Fingers Crossed! An Investigation of Somatotopic Representations Using Spatial Directional Judgements

Processing of tactile stimuli requires both localising the stimuli on the body surface and combining this information with a representation of the current posture. When tactile stimuli are applied to crossed hands, the system first assumes a prototypical (e.g. uncrossed) positioning of the limbs. Remapping to include the crossed posture occurs within about 300 ms. Since fingers have been suggested to be represented in a mainly somatotopic reference frame we were interested in how the processing of tactile stimuli applied to the fingers would be affected by an unusual posture of the fingers. We asked participants to report the direction of movement of two tactile stimuli, applied successively to the crossed or uncrossed index and middle fingers of one hand at different inter-stimulus intervals (15 to 700 ms). Participants almost consistently reported perceiving the stimulus direction as opposite to what it was in the fingers crossed condition, even with SOAs of 700 ms, suggesting that on average they did not incorporate the unusual relative finger positions. Therefore our results are in agreement with the idea that, by default, the processing of tactile stimuli assumes a prototypical positioning of body parts. However, in contrast to what is generally found with tactile perception with crossed hands, performance did not improve with SOAs as long as 700 ms. This suggests that the localization of stimuli in a somatotopic reference and the integration of this representation with postural information are two separate processes that apply differently to the hands and fingers.     

Perceptual auditory aftereffects on voice identity using brief vowel stimuli

Humans can identify individuals from their voice, suggesting the existence of a perceptual representation of voice identity. We used perceptual aftereffects - shifts in perceived stimulus quality after brief exposure to a repeated adaptor stimulus - to further investigate the representation of voice identity in two experiments. Healthy adult listeners were familiarized with several voices until they reached a recognition criterion. They were then tested on identification tasks that used vowel stimuli generated by morphing between the different identities, presented either in isolation (baseline) or following short exposure to different types of voice adaptors (adaptation). Experiment 1 showed that adaptation to a given voice induced categorization shifts away from that adaptor's identity even when the adaptors consisted of vowels different from the probe stimuli. Moreover, original voices and caricatures resulted in comparable aftereffects, ruling out an explanation of identity aftereffects in terms of adaptation to low-level features. In Experiment 2, we show that adaptors with a disrupted configuration, i.e., altered fundamental frequency or formant frequencies, failed to produce perceptual aftereffects showing the importance of the preserved configuration of these acoustical cues in the representation of voices. These two experiments indicate a high-level, dynamic representation of voice identity based on the combination of several lower-level acoustical features into a specific voice configuration.     

Perception of fluids with diverse rheology applied to the underarm versus forearm skin

Little is known about the tactile–perceptual structure of fluids. Therefore, ten fluids with diverse, characterized rheologies were rated by 16 females, on 27 sensory attributes (e.g., “slippery”) and 14 emotional attributes (e.g., “enjoyable”) via five-point categorical scales. Fluids were assessed against the volar forearm and underarm, sites that commonly experience contact with fluids during the use of personal care products. Application of fluids was either by the participant to their own body (“self-applied”) or by the experimenter to the participant's body (“experimenter-applied”). Separate factor analyses of the sensory and emotional attributes for different body sites and modes of touch suggested approximately the same factorial structure in each case. Four general sensory factors emerged, labeled Lubricating, Textured, Silken, and Viscous, and two emotional factors, Comfortable and Arousing. These factors resembled those from equivalent work that used solid materials as stimuli, emphasizing that despite the differences in perceptual structure between fluid-coated and dry, solid surfaces, different body sites, and modes of touch influence the perception of fluid and dry stimuli similarly. As expected, fluids varied widely in how they scored on the factors. Site-wise differences were found, whereby stimuli assessed against the forearm were rated as more Lubricating, less Textured, more Silken, and more Comfortable than they were against the underarm. Self-applied touch was less Comfortable than experimenter-applied. The physical and perceptual were linked insofar as greater measured viscosity at low shear rates was associated with perceptions of cold and wet, whereas at high shear rates, greater viscosity was associated with greater perceived thickness.     

From a Somatotopic to a Spatiotopic Frame of Reference for the Localization of Nociceptive Stimuli

To react efficiently to potentially threatening stimuli, we have to be able to localize these stimuli in space. In daily life we are constantly moving so that our limbs can be positioned at the opposite side of space. Therefore, a somatotopic frame of reference is insufficient to localize nociceptive stimuli. Here we investigated whether nociceptive stimuli are mapped into a spatiotopic frame of reference, and more specifically a peripersonal frame of reference, which takes into account the position of the body limbs in external space, as well as the occurrence of external objects presented near the body. Two temporal order judgment (TOJ) experiments were conducted, during which participants had to decide which of two nociceptive stimuli, one applied to either hand, had been presented first while their hands were either uncrossed or crossed over the body midline. The occurrence of the nociceptive stimuli was cued by uninformative visual cues. We found that the visual cues prioritized the perception of nociceptive stimuli applied to the hand laying in the cued side of space, irrespective of posture. Moreover, the influence of the cues was smaller when they were presented far in front of participants’ hands as compared to when they were presented in close proximity. Finally, participants’ temporal sensitivity was reduced by changing posture. These findings are compatible with the existence of a peripersonal frame of reference for the localization of nociceptive stimuli. This allows for the construction of a stable representation of our body and the space closely surrounding our body, enabling a quick and efficient reaction to potential physical threats.     

Neural substrate of body size: illusory feeling of shrinking of the waist

The perception of the size and shape of one's body (body image) is a fundamental aspect of how we experience ourselves. We studied the neural correlates underlying perceived changes in the relative size of body parts by using a perceptual illusion in which participants felt that their waist was shrinking. We scanned the brains of the participants using functional magnetic resonance imaging. We found that activity in the cortices lining the left postcentral sulcus and the anterior part of the intraparietal sulcus reflected the illusion of waist shrinking, and that this activity was correlated with the reported degree of shrinking. These results suggest that the perceived changes in the size and shape of body parts are mediated by hierarchically higher-order somatosensory areas in the parietal cortex. Based on this finding we suggest that relative size of body parts is computed by the integration of more elementary somatic signals from different body segments.     

Illusory Changes in Body Size Modulate Body Satisfaction in a Way That Is Related to Non-Clinical Eating Disorder Psychopathology

Historically, body size overestimation has been linked to abnormal levels of body dissatisfaction found in eating disorders. However, recently this relationship has been called into question. Indeed, despite a link between how we perceive and how we feel about our body seeming intuitive, until now lack of an experimental method to manipulate body size has meant that a causal link, even in healthy participants, has remained elusive. Recent developments in body perception research demonstrate that the perceptual experience of the body can be readily manipulated using multisensory illusions. The current study exploits such illusions to modulate perceived body size in an attempt to influence body satisfaction. Participants were presented with stereoscopic video images of slimmer and wider mannequin bodies viewed through head-mounted displays from first person perspective. Illusory ownership was induced by synchronously stroking the seen mannequin body with the unseen real body. Pre and post-illusion affective and perceptual measures captured changes in perceived body size and body satisfaction. Illusory ownership of a slimmer body resulted in participants perceiving their actual body as slimmer and giving higher ratings of body satisfaction demonstrating a direct link between perceptual and affective body representations. Change in body satisfaction following illusory ownership of a wider body, however, was related to degree of (non-clinical) eating disorder psychopathology, which can be linked to fluctuating body representations found in clinical samples. The results suggest that body perception is linked to body satisfaction and may be of importance for eating disorder symptomology.     

Visual Working Memory Contents Bias Ambiguous Structure from Motion Perception

The way we perceive the visual world depends crucially on the state of the observer. In the present study we show that what we are holding in working memory (WM) can bias the way we perceive ambiguous structure from motion stimuli. Holding in memory the percept of an unambiguously rotating sphere influenced the perceived direction of motion of an ambiguously rotating sphere presented shortly thereafter. In particular, we found a systematic difference between congruent dominance periods where the perceived direction of the ambiguous stimulus corresponded to the direction of the unambiguous one and incongruent dominance periods. Congruent dominance periods were more frequent when participants memorized the speed of the unambiguous sphere for delayed discrimination than when they performed an immediate judgment on a change in its speed. The analysis of dominance time-course showed that a sustained tendency to perceive the same direction of motion as the prior stimulus emerged only in the WM condition, whereas in the attention condition perceptual dominance dropped to chance levels at the end of the trial. The results are explained in terms of a direct involvement of early visual areas in the active representation of visual motion in WM.     

The relationship between visual working memory and attention: retention of precise colour information in the absence of effects on perceptual selection

We examined the conditions under which a feature value in visual working memory (VWM) recruits visual attention to matching stimuli. Previous work has suggested that VWM supports two qualitatively different states of representation: an active state that interacts with perceptual selection and a passive (or accessory) state that does not. An alternative hypothesis is that VWM supports a single form of representation, with the precision of feature memory controlling whether or not the representation interacts with perceptual selection. The results of three experiments supported the dual-state hypothesis. We established conditions under which participants retained a relatively precise representation of a parcticular colour. If the colour was immediately task relevant, it reliably recruited attention to matching stimuli. However, if the colour was not immediately task relevant, it failed to interact with perceptual selection. Feature maintenance in VWM is not necessarily equivalent with feature-based attentional selection.     

Visual clutter causes high-magnitude errors

Perceptual decisions are often made in cluttered environments, where a target may be confounded with competing “distractor” stimuli. Although many studies and theoretical treatments have highlighted the effect of distractors on performance, it remains unclear how they affect thequality of perceptual decisions. Here we show that perceptual clutter leads not only to an increase in judgment errors, but also to an increase in perceived signal strength and decision confidence on erroneous trials. Observers reported simultaneously the direction and magnitude of the tilt of a target grating presented either alone, or together with vertical distractor stimuli. When presented in isolation, observers perceived isolated targets as only slightly tilted on error trials, and had little confidence in their decision. When the target was embedded in distractors, however, they perceived it to be strongly tilted on error trials, and had high confidence of their (erroneous) decisions. The results are well explained by assuming that the observers' internal representation of stimulus orientation arises from a nonlinear combination of the outputs of independent noise-perturbed front-end detectors. The implication that erroneous perceptual decisions in cluttered environments are made with high confidence has many potential practical consequences, and may be extendable to decision-making in general.     

Perceptual Grouping Enhances Visual Plasticity

Visual perceptual learning, a manifestation of neural plasticity, refers to improvements in performance on a visual task achieved by training. Attention is known to play an important role in perceptual learning, given that the observer''s discriminative ability improves only for those stimulus feature that are attended. However, the distribution of attention can be severely constrained by perceptual grouping, a process whereby the visual system organizes the initial retinal input into candidate objects. Taken together, these two pieces of evidence suggest the interesting possibility that perceptual grouping might also affect perceptual learning, either directly or via attentional mechanisms. To address this issue, we conducted two experiments. During the training phase, participants attended to the contrast of the task-relevant stimulus (oriented grating), while two similar task-irrelevant stimuli were presented in the adjacent positions. One of the two flanking stimuli was perceptually grouped with the attended stimulus as a consequence of its similar orientation (Experiment 1) or because it was part of the same perceptual object (Experiment 2). A test phase followed the training phase at each location. Compared to the task-irrelevant no-grouping stimulus, orientation discrimination improved at the attended location. Critically, a perceptual learning effect equivalent to the one observed for the attended location also emerged for the task-irrelevant grouping stimulus, indicating that perceptual grouping induced a transfer of learning to the stimulus (or feature) being perceptually grouped with the task-relevant one. Our findings indicate that no voluntary effort to direct attention to the grouping stimulus or feature is necessary to enhance visual plasticity.     

Neural modelling of the encoding of fast frequency modulation

Frequency modulation (FM) is a basic constituent of vocalisation in many animals as well as in humans. In human speech, short rising and falling FM-sweeps of around 50 ms duration, called formant transitions, characterise individual speech sounds. There are two representations of FM in the ascending auditory pathway: a spectral representation, holding the instantaneous frequency of the stimuli; and a sweep representation, consisting of neurons that respond selectively to FM direction. To-date computational models use feedforward mechanisms to explain FM encoding. However, from neuroanatomy we know that there are massive feedback projections in the auditory pathway. Here, we found that a classical FM-sweep perceptual effect, the sweep pitch shift, cannot be explained by standard feedforward processing models. We hypothesised that the sweep pitch shift is caused by a predictive feedback mechanism. To test this hypothesis, we developed a novel model of FM encoding incorporating a predictive interaction between the sweep and the spectral representation. The model was designed to encode sweeps of the duration, modulation rate, and modulation shape of formant transitions. It fully accounted for experimental data that we acquired in a perceptual experiment with human participants as well as previously published experimental results. We also designed a new class of stimuli for a second perceptual experiment to further validate the model. Combined, our results indicate that predictive interaction between the frequency encoding and direction encoding neural representations plays an important role in the neural processing of FM. In the brain, this mechanism is likely to occur at early stages of the processing hierarchy.     

Dynamic Sounds Capture the Boundaries of Peripersonal Space Representation in Humans

Background

We physically interact with external stimuli when they occur within a limited space immediately surrounding the body, i.e., Peripersonal Space (PPS). In the primate brain, specific fronto-parietal areas are responsible for the multisensory representation of PPS, by integrating tactile, visual and auditory information occurring on and near the body. Dynamic stimuli are particularly relevant for PPS representation, as they might refer to potential harms approaching the body. However, behavioural tasks for studying PPS representation with moving stimuli are lacking. Here we propose a new dynamic audio-tactile interaction task in order to assess the extension of PPS in a more functionally and ecologically valid condition.

Methodology/Principal Findings

Participants vocally responded to a tactile stimulus administered at the hand at different delays from the onset of task-irrelevant dynamic sounds which gave the impression of a sound source either approaching or receding from the subject’s hand. Results showed that a moving auditory stimulus speeded up the processing of a tactile stimulus at the hand as long as it was perceived at a limited distance from the hand, that is within the boundaries of PPS representation. The audio-tactile interaction effect was stronger when sounds were approaching compared to when sounds were receding.

Conclusion/Significance

This study provides a new method to dynamically assess PPS representation: The function describing the relationship between tactile processing and the position of sounds in space can be used to estimate the location of PPS boundaries, along a spatial continuum between far and near space, in a valuable and ecologically significant way.     

Distortions of subjective time perception within and across senses

Background

The ability to estimate the passage of time is of fundamental importance for perceptual and cognitive processes. One experience of time is the perception of duration, which is not isomorphic to physical duration and can be distorted by a number of factors. Yet, the critical features generating these perceptual shifts in subjective duration are not understood.

Methodology/Findings

We used prospective duration judgments within and across sensory modalities to examine the effect of stimulus predictability and feature change on the perception of duration. First, we found robust distortions of perceived duration in auditory, visual and auditory-visual presentations despite the predictability of the feature changes in the stimuli. For example, a looming disc embedded in a series of steady discs led to time dilation, whereas a steady disc embedded in a series of looming discs led to time compression. Second, we addressed whether visual (auditory) inputs could alter the perception of duration of auditory (visual) inputs. When participants were presented with incongruent audio-visual stimuli, the perceived duration of auditory events could be shortened or lengthened by the presence of conflicting visual information; however, the perceived duration of visual events was seldom distorted by the presence of auditory information and was never perceived shorter than their actual durations.

Conclusions/Significance

These results support the existence of multisensory interactions in the perception of duration and, importantly, suggest that vision can modify auditory temporal perception in a pure timing task. Insofar as distortions in subjective duration can neither be accounted for by the unpredictability of an auditory, visual or auditory-visual event, we propose that it is the intrinsic features of the stimulus that critically affect subjective time distortions.     

Infants’ Somatotopic Neural Responses to Seeing Human Actions: I’ve Got You under My Skin

Human infants rapidly learn new skills and customs via imitation, but the neural linkages between action perception and production are not well understood. Neuroscience studies in adults suggest that a key component of imitation–identifying the corresponding body part used in the acts of self and other–has an organized neural signature. In adults, perceiving someone using a specific body part (e.g., hand vs. foot) is associated with activation of the corresponding area of the sensory and/or motor strip in the observer’s brain–a phenomenon called neural somatotopy. Here we examine whether preverbal infants also exhibit somatotopic neural responses during the observation of others’ actions. 14-month-old infants were randomly assigned to watch an adult reach towards and touch an object using either her hand or her foot. The scalp electroencephalogram (EEG) was recorded and event-related changes in the sensorimotor mu rhythm were analyzed. Mu rhythm desynchronization was greater over hand areas of sensorimotor cortex during observation of hand actions and was greater over the foot area for observation of foot actions. This provides the first evidence that infants’ observation of someone else using a particular body part activates the corresponding areas of sensorimotor cortex. We hypothesize that this somatotopic organization in the developing brain supports imitation and cultural learning. The findings connect developmental cognitive neuroscience, adult neuroscience, action representation, and behavioral imitation.     

Discrimination and scaling of velocity of stimulus motion across the skin

The capacity of human subjects to discriminate and to scale the velocity of tactile brushing stimuli was assessed. Signal detection and classical psychophysical techniques were employed to estimate the Weber fraction over a wide range of velocities (from 1.5 to 140 cm/sec). In addition, free magnitude estimates of (1) the velocity and (2) the duration of moving tactile stimuli were obtained. It was found that human capacity to discriminate stimuli delivered to a 4 to 6-cm chord of skin on the dorsal forearm and differing in velocity remains grossly constant over the range of velocities tested and is relatively poor (i.e., the Weber fraction = 0.2-0.25). A simple power function (exponent = 0.6) satisfactorily describes the psychophysical relation (1) between the perceived and actual velocity and (2) between the perceived and actual duration of these stimuli. Since a direct proportionality between the reciprocal of a subject's estimate of duration and his or her estimate of velocity was observed, it is suggested that these two sensory attributes may reflect the operation of a neural mechanism sensitive to the duration of stimulation. Moreover, the data are inconsistent with the hypothesis that the subjects computed estimates of mean velocity from the ratio of perceived distance to perceived duration.     

Itch Relief by Mirror Scratching. A Psychophysical Study

Objective

The goal of this study was to test whether central mechanisms of scratching-induced itch attenuation can be activated by scratching the limb contralateral to the itching limb when the participant is made to visually perceive the non-itching limb as the itching limb by means of mirror images.

Methods

Healthy participants were asked to assess the intensity of an experimentally induced itch at their right forearm while they observed externally guided scratch movements either at their right (itching) or left (non-itching) forearm which were either mirrored or not mirrored. In the first experiment, a mirror placed between the participant’s forearms was used to create the visual illusion that the participant’s itching (right) forearm was being scratched while in fact the non-itching (left) forearm was scratched. To control visibility of the left (non-mirrored) forearm, a second experiment was performed in which unflipped and flipped real-time video displays of the participant’s forearms were used to create experimental conditions in which the participant visually perceived scratching either on one forearm only, on both forearms, or no scratching at all.

Results

In both experiments, scratching the non-itching limb attenuated perceived itch intensity significantly and selectively in the mirror condition, i.e., when the non-itching forearm was visually perceived as the itching limb.

Discussion

These data provide evidence that the visual illusion that an itching limb is being scratched while in fact the non-itching limb contralateral to the itching limb is scratched, can lead to significant itch relief. This effect might be due to a transient illusionary intersensory perceptual congruency of visual, tactile and pruriceptive signals. “Mirror scratching” might provide an alternative treatment to reduce itch perception in focal skin diseases with persistent pruritus without causing additional harm to the affected skin and might therefore have significant clinical impact.     

Shifting attention within memory representations involves early visual areas

Prior studies have shown that spatial attention modulates early visual cortex retinotopically, resulting in enhanced processing of external perceptual representations. However, it is not clear whether the same visual areas are modulated when attention is focused on, and shifted within a working memory representation. In the current fMRI study participants were asked to memorize an array containing four stimuli. After a delay, participants were presented with a verbal cue instructing them to actively maintain the location of one of the stimuli in working memory. Additionally, on a number of trials a second verbal cue instructed participants to switch attention to the location of another stimulus within the memorized representation. Results of the study showed that changes in the BOLD pattern closely followed the locus of attention within the working memory representation. A decrease in BOLD-activity (V1-V3) was observed at ROIs coding a memory location when participants switched away from this location, whereas an increase was observed when participants switched towards this location. Continuous increased activity was obtained at the memorized location when participants did not switch. This study shows that shifting attention within memory representations activates the earliest parts of visual cortex (including V1) in a retinotopic fashion. We conclude that even in the absence of visual stimulation, early visual areas support shifting of attention within memorized representations, similar to when attention is shifted in the outside world. The relationship between visual working memory and visual mental imagery is discussed in light of the current findings.     

How do we select perceptions and actions? Human brain imaging studies.

The selective nature of human perception and action implies a modulatory interaction between sensorimotor processes and attentional processes. This paper explores the use of functional imaging in humans to explore the mechanisms of perceptual selection and the fate of irrelevant stimuli that are not selected. Experiments with positron emission tomography show that two qualitatively different patterns of modulation of cerebral blood flow can be observed in experiments where non-spatial visual attention and auditory attention are manipulated. These patterns of modulation of cerebral blood flow modulation can be described as gain control and bias signal mechanisms. In visual and auditory cortex, the dominant change in cerebral blood flow associated with attention to either modality is related to a bias signal. The relation of these patterns of modulation to attentional effects that have been observed in single neurons is discussed. The existence of mechanisms for selective perception raises the more general question of whether irrelevant ignored stimuli are nevertheless perceived. Lavie''s theory of attention proposes that the degree to which ignored stimuli are processed varies depending on the perceptual load of the current task. Evidence from behavioural and functional magnetic resonance imaging studies of ignored visual motion processing is presented in support of this proposal.     

Expansion of direction space around the cardinal axes revealed by smooth pursuit eye movements

It is well established that perceptual direction discrimination shows an oblique effect; thresholds are higher for motion along diagonal directions than for motion along cardinal directions. Here, we compare simultaneous direction judgments and pursuit responses for the same motion stimuli and find that both pursuit and perceptual thresholds show similar anisotropies. The pursuit oblique effect is robust under a wide range of experimental manipulations, being largely resistant to changes in trajectory (radial versus tangential motion), speed (10 versus 25 deg/s), directional uncertainty (blocked versus randomly interleaved), and cognitive state (tracking alone versus concurrent tracking and perceptual tasks). Our data show that the pursuit oblique effect is caused by an effective expansion of direction space surrounding the cardinal directions and the requisite compression of space for other directions. This expansion suggests that the directions around the cardinal directions are in some way overrepresented in the visual cortical pathways that drive both smooth pursuit and perception.