Neural bases of loss aversion when choosing for oneself versus known or unknown others.

Despite the ubiquitous interdependence between one's own decisions and others' welfare, and the controversial evidence on the behavioral effect of choosing for others, the neural bases of making decisions for another versus oneself remain unexplored. We investigated whether loss aversion (LA; the tendency to avoid losses over approaching equivalent gains) is modulated by (i) choosing for oneself, other individuals, or both; (ii) knowing or not knowing the other recipients; or (iii) an interaction between these factors. We used fMRI to assess the brain activations associated with choosing whether to accept or reject mixed gambles, either for oneself, for another player, or both, in 2 groups of 28 participants who had or had not briefly interacted with the other players before scanning. Participants displayed higher LA for choices involving their payoff compared with those affecting only the payoff of other, known, players. This "social" modulation of decision-making was found to engage the dorsomedial prefrontal cortex and its inhibitory connectivity to the middle cingulate cortex. This pattern might underpin decision-making for known others via self-other distinction processes associated with dorsomedial prefrontal areas, with this in turn promoting the inhibition of socially oriented responses through the downregulation of the midcingulate node of the empathy network.

Evidence for a cost of time in the invigoration of isometric reaching movements.

How the brain determines the vigor of goal-directed movements is a fundamental question in neuroscience. Recent evidence has suggested that vigor results from a trade-off between a cost related to movement production (cost of movement) and a cost related to our brain's tendency to temporally discount the value of future reward (cost of time). However, whether it is critical to hypothesize a cost of time to explain the vigor of basic reaching movements with intangible reward is unclear because the cost of movement may be theoretically sufficient for this purpose. Here we directly address this issue by designing an isometric reaching task whose completion can be accurate and effortless in prefixed durations. The cost of time hypothesis predicts that participants should be prone to spend energy to save time even if the task can be accomplished at virtually no motor cost. Accordingly, we found that all participants generated substantial amounts of force to invigorate task accomplishment, especially when the prefixed duration was long enough. Remarkably, the time saved by each participant was linked to their original vigor in the task and predicted by an optimal control model balancing out movement and time costs. Taken together, these results support the existence of an idiosyncratic, cognitive cost of time that underlies the invigoration of basic isometric reaching movements.NEW & NOTEWORTHY Movement vigor is generally thought to result from a trade-off between time and motor costs. However, it remains unclear whether the time cost only modulates vigor around some nominal value explained by a minimal motor cost or whether it determines movement invigoration more broadly. Here, we present an original paradigm allowing us to neutralize the cost of movement and provide new evidence that a cost of time must underlie the invigoration of isometric reaching movements.

Encoding of Serial Order in Working Memory: Neuronal Activity in Motor, Premotor, and Prefrontal Cortex during a Memory Scanning Task.

We have adapted Sternberg's context-recall task to investigate the neural mechanisms of encoding serial order information in working memory, in 2 male rhesus monkeys. We recorded from primary motor, premotor, and dorsolateral prefrontal cortex while the monkeys performed the task. In each cortical area, most neurons displayed marked modulation of activity during the list presentation period of the task, whereas the serial order of the stimuli needed to be encoded in working memory. The activity of many neurons changed in a consistent manner over the course of the list presentation period, without regard to the location of the stimuli presented. Remarkably, these neurons encoded serial position information in a relative (rather than absolute) manner across different list lengths. In addition, many neurons showed activity related to both location and serial position, in the form of an interaction effect. Surprisingly, the activity of these neurons was often modulated by the location of stimuli presented before the epoch in which the activity changes occurred. In motor and premotor areas, a large proportion of neurons with list presentation activity also showed direction-related activity during the response phase, whereas in prefrontal cortex most cells showed only list presentation effects. These results show that many neurons had a heterogeneous functionality by representing distinct task variables at different periods of the task. Finally, potential confounds could not account for the effects observed. For these reasons, we conclude that these neurons were indeed participating in sequence encoding in working memory.SIGNIFICANCE STATEMENT Traditionally, primary motor, premotor, and prefrontal areas have been considered to be mainly engaged in motor output, visuomotor transformation, and higher cognitive functions, respectively. Here we show that neurons in all three cortical regions participate in the encoding of a sequence of spatial stimuli in working memory. Furthermore, a central question in cognitive neuroscience has been the manner in which the position of an item within a sequence is encoded in the brain. Our findings provide direct neurophysiological support for a specific hypothesis from cognitive psychology: that of relative coding of serial order.

Neural markers of loss aversion in resting-state brain activity.

Neural responses in striatal, limbic and somatosensory brain regions track individual differences in loss aversion, i.e. the higher sensitivity to potential losses compared with equivalent gains in decision-making under risk. The engagement of structures involved in the processing of aversive stimuli and experiences raises a further question, i.e. whether the tendency to avoid losses rather than acquire gains represents a transient fearful overreaction elicited by choice-related information, or rather a stable component of one's own preference function, reflecting a specific pattern of neural activity. We tested the latter hypothesis by assessing in 57 healthy human subjects whether the relationship between behavioral and neural loss aversion holds at rest, i.e. when the BOLD signal is collected during 5minutes of cross-fixation in the absence of an explicit task. Within the resting-state networks highlighted by a spatial group Independent Component Analysis (gICA), we found a significant correlation between strength of activity and behavioral loss aversion in the left ventral striatum and right posterior insula/supramarginal gyrus, i.e. the very same regions displaying a pattern of neural loss aversion during explicit choices. Cross-study analyses confirmed that this correlation holds when voxels identified by gICA are used as regions of interest in task-related activity and vice versa. These results suggest that the individual degree of (neural) loss aversion represents a stable dimension of decision-making, which reflects in specific metrics of intrinsic brain activity at rest possibly modulating cortical excitability at choice.

Tactile perception during action observation.

It has been suggested that tactile perception becomes less acute during movement to optimize motor control and to prevent an overload of afferent information generated during action. This empirical phenomenon, known as "tactile gating effect," has been associated with mechanisms of sensory feedback prediction. However, less attention has been given to the tactile attenuation effect during the observation of an action. The aim of this study was to investigate whether and how the observation of a goal-directed action influences tactile perception as during overt action. In a first experiment, we recorded vocal reaction times (RTs) of participants to tactile stimulations during the observation of a reach-to-grasp action. The stimulations were delivered on different body parts that could be either congruent or incongruent with the observed effector (the right hand and the right leg, respectively). The tactile stimulation was contrasted with a no body-related stimulation (an auditory beep). We found increased RTs for tactile congruent stimuli compared to both tactile incongruent and auditory stimuli. This effect was reported only during the observation of the reaching phase, whereas RTs were not modulated during the grasping phase. A tactile two-alternative forced-choice (2AFC) discrimination task was then conducted in order to quantify the changes in tactile sensitivity during the observation of the same goal-directed actions. In agreement with the first experiment, the tactile perceived intensity was reduced only during the reaching phase. These results suggest that tactile processing during action observation relies on a process similar to that occurring during action execution.

Integrating force and position: testing model predictions.

In this study, we investigated the integration of force and position information in a task in which participants were asked to estimate the center of a weak force field. Two hypotheses, describing how participants solved this task, were tested: (1) by only using the position(s) where the force reaches the detection threshold, and (2) by extrapolating the force field based on perceived stiffness. Both hypotheses were also described formally, assuming a psychophysical function obeying a power law with an exponent smaller than one. The hypotheses were tested in two psychophysical experiments, in which 12 participants took part. In Experiment 1, an asymmetric force field was used and the presence of visual feedback about hand position was varied. In Experiment 2, a unilateral force field was used. For both experiments, hypothesis 1 predicts biases between (Experiment 1) or at (Experiment 2) the position(s) of the force detection threshold, while hypothesis 2 predicts smaller biases. The measured data show significant biases in both experiments that coincide with the biases predicted by using force detection thresholds from the literature. The average measured responses and their variabilities also fitted very well with the mathematical model of hypothesis 1. These results underline the validity of hypothesis 1. So, participants did not use a percept of the stiffness of the force field, but based their estimation of the center of the force field on the position(s) where the force reached the detection threshold. This shows that force and position information were not integrated in this task.

Motor commands induce time compression for tactile stimuli.

Saccades cause compression of visual space around the saccadic target, and also a compression of time, both phenomena thought to be related to the problem of maintaining saccadic stability (Morrone et al., 2005; Burr and Morrone, 2011). Interestingly, similar phenomena occur at the time of hand movements, when tactile stimuli are systematically mislocalized in the direction of the movement (Dassonville, 1995; Watanabe et al., 2009). In this study, we measured whether hand movements also cause an alteration of the perceived timing of tactile signals. Human participants compared the temporal separation between two pairs of tactile taps while moving their right hand in response to an auditory cue. The first pair of tactile taps was presented at variable times with respect to movement with a fixed onset asynchrony of 150 ms. Two seconds after test presentation, when the hand was stationary, the second pair of taps was delivered with a variable temporal separation. Tactile stimuli could be delivered to either the right moving or left stationary hand. When the tactile stimuli were presented to the motor effector just before and during movement, their perceived temporal separation was reduced. The time compression was effector-specific, as perceived time was veridical for the left stationary hand. The results indicate that time intervals are compressed around the time of hand movements. As for vision, the mislocalizations of time and space for touch stimuli may be consequences of a mechanism attempting to achieve perceptual stability during tactile exploration of objects, suggesting common strategies within different sensorimotor systems.

The functional and structural neural basis of individual differences in loss aversion.

Decision making under risk entails the anticipation of prospective outcomes, typically leading to the greater sensitivity to losses than gains known as loss aversion. Previous studies on the neural bases of choice-outcome anticipation and loss aversion provided inconsistent results, showing either bidirectional mesolimbic responses of activation for gains and deactivation for losses, or a specific amygdala involvement in processing losses. Here we focused on loss aversion with the aim to address interindividual differences in the neural bases of choice-outcome anticipation. Fifty-six healthy human participants accepted or rejected 104 mixed gambles offering equal (50%) chances of gaining or losing different amounts of money while their brain activity was measured with functional magnetic resonance imaging (fMRI). We report both bidirectional and gain/loss-specific responses while evaluating risky gambles, with amygdala and posterior insula specifically tracking the magnitude of potential losses. At the individual level, loss aversion was reflected both in limbic fMRI responses and in gray matter volume in a structural amygdala-thalamus-striatum network, in which the volume of the "output" centromedial amygdala nuclei mediating avoidance behavior was negatively correlated with monetary performance. We conclude that outcome anticipation and ensuing loss aversion involve multiple neural systems, showing functional and structural individual variability directly related to the actual financial outcomes of choices. By supporting the simultaneous involvement of both appetitive and aversive processing in economic decision making, these results contribute to the interpretation of existing inconsistencies on the neural bases of anticipating choice outcomes.

Enhancing Visual Exploration through Augmented Gaze: High Acceptance of Immersive Virtual Biking by Oldest Olds.

The diffusion of virtual reality applications dedicated to aging urges us to appraise its acceptance by target populations, especially the oldest olds. We investigated whether immersive virtual biking, and specifically a visuomotor manipulation aimed at improving visual exploration (augmented gaze), was well accepted by elders living in assisted residences. Twenty participants (mean age 89.8 years, five males) performed three 9 min virtual biking sessions pedalling on a cycle ergometer while wearing a Head-Mounted Display which immersed them inside a 360-degree pre-recorded biking video. In the second and third sessions, the relationship between horizontal head rotation and contingent visual shift was experimentally manipulated (augmented gaze), the visual shift being twice (gain = 2.0) or thrice (gain = 3.0) the amount of head rotation. User experience, motion sickness and visual exploration were measured. We found (i) very high user experience ratings, regardless of the gain; (ii) no effect of gain on motion sickness; and (iii) increased visual exploration (slope = +46%) and decreased head rotation (slope = -18%) with augmented gaze. The improvement in visual exploration capacity, coupled with the lack of intolerance signs, suggests that augmented gaze can be a valuable tool to improve the "visual usability" of certain virtual reality applications for elders, including the oldest olds.

Tapping Force Encodes Metrical Aspects of Rhythm.

Humans possess the ability to extract highly organized perceptual structures from sequences of temporal stimuli. For instance, we can organize specific rhythmical patterns into hierarchical, or metrical, systems. Despite the evidence of a fundamental influence of the motor system in achieving this skill, few studies have attempted to investigate the organization of our motor representation of rhythm. To this aim, we studied-in musicians and non-musicians-the ability to perceive and reproduce different rhythms. In a first experiment participants performed a temporal order-judgment task, for rhythmical sequences presented via auditory or tactile modality. In a second experiment, they were asked to reproduce the same rhythmic sequences, while their tapping force and timing were recorded. We demonstrate that tapping force encodes the metrical aspect of the rhythm, and the strength of the coding correlates with the individual's perceptual accuracy. We suggest that the similarity between perception and tapping-force organization indicates a common representation of rhythm, shared between the perceptual and motor systems.

The effects of a 'pretend play-based training' designed to promote the development of emotion comprehension, emotion regulation, and prosocial behaviour in 5- to 6-year-old Swiss children.

The objective of this study was to evaluate the effects of a pretend play-based training designed to promote the development of socio-emotional competences. 79 children aged 5 to 6 years were evaluated before and after a pretend play-based training. The experimental group (39 children) received this programme on emotion comprehension, negative emotion regulation, and prosocial behaviour one hour a week for eleven weeks during class hours, while the control group (40 children) received no specific intervention. The programme was implemented by 5 teachers. The results show improvements in the ability to understand emotions in children who benefited from the training. These findings are discussed in the broader context of using this form of play as a privileged pedagogical tool to allow children to develop these competences.

Delayed visual feedback affects both manual tracking and grip force control when transporting a handheld object.

When we manipulate an object, grip force is adjusted in anticipation of the mechanical consequences of hand motion (i.e., load force) to prevent the object from slipping. This predictive behavior is assumed to rely on an internal representation of the object dynamic properties, which would be elaborated via visual information before the object is grasped and via somatosensory feedback once the object is grasped. Here we examined this view by investigating the effect of delayed visual feedback during dextrous object manipulation. Adult participants manually tracked a sinusoidal target by oscillating a handheld object whose current position was displayed as a cursor on a screen along with the visual target. A delay was introduced between actual object displacement and cursor motion. This delay was linearly increased (from 0 to 300 ms) and decreased within 2-min trials. As previously reported, delayed visual feedback altered performance in manual tracking. Importantly, although the physical properties of the object remained unchanged, delayed visual feedback altered the timing of grip force relative to load force by about 50 ms. Additional experiments showed that this effect was not due to task complexity nor to manual tracking. A model inspired by the behavior of mass-spring systems suggests that delayed visual feedback may have biased the representation of object dynamics. Overall, our findings support the idea that visual feedback of object motion can influence the predictive control of grip force even when the object is grasped.

Blind saccades: an asynchrony between seeing and looking.

Saccades may not always wait for the completion of the perceptual analysis. By taking advantage of a motion-induced illusion of position and of the spontaneous scatter of saccade latency, we showed that in normal observers, regular saccades (latency, approximately 200 ms) were accurately directed to the target, whereas at higher latencies, saccades were increasingly biased by visual motion until they reflected the perceptual illusion. We reconstructed the time course of saccadic direction coding and identified an early phase in which saccades are mostly predictive (latencies less than approximately 100 ms), followed by a phase in which saccades are guided by the target position signal (latencies approximately 100-250 ms), and a later phase associated with the buildup of mislocalization (approximately 250-450 ms). This transient dissociation between action and perception indicates that seeing and looking are based on asynchronous processes, possibly because of independent thresholds for saccades and perceptual localization. The metrics of a saccade would then reflect the evolution of cortical visual signals from a predictive state to a perceptual state, passing through an intermediate visuomotor state. If saccades occur during the visuomotor state, they escape the tricks of perception.

Audio motor training improves mobility and spatial cognition in visually impaired children.

Since it has been demonstrated that spatial cognition can be affected in visually impaired children, training strategies that exploit the plasticity of the human brain should be early adopted. Here we developed and tested a new training protocol based on the reinforcement of audio-motor associations and thus supporting spatial development in visually impaired children. The study involved forty-four visually impaired children aged 6-17 years old assigned to an experimental (ABBI training) or a control (classical training) rehabilitation conditions. The experimental training group followed an intensive but entertaining rehabilitation for twelve weeks during which they performed ad-hoc developed audio-spatial exercises with the Audio Bracelet for Blind Interaction (ABBI). A battery of spatial tests administered before and after the training indicated that children significantly improved in almost all the spatial aspects considered, while the control group didn't show any improvement. These results confirm that perceptual development in the case of blindness can be enhanced with naturally associated auditory feedbacks to body movements. Therefore the early introduction of a tailored audio-motor training could potentially prevent spatial developmental delays in visually impaired children.

Contact forces evoked by transcranial magnetic stimulation of the motor cortex in a multi-finger grasp.

Transcranial magnetic stimulation (TMS) is a tool of choice to study the functionality of the corticospinal pathway. In this study, we used single-pulse TMS at different intensities and during different levels of grasping force, to stimulate the hand area of the left primary motor cortex (M1). We measured, the TMS-evoked forces, or motor evoked forces (MEFs) in a multi-fingered three-point grasp in addition of the conventional motor evoked potentials (MEPs) from the right forearm and intrinsic hand muscles. This paper aims at presenting the viability of this innovative approach and some preliminary results. The timing (i.e., latencies and peak times), amplitudes and directions of the MEF were analyzed. We found that the TMS evoked synergistic increases of the force magnitudes, akin to those observed when participants voluntarily increased the grip force. The MEF sizes and MEP amplitudes increased with TMS intensity in most cases. The grip force (which measures the overall force involved in the grasp) and the net force (which measures the net effect of all contact forces exerted on the object) seem to be differently affected by single TMS pulses of the motor cortex.

The haptic perception of spatial orientations.

This review examines the isotropy of the perception of spatial orientations in the haptic system. It shows the existence of an oblique effect (i.e., a better perception of vertical and horizontal orientations than oblique orientations) in a spatial plane intrinsic to the haptic system, determined by the gravitational cues and the cognitive resources and defined in a subjective frame of reference. Similar results are observed from infancy to adulthood. In 3D space, the haptic processing of orientations is also anisotropic and seems to use both egocentric and allocentric cues. Taken together, these results revealed that the haptic oblique effect occurs when the sensory motor traces associated with exploratory movement are represented more abstractly at a cognitive level.

Force sharing and other collaborative strategies in a dyadic force perception task.

When several persons perform a physical task jointly, such as transporting an object together, the interaction force that each person experiences is the sum of the forces applied by all other persons on the same object. Therefore, there is a fundamental ambiguity about the origin of the force that each person experiences. This study investigated the ability of a dyad (two persons) to identify the direction of a small force produced by a haptic device and applied to a jointly held object. In this particular task, the dyad might split the force produced by the haptic device (the external force) in an infinite number of ways, depending on how the two partners interacted physically. A major objective of this study was to understand how the two partners coordinated their action to perceive the direction of the third force that was applied to the jointly held object. This study included a condition where each participant responded independently and another one where the two participants had to agree upon a single negotiated response. The results showed a broad range of behaviors. In general, the external force was not split in a way that would maximize the joint performance. In fact, the external force was often split very unequally, leaving one person without information about the external force. However, the performance was better than expected in this case, which led to the discovery of an unanticipated strategy whereby the person who took all the force transmitted this information to the partner by moving the jointly held object. When the dyad could negotiate the response, we found that the participant with less force information tended to switch his or her response more often.

Corrigendum: Shape Perception and Navigation in Blind Adults.

[This corrects the article on p. 10 in vol. 8, PMID: 28144226.].

Low-Cost Robotic Assessment of Visuo-Motor Deficits in Alzheimer's Disease.

A low-cost robotic interface was used to assess the visuo-motor performance of patients with Alzheimer's disease (AD). Twenty AD patients and twenty age-matched controls participated in this work. The battery of tests included simple reaction times, position tracking, and stabilization tasks performed with both hands. The regularity, velocity, visual and haptic feedback were manipulated to vary movement complexity. Reaction times and movement tracking error were analyzed. Results show a marked group effect on a subset of conditions, in particular when the patients could not rely on the visual feedback of hand movement. The visuo-motor performance correlated with the measures of global cognitive functioning and with different memory-related abilities. Our results support the hypothesis that the ability to recall and use visuo-spatial associations might underlie the impairment in complex motor behavior that has been reported in AD patients. Importantly, the patients had preserved learning effects across sessions, which might relate to visuo-motor deficits being less evident in every-day life and clinical assessments. This robotic assessment, lasting less than 1 h, provides detailed information about the integrity of visuo-motor abilities. The data can aid the understanding of the complex pattern of deficits that characterizes this pervasive disease.

Shape Perception and Navigation in Blind Adults.

Different sensory systems interact to generate a representation of space and to navigate. Vision plays a critical role in the representation of space development. During navigation, vision is integrated with auditory and mobility cues. In blind individuals, visual experience is not available and navigation therefore lacks this important sensory signal. In blind individuals, compensatory mechanisms can be adopted to improve spatial and navigation skills. On the other hand, the limitations of these compensatory mechanisms are not completely clear. Both enhanced and impaired reliance on auditory cues in blind individuals have been reported. Here, we develop a new paradigm to test both auditory perception and navigation skills in blind and sighted individuals and to investigate the effect that visual experience has on the ability to reproduce simple and complex paths. During the navigation task, early blind, late blind and sighted individuals were required first to listen to an audio shape and then to recognize and reproduce it by walking. After each audio shape was presented, a static sound was played and the participants were asked to reach it. Movements were recorded with a motion tracking system. Our results show three main impairments specific to early blind individuals. The first is the tendency to compress the shapes reproduced during navigation. The second is the difficulty to recognize complex audio stimuli, and finally, the third is the difficulty in reproducing the desired shape: early blind participants occasionally reported perceiving a square but they actually reproduced a circle during the navigation task. We discuss these results in terms of compromised spatial reference frames due to lack of visual input during the early period of development.