Saccade-related and visual activities in the pulvinar nuclei of the behaving rhesus monkey

Summary We studied three subdivisions of the pulvinar: a retinotopically organized inferior area (PI), a retinotopically mapped region of the lateral pulvinar (PL), and a separate, visually responsive component of the lateral pulvinar (Pdm). Single neurons were recorded in these regions from awake, trained rhesus monkeys, and we correlated the discharge patterns of the cells with eye movements. About 60% of the neurons discharged after saccadic eye movements in an illuminated environment and had either excitatory, inhibitory, or biphasic (inhibitory-excitatory) response patterns. These responses were most often transient in nature. Neurons with excitatory activity had a mean onset latency of 72 ms after the termination of the eye movement. Latencies for cells with inhibitory responses averaged 58 ms. In sharp contrast, the cells with biphasic response patterns became active before the termination of the eye movement. A unique set of these neurons termed saccade cells, were active with visually guided eye movements in the light, with the same eye movements made to a briefly pulsed target in the dark, and for similar eye movements made spontaneously in total darkness. The activity was present with the appropriate saccade, independent of the beginning eye position. Biphasic response patterns were typical of these saccade cells. Saccade cells were most common in Pdm and PI. About half of the saccade cells also had some visual response that was independent of eye movement. A second group of cells was active with saccadic eye movements in the light but not in the dark. Some of these cells had clear visual responses that could account for their activity following eye movements; others had no clear visual receptive field. Because of these and other physiological data, we propose that the saccade cells found in Pdm may function in a system dealing with visual spatial attention, while those found in PI may have a role in dealing with the visual consequences of eye movements.     

Visual responses of pulvinar and collicular neurons during eye movements of awake, trained macaques.

1. We recorded from single neurons in awake, trained rhesus monkeys in a lighted environment and compared responses to stimulus movement during periods of fixation with those to motion caused by saccadic or pursuit eye movements. Neurons in the inferior pulvinar (PI), lateral pulvinar (PL), and superior colliculus were tested. 2. Cells in PI and PL respond to stimulus movement over a wide range of speeds. Some of these cells do not respond to comparable stimulus motion, or discharge only weakly, when it is generated by saccadic or pursuit eye movements. Other neurons respond equivalently to both types of motion. Cells in the superficial layers of the superior colliculus have similar properties to those in PI and PL. 3. When tested in the dark to reduce visual stimulation from the background, cells in PI and PL still do not respond to motion generated by eye movements. Some of these cells have a suppression of activity after saccadic eye movements made in total darkness. These data suggest that an extraretinal signal suppresses responses to visual stimuli during eye movements. 4. The suppression of responses to stimuli during eye movements is not an absolute effect. Images brighter than 2.0 log units above background illumination evoke responses from cells in PI and PL. The suppression appears stronger in the superior colliculus than in PI and PL. 5. These experiments demonstrate that many cells in PI and PL have a suppression of their responses to stimuli that cross their receptive fields during eye movements. These cells are probably suppressed by an extraretinal signal. Comparable effects are present in the superficial layers of the superior colliculus. These properties in PI and PL may reflect the function of the ascending tectopulvinar system.     

Effect of attentive fixation in macaque thalamus and cortex

Attentional modulation of neuronal responsiveness is common in many areas of visual cortex. We examined whether attentional modulation in the visual thalamus was quantitatively similar to that in cortex. Identical procedures and apparatus were used to compare attentional modulation of single neurons in seven different areas of the visual system: the lateral geniculate, three visual subdivisions of the pulvinar [inferior, lateral, dorsomedial part of lateral pulvinar (Pdm)], and three areas of extrastriate cortex representing early, intermediate, and late stages of cortical processing (V2, V4/PM, area 7a). A simple fixation task controlled transitions among three attentive states. The animal waited for a fixation point to appear (ready state), fixated the point until it dimmed (fixation state), and then waited idly to begin the next trial (idle state). Attentional modulation was estimated by flashing an identical, irrelevant stimulus in a neuron's receptive field during each of the three states; the three responses defined a "response vector" whose deviation from the line of equal response in all three states (the main diagonal) indicated the character and magnitude of attentional modulation. Attentional modulation was present in all visual areas except the lateral geniculate, indicating that modulation was of central origin. Prevalence of modulation was modest (26%) in pulvinar, and increased from 21% in V2 to 43% in 7a. Modulation had a push-pull character (as many cells facilitated as suppressed) with respect to the fixation state in all areas except Pdm where all cells were suppressed during fixation. The absolute magnitude of attentional modulation, measured by the angle between response vector and main diagonal expressed as a percent of the maximum possible angle, differed among brain areas. Magnitude of modulation was modest in the pulvinar (19-26%), and increased from 22% in V2 to 41% in 7a. However, average trial-to-trial variability of response, measured by the coefficient of variation, also increased across brain areas so that its difference among areas accounted for more than 90% of the difference in modulation magnitude among areas. We also measured attentional modulation by the ratio of cell discharge due to attention divided by discharge variability. The resulting signal-to-noise ratio of attention was small and constant, 1.3 +/- 10%, across all areas of pulvinar and cortex. We conclude that the pulvinar, but not the lateral geniculate, is as strongly affected by attentional state as any area of visual cortex we studied and that attentional modulation amplitude is closely tied to intrinsic variability of response.     

Both striate cortex and superior colliculus contribute to visual properties of neurons in superior temporal polysensory area of macaque monkey

Although the tectofugal system projects to the primate cerebral cortex by way of the pulvinar, previous studies have failed to find any physiological evidence that the superior colliculus influences visual activity in the cortex. We studied the relative contributions of the tectofugal and geniculostriate systems to the visual properties of neurons in the superior temporal polysensory area (STP) by comparing the effects of unilateral removal of striate cortex, the superior colliculus, or of both structures. In the intact monkey, STP neurons have large, bilateral receptive fields. Complete unilateral removal of striate cortex did not eliminate visual responses of STP neurons in the contralateral visual hemifield; rather, nearly half the cells still responded to visual stimuli in the hemifield contralateral to the lesion. Thus the visual properties of STP neurons are not completely dependent on the geniculostriate system. Unilateral striate lesions did affect the response properties of STP neurons in three ways. Whereas most STP neurons in the intact monkey respond similarly to stimuli in the two visual hemifields, responses to stimuli in the hemifield contralateral to the striate lesion were usually weaker than responses in the ipsilateral hemifield. Whereas the responses of many STP neurons in the intact monkey were selective for the direction of stimulus motion or for stimulus form, responses in the hemifield contralateral to the striate lesion were not selective for either motion or form. Whereas the median receptive field in the intact monkey extended 80 degrees into the contralateral visual field, the receptive fields of cells with responses in the contralateral field that survived the striate lesions had a median border that extended only 50 degrees into the contralateral visual field. Removal of both striate cortex and the superior colliculus in the same hemisphere abolished the responses of STP neurons to visual stimuli in the hemifield contralateral to the combined lesion. Nearly 80% of the cells still responded to visual stimuli in the hemifield ipsilateral to the lesion. Unilateral removal of the superior colliculus alone had only small effects on visual responses in STP. Receptive-field size and visual response strength were slightly reduced in the hemifield contralateral to the collicular lesion. As in the intact monkey, selectivity for stimulus motion or form were similar in the two visual hemifields. We conclude that both striate cortex and the superior colliculus contribute to the visual responses of STP neurons. Striate cortex is crucial for the movement and stimulus specificity of neurons in STP.(ABSTRACT TRUNCATED AT 400 WORDS)     

Corticogeniculate neurons, corticotectal neurons, and suspected interneurons in visual cortex of awake rabbits: receptive-field properties, axonal properties, and effects of EEG arousal

The intrinsic stability of the rabbit eye was exploited to enable receptive-field analysis of antidromically identified corticotectal (CT) neurons (n = 101) and corticogeniculate (CG) neurons (n = 124) in visual area I of awake rabbits. Eye position was monitored to within 1/5 degrees. We also studied the receptive-field properties of neurons synaptically activated via electrical stimulation of the dorsal lateral geniculate nucleus (LGNd). Whereas most CT neurons had either complex (59%) or motion/uniform (15%) receptive fields, we also found CT neurons with simple (9%) and concentric (4%) receptive fields. Most complex CT cells were broadly tuned to both stimulus orientation and velocity, but only 41% of these cells were directionally selective. We could elicit no visual responses from 6% of CT cells, and these cells had significantly lower conduction velocities than visually responsive CT cells. The median spontaneous firing rates for all classes of CT neurons were 4-8 spikes/s. CG neurons had primarily simple (60%) and concentric (9%) receptive fields, and none of these cells had complex receptive fields. CG simple cells were more narrowly tuned to both stimulus orientation and velocity than were complex CT cells, and most (85%) were directionally selective. Axonal conduction velocities of CG neurons (mean = 1.2 m/s) were much lower than those of CT neurons (mean = 6.4 m/s), and CG neurons that were visually unresponsive (23%) had lower axonal conduction velocities than did visually responsive CG neurons. Some visually unresponsive CG neurons (14%) responded with saccadic eye movements. The median spontaneous firing rates for all classes of CG neurons were less than 1 spike/s. All neurons synaptically activated via LGNd stimulation at latencies of less than 2.0 ms had receptive fields that were not orientation selective (89% motion/uniform, 11% concentric), whereas most cells with orientation-selective receptive fields had considerably longer synaptic latencies. Most short-latency motion/uniform neurons responded to electrical stimulation of the LGNd (and visual area II) with a high-frequency burst (500-900 Hz) of three or more spikes. Action potentials of these neurons were of short duration, thresholds of synaptic activation were low, and spontaneous firing rates were the highest seen in rabbit visual cortex. These properties are similar to those reported for interneurons in several regions in mammalian central nervous system. Nonvisual sensory stimuli that resulted in electroencephalographic arousal (hippocampal theta activity) had a profound effect on the visual responses of many visual cortical neurons.(ABSTRACT TRUNCATED AT 400 WORDS)     

Response characteristics of neurons in the pulvinar of awake cats to saccades and to visual stimulation

We studied responses of pulvinar neurons in awake cats that were allowed to execute spontaneous eye movements. Extracellular cell activity during saccades, saccade-like image shifts, and various stationary visual stimuli was recorded together with the animals' eye positions. All neurons analyzed had receptive fields that covered most of the central 80x80 degrees of the animals' visual field and did only respond to large (>20 degrees) visual stimuli. According to their response properties, recorded neurons were divided into three populations. The first group, termed "S neurons" (16%), responded when the animals performed saccades but were unresponsive to any of the visual stimuli tested. These neurons do not seem to receive a visual input that is strong enough to drive them. The second group, termed "V neurons" (51%), responded to various visual stimuli including saccade-like image motion when the eyes were stationary, but not when the animals executed saccades. V neurons therefore distinguish retinal image movements that are generated externally from internally generated image motion. Finally, "SV neurons" (31%) responded when the animals made saccades as well as to saccade-like image motion or to stationary stimuli. Although these neurons do not distinguish self-induced retinal image motion from motion generated by external stimulus movements, they must receive non-retinal motion-related input, because responses elicited by saccades had shorter latencies than responses to saccade-like stimulus movements. Only SV neurons resemble response properties of pretectal neurons that project to the pulvinar and that comprise the major subcortical visual input. The functional significance of pulvinar neuronal populations for visual and visuomotor information processing is discussed.     

Gating and control of primary visual cortex by pulvinar

The primary visual cortex (V1) receives its driving input from the eyes via the lateral geniculate nucleus (LGN) of the thalamus. The lateral pulvinar nucleus of the thalamus also projects to V1, but this input is not well understood. We manipulated lateral pulvinar neural activity in prosimian primates and assessed the effect on supra-granular layers of V1 that project to higher visual cortex. Reversibly inactivating lateral pulvinar prevented supra-granular V1 neurons from responding to visual stimulation. Reversible, focal excitation of lateral pulvinar receptive fields increased the visual responses in coincident V1 receptive fields fourfold and shifted partially overlapping V1 receptive fields toward the center of excitation. V1 responses to regions surrounding the excited lateral pulvinar receptive fields were suppressed. LGN responses were unaffected by these lateral pulvinar manipulations. Excitation of lateral pulvinar after LGN lesion activated supra-granular layer V1 neurons. Thus, lateral pulvinar is able to powerfully control and gate information outflow from V1.     

Receptive fields and response properties of neurons in layer 4 of ferret visual cortex

The ferret has become a model animal for studies exploring the development of the visual system. However, little is known about the receptive-field structure and response properties of neurons in the adult visual cortex of the ferret. We performed single-unit recordings from neurons in layer 4 of adult ferret primary visual cortex to determine the receptive-field structure and visual-response properties of individual neurons. In particular, we asked what is the spatiotemporal structure of receptive fields of layer 4 neurons and what is the orientation selectivity of layer 4 neurons? Receptive fields of layer 4 neurons were mapped using a white-noise stimulus; orientation selectivity was determined using drifting, sine-wave gratings. Our results show that most neurons (84%) within layer 4 are simple cells with elongated, spatially segregated, ON and OFF subregions. These neurons are also selective for stimulus orientation; peaks in orientation-tuning curves have, on average, a half-width at half-maximum response of 21.5 +/- 1.2 degrees (mean +/- SD). The remaining neurons in layer 4 (16%) lack orientation selectivity and have center/surround receptive fields. Although the organization of geniculate inputs to layer 4 differs substantially between ferret and cat, our results demonstrate that, like in the cat, most neurons in ferret layer 4 are orientation-selective simple cells.     

Orbital position and eye movement influences on visual responses in the pulvinar nuclei of the behaving macaque

Summary We studied the influences of eye movements on the visual responses of neurons in two retinotopically organized areas of the pulvinar of the macaque. Cells were recorded from awake, trained monkeys, and visual responses were characterized immediately before and after the animals made saccadic eye movements. A significant proportion of the cells were more responsive to stimuli around the time of eye movements than they were at other intervals. Other cells had response reduction. For some neurons, the change in excitability was associated with orbital position and not the eye movement. For other cells the change was present with eye movements of similar amplitude and direction but with different starting and ending positions. Here it appears that the eye movement is the important parameter. Other cells had effects related to both eye position and eye movements. In all cells tested, the changes in excitability were present when the experiments were conducted in the dark as well as in the light. This suggests that the mechanism of the effect is related to the eye position or eye movement and not visual-visual interactions. For about half of the neurons with modulations, the response showed facilitation for stimuli presented in the most responsive region of the receptive field but not for those at the edge of the field. For the other cells there was facilitation throughout the field. Thus, a gradient of modulation in the receptive field may vary among cells. These experiments demonstrate modulations of visual responses in the pulvinar by eye movements. Such effects may be part of the visual-behavioral improvements at the end of eye movements and/or contribute to spatial constancy.     

Functional properties of monkey caudate neurons. II. Visual and auditory responses

1. Visual responses of caudate neurons were studied in monkeys trained to fixate on a small spot of light. A visual stimulus (another spot of light) was presented in various contexts of behavior using different behavioral paradigms. Visual receptive fields were usually large and centered on the contralateral hemifield. Among 217 neurons with visual responses, 184 were further classified into subtypes. 2. Visual responses in 64 neurons were not modulated by changing the paradigms (unconditional visual responses). In the other neurons, visual responses were dependent on the behavioral contexts in which the stimulus was presented. Three types of behavioral modulation were found. 3. A saccade-enhanced visual response (n = 37) was the one that was enhanced if the monkey made a saccade to the stimulus on its appearance. The enhancement was spatially selective: the response was depressed if the saccade was directed away from the stimulus. 4. Memory-contingent visual responses (n = 36) were present preferentially when the monkey remembered the location of the stimulus and a few seconds later made a saccade to the remembered location. Responses were greater when the location of the stimulus was randomized between trials. 5. Expectation-contingent visual responses (n = 46) were present preferentially when the stimulus came on while the monkey was not fixating another spot, and the stimulus was related directly to a reward. Unlike the other types, its receptive field included both contralateral and ipsilateral hemifields without a particular preference. 6. A small number of neurons (n = 16) showed a visual response that easily habituated. 7. Latencies of visual responses were usually between 100 and 200 ms. The latencies of the memory-contingent, expectation-contingent, and habituated visual responses tended to be longer than the others and tended to be more variable between trials. 8. Among auditory responsive neurons only a small proportion were related to the tasks. The response was greater to a contralateral sound. It was enhanced if the monkey used the sound as the cue for the future target location. 9. The results suggest that sensory responses of caudate neurons could be used to guide a subsequent sequence of learned behaviors by confirming predicted environmental states, renewing memory, or establishing a motor set.     

Characterization of pretectal-nuclear-complex afferents to the pulvinar in the cat

We investigated anatomical and physiological properties of the projection from the pretectal nuclear complex (PNC) to the ipsilateral lateral posterior–pulvinar complex in the cat. After Phaseolus vulgaris leucoagglutinin injections into the PNC, the majority (70%) of anterogradely labeled terminals was localized in the pulvinar proper, the remaining 30% were scattered in the lateral and medial portions of the LP. No PNC neuron retrogradely labeled from the pulvinar was found to also express glutamic acid decarboxylase (GAD) mRNA, although a large number of neurons carrying the GAD label were found in close vicinity. In contrast, 69% of retrogradely labeled PNC cells also displayed glutamate-like immunoreactivity. Twenty-six out of 96 (27%) visually responsive pulvinar neurons were orthodromically activated by electrical stimulation of the ipsilateral PNC at latencies between 1 and 10 ms (median 1.9 ms). All orthodromically activated neurons responded well to the onset and offset of large visual stimuli and to sudden stimulus shifts. Whenever a saccadic eye movement was executed, these neurons were also activated, except during saccades in darkness. The comparison of saccade-evoked response with responses to visual stimuli that elicit similar retinal image shifts revealed that pretectorecipient pulvinar neurons also seem to receive a saccade-related non-visual input. All response properties correspond to those of a specific class of pulvinar neurons that have been termed "SV" neurons because they respond to visual stimulation as well as during saccades. They also closely resemble response properties of PNC neurons that project to the ipsilateral pulvinar. The results support the proposal that PNC cells not only directly activate their postsynaptic target neurons in the pulvinar, but that they also provide a visual input to these neurons that greatly contributes to their response characteristics. Electronic Publication     

Structure of receptive fields of cat pulvinar neurons sensitive to photic stimulation

Receptive fields of 262 pulvinar neurons were studied. Receptive fields of 142 of these neurons were studied in detail with the aid of a stationary spot of light, flashing in different parts of the receptive field. Depending on responses to presentation of the stationary stimulus the neurons were divided into six groups. The first group included neurons with on-off responses to photic stimulation (44 of 142), the second group neurons with off responses only (42 of 142). In cells of the third group (19 of 142) an on response only was recorded in all structures of the receptive field tested. Neurons of the fourth group (eight of 142) had a receptive field of similar structure to that of the simple receptive fields of neurons in cortical area 17. The fifth group (10 of 142) included neurons with a receptive field of concentric structure, the sixth (19 of 142) consisted of neurons with receptive fields with multiple discharge centers. The structure of the receptive field of these neurons was mosaic, with an irregular distribution of exciting and silent zones. The mean response latency of the pulvinar neurons was 40–70 msec. Responses of neurons with shorter (20 msec) and longer (130–160 msec) latent periods also were recorded.Translated from Neirofiziologiya, Vol. 11, No. 1, pp. 3–10, January–February, 1979.     

Visual response characteristics of neurons in the nucleus isthmi magnocellularis and nucleus isthmi parvocellularis of pigeons

Summary The visual response characteristics of single cells in the nucleus isthmi (NI) of pigeons were investigated using standard extracellular recording techniques. The results show that both major components of NI, the parvocellular NI (Ipc) and the magnocellular NI (Imc), have a tight retinotopic organization where nasal regions of the visual field are mapped onto the rostral poles of Ipc and Imc, and temporal regions of the visual field are mapped onto the caudal poles. The more ventral regions of these nuclei receive input from more inferior regions of visual space. The receptive fields (RFs) of both Ipc and Imc are large and oval-shaped, and their long axis is oriented vertically in the visual field. Most RFs are distributed on the contralateral visual horizon, and no binocular responses were found in either Ipc or Imc. All of the excitatory RFs of NI cells were surrounded by large inhibitory regions which participated in the dramatic modulation of the driven visual response when a large background pattern was moved across this zone. Although both Ipc and Imc neurons are driven best by small dark spots, some of them also show orientation selectivity to bars which may result from their ovalshaped RF (74% of Imc cells, 20/38, were orientation selective as compared to 10% of the Ipc cells, 3/30). It is suggested that some tectal cells with small RFs, and which originate from a vertically oriented zone may converge onto a single NI neuron to produce the elliptical shaped receptive fields.     

Receptive field center size decreases and firing properties mature in ON and OFF retinal ganglion cells after eye opening in the mouse

Development of the mammalian visual system is not complete at birth but continues postnatally well after eye opening. Although numerous studies have revealed changes in the development of the thalamus and visual cortex during this time, less is known about the development of response properties of retinal ganglion cells (RGCs). Here, we mapped functional receptive fields of mouse RGCs using a Gaussian white noise checkerboard stimulus and a multielectrode array to record from retinas at eye opening, 3 days later, and 4 wk after birth, when visual responses are essentially mature. Over this time, the receptive field center size of ON and OFF RGC populations decreased. The average receptive field center size of ON RGCs was larger than that of OFF RGCs at eye opening, but they decreased to the same size in the adult. Firing properties were also immature at eye opening. RGCs had longer latencies, lower frequencies of firing, and lower sensitivity than in the adult. Hence, the dramatic maturation of the visual system during the first weeks of visual experience includes the retina.     

Connections of calbindin-D28k-defined subdivisions in inferior pulvinar with visual areas V2, V4 and MT in macaque monkeys

We examined the relationship between calbindin-D28k-defined subdivisions in the macaque inferior pulvinar and the patterns of afferent connections to cortical visual areas MT, V4 and V2. The subdivisions identified included the posterior (PIp), medial (PIm), central (PIc) and lateral (PI1) subdivisions. Projections to MT and V4 were largely segregated in the inferior pulvinar: projections to MT originated mainly from PIm, while projections to V4 originated mainly from PI1. In addition, weaker projections to MT were observed from PIp and PIc, and some projections from PIp to V4 were observed in one of two cases. Projections to V2 originated preferentially from PI1, with a lesser projection from PIc. No labeled cells were observed in PIm in five monkeys injected with various tracers into different regions of V2. Since most V2 injections were large enough to involve several neighboring stripe-like compartments, these findings suggest that PIm does not project to V2 compartments associated with neither dorsal nor ventral cortical processing streams. Cells projecting to V4 were not strictly segregated from those projecting to V2 in neither PI1 nor PIc, suggesting that inferior pulvinar projections do not map the position of visual areas in the cortical mantle.     

Visual responses with and without fixation: neurons in premotor cortex encode spatial locations independently of eye position

 The ventral premotor cortex (PMv) of the macaque monkey contains neurons that respond both to visual and to tactile stimuli. For almost all of these “bimodal” cells, the visual receptive field is anchored to the tactile receptive field on the head or the arms, and remains stationary when the eyes fixate different locations. This study compared the responses of bimodal PMv neurons to a visual stimulus when the monkey was required to fixate a spot of light and when no fixation was required. Even when the monkey was not fixating and the eyes were moving, the visual receptive fields remained in the same location, near the associated tactile receptive field. For many of the neurons, the response to the visual stimulus was significantly larger when the monkey was not performing the fixation task. In control tests, the presence or absence of the fixation spot itself had little or no effect on the response to the visual stimulus. These results show that even when the monkey’s eye position is continuously changing, the neurons in PMv have visual receptive fields that are stable and fixed to the relevant body part. The reduction in response during fixation may reflect a shift of attention from the visual stimulus to the demands of the fixation task. Received: 8 April 1997 / Accepted: 16 July 1997     

Reference frames for representing visual and tactile locations in parietal cortex

The ventral intraparietal area (VIP) receives converging inputs from visual, somatosensory, auditory and vestibular systems that use diverse reference frames to encode sensory information. A key issue is how VIP combines those inputs together. We mapped the visual and tactile receptive fields of multimodal VIP neurons in macaque monkeys trained to gaze at three different stationary targets. Tactile receptive fields were found to be encoded into a single somatotopic, or head-centered, reference frame, whereas visual receptive fields were widely distributed between eye- to head-centered coordinates. These findings are inconsistent with a remapping of all sensory modalities in a common frame of reference. Instead, they support an alternative model of multisensory integration based on multidirectional sensory predictions (such as predicting the location of a visual stimulus given where it is felt on the skin and vice versa). This approach can also explain related findings in other multimodal areas.     

Visual responses of thalamic neurons depending on the direction of gaze and the position of targets in space

Summary Visual receptive field properties of neurons in the region of the thalamic internal medullary lamina were studied in alert cats while they fixated in various directions. In slightly more than 50% of the cells, the responsiveness of the cells was found to depend on the location of the stimulus with respect to the head-body axis (stimulus absolute position). A cell could ignore a stimulus outside its absolute field even if it was well placed within its receptive field.Three types of neurons were distinguished. Neurons with small central receptive fields were tonically activated when the animal fixated the stimulus in one half of the screen (usually contralateral). The firing rate of these cells was related to the stimulus absolute position measured along a preferred axis. Similarly, neurons with large receptive fields fired as a function of stimulus absolute position but stimulus fixation was not required. Neurons with eccentric fields responded to stimuli located in a target area defined in head-body coordinates. Such cells gave presaccadic bursts with eye movements terminating in the target area.The conclusion proposed is that neurons exist which code visual spatial information in a non-retinal frame of reference. This coding takes place at the time of stimulus presentation. Its role may be seen in the initiation of visually guided movements.     

Contralateral visual hemifield representations in the human pulvinar nucleus

The pulvinar is a major nucleus of the thalamus. Macaque pulvinar includes two subregions that are connected to the visual cortex and are retinotopically organized, but the organizing principles of the visual portions of the human pulvinar are unknown. We employed two tasks to address the question of whether human pulvinar exhibits spatial organization using event-related functional magnetic resonance imaging. The first was a global-motion discrimination with a rich visual stimulus and the second a luminance-discrimination task of similar difficulty that used a minimal visual stimulus. Both tasks required central fixation and covert peripheral attention. A group analysis of blood-oxygen-level-dependent responses elicited in the motion-discrimination task revealed activity bilaterally in the ventral pulvinar (z = 2 in Talairach space). Clear position specificity was observed with activity elicited only by contralateral stimuli. Ipsilateral stimuli caused suppression. This locus of activity is distinct from the more dorsal (z = 10) region of the pulvinar that has previously been reported to be visually responsive but not retinotopic. In the luminance-discrimination task, similar activity was seen, but it was weaker and detectable only in the left pulvinar. In additional experiments with no task, passively viewed global-motion stimuli also activated the ventral pulvinar bilaterally. Our results show for the first time a distinct, bilateral visual representation in human inferior pulvinar that appears to be contralaterally organized.     

Sensitivity of MST neurons to optic flow stimuli. II. Mechanisms of response selectivity revealed by small-field stimuli

1. In these experiments we examined the receptive field mechanisms that support the optic flow field selective responses of neurons in the dorsomedial region of the medial superior temporal area (MSTd). Our experiments tested the predictions of two hypotheses of optic flow field selectivity. The direction mosaic hypothesis states that these receptive fields contain a set of planar direction-selective subfields that match the local directions of motion within optic flow fields. The vector field hypothesis states that these receptive fields are uniquely sensitive to distributed properties of planar, circular, or radial optic flow fields. 2. Experiments using large-field stimuli revealed that some neurons showed changes in optic flow field selectivity depending on the position of the stimulus in the receptive field; these are position-dependent responses. However, other neurons maintained the same optic flow field selectivities in spite of changes in stimulus position; these are position-invariant responses. We have used the position dependence or invariance of optic flow field selectivity as a way of testing the direction mosaic and vector field hypotheses. Position dependence is more consistent with the direction mosaic hypothesis, whereas position invariance is more consistent with the vector field hypothesis. 3. To test for position effects, we examined the optic flow field selectivity of small subfields within the large receptive fields of 160 MSTd neurons. First, we centered small-field optic flow stimuli of various sizes over the same position in the receptive field. Most MSTd neurons showed decreasing response amplitude with decreasing stimulus size but maintained optic flow field selectivity. 4. We then placed small-field stimuli at various positions within the large receptive field of these MSTd neurons. Position-invariant response selectivity was most prominent in single-component neurons, suggesting that they were more consistent with the vector field hypothesis. Position-dependent response selectivity was most prominent in triple-component neurons, suggesting that they were more consistent with the direction mosaic hypothesis. However, the variations in planar direction preference throughout the receptive field of these triple-component neurons were not consistent with a direction mosaic explanation of the large-field circular or radial selectivity observed. 5. Small-field position studies also demonstrated the existence of zones within the receptive field in which either direction-selective inhibitory or direction-selective excitatory responses predominated. The degree of overlap between these zones increased from nonselective to triple- to double- and finally to single-component neurons.(ABSTRACT TRUNCATED AT 400 WORDS)