Top-down attention does not modulate mechanical hypersensitivity consecutive to central sensitization: insights from an experimental analysis.

ABSTRACT: According to the neurocognitive model of attention to pain, when the attentional resources invested in a task unrelated to pain are high, limited cognitive resources can be directed toward the pain. This is supported by experimental studies showing that diverting people's attention away from acute pain leads to experiencing less pain. Theoretical work has suggested that this phenomenon may present a top-down modulatory mechanism for persistent pain as well. However, conclusive empirical evidence is lacking. To fill this gap, we used a preregistered, double-blind, between-subject study design to investigate whether performing a tailored, demanding, and engaging working memory task unrelated to pain (difficult) vs a task that requires less mental effort to be performed (easy), could lead to lower development of secondary hypersensitivity-a hallmark of central sensitization. Eighty-five healthy volunteers, randomly assigned to one of the 2 conditions, performed a visual task with a different cognitive load (difficult vs easy), while secondary hypersensitivity was induced on their nondominant forearm using high-frequency stimulation. To assess the development of secondary hypersensitivity, sensitivity to mechanical stimuli was measured 3 times: T0, for baseline and 20 (T1) and 40 (T2) minutes after the procedure. We did not observe any significant difference in the development of secondary hypersensitivity between the 2 groups, neither in terms of the intensity of mechanical sensitivity nor its spatial extent. Our results suggest that a top-down modulation through attention might not be sufficient to affect pain sensitization and the development of secondary hypersensitivity.

Perceptual simultaneity between nociceptive and visual stimuli depends on their spatial congruence.

To protect our body against physical threats, it is important to integrate the somatic and extra-somatic inputs generated by these stimuli. Temporal synchrony is an important parameter determining multisensory interaction, and the time taken by a given sensory input to reach the brain depends on the length and conduction velocity of the specific pathways through which it is transmitted. Nociceptive inputs are transmitted through very slow conducting unmyelinated C and thinly myelinated Aδ nociceptive fibers. It was previously shown that to perceive a visual stimulus and a thermo-nociceptive stimulus applied on the hand as coinciding in time, the nociceptive stimulus must precede the visual one by 76 ms for nociceptive inputs conveyed by Aδ fibers and 577 ms for inputs conveyed by C fibers. Since spatial proximity is also hypothesized to contribute to multisensory interaction, the present study investigated the effect of spatial congruence between visual and nociceptive stimuli. Participants judged the temporal order of visual and nociceptive stimuli, with the visual stimuli flashed either next to the stimulated hand or next to the opposite unstimulated hand, and with nociceptive stimuli evoking responses mediated by either Aδ or C fibers. The amount of time by which the nociceptive stimulus had to precede the visual stimulus for them to be perceived as appearing concomitantly was smaller when the visual stimulus occurred near the hand receiving the nociceptive stimulus as compared to when it occurred near the contralateral hand. This illustrates the challenge for the brain to process the synchrony between nociceptive and non-nociceptive stimuli to enable their efficient interaction to optimize defensive reaction against physical dangers.

Characterizing biased visuospatial perception in complex regional pain syndrome.

BACKGROUND: Patients suffering from complex regional pain syndrome (CRPS) are increasingly shown to be affected by cognitive difficulties. While these cognitive symptoms were initially described as limited to the perception, representation and use of the body, that is, the somatic space, they were recently shown to also extend to the perception of extra-somatic space. CRPS patients seem indeed to pay less attention to visual stimuli occurring in the same side of space as their affected limb and especially those occurring close to that limb. The aim of the present study was to more precisely characterize these visuospatial biases, by investigating whether they may be dependent on the visually perceived proximity between the visual stimuli and the affected limb. METHODS: Upper-limb CRPS patients and matched control participants performed temporal order judgements on visual stimuli, one presented in either side of space, while they could either see their hands near the visual stimuli or not. RESULTS: Visuospatial biases were not modulated by the availability of visual feedback about the hands. However, secondary analyses revealed that these biases depended on the type of rehabilitation program that the patients followed: whereas patients who did not follow any specific program presented significant biases to the detriment of visual stimuli in the affected side of space, patients who did follow a CRPS-specialized program did not. CONCLUSIONS: Patients' cognitive strategies will be important to consider when studying inter-individual differences in the cognitive symptomatology and associated cognitive-based rehabilitation procedures in CRPS. SIGNIFICANCE: The existence of biases in visuospatial perception in Complex Regional Pain Syndrome has been reported but not always systematically replicated. We show that these biases might depend on the type of general rehabilitation program that the patients follow. Patients' individual cognitive strategies will be important to consider when studying the cognitive symptomatology of CRPS.

Biological and psychological early prognostic factors in complex regional pain syndrome: A systematic review.

BACKGROUND AND OBJECTIVE: Several risk factors for the onset of CRPS have been found, but evidence for prognostic factors associated with the progression of this condition remains sparse. However, the detection and management of these factors are necessary to design secondary prevention strategies. The objective of this systematic review was to identify prognostic factors in adult individuals with early CRPS. DATABASE AND DATA TREATMENT: PubMed, Embase, PsycINFO, Cochrane Library and Scopus, were published between January 1990 and November 2021. Two independent investigators selected cross-sectional and longitudinal studies looking at early (<12 weeks from onset) prognostic factors for pain, CRPS severity score, disability, return to work, or quality of life. The quality in prognostic studies (QUIPS) tool was used to assess the risk of bias. A qualitative meta-synthesis was performed. RESULTS: Out of 4652 different articles, six studies met the inclusion criteria. We identified 21 early factors associated with a poorer prognosis in type I CRPS. We found moderate evidence to support six of them: higher pain intensity, self-rated disability, anxiety, pain-related fear, being a female and high-energy triggering event. Only two studies had an overall low risk of bias. CONCLUSIONS: This study showed an important lack of information on early prognostic factors in CRPS. Only one article investigated the link with psychological characteristics. There is a crucial need for larger studies, with a well-defined population using validated measures. SIGNIFICANCE: This systematic review highlights the lack of knowledge about early prognostic factors in CRPS. A few putative prognostic factors were identified. Most of the moderate evidence is related to a single cohort. Future research is required to find out which patients are vulnerable to chronification.

No evidence for an effect of selective spatial attention on the development of secondary hyperalgesia: A replication study.

Central sensitization refers to the increased responsiveness of nociceptive neurons in the central nervous system after repeated or sustained peripheral nociceptor activation. It is hypothesized to play a key role in the development of chronic pain. A hallmark of central sensitization is an increased sensitivity to noxious mechanical stimuli extending beyond the injured location, known as secondary hyperalgesia. For its ability to modulate the transmission and the processing of nociceptive inputs, attention could constitute a promising target to prevent central sensitization and the development of chronic pain. It was recently shown that the experimental induction of central sensitization at both forearms of healthy volunteers using bilateral high-frequency electrocutaneous stimulation (HFS), can be modulated by encouraging participants to selectively focus their attention to one arm, to the detriment of the other arm, resulting in a greater secondary hyperalgesia on the attended arm as compared to the unattended one. Given the potential value of the question being addressed, we conducted a preregistered replication study in a well-powered independent sample to assess the robustness of the effect, i.e., the modulatory role of spatial attention on the induction of central sensitization. This hypothesis was tested using a double-blind, within-subject design. Sixty-seven healthy volunteers performed a task that required focusing attention toward one forearm to discriminate innocuous vibrotactile stimuli while HFS was applied on both forearms simultaneously. Our results showed a significant increase in mechanical sensitivity directly and 20 min after HFS. However, in contrast to the previous study, we did not find a significant difference in the development of secondary hyperalgesia between the attended vs. unattended arms. Our results question whether spatial selective attention affects the development of secondary hyperalgesia. Alternatively, the non-replication could be because the bottom-up capture of attention caused by the HFS-mediated sensation was too strong in comparison to the top-down modulation exerted by the attentional task. In other words, the task was not engaging enough and the HFS pulses, including those on the unattended arm, were too salient to allow a selective focus on one arm and modulate nociceptive processing.

Measuring the sensitivity of tactile temporal order judgments in sighted and blind participants using the adaptive psi method.

Spatial locations of somatosensory stimuli are coded according to somatotopic (anatomical distribution of the sensory receptors on the skin surface) and spatiotopic (position of the body parts in external space) reference frames. This was mostly evidenced by means of temporal order judgment (TOJ) tasks in which participants discriminate the temporal order of two tactile stimuli, one applied on each hand. Because crossing the hands generates a conflict between anatomical and spatial responses, TOJ performance is decreased in such posture, except for congenitally blind people, suggesting a role of visual experience in somatosensory perception. In previous TOJ studies, stimuli were generally presented using the method of constant stimuli-that is, the repetition of a predefined sample of stimulus-onset asynchronies (SOA) separating the two stimuli. This method has the disadvantage that a large number of trials is needed to obtain reliable data when aiming at dissociating performances of groups characterized by different cognitive abilities. Indeed, each SOA among a large variety of different SOAs should be presented the same number of times irrespective of the participant's performance. This study aimed to replicate previous tactile TOJ data in sighted and blind participants with the adaptive psi method in order to validate a novel method that adapts the presented SOA according to the participant's performance. This allows to precisely estimate the temporal sensitivity of each participant while the presented stimuli are adapted to the participant's individual discrimination threshold. We successfully replicated previous findings in both sighted and blind participants, corroborating previous data using a more suitable psychophysical tool.

No Evidence for an Effect of the Distance Between the Hands on Tactile Temporal Order Judgments.

Localizing somatosensory stimuli is an important process, as it allows us to spatially guide our actions toward the object entering in contact with the body. Accordingly, the positions of tactile inputs are coded according to both somatotopic and spatiotopic representations, the latter one considering the position of the stimulated limbs in external space. The spatiotopic representation has often been evidenced by means of temporal order judgment (TOJ) tasks. Participants' judgments about the order of appearance of two successive somatosensory stimuli are less accurate when the hands are crossed over the body midline than uncrossed but also when participants' hands are placed close together when compared with farther away. Moreover, these postural effects might depend on the vision of the stimulated limbs. The aim of this study was to test the influence of seeing the hands, on the modulation of tactile TOJ by the spatial distance between the stimulated limbs. The results showed no influence of the distance between the stimulated hands on TOJ performance and prevent us from concluding whether vision of the hands affects TOJ performance, or whether these variables interact. The reliability of such distance effect to investigate the spatial representations of tactile inputs is questioned.

Visuomotor impairments in complex regional pain syndrome during pointing tasks.

ABSTRACT: Complex regional pain syndrome (CRPS) is thought to be characterized by cognitive deficits affecting patients' ability to represent, perceive, and use their affected limb as well as its surrounding space. This has been tested, among others, by straight-ahead tasks testing oneself's egocentric representation, but such experiments lead to inconsistent results. Because spatial cognitive abilities encompass various processes, we completed such evaluations by varying the sensory inputs used to perform the task. Complex regional pain syndrome and matched control participants were asked to assess their own body midline either visually (ie, by means of a moving visual cue) or manually (ie, by straight-ahead pointing with one of their upper limbs) and to reach and point to visual targets at different spatial locations. Although the 2 former tasks only required one single sensory input to be performed (ie, either visual or proprioceptive), the latter task was based on the ability to coordinate perception of the position of one's own limb with visuospatial perception. However, in this latter task, limb position could only be estimated by proprioception, as vision of the limb was prevented. Whereas in the 2 former tasks CRPS participants' performance was not different from that of controls, they made significantly more deviations errors during the visuospatial task, regardless of the limb used to point or the direction of pointing. Results suggest that CRPS patients are not specifically characterized by difficulties in representing their body but, more particularly, in integrating somatic information (ie, proprioception) during visually guided movements of the limb.

Investigating perceptual simultaneity between nociceptive and visual stimuli by means of temporal order judgments.

Multisensory interactions between pain and vision allow us to adapt our behavior to optimize detection and reaction against bodily threats. Interactions between different sensory inputs are enhanced when they are perceived closely in space and time. However, thermo-nociceptive and visual stimuli are conveyed to the cortex through specific pathways with their own conduction velocity. The present experiment aims to measure the necessary asynchrony between a nociceptive stimulus and a visual stimulus for both to be perceived as occurring simultaneously. Healthy volunteers performed a temporal order judgment task during which they discriminated the temporal order between a laser-induced nociceptive stimulus applied on one hand dorsum and a visual stimulus presented next to the stimulated hand. Laser stimulus temperature selectively activated Aδ- and/or C- fiber afferents. In order to be perceived as occurring simultaneously with a visual stimulus, a thermo-nociceptive input selectively conveyed by C-fiber afferents must precede the visual stimulus by 577 ms on average, while the stimulus-evoked input conveyed by Aδ-fiber afferents must precede it by 76 ms on average. This experiment focuses on the necessary asynchrony between thermo-nociceptive and visual inputs for them to be perceived simultaneously, to optimize the conditions under which they interact closely. Since C-fibers are unmyelinated, the asynchrony between a C-fiber stimulus and a visual stimulus is much greater than the asynchrony between a nociceptive stimulus additionally activating Aδ-fibers and that same visual stimulus. It is crucial to consider these discrepancies in further studies interested in multisensory interactions.

Rating the Intensity of a Laser Stimulus, but Not Attending to Changes in Its Location or Intensity Modulates the Laser-Evoked Cortical Activity.

Top-down attention towards nociceptive stimuli can be modulated by asking participants to pay attention to specific features of a stimulus, or to provide a rating about its intensity/unpleasantness. Whether and how these different top-down processes may lead to different modulations of the cortical response to nociceptive stimuli remains an open question. We recorded electroencephalographic (EEG) responses to brief nociceptive laser stimuli in 24 healthy participants while they performed a task in which they had to compare two subsequent stimuli on their Spatial location (Location task) or Intensity (Intensity Task). In two additional blocks (Location + Ratings, and Intensity + Ratings) participants had to further provide a rating of the perceived intensity of the stimulus. Such a design allowed us to investigate whether focusing on spatial or intensity features of a nociceptive stimulus and rating its intensity would exert different effects on the EEG responses. We did not find statistical evidence for an effect on the signal while participants were focusing on different features of the signal. We only observed a significant cluster difference in frontoparietal leads at approximately 300-500 ms post-stimulus between the magnitude of the signal in the Intensity and Intensity + Rating conditions, with a less negative response in the Intensity + Rating condition in frontal electrodes, and a less positive amplitude in parietal leads. We speculatively propose that activity in those electrodes and time window reflects magnitude estimation processes. Moreover, the smaller frontal amplitude in the Intensity + Rating condition can be explained by greater working memory engagement known to reduce the magnitude of the EEG signal. We conclude that different top-down attentional processes modulate responses to nociceptive laser stimuli at different electrodes and time windows depending on the underlying processes that are engaged.

An investigation of perceptual biases in complex regional pain syndrome.

Patients with complex regional pain syndrome (CRPS) report cognitive difficulties, affecting the ability to represent, perceive and use their affected limb. Moseley, Gallace & Spence (2009) observed that CRPS patients tend to bias the perception of tactile stimulation away from the pathological limb. Interestingly, this bias was reversed when CRPS patients were asked to cross their arms, implying that this bias is embedded in a complex representation of the body that takes into account the position of body-parts. Other studies have failed to replicate this finding (Filbrich et al., 2017) or have even found a bias in the opposite direction (Sumitani et al., 2007). Moreover, perceptual biases in CRPS patients have not often been compared to these of other chronic pain patients. Chronic pain patients are often characterized by an excessive focus of attention for bodily sensations. We might therefore expect that non-CRPS pain patients would show a bias towards instead of away from their affected limb. The aim of this study was to replicate the study of Moseley, Gallace & Spence (2009) and to extend it by comparing perceptual biases in a CRPS group with two non-CRPS pain control groups (i.e., chronic unilateral wrist and shoulder pain patients). In a temporal order judgment (TOJ) task, participants reported which of two tactile stimuli, one applied to either hand at various intervals, was perceived as occurring first. TOJs were made, either with the arms in a normal (uncrossed) position, or with the arms crossed over the body midline. We found no consistent perceptual biases in either of the patient groups and in either of the conditions (crossed/uncrossed). Individual differences were large and might, at least partly, be explained by other variables, such as pain duration and temperature differences between the pathological and non-pathological hand. Additional studies need to take these variables into account by, for example, comparing biases in CRPS (and non-CRPS) patients in an acute versus a chronic pain state.

Testing the exteroceptive function of nociception: The role of visual experience in shaping the spatial representations of nociceptive inputs.

Adequately localizing pain is crucial to protect the body against physical damage and react to the stimulus in external space having caused such damage. Accordingly, it is hypothesized that nociceptive inputs are remapped from a somatotopic reference frame, representing the skin surface, towards a spatiotopic frame, representing the body parts in external space. This ability is thought to be developed and shaped by early visual experience. To test this hypothesis, normally sighted and early blind participants performed temporal order judgment tasks during which they judged which of two nociceptive stimuli applied on each hand's dorsum was perceived as first delivered. Crucially, tasks were performed with the hands either in an uncrossed posture or crossed over body midline. While early blinds were not affected by the posture, performances of the normally sighted participants decreased in the crossed condition relative to the uncrossed condition. This indicates that nociceptive stimuli were automatically remapped into a spatiotopic representation that interfered with somatotopy in normally sighted individuals, whereas early blinds seemed to mostly rely on a somatotopic representation to localize nociceptive inputs. Accordingly, the plasticity of the nociceptive system would not purely depend on bodily experiences but also on crossmodal interactions between nociception and vision during early sensory experience.

The focus of spatial attention during the induction of central sensitization can modulate the subsequent development of secondary hyperalgesia.

Intense or sustained activation of peripheral nociceptors can induce central sensitization. This enhanced responsiveness to nociceptive input of the central nervous system primarily manifests as an increased sensitivity to painful mechanical pinprick stimuli extending beyond the site of injury (secondary mechanical hyperalgesia) and is thought to be a key mechanism in the development of chronic pain, such as persistent post-operative pain. It is increasingly recognized that emotional and cognitive factors can strongly influence the pain experience. Furthermore, through their potential effects on pain modulation circuits including descending pathways to the spinal cord, it has been hypothesized that these emotional and cognitive factors could constitute risk factors for the susceptibility to develop chronic pain. Here, we tested whether, in healthy volunteers, the experimental induction of central sensitization by peripheral nociceptive input can be modulated by selective spatial attention. While participants performed a somatosensory detection task that required focusing attention towards one of the forearms, secondary hyperalgesia was induced at both forearms using bilateral and simultaneous high-frequency electrical stimulation (HFS) of the skin. HFS induced an increased sensitivity to mechanical pinprick stimuli at both forearms, directly (T1) and 20 min (T2) after HFS, confirming the successful induction of secondary hyperalgesia at both forearms. Most importantly, at T2, the HFS-induced increase in pinprick sensitivity as well as the area of secondary hyperalgesia was greater at the attended arm as compared to the non-attended arm. This indicates that top-down attentional factors can modulate the development of central sensitization by peripheral nociceptive input, and that the focus of spatial attention, besides its modulatory effects on perception, can affect activity-dependent neuroplasticity.

Seeing or not Seeing Where Your Hands Are. The Influence of Visual Feedback About Hand Position on the Interaction Between Nociceptive and Visual Stimuli.

Examining the mechanisms underlying crossmodal interaction between nociceptive and visual stimuli is crucial to understand how humans handle potential bodily threats in their environment. It has recently been shown that nociceptive stimuli can affect the perception of visual stimuli, provided that they occur close together in external space. The present study addresses the question whether these crossmodal interactions between nociceptive and visual stimuli are mediated by the visually perceived proximity between the visual stimuli and the limb on which nociceptive stimuli are applied, by manipulating the presence vs. absence of visual feedback about the position of the stimulated limb. Participants performed temporal order judgments on pairs of visual stimuli, shortly preceded by nociceptive stimuli, either applied on one hand or both hands simultaneously. The hands were placed near the visual stimuli and could either be seen directly, seen through a glass barrier, or hidden from sight with a wooden board. Unilateral nociceptive stimuli induced spatial biases to the advantage of visual stimuli presented near the stimulated hand, which were greater in the conditions in which the hands were seen than in the condition in which vision was prevented. Spatial biases were not modulated by the presence of the glass barrier, minimizing the possibility that the differential effect between the vision and no-vision conditions is solely due to the presence of the barrier between the hands and the visual stimuli. These findings highlight the importance of visual feedback for determining spatial mapping between nociceptive and visual stimuli for crossmodal interaction.

The influence of visual experience and cognitive goals on the spatial representations of nociceptive stimuli.

Localizing pain is crucial because it allows for detecting which part of the body is being hurt and identifying in its surrounding which stimulus is producing the damage. Nociceptive inputs should therefore be mapped according to somatotopic ("which limb is stimulated?") and spatiotopic representations ("where is the stimulated limb?"). Because the body posture constantly changes, the brain has to realign the different spatial representations, for instance when the arms are crossed with the left hand in the right space and vice versa, to adequately guide actions towards the threatening object. Such ability is thought to be dependent on past sensory experience and contextual factors. We compared performances of early blind and normally sighted participants during temporal order judgement tasks. Two nociceptive stimuli were applied, one on each hand, with the hands either uncrossed or crossed. Participants reported which stimulus they perceived as first presented, according to either its location on the body or the position of the stimulated hand, respectively, prioritizing anatomy or external space as task-relevant reference frame. Relative to the uncrossed posture, sighted participants' performances were decreased when the hands were crossed, whatever the instruction be. Early blind participants' performances were affected by crossing the hands during spatial instruction, but not during anatomical instruction. These results indicate that nociceptive stimuli are automatically coded according to both somatotopic and spatiotopic representations, but the integration of the different spatial reference frames depends on early visual experience and ongoing cognitive goals, illustrating the plasticity and the flexibility of the nociceptive system.

Robot-assisted line bisection in patients with Complex Regional Pain Syndrome.

Complex Regional Pain Syndrome (CRPS) is characterized by pain, motor and inflammatory symptoms usually affecting one limb. Cognitive difficulties have been reported to affect patients' ability to represent, perceive and use their affected limb. It is debated whether these difficulties result from deficits in controlling goal-directed movements in space or from a learned strategy to protect the affected limb. In order to dissociate the two hypotheses, patients with upper-limb CRPS were asked to move with their unaffected hand towards visual targets projected at different positions on a horizontal semi-reflexive mirror. By means of a robotic handle placed below the screen, they were asked to move a cursor, to reach and cross lines at their estimated midpoint. In some of the stimulation series, the affected hand was placed below the mirror so that some lines appeared projected onto that hand. Vision of the hands and the robotic handle was preserved or prevented by opening or closing a shutter below the mirror. Lines were displayed on the mirror according to which part of the body was affected (ispi- vs. contralateral) and the actual position of the affected hand (inside vs. outside the workspace). Comparatively to control participants, CRPS patients generally biased their estimation by bisecting the lines towards their left side, irrelative of which part of the body was affected and the position of the affected hand, both in ipsi- and contralateral space, with only a few exceptions. Our results are in line with previous studies having described a visuospatial deficit in CRPS patients and discard the explanation of observed symptoms in terms of learned nonuse strategies, as only the unaffected hand was used to perform the task. It is suggested that CRPS patients can display difficulties to perform tasks requesting visuo-motor coordination, reflecting the complex cortical reorganization occurring in CRPS.

Unimodal and crossmodal extinction of nociceptive stimuli in healthy volunteers.

Nociception, the physiological mechanisms specifically processing information about noxious and potentially painful stimuli, has the double function to warn about potential body damages (interoception) and about the cause of such potential damages (exteroception). The exteroceptive function is thought to rely on multisensory integration between somatic and extra-somatic stimuli, provided that extra-somatic stimuli occur near the stimulated body area. To corroborate this hypothesis, we succeeded to show in healthy volunteers that the perception of nociceptive stimuli applied on one hand can be extinguished, as compared to single presentation, by the simultaneous application of nociceptive stimuli on the opposite hand, as well as by the presentation of visual stimuli near the opposite hand. On the contrary, visual stimuli presented near the same stimulated hand facilitated the perception of nociceptive stimuli. This nociceptive extinction phenomenon indicates that the perception of noxious events does not merely rely on the specific activation of the nociceptive system, but also depends on other sensory experiences about the body and the space around it.

Fast periodic visual stimulation to study tool-selective processing in the human brain.

Because tools are manipulated for the purpose of action, they are often considered to be a specific object category that associates perceptual and motor properties. Their neural processing has been studied extensively by comparing the cortical activity elicited by the separate presentation of tool and non-tool objects, assuming that observed differences are solely due to activity selective for processing tools. Here, using a fast periodic visual stimulation (FPVS) paradigm, we isolated EEG activity selectively related to the processing of tool objects embedded in a stream of non-tool objects. Participants saw a continuous sequence of tool and non-tool images at a 3.7 Hz presentation rate, arranged as a repeating pattern of four non-tool images followed by one tool image. We expected the stimulation to generate an EEG response at the frequency of image presentation (3.7 Hz) and its harmonics, reflecting activity common to the processing of tool and non-tool images. Most importantly, if tool and non-tool images evoked different neural responses, we expected this differential activity to generate an additional response at the frequency of tool images (3.7 Hz/5 = 0.74 Hz). To ensure that this response was not due to unaccounted for systematic differences in low-level visual features, we also tested a phase-scrambled version of the sequence. The periodic insertion of tool stimuli within a stream of non-tool stimuli elicited a significant EEG response at the tool-selective frequency and its harmonics. This response was reduced when the images were phase-scrambled. We conclude that FPVS is a promising technique to selectively measure tool-related activity.

Does Body Perception Shape Visuospatial Perception?

How we perceive our body is shaped by sensory experiences with our surrounding environment, as witnessed by poor performance in tasks during which participants judge with their hands crossed the temporal order between two somatosensory stimuli, one applied on each hand. This suggests that somatosensory stimuli are not only processed according to a somatotopic representation but also a spatiotopic representation of the body. We investigated whether the perception of stimuli occurring in external space, such as visual stimuli, can also be influenced by the body posture and somatosensory stimuli. Participants performed temporal order judgements on pairs of visual stimuli, one in each side of space, with their hands uncrossed or crossed. In Experiment 1, participants' hands were placed either near or far from the visual stimuli. In Experiment 2, the visual stimuli were preceded, either by 60 ms or 360 ms, by tactile stimuli applied on the hands placed near the visual stimuli. Manipulating the time interval was intended to activate either a somatotopic or a spatiotopic representation of somatic inputs. We did not obtain any evidence for an influence of body posture on visual temporal order judgment, suggesting that body perception is less relevant for processing extrabody stimuli than the reverse.