The hands' default location guides tactile spatial selectivity.

Our skin is a two-dimensional sheet that can be folded into a multitude of configurations due to the mobility of our body parts. Parts of the human tactile system might account for this flexibility by being tuned to locations in the world rather than on the skin. Using adaptation, we scrutinized the spatial selectivity of two tactile perceptual mechanisms for which the visual equivalents have been reported to be selective in world coordinates: tactile motion and the duration of tactile events. Participants' hand position-uncrossed or crossed-as well as the stimulated hand varied independently across adaptation and test phases. This design distinguished among somatotopic selectivity for locations on the skin and spatiotopic selectivity for locations in the environment, but also tested spatial selectivity that fits neither of these classical reference frames and is based on the default position of the hands. For both features, adaptation consistently affected subsequent tactile perception at the adapted hand, reflecting skin-bound spatial selectivity. Yet, tactile motion and temporal adaptation also transferred across hands but only if the hands were crossed during the adaptation phase, that is, when one hand was placed at the other hand's typical location. Thus, selectivity for locations in the world was based on default rather than online sensory information about the location of the hands. These results challenge the prevalent dichotomy of somatotopic and spatiotopic selectivity and suggest that prior information about the hands' default position -right hand at the right side-is embedded deep in the tactile sensory system.

Prior Movement of One Arm Facilitates Motor Adaptation in the Other.

Many movements in daily life are embedded in motion sequences that involve more than one limb, demanding the motor system to monitor and control different body parts in quick succession. During such movements, systematic changes in the environment or the body might require motor adaptation of specific segments. However, previous motor adaptation research has focused primarily on motion sequences produced by a single limb, or on simultaneous movements of several limbs. For example, adaptation to opposing force fields is possible in unimanual reaching tasks when the direction of a prior or subsequent movement is predictive of force field direction. It is unclear, however, whether multilimb sequences can support motor adaptation processes in a similar way. In the present study (38 females, 38 males), we investigated whether reaches can be adapted to different force fields in a bimanual motor sequence when the information about the perturbation is associated with the prior movement direction of the other arm. In addition, we examined whether prior perceptual (visual or proprioceptive) feedback of the opposite arm contributes to force field-specific motor adaptation. Our key finding is that only active participation in the bimanual sequential task supports pronounced adaptation. This result suggests that active segments in bimanual motion sequences are linked across limbs. If there is a consistent association between movement kinematics of the linked and goal movement, the learning process of the goal movement can be facilitated. More generally, if motion sequences are repeated often, prior segments can evoke specific adjustments of subsequent movements.SIGNIFICANCE STATEMENT Movements in a limb's motion sequence can be adjusted based on linked movements. A prerequisite is that kinematics of the linked movements correctly predict which adjustments are needed. We show that use of kinematic information to improve performance is even possible when a prior linked movement is performed with a different limb. For example, a skilled juggler might have learned how to correctly adjust his catching movement of the left hand when the right hand performed a throwing action in a specific way. Linkage is possibly a key mechanism of the human motor system for learning complex bimanual skills. Our study emphasizes that learning of specific movements should not be studied in isolation but within their motor sequence context.

Tactile training facilitates infants' ability to reach to targets on the body.

This longitudinal study investigated the effect of experience with tactile stimulation on infants' ability to reach to targets on the body, an important adaptive skill. Infants were provided weekly tactile stimulation on eight body locations from 4 to 8 months of age (N = 11), comparing their ability to reach to the body to infants in a control group who did not receive stimulation (N = 10). Infants who received stimulation were more likely to successfully reach targets on the body than controls by 7 months of age. These findings indicate that tactile stimulation facilitates the development of reaching to the body by allowing infants to explore the sensorimotor correlations emerging from the stimulation.

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.

Repetition effects in action planning reflect effector- but not hemisphere-specific coding.

Action choices are influenced by future and recent past action states. For example, when performing two actions in succession, response times (RTs) to initiate the second action are reduced when the same hand is used. These findings suggest the existence of effector-specific processing for action planning. However, given that each hand is primarily controlled by the contralateral hemisphere, the RT benefit might actually reflect effector-independent, hemisphere-specific rather than effector-specific repetition effects. Here, participants performed two consecutive movements, each with a hand or a foot, in one of two directions. Direction was specified in an egocentric reference frame (inward, outward) or in an allocentric reference frame (left, right). Successive actions were initiated faster when the same limb (e.g., left hand-left hand), but not the other limb of the same body side (e.g., left foot-left hand), executed the second action. The same-limb advantage was evident even when the two movements involved different directions, whether specified egocentrically or allocentrically. Corroborating evidence from computational modeling lends support to the claim that repetition effects in action planning reflect persistent changes in baseline activity within neural populations that encode effector-specific action plans.NEW & NOTEWORTHY Repeated hand use facilitates the initiation of successive actions (repetition effect). This finding has been interpreted as evidence for effector-specific action plans. However, given that each hand is primarily controlled by the contralateral hemisphere, any differences might reflect effector-independent, hemisphere-specific rather than effector-specific processing. We dissociated these alternatives by asking participants to perform successive actions with hands and feet and provide novel evidence that repetition effects in limb use truly reflect effector-specific coding.

Reach Trajectories Characterize Tactile Localization for Sensorimotor Decision Making.

Spatial target information for movement planning appears to be coded in a gaze-centered reference frame. In touch, however, location is initially coded with reference to the skin. Therefore, the tactile spatial location must be derived by integrating skin location and posture. It has been suggested that this recoding is impaired when the limb is placed in the opposite hemispace, for example, by limb crossing. Here, human participants reached toward visual and tactile targets located at uncrossed and crossed feet in a sensorimotor decision task. We characterized stimulus recoding by analyzing the timing and spatial profile of hand reaches. For tactile targets at crossed feet, skin-based information implicates the incorrect side, and only recoded information points to the correct location. Participants initiated straight reaches and redirected the hand toward a target presented in midflight. Trajectories to visual targets were unaffected by foot crossing. In contrast, trajectories to tactile targets were redirected later with crossed than uncrossed feet. Reaches to crossed feet usually continued straight until they were directed toward the correct tactile target and were not biased toward the skin-based target location. Occasional, far deflections toward the incorrect target were most likely when this target was implicated by trial history. These results are inconsistent with the suggestion that spatial transformations in touch are impaired by limb crossing, but are consistent with tactile location being recoded rapidly and efficiently, followed by integration of skin-based and external information to specify the reach target. This process may be implemented in a bounded integrator framework. SIGNIFICANCE STATEMENT: How do you touch yourself, for instance, to scratch an itch? The place you need to reach is defined by a sensation on the skin, but our bodies are flexible, so this skin location could be anywhere in 3D space. The movement toward the tactile sensation must therefore be specified by merging skin location and body posture. By investigating human hand reach trajectories toward tactile stimuli on the feet, we provide experimental evidence that this transformation process is quick and efficient, and that its output is integrated with the original skin location in a fashion consistent with bounded integrator decision-making frameworks.

Functional versus effector-specific organization of the human posterior parietal cortex: revisited.

It has been proposed that the posterior parietal cortex (PPC) is characterized by an effector-specific organization. However, strikingly similar functional MRI (fMRI) activation patterns have been found in the PPC for hand and foot movements. Because the fMRI signal is related to average neuronal activity, similar activation levels may result either from effector-unspecific neurons or from intermingled subsets of effector-specific neurons within a voxel. We distinguished between these possibilities using fMRI repetition suppression (RS). Participants made delayed, goal-directed eye, hand, and foot movements to visual targets. In each trial, the instructed effector was identical or different to that of the previous trial. RS effects indicated an attenuation of the fMRI signal in repeat trials. The caudal PPC was active during the delay but did not show RS, suggesting that its planning activity was effector independent. Hand and foot-specific RS effects were evident in the anterior superior parietal lobule (SPL), extending to the premotor cortex, with limb overlap in the anterior SPL. Connectivity analysis suggested information flow between the caudal PPC to limb-specific anterior SPL regions and between the limb-unspecific anterior SPL toward limb-specific motor regions. These results underline that both function and effector specificity should be integrated into a concept of PPC action representation not only on a regional but also on a fine-grained, subvoxel level.

Towards explaining spatial touch perception: Weighted integration of multiple location codes.

Touch is bound to the skin - that is, to the boundaries of the body. Yet, the activity of neurons in primary somatosensory cortex just mirrors the spatial distribution of the sensors across the skin. To determine the location of a tactile stimulus on the body, the body's spatial layout must be considered. Moreover, to relate touch to the external world, body posture has to be evaluated. In this review, we argue that posture is incorporated, by default, for any tactile stimulus. However, the relevance of the external location and, thus, its expression in behaviour, depends on various sensory and cognitive factors. Together, these factors imply that an external representation of touch dominates over the skin-based, anatomical when our focus is on the world rather than on our own body. We conclude that touch localization is a reconstructive process that is adjusted to the context while maintaining all available spatial information.

Neural correlates of tactile perception during pre-, peri-, and post-movement.

Tactile information is differentially processed over the various phases of goal-directed movements. Here, event-related potentials (ERPs) were used to investigate the neural correlates of tactile and visual information processing during movement. Participants performed goal-directed reaches for an object placed centrally on the table in front of them. Tactile and visual stimulation (100 ms) was presented in separate trials during the different phases of the movement (i.e. preparation, execution, and post-movement). These stimuli were independently delivered to either the moving or resting hand. In a control condition, the participants only performed the movement, while omission (i.e. movement-only) ERPs were recorded. Participants were instructed to ignore the presence or absence of any sensory events and to concentrate solely on the execution of the movement. Enhanced ERPs were observed 80-200 ms after tactile stimulation, as well as 100-250 ms after visual stimulation: These modulations were greatest during the execution of the goal-directed movement, and they were effector based (i.e. significantly more negative for stimuli presented to the moving hand). Furthermore, ERPs revealed enhanced sensory processing during goal-directed movements for visual stimuli as well. Such enhanced processing of both tactile and visual information during the execution phase suggests that incoming sensory information is continuously monitored for a potential adjustment of the current motor plan. Furthermore, the results reported here also highlight a tight coupling between spatial attention and the execution of motor actions.

Understanding effector selectivity in human posterior parietal cortex by combining information patterns and activation measures.

The posterior parietal cortex (PPC) has traditionally been viewed as containing separate regions for the planning of eye and limb movements, but recent neurophysiological and neuroimaging observations show that the degree of effector specificity is limited. This has led to the hypothesis that effector specificity in PPC is part of a more efficient than strictly modular organization, characterized by both distinct and common activations for different effectors. It is unclear, however, what differentiates the distinctions and commonalities in effector representations. Here, we used fMRI in humans to study the cortical representations involved in the planning of eye, hand, and foot movements. We used a novel combination of fMRI measures to assess the effector-related representational content of the PPC: a multivariate information measure, reflecting whether representations were distinct or common across effectors and a univariate activation measure, indicating which representations were actively involved in movement preparation. Active distinct representations were evident in areas previously reported to be effector specific: eye specificity in the posterior intraparietal sulcus (IPS), hand tuning in anterior IPS, and a foot bias in the anterior precuneus. Crucially, PPC regions responding to a particular effector also contained an active representation common across the other two effectors. We infer that rostral PPC areas do not code single effectors, but rather dichotomies of effectors. Such combinations of representations could be well suited for active effector selection, efficiently coding both a selected effector and its alternatives.

Oscillatory activity reflects differential use of spatial reference frames by sighted and blind individuals in tactile attention.

Touch can be localized either on the skin in anatomical coordinates, or, after integration with posture, in external space. Sighted individuals are thought to encode touch in both coordinate systems concurrently, whereas congenitally blind individuals exhibit a strong bias for using anatomical coordinates. We investigated the neural correlates of this differential dominance in the use of anatomical and external reference frames by assessing oscillatory brain activity during a tactile spatial attention task. The EEG was recorded while sighted and congenitally blind adults received tactile stimulation to uncrossed and crossed hands while detecting rare tactile targets at one cued hand only. In the sighted group, oscillatory alpha-band activity (8-12Hz) in the cue-target interval was reduced contralaterally and enhanced ipsilaterally with uncrossed hands. Hand crossing attenuated the degree of posterior parietal alpha-band lateralization, indicating that attention deployment was affected by external spatial coordinates. Beamforming suggested that this posture effect originated in the posterior parietal cortex. In contrast, cue-related lateralization of central alpha-band as well as of beta-band activity (16-24Hz) were unaffected by hand crossing, suggesting that these oscillations exclusively encode anatomical coordinates. In the blind group, central alpha-band activity was lateralized, but did not change across postures. The pattern of beta-band activity was indistinguishable between groups. Because the neural mechanisms for posterior alpha-band generation seem to be linked to developmental vision, we speculate that the lack of this neural mechanism in blind individuals is related to their preferred use of anatomical over external spatial codes in sensory processing.

Exploring the response code in a compatibility effect between physical size and left/right responses: The hand is more important than location.

The spatial-size association of response codes (SSARC) effect refers to the finding of better performance with the left hand to small stimuli and with the right hand to large stimuli, as compared to the reverse mapping. In the present study, we investigated which response coding is responsible for the emergence of the SSARC effect. We observed a SSARC effect only with response selection between hands but not between fingers of one hand, indicating that the responses are coded relative to the body midline. Furthermore, we observed a SSARC effect with parallel arms but not with crossed arms, suggesting that both the anatomical side of the effector and its external spatial position contribute to the response code. However, using a reaching task as compared to keypresses, the SSARC effect followed the arms, suggesting that the crucial spatial response code refers more strongly to the anatomical side of the effector rather than to the external spatial response position. These findings document a strong influence of anatomically- or body-based coding on the SSARC effect, are at odds with the proposition of a generalized magnitude system that utilizes a common, external spatial metric, and point toward a categorical nature of response codes underlying the SSARC effect. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Dynamic spatial coding in parietal cortex mediates tactile-motor transformation.

Movements towards touch on the body require integrating tactile location and body posture information. Tactile processing and movement planning both rely on posterior parietal cortex (PPC) but their interplay is not understood. Here, human participants received tactile stimuli on their crossed and uncrossed feet, dissociating stimulus location relative to anatomy versus external space. Participants pointed to the touch or the equivalent location on the other foot, which dissociates sensory and motor locations. Multi-voxel pattern analysis of concurrently recorded fMRI signals revealed that tactile location was coded anatomically in anterior PPC but spatially in posterior PPC during sensory processing. After movement instructions were specified, PPC exclusively represented the movement goal in space, in regions associated with visuo-motor planning and with regional overlap for sensory, rule-related, and movement coding. Thus, PPC flexibly updates its spatial codes to accommodate rule-based transformation of sensory input to generate movement to environment and own body alike.

Functional rather than effector-specific organization of human posterior parietal cortex.

Neurophysiological and neuroimaging studies have shown that the posterior parietal cortex (PPC) distinguishes between the planning of eye and hand movements. This distinction has usually been interpreted as evidence for a modular, effector-specific organization of this cerebral region. However, the eyes differ markedly from other body parts both in terms of their functional purpose and with regard to the spatial transformations required to plan goal-directed movements. PPC may therefore provide specialized subregions for eye movements, but distinguish less for other effectors. Using functional magnetic resonance imaging, we compared activity during memory-guided eye, hand, and foot movements in human participants. The results did not reveal any significant activation differences during the planning of hand and foot movements, except in the most anterior part of PPC [Brodmann's area (BA) 5], marginally extending into anterior BA 7/40. This region showed a lateral-to-medial gradient for hand versus foot movement planning. The limb-unspecific PPC regions were functionally connected with hand and foot motor regions. In contrast, a gradient-like organization was found for all of PPC for the planning of eye versus hand and foot movements. Although planning-related activity across the three effectors considerably overlapped, saccade planning activated occipitoparietal regions more than limb movements, whereas limb movements activated anterior regions of the superior parietal lobule more than saccades. We infer that PPC does not follow a strict effector-specific organization. Rather, the large-scale organization of this region might reflect the different computational constraints that need to be satisfied when planning eye and limb movements.

Allocation of Visuospatial Attention Indexes Evidence Accumulation for Reach Decisions.

Visuospatial attention is a prerequisite for the performance of visually guided movements: perceptual discrimination is regularly enhanced at target locations before movement initiation. It is known that this attentional prioritization evolves over the time of movement preparation; however, it is not clear whether this build-up simply reflects a time requirement of attention formation or whether, instead, attention build-up reflects the emergence of the movement decision. To address this question, we combined behavioral experiments, psychophysics, and computational decision-making models to characterize the time course of attention build-up during motor preparation. Participants (n = 46, 29 female) executed center-out reaches to one of two potential target locations and reported the identity of a visual discrimination target (DT) that occurred concurrently at one of various time-points during movement preparation and execution. Visual discrimination increased simultaneously at the two potential target locations but was modulated by the experiment-wide probability that a given location would become the final goal. Attention increased further for the location that was then designated as the final goal location, with a time course closely related to movement initiation. A sequential sampling model of decision-making faithfully predicted key temporal characteristics of attentional allocation. Together, these findings provide evidence that visuospatial attentional prioritization during motor preparation does not simply reflect that a spatial location has been selected as movement goal, but rather indexes the time-extended, cumulative decision that leads to the selection, hence constituting a link between perceptual and motor aspects of sensorimotor decisions.

The macaque ventral intraparietal area has expanded into three homologue human parietal areas.

The macaque ventral intraparietal area (VIP) in the fundus of the intraparietal sulcus has been implicated in a diverse range of sensorimotor and cognitive functions such as motion processing, multisensory integration, processing of head peripersonal space, defensive behavior, and numerosity coding. Here, we exhaustively review macaque VIP function, cytoarchitectonics, and anatomical connectivity and integrate it with human studies that have attempted to identify a potential human VIP homologue. We show that human VIP research has consistently identified three, rather than one, bilateral parietal areas that each appear to subsume some, but not all, of the macaque area's functionality. Available evidence suggests that this human "VIP complex" has evolved as an expansion of the macaque area, but that some precursory specialization within macaque VIP has been previously overlooked. The three human areas are dominated, roughly, by coding the head or self in the environment, visual heading direction, and the peripersonal environment around the head, respectively. A unifying functional principle may be best described as prediction in space and time, linking VIP to state estimation as a key parietal sensorimotor function. VIP's expansive differentiation of head and self-related processing may have been key in the emergence of human bodily self-consciousness.

Illusory tactile movement crosses arms and legs and is coded in external space.

Humans often misjudge where on the body a touch occurred. Theoretical accounts have ascribed such misperceptions to local interactions in peripheral and primary somatosensory neurons, positing that spatial-perceptual mechanisms adhere to limb boundaries and skin layout. Yet, perception often reflects integration of sensory signals with prior experience. On their trajectories, objects often touch multiple limbs; therefore, body-environment interactions should manifest in perceptual mechanisms that reflect external space. Here, we demonstrate that humans perceived the cutaneous rabbit illusion - the percept of multiple identical stimuli as hopping across the skin - along the Euclidian trajectory between stimuli on two body parts and regularly mislocalized stimuli from one limb to the other. A Bayesian model based on Euclidian, as opposed to anatomical, distance faithfully reproduced key aspects of participants' localization behavior. Our results suggest that prior experience of touch in space critically shapes tactile spatial perception and illusions beyond anatomical organization.

Common anatomical and external coding for hands and feet in tactile attention: evidence from event-related potentials.

Recent studies have suggested that the location of tactile stimuli is automatically recoded from anatomical into external coordinates, independent of the task requirements. However, research has mainly involved the two hands, which may not be representative for the whole body because they are excessively used for the visually guided manipulation of objects and tools. We recorded event-related potentials (ERPs) while participants received tactile stimuli to the hands and feet, but attended only one limb. The hands were placed near the feet either in an uncrossed or a crossed posture, thus varying the spatial distance of each hand from each foot. Centro-parietal ERPs 100-140 msec poststimulus were more positive when stimulating the anatomically same-side hand while attending a foot. They were also more positive when the Euclidean distance between the stimulated hand and the attended foot was small rather than large. When a foot was stimulated and a hand attended, a similar modulation of foot ERPs was observed for the right foot. To assess the spatial distance between two limbs in space, the external location of both must be known. The present ERP results therefore suggest that not only the hands but also other body parts are remapped into external coordinates. The use of both anatomical and external coordinates may facilitate the control of actions toward tactile events and the choice of the most suitable effector.

Presaccadic attention interferes with feature detection.

Preparing a saccadic eye movement to a particular spatial location enhances the perception of visual targets at this location and decreases perception of nearby targets prior to movement onset. This effect has been termed the orientation of pre-saccadic attention. Here, we investigated whether pre-saccadic attention influenced the detection of a simple visual feature-a process that has been hypothesized to occur without the need for attention. Participants prepared a saccade to a cued location and detected the occurrence of a "pop-out" feature embedded in distracters at the same or different location. The results show that preparing a saccade to a given location decreased detection of features at non-aimed-for locations, suggesting that the selection of a location as the next saccade endpoint influences sensitivity to basic visual features across the visual field.