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Auditory cues are integrated with vision and body-based self-motion cues for motion perception, balance, and gait, though limited research has evaluated their effectiveness for navigation. Here, we tested whether an auditory cue co-localized with a visual target could improve spatial updating in a virtual reality homing task. Participants navigated a triangular homing task with and without an easily localizable spatial audio signal co-located with the home location. The main outcome was unsigned angular error, defined as the absolute value of the difference between the participant's turning response and the correct response towards the home location. Angular error was significantly reduced in the presence of spatial sound compared to a head-fixed identical auditory signal. Participants' angular error was 22.79° in the presence of spatial audio and 30.09° in its absence. Those with the worst performance in the absence of spatial sound demonstrated the greatest improvement with the added sound cue. These results suggest that auditory cues may benefit navigation, particularly for those who demonstrated the highest level of spatial updating error in the absence of spatial sound.

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Older adults demonstrate impairments in navigation that cannot be explained by general cognitive and motor declines. Previous work has shown that older adults may combine sensory cues during navigation differently than younger adults, though this work has largely been done in dark environments where sensory integration may differ from full-cue environments. Here, we test whether aging adults optimally combine cues from two sensory systems critical for navigation: vision (landmarks) and body-based self-motion cues. Participants completed a homing (triangle completion) task using immersive virtual reality to offer the ability to navigate in a well-lit environment including visibility of the ground plane. An optimal model, based on principles of maximum-likelihood estimation, predicts that precision in homing should increase with multisensory information in a manner consistent with each individual sensory cue's perceived reliability (measured by variability). We found that well-aging adults (with normal or corrected-to-normal sensory acuity and active lifestyles) were more variable and less accurate than younger adults during navigation. Both older and younger adults relied more on their visual systems than a maximum likelihood estimation model would suggest. Overall, younger adults' visual weighting matched the model's predictions whereas older adults showed sub-optimal sensory weighting. In addition, high inter-individual differences were seen in both younger and older adults. These results suggest that older adults do not optimally weight each sensory system when combined during navigation, and that older adults may benefit from interventions that help them recalibrate the combination of visual and self-motion cues for navigation.

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Keeping track of locations across self-motion is possible by continuously updating spatial representations or by encoding and later instantaneously retrieving spatial representations. In virtual reality (VR), sensory cues to self-motion used in continuous updating are typically reduced. In passive translation compared to real walking in VR, optic flow is available but body-based (idiothetic) cues are missing. With both kinds of translation, boundaries and landmarks as static visual cues can be used for instantaneous updating. In two experiments, we let participants encode two target locations, one of which had to be reproduced by pointing after forward translation in immersive VR (HMD). We increased sensory cues to self-motion in comparison to passive translation either by strengthening optic flow or by real walking. Furthermore, we varied static visual cues in the form of boundaries and landmarks inside boundaries. Increased optic flow and real walking did not reliably increase performance suggesting that optic flow even in a sparse environment was sufficient for continuous updating or that merely instantaneous updating took place. Boundaries and landmarks, however, did support performance as quantified by decreased bias and increased precision, particularly if they were close to or even enclosed target locations. Thus, enriched spatial context is a viable method to support spatial updating in VR and synthetic environments (teleoperation). Spatial context does not only provide a static visual reference in offline updating and continuous allocentric self-location updating but also, according to recent neuroscientific evidence on egocentric bearing cells, contributes to continuous egocentric location updating as well.

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Updating spatial representations in visual and auditory working memory relies on common processes, and the modalities should compete for attentional resources. If competition occurs, one type of spatial information is presumably weighted over the other, irrespective of sensory modality. This study used incompatible spatial information conveyed from two different cue modalities to examine relative dominance in memory updating. Participants mentally manoeuvred a designated target in a matrix according to visual or auditory stimuli that were presented simultaneously, to identify a terminal location. Prior to the navigation task, the relative perceptual saliences of the visual cues were manipulated to be equal, superior, or inferior to the auditory cues. The results demonstrate that visual and auditory information competed for attentional resources, such that visual/auditory guidance was impaired by incongruent cues delivered from the other modality. Although visual bias was generally observed in working-memory navigation, stimuli of relatively high salience interfered with or facilitated other stimuli regardless of modality, demonstrating the processing symmetry of spatial updating in visual and auditory spatial working memory. Furthermore, this processing symmetry can be identified during the encoding of sensory inputs into working-memory representations. The results imply that auditory spatial updating is comparable to visual spatial updating in that salient stimuli receive a high priority when selecting inputs and are used when tracking spatial representations.

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As we move through an environment, we update positions of our body relative to other objects, even when some objects temporarily or permanently leave our field of view-this ability is termed egocentric spatial updating and plays an important role in everyday life. Still, our knowledge about its representation in the brain is still scarce, with previous studies using virtual movements in virtual environments or patients with brain lesions suggesting that the precuneus might play an important role. However, whether this assumption is also true when healthy humans move in real environments where full body-based cues are available in addition to the visual cues typically used in many VR studies is unclear. Therefore, in this study we investigated the role of the precuneus in egocentric spatial updating in a real environment setting in 20 healthy young participants who underwent two conditions in a cross-over design: (a) stimulation, achieved through applying continuous theta-burst stimulation (cTBS) to inhibit the precuneus and (b) sham condition (activated coil turned upside down). In both conditions, participants had to walk back with blindfolded eyes to objects they had previously memorized while walking with open eyes. Simplified trials (without spatial updating) were used as control condition, to make sure the participants were not affected by factors such as walking blindfolded, vestibular or working memory deficits. A significant interaction was found, with participants performing better in the sham condition compared to real stimulation, showing smaller errors both in distance and angle. The results of our study reveal evidence of an important role of the precuneus in a real-environment egocentric spatial updating; studies on larger samples are necessary to confirm and further investigate this finding.

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Reaching toward a target viewed through laterally refracting prisms results in adaptation of both visual and (limb) proprioceptive spatial representations. Common ways to measure adaptation after-effect are to ask a person to point straight ahead with their eyes closed ("manual straight ahead", MSA), or to a seen target using their unseen hand ("open-loop pointing", OLP). MSA measures changes in proprioception only, whereas OLP measures the combined visual and proprioceptive shift. The behavioural and neurological mechanisms of prism adaptation have come under scrutiny following reports of reduced hemispatial neglect in patients following this procedure. We present evidence suggesting that shifts in proprioceptive spatial representations induced by prism adaptation are larger following lesions to the intraparietal cortex - a brain region that integrates retinotopic visual signals with signals of eye position in the orbit and that is activated during prism adaptation. Six healthy participants and six patients with unilateral intraparietal cortex lesions underwent prism adaptation. After-effects were measured with OLP and MSA. After-effects of control participants were larger when measured with OLP than with MSA, consistent with previous research and with the additional contribution of visual shift to OLP after-effects. However, patients' OLP shifts were not significantly different to their MSA shifts. We conclude that, for the patients, correction of pointing errors during prism adaptation involved proportionally more changes to arm proprioception than for controls. Since lesions to intraparietal cortex led to enhanced realignment of arm proprioceptive representations, our results indirectly suggest that the intraparietal cortex could be key for visual realignment.

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Sensory cues enable navigation through space, as they inform us about movement properties, such as the amount of travelled distance and the heading direction. In this study, we focused on the ability to spatially update one's position when only proprioceptive and vestibular information is available. We aimed to investigate the effect of yaw rotation on path integration across development in the absence of visual feedback. To this end, we utilized the triangle completion task: participants were guided through two legs of a triangle and asked to close the shape by walking along its third imagined leg. To test the influence of yaw rotation across development, we tested children between 6 and 11 years old (y.o.) and adults on their perceptions of angles of different degrees. Our results demonstrated that the amount of turn while executing the angle influences performance at all ages, and in some aspects, also interacted with age. Indeed, whilst adults seemed to adjust their heading towards the end of their walked path, younger children took less advantage of this strategy. The amount of disorientation the path induced also affected participants' full maturational ability to spatially navigate with no visual feedback. Increasing induced disorientation required children to be older to reach adult-level performance. Overall, these results provide novel insights on the maturation of spatial navigation-related processes.

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Although visual and auditory inputs are initially processed in separate perception systems, studies have built on the idea that to maintain spatial information these modalities share a component of working memory. The present study used working memory navigation tasks to examine functional similarities and dissimilarities in the performance of updating tasks. Participants mentally updated the spatial location of a target in a virtual array in response to sequential pictorial and sonant directional cues before identifying the target's final location. We predicted that if working memory representations are modality-specific, mixed-modality cues would demonstrate a cost of modality switching relative to unimodal cues. The results indicate that updating performance using visual unimodal cues positively correlated with that using auditory unimodal cues. Task performance using unimodal cues was comparable to that using mixed modality cues. The results of a subsequent experiment involving updating of target traces were consistent with those of the preceding experiments and support the view of modality-nonspecific memory.

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The relative contribution of different sources of information for spatial updating - keeping track of one's position in an environment - has been highly debated. Further, children and adults may differ in their reliance on visual versus body-based information for spatial updating. In two experiments, we tested children (age 10-12 years) and young adult participants on a virtual point-to-origin task that varied the types of self-motion information available for translation: full-dynamic (walking), visual-dynamic (controller induced), and no-dynamic (teleporting). In Experiment 1, participants completed the three conditions in an indoor virtual environment with visual landmark cues. Adults were more accurate in the full- and visual-dynamic conditions (which did not differ from each other) compared to the no-dynamic condition. In contrast, children were most accurate in the visual-dynamic condition and also least accurate in the no-dynamic condition. Adults outperformed children in all conditions. In Experiment 2, we removed the potential for relying on visual landmarks by running the same paradigm in an outdoor virtual environment with no geometrical room cues. As expected, adults' errors increased in all conditions, but performance was still relatively worse in teleporting. Surprisingly, children showed overall similar accuracy and patterns across locomotion conditions to adults. Together, the results support the importance of dynamic translation information (either visual or body-based) for spatial updating across both age groups, but suggest children may be more reliant on visual information than adults.

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This study investigated to what extent humans can encode spatial relations between different surfaces (i.e., floor, walls, and ceiling) in a three-dimensional (3D) space and extend their headings on the floor to other surfaces when locomoting to walls (pitch 90°) and the ceiling (pitch 180°). In immersive virtual reality environments, participants first learned a layout of objects on the ground. They then navigated to testing planes: south (or north) walls facing Up, or the ceiling via walls facing North (or South). Participants locomoted to the walls with pitch rotations indicated by visual and idiothetic cues (Experiment 1) and only by visual cues (Experiment 2) and to the ceiling with visual pitch rotations only (Experiment 3). Using the memory of objects' locations, they either reproduced the object layout on the testing plane or did a Judgements of Relative Direction (JRD) task ("imagine standing at object A, facing B, point to C") with imagined headings of south and north on the ground. The results showed that participants who locomoted onto the wall with idiothetic cues showed a better performance in JRD for an imagined heading from which their physical heading was extended (e.g., imagined heading of North at the north wall). In addition, the participants who reproduced the layout of objects on the ceiling from a perspective extended from the ground also showed a sensorimotor alignment effect predicted by an extended heading. These results indicate that humans encode spatial relations between different surfaces and extend headings via pitch rotations three-dimensionally, especially with idiothetic cues.

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In six experiments, reading times and probe naming times were measured in order to examine the conditions under which spatial information became accessible and/or reactivated. In Experiments 1-4, reading times were measured for target sentences containing spatial inconsistencies. Spatial inconsistencies did not disrupt processing (Experiment 1) unless there were increases in task demands (Experiment 2), elaboration of the protagonist's location (Experiment 3), or both (Experiment 4). In Experiments 5 and 6, naming times were measured to directly assess the activation of spatial information, specifically objects associated with a protagonist. Spatial information was highly active in memory immediately after being read and less active after four intervening sentences (Experiment 5), but explicit cues (e.g., location or object) as well as references to the current situation model were effective in reactivating previously mentioned spatial information (Experiment 6). The combined results of six experiments are discussed within the context of the RI-Val model.

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As we move through an environment the positions of surrounding objects relative to our body constantly change. Maintaining orientation requires spatial updating, the continuous monitoring of self-motion cues to update external locations. This ability critically depends on the integration of visual, proprioceptive, kinesthetic, and vestibular information. During weightlessness gravity no longer acts as an essential reference, creating a discrepancy between vestibular, visual and sensorimotor signals. Here, we explore the effects of repeated bouts of microgravity and hypergravity on spatial updating performance during parabolic flight. Ten healthy participants (four women, six men) took part in a parabolic flight campaign that comprised a total of 31 parabolas. Each parabola created about 20-25 s of 0 g, preceded and followed by about 20 s of hypergravity (1.8 g). Participants performed a visual-spatial updating task in seated position during 15 parabolas. The task included two updating conditions simulating virtual forward movements of different lengths (short and long), and a static condition with no movement that served as a control condition. Two trials were performed during each phase of the parabola, i.e., at 1 g before the start of the parabola, at 1.8 g during the acceleration phase of the parabola, and during 0 g. Our data demonstrate that 0 g and 1.8 g impaired pointing performance for long updating trials as indicated by increased variability of pointing errors compared to 1 g. In contrast, we found no support for any changes for short updating and static conditions, suggesting that a certain degree of task complexity is required to affect pointing errors. These findings are important for operational requirements during spaceflight because spatial updating is pivotal for navigation when vision is poor or unreliable and objects go out of sight, for example during extravehicular activities in space or the exploration of unfamiliar environments. Future studies should compare the effects on spatial updating during seated and free-floating conditions, and determine at which g-threshold decrements in spatial updating performance emerge.

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Both visual and body-based (vestibular and proprioceptive) information contribute to spatial updating, or the way a navigator keeps track of self-position during movement. Research has tested the relative contributions of these sources of information and found mixed results, with some studies demonstrating the importance of body-based information, especially for translation, and some demonstrating the sufficiency of visual information. Here, we invoke an individual differences approach to test whether some individuals may be more dependent on certain types of information compared to others. Movement experts tend to be dependent on motor processes in small-scale spatial tasks, which can help or hurt performance, but it is unknown if this effect extends into large-scale spatial tasks like spatial updating. In the current study, expert dancers and non-dancers completed a virtual reality point-to-origin task with three locomotion methods that varied the availability of body-based and visual information for translation: walking, joystick, and teleporting. We predicted decrements in performance in both groups as self-motion information was reduced, and that dancers would show a larger cost. Surprisingly, both dancers and non-dancers performed with equal accuracy in walking and joystick and were impaired in teleporting, with no large differences between groups. We found slower response times for both groups with reductions in self-motion information, and minimal evidence for a larger cost for dancers. While we did not see strong dance effects, more participation in spatial activities related to decreased angular error. Together, the results suggest a flexibility in reliance on visual or body-based information for translation in spatial updating that generalizes across dancers and non-dancers, but significant decrements associated with removing both of these sources of information.

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A growing body of research suggests that performing actions can distort the perception of size, distance, and other visual information. These distortions have been observed under a variety of circumstances, and appear to persist in both perception and memory. However, it is unclear whether these distortions persist as observers move to new viewpoints. To address this issue, the present study assessed whether action-specific distortions persist across changes in viewpoint. Participants viewed an object that was projected onto a table, then reached for it with their index finger or a reach-extending tool. After reaching for the object, participants remained stationary or moved to a new viewpoint, then estimated the object's distance from their current viewpoint. When participants remained stationary, using a reach-extending tool led them to report shorter distance estimates. However, when participants moved to a new viewpoint, these distortions were eliminated. Similar effects were observed when participants produced different types of movement, including when participants rotated in place, moved to a new location, or simply walked in place. Together, these findings suggest that action-specific distortions are eliminated when observers move and perform other actions.

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Navigation can be haptically guided. In specific, tissue deformations arising from both limb motions during locomotion (i.e., gait patterns) and mechanical interactions between the limbs and the environment can convey information, detected by the haptic perceptual system, about how the body is moving relative to the environment. Here, we test hypotheses concerning the properties of mechanically contacted environments relevant to navigation of this kind. We studied blindfolded participants implicitly learning to perceive their location within environments that were physically encountered via walking on, stepping on, and probing ground surfaces with a cane. Environments were straight-line paths with elevated sections where the path either narrowed or remained the same width. We formed hypotheses concerning how these two environments would affect spatial updating and reorientation processes. In the constant pathwidth environment, homing task accuracy was higher and a manipulation of the elevated surface, to be either unchanged or (unbeknown to participants) shortened, biased the performance. This was consistent with our hypothesis of a metric recalibration scaled to elevated surface extent. In the narrowing pathwidth environment, elevated surface shortening did not bias performance. This supported our hypothesis of positional recalibration resulting from contact with the leading edge of the elevated surface. We discuss why certain environmental properties, such as path-narrowing, have significance for how one becomes implicitly oriented the surrounding environment.

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The continuous pointing task uses target-directed pointing responses to determine how perceived distance traveled is estimated during forward linear walking movements. To more precisely examine the regulation of this online process, the current study measured upper extremity joint angles and step-cycle kinematics in full vision and no-vision continuous pointing movements. Results show perceptual under-estimation of traveled distance in no-vision trials compared to full vision trials. Additionally, parsing of the shoulder plane of elevation trajectories revealed discontinuities that reflected this perceptual under-estimation and that were most frequently coupled with the early portion of the right foot swing phase of the step-cycle. This suggests that spatial updating may be composed of discrete iterations that are associated with gait parameters.

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People make use of different frames of reference (north-south; left-right) to talk about space. To explore the cognitive capacity that children bring to learning spatial language, Haun, Rapold, Call, Janzen, and Levinson (2006) examined children's ability to notice and abstract invariant frames of references across instances. They found that 4-year-olds and non-human great apes often noticed environment-defined allocentric relations and not body-defined egocentric ones, leading them to conclude that preschoolers are ready to learn environment-defined terms (e.g. "uphill"), but not body-defined ones (e.g., "left"). However, such a conclusion may be premature. In four new experiments we demonstrate that the previous findings could be an artifact of specific task constraints. With minor experiment modifications, similar-aged children readily noticed egocentric relations. Reviewing additional research, we provide an account of what makes acquiring frames of reference easy or difficult, and why full mastery of terms like "left" and "right" may take many years under normal circumstances.

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When walking or driving, it is of the utmost importance to continuously track the spatial relationship between objects in the environment and the moving body in order to prevent collisions. Although this process of spatial updating occurs naturally, it involves the processing of a myriad of noisy and ambiguous sensory signals. Here, using a psychometric approach, we investigated the integration of visual optic flow and vestibular cues in spatially updating a remembered target position during a linear displacement of the body. Participants were seated on a linear sled, immersed in a stereoscopic virtual reality environment. They had to remember the position of a target, briefly presented before a sideward translation of the body involving supra-threshold vestibular cues and whole-field optic flow that provided slightly discrepant motion information. After the motion, using a forced response participants indicated whether the location of a brief visual probe was left or right of the remembered target position. Our results show that in a spatial updating task involving passive linear self-motion humans integrate optic flow and vestibular self-displacement information according to a weighted-averaging process with, across subjects, on average about four times as much weight assigned to the visual compared to the vestibular contribution (i.e., 79% visual weight). We discuss our findings with respect to previous literature on the effect of optic flow on spatial updating performance.

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The ability to update spatial memories is important for everyday situations, such as remembering where we left our keys or parked our car. Although rodent studies have suggested that old age might impair spatial updating, direct evidence for such a deficit in humans is missing. Here, we tested whether spatial updating deficits occur in human aging, whether the learning mode influences spatial updating, and what mnemonic mechanism underlies the presumed deficits. To address these questions, younger and older participants had to indicate the latest location of relocated items, following either incidental or intentional learning. Using eye tracking, we further quantified memory traces of the original and updated locations. We found that older participants were selectively impaired in recalling locations of relocated items. Furthermore, they depicted relatively stronger representations of the original locations, which were correlated with their spatial updating deficits. The findings demonstrate that stronger representations of former spatial contexts can impair spatial updating in aging, a mechanism that can help explain the commonly observed age-related decline in spatial memory.

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The brain uses self-motion information to internally update egocentric representations of locations of remembered world-fixed visual objects. If a discrepancy is observed between this internal update and reafferent visual feedback, this could be either due to an inaccurate update or because the object has moved during the motion. To optimally infer the object's location it is therefore critical for the brain to estimate the probabilities of these two causal structures and accordingly integrate and/or segregate the internal and sensory estimates. To test this hypothesis, we designed a spatial updating task involving passive whole body translation. Participants, seated on a vestibular sled, had to remember the world-fixed position of a visual target. Immediately after the translation, the reafferent visual feedback was provided by flashing a second target around the estimated "updated" target location, and participants had to report the initial target location. We found that the participants' responses were systematically biased toward the position of the second target position for relatively small but not for large differences between the "updated" and the second target location. This pattern was better captured by a Bayesian causal inference model than by alternative models that would always either integrate or segregate the internally updated target location and the visual feedback. Our results suggest that the brain implicitly represents the posterior probability that the internally updated estimate and the visual feedback come from a common cause and uses this probability to weigh the two sources of information in mediating spatial constancy across whole body motion. NEW & NOTEWORTHY When we move, egocentric representations of object locations require internal updating to keep them in register with their true world-fixed locations. How does this mechanism interact with reafferent visual input, given that objects typically do not disappear from view? Here we show that the brain implicitly represents the probability that both types of information derive from the same object and uses this probability to weigh their contribution for achieving spatial constancy across whole body motion.

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