Infant brain responses to felt and observed touch of hands and feet: an MEG study

There is growing interest concerning the ways in which the human body, both one's own and that of others, is represented in the developing human brain. In two experiments with 7‐month‐old infants, we employed advances in infant magnetoencephalography (MEG) brain imaging to address novel questions concerning body representations in early development. Experiment 1 evaluated the spatiotemporal organization of infants’ brain responses to being touched. A punctate touch to infants’ hands and feet produced significant activation in the hand and foot areas of contralateral primary somatosensory cortex as well as in other parietal and frontal areas. Experiment 2 explored infant brain responses to visually perceiving another person's hand or foot being touched. Results showed significant activation in early visual regions and also in regions thought to be involved in multisensory body and self–other processing. Furthermore, observed touch of the hand and foot activated the infant's own primary somatosensory cortex, although less consistently than felt touch. These findings shed light on aspects of early social cognition, including action imitation, which may build, at least in part, on infant neural representations that map equivalences between the bodies of self and other.     

Preliminary evidence for a fronto‐parietal dysfunction in able‐bodied participants with a desire for limb amputation

Background. Reports of able‐bodied participants with the persisting desire for limb amputation raise legal and ethical questions that are partly due to insufficient empirical knowledge about the condition. Here, we searched for potential neurological mechanisms in participants with desire for limb amputation in order to help develop adequate nosological classifications, diagnosis, and treatment. Methods. Semi‐structured interviews were carried out with 20 participants who self‐identified themselves as able‐bodied individuals desiring amputation of a limb. Results. The results suggest that amputation desire is not unspecific, but in most cases specific for a circumscribed part of the body. Most frequently affected was the leg, mostly on the left, non‐dominant side. Left‐sidedness and limb specificity was associated with elementary and complex somatosensory disturbances of the affected limb akin to those reported by neurological patients. The most frequent neurological co‐morbidity was migraine headache. Conclusions. These results document the existence of an unusual condition in able‐bodied participants characterized by a person's desire for the amputation of one or more particular limbs. Left‐sidedness, limb specificity and somatosensory disturbances of the affected limb are suggestive of abnormal brain mechanisms in right fronto‐parietal cortex. Based on this association we suggest that desire for limb amputation may be conceptualized as asomatognosia due to disturbed integration of multi‐sensory information of the affected body parts into a coherent cerebral representation of the own body. This suggestion has to be regarded with caution as we did not perform any neurological examination.     

Functional brain organization for number processing in pre‐verbal infants

Humans are born with the ability to mentally represent the approximate numerosity of a set of objects, but little is known about the brain systems that sub‐serve this ability early in life and their relation to the brain systems underlying symbolic number and mathematics later in development. Here we investigate processing of numerical magnitudes before the acquisition of a symbolic numerical system or even spoken language, by measuring the brain response to numerosity changes in pre‐verbal infants using functional near‐infrared spectroscopy (fNIRS). To do this, we presented infants with two types of numerical stimulus blocks: number change blocks that presented dot arrays alternating in numerosity and no change blocks that presented dot arrays all with the same number. Images were carefully constructed to rule out the possibility that responses to number changes could be due to non‐numerical stimulus properties that tend to co‐vary with number. Interleaved with the two types of numerical blocks were audio‐visual animations designed to increase attention. We observed that number change blocks evoked an increase in oxygenated hemoglobin over a focal right parietal region that was greater than that observed during no change blocks and during audio‐visual attention blocks. The location of this effect was consistent with intra‐parietal activity seen in older children and adults for both symbolic and non‐symbolic numerical tasks. A distinct set of bilateral occipital and middle parietal channels responded more to the attention‐grabbing animations than to either of the types of numerical stimuli, further dissociating the specific right parietal response to number from a more general bilateral visual or attentional response. These results provide the strongest evidence to date that the right parietal cortex is specialized for numerical processing in infancy, as the response to number is dissociated from visual change processing and general attentional processing.     

Cerebral blood flow in 5‐ to 8‐month‐olds: Regional tissue maturity is associated with infant affect

Infancy is marked by rapid neural and emotional development. The relation between brain function and emotion in infancy, however, is not well understood. Methods for measuring brain function predominantly rely on the BOLD signal; however, interpretation of the BOLD signal in infancy is challenging because the neuronal‐hemodynamic relation is immature. Regional cerebral blood flow (rCBF) provides a context for the infant BOLD signal and can yield insight into the developmental maturity of brain regions that may support affective behaviors. This study aims to elucidate the relations among rCBF, age, and emotion in infancy. One hundred and seven mothers reported their infants' (infant age M ± SD = 6.14 ± 0.51 months) temperament. A subsample of infants completed MRI scans, 38 of whom produced usable perfusion MRI during natural sleep to quantify rCBF. Mother‐infant dyads completed the repeated Still‐Face Paradigm, from which infant affect reactivity and recovery to stress were quantified. We tested associations of infant age at scan, temperament factor scores, and observed affect reactivity and recovery with voxel‐wise rCBF. Infant age was positively associated with CBF in nearly all voxels, with peaks located in sensory cortices and the ventral prefrontal cortex, supporting the formulation that rCBF is an indicator of tissue maturity. Temperamental Negative Affect and recovery of positive affect following a stressor were positively associated with rCBF in several cortical and subcortical limbic regions, including the orbitofrontal cortex and inferior frontal gyrus. This finding yields insight into the nature of affective neurodevelopment during infancy. Specifically, infants with relatively increased prefrontal cortex maturity may evidence a disposition toward greater negative affect and negative reactivity in their daily lives yet show better recovery of positive affect following a social stressor.     

Parietal versus temporal lobe components in spatial cognition: Setting the mid‐point of a horizontal line

Recent anatomo‐clinical correlation studies have extended to the superior temporal gyrus, the right hemisphere lesion sites associated with the left unilateral spatial neglect, in addition to the traditional posterior‐inferior‐parietal localization of the responsible lesion (supramarginal gyrus, at the temporo‐parietal junction). The study aimed at teasing apart, by means of repetitive transcranial magnetic stimulation (rTMS), the contribution of the inferior parietal lobule (angular gyrus versus supramarginal gyrus) and of the superior temporal gyrus of the right hemisphere, in making judgments about the mid‐point of a horizontal line, a widely used task for detecting and investigating spatial neglect. rTMS trains at 25 Hz frequency were delivered over the inferior parietal lobule (angular gyrus and supramarginal gyrus), the superior temporal gyrus and the anterior parietal lobe of the right hemisphere, in 10 neurologically unimpaired participants, performing a line bisection judgment task. rTMS of the inferior parietal lobule at the level of the supramarginal gyrus brought about a rightward error in the bisection judgment, ipsilateral to the side of the rTMS, with stimulation over the other sites being ineffective. The neural correlates of computing the mid‐point of a horizontal segment include the right supramarginal gyrus in the inferior parietal lobule and do not extend to the angular gyrus and the superior temporal gyrus. These rTMS data in unimpaired subjects constrain the evidence from lesion studies in brain‐damaged patients, emphasizing the major role of a subset of relevant regions.     

The role of body image and self‐perception in anorexia nervosa: the neuroimaging perspective

Anorexia nervosa is a severe psychiatric illness characterized by intense fear of gaining weight, relentless pursuit of thinness, deep concerns about food and a pervasive disturbance of body image. Functional magnetic resonance imaging tries to shed light on the neurobiological underpinnings of anorexia nervosa. This review aims to evaluate the empirical neuroimaging literature about self‐perception in anorexia nervosa. This narrative review summarizes a number of task‐based and resting‐state functional magnetic resonance imaging studies in anorexia nervosa about body image and self‐perception. The articles listed in references were searched using electronic databases (PubMed and Google Scholar) from 1990 to February 2016 using specific key words. All studies were reviewed with regard to their quality and eligibility for the review. Differences in brain activity were observed using body image perception and body size estimation tasks showing significant modifications in activity of specific brain areas (extrastriate body area, fusiform body area, inferior parietal lobule). Recent studies highlighted the role of emotions and self‐perception in anorexia nervosa and their neural substrate involving resting‐state networks and particularly frontal and posterior midline cortical structures within default mode network and insula. These findings open new horizons to understand the neural substrate of anorexia nervosa.     

Functional brain organization of working memory in adolescents varies in relation to family income and academic achievement

Working memory (WM) capacity reflects executive functions associated with performance on a wide range of cognitive tasks and education outcomes, including mathematics achievement, and is associated with dorsolateral prefrontal and parietal cortices. Here we asked if family income is associated with variation in the functional brain organization of WM capacity among adolescents, and whether that variation is associated with performance on a statewide test of academic achievement in mathematics. Participants were classified into higher‐income and lower‐income groups based on family income, and performed a WM task with a parametric manipulation of WM load (N‐back task) during functional magnetic resonance imaging (fMRI). Behaviorally, the higher‐income group had greater WM capacity and higher mathematics achievement scores. Neurally, the higher‐income group showed greater activation as a function of WM load in bilateral prefrontal, parietal, and other regions, although the lower‐income group exhibited greater activation at the lowest load. Both groups exhibited positive correlations between parietal activations and mathematics achievement scores, but only the higher‐income group exhibited a positive correlation between prefrontal activations and mathematics scores. Most of these findings were maintained when higher‐ and lower‐income groups were matched on WM task performance or nonverbal IQ. Findings indicate that the functional neural architecture of WM varies with family income and is associated with education measures of mathematics achievement.     

Spatial stimulus-response compatibility and coding of tactile motor events: influence of distance between stimulated and responding body parts, spatial complexity of the task and sex of subject

We studied spatial stimulus response compatibility in the somatosensory modality by instructing 16 men and 16 women to press a key using the left or the right thumb in response to a nonnoxious electric stimulation delivered either to the left or to the right little finger or, in different blocks, to the left or to the right malleolar region. The task was performed in compatible (stimulus and key-response on the same side of the corporeal midline) and in incompatible conditions (stimulus and key-response on opposite sides of the corporeal midline). In Exp. 1 subjects were tested while keeping their limbs in anatomic position; in Exp. 2 subjects performed the task while keeping the left upper and lower limbs on the right side and the right limbs on the left side of the bodily midline (crossed position). The compatibility effect was observed in both experiments and was higher for stimuli delivered to the little finger than to the malleolar region. This suggests that the cost of inhibiting compatible responses is maximal when stimulated and responding body parts are contiguous. Moreover, in the spatially most demanding task (Exp. 2) men outperformed women for both speed and accuracy suggesting a sex related specialisation in the spatial processing of somatosensory information.     

Infants' mu suppression during the observation of real and mimicked goal‐directed actions

Since their discovery in the early 1990s, mirror neurons have been proposed to be related to many social‐communicative abilities, such as imitation. However, research into the early manifestations of the putative neural mirroring system and its role in early social development is still inconclusive. In the current EEG study, mu suppression, generally thought to reflect activity in neural mirroring systems was investigated in 18‐ to 30‐month‐olds during the observation of object manipulations as well as mimicked actions. EEG power data recorded from frontal, central, and parietal electrodes were analysed. As predicted, based on previous research, mu wave suppression was found over central electrodes during action observation and execution. In addition, a similar suppression was found during the observation of intransitive, mimicked hand movements. To a lesser extent, the results also showed mu suppression at parietal electrode sites, over all three conditions. Mu wave suppression during the observation of hand movements and during the execution of actions was significantly correlated with quality of imitation, but not with age or language level.     

Children’s coding of human action: cognitive factors influencing imitaation in 3‐year‐old

We used imitation as a tool for investigating how young children code action. The study was designed to examine the errors children make in re‐enacting manual gestures they see. Thirty‐two 3‐year‐old children served as subjects. Each child was shown 24 gestures, generated by systematically crossing four factors: visual monitoring, spatial endpoint, movement path, and number of hands. The results showed no difference as a function of whether the children could visually monitor their own responses. Interestingly, children made significantly more errors when the adult's action terminated on a body part than they did when the same movement terminated near the body part. There were also significantly more errors when the demonstrated act involved crossing midline than when it did not, and more errors when it involved one hand rather than two hands. Our hypothesis is that human acts are coded in terms of goals. The goals are hierarchically organized, and because young children have difficulty simultaneously integrating multiple goals into one act they often re‐enact the goals that are ranked higher, which leads to the errors observed. We argue that imitation is an active reconstruction of perceived events and taps cognitive processing. We suggest that the goal‐based imitation in 3‐year‐olds is a natural developmental outgrowth of the perceptual–motor mapping and goal‐directed coding of human acts found in infancy.     

Hearing visuo‐tactile synchrony – Sound‐induced proprioceptive drift in the invisible hand illusion

The rubber hand illusion (RHI) and its variant the invisible hand illusion (IHI) are useful for investigating multisensory aspects of bodily self‐consciousness. Here, we explored whether auditory conditioning during an RHI could enhance the trisensory visuo‐tactile‐proprioceptive interaction underlying the IHI. Our paradigm comprised of an IHI session that was followed by an RHI session and another IHI session. The IHI sessions had two parts presented in counterbalanced order. One part was conducted in silence, whereas the other part was conducted on the backdrop of metronome beats that occurred in synchrony with the brush movements used for the induction of the illusion. In a first experiment, the RHI session also involved metronome beats and was aimed at creating an associative memory between the brush stroking of a rubber hand and the sounds. An analysis of IHI sessions showed that the participants’ perceived hand position drifted more towards the body‐midline in the metronome relative to the silent condition without any sound‐related session differences. Thus, the sounds, but not the auditory RHI conditioning, influenced the IHI. In a second experiment, the RHI session was conducted without metronome beats. This confirmed the conditioning‐independent presence of sound‐induced proprioceptive drift in the IHI. Together, these findings show that the influence of visuo‐tactile integration on proprioceptive updating is modifiable by irrelevant auditory cues merely through the temporal correspondence between the visuo‐tactile and auditory events.     

具身认知视角下语义理解中的身体角色:以研究方法为线索

具身认知认为,概念形成和语言理解等高级心理过程本质上是以感知觉和运动经验为基础的。神经影像学研究发现,理解身体动作词激活了支配该部分肢体的感觉运动脑区。理解手部、脚部和面部动作词能够相应激活支配手部、脚部和面部的感觉运动脑区,体现出一种身体动作词语义理解激活的脑区与真实身体动作激活脑区的耦合效应。临床和经颅磁刺激研究结果表明,感觉运动皮层的激活与身体动作词语义处理具有因果性作用。未来研究应关注身体动作词语义理解的具身程度以及相关语言疗法在临床患者机能恢复中所起的作用。     

Sex differences in pain and thermal sensitivity: the role of body size

Sex differences in heat-pain and thermal sensitivity were investigated in 32 women (20 to 60 years of age) and 32 men (17 to 63 years of age) who had no somatosensory impairments. Pain thresholds were measured with stimuli of two different durations (phasic and tonic). Warmth and cold thresholds were assessed as indices of thermal sensitivity. Stimulation was applied to the hand and to the foot by an apparatus containing a Peltier thermode. There were no sex differences in heat-pain thresholds. Women had significantly lower warmth thresholds than men (more pronounced on the foot than on the hand), but similar cold thresholds. Measures of body size (weight, height) correlated much more strongly with thermal than with pain sensitivity, and helped to explain the sex difference in the warmth threshold. A reduction of sex differences to body-measure differences appears likely, but could not be demonstrated unequivocally.     

Brain hyper‐connectivity and operation‐specific deficits during arithmetic problem solving in children with developmental dyscalculia

Developmental dyscalculia (DD) is marked by specific deficits in processing numerical and mathematical information despite normal intelligence (IQ) and reading ability. We examined how brain circuits used by young children with DD to solve simple addition and subtraction problems differ from those used by typically developing (TD) children who were matched on age, IQ, reading ability, and working memory. Children with DD were slower and less accurate during problem solving than TD children, and were especially impaired on their ability to solve subtraction problems. Children with DD showed significantly greater activity in multiple parietal, occipito‐temporal and prefrontal cortex regions while solving addition and subtraction problems. Despite poorer performance during subtraction, children with DD showed greater activity in multiple intra‐parietal sulcus (IPS) and superior parietal lobule subdivisions in the dorsal posterior parietal cortex as well as fusiform gyrus in the ventral occipito‐temporal cortex. Critically, effective connectivity analyses revealed hyper‐connectivity, rather than reduced connectivity, between the IPS and multiple brain systems including the lateral fronto‐parietal and default mode networks in children with DD during both addition and subtraction. These findings suggest the IPS and its functional circuits are a major locus of dysfunction during both addition and subtraction problem solving in DD, and that inappropriate task modulation and hyper‐connectivity, rather than under‐engagement and under‐connectivity, are the neural mechanisms underlying problem solving difficulties in children with DD. We discuss our findings in the broader context of multiple levels of analysis and performance issues inherent in neuroimaging studies of typical and atypical development.     

Enhancement of phonological memory following transcranial magnetic stimulation (TMS)

Phonologically similar items (mell, rell, gell) are more difficult to remember than dissimilar items (shen, floy, stap), likely because of mutual interference of the items in the phonological store. Low-frequency transcranial magnetic stimulation (TMS), guided by functional magnetic resonance imaging (fMRI) was used to disrupt this phonological confusion by stimulation of the left inferior parietal (LIP) lobule. Subjects received TMS or placebo stimulation while remembering sets of phonologically similar or dissimilar pseudo-words. Consistent with behavioral performance of patients with neurological damage, memory for phonologically similar, but not dissimilar, items was enhanced following TMS relative to placebo stimulation. Stimulation of a control region of the brain did not produce any changes in memory performance. These results provide new insights into how the brain processes verbal information by establishing the necessity of the inferior parietal region for optimal phonological storage. A mechanism is proposed for how TMS reduces phonological confusion and leads to facilitation of phonological memory.     

Cultivating a healthy neuro‐immune network: A health psychology approach

The field of psychoneuroimmunology (PNI) examines interactions among psychological and behavioral states, the brain, and the immune system. Research in PNI has elegantly documented effects of stress at multiple levels of the neuro‐immune network, with profound implications for both physical and mental health. In this review, we consider how the neuro‐immune network might be influenced by “positive” psychological and behavioral states, focusing on positive affect, eudaimonic well‐being, physical activity, and sleep. There is compelling evidence that these positive states and behaviors are associated with changes in immune activity in the body, including reductions in peripheral inflammatory processes relevant for physical health. Growing evidence from animal models also suggests effects of positive states on immune cells in the brain and the blood‐brain barrier, which then impact critical aspects of mood, cognition, and behavior. Tremendous advances are being made in our understanding of neuro‐immune dynamics; one of the central goals of this review is to highlight recent preclinical research in this area and consider how we can leverage these findings to investigate and cultivate a healthy neuro‐immune network in humans.     

The brain adapts to orthography with experience: evidence from English and Chinese

Using functional magnetic resonance imaging (fMRI), we examined the process of language specialization in the brain by comparing developmental changes in two contrastive orthographies: Chinese and English. In a visual word rhyming judgment task, we found a significant interaction between age and language in left inferior parietal lobule and left superior temporal gyrus, which was due to greater developmental increases in English than in Chinese. Moreover, we found that higher skill only in English children was correlated with greater activation in left inferior parietal lobule. These findings suggest that the regions associated with phonological processing are essential in English reading development. We also found greater developmental increases in English than in Chinese in left inferior temporal gyrus, suggesting refinement of this region for fine‐grained word form recognition. In contrast, greater developmental increases in Chinese than in English were found in right middle occipital gyrus, suggesting the importance of holistic visual‐orthographic analysis in Chinese reading acquisition. Our results suggest that the brain adapts to the special features of the orthography by engaging relevant brain regions to a greater degree over development.     

Updating of locations during whole-body rotations in patients with hemispatial neglect

Posterior parietal cortex lesions have been associated with both hemispatial neglect and spatialupdating deficits. Currently, the relation between these processes remains poorly understood. We tested the ability of parietal patients with neglect to update remembered target locations during passive whole-body rotations. The rotations and manual pointing responses were executed with and without vision. During the rotation, the remembered location stayed on the same side of the body midline or crossed the midline. Parietal patients generally underestimated rotations, as compared with control groups, but updated targets equally well on either side of the body midline, regardless of the amount of updating required. Once parietal patients have localized a target, they can use self-motion information to update its location, even if it passes into the region they typically neglect. This lack of contralesional updating effects contrasts with impairments in eye position updating found in previous work with parietal patients.     

Effects of motor intention on the perception of somatosensory events: A behavioural and functional magnetic resonance imaging study

The intention to execute a movement can modulate our perception of sensory events, and this modulation is observed ahead of both ocular and upper limb movements. However, theoretical accounts of these effects, and also the empirical data, are often contradictory. Accounts of “active touch”, and the premotor theory of attention, have emphasized how movement intention leads to enhanced perceptual processing at the target of a movement, or on the to-be-moved effector. By contrast, recent theories of motor control emphasize how internal “forward” model (FM) estimates may be used to cancel or attenuate sensory signals that arise as a result of self-generated movements. We used behavioural and functional brain imaging (functional magnetic resonance imaging, fMRI) to investigate how perception of a somatosensory stimulus differed according to whether it was delivered to a hand that was about to execute a reaching movement or the alternative, nonmoving, hand. The results of our study demonstrate that a somatosensory stimulus delivered to a hand that is being prepared for movement is perceived to have occurred later than when that same stimulus is delivered to a nonmoving hand. This result indicates that it takes longer for a tactile stimulus to be detected when it is delivered to a moving limb and may correspond to a change in perceptual threshold. Our behavioural results are paralleled by the results of our fMRI study that demonstrated that there were significantly reduced blood-oxygen-level-dependent (BOLD) responses within the parietal operculum and insula following somatosensory stimulation of the hand being prepared for movement, compared to when an identical stimulus was delivered to a nonmoving hand. These findings are consistent with the prediction of FM accounts of motor control that postulate that central sensory suppression of somatosensation accompanies self-generated limb movements, and with previous reports indicating that effects of sensory suppression are observed in higher order somatosensory regions.     

Effects of motor intention on the perception of somatosensory events: a behavioural and functional magnetic resonance imaging study

The intention to execute a movement can modulate our perception of sensory events, and this modulation is observed ahead of both ocular and upper limb movements. However, theoretical accounts of these effects, and also the empirical data, are often contradictory. Accounts of "active touch", and the premotor theory of attention, have emphasized how movement intention leads to enhanced perceptual processing at the target of a movement, or on the to-be-moved effector. By contrast, recent theories of motor control emphasize how internal "forward" model (FM) estimates may be used to cancel or attenuate sensory signals that arise as a result of self-generated movements. We used behavioural and functional brain imaging (functional magnetic resonance imaging, fMRI) to investigate how perception of a somatosensory stimulus differed according to whether it was delivered to a hand that was about to execute a reaching movement or the alternative, nonmoving, hand. The results of our study demonstrate that a somatosensory stimulus delivered to a hand that is being prepared for movement is perceived to have occurred later than when that same stimulus is delivered to a nonmoving hand. This result indicates that it takes longer for a tactile stimulus to be detected when it is delivered to a moving limb and may correspond to a change in perceptual threshold. Our behavioural results are paralleled by the results of our fMRI study that demonstrated that there were significantly reduced blood-oxygen-level-dependent (BOLD) responses within the parietal operculum and insula following somatosensory stimulation of the hand being prepared for movement, compared to when an identical stimulus was delivered to a nonmoving hand. These findings are consistent with the prediction of FM accounts of motor control that postulate that central sensory suppression of somatosensation accompanies self-generated limb movements, and with previous reports indicating that effects of sensory suppression are observed in higher order somatosensory regions.