Evidence of midline retinal nasotemporal overlap in healthy humans: A model for foveal sparing in hemianopia?

The existence of midline retinal nasotemporal overlap in humans is controversial. Here we used the Poffenberger paradigm and monocular vision to assess the existence of a midline retinal area projecting to both hemispheres and of a possible differential contribution of the two hemiretinae. When brief visual stimuli were presented at 1° eccentricity they were responded to equally quickly with either hand while at 6° the hand on the same side as the stimulated hemifield was consistently faster than the contralateral hand. This pattern of results is consistent with a nasotemporal overlap at 1° and a complete lateralization at 6°. Both hemiretinae contribute to the overlap area which can be considered as responsible for foveal sparing in hemianopic patients.     

Shifting attention in viewer- and object-based reference frames after unilateral brain injury

The aims of the present study were to investigate the respective roles that object- and viewer-based reference frames play in reorienting visual attention, and to assess their influence after unilateral brain injury. To do so, we studied 16 right hemisphere injured (RHI) and 13 left hemisphere injured (LHI) patients. We used a cueing design that manipulates the location of cues and targets relative to a display comprised of two rectangles (i.e., objects). Unlike previous studies with patients, we presented all cues at midline rather than in the left or right visual fields. Thus, in the critical conditions in which targets were presented laterally, reorienting of attention was always from a midline cue. Performance was measured for lateralized target detection as a function of viewer-based (contra- and ipsilesional sides) and object-based (requiring reorienting within or between objects) reference frames. As expected, contralesional detection was slower than ipsilesional detection for the patients. More importantly, objects influenced target detection differently in the contralesional and ipsilesional fields. Contralesionally, reorienting to a target within the cued object took longer than reorienting to a target in the same location but in the uncued object. This finding is consistent with object-based neglect. Ipsilesionally, the means were in the opposite direction. Furthermore, no significant difference was found in object-based influences between the patient groups (RHI vs. LHI). These findings are discussed in the context of reference frames used in reorienting attention for target detection.     

Nasotemporal overlap of crossed and uncrossed retinal ganglion cell projections in the Japanese monkey (Macaca fuscata)

The nasotemporal overlap of crossed and uncrossed retinal ganglion cell projections were studied in 11 Japanese monkeys (Macaca fuscata) using HRP and fluorescent dyes (DAPI and RITC) as retrograde tracers and by physiological recordings of antidromic field potentials. A strip of nasotemporal overlap ran orthogonal to the horizontal meridian in all the whole-mount retinas studied. In HRP-labeled retinas of 6 monkeys, the width of the overlap gradually increased from 0.6 degrees in the central retina up to 5 degrees at eccentricity of 5 mm, and to 15 degrees at the extreme periphery. We also noted a clear asymmetric distribution of crossed and uncrossed retinal ganglion cell projections particularly in the perifoveal region; ipsilaterally projecting cells encircled the nasal edge of the fovea, whereas few contralaterally projecting cells were observed in the temporal edge. Soma-size analysis revealed that crossed projections in the temporal portion of the overlap arose mainly from large and small cells (presumably P alpha and P gamma cells, respectively); uncrossed projections in its nasal portion arose from medium-sized cells (presumably P beta cell). Direct evidence of the overlap as well as of the asymmetry was obtained in subsequent fluorescent dye experiments in 3 monkeys. Physiological studies on 2 additional monkeys confirmed the widening of the nasotemporal overlap towards the upper and lower parts of the retina. Moreover, in the nasal portion of the overlap, only slow potentials, which presumably reflect activities of P beta cells, were recorded after stimulation of the ipsilateral LGN as expected from the morphological study. The findings are discussed in relation to clinical observations of macular sparing and splitting, and with regard to the functional differences between P alpha and P beta cell systems on which binocular stereoscopic vision along the midsagittal plane may be based.     

The cortical representation of foveal stimuli: evidence from quadrantanopia and TMS-induced suppression

To address the extent to which the visual foveal representation is split, we examined a 29-year-old patient with a lower right quadrantanopia following surgical removal of the left occipital cortex above the calcarine sulcus and compared her performance with subjects receiving transcranial magnetic stimulation (TMS) over the occipital lobes. In a letter/digit classification task, the patient responded accurately to targets presented in the upper visual field, for all horizontal eccentricities. In the lower visual field, she failed to discriminate letters from digits when targets were presented in the right, but not the left visual field (RVF and LVF, respectively). This pattern was also true for the foveal targets, with poor performance to foveal-RVF (0.5 degrees to the right of fixation) but not foveal-LVF (0.5 degrees to the left of fixation) targets. Similar patterns of normal performance to LVF but not RVF or foveal-RVF targets were observed in a group of nine normal observers when TMS was applied over their left occipital cortex. Complementary impairments to LVF and foveal-LVF target classification were induced with TMS over the right occipital cortex. Thus, we have induced an hemianopic pattern in normal observers contralateral to the magnetically stimulated hemisphere. This correspondence between real and TMS-induced visual field defects is further evidence, in neurologically intact subjects, that the cortical representation of the fovea is split between the two hemispheres along the vertical meridian.     

Single representation of the visual midline in humans

A choice reaction time experiment was performed to find evidence of dual representation of the visual midline in normal humans. Visual targets were presented either to the left or right of fixation and subjects responded by pressing one of two keys. Major variables were: retinal locus (4, 3.5, 3, 2.5, 2, 1.5, 1, 0.75, 0.5 and 0.25° from fixation); visual field (left or right), eye (left or right) and responding hand (left or right). RT was 25 msec faster when the target projected to the hemisphere controlling the responding hand (uncrossed) than when the two separate hemispheres were involved (crossed). The RT difference was maintained even for targets 15 min arc from fixation, provided no evidence for dual representation.     

Visual-field asymmetry in dual-stream RSVP

The attentional blink (AB) refers to a decrement in detecting the occurrence of a probe item if it closely follows a previous target item in rapid serial visual presentation (RSVP). In the present study we presented target and probe stimuli in two parallel RSVP streams, one in each visual field, in order to address the question of whether the AB might differ between the cerebral hemispheres. The characteristic AB, with reduced detection of the probe at post-target Lags 2-5, but no such deficit at Lag 1 (Lag 1 sparing), was observed when target and probe were both in the right visual field. When they were both presented in the left visual field the AB was attenuated. When the target and probe were in opposite visual fields, probe detection was again reduced when it was in the left visual field, and there was no Lag 1 sparing. The left-visual-field advantage in performing the AB task may reflect a general right-hemispheric specialization for attentional processing.     

Assessing effects of fixation demands on perception of lateralized words: A visual window technique for studying hemispheric asymmetry

A major concern when using lateralized words to study hemispheric asymmetry is that the retinal eccentricity of targets is matched across visual hemifields. The standard technique is to fixate a point fixed at the centre of the visual field. However, the demands of this fixation task are substantial and so may confound performance with lateralized targets. To investigate this possibility, words were presented unilaterally in each visual hemifield and retinal eccentricity was controlled using (a) a fixed central point or (b) a window technique that permitted small shifts in fixation while maintaining accurate retinal eccentricity by using automatic adjustments to target location. Fixation errors and time to fixate indicated that the demands of the standard technique were considerable and far greater than those of the window technique. Nevertheless, both techniques produced the same pattern of visual field effects, indicating that the demands of fixating a fixed central point do not confound performance with lateralized words. However, the window technique was more efficient and easier for participants to use and so offers a new improved methodology for studying hemispheric asymmetry.     

Visual search training for a case of homonymous field defect with multiple visual dysfunctions]

A 72-year-old right handed man developed right homonymous hemianopia without macular sparing, left homonymous lower quadranopia with macular sparing, cerebral amblyopia, cerebral achromatopsia, impaired form vision, and mild right hemispatial neglect, after multiple cerebral infarctions, involving bilateral occipital cortices. His intelligence and memory were deteriorated moderately. He failed to notice objects located in the affected visual field, because of his severely impaired visual search. When ordinary lighting was used, he showed severe right-sided omissions on the line cancellation test. However, omissions were less marked under the brighter lighting. By using a modified method of Kerkhoff and Vianen (1994), he was trained to make saccadic eye movements toward affected regions to find a target and to search and point at targets arranged randomly. As the sensitivity for contrast of isoluminante red and green stimuli was preserved well at high spatial frequencies despite the decreaced contrast sensitivity for brightness, we used green targets as the training stimuli. After the training, search field and pointing range that could be covered by the patient increased in size for both green and white targets, and daily activities improved. Moreover, after the training, he no longer showed discrepancy in line cancellation performances between ordinary and brighter lighting conditions. In the follow up period, the search field and the performance on the line cancellation test were maintained, while the performance of pointing targets array declined. The family members complained of mild re-deterioration of daily activities. Then, the training for searching and pointing re-introduced at home. After the training, his pointing performance and daily activities, evaluated by questionnaires to his family members, improved again. In conclusion, it was suggested that disordered visual search after a homonymous field defect can be treated effectively, even if multiple visual dysfunctions were associated.     

The mapping of the visual field onto the dorso-lateral tectum of the pigeon (Columba livia) and its relations with retinal specializations

Most of the physiological studies of the pigeon retino-tectal visual pathway have investigated the accessible tectum, a small dorso-lateral tectal section that can be easily accessed by a simple craniotomy. However, at present we lack a detailed study of the topographical arrangement between the visual field, the retina and the accessible tectum. In particular, it is not known which section of the visual field is mapped onto the accessible tectum, and which of the specialized retinal areas mediates this projection. Here we determined, using local field potential (LFP) recordings and reverse retinoscopy, the shape, size and position in the visual space of the portion of the visual field mapped onto the accessible tectum (called here the accessible visual field, or AVF). Using this data and the mapping of Nalbach et al. [Vis. Res. 30 (4) (1990) 529], the retinal area corresponding to the AVF was determined. Such retinal area was also directly delimited by means of retrograde transport of DiI. The results indicate that the AVF is a triangular perifoveal zone encompassing only 15% of total visual field. The retinal region corresponding to the AVF has the shape of an elongated triangle that runs parallel to the visual equator and contains the fovea, the tip of the pecten, a perifoveal region of the yellow field and a small crescent of the red field. In agreement with this anatomical heterogeneity, visual evoked potentials measured in different parts of the accessible tectum present steep variations in shape and size. These results are helpful to better design and interpret anatomical and physiological experiments involving the pigeon's visual system.     

Interhemispheric integration of letter stimuli presented foveally or extra-foveally

Two experiments are reported in which participants decided whether a single target letter presented below and to the left or right of fixation matched either of two probe letters presented above the fixation cross to the left and right. The level of matching required was either physical (A-A) or abstract (A-a). All three letters were presented either extra-foveally (Experiment 1) or within the fovea (Experiment 2). In both experiments, physical matching was faster than abstract matching. Physical matching was faster within than across visual fields while abstract matching showed the opposite pattern. Matching was faster when the matching probe letter was in the LVF than when it was in the RVF. Importantly, the pattern of results was the same for extra-foveal and foveal presentations, supporting the theory that the representation of the fovea is split down the middle into two visual fields rather then being bilateral. The practical implication following this study is that lateralization studies can be performed with closer to fixation stimuli presentation.     

Transient interference of right hemispheric function due to automatic emotional processing

We examined the effects of emotional stimuli on right and left hemisphere detection performance in a hemifield visual discrimination task. A group of 18 healthy subjects were asked to discriminate between upright and inverted triangles (target). Targets were randomly presented in the left or right visual hemifield (150 ms target duration). A brief emotional picture (pleasant or unpleasant; 150 ms stimulus duration) or neutral picture selected from the International Affective Picture System was randomly presented either in the same (47%) or the opposite (47%) spatial location to the subsequent target. Emotional or neutral stimuli offset 150 ms prior to the subsequent target. Subjects were instructed to ignore the pictures and respond to the targets as quickly and accurately as possible. Independent of field of presentation, emotional stimuli prolonged reaction times (P < 0.01) to LVF targets, with unpleasant stimuli showing a greater effect than pleasant stimuli. The current study shows that brief emotional stimuli selectively impair right hemispheric visual discrimination capacity. The findings suggest automatic processing of emotional stimuli captures right hemispheric processing resources and transiently interferes with other right hemispheric functions.     

Detection and discrimination of chromatic targets in hemianopic macaque monkeys and humans

We measured the ability of three macaque monkeys with unilateral removal of primary visual cortex to detect 9 degrees, 200-ms targets presented at random in the upper or lower quadrants of the normal and hemianopic visual fields. The white or variously coloured target could differ from the background in both colour and luminance, or in either of them. Blue and red targets were detectable at any luminance contrast, but green and white targets were barely or not at all detectable at and near isoluminance in the hemianopic field. Blue and red targets were better detected than white targets at the same luminance difference. However, when both the target and the background were dynamically luminance-masked, detection in the hemianopic field failed at isoluminance whatever the colour. In addition, the monkeys were unable to discriminate between simultaneously presented red and green or blue and green targets in the hemianopic field when both targets had similar luminance contrast with the background. Two hemianopic patients tested on a subset of the tasks performed similarly to the monkeys. Together, the results indicate that the residual colour-opponent system that survives damage to V1 is involved in the detection of chromatic changes but cannot sustain simultaneous discrimination between pairs of colours.     

Visual localization of sounds

Circumscribed hemispheric lesions in the right hemisphere have been shown to impair auditory spatial functions. Due to a strong crossmodal links that exist between vision and audition, in the present study, we have hypothesized that multisensory integration can play a specific role in recovery from spatial representational deficits. To this aim, a patient with severe auditory localization defect was asked to indicate verbally the spatial position where the sound was presented. The auditory targets were presented at different spatial locations, at 8 degrees, 24 degrees, 40 degrees, 56 degrees to either sides of the central fixation point. The task was performed either in a unimodal condition (i.e., only sounds were presented) or in crossmodal conditions (i.e., a visual stimulus was presented simultaneously to the auditory target). In the crossmodal conditions, the visual cue was presented either at the same spatial position as the sound or at 16 degrees or 32 degrees, nasal or temporal, of spatial disparity from the auditory target. The results showed that a visual stimulus strongly improves the patient's ability to localize the sounds, but only when it was presented in the same spatial position of the auditory target.     

The role of striate cortex in the guidance of eye movements in the monkey

We studied the effect of unilateral striate cortical ablations on smooth pursuit and saccadic eye movements in the monkey. The monkeys made quite accurate saccades to stationary stimuli in the field contralateral to the lesion, and they readily pursued foveal targets moving in all directions. However, when visual stimuli were stepped into the field contralateral to the lesion and then began to move, thus insuring that the moving stimulus was confined to the impaired visual hemifield, several oculomotor abnormalities emerged. Saccades to moving stimuli presented in the impaired field consistently undershot targets that moved away from the central fixation point after the step, and overshot targets that moved back towards the central fixation point. There was little or no smooth pursuit eye velocity generated in any direction to moving stimuli in the impaired field, and the monkeys could not generate smooth pursuit to stimuli maintained a few degrees from the fovea in the impaired field, although they were able to pursue such stimuli held in the normal field. Ablation of striate cortex also affected the latencies of saccades. When step-ramp stimuli were presented in the normal field, the monkeys delayed the initiation of saccades to targets moving towards the central fixation point, and hastened the initiation of saccades to targets moving away from the central fixation point. By contrast, changes in the direction of target movement did not affect the latencies of saccades into the impaired field. The deficits seemed permanent, lasting as long as the monkeys were tested--over 2 years in one case--but they were not total. Each monkey could use stimuli moving into the affected field to develop some eye velocity, although this residual ability had a much longer latency and lower gain than that provided by the intact visual system. These results show that striate cortex is intimately involved in the estimation of stimulus velocity critical to the genesis of smooth pursuit and saccadic eye movements.     

The visual system of a Palaeognathous bird: Visual field,retinal topography and retino‐central connections in the Chilean Tinamou (Nothoprocta perdicaria)

Most systematic studies of the avian visual system have focused on Neognathous species, leaving virtually unexplored the Palaeognathae, comprised of the flightless ratites and the South American tinamous. We investigated the visual field, the retinal topography, and the pattern of retinal and centrifugal projections in the Chilean tinamou, a small Palaeognath of the family Tinamidae. The tinamou has a panoramic visual field with a small frontal binocular overlap of 20°. The retina possesses three distinct topographic specializations: a horizontal visual streak, a dorsotemporal area, and an area centralis with a shallow fovea. The maximum ganglion cell density is 61,900/ mm², comparable to Falconiformes. This would provide a maximal visual acuity of 14.0 cycles/degree, in spite of relatively small eyes. The central retinal projections generally conform to the characteristic arrangement observed in Neognathae, with well‐differentiated contralateral targets and very few ipsilateral fibers. The centrifugal visual system is composed of a considerable number of multipolar centrifugal neurons, resembling the “ectopic” neurons described in Neognathae. They form a diffuse nuclear structure, which may correspond to the ancestral condition shared with other sauropsids. A notable feature is the presence of terminals in deep tectal layers 11–13. These fibers may represent either a novel retinotectal pathway or collateral branches from centrifugal neurons projecting to the retina. Both types of connections have been described in chicken embryos. Our results widen the basis for comparative studies of the vertebrate visual system, stressing the conserved character of the visual projections' pattern within the avian clade. J. Comp. Neurol. 523:226–250, 2015. © 2014 Wiley Periodicals, Inc.     

The effects of parvocellular lateral geniculate lesions on the acuity and contrast sensitivity of macaque monkeys

The effects of ablating the visual pathway that passes through the parvocellular (dorsal) LGN were tested in 2 macaque monkeys by measuring acuity and both luminance and chromatic contrast sensitivity. Thresholds were tested monocularly before and after ibotenic acid was used to lesion parvocellular layers 4 and 6 of the contralateral geniculate. The injections were centered at the representation of 6 degrees in the temporal field on the horizontal meridian, and vision was tested with localized stimuli at this location. In addition, in one of the monkeys, a lesion was made in magnocellular layer 1 of the opposite geniculate, and the same thresholds were tested. Physiological and anatomical reconstructions demonstrated complete destruction of the target layers in 1 parvocellular lesions and in the magnocellular lesion, and sparing of the nontarget layers in the tested region. Parvocellular lesions caused a 3-4-fold reduction in visual acuity within the affected part of the visual field, while the magnocellular lesion did not affect acuity. Both luminance and chromatic contrast sensitivity, tested with stationary gratings of 2 c/degree, were severely reduced by parvocellular lesions, but not affected by the magnocellular lesion. However, when luminance contrast sensitivity was tested with 1 c/degree gratings, counterphase modulated at 10 Hz, it was reduced by both parvocellular and magnocellular lesions. This study demonstrates that the parvocellular pathway dominates chromatic vision, acuity, and contrast detection at low temporal and high spatial frequencies, while the magnocellular pathway may mediate contrast detection at higher temporal and lower spatial frequencies.     

The role of perceptual reference frames in visual field asymmetries

The direct access model of hemispheric asymmetry was tested in a letter reflection (normal of reflected) judgement task. In the baseline condition letters were presented in their upright orientation, and reaction times were faster to letters presented in the right visual field than to those presented in the left visual field. In two other conditions the slides, and thus the letters, were rotated +90 degrees clockwise (Rotated +90) from upright or -90 degrees counterclockwise (Rotated -90). This resulted in the letters in both rotated conditions being shown in the upper or lower visual field through the sagittal plane but rotated 90 degrees. Despite this fact a "right field" advantage was again found when the right field was defined relative to the orientation of the tops of the rotated letters (lower visual field in the Rotated +90 condition and upper visual field in the Rotated -90 condition). These results demonstrate that the internal representation of locations in space is more important in predicting visual field asymmetries, at least in the present task, than the field of stimulus presentation relative to the fovea.     

Ambulatory autonomy and visual motion perception in a case of almost total cortical blindness]

A 37-year-old man experienced cortical blindness following a bilateral stroke in the territory of the posterior cerebral arteries. Four years later, the measurement of visual field defects (Goldmann perimeter) showed persistence of bilateral blindness with a 2-degree preservation of macular vision and a perifoveal sparing between 10 to 30 degrees of eccentricity in the left inferior quadrant. Despite this visual impairment, the subject was able to perform visually-guided locomotion. Moreover he consciously perceived visual motion in the blind parts of his visual field. CT and MRI showed a lesion involving most of the striated cortex. The visual cortex located in the internal occipito-parital regions was relatively spared. The contribution of this structure to extra-striated vision of motion is discussed.     

Saccadic adaptation without retinal postsaccadic error

Primary saccades undershoot their target. Corrective saccades are then triggered by retinal postsaccadic information. We tested whether primary saccades still undershoot when no postsaccadic visual information is available. Participants saccaded to five targets (10-34 degrees) that were either constantly illuminated (ON) or extinguished at saccade onset (OFF(Onset)). In OFF(Onset), few corrective saccades were observed. The saccadic gain increased over trials for the furthest (34 degrees) target. Terminal eye position after glissades or microsaccades progressively converged to the values observed in ON (targets over 16 degrees). Target extinction during the saccade only did not elicit any change. The results show that (i) postsaccadic retinal signals stabilize the saccadic gain and (ii) adaptive changes that reduce terminal error can take place without visual information.