Increasing striatal dopamine release through repeated bouts of theta burst transcranial magnetic stimulation of the left dorsolateral prefrontal cortex. A 18F-desmethoxyfallypride positron emission tomography study.

Introduction: Transcranial Magnetic Stimulation (TMS) can modulate fronto-striatal connectivity in the human brain. Here Positron Emission Tomography (PET) and neuro-navigated TMS were combined to investigate the dynamics of the fronto-striatal connectivity in the human brain. Employing 18F-DesmethoxyFallypride (DMFP) - a Dopamine receptor-antagonist - the release of endogenous dopamine in the striatum in response to time-spaced repeated bouts of excitatory, intermittent theta burst stimulation (iTBS) of the Left-Dorsolateral Prefrontal Cortex (L-DLPFC) was measured. Methods: 23 healthy participants underwent two PET sessions, each one with four blocks of iTBS separated by 30 minutes: sham (control) and verum (90% of individual resting motor threshold). Receptor Binding Ratios were collected for sham and verum sessions across 37 time frames (about 130 minutes) in striatal sub-regions (Caudate nucleus and Putamen). Results: Verum iTBS increased the dopamine release in striatal sub-regions, relative to sham iTBS. Dopamine levels in the verum session increased progressively across the time frames until frame number 28 (approximately 85 minutes after the start of the session and after three iTBS bouts) and then essentially remained unchanged until the end of the session. Conclusion: Results suggest that the short-timed iTBS protocol performed in time-spaced blocks can effectively induce a dynamic dose dependent increase in dopaminergic fronto-striatal connectivity. This scheme could provide an alternative to unpleasant and distressing, long stimulation protocols in experimental and therapeutic settings. Specifically, it was demonstrated that three repeated bouts of iTBS, spaced by short intervals, achieve larger effects than one single stimulation. This finding has implications for the planning of therapeutic interventions, for example, treatment of major depression.

Anthropomorphic or non-anthropomorphic? Effects of biological sex in observation of actions in a digital human model and a gantry robot model.

Robots are ever more relevant for everyday life, such as healthcare or rehabilitation, as well as for modern industrial environment. One important issue in this context is the way we perceive robots and their actions. From our previous study, evidence exists that sex can affect the way people perceive certain robot's actions. In our fMRI study, we analyzed brain activations of female and male participants, while they observed anthropomorphic and robotic movements performed by a human or a robot model. While lying in the scanner, participants rated the perceived level of anthropomorphic and robotic likeness of movements in the two models. The observation of the human model and the anthropomorphic movements similarly activated the biological motion coding areas in posterior temporal and parietal areas. The observation of the robot model activated predominantly areas of the ventral stream, whereas the observation of robotic movements activated predominantly the primary and higher order motor areas. To note, this later activation originated mainly from female participants, whereas male participants activated, in both robot model and robotic movements contrasts, areas in the posterior parietal cortex. Accordingly, the general contrast of sex suggests that men tend to use the ventro-dorsal stream most plausibly to rely on available previous knowledge to analyze the movements, whereas female participants use the dorso-dorsal and the ventral streams to analyze online the differences between the movement types and between the different models. The study is a first step toward the understanding of sex differences in the processing of anthropomorphic and robotic movements.

Respiratory function modulated during execution, observation, and imagination of walking via SII.

The Mirror Neurons System (MNS) consists of brain areas active during actions execution, as well as observation-imagination of the same actions. MNS represents a potential mechanism by which we understand other's action goals. We investigated MNS activation for legs actions, and its interaction with the autonomic nervous system. We performed a physiological and fMRI investigation on the common neural structures recruited during the execution, observation, and imagination of walking, and their effects on respiratory activity. Bilateral SMA were activated by all three tasks, suggesting that these areas are responsible for the core of the MNS effect for walking. Moreover, we observed in bilateral parietal opercula (OP1, secondary somatosensory cortex-SII) evidence of an MNS subtending walking execution-observation-imagination that also modulated the respiratory function. We suggest that SII, in modulating the vegetative response during motor activity but also during observation-imagination, consists of a re-enacting function which facilitates the understanding of motor actions.

Motor sequence learning in patients with ideomotor apraxia: Effects of long-term training.

Recent studies show that limb apraxia is a quite frequent, yet often underdiagnosed, higher motor impairment following stroke. Because it adversely affects every-day life and personal independence, successful rehabilitation of apraxia is essential for personal well-being. Nevertheless, evidence of long-term efficacy of training schemes and generalization to untrained actions is still scarce. One possible reason for the tendency of this neurological disorder to persist may be a deficit in planning, conceptualisation and storage of complex motor acts. This pilot study aims at investigating explicit motor learning in apractic stroke patients. In particular, we addressed the ability of apractic patients to learn and to retain new explicit sequential finger movements across 10 training sessions over a 3-week interval. Nine stroke patients with ideomotor apraxia in its chronic stage participated in a multi-session training regimen and were included in data analyses. Patients performed an explicit finger sequence learning task (MSLT - motor sequence learning task), which is a well-established paradigm to investigate motor learning and memory processes. Patients improved task performance in terms of speed and accuracy across sessions. Specifically, they showed a noticeable reduction in the mean time needed to perform a correct sequence and the number of erroneous sequences. We found also a trend for improved performance at the Goldenberg apraxia test protocol: "imitation of meaningless hand and finger gestures" relative to when assessed before the MSLT training. Patients with ideomotor apraxia demonstrated the ability to acquire and maintain a novel sequence of movements; and, this training was associated with hints towards improvement of apraxia symptoms.

Embodied negation and levels of concreteness: A TMS study on German and Italian language processing.

According to the embodied cognition perspective, linguistic negation may block the motor simulations induced by language processing. Transcranial magnetic stimulation (TMS) was applied to the left primary motor cortex (hand area) of monolingual Italian and German healthy participants during a rapid serial visual presentation of sentences from their own language. In these languages, the negative particle is located at the beginning and at the end of the sentence, respectively. The study investigated whether the interruption of the motor simulation processes, accounted for by reduced motor evoked potentials (MEPs), takes place similarly in two languages differing on the position of the negative marker. Different levels of sentence concreteness were also manipulated to investigate if negation exerts generalized effects or if it is affected by the semantic features of the sentence. Our findings indicate that negation acts as a block on motor representations, but independently from the language and words concreteness level.

The prominent role of perceptual salience in object discrimination: overt discrimination of graspable side does not activate grasping affordances.

Responses to object stimuli are often faster when jutting handles are aligned with responding hands, than when they are not: handle-to-hand correspondence effects. According to a location coding account, locations of visually salient jutting parts determine the spatial coding of objects. This asymmetry then facilitates same-sided responses compared to responses on the opposite side. Alternatively, this effect has been attributed to grasping actions of the left or the right hand afforded by the handle orientation and independent of its salience (affordance activation account). Our experiments were designed to disentangle the effects of pure salience from those of affordance activations. We selected pictures of tools with one salient and non-graspable side, and one graspable and non-salient side (non-jutting handle). Two experiments were run. Each experiment had two groups of participants: one group discriminated the location of the salient side of the object stimuli; the other group discriminated the location of the graspable side of them. In Experiment 1, responses were left and right button presses; in Experiment 2, they were left and right button presses plus reach-and-grasp actions. When visual salience was removed from graspable sides, no correspondence effect was observed between their orientation and the responding hands in both the experiments. Conversely, when salience depended on non-graspable portions, a correspondence effect was produced between their orientation and the responding hand. Overt attention to graspable sides did not potentiate any grasping affordance even when participants executed grasping responses in addition to button presses. Results support the location coding account: performance was influenced by the spatial coding of visually salient properties of objects.

Gender Effects in Observation of Robotic and Humanoid Actions.

Robots are gaining an increasingly important role in industrial production. Notably, a high level of acceptance is an important factor for co-working situation between human and robot. The aim of the present study was to investigate the differences in the perception of anthropomorphic and robotic movements using models consisting of a virtual robot and a digital human. Videos of each model displayed different degrees of human likeness or robot likeness in speed and trajectories of placing movements. Female and male participants were asked to rate on a Likert scale the perceived levels of human likeness or robot likeness in the two models. Overall, results suggest that males were sensitive to the differences between robotic and anthropomorphic movements, whereas females showed no difference between them. However, compared to males, female participants attributed more anthropomorphic features to robotic movements. The study is a first step toward a more comprehensive understanding of the human ability to differentiate between anthropomorphic and robotic movements and suggests a crucial role of gender in the human-robot interaction.

Correspondence effect driven by salient visual asymmetries in integral object stimuli.

The handle-to-hand correspondence effect consists of faster and more accurate responses when the responding hand is aligned with the handle side of an object tool, compared to when they lay on opposite sides. This effect has been attributed to the activation of affordances. Recent studies, however, claimed that it may depend on the spatial coding of the object on the basis of its visual asymmetry (location-coding account). Affordances are namely direct and meaningful relations between recognized objects and the observers' action system. Therefore, any manipulation that disrupts the body structure of object tools could potentially affect their identification and prevent the activation of affordances. The present study investigated the nature of the handle-to-hand correspondence effects by manipulating structural asymmetry and visual salience of object tools, while preserving their integrity that is, leaving unaltered the original possibilities to activate grasping affordances. Three experiments were run. Results were consistent with the location-coding account and claim for accurate control of visual asymmetries in object stimuli during investigation of affordance effects.

Proton Magnetic Resonance Spectroscopy of the motor cortex reveals long term GABA change following anodal Transcranial Direct Current Stimulation.

Anodal transcranial direct current stimulation (tDCS) over the primary motor cortex (M1) has been reported to increase the firing rates of neurons and to modulate the gamma-aminobutyric acid (GABA) concentration. To date, knowledge about the nature and duration of these tDCS induced effects is incomplete. We aimed to investigate long-term effects of anodal tDCS over M1 on GABA dynamics in humans. Repeated magnetic resonance spectroscopy (MRS) was employed to measure relative GABA concentration in M1 for approximately 64 minutes after stimulation. The study was performed on 32 healthy subjects. Either anodal or sham tDCS were applied for 10 minutes with the active electrode over the left M1 and the reference electrode over the right supra-orbital region. Pre and post-tDCS MRS scans were performed to acquire GABA-edited spectra using 3 T Prisma Siemens scanner. GABA signals showed no change over time in the sham tDCS group, whereas anodal tDCS resulted in a significant early decrease within 25 minutes after tDCS and then significant late decrease after 66 minutes which continued until the last test measurements. The late changes in GABA concentration might be related to long-term plasticity mechanism. These results contribute to a better understanding of the neurochemical mechanism underlying long-term cortical plasticity following anodal tDCS.

The unimanual handle-to-hand correspondence effect: evidence for a location coding account.

The handle-to-hand correspondence effect refers to faster and more accurate responses when the responding hand is aligned with the graspable part of an object tool, compared to when they lay on opposite sides. We performed four behavioral experiments to investigate whether this effect depends on the activation of grasping affordances (affordance activation account) or is to be traced back to a Simon effect, resulting from the spatial coding of stimuli and responses and from their dimensional overlap (location coding account). We manipulated the availability of a response alternative by requiring participants to perform either a unimanual go/no-go task (absence of a response alternative) or a joint go/no-go task (available response alternative) and the type of response required (button-press or grasping response). We found no handle-to-hand correspondence effect in the individual go/no-go task either when a button-press (Experiment 1A) or a grasping (Experiment 2A) response was required, whereas a significant effect emerged in the joint go/no-go task, irrespective of response modality (Experiments1B and 2B). These results do not support the idea that complex motor affordances are activated for meaningful objects, but are rather consistent with the more parsimonious location coding account.

Location-coding account versus affordance-activation account in handle-to-hand correspondence effects: Evidence of Simon-like effects based on the coding of action direction.

An increasing number of studies have shown a close link between perception and action, which is supposed to be responsible for the automatic activation of actions compatible with objects' properties, such as the orientation of their graspable parts. It has been observed that left and right hand responses to objects (e.g., cups) are faster and more accurate if the handle orientation corresponds to the response location than when it does not. Two alternative explanations have been proposed for this handle-to-hand correspondence effect: location coding and affordance activation. The aim of the present study was to provide disambiguating evidence on the origin of this effect by employing object sets for which the visually salient portion was separated from, and opposite to the graspable 1, and vice versa. Seven experiments were conducted employing both single objects and object pairs as visual stimuli to enhance the contextual information about objects' graspability and usability. Notwithstanding these manipulations intended to favor affordance activation, results fully supported the location-coding account displaying significant Simon-like effects that involved the orientation of the visually salient portion of the object stimulus and the location of the response. Crucially, we provided evidence of Simon-like effects based on higher-level cognitive, iconic representations of action directions rather than based on lower-level spatial coding of the pure position of protruding portions of the visual stimuli. (PsycINFO Database Record

Interactive effects between gaze direction and facial expression on attentional resources deployment: the task instruction and context matter.

In three experiments, we tested whether the amount of attentional resources needed to process a face displaying neutral/angry/fearful facial expressions with direct or averted gaze depends on task instructions, and face presentation. To this end, we used a Rapid Serial Visual Presentation paradigm in which participants in Experiment 1 were first explicitly asked to discriminate whether the expression of a target face (T1) with direct or averted gaze was angry or neutral, and then to judge the orientation of a landscape (T2). Experiment 2 was identical to Experiment 1 except that participants had to discriminate the gender of the face of T1 and fearful faces were also presented randomly inter-mixed within each block of trials. Experiment 3 differed from Experiment 2 only because angry and fearful faces were never presented within the same block. The findings indicated that the presence of the attentional blink (AB) for face stimuli depends on specific combinations of gaze direction and emotional facial expressions and crucially revealed that the contextual factors (e.g., explicit instruction to process the facial expression and the presence of other emotional faces) can modify and even reverse the AB, suggesting a flexible and more contextualized deployment of attentional resources in face processing.

The Simon effect in action: planning and/or on-line control effects?

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.

Interhemispheric vs. stimulus-response spatial compatibility effects in bimanual reaction times to lateralized visual stimuli.

In the present study, we tested right- and left-handed participants in a Poffenberger paradigm with bimanual responses and hands either in an anatomical or in a left-right inverted posture. We observed a significant positive crossed-uncrossed difference (CUD) in RTs for both manual dominance groups and both response postures. These results rule out an explanation of the CUD in terms of stimulus-response spatial compatibility (SRSC) and provide convincing evidence on the important role of interhemispheric callosal transfer in bimanual responding in right- as well as left-handed individuals.

Skill memory escaping from distraction by sleep--evidence from dual-task performance.

BACKGROUND: Sleep facilitates off-line consolidation of memories, as shown for learning of motor skills in the absence of concomitant distractors. We often perform complex tasks focusing our attention mostly on one single part of them. However, we are equally able to skillfully perform other concurrent tasks. One may even improve performance on disregarded parts of complex tasks, which were learned implicitly. In the present study we investigated the role of sleep in the off-line consolidation of procedural skills when attention is diverted from the procedural task because of interference from a concurrent task. METHODOLOGY/PRINCIPAL FINDINGS: We used a dual-task paradigm containing (i) procedural serial reaction time task (SRTT), which was labeled as subordinate and unimportant and (ii) declarative word-pair association task (WPAT), performed concomitantly. The WPAT served as a masked distractor to SRTT and was strongly reinforced by the instructions. One experimental and three control groups were tested. The experimental group was re-tested after two nights of sleep (sleep group, SG). The first control group had sleep deprivation on the first post-learning night (nighttime-awake group, NA), the second control group was tested in the morning and then re-tested after 12-hours (daytime-awake group, DA); the third one had the same assignments as DA but with a subsequent, instead of a concomitant, WPAT (daytime-awake-subsequent-WPAT group, DAs). We found SRTT performance gains in SG but not in NA and DA groups. Furthermore, SG reached similar learning gains in SRTT as the DAs group, which gained in SRTT performance because of post-training interference from the declarative task. CONCLUSIONS/SIGNIFICANCE: The results demonstrate that sleep allows off-line consolidation, which is resistant to deteriorating effects of a reinforced distractor on the implicit procedural learning and allowing for gains which are consistent with those produced when inhibited declarative memories of SRTT do not compete with procedural ones.

Gaze direction and facial expressions exert combined but different effects on attentional resources.

Gaze direction and facial expressions are critical components of face processing and have been shown to influence attention deployment. We investigated whether gaze direction (direct vs. averted) combined with a neutral or angry expression modulates the deployment of attentional resources over time. In a rapid serial visual presentation (RSVP) paradigm participants had to decide the gender of a neutral or an angry target face with direct or averted gaze (T1) and then to judge the orientation of a target picture of a landscape (T2), following the face at different time intervals. Results showed no attentional blink effect (i.e., no deterioration in T2 accuracy) when T1 was an angry face with direct gaze, whereas it was present for angry faces with averted gaze or neutral faces with either averted or direct gaze. These findings are consistent with appraisal theories and are discussed against the background of automatic processing of threat stimuli.

Influence of temporal overlap on time course of the Simon effect.

Two experiments are reported in which we manipulated relevant and irrelevant stimulus dimensions to assess whether an increase in temporal overlap would influence the time course of a "standard" Simon effect (obtained when visual stimuli are presented on the left/right of the screen and left/right responses are performed with uncrossed hands). This procedure is new in two ways: First, the manipulations were intended to reduce, instead of increase, the distance between conditional and unconditional response-activation processes. Second, we manipulated the relevant and irrelevant stimulus dimensions in a manner that did not vary stimulus onset asynchronies, precues, or go/no go trials, or alter the stimulus quality. Results were consistent with the hypothesis that when the two response processes are shifted closer to each other, the Simon effect would be sustained across time, instead of decreasing as typically found. These findings are discussed in line with the temporal overlap hypothesis and with an automatic activation account.

Simon-like and functional affordance effects with tools: the effects of object perceptual discrimination and object action state.

In the present study two separate stimulus-response compatibility effects (functional affordance and Simon-like effects) were investigated with centrally presented pictures of an object tool (a torch) characterized by a structural separation between the graspable portion and the goal-directed portion. In Experiment 1, participants were required to decide whether the torch was red or blue, while in Experiment 2 they were required to decide whether the torch was upright or inverted. Our results showed that with the same stimulus two types of compatibility effect emerged: one based on the direction signalled by the goal-directed portion of the tool (a Simon-like effect as observed in Experiment 1), and the other based on the actions associated with an object (a functional affordance effect as observed in Experiment 2). Both effects emerged independently of the person's intention to act on the stimulus, but depended on the stimulus properties that were processed in order to perform the task.