A visual bias for falling objects.

Aristotle believed that objects fell at a constant velocity. However, Galileo Galilei showed that when an object falls, gravity causes it to accelerate. Regardless, Aristotle's claim raises the possibility that people's visual perception of falling motion might be biased away from acceleration towards constant velocity. We tested this idea by requiring participants to judge whether a ball moving in a simulated naturalistic setting appeared to accelerate or decelerate as a function of its motion direction and the amount of acceleration/deceleration. We found that the point of subjective constant velocity (PSCV) differed between up and down but not between left and right motion directions. The PSCV difference between up and down indicated that more acceleration was needed for a downward-falling object to appear at constant velocity than for an upward "falling" object. We found no significant differences in sensitivity to acceleration for the different motion directions. Generalized linear mixed modeling determined that participants relied predominantly on acceleration when making these judgments. Our results support the idea that Aristotle's belief may in part be due to a bias that reduces the perceived magnitude of acceleration for falling objects, a bias not revealed in previous studies of the perception of visual motion.

Vestibular Perceptual Thresholds in Older Adults With and Without Age-related Hearing Loss.

OBJECTIVES: Older adults with age-related hearing loss (ARHL) are at greater risk of falling and have greater mobility problems than older adults with normal hearing (NH). The underlying cause of these associations remains unclear. One possible reason is that age-related declines in the vestibular system could parallel those observed in the auditory system within the same individuals. Here, we compare the sensitivity of vestibular perceptual abilities (psychophysics), vestibular end-organ functioning (vestibular evoked myogenic potentials and video head impulse tests), and standing balance (posturography) in healthy older adults with and without ARHL. DESIGN: A total of 46 community-dwelling older adults, 23 with ARHL and 23 with NH, were passively translated in heave (up and down) and rotated in pitch (tilted forward and backward) in the dark using a motion platform. Using an adaptive staircase psychophysical procedure, participants' heave and pitch detection and discrimination thresholds were determined. In a posturography task, participants' center of pressure (COP) path length was measured as they stood on a forceplate with eyes open and closed, on firm and compliant surfaces, with and without sound suppression. Baseline motor, cognitive, and sensory functioning, including vestibular end-organ function, were measured. RESULTS: Individuals with ARHL were less sensitive at discriminating pitch movements compared to older adults with NH. Poorer self-reported hearing abilities were also associated with poorer pitch discrimination. In addition to pitch discrimination thresholds, lower pitch detection thresholds were significantly associated with hearing loss in the low-frequency range. Less stable standing balance was significantly associated with poorer vestibular perceptual sensitivity. DISCUSSION: These findings provide evidence for an association between ARHL and reduced vestibular perceptual sensitivity.

Body Orientation Affects the Perceived Size of Objects.

Here, we investigate how body orientation relative to gravity affects the perceived size of visual targets. When in virtual reality, participants judged the size of a visual target projected at simulated distances of between 2 and 10 m and compared it to a physical reference length held in their hands while they were standing or lying prone or supine. Participants needed to make the visual size of the target 5.4% larger when supine and 10.1% larger when prone, compared to when they were in an upright position to perceive that it matched the physical reference length. Needing to make the target larger when lying compared to when standing suggests some not mutually exclusive possibilities. It may be that while tilted participants perceived the targets as smaller than when they were upright. It may be that participants perceived the targets as being closer while tilted compared to when upright. It may also be that participants perceived the physical reference length as longer while tilted. Misperceiving objects as larger and/or closer when lying may provide a survival benefit while in such a vulnerable position.

Seeing your own or someone else's hand moving in accordance with your action: The neural interaction of agency and hand identity.

Forward models can predict sensory consequences of self-action, which is reflected by less neural processing for actively than passively generated sensory inputs (BOLD suppression effect). However, it remains open whether forward models take the identity of a moving body part into account when predicting the sensory consequences of an action. In the current study, fMRI was used to investigate the neural correlates of active and passive hand movements during which participants saw either an on-line display of their own hand or someone else's hand moving in accordance with their movement. Participants had to detect delays (0-417 ms) between their movement and the displays. Analyses revealed reduced activation in sensory areas and higher delay detection thresholds for active versus passive movements. Furthermore, there was increased activation in the hippocampus, the amygdala, and the middle temporal gyrus when someone else's hand was seen. Most importantly, in posterior parietal (angular gyrus and precuneus), frontal (middle, superior, and medial frontal gyrus), and temporal (middle temporal gyrus) regions, suppression for actively versus passively generated feedback was stronger when participants were viewing their own compared to someone else's hand. Our results suggest that forward models can take hand identity into account when predicting sensory action consequences.

The influence of rhythm on detection of auditory and vibrotactile asynchrony.

The perception of an event is strongly influenced by the context in which it occurs. Here, we examined the effect of a rhythmic context on detection of asynchrony in both the auditory and vibrotactile modalities. Using the method of constant stimuli and a two-alternative forced choice (2AFC), participants were presented with pairs of pure tones played either simultaneously or with various levels of stimulus onset asynchrony (SOA). Target stimuli in both modalities were nested within either: (i) a regularly occurring, predictable rhythm (ii) an irregular, unpredictable rhythm, or (iii) no rhythm at all. Vibrotactile asynchrony detection had higher thresholds and showed greater variability than auditory asynchrony detection in general. Asynchrony detection thresholds for auditory targets but not vibrotactile targets were significantly reduced when the target stimulus was embedded in a regular rhythm as compared to no rhythm. Embedding within an irregular rhythm produced no such improvement. The observed modality asymmetries are interpreted with regard to the superior temporal resolution of the auditory system and specialized brain circuitry supporting auditory-motor coupling.

The effect of training on the perceived approach angle in visual vertical heading judgements in a virtual environment.

Past studies have found poorer performance on vertical heading judgement accuracy compared to horizontal heading judgement accuracy. In everyday life, precise vertical heading judgements are used less often than horizontal heading judgements as we cannot usually control our vertical direction. However, pilots judging a landing approach need to consistently discriminate vertical heading angles to land safely. This study addresses the impact of training on participants' ability to judge their touchdown point relative to a target in a virtual environment with a clearly defined ground plane and horizon. Thirty-one participants completed a touchdown point estimation task twice, using three angles of descent (3°, 6° and 9°). In between the two testing tasks, half of the participants completed a flight simulator landing training task which provided feedback on their vertical heading performance; while, the other half completed a two-dimensional puzzle game as a control. Overall, participants were more precise in their responses in the second testing compared to the first (from a SD of ± 0.91° to ± 0.67°), but only the experimental group showed improvement in accuracy (from a mean error of - 2.1° to - 0.6°). Our results suggest that with training, vertical heading judgments can be as accurate as horizontal heading judgments. This study is the first to show the effectiveness of training in vertical heading judgement in naïve individuals. The results are applicable in the field of aviation, informing possible strategies for pilot training.

Visual feedback is not necessary for recalibrating the vestibular contribution to the dynamic phase of a perturbation recovery response.

Our recent work demonstrated that vision can recalibrate the vestibular signal used to re-establish equilibrium following a platform perturbation. Here, we investigate whether vision provided during a platform perturbation can recalibrate the use of vestibular reafference during the dynamic phase of the perturbation response. Dynamic postural responses were examined during a series of five forward perturbations to the body, while galvanic vestibular stimulation (GVS) selectively altered vestibular feedback and LCD occlusion spectacles controlled visual availability. Responses with and without vision were compared. The presence of GVS caused 1.78 ± 0.19 cm of medio-lateral (ML) body motion toward the anode during the initial 3 s of the dynamic postural response across perturbations. This dynamic ML response was attenuated across perturbations 1-3 independent of visual availability, resulting in a significant reduction of ML center of mass and pressure deviations (p < 0.01, ƞ2 = 0.27). That is, the vestibular influence on the ML perturbation response could be altered but vision was not necessary for this adaptation. After removing GVS, the ML response component reversed in direction toward the cathode with a magnitude that was not significantly different to the amount of response attenuation seen when GVS was present (- 0.95 ± 0.19 cm; p = 0.99, ƞ2 = 0.00). This suggested that the use of a GVS-altered vestibular signal during dynamic perturbation responses could be recalibrated, but that visual feedback was likely not responsible. Alternative mechanisms to explain the recalibration process are discussed.

Steady-state visually evoked potentials reveal partial size constancy in early visual cortex.

Our visual system maintains a stable representation of object size when viewing distance, and thus retinal size, changes. Previous studies have revealed that the extent of an object's representation in V1 shows systematic deviations from strict retinotopy when the object is perceived to be at different distances. It remains unknown, however, to what degree V1 activity accounts for perceptual size constancy. We investigated the neural correlates of size-constancy using steady-state visually evoked potentials (SSVEP) known to originate in early visual cortex. Flickering stimuli of various sizes were placed at a viewing distance of 40 cm and stimuli twice as large were shown at 80 cm. Thus both sets of stimuli had identical retinal sizes. At a constant viewing distance, SSVEP amplitude increased as a function of increasing retinal size. Crucially, SSVEP was larger when stimuli of a given retinal size were presented at 80 cm compared with at 40 cm independent of flicker frequency. Experiments were repeated and extended in virtual reality. Our results agree with previous findings showing that V1 activity plays a role in size constancy. Furthermore, we estimated the degree of the neural correction for the SSVEP as being close to 50% of the perceptual size constancy. This was the case in all experiments, independent of the effectiveness of perceptual size constancy. We conclude that retinotopy in V1 does get quite massively adjusted by perceived size, but not to the same extent as perceptual judgments.

Vestibular-somatosensory interactions affect the perceived timing of tactile stimuli.

Passive rotation has been shown to alter temporal-order judgments for tactile stimuli delivered to the hands giving an advantage to the leading hand. Here we measure thresholds for detecting stimulus onset asynchrony for touches on the hands during tilt to the left or right and during galvanic vestibular stimulation (GVS) that evoked illusory tilt. During tilt to one side, the effect of gravity on the otoliths is equivalent to a physical acceleration away from that side (e.g., tilt left is equivalent to accelerating rightwards). We therefore predicted a "leading hand advantage" for the hand opposite to the tilt direction. Thresholds for detecting asynchronicity for left-hand-first and right-hand-first touches (defined as correct detection 75% of the time) were measured separately using interleaved adaptive staircases for 15 participants. For both physical and illusory tilt there was a temporal advantage for stimuli presented to the hand contralateral to the tilt-equivalent to the "leading hand" during passive rotation. That is, there was a temporal advantage for the upward hand (for physical tilt) and for the anodal-side hand (for illusory tilt caused by GVS). These results are discussed in terms of attention and direct sensory components evoking the "leading hand" bias. These findings add to the emerging understanding of the pervasive role of vestibular activity in many aspects of cognitive processing.

The Weighting of Cues to Upright Following Stroke With and Without a History of Pushing.

OBJECTIVE: Perceived upright depends on three main factors: vision, graviception, and the internal representation of the long axis of the body. We assessed the relative contributions of these factors in individuals with sub-acute and chronic stroke and controls using a novel tool; the Oriented Character Recognition Test (OCHART). We also considered whether individuals who displayed active pushing or had a history of pushing behaviours had different weightings than those with no signs of pushing. METHOD: Three participants experienced a stroke 6 months prior: eight with a history of pushing. In total, 12 participants served as healthy aged-matched controls. Visual and graviceptive cues were dissociated by orienting the visual background left, right, or upright relative to the body, or by orienting the body left, right, or upright relative to gravity. A three-vector model was used to quantify the weightings of vision, graviception, and the body to the perceptual upright. RESULTS: The control group showed weightings of 13% vision, 25% graviception, and 62% body. Some individuals with stroke showed a similar pattern; others, particularly those with recent stroke, showed different patterns, for example, being unaffected by one of the three factors. The participant with active pushing behaviour displayed an ipsilesional perceptual bias (>30°) and was not affected by visual cues to upright. CONCLUSION: The results of OCHART may be used to quantify the weightings of multisensory inputs in individuals post-stroke and may help characterize perceptual sources of pushing behaviours.

The effect of hand position on perceived finger orientation in left- and right-handers.

In the absence of visual feedback, the perceived orientation of the fingers is systematically biased. In right-handers these biases are asymmetrical between the left and right hands in the horizontal plane and may reflect common functional postures for the two hands. Here we compared finger orientation perception in right- and left-handed participants for both hands, across various hand positions in the horizontal plane. Participants rotated a white line on a screen optically superimposed over their hand to indicate the perceived position of the finger that was rotated to one of seven orientations with the hand either aligned with the body midline, aligned with the shoulder, or displaced by twice the shoulder-to-midline distance from the midline. We replicated the asymmetric pattern of biases previously reported in right-handed participants (left hand biased towards an orientation ~30° inward, right hand ~10° inward). However, no such asymmetry was found for left-handers, suggesting left-handers may use different strategies when mapping proprioception to body or space coordinates and/or have less specialization of function between the hands. Both groups' responses rotated further outward as distance of the hand from the body midline increased, consistent with other research showing spatial orientation estimates diverge outward in the periphery. Finally, for right-handers, precision of responses was best when the hand was aligned with the shoulder compared to the other two conditions. These results highlight the unique role of hand dominance and hand position in perception of finger orientation, and provide insight into the proprioceptive position sense of the upper limbs.

Multisensory integration is independent of perceived simultaneity.

The importance of multisensory integration for perception and action has long been recognised. Integrating information from individual senses increases the chance of survival by reducing the variability in the incoming signals, thus allowing us to respond more rapidly. Reaction times (RTs) are fastest when the components of the multisensory signals are simultaneous. This response facilitation is traditionally attributed to multisensory integration. However, it is unclear if facilitation of RTs occurs when stimuli are perceived as synchronous or are actually physically synchronous. Repeated exposure to audiovisual asynchrony can change the delay at which multisensory stimuli are perceived as simultaneous, thus changing the delay at which the stimuli are integrated-perceptually. Here we set out to determine how such changes in multisensory integration for perception affect our ability to respond to multisensory events. If stimuli perceived as simultaneous were reacted to most rapidly, it would suggest a common system for multisensory integration for perception and action. If not, it would suggest separate systems. We measured RTs to auditory, visual, and audiovisual stimuli following exposure to audiovisual asynchrony. Exposure affected the variability of the unisensory RT distributions; in particular, the slowest RTs were either speed up or slowed down (in the direction predicted from shifts in perceived simultaneity). Additionally, the multisensory facilitation of RTs (beyond statistical summation) only occurred when audiovisual onsets were physically synchronous, rather than when they appeared simultaneous. We conclude that the perception of synchrony is therefore independent of multisensory integration and suggest a division between multisensory processes that are fast (automatic and unaffected by temporal adaptation) and those that are slow (perceptually driven and adaptable).

Vision can recalibrate the vestibular reafference signal used to re-establish postural equilibrium following a platform perturbation.

Visuo-vestibular recalibration, in which visual information is used to alter the interpretation of vestibular signals, has been shown to influence both oculomotor control and navigation. Here we investigate whether vision can recalibrate the vestibular feedback used during the re-establishment of equilibrium following a perturbation. The perturbation recovery responses of nine participants were examined following exposure to a period of 11 s of galvanic vestibular stimulation (GVS). During GVS in VISION trials, occlusion spectacles provided 4 s of visual information that enabled participants to correct for the GVS-induced tilt and associate this asymmetric vestibular signal with a visually provided 'upright'. NoVISION trials had no such visual experience. Participants used the visual information to assist in realigning their posture compared to when visual information was not provided (p < 0.01). The initial recovery response to a platform perturbation was not impacted by whether vision had been provided during the preceding GVS, as determined by peak centre of mass and pressure deviations (p = 0.09). However, after using vision to reinterpret the vestibular signal during GVS, final centre of mass and pressure equilibrium positions were significantly shifted compared to trials in which vision was not available (p < 0.01). These findings support previous work identifying a prominent role of vestibular input for re-establishing postural equilibrium following a perturbation. Our work is the first to highlight the capacity for visual feedback to recalibrate the vertical interpretation of vestibular reafference for re-establishing equilibrium following a perturbation. This demonstrates the rapid adaptability of the vestibular reafference signal for postural control.

Using optic flow in the far peripheral field.

Self-motion information can be used to update spatial memory of location through an estimate of a change in position. Viewing optic flow alone can create Illusory self-motion or "vection." Early studies suggested that peripheral vision is more effective than central vision in evoking vection, but controlling for retinal area and perceived distance suggests that all retinal areas may be equally effective. However, the contributions of the far periphery, beyond 90°, have been largely neglected. Using a large-field Edgeless Graphics Geometry display (EGG, Christie, Canada, field of view ±112°) and systematically blocking central (±20° to ±90°) or peripheral (viewing through tunnels ±20° to ±40°) parts of the field, we compared the effectiveness of different retinal regions at evoking forwards linear vection. Fifteen participants indicated when they had reached the position of a previously presented target after visually simulating motion down a simulated corridor. The amount of simulated travel needed to match a given target distance was modelled with a leaky spatial integrator model to estimate gains (perceived/actual distance) and a spatial decay factor. When optic flow was presented only in the far periphery (beyond 90°) gains were significantly higher than for the same motion presented full field or in only the central field, resulting in accurate performance in the range of speeds associated with normal walking. The increased effectiveness of optic flow in the peripheral field alone compared to full-field motion is discussed in terms of emerging neurophysiological studies that suggest brain areas dedicated to processing information from the far peripheral field.

Long-range tactile masking occurs in the postural body schema.

Long-range tactile masking has been reported between mirror symmetric body locations. This suggests a general principle of contralateral inhibition between corresponding points on each side of the body that may serve to enhance distinguishing touches on the two halves of the body. Do such effects occur before or after posture is added to the body schema? Here, we address this question by exploring the effect of arm position on long-range tactile masking. The influence of arm position was investigated using different positions of both the test and masking arms. Tactile sensitivity was measured on one forearm, while vibrotactile-masking stimulation was applied to the opposite arm or to a control site on the shoulder. No difference was found in sensitivity when test arm position was varied. Physical contact between the arms significantly increased the effectiveness of a masking stimulus applied to the other arm. Long-range masking between the arms was strongest when the arms were held parallel to each other and was abolished if the position of either the test arm or the masking arm was moved from this position. Modulation of the effectiveness of masking by the position of both the test and masking arms suggests that these effects occur after posture information is added to the body's representation in the brain.

Perceived finger orientation is biased towards functional task spaces.

In the absence of visual feedback, the perceived position of the hands is systematically biased towards the plausible manual task space. Here we tested whether perceived orientation of the finger is similarly misperceived in right-handed individuals. Participants' index fingers were passively rotated about the middle joint to a range of test angles, either in the frontoparallel plane (Experiment 1) or the horizontal plane (Experiment 2); they reported perceived orientation of the finger by rotating a visual line presented on a screen optically superimposed on the location of their unseen finger. Perceived finger orientations were biased towards positions that varied across hands and planes. Both hands were biased towards 10° inward in the frontoparallel plane and, in the horizontal plane, the left hand was biased towards 25° inward, whereas the right hand was biased towards 2° inwards. In a third experiment, participants reported finger orientation with respect to non-visual targets: gravitational vertical or straight ahead. Biases in perceived finger orientation to non-visual targets were similar to those found in the visual line task. The asymmetrical nature of biases across hands and planes reflects the typical orientation of the hands while working and supports the theory of a functional rather than anatomical representation of the fingers and hands in space.

Left-handers show no self-advantage in detecting a delay in visual feedback concerning an active movement.

Right-handed people show an advantage in detecting a delay in visual feedback concerning an active movement of their right hand when it is viewed in a natural perspective compared to when it is seen as if viewing another person's hand (Hoover and Harris in Exp Brain Res 233:1053-1060, 2012. doi: 10.1007/s00221-014-4181-9 ; Exp Brain Res 222:389-397, 2015a. doi: 10.1007/s00221-012-3224-3 ). This self-advantage is unique to their dominant hand and may reflect an enhanced sense of ownership which contributes to how right-handed people relate to the world. Here we asked whether left-handers show the same pattern of performance for their dominant hand. We measured the minimum delay that could be detected by 29 left-handers when viewing either their dominant or non-dominant hand from 'self' or 'other' perspectives and compared their thresholds to an age-matched sample of 22 right-handers. Right-handers showed a significant signature self-advantage of 19 ms when viewing their dominant hand in an expected 'self' perspective compared to 'other' perspectives. Left-handers, however, showed no such advantage for either their dominant or non-dominant hand. This lack of self-advantage in detecting delayed visual feedback might indicate a less secure sense of body ownership amongst left-handers.

Inducing ownership over an 'other' perspective with a visuo-tactile manipulation.

Seeing our body from a 'self' perspective while performing a movement improves our ability to detect asynchrony between the visual and proprioceptive information concerning that movement: a signature of enhanced body ownership referred to as the 'self-advantage'. We consequently experience no self-advantage when seeing our body from an 'other' perspective. Here we ask whether introducing visuo-tactile stimulation (VTS), similar to that used in the rubber hand illusion to invoke ownership over a dummy hand, would produce a self-advantage when viewing the body from a typically 'other' perspective. Prior to the experiment, participants watched a live video of their own back using a camera mounted behind them while their back was tapped with a rod for 2 min. The video was either synchronous (sVTS) or asynchronous (aVTS) with the tapping. Participants then raised their hands and made a stereotyped finger movement that they watched from the same camera either in the original, natural perspective or upside down. Participants indicated which of two periods (one with minimum delay and one with an added delay of 33-264 ms) appeared delayed. Sensitivity was calculated using psychometric functions. The sVTS group showed a self-advantage of about 45 ms in the natural visual condition compared to the upside down condition, whereas the aVTS group showed no difference between the two conditions. Synchronous visuo-tactile experience increased the feeling of ownership over a typically 'other' perspective in a quantifiable way indicating the multisensory and malleable nature of body representation.

The role of the viewpoint on body ownership.

People are more sensitive to detecting asynchrony between a self-generated movement of the hand and delayed visual feedback when what they see matches the expected "self" perspective rather than an "other" perspective (Hoover and Harris in Exp Brain Res 222:389-397, 2012). We take this as corresponding to the ability to distinguish self from others and call it the "self-advantage": a measure of body ownership. What about views of the body that cannot be seen directly? Here, we assessed the effect of familiarity of the view of the body on the self-advantage. Participants performed self-generated hand and head movements viewed directly, in a mirror, and from behind with a variable delay added to the visual feedback. Each view was shown either in the natural perspective or flipped about the vertical or horizontal axes to provide a view from another perspective. Thresholds for detecting a delay in visual feedback were calculated. Dependency of the self-advantage on perspective was most evident for views of the body that are seen most often. Results support the importance of correlating visual feedback with movement information in creating the sense of body ownership.