Visual exploration of emotional body language: a behavioural and eye-tracking study.

Bodily postures are essential to correctly comprehend others' emotions and intentions. Nonetheless, very few studies focused on the pattern of eye movements implicated in the recognition of emotional body language (EBL), demonstrating significant differences in relation to different emotions. A yet unanswered question regards the presence of the "left-gaze bias" (i.e. the tendency to look first, to make more fixations and to spend more looking time on the left side of centrally presented stimuli) while scanning bodies. Hence, the present study aims at exploring both the presence of a left-gaze bias and the modulation of EBL visual exploration mechanisms, by investigating the fixation patterns (number of fixations and latency of the first fixation) of participants while judging the emotional intensity of static bodily postures (Angry, Happy and Neutral, without head). While results on the latency of first fixations demonstrate for the first time the presence of the left-gaze bias while scanning bodies, suggesting that it could be related to the stronger expressiveness of the left hand (from the observer's point of view), results on fixations' number only partially fulfil our hypothesis. Moreover, an opposite viewing pattern between Angry and Happy bodily postures is showed. In sum, the present results, by integrating the spatial and temporal dimension of gaze exploration patterns, shed new light on EBL visual exploration mechanisms.

Schizotypy and individual differences in peripersonal space plasticity.

The space surrounding our body, defined as peripersonal space (PPS), is dynamically shaped by our motor experiences. For instance, PPS extends after using a tool to reach far objects. Several studies have demonstrated how PPS size varies across people, depending on different individual characteristics, including schizotypy. Coherently, narrower PPS boundaries have been reported among high schizotypal individuals and schizophrenia patients. However, little is known about the relationship between PPS plasticity and personality traits like schizotypy. To this purpose, the present study has investigated the individual PPS plasticity, after two different motor trainings, along the schizotypal continuum. Specifically, PPS plasticity was tested after using a tool (Experiment 1) and after the mere observation of another person using the same tool (Experiment 2). Indeed, previous evidence has shown that tool-use observation influences visual distance judgments, extending the representation of PPS. To date, however, there is no study investigating whether observation of tools action could also affect multisensory PPS tasks. Experiment 1 has shown that PPS boundaries extended after using the tool; on the other hand, Experiment 2 has revealed the absence of PPS expansion. Moreover, greater PPS expansion emerged in the relatively-low schizotypal group than in the relatively-high one, regardless of the type of motor training performed. The absence of PPS modulation after the observation task is discussed in relation to recent findings showing that intentional action and/or the goal of the action represent potentially crucial elements to trigger PPS plasticity. Finally, these new results extend previous evidence underlining a potential general functional alteration of PPS with the increase of schizotypal level.

Beholders' sensorimotor engagement enhances aesthetic rating of pictorial facial expressions of pain.

The present study addresses a novel issue by investigating whether beholders' sensorimotor engagement with the emotional content of works of art contributes to the formation of their objective aesthetic judgment of beauty. To this purpose, participants' sensorimotor engagement was modulated by asking them to overtly contract the Corrugator Supercilii facial muscles or to refrain from any voluntary facial movement while judging the aesthetic value of painful and neutral facial expressions in select examples of Renaissance and Baroque paintings. Results demonstrated a specific increase in the aesthetic rating of paintings showing painful facial expressions during the congruent activation of the Corrugator Supercilii muscles. Furthermore, participants' empathetic traits and expertise in art were found to correlate directly with the amplitude of the motor enactment effect on aesthetic judgments. For the first time, we show the role of bottom-up bodily driven sensorimotor processes in the objective aesthetic evaluation of works of art.

Shared multisensory experience affects Others' boundary: The enfacement illusion in schizophrenia.

Schizophrenia has been described as a psychiatric condition characterized by deficits in one's own and others' face recognition, as well as by a disturbed sense of body-ownership. To date, no study has integrated these two lines of research with the aim of investigating Enfacement Illusion (EI) proneness in schizophrenia. To accomplish this goal, the classic EI protocol was adapted to test the potential plasticity of both Self-Other and Other-Other boundaries. Results showed that EI induced the expected malleability of Self-Other boundary among both controls and patients. Interestingly, for the first time, the present study demonstrates that also the Other-Other boundary was influenced by EI. Furthermore, comparing the two groups, the malleability of the Other-Other boundary showed an opposite modulation. These results suggest that, instead of greater Self-Other boundary plasticity, a qualitative difference can be detected between schizophrenia patients and controls in the malleability of the Other-Other boundary. The present study points out a totally new aspect about body-illusions and schizophrenia disorder, demonstrating that EI is not only confined to self-sphere but it also affects the way we discriminate others, representing a potential crucial aspect in the social domain.

Decoding the activity of grasping neurons recorded from the ventral premotor area F5 of the macaque monkey.

Many neurons in the monkey ventral premotor area F5 discharge selectively when the monkey grasps an object with a specific grip. Of these, the motor neurons are active only during grasping execution, whereas the visuomotor neurons also respond to object presentation. Here we assessed whether the activity of 90 task-related F5 neurons recorded from two macaque monkeys during the performance of a visually-guided grasping task can be used as input to pattern recognition algorithms aiming to decode different grips. The features exploited for the decoding were the mean firing rate and the mean interspike interval calculated over different time spans of the movement period (all neurons) or of the object presentation period (visuomotor neurons). A support vector machine (SVM) algorithm was applied to the neural activity recorded while the monkey grasped two sets of objects. The original set contained three objects that were grasped with different hand shapes, plus three others that were grasped with the same grip, whereas the six objects of the special set were grasped with six distinctive hand configurations. The algorithm predicted with accuracy greater than 95% all the distinct grips used to grasp the objects. The classification rate obtained using the first 25% of the movement period was 90%, whereas it was nearly perfect using the entire period. At least 16 neurons were needed for accurate performance, with a progressive increase in accuracy as more neurons were included. Classification errors revealed by confusion matrices were found to reflect similarities of hand grips used to grasp the objects. The use of visuomotor neurons' responses to object presentation yielded grip classification accuracy similar to that obtained from actual grasping execution. We suggest that F5 grasping-related activity might be used by neural prostheses to tailor hand shape to the specific object to be grasped even before movement onset.

Multimodal communication in animals, humans and robots: an introduction to perspectives in brain-inspired informatics.

Recent years have seen convergence in research on brain mechanisms and neurocomputational approaches, culminating in the creation of a new generation of robots whose artificial "brains" respect neuroscience principles and whose "cognitive" systems venture into higher cognitive domains such as planning and action sequencing, complex object and concept processing, and language. The present article gives an overview of selected projects in this general multidisciplinary field. The work reviewed centres on research funded by the EU in the context of the New and Emergent Science and Technology, NEST, funding scheme highlighting the topic "What it means to be human". Examples of such projects include learning by imitation (Edici project), examining the origin of human rule-based reasoning (Far), studying the neural origins of language (Neurocom), exploring the evolutionary origins of the human mind (Pkb140404), researching into verbal and non-verbal communication (Refcom), using and interpreting signs (Sedsu), characterising human language by structural complexity (Chlasc), and representing abstract concepts (Abstract). Each of the communication-centred research projects revealed individual insights; however, there had been little overall analysis of results and hypotheses. In the Specific Support Action Nestcom, we proposed to analyse some NEST projects focusing on the central question "What it means to communicate" and to review, understand and integrate the results of previous communication-related research, in order to develop and communicate multimodal experimental hypotheses for investigation by future projects. The present special issue includes a range of papers on the interplay between neuroinformatics, brain science and robotics in the general area of higher cognitive functions and multimodal communication. These papers extend talks given at the NESTCOM workshops, at ICANN (http://www.his.sunderland.ac.uk/nestcom/workshop/icann.html) in Porto and at the first meeting of the Federation of the European Societies of Neuropsychology in Edinburgh in 2008 (http://www.his.sunderland.ac.uk/nestcom/workshop/esn.html). We hope that the collection will give a vivid insight into current trends in the field.

When pliers become fingers in the monkey motor system.

The capacity to use tools is a fundamental evolutionary achievement. Its essence stands in the capacity to transfer a proximal goal (grasp a tool) to a distal goal (e.g., grasp food). Where and how does this goal transfer occur? Here, we show that, in monkeys trained to use tools, cortical motor neurons, active during hand grasping, also become active during grasping with pliers, as if the pliers were now the hand fingers. This motor embodiment occurs both for normal pliers and for "reverse pliers," an implement that requires finger opening, instead of their closing, to grasp an object. We conclude that the capacity to use tools is based on an inherently goal-centered functional organization of primate cortical motor areas.

Listening to action-related sentences modulates the activity of the motor system: a combined TMS and behavioral study.

Transcranial magnetic stimulation (TMS) and a behavioral paradigm were used to assess whether listening to action-related sentences modulates the activity of the motor system. By means of single-pulse TMS, either the hand or the foot/leg motor area in the left hemisphere was stimulated in distinct experimental sessions, while participants were listening to sentences expressing hand and foot actions. Listening to abstract content sentences served as a control. Motor evoked potentials (MEPs) were recorded from hand and foot muscles. Results showed that MEPs recorded from hand muscles were specifically modulated by listening to hand-action-related sentences, as were MEPs recorded from foot muscles by listening to foot-action-related sentences. This modulation consisted of an amplitude decrease of the recorded MEPs. In the behavioral task, participants had to respond with the hand or the foot while listening to actions expressing hand and foot actions, as compared to abstract sentences. Coherently with the results obtained with TMS, when the response was given with the hand, reaction times were slower during listening to hand-action-related sentences, while when the response was given with the foot, reaction times were slower during listening to foot-action-related sentences. The present data show that processing verbally presented actions activates different sectors of the motor system, depending on the effector used in the listened-to action.

Audiovisual mirror neurons and action recognition.

Many object-related actions can be recognized both by their sound and by their vision. Here we describe a population of neurons in the ventral premotor cortex of the monkey that discharge both when the animal performs a specific action and when it hears or sees the same action performed by another individual. These 'audiovisual mirror neurons' therefore represent actions independently of whether these actions are performed, heard or seen. The magnitude of auditory and visual responses did not differ significantly in half the neurons. A neurometric analysis revealed that based on the response of these neurons, two actions could be discriminated with 97% accuracy.

I know what you are doing. a neurophysiological study.

In the ventral premotor cortex of the macaque monkey, there are neurons that discharge both during the execution of hand actions and during the observation of the same actions made by others (mirror neurons). In the present study, we show that a subset of mirror neurons becomes active during action presentation and also when the final part of the action, crucial in triggering the response in full vision, is hidden and can therefore only be inferred. This implies that the motor representation of an action performed by others can be internally generated in the observer's premotor cortex, even when a visual description of the action is lacking. The present findings support the hypothesis that mirror neuron activation could be at the basis of action recognition.

Cortical mechanism for the visual guidance of hand grasping movements in the monkey: A reversible inactivation study.

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.

Action observation activates premotor and parietal areas in a somatotopic manner: an fMRI study.

Functional magnetic resonance imaging (fMRI) was used to localize brain areas that were active during the observation of actions made by another individual. Object- and non-object-related actions made with different effectors (mouth, hand and foot) were presented. Observation of both object- and non-object-related actions determined a somatotopically organized activation of premotor cortex. The somatotopic pattern was similar to that of the classical motor cortex homunculus. During the observation of object-related actions, an activation, also somatotopically organized, was additionally found in the posterior parietal lobe. Thus, when individuals observe an action, an internal replica of that action is automatically generated in their premotor cortex. In the case of object-related actions, a further object-related analysis is performed in the parietal lobe, as if the subjects were indeed using those objects. These results bring the previous concept of an action observation/execution matching system (mirror system) into a broader perspective: this system is not restricted to the ventral premotor cortex, but involves several somatotopically organized motor circuits.

The ability to follow eye gaze and its emergence during development in macaque monkeys.

The ability of monkeys to follow the gaze of other individuals is a matter of debate in many behavioral studies. Physiological studies have shown that in monkeys, as in humans, there are neural correlates of eye direction detection. There is little evidence at the behavioral level, however, of the presence and development of such abilities in monkeys. The aim of the present study was to assess in juveniles and adult pig-tailed macaques (Macaca nemestrina) the capacity to use eye cues only to follow the gaze of an experimenter. Biological stimuli (head, eye, and trunk movements) were presented by an experimenter to 2 adult monkeys with their heads restrained (Experiment 1) and to 11 monkeys of different ages, free to move in their home cages (Experiment 2). A nonbiological stimulus served as a control. Results showed that macaques can follow the gaze of the experimenter by using head/eye and eye cues alone. Trunk movements and nonbiological stimuli did not significantly elicit similar reactions. Juvenile monkeys were not able to orient their attention on the basis of eye cues alone. In general, gaze following was more frequent in adults than in juveniles. Like in humans, however, such abilities in macaques dramatically improve with age suggesting that the transition to adulthood is a crucial period in the development of gaze-following behavior.

Selectivity for the shape, size, and orientation of objects for grasping in neurons of monkey parietal area AIP.

In this study, we mainly investigated the visual selectivity of hand-manipulation-related neurons in the anterior intraparietal area (area AIP) while the animal was grasping or fixating on three-dimensional (3D) objects of different geometric shapes, sizes, and orientations. We studied the activity of 132 task-related neurons during the hand-manipulation tasks in the light and in the dark, as well as during object fixation. Seventy-seven percent (101/132) of the hand-manipulation-related neurons were visually responsive, showing either lesser activity during manipulation in the dark than during that in the light (visual-motor neurons) or no activation in the dark (visual-dominant neurons). Of these visually responsive neurons, more than half (n = 66) responded during the object-fixation task (object-type). Among these, 55 were tested for their shape selectivity during the object-fixation task, and many (n = 25) were highly selective, preferring one particular shape of the six different shapes presented (ring, cube, cylinder, cone, sphere, and square plate). For 28 moderately selective object-type neurons, we performed multidimensional scaling (MDS) to examine how the neurons encode the similarity of objects. The results suggest that some moderately selective neurons responded preferentially to common geometric features shared by similar objects (flat, round, elongated, etc.). Moderately selective nonobject-type visually responsive neurons, which did not respond during object fixation, were found by MDS to be more closely related to the handgrip than to the object shape. We found a similar selectivity for handgrip in motor-dominant neurons that did not show any visual response. With regard to the size of the objects, 16 of 26 object-type neurons tested were selective for both size and shape, whereas 9 object-type neurons were selective for shape but not for size. Seven of 12 nonobject-type and all (8/8) of the motor-dominant neurons examined were selective for size, and almost all of them were also selective for objects. Many hand-manipulation-related neurons that preferred the plate and/or ring were selective for the orientation of the objects (17/20). These results suggest that the visual responses of object-type neurons represent the shape, size, and/or orientation of 3D objects, whereas those of the nonobject-type neurons probably represent the shape of the handgrip, grip size, or hand-orientation. The activity of motor-dominant neurons was also, in part, likely to represent these parameters of hand movement. This suggests that the dorsal visual pathway is concerned with the aspect of form, orientation, and/or size perception that is relevant for the visual control of movements.

Visuomotor neurons: ambiguity of the discharge or 'motor' perception?

The cortical motor system has been classically considered as the unitary, output stage of the brain processing of sensory information. According to this idea, the motor cortex - the acting brain - receives the result of the perceptual processing (visual, acoustical, tactile, etc.) elaborated by the 'associative cortex'. During the last two decades this perspective has been challenged by a series of anatomical, hodological, and neurophysiological data. This converging evidence delineates a dramatically changed picture. Far from being unitary, the cortical motor system appears to be constituted by a constellation of distinct areas, each of those endowed with specific functional properties and linked by reciprocal connections with distinct sectors of the parietal cortex. Furthermore, several 'motor' neurons in addition to their motor discharge, are also activated by somatosensory and visual stimulation (somatomotor and visuomotor neurons). In the present paper we will discuss the functional properties of those sensorimotor neurons located in the ventral part of the monkey premotor cortex. On the basis of electrophysiological data, we will propose that the apparent parodox stemming from the coexistence within the same neuron of motor and sensory properties can be solved by postulating that the motor system not only executes actions but also internally represents them in terms of 'motor ideas'. These motor ideas may provide the neurobiological basis for space representation, understanding of actions made by others and, possibly, semantic categorization of objects.

Visual responses in the dorsal premotor area F2 of the macaque monkey.

This study aimed to determine the presence of neurons responding to visual stimuli in area F2 of the dorsal premotor cortex of the macaque monkey. In order to delimit the sector in which visually responsive neurons are located, the somatotopic organization of area F2 was studied with intracortical microstimulation and single neuron recording. The results showed that: (1) in area F2 there is a significant percentage of visually responsive neurons (15.9% of all recorded neurons); (2) area F2 is excitable with a low-threshold current (average 28.1 microA) and has a somatotopic representation of the whole body, except the face; and (3) most visually driven neurons (n=130 out of 169) are concentrated within the rostrolateral sector of the forelimb representation of area F2, thus providing for the first time functional support for the neuroanatomical evidence that the visual input to area F2 is mostly restricted to this sector.

Resonance behaviors and mirror neurons.

This article is subdivided into two parts. In the first part we review the properties of a particular class of premotor neurons, the "mirror" neurons. With this term we define neurons that discharge both when the monkey makes a particular action and when it observes another individual (monkey or human) making a similar action. The second part is an attempt to give a neurophysiological account of the mechanisms underlying behaviors where an individual reproduces, overtly or internally, movements or actions made by another individual. We will refer to these behaviors as "resonance behaviors". We distinguish two types of resonance behavior. The first type is characterized by imitation, immediate or with delay, of movements made by other individuals. Examples of resonance behavior of this type are the "imitative" behaviors observed in birds, young infants and patients with frontal lesions. The second type of resonance behavior is characterized by the occurrence, at the observation of an action, of a neural pattern, which, when internally generated, determines the making of the observed action. In this type of resonance behavior the observed action is, typically, not repeated (overtly). We argue that resonance behavior of the second type is at the basis of the understanding of actions made by others. At the end of the article we review evidence of mirror mechanisms in humans and discuss their anatomical localizations.