Location In External Space Research Articles

No Evidence for a Role of Spatially Modulated α-Band Activity in Tactile Remapping and Short-Latency, Overt Orienting Behavior.

Oscillatory α-band activity is commonly associated with spatial attention and multisensory prioritization. It has also been suggested to reflect the automatic transformation of tactile stimuli from a skin-based, somatotopic reference frame into an external one. Previous research has not convincingly separated these two possible roles of α-band activity. Previous experimental paradigms have used artificially long delays between tactile stimuli and behavioral responses to aid relating oscillatory activity to these different events. However, this strategy potentially blurs the temporal relationship of α-band activity relative to behavioral indicators of tactile-spatial transformations. Here, we assessed α-band modulation with massive univariate deconvolution, an analysis approach that disentangles brain signals overlapping in time and space. Thirty-one male and female human participants performed a delay-free, visual search task in which saccade behavior was unrestricted. A tactile cue to uncrossed or crossed hands was either informative or uninformative about visual target location. α-Band suppression following tactile stimulation was lateralized relative to the stimulated hand over central-parietal electrodes but relative to its external location over parieto-occipital electrodes. α-Band suppression reflected external touch location only after informative cues, suggesting that posterior α-band lateralization does not index automatic tactile transformation. Moreover, α-band suppression occurred at the time of, or after, the production of the saccades guided by tactile stimulation. These findings challenge the idea that α-band activity is directly involved in tactile-spatial transformation and suggest instead that it reflects delayed, supramodal processes related to attentional reorienting.SIGNIFICANCE STATEMENT Localizing a touch in space requires integrating somatosensory information about skin location and proprioceptive or visual information about posture. The automatic remapping between skin-based tactile information to a location in external space has been proposed to rely on the modulation of oscillatory brain activity in the α-band range, across the multiple cortical areas that are involved in tactile, multisensory, and spatial processing. We report two findings that are inconsistent with this view. First, α-band activity reflected the remapped stimulus location only when touch was task relevant. Second, α-band modulation occurred too late to account for spatially directed behavioral responses and, thus, only after remapping must have taken place. These characteristics contradict the idea that α-band directly reflects automatic tactile remapping processes.

External location of touch is constructed post-hoc based on limb choice.

When humans indicate on which hand a tactile stimulus occurred, they often err when their hands are crossed. This finding seemingly supports the view that the automatically determined touch location in external space affects limb assignment: the crossed right hand is localized in left space, and this conflict presumably provokes hand assignment errors. Here, participants judged on which hand the first of two stimuli, presented during a bimanual movement, had occurred, and then indicated its external location by a reach-to-point movement. When participants incorrectly chose the hand stimulated second, they pointed to where that hand had been at the correct, first time point, though no stimulus had occurred at that location. This behavior suggests that stimulus localization depended on hand assignment, not vice versa. It is, thus, incompatible with the notion of automatic computation of external stimulus location upon occurrence. Instead, humans construct external touch location post-hoc and on demand.

The perceptual continuity field is retinotopic

Visual perception is systematically biased towards input from the recent past: perceived orientation, numerosity, and face identity are pulled towards previously seen stimuli. To better understand the brain level at which serial dependence occurs, the present study examined its spatial tuning. In three experiments, serial dependence occurred between stimuli occupying the same retinal position. Serial dependence between stimuli at distant retinal locations was smaller, even when the stimuli occupied the same location in external space. The spatial window over which serial dependence occurs is thus retinotopic, but wide, suggesting that serial dependence occurs at late stages of visual processing.

Visual and Proprioceptive Influences on Tactile Spatial Processing in Adults with Autism Spectrum Disorders.

Children with autism spectrum disorders (ASDs) often exhibit altered representations of the external world. Consistently, when localizing touch, children with ASDs were less influenced than their peers by changes of the stimulated limb's location in external space [Wada et al., Scientific Reports 2015, 4(1), 5985]. However, given the protracted development of an external-spatial dominance in tactile processing in typically developing children, this difference might reflect a developmental delay rather than a set suppression of external space in ASDs. Here, adults with ASDs and matched control-participants completed (a) the tactile temporal order judgment (TOJ) task previously used to test external-spatial representation of touch in children with ASDs and (b) a tactile-visual cross-modal congruency (CC) task which assesses benefits of task-irrelevant visual stimuli on tactile localization in external space. In both experiments, participants localized tactile stimuli to the fingers of each hand, while holding their hands either crossed or uncrossed. Performance differences between hand postures reflect the influence of external-spatial codes. In both groups, tactile TOJ-performance markedly decreased when participants crossed their hands and CC-effects were especially large if the visual stimulus was presented at the same side of external space as the task-relevant touch. The absence of group differences was statistically confirmed using Bayesian statistical modeling: adults with ASDs weighted external-spatial codes comparable to typically developed adults during tactile and visual-tactile spatio-temporal tasks. Thus, atypicalities in the spatial coding of touch for children with ASDs appear to reflect a developmental delay rather than a stable characteristic of ASD. Autism Res 2019, 12: 1745-1757. © 2019 International Society for Autism Research, Wiley Periodicals, Inc. LAY SUMMARY: A touched limb's location can be described twofold, with respect to the body (right hand) or the external world (right side). Children and adolescents with autism spectrum disorder (ASD) reportedly rely less than their peers on the external world. Here, adults with and without ASDs completed two tactile localization tasks. Both groups relied to the same degree on external world locations. This opens the possibility that the tendency to relate touch to the external world is typical in individuals with ASDs but emerges with a delay.

Feeling a Touch to the Hand on the Foot

Where we perceive a touch putatively depends on topographic maps that code the touch's location on the skin [1] as well as its position in external space [2-5]. However, neither somatotopic nor external-spatial representations can account for atypical tactile percepts in some neurological patients and amputees; referral of touch to an absent or anaesthetized hand after stimulation of a foot [6, 7] or the contralateral hand [8-10] challenges the role of topographic representations when attributing touch to the limbs. Here, we show that even healthy adults systematically misattribute touch to other limbs. Participants received two tactile stimuli, each to a different limb-hand or foot-and reported which of all four limbs had been stimulated first. Hands and feet were either uncrossed or crossed to dissociate body-based and external-spatial representations [11-14]. Remarkably, participants regularly attributed the first touch to a limb that had received neither of the two stimuli. The erroneously reported, non-stimulated limb typically matched the correct limb with respect to limb type or body side. Touch was misattributed to non-stimulated limbs of the other limb type and body side only if they were placed at the correct limb's canonical (default) side of space. The touch's actual location in external space was irrelevant. These errors replicated across several contexts, and modeling linked them to incoming sensory evidence rather than to decision strategies. The results highlight the importance of the touched body part's identity and canonical location but challenge the role of external-spatial tactile representations when attributing touch to a limb.

Hallucinations: A Systematic Review of Points of Similarity and Difference Across Diagnostic Classes.

Hallucinations constitute one of the 5 symptom domains of psychotic disorders in DSM-5, suggesting diagnostic significance for that group of disorders. Although specific featural properties of hallucinations (negative voices, talking in the third person, and location in external space) are no longer highlighted in DSM, there is likely a residual assumption that hallucinations in schizophrenia can be identified based on these candidate features. We investigated whether certain featural properties of hallucinations are specifically indicative of schizophrenia by conducting a systematic review of studies showing direct comparisons of the featural and clinical characteristics of (auditory and visual) hallucinations among 2 or more population groups (one of which included schizophrenia). A total of 43 articles were reviewed, which included hallucinations in 4 major groups (nonclinical groups, drug- and alcohol-related conditions, medical and neurological conditions, and psychiatric disorders). The results showed that no single hallucination feature or characteristic uniquely indicated a diagnosis of schizophrenia, with the sole exception of an age of onset in late adolescence. Among the 21 features of hallucinations in schizophrenia considered here, 95% were shared with other psychiatric disorders, 85% with medical/neurological conditions, 66% with drugs and alcohol conditions, and 52% with the nonclinical groups. Additional differences rendered the nonclinical groups somewhat distinctive from clinical disorders. Overall, when considering hallucinations, it is inadvisable to give weight to the presence of any featural properties alone in making a schizophrenia diagnosis. It is more important to focus instead on the co-occurrence of other symptoms and the value of hallucinations as an indicator of vulnerability.

Response speed advantage for vision does not extend to touch in early deaf adults

Early deaf adults typically respond faster than hearing controls when performing a speeded simple detection on visual targets. Whether this response time advantage can generalise to another intact modality (touch) or it is instead specific to visual processing remained unexplored. We tested eight early deaf adults and twelve hearing controls in a simple detection task, with visual or tactile targets delivered on the arms and occupying the same locations in external space. Catch trials were included in the experimental paradigm. Results revealed a response time advantage in deaf adults compared to hearing controls, selectively for visual targets. This advantage did not extend to touch. The number of anticipation errors was negligible and comparable in both groups. The present findings strengthen the notion that response time advantage in deaf adults emerges as a consequence of changes specific to visual processing. They also exclude the involvement of sensory-unspecific cognitive mechanisms in this improvement (e.g. increased impulsivity in initiation of response, longer-lasting sustained attention or higher motivation to perform the task). Finally, they provide initial evidence that the intact sensory modalities can reorganise independently from each other following early auditory deprivation.

The effects of immediate vision on implicit hand maps

Perceiving the external spatial location of the limbs using position sense requires that immediate proprioceptive afferent signals be combined with a stored body model specifying the size and shape of the body. Longo and Haggard (Proc Natl Acad Sci USA 107:11727-11732, 2010) developed a method to isolate and measure this body model in the case of the hand in which participants judge the perceived location in external space of several landmarks on their occluded hand. The spatial layout of judgments of different landmarks is used to construct implicit hand maps, which can then be compared with actual hand shape. Studies using this paradigm have revealed that the body model of the hand is massively distorted, in a highly stereotyped way across individuals, with large underestimation of finger length and overestimation of hand width. Previous studies using this paradigm have allowed participants to see the locations of their judgments on the occluding board. Several previous studies have demonstrated that immediate vision, even when wholly non-informative, can alter processing of somatosensory signals and alter the reference frame in which they are localised. The present study therefore investigated whether immediate vision contributes to the distortions of implicit hand maps described previously. Participants judged the external spatial location of the tips and knuckles of their occluded left hand either while being able to see where they were pointing (as in previous studies) or while blindfolded. The characteristic distortions of implicit hand maps reported previously were clearly apparent in both conditions, demonstrating that the distortions are not an artefact of immediate vision. However, there were significant differences in the magnitude of distortions in the two conditions, suggesting that vision may modulate representations of body size and shape, even when entirely non-informative.

A 2.5-D representation of the human hand.

Primary somatosensory maps in the brain represent the body as a discontinuous, fragmented set of two-dimensional (2-D) skin regions. We nevertheless experience our body as a coherent three-dimensional (3-D) volumetric object. The links between these different aspects of body representation, however, remain poorly understood. Perceiving the body's location in external space requires that immediate afferent signals from the periphery be combined with stored representations of body size and shape. At least for the back of the hand, this body representation is massively distorted, in a highly stereotyped manner. Here we test whether a common pattern of distortions applies to the entire hand as a 3-D object, or whether each 2-D skin surface has its own characteristic pattern of distortion. Participants judged the location in external space of landmark points on the dorsal and palmar surfaces of the hand. By analyzing the internal configuration of judgments, we produced implicit maps of each skin surface. Qualitatively similar distortions were observed in both cases. The distortions were correlated across participants, suggesting that the two surfaces are bound into a common underlying representation. The magnitude of distortion, however, was substantially smaller on the palmar surface, suggesting that this binding is incomplete. The implicit representation of the human hand may be a hybrid, intermediate between a 2-D representation of individual skin surfaces and a 3-D representation of the hand as a volumetric object.

Visual context and practice change the distribution of attention in touch

It has often been shown that people experience difficulty restricting their tactile attention to one hand when both hands are touched simultaneously, especially if the hands are in close spatial proximity. Taken in conjunction with a growing body of evidence that the sensations arising from stimulation of the skin are referred to locations in external space rather than their positions on the body, these tactile attention findings provide evidence for the spatial distribution of attention in touch. However, the effect of spatial distance between the arms on the distribution of attention has been shown in only a limited number of studies and in single testing sessions. In a set of experiments we examined the stability of the arm separation effect by measuring performance over two sessions and with varying visual information about arm location. Tactile selection was more efficient when the stimulated hands were placed far apart compared with when they were adjacent but the effect was short lived (session 1 only) and only occurred when participants were tested with their eyes open. These findings suggest that somatosensory attention is a flexible, dynamic process, based on coordinates that can vary over time.

An implicit body representation underlying human position sense

Knowing the body's location in external space is a fundamental perceptual task. Perceiving the location of body parts through proprioception requires that information about the angles of each joint (i.e., body posture) be combined with information about the size and shape of the body segments between joints. Although information about body posture is specified by on-line afferent signals, no sensory signals are directly informative about body size and shape. Thus, human position sense must refer to a stored body model of the body's metric properties, such as body part size and shape. The need for such a model has long been recognized; however, the properties of this model have never been systematically investigated. We developed a technique to isolate and measure this body model. Participants judged the location in external space of 10 landmarks on the hand. By analyzing the internal configuration of the locations of these points, we produced implicit maps of the mental representation of hand size and shape. We show that this part of the body model is massively distorted, in a reliable and characteristic fashion, featuring shortened fingers and broadened hands. Intriguingly, these distortions appear to retain several characteristics of primary somatosensory representations, such as the Penfield homunculus.

Changing Reference Frames during the Encoding of Tactile Events

The mindless act of swatting a mosquito on the hand poses a remarkable challenge for the brain. Given that the primary somatosensory cortex maps skin location independently of arm posture [1, 2], the brain must realign tactile coordinates in order to locate the origin of the stimuli in extrapersonal space. Previous studies have highlighted the behavioral relevance of such an external mapping of touch, which results from combining somatosensory input with proprioceptive and visual cues about body posture [3-7]. However, despite the widely held assumption about the existence of this remapping process from somatotopic to external space and various findings indirectly suggesting its consequences [8-11], a demonstration of its changing time course and nature was lacking. We examined the temporal course of this multisensory interaction and its implications for tactile awareness in humans using a crossmodal cueing paradigm [12, 13]. What we show is that before tactile events are referred to external locations [12-15], a fleeting, unconscious image of the tactile sensation abiding to a somatotopic frame of reference rules performance. We propose that this early somatotopic "glimpse" arises from the initial feed-forward sweep of neural activity to the primary somatosensory cortex, whereas the later externally-based, conscious experience reflects the activity of a somatosensory network involving recurrent connections from association areas.

Retinotopic effects during spatial audio-visual integration

The successful integration of visual and auditory stimuli requires information about whether visual and auditory signals originate from corresponding places in the external world. Here we report crossmodal effects of spatially congruent and incongruent audio-visual (AV) stimulation. Visual and auditory stimuli were presented from one of four horizontal locations in external space. Seven healthy human subjects had to assess the spatial fit of a visual stimulus (i.e. a gray-scaled picture of a cartoon dog) and a simultaneously presented auditory stimulus (i.e. a barking sound). Functional magnetic resonance imaging (fMRI) revealed two distinct networks of cortical regions that processed preferentially either spatially congruent or spatially incongruent AV stimuli. Whereas earlier visual areas responded preferentially to incongruent AV stimulation, higher visual areas of the temporal and parietal cortex (left inferior temporal gyrus [ITG], right posterior superior temporal gyrus/sulcus [pSTG/STS], left intra-parietal sulcus [IPS]) and frontal regions (left pre-central gyrus [PreCG], left dorsolateral pre-frontal cortex [DLPFC]) responded preferentially to congruent AV stimulation. A position-resolved analysis revealed three robust cortical representations for each of the four visual stimulus locations in retinotopic visual regions corresponding to the representation of the horizontal meridian in area V1 and at the dorsal and ventral borders between areas V2 and V3. While these regions of interest (ROIs) did not show any significant effect of spatial congruency, we found subregions within ROIs in the right hemisphere that showed an incongruency effect (i.e. an increased fMRI signal during spatially incongruent compared to congruent AV stimulation). We interpret this finding as a correlate of spatially distributed recurrent feedback during mismatch processing: whenever a spatial mismatch is detected in multisensory regions (such as the IPS), processing resources are re-directed to low-level visual areas.

Spatiotopic Transfer of Visual-Form Adaptation across Saccadic Eye Movements

Although conscious perception is smooth and continuous, the input to the visual system is a series of short, discrete fixations interleaved with rapid shifts of the eye. One possible explanation for visual stability is that internal maps of objects and their visual properties are remapped around the time of saccades, but numerous studies have demonstrated that visual patterns are not combined across saccades. Here, we report that visual-form aftereffects transfer across separate fixations when adaptor and test are presented in the same spatial position. The magnitude of the transsaccadic adaptation increased with stimulus complexity, suggesting a progressive construction of spatiotopic receptive fields along the visual-form pathway. These results demonstrate that basic shape information is combined across saccades, allowing for predictive and consistent information from the past to be incorporated into each new fixation.

Influence of gaze direction on crossmodal modulation of visual ERPS by endogenous tactile spatial attention

Recent evidence indicates that the spatial direction of endogenous covert spatial attention in one sensory modality can crossmodally influence early processing of stimuli in a different modality. However, spatial locations are initially coded according to different frames of reference for different modalities (e.g., body-centered for touch versus retinocentric vision) and postural changes (e.g., gaze shifts) will realign these. Here, we used event-related potentials (ERPs) to investigate how the direction of endogenous tactile attention affects sensory-specific visual ERP components. Critically, by manipulating direction of gaze, we were able to test whether any crossmodal effects depend on visual and tactile projections to a common hemisphere, on common locations in external space, or on some combination of the two. We found that both P1 and N1 visual components were modulated according to the direction of endogenous tactile attention. While the P1 crossmodal effect followed purely hemispheric constraints, the attentional modulation of N1 appeared to combine both anatomical and external spatial constraints.

Effects of hand posture on preparatory control processes and sensory modulations in tactile-spatial attention.

Event-related brain potentials (ERPs) were measured to investigate spatial coordinate systems involved in the control of preparatory tactile-spatial orienting, and in subsequent attentional modulations of somatosensory processing. On each trial, a visual precue directed attention to the left or right hand, where infrequent tactile targets had to be detected. Hands were positioned either close together or wide apart. ERPs were recorded in the cue-target interval and in response to attended and unattended tactile non-targets. A frontal anterior directing attention negativity (ADAN) and a posterior late directing attention positivity (LDAP) were elicited in the cue-target interval contralateral to the direction of an attentional shift. The ADAN was unaffected by hand posture, but the LDAP was attenuated when hands were close together. N140 amplitudes were enhanced in response to tactile stimuli presented to the attended hand, and this effect was more pronounced when hands were wide apart. ADAN and LDAP are linked to separable anterior and posterior attentional control systems, which use coordinate systems based on somatotopic and external space, respectively. Effects of spatial attention on somatosensory stimulus processing are affected by variations in body posture. Our results demonstrate that representations of body locations in external space play a central role in the control of tactile attention.

Context effects in visual length perception: role of ocular, retinal, and spatial location.

In three experiments, we examined the transfer of orientation-contingent context effects between the eyes and across portions of the retina with or without variation in external spatial location. Previous research had shown that vertical lines are judged long, relative to horizontal lines, when the stimulus set comprises relatively long horizontals and short verticals (Contextual Condition B), as compared with the reverse when the stimulus set comprises relatively short horizontals and long verticals (Contextual Condition A). Consequently, the contextual set of stimuli influences the magnitude of the horizontal-vertical illusion (HVI), decreasing its size under Contextual Condition A and increasing its size under Contextual Condition B. Experiment 1 showed that exposing one eye to different stimulus contexts modulated the size of the HVI at the exposed eye but had little or no effect at the other eye. Experiments 2 and 3 showed that the effect of the contextual sets generalized poorly across adjacent portions of the retina but transferred almost perfectly across different locations in external space when retinal location was constant. Thus, orientation-contingent context effects in visual length perception appear to be specific to the eye and to the region of the retina stimulated, suggesting that these effects reflect relatively early and local changes in sensitivity, rather than relatively late and general shifts in response criteria.