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According to available evidence, after making an erroneous decision people tend to slow down on the next decision. This empirical regularity, known as "post error slowing" (PES), has been traditionally interpreted as the result of a conservative response criterion adopted to avoid future errors and it is supposed to be driven by changes in the excitability of the motor system. However, the consequences of errors have been almost exclusively investigated by means of button-press tasks, which have been criticized because of their limited ecological validity and it is still unclear to what extent errors bias the motor system activity during the planning and the on-line control of complex and realistic goal-directed actions. To overcome these potential limitations, in the present study, we investigated the effect of errors on the preparation and execution of the reach-to-grasp movement, one of the most significant daily life actions. In addition to reaction times (RTs), we measured motor-evoked potential (MEP) to explore the influence of errors on corticospinal (CS) excitability, and we applied kinematical analysis to examine the underlying reorganization of the movement following an error. The results of the present study showed that MEPs tend to be reduced after the failure to reach and grasp an object, supporting the traditional interpretation of PES. Furthermore, in addition to RTs, we found that error-reactivity strategically influences the grasping component of the action, whereas the reaching component appears to be impermeable to PES. These findings demonstrate that the error-reactivity is a strong empirical phenomenon, which spreads into the motor system at the level of both movement preparation and execution, even when more realistic and ecologically valid tasks are used., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

For human beings, monitoring others' errors is essential for efficient goal-directed behavior. Indeed, the mere observation of other individuals' errors provides a rich source of information that can be used to avoid potential errors and improve our performance without direct experience. Recent studies have outlined that vicarious experience of errors influences the observer's overt behavior. This observational posterror slowing (oPES) is supposed to reflect a strategic increase in control aimed at reducing the probability of an error. Because the consequences of error observation have been exclusively investigated by means of arbitrary button-press responses, which limit the investigation to premovement processes, it is unclear whether the observation of an error also influences the online control of goal-directed actions. In the present study, for the first time, we investigated the effect of error observation on the reach-to-grasp movement by means of kinematical analysis. The results revealed that error-observation effects are not confined to premovement processes-they also strategically affect spatial movement trajectories. Our findings add substantially to previous literature, showing that the oPES spreads to movement execution when a more realistic, ecologically valid task is employed.

A common finding across many speeded reaction time (RT) tasks is that people tend to respond more slowly after making an error. This phenomenon, known as post-error slowing (PES), has been traditionally hypothesized to reflect a strategic increase in response caution, aimed at preventing the occurrence of new errors. However, this interpretation of PES has been challenged on multiple fronts. Firstly, recent investigations have suggested that errors may produce a decrement in performance accuracy and that PES might occur because error processing has a detrimental effect on subsequent information processing. Secondly, previous research has been criticized because of the limited ecological validity of speeded RT tasks. In the present study, we investigated error-reactivity in the context of goal-directed actions, in order to examine the extent to which PES effects impact on realistic and complex movements. Specifically, we investigated the effect of errors on the reach to grasp movement (Experiment 1). In addition to RTs, we performed a kinematical analysis in order to explore the underlying reorganization of the movements after an error. The results of the present study showed that error reactivity strategically influences the grasping component of the action, whereas the reaching component appears to be impermeable to PES. The resistance of the reaching component to PES was confirmed in a second 'only reaching' experiment (Experiment 2). These finding supports the hypothesis that error reactivity is a flexible process whose effects on behavior also depend on the motor components involved in the action., (Copyright © 2018 Elsevier B.V. All rights reserved.)

According to the dual-stream theory, the processing of visual information is divided into a ventral pathway mediating object recognition, and a dorsal pathway supporting visuomotor control. Increasing evidence suggests that these streams are not independent, but where this dorsal-ventral stream integration occurs remains unknown. We explored the candidate white matter pathways linking dorsal and ventral visual streams in 30 right-handed participants performing object-oriented hand movements of varying complexity (reaching, grasping and lifting), using advanced diffusion imaging tractography based on the spherical deconvolution and kinematical analysis. We provided for the first time a direct evidence of cross-communication between dorsal and ventral visual streams in humans, through vertical occipital fasciculus and temporo-parietal fibers of the arcuate fasciculus during on-line control of grasping and lifting actions. We showed that individual differences in the microstructure of these cross-talk connections were associated with the variability of the arm deceleration, the grip opening and the grasp accuracy. This study suggests that hand kinematics, in skilled hand actions where high degree of online control is required, is related to the anatomy of the cross-talk networks between dorsal and ventral streams, bringing important insights to the dual-stream theory and the sensorimotor organization of hand actions., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Functional neuroimaging and brain lesion studies demonstrate that secondary motor areas of the frontal lobe play a crucial role in the cortical control of hand movements. However, no study so far has examined frontal white matter connections of the secondary motor network, namely the frontal aslant tract, connecting the supplementary motor complex and the posterior inferior frontal regions, and the U-shaped dorsal and ventral premotor fibers running through the middle frontal gyrus. The aim of the current study is to explore the involvement of the short frontal lobe connections in reaching and reach-to-grasp movements in 32 right-handed healthy subjects by correlating tractography data based on spherical deconvolution approach with kinematical data. We showed that individual differences in the microstructure of the bilateral frontal aslant tract, bilateral ventral and left dorsal premotor tracts were associated with kinematic features of hand actions. Furthermore, bilateral ventral premotor connections were also involved in the closing grip phase necessary for determining efficient and stable grasping of the target object. This work suggests for the first time that hand kinematics and visuomotor processing are associated with the anatomy of the short frontal lobe connections., (Copyright © 2016 Elsevier Inc. All rights reserved.)

Appropriate communication is at the heart of successful, healthy social interactions in humans. Deficits in social communication are a hallmark of several neurological and psychiatric disorders. Yet, very little research has been devoted to understanding the mechanisms underlying these issues. It has been suggested that dopamine is a candidate neurotransmitter system involved in stimulating communication in individuals that are not highly motivated to communicate. A typical model to study dopaminergic dysfunctions in humans is represented by Parkinson's disease (PD) patients, who show motor, cognitive and motivational symptoms. Our study aimed to investigate the effects of social communication on actions in non-demented PD patients receiving dopamine replacement therapy (Levodopa = l-Dopa) and in neurologically healthy control participants. Patients' ability to modulate motor patterning depending on the communicative intention motivating the action to be performed was evaluated both in "on" (with l-Dopa) and "off" (without l-Dopa) states. In two main conditions, participants were requested to reach towards, grasp an object, and either simply lift it (individual condition) or lift it with the intent to communicate a meaning to a partner (communicative condition). Movements' kinematics was recorded using a three-dimensional motion analysis system. The results indicate that kinematics is sensitive to communicative intention and that l-Dopa treatment has positive effects on translating communicative intentions into specific motor patterns in PD patients. Although the to-be-grasped object remained the same both the controls and the PD patients in an 'on' state adopted different kinematic patterning for the 'individual' and the 'communication' conditions. The PD patients in the 'off' state, instead, were unable to kinematically differentiate between the two conditions. We contend that social and communicative impairments are associated with abnormalities in dopaminergic pathways., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

A large body of research indicates that observing actions made by others is associated with corresponding motor facilitation of the observer's corticospinal system. However, it is still controversial whether this matching mechanism strictly reflects the kinematics of the observed action or its meaning. To test this issue, motor evoked potentials induced by single-pulse transcranial magnetic stimulation were recorded from hand and leg muscles while participants observed a symbolic action carried out with the index finger, but classically performed with the leg (i.e., a soccer penalty kick). A control condition in which participants observed a similar (but not symbolic) hand movement was also included. Results showed that motor facilitation occurs both in the observer's hand (first dorsal interosseous) and leg (quadriceps femoris) muscles. The present study provides evidence that both the kinematics and the symbolic value of an observed action are able to modulate motor cortex excitability. The human motor system is thus not only involved in mirroring observed actions but is also finely tuned to their symbolic value., (© 2015 Federation of European Neuroscience Societies and John Wiley & Sons Ltd.)

Dysfunction of the dopaminergic system leads to motor, cognitive and motivational symptoms in brain disorders such as Parkinson's disease (PD). Moreover, the dopaminergic system plays an important role in social interactions. The dopaminergic input to the basal ganglia (BG) thought to integrate social cues during the planning and execution of voluntary movements remains, however, largely unexplored. Since PD provides a model to assess this function in humans, our study aimed to investigate the effects of social intentions on actions in non-demented PDpatients receiving dopamine replacement therapy (Levodopa = l-Dopa) and in neurologically healthy control participants. Patients' ability to modulate motor patterning depending on the intention motivating the action to be performed was evaluated both in "on" (with l-Dopa) and "off" (without l-Dopa) states. Participants were instructed to reach for and to grasp an object; they were then told to hand it to another person (social condition) or to place it on a concave frame (individual condition). A 'passive-observer' condition, which was similar to the 'individual' condition except for the presence of an onlooker who simply observed the scene, was also assessed to exclude the possibility that differences might be due to the presence of another person. Movement kinematics were recorded using a three-dimensional motion analysis system. Study results demonstrated that the controls and the PD patients in an 'on' state adopted different kinematic patterning for the 'social' and the 'individual' conditions; the PD patients in the 'off' state, instead, were unable to kinematically differentiate between the two conditions. These results suggest that l-Dopa treatment has positive effects on translating social intentions into specific motor patterns in PD patients., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

The most popular model to explain how prehensile movements are organized assumes that they comprise two "components", the reaching component encoding information regarding the object's spatial location and the grasping component encoding information on the object's intrinsic properties such as size and shape. Comparative kinematic studies on grasping behavior in the humans and in macaques have been carried out to investigate the similarities and differences existing across the two species. Although these studies seem to favor the hypothesis that macaques and humans share a number of kinematic features it remains unclear how the reaching and grasping components are coordinated during prehension movements in free-ranging macaque monkeys. Twelve hours of video footage was filmed of the monkeys as they snatched food items from one another (i.e., snatching) or collect them in the absence of competitors (i.e., unconstrained). The video samples were analyzed frame-by-frame using digitization techniques developed to perform two-dimensional post-hoc kinematic analyses of the two types of actions. The results indicate that only for the snatching condition when the reaching variability increased there was an increase in the amplitude of maximum grip aperture. Besides, the start of a break-point along the deceleration phase of the velocity profile correlated with the time at which maximum grip aperture occurred. These findings suggest that macaques can spatially and temporally couple the reaching and the grasping components when there is pressure to act quickly. They offer a substantial contribution to the debate about the nature of how prehensile actions are programmed.

Choice reaction tasks are performed faster when stimulus location corresponds to response location (Simon effect). This spatial stimulus-response compatibility effect affects performance at the level of action planning and execution. However, when response selection is completed before movement initiation, the Simon effect arises only at the planning level. The aim of this study was to ascertain whether when a precocious response selection is requested, the Simon effect can be detected on the kinematics characterizing the online control phase of a non-ballistic movement. Participants were presented with red or green colored squares, which could appear on the right, left, above, or below a central cross. Depending on the square's color, participants had to release one of two buttons (right/left), then reach toward and press a corresponding lateral pad. We found evidence of the Simon effect on both action planning and on-line control. Moreover, the investigation of response conflict at the level of previous trials (i.e., n-1), a factor that might determine interference at the level of the current response, revealed a conflict adaptation process across trials. Results are discussed in terms of current theories concerned with the Simon effect and the distinction between action planning and control., (Copyright © 2014 Cognitive Science Society, Inc.)

Research in the field of psychology and cognitive neuroscience has begun to explore the functional underpinnings of voluntary actions and how they differ from stimulus-driven actions. From these studies one can conclude that the two action modes differ with respect to their neural and behavioural correlates. So far, however, no study has investigated whether the voluntary and stimulus-driven actions also differ in terms of motor programming. We report two experiments in which participants had to perform either voluntary or stimulus-driven reach-to-grasp actions upon the same stimulus. Using kinematic methods, in Experiment 1 we obtained evidence that voluntary actions and stimulus-driven actions translate into differential movement patterns. Results for Experiments 2 suggest that selecting what to do, when to act, and whether to act are characterized by specific kinematic signatures and affect different aspects of the reach-to-grasp movement in a selective fashion. These findings add to current models of volition suggesting that voluntary action control results from an interplay of dissociable subfunctions related to specific decision components: what action execute, when to execute an action, and whether to execute any action., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

The goal of the present experiment was twofold: identifying similarities and differences between flavour memory and visual memory mechanisms and investigating whether kinematics could serve as an implicit measure for food selection. To test flavour and visual memory an 'implicit' paradigm to represent real-life situations in a controlled lab setting was implemented. A target, i.e., a piece of cake shaped like either an orange or a tangerine, covered with either orange- or a tangerine-flavoured icing, was provided to participants on Day 1. On Day 2, without prior notice, participants were requested to recognize the target amongst a set of distractors, characterized by various flavours (orange vs. tangerine) and/or sizes (orange-like vs. tangerine-like). Similarly, targets and distractors consisting of 2D figures varying in shape and size were used to assess visual memory. Reach-to-grasp kinematics towards the targets were recorded and analysed by means of digitalization techniques. Correlations between kinematic parameters, memory and liking for each food item were also calculated. Results concerned with memory recollection indices provided evidence of different key mechanisms which could be based either on novelty of flavour memory or visual memory, respectively. To a moderate extent, kinematics may serve as an implicit index of food selection processes., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

In both monkeys and humans, reaching-related sensorimotor transformations involve the activation of a wide fronto-parietal network. Recent neurophysiological evidence suggests that some components of this network host not only neurons encoding the direction of arm reaching movements, but also neurons whose involvement is modulated by the intrinsic features of an object (e.g. size and shape). To date, it has yet to be investigated whether a similar modulation is evident in the human reaching-related areas. To fill this gap, we asked participants to reach towards either a small or a large object while kinematic and electroencephalographic signals were recorded. Behavioral results showed that the precision requirements were taken into account and the kinematics of reaching was modulated depending on the object size. Similarly, reaching-related neural activity at the level of the posterior parietal and premotor cortices was modulated by the level of accuracy determined by object size. We therefore conclude that object size is engaged in the neural computations for reach planning and execution, consistent with the results from physiological studies in non-human primates., (© 2014 Federation of European Neuroscience Societies and John Wiley & Sons Ltd.)

Although there is a wealth of behavioral data regarding grasping movements in non-human primates, how posture influences the kinematics of prehensile behavior is not yet clearly understood. The purpose of this study was to examine and compare kinematic descriptions of grip behaviors while primates (macaque monkeys) were in a sitting posture or when stopping after quadrupedal locomotion (i.e., tripedal stance). Video footage taken while macaques grasped objects was analyzed frame-by-frame using digitalization techniques. Each of the two grip types considered (power and precision grips) was found to be characterized by specific, distinct kinematic signatures for both the reaching and the grasping components when those actions were performed in a sitting position. The grasping component did not differentiate in relation to the type of grip that was needed when, instead, the prehensile action took place in a tripedal stance. Quadrupedal locomotion affected the concomitant organization of prehensile activities determining in fact a similar kinematic patterning for the two grips regardless of the size of the object to be grasped. It is suggested that using a single kinematic grip patterning for all prehensile activities might be both the by-product of planning a grasping action while walking and a way to simplify motor programming during unstable tripedal stance.

Highly efficient systems are needed to link perception with action in the context of the highly complex environments in which primates move and interact. Another important component is, nonetheless, needed for action: selection. When one piece of fruit from a branch is being chosen by a monkey, many other pieces are within reach and visible: do the perceptual features of the objects surrounding a target determine interference effects? In humans, reaching to grasp a desired object appears to integrate the motor features of the objects which might become potential targets - a process which seems to be driven by inhibitory attention mechanisms. Here we show that non-human primates use similar mechanisms when carrying out goal-directed actions. The data indicate that the volumetric features of distractors are internally represented, implying that the basic cognitive operations allowing for action selection have deep evolutionary roots.

Background: The aim of this longitudinal study was to investigate how the kinematic organization of upper limb movements changes from fetal to post-natal life. By means of off-line kinematical techniques we compared the kinematics of hand-to-mouth and hand-to-eye movements, in the same individuals, during prenatal life and early postnatal life, as well as the kinematics of hand-to-mouth and reaching-toward-object movements in the later age periods., Methodology/principal Findings: Movements recorded at the 14(th), 18(th) and 22(nd) week of gestation were compared with similar movements recorded in an ecological context at 1, 2, 3, 4, 8, and 12 months after birth. The results indicate a similar kinematic organization depending on movement type (i.e., eye, mouth) for the infants at one month and for the fetuses at 22 weeks of gestation. At two and three months such differential motor planning depending on target is lost and no statistical differences emerge. Hand to eye movements were no longer observed after the fourth month of life, therefore we compared kinematics for hand to mouth with hand to object movements. Results of these analyses revealed differences in the performance of hand to mouth and reaching to object movements in the length of the deceleration phase of the movement, depending on target., Conclusion/significance: Data are discussed in terms of how the passage from intrauterine to extra-uterine environments modifies motor planning. These results provide novel evidence of how different types of upper extremity movements, those directed towards one's own face and those directed to external objects, develop.

Although facilitation of the cortico-spinal system during action observation is widely accepted, it remains controversial whether this facilitation reflects a replica of the observed movements or the goal of the observed motor acts. In the present transcranial magnetic stimulation study, we recorded motor evoked potentials from two hand muscles (first dorsal interosseous and abductor digiti minimi) while 22 healthy participants observed a hand reaching towards and grasping a bottle. To test for alternative coding levels (goal vs. movement), three relevant aspects were systematically manipulated: the type of observed movement (precision grip or whole hand grasping), situational context (bottle positioned in front of or behind a wall-like barrier), and processing stage (transcranial magnetic stimulation pulse delivered at the onset of the movement or at the moment of contact between the fingers and the object). At movement onset, motor evoked potential responses reflected the program necessary to achieve the action goal within the situational context. During movement observation, however, the type of observed movement was taken into account and a transition towards a movement-related modulation was observed. These results suggest that, rather than being exclusive alternatives, goal coding and movement coding may relate to different processing stages., (© 2013 Federation of European Neuroscience Societies and John Wiley & Sons Ltd.)

Observing the actions of others prompts the motor system to perform a similar action. However, visual cues are not the only source of sensory information for the motor system, which is affected by stimuli presented in all modalities even when they are irrelevant for action completion. The current experiment explored whether (and how) olfactory stimuli can influence the performance of a reach-to-grasp movement to visual objects differing in size (small and large) in the context of an automatic imitation task. Odours could match-or not- the size of a to-be-grasped visual target, or be nonexistent. Movement duration, an integral index of motor control, was significantly shorter when participants previously observed the same action. Addition of the odour component suggested that when the odour matched the size of a small target, a facilitation effect was found. Results are discussed in terms of olfactory-visual integration mechanisms and how they relate to embodied cognition.

Background: Anticipatory planning, the ability to anticipate future perceptual-motor demands of a goal-oriented action sequence, is essential for flexible, purposeful behavior. Once an action goal has been defined, movement details necessary to achieve that goal can be selected. Here, we investigate if anticipatory planning takes place even when multi-step actions are being carried out. How, we may ask, are the cerebral circuits involved in movement selection influenced by anticipated object-center task demands? Transcranial magnetic stimulation (TMS) was used to investigate how changes in corticospinal excitability (CSE) are dependent on anticipated task variables of intended future actions. Specifically, single- and paired-pulse TMS was used to evaluate corticospinal excitability during the action selection phase preparatory to grasp execution., Results: We found that during the premovement phase, there is an object- and muscle-specific modulation in the intrinsic hand muscle that will be used during a forthcoming grasping action. Depending on whether the participants were instructed to perform a single- or double-step movement sequence, modulation of the corticospinal output to the appropriate hand muscles was dependent on what object was to be grasped and what type of movement was being prepared. No modulation in excitability was observed during one-step movements., Conclusions: Anticipation of intended task demands plays an important role in controlling multi- step actions during which ongoing behavior may need to be adjusted. This finding supports the notion that the cortico-cortical mechanism involving movement planning is specific for an object's properties as well as for the goal of the movement sequence.

It is well known that perceiving another's body movements activates corresponding motor representations in an observer's brain. It is nevertheless true that in many situations simply imitating another's actions would not be an effective or appropriate response, as successful interaction often requires complementary rather than emulative movements. At what point does the automatic tendency to mirror another's actions become the inclination to carry out appropriate, complementary movements? In the present study, single-pulse transcranial magnetic stimulation (TMS) was used to explore corticospinal excitability in participants observing action sequences evoking imitative or complementary movements. TMS was delivered at five time points corresponding to different moments in time when key kinematic landmarks characterizing an observed action occurred. A variation in motor evoked potentials (MEPs) confirmed that the motor system flexibly shifts from imitative to complementary action tendencies. That shift appears to take place very precociously in time. Observers are attuned to advance movement information and can use it to anticipate a future course of action and to prepare for an appropriate, complementary action. Altogether, these findings represent a step forward in research concerning social action-perception coupling mechanisms providing important data to better understand the role of predictive simulation in social contexts.

Background: In non-human primates grasp-related sensorimotor transformations are accomplished in a circuit involving the anterior intraparietal sulcus (area AIP) and both the ventral and the dorsal sectors of the premotor cortex (vPMC and dPMC, respectively). Although a human homologue of such a circuit has been identified, the time course of activation of these cortical areas and how such activity relates to specific kinematic events has yet to be investigated., Methodology/principal Findings: We combined kinematic and event-related potential techniques to explicitly test how activity within human grasping-related brain areas is modulated in time. Subjects were requested to reach towards and grasp either a small stimulus using a precision grip (i.e., the opposition of index finger and thumb) or a large stimulus using a whole hand grasp (i.e., the flexion of all digits around the stimulus). Results revealed a time course of activation starting at the level of parietal regions and continuing at the level of premotor regions. More specifically, we show that activity within these regions was tuned for specific grasps well before movement onset and this early tuning was carried over--as evidenced by kinematic analysis--during the preshaping period of the task., Conclusions/significance: Data are discussed in terms of recent findings showing a marked differentiation across different grasps during premovement phases which was carried over into subsequent movement phases. These findings offer a substantial contribution to the current debate about the nature of the sensorimotor transformations underlying grasping. And provide new insights into the detailed movement information contained in the human preparatory activity for specific hand movements.

Many reports in the literature indicate that idiopathic Parkinson's disease (IPD) patients have substantial olfactory dysfunctions even before motor symptoms become evident. It has not yet been clarified, however, if some form of implicit olfactory processing is preserved in this population. An olfactory visuomotor priming paradigm, which detects implicit olfactory processing in neurologically healthy participants, was utilized to investigate motor control in relation to olfactory signals in a group of IPD patients. Two control groups were also considered: 12 vascular Parkinson's disease (VPD) in whom normal olfactory abilities are typically reported and 12 neurologically healthy participants. All of the participants were asked to perform reach-to-grasp movements toward large or small targets following olfactory cues delivered by a computer-controlled olfactometer. The odor was either 'size' congruent with the target (e.g., strawberry or apple, respectively) or incongruent (e.g., apple or strawberry, respectively). A bend sensor glove (CyberGlove) was used to measure the hand kinematics. Facilitation effects were noted in all the groups with regard to movement time. If a congruent rather than an incongruent odor was delivered, the movement time of the reach-to-grasp was shortened and facilitation effects in maximum grip amplitude were noted in both the IPD and the VPD groups. The maximum grip amplitude was smaller when no odor, as compared to a congruent odor, was delivered. The present results suggest that implicit olfactory processing affects motor control in IPD patients favoring less severe bradykinesia and hand movement hypometria. Once confirmed, these findings could be useful when rehabilitation strategies are being hypothesized for these patients., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

The prehensile hand is one of the major traits distinguishing primates from other mammal species. All primates, in fact, are able to grasp an object and hold it in part or entirely using a single hand. Although there is a wealth of behavioral data regarding grasping movements in humans and apes, there is relatively little material on macaques, the animal model often used to investigate neuronal mechanisms responsible for grip control in humans. To date, evidence regarding free-ranging macaques is confined to observational data, while quantitative reports describe studies carried out in laboratory settings or in captivity. The purpose of the present study was to provide the first kinematic descriptions of basic grip behavior with regard to precision and power grips in free-ranging macaque monkeys. Video footage of those animals grasping objects was analyzed frame-by-frame using digitalization techniques. The results revealed that the two types of grips considered are each characterized by specific kinematic signatures. It was also found that hand kinematics was scaled depending on the type of grasp needing to be adopted and the intrinsic properties of the object to be grasped. In accordance with data concerning humans, these findings indicate that the intrinsic features of an object affect the planning and control of reach-to-grasp movements even in free-ranging macaques. The data presented here take research in the field of comparative reach-to-grasp kinematics in human and non-human primates another step forward as they are based on precise measurements of spontaneous grasping movements by animals living/acting in their natural environment.

When someone is choosing one piece from a bowl full of fruit, many pieces are within reach and visible. Although the desired piece seems to govern the particular pattern and direction of that person's reaching movement, the selection process is not impervious to the presence of task-irrelevant information (i.e. the other fruits). Evidence suggests that the kinematics of reach-to-grasp actions for a desired object integrates the motor features of all the objects which might become potential targets. Transcranial magnetic stimulation (TMS) and motor-evoked potentials (MEPs) were used by us to establish if that motor integration process can be transferred to an onlooker. Our results indicate that observation of hybrid reach-to-grasp movement kinematics is reflected in the observer's pattern of MEP amplitudes. This effect can be defined as a form of motor resonance which operates by 'reading' the kinematics of an observed action. The brain's ability to mirror motor integration processes while observing someone else's action helps an onlooker to understand what the other person is doing and to predict his/her motor alternatives.

In the present study, we used functional magnetic resonance imaging to investigate the influence that the presence of an object, intended as the target for an action, might have on the neural circuit underlying the observation of the movement of others (action observation network [AON]). This system entails a simulation process of the observed movement occurring via the recruitment of the motor and somatosensory cortices involved in the execution of such movement. Here, participants were requested to observe grasping actions ending behind a partition knowing in advance whether a target-object would be present (i.e., object-directed action) or absent (i.e., non object-directed action). These "action" conditions were compared with "control" conditions in which a stationary hand was presented either alone or together with the target-object hidden behind the partition. Activation within most regions of the AON was similar for observing partially occluded object-related and non object-related actions. However, within one of the regions belonging to the AON, namely in the somatosensory cortices, blood oxygen level-dependent activity was modulated by the presence of a target-object. We interpreted such modulation as demonstration of the differential involvement of the somatosensory component of the AON for the coding of these 2 types of actions.

To investigate implicit olfactory abilities in a group of anosmic traumatic brain injured (TBI) patients, an olfactomotor priming paradigm was administered. A group of matched normosmic/mildly microsmic TBI patients and a group of neurologically healthy participants served as controls. For all the groups, an interference effect was evident on the peak velocity of grip aperture when participants grasped a large target preceded by a "small" odor. The present results suggest that some form of implicit olfactory processing is preserved in TBI patients even when diagnosed as anosmic on the basis of explicit olfactory testing.

Research on multisensory integration during natural tasks has revealed how chemical senses contribute to plan and control movements. An aspect which has yet to be investigated regards whether the motor representations evoked by chemosensory stimuli, once established for a particular movement, can be used to control different effectors. Here, we investigate this issue by asking participants to drink a sip of flavored solution, grasp with the hand a visual target, and then bring it to the mouth, miming the action of biting. Results show that hand and lip apertures were scaled according to the size of the object evoked by the flavor. Maximum hand and lip apertures were greater when the action toward a small visual target (e.g., strawberry) was preceded by a sip of a "large" (e.g., orange) than a "small" (e.g., almond) flavor solution. Conversely, maximum hand and lip apertures were smaller when the action toward a large visual target (e.g., apple) was preceded by the presentation of a "small" (e.g., strawberry) rather than a "large" flavor solution. These findings support previous evidence on the presence of a unique motor plan underlying the act of grasping with-the-hand and with-the-mouth, extending the knowledge of chemosensorimotor transformations to motor equivalence.

A large body of research reports that perceiving body movements of other people activates motor representations in the observer's brain. This automatic resonance mechanism appears to be imitative in nature. However, action observation does not inevitably lead to symmetrical motor facilitation: mirroring the observed movement might be disadvantageous for successfully performing joint actions. In two experiments, we used transcranial magnetic stimulation (TMS) to investigate whether the excitability of the corticospinal system was selectively modulated by the social dimension of an observed action. We recorded motor-evoked potentials (MEPs) from right-hand muscles during the observation of an action sequence which, depending on context, might or might not elicit a complementary response. The results demonstrate a differential motor facilitation depending on action context. Specifically, when the context called for a complementary action, the excitability pattern reflected the under-threshold activation of a complementary action, whereas when the context did not imply acting in a complementary manner, the observer's corticospinal activity reflected symmetrical motor resonance. We contend that the mechanisms underlying action observation are flexible and respond to contextual factors that guide the social interaction between individuals beyond emulation.

Previous research on multisensory integration during goal-directed natural actions reported that visual, proprioceptive, auditory and orthonasal olfactory stimulation has the ability to influence motor control. In this study, we used kinematics to investigate the integration between vision and flavour perception during reach-to-grasp movements. Participants were requested to drink a sip of flavoured solution and then grasp an object presented in central vision. The results indicate that when the objects evoked by the flavour and by the visual target were of a similar size (i.e., large or small) and evoked the same kind of hand shaping in order to be grasped (i.e., congruent condition) facilitation effects emerged. Conversely, when the object evoked by the flavour and by the visual target was of a different size and evoked a different kind of hand shaping in order to be grasped (i.e., incongruent condition) interference effects emerged. Interference effects, however, were only evident for the combination involving a large visual target and a 'small' flavour. When comparing hand kinematics between the congruent and a 'no flavour' condition (i.e., water), facilitation effects emerged in favour of the former condition. Taken together, these results indicate the contribution of complex chemosensory stimuli for the planning and execution of visually guided reach to grasp movements. And, contribute to the current debate regarding the multisensory nature of the sensorimotor transformations underlying motor performance., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

Previous evidence indicates that we understand others' actions not only by perceiving their visual features but also by their sound. This raises the possibility that brain regions responsible for action understanding respond to cues coming from different sensory modalities. Yet no studies, to date, have examined if this extends to olfaction. Here we addressed this issue by using functional magnetic resonance imaging. We searched for brain activity related to the observation of an action executed towards an object that was smelled rather than seen. The results show that temporal, parietal, and frontal areas were activated when individuals observed a hand grasping a smelled object. This activity differed from that evoked during the observation of a mimed grasp. Furthermore, superadditive activity was revealed when the action target-object was both seen and smelled. Together these findings indicate the influence of olfaction on action understanding and its contribution to multimodal action representations.

Previous evidence suggests that hand shaping during reaching is modulated by the presence and the nature of the end-goal following object's grasp. Here we test whether such modulation is maintained in Parkinson's disease (PD). Six participants with PD and six healthy participants took part in the study. Participants were requested to reach towards a bottle filled with water, and then: (1) grasp it without performing any subsequent action; (2) grasp it and place it accurately on a target area; (3) grasp it and pour its contents within a container. The results showed that participants shaped their hand differently depending on the presence or absence of an action following object's grasp. However, the request to perform an action after grasp determined a modulation of hand kinematics which was delayed for PD than for control participants. Further, whereas for control participants the nature of the end-goal determined a modulation of hand shaping, for PD patients such modulation was not evident. Data are discussed in terms of the role played by basal ganglia in implementing anticipatory mechanisms for the control of manipulative activities. We contend that in PD patients these mechanisms are not totally compromised, but their implementation depends on the action information that has to be anticipated., (Copyright 2010 Elsevier Inc. All rights reserved.)

Four experiments investigated the influence of a sudden social request on the kinematics of a preplanned action. In Experiment 1, participants were requested to grasp an object and then locate it within a container (unperturbed trials). On 20% of trials, a human agent seated nearby the participant unexpectedly stretched out her arm and unfolded her hand as if to ask for the object (perturbed trials). In the remaining 3 experiments, similar procedures were adopted except that (a) the human was replaced by a robotic agent, (b) the gesture performed by the human agent did not imply a social request, and (c) the gaze of the human agent was not available. Only when the perturbation was characterized by a social request involving a human agent were there kinematic changes to the action directed toward the target. Conversely, no effects on kinematics were evident when the perturbation was caused by the robotic agent or by a human agent performing a nonsocial gesture. These findings are discussed in the light of current theories proposed to explain the effects of social context on the control of action., (PsycINFO Database Record (c) 2009 APA, all rights reserved.)

Recent fMRI evidence indicates that both the execution and the observation of hand actions in multiple sclerosis (MS) patients increase recruitment of a portion of the so-called mirror neuron system. However, it remains unclear whether this is the expression of a compensatory mechanism for the coding of observed action or whether such a mechanism represents a rather unspecific functional adaptation process. Here we used fMRI on early relapsing remitting MS (RRMS) patients to clarify this issue. Functional images of 15 right-handed early RRMS patients and of 15 sex- and age-matched right-handed healthy controls were acquired using a 1.5 T scanner. During scanning, participants simply observed images depicting a human hand either grasping an object or resting alongside an object. As shown by a between-group analysis, when compared to controls, RRMS patients revealed a robust increase of activation in an extensive network of brain regions including frontal, parietal, temporal and visual areas usually activated during action observation. However, this pattern of hemodynamic activity was completely independent of the type of observed hand-object interaction as revealed by the lack of any significant between-group interaction. Our findings are in line with previous fMRI evidence demonstrating cortical reorganization in MS patients during action observation. However, based on our findings we go one step further and suggest that such functional cortical changes may be the expression of a generalized and unspecific compensatory mechanism, that is not necessarily involved in action understanding.

Previous neuroimaging research on healthy humans has provided evidence for a neural system underlying the observation of another person's hand actions. However, whether the neural processes involved in this capacity are activated by the observation of other transitive hand actions such as pointing remains unknown. Therefore, using functional magnetic resonance imaging we investigated the neural mechanisms underlying the observation of static images representing the hand of a human model pointing to an object (pointing condition), grasping an object (grasping condition), or resting in proximity of an object (control condition). The results indicated that activity within portions of the lateral occipitotemporal and the somatosensory cortices modulates according to the type of observed transitive actions. Specifically, these regions were more activated for the grasping than for the pointing condition. In contrast, the premotor cortex, a neural marker of action observation, did not show any differential activity when contrasting the considered experimental conditions. Our findings may provide novel insights regarding a possible role of extrastriate and somatosensory brain areas for the perception of distinct types of human hand-object interactions.

Objects can be grasped in several ways due to their physical properties, the context surrounding the object, and the goal of the grasping agent. The aim of the present study was to investigate whether the prior-to-contact grasping kinematics of the same object vary as a result of different goals of the person grasping it. Subjects were requested to reach toward and grasp a bottle filled with water, and then complete one of the following tasks: (1) Grasp it without performing any subsequent action; (2) Lift and throw it; (3) Pour the water into a container; (4) Place it accurately on a target area; (5) Pass it to another person. We measured the angular excursions at both metacarpal-phalangeal (mcp) and proximal interphalangeal (pip) joints of all digits, and abduction angles of adjacent digit pairs by means of resistive sensors embedded in a glove. The results showed that the presence and the nature of the task to be performed following grasping affect the positioning of the fingers during the reaching phase. We contend that a one-to-one association between a sensory stimulus and a motor response does not capture all the aspects involved in grasping. The theoretical approach within which we frame our discussion considers internal models of anticipatory control which may provide a suitable explanation of our results.

The ability to understand another person's action and, if needed, to imitate that action, is a core component of human social behaviour. Imitation skills have attracted particular attention in the search for the underlying causes of the social difficulties that characterize autism. In recent years, it has been reported that people with autism can bypass some of their social deficits by interacting with robots. However, the robot preference in terms of imitation has yet to be proved. Here we provide empirical evidence that interaction with robots can trigger imitative behaviour in children with autism. We compared a group of high functioning children with autism with a group of typically developing children in a visuomotor priming experiment. Participants were requested to observe either a human or a robotic arm model performing a reach-to-grasp action towards a spherical object. Subsequently, the observers were asked to perform the same action towards the same object. Two 'control' conditions in which participants performed the movement in the presence of either the static human or robot model were also included. Kinematic analysis was conducted on the reach-to-grasp action performed by the observer. Our results show that children with autism were facilitated - as revealed by a faster movement duration and an anticipated peak velocity - when primed by a robotic but not by a human arm movement. The opposite pattern was found for normal children. The present results show that interaction with robots has an effect on visuomotor priming processes. These findings suggest that in children with autism the neural mechanism underlying the coding of observed actions might be tailored to process socially simpler stimuli.

Event-related functional magnetic resonance imaging (fMRI) was used to explore how the human brain models gaze-object relations. During scanning participants observed a human model gazing towards or away a target object presented either in isolation or flanked by a distractor object. In two further conditions the model's gaze was shifted and subsequently maintained away from the stimulus/i. These four conditions were implemented within a factorial design in which the main factors were "type of observed behavior" (gaze vs. gaze-away) and "context" (target alone vs. target flanked by a distractor). Results revealed that premotor, parietal and temporal areas, known to sub-serve the understanding of other people actions, were significantly more activated by the observation of the model gazing towards rather than away from the stimulus/i. In addition, a significant interaction indicated that, when the target was presented in isolation, neural activity within the inferior frontal gyrus, another key area for action understanding, was influenced by gaze-object relations. Our findings suggest that this area is important for the establishment of intentional gaze-object relations and indicate that the presence of a distractor interferes with the representation of such relations.

This study assessed how hand shaping responds to a perturbation of object shape. In blocked trials (80% of total), subjects were instructed to reach, to grasp and lift a concave or a convex object. In perturbed trials (20% of total), a rotating device allowed for the rapid change from the concave to the convex object or vice versa. In this situation subjects grasped the last presented object. Flexion/extension at the metacarpal-phalangeal and proximal interphalangeal joints of all digits was measured by resistive sensors embedded in a glove. In the blocked condition we found that most joints of the fingers were modulated by the type of the to-be-grasped object during the reach. When object shape was perturbed, reach duration was longer and angular excursion of all fingers differed with respect to blocked trials. For the 'convex --> concave' perturbation, a greater degree of finger extension was found than during the blocked 'concave' trials. In contrast, for the 'concave --> convex' perturbation, fingers were more flexed than for the blocked 'convex' trials. The thumb reacted to the perturbation showing a similar pattern (i.e., over-flexion with respect to the blocked trials) regardless the 'direction' of the perturbation. The present results suggest that applying an object shape perturbation during a reach-to-grasp action determines a reorganization of all digits. This pattern is suggestive of a control strategy, which assigns to opposing digits different roles.

The selection of objects in the visual environment is important in everyday life when acting in a goal-directed manner. Here we used functional magnetic resonance imaging (fMRI) to investigate brain activity while healthy subjects (N=15) selectively reached to grasp a three-dimensional (3D) target stimulus presented either in isolation or in the presence of 3D non-target stimuli. A pneumatic MRI compatible apparatus was designed to precisely control the presentation of 3D graspable stimuli within the scanner. During scanning subjects were instructed to reach and grasp towards a target presented at an unknown location either in isolation or flanked by two distractor objects. Results indicated that reaching towards and grasping the target object in the presence of other non-target stimuli was associated with greater activation within the contralateral primary motor cortex and the precuneus as compared to the execution of reach-to-grasp movements towards the target presented in isolation. We conclude that the presence of non-targets evokes a differential level of neural activity within areas responsible for the planning and execution of selective reach-to-grasp movement.

The aim of the present study is to elucidate the influence of context on the kinematics of the reach-to-grasp movement. In particular, we consider two basic modes of social cognition, namely cooperation and competition. In two experiments kinematics of the very same action - reaching-to-grasp a wooden block - were analyzed in two different contexts provided by a cooperative task and competitive task. For the 'cooperation' tasks two participants were required to reach and grasp their respective objects and to cooperate to join the two objects in specific configurations in the middle of the working surface. For the 'competition' tasks, the two participants had to compete to place their own object first in the middle of the working surface. Results revealed specific kinematic patterns for cooperation and competition which were distinct from similar actions performed by each participant in isolation. Further, during the cooperation tasks, a high level of correlation between key kinematical parameters of the two participants was found. In accordance with evidence from neuroimaging, developmental and social psychology our results suggest the existence of motor patterns which reflect the intention to act in a social context.

The aim of the present study was to investigate whether foetal hand movements are planned and how they are executed. We performed a kinematic analysis of hand movements directed towards the mouth and the eyes in the foetuses of eight women with normally evolving pregnancies. At 14, 18 and 22 weeks of gestation, eight foetuses underwent a 20-min four-dimensional-ultrasound session. The video recordings for these movements were then imported into in-house software developed to perform kinematic analysis. We found that spatial and temporal characteristics of foetal movements are by no means uncoordinated or unpatterned. By 22 weeks of gestation the movements seem to show the recognizable form of intentional actions, with kinematic patterns that depend on the goal of the action, suggesting a surprisingly advanced level of motor planning.

Previous research has provided evidence for a neural system underlying the observation of another person's hand actions. Is the neural system involved in this capacity also important in inferring another person's motor intentions toward an object from their eye gaze? In real-life situations, humans use eye movements to catch and direct the attention of others, often without any accompanying hand movements or speech. In an event-related functional magnetic resonance imaging study, subjects observed videos showing a human model either grasping a target object (grasping condition) or simply gazing (gaze condition) at the same object. These two conditions were contrasted with each other and against a control condition in which the human model was standing behind the object without performing any gazing or grasping action. The results revealed activations within the dorsal premotor cortex, the inferior frontal gyrus, the inferior parietal lobule, and the superior temporal sulcus in both "grasping" and "gaze" conditions. These findings suggest that signaling the presence of an object through gaze elicits in an observer a similar neural response to that elicited by the observation of a reach-to-grasp action performed on the same object.

People can adjust their reach-to-grasp movements online to sudden changes in the spatial properties of a target. We investigated whether they can also do this when a non-spatial property, weight, suddenly changes. Guiding your movement by using visual cues about an object's weight depends heavily on experience and is expected to be processed by the (slow) ventral stream rather than the (fast) 'online control' dorsal stream. In the first experiment, participants reached out and lifted an object with an expected or an unexpected weight. As predicted, there was an effect of expected weight on the time between the end of the reaching phase and the object's lift-off. In the second experiment, the object sometimes visibly changed weight after the participants had started their movement. The lifting time did not depend on whether the object had changed weight. Thus, participants can make online adjustments to a visually indicated change in weight. These results are interpreted as being contrary to existing theories of online control.

The aim of the present study was to determine whether hand shaping was affected by planning of an action subsequent to object contact. Ten subjects (5 females and 5 males, ages 19-33) were requested to reach toward and grasp a convex object between the thumb and the four fingers of the right hand and to perform one of the following actions: 1) lift up the object; 2) insert the object into a niche of a similar shape and size as the object, or 3) insert the object into a rectangular niche much larger than the object. Flexion/extension at the metacarpal-phalangeal and proximal interphalangeal joints of all digits were measured using resistive sensors embedded in a glove. Although all experimental conditions required grasping the same object, we found different covariation patterns among finger joint angles across conditions. Gradual preshaping of the hand occurred only when planning object lift or when the end-goal required object placement into the tight niche. In contrast, for the larger niche, gradual preshaping was not evident for the ring and the little finger. Further, reaching movements were faster for movements ending with the larger niche than for the other movement conditions. The present results suggest that hand shaping takes into account end-goal in addition to object geometry. We discuss these findings in the context of forward internal models that allow the prediction of the sensorimotor consequences of motor commands in advance to their execution.

Previous evidence based on perceptual integration and arbitrary responses suggests extensive cross-modal links in attention across the various modalities. Attention typically shifts to a common location across the modalities, despite the vast differences in their initial coding of space. An issue that remains unclear is whether or not these effects of multisensory coding occur during more natural tasks, such as grasping and manipulating three-dimensional objects. Using kinematic measures, we found strong effects of the diameter of a grasped distractor object on the aperture used to grasp a target object at both coincident and non-coincident locations. These results suggest that interference effects can occur between proprioceptive and visuomotor signals in grasping. Unlike other interference effects in cross-modal attention, these effects do not depend on the spatial relation between target and distractor, but occur within an object-based frame of reference.

In this study, the reach-to-grasp movement of 5-year-old children was compared to that of adults. Participants were required to reach out and grasp objects, with and without on-line visual feedback. Object size and distance were covaried in a within-subjects design and it was found that for both groups, grip formation and reach kinematics were affected by the manipulation of either variable. Although there are a large number of similarities, a few differences between the two groups emerge. For the reaching component, the children revealed a longer movement duration and deceleration time and a lower maximum height of wrist trajectory than in adults. For the grasp component, the children, in both the vision and no-vision condition, show a maximal finger aperture larger than the adults. Further, the children of this study were able to scale their grip aperture according to object size when visual feedback during the movement was lacking. These findings suggest that children adopt different strategies than adults when planning a reach-to-grasp movement on the basis of object size, distance, and the predictability of visual feedback. The results are discussed in terms of the neural mechanisms underlying hand action and how these mechanisms may not be fully developed by the age of 5.

Although the effect of visual illusions on overt actions has been an area of keen interest in motor performance, no study has yet examined whether illusions have similar or different effects on overt and imagined movements. Two experiments were conducted that compared the effects of an orientation illusion on an overt posture selection task and an imagined posture selection task. In Experiment 1 subjects were given a choice of grasping a bar with the thumb on the left side or right side of the bar. In Experiment 2 subjects were instructed to only imagine grasping the bar while remaining motionless. Subjects then reported which side of the bar their thumb had been placed in imagined grasping. Both the overt selection and imagined selection tasks were found to be sensitive to the orientation illusion, suggesting that similar visual information is used for overt and imagined movements, with both being sensitive to an orientation illusion. The results are discussed in terms of the visual processing and representation of real and imagined actions.

The aim of this study is to describe the reaching action of children with developmental coordination disorder (DCD) and to investigate whether their use of visual feedback during the time course of this action differs from that of normally developing children. Fifty-two children subdivided into 2 age bands (7-8 and 9-10 years) within 2 groups (with and without DCD) participated in this experiment. They were asked to reach for a target positioned either ipsilaterally or contralaterally to the reaching hand in 2 visual conditions: a condition where vision was unrestrained (normal vision) and a condition where they wore glasses with prismatic lenses (perturbed vision). An analysis of the experimental data indicates that the trajectories followed by the DCD group were longer and more curved than those of the control group. Further, the deceleration times were longer for the DCD group than for the normally developing children. The introduction of the prismatic lenses supports the idea that the use of visual feedback by children with DCD may be different from that in children without DCD.

Previous work has shown that both human adults and children attend to grasping actions performed by another person but not necessarily to those made by a mechanical device. According to recent neurophysiological data, the monkey premotor cortex contains "mirror" neurons that discharge both when the monkey performs specific manual grasping actions and when it observes another individual performing the same or similar actions. However, when a human model uses tools to perform grasping actions, the mirror neurons are not activated. A similar "mirror" system has been described in humans, but whether or not it is also tuned specifically to biological actions has never been tested. Here we show that when subjects observed manual grasping actions performed by a human model a significant neural response was elicited in the left premotor cortex. This activation was not evident for the observation of grasping actions performed by a robot model commanded by an experimenter. This result indicates for the first time that in humans the mirror system is biologically tuned. This system appears to be the neural substrate for biological preference during action coding.

Shadows in visual scenes can have profound effects on visual perception. Here we have found that visual distracters distant from the body interfere with human spatial discrimination of tactile targets at a hand, particularly when the shadow of the stimulated hand stretches toward them in extrapersonal space. These findings suggest that shadows cast by a person's own body parts can bridge the gap between personal and extrapersonal space.