The directional effects of passive eye movement on the directional visual responses of single units in the pigeon optic tectum

 We have investigated the visual responses of 184 single units located in the superficial layers of the optic tectum (OT) of the decerebrate, paralysed pigeon. Visual responses were similar to those reported in non-decerebrate preparations; most units responded best to moving visual stimuli, 18% were directionally selective (they had a clear preference for a particular direction of visual stimulus movement), 76% were plane-selective (they responded to movement in either direction in a particular plane). However, we also found that a high proportion of units showed some sensitivity to the orientation of visual stimuli. We examined the effects of extraocular muscle (EOM) afferent signals, induced by passive eye movement (PEM), on the directional visual responses of units. Visual responses were most modified by particular directions of eye movement, although there was no unique relationship between the direction of visual stimulus movement to which an individual unit responded best and the direction of eye movement that caused the greatest modification of that visual response. The results show that EOM afferent signals, carrying information concerning the direction of eye movement, reach the superficial layers of the OT in the pigeon and there modify the visual responses of units in a manner that suggests some role for these signals in the processing of visual information. Received: 17 June 1996 / Accepted: 29 April 1997     

Functional properties of neurons in middle temporal visual area of the macaque monkey. I. Selectivity for stimulus direction, speed, and orientation

1. Recordings were made from single units in the middle temporal visual area (MT) of anesthetized, paralyzed macaque monkeys. A computer-driven stimulator was used to make quantitative tests of selectivity for stimulus direction, speed, and orientation. The data were taken from 168 units that were histologically identified as being in MT. 2. The results confirm previous reports of a high degree of direction selectivity in MT. The response above background to stimuli moving in a unit's preferred direction was, an average, 10.9 times that to stimuli moving in the opposite direction. There was a marked tendency for nearby units to have similar preferred directions. 3. Most units were also sharply tuned for the speed of stimulus motion. For some cells the response fell to less than half-maximal at speeds only a factor of two from the optimum; on average, responses were greater than half-maximal only over a 7.7-fold range of speed. The distribution of preferred speeds for different units was unimodal, with a peak near 32 degrees/s; the total range of preferred speeds extended from 2 to 256 degrees/s. Nearby units generally responded best to similar speeds of motion. 4. Most units in MT showed selectivity for stimulus orientation when tested with stationary, flashed bars. However, stationary stimuli generally elicited only brief responses; when averaged over the duration of the stimulus, the responses were much less than those to moving stimuli. The preferred orientation was usually, but not always, perpendicular to the preferred direction of movement. 5. A comparison of the results of the present study with a previous quantitative investigation in the owl monkey shows a striking similarity in response properties in MT of the two species. 6. The presence of both direction and speed selectivity in MT of the macaque suggests that this area is more specialized for the analysis of visual motion than has been previously recognized.     

The effect of a chronic lesion in cortical area 17 on the visual responses of units in area 18 of the cat.

1. Lesions were made in cortical Area 17 (Visual I) of eight cats which were then allowed to recover. 2. During acute experiments between 1 and 11 weeks after the lesion the activity of Area 18 (Visual II) units was recorded and the results were compared with those obtained in normal cats. 3. The receptive fields were similar in size and distribution in the two groups but the lesioned animals had a much higher proportion of units unaffected by a visual stimulus and a higher proportion of the visually responsive units lacked specific direction or orientation preference. 4. Of six units which were tested in Area 18 of cats with lesions five showed variability of their direction or orientation preference with time. 5. The effects described above are most probably due to destruction of the corticocortical pathway which connects Areas 17 and 18. Some units in Area 18 appear to be driven by visual stimuli via their geniculate input alone. The corticocortical (Area 17 to 18) pathway may normally confer additional specificity on some of these units.     

Selectivity for orientation and direction of motion of single neurons in cat striate and extrastriate visual cortex

1. We consider the consequences of the orientation selectivity shown by most cortical neurons for the nature of the signals they can convey about the direction of stimulus movement. On theoretical grounds we distinguish component direction selectivity, in which cells are selective for the direction of movement of oriented components of a complex stimulus, from pattern direction selectivity, or selectivity for the overall direction of movement of a pattern irrespective of the directions of its components. We employed a novel test using grating and plaid targets to distinguish these forms of direction selectivity. 2. We studied the responses of 280 cells from the striate cortex and 107 cells from the lateral suprasylvian cortex (LS) to single sinusoidal gratings to determine their orientation preference and directional selectivity. We tested 73 of these with sinusoidal plaids, composed of two sinusoidal gratings at different orientations, to study the organization of the directional mechanisms within the receptive field. 3. When tested with single gratings, the directional tuning of 277 oriented cells in area 17 had a mean half width of 20.6 degrees, a mode near 13 degrees, and a range of 3.8-58 degrees. Simple cells were slightly more narrowly tuned than complex cells. The selectivity of LS neurons for the direction of moving gratings is not markedly different from that of neurons in area 17. The mean direction half width was 20.7 degrees. 4. We evaluated the directional selectivity of these neurons by comparing responses to stimuli moved in the optimal direction with those elicited by a stimulus moving in the opposite direction. In area 17 about two-thirds of the neurons responded less than half as well to the non-preferred direction as to the preferred direction; two-fifths of the units responded less than one-fifth as well. Complex cells showed a somewhat greater tendency to directional bias than simple cells. LS neurons tended to have stronger directional asymmetries in their response to moving gratings: 83% of LS neurons showed a significant directional asymmetry. 5. Neurons in both areas responded independently to each component of the plaid. Thus cells giving single-lobed directional-tuning curves to gratings showed bilobed plaid tuning curves, with each lobe corresponding to movement in an effective direction by one of the two component gratings within the plaid. The two best directions for the plaids were those at which one or other single grating would have produced an optimal response when presented alone.(ABSTRACT TRUNCATED AT 400 WORDS)     

Organization of orientation and direction selectivity in areas 17 and 18 of cat cerebral cortex

1. The organization of subunits and sequences subserving preferred stimulus orientation and preferred direction of stimulus motion in cat cerebral cortical areas 17 and 18 was determined by making vertical, tangential, and oblique microelectrode penetrations into those areas. 2. Quantitative measurements of direction selectivity indicated that not all shades of direction selectivity are equally represented in area 17. Peaks in the distribution of direction indices may correspond to the bidirectional, direction biased, and direction selective categories used in qualitative studies. 3. The relationship between preferred direction and location in the visual field was examined for units with receptive fields centered more than 15 degrees from the area centralis. Simple cells had orientation preferences that tended to be parallel to radii extending out from the area centralis. Wide-field complex cells had orientation preferences that tended to be parallel to concentric circles centered on the area centralis; the direction preferences of this group were biased toward motion away from the area centralis. 4. Unit pairs separated by 200 microns or less were 4.2 times as likely to have the same preferred direction as to have opposite preferred directions, indicating that, on average, strings of five neurons have similar direction preferences. 5. Tracks in area 18 showed a similar pattern to those in area 17. 6. In the vertical tracks in area 17 a small proportion (12%) of the units recorded in infragranular layers had preferred orientations that deviated 30 degrees or more from the first unit recorded in the same column. The presence of these cells most likely reflects the relative crowding of columns in infragranular layers, which occurs at the crown of the lateral gyrus. Columns with such large jumps in preferred orientation were not observed in area 18, which occupies a relatively flat region of cortex. 7. In both areas 17 and 18 direction preference in vertical tracks usually reversed at least once, either between supra- and infragranular layers or within infragranular layers. Along these same tracks, orientation preference usually did not change. 8. In tangential tracks, preferred direction and orientation preferences changed together in small increments. Occasionally a large jump in preferred direction would occur with only a small change in preferred orientation. These large jumps were considered to mark the boundaries of the direction sequences. Most frequently these boundaries were separated by 400-600 microns. This value is approximately half the size of a complete set of orientation preferences (700-1,200 microns).(ABSTRACT TRUNCATED AT 400 WORDS)     

Quantitative analysis of visual receptive fields of neurons in nucleus of the optic tract and dorsal terminal nucleus of the accessory optic tract in macaque monkey

1. The visual receptive field properties of neurons in the nucleus of the optic tract (NOT) in the pretectum and the dorsal terminal nucleus (DTN) of the accessory optic tract were analyzed quantitatively in anesthetized, paralyzed macaque monkeys. 2. Visual latencies to reversals in direction of stimulus movement ranged from 40 to 80 ms [61 +/- 13.5 (SD) ms]. 3. All neurons increased their discharge rate during ipsiversive movement and decreased their ongoing activity during contraversive movement of single stimuli or whole-field random dot patterns. The population of neurons in the left NOT-DTN was excited most strongly by leftward movement pointing 4 degrees down; neurons in the right NOT-DTN were excited most strongly by rightward movement pointing 6 degrees down. The mean angle between the directions yielding the highest and the lowest discharge rate in the two populations of NOT-DTN neurons was 177 degrees. 4. The deviation of the preferred excitatory directions from the horizon in individual neurons varied with recording depth. Within the first 500 microns below the midbrain surface, neurons preferred near-horizontal directions, whereas neurons recorded more deeply preferred more oblique directions of stimulus movement. 5. The tuning widths of NOT-DTN neurons in the preferred excitatory direction were very broad. The mean halfwidth defined as the range of directions eliciting responses greater than 50% of the maximum was 127 +/- 25 degrees. 6. Moving a random dot pattern and a single bar of light simultaneously but in opposite directions caused NOT-DTN neurons to increase their discharge rate as soon as one of the two stimuli moved in the ipsiversive direction. The reduction in overall discharge rates when two stimuli moved in opposite directions indicates mainly inhibitory interactions. 7. All NOT-DTN neurons could be activated from both eyes. Interactions between the two eyes were modest and unspecific. Misalignment of the visual axes of the two eyes had no influence on response strength. 8. Optimal speeds of stimulus movement varied widely for different NOT-DTN neurons. The effective range of speeds to elicit direction-selective responses in the total population was very broad (0.1400 degrees/s. With oscillating horizontal stimulation, NOT-DTN neurons followed repetition rates up to 4 Hz at excursions of 40 degrees. Speeds greater than 500 degrees/s were either not effective or resulted in a suppression of ongoing activity in all directions of movement.(ABSTRACT TRUNCATED AT 400 WORDS)     

Receptive-field characteristics of single neurons in lateral suprasylvian visual area of the cat.

The visual receptive fields of 213 cells in the lateral suprasylvian visual cortex (LS, or Clare-Bishop area) were studied in cats anesthetized with nitrous oxide. Eighty-one percent of the cells were directionally selective. They responded poorly to stationary stimuli flashed on or off, but gave a directionally selective response to stimuli moving through the receptive field. Most of these had a single preferred direction and an opposite null direction. They typically responded to a range of directions of stimulus movement from 45 to 90 degrees to either side of the preferred direction. Small stimuli (1-2 degrees or smaller) typically were effective and 87% of the directionally selective cells showed spatial summation. About 32% had inhibitory mechanisms which decreased the response of the cell if the stimulus exceeded a maximum size. There was little or no evidence that LS area cells were orientation selective or sensitive to variations in stimulus shape independent of size.     

Neuronal responsiveness in areas 19 and 21a,and the posteromedial lateral suprasylvian cortex of the cat

Responsiveness to slits and pattern stimuli was quantified in a total of 68 cells sampled in the posterior extreme of the lateral suprasylvian (PS) cortex as response indices. The cells were studied in relationship to their locations in several subareas of the PS cortex, including areas 19 (n=15) and 21a (n=32) and the posteromedial lateral suprasylvian cortex (PMLS; n=21). These subareas were identified based on retrograde labelling from area 17 and also supplemented with photic responsiveness. This analysis revealed that each cortical area contains cells expressing different combinations of stimulus features. Area 19 contained two major groups of cells: (1) those with strong end-stop selectivity combined with moderate orientation or direction selectivity, and (2) those with weak end-stop selectivity combined with strong orientation selectivity. The groups of cells with strong or moderate orientation selectivity showed a strong preference for stripe over visual noise patterns and relatively large modulatory responses to motion of individual stripes. The PMLS contained one major group of cells with strong end-stop and direction selectivities and with poor orientation selectivity. They also showed stronger preference for visual noise than cells in the other cortical areas and rather weak modulatory responses. Area 21a contained only one group of cells with strong orientation selectivity and length summation property rather than end-stop selectivity, and they also lacked direction selectivity. These cells exhibited a strong preference for stripe patterns and moderate or weak modulatory responses. Altogether, these findings indicate that each cortical area is specialized in expressing different stimulus features. The two groups of cells in area 19 may encode the position and motion of discontinuous visual elements such as corners and line ends and continuous elements such as lines and edges. PMLS cells may encode the motion of single elements or associated motion of multiple discontinuous elements such as textures and backgrounds. Area 21a cells may specifically encode the orientation of long, continuous elements such as lines and edges. In support of this view, two types of statistical analyses demonstrated that the combinations of the response properties expressed in individual PS cells are highly correlated with their locations in cortical areas and that the anatomical locations of individual PS cells are reliably predicted from the sets of response indices expressed in these cells.     

A novel mechanism of response selectivity of neurons in cat visual cortex

The spiking of cortical neurons critically depends on properties of the afferent stimuli. In the visual cortex, neurons respond selectively to the orientation and direction of movement of an object. The orientation and direction selectivity is improved upon transformation of the membrane potential changes into trains of action potentials. To address the question of whether the transformation of the membrane potential changes into spiking of a cell depends on the stimulus orientation and the direction of movement, we made intracellular recordings from the cat visual cortex in vivo during presentation of moving gratings of different orientations. We found that the relationship between the membrane polarization and the firing rate (input-output transfer function) depended on the stimulus orientation. The input-output transfer function was steepest during responses to the optimal stimulus; membrane depolarization of a given amplitude led to generation of more action potentials when evoked by an optimal stimulus than during non-optimal stimulation. The threshold for the action potential generation did not depend on stimulus orientation, and thus could not account for the observed difference in the transfer function. Oscillations of the membrane potential in the γ-frequency range (25–70 Hz) were most pronounced during optimal stimulation and their strength changed in parallel with the changes in the transfer function, suggesting a possible relationship between the two parameters. We suggest that the improved input-output relationship of neurons during optimal stimulation represents a novel mechanism that may contribute to the final sharp orientation selectivity of spike responses in the cortical cells.     

Quantitative study of cortical orientation selectivity in visually inexperienced kitten.

1. Extracellular recordings were made from single units in the visual cortices of six kittens deprived of experience with pattern vision by binocular lid suture. 2. Selectivity for stimulus orientation was quantitatively assessed in 98 units; 90 responded selectively to the orientation of a moving bar stimulus, the remainder responding nonselectively or too poorly to classify. Cells in these visually inexperienced kittens were similar in their degree of selectivity for orientation to cells tested in adult cats. However, responses tended to be weaker and somewhat more erratic. 3. About half the cells in this simple responded to both directions of stimulus motion at the optimal orientation. Most of those responding to only one direction of motion were considered orientation rather than direction selective because they responded more strongly or more selectively to a moving bar than to a moving spot. 4. Cells appeared to be organized within the cortex in a pattern similar to that found in adult cats, with cells in one column selective for the same orientation, and adjacent column having similar preferred orientations. 5. It is concluded that selectivity for stimulus orientation in the cat's visual cortex is innately determined.     

Visual-response properties of neurons in turtle basal optic nucleus in vitro

1. The spike responses of 105 cells to visual-stimulus movement were analyzed in the turtle's basal optic nucleus (BON) in vitro in the absence of the telencephalon. All cells were direction sensitive (DS) and were driven solely by stimulation of the contralateral eye. These cells had large receptive fields and had vigorous responses to moving, textured patterns. Small moving spots generated only weak responses from these cells, as did the onset or offset of diffuse light flashes. 2. The direction tuning of BON cells was quite broad with most back and forth responses being DS. In fact, for 86% of the cells, there were seven to nine axes (out of 9 total, in 20 degrees increments) for which response to movement in one direction was at least twice that for the opposite direction. In instances where spontaneous activity was relatively high, a suppression of that spike firing was evident when the stimulus moved in directions opposite to preferred stimulus directions. 3. Cells preferring many different directions are found in the BON. More cells preferred inferior-temporal directed motion (49%), compared to superior-temporal (35%) and nasal stimuli (13%). 4. BON cells remained DS over 3 log units of velocity, with their strongest responses between 1 and 50 degrees/s. Responses were often non-DS for stimuli moving slower than 0.1 degrees/s. 5. The receptive fields of BON cells were large and occupied different parts of the retina. When different subregions of a receptive field were stimulated, the cell's directional tuning always remained the same as the full field direction tuning. 6. Thus, BON cells seem well-suited for the analysis of global, visual-field motion in any direction, performed by the accessory optic system. Other brain stem pathways necessary for optokinetic reflexes can be elucidated with the use of this whole-brain, eyes-attached in vitro preparation.     

The analysis of image motion by the rabbit retina

1. Micro-electrode recordings were made from rabbit retinal ganglion cells or their axons. Of particular interest were direction-selective units; the common on-off type represented 20.6% of the total sample (762 units), and the on-type comprised 5% of the total.2. From the large sample of direction-selective units, it was found that on-off units were maximally sensitive to only four directions of movement; these directions, in the visual field, were, roughly, anterior, superior, posterior and inferior. The on-type units were maximally sensitive to only three directions: anterior, superior and inferior.3. The direction-selective unit's responses vary with stimulus velocity; both unit types are more sensitive to velocity change than to absolute speed. On-off units respond to movement at speeds from 6'/sec to 10 degrees /sec; the on-type units responded as slowly as 30'/sec up to about 2 degrees /sec. On-type units are clearly slow-movement detectors.4. The distribution of direction-selective units depends on the retinal locality. On-off units are more common outside the ;visual streak' (area centralis) than within it, while the reverse is true for the on-type units.5. A stimulus configuration was found which would elicit responses from on-type units when the stimulus was moved in the null direction. This ;paradoxical response' was shown to be associated with the silent receptive field surround.6. The four preferred directions of the on-off units were shown to correspond to the directions of retinal image motion produced by contractions of the four rectus eye muscles. This fact, combined with data on velocity sensitivity and retinal distribution of on-off units, suggests that the on-off units are involved in control of reflex eye movements.7. The on-off direction-selective units may provide error signals to a visual servo system which minimizes retinal image motion. This hypothesis agrees with the known characteristics of the rabbit's visual following reflexes, specifically, the slow phase of optokinetic nystagmus.     

Directional selectivity in the responses of units in cat primary visual cortex to passive eye movement

The responses of single units in the primary visual cortex (Area 17) of anaesthetized, paralysed cats, to passive movement of the ipsilateral eye were studied. Responses to passive eye movement were found in about one-third of the cortical units isolated. Appropriate control experiments excluded visual, auditory and cutaneous inputs as the source of the effective signal during passive eye movement. The magnitudes of the responses to a number (usually four) of radial directions of passive eye movement were estimated from sets of peristimulus time histograms "interleaved" in time. Units were defined as "radially selective" if the responses to movement along one radius (e.g. vertically upwards) exceeded that along at least one other orthogonal radius (e.g. horizontal-temporal). Of 60 units tested, 53 (88%) were "radially selective" according to this definition. Some of the "radially selective" units showed an additional type of specificity to passive eye movement: (a) Some units responded preferentially to movement along one of the arcs of passive eye movement which were tested (e.g. vertical movement above the equator of the orbit). These units we have called "arc selective". (b) Other units were sensitive to the direction of movement and preferred movement in a particular direction over more than one arc (e.g. horizontal movement towards the temporal side in both nasal and temporal halves of the orbit). These we have called "direction selective". Twenty-one "radially selective" units showed one of these additional properties, nine were arc selective and twelve were direction selective. The implications of these results for the understanding of the function of orbital proprioceptive signals in the cortex are discussed briefly. Responses to passive eye movement were found in all of layers II-VI in Area 17 and the implications of this for the understanding of the pathway by which orbital proprioceptive signals reach the primary visual cortex are discussed. The experiments have shown that many units in cat visual cortex respond to passive eye movement and that most of these units have some specificity for particular radial directions of movement while some have additional specific properties. We believe that these properties of radial, directional and arc sensitivity are likely to be important in understanding the function of the orbital proprioceptive signal which arises during eye movement and they are particularly interesting in relation to the findings of others that this proprioceptive signal appears to be concerned in the normal development of visual properties in the cortex and in the control of visually guided movement in adult cats.     

Response of cat cutaneous mechanoreceptors to punctate and grating stimuli

Cat hairy skin type I, type II, and field mechanoreceptor response characteristics were studied by drawing punctate and grating stimuli across a unit's receptive field (RF). Area RF maps were generated with scans covering approximately 1 X 1 cm using different vertical loadings, scan velocities, and scan orientations. The results from stimulating with a 1 g mass, 1 mm diam, rounded-tip punctate probe indicate that type I units display essentially invariant response topographies as a function of stimulus parameters. Type II units, presumably because of their unidirectional stretch sensitivity, showed response differences that were a function of the scan orientation. Field units, although not considered to be important for stimulus feature extraction, also displayed a directionally sensitive response profile. Changing the vertical loading or the scanning velocity of the stimulus had a minimal affect on the resulting RF profiles for any of the units studied. Grating stimuli with periods of 0.25-2.0 mm were scanned over receptors to study their grating discrimination ability. Different scan directions were used to study the directional sensitivity of a unit. Both type I and II units had optimal scan orientations for minimum grating period discrimination. For type I units, the dome orientation pattern and interdome sensitivities seemed to be a factor in determining the scan direction for minimum grating period detection. For type II units it was equivocal whether scanning parallel or orthogonal to the direction of maximum stretch sensitivity was better for grating detection. Grating response results were qualitatively similar to the results reported by others for receptors in the glabrous skin of primates. Both type I and II units could reliably discriminate gratings with periods of 0.75 mm and could marginally discriminate gratings with 0.5-mm periods. Field units were not well represented in this study, but those units that were studied were unable to unambiguously discriminate a 2.0-mm period grating, the largest one used.     

Visual motion response properties of neurons in dorsolateral pontine nucleus of alert monkey

1. In this study we sought to characterize the visual motion processing that exists in the dorsolateral pontine nucleus (DLPN) and make a comparison with the reported visual responses of the middle temporal (MT) and medial superior temporal (MST) areas of the monkey cerebral cortex. The DLPN is implicated as a component of the visuomotor interface involved with the regulation of smooth-pursuit eye movements, because it is a major terminus for afferents from MT and MST and also the source of efferents to cerebellar regions involved with eye-movement control. 2. Some DLPN cells were preferentially responsive to discrete (spot and bar) visual stimuli, or to large-field, random-dot pattern motion, or to both discrete and large-field visual motion. The results suggest differential input from localized regions of MT and MST. 3. The visual-motion responses of DLPN neurons were direction selective for 86% of the discrete visual responses and 95% of the large-field responses. Direction tuning bandwidths (full-width at 50% maximum response amplitude) averaged 107 degrees and 120 degrees for discrete and large-field visual motion responses, respectively. For the two visual response types, the direction index averaged 0.95 and 1.02, indicating that responses to stimuli moving in preferred directions were, on average, 20 and 50 times greater than responses to discrete or large-field stimulus movement in the opposite directions, respectively. 4. Most of the DLPN visual responses to movements of discrete visual stimuli exhibited increases in amplitude up to preferred retinal image speeds between 20 and 80 degrees/s, with an average preferred speed of 39 degrees/s. At higher speeds, the response amplitude of most units decreased, although a few units exhibited a broad saturation in response amplitude that was maintained up to at least 150 degrees/s before the response decreased. Over the range of speeds up to the preferred speeds, the sensitivity of DLPN neurons to discrete stimulus-related, retinal-image speed averaged 3.0 spikes/s per deg/s. The responses to large-field visual motion were less sensitive to retinal image speed and exhibited an average sensitivity of 1.4 spikes/s per deg/s before the visual response saturated. 5. DLPN and MT were quantitatively comparable with respect to degree of direction selectivity, retinal image speed tuning, and distribution of preferred speeds. Many DLPN receptive fields contained the fovea and were larger than those of MT and more like MST receptive fields in size.(ABSTRACT TRUNCATED AT 400 WORDS)     

How moving visual stimuli modulate the activity of the substantia nigra pars reticulata

The orientation of spatial attention via saccades is modulated by a pathway from the substantia nigra pars reticularis (SNr) to the superior colliculus, which enhances the ability to respond to novel stimuli. However, the algorithm whereby the SNr translates visual input to saccade-related information is still unknown. We recorded extracellular single-unit responses of 343 SNr cells to visual stimuli in anesthetized cats. Depending on the size, velocity and direction of the visual stimulus, SNr neurons responded by either increasing or decreasing their firing rate. Using artificial neuronal networks, visual SNr neurons could be classified into distinct groups. Some of the units showed a clear preference for one specific combination of direction and velocity (simple neurons), while other SNr neurons were sensitive to the direction (direction-tuned neurons) or the velocity (velocity-tuned neurons) of the movement. Furthermore, a subset of SNr neurons exhibited a narrow inhibitory/excitatory domain in the velocity/direction plane with an opposing surround (concentric neurons). According to our results, spatiotemporally represented visual information may determine the discharge pattern of the SNr. We suggest that the SNr utilizes spatiotemporal properties of the visual information to generate vector-based commands, which could modulate the activity of the superior colliculus and enhance or inhibit the reflexive initiation of complex and accurate saccades.     

Visual cells in the pontine nuclei of the cat.

Two hundred and thirty-two visually activated neurones were recorded in a small area of the rostral pontine nuclei of cats. The location of visually activated neurones was coextensive with the input from visual areas of cat's cortex as determined by degeneration studies. 2. Pontine visual cells could only be driven by visual stimuli. Cells responsive to somatosensory or auditory stimuli were also found in different regions in rostral pontine nuclei. They too responded to only one modality. 3. 96% of the cells were directionally selective. 4. Pontine visual cells were responsive to a wide range of stimulus speeds. Some cells responded to targets moving as fast as 1000 degrees/sec without losing directional selectivity. No pontine visual cells gave a clearly sustained response to a stationary stimulus. 5. Exact stimulus configurations were not critical. Large fields containing many spots were the most effective stimuli for 50% of the cells. Inhibition of responses depending upon stimulus dimensions, direction of movement, or location in the visual field was found for many cells. 6. Receptive field dimensions were large, ranging in size from 3 degrees X 4 degrees to more than an entire hemifield. 7. 94% of the cells had receptive fields which were centred in the contralateral hemifield. 8. 98% of the cells could be driven from both eyes. 9. The properties of the pontine visual cells suggest a corticopontocerebellar pathway sensitive to a wide range of speeds and directions of movement, but not sensitive to precise form.     

Response properties of single units in the lateral terminal nucleus of the accessory optic system in the behaving primate

1. To determine the potential role of the primate accessory optic system (AOS) in optokinetic and smooth-pursuit eye movements, we recorded the activity of 110 single units in a subdivision of the AOS, the lateral terminal nucleus (LTN), in five alert rhesus macaques. All monkeys were trained to fixate a stationary target spot during visual testing and to track a small spot moving in a variety of visual environments. 2. LTN units formed a continuum of types ranging from purely visual to purely oculomotor. Visual units (50%) responded best for large-field (70 x 50 degrees), moving visual stimuli and had no response associated with smooth-pursuit eye movement; some responded during smooth pursuit in the dark, but the response disappeared if the target was briefly extinguished, indicating that their smooth-pursuit-related response reflected activation of a parafoveal receptive field. Eye movement and visual units (36%) responded both for large, moving visual stimuli and during smooth-pursuit eye movements made in the dark. Eye movement units (14%) discharged during smooth-pursuit or other eye movements but showed no evidence of visual sensitivity. 3. Essentially all (98%) LTN units were direction selective, responding preferentially during vertical background and/or smooth-pursuit movement. The vast majority (88%) preferred upward background and/or eye movement. During periodic movement of the large-field visual background while the animal fixated, their firing rates were modulated above and below rather high resting rates. Although LTN units typically responded best to movement of large-field stimuli, some also responded well to small moving stimuli (0.25 degrees diam). 4. LTN units could be separated into two populations according to their dependence on visual stimulus velocity. For periodic triangle wave stimuli, both types had velocity thresholds less than 3 degrees/s. As stimulus velocity increased above threshold, the activity of one type reached peak firing rates over a very narrow velocity range and remained nearly at peak firing for velocities from approximately 4-80 degrees/s. The firing rates of the other type exhibited velocity tuning in which the firing rate peaked at an average preferred velocity of 13 degrees/s and decreased for higher velocities. 5. A close examination of firing rates to sinusoidal background stimuli revealed that both unit types exhibited unusual behaviors at the extremes of stimulus velocity.(ABSTRACT TRUNCATED AT 400 WORDS)     

The primary visual cortex in the mouse: Receptive field properties and functional organization

Summary Receptive field (RF) characteristics of cells in primary visual cortex of the mouse (C57B16 strain) were studied by single unit recording. We have studied the functional organization of area 17 along both the radial and tangential dimensions of the cortex. Eighty seven percent of the visual neurons could be classified according to their responses to oriented stimuli and to moving stimuli. Cells which preferred a flashed or moving bar of a particular orientation and responded less well to bars of other orientations or to spots, were classified as orientation selective (simple RF 23%, complex RF 18%). The majority of them were, moreover, unidirectional (24%). All orientations were roughly equally represented. Cells with oriented RFs were recorded mostly in the upper part of cortical layers II–III, where they appeared to be clustered according to their preferred orientation. Neurons that responded equally well to spots and bars of all orientations (46%) were classified as non-oriented; among these neurons there were several subcategories. Cells which responded equally well to spots and bars but preferred stimuli moving along one or both directions of a particular axis were classified as non oriented asymmetric cells (unidirectional 14%, bidirectional 4%). They were recorded mainly in supra- and infra-granular layers. Cells unaffected by stimulus shape and orientation which responded equally well to all directions of movement were classified as symmetric units. They had receptive field classified as ON (11%), OFF (1%), ON/ OFF (11%), or were unresponsive to stationary stimuli (5%). These cells were mostly found in layer IV, in which they constituted the majority of recorded cells. There was no apparent correlation between the functional type and size of RFs. However, the greatest proportion of small RFs was found in layer IV. In the binocular segment of the mouse striate cortex, the influence of the contralateral eye predominated. Ninety five percent of cells in this segment were driven through the contralateral eye. However, 70% of cells were binocularly activated, showing that considerable binocular integration occured in this cortical segment. Ocular dominance varied less along the radial than along the tangential dimension of the cortex.     

Processing of kinetically defined boundaries in areas V1 and V2 of the macaque monkey

We recorded responses in 107 cells in the primary visual area V1 and 113 cells in the extrastriate visual area V2 while presenting a kinetically defined edge or a luminance contrast edge. Cells meeting statistical criteria for responsiveness and orientation selectivity were classified as selective for the orientation of the kinetic edge if the preferred orientation for a kinetic boundary stimulus remained essentially the same even when the directions of the two motion components defining that boundary were changed by 90 degrees. In area V2, 13 of the 113 cells met all three requirements, whereas in V1, only 4 cells met the criteria of 107 that were tested, and even these demonstrated relatively weak selectivity. Correlation analysis showed that V1 and V2 populations differed greatly (P < 1.0 x 10(-6), Student's t-test) in their selectively for specific orientations of kinetic edge stimuli. Neurons in V2 that were selective for the orientation of a kinetic boundary were further distinguished from their counterparts in V1 in displaying a strong, sharply tuned response to a luminance edge of the same orientation. We concluded that selectivity for the orientation of kinetically defined boundaries first emerges in area V2 rather than in primary visual cortex. An analysis of response onset latencies in V2 revealed that cells selective for the orientation of the motion-defined boundary responded about 40 ms more slowly, on average, to the kinetic edge stimulus than to a luminance edge. In nonselective cells, that is, those presumably responding only to the local motion in the stimulus, this difference was only about 20 ms. Response latencies for the luminance edge were indistinguishable in KE-selective and -nonselective neurons. We infer that while responses to luminance edges or local motion are indigenous to V2, KE-selective responses may involve feedback entering the ventral stream at a point downstream with respect to V2.