Temporal dynamics of the Rubber Hand Illusion.

It is widely accepted that the representation of the body is not fixed and immutable, but rather flexible and constantly updated based on a continuous stream of multisensory information. This mechanism can be very useful to adapt to several situations, but it would not be adaptive if the body representation was too malleable or if it wasn't capable of restoring its integrity after a transient modification. Here we used the Rubber Hand Illusion (RHI) to investigate how quickly the body representation can be modified. Previous studies have investigated the timing of the onset and offset of the illusion, however, they did not assess a fine temporal resolution. Here, we used a potentiometer to record a moment-by-moment rating of the feeling of owning the RH for two minutes during the visuo-tactile stimulation and two minutes following the stimulation. Our results suggest that the feeling of Ownership is already established during the first 19 s of stimulation then it continues to grow, but at a much slower pace. The feeling of Ownership disappears within 66 s from the end of the stimulation. This work sheds new light on the temporal dynamics of the RHI and the malleability of the body self-consciousness.

Expertise in Tool Use Promotes Tool Embodiment.

Body representations are known to be dynamically modulated or extended through tool use. Here, we review findings that demonstrate the importance of a user's tool experience or expertise for successful tool embodiment. Examining expert tool users, such as individuals who use tools in professional sports, people who use chopsticks at every meal, or spinal injury patients who use a wheelchair daily, offers new insights into the role of expertise in tool embodiment: Not only does tool embodiment differ between novices and experts, but experts may experience enhanced changes to their body representation when interacting with their own, personal tool. The findings reviewed herein reveal the importance of assessing tool skill in future studies of tool embodiment.

Choosing efficient actions: Deciding where to walk.

Humans evolved to be endurance animals. Our ancestors were persistence hunters; they would chase animals, including gazelles, until they ran them into exhaustion. Put simply, people evolved in an ecological niche that selected for endurance and efficiency of locomotion. To locomote to any destination, one could take countless different paths, each requiring different amounts of energy. Because the ground is typically not flat or homogeneous, the straight direct path is often not the most energetically efficient. For hills below 14°, the direct straight path up the hill is the most energetically efficient. However, for hills above 14°, walkers would minimize their absolute energy expenditure by taking a zigzagged path so that their gradient of ascension is 14° [1]. In three experiments, we assessed the degree to which people make bioenergetically efficient decisions about locomotion through path selection. In Experiment 1, people were immersed into a virtual environment and adjusted the angle of ascension of a virtual path up hills of various gradients so that when taking the path, they would expend the least amount of energy when they reached the top. The second experiment was of a similar design, but was conducted in the real word. In the last experiment, in a virtual environment, participants choose between two paths up hills of various gradient, where these paths varied in the energy required for ascent. Participants made these judgements both before and after motor experience with gradient climbing on an incline trainer. For steep hills, we found that people choose much straighter paths over the bioenergetically optimal zigzagged paths. Motor experience did lead to higher probability for choosing optimal paths for steep hills, but lead to less optimal paths for shallower ones. These results show clearly that individuals show a straight path bias when deciding how to ascend hills.

Can I reach that? Blind reaching as an accurate measure of estimated reachable distance.

Judgments of one's reach extent have been repeatedly found to be overestimated by about 10%. In 3 studies, a new dependent measure was employed in which participants viewed targets, closed their eyes, and then touched the location of the remembered target or pointed to its location if out of reach. This experimental paradigm yielded a much smaller but still present bias to over-estimate by about 2%. In addition, participants often reached for and touched target locations that were actually out of reach in a manner indicative of the typical 10% over-estimation bias. Surprisingly, participant response accuracy improved significantly and consistently across experimental trials even without visual or tactile feedback. This suggests that the proprioceptive information about the arm in space coupled with the remembered visual information about target location were sufficient to facilitate learning.

Tool Embodiment: The Tool's Output Must Match the User's Input.

The embodiment of tools and rubber hands is believed to involve the modification of two separate body representations: the body schema and the body image, respectively. It is thought that tools extend the capabilities of the body's action schema, whereas prosthetics like rubber hands are incorporated into the body image itself. Contrary to this dichotomy, recent research demonstrated that chopsticks can be embodied perceptually during a modified version of the rubber hand illusion (RHI) in which tools are held by the rubber hand and by the participant. In the present research, two experiments examined tool morpho-functional (tool output affordance, e.g., precision grasping) and sensorimotor (tool input, e.g., precision grip) match as a mechanism for this tool-use dependent change to the body image. Proprioceptive drift in the RHI occurred when the tool's output and the user's input matched, but not when this match was absent. This suggests that this factor may be necessary for tools to interact with the body image in the RHI.

Multisensory integration induces body ownership of a handtool, but not any handtool.

Bodily boundaries are computed by integrating multisensory bodily signals and can be experimentally manipulated using bodily illusions. Research on tool use demonstrates that tools alter body representations motorically to account for changes in a user's action repertoire. The present experiment sought to unify perceptual and motoric accounts of tool embodiment using a modified Rubber Hand Illusion (RHI) that also addressed the skill and practice aspects of the tool use literature. In Experiment 1, synchronous multisensory stimulation induced perceptual embodiment of a tool, chopsticks. The embodiment of chopsticks was stronger for more skilled participants, and if the illusion was preceded by tool use. In Experiment 2, the illusion was not elicited with a different type of tool, a teacup, showing that not all objects can be incorporated. This experiment helps to clarify the role of perceptual and motoric embodiment and suggests future avenues for research into tools embodiment using this method.

Optic flow is calibrated to walking effort.

Through experience, people learn that a given magnitude of walking produces an associated magnitude of optic flow. Artificially altering this relationship has both behavioral and perceptual consequences: walking on a treadmill results in zero translational optic flow and causes people to subsequently drift forward when attempting to walk in place while blindfolded (they have learned that forward walking is required to remain stationary). Similarly, after walking on a treadmill people perceive the walking distance to targets to be greater (they have recalibrated the magnitude of walking required to reach the target). While the measurement unit for walking magnitude in this relationship has been treated as walking speed (stride length * [steps / time]), recent experiments suggest that walkable distances may instead be measured in bioenergetic units (i.e., the magnitude of energy required to produce a given magnitude of optic flow). In the first experiment, zero translational optic flow was paired with a constant walking speed, and walking energy was manipulated by varying the incline of the treadmill. Participants who walked on an inclined treadmill drifted farther while attempting to walk in place than participants who walked on a flat treadmill. A follow-up experiment presented optic flow via an immersive virtual environment, and no difference between flat and inclined treadmill walking was found, thereby showing that the effect found in the first experiment was not an artifact of biomechanical differences associated with flat versus inclined treadmill walking. The results support the hypothesis that walking magnitude is scaled by bioenergetic units.

Designed Natural Spaces: Informal Gardens Are Perceived to Be More Restorative than Formal Gardens.

Experimental research shows that there are perceived and actual benefits to spending time in natural spaces compared to urban spaces, such as reduced cognitive fatigue, improved mood, and reduced stress. Whereas past research has focused primarily on distinguishing between distinct categories of spaces (i.e., nature vs. urban), less is known about variability in perceived restorative potential of environments within a particular category of outdoor spaces, such as gardens. Conceptually, gardens are often considered to be restorative spaces and to contain an abundance of natural elements, though there is great variability in how gardens are designed that might impact their restorative potential. One common practice for classifying gardens is along a spectrum ranging from "formal or geometric" to "informal or naturalistic," which often corresponds to the degree to which built or natural elements are present, respectively. In the current study, we tested whether participants use design informality as a cue to predict perceived restorative potential of different gardens. Participants viewed a set of gardens and rated each on design informality, perceived restorative potential, naturalness, and visual appeal. Participants perceived informal gardens to have greater restorative potential than formal gardens. In addition, gardens that were more visually appealing and more natural-looking were perceived to have greater restorative potential than less visually appealing and less natural gardens. These perceptions and precedents are highly relevant for the design of gardens and other similar green spaces intended to provide relief from stress and to foster cognitive restoration.

Walkable distances are bioenergetically scaled.

In perceiving spatial layout, the angular units of visual information are transformed into linear units appropriate for specifying size and extent. This derivation of linear units from angular ones requires geometry and a ruler. Numerous studies suggest that the requisite perceptual rulers are derived from the observer's body. In the case of walkable extents, it has been proposed that people scale distances to the bioenergetic resources required to traverse the extents relative to the bioenergetic resources currently available. The current study sought to rigorously test this proposal. Using methods from exercise physiology, a host of physiological measures were recorded as participants engaged in exercise on 2 occasions: once while provided with a carbohydrate supplement and once with a placebo. Distance estimates were made before and after exercise on both occasions. As in previous studies, the carbohydrate manipulation caused decreased distance estimates relative to the placebo condition. More importantly, individual differences in physiological measures that are associated with physical fitness predicted distance estimates both before and after the experimental manipulations. Results suggest that walkable distances are bioenergetically scaled.

The myth of pure perception.

Firestone & Scholl (F&S) assume that pure perception is unaffected by cognition. This assumption is untenable for definitional, anatomical, and empirical reasons. They discount research showing nonoptical influences on visual perception, pointing out possible methodological "pitfalls." Results generated in multiple labs are immune to these "pitfalls," suggesting that perceptions of physical layout do indeed reflect bioenergetic resources.

The perceptual homunculus: the perception of the relative proportions of the human body.

Given that observing one's body is ubiquitous in experience, it is natural to assume that people accurately perceive the relative sizes of their body parts. This assumption is mistaken. In a series of studies, we show that there are dramatic systematic distortions in the perception of bodily proportions, as assessed by visual estimation tasks, where participants were asked to compare the lengths of two body parts. These distortions are not evident when participants estimate the extent of a body part relative to a noncorporeal object or when asked to estimate noncorporal objects that are the same length as their body parts. Our results reveal a radical asymmetry in the perception of corporeal and noncorporeal relative size estimates. Our findings also suggest that people visually perceive the relative size of their body parts as a function of each part's relative tactile sensitivity and physical size.

Judgments of others' heights are biased toward the height of the perceiver.

We examined how observers use one aspect of their own morphology, height, when judging the physical characteristics of other people. To address this, participants judged the heights of people as they walked past. We tested the hypothesis that differences between participant and target height account for systematic patterns of variability and bias in height estimation. Height estimate error and error variability increased as the difference between participant height and target height increased, suggesting that estimates are scaled to observers' heights. Furthermore, participants' height estimates were biased toward two standards, demonstrating classic category effects. First, estimates were biased toward participants' own heights. Second, participants biased height estimates toward the average height of the target distribution. These results support past research on using both the body and categorical information to estimate target properties but extend to real-world situations involving interactions with moving people, such as height judgments provided during eyewitness testimony.

Implicit associations have a circadian rhythm.

The current study shows that people's ability to inhibit implicit associations that run counter to their explicit views varies in a circadian pattern. The presence of this rhythmic variation suggests the involvement of a biological process in regulating automatic associations--specifically, with the current data, associations that form undesirable social biases. In 1998, Greenwald, McGhee, and Schwartz introduced the Implicit Association Test as a means of measuring individual differences in implicit cognition. The IAT is a powerful tool that has become widely used. Perhaps most visibly, studies employing the IAT demonstrate that people generally hold implicit biases against social groups, which often conflict with their explicitly held views. The IAT engages inhibitory processes similar to those inherent in self-control tasks. Because the latter processes are known to be resource-limited, we considered whether IAT scores might likewise be resource dependent. Analyzing IAT performance from over a million participants across all times of day, we found a clear circadian pattern in scores. This finding suggests that the IAT measures not only the strength of implicit associations, but also the effect of variations in the physiological resources available to inhibit their undesirable influences on explicit behavior.

Evidence for hand-size constancy: the dominant hand as a natural perceptual metric.

The hand is a reliable and ecologically useful perceptual ruler that can be used to scale the sizes of close, manipulatable objects in the world in a manner similar to the way in which eye height is used to scale the heights of objects on the ground plane. Certain objects are perceived proportionally to the size of the hand, and as a result, changes in the relationship between the sizes of objects in the world and the size of the hand are attributed to changes in object size rather than hand size. To illustrate this notion, we provide evidence from several experiments showing that people perceive their dominant hand as less magnified than other body parts or objects when these items are subjected to the same degree of magnification. These findings suggest that the hand is perceived as having a more constant size and, consequently, can serve as a reliable metric with which to measure objects of commensurate size.

An Embodied Approach to Perception: By What Units Are Visual Perceptions Scaled?

When humans perceive the environment, angular units of visual information must be transformed into units appropriate for the specification of such parameters of surface layout as extent, size, and orientation. Our embodied approach to perception proposes that these scaling units derive from the body. For example, hand size is relevant for scaling the size of a strawberry, whereas an extent across a meadow is scaled by the amount of walking required to traverse it. In his article, Firestone (2013, this issue) argued that our approach is wrong; in fact, he argued that it must be wrong. This reply to Firestone's critique is organized into three parts, which address the following questions: (a) What is the fundamental question motivating our approach? (b) How does our approach answer this question? (c) How can we address Firestone's arguments against our approach? A point-by-point critique of Firestone's arguments is presented. Three conclusions are drawn: (a) Most of Firestone's arguments reflect a misunderstanding of our approach, (b) none of his arguments are the fatal flaws in our approach that he believes them to be, and (c) there are good reasons to believe that perception-just like any other biological function-is a phenotypic expression.

A perceptual-motor deficit predicts social and communicative impairments in individuals with autism spectrum disorders.

Individuals with autism spectrum disorders (ASDs) have known impairments in social and motor skills. Identifying putative underlying mechanisms of these impairments could lead to improved understanding of the etiology of core social/communicative deficits in ASDs, and identification of novel intervention targets. The ability to perceptually integrate one's physical capacities with one's environment (affordance perception) may be such a mechanism. This ability has been theorized to be impaired in ASDs, but this question has never been directly tested. Crucially, affordance perception has shown to be amenable to learning; thus, if it is implicated in deficits in ASDs, it may be a valuable unexplored intervention target. The present study compared affordance perception in adolescents and adults with ASDs to typically developing (TD) controls. Two groups of individuals (adolescents and adults) with ASDs and age-matched TD controls completed well-established action capability estimation tasks (reachability, graspability, and aperture passability). Their caregivers completed a measure of their lifetime social/communicative deficits. Compared with controls, individuals with ASDs showed unprecedented gross impairments in relating information about their bodies' action capabilities to visual information specifying the environment. The magnitude of these deficits strongly predicted the magnitude of social/communicative impairments in individuals with ASDs. Thus, social/communicative impairments in ASDs may derive, at least in part, from deficits in basic perceptual-motor processes (e.g. action capability estimation). Such deficits may impair the ability to maintain and calibrate the relationship between oneself and one's social and physical environments, and present fruitful, novel, and unexplored target for intervention.

Scared stiff: the influence of anxiety on the perception of action capabilities.

Influences on the perception of affordances (i.e., opportunities for actions) have been primarily studied by manipulating the functional morphology of the body. However, affordances are not just determined by the functional morphology of the perceiver, but also by the physiological state of the perceiver. States of anxiety have been shown to lead to marked changes in individuals' physiological state and their behaviour. To assess the influence of emotional state on affordance perception, the perception of action capabilities in near space was examined after participants completed an anxiety-provoking task. Anxiety was induced immediately prior to tasks that assessed participants' perceived reaching ability in Experiment 1, grasping ability in Experiment 2, and the ability to pass their hands through apertures in Experiment 3. Results indicated that those participants who experienced changes in anxiety underestimated their reaching, grasping, and passing ability compared to non-anxious participants. In other words, anxious participants were more conservative in their estimations of their action capabilities. These results suggest that anxiety influences the perception for affordances in near space and are consistent with the notion that anxiety induces withdrawal behaviours.

Memory for target height is scaled to observer height.

According to the embodied approach to visual perception, individuals scale the environment to their bodies. This approach highlights the central role of the body for immediate, situated action. The present experiments addressed whether body scaling--specifically, eye-height scaling--occurs in memory when action is not immediate. Participants viewed standard targets that were either the same height as, taller than, or shorter than themselves. Participants then viewed a comparison target and judged whether the comparison was taller or shorter than the standard target. Participants were most accurate when the standard target height matched their own heights, taking into account postural changes. Participants were biased to underestimate standard target height, in general, and to push standard target height away from their own heights. These results are consistent with the literature on eye-height scaling in visual perception and suggest that body scaling is not only a useful metric for perception and action, but is also preserved in memory.

Treadmill experience mediates the perceptual-motor aftereffect of treadmill walking.

People have a lifetime of experience in which to calibrate their self-produced locomotion with the resultant optical flow. Contrary to walking across the ground, however, walking on a treadmill produces minimal optical flow, and consequentially, a perceptual-motor aftereffect results. We demonstrate that the magnitude of this perceptual-motor aftereffect-measured by forward drift while attempting to march in-place following treadmill walking-decreases as experience walking on a treadmill is acquired over time. Experience with treadmill walking enables walking in this context to become sufficiently distinguished from walking in other contexts. Consequently, two distinct perceptual-motor calibration states are maintained, each linked to the context in which walking occurs. Experience with treadmill walking maintains perceptual-motor calibration accuracy in both walking contexts, despite changes to the relationship between perception and action.