External Spatial Representations Research Articles

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.

Tactile localization performance in children with developmental coordination disorder (DCD) corresponds to their motor skill and not their cognitive ability

When localizing touches to the hands, typically developing children and adults show a “crossed hands effect” whereby identifying which hand received a tactile stimulus is less accurate when the hands are crossed than uncrossed. This demonstrates the use of an external frame of reference for locating touches to one’s own body. Given that studies indicate that developmental vision plays a role in the emergence of external representations of touch, and reliance on vision for representing the body during action is atypical in developmental coordination disorder (DCD), we investigated external spatial representations of touch in children with DCD using the “crossed hands effect”. Nineteen children with DCD aged 7–11years completed a tactile localization task in which posture (uncrossed, crossed) and view (hands seen, unseen) were varied systematically. Their performance was compared to that of 35 typically developing controls (19 of a similar age and cognitive ability, and 16 of a younger age but similar fine motor ability). Like controls, the DCD group exhibited a crossed hands effect, whilst their overall tactile localization performance was weaker than their peers of similar age and cognitive ability, but in line with younger controls of similar motor ability. For children with movement difficulties, these findings indicate tactile localization impairments in relation to age expectations, but apparently typical use of an external reference frame for localizing touch.

A three-dimensional spatial characterization of the crossed-hands deficit

To perceive the location of touch in space, we integrate information about skin-location with information about the location of that body part in space. Most research investigating this process of tactile spatial remapping has used the so-called crossed-hands deficit, in which the ability to judge the temporal order of touches on the two hands is impaired when the arms are crossed. This posture induces a conflict between skin-based and tactile external spatial representations, specifically in the left-right dimension. Thus, it is unknown whether touch is affected by posture when spatial relations other than the right-left dimension are available. Here, we tested the extent to which the crossed-hands deficit is a measure of tactile remapping, reflecting tactile encoding in three-dimensional space. Participants judged the temporal order of tactile stimuli presented to crossed and uncrossed hands. The arms were placed at different elevations (up-down dimension; Experiments 1 and 2), or at different distances from the body in the depth plane (close-far dimension; Experiment 3). The crossed-hands deficit was reduced when other sources of spatial information, orthogonal to the left-right dimension (i.e., close-far, up-down), were available. Nonetheless, the deficit persisted in all conditions, even when processing of non-conflicting information in the close-far or up-down dimensions was enough to solve the task. Together, these results demonstrate that the processing underlying the crossed-hands deficit is related to the encoding of tactile localization in three-dimensional space, rather than related uniquely to the cost of processing information in the right-left dimension. Furthermore, the persistence of the crossing effect provides evidence for automatic integration of all available information during the encoding of tactile information.

Dynamic Tuning of Tactile Localization to Body Posture

Localizing touch in space is essential for goal-directed action. Because body posture changes, the brain must transform tactile coordinates from an initial skin-based representation to external space by integrating information about current posture. This process, referred to as tactile remapping, generally results in accurate localization, but accuracy drops when skin-based and external spatial representations of touch are conflicting, e.g., after crossing the limbs. Importantly, frequent experience of such postures can improve localization. This suggests that remapping may not only integrate current sensory input but also prior experience. Here, we demonstrate that this can result in rapid changes in localization performance over the course of few trials. We obtained an implicit measure of tactile localization by studying the perceived temporal order of two touches, one on each hand. Crucially, we varied the number of consecutive trials during which participants held their arms crossed or uncrossed. As expected, accuracy dropped immediately after the arms had been crossed. Importantly, this was followed by a progressive recovery if posture was maintained, despite the absence of performance feedback. Strikingly, a significant improvement was already evident in the localization of the second pair of touches. This rapid improvement required preceding touch in the same posture and did not occur merely as a function of time. Moreover, even touches that were not task relevant led to improved localization of subsequent touch. Our findings show that touches are mapped from skin to external space as a function of recent tactile experience.

Landmarks in nature to support wayfinding: the effects of seasons and experimental methods

Landmarks constitute an essential basis for a structural understanding of the spatial environment. Therefore, they are crucial factors in external spatial representations such as maps and verbal route descriptions, which are used to support wayfinding. However, selecting landmarks for these representations is a difficult task, for which an understanding of how people perceive and remember landmarks in the environment is needed. We investigated the ways in which people perceive and remember landmarks in nature using the thinking aloud and sketch map methods during both the summer and the winter seasons. We examined the differences between methods to identify those landmarks that should be selected for external spatial representations, such as maps or route descriptions, in varying conditions. We found differences in the use of landmarks both in terms of the methods and also between the different seasons. In particular, the participants used passage and tree-related landmarks at significantly different frequencies with the thinking aloud and sketch map methods. The results are likely to reflect the different roles of the landmark groups when using the two methods, but also the differences in counting landmarks when using both methods. Seasonal differences in the use of landmarks occurred only with the thinking aloud method. Sketch maps were drawn similarly in summertime and wintertime; the participants remembered and selected landmarks similarly independent of the differences in their perceptions of the environment due to the season. The achieved results may guide the planning of external spatial representations within the context of wayfinding as well as when planning further experimental studies.

What decision-making models can tell us about tactile remapping

To locate an object by touch, for example when fumbling around in the dark, or to react to a tactile event on the skin, somatosensory information must be quickly combined with the current position of the hand relative to the rest of the body. Not much is know yet about the time course of this integration between touch and body posture (e.g., proprioception), called tactile remapping. A tactile localization task in which there is a spatial conflict between the somatotopic and the external spatial representations can provide some constraints on the timing of the proprioceptive signal (necessary for the remapping) relative to the tactile input. Here, we construe this localization task as a perceptual discrimination problem. By making use of established models of decision-making, we aim to shed light on the internal processes during tactile remapping. In the experiment, participants had to make a speeded saccade towards a brief somatosensory stimulus applied to the finger of one hand, while their arms were either crossed or uncrossed about the body midline. The eye movement trajectories were recorded and revealed that especially in the crossed condition fast saccades occasionally started toward the incorrect (anatomically congruent) side and were corrected in-flight, resulting in a turn-around saccade (about 250 ms after tactile onset, which might reflect the completion of tactile remapping). Similar curved traces have been observed recently in a random-dot motion (RDM) discrimination task were subjects had to report their decision by moving a handle to a left or right target [1]. While in the RDM task the curved traces were interpreted as a change of mind, direction-reversal here is thought to be induced by the (delayed) proprioceptive signal. With crossed hands, the current body posture needs to be taken into account and the representation of touch changes through time due to the remapping [2]. In the uncrossed posture remapping is trivial as the anatomical and external reference frames are aligned. In the framework of decision-making and in line with [1], we treat the initial direction of a saccade as a first choice, caused by crossing a decision-threshold. Accordingly, we apply two computational models able to account for changes of mind, a simple drift-diffusion model [1] and a biophysically-realistic nonlinear attractor model [3], to fit experimental reaction-time distributions and performance. In the models, turn-around saccades originate from a second threshold crossing if the accumulated evidence of the initially losing alternative overtakes the first choice. Remapping only the outcome of the decision to a different motor response in the crossed-hand condition proved insufficient to explain the differences in reaction times and performance to the uncrossed case. This suggests that internal differences between the two conditions are already present prior to the decision-making stage. From the modeling perspective the proprioceptive signal thus changes the input (evidence) for deciding on a saccade to the left or right. Allowing only the model inputs to vary between crossed and uncrossed conditions, we gain predictions on the relative size and timing of the internal proprioceptive signal for tactile remapping.

The Posterior Parietal Cortex Remaps Touch into External Space

Localizing tactile events in external space is required for essential functions such as orienting, haptic exploration, and goal-directed action in peripersonal space. In order to map somatosensory input into a spatiotopic representation, information about skin location must be integrated with proprioceptive information about body posture. We investigated the neural bases of this tactile remapping mechanism in humans by disrupting neural activity in the putative human homolog of the monkey ventral intraparietal area (hVIP), within the right posterior parietal cortex (rPPC), which is thought to house external spatial representations. Participants judged the elevation of touches on their (unseen) forearm relative to touches on their face. Arm posture was passively changed along the vertical axis, so that elevation judgments required the use of an external reference frame. Single-pulse transcranial magnetic stimulation (TMS) over the rPPC significantly impaired performance compared to a control site (vertex). Crucially, proprioceptive judgments of arm elevation or tactile localization on the skin remained unaffected by rPPC TMS. This selective disruption of tactile remapping suggests a distinct computational process dissociable from pure proprioceptive and somatosensory localization. Furthermore, this finding highlights the causal role of human PPC, putatively VIP, in remapping touch into external space.