Position- and scale-invariant object-centered spatial localization in monkey frontoparietal cortex dynamically adapts to cognitive demand.

Egocentric encoding is a well-known property of brain areas along the dorsal pathway. Different to previous experiments, which typically only demanded egocentric spatial processing during movement preparation, we designed a task where two male rhesus monkeys memorized an on-the-object target position and then planned a reach to this position after the object re-occurred at variable location with potentially different size. We found allocentric (in addition to egocentric) encoding in the dorsal stream reach planning areas, parietal reach region and dorsal premotor cortex, which is invariant with respect to the position, and, remarkably, also the size of the object. The dynamic adjustment from predominantly allocentric encoding during visual memory to predominantly egocentric during reach planning in the same brain areas and often the same neurons, suggests that the prevailing frame of reference is less a question of brain area or processing stream, but more of the cognitive demands.

Task demand mediates the interaction of spatial and temporal attention.

Psychophysical studies typically test attentional mechanisms in isolation, but in everyday life they interact to optimize human behavior. We investigated whether spatial and temporal attention interact in two orientation discrimination experiments that vary in task demand. We manipulated temporal and spatial attention separately and conjointly with well-established methods for testing each spatial or temporal attention. We assessed sensitivity (d') and reaction time for every combination of spatial and timing cues, each of which was valid, neutral, or invalid. Spatial attention modulated sensitivity (d') and speed (reaction time) across temporal attention conditions. Temporal attention modulated sensitivity and speed under high- but not low- task demands. Furthermore, spatial and temporal attention interacted for the high-demand task. This study reveals that task demand matters; in a simple task spatial attention suffices to improve performance, whereas in a more demanding task both spatial and temporal attention interact to boost performance, albeit in a subadditive fashion.

Toward a theory of perspective perception in pictures.

This paper reviews projection models and their perception in realistic pictures, and proposes hypotheses for three-dimensional (3D) shape and space perception in pictures. In these hypotheses, eye fixations, and foveal vision play a central role. Many past theories and experimental studies focus solely on linear perspective. Yet, these theories fail to explain many important perceptual phenomena, including the effectiveness of nonlinear projections. Indeed, few classical paintings strictly obey linear perspective, nor do the best distortion-avoidance techniques for wide-angle computational photography. The hypotheses here employ a two-stage model for 3D human vision. When viewing a picture, the first stage perceives 3D shape for the current gaze. Each fixation has its own perspective projection, but, owing to the nature of foveal and peripheral vision, shape information is obtained primarily for a small region of the picture around the fixation. As a viewer moves their eyes, the second stage continually integrates some of the per-gaze information into an overall interpretation of a picture. The interpretation need not be geometrically stable or consistent over time. It is argued that this framework could explain many disparate pictorial phenomena, including different projection styles throughout art history and computational photography, while being consistent with the constraints of human 3D vision. The paper reviews open questions and suggests new studies to explore these hypotheses.

Semantic partitioning facilitates memory for object location through category-partition cueing.

In our lived environments, objects are often semantically organised (e.g., cookware and cutlery are placed close together in the kitchen). Across four experiments, we examined how semantic partitions (that group same-category objects in space) influenced memory for object locations. Participants learned the locations of items in a semantically partitioned display (where each partition contained objects from a single category) as well as a purely visually partitioned display (where each partition contained a scrambled assortment of objects from different categories). Semantic partitions significantly improved location memory accuracy compared to the scrambled display. However, when the correct partition was cued (highlighted) to participants during recall, performance on the semantically partitioned display was similar to the scrambled display. These results suggest that semantic partitions largely benefit memory for location by enhancing the ability to use the given category as a cue for a visually partitioned area (e.g., toys - top left). Our results demonstrate that semantically structured spaces help location memory across partitions, but not items within a partition, providing new insights into the interaction between meaning and memory.

Conjunctive encoding of exploratory intentions and spatial information in the hippocampus.

The hippocampus creates a cognitive map of the external environment by encoding spatial and self-motion-related information. However, it is unclear whether hippocampal neurons could also incorporate internal cognitive states reflecting an animal's exploratory intention, which is not driven by rewards or unexpected sensory stimuli. In this study, a subgroup of CA1 neurons was found to encode both spatial information and animals' investigatory intentions in male mice. These neurons became active before the initiation of exploration behaviors at specific locations and were nearly silent when the same fields were traversed without exploration. Interestingly, this neuronal activity could not be explained by object features, rewards, or mismatches in environmental cues. Inhibition of the lateral entorhinal cortex decreased the activity of these cells during exploration. Our findings demonstrate that hippocampal neurons may bridge external and internal signals, indicating a potential connection between spatial representation and intentional states in the construction of internal navigation systems.

Vestibulospinal reflexes elicited with a tone burst method are dependent on spatial orientation.

Balance involves several sensory modalities including vision, proprioception and the vestibular system. This study aims to investigate vestibulospinal activation elicited by tone burst stimulation in various muscles and how head position influences these responses. We recorded electromyogram (EMG) responses in different muscles (sternocleidomastoid-SCM, cervical erector spinae-ES-C, lumbar erector spinae-ES-L, gastrocnemius-G, and tibialis anterior-TA) of healthy participants using tone burst stimulation applied to the vestibular system. We also evaluated how head position affected the responses. Tone burst stimulation elicited reproducible vestibulospinal reflexes in the SCM and ES-C muscles, while responses in the distal muscles (ES-L, G, and TA) were less consistent among participants. The magnitude and polarity of the responses were influenced by the head position relative to the cervical spine. When the head was rotated or tilted, the polarity of the vestibulospinal responses changed, indicating the integration of vestibular and proprioceptive inputs in generating these reflexes. Overall, our study provides valuable insights into the complexity of vestibulospinal reflexes and their modulation by head position. However, the high variability in responses in some muscles limits their clinical application. These findings may have implications for future research in understanding vestibular function and its role in posture and movement control.

The generative neural microdynamics of cognitive processing.

The entorhinal cortex and hippocampus form a recurrent network that informs many cognitive processes, including memory, planning, navigation, and imagination. Neural recordings from these regions reveal spatially organized population codes corresponding to external environments and abstract spaces. Aligning the former cognitive functionalities with the latter neural phenomena is a central challenge in understanding the entorhinal-hippocampal circuit (EHC). Disparate experiments demonstrate a surprising level of complexity and apparent disorder in the intricate spatiotemporal dynamics of sequential non-local hippocampal reactivations, which occur particularly, though not exclusively, during immobile pauses and rest. We review these phenomena with a particular focus on their apparent lack of physical simulative realism. These observations are then integrated within a theoretical framework and proposed neural circuit mechanisms that normatively characterize this neural complexity by conceiving different regimes of hippocampal microdynamics as neuromarkers of diverse cognitive computations.

How three-dimensional sketching environments affect spatial thinking: A functional magnetic resonance imaging study of virtual reality.

Designers rely on sketching to visualize and refine their initial ideas, and virtual reality (VR) tools now facilitate sketching in immersive 3D environments. However, little research has been conducted on the differences in the visual and spatial processes involved in 3D versus 2D sketching and their effects on cognition. This study investigated potential differences in spatial and visual functions related to the use of 3D versus 2D sketching media by analyzing functional magnetic resonance imaging (fMRI) data. We recruited 20 healthy, right-handed students from the Department of Horticulture and Landscape Architecture with at least three years of experience in freehand landscape drawing. Using an Oculus Quest VR headset controller and a 12.9-inch iPad Pro with an Apple Pencil, we tested participants individually with 3D and 2D sketching, respectively. When comparing 2D and 3D sketches, our fMRI results revealed significant differences in the activation of several brain regions, including the right middle temporal gyrus, both sides of the parietal lobe, and the left middle occipital gyrus. We also compared different sketching conditions, such as lines, geometrical objects (cube), and naturalistic objects (perspective view of a tree), and found significant differences in the spatial and visual recognition of brain areas that support visual recognition, composition, and spatial perception. This finding suggests that 3D sketching environments, such as VR, may activate more visual-spatial functions during sketching compared to 2D environments. The result highlights the potential of immersive sketching environments for design-related processes and spatial thinking.

Disengagement of attention with spatial neglect: A systematic review of behavioral and anatomical findings.

The present review examined the consequences of focal brain injury on spatial attention studied with cueing paradigms, with a particular focus on the disengagement deficit, which refers to the abnormal slowing of reactions following an ipsilesional cue. Our review supports the established notion that the disengagement deficit is a functional marker of spatial neglect and is particularly pronounced when elicited by peripheral cues. Recent research has revealed that this deficit critically depends on cues that have task-relevant characteristics or are associated with negative reinforcement. Attentional capture by task-relevant cues is contingent on damage to the right temporo-parietal junction (TPJ) and is modulated by functional connections between the TPJ and the right insular cortex. Furthermore, damage to the dorsal premotor or prefrontal cortex (dPMC/dPFC) reduces the effect of task-relevant cues. These findings support an interactive model of the disengagement deficit, involving the right TPJ, the insula, and the dPMC/dPFC. These interconnected regions play a crucial role in regulating and adapting spatial attention to changing intrinsic values of stimuli in the environment.

Task-anchored grid cell firing is selectively associated with successful path integration-dependent behaviour.

Grid firing fields have been proposed as a neural substrate for spatial localisation in general or for path integration in particular. To distinguish these possibilities, we investigate firing of grid and non-grid cells in the mouse medial entorhinal cortex during a location memory task. We find that grid firing can either be anchored to the task environment, or can encode distance travelled independently of the task reference frame. Anchoring varied between and within sessions, while spatial firing of non-grid cells was either coherent with the grid population, or was stably anchored to the task environment. We took advantage of the variability in task-anchoring to evaluate whether and when encoding of location by grid cells might contribute to behaviour. We find that when reward location is indicated by a visual cue, performance is similar regardless of whether grid cells are task-anchored or not, arguing against a role for grid representations when location cues are available. By contrast, in the absence of the visual cue, performance was enhanced when grid cells were anchored to the task environment. Our results suggest that anchoring of grid cells to task reference frames selectively enhances performance when path integration is required.

Child development and the role of visual experience in the use of spatial and non-spatial features in haptic object perception.

Previous work has suggested a different developmental timeline and role of visual experience for the use of spatial and non-spatial features in haptic object recognition. To investigate this conjecture, we used a haptic ambiguous odd-one-out task in which one object needed to be selected as being different from two other objects. The odd-one-out could be selected based on four characteristics: size, shape (spatial), texture, and weight (non-spatial). We tested sighted children from 4 to 12 years of age; congenitally blind, late blind, and adult participants with low vision; and normally sighted adults. Given the protracted developmental time course for spatial perception, we expected a shift from a preference for non-spatial features toward spatial features during typical development. Due to the dominant influence of vision for spatial perception, we expected congenitally blind adults to show a similar preference for non-spatial features as the youngest children. The results confirmed our first hypothesis; the 4-year-olds demonstrated a lower dominance for spatial features for object classification compared with older children and sighted adults. In contrast, our second hypothesis was not confirmed; congenitally blind adults' preferred categorization criteria were indistinguishable from those of sighted controls. These findings suggest an early development, but late maturation, of spatial processing in haptic object recognition independent of visual experience.

Repetition Suppression Reveals Cue-Specific Spatial Representations for Landmarks and Self-Motion Cues in the Human Retrosplenial Cortex.

The efficient use of various spatial cues within a setting is crucial for successful navigation. Two fundamental forms of spatial navigation, landmark-based and self-motion-based, engage distinct cognitive mechanisms. The question of whether these modes invoke shared or separate spatial representations in the brain remains unresolved. While nonhuman animal studies have yielded inconsistent results, human investigation is limited. In our previous work (Chen et al., 2019), we introduced a novel spatial navigation paradigm utilizing ultra-high field fMRI to explore neural coding of positional information. We found that different entorhinal subregions in the right hemisphere encode positional information for landmarks and self-motion cues. The present study tested the generalizability of our previous finding with a modified navigation paradigm. Although we did not replicate our previous finding in the entorhinal cortex, we identified adaptation-based allocentric positional codes for both cue types in the retrosplenial cortex (RSC), which were not confounded by the path to the spatial location. Crucially, the multi-voxel patterns of these spatial codes differed between the cue types, suggesting cue-specific positional coding. The parahippocampal cortex exhibited positional coding for self-motion cues, which was not dissociable from path length. Finally, the brain regions involved in successful navigation differed from our previous study, indicating overall distinct neural mechanisms recruited in our two studies. Taken together, the current findings demonstrate cue-specific allocentric positional coding in the human RSC in the same navigation task for the first time and that spatial representations in the brain are contingent on specific experimental conditions.

Modeling the Impact of Head-Body Rotations on Audio-Visual Spatial Perception for Virtual Reality Applications.

Humans perceive the world by integrating multimodal sensory feedback, including visual and auditory stimuli, which holds true in virtual reality (VR) environments. Proper synchronization of these stimuli is crucial for perceiving a coherent and immersive VR experience. In this work, we focus on the interplay between audio and vision during localization tasks involving natural head-body rotations. We explore the impact of audio-visual offsets and rotation velocities on users' directional localization acuity for various viewing modes. Using psychometric functions, we model perceptual disparities between visual and auditory cues and determine offset detection thresholds. Our findings reveal that target localization accuracy is affected by perceptual audio-visual disparities during head-body rotations, but remains consistent in the absence of stimuli-head relative motion. We then showcase the effectiveness of our approach in predicting and enhancing users' localization accuracy within realistic VR gaming applications. To provide additional support for our findings, we implement a natural VR game wherein we apply a compensatory audio-visual offset derived from our measured psychometric functions. As a result, we demonstrate a substantial improvement of up to 40% in participants' target localization accuracy. We additionally provide guidelines for content creation to ensure coherent and seamless VR experiences.

The line bisection bias as a deficit of proportional reasoning - evidence from number line estimation in neglect.

This study aimed to investigate whether neurological patients presenting with a bias in line bisection show specific problems in bisecting a line into two equal parts or their line bisection bias rather reflects a special case of a deficit in proportional reasoning more generally. In the latter case, the bias should also be observed for segmentations into thirds or quarters. To address this question, six neglect patients with a line bisection bias were administered additional tasks involving horizontal lines (e.g., segmentation into thirds and quarters, number line estimation, etc.). Their performance was compared to five neglect patients without a line bisection bias, 10 patients with right hemispheric lesions without neglect, and 32 healthy controls. Most interestingly, results indicated that neglect patients with a line bisection bias also overestimated segments on the left of the line (e.g., one third, one quarter) when dissecting lines into parts smaller than halves. In contrast, such segmentation biases were more nuanced when the required line segmentation was framed as a number line estimation task with either fractions or whole numbers. Taken together, this suggests a generalization of line bisection bias towards a segmentation or proportional processing bias, which is congruent with attentional weighting accounts of line bisection/neglect. As such, patients with a line bisection bias do not seem to have specific problems bisecting a line, but seem to suffer from a more general deficit processing proportions.

Individual differences in emerging adults' spatial abilities: What role do affective factors play?

Spatial ability is defined as a cognitive or intellectual skill used to represent, transform, generate, and recall information of an object or the environment. Individual differences across spatial tasks have been strongly linked to science, technology, engineering, and mathematics (STEM) interest and success. Several variables have been proposed to explain individual differences in spatial ability, including affective factors such as one's confidence and anxiety. However, research is lacking on whether affective variables such as confidence and anxiety relate to individual differences in both a mental rotation task (MRT) and a perspective-taking and spatial orientation task (PTSOT). Using a sample of 100 college students completing introductory STEM courses, the present study investigated the effects of self-reported spatial confidence, spatial anxiety, and general anxiety on MRT and PTSOT. Spatial confidence, after controlling for effects of general anxiety and biological sex, was significantly related to performance on both the MRT and PTSOT. Spatial anxiety, after controlling for effects of general anxiety and biological sex, was not related to either PTSOT or MRT scores. Together these findings suggest some affective factors, but not others, contribute to spatial ability performance to a degree that merits advanced investigation in future studies.

Multisensory peripersonal space: Visual looming stimuli induce stronger response facilitation to tactile than auditory and visual stimulations.

Anticipating physical contact with objects in the environment is a key component of efficient motor performance. Peripersonal neurons are thought to play a determinant role in these predictions by enhancing responses to touch when combined with visual stimuli in peripersonal space (PPS). However, recent research challenges the idea that this visuo-tactile integration contributing to the prediction of tactile events occurs strictly in PPS. We hypothesised that enhanced sensory sensitivity in a multisensory context involves not only contact anticipation but also heightened attention towards near-body visual stimuli. To test this hypothesis, Experiment 1 required participants to respond promptly to tactile (probing contact anticipation) and auditory (probing enhanced attention) stimulations presented at different moments of the trajectory of a (social and non-social) looming visual stimulus. Reduction in reaction time as compared to a unisensory baseline was observed from an egocentric distance of around 2 m (inside and outside PPS) for all multisensory conditions and types of visual stimuli. Experiment 2 tested whether these facilitation effects still occur in the absence of a multisensory context, i.e., in a visuo-visual condition. Overall, facilitation effects induced by the looming visual stimulus were comparable in the three sensory modalities outside PPS but were more pronounced for the tactile modality inside PPS (84 cm from the body as estimated by a reachability judgement task). Considered together, the results suggest that facilitation effects induced by visual looming stimuli in multimodal sensory processing rely on the combination of attentional factors and contact anticipation depending on their distance from the body.

Hierarchical processing of feature, egocentric and relational information for spatial orientation in domestic chicks.

Animals can use different types of information for navigation. Domestic chicks (Gallus gallus) prefer to use local features as a beacon over spatial relational information. However, the role of egocentric navigation strategies is less understood. Here, we tested domestic chicks' egocentric and allocentric orientation abilities in a large circular arena. In experiment 1, we investigated whether domestic chicks possess a side bias during viewpoint-dependent egocentric orientation, revealing facilitation for targets on the chicks' left side. Experiment 2 showed that local features are preferred over viewpoint-dependent egocentric information when the two conflict. Lastly, in experiment 3, we found that in a situation where there is a choice between egocentric and allocentric spatial relational information provided by free-standing objects, chicks preferentially rely on egocentric information. We conclude that chicks orient according to a hierarchy of cues, in which the use of the visual appearance of an object is the dominant strategy, followed by viewpoint-dependent egocentric information and finally by spatial relational information.

Environment geometry alters subiculum boundary vector cell receptive fields in adulthood and early development.

Boundaries to movement form a specific class of landmark information used for navigation: Boundary Vector Cells (BVCs) are neurons which encode an animal's location as a vector displacement from boundaries. Here we characterise the prevalence and spatial tuning of subiculum BVCs in adult and developing male rats, and investigate the relationship between BVC spatial firing and boundary geometry. BVC directional tunings align with environment walls in squares, but are uniformly distributed in circles, demonstrating that environmental geometry alters BVC receptive fields. Inserted barriers uncover both excitatory and inhibitory components to BVC receptive fields, demonstrating that inhibitory inputs contribute to BVC field formation. During post-natal development, subiculum BVCs mature slowly, contrasting with the earlier maturation of boundary-responsive cells in upstream Entorhinal Cortex. However, Subiculum and Entorhinal BVC receptive fields are altered by boundary geometry as early as tested, suggesting this is an inherent feature of the hippocampal representation of space.

Manipulating objects during learning shrinks the global scale of spatial representations in memory: a virtual reality study.

Goal-directed approaches to perception usually consider that distance perception is shaped by the body and its potential for interaction. Although this phenomenon has been extensively investigated in the field of perception, little is known about the effect of motor interactions on memory, and how they shape the global representation of large-scale spaces. To investigate this question, we designed an immersive virtual reality environment in which participants had to learn the positions of several items. Half of the participants had to physically (but virtually) grab the items with their hand and drop them at specified locations (active condition). The other half of the participants were simply shown the items which appeared at the specified position without interacting with them (passive condition). Half of the items used during learning were images of manipulable objects, and the other half were non manipulable objects. Participants were subsequently asked to draw a map of the virtual environment from memory, and to position all the items in it. Results show that active participants recalled the global shape of the spatial layout less precisely, and made more absolute distance errors than passive participants. Moreover, global scaling compression bias was higher for active participants than for passive participants. Interestingly, manipulable items showed a greater compression bias compared to non-manipulable items, yet they had no effect on correlation scores and absolute non-directional distance errors. These results are discussed according to grounded approaches of spatial cognition, emphasizing motor simulation as a possible mechanism for position retrieval from memory.

Assessing Human Spatial Navigation in a Virtual Space and its Sensitivity to Exercise.

Spatial navigation (SN) is the ability to locomote through the environment, which requires an understanding of where one is located in time and space. This capacity is known to rely on the sequential firing of place cells within the hippocampus. SN is an important behavior to investigate as this process deteriorates with age, especially in neurodegenerative disorders. However, the investigation of SN is limited by the lack of sophisticated behavioral techniques to assess this hippocampal-dependent task. Therefore, the goal of this protocol was to develop a novel, real-world approach to studying SN in humans. Specifically, an active virtual SN task was developed using a cross-platform game engine. During the encoding phase, participants navigated their way through a virtual city to locate landmarks. During the remembering phase, participants remembered where these reward locations were and delivered items to these locations. Time to find each location was captured and episodic memory was assessed by a free recall phase, including aspects of place, order, item, and association. Movement behavior (x, y, and z coordinates) was assessed through an asset available in the game engine. Importantly, results from this task demonstrate that it accurately captures both spatial learning and memory abilities as well as episodic memory. Further, findings indicate that this task is sensitive to exercise, which improves hippocampal functioning. Overall, the findings suggest a novel way to track human hippocampal functioning over the course of time, with this behavior being sensitive to physical activity training paradigms.