Associative learning and facial expression recognition in schizophrenic patients: Effects of social presence.

Schizophrenia (SCZ) is a psychiatric disorder that alters both general and social cognition. However, the exact mechanisms that are altered remain to be elucidated. In this study, we investigated associative learning (AL) and facial expression recognition (FER) in the same patients, using emotional expressions and abstract images. Our main aim was to investigate how these cognitive abilities are affected by SCZ and to assess the role of mere social presence, a factor that has not been considered before. The study compared the behavioral performance of 60 treated outpatients with SCZ and 103 demographically matched healthy volunteers. In the AL task, participants had to associate abstract images or facial expressions with key presses, guided by feedback on each trial. In the FER task, they had to report whether two successively presented facial expressions were the same or different. All participants performed the two tasks under two social context conditions: alone or in the presence of a passive but attentive audience. The results showed a severe learning impairment in patients compared to controls, with a slight advantage for facial expressions compared to abstract images, and a gender-dependent effect of social presence. In contrast, facial expression recognition was partially spared in patients and facilitated by social presence. We conclude that cognitive abilities are impaired in patients with SCZ, but their investigation needs to take into account the social context in which they are assessed.

Social facilitation and bilingual cognitive advantage: Bridging social psychology and psycholinguistics.

This study examined the role of social context in the expression of the bilingual cognitive advantage in 145 bilingual university students. All participants mastered Arabic as their native language (L1), but half were highly proficient in French (high L2 group), whereas half were less proficient (low L2 group). A color-word Stroop test with incongruent, congruent and neutral stimuli was administered in single language blocks (Arabic or French words) or in a mixed block (Arabic and French words), either under social presence, or alone. Stroop interference was analyzed to assess the cost of resolving conflict in incongruent trials and was compared across groups and experimental conditions. If bilingualism comes with a cognitive advantage, a reduction of interference in high (vs. low) L2 proficient subjects is to be expected. Analysis revealed that interference was significantly reduced in high L2 group, but only under the single language condition. Furthermore, whereas social context and sex had no main effects, analysis revealed a significant 4-factor interaction between L2 proficiency, linguistic context, social context, and sex. Social presence further reduced interference (social facilitation) in high L2 proficient females, but not in males. Overall, the results suggest that mastering a second language comes with cognitive advantages which adapt dynamically to social and linguistic contexts in a sex-dependent manner. We argue that advancing bilingualism research requires more attention to the social environment.

Hand modulation of visual, preparatory, and saccadic activity in the monkey frontal eye field.

Behavioral studies have shown that hand position influences saccade characteristics. This study examined the neuronal changes that could underlie this behavioral observation. Single neurons were recorded in the frontal eye field (FEF) of 2 monkeys as they executed a visually guided saccade task, while holding their hand at given locations on a touch screen. The task was performed with the hand either visible or invisible, in order to assess the relative contribution of visual and proprioceptive information on hand position. Among the 224 neurons tested, the visual, saccadic and/or preparatory activity of more than half of them was modulated by hand position, whether the hand was visible or invisible. Comparison of lower (hand's workspace) and upper (out of reach) visual targets showed that hand modulation was predominant in the hand's workspace. Finally, some cells preferred congruency of hand and target in space, others preferred incongruency. Interestingly, hand modulation of saccadic activity correlated with hand position effects on saccade reaction times. We conclude that visual and proprioceptive signals derived from the hand are integrated by FEF neurons. These signals can modulate target selection through attention and allow the oculomotor system to use hand-related somatosensory signals for the initiation of visually guided saccades.

Insight in schizophrenia: from conceptualization to neuroscience.

Lack of insight into illness is a prevalent and distinguishing feature of schizophrenia, which has a complex history and has been given a variety of definitions. Currently, insight is measured and treated as a multidimensional phenomenon, because it is believed to result from psychological, neuropsychological and organic factors. Thus, schizophrenia patients may display dramatic disorders including demoralization, depression and a higher risk of suicide, all of which are directly or indirectly related to a lack of insight into their illness, and make the treatment difficult. To improve the treatment of people with schizophrenia, it is thus crucial to advance research on insight into their illness. Insight is studied in a variety of ways. Studies may focus on the relationship between insight and psychopathology, may view behavioral outcomes or look discretely at the cognitive dysfunction versus anatomy level of insight. All have merit but they are dispersed across a wide body of literature and rarely are the findings integrated and synthesized in a meaningful way. The aim of this study was to synthesize findings across the large body of literature dealing with insight, to highlight its multidimensional nature, measurement, neuropsychology and social impact in schizophrenia. The extensive literature on the cognitive consequences of lack of insight and the contribution of neuroimaging techniques to elucidating neurological etiology of insight deficits, is also reviewed.

Differential roles of caudate nucleus and putamen during instrumental learning.

The dorsal striatum is crucial for the acquisition and consolidation of instrumental behaviour, but the underlying computations and internal dynamics remain elusive. To address this issue, we combined a model of key computations supporting decision-making during instrumental learning with human behavioural and functional magnetic resonance imaging (fMRI) data. The results showed that the associative and sensorimotor dorsal striatum host complementary computations that, we suggest, may differentially support goal-directed and habitual processes. The anterior caudate nucleus integrates information about performance and cognitive control demands, whereas the putamen tracks how likely the conditioning stimuli lead to correct response. Contrary to current models, the putamen is recruited during initial acquisition. As the exploratory phase proceeds, the relative contribution of the caudate nucleus becomes dominant over the putamen. During early consolidation, caudate nucleus and putamen settle to asymptotic values and share control. We then investigated how dorsal striatal computations may affect decision-making. We found that portion of reaction times' variance parallels the combined cost associated with the dorsal striatal computations. Overall, our findings provide a deeper insight into the functional heterogeneity within the dorsal striatum and suggest that the dynamic interplay between caudate nucleus and putamen, rather than their serial recruitment, underlies the acquisition and early consolidation of instrumental behaviours.

Understanding the neural computations of arbitrary visuomotor learning through fMRI and associative learning theory.

Associative theory postulates that learning the consequences of our actions in a given context is represented in the brain as stimulus-response-outcome associations that evolve according to prediction-error signals (the discrepancy between the observed and predicted outcome). We tested the theory on brain functional magnetic resonance imaging data acquired from human participants learning arbitrary visuomotor associations. We developed a novel task that systematically manipulated learning and induced highly reproducible performances. This granted the validation of the model-based results and an in-depth analysis of the brain signals in representative single trials. Consistent with the Rescorla-Wagner model, prediction-error signals are computed in the human brain and selectively engage the ventral striatum. In addition, we found evidence of computations not formally predicted by the Rescorla-Wagner model. The dorsal fronto-parietal network, the dorsal striatum, and the ventrolateral prefrontal cortex are activated both on the incorrect and first correct trials and may reflect the processing of relevant visuomotor mappings during the early phases of learning. The left dorsolateral prefrontal cortex is selectively activated on the first correct outcome. The results provide quantitative evidence of the neural computations mediating arbitrary visuomotor learning and suggest new directions for future computational models.

Emotion processing in Parkinson's disease: a blood oxygenation level-dependent functional magnetic resonance imaging study.

Parkinson's disease is a neurodegenerative disorder caused by loss of dopamine neurons in the substantia nigra pars compacta. Tremor, rigidity, and bradykinesia are the major symptoms of the disease. These motor impairments are often accompanied by affective and emotional dysfunctions which have been largely studied over the last decade. The aim of this study was to investigate emotional processing organization in the brain of patients with Parkinson's disease and to explore whether there are differences between recognition of different types of emotions in Parkinson's disease. We examined 18 patients with Parkinson's disease (8 men, 10 women) with no history of neurological or psychiatric comorbidities. All these patients underwent identical brain blood oxygenation level-dependent functional magnetic resonance imaging for emotion evaluation. Blood oxygenation level-dependent functional magnetic resonance imaging results revealed that the occipito-temporal cortices, insula, orbitofrontal cortex, basal ganglia, and parietal cortex which are involved in emotion processing, were activated during the functional control. Additionally, positive emotions activate larger volumes of the same anatomical entities than neutral and negative emotions. Results also revealed that Parkinson's disease associated with emotional disorders are increasingly recognized as disabling as classic motor symptoms. These findings help clinical physicians to recognize the emotional dysfunction of patients with Parkinson's disease.

I learned from what you did: Retrieving visuomotor associations learned by observation.

Observational learning allows individuals to acquire knowledge without incurring in the costs and risks of discovering and testing. The neural mechanisms mediating the retrieval of rules learned by observation are currently unknown. To explore this fundamental cognitive ability, we compared the brain responses when retrieving visuomotor associations learned either by observation or by individual learning. To do so, we asked eleven adults to learn two sets of arbitrary visuomotor associations: one set was learned through the observation of an expert actor while the other was learned by trial and error. During fMRI scanning, subjects were requested to retrieve the visuomotor associations previously learned under the two modalities. The conjunction analysis between the two learning conditions revealed a common brain network that included the ventral and dorsal lateral prefrontal cortices, the superior parietal lobe and the pre-SMA. This suggests the existence of a mirror-like system responsible for the storage of rules learned either by trial and error or by observation of others' actions. In addition, the pars triangularis in the right prefrontal cortex (BA45) was found to be selective for rules learned by observation. This suggests a preferential role of this area in the storage of rules learned in a social context.

Hand position modulates saccadic activity in the frontal eye field.

Recent neurophysiological studies have begun to uncover the neuronal correlates of eye hand coordination. This study was designed to test whether the frontal eye field (FEF) saccadic activity is modulated by hand position. Single neurons were recorded in two macaque monkeys performing visually guided saccades while holding their hand at given locations on a touchscreen. To determine the relative contributions of hand vision and its proprioception, monkeys executed the task with or without vision of the hand. We found that saccadic activity of more than half of the neuronal sample (54%; n=130) was dependent on hand position relative to the saccade end point. Both visual and proprioceptive signals contributed to this modulation. These data demonstrate that the oculomotor function of the FEF takes into account hand position in space.

Visuo-motor learning with combination of different rates of motor imagery and physical practice.

Sports psychology suggests that mental rehearsal facilitates physical practice in athletes and clinical rehabilitation attempts to use mental rehearsal to restore motor function in hemiplegic patients. Our aim was to examine whether mental rehearsal is equivalent to physical learning, and to determine the optimal proportions of real execution and rehearsal. Subjects were asked to grasp an object and insert it into an adapted slot. One group (G0) practiced the task only by physical execution (240 trials); three groups imagined performing the task in different rates of trials (25%, G25; 50%, G50; 75%, G75), and physically executed movements for the remaining trials; a fourth, control group imagined a visual rotation task in 75% of the trials and then performed the same motor task as the others groups. Movement time (MT) was compared for the first and last physical trials, together with other key trials, across groups. All groups learned, suggesting that mental rehearsal is equivalent to physical motor learning. More importantly, when subjects rehearsed the task for large numbers of trials (G50 and G75), the MT of the first executed trial was significantly shorter than the first executed trial in the physical group (G0), indicating that mental practice is better than no practice at all. Comparison of the first executed trial in G25, G50 and G75 with the corresponding trials in G0 (61, 121 and 181 trials), showed equivalence between mental and physical practice. At the end of training, the performance was much better with high rates of mental practice (G50/G75) compared to physical practice alone (G0), especially when the task was difficult. These findings confirm that mental rehearsal can be beneficial for motor learning and suggest that imagery might be used to supplement or partly replace physical practice in clinical rehabilitation.

Estimating the hidden learning representations.

Successful adaptation relies on the ability to learn the consequence of our actions in different environments. However, understanding the neural bases of this ability still represents one of the great challenges of system neuroscience. In fact, the neuronal plasticity changes occurring during learning cannot be fully controlled experimentally and their evolution is hidden. Our approach is to provide hypotheses about the structure and dynamics of the hidden plasticity changes using behavioral learning theory. In fact, behavioral models of animal learning provide testable predictions about the hidden learning representations by formalizing their relation with the observables of the experiment (stimuli, actions and outcomes). Thus, we can understand whether and how the predicted learning processes are represented at the neural level by estimating their evolution and correlating them with neural data. Here, we present a bayesian model approach to estimate the evolution of the internal learning representations from the observations of the experiment (state estimation), and to identify the set of models' parameters (parameter estimation) and the class of behavioral model (model selection) that are most likely to have generated a given sequence of actions and outcomes. More precisely, we use Sequential Monte Carlo methods for state estimation and the maximum likelihood principle (MLP) for model selection and parameter estimation. We show that the method recovers simulated trajectories of learning sessions on a single-trial basis and provides predictions about the activity of different categories of neurons that should participate in the learning process. By correlating the estimated evolutions of the learning variables, we will be able to test the validity of different models of instrumental learning and possibly identify the neural bases of learning.

Social and asocial prefrontal cortex neurons: a new look at social facilitation and the social brain.

A fundamental aspect of behavior in many animal species is 'social facilitation', the positive effect of the mere presence of conspecifics on performance. To date, the neuronal counterpart of this ubiquitous phenomenon is unknown. We recorded the activity of single neurons from two prefrontal cortex regions, the dorsolateral part and the anterior cingulate cortex in monkeys as they performed a visuomotor task, either in the presence of a conspecific (Presence condition) or alone. Monkeys performed better in the presence condition than alone (social facilitation), and analyses of outcome-related activity of 342 prefrontal neurons revealed that most of them (86%) were sensitive to the performance context. Two populations of neurons were discovered: 'social neurons', preferentially active under social presence and 'asocial neurons', preferentially active under social isolation. The activity of these neurons correlated positively with performance only in their preferred context (social neurons under social presence; asocial neurons under social isolation), thereby providing a potential neuronal mechanism of social facilitation. More generally, the fact that identical tasks recruited either social or asocial neurons depending on the presence or absence of a conspecific also brings a new look at the social brain hypothesis.

Role of Anterior Cingulate Cortex in Instrumental Learning: Blockade of Dopamine D1 Receptors Suppresses Overt but Not Covert Learning.

HIGHLIGHTS Blockade of dopamine D1 receptors in ACC suppressed instrumental learning when overt responding was required.Covert learning through observation was not impaired.After treatment with a dopamine antagonist, instrumental learning recovered but not the rat's pretreatment level of effort tolerance.ACC dopamine is not necessary for acquisition of task-relevant cues during learning, but regulates energy expenditure and effort based decision. Dopamine activity in anterior cingulate cortex (ACC) is essential for various aspects of instrumental behavior, including learning and effort based decision making. To dissociate learning from physical effort, we studied both observational (covert) learning, and trial-and-error (overt) learning. If ACC dopamine activity is required for task acquisition, its blockade should impair both overt and covert learning. If dopamine is not required for task acquisition, but solely for regulating the willingness to expend effort for reward, i.e., effort tolerance, blockade should impair overt learning but spare covert learning. Rats learned to push a lever for food rewards either with or without prior observation of an expert conspecific performing the same task. Before daily testing sessions, the rats received bilateral ACC microinfusions of SCH23390, a dopamine D1 receptor antagonist, or saline-control infusions. We found that dopamine blockade suppressed overt responding selectively, leaving covert task acquisition through observational learning intact. In subsequent testing sessions without dopamine blockade, rats recovered their overt-learning capacity but not their pre-treatment level of effort tolerance. These results suggest that ACC dopamine is not required for the acquisition of conditioned behaviors and that apparent learning impairments could instead reflect a reduced level of willingness to expend effort due to cortical dopamine blockade.

Prehension movements in the macaque monkey: effects of perturbation of object size and location.

While the neural bases of prehension have been extensively studied in monkeys, a few kinematic studies have examined their prehension behavior. Recently (Roy et al. 2000, 2002), we have described the kinematics of reaching and grasping in freely behaving monkeys under normal conditions by applying the high-resolution recording techniques (Optotrak system) and behavioral paradigms used in humans. Here we determined whether online movement reorganization observed in monkeys following sudden changes of either object size or location at movement onset is similar to that observed in humans. We found that changing object size led to rapid on-flight re-calibration of the different movement parameters, eventually preserving the unitary aspect of the movement with a minor time cost. By contrast, a shift in object location triggered a massive time-consuming reorganization. Re-directed movements appeared as a concatenation of two sub-movements: a first one directed to the initial object and a second one directed to the new object location. These findings first complement our earlier studies in providing further evidence of the similarities between monkey and human prehension. Second, they suggest that the two components of prehension, reaching and grasping, interact through coordination mechanisms that are more efficient to correct for size than for location perturbation. This difference may reflect a hierarchical organization in which reaching would be the subordinate of grasping in both primate species.

Callosal connections of dorsal versus ventral premotor areas in the macaque monkey: a multiple retrograde tracing study.

BACKGROUND: The lateral premotor cortex plays a crucial role in visually guided limb movements. It is divided into two main regions, the dorsal (PMd) and ventral (PMv) areas, which are in turn subdivided into functionally and anatomically distinct rostral (PMd-r and PMv-r) and caudal (PMd-c and PMv-c) sub-regions. We analyzed the callosal inputs to these premotor subdivisions following 23 injections of retrograde tracers in eight macaque monkeys. In each monkey, 2-4 distinct tracers were injected in different areas allowing direct comparisons of callosal connectivity in the same brain. RESULTS: Based on large injections covering the entire extent of the corresponding PM area, we found that each area is strongly connected with its counterpart in the opposite hemisphere. Callosal connectivity with the other premotor areas, the primary motor cortex, prefrontal cortex and somatosensory cortex varied from one area to another. The most extensive callosal inputs terminate in PMd-r and PMd-c, with PMd-r strongly connected with prefrontal cortex. Callosal inputs to PMv-c are more extensive than those to PMv-r, whose connections are restricted to its counterpart area. Quantitative analysis of labelled cells confirms these general findings, and allows an assessment of the relative strength of callosal inputs. CONCLUSION: PMd-r and PMv-r receive their strongest callosal inputs from their respective counterpart areas, whereas PMd-c and PMv-c receive strong inputs from heterotopic areas as well (namely from PMd-r and PMv-r, respectively). Finally, PMd-r stands out as the lateral premotor area with the strongest inputs from the prefrontal cortex, and only the PMd-c and PMv-c receive weak callosal inputs from M1.

Learning by observation in the macaque monkey under high experimental constraints.

While neuroscience research has tremendously advanced our knowledge about the neural mechanisms of individual learning, i.e. through trial-and-error, it is only recently that neuroscientists have begun to study observational learning, and thus little is known about its neural mechanisms. One limitation is that observational learning has been addressed under unconstrained experimental conditions, not compatible with neuronal recordings. This study examined observational learning in macaque monkeys under the constraining conditions of behavioral neurophysiology. Two animals sat in primate chairs facing each other, with their head fixed. A touch screen was placed face up between the chairs at arm's reach, and the monkeys were trained on an abstract visuomotor associative task. In one experiment, the monkeys alternated the roles of "actor" and "observer". The actor learned to associate visual cues with reaching targets, while the observer "watched" freely. Then, the observer was given the same cue-target associations just performed by the actor, or had to learn new, not previously observed ones. The results show that learning performance is better after observation. In experiment 2, one monkey learned from a human actor who performed the task with errors only, or with successes only in separate blocks. The monkey's gain in performance was higher after observation of errors than after successes. The findings suggest that observational learning can occur even under highly constraining conditions, and open the way for investigating the neuronal correlates of social learning using the methods of behavioral neurophysiology.

Projections of the claustrum to the primary motor, premotor, and prefrontal cortices in the macaque monkey.

The claustrum is interconnected with the frontal lobe, including the motor cortex, prefrontal cortex, and cingulate cortex. The goal of the present study was to assess whether the claustral projections to distinct areas within the frontal cortex arise from separate regions within the claustrum. Multiple injections of tracers were performed in 15 macaque monkeys, aimed toward primary motor area (M1), pre-supplementary motor area (pre-SMA), SMA-proper, rostral (PMd-r) and caudal (PMd-c) parts of the dorsal premotor cortex (PM), rostral (PMv-r) and caudal (PMv-c) parts of the ventral PM, and superior and inferior parts of area 46. The distribution of retrogradely labeled neurons showed no clear segregation along the rostrocaudal axis of the claustrum; they were usually located along the entire anteroposterior extent of the claustrum. For all motor cortical areas, there was a general trend of the labeled neurons to occupy the dorsal and intermediate parts of the claustrum along the dorsoventral axis. The same territories were labeled after injection in area 46, but in addition numerous labeled neurons were found in the most ventral part of the claustrum. At higher resolution, however, there was clear evidence that the territories projecting to pre-SMA and SMA-proper formed separate, interdigitating, clusters along the dorsoventral axis. A comparable local segregation was observed for the two subdivisions of area 46, whereas there was more local overlap among the subareas of PM. The projections from the claustrum to the multiple subareas of the motor cortex and to area 46 arise from largely overlapping territories, with, however, some degree of local segregation.

Functional connectivity during real vs imagined visuomotor tasks: an EEG study.

It is proposed that real and imagined movements activate identical neural networks. Cortical oscillatory activity is proposed as a mechanism through which distributed neuronal networks may bind into coherent ensembles and coupling of oscillators is used as a tool to investigate modulations of cortical connectivity. The aim of the present study was to test the hypothesis that, although the same brain network is involved in both real and imagined movements, the functional connectivity within the network differs. To do so, we measured interregional coupling, quantified using coherence between scalp EEG electrodes, during different periods of a prehension task during real and imagined movements. The results demonstrated a different pattern of coupling in the beta frequency range between electrodes overlying occipital and motor cortices during executed and imagined movements. These findings are consistent with the hypothesis that the neural networks during real and imagined movements are not identical.

Conditional visuo-motor learning in primates: a key role for the basal ganglia.

Sensory guidance of behavior often involves standard visuo-motor mapping of body movements onto objects and spatial locations. For example, looking at and reaching to grasp a glass of wine requires the mapping of the eyes and hand to the location of the glass in space, as well as the formation of a hand configuration appropriate to the shape of the glass. But our brain is far more than just a standard sensorimotor mapping machine. Through evolution, the brain of advanced mammals, in particular human and non-human primates, has acquired a formidable capacity to construct non-standard, arbitrary mapping using associations between external events and behavioral responses that bear no direct relationship. For example, we have all learned to stop at a red traffic light and to go at a green one, or to wait for a specific tone before dialing a phone number and to hang up when hearing a busy signal. These arbitrary associations are acquired through experience, thereby providing primates with a rich and flexible sensorimotor repertoire. Understanding how they are learned, and how they are recalled and used when the context requires them, has been one of the challenging issues for cognitive neuroscience. Valuable insights have been gained over the last two decades through the convergence of multiple complementary approaches. Human neuropsychology and experimental lesions in monkeys have identified a network of brain structures important for conditional sensorimotor associations, whereas imaging studies in healthy human subjects and electrophysiological recordings in awake monkeys have sought to identify the different functional processes underlying the overall function. The present review focuses on the contribution of a network linking the prefrontal cortex, basal ganglia, and dorsal premotor cortex, with special emphasis on results from recording experiments in monkeys. We will first review data pointing to a specific contribution of each component of the network to the performance of well-learned arbitrary visuo-motor associations, as well as data suggesting how novel associations are formed. Then we will propose a model positing that each component of the fronto-striatal network makes a specific contribution to the formation and/or execution of sensorimotor associations. In this model, the basal ganglia are thought to play a key role in linking the sensory, motor, and reward information necessary for arbitrary mapping.

[The planning of action: can one separate attention from intention?]. / Le codage de l'action: peut-on dissocier l'attention de l'intention?

Attention and motor preparation are two intimately linked processes. However, they can be dissociated in the laboratory in order to study their neuronal basis. Behavioral neurophysiology has thus shown that neurons that discharge in relation with attention or with motor preparation (or intention) exist in a variety of brain regions in the monkey, especially the prefrontal and premotor cortices. When examined more carefully, these two regions appear different in both the proportion of cells that respond during attention versus intention, and in the information coded in the so-called "preparatory activity". This activity reflects sensory selection in the prefrontal cortex (spatial attention/memory), motor selection in the premotor cortex. Furthermore, two regions in the dorsal aspect of premotor cortex can be distinguished on the basis of their relative involvement in attention: a rostral (anterior) region, functionally close to prefrontal cortex, and a caudal one, which appears functionally close to motor cortex. Using an experimental design derived from monkey experiments, a functional magnetic resonance imaging (fMRI) study recently indicated that the functional specialization within the premotor cortex is similar in monkey and man.