Visual perceptual learning is enhanced by training in the illusory far space.

Visual objects in the peripersonal space (PPS) are perceived faster than farther ones appearing in the extrapersonal space (EPS). This shows preferential processing for visual stimuli near our body. Such an advantage should favour visual perceptual learning occurring near, as compared with far from observers, but opposite evidence has been recently provided from online testing protocols, showing larger perceptual learning in the far space. Here, we ran two laboratory-based experiments investigating whether visual training in PPS and EPS has different effects. We used the horizontal Ponzo Illusion to create a lateralized depth perspective while participants completed a visual search task in which they reported whether or not a specific target object orientation (e.g., a triangle pointing upwards) was present among distractors. This task was completed before and after a training phase in either the (illusory) near or far space for 1 h. In Experiment 1, the near space was in the left hemispace, whereas in Experiment 2, it was in the right. Results showed that, in both experiments, participants were more accurate after training in the far space, whereas training in the near space led to either improvement in the far space (Experiment 1), or no change (Experiment 2). Moreover, we found a larger visual perceptual learning when stimuli were presented in the left compared with the right hemispace. Differently from visual processing, visual perceptual learning is more effective in the far space. We propose that depth is a key dimension that can be used to improve human visual learning.

Visual perceptual learning is effective in the illusory far but not in the near space.

Visual shape discrimination is faster for objects close to the body, in the peripersonal space (PPS), compared with objects far from the body. Visual processing enhancement in PPS occurs also when perceived depth is based on 2D pictorial cues. This advantage has been observed from relatively low-level (detection, size, orientation) to high-level visual features (face processing). While multisensory association also displays proximal advantages, whether PPS influences visual perceptual learning remains unclear. Here, we investigated whether perceptual learning effects vary according to the distance of visual stimuli (near or far) from the observer, illusorily induced by leveraging the Ponzo illusion. Participants performed a visual search task in which they reported whether a specific target object orientation (e.g., triangle pointing downward) was present among distractors. Performance was assessed before and after practicing the visual search task (30 minutes/day for 5 days) at either the close (near group) or far (far group) distance. Results showed that participants that performed the training in the near space did not improve. By contrast, participants that performed the training in the far space showed an improvement in the visual search task in both the far and near spaces. We suggest that such improvement following the far training is due to a greater deployment of attention in the far space, which could make the learning more effective and generalize across spaces.

Size Constancy Mechanisms: Empirical Evidence from Touch.

Several studies have shown the presence of large anisotropies for tactile distance perception across several parts of the body. The tactile distance between two touches on the dorsum of the hand is perceived as larger when they are oriented mediolaterally (across the hand) than proximodistally (along the hand). This effect can be partially explained by the characteristics of primary somatosensory cortex representations. However, this phenomenon is significantly attenuated relative to differences in acuity and cortical magnification, suggesting a process of tactile size constancy. It is unknown whether the same kind of compensation also takes place when estimating the size of a continuous object. Here, we investigate whether the tactile anisotropy that typically emerges when participants have to estimate the distance between two touches is also present when a continuous object touches the skin and participants have to estimate its size. In separate blocks, participants judged which of two tactile distances or objects on the dorsum of their hand felt larger. One stimulation (first or second) was aligned with the proximodistal axis (along the hand) and the other with the mediolateral axis (across the hand). Results showed a clear anisotropy for distances between two distinct points, with across distances consistently perceived as larger than along distances, as in previous studies. Critically, however, this bias was significantly reduced or absent for judgments of the length of continuous objects. These results suggest that a tactile size constancy process is more effective when the tactile size of an object has to be approximated compared to when the distance between two touches has to be determined. The possible mechanism subserving these results is described and discussed. We suggest that a lateral inhibition mechanism, when an object touches the skin, provides information through the distribution of the inhibitory subfields of the RF about the shape of the tactile RF itself. Such a process allows an effective tactile size compensatory mechanism where a good match between the physical and perceptual dimensions of the object is achieved.

Tactile distance anisotropy on the feet.

Perception of distance between two touches varies with orientation on the hand, with distances aligned with hand width perceived as larger than those aligned with hand length. Similar anisotropies are found on other body parts (e.g., the face), suggesting they may reflect a general feature of tactile organization, but appear absent on other body parts (e.g., the belly). Here, we investigated tactile-distance anisotropy on the foot, a body part structurally and embryologically similar to the hand, but with very different patterns of functional usage in humans. In three experiments, we compared the perceived distance between pairs of touches aligned with the medio-lateral and proximal-distal foot axes. On the hairy skin of the foot dorsum, anisotropy was consistently found, with distances aligned with the medio-lateral foot axis perceived as larger than those in the proximo-distal axis. In contrast, on the glabrous skin of the sole, inconsistent results were found across experiments, with no overall evidence for anisotropy. This shows a pattern of anisotropy on the foot broadly similar to that on the hand, adding to the list of body parts showing tactile-distance anisotropy, and providing further evidence that such biases are a general aspect of tactile spatial organization across the body. Significance: The perception of tactile distance has been widely used to understand the spatial structure of touch. On the hand, anisotropy of tactile distance perception is well established, with distances oriented across hand width perceived larger than those oriented along hand length. We investigated tactile-distance anisotropy on the feet, a body part structurally, genetically, and developmentally homologous to the hands, but with strikingly different patterns of functional usage. We report highly similar patterns of anisotropy on the hairy skin of the hand dorsum and foot dorsum. This suggests that anisotropy arises from the general organization of touch across the body.

Whole-hand perceptual maps of joint location.

Hands play a fundamental role in everyday behaviour. Nevertheless, healthy adults show striking misrepresentations of their hands which have been documented by a wide range of studies addressing various aspects of body representation. For example, when asked to indicate the location within the hand of the knuckles, people place them substantially farther forward than they actually are. Previous research, however, has focused exclusively on the knuckles at the base of each finger, not considering the other knuckles in the fingers. This study, therefore, aimed to investigate conceptual knowledge of the structure of the whole hand, by investigating judgements of the location of all 14 knuckle joints in the hand. Participants localised each of the 14 knuckles of their own hand (Experiment 1) or of the experimenter's hand (Experiment 2) on a hand silhouette. We measured whether there are systematic localisation biases. The results showed highly similar pattern of mislocalisation for the knuckles of one's own hand and those of another person's hand, suggesting that people share an abstract conceptual knowledge about the hand structure. In line with previous reports, we showed that the metacarpophalangeal joints at the base of the fingers are judged as substantially father forward in the hand than they actually are. Moreover, for the first time we showed a gradient of this bias, with progressive reduction of distal bias from more proximal to more distal joints. In sum, people think their finger segments are roughly the same, and that their fingers are shorter than they are.

Reconstructing neural representations of tactile space.

Psychophysical experiments have demonstrated large and highly systematic perceptual distortions of tactile space. Such a space can be referred to our experience of the spatial organisation of objects, at representational level, through touch, in analogy with the familiar concept of visual space. We investigated the neural basis of tactile space by analysing activity patterns induced by tactile stimulation of nine points on a 3 × 3 square grid on the hand dorsum using functional magnetic resonance imaging. We used a searchlight approach within pre-defined regions of interests to compute the pairwise Euclidean distances between the activity patterns elicited by tactile stimulation. Then, we used multidimensional scaling to reconstruct tactile space at the neural level and compare it with skin space at the perceptual level. Our reconstructions of the shape of skin space in contralateral primary somatosensory and motor cortices reveal that it is distorted in a way that matches the perceptual shape of skin space. This suggests that early sensorimotor areas critically contribute to the distorted internal representation of tactile space on the hand dorsum.

Tactile interactions in the path of tactile apparent motion.

Perceptual completion is a fundamental perceptual function serving to maintain robust perception against noise. For example, we can perceive a vivid experience of motion even for the discrete inputs across time and space (apparent motion: AM). In vision, stimuli irrelevant to AM perception are suppressed to maintain smooth AM perception along the AM trajectory where no physical inputs are applied. We investigated whether such perceptual masking induced by perceptual completion of dynamic inputs is general across sensory modalities by focusing on touch. Participants tried to detect a vibro-tactile target stimulus presented along the trajectory of AM induced by two other tactile stimuli on the forearm. In a control condition, the inducing stimuli were applied simultaneously, resulting in no motion percept. Tactile target detection was impaired with tactile AM. Our findings support the notion that the perceptual masking induced by perceptual completion mechanism of AM is a general function rather than a sensory specific effect.

Fingers hold spatial information that toes do not.

Fingers have preferential associations with relative spatial locations. Tactile localisation is faster when the fingers are in these locations, such as when the index finger is in a relatively higher spatial position, and the thumb in a relatively lower position. However, it is unclear whether these associations are related to hands specifically, or are a more general characteristic of limbs. The present study therefore investigated whether toes have similar spatial associations. If these associations reflect the statistics of natural limb usage, very different patterns of association would be expected for the fingers and toes, given their different functional roles in daily behaviour. We measured reaction time (RT) and error rates of responses to tactile stimuli applied to the middle finger/toe or thumb/big toe, when they were positioned in a relative upper or lower location. We replicated the finding that fingers have preferential associations that facilitates localisation-RT and error rate were lower when the index finger was in the top position, and the thumb in the bottom position. We found that toes do not hold the same spatial information, though it remains unclear whether toes hold different spatial information or none at all. These results demonstrate spatial information held by the fingers is stronger and more reliable than for the toes, so is not a general characteristic of limbs, but possibly related to hand use.

Anisotropy in tactile time perception.

Spatial distortions in touch have been investigated since the 19th century. For example, two touches applied to the hand dorsum feel farther apart when aligned with the mediolateral axis (i.e., across the hand) than when aligned with the proximodistal axis (along the hand). Stimulations to our sensory receptors are usually dynamic, where spatial and temporal inputs closely interact to establish our percept. For example, physically bigger tactile stimuli are judged to last longer than smaller stimuli. Given such links between space and time in touch, we investigated whether there is a tactile anisotropy in temporal perception analogous to the anisotropy described above. In this case, the perceived duration between the onset of two touches should be larger when they are aligned with the mediolateral than with the proximodistal axis of the hand dorsum. To test this hypothesis, we asked participants to judge which of two tactile temporal sequences, having the same spatial separation along and across the dorsum, felt longer. A clear anisotropy of the temporal perception was observed: temporal intervals across the hand were perceived as longer than those along the hand. Consistent with the spatial anisotropy, the temporal anisotropy did not appear on the palm side of the hand, indicating that the temporal anisotropy was based on perceptual processes rather than top-down modulations such as attentional or decisional/response biases. Contrary to our predictions, however, we found no correlation between the magnitudes of the temporal and spatial anisotropies. Our results demonstrated a novel type of temporal illusion in touch, which is strikingly similar in nature to the previously reported spatial anisotropy. Thus, qualitatively similar distorted somatosensory representations appear to underlie both temporal and spatial processing of touch.

Structural representations of fingers rely on both anatomical and spatial reference frames.

Finger agnosia refers to a neurological condition in which patients with left posterior parietal lesions fail to identify their fingers, despite having relatively preserved abilities in sensation and skilled action. This dissociation suggests that the structural body representations (BSRs) may be distinct from sensorimotor representations. However, recent research has reported that postural changes modulate representation of hand structure, revealing dynamic interactions between structural and sensorimotor body representations. However, it is unknown how and to what extent anatomical and spatial proximity contribute to shape the hand structural representation. We investigate this question using the "in-between" test in which participants estimate how many unstimulated fingers are in between 2 touched fingers of the left hand placed palm down. The first phalange of the participants' fingers was touched on the left or right side. Judged finger numerosity was greater when fingers were stimulated on far sides (i.e., opposite sides of the 2 fingers) compared to when they were stimulated on close (i.e., sides facing each other's) or middistance (i.e., sides facing in the same direction) sides. Therefore, finger identification was modulated by anatomical and spatial proximity in external space between touches. This demonstrates that BSRs rely on both anatomical and external reference frames. (PsycINFO Database Record (c) 2020 APA, all rights reserved).

A common representation of fingers and toes.

There are many similarities and differences between the human hands and feet. On a psychological level, there is some evidence from clinical disorders and studies of tactile localisation in healthy adults for deep functional connections between the hands and feet. One form these connections may take is in common high-level mental representations of the hands and feet. Previous studies have shown that there are systematic, but distinct patterns of confusion found between both the fingers and toes. Further, there are clear individual differences between people in the exact patterns of mislocalisations. Here, we investigated whether these idiosyncratic differences in tactile localisation are shared between the fingers and toes, which may indicate a shared high-level representation. We obtained confusion matrices showing the pattern of mislocalisation on the hairy skin surfaces of both the fingers and toes. Using a decoding approach, we show that idiosyncratic differences in individuals' pattern of confusions are shared across the fingers and toes, despite different overall patterns of confusions. These results suggest that there is a common representation of the fingers and toes.

The standard posture of the hand.

Perceived limb position is known to rely on sensory signals and motor commands. Another potential source of input is a standard representation of body posture, which may bias perceived limb position toward more stereotyped positions. Recent results show that tactile stimuli are processed more efficiently when delivered to a thumb in a relatively low position or an index finger in a relatively high position. This observation suggests that we may have a standard posture of the body that promotes a more efficient interaction with the environment. In this study, we mapped the standard posture of the entire hand by characterizing the spatial associations of all 5 digits. Moreover, we show that the effect is not an artifact of intermanual integration. Results showed that the thumb is associated with low positions, while the other fingers are associated with upper locations. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

A Conceptual Model of Tactile Processing across Body Features of Size, Shape, Side, and Spatial Location.

The processing of touch depends of multiple factors, such as the properties of the skin and type of receptors stimulated, as well as features related to the actual configuration and shape of the body itself. A large body of research has focused on the effect that the nature of the stimuli has on tactile processing. Less research, however, has focused on features beyond the nature of the touch. In this review, we focus on some features related to the body that have been investigated for less time and in a more fragmented way. These include the symmetrical quality of the two sides of the body, the postural configuration of the body, as well as the size and shape of different body parts. We will describe what we consider three key aspects: (1) how and at which stages tactile information is integrated between different parts and sides of the body; (2) how tactile signals are integrated with online and stored postural configurations of the body, regarded as priors; (3) and how tactile signals are integrated with representations of body size and shape. Here, we describe how these different body dimensions affect integration of tactile information as well as guide motor behavior by integrating them in a single model of tactile processing. We review a wide range of neuropsychological, neuroimaging, and neurophysiological data and suggest a revised model of tactile integration on the basis of the one proposed previously by Longo et al.

Locating primary somatosensory cortex in human brain stimulation studies: systematic review and meta-analytic evidence.

Transcranial magnetic stimulation (TMS) over human primary somatosensory cortex (S1), unlike over primary motor cortex (M1), does not produce an immediate, objective output. Researchers must therefore rely on one or more indirect methods to position the TMS coil over S1. The "gold standard" method of TMS coil positioning is to use individual functional and structural magnetic resonance imaging (f/sMRI) alongside a stereotactic navigation system. In the absence of these facilities, however, one common method used to locate S1 is to find the scalp location that produces twitches in a hand muscle (e.g., the first dorsal interosseus, M1-FDI) and then move the coil posteriorly to target S1. There has been no systematic assessment of whether this commonly reported method of finding the hand area of S1 is optimal. To do this, we systematically reviewed 124 TMS studies targeting the S1 hand area and 95 fMRI studies involving passive finger and hand stimulation. Ninety-six TMS studies reported the scalp location assumed to correspond to S1-hand, which was on average 1.5-2 cm posterior to the functionally defined M1-hand area. Using our own scalp measurements combined with similar data from MRI and TMS studies of M1-hand, we provide the estimated scalp locations targeted in these TMS studies of the S1-hand. We also provide a summary of reported S1 coordinates for passive finger and hand stimulation in fMRI studies. We conclude that S1-hand is more lateral to M1-hand than assumed by the majority of TMS studies.

Locating primary somatosensory cortex in human brain stimulation studies: experimental evidence.

Transcranial magnetic stimulation (TMS) over human primary somatosensory cortex (S1) does not produce immediate outputs. Researchers must therefore rely on indirect methods for TMS coil positioning. The "gold standard" is to use individual functional and structural magnetic resonance imaging (MRI) data, but the majority of studies don't do this. The most common method to locate the hand area of S1 (S1-hand) is to move the coil posteriorly from the hand area of primary motor cortex (M1-hand). Yet, S1-hand is not directly posterior to M1-hand. We localized the index finger area of S1-hand (S1-index) experimentally in four ways. First, we reanalyzed functional MRI data from 20 participants who received vibrotactile stimulation to their 10 digits. Second, to assist the localization of S1-hand without MRI data, we constructed a probabilistic atlas of the central sulcus from 100 healthy adult MRIs and measured the likely scalp location of S1-index. Third, we conducted two experiments mapping the effects of TMS across the scalp on tactile discrimination performance. Fourth, we examined all available neuronavigation data from our laboratory on the scalp location of S1-index. Contrary to the prevailing method, and consistent with systematic review evidence, S1-index is close to the C3/C4 electroencephalography (EEG) electrode locations on the scalp, ~7-8 cm lateral to the vertex, and ~2 cm lateral and 0.5 cm posterior to the M1-hand scalp location. These results suggest that an immediate revision to the most commonly used heuristic to locate S1-hand is required. The results of many TMS studies of S1-hand need reassessment. NEW & NOTEWORTHY Noninvasive human brain stimulation requires indirect methods to target particular brain areas. Magnetic stimulation studies of human primary somatosensory cortex have used scalp-based heuristics to find the target, typically locating it 2 cm posterior to the motor cortex. We measured the scalp location of the hand area of primary somatosensory cortex and found that it is ~2 cm lateral to motor cortex. Our results suggest an immediate revision of the prevailing method is required.

Dissociation of feeling and belief in the rubber hand illusion.

The Rubber Hand Illusion (RHI) has been widely used to investigate the perception of the bodily self. Commonly used measures of the illusion are self-report questionnaires and proprioceptive drift of the participants' hands towards the rubber hand. Recent studies have shown that these measures can be dissociated, suggesting they may arise from distinct mechanisms. In previous studies using questionnaires, participants were asked to base responses on their subjective feelings of body ownership, rather than their beliefs. This makes sense given the obvious fact that whereas participants may feel like the rubber hand is part of their body, they do not believe that it is. It is not clear, however, whether a similar dissociation between feelings and beliefs also exists for proprioceptive drift. Here, we investigated the presence of a dissociation between feeling and belief in the context of the RHI. When participants reported their feelings there was an increase both in the sense of body ownership over the fake hand as well as in the proprioceptive drift, compared to when they reported their beliefs. Strikingly, unlike the sense of ownership, proprioceptive drift was unaffected by the synchrony of stimulation. This may be an important way in which the two measures of the RHI differ.

Tactile confusions of the fingers and toes.

Recent research has shown systematic patterns of confusions between digits of the hands and feet. The present study addressed whether such confusions arise from early somatosensory maps or higher level body representations. As the glabrous and hairy skin of the hands and feet have distinct representations in somatosensory cortex, an effect arising from early somatotopic maps may show distinct patterns on each skin surface. In contrast, if the effect arises from higher level body representations which represent the digits as volumetric units, similar patterns should be apparent regardless of which side of the digit is touched. We obtained confusion matrices showing the pattern of mislocalization on the glabrous and hairy skin surfaces of the toes (Experiment 1) and fingers (Experiment 2). Our results replicated the characteristic pattern of mislocalizations found on the glabrous skin reported in previous studies. Critically, these effects were highly similar on the hairy skin surface of both the toes and fingers. Despite the pattern of mislocalizations being highly stereotyped across participants, there were consistent individual differences in the pattern of confusions across the two skin surfaces. These results suggest that mislocalizations occur at the level of individual digits, consistent with their resulting from higher level body representations. (PsycINFO Database Record (c) 2018 APA, all rights reserved).

More than skin-deep: Integration of skin-based and musculoskeletal reference frames in localization of touch.

The skin of the forearm is, in one sense, a flat 2-dimensional (2D) sheet, but in another sense approximately cylindrical, mirroring the 3-dimensional (3D) volumetric shape of the arm. The role of frames of reference based on the skin as a 2D sheet versus based on the musculoskeletal structure of the arm remains unclear. When we rotate the forearm from a pronated to a supinated posture, the skin on its surface is displaced. Thus, a marked location will slide with the skin across the underlying flesh, and the touch perceived at this location should follow this displacement if it is localized within a skin-based reference frame. We investigated, however, if the perceived tactile locations were also affected by the rearrangement in underlying musculoskeletal structure, that is, displaced medially and laterally on a pronated and supinated forearm, respectively. Participants pointed to perceived touches (Experiment 1), or marked them on a (3D) size-matched forearm on a computer screen (Experiment 2). The perceived locations were indeed displaced medially after forearm pronation in both response modalities. This misperception was reduced (Experiment 1), or absent altogether (Experiment 2) in the supinated posture when the actual stimulus grid moved laterally with the displaced skin. The grid was perceptually stretched at medial-lateral axis, and it was displaced distally, which suggest the influence of skin-based factors. Our study extends the tactile localization literature focused on the skin-based reference frame and on the effects of spatial positions of body parts by implicating the musculoskeletal factors in localization of touch on the body. (PsycINFO Database Record (c) 2018 APA, all rights reserved).

Multisensory Perception: Magnetic Disruption of Attention in Human Parietal Lobe.

Paying attention to sounds and touches at the same time is demanding. New research shows how the parietal lobe of the human brain mediates multisensory perception of stimulus frequency and intensity.