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1.  Counterproductive Effect of Saccadic Suppression during Attention Shifts 
PLoS ONE  2014;9(1):e86633.
During saccadic eye movements, the processing of visual information is transiently interrupted by a mechanism known as “saccadic suppression” [1] that is thought to ensure perceptual stability [2]. If, as proposed in the premotor theory of attention [3], covert shifts of attention rely on sub-threshold recruitment of oculomotor circuits, then saccadic suppression should also occur during covert shifts. In order to test this prediction, we designed two experiments in which participants had to orient towards a cued letter, with or without saccades. We analyzed the time course of letter identification score in an “attention” task performed without saccades, using the saccadic latencies measured in the “saccade” task as a marker of covert saccadic preparation. Visual conditions were identical in all tasks. In the “attention” task, we found a drop in perceptual performance around the predicted onset time of saccades that were never performed. Importantly, this decrease in letter identification score cannot be explained by any known mechanism aligned on cue onset such as inhibition of return, masking, or microsaccades. These results show that attentional allocation triggers the same suppression mechanisms as during saccades, which is relevant during eye movements but detrimental in the context of covert orienting.
PMCID: PMC3900577  PMID: 24466181
2.  Pupil size variations correlate with physical effort perception 
It has long been established that the pupil diameter increases during mental activities in proportion to the difficulty of the task at hand. However, it is still unclear whether this relationship between the pupil size and effort applies also to physical effort. In order to address this issue, we asked healthy volunteers to perform a power grip task, at varied intensity, while evaluating their effort both implicitly and explicitly, and while concurrently monitoring their pupil size. Each trial started with a contraction of imposed intensity, under the control of a continuous visual feedback. Upon completion of the contraction, participants had to choose whether to replicate, without feedback, the first contraction for a variable monetary reward, or whether to skip this step and go directly to the next trial. The rate of acceptance of effort replication and the amount of force exerted during the replication were used as implicit measures of the perception of the effort exerted during the first contraction. In addition, the participants were asked to rate on an analog scale, their explicit perception of the effort for each intensity condition. We found that pupil diameter increased during physical effort and that the magnitude of this response reflected not only the actual intensity of the contraction but also the subjects' perception of the effort. This finding indicates that the pupil size signals the level of effort invested in a task, irrespective of whether it is physical or mental. It also helps refining the potential brain circuits involved since the results of the current study imply a convergence of mental and physical effort information at some level along this pathway.
PMCID: PMC4142600  PMID: 25202247
pupillometry; effort; psychophysiology; effort-based decision making; pupil dilation; physical exertion
3.  The Role of Left Supplementary Motor Area in Grip Force Scaling 
PLoS ONE  2013;8(12):e83812.
Skilled tool use and object manipulation critically relies on the ability to scale anticipatorily the grip force (GF) in relation to object dynamics. This predictive behaviour entails that the nervous system is able to store, and then select, the appropriate internal representation of common object dynamics, allowing GF to be applied in parallel with the arm motor commands. Although psychophysical studies have provided strong evidence supporting the existence of internal representations of object dynamics, known as “internal models”, their neural correlates are still debated. Because functional neuroimaging studies have repeatedly designated the supplementary motor area (SMA) as a possible candidate involved in internal model implementation, we used repetitive transcranial magnetic stimulation (rTMS) to interfere with the normal functioning of left or right SMA in healthy participants performing a grip-lift task with either hand. TMS applied over the left, but not right, SMA yielded an increase in both GF and GF rate, irrespective of the hand used to perform the task, and only when TMS was delivered 130–180 ms before the fingers contacted the object. We also found that both left and right SMA rTMS led to a decrease in preload phase durations for contralateral hand movements. The present study suggests that left SMA is a crucial node in the network processing the internal representation of object dynamics although further experiments are required to rule out that TMS does not affect the GF gain. The present finding also further substantiates the left hemisphere dominance in scaling GF.
PMCID: PMC3877107  PMID: 24391832
4.  Theta Burst Stimulation Applied over Primary Motor and Somatosensory Cortices Produces Analgesia Unrelated to the Changes in Nociceptive Event-Related Potentials 
PLoS ONE  2013;8(8):e73263.
Continuous theta burst stimulation (cTBS) applied over the primary motor cortex (M1) can alleviate pain although the neural basis of this effect remains largely unknown. Besides, the primary somatosensory cortex (S1) is thought to play a pivotal role in the sensori-discriminative aspects of pain perception but the analgesic effect of cTBS applied over S1 remains controversial. To investigate cTBS-induced analgesia we characterized, in two separate experiments, the effect of cTBS applied either over M1 or S1 on the event-related brain potentials (ERPs) and perception elicited by nociceptive (CO2 laser stimulation) and non-nociceptive (transcutaneous electrical stimulation) somatosensory stimuli. All stimuli were delivered to the ipsilateral and contralateral hand. We found that both cTBS applied over M1 and cTBS applied over S1 significantly reduced the percept elicited by nociceptive stimuli delivered to the contralateral hand as compared to similar stimulation of the ipsilateral hand. In contrast, cTBS did not modulate the perception of non-nociceptive stimuli. Surprisingly, this side-dependent analgesic effect of cTBS was not reflected in the amplitude modulation of nociceptive ERPs. Indeed, both nociceptive (N160, N240 and P360 waves) and late-latency non-nociceptive (N140 and P200 waves) ERPs elicited by stimulation of the contralateral and ipsilateral hands were similarly reduced after cTBS, suggesting an unspecific effect, possibly due to habituation or reduced alertness. In conclusion, cTBS applied over M1 and S1 reduces similarly the perception of nociceptive inputs originating from the contralateral hand, but this analgesic effect is not reflected in the magnitude of nociceptive ERPs.
PMCID: PMC3748010  PMID: 23977382
5.  Deficit in Complex Sequence Processing after a Virtual Lesion of Left BA45 
PLoS ONE  2013;8(6):e63722.
Although the contribution of Broca's area to motor cognition is generally accepted, its exact role remains controversial. A previous functional imaging study has suggested that Broca's area implements hierarchically organised motor behaviours and, in particular, that its anterior (Brodmann area 45, BA45) and posterior (BA44) parts process, respectively, higher and lower-level hierarchical elements. This function of Broca's area could generalize to other cognitive functions, including language. However, because of the correlative nature of functional imaging data, the causal relationship between Broca's region activation and its behavioural significance cannot be ascertained. To circumvent this limitation, we used on-line repetitive transcranial magnetic stimulation to disrupt neuronal processing in left BA45, left BA44 or left dorsal premotor cortex, three areas that have been shown to exhibit a phasic activation when participants performed hierarchically organised motor behaviours. The experiment was conducted in healthy volunteers performing the same two key-press sequences as those used in a previous imaging study, and which differed in terms of hierarchical organisation. The performance of the lower-order hierarchical task (Experiment #1) was unaffected by magnetic stimulation. In contrast, in the higher-order hierarchical task (Experiment #2, “superordinate” task), we found that a virtual lesion of the anterior part of Broca's area (left BA45) delayed the processing of the cue initiating the sequence in an effector-independent way. Interestingly, in this task, the initiation cue only informed the subjects about the rules to be applied to produce the appropriate response but did not allow them to anticipate the entire motor sequence. A second important finding was a RT decrease following left PMd virtual lesions in the superordinate task, a result compatible with the view that PMd plays a critical role in impulse control. The present study therefore demonstrates the role of left BA45 in planning the higher-order hierarchical levels of motor sequences.
PMCID: PMC3677864  PMID: 23762232
6.  Dissociating the role of prefrontal and premotor cortices in controlling inhibitory mechanisms during motor preparation 
The Journal of Neuroscience  2012;32(3):806-816.
Top-down control processes are critical to select goal-directed actions in flexible environments. In humans, these processes include two inhibitory mechanisms that operate during response selection: one is involved in solving a competition between different response options, the other ensures that a selected response is initiated timely. Here, we evaluated the role of dorsal premotor cortex (PMd) and lateral prefrontal cortex (LPF) of healthy subjects in these two forms of inhibition by using an innovative transcranial magnetic stimulation (TMS) protocol combining repetitive TMS (rTMS) over PMd or LPF and a single pulse TMS (sTMS) over primary motor cortex (M1). sTMS over M1 allowed us to assess inhibitory changes in corticospinal excitability, while rTMS was used to produce transient disruption of PMd or LPF. We found that rTMS over LPF reduces inhibition associated with competition resolution whereas rTMS over PMd decreases inhibition associated with response impulse control. These results emphasize the dissociable contributions of these two frontal regions to inhibitory control during motor preparation. The association of LPF with competition resolution is consistent with the role of this area in relatively abstract aspects of control related to goal maintenance, ensuring that the appropriate response is selected in a variable context. In contrast, the association of PMd with impulse control is consistent with the role of this area in more specific processes related to motor preparation and initiation.
PMCID: PMC3304578  PMID: 22262879
inhibition; action selection; competition; impulse control; transcranial magnetic stimulation; motor preparation
7.  Differential effects of parietal and frontal inactivations on reaction times distributions in a visual search task 
The posterior parietal cortex participates to numerous cognitive functions, from perceptual to attentional and decisional processes. However, the same functions have also been attributed to the frontal cortex. We previously conducted a series of reversible inactivations of the lateral intraparietal area (LIP) and of the frontal eye field (FEF) in the monkey which showed impairments in covert visual search performance, characterized mainly by an increase in the mean reaction time (RT) necessary to detect a contralesional target. Only subtle differences were observed between the inactivation effects in both areas. In particular, the magnitude of the deficit was dependant of search task difficulty for LIP, but not for FEF. In the present study, we re-examine these data in order to try to dissociate the specific involvement of these two regions, by considering the entire RT distribution instead of mean RT. We use the LATER model to help us interpret the effects of the inactivations with regard to information accumulation rate and decision processes. We show that: (1) different search strategies can be used by monkeys to perform visual search, either by processing the visual scene in parallel, or by combining parallel and serial processes; (2) LIP and FEF inactivations have very different effects on the RT distributions in the two monkeys. Although our results are not conclusive with regards to the exact functional mechanisms affected by the inactivations, the effects we observe on RT distributions could be accounted by an involvement of LIP in saliency representation or decision-making, and an involvement of FEF in attentional shifts and perception. Finally, we observe that the use of the LATER model is limited in the context of a visual search as it cannot fit all the behavioral strategies encountered. We propose that the diversity in search strategies observed in our monkeys also exists in individual human subjects and should be considered in future experiments.
PMCID: PMC3386550  PMID: 22754512
LIP; FEF; visual search; inactivation; manual reaction times; LATER model; distribution
8.  Role of the Primary Motor Cortex in the Early Boost in Performance Following Mental Imagery Training 
PLoS ONE  2011;6(10):e26717.
Recently, it has been suggested that the primary motor cortex (M1) plays a critical role in implementing the fast and transient post-training phase of motor skill consolidation, known to yield an early boost in performance. Whether a comparable early boost in performance occurs following motor imagery (MIM) training is still unknown. To address this issue, two groups of subjects learned a finger tapping sequence either by MIM or physical practice (PP). In both groups, performance increased significantly in the post-training phase when compared with the pre-training phase and further increased after a 30 min resting period, indicating that both MIM and PP trainings were equally efficient and induced an early boost in motor performance. This conclusion was corroborated by the results of an additional control group. In a second experiment, we then investigated the causal role of M1 in implementing the early boost process resulting from MIM training. To do so, we inhibited M1 by applying a continuous theta-burst stimulation (cTBS) in healthy volunteers just after they learnt, by MIM, the same finger-tapping task as in Experiment #1. As a control, cTBS was applied over the vertex of subjects who underwent the same experiment. We found that cTBS applied over M1 selectively abolished the early boost process subsequent to MIM training. Altogether, the present study provides evidence that MIM practice induces an early boost in performance and demonstrates that M1 is causally involved in this process. These findings further divulge some degree of behavioral and neuronal similitude between MIM and PP.
PMCID: PMC3202558  PMID: 22046337
Despite the prominence of parietal activity in human neuromaging investigations of sensorimotor and cognitive processes there remains uncertainty about basic aspects of parietal cortical anatomical organization. Descriptions of human parietal cortex draw heavily on anatomical schemes developed in other primate species but the validity of such comparisons has been questioned by claims that there are fundamental differences between the parietal cortex in humans and other primates. A scheme is presented for parcellation of human lateral parietal cortex into component regions on the basis of anatomical connectivity and the functional interactions of the resulting clusters with other brain regions. Anatomical connectivity was estimated using diffusion-weighted magnetic resonance image (MRI) based tractography and functional interactions were assessed by correlations in activity measured with functional MRI (fMRI) at rest. Resting state functional connectivity was also assessed directly in the rhesus macaque lateral parietal cortex in an additional experiment and the patterns found reflected known neuroanatomical connections. Cross-correlation in the tractography-based connectivity patterns of parietal voxels reliably parcellated human lateral parietal cortex into ten component clusters. The resting state functional connectivity of human superior parietal and intraparietal clusters with frontal and extrastriate cortex suggested correspondences with areas in macaque superior and intraparietal sulcus. Functional connectivity patterns with parahippocampal cortex and premotor cortex again suggested fundamental correspondences between inferior parietal cortex in humans and macaques. In contrast, the human parietal cortex differs in the strength of its interactions between the central inferior parietal lobule region and the anterior prefrontal cortex.
PMCID: PMC3091022  PMID: 21411650
10.  Evidence for two concurrent inhibitory mechanisms during response preparation 
Inhibitory mechanisms are critically involved in goal-directed behaviors. To gain further insight into how such mechanisms shape motor representations during response preparation, motor evoked potentials (MEPs) elicited by transcranial magnetic stimulation (TMS) and H-reflexes were recorded from left hand muscles during choice reaction time tasks. The imperative signal, which indicated the required response, was always preceded by a preparatory cue. During the post-cue delay period, left MEPs were suppressed when the left hand had been cued for the forthcoming response, suggestive of a form of inhibition specifically directed at selected response representations. H-reflexes were also suppressed on these trials, indicating that the effects of this inhibition extend to spinal circuits. In addition, left MEPs were suppressed when the right hand was cued, but only when left hand movements were a possible response option before the onset of the cue. Notably, left hand H-reflexes were not modulated on these trials, consistent with a cortical locus of inhibition that lowers the activation of task-relevant, but non-selected responses. These results suggest the concurrent operation of two inhibitory mechanisms during response preparation: one decreases the activation of selected responses at the spinal level, helping to control when selected movements should be initiated by preventing their premature release; a second, upstream mechanism helps to determine what response to make during a competitive selection process.
PMCID: PMC2852647  PMID: 20220014
cognitive control; competition; corticospinal excitability; decision-making; response selection; transcranial magnetic stimulation; H-reflex
11.  Ventral premotor to primary motor cortical interactions during object-driven grasp in humans 
Interactions between the ventral premotor (PMv) and the primary motor cortex (M1) are crucial for transforming an object's geometrical properties, such as its size and shape, into a motor command suitable for grasp of the object. Recently, we showed that PMv interacts with M1 in a specific fashion, depending on the hand posture. However, the functional connectivity between PMv and M1 during the preparation of an actual grasp is still unknown.
To address this issue, PMv–M1 interactions were tested while subjects were preparing to grasp different visible objects requiring either a precision grip or a whole hand grasp. A conditioning–test transcranial magnetic stimulation (TMS) paradigm was used: a test stimulus was applied over M1 either in isolation or after a conditioning stimulus delivered, at different delays, over the ipsilateral PMv. Motor evoked potentials (MEPs) were recorded in the first dorsal interosseus and abductor digiti minimi muscles, which show highly differentiated activity according to grasp.
While subjects prepared to grasp, delivering a conditioning PMv pulse 6 or 8 msec before a test pulse over M1 strikingly facilitated MEPs in the specific muscles that were used in the upcoming grasp. This degree of facilitation correlated with the amount of muscle activity used later in the trial to grasp the objects.
The present results demonstrate that, during grasp preparation, the PMv–M1 interactions are muscle-specific. PMv appears to process the object geometrical properties relevant for the upcoming grasp, and transmits this information to M1, which in turn generates a motor command appropriate for the grasp. We also reveal that the grasp-specific facilitation resulting from PMv–M1 interactions is differently related to the upcoming grasp muscle activity than is that from paired-pulse stimulation over M1, suggesting that these two TMS paradigms assess the excitability of cortico-cortical pathways devoted to the control of grasp at two different levels.
PMCID: PMC2730595  PMID: 19345344
Functional connectivity; EMG; Motor control; Hand shaping; Corticospinal
12.  Spatial and Temporal Dynamics of Attentional Guidance during Inefficient Visual Search 
PLoS ONE  2008;3(5):e2219.
Spotting a prey or a predator is crucial in the natural environment and relies on the ability to extract quickly pertinent visual information. The experimental counterpart of this behavior is visual search (VS) where subjects have to identify a target amongst several distractors. In difficult VS tasks, it has been found that the reaction time (RT) is influenced by salience factors, such as the target-distractor similarity, and this finding is usually regarded as evidence for a guidance of attention by preattentive mechanisms. However, the use of RT measurements, a parameter which depends on multiple factors, allows only very indirect inferences about the underlying attentional mechanisms. The purpose of the present study was to determine the influence of salience factors on attentional guidance during VS, by measuring directly attentional allocation. We studied attention allocation by using a dual covert VS task in subjects who had 1) to detect a target amongst different items and 2) to report letters briefly flashed inside those items at different delays. As predicted, we showed that parallel processes guide attention towards the most relevant item by virtue of both goal-directed and stimulus-driven factors, and we demonstrated that this attentional selection is a prerequisite for target detection. In addition, we show that when the target is characterized by two features (conjunction VS), the goal-directed effects of both features are initially combined into a unique salience value, but at a later stage, grouping phenomena interact with the salience computation, and lead to the selection of a whole group of items. These results, in line with Guided Search Theory, show that efficient and rapid preattentive processes guide attention towards the most salient item, allowing to reduce the number of attentional shifts needed to find the target.
PMCID: PMC2375050  PMID: 18493320

Results 1-12 (12)