A comparison of magnification functions in area 19 and the lateral suprasylvian visual area in the cat

A retinotopic map can be described by a magnification function that relates magnification factor to visual field eccentricity. Magnification factor for primary visual cortex (VI) in both the cat and the macaque monkey is directly proportional to retinal ganglion cell density. However, among those extrastriate areas for which a magnification function has been described, this is often not the case. Deviations from the pattern established in V1 are of considerable interest because they may provide insight into an extrastriate area's role in visual processing. The present study explored the magnification function for the lateral suprasylvian area (LS) in the cat. Because of its complex retinotopic organization, magnification was calculated indirectly using the known magnification function for area 19. Small tracer injections were made in area 17, and the extent of anterograde label in LS and in area 19 was measured. Using the ratio of cortical area labeled in LS to that in area 19, and the known magnification factor for area 19 at the corresponding retinotopic location, we were able to calculate magnification factor for LS. We found that the magnification function for LS differed substantially from that for area 19: central visual field was expanded, and peripheral field compressed in LS compared with area 19. Additionally, we found that the lower vertical meridian's representation was compressed relative to that of the horizontal meridian. We also examined receptive field size in areas 17, 19, and LS and found that, for all three areas, receptive field size was inversely proportional to magnification factor.     

Spatial interactions in the rhesus monkey retina: a behavioural study using the Westheimer paradigm

Summary For two trained rhesus monkeys, increment thresholds for a small test-spot of 100 ms duration were determined as a function of background size, at 10 retinal eccentricities along the horizontal meridian. Typical Westheimer-functions were obtained, i.e. threshold first increases with increasing background size, reaches a maximum, then decreases with further increasing backgrounds and finally reaches a plateau. With increasing retinal eccentricities, the position of the peak of the functions is shifted towards larger background sizes, indicating an increase of perceptive field centre size from 0.25° at 5° eccentricity to 1.5° at 40° eccentricity. The perceptive field centres tend to be slightly smaller in the nasal retina. Total perceptive field sizes, as indicated by the beginnings of the plateaus, increase from about 1° near the fovea to about 3° at 40° eccentricity. The perceptive field centre sizes of two human observers, tested under the same experimental conditions, closely resemble those of the monkeys. The total perceptive fields are larger in the human subjects. The retinal ganglion cells determining threshold in this experiment are most likely the broad-band cells. The agreement between the behaviourally determined perceptive field centre sizes and the receptive field centre sizes of broadband cells (measured by DeMonasterio and Gouras 1975) is excellent. The dendritic fields of P--ganglion cells, most likely the morphological substrates of the broad-band cells (Perry, Oehler and Cowey 1984) are somewhat smaller at all eccentricities. The close agreement between monkey and human psychophysical data and between monkey psychophysical and neurophysiological data indicates that the Westheimer paradigm actually provides a tool to study behaviourally the receptive field organization of the human retina.Part of this work was presented at the Spring Meeting of the Deutsche Physiologische Gesellschaft, 1982, Giessen (Pflügers Arch. 392, R 49, 1982) and on the Annual Spring Meeting of the Association for Research in Vision and Ophthalmology, Sarasota, Florida, 1982     

Visual field projection columns and magnification factors in the lateral geniculate nucleus of the cat

Summary The precision of the projection of the visual field to the dorsal lateral geniculate nucleus (LGNd) of the cat was studied by plotting the receptive fields of single neurons recorded extra-cellularly in the nucleus. The concepts of a projection column and of random scatter in the location of receptive fields have been defined in relation to cells in the LGNd. A projection column contains 90% of all the cells in the LGNd that have receptive fields with a common visual direction, the central axis of the column being the projection line for the given visual direction. In the region of the LGNd devoted to central vision the columns have a circular cross section and are about 1 mm in diameter.The projections of adjacent areas of visual field (and hence of the retina) overlap extensively in the LGNd. In this study, the overlap of retinal afferents in the LGNd was measured in terms of the random scatter of receptive field positions for cells recorded in a given electrode penetration parallel to projection columns in the nucleus. The monocular receptive field scatter within a column in the LGNd is about of the same magnitude as both the monocular receptive field scatter within a cortical column and the binocular receptive field disparities of cortical units.The differential magnification of the visual field on the LGNd is a reflection of the ganglion cell density differences in the retina.     

Magnification factor and receptive field size in foveal striate cortex of the monkey

Summary Receptive field size and magnification have been studied in striate cortex of awake, behaving rhesus monkeys at visual eccentricities in the range of 5–160 min. The major findings that emerge are (1) magnification in the foveola achieves values in the range of 30 mm/deg, (2) mean field size is not proportional to inverse magnification in contrast with previous reports, and (3) the product, magnification X aggregate field size, is greater in central vision than in peripheral vision. Thus, a point of light projected onto foveal retina is seen by larger numbers of striate cortical cells than a point of light projected onto peripheral retina.Implications of these findings for visual localization and two-point discrimination are discussed.Dedicated to Hermann RahnSupported by NIH grants EY02349 and 5 T32 EY07019     

Representation of the fovea in the superior temporal sulcus of the macaque monkey

Summary The response properties of 633 neurons from striate and prestriate cortex were recorded in 3 hemispheres of two awake cynomolgus monkeys while they fixated or tracked a small spot of light. Of 254 penetrations located at 1 mm intervals, 39% were identifiable from visible electrolytic lesions or electrode tracks and were used to reconstruct the positions of all recording sites. A total of 226 cells were located in the superior temporal sulcus and 81 cells in area V1. The location and visuotopic organization of the foveal portion of the middle temporal (MT) visual area were determined in three hemispheres. MT was defined physiologically on the basis of direction-selectivity, receptive field size, and retinotopic organization. Of 170 MT neurons, most were motion sensitive, and 65% had a directionality index, (best — opposite)/best, of 0.6 or higher. MT was defined anatomically on the basis of myelin staining within the superior temporal sulcus (STS). On the posterior bank of the STS the physiologically defined border corresponded closely to a myelin border visible on our sections. Distinct myelin borders were not consistently identifiable on the anterior bank. The representation of the central fovea (eccentricities of 0–1 deg) was located partly on the floor, but mostly on the posterior bank of the STS at the extreme postero-lateral edge of MT. In all three hemispheres foveal MT extended onto the roof of a cleft formed between the posterior bank and a wide flattened area on the floor of the STS. This region lies 10–12 mm below the brain surface, measuring along a line normal to the surface at a point 2–3 mm antero-lateral to foveal V1. The area of MT was 6–9 mm² for the central fovea (0–1 deg), 15–24 mm² for the entire fovea (0–3 deg), and 28–40 mm² including the fovea and parafovea (0–10 deg). A visuotopic map of central foveal V1 (0–1 deg) was obtained in one animal. The measured area of this representation was 116 mm². Using published estimates of the total areas of cynomolgus MT and V1 (73 and 1200 mm² respectively) the ratio of central foveal to total area was calculated to be 0.10 for both MT (7.5/73) and V1 (116/1200), indicating that the relative magnification of the foveal versus the peripheral visual field is preserved in the mapping of V1 onto MT. A separate representation of the central visual field was found immediately adjacent to foveal MT. This region, the FST area (Ungerleider et al. 1982; Ungerleider and Desimone 1986a, b), was distinguishable from MT in three ways: 1) by the presence of occasional visually unresponsive cells, 2) by the presence of cells with very large receptive fields intermingled with cells whose receptive fields are comparable in size to those found in foveal MT, and 3) by an increased incidence of cells responding during tracking. Of 34 FST neurons, 53% had a directionality index of 0.6 or higher. An additional 22 cells recorded in the superior temporal sulcus were judged to be outside both MT and FST.     

Magnification factors,receptive field images and point-image size in the superior colliculus of flying foxes: comparison with the primary visual cortex

The magnification factor (MF) of the stratum griseum superficialle (SGS) of the superior colliculus (SC) was calculated based on visual receptive fields recorded from anaesthetised and paralysed flying foxes (Pteropus spp.). In areal terms, the MF at the representation of central vision was 4–6 times larger than that in the peripheral representation. This variation is less marked than that observed in the primary visual area (VI), but is roughly that expected if the retinotopic map in the SC was defined by the distribution of ganglion cells in the retina. Two measures of the functional spread of activity in the SC, the receptive field images and the point-image size, were calculated. Receptive field images are remarkably similar throughout the SC. As in VI, the point-image size in the SGS of flying foxes is 0.5–0.6 mm and varies little with eccentricity. Bilateral ablation of the visual cortex results in a reduction of the mean receptive field size of neurones in the SGS, and the point-image size is reduced by half. However, the shape of the point-image function is not affected. These results demonstrate that the spread of activity in the SC is nearly constant throughout the retinotopic map and that this is primarily a result of the direct retinal projection. Although the visual cortex has an expanded central representation in comparison with the SC, the corticotectal pathway does not exert a preferential influence on the central representation of the SC.     

Orientation bias of cat retinal ganglion cells: a reassessment

Summary For cat retinal ganglion cells whose receptive field centres were distributed in specified sections of the left visual field, the deviations of the major axis from the radial, horizontal, and circumferential directions were determined. The percentage of cells with deviations within ± 20° from the radial, horizontal, and circumferential directions were, respectively, 33%, 68%, 16%. In addition, comparison between values of deviation from the horizontal direction for cells located at eccentricities of 10° and 20° from the area centralis showed a statistically significant trend: the bias for the horizontal increased with eccentricity.     

Spatial summation in lateral geniculate nucleus and visual cortex

We have compared the spatial summation characteristics of cells in the primary visual cortex with those of cells in the dorsal lateral geniculate nucleus (LGN) that provide the input to the cortex. We explored the influence of varying the diameter of a patch of grating centred over the receptive field and quantitatively determined the optimal summation diameter and the degree of surround suppression for cells at both levels of the visual system using the same stimulus parameters. The mean optimal summation size for LGN cells (0.90 degrees) was much smaller than that of cortical cells (3.58 degrees). Virtually all LGN cells exhibited strong surround suppression with a mean value of 74%+/-1.61% SEM for the population as a whole. This potent surround suppression in the cells providing the input to the cortex suggests that cortical cells must integrate their much larger summation fields from the low firing rates associated with the suppression plateau of the LGN cell responses. Our data suggest that the strongest input to cortical cells will arise from geniculate cells representing areas of visual space located at the borders of a visual stimulus. We suggest that analysis of response properties by patterns centred over the receptive fields of cells may give a misleading impression of the process of the representation. Analysis of pattern terminations or salient borders over the receptive field may provide much more insight into the processing algorithms involved in stimulus representation.     

Representation of the visual field in the lateral intraparietal area of macaque monkeys: a quantitative receptive field analysis

The representation of the visual field in the primate lateral intraparietal area (LIP) was examined, using a rapid, computer-driven receptive field (RF) mapping procedure. RF characteristics of single LIP neurons could thus be measured repeatedly under different behavioral conditions. Here we report data obtained using a standard ocular fixation task during which the animals were required to monitor small changes in color of the fixated target. In a first step, statistical analyses were conducted in order to establish the experimental limits of the mapping procedure on 171 LIP neurons recorded from three hemispheres of two macaque monkeys. The characteristics of the receptive fields of LIP neurons were analyzed at the single cell and at the population level. Although for many neurons the assumption of a simple two-dimensional gaussian profile with a central area of maximal excitability at the center and progressively decreasing response strength at the periphery can represent relatively accurately the spatial structure of the RF, about 19% of the cells had a markedly asymmetrical shape. At the population level, we observed, in agreement with prior studies, a systematic relation between RF size and eccentricity. However, we also found a more accentuated overrepresentation of the central visual field than had been previously reported and no marked differences between the upper and lower visual representation of space. This observation correlates with an extension of the definition of LIP from the posterior third of the lateral intraparietal sulcus to most of the middle and posterior thirds. Detailed histological analyses of the recorded hemispheres suggest that there exists, in this newly defined unitary functional cortical area, a coarse but systematic topographical organization in area LIP that supports the distinction between its dorsal and ventral regions, LIPd and LIPv, respectively. Paralleling the physiological data, the central visual field is mostly represented in the middle dorsal region and the visual periphery more ventral and posterior. An anteroposterior gradient from the lower to the upper visual field representations can also be identified. In conclusion, this study provides the basis for a reliable mapping method in awake monkeys and a reference for the organization of the properties of the visual space representation in an area LIP extended with respect to the previously described LIP and showing a relative emphasis of central visual field. Electronic Publication     

Limitation of sensitization to injured parts of receptive fields in human skin C-nociceptors

Unmyelinated cutaneous mechano-heat fibers (CMH) in the peroneal nerve of healthy human volunteers were studied by means of a marking technique which allows stable recordings from identified single units over extended periods. Mechanoreceptive field sizes were 105±13 mm² in 25 units. These large receptive fields indicate extensive terminal branching of C fibers in the skin of foot and lower leg. Sensitization of CMHs was tested by assessment of thresholds for mechanical (von Frey hair) and heat stimuli before and after topical application of mustard oil (allyl isothiocyanate) and capsaicin (8-methyl-N-vanillyl-6-noneamide). While in a group of 14 CMHs the entire receptive field was treated with these irritant substances, in another group of 11 CMH units only parts of the receptive field were treated to check for signs of spreading sensitization through axon collaterals. Mustard oil application did not change mechanical thresholds, regardless of whether parts of or complete receptive fields were treated. However, mean heat thresholds dropped by 5.6° C to 36.5±1.5°C in completely treated receptive fields and by 5.7° C to 37.3±3.4° C in treated parts of receptive fields (primary sensitization). In contrast, heat thresholds in the non-treated parts did not change significantly (42.1±3.4° C vs 41.2±3.9° C), i.e. secondary sensitization to heat was lacking. The absence of primary sensitization to probing with von Frey hairs indicates that sensitization of insensitive C fibers and recruitment of insensitive axon collaterals may be more important for mechanical hyperalgesia than sensitization of conventional CMH units — apart from the contribution of central mechanisms. The lack of spread of sensitization to untreated parts of the receptive fields o CMHs (secondary sensitization) indicates that this fiber group is probably not involved in any form of secondary hyperalgesia to heating.     

The size and shape of rod and cone centres of cat retinal ganglion cells

Summary Receptive field centres of cat retinal ganglion cells, as mediated by rod and by cone inputs, were mapped as contours of iso-sensitivity at a mid-mesopic adapting luminance using, respec-tively, 452 nm-blue and 578 nm-yellow narrow-band lights at an intensity 1 log unit above threshold for the most sensitive locus. Based on the sizes and shapes of mapped rod and cone centres for 74 ganglion cells, four receptive field centre categories were distinguished. Cone and rod centres were usually elliptical, and in almost 60% of cells the major axis through the receptive field centre was oriented within ±20 ° of horizontal. In 69%, rod and cone centres were non-concentric, 66% had larger rod than cone centres — area ratios ranging from 0.6 1 to 2.9 1, and in only two cases was the rod centre actually smaller than the cone centre.     

‘Real-motion’ cells in visual area V2 of behaving macaque monkeys

Summary Extracellular recordings were made in area V2 of behaving macaque monkeys. Neurons were classified into three groups: non-oriented cells, oriented cells with antagonistic areas and oriented cells without antagonistic areas in their receptive field. All neurons were tested with standard visual stimulations in order to assess whether they gave different responses to the movement of a stimulus and to the movement of its retinal image alone, when the stimulus was motionless and the animal voluntarily moved its eyes. To do this, neuronal responses obtained when a moving stimulus swept a stationary receptive field (during steady fixation) and when a moving receptive field swept a stationary stimulus (during tracking eye movements), were compared. The receptive field stimulation at retinal level was physically the same in both cases, but only in the first was there actual movement of the visual stimulus. Control trials, where the monkeys performed tracking eye movements without any intentional receptive field stimulation, were also carried out. Out of a total of 263 neurons isolated in the central 10 deg representation of area V2, 101 were fully studied with the visual stimulation described above. Most of these (83/ 101; 82%) gave about the same response to the two situations. About 14% (14/101) gave a good response to stimulus movements during steady fixation and a very weak one to retinal image displacements of stationary stimuli during visual tracking. We have called neurons of this type real-motion cells (cf. Galletti et al. 1984). None of the non-oriented cells was a real-motion one, while about an equal percentage of real-motion cells was found among the oriented cells with and without antagonistic areas. Finally, we found only 4 neurons which showed behaviour opposite to that of real-motion cells, i.e. they showed a better response to displacement of the retinal image of stationary stimuli than to actual movement of stimuli. We suggest that real-motion cells might contribute to correctly evaluating movement in the visual field in spite of eye movements and that they might allow recognition of the movement of an object even if it moves across a non-patterned visual background. Present data on area V2, together with similar results observed in area V1 (Galletti et al. 1984; Battaglini et al. 1986), support the view that these two cortical areas analyse the movement in a parallel fashion along with many other characteristics of the visual stimulus.     

Analysis of retinal correspondence by studying receptive fields of rinocular single units in cat striate cortex

Summary The concept of corresponding retinal points was examined in terms of the binocular receptive fields of neurons in Area 17 of the cerebral cortex of the cat. Only a proportion of the binocular receptive field pairs can be accurately superimposed at the one time in a given plane. The fields which are not corresponding are said to show receptive field disparity. The attempt has been made to establish, on a quantitative basis, the parameters of the receptive field disparities that occur within 5° of the visual axis. A new method was used for defining the zero (vertical) meridian. Very effective paralysis of the extraocular muscles was achieved and the very small residual eye movements that occurred were regularly monitored so that corrections could be applied to the plotted positions of the receptive field pairs. The distribution of the receptive field disparities about the position of maximal correspondence has a range of about ±1.2° (S.D. 0.6°) in both the horizontal and vertical directions for fields in the vicinity of the visual axis. Panum's fusional area may represent the extent to which receptive fields in the one eye, all with the same visual direction, are linked to fellow members of a pair in the other eye over a range of receptive field disparities. A naso-temporal overlap of receptive fields occurs which is probably little if any more than can be accounted for on the basis of the disparity of receptive fields lying along the zero (vertical) meridian. When the extraocular muscles are paralyzed the eyes diverge and the binocular receptive field pairs are separated on the tangent screen. The distribution of the horizontal and vertical separations of the receptive field pairs have been examined.Selby Fellow of the Australian Academy of Sciences.     

A visual field dependent architecture for second order motion processing

The visual system exploits a cortical hierarchy to process complex inputs such as those defined by modulations of motion and/or texture. One class of visual stimuli, composed of alternate stripes of opposing motion requires at least 2 separate stages of computation within this cortical hierarchy, thought to involve cortical area V1 and extra-striate regions like global motion area MT respectively. Using a psychophysical task, we characterise sensitivity to such stimuli containing periodic spatial modulations of motion gradients as a function of the ratio of the spatial parameters at the two processing levels by manipulating the spatial properties of the carrier and modulator. We find band-passed functions for foveal stimulus presentations showing an optimum sensitivity at ratios in the range of r ≤ 10, informative of the coupling relationship between frequency channels at the carrier and modulator levels. An annulus stimulus (excluding the fovea) with a radius of 15.5° exhibited optima of sensitivity at r > 15. This difference in the optimal coupling between filtering stages reflects a processing architecture that changes with eccentricity, consistent with the previously observed smaller differences between mean receptive field sizes in striate and extra-striate filtering stages in the fovea compared to the periphery. This is also important for visual psychophysics when comparing sensitivity for first and second order stimuli across retinal eccentricity.     

Cortical magnification factor and contrast sensitivity to luminance-modulated chromatic gratings

Contrast sensitivity to luminance-modulated blue, green, red, and neutral gratings was measured at different spatial frequencies and eccentricities within 0-86 deg in the temporal visual field. Contrast sensitivity to gratings of constant area and spatial frequency was independent of wavelength composition but decreased with increasing eccentricity. When the gratings were scaled by the magnification factor of the human striate cortex to produce cortically similar stimulus conditions at different eccentricities (M-scaling), contrast sensitivities became independent of visual field location irrespective of grating colour. Using colour naming we found, in accordance with previous studies, that hue changed and saturation decreased when the eccentricity of a constant-size grating field increased. In contrast, the hue and saturation of M-scaled grating fields were independent of eccentricity. The results suggest that the effects of eccentricity on photopic colour vision can largely be counteracted by M-scaling which adjusts the spatial aspects of stimuli with respect to the decrease in ganglion cell density and increase in receptive field size towards the periphery of the visual field.     

Differences of visual field representation in the medial and lateral banks of the suprasylvian cortex (PMLS/PLLS) of the cat

Summary We have studied the orderliness of representation of visual space in the medial and lateral banks of the middle suprasylvian sulcus. Penetrations were made either parallel to the sulcus, in one bank or the other, or vertical, thus crossing the sulcus between the postero-medial (PMLS) and posterolateral (PLLS) divisions of this area. In some cases we found clear evidence for topographical order in the representation of the visual field with a tendency (greater in PMLS than in PLLS) for the receptive fields of cells recorded deeper in the walls of the sulcus to lie closer to the area centralis, but along many penetrations the receptive fields were so large and so scattered that no retinotopic arrangement could be discerned. In PMLS the receptive fields of the majority of units we studied were centred below and close to the horizontal meridian, whereas in PLLS they were distributed over both the upper and lower visual fields with an over-representation of the upper field. Receptive fields were significantly larger in PLLS (mean field area = 442.2 deg²) than in PMLS (mean area = 154.4 deg²); there was also less clear correlation between receptive field size and eccentricity in PLLS (correlation coefficient = +0.25) than in PMLS (corr. coeff. = +0.72). Analysis of the distance between the receptive field centres of consecutively recorded units demonstrated that the mean scatter in both PMLS and PLLS amounts to about half the average receptive field diameter. In summary the topographical representation of visual space is less orderly in PLLS, and may involve a wider area of the visual field. These findings may relate to the segregated visual cortical and extrageniculate thalamic connections that the medial and lateral banks of the LS receive.     

Response variability and orientation discrimination of single cells in striate cortex of cat

Summary The response of single cells in the striate cortex of cat to a moving light bar of variable orientation was measured by a method providing data on the mean response as well as the standard deviation (SD) at the different stimulus orientations.At the optimal stimulus orientation the SD was about 1/3 of the mean response. Marked differences in this respect were found between simple and complex cells, i.e., the SD for the simple cells was about 1/2 of the mean response and about 1/4 for the complex cells.The variation coefficient (Vc = SD/mean) was minimal at the optimal orientation and increased relatively in the same manner for simple and complex cells as the stimulus orientation was varied away from optimal orientation. The Vc varied with the mean response at optimal orientation in a nonlinear manner. A function is proposed which fits this relationship and which is equally applicable for both simple and complex cells.The mean orientation discrimination (MOD) was defined as that change in orientation angle away from the optimal which produced a response statistically different — on the 1 % level — from the response to the optimal orientation. There were differences in MOD between the two sides of the orientation tuning curve: the mean of the smaller of the two values was 13.5 deg and of the larger 19.7 deg. No significant difference in MOD was found between simple and complex cells despite the fact that the halfwidth of the tuning curves for the two cell types was 19.5 deg and 31.6 deg, respectively.The preciseness in localization of the most sensitive part within the receptive field of single cells was calculated from the variability in time of occurrence of the smallest interspike interval. The degree of preciseness was found to be of the order of 1/4 of the receptive field diameter in both simple and complex cells. When nonoptimal stimulus orientations were presented, the preciseness significantly decreased in complex cells whereas it remained unchanged in simple cells.It is suggested that the same type of intracortical wiring produces orientation selectivity in simple and complex cells, and that the differences in tuning width are mainly due to a larger extension of inhibitory fields in the simple cells. Considering the cortical visual cells as elementary units in a network built for orientation detection and discrimination, the tuning width seems of minor importance for that function.     

Retrograde transneuronal transport of wheat-germ agglutinin to the retina from visual cortex in the cat

Summary The stability of visual perception despite eye movements suggests the existence, in the visual system, of neural elements able to recognize whether a movement of an image occurring in a particular part of the retina is the consequence of an actual movement that occurred in the visual field, or self-induced by an ocular movement while the object was still in the field of view. Recordings from single neurons in area V3A of awake macaque monkeys were made to check the existence of such a type of neurons (called real-motion cells; see Galletti et al. 1984, 1988) in this prestriate area of the visual cortex. A total of 119 neurons were recorded from area V3A. They were highly sensitive to the orientation of the visual stimuli, being on average more sensitive than V1 and V2 neurons. Almost all of them were sensitive to a large range of velocities of stimulus movement and about one half to the direction of it. In order to assess whether they gave different responses to the movement of a stimulus and to that of its retinal image alone (self-induced by an eye movement while the stimulus was still), a comparison was made between neuronal responses obtained when a moving stimulus swept a stationary receptive field (during steady fixation) and when a moving receptive field swept a stationary stimulus (during tracking eye movement). The receptive field stimulation at retinal level was physically the same in both cases, but only in the first was there actual movement of the visual stimulus. Control trials, where the monkeys performed tracking eye movements without any intentional receptive field stimulation, were also carried out. For a number of neurons, the test was repeated in darkness and against a textured visual background. Eighty-seven neurons were fully studied to assess whether they were real-motion cells. About 48% of them (42/87) showed significant differences between responses to stimulus versus eye movement. The great majority of these cells (36/42) were real-motion cells, in that they showed a weaker response to visual stimulation during tracking than to the actual stimulus movement during steady fixation. On average, the reduction in visual response during eye movement was 64.0 ± 15.7% (SD). Data obtained with a uniform visual background, together with those obtained in darkness and with textured background, indicate that real-motion cells receive an eye-motion input, either retinal or extraretinal in nature, probably acting presynaptically on the cell's visual input. In some cases, both retinal and extraretinal eye-motion inputs converge on the same real-motion cell. No correlation was observed between the real-motion behaviour and the sensitivity to either orientation or direction of movement of the visual stimulus used to activate the receptive field, nor with the retinotopic location of the receptive field. We suggest that the visual system uses real-motion cells in order to distinguish real from self-induced movements of retinal images, hence to recognize the actual movement in the visual field. Based on psychophysical data, the hypothesis has been advanced of an internal representation of the field of view, stable despite eye movement (cf. MacKay 1973). The real-motion cells may be neural elements of this network and we suggest that the visual system uses the output of this network to properly interpret the large number of sensory changes resulting from exploratory eye movements in a stable visual world.     

Spatio-temporal interactions and the spatial phase preferences of visual neurons

Summary We recorded single neuron responses in the cat's lateral geniculate nucleus (LGN) and visual cortex to compound stimuli composed of two sinusoidal gratings in a 21 frequency ratio. To probe visual receptive field symmetry, we varied the relative spatial phase of the two components and measured the effect on neuronal responses. We expected that on-center LGN neurons would respond best to gratings combined in positive cosine (bright bar) phase, while off-center LGN neurons would respond best to gratings combined in negative cosine (dark bar) phase. When drifting stimuli were used, cells' phase preferences were roughly 90 deg away from the expected values; when stationary, contrast-modulated stimuli were used, phase preferences were as originally predicted. Computer simulations showed that this discrepancy could be explained by taking into account the cells' temporal properties. Thus, tests using drifting stimuli confound the spatial structure of visual neural receptive fields with their temporal response characteristics. A small sample of data from cortical neurons reveals the same confound.     

Classification of receptive field properties in cat visual cortex

Summary The properties of the receptive fields of visual cortex neurons of cats were studied manually and by a computer controlled system using single lines, double lines and multiple lines (gratings). The multiple selectivities of each of the receptive fields studied make it necessary to abandon the concept that each cell functions as a feature detector. Instead, an attempt was made to classify the receptive field properties with the aim to delineate the transfer functions (of the total networks) served by each property. When tested with one-line stimulus, cells with simple receptive field properties diffefed from cells with complex receptive field properties as to their velocity selectivity (simple: 1 ° to 3 °/s; complex: 4 ° to 10 °/s), spontaneous activity (lower for cells with simple properties), optimal firing rate (lower for cells with simple properties) and receptive field size (smaller for cells with simple properties) but not for orientation and direction selectivity. When tested with a 2-lines moving stimulus, the responses of cells with simple properties were facilitated by the progressive separation of the lines whereas the responses of cells with complex receptive field properties were inhibited. When multiple lines, i.e. gratings, were used, an equivalence between simple and X properties and complex and Y properties was shown, while the sustained/transient classification proved to be independent of the simple/complex (X/Y) classification. Thus, receptive field properties can be classified into three categories: one reflects the input to the receptive fields; a second deals with the interactive properties of the fields; while a third appears more related to the overall properties of the network.This research was supported by a postdoctoral fellowship from the Medical Research Council of Canada to Maurice Ptito, a predoctoral fellowship from the National Research Council of Canada to Maryse C. Lassende and NIMH Grant MH12970 and NIMH Career Research Award MH15214 to Karl H. Pribram