehicle only, and the role of afferent C-fibres in sensory mechanisms is discussed.