Responses to visual stimulation and relationship between visual, auditory, and somatosensory inputs in mouse superior colliculus

1975 ◽  
Vol 38 (3) ◽  
pp. 690-713 ◽  
Author(s):  
U. C. Drager ◽  
D. H. Hubel
Keyword(s):  
Visual Field ◽  
Receptive Field ◽  
Superior Colliculus ◽  
Single Cells ◽  
Receptive Fields ◽  
Auditory Stimuli ◽  
Optimal Size ◽  
Visual Cell ◽  
Temporal Part ◽  
Cell Type

The superior colliculus was studied in anesthetized mice by recording from single cells and from unit clusters. The topographic representation of the visual filed was similar to what has been found in other mammals, with the temporal part of the contralateral visual field projecting posteriorly and the inferior visual field projecting laterally. At the anterior margin of the tectum receptive fields recorded through the contralateral eye and invaded the ipsilateral visual hemifield for up to 35 degrees, suggesting that the entire visual field through one eye is represented on the contralateral superior colliculus. Cells located closest to the tectal surface had relatively small receptive fields, averaging 9 degrees in center diameter; field sizes increased steadily with depth. The prevailing cell type in the stratum zonal and superficial gray responded best to a small dark or light object of any shape moved slowly through the receptive-field center or to turning a small stationary spot on or off. Large objects or diffuse light were usually much less effective. Less than one-quarter of superficial layer cells showed directional selectivity to a moving object, the majority of these favoring up and nasal movement. The chief visual cell type in the stratum opticum and upper part of the intermediate gray resembled in the newness neurons described for many other vertebrates: they had large receptive fields and responded best to up and nasal movement of a small dark or light object, whose optimal size was similar to the optimum for upper-layer cells. If the same part of the receptive field was repeatedly stimulated there was a marked tendency to habituate. Only very few cels responded to the ipsilateral eye. Intermixed with visual cells in the upper part of the intermediate gray were cells that responded to somatosensory or auditory stimuli. Here bimodal and trimodal cells were also seen. In deeper layers somatosensory and auditory modalities tended to take over. These two modalities were not segregated into sublayers but rather seemed to be arranged in clusters. Responses to somatosensory and auditory stimuli were brisk, showing little habituation to repeated stimulation.

Ferrier lecture - Functional architecture of macaque monkey visual cortex

1977 ◽  
Vol 198 (1130) ◽  
pp. 1-59 ◽  
Keyword(s):  
Visual Cortex ◽  
Visual Field ◽  
Receptive Field ◽  
Striate Cortex ◽  
Single Cells ◽  
Ocular Dominance ◽  
Receptive Fields ◽  
Macaque Monkey ◽  
Area 17 ◽  
Orientation Specificity

Of the many possible functions of the macaque monkey primary visual cortex (striate cortex, area 17) two are now fairly well understood. First, the incoming information from the lateral geniculate bodies is rearranged so that most cells in the striate cortex respond to specifically oriented line segments, and, second, information originating from the two eyes converges upon single cells. The rearrangement and convergence do not take place immediately, however: in layer IVc, where the bulk of the afferents terminate, virtually all cells have fields with circular symmetry and are strictly monocular, driven from the left eye or from the right, but not both; at subsequent stages, in layers above and below IVc, most cells show orientation specificity, and about half are binocular. In a binocular cell the receptive fields in the two eyes are on corresponding regions in the two retinas and are identical in structure, but one eye is usually more effective than the other in influencing the cell; all shades of ocular dominance are seen. These two functions are strongly reflected in the architecture of the cortex, in that cells with common physiological properties are grouped together in vertically organized systems of columns. In an ocular dominance column all cells respond preferentially to the same eye. By four independent anatomical methods it has been shown that these columns have the form of vertically disposed alternating left-eye and right-eye slabs, which in horizontal section form alternating stripes about 400 μm thick, with occasional bifurcations and blind endings. Cells of like orientation specificity are known from physiological recordings to be similarly grouped in much narrower vertical sheeet-like aggregations, stacked in orderly sequences so that on traversing the cortex tangentially one normally encounters a succession of small shifts in orientation, clockwise or counterclockwise; a 1 mm traverse is usually accompanied by one or several full rotations through 180°, broken at times by reversals in direction of rotation and occasionally by large abrupt shifts. A full complement of columns, of either type, left-plus-right eye or a complete 180° sequence, is termed a hypercolumn. Columns (and hence hypercolumns) have roughly the same width throughout the binocular part of the cortex. The two independent systems of hypercolumns are engrafted upon the well known topographic representation of the visual field. The receptive fields mapped in a vertical penetration through cortex show a scatter in position roughly equal to the average size of the fields themselves, and the area thus covered, the aggregate receptive field, increases with distance from the fovea. A parallel increase is seen in reciprocal magnification (the number of degrees of visual field corresponding to 1 mm of cortex). Over most or all of the striate cortex a movement of 1-2 mm, traversing several hypercolumns, is accompanied by a movement through the visual field about equal in size to the local aggregate receptive field. Thus any 1-2 mm block of cortex contains roughly the machinery needed to subserve an aggregate receptive field. In the cortex the fall-off in detail with which the visual field is analysed, as one moves out from the foveal area, is accompanied not by a reduction in thickness of layers, as is found in the retina, but by a reduction in the area of cortex (and hence the number of columnar units) devoted to a given amount of visual field: unlike the retina, the striate cortex is virtually uniform morphologically but varies in magnification. In most respects the above description fits the newborn monkey just as well as the adult, suggesting that area 17 is largely genetically programmed. The ocular dominance columns, however, are not fully developed at birth, since the geniculate terminals belonging to one eye occupy layer IVc throughout its length, segregating out into separate columns only after about the first 6 weeks, whether or not the animal has visual experience. If one eye is sutured closed during this early period the columns belonging to that eye become shrunken and their companions correspondingly expanded. This would seem to be at least in part the result of interference with normal maturation, though sprouting and retraction of axon terminals are not excluded.

Indirect, across-the-midline retinotectal projections and representation of ipsilateral visual field in superior colliculus of the cat

1978 ◽  
Vol 41 (2) ◽  
pp. 285-304 ◽  
Author(s):  
A. Antonini ◽  
G. Berlucchi ◽  
J. M. Sprague
Keyword(s):  
Visual Field ◽  
Receptive Field ◽  
Superior Colliculus ◽  
Visual Information ◽  
Receptive Fields ◽  
Optic Chiasm ◽  
Anterior Portion ◽  
Vertical Meridian ◽  
Contralateral Eye ◽  
Rostral Portion

1. In agreement with previous work, we have found that the ipsilateral visual field is represented in an extensive rostral portion--from one-third to one-half--of the superior colliculus (SC) of the cat. This representation is binocular. The SC representation of the ipsilateral visual field can be mediated both directly, by crossed retinotectal connections originating from temporal hemiretina, and indirectly, by across-the-midline connections relaying visual information from one-half of the brain to contralateral SC. 2. In order to study the indirect, across-the-midline visual input to the SC, we have recorded responses of SC neurons to visual stimuli presented to either the ipsilateral or the contralateral eye of cats with a midsagittal splitting of the optic chiasm. Units driven by the ipsilateral eye, presumably through the direct retinotectal input and/or corticotectal connections from ipsilateral visual cortex, were found throughout the SC, except at its caudal pole, which normally receives fibers from the extreme periphery of the contralateral nasal hemiretina. Units driven by the contralateral eye, undoubtedly through an indirect across-the-midline connection, were found only in the anterior portion of the SC, in which is normally represented the ipsilateral visual field. Receptive fields in both ipsilateral and contralateral eye had properties typical of SC receptive fields in cats with intact optic pathways. 3. All units having a receptive field in the contralateral eye had also a receptive field in the ipsilateral eye; for each of these units, the receptive fields in both eyes invariably abutted the vertical meridian of the visual field. The receptive field in one eye had about the same elevation relative to the horizontal meridian and the same vertical extension as the receptive field in the other eye; the two receptive fields of each binocular unit matched each other at the vertical meridian and formed a combined receptive field straddling the vertical midline of the horopter...

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

1986 ◽  
Vol 55 (5) ◽  
pp. 1057-1075 ◽  
Author(s):  
C. J. Bruce ◽  
R. Desimone ◽  
C. G. Gross
Keyword(s):  
Visual Field ◽  
Receptive Field ◽  
Superior Colliculus ◽  
Striate Cortex ◽  
Visual Stimuli ◽  
Receptive Fields ◽  
Visual Responses ◽  
Stimulus Motion ◽  
Visual Hemifields ◽  
Visual Properties

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)

Receptive-field properties in reptilian optic tectum: some comparisons with mammals

1983 ◽  
Vol 50 (1) ◽  
pp. 102-124 ◽  
Author(s):  
B. E. Stein ◽  
N. S. Gaither
Keyword(s):  
Receptive Field ◽  
Superior Colliculus ◽  
Optic Tectum ◽  
Single Cells ◽  
Receptive Fields ◽  
Spatial Summation ◽  
Similar Distribution ◽  
Moving Stimuli ◽  
Receptive Field Properties ◽  
Sensory Processes

The receptive-field properties of single cells in the optic tectum of Iguana iguana were studied using the same procedures as have been used in this laboratory in studying its mammalian homologue, the superior colliculus. Surprisingly, despite some species-specific characteristics, the range of physiological properties of tectal and superior collicular cells appeared to be strikingly similar. This observation is not consistent with the notion that functional differences between these structures evolved as a consequence of the tremendous elaboration of mammalian neocortex and its involvement in sensory processes. The internal organization of visual tectal receptive fields was observed to be very much like that described in mammals. This included a similar distribution of on-off areas, the presence of both spatial summation and spatial inhibition within the excitatory receptive-field borders, suppressive areas just beyond these borders, and a marked tendency for habituation. In addition, many cells showed distinct directional preferences that were strongly influenced by the velocity of movement through the receptive field. Furthermore, the receptive fields of bimodal and trimodal cells showed topographic correspondences as in mammals, although the sizes of the fields were often large, thereby emphasizing the lack of an exact register between modalities. In contrast to the findings in mammals, however, a preference for stationary over moving stimuli was observed in most neurons, and specializations in iguana tectal cells representing the fovea were noted that have not been described in other species. These foveal specializations include a distinct preference for stationary over moving stimuli, the absence of directional selectivity, and the presence of a majority of cells responding at light onset only. It is suggested that the similarities in the organization and response properties of cells of the optic tectum and superior colliculus reflect the retention of ancestral characteristics through various levels of vertebrate evolution. Furthermore, the subtle species differences in the properties of these cells appear to reflect adaptations to specific ecological pressures rather than general evolutionary trends.

scholarly journals Visual performance in behaving cats after prenatal unilateral enucleation

1993 ◽  
Vol 90 (23) ◽  
pp. 11142-11146 ◽  
Author(s):  
S Bisti ◽  
C Trimarchi
Keyword(s):  
Visual Acuity ◽  
Visual Field ◽  
Receptive Field ◽  
Receptive Fields ◽  
Direction Selectivity ◽  
Visual Performance ◽  
Field Size ◽  
Electrophysiological Recordings ◽  
Severe Impairment ◽  
Orientation Columns

Prenatal unilateral enucleation in mammals causes an extensive anatomical reorganization of visual pathways. The remaining eye innervates the entire extent of visual subcortical and cortical areas. Electrophysiological recordings have shown that the retino-geniculate connections are retinotopically organized and geniculate neurones have normal receptive field properties. In area 17 all neurons respond to stimulation of the remaining eye and retinotopy, orientation columns, and direction selectivity are maintained. The only detectable change is a reduction in receptive field size. Are these changes reflected in the visual behavior? We studied visual performance in cats unilaterally enucleated 3 weeks before birth (gestational age at enucleation, 39-42 days). We tested behaviorally the development of visual acuity and, in the adult, the extension of the visual field and the contrast sensitivity. We found no difference between prenatal monocularly enucleated cats and controls in their ability to orient to targets in different positions of the visual field or in their visual acuity (at any age). The major difference between enucleated and control animals was in contrast sensitivity:prenatal enucleated cats present a loss in sensitivity for gratings of low spatial frequency (below 0.5 cycle per degree) as well as a slight increase in sensitivity at middle frequencies. We conclude that prenatal unilateral enucleation causes a selective change in the spatial performance of the remaining eye. We suggest that this change is the result of a reduction in the number of neurones with large receptive fields, possibly due to a severe impairment of the Y system.

Context dependence of receptive field remapping in superior colliculus

2011 ◽  
Vol 106 (4) ◽  
pp. 1862-1874 ◽  
Author(s):  
Jan Churan ◽  
Daniel Guitton ◽  
Christopher C. Pack
Keyword(s):  
Receptive Field ◽  
Superior Colliculus ◽  
Visual Stimulation ◽  
Receptive Fields ◽  
Spatial Location ◽  
Macaque Monkey ◽  
Visual Signals ◽  
Perceptual Stability ◽  
Visual Background ◽  
The Brain

Our perception of the positions of objects in our surroundings is surprisingly unaffected by movements of the eyes, head, and body. This suggests that the brain has a mechanism for maintaining perceptual stability, based either on the spatial relationships among visible objects or internal copies of its own motor commands. Strong evidence for the latter mechanism comes from the remapping of visual receptive fields that occurs around the time of a saccade. Remapping occurs when a single neuron responds to visual stimuli placed presaccadically in the spatial location that will be occupied by its receptive field after the completion of a saccade. Although evidence for remapping has been found in many brain areas, relatively little is known about how it interacts with sensory context. This interaction is important for understanding perceptual stability more generally, as the brain may rely on extraretinal signals or visual signals to different degrees in different contexts. Here, we have studied the interaction between visual stimulation and remapping by recording from single neurons in the superior colliculus of the macaque monkey, using several different visual stimulus conditions. We find that remapping responses are highly sensitive to low-level visual signals, with the overall luminance of the visual background exerting a particularly powerful influence. Specifically, although remapping was fairly common in complete darkness, such responses were usually decreased or abolished in the presence of modest background illumination. Thus the brain might make use of a strategy that emphasizes visual landmarks over extraretinal signals whenever the former are available.

Neuronal Activity in Somatosensory Cortex of Monkeys Using a Precision Grip. I. Receptive Fields and Discharge Patterns

1999 ◽  
Vol 81 (2) ◽  
pp. 825-834 ◽  
Author(s):  
Iran Salimi ◽  
Thomas Brochier ◽  
Allan M. Smith
Keyword(s):  
Receptive Field ◽  
Somatosensory Cortex ◽  
Neuronal Activity ◽  
Single Cells ◽  
Precision Grip ◽  
Receptive Fields ◽  
Pressure Change ◽  
Discharge Pattern ◽  
Cortical Cells ◽  
Discharge Patterns

Neuronal activity in somatosensory cortex of monkeys using a precision grip. I. Receptive fields and discharge patterns. Three adolescent Macaca fascicularis monkeys weighing between 3.5 and 4 kg were trained to use a precision grip to grasp a metal tab mounted on a low friction vertical track and to lift and hold it in a 12- to 25-mm position window for 1 s. The surface texture of the metal tab in contact with the fingers and the weight of the object could be varied. The activity of 386 single cells with cutaneous receptive fields contacting the metal tab were recorded in Brodmann’s areas 3b, 1, 2, 5, and 7 of the somatosensory cortex. In this first of a series of papers, we describe three types of discharge pattern, the receptive-field properties, and the anatomic distribution of the neurons. The majority of the receptive fields were cutaneous and covered less than one digit, and a χ2 test did not reveal any significant differences in the Brodmann’s areas representing the thumb and index finger. Two broad categories of discharge pattern cells were identified. The first category, dynamic cells, showed a brief increase in activity beginning near grip onset, which quickly subsided despite continued pressure applied to the receptive field. Some of the dynamic neurons responded to both skin indentation and release. The second category, static cells, had higher activity during the stationary holding phase of the task. These static neurons demonstrated varying degrees of sensitivity to rates of pressure change on the skin. The percentage of dynamic versus static cells was about equal for areas 3b, 2, 5, and 7. Only area 1 had a higher proportion of dynamic cells (76%). A third category was identified that contained cells with significant pregrip activity and included cortical cells with both dynamic or static discharge patterns. Cells in this category showed activity increases before movement in the absence of receptive-field stimulation, suggesting that, in addition to peripheral cutaneous input, these cells also receive strong excitation from movement-related regions of the brain.

Spatial determinants of multisensory integration in cat superior colliculus neurons

1996 ◽  
Vol 75 (5) ◽  
pp. 1843-1857 ◽  
Author(s):  
M. A. Meredith ◽  
B. E. Stein
Keyword(s):  
Receptive Field ◽  
Superior Colliculus ◽  
Neuronal Response ◽  
Receptive Fields ◽  
Spatial Relationship ◽  
Physiological Role ◽  
Extracellular Recording ◽  
Average Proportion ◽  
Relationship Of ◽  
High Degree

1. Although a representation of multisensory space is contained in the superior colliculus, little is known about the spatial requirements of multisensory stimuli that influence the activity of neurons here. Critical to this problem is an assessment of the registry of the different receptive fields within individual multisensory neurons. The present study was initiated to determine how closely the receptive fields of individual multisensory neurons are aligned, the physiological role of that alignment, and the possible functional consequences of inducing receptive-field misalignment. 2. Individual multisensory neurons in the superior colliculus of anesthetized, paralyzed cats were studied with the use of standard extracellular recording techniques. The receptive fields of multisensory neurons were large, as reported previously, but exhibited a surprisingly high degree of spatial coincidence. The average proportion of receptive-field overlap was 86% for the population of visual-auditory neurons sampled. 3. Because of this high degree of intersensory receptive-field correspondence, combined-modality stimuli that were coincident in space tended to fall within the excitatory regions of the receptive fields involved. The result was a significantly enhanced neuronal response in 88% of the multisensory neurons studied. If stimuli were spatially disparate, so that one fell outside its receptive field, either a decreased response occurred (56%), or no intersensory effect was apparent (44%). 4. The normal alignment of the different receptive fields of a multisensory neuron could be disrupted by passively displacing the eyes, pinnae, or limbs/body. In no case was a shift in location or size observed in a neuron's other receptive field(s) to compensate for this displacement. The physiological result of receptive-field misalignment was predictable and based on the location of the stimuli relative to the new positions of their respective receptive fields. Now, for example, one component of a spatially coincident pair of stimuli might fall outside its receptive field and inhibit the other's effects. 5. These data underscore the dependence of multisensory integrative responses on the relationship of the different stimuli to their corresponding receptive fields rather than to the spatial relationship of the stimuli to one another. Apparently, the alignment of different receptive fields for individual multisensory neurons ensures that responses to combinations of stimuli derived from the same event are integrated to increase the salience of that event. Therefore the maintenance of receptive-field alignment is critical for the appropriate integration of converging sensory signals and, ultimately, elicitation of adaptive behaviors.

Receptive-field characteristics of neurons in cat striate cortex: Changes with visual field eccentricity

1976 ◽  
Vol 39 (3) ◽  
pp. 512-533 ◽  
Author(s):  
J. R. Wilson ◽  
S. M. Sherman
Keyword(s):  
Visual Field ◽  
Receptive Field ◽  
Striate Cortex ◽  
Ocular Dominance ◽  
Receptive Fields ◽  
Direction Selectivity ◽  
Orientation Selectivity ◽  
Simple Cells ◽  
Complex Cells ◽  
Cat Striate Cortex

1. Receptive-field properties of 214 neurons from cat striate cortex were studied with particular emphasis on: a) classification, b) field size, c) orientation selectivity, d) direction selectivity, e) speed selectivity, and f) ocular dominance. We studied receptive fields located throughtout the visual field, including the monocular segment, to determine how receptivefield properties changed with eccentricity in the visual field.2. We classified 98 cells as "simple," 80 as "complex," 21 as "hypercomplex," and 15 in other categories. The proportion of complex cells relative to simple cells increased monotonically with receptive-field eccenticity.3. Direction selectivity and preferred orientation did not measurably change with eccentricity. Through most of the binocular segment, this was also true for ocular dominance; however, at the edge of the binocular segment, there were more fields dominated by the contralateral eye.4. Cells had larger receptive fields, less orientation selectivity, and higher preferred speeds with increasing eccentricity. However, these changes were considerably more pronounced for complex than for simple cells.5. These data suggest that simple and complex cells analyze different aspects of a visual stimulus, and we provide a hypothesis which suggests that simple cells analyze input typically from one (or a few) geniculate neurons, while complex cells receive input from a larger region of geniculate neurons. On average, this region is invariant with eccentricity and, due to a changing magnification factor, complex fields increase in size with eccentricity much more than do simple cells. For complex cells, computations of this geniculate region transformed to cortical space provide a cortical extent equal to the spread of pyramidal cell basal dendrites.

scholarly journals Beyond the labeled line: variation in visual reference frames from intraparietal cortex to frontal eye fields and the superior colliculus

2017 ◽  
Author(s):  
V. C. Caruso ◽  
D. S. Pages ◽  
M. A. Sommer ◽  
J. M. Groh
Keyword(s):  
Receptive Field ◽  
Superior Colliculus ◽  
Reference Frames ◽  
Receptive Fields ◽  
Visual Signals ◽  
Eye Position ◽  
Frontal Eye Fields ◽  
Response Patterns ◽  
The Stability ◽  
Retinal Information

ABSTRACTWe accurately perceive the visual scene despite moving our eyes ~3 times per second, an ability that requires incorporation of eye position and retinal information. We assessed how this neural computation unfolds across three interconnected structures: frontal eye fields (FEF), intraparietal cortex (LIP/MIP), and the superior colliculus (SC). Single unit activity was assessed in head-restrained monkeys performing visually-guided saccades from different initial fixations. As previously shown, the receptive fields of most LIP/MIP neurons shifted to novel positions on the retina for each eye position, and these locations were not clearly related to each other in either eye- or head-centered coordinates (hybrid coordinates). In contrast, the receptive fields of most SC neurons were stable in eye-centered coordinates. In FEF, visual signals were intermediate between those patterns: around 60% were eye-centered, whereas the remainder showed changes in receptive field location, boundaries, or responsiveness that rendered the response patterns hybrid or occasionally head-centered. These results suggest that FEF may act as a transitional step in an evolution of coordinates between LIP/MIP and SC. The persistence across cortical areas of hybrid representations that do not provide unequivocal location labels in a consistent reference frame has implications for how these representations must be read-out.New & NoteworthyHow we perceive the world as stable using mobile retinas is poorly understood. We compared the stability of visual receptive fields across different fixation positions in three visuomotor regions. Irregular changes in receptive field position were ubiquitous in intraparietal cortex, evident but less common in the frontal eye fields, and negligible in the superior colliculus (SC), where receptive fields shifted reliably across fixations. Only the SC provides a stable labelled-line code for stimuli across saccades.

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