Effects of optokinetic velocity and medial vestibulo-cerebellum lesions on vestibulo-ocular reflex direction adaptation in the cat

Adaptive horizontal vestibulo-ocular reflex eye movements in response to vertical (pitch) rotations were produced by exposing three cats to synchronized horizontal optokinetic and vertical vestibular oscillations at 0.25 Hz. The effects of optokinetic stimulus velocity (6–80°/s peak) and nodulo-uvular cerebellum lesions were studied. All optokinetic velocities elicited vestibulo-ocular reflex direction adaptation, though 6°/s stimuli were somewhat less effective. Restricted aspirations of cerebellar vermis lobules IX and X in two cats resulted in reduced but still clearly evident adaptation.     

Video-oculography in the gerbil

Normative vestibulo-ocular and optokinetic reflexes (VOR and OKR) and pupil diameter were measured in young adult gerbils using infrared video-oculography with 60 Hz sampling during head-fixed binocular recordings. The pupillary light-sink technique was preferred over a single-beam retinal reflection method because its measurements were less affected by pupil size. Eye movements were generally conjugate with occasional independent saccadic movements, and independent drifting movements in the dark. The horizontal optokinetic response to sinusoidal motion of a randomly spaced white dot pattern was maximal at low velocities (5 degrees/s), stronger temporonasally, and dropped off quickly at approximately 20 degrees/s. Constant velocity gain was near unity through 60-80 degrees/s with a sharp drop-off. Monocular viewing revealed almost no nasotemporal optokinetic response. Pupil diameter was found to vary as a saddle function with optokinetic gain from cycle to cycle, but also have a circadian rhythm (smaller at dusk) that related inversely to mean horizontal VOR gain. Gerbils with eyes open sometimes had no optokinetic response during long stimulus periods, which then resumed after a brief vestibular stimulus. The horizontal angular VOR gain was relatively flat across 0.1-1.0 Hz and 30-120 d/s sines (phase near zero), with a mean gain of approximately 0.78 in the dark, and 1.0 with the fixed pattern surround (n=15, for both raw calibrated and normalized data). Most animals also revealed a strong slow phase eye velocity asymmetry (dominant during ipsilateral rotation) in the half-cycle gain of their horizontal angular VOR response in the dark. A constant velocity horizontal optokinetic bias velocity did not change the gain or symmetry of the sinusoidal VOR response, but shifted the VOR response velocity in an additive (linear) fashion. Both cross-coupling (pitch or roll while rotating) and pseudo-OVAR (off-axis counter-rotation) stimuli generated horizontal nystagmus. The findings suggest that the gerbil, like other lateral-eyed rodents, relies on otolith cues to interpret angular motion.     

Functional properties of area 19 as compared to area 17 of the cat

Out of a total of 139 area 19 cells, 87 were examined quantitatively for their responses to stimuli moving at a wide range of velocities. The results were compared with those obtained on a sample of 106 area 17 cells (out of a total of 172) tested in the same way.It was found that both areas 19 and 17 prefer slow stimulus movement. However, area 17 responded well to velocities up to 4%/s while response amplitudes of area 19 cells started to decrease for stimulus velocities over 0.3%/s. In both areas, responsiveness to fast stimuli improved at higher eccentricities. The most frequently encountered velocity type in area 19 was the ‘velocity broad band’ (VBB) type, lacking velocity preference, while the rarest type was the ‘velocity tuned’ (VT) type. As in area 17 very few area 19 cells were found to prefer high velocities (velocity high pass type, VHP) and those that were encountered had peripheral receptive fields (RFs). Cells preferring exclusively low velocities (velocity low pass type, VLP) were less frequent in area 19 than in area 17 and were considerably more sluggish.Area 19 was also less direction and orientation selective than area 17. In contrast, end-stopping was very common in area 19 (66%) and more units were binocular as compared to area 17. On average firing rate during optimal stimulation was lower in area 19 than in area 17.These results are consistent with the notion that area 19 receives predominantly W-type input and is involved in form discrimination (low spatial frequencies) during fixation.     

Visual receptive fields in the lateral suprasylvian area (Clare-Bishop area) of the cat.

Single units were recorded from the visual area of the lateral suprasylvian gyrus (LSSA or Clare-Bishop area) in 20 unanesthetized cats. Most LSSA units were poorly responsive to stationary visual stimuli, but they responded vigorously to moving visual stimuli. Their receptive fields appeared to be constituted of a large activating region (discharge area) often surrounded by inhibitory flanks. Relating unit behavior to changes of stimulus length, the LSSA neurons could be subdivided into 5 categories. The first category (22 out of 95 units tested, 23.16%) consisted of units showing summation inside the discharge area. Expanding the stimulus outside the discharge area did not affect the response. The second category (7.37%) was formed by units which showed summation inside the discharge area and inhibition when the stimulus was extended outside the discharge area. The third category (21.05%) consisted of units largely insensitive to the stimulus length inside the discharge area, but surrounded by inhibitory flanks. The fourth category (41.05%) consisted of units which showed inhibition of the response when the stimulus, well inside the discharge area, became longer than a certain optimal lenght. They were surrounded by inhibitory flanks. The fifth category (7.37%) was formed by units insensitive to variations of the stimulus length inside as well as outside the discharge area. Almost all units, independent of their category, were directionally specific, that is their response could be decreased 50% or more by varying the direction of movement away from that which gave the maximal response (preferred direction). Typically the response was halved when the stimulus was moved +/- 50 degrees from the preferred direction. Among the directionally specific units, 71% showed the minimal response 180 degrees away from the preferred direction (direction specificity curve type 1), 20% had the minimal response 90 degrees from the preferred direction (direction specificity curve type 2); the remaining could not be classified in this respect. Of LSSA units, 87% (all those of type 1 and many of those of type 2) were directionally selective, that is their response to movement in the preferred direction was at least double that in the opposite direction. The LSSA units usually preferred stimuli moving at rather high speeds. The optimal speed for 71% of units was 20 degrees/sec or greater. Almost all units responded over a wide range of speeds, many of them from 5-10 degrees/sec to over 100 degrees/sec. Most neurons had a low spontaneous activity and some of them remained completely silent for seconds.     

Responses of Neurons of the Nucleus of the Optic Tract and the Dorsal Terminal Nucleus of the Accessory Optic Tract in the Awake Monkey

The nucleus of the optic tract (NOT) and the dorsal terminal nucleus of the accessory optic tract (DTN) are essential nuclei for the generation of slow-phase eye movements during horizontal optokinetic nystagmus. We recorded from 101 neurons (all directionally selective) in four NOT/DTN of three trained and behaving rhesus monkeys. Neuronal activity increased when stimuli moved ipsiversively with respect to the recording site and decreased below spontaneous activity when stimuli moved contraversively. While the monkey fixated a small spot, some NOT/DTN neurons did not respond at all to the retinal image slip of a whole-field random dot pattern; others showed a monotonic increase of activity to increasing velocities of that stimulus. The velocity range tested was up to 100°/s. During the execution of optokinetic nystagmus, 39 of 73 cells tested showed a velocity-tuned response with an average optimum at 21°/s retinal image slip. Following saccades during optokinetic nystagmus (quick phases), the NOT/DTN neuronal activity briefly attained the level of spontaneous activity, as predicted from the velocity selectivity during optokinetic nystagmus. Immediately upon cessation of optokinetic stimulation in the preferred direction, NOT/DTN activity returned to the spontaneous level and did not reflect the ongoing optokinetic afternystagmus in darkness. Most NOT/DTN neurons displayed direction selectivity also during smooth pursuit. Twenty-one of 50 cells tested (42%) always responded to the retinal slip of the target (target velocity cells), 16 cells (32%) responded to the retinal slip of the background (background velocity cells), and 13 cells (26%) did not respond at all during smooth pursuit. We conclude from our results that the NOT/DTN is an essential structure for the processing of the direction and speed of retinal image slip. This information is then used for the generation and maintenance of slow eye movements, preferentially during horizontal optokinetic nystagmus but also during pursuit eye movements.     

Parallel and serial visual search after closed head injury: electrophysiological evidence for perceptual dysfunctions.

Event-related potentials (ERPs) were recorded from closed head injury (CHI) patients at least 2 years postinjury and from controls in order to assess their parallel and serial processing abilities during visual search. In Experiment 1, stimuli consisted of arrays of eight triangles; half of the arrays contained a target item. In the "feature-present" condition, the target item was a triangle with an additional horizontal line that could be detected automatically and in parallel, while in the "feature-absent" condition all items except for the target triangle had an additional horizontal line, thus requiring a serial search. In Experiment 2, stimuli consisted of eight solid bars (50%), seven solid bars and a vertical open bar (25%), and seven solid bars and a horizontal open bar (25%): the array containing the horizontal bar served as a target. By recording ERPs to the arrays containing vertical open bars, which were similar to the target items, parallel processing of "pop-out" stimuli could be studied in the absence of any overt response. ERP data were compared with the results of neuropsychological and neuroimaging (MRI, CAT) examination. Patient exhibited a decreased behavioral performance both in the parallel and in the serial processing mode. Furthermore, abnormalities of early and intermediate ERP components (P1, N1, P2, N2) were found, whereas the late component (P3) was less affected by CHI. The results were interpreted as an index of CHI-induced dysfunctions in perceptual processes such as simple feature registration and early target discrimination. It was suggested that these dysfunctions contribute to impairments of parallel as well as serial processes in visual search.     

Incidence of F waves in single human thenar motor units.

F-wave generation, axon conduction velocities, and contractile properties were compared in 44 healthy individual human thenar motor units. Force and muscle action potentials were recorded when single motor axons were stimulated intraneurally about 10 cm proximal to the elbow. Each stimulus usually evoked only one electromyographic (EMG) potential. However, in seven units (16%), a single stimulus elicited an F wave in response to 1.7 +/- 1.6% (mean +/- SD) of the stimuli applied. Axon conduction velocity proximal to the site of stimulation was faster than distal conduction velocity (72.7 +/- 8.0 m/s versus 64.2 +/- 10.5 m/s). Distal conduction velocities, twitch forces, and contraction times were similar for units that did and did not generate F waves. Thus, no obvious subset of thenar motor units generated F waves. These results provide valuable baseline information on F waves that can be used to assess changes in axon conduction, motor unit contractile properties, and motoneuron excitability in disease.     

Tectal neurons signal impending collision of looming objects in the pigeon

Although the optic tectum in non-mammals and its mammalian homolog, the superior colliculus, are involved in avoidance behaviors, whether and how tectal neurons respond to an object approaching on a collision course towards the animal remain unclear. Here we show by single unit recording that there exist three classes of looming-sensitive neurons in the pigeon tectal layer 13, which sends looming information to the nucleus rotundus or to the tectopontine system. The response onset time of tau cells is approximately constant whereas that for rho and eta cells depends on the square root of the diameter/velocity ratio of objects looming towards the animal, the cardioacceleration of which is also linearly related to the square root of this ratio. The receptive field of tectal cells is composed of an excitatory center and an inhibitory periphery, and this periphery does not inhibit responses to looming stimuli. These results suggest that three classes of tectal neurons are specified for detecting an object approaching on a collision course towards the animal, and that rho and eta cells may signal early warning of impending collision whereas tau cells initiate avoidance responses at a constant time before collision through the tectopontine system.     

Visual direction-selective neurons in the pretectum of the rainbow trout

In mammals, the essential neuronal substrate for the generation of the horizontal optokinetic nystagmus (hOKN) are the nucleus of optical tract (NOT) and the dorsal terminal nucleus (DTN). The medial terminal nucleus (MTN) is thought to be involved in vertical OKN control. Characteristic for all of these neurons is a high-direction selectivity. Although behavioural hOKN experiments in different fish species show comparability to mammals, little is known about the neuronal OKN control in fish. In preceding studies, we demonstrated that the rainbow trout has a nearly symmetrical monocular hOKN at low stimulus speeds. With increasing visual stimulus speeds (>14 degrees /s), the monocular hOKN becomes asymmetrical with a temporo to nasal preferred direction. For visual stimulation, we presented random-dot-patterns projected by a planetarium inside a perimeter. We tested four rotation axes of the planetarium, yaw (0 degrees -180 degrees ), roll (90 degrees -270 degrees ), diagonal (45 degrees -225 degrees ) and anti-diagonal (135 degrees -315 degrees ). In every position, the visual stimulus turned in clockwise and counter-clockwise direction. In a subregion of the pretectum of nine fish, we recorded 47 direction-selective neurons. Analysis of tuning-curves and preferred direction vectors show that these neurons encode both horizontal (yaw) and vertical (roll) visual stimulus directions. These results suggest that the control of horizontal and vertical OKN might not segregate into different nuclei in fish.     

Anatomy and physiology of a binocular system in the frograna pipiens

The locations of tectal neurons projecting to nucleus isthmi (n. isthmi) were found by iontophoretic injection of horseradish peroxidase (HRP) into n. isthmi. After retrograde transport, stained tectal somata are found to lie almost exclusively in layer 6 and below of the ipsilateral tectum. Many cells are colored throughout the extent of their dendrites into the fine rami, giving the appearance of a Golgi stain. Nucleus isthmi receives projections from the ipsilateral tectum and from no other region. Nucleus isthmi units recorded electrically respond to visual stimuli and are arranged in a topographic map of the visual field. There are two types of receptive fields, those with small centers and those with large centers. The small centers are about 3-5 degrees in diameter, similar to type 2 optic nerve fibers. Their response is to many of the same geometric features of stimulus as excite type 2 fibers. The large centers are at least 7-10 degrees in diameter and respond to many of the same features as excite types 3 and 4 optic nerve fibers. The responsiveness of small and large center n. isthmi units is very similar to the elements of the ipsilateral visual field projection onto tectum, i.e. the neuropilar units recorded in layers A and 8 of the tectum when the contralateral eye is occluded. These are in strong contrast to those of tectal cells of layer 6 and below, which have large receptive fields, show far less vivacious response, adapt extremely rapidly to repeated stimuli and are hard to describe in terms of characteristic stimuli because they are unresponsive most of the time. We suggest, therefore, that the axons of tecto-isthmic cells are quite active and that their cell bodies, located in layer 6 and below, only fire occasionally on the firing of their axons.     

Tilt discrimination in the mouse

We tested the visual discrimination of mice in a two-choice discrimination box. After reaching criterion in a brightness discrimination the animals were trained on an orientation discrimination (vertical vs horizontal). Training was continued with the discrimination of vertical striations (S+) vs oblique ones (S-) of 10 different angles (80 to 15 degrees). The tilt discrimination for the last 5 days of training we found to be 30.5 (S.E.M. 3.2) degrees in albino and 19.9 (S.E.M. 0.7) degrees in pigmented mice. This is much higher than the 9.55 (S.E.M. 0.95) degrees found in pigmented rabbits but is comparable to the 17 degrees found in hooded rats by others.     

Comparisons of visual properties between tectal and thalamic neurons with overlapping receptive fields in the pigeon

The present study is the first attempt to make comparisons of the visual response properties between tectal and thalamic neurons with spatially overlapping receptive fields by using extracellular recording and computer mapping techniques. The results show that in neuronal pairs about 70% of thalamic cells have excitatory receptive field alone, whereas 85% of tectal cells possess an excitatory receptive field surrounded by an inhibitory receptive field. In 70% of pairs the tectal cells are selective for direction of motion different from that which the thalamic cells prefer. Most thalamic cells prefer high speeds (80-160 degrees/s), whereas tectal cells prefer intermediate (40 degrees/s) or low (10-20 degrees/s) speeds. Photergic and scotergic cells exist in the thalamus but not in the tectum. These results provide evidence that tectal and thalamic cells extract different visual information from the same region of the visual field. The functional significance of these differences is discussed.     

Direction discrimination of moving gratings and plaids and coherence in dot displays without primary visual cortex (V1)

We present new experimental observations of G.Y., a well-tested patient with unilateral loss of primary visual cortex. We stimulated G.Y.'s blind hemifield using first- and second-order motion stimuli at velocities around psychophysical threshold. Using a dual response paradigm (awareness level of visual motion, motion direction discrimination) psychophysical performance improved with increasing velocity and dot coherence. We were also able to influence directly G.Y.'s performance for the better and at will, by placing the emphasis solely on direction discrimination. In the absence of V1, graduated detection and discrimination of stimuli known to activate both V1 and extrastriate motion areas MT/V5 and MST is still possible. These results are in line with residual visual processing but did not show evidence of unconscious processing of motion stimuli characteristic of ‘blindsight’.     

Lesions of the dorsal striatum impair orienting behaviour of salamanders without affecting visual processing in the tectum

In amphibians, visual information in the midbrain tectum is relayed via the thalamus to telencephalic centres. Lesions of the dorsal thalamus of the salamander Plethodon shermani result in impairment of orienting behaviour and in modulation of spike pattern of tectal neurons. These effects may be induced by an interruption of a tectum‐thalamus‐telencephalon‐tectum feedback loop enabling spatial attention and selection of visual objects. The striatum is a potential candidate for involvement in this pathway; accordingly, we investigated the effects of lesioning the dorsal striatum. Compared to controls and sham lesioned salamanders, striatum‐lesioned animals exhibited a significantly lower number of orienting responses toward one of two competing prey stimuli. Orienting towards stimuli was impaired, while the spike pattern of tectal cells was unaffected, because both in controls and striatum‐lesioned salamanders the spike number significantly decreased at presentation of one prey stimulus inside the excitatory receptive field and another one in the surround compared to that at single presentation inside the excitatory receptive field. We conclude that the dorsal striatum contributes to orienting behaviour, but not to an inhibitory feedback signal onto tectal neurons. The brain area engaged in the feedback loop during visual object discrimination and selection has yet to be identified. Information processing in the amphibian striatum includes multisensory integration; the striatum generates behavioural patterns that influence (pre)motor processing in the brainstem. This situation resembles the situation found in rats, in which the dorsolateral striatum is involved in stimulus‐response learning regardless of the sensory modality, as well as in habit formation.     

Receptive fields of single cells and topography in mouse visual cortex

The visual cortex was studied in the mouse (C57 Black/6J strain) by recording from single units, and a topographic map of the visual field was constructed. Forty-five percent of the neurons in striate cortex responded best to oriented line stimuli moving over their receptive fields; they were classified as simple (17%), complex (25%) and hypercomplex (3%). Of all preferred orientations horizontal was most common. Fifty-five percent of receptive fields were cicularly symmetric: these were on-center (25%), off-center (7%) and homogeneous on-off in type (23%). Optimal stimulus velocities were much higher than those reported in the cat, mostly varying between 20° and 300°/sec. The field of vision common to the two eyes projected to more than one-third of the striate cortex. Although the contralateral eye provided the dominating influence on cells in this binocular area, more than two-thirds of cells could also be driven through the ipsilateral eye. The topography of area 17 was similar to that found in other mammals: the upper visual field projected posteriorly, the most nasal part mapped onto the lateral border. Here the projection did not end at the vertical meridian passing through the animal's long axis, but proceeded for at least 10° into the ipsilateral hemifield of vision, so that at least 20° of visual field were represented in both hemispheres. The magnification in area 17 was rather uniform throughout the visual field. In an area latral to area 17 (18a) the fields were projected in condensed mirror image fashion with respect to the arrangement of area 17. Medial to area 17 a third visual area (area 18) was again related to 17 as a condensed mirror image.     

The optic tectum of the dogfish Scyliorhinus canicula L.: a Golgi study.

The optic tectum of the dogfish Scyliorhinus canicula L. was studied by using the methods of Nissl, reduced silver nitrate, Golgi-aldehyde, and Golgi-Cox. Six layers and eight types of neurons were recognized. These cell types are not restricted to one layer; in fact some are found in all six tectal layers. The types of cells found are A) monopolar, B) triangular, C) radial bipolar, D) horizontal fusiform, E) large tectal, F) small tectal, G) pyriform, and H) stellate cells. In at least six of the cell types a series of dendritic specializations can be observed, such as spines in the form of "drumsticks" and thin varicose appendages, similar to those reported previously in the optic tecta of amphibians and teleosts. The optic tectum of the dogfish shows a degree of complexity comparable to that of amphibians and teleosts.     

The effects of collinearity and orientation on texture visual evoked potentials.

OBJECTIVE: The aim of the research was to study the effects of stimulus orientation at both the local (textons) and the global (segregated elements) level on texture visual evoked potentials (tVEPs). METHODS: Two tVEP paradigms were presented to 10 volunteers. The paradigms were characterized by alternating uniform textures (random mixture of square dots and lines) and textures in which stripes of randomly disposed lines segregated from a square dots' background. In one paradigm, the stripes were horizontal and in the other, vertical. The lines could be either horizontal or vertical in single stimuli of both paradigms. Thus, two stimuli with local/global collinearity and two stimuli without local/global collinearity were available. tVEPs were derived from Oz referenced to the left earlobe and averaged separately for each condition. Segregation-related components were obtained subtracting the traces without segregation from the traces with segregation. RESULTS: A negative segregation component starting at the latency of P1 and extending until the end of N2 characterized the tVEPs, without significant differences among the 4 stimulus conditions. In the presence of local/global collinearity, we found an early modulation of N1 amplitude. This modulation was orientation-dependent, as vertical collinearity increased N1 negativity and horizontal collinearity reduced N1 negativity. CONCLUSIONS: Our experiment confirms previous findings about the segregation negativity, which may depend on contextual modulation of V1 neurons by long-range horizontal and feed-back connections. The early effect of collinearity may depend on more local modulatory connections.     

The accessory optic system of the monocularly deprived cat

Single-unit extracellular recordings were made in the lateral (LTN) and dorsal (DTN) terminal nuclei of the accessory optic system (AOS) of 10 monocularly deprived cats. The separate effects of monocular deprivation (MD) observed in each hemisphere are outlined below. Unlike many units in normal animals, LTN and DTN cells in the hemisphere contralateral to the non-deprived (open) eye, were no longer activated through visual stimulation of the ipsilateral (deprived) eye. In both nuclei, cells were driven effectively only by stimuli presented via the contralateral eye. The distribution of preferred directions was considerably altered in the LTN but not in the DTN. Almost every LTN unit encountered in MD cats preferred downward stimulus motion, in contrast to normal animals where equal numbers of LTN cells show preferences for upward and downward movement (J. Neurophysiol., 51 (1984) 276-293). DTN units showed the usual preference for horizontal motion toward the recorded hemisphere. Velocity preferences were slower on average in the DTN, and unaffected in the LTN. In the hemisphere contralateral to the deprived eye, the ocular dominance distribution of LTN and DTN cells showed a distinct shift in favor of the contralateral (deprived) eye. This effect was not as complete as that observed in the other hemisphere. Cells in both nuclei displayed a small influence from the ipsilateral (exposed) eye in some animals, but this input was much less than that observed in normally reared cats. Average velocity preferences among DTN units were slower than normal, and slower relative to the DTN population in the opposite hemisphere. No pronounced changes were observed in LTN velocity tuning. The distributions of preferred directions for both nuclei were similar to those obtained in the other hemisphere: DTN cells were found to prefer horizontal motion, while most LTN units were activated best by stimuli moving vertically and down within their receptive fields.     

Visual discrimination in the absence of visual cortex

Normal rats and rats with devascularization lesions ranging from subtotal removals of striate cortex (Area 17) to complete removal of neocortex were trained in a horizontal/vertical stripe discrimination for a liquid reinforcer. Subgroups of animals were identified on the basis of size and location of lesion (with particular reference to striate cortex) as Subtotal, Striate, Posterior and Decorticate. Some animals in all of the lesion groups were able to acquire the discrimination, but there was a direct relationship between lesion size and number of training trials. Those animals which reached criterion on the original discrimination were trained on a second horizontal/vertical discrimination under either transfer or reversal conditions using 'rotated obliques' stimuli. Performance on this second discrimination indicated that animals from all lesion groups had been using visual stimuli based on stripe orientation in the original problem. Members of all lesion groups solved the rotated obliques problem under the transfer condition, though the speed and completeness with which they did so was again inversely related to lesion size. These data show high levels of visual competence in the absence of visual cortex even when stimuli thought to detect form discrimination are used and thus reinforce the view that superior colliculus may be a more significant visual area for the rat than was previously assumed. They also support other observations that animals do not use residual visual capacities without extensive experience and appropriate motivation.     

An investigation of the relationship between free-viewing perceptual asymmetries for vertical and horizontal stimuli

Two experiments examine the relationship between free-viewing vertical and horizontal perceptual biases. In Experiment 1, normal participants (n=24) made forced-choice luminance judgments on two mirror-reversed luminance gradients (the 'grayscales' task). The stimuli were presented in vertical, horizontal and oblique (+/-45 degrees ) orientations. Leftward and upward biases were observed in the horizontal and vertical conditions, respectively. In the oblique conditions, leftward and upward biases combined to produce a strong shift of attention away from the lower/right space toward the upper/left. Regression analyses revealed that the oblique biases were the combined product of the vertical and horizontal biases. A lack of correlation between the vertical and horizontal biases, however, suggests they reflect the operation of independent cognitive/neural mechanisms. In Experiment 2, the same stimuli were given to right-hemisphere-lesioned patients with spatial neglect (n=4). Rightward and upward biases were observed for horizontal and vertical stimuli, respectively. The biases combined to produce a strong shift of attention away from the lower/left space toward the upper/right. While our research demonstrates that vertical and horizontal attentional biases are additive, it also appears that they reflect the operation of independent cognitive/neural mechanisms. Potential applications of these findings to the remediation of spatial neglect are discussed.