Our body is a unique entity by which we interact with the external world. Consequently, the way we represent our body has profound implications in the way we process and locate sensations and in turn perform appropriate actions. The body can be the subject, but also the object of our experience, providing information from sensations on the body surface and viscera, but also knowledge of the body as a physical object. However, the extent to which different senses contribute to constructing the rich and unified body representations we all experience remains unclear. In this review, we aim to bring together recent research showing important roles for several different sensory modalities in constructing body representations. At the same time, we hope to generate new ideas of how and at which level the senses contribute to generate the different levels of body representations and how they interact. We will present an overview of some of the most recent neuropsychological evidence about multisensory control of pain, and the way that visual, auditory, vestibular and tactile systems contribute to the creation of coherent representations of the body. We will focus particularly on some of the topics discussed in the symposium on Multimodal Contributions to Body Representation held on the 15th International Multisensory Research Forum (2015, Pisa, Italy).

According to current textbook knowledge, the primary somatosensory cortex (SI) supports unilateral tactile representations, whereas structures beyond SI, in particular the secondary somatosensory cortex (SII), support bilateral tactile representations. However, dexterous and well-coordinated bimanual motor tasks require early integration of bilateral tactile information. Sequential processing, first of unilateral and subsequently of bilateral sensory information, might not be sufficient to accomplish these tasks. This view of sequential processing in the somatosensory system might therefore be questioned, at least for demanding bimanual tasks. Evidence from the last 15 years is forcing a revision of this textbook notion. Studies in animals and humans indicate that SI is more than a simple relay for unilateral sensory information and, together with SII, contributes to the integration of somatosensory inputs from both sides of the body. Here, we review a series of recent works from our own and other laboratories in favour of interactions between tactile stimuli on the two sides of the body at early stages of processing. We focus on tactile processing, although a similar logic may also apply to other aspects of somatosensation. We begin by describing the basic anatomy and physiology of interhemispheric transfer, drawing on neurophysiological studies in animals and behavioural studies in humans that showed tactile interactions between body sides, both in healthy and in brain-damaged individuals. Then we describe the neural substrates of bilateral interactions in somatosensation as revealed by neurophysiological work in animals and neuroimaging studies in humans (i.e., functional magnetic resonance imaging, magnetoencephalography, and transcranial magnetic stimulation). Finally, we conclude with considerations on the dilemma of how efficiently integrating bilateral sensory information at early processing stages can coexist with more lateralized representations of somatosensory input, in the context of motor control.

The processing of touch depends of multiple factors, such as the properties of the skin and type of receptors stimulated, as well as features related to the actual configuration and shape of the body itself. A large body of research has focused on the effect that the nature of the stimuli has on tactile processing. Less research, however, has focused on features beyond the nature of the touch. In this review, we focus on some features related to the body that have been investigated for less time and in a more fragmented way. These include the symmetrical quality of the two sides of the body, the postural configuration of the body, as well as the size and shape of different body parts. We will describe what we consider three key aspects: (1) how and at which stages tactile information is integrated between different parts and sides of the body; (2) how tactile signals are integrated with online and stored postural configurations of the body, regarded as priors; (3) and how tactile signals are integrated with representations of body size and shape. Here, we describe how these different body dimensions affect integration of tactile information as well as guide motor behavior by integrating them in a single model of tactile processing. We review a wide range of neuropsychological, neuroimaging, and neurophysiological data and suggest a revised model of tactile integration on the basis of the one proposed previously by Longo et al.

Although the somatosensory homunculus is a classically used description of the way somatosensory inputs are processed in the brain, the actual contributions of primary (SI) and secondary (SII) somatosensory cortices to the spatial coding of touch remain poorly understood. We studied adaptation of the fMRI BOLD response in the somatosensory cortex by delivering pairs of vibrotactile stimuli to the finger tips of the index and middle fingers. The first stimulus (adaptor) was delivered either to the index or to the middle finger of the right or left hand, and the second stimulus (test) was always administered to the left index finger. The overall BOLD response evoked by the stimulation was primarily contralateral in SI and was more bilateral in SII. However, our fMRI adaptation approach also revealed that both somatosensory cortices were sensitive to ipsilateral as well as to contralateral inputs. SI and SII adapted more after subsequent stimulation of homologous as compared with nonhomologous fingers, showing a distinction between different fingers. Most importantly, for both somatosensory cortices, this finger-specific adaptation occurred irrespective of whether the tactile stimulus was delivered to the same or to different hands. This result implies integration of contralateral and ipsilateral somatosensory inputs in SI as well as in SII. Our findings suggest that SI is more than a simple relay for sensory information and that both SI and SII contribute to the spatial coding of touch by discriminating between body parts (fingers) and by integrating the somatosensory input from the two sides of the body (hands).