Kosketuksen käsittely ihmisen tunto- ja kuuloaivokuorella

Tactile sensation plays an important role in everyday life. While the somatosensory system has been studied extensively, the majority of information has come from studies using animal models. Recent development of high-resolution anatomical and functional imaging techniques has enabled the non-invasive study of human somatosensory cortex and thalamus. This thesis provides new insights into the functional organization of the human brain areas involved in tactile processing using magnetoencephalography (MEG) and functional magnetic resonance imaging (fMRI). The thesis also demonstrates certain optimizations of MEG and fMRI methods. Tactile digit stimulation elicited stimulus-specific responses in a number of brain areas. Contralateral activation was observed in somatosensory thalamus (Study II), primary somatosensory cortex (SI; I, III, IV), and post-auditory belt area (III). Bilateral activation was observed in secondary somatosensory cortex (SII; II, III, IV). Ipsilateral activation was found in the post-central gyrus (area 2 of SI cortex; IV). In addition, phasic deactivation was observed within ipsilateral SI cortex and bilateral primary motor cortex (IV). Detailed investigation of the tactile responses demonstrated that the arrangement of distal-proximal finger representations in area 3b of SI in humans is similar to that found in monkeys (I). An optimized MEG approach was sufficient to resolve such fine detail in functional organization. The SII region appeared to contain double representations for fingers and toes (II). The detection of activations in the SII region and thalamus improved at the individual and group levels when cardiac-gated fMRI was used (II). Better detection of body part representations at the individual level is an important improvement, because identification of individual representations is crucial for studying brain plasticity in somatosensory areas. The posterior auditory belt area demonstrated responses to both auditory and tactile stimuli (III), implicating this area as a physiological substrate for the auditory-tactile interaction observed in earlier psychophysical studies. Comparison of different smoothing parameters (III) demonstrated that proper evaluation of co-activation should be based on individual subject analysis with minimal or no smoothing. Tactile input consistently influenced area 3b of the human ipsilateral SI cortex (IV). The observed phasic negative fMRI response is proposed to result from interhemispheric inhibition via trans-callosal connections. This thesis contributes to a growing body of human data suggesting that processing of tactile stimuli involves multiple brain areas, with different spatial patterns of cortical activation for different stimuli

Intra- and inter-hemispheric effective connectivity in the human somatosensory cortex during pressure stimulation

Background: Slow-adapting type I (SA-I) afferents deliver sensory signals to the somatosensory cortex during low-frequency (or static) mechanical stimulation. It has been reported that the somatosensory projection from SA-I afferents is effective and reliable for object grasping and manipulation. Despite a large number of neuroimaging studies on cortical activation responding to tactile stimuli mediated by SA-I afferents, how sensory information of such tactile stimuli flows over the somatosensory cortex remains poorly understood. In this study, we investigated tactile information processing of pressure stimuli between the primary (SI) and secondary (SII) somatosensory cortices by measuring effective connectivity using dynamic causal modeling (DCM). We applied pressure stimuli for 3 s to the right index fingertip of healthy participants and acquired functional magnetic resonance imaging (fMRI) data using a 3T MRI system. Results: DCM analysis revealed intra-hemispheric effective connectivity between the contralateral SI (cSI) and SII (cSII) characterized by both parallel (signal inputs to both cSI and cSII) and serial (signal transmission from cSI to cSII) pathways during pressure stimulation. DCM analysis also revealed inter-hemispheric effective connectivity among cSI, cSII, and the ipsilateral SII (iSII) characterized by serial (from cSI to cSII) and SII-level (from cSII to iSII) pathways during pressure stimulation. Conclusions: Our results support a hierarchical somatosensory network that underlies processing of low-frequency tactile information. The network consists of parallel inputs to both cSI and cSII (intra-hemispheric), followed by serial pathways from cSI to cSII (intra-hemispheric) and from cSII to iSII (inter-hemispheric). Importantly, our results suggest that both serial and parallel processing take place in tactile information processing of static mechanical stimuli as well as highlighting the contribution of callosal transfer to bilateral neuronal interactions in SII.open1

Functional Connectivity Evoked by Orofacial Tactile Perception of Velocity

The cortical representations of orofacial pneumotactile stimulation involve complex neuronal networks, which are still unknown. This study aims to identify the characteristics of functional connectivity (FC) evoked by three different saltatory velocities over the perioral and buccal surface of the lower face using functional magnetic resonance imaging in twenty neurotypical adults. Our results showed a velocity of 25 cm/s evoked stronger connection strength between the right dorsolateral prefrontal cortex and the right thalamus than a velocity of 5 cm/s. The decreased FC between the right secondary somatosensory cortex and right posterior parietal cortex for 5-cm/s velocity versus all three velocities delivered simultaneously (“All ON”) and the increased FC between the right thalamus and bilateral secondary somatosensory cortex for 65 cm/s vs “All ON” indicated that the right secondary somatosensory cortex might play a role in the orofacial tactile perception of velocity. Our results have also shown different patterns of FC for each seed (bilateral primary and secondary somatosensory cortex) at various velocity contrasts (5 vs 25 cm/s, 5 vs 65 cm/s, and 25 vs 65 cm/s). The similarities and differences of FC among three velocities shed light on the neuronal networks encoding the orofacial tactile perception of velocity

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

Brain mechanisms of audiotactile and audiomotor interactions

This thesis focuses on audiotactile integration, brain areas activated by vibrotactile stimulation, transfer of vibrotactile information to motor output, and reactivity of the human primary motor and somatosensory cortices in action observation. Human experience of the outside world results from integration of information obtained simultaneously via multiple senses. Accumulating evidence, from studies in both primates and humans, suggests that integration between different sensory modalities also occurs at early stages of cortical processing, in areas classically considered as purely unisensory. In Study I we studied integration between auditory and somatosensory systems. We showed, in a loudness-matching task, that subjects chose lower intensities for the probe than for the reference tone, when auditory and vibrotactile stimuli were presented simultaneously. In Studies II and III we explored brain areas involved in processing vibrotactile and tactile information, respectively. We showed that, besides primary and secondary somatosensory areas, auditory areas are also activated. In Study II we characterized the time course of brain activations and showed convergence of vibrotactile information to auditory areas. On the other hand, in Study III we identified, with good spatial accuracy, common neural substrates that process auditory and tactile information in auditory belt areas. In Study IV we assessed whether frequency information transfers from touch to vocal utterance in normal-hearing female adults. We demonstrated that such information transfer occurs clearly between 150–400 Hz. Based on findings in Studies II and III, we hypothesized that this transfer may involve at least primary and secondary somatosensory and auditory areas. Our social skills rely on the capability to understand others. In the human brain, the mirror-neuron system matches observation and execution of actions. This system comprises at least the inferior frontal gyrus, premotor areas, primary motor cortex, and the inferior parietal lobule. In Study V we investigated similarities in sensorimotor oscillatory activity between own, observed, and heard actions. We demonstrated that the primary motor cortex is activated before own and observed actions and stabilizes similarly. We also showed that rhythmic activity in the primary somatosensory cortex recovers later during own actions, which may be related to proprioceptive input and contribute to maintaining the sense of agency.reviewe

Event-related fMRI at 7T reveals overlapping cortical representations for adjacent fingertips in S1 of individual subjects

Recent fMRI studies of the human primary somatosensory cortex have been able to differentiate the cortical representations of different fingertips at a single-subject level. These studies did not, however, investigate the expected overlap in cortical activation due to the stimulation of different fingers. Here, we used an event-related design in six subjects at 7 Tesla to explore the overlap in cortical responses elicited in S1 by vibrotactile stimulation of the five fingertips. We found that all parts of S1 show some degree of spatial overlap between the cortical representations of adjacent or even nonadjacent fingertips. In S1, the posterior bank of the central sulcus showed less overlap than regions in the post-central gyrus, which responded to up to five fingertips. The functional properties of these two areas are consistent with the known layout of cytoarchitectonically defined subareas, and we speculate that they correspond to subarea 3b (S1 proper) and subarea 1, respectively. In contrast with previous fMRI studies, however, we did not observe discrete activation clusters that could unequivocally be attributed to different subareas of S1. Venous maps based on T2*-weighted structural images suggest that the observed overlap is not driven by extra-vascular contributions from large vein

Event-related fMRI at 7T reveals overlapping cortical representations for adjacent fingertips in S1 of individual subjects

Recent fMRI studies of the human primary somatosensory cortex have been able to differentiate the cortical representations of different fingertips at a single-subject level. These studies did not, however, investigate the expected overlap in cortical activation due to the stimulation of different fingers. Here, we used an event-related design in six subjects at 7 Tesla to explore the overlap in cortical responses elicited in S1 by vibrotactile stimulation of the five fingertips. We found that all parts of S1 show some degree of spatial overlap between the cortical representations of adjacent or even nonadjacent fingertips. In S1, the posterior bank of the central sulcus showed less overlap than regions in the post-central gyrus, which responded to up to five fingertips. The functional properties of these two areas are consistent with the known layout of cytoarchitectonically defined subareas, and we speculate that they correspond to subarea 3b (S1 proper) and subarea 1, respectively. In contrast with previous fMRI studies, however, we did not observe discrete activation clusters that could unequivocally be attributed to different subareas of S1. Venous maps based on T2*-weighted structural images suggest that the observed overlap is not driven by extra-vascular contributions from large vein

FMRI evidence of memory representations of somatosensory stimuli in the human brain

Distinct brain regions process innocuous vibration and cutaneous heat pain. The role of these areas in the perception of pain is still a matter of debate; and the role of these areas in the mediation of memory of somatosensory stimuli is uncertain and has not been studied with brain imaging in healthy human volunteers. All experiments described here, involved an experimental design, which included a delayed-discrimination paradigm and functional magnetic resonance imaging (fMRI). In manuscript #1, we aimed at unraveling the cerebral correlates of attention and spatial localization of innocuous vibrotactile stimuli applied to the right volar surface of the forearm. In this study, we report that increased degrees of attention to the vibrotactile stimuli were associated with heightened levels of activation in several brain areas. In manuscript #2, we investigated the short-term memory for sensory aspects (intensity and location) of cutaneous heat pain delivered to two areas (thenar and hypothenar eminences) of the palm of the right hand. In this experiment, the memory and control trials were presented in blocks, whereby the subjects could predict what trials were going to follow. This study revealed that the presentation of painful stimuli evoked activation in different brain regions than those activated during the online maintenance (interstimulus interval or ISI) of the intensity and spatial features of those stimuli; a process, which I will refer to short-term memory. In manuscript #3, we investigated again short-term memory for sensory aspects of heat pain (as in manuscript #2), but in this case, the memory and control trials were presented in a randomized order. In this study, we found that the perception and short-term memory of pain were processed by a comparable network of areas. The predictability of the memory and control trials may have contributed to these findings.La vibration inoffensive ainsi que la chaleur douloureuse cutanée sont traitées pardifférentes régions du cerveau. Le rôle de ces régions dans la perception de la douleurest controversé; et le rôle de ces régions dans la mémoire des stimuli somatosensorielsest incertain et n'a jamais encore été étudié en imagerie cérébrale chez des sujetshumains sains. Le design expérimental de toutes les études décrites ici comprenait unparadigme de 'delayed-discrimination' et l'imagerie par résonance magnétiquefonctionnelle (IRMf). L'étude #1 visait à élucider les corrélats cérébraux de l'attention etde la localisation spatiale des stimuli vibrotactiles inoffensifs présentés à la faceantérieure de l'avant-bras droit. Dans cette étude, nous avons trouvé que des degrésélevés d'attention portée aux stimuli vibrotactiles étaient associés à des niveaux accrusd'activation dans plusieurs zones du cerveau. Dans l'étude #2, nous avons enquêté surla mémoire à court-terme des caractéristiques sensorielles (intensité et emplacement)de la chaleur douloureuse cutanée présentée à deux endroits (éminences thénar ethypothénar) de la paume de la main droite. Dans cette étude, les essais mémoire etcontrôle étaient présentés en bloc, ou de sorte que les participants pouvaient prévoir dequel type serait le prochain essai. Cette étude a révélé que la présentation des stimulidouloureux a évoqué une activation de différentes régions cérébrales que celles quiétaient activées lors de la rétention de l'intensité et de l'emplacement des stimulationsdurant l'intervalle inter-stimuli (liS); un processus que je qualifierai de mémoire à courtterme.Dans l'étude #3, nous avons également enquêté sur la 'mémoire à court-termedes aspects sensoriels de la chaleur douloureuse (tout comme dans l'étude #2), maisdans ce cas, les essais mémoire et contrôle étaient présentés de façon aléatoire. Danscette étude, nous avons trouvé que la perception de la douleur ainsi que la mémoire àcourt-terme de la douleur étaient traitées par un réseau de régions semblable. Laprévisibilité des essais mémoire et contrôle peut avoir contribué à ce résultat

The in vivo functional neuroanatomy and neurochemistry of vibrotactile processing

Touch is a sense with which humans are able to actively explore the world around them. Primary somatosensory cortex (S1) processing has been studied to differing degrees at both the macroscopic and microscopic levels in both humans and animals. Both levels of enquiry have their advantages, but attempts to combine the two approaches are still in their infancy. One mechanism that is possibly involved in determining the reponse properties of neurons that are involved in sensory discrimination is inhibition by γ-aminobutyric acid (GABA). Several studies have shown that inhibition is an important mechanism to “tune” the response of neurons. Recently it has become possible to measure the concentration of GABA in vivo using edited Magnetic Resonance Spectroscopy (MRS), whereas magnetoencephalography (MEG) offers the possibility to look at changes in neuromagnetic activation with millisecond accuracy. With these methods we aimed to establish whether in vivo non-invasive neuroimaging can elucidate the underlying neuronal mechanisms of human tactile behaviour and to determine how such findings can be integrated with what is currently known from invasive methods. Edited GABA-MRS has shown that individual GABA concentration in S1 correlates strongly with tactile frequency discrimination. MEG was used to investigate the neuromagnetic correlates of a frequency discrimination paradigm in which we induced adaptation to a 25 Hz frequency. We showed that S1 is driven by the adapting stimulus and shows that neural rhythms are modulated as a result of adaptation. This is the first time that behavioural psychophysics of tactile adaptation has been investigated using complimentary neuroimaging methods. We combined different methods to complement both physiological and behavioural studies of tactile processing in S1 to investigate the factors involved in the neural dynamics of tactile processing and we show that non-invasive studies on humans can be used to understand physiological underpinnings of somatosensory processing.EThOS - Electronic Theses Online ServiceGBUnited Kingdo