Unreliable association between self-reported sense of direction and peripheral vestibular function.

BACKGROUND: Uni- or bilateral peripheralvestibular impairment causes objective spatial orientation deficits, which can be measured using pen-and-paper-tests or sensorimotor tasks (navigation or pointing). For patients' subjective orientation abilities, questionnaires are commonly used (e.g., Santa Barbara sense of direction scale [SBSODS]). However, the relationship between subjective assessment of spatial skills and objective vestibular function has only been scarcely investigated. METHODS: A total of 177 patients (mean age 57.86 ± 17.53 years, 90 females) who presented in our tertiary Center for Vertigo and Balance Disorders underwent neuro-otological examinations, including bithermal water calorics, video head impulse test (vHIT), and testing of the subjective visual vertical (SVV), and filled out the SBSODS (German version). Correlation analyses and linear multiple regression model analyses were performed between vestibular test results and self-assessment scores. Additionally, groupwise vestibular function for patients with low, average, and high self-report scores was analyzed. RESULTS: Forty-two patients fulfilled the diagnostic criteria for bilateral vestibulopathy, 93 for chronic unilateral vestibulopathy (68 unilateral caloric hypofunction and 25 isolated horizontal vestibulo-ocular reflex deficits), and 42 patients had normal vestibular test results. SBSODS scores showed clear sex differences with higher subjective skill levels in males (mean score males: 4.94 ± 0.99, females 4.40 ± 0.94; Student's t-test: t-3.78, p < .001***). No stable correlation between objective vestibular function and subjective sense of spatial orientation was found. A multiple linear regression model could not reliably explain the self-reported variance. The three patient groups with low, average, and high self-assessment-scores showed no significant differences of vestibular function. CONCLUSION: Self-reported assessment of spatial orientation does not robustly correlate with objective peripheral vestibular function. Therefore, other methods of measuring spatial skills in real-world and virtual environments are required to disclose orientation deficits due to vestibular hypofunction.

Right visual field advantage in orientation discrimination is influenced by biased suppression.

Visual input is not equally processed over space. In recent years, a right visual field advantage during free walking and standing in orientation discrimination and contrast detection task was reported. The current study investigated the underlying mechanism of the previously reported right visual field advantage. It particularly tested if the advantage is driven by a stronger suppression of distracting input from the left visual field or improved processing of targets from the right visual field. Combing behavioural and electrophysiological measurements in a mobile EEG and augmented reality setup, human participants (n = 30) in a standing and a walking condition performed a line orientation discrimination task with stimulus eccentricity and distractor status being manipulated. The right visual field advantage, as demonstrated in accuracy and reaction time, was influenced by the distractor status. Specifically, the right visual field advantage was only observed when the target had an incongruent line orientation with the distractor. Neural data further showed that the right visual field advantage was paralleled by a strong modulation of neural activity in the right hemisphere (i.e. contralateral to the distractor). A significant positive correlation between this right hemispheric event related potential (ERP) and behavioural measures (accuracy and reaction time) was found exclusively for trials in which a target was presented on the right and an incongruent distractor was presented on the left. The right hemispheric ERP component further predicted the strength of the right visual field advantage. Notably, the lateralised brain activity and the right visual field advantage were both independent of stimulus eccentricity and the movement state of participants. Overall, our findings suggest an important role of spatially biased suppression of left distracting input in the right visual field advantage as found in orientation discrimination.

Encoding of 2D Self-Centered Plans and World-Centered Positions in the Rat Frontal Orienting Field.

The neural mechanisms of motor planning have been extensively studied in rodents. Preparatory activity in the frontal cortex predicts upcoming choice, but limitations of typical tasks have made it challenging to determine whether the spatial information is in a self-centered direction reference frame or a world-centered position reference frame. Here, we trained male rats to make delayed visually guided orienting movements to six different directions, with four different target positions for each direction, which allowed us to disentangle direction versus position tuning in neural activity. We recorded single unit activity from the rat frontal orienting field (FOF) in the secondary motor cortex, a region involved in planning orienting movements. Population analyses revealed that the FOF encodes two separate 2D maps of space. First, a 2D map of the planned and ongoing movement in a self-centered direction reference frame. Second, a 2D map of the animal's current position on the port wall in a world-centered reference frame. Thus, preparatory activity in the FOF represents self-centered upcoming movement directions, but FOF neurons multiplex both self- and world-reference frame variables at the level of single neurons. Neural network model comparison supports the view that despite the presence of world-centered representations, the FOF receives the target information as self-centered input and generates self-centered planning signals.

Estimating orientation in natural scenes: A spiking neural network model of the insect central complex.

The central complex of insects contains cells, organised as a ring attractor, that encode head direction. The 'bump' of activity in the ring can be updated by idiothetic cues and external sensory information. Plasticity at the synapses between these cells and the ring neurons, that are responsible for bringing sensory information into the central complex, has been proposed to form a mapping between visual cues and the heading estimate which allows for more accurate tracking of the current heading, than if only idiothetic information were used. In Drosophila, ring neurons have well characterised non-linear receptive fields. In this work we produce synthetic versions of these visual receptive fields using a combination of excitatory inputs and mutual inhibition between ring neurons. We use these receptive fields to bring visual information into a spiking neural network model of the insect central complex based on the recently published Drosophila connectome. Previous modelling work has focused on how this circuit functions as a ring attractor using the same type of simple visual cues commonly used experimentally. While we initially test the model on these simple stimuli, we then go on to apply the model to complex natural scenes containing multiple conflicting cues. We show that this simple visual filtering provided by the ring neurons is sufficient to form a mapping between heading and visual features and maintain the heading estimate in the absence of angular velocity input. The network is successful at tracking heading even when presented with videos of natural scenes containing conflicting information from environmental changes and translation of the camera.

Orienting of attention and spatial cognition.

Humans orient to their sensory world through foveation of target location or through covert shifts of attention. Orienting provides primacy to the selected location and in humans improves the precision of discrimination. Covert orienting appears to arise separately from the mechanisms involved in saccadic eye movements. Covert orienting can serve to prioritize processing the target even increasing its subjective intensity and its acuity. However, this network does not appear to be involved in the operations related to binding and segmentation. Cells exist in the early visual cortex that are activated by both color and form features without attention, however, color and form appear to remain independent even when oriented to the target that is required to be reported. An understanding of the pathways that connect attention networks to memory networks may allow us to understand more complex aspects of spatial cognition and enhance orienting and thus improve spatial cognition.

Hue and orientation pinwheels in macaque area V4.

Area V4 is an intermediate-level area of the macaque visual cortical hierarchy that serves key functions in visual processing by integrating inputs from lower areas such as V1 and V2 and providing feedforward inputs to many higher cortical areas. Previous V4 imaging studies have focused on differential responses to color, orientation, disparity, and motion stimuli, but many details of the spatial organization of significant hue and orientation tuning have not been fully described. We used support vector machine (SVM) decoding of intrinsic cortical single-condition responses to generate high-resolution maps of hue and orientation tuning and to describe the organization of hue and orientation pinwheels in V4. Like V1 and V2, V4 contains maps of orientation that are organized as pinwheels. V4 also contains maps of hue that are organized as pinwheels, whose circular organization more closely represents the perception of hue than is observed in antecedent cortical areas. Unlike V1, where orientation is continuously mapped across the surface, V4 hue and orientation pinwheels are organized in limited numbers of pinwheel sequences. The organization of these sequences and the distance between pinwheels may provide insight into the functional organization of V4. Regions significantly tuned for hue occupy roughly four times that of the orientation, are largely separated from each other, and overlap by roughly 5%. This spatial organization is largely consistent with segregated inputs arising from V2 thin and interstripes. This modular organization of V4 suggests that further integration of color and shape might occur in higher areas in inferotemporal cortical.NEW & NOTEWORTHY The representation of hue and orientation in macaque monkey area V4 was determined by intrinsic cortical imaging of responses to isoluminant hues and achromatic grating stimuli. Vector summation of support vector machine (SVM) decoded single-condition responses was used to generate hue and orientation maps that, like V1 orientation maps, were both characterized by distinct pinwheel patterns. These data suggest that pinwheels are an important structure to represent different stimulus features across multiple visual cortical areas.

Extension of a computational model of a class of orientation illusions.

Polarity-dependent orientation illusions constitute a class of illusions in which the impression of orientation does not depend only on geometrical relations between its elements, but also on the relations between their luminances. Several examples of such figures are presented in the paper. Todorovic (2021a) presented a simple computational model of such phenomena. Simulations of the model indicated that a common feature of the neural basis of these illusions is the presence of certain neural structures called 'oblique clusters'. A limitation of the model was that it used a restricted set of parameters. In this paper a generalization of the model is introduced involving types of receptive fields, their orientation sensitivity and their size or spatial frequency tuning. The simulations of the new model indicated that oblique clusters were present in the reaction patterns under a much wider set of conditions, though not all. The original hypothesis that oblique clusters constituted the neural foundations of impressions of tilt in this class of illusions was vindicated.

Distinct neural mechanisms for heading retrieval and context recognition in the hippocampus during spatial reorientation.

Reorientation, the process of regaining one's bearings after becoming lost, requires identification of a spatial context (context recognition) and recovery of facing direction within that context (heading retrieval). We previously showed that these processes rely on the use of features and geometry, respectively. Here, we examine reorientation behavior in a task that creates contextual ambiguity over a long timescale to demonstrate that male mice learn to combine both featural and geometric cues to recover heading. At the neural level, most CA1 neurons persistently align to geometry, and this alignment predicts heading behavior. However, a small subset of cells remaps coherently in a context-sensitive manner, which serves to predict context. Efficient heading retrieval and context recognition correlate with rate changes reflecting integration of featural and geometric information in the active ensemble. These data illustrate how context recognition and heading retrieval are coded in CA1 and how these processes change with experience.

Influence of scene aspect ratio and depth cues on verticality perception bias.

Perceiving verticality is crucial for accurate spatial orientation. Previous research has revealed that tilted scenes can bias verticality perception. Verticality perception bias can be represented as the sum of multiple periodic functions that play a role in the perception of visual orientation, where the specific factors affecting each periodicity remain uncertain. This study investigated the influence of the width and depth of an indoor scene on each periodic component of the bias. The participants were presented with an indoor scene showing a rectangular checkerboard room (Experiment 1), a rectangular aperture on the wall (Experiment 2), or a rectangular dotted room (Experiment 3), with various aspect ratios. The stimuli were presented with roll orientations ranging from 90° clockwise to 90° counterclockwise. The participants were asked to report their subjective visual vertical (SVV) perceptions. The contributions of 45°, 90°, and 180° periodicities to the SVV error were assessed by the weighted vector sum model. In Experiment 1, the periodic components of the SVV error increased with the aspect ratio. In Experiments 2 and 3, only the 90° component increased with the aspect ratio. These findings suggest that extended transverse surfaces may modulate the periodic components of verticality perception.

Flock2: A model for orientation-based social flocking.

The aerial flocking of birds, or murmurations, has fascinated observers while presenting many challenges to behavioral study and simulation. We examine how the periphery of murmurations remain well bounded and cohesive. We also investigate agitation waves, which occur when a flock is disturbed, developing a plausible model for how they might emerge spontaneously. To understand these behaviors a new model is presented for orientation-based social flocking. Previous methods model inter-bird dynamics by considering the neighborhood around each bird, and introducing forces for avoidance, alignment, and cohesion as three dimensional vectors that alter acceleration. Our method introduces orientation-based social flocking that treats social influences from neighbors more realistically as a desire to turn, indirectly controlling the heading in an aerodynamic model. While our model can be applied to any flocking social bird we simulate flocks of starlings, Sturnus vulgaris, and demonstrate the possibility of orientation waves in the absence of predators. Our model exhibits spherical and ovoidal flock shapes matching observation. Comparisons of our model to Reynolds' on energy consumption and frequency analysis demonstrates more realistic motions, significantly less energy use in turning, and a plausible mechanism for emergent orientation waves.

Influence of spatial orientation training in a centrifuge on the ability of fighter pilots to assess the bank angle during flight without visual references.

Without visual references, nonpilots exposed to coordinated flight turns underestimate the bank angle, because of discordant information of the roll-angular displacement from the otoliths, consistently signaling vertical position, versus the semicircular canals, enabling detection of the displacement. Pilots may also use their ability to perceive the G load and knowledge of the relation between load and angle to assess the bank angle. Our aim was to investigate whether the perception of bank angle can be improved by spatial orientation training in a centrifuge. Sixteen pilots/pilot students assessed their roll tilt, in complete darkness, during both real coordinated flight turns and gondola centrifugation, at roll tilts of 30° and 60°. The experiments were repeated after a 3-wk period, during which eight of the subjects performed nine training sessions in the centrifuge, comprising feedback on roll angle vs. G load, and on indicating requested angles. Before training, the subjects perceived in the aircraft and centrifuge, respectively: 37 (17)°, 38 (14)° during 60° turns and 19 (12)°, 20 (10)° during 30° turns. Training improved the perception of angle during the 60° [to 60 (7)°, 55 (10)°; P ≤ 0.04] but not the 30° [21 (10)°, 15 (9)°; P ≥ 0.30] turns; the improvement disappeared within 2 yr after training. Angle assessments did not change in the untrained group. The results suggest that it is possible to, in a centrifuge, train a pilot's ability to perceive large but not discrete-to-moderate roll-angular displacements. The transient training effect is attributable to improved capacity to perceive and translate G load into roll angle and/or to increased reliance on semicircular canal signals.NEW & NOTEWORTHY Spatial disorientation is a major problem in aviation. When performing coordinated flight turns without external visual cues (e.g., flying in clouds or darkness), the pilot underestimates the aircraft bank angle because the vestibular system provides unreliable information of roll tilt. The present study demonstrates that it is possible to, in a long-arm centrifuge, train a pilot's ability to perceive large but not discrete-to-moderate roll-angular displacements.

Development and organization of the retinal orientation selectivity map.

Orientation or axial selectivity, the property of neurons in the visual system to respond preferentially to certain angles of visual stimuli, plays a pivotal role in our understanding of visual perception and information processing. This computation is performed as early as the retina, and although much work has established the cellular mechanisms of retinal orientation selectivity, how this computation is organized across the retina is unknown. Using a large dataset collected across the mouse retina, we demonstrate functional organization rules of retinal orientation selectivity. First, we identify three major functional classes of retinal cells that are orientation selective and match previous descriptions. Second, we show that one orientation is predominantly represented in the retina and that this predominant orientation changes as a function of retinal location. Third, we demonstrate that neural activity plays little role on the organization of retinal orientation selectivity. Lastly, we use in silico modeling followed by validation experiments to demonstrate that the overrepresented orientation aligns along concentric axes. These results demonstrate that, similar to direction selectivity, orientation selectivity is organized in a functional map as early as the retina.

Transfer of visual perceptual learning over a task-irrelevant feature through feature-invariant representations: Behavioral experiments and model simulations.

A large body of literature has examined specificity and transfer of perceptual learning, suggesting a complex picture. Here, we distinguish between transfer over variations in a "task-relevant" feature (e.g., transfer of a learned orientation task to a different reference orientation) and transfer over a "task-irrelevant" feature (e.g., transfer of a learned orientation task to a different retinal location or different spatial frequency), and we focus on the mechanism for the latter. Experimentally, we assessed whether learning a judgment of one feature (such as orientation) using one value of an irrelevant feature (e.g., spatial frequency) transfers to another value of the irrelevant feature. Experiment 1 examined whether learning in eight-alternative orientation identification with one or multiple spatial frequencies transfers to stimuli at five different spatial frequencies. Experiment 2 paralleled Experiment 1, examining whether learning in eight-alternative spatial-frequency identification at one or multiple orientations transfers to stimuli with five different orientations. Training the orientation task with a single spatial frequency transferred widely to all other spatial frequencies, with a tendency to specificity when training with the highest spatial frequency. Training the spatial frequency task fully transferred across all orientations. Computationally, we extended the identification integrated reweighting theory (I-IRT) to account for the transfer data (Dosher, Liu, & Lu, 2023; Liu, Dosher, & Lu, 2023). Just as location-invariant representations in the original IRT explain transfer over retinal locations, incorporating feature-invariant representations effectively accounted for the observed transfer. Taken together, we suggest that feature-invariant representations can account for transfer of learning over a "task-irrelevant" feature.

Tactile cues compensate for unbalanced vestibular cues during progression on inclined surfaces.

A previous study demonstrated that rodents on an inclined square platform traveled straight vertically or horizontally and avoided diagonal travel. Through behavior they aligned their head with the horizontal plane, acquiring similar bilateral vestibular cues - a basic requirement for spatial orientation and a salient feature of animals in motion. This behavior had previously been shown to be conspicuous in Tristram's jirds. Here, therefore jirds were challenged by testing their travel behavior on a circular arena inclined at 0°-75°. Our hypothesis was that if, as typical to rodents, the jirds would follow the curved arena wall, they would need to display a compensating mechanism to enable traveling in such a path shape, which involves a tilted frontal head axis and unbalanced bilateral vestibular cues. We found that with the increase in inclination, the jirds remained more in the lower section of the arena (geotaxis). When tested on the steep inclinations, however, their travel away from the arena wall was strictly straight up or down, in contrast to the curved paths that followed the circular arena wall. We suggest that traveling along a circular path while maintaining contact with the wall (thigmotaxis), provided tactile information that compensated for the unbalanced bilateral vestibular cues present when traveling along such curved inclined paths. In the latter case, the frontal plane of the head was in a diagonal posture in relation to gravity, a posture that was avoided when traveling away from the wall.

Albatrosses employ orientation and routing strategies similar to yacht racers.

The way goal-oriented birds adjust their travel direction and route in response to wind significantly affects their travel costs. This is expected to be particularly pronounced in pelagic seabirds, which utilize a wind-dependent flight style called dynamic soaring. Dynamic soaring seabirds in situations without a definite goal, e.g. searching for prey, are known to preferentially fly with crosswinds or quartering-tailwinds to increase the speed and search area, and reduce travel costs. However, little is known about their reaction to wind when heading to a definite goal, such as homing. Homing tracks of wandering albatrosses (Diomedea exulans) vary from beelines to zigzags, which are similar to those of sailboats. Here, given that both albatrosses and sailboats travel slower in headwinds and tailwinds, we tested whether the time-minimizing strategies used by yacht racers can be compared to the locomotion patterns of wandering albatrosses. We predicted that when the goal is located upwind or downwind, albatrosses should deviate their travel directions from the goal on the mesoscale and increase the number of turns on the macroscale. Both hypotheses were supported by track data from albatrosses and racing yachts in the Southern Ocean confirming that albatrosses qualitatively employ the same strategy as yacht racers. Nevertheless, albatrosses did not strictly minimize their travel time, likely making their flight robust against wind fluctuations to reduce flight costs. Our study provides empirical evidence of tacking in albatrosses and demonstrates that man-made movement strategies provide a new perspective on the laws underlying wildlife movement.

Influence of postural control difficulty on changes in spatial orienting of attention after leftward prism adaptation.

Prism adaptation (PA) affects visuospatial attention such as spatial orienting in both the right and left hemifields; however, the systematic after-effects of PA on visuospatial attention remain unclear. Visuospatial attention can be affected by non-spatial attentional factors, and postural control difficulty, which delays the reaction time (RT) to external stimulation, may be one such factor. Therefore, we aimed to investigate the influence of postural control difficulty on changes in spatial orienting of attention after leftward PA. Seventeen healthy young adults underwent 15-min and 5-min PA procedures for a leftward visual shift (30 diopters). Participants underwent the Posner cueing test immediately before (pre-evaluation) and in between and after the PA procedures (post-evaluations) while standing barefoot on the floor (normal standing condition) and on a balance-disc (balance standing condition). In the pre-evaluation, RTs in the balance standing condition were significantly longer compared to those in the normal standing condition for targets appearing in both the right and left hemifields. Leftward PA improved the RT for targets appearing in the right, but no left, hemifield in the balance standing condition, such that RTs for targets in the right hemifield in the post-evaluation were not significantly different between the two standing conditions. However, leftward PA did not significantly change RTs for targets in both hemifields in the normal standing condition. Therefore, postural control difficulty may enhance sensitivity to the features of the visuospatial cognitive after-effects of leftward PA.

Feature variability determines specificity and transfer in multiorientation feature detection learning.

Historically, in many perceptual learning experiments, only a single stimulus is practiced, and learning is often specific to the trained feature. Our prior work has demonstrated that multi-stimulus learning (e.g., training-plus-exposure procedure) has the potential to achieve generalization. Here, we investigated two important characteristics of multi-stimulus learning, namely, roving and feature variability, and their impacts on multi-stimulus learning and generalization. We adopted a feature detection task in which an oddly oriented target bar differed by 16° from the background bars. The stimulus onset asynchrony threshold between the target and the mask was measured with a staircase procedure. Observers were trained with four target orientation search stimuli, either with a 5° deviation (30°-35°-40°-45°) or with a 45° deviation (30°-75°-120°-165°), and the four reference stimuli were presented in a roving manner. The transfer of learning to the swapped target-background orientations was evaluated after training. We found that multi-stimulus training with a 5° deviation resulted in significant learning improvement, but learning failed to transfer to the swapped target-background orientations. In contrast, training with a 45° deviation slowed learning but produced a significant generalization to swapped orientations. Furthermore, a modified training-plus-exposure procedure, in which observers were trained with four orientation search stimuli with a 5° deviation and simultaneously passively exposed to orientations with high feature variability (45° deviation), led to significant orientation learning generalization. Learning transfer also occurred when the four orientation search stimuli with a 5° deviation were presented in separate blocks. These results help us to specify the condition under which multistimuli learning produces generalization, which holds potential for real-world applications of perceptual learning, such as vision rehabilitation and expert training.

Converting an allocentric goal into an egocentric steering signal.

Neuronal signals that are relevant for spatial navigation have been described in many species1-10. However, a circuit-level understanding of how such signals interact to guide navigational behaviour is lacking. Here we characterize a neuronal circuit in the Drosophila central complex that compares internally generated estimates of the heading and goal angles of the fly-both of which are encoded in world-centred (allocentric) coordinates-to generate a body-centred (egocentric) steering signal. Past work has suggested that the activity of EPG neurons represents the fly's moment-to-moment angular orientation, or heading angle, during navigation2,11. An animal's moment-to-moment heading angle, however, is not always aligned with its goal angle-that is, the allocentric direction in which it wishes to progress forward. We describe FC2 cells12, a second set of neurons in the Drosophila brain with activity that correlates with the fly's goal angle. Focal optogenetic activation of FC2 neurons induces flies to orient along experimenter-defined directions as they walk forward. EPG and FC2 neurons connect monosynaptically to a third neuronal class, PFL3 cells12,13. We found that individual PFL3 cells show conjunctive, spike-rate tuning to both the heading angle and the goal angle during goal-directed navigation. Informed by the anatomy and physiology of these three cell classes, we develop a model that explains how this circuit compares allocentric heading and goal angles to build an egocentric steering signal in the PFL3 output terminals. Quantitative analyses and optogenetic manipulations of PFL3 activity support the model. Finally, using a new navigational memory task, we show that flies expressing disruptors of synaptic transmission in subsets of PFL3 cells have a reduced ability to orient along arbitrary goal directions, with an effect size in quantitative accordance with the prediction of our model. The biological circuit described here reveals how two population-level allocentric signals are compared in the brain to produce an egocentric output signal that is appropriate for motor control.

Transforming a head direction signal into a goal-oriented steering command.

To navigate, we must continuously estimate the direction we are headed in, and we must correct deviations from our goal1. Direction estimation is accomplished by ring attractor networks in the head direction system2,3. However, we do not fully understand how the sense of direction is used to guide action. Drosophila connectome analyses4,5 reveal three cell populations (PFL3R, PFL3L and PFL2) that connect the head direction system to the locomotor system. Here we use imaging, electrophysiology and chemogenetic stimulation during navigation to show how these populations function. Each population receives a shifted copy of the head direction vector, such that their three reference frames are shifted approximately 120° relative to each other. Each cell type then compares its own head direction vector with a common goal vector; specifically, it evaluates the congruence of these vectors via a nonlinear transformation. The output of all three cell populations is then combined to generate locomotor commands. PFL3R cells are recruited when the fly is oriented to the left of its goal, and their activity drives rightward turning; the reverse is true for PFL3L. Meanwhile, PFL2 cells increase steering speed, and are recruited when the fly is oriented far from its goal. PFL2 cells adaptively increase the strength of steering as directional error increases, effectively managing the tradeoff between speed and accuracy. Together, our results show how a map of space in the brain can be combined with an internal goal to generate action commands, via a transformation from world-centric coordinates to body-centric coordinates.

Interpreting the orientation of objects: A cross-disciplinary review.

Is object orientation an inherent aspect of the shape of the object or is it represented separately and bound to the object shape in a similar way to other features, such as colour? This review brings together findings from neuropsychological studies of patients with agnosia for object orientation and experimental studies of object perception in healthy individuals that provide converging evidence of separate processing of object identity and orientation. Individuals with agnosia for object orientation, which typically results from damage to the right parietal lobe, can recognize objects presented in a range of orientations yet are unable to interpret or discriminate the objects' orientation. Healthy individuals tested with briefly presented objects demonstrate a similar dissociation: object identity is extracted rapidly in an orientation-invariant way, whereas processing the object's orientation is slower, requires attention and is influenced by the degree of departure from the canonical orientation. This asymmetry in processing can sometimes lead to incorrect bindings between the identity and orientation of objects presented in close temporal proximity. Overall, the available evidence indicates that object recognition is achieved in a largely orientation-invariant manner and that interpreting the object's orientation requires an additional step of mapping this orientation-invariant representation to a spatial reference frame.