Two distinct networks for encoding goals and forms of action: An effective connectivity study.

Goal-directed actions are characterized by two main features: the content (i.e., the action goal) and the form, called vitality forms (VF) (i.e., how actions are executed). It is well established that both the action content and the capacity to understand the content of another's action are mediated by a network formed by a set of parietal and frontal brain areas. In contrast, the neural bases of action forms (e.g., gentle or rude actions) have not been characterized. However, there are now studies showing that the observation and execution of actions endowed with VF activate, in addition to the parieto-frontal network, the dorso-central insula (DCI). In the present study, we established-using dynamic causal modeling (DCM)-the direction of information flow during observation and execution of actions endowed with gentle and rude VF in the human brain. Based on previous fMRI studies, the selected nodes for the DCM comprised the posterior superior temporal sulcus (pSTS), the inferior parietal lobule (IPL), the premotor cortex (PM), and the DCI. Bayesian model comparison showed that, during action observation, two streams arose from pSTS: one toward IPL, concerning the action goal, and one toward DCI, concerning the action vitality forms. During action execution, two streams arose from PM: one toward IPL, concerning the action goal and one toward DCI concerning action vitality forms. This last finding opens an interesting question concerning the possibility to elicit VF in two distinct ways: cognitively (from PM to DCI) and affectively (from DCI to PM).

Investigating form and content of emotional and non-emotional laughing.

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.

Expressing our internal states and understanding those of others.

Vitality form is a term that describes the style with which motor actions are performed (e.g., rude, gentle, etc.). They represent one characterizing element of conscious and unconscious bodily communication. Despite their importance in interpersonal behavior, vitality forms have been, until now, virtually neglected in neuroscience. Here, using the functional MRI (fMRI) technique, we investigated the neural correlates of vitality forms in three different tasks: action observation, imagination, and execution. Conjunction analysis showed that, in all three tasks, there is a common, consistent activation of the dorsocentral sector of the insula. In addition, a common activation of the parietofrontal network, typically active during arm movements production, planning, and observation, was also found. We conclude that the dorsocentral part of the insula is a key element of the system that modulates the cortical motor activity, allowing individuals to express their internal states through action vitality forms. Recent monkey anatomical data show that the dorsocentral sector of the insula is, indeed, connected with the cortical circuit involved in the control of arm movements.

The neural correlates of velocity processing during the observation of a biological effector in the parietal and premotor cortex.

While there have been several studies investigating the neural correlates of action observation associated with hand grasping movements, comparatively little is known about the neural bases of observation of reaching movements. In two experiments, using functional magnetic resonance imaging (fMRI), we defined the cortical areas encoding reaching movements and assessed their sensitivity to biological motion and to movement velocity. In the first experiment, participants observed video-clips showing either a biological effector (an arm) or a non-biological object (rolling cylinder) reaching toward a target with a biological and a non-biological motion, respectively. In the second experiment, participants observed video-clips showing either a biological effector (an arm) or a non-biological object (an arrow) reaching toward a target with the same biological motion profiles. The results of the two experiments revealed activation of superior parietal and dorsal premotor sites during observation of the biological motion only, independent of whether it was performed by a biological effector (reaching arm) or a non-biological object (reaching arrow). These areas were not activated when participants observed the non-biological movement (rolling cylinder). To assess the responsiveness of parietal and frontal sites to movement velocity, the fMRI repetition-suppression (RS) technique was used, in which movement was shown with same or different velocities between consecutive videos, and observation of identical stimuli was contrasted with observation of different stimuli. Regions of interest were defined in the parietal and frontal cortices, and their response to stimulus repetition was analyzed (same vs. different velocities). The results showed an RS effect for velocity only during the observation of movements performed by the biological effector and not by the non-biological object. These data indicate that dorsal premotor and superior parietal areas represent a neural substrate involved in the encoding of reaching movements and that their responsiveness to movement velocity of a biological effector could be instrumental to the discrimination of movements performed by others.

Affective Contagion: How Attitudes Expressed by Others Influence Our Perception of Actions.

Vitality forms represent a fundamental aspect of social interactions by characterizing how actions are performed and how words are pronounced on the basis of the attitude of the agent. Same action, such as a handshake, may have a different impact on the receiver when it is performed kindly or vigorously, and similarly, a gentle or rude tone of voice may have a different impact on the listener. In the present study, we carried out two experiments that aimed to investigate whether and how vocal requests conveying different vitality forms can influence the perception of goal-directed actions and to measure the duration of this effect over time. More specifically, participants were asked to listen to the voice of an actor pronouncing "give me" in a rude or gentle way. Then, they were asked to observe the initial part of a rude or a gentle passing action, continue it mentally, and estimate the time of its completion. Results showed that the perception of different vitality forms expressed by vocal requests influenced the estimation of action duration. Moreover, we found that this effect was limited to a certain time interval (800 ms), after which it started to decay.

The neural bases of vitality forms.

Unlike emotions, which are short-lasting events accompanied by viscero-motor responses, vitality forms are continuous internal states that modulate the motor behaviors of individuals and are devoid of the autonomic modifications that characterize real emotions. Despite the importance of vitality forms in social life, only recently have neurophysiological studies been devoted to this issue. The first part of this review describes fMRI experiments, showing that the dorso-central insula is activated during the execution, the perception and the imagination of arm actions endowed with different vitality forms as well as during the hearing and the production of speech conveying vitality forms. In the second part, we address the means by which the dorso-central insula modulates the networks for controlling action execution and how the sensory and interoceptive information is conveyed to this insular sector. Finally, we present behavioral data showing the importance of vitality forms in social interactions.

Mirror Neuron System Activation in Children With Unilateral Cerebral Palsy During Observation of Actions Performed by a Pathological Model.

Background. Recent evidence suggested that Action Observation Therapy (AOT), based on observation of actions followed by immediate reproduction, could be a useful rehabilitative strategy for promoting functional recovery of children affected by unilateral cerebral palsy (UCP). AOT most likely exploits properties of the parieto-premotor mirror neuron system (MNS). This is more intensely activated when participants observe actions belonging to their own motor repertoire. Objective. The aim of the present study was to investigate the issue of whether MNS of UCP children is better activated by actions performed by a paretic hand rather than a healthy one. Methods. Using functional magnetic resonance imaging, we assessed brain activation in a homogeneous group of 10 right UCP children compared with that of 10 right-handed typically developing (TD) children, during observation of grasping actions performed by a healthy or a paretic hand. Results. The results revealed a significant activation within the MNS in both UCP and TD children, more lateralized to the left hemisphere in the TD group. Most important, region of interest (ROI) analysis on parietal and premotor regions showed that, in UCP, the MNS was more strongly activated by observation of actions performed by the paretic hand, a motor model more similar to the observer's motor repertoire. Conclusions. This study shows that children affected by spastic UCP exhibit enhanced activation of the MNS during observation of goal-directed actions performed by a pathological model with respect to a healthy one.

Vitality Forms Expressed by Others Modulate Our Own Motor Response: A Kinematic Study.

During social interaction, actions, and words may be expressed in different ways, for example, gently or rudely. A handshake can be gentle or vigorous and, similarly, tone of voice can be pleasant or rude. These aspects of social communication have been named vitality forms by Daniel Stern. Vitality forms represent how an action is performed and characterize all human interactions. In spite of their importance in social life, to date it is not clear whether the vitality forms expressed by the agent can influence the execution of a subsequent action performed by the receiver. To shed light on this matter, in the present study we carried out a kinematic study aiming to assess whether and how visual and auditory properties of vitality forms expressed by others influenced the motor response of participants. In particular, participants were presented with video-clips showing a male and a female actor performing a "giving request" (give me) or a "taking request" (take it) in visual, auditory, and mixed modalities (visual and auditory). Most importantly, requests were expressed with rude or gentle vitality forms. After the actor's request, participants performed a subsequent action. Results showed that vitality forms expressed by the actors influenced the kinematic parameters of the participants' actions regardless to the modality by which they are conveyed.

Differences in Action Style Recognition in Children with Autism Spectrum Disorders.

Vitality form is a term, originally introduced by Stern (2010), to describe "how" an action is performed. The capacity to perceive the vitality form of others' actions is a fundamental element of social interactions and a basic way of relating to and understanding others' behaviors. Although vitality forms characterize all human interactions, few studies have addressed their role in social and communicative disorders such as autism. The aim of the present study is to evaluate the ability to recognize different vitality forms during the observation of different motor actions in a group of children with autism spectrum disorders (ASD) compared to typically developing controls (TD). Results show a significant difference between children with ASD and TD in vitality forms recognition. This finding sheds new light on how children with ASD understand others' actions providing new ideas on overall social understanding as well as useful insights for professionals and caregivers alike.

Vitality Forms Processing in the Insula during Action Observation: A Multivoxel Pattern Analysis.

Observing the style of an action done by others allows the observer to understand the cognitive state of the agent. This information has been defined by Stern "vitality forms". Previous experiments showed that the dorso-central insula is selectively active both during vitality form observation and execution. In the present study, we presented participants with videos showing hand actions performed with different velocities and asked them to judge either their vitality form (gentle, neutral, rude) or their velocity (slow, medium, fast). The aim of the present study was to assess, using multi-voxel pattern analysis, whether vitality forms and velocities of observed goal-directed actions are differentially processed in the insula, and more specifically whether action velocity is encoded per se or it is an element that triggers neural populations of the insula encoding the vitality form. The results showed that, consistently across subjects, in the dorso-central sector of the insula there were voxels selectively tuned to vitality forms, while voxel tuned to velocity were rare. These results indicate that the dorso-central insula, which previous data showed to be involved in the vitality form processing, contains voxels specific for the action style processing.

Human, Nature, Dynamism: The Effects of Content and Movement Perception on Brain Activations during the Aesthetic Judgment of Representational Paintings.

Movement perception and its role in aesthetic experience have been often studied, within empirical aesthetics, in relation to the human body. No such specificity has been defined in neuroimaging studies with respect to contents lacking a human form. The aim of this work was to explore, through functional magnetic imaging (f MRI), how perceived movement is processed during the aesthetic judgment of paintings using two types of content: human subjects and scenes of nature. Participants, untutored in the arts, were shown the stimuli and asked to make aesthetic judgments. Additionally, they were instructed to observe the paintings and to rate their perceived movement in separate blocks. Observation highlighted spontaneous processes associated with aesthetic experience, whereas movement judgment outlined activations specifically related to movement processing. The ratings recorded during aesthetic judgment revealed that nature scenes received higher scored than human content paintings. The imaging data showed similar activation, relative to baseline, for all stimuli in the three tasks, including activation of occipito-temporal areas, posterior parietal, and premotor cortices. Contrast analyses within aesthetic judgment task showed that human content activated, relative to nature, precuneus, fusiform gyrus, and posterior temporal areas, whose activation was prominent for dynamic human paintings. In contrast, nature scenes activated, relative to human stimuli, occipital and posterior parietal cortex/precuneus, involved in visuospatial exploration and pragmatic coding of movement, as well as central insula. Static nature paintings further activated, relative to dynamic nature stimuli, central and posterior insula. Besides insular activation, which was specific for aesthetic judgment, we found a large overlap in the activation pattern characterizing each stimulus dimension (content and dynamism) across observation, aesthetic judgment, and movement judgment tasks. These findings support the idea that the aesthetic evaluation of artworks depicting both human subjects and nature scenes involves a motor component, and that the associated neural processes occur quite spontaneously in the viewer. Furthermore, considering the functional roles of posterior and central insula, we suggest that nature paintings may evoke aesthetic processes requiring an additional proprioceptive and sensori-motor component implemented by "motor accessibility" to the represented scenario, which is needed to judge the aesthetic value of the observed painting.

The neural correlates of 'vitality form' recognition: an fMRI study: this work is dedicated to Daniel Stern, whose immeasurable contribution to science has inspired our research.

The observation of goal-directed actions performed by another individual allows one to understand what that individual is doing and why he/she is doing it. Important information about others' behaviour is also carried out by the dynamics of the observed action. Action dynamics characterize the 'vitality form' of an action describing the cognitive and affective relation between the performing agent and the action recipient. Here, using the fMRI technique, we assessed the neural correlates of vitality form recognition presenting participants with videos showing two actors executing actions with different vitality forms: energetic and gentle. The participants viewed the actions in two tasks. In one task (what), they had to focus on the goal of the presented action; in the other task (how), they had to focus on the vitality form. For both tasks, activations were found in the action observation/execution circuit. Most interestingly, the contrast how vs what revealed activation in right dorso-central insula, highlighting the involvement, in the recognition of vitality form, of an anatomical region connecting somatosensory areas with the medial temporal region and, in particular, with the hippocampus. This somatosensory-insular-limbic circuit could underlie the observers' capacity to understand the vitality forms conveyed by the observed action.