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As cold actions (i.e. actions devoid of an emotional content), also emotions are expressed with different vitality forms. For example, when an individual experiences a positive emotion, such as laughing as expression of happiness, this emotion can be conveyed to others by different intensities of face expressions and body postures. In the present study, we investigated whether the observation of emotions, expressed with different vitality forms, activates the same neural structures as those involved in cold action vitality forms processing. To this purpose, we carried out a functional magnetic resonance imaging study in which participants were tested in 2 conditions: emotional and non-emotional laughing both conveying different vitality forms. There are 3 main results. First, the observation of emotional and non-emotional laughing conveying different vitality forms activates the insula. Second, the observation of emotional laughing activates a series of subcortical structures known to be related to emotions. Furthermore, a region of interest analysis carried out in these structures reveals a significant modulation of the blood-oxygen-leveldependent (BOLD) signal during the processing of different vitality forms exclusively in the right amygdala, right anterior thalamus/hypothalamus, and periaqueductal gray. Third, in a subsequent electromyography study, we found a correlation between the zygomatic muscles activity and BOLD signal in the right amygdala only., (© The Author(s) 2022. Published by Oxford University Press.)

In previous studies on auditory vitality forms, we found that listening to action verbs pronounced gently or rudely, produced, relative to a neutral robotic voice, activation of the dorso-central insula. One might wonder whether this insular activation depends on the conjunction of action verbs and auditory vitality forms, or whether auditory vitality forms are sufficient per se to activate the insula. To solve this issue, we presented words not related to actions such as concrete nouns (e.g.,"ball"), pronounced gently or rudely. No activation of the dorso-central insula was found. As a further step, we examined whether interjections, i.e., speech stimuli conveying communicative intention (e.g., "hello"), pronounced with different vitality forms, would be able to activate, relative to control, the insula. The results showed that stimuli conveying a communicative intention, pronounced with different auditory vitality forms activate the dorsal-central insula. These data deepen our understanding of the vitality forms processing, showing that insular activation is not specific to action verbs, but can be also activated by speech acts conveying communicative intention such as interjections. These findings also show the intrinsic social nature of vitality forms because activation of the insula was not observed in the absence of a communicative intention., (© The Author(s) 2021. Published by Oxford University Press.)

Although clinical neuroscience and the neuroscience of consciousness have long sought mechanistic explanations of tactile-awareness disorders, mechanistic insights are rare, mainly because of the difficulty of depicting the fine-grained neural dynamics underlying somatosensory processes. Here, we combined the stereo-EEG responses to somatosensory stimulation with the lesion mapping of patients with a tactile-awareness disorder, namely tactile extinction. Whereas stereo-EEG responses present different temporal patterns, including early/phasic and long-lasting/tonic activities, tactile-extinction lesion mapping co-localizes only with the latter. Overlaps are limited to the posterior part of the perisylvian regions, suggesting that tonic activities may play a role in sustaining tactile awareness. To assess this hypothesis further, we correlated the prevalence of tonic responses with the tactile-extinction lesion mapping, showing that they follow the same topographical gradient. Finally, in parallel with the notion that visuotactile stimulation improves detection in tactile-extinction patients, we demonstrated an enhancement of tonic responses to visuotactile stimuli, with a strong voxel-wise correlation with the lesion mapping. The combination of these results establishes tonic responses in the parietal operculum as the ideal neural correlate of tactile awareness., (© The Author(s) (2021). Published by Oxford University Press on behalf of the Guarantors of Brain.)

Laughter is a complex motor behavior occurring in both emotional and nonemotional contexts. Here, we investigated whether the different functions of laughter are mediated by distinct networks and, if this is the case, which are the white matter tracts sustaining them. We performed a multifiber tractography investigation placing seeds in regions involved in laughter production, as identified by previous intracerebral electrical stimulation studies in humans: the pregenual anterior cingulate (pACC), ventral temporal pole (TPv), frontal operculum (FO), presupplementary motor cortex, and ventral striatum/nucleus accumbens (VS/NAcc). The primary motor cortex (M1) and two subcortical territories were also studied to trace the descending projections. Results provided evidence for the existence of two relatively distinct networks. A first network, including pACC, TPv, and VS/NAcc, is interconnected through the anterior cingulate bundle, the accumbofrontal tract, and the uncinate fasciculus, reaching the brainstem throughout the mamillo-tegmental tract. This network is likely involved in the production of emotional laughter. A second network, anchored to FO and M1, projects to the brainstem motor nuclei through the internal capsule. It is most likely the neural basis of nonemotional and conversational laughter. The two networks interact throughout the pre-SMA that is connected to both pACC and FO., (© The Author(s) 2020. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

It has been recently found that the human dorso-central insular cortex contributes to the execution and recognition of the affective component of hand actions, most likely through modulation of the activity of the parieto-frontal circuits. While the anatomical connections between the hand representation of the insula and, the parietal and frontal regions controlling reaching/grasping actions is well assessed in the monkey, it is unknown the existence of a homolog circuit in humans. In the present study, we performed a multifiber tractography investigation to trace the tracts possibly connecting the insula to the parieto-frontal circuits by locating seeds in the parietal, premotor, and prefrontal nodes of the reaching/grasping network, in both humans and monkeys. Results showed that, in both species, the insula is connected with the cortical action execution/recognition circuit by similar white matter tracts, running in parallel to the third branch of the superior longitudinal fasciculus and the anterior segment of the arcuate fasciculus., (© The Author(s) 2018. Published by Oxford University Press. All rights reserved. For Permissions, please e-mail: journals.permissions@oup.com.)

The cingulate cortex is a mosaic of different anatomical fields, whose functional characterization is still a matter of debate. In humans, one method that may provide useful insights on the role of the different cingulate regions, and to tackle the issue of the functional differences between its anterior, middle and posterior subsectors, is intracortical electrical stimulation. While previous reports showed that a variety of integrated behaviours could be elicited by stimulating the midcingulate cortex, little is known about the effects of the electrical stimulation of anterior and posterior cingulate regions. Moreover, the internal arrangement of different behaviours within the midcingulate cortex is still unknown. In the present study, we extended previous stimulation studies by retrospectively analysing all the clinical manifestations induced by intracerebral high frequency electrical stimulation (50 Hz, pulse width: 1 ms, 5 s, current intensity: average intensity of 2.7 ± 0.7 mA, biphasic) of the entire cingulate cortex in a cohort of 329 drug-resistant epileptic patients (1789 stimulation sites) undergoing stereo-electroencephalography for a presurgical evaluation. The large number of patients, on one hand, and the accurate multimodal image-based localization of stereo-electroencephalography electrodes, on the other hand, allowed us to assign specific functional properties to modern anatomical subdivisions of the cingulate cortex. Behavioural or subjective responses were elicited from the 32.3% of all cingulate sites, mainly located in the pregenual and midcingulate regions. We found clear functional differences between the pregenual part of the cingulate cortex, hosting the majority of emotional, interoceptive and autonomic responses, and the anterior midcingulate sector, controlling the majority of all complex motor behaviours. Particularly interesting was the 'actotopic' organization of the anterior midcingulate sector, arranged along the ventro-dorsal axis: (i) whole-body behaviours directed to the extra-personal space, such as getting-up impulses, were elicited ventrally, close to the corpus callosum; (ii) hand actions in the peripersonal space were evoked by the stimulation of the intermediate position; and (iii) body-directed actions were induced by the stimulation of the dorsal branch of the cingulate sulcus. The caudal part of the midcingulate cortex and the posterior cingulate cortex were, in contrast, poorly excitable, and mainly devoted to sensory modalities. In particular, the caudal part of the midcingulate cortex hosted the majority of vestibular responses, while posterior cingulate cortex was the principal recipient of visual effects. We will discuss our data in the light of current controversies on the role of the cingulate cortex in cognition and emotion.

Action and speech may take different forms, being expressed, for example, gently or rudely. These aspects of social communication, named vitality forms, have been little studied in neuroscience. In the present functional magnetic resonance imaging study, we investigated the role of insula in processing action and speech vitality forms. In speech runs, participants were asked to listen or imaging themselves to pronounce action verbs gently or rudely. In action runs, they were asked to observe or imaging themselves to perform actions gently or rudely. The results showed that, relative to controls, there was an activation of the dorso-central insula in both tasks of speech and action runs. The insula sector specific for action vitality form was located slightly more dorsally than that of speech with a large overlap of their activations. The psycho-physiological interaction analysis showed that the insular sector involved in action vitality forms processing is connected with the left hemisphere areas controlling arm actions, whereas the sector involved in speech vitality forms processing is linked with right hemisphere areas related to speech prosody. We conclude that the central part of the insula is a key region for vitality forms processing regardless of the modality by which they are conveyed or expressed.

A description of the spatiotemporal dynamics of human cortical activity during cognitive tasks is a fundamental goal of neuroscience. In the present study, we employed stereo-EEG in order to assess the neural activity during tool-action observation. We recorded from 49 epileptic patients (5502 leads) implanted with intracerebral electrodes, while they observed tool and hand actions. We deconstructed actions into 3 events-video onset, action onset, and tool-object contact-and assessed how different brain regions respond to these events. Video onset, with actions not yet visible, recruited only visual areas. Aligning the responses at action onset, yielded activity in the parietal-frontal manipulation circuit and, selectively for tool actions, in the left anterior supramarginal gyrus (aSMG). Finally, by aligning to the tool-object contact that signals the achievement of the main goal of the observed action, activations were found in SII and dorsal premotor cortex. In conclusion, our data show that during tool-action observation, in addition to the general action observation network there is a selective activation of aSMG, which exhibits internally different patterns of responsiveness. In addition, neural responses selective for the contact between the tool and the object were also observed., (© The Author 2017. Published by Oxford University Press.)

Grasping relies on a network of parieto-frontal areas lying on the dorsolateral and dorsomedial parts of the hemispheres. However, the initiation and sequencing of voluntary actions also requires the contribution of mesial premotor regions, particularly the pre-supplementary motor area F6. We recorded 233 F6 neurons from 2 monkeys with chronic linear multishank neural probes during reaching-grasping visuomotor tasks. We showed that F6 neurons play a role in the control of forelimb movements and some of them (26%) exhibit visual and/or motor specificity for the target object. Interestingly, area F6 neurons form 2 functionally distinct populations, showing either visually-triggered or movement-related bursts of activity, in contrast to the sustained visual-to-motor activity displayed by ventral premotor area F5 neurons recorded in the same animals and with the same task during previous studies. These findings suggest that F6 plays a role in object grasping and extend existing models of the cortical grasping network., (© The Author 2016. Published by Oxford University Press.)

The purpose of this study was to investigate the contribution of stereopsis to the processing of observed manipulative actions. To this end, we first combined the factors "stimulus type" (action, static control, and dynamic control), "stereopsis" (present, absent) and "viewpoint" (frontal, lateral) into a single design. Four sites in premotor, retro-insular (2) and parietal cortex operated specifically when actions were viewed stereoscopically and frontally. A second experiment clarified that the stereo-action-specific regions were driven by actions moving out of the frontoparallel plane, an effect amplified by frontal viewing in premotor cortex. Analysis of single voxels and their discriminatory power showed that the representation of action in the stereo-action-specific areas was more accurate when stereopsis was active. Further analyses showed that the 4 stereo-action-specific sites form a closed network converging onto the premotor node, which connects to parietal and occipitotemporal regions outside the network. Several of the specific sites are known to process vestibular signals, suggesting that the network combines observed actions in peripersonal space with gravitational signals. These findings have wider implications for the function of premotor cortex and the role of stereopsis in human behavior., (© The Author 2016. Published by Oxford University Press.)

Picking up an object requires two basic motor operations: reaching and grasping. Neurophysiological studies in monkeys have suggested that the visuomotor transformations necessary for these two operations are carried out by separate parietofrontal circuits and that, for grasping, a key role is played by a specific sector of the ventral premotor cortex: area F5. The aim of the present study was to test the validity of this hypothesis by reversibly inactivating area F5 in monkeys trained to grasp objects of different shape, size and orientation. In separate sessions, the hand field of the primary motor cortex (area F1 or area 4) was also reversibly inactivated. The results showed that after inactivation of area F5 buried in the bank of the arcuate sulcus (the F5 sector where visuomotor neurones responding to object presentation are located), the hand shaping preceding grasping was markedly impaired and the hand posture was not appropriate for the object size and shape. The monkeys were eventually able to grasp the objects, but only after a series of corrections made under tactile control. With small inactivations the deficits concerned the contralesional hand, with larger inactivations the ipsilateral hand as well. In addition, there were signs of peripersonal neglect in the hemispace contralateral to the inactivation site. Following inactivation of area F5 lying on the cortical convexity (the F5 sector where visuomotor neurones responding to action observation, 'mirror neurones', are found) only a motor slowing was observed, the hand shaping being preserved. The inactivation of the hand field of area F1 produced a severe paralysis of contralateral finger movements with hypotonia. The results of this study indicate the crucial role of the ventral premotor cortex in visuomotor transformations for grasping movements. More generally, they provide strong support for the notion that distal and proximal movement organization relies upon distinct cortical circuits. Clinical data on distal movement deficits in humans are re-examined in the light of the present findings.

We recorded electrical activity from 532 neurons in the rostral part of inferior area 6 (area F5) of two macaque monkeys. Previous data had shown that neurons of this area discharge during goal-directed hand and mouth movements. We describe here the properties of a newly discovered set of F5 neurons ("mirror neurons', n = 92) all of which became active both when the monkey performed a given action and when it observed a similar action performed by the experimenter. Mirror neurons, in order to be visually triggered, required an interaction between the agent of the action and the object of it. The sight of the agent alone or of the object alone (three-dimensional objects, food) were ineffective. Hand and the mouth were by far the most effective agents. The actions most represented among those activating mirror neurons were grasping, manipulating and placing. In most mirror neurons (92%) there was a clear relation between the visual action they responded to and the motor response they coded. In approximately 30% of mirror neurons the congruence was very strict and the effective observed and executed actions corresponded both in terms of general action (e.g. grasping) and in terms of the way in which that action was executed (e.g. precision grip). We conclude by proposing that mirror neurons form a system for matching observation and execution of motor actions. We discuss the possible role of this system in action recognition and, given the proposed homology between F5 and human Brocca's region, we posit that a matching system, similar to that of mirror neurons exists in humans and could be involved in recognition of actions as well as phonetic gestures.

In the present study we investigated the possibility of a dissociation between the visual control of reaching and the visual control of grasping in a prehension task. To this purpose we studied the kinematics of prehension movements in a patient with a right parietal lesion and in six right-handed healthy control subjects. The task we used was one in which the subjects had to reach and grasp target objects in the presence or absence of a simultaneously presented distractor object. All stimuli were presented in the space ipsilateral to the lesion. The distractor could be either of the same or different size to the target object and was presented either to the right or to the left of the target. The following parameters of the prehension 'transport' component were analysed: wrist trajectory, transport time, tangential peak velocity, acceleration. Maximal finger aperture, time to maximal finger aperture, peak acceleration and time to peak acceleration of grip aperture were the parameters of the 'grasping' component analysed. The results showed that, although the patient had no misreaching, her hand trajectory deviated abnormally towards the distractor position when the distractor was to the right (ipsilateral) side of the target. In contrast, the grasp kinematics was not affected by the distractors, even when the size of the right distractor was different from the target. It appears, therefore, that the attentional shift towards the ipsilesional side, typical of neglect patients, determines a surprising dissociation in motor control. In the presence of a right distractor, the patient plans and partially executes a reaching movement towards that object and simultaneously performs a grasping movement towards a second object, i.e. the centrally located target. The presentation of distractors had no effects on the prehension kinematics of the control subjects.

The effect of unilateral surgical ablations of the postarcuate cortex (area 6) has been studied in the macaque monkey (Macaca irus). Two series of neurological deficits were found: (1) a failure to grasp food with the mouth when presented contralaterally to the lesion and a reluctance to use the contralateral hand; (2) a severe hemi-inattention. This was present in both the somatosensory and visual modalities but, in the latter, was limited to the peripersonal space. Both motor and attentional deficits improved with time. However, when the symptomatology was stabilized, the animals tended to use the ipsilateral hand and, when presented with two stimuli, preferred that ipsilateral to the lesion. Control experiments in which area 8 (frontal eye fields) was ablated unilaterally showed a decrease of eye movements contralateral to the lesion and a neglect of the contralateral hemispace. No somatosensory deficits were found. The visual neglect was more prominent for 'far' space than for that near the animal. The role of area 6 in organizing motor acts and the strict relation between the type of motor deficit and the type of neglect are discussed.