The neural circuits underlying initial sensory processing of somatic information are relatively well understood. In contrast, the processes that go beyond primary somatosensation to create more abstract representations related to the body are less clear. In this review, we focus on two classes of higher-order processing beyond somatosensation. Somatoperception refers to the process of perceiving the body itself, and particularly of ensuring somatic perceptual constancy.We review three key elements of somatoperception: (a) remapping information from the body surface into an egocentric reference frame (b) exteroceptive perception of objects in the external world through their contact with the body and (c) interoceptive percepts about the nature and state of the body itself. Somatorepresentation, in contrast, refers to the essentially cognitive process of constructing semantic knowledge and attitudes about the body, including: (d) lexicalsemantic knowledge about bodies generally and one's own body specifically, (e) configural knowledge about the structure of bodies, (f) emotions and attitudes directed towards one's own body, and (g) the link between physical body and psychological self. We review a wide range of neuropsychological, neuroimaging and neurophysiological data to explore the dissociation between these different aspects of higher somatosensory function.Body Beyond SI 3

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.

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.