Fine-grained nociceptive maps in primary somatosensory cortex

Topographic maps of the receptive surface are a fundamental feature of neural organization in many sensory systems. While touch is finely mapped in the cerebral cortex, it remains controversial how precise any cortical nociceptive map may be. Given that nociceptive innervation density is relatively low on distal skin regions such as the digits, one might conclude that the nociceptive system lacks fine representation of these regions. Indeed, only gross spatial organization of nociceptive maps has been reported so far. However, here we reveal the existence of fine-grained somatotopy for nociceptive inputs to the digits in human primary somatosensory cortex (SI). Using painful nociceptive-selective laser stimuli to the hand, and phase-encoded fMRI analysis methods, we observed somatotopic maps of the digits in contralateral SI. These nociceptive maps were highly aligned with maps of non-painful tactile stimuli, suggesting comparable cortical representations for, and possible interactions between, mechanoreceptive and nociceptive signals. Our findings may also be valuable for future studies tracking the timecourse and the spatial pattern of plastic changes in cortical organization involved in chronic pain

Symmetric Sensorimotor Somatotopy

BACKGROUND: Functional imaging has recently been used to investigate detailed somatosensory organization in human cortex. Such studies frequently assume that human cortical areas are only identifiable insofar as they resemble those measured invasively in monkeys. This is true despite the electrophysiological basis of the latter recordings, which are typically extracellular recordings of action potentials from a restricted sample of cells. METHODOLOGY/PRINCIPAL FINDINGS: Using high-resolution functional magnetic resonance imaging in human subjects, we found a widely distributed cortical response in both primary somatosensory and motor cortex upon pneumatic stimulation of the hairless surface of the thumb, index and ring fingers. Though not organized in a discrete somatotopic fashion, the population activity in response to thumb and index finger stimulation indicated a disproportionate response to fingertip stimulation, and one that was modulated by stimulation direction. Furthermore, the activation was structured with a line of symmetry through the central sulcus reflecting inputs both to primary somatosensory cortex and, precentrally, to primary motor cortex. CONCLUSIONS/SIGNIFICANCE: In considering functional activation that is not somatotopically or anatomically restricted as in monkey electrophysiology studies, our methodology reveals finger-related activation that is not organized in a simple somatotopic manner but is nevertheless as structured as it is widespread. Our findings suggest a striking functional mirroring in cortical areas conventionally ascribed either an input or an output somatotopic function

Effects of Fusion between Tactile and Proprioceptive Inputs on Tactile Perception

Tactile perception is typically considered the result of cortical interpretation of afferent signals from a network of mechanical sensors underneath the skin. Yet, tactile illusion studies suggest that tactile perception can be elicited without afferent signals from mechanoceptors. Therefore, the extent that tactile perception arises from isomorphic mapping of tactile afferents onto the somatosensory cortex remains controversial. We tested whether isomorphic mapping of tactile afferent fibers onto the cortex leads directly to tactile perception by examining whether it is independent from proprioceptive input by evaluating the impact of different hand postures on the perception of a tactile illusion across fingertips. Using the Cutaneous Rabbit Effect, a well studied illusion evoking the perception that a stimulus occurs at a location where none has been delivered, we found that hand posture has a significant effect on the perception of the illusion across the fingertips. This finding emphasizes that tactile perception arises from integration of perceived mechanical and proprioceptive input and not purely from tactile interaction with the external environment

Stroking or Buzzing? A Comparison of Somatosensory Touch Stimuli Using 7 Tesla fMRI.

Studying body representations in the brain helps us to understand how we humans relate to our own bodies. The in vivo mapping of the somatosensory cortex, where these representations are found, is greatly facilitated by the high spatial resolution and high sensitivity to brain activation available at ultra-high field. In this study, the use of different stimulus types for somatotopic mapping of the digits at ultra-high field, specifically manual stroking and mechanical stimulation, was compared in terms of sensitivity and specificity of the brain responses. Larger positive responses in digit regions of interest were found for manual stroking than for mechanical stimulation, both in terms of average and maximum t-value and in terms of number of voxels with significant responses to the tactile stimulation. Responses to manual stroking were higher throughout the entire post-central sulcus, but the difference was especially large on its posterior wall, i.e. in Brodmann area 2. During mechanical stimulation, cross-digit responses were more negative than during manual stroking, possibly caused by a faster habituation to the stimulus. These differences indicate that manual stroking is a highly suitable stimulus for fast somatotopic mapping procedures, especially if Brodmann area 2 is of interest

tDCS modulation of visually induced analgesia.

Multisensory interactions can produce analgesic effects. In particular, viewing one's own body reduces pain levels, perhaps because of changes in connectivity between visual areas specialized for body representation, and sensory areas underlying pain perception. We tested the causal role of the extrastriate visual cortex in triggering visually induced analgesia by modulating the excitability of this region with transcranial direct current stimulation (tDCS). Anodal, cathodal, or sham tDCS (2 mA, 10 min) was administered to 24 healthy participants over the right occipital or over the centro-parietal areas thought to be involved in the sensory processing of pain. Participants were required to rate the intensity of painful electrical stimuli while viewing either their left hand or an object occluding the left hand, both before and immediately after tDCS. We found that the analgesic effect of viewing the body was enhanced selectively by anodal stimulation of the occipital cortex. The effect was specific for the polarity and the site of stimulation. The present results indicate that visually induced analgesia may depend on neural signals from the extrastriate visual cortex

Brain (re)organisation following amputation:implications for phantom limb pain

Following arm amputation the region that represented the missing hand in primary somatosensory cortex (S1) becomes deprived of its primary input, resulting in changed boundaries of the S1 body map. This remapping process has been termed ‘reorganisation’ and has been attributed to multiple mechanisms, including increased expression of previously masked inputs. In a maladaptive plasticity model, such reorganisation has been associated with phantom limb pain (PLP). Brain activity associated with phantom hand movements is also correlated with PLP, suggesting that preserved limb functional representation may serve as a complementary process. Here we review some of the most recent evidence for the potential drivers and consequences of brain (re)organisation following amputation, based on human neuroimaging. We emphasise other perceptual and behavioural factors consequential to arm amputation, such as non-painful phantom sensations, perceived limb ownership, intact hand compensatory behaviour or prosthesis use, which have also been related to both cortical changes and PLP. We also discuss new findings based on interventions designed to alter the brain representation of the phantom limb, including augmented/virtual reality applications and brain computer interfaces. These studies point to a close interaction of sensory changes and alterations in brain regions involved in body representation, pain processing and motor control. Finally, we review recent evidence based on methodological advances such as high field neuroimaging and multivariate techniques that provide new opportunities to interrogate somatosensory representations in the missing hand cortical territory. Collectively, this research highlights the need to consider potential contributions of additional brain mechanisms, beyond S1 remapping, and the dynamic interplay of contextual factors with brain changes for understanding and alleviating PLP

A probabilistic atlas of finger dominance in the primary somatosensory cortex

With the advent of ultra-high field (7T), high spatial resolution functional MRI (fMRI) has allowed the differentiation of the cortical representations of each of the digits at an individual-subject level in human primary somatosensory cortex (S1). Here we generate a probabilistic atlas of the contralateral SI representations of the digits of both the left and right hand in a group of 22 right-handed individuals. The atlas is generated in both volume and surface standardised spaces from somatotopic maps obtained by delivering vibrotactile stimulation to each distal phalangeal digit using a travelling wave paradigm. Metrics quantify the likelihood of a given position being assigned to a digit (full probability map) and the most probable digit for a given spatial location (maximum probability map). The atlas is validated using a leave-one-out cross validation procedure. Anatomical variance across the somatotopic map is also assessed to investigate whether the functional variability across subjects is coupled to structural differences. This probabilistic atlas quantifies the variability in digit representations in healthy subjects, finding some quantifiable separability between digits 2, 3 and 4, a complex overlapping relationship between digits 1 and 2, and little agreement of digit 5 across subjects. The atlas and constituent subject maps are available online for use as a reference in future neuroimaging studies

Unraveling the spatiotemporal brain dynamics during a simulated reach-to-eat task

The reach-to-eat task involves a sequence of action components including looking, reaching, grasping, and feeding. While cortical representations of individual action components have been mapped in human functional magnetic resonance imaging (fMRI) studies, little is known about the continuous spatiotemporal dynamics among these representations during the reach-to-eat task. In a periodic event-related fMRI experiment, subjects were scanned while they reached toward a food image, grasped the virtual food, and brought it to their mouth within each 16-s cycle. Fourier-based analysis of fMRI time series revealed periodic signals and noise distributed across the brain. Independent component analysis was used to remove periodic or aperiodic motion artifacts. Timefrequency analysis was used to analyze the temporal characteristics of periodic signals in each voxel. Circular statistics was then used to estimate mean phase angles of periodic signals and select voxels based on the distribution of phase angles. By sorting mean phase angles across regions, we were able to show the real-time spatiotemporal brain dynamics as continuous traveling waves over the cortical surface. The activation sequence consisted of approximately the following stages: (1) stimulus related activations in occipital and temporal cortices; (2) movement planning related activations in dorsal premotor and superior parietal cortices; (3) reaching related activations in primary sensorimotor cortex and supplementary motor area; (4) grasping related activations in postcentral gyrus and sulcus; (5) feeding related activations in orofacial areas. These results suggest that phase-encoded design and analysis can be used to unravel sequential activations among brain regions during a simulated reach-to-eat task