No support for a causal role of primary motor cortex in construing meaning from language: An rTMS study.

Embodied cognition theories predict a functional involvement of sensorimotor processes in language understanding. In a preregistered experiment, we tested this idea by investigating whether interfering with primary motor cortex (M1) activation can change how people construe meaning from action language. Participants were presented with sentences describing actions (e.g., "turning off the light") and asked to choose between two interpretations of their meaning, one more concrete (e.g., "flipping a switch") and another more abstract (e.g., "going to sleep"). Prior to this task, participants' M1 was disrupted using repetitive transcranial magnetic stimulation (rTMS). The results yielded strong evidence against the idea that M1-rTMS affects meaning construction (BF01 > 30). Additional analyses and control experiments suggest that the absence of effect cannot be accounted for by failure to inhibit M1, lack of construct validity of the task, or lack of power to detect a small effect. In sum, these results do not support a causal role for primary motor cortex in building meaning from action language.

Neural basis of social attention: common and distinct mechanisms for social and nonsocial orienting stimuli.

Social and nonsocial directional stimuli (such as gaze and arrows, respectively) share their ability to trigger attentional processes, although the issue of whether social stimuli generate other additional (and unique) attentional effects is still under debate. In this study, we used the spatial interference paradigm to explore, using functional magnetic resonance imaging, shared and dissociable brain activations produced by gaze and arrows. Results showed a common set of regions (right parieto-temporo-occipital) similarly involved in conflict resolution for gaze and arrows stimuli, which showed stronger co-activation for incongruent than congruent trials. The frontal eye field showed stronger functional connectivity with occipital regions for congruent as compared with incongruent trials, and this effect was enhanced for gaze as compared with arrow stimuli in the right hemisphere. Moreover, spatial interference produced by incongruent (as compared with congruent) arrows was associated with increased functional coupling between the right frontal eye field and a set of regions in the left hemisphere. This result was not observed for incongruent (as compared with congruent) gaze stimuli. The right frontal eye field also showed greater coupling with left temporo-occipital regions for those conditions in which larger conflict was observed (arrow incongruent vs. gaze incongruent trials, and gaze congruent vs. arrow congruent trials). These findings support the view that social and nonsocial stimuli share some attentional mechanisms, while at the same time highlighting other differential effects. Highlights Attentional orienting triggered by social (gaze) and nonsocial (arrow) cues is comparable. When social and nonsocial stimuli are used as targets, qualitatively different behavioral effects are observed. This study explores the neural bases of shared and dissociable neural mechanisms for social and nonsocial stimuli. Shared mechanisms were found in the functional coupling between right parieto-temporo-occipital regions. Dissociable mechanisms were found in the functional coupling between right frontal eye field and ipsilateral and contralateral occipito-temporal regions.

Attention to space and time: Independent or interactive systems? A narrative review.

While there is ample evidence for the ability to selectively attend to where in space and when in time a relevant event might occur, it remains poorly understood whether spatial and temporal attention operate independently or interactively to optimize behavior. To elucidate this important issue, we provide a narrative review of the literature investigating the relationship between the two. The studies were organized based on the attentional manipulation employed (endogenous vs. exogenous) and the type of task (detection vs. discrimination). Although the reviewed findings depict a complex scenario, three aspects appear particularly important in promoting independent or interactive effects of spatial and temporal attention: task demands, attentional manipulation, and their combination. Overall, the present review provides key insights into the relationship between spatial and temporal attention and identifies some critical gaps that need to be addressed by future research.

A transcranial magnetic stimulation study on the role of the left intraparietal sulcus in temporal orienting of attention.

Adaptive behavior requires the ability to orient attention to the moment in time at which a relevant event is likely to occur. Temporal orienting of attention has been consistently associated with activation of the left intraparietal sulcus (IPS) in prior fMRI studies. However, a direct test of its causal involvement in temporal orienting is still lacking. The present study tackled this issue by transiently perturbing left IPS activity with either online (Experiment 1) or offline (Experiment 2) transcranial magnetic stimulation (TMS). In both experiments, participants performed a temporal orienting task, alternating between blocks in which a temporal cue predicted when a subsequent target would appear and blocks in which a neutral cue provided no information about target timing. In Experiment 1 we used an online TMS protocol, aiming to interfere specifically with cue-related temporal processes, whereas in Experiment 2 we employed an offline protocol whereby participants performed the temporal orienting task before and after receiving TMS. The right IPS and/or the vertex were stimulated as active control regions. While results replicated the canonical pattern of temporal orienting effects on reaction time, with faster responses for temporal than neutral trials, these effects were not modulated by TMS over the left IPS (as compared to the right IPS and/or vertex regions) regardless of the online or offline protocol used. Overall, these findings challenge the causal role of the left IPS in temporal orienting of attention inviting further research on its underlying neural substrates.

Impaired attention mechanisms in confabulating patients: A VLSM and DWI study.

Attention is one of the most studied cognitive functions in brain-damaged populations or neurological syndromes, as its malfunction can be related to deficits in other higher cognitive functions. In the present study, we aimed at delimiting the attention deficits of a sample of brain-injured patients presenting confabulations by assessing their performance on alertness, spatial orienting, and executive control tasks. Confabulating patients, who present false memories or beliefs without intention to deceive, usually show memory deficits and/or executive dysfunction. However, it is also likely that attention processes may be impaired in patients showing confabulations. Here, we compared confabulating patients' attention performance to a lesion control group and a healthy control group. Confabulating patients' mean overall accuracy was lower than the one of healthy and lesion controls along the three experimental tasks. Importantly, confabulators presented a greater Simon congruency effect than both lesion controls and healthy controls in the presence of predictive spatial cues, besides a lower percentage of hits and longer RTs in the Go-NoGo task, demonstrating deficits in executive control. They also showed a higher reliance on alerting and spatially predictive orienting cues in the context of a deficient performance. Grey and white matter analyses showed that patients' percentage of hits in the Go-NoGo task was related to damage to the right inferior frontal gyrus (pars triangularis and pars opercularis), whereas the integrity of the right inferior fronto-occipital fasciculus was negatively correlated with their alertness effect. These results are consistent with previous literature highlighting an executive dysfunction in confabulating patients, and suggest that some additional forms of attention, such as alertness and spatial orienting, could be selectively impaired in this clinical syndrome.

Explicit vs. implicit spatial processing in arrow vs. eye-gaze spatial congruency effects.

Arrows and gaze stimuli lead to opposite spatial congruency effects. While standard congruency effects are observed for arrows (faster responses for congruent conditions), responses are faster when eye-gaze stimuli are presented on the opposite side of the gazed-at location (incongruent trials), leading to a reversed congruency effect (RCE). Here, we explored the effects of implicit vs. explicit processing of arrows and eye-gaze direction. Participants were required to identify the direction (explicit task) or the colour (implicit task) of left or right looking/pointing gaze or arrows, presented to either the left or right of the fixation point. When participants responded to the direction of stimuli, standard congruency effects for arrows and RCE for eye-gaze stimuli were observed. However, when participants responded to the colour of stimuli, no congruency effects were observed. These results suggest that it is necessary to explicitly pay attention to the direction of eye-gaze and arrows for the congruency effect to occur. The same pattern of data was observed when participants responded either manually or verbally, demonstrating that manual motor components are not responsible for the results observed. These findings are not consistent with some hypotheses previously proposed to explain the RCE observed with eye-gaze stimuli and, therefore, call for an alternative plausible hypothesis.

Functional characterization of correct and incorrect feature integration.

Our sensory system constantly receives information from the environment and our own body. Despite our impression to the contrary, we remain largely unaware of this information and often cannot report it correctly. Although perceptual processing does not require conscious effort on the part of the observer, it is often complex, giving rise to errors such as incorrect integration of features (illusory conjunctions). In the present study, we use functional magnetic resonance imaging to study the neural bases of feature integration in a dual task that produced ~30% illusions. A distributed set of regions demonstrated increased activity for correct compared to incorrect (illusory) feature integration, with increased functional coupling between occipital and parietal regions. In contrast, incorrect feature integration (illusions) was associated with increased occipital (V1-V2) responses at early stages, reduced functional connectivity between right occipital regions and the frontal eye field at later stages, and an overall decrease in coactivation between occipital and parietal regions. These results underscore the role of parietal regions in feature integration and highlight the relevance of functional occipito-frontal interactions in perceptual processing.

The cost of attentional reorienting on conscious visual perception: an MEG study.

How do attentional networks influence conscious perception? To answer this question, we used magnetoencephalography in human participants and assessed the effects of spatially nonpredictive or predictive supra-threshold peripheral cues on the conscious perception of near-threshold Gabors. Three main results emerged. (i) As compared with invalid cues, both nonpredictive and predictive valid cues increased conscious detection. Yet, only predictive cues shifted the response criterion toward a more liberal decision (i.e. willingness to report the presence of a target under conditions of greater perceptual uncertainty) and affected target contrast leading to 50% detections. (ii) Conscious perception following valid predictive cues was associated to enhanced activity in frontoparietal networks. These responses were lateralized to the left hemisphere during attentional orienting and to the right hemisphere during target processing. The involvement of frontoparietal networks occurred earlier in valid than in invalid trials, a possible neural marker of the cost of re-orienting attention. (iii) When detected targets were preceded by invalid predictive cues, and thus reorienting to the target was required, neural responses occurred in left hemisphere temporo-occipital regions during attentional orienting, and in right hemisphere anterior insular and temporo-occipital regions during target processing. These results confirm and specify the role of frontoparietal networks in modulating conscious processing and detail how invalid orienting of spatial attention disrupts conscious processing.

Attention does not always help: The role of expectancy, divided, and spatial attention on illusory conjunctions.

Humans have the subjective impression of a rich perceptual experience, but this perception is riddled with errors that might be produced by top-down expectancies or failures in feature integration. The role of attention in feature integration is still unclear. Some studies support the importance of attention in feature integration, whereas others suggest that feature integration does not require attention. Understanding attention as a heterogeneous system, in this study, we explored the role of divided (as opposed to focused-Experiment 1) attention, and endogenous-exogenous spatial orienting (Experiments 2 and 3) in feature integration. We also explored the role of feature expectancy, by presenting stimulus features that were completely unexpected to the participants. Results demonstrated that both endogenous and exogenous orienting improved feature integration whereas divided attention did not. Moreover, a strong and consistent feature expectancy effect was observed, demonstrating perceptual completion when an unexpected perceptual feature was presented in the scene. These results support the feature confirmation account, which proposes that attention is important for top-down matching of stable representations.

Visual attention in realistic driving situations: Attentional capture and hazard prediction.

In real life, many objects catch our attention involuntarily or exogenously. Exogenous attention occurs fast and its effects are short-lived. In the laboratory, when attentional orientation is studied, both valid and invalid attentional signals are used: the valid ones direct the attention to a location where something relevant is going to appear. The invalid ones occur in a location where nothing relevant is going to happen. Usually, performance is improved when valid signals rather than invalid ones are presented. This work is novel in that it explores the effects of attentional capture and driving experience in situations of day-to-day driving while participants carry out a Hazard Prediction task. We created new Hazard Prediction (HPr) and Risk Estimation (RE) tests when driving by selecting 48 short videos recorded in a realistic way from the perspective of a car driver. We created valid and invalid trials by selecting videos in which a what?? was presented in the same spatial location as the one where the hazard was beginning to develop or in a different location. Simple situations, with only one developing hazard, were also presented. A total of 92 participants (30 experienced drivers, 32 novices and 30 with no experience) were placed in the position of the driver and answered the questions: 1) What will happen after the video is cut? 2) To what extent do you consider this situation risky? The results from the Hazard Prediction test replicate the attentional capture effect in complex driving situations, with invalid trials obtaining the worst results, followed by valid and simple ones. Participants with experience obtained better scores than novices, and novices were better than drivers without experience. No interaction between attentional orientation and experience was found, suggesting the obligatory and automatic nature of orientation processes, which do not appear to be compensated for by driving experience. No significant differences were found for the Risk Estimation test.

Fronto-parietal networks underlie the interaction between executive control and conscious perception: Evidence from TMS and DWI.

The executive control network is involved in novel situations or those in which prepotent responses need to be overridden. Previous studies have demonstrated that when control is exerted, conscious perception is impaired, and this effect is related to the functional connectivity of fronto-parietal regions. In the present study, we explored the causal involvement of one of the nodes of this fronto-parietal network (the right Supplementary Motor Area, SMA) in the interaction between executive control and conscious perception. Participants performed a dual task in which they responded to a Stroop task while detecting the presence/absence of a near-threshold Gabor stimulus. Concurrently, transcranial magnetic stimulation (TMS) was applied over the right SMA or a control site (vertex; Experiment 1). As a further control, the right Frontal Eye Field (FEF) was stimulated in Experiment 2. Diffusion-weighted imaging (DWI) tractography was used to isolate the three branches of the superior longitudinal fasciculus (SLF I, II and III), and the frontal aslant tract (FAT), and to explore if TMS effects were related to their micro- and macrostructural characteristics. Results demonstrated reduced perceptual sensitivity on incongruent as compared to congruent Stroop trials. A causal role of the right SMA on the modulation of perceptual sensitivity by executive control was only demonstrated when the microstructure of the right SLF III or the left FAT were taken into account. The volume of the right SLF III was also related to the modulation of response criterion by executive control when the right FEF was stimulated. These results add evidence in favor of shared neural correlates for attention and conscious perception in fronto-parietal regions and highlight the role of white matter in TMS effects.

The causal role of the left parietal lobe in facilitation and inhibition of return.

Following non-informative peripheral cues, responses are facilitated at the cued compared to the uncued location at short cue-target intervals. This effect reverses at longer intervals, giving rise to Inhibition of Return (IOR). The integration-segregation hypothesis (Lupiáñez, 2010) suggests that peripheral cues always produce an onset-detection cost regardless the behavioral cueing effect that is measured - either facilitation or IOR. In the present study, we used transcranial magnetic stimulation (TMS) to investigate the causal contribution of this detection cost to performance. We used a cueing paradigm with a target discrimination task that was preceded by a non-informative peripheral cue. The presence-absence of a central intervening event was manipulated. Online TMS to the left superior parietal lobe (compared to an active vertex stimulation) lead to an overall more positive effect (faster responses for cued as compared to uncued trials), by putatively impairing the detection cost contribution to performance. The data revealed a strong association between overall RT and the TMS effect, and also between overall RT and the integrity of the first branch of the left superior longitudinal fascicule. These results have critical implications not only for the open debate about the mechanism/s underlying spatial orienting effects, but also for the growing literature demonstrating that white matter connectivity is crucial for explaining inter-individual behavioral variability.

Conscious perception in patients with prefrontal damage.

We are conscious and verbally report some of the information reaching our senses, although a big amount of information is processed unconsciously. There is no agreement about the neural correlates of consciousness, with low-level theories proposing that neural processing on primary sensory brain regions is the most important neural correlate of consciousness, while high-level theories propose that activity within the fronto-parietal network is the key component of conscious processing (Block, 2009). Contrary to the proposal of high-level theories, patients with prefrontal lobe damage do not present clinical symptoms associated to consciousness deficits. In the present study, we explored the conscious perception of near-threshold stimuli in a group of patients with right prefrontal damage and a group of matched healthy controls. Results demonstrated that perceptual contrast to perceive the near-threshold targets was related to damage to the right dorsolateral prefrontal cortex, and with reduced integrity of the ventral branch of the right superior longitudinal fascicule (SLF III). These results suggest a causal role of the prefrontal lobe in conscious processing.

Causal Contributions of the SMA to Alertness and Consciousness Interactions.

Phasic alertness facilitates conscious perception through a fronto-striatal network, including the supplementary motor area (SMA). The functioning of the ventral attentional network has been related to the alerting system, overlapping with the ventral branch of the superior longitudinal fasciculus (SLF III). In this study, we use repetitive transcranial magnetic stimulation (rTMS) and a conscious detection task with near-threshold stimuli that could be preceded by an alerting tone to explore the causal implication of the SMA in the relationship between phasic alertness and conscious perception. Complementary to SMA stimulation, a sham and an active condition (left inferior parietal lobe; IPL) were included. Deterministic tractography was used to isolate the right and left SLF III. Behaviorally, the alerting tone enhanced conscious perception and confidence ratings. rTMS over the SMA reduced the alerting effect on the percentage of perceived stimuli while rTMS over the left IPL produced no modulations, demonstrating a region-specific effect. Additionally, a correlation between the rTMS effect and the integrity of the right SLF III was found. Our results highlight the causal implication of a frontal region, the SMA, in the relationship between phasic alertness and conscious perception, which is related to the white matter microstructure of the SLF III.

Heart-rate modulations reveal attention and consciousness interactions.

Our environment is constantly overloaded with information, although we cannot consciously process all the stimulation reaching our senses. Current theoretical models are focused on the cognitive and neural processes underlying conscious perception. However, cognitive processes do not occur in an isolated brain but in a complex interaction between the environment, the brain, and the organism. The brain-body interaction has largely been neglected in the study of conscious perception. The aim of the present study was to explore if heart rate and skin conductance (SC) are modulated by the interaction between phasic alertness and conscious perception. We presented near-threshold visual stimuli that could be preceded by an alerting tone on 50% of the trials. Behaviorally, phasic alerting improved perceptual sensitivity for detecting a near-threshold stimulus (along with changes in response criterion). Following the alerting tone, a cardiac deceleration-acceleration pattern was observed, which was more pronounced when the near-threshold stimulus was consciously perceived in comparison with unconsciously perceived stimuli. SC results further showed some degree of subliminal processing of unseen stimuli. These results reveal that cardiac activity could be a marker of attention and consciousness interactions, emphasizing the need for taking into account brain-body interactions for current theoretical models of consciousness.

Connectivity of Frontoparietal Regions Reveals Executive Attention and Consciousness Interactions.

The executive control network is involved in the voluntary control of novel and complex situations. Solving conflict situations or detecting errors have demonstrated to impair conscious perception of near-threshold stimuli. The aim of this study was to explore the neural mechanisms underlying executive control and its interaction with conscious perception using functional magnetic resonance imaging and diffusion-weighted imaging. To this end, we used a dual-task paradigm involving Stroop and conscious detection tasks with near-threshold stimuli. A set of prefrontal and frontoparietal regions were more strongly engaged for incongruent than congruent trials while a distributed set of frontoparietal regions showed stronger activation for consciously than nonconsciously perceived trials. Functional connectivity analysis revealed an interaction between executive control and conscious perception in frontal and parietal nodes. The microstructural properties of the middle branch of the superior longitudinal fasciculus were associated with neural measures of the interaction between executive control and consciousness. These results demonstrate that conscious perception and executive control share neural resources in frontoparietal networks, as proposed by some influential models.

Neural modulations of interference control over conscious perception.

The relation between attention and consciousness is a highly debated topic in Cognitive Neuroscience. Although there is an agreement about their relationship at the functional level, there is still no consensus about how these two cognitive processes interact at the neural level. According to the gateway hypothesis (Posner, 1994), attention filters the information accessing to consciousness, resulting in both neural and functional modulations. Contrary to this idea, the cumulative influence hypothesis (Tallon-Baudry, 2012) proposes that both attention and consciousness independently impact decision processes about the perception of stimuli. Accordingly, we could observe an interaction between attention and consciousness at the behavioral level, but not at the neural level. Previous studies have shown that alerting and orienting networks of attention modulate participants' ability to verbally report near-threshold visual stimuli both at behavioral and neural levels, supporting the gateway hypothesis over the cumulative influence hypothesis. The impact of the executive control network of attention on conscious perception, however, has only been explored behaviorally (Colás et al., 2017). In the present study, we employed high-density encephalography to investigate the neural basis of the interaction between executive attention and conscious perception. We presented a classical Stroop task concurrently with a detection task of near-threshold stimuli. In two separate sessions, we manipulated the proportion of congruent and incongruent Stroop stimuli. We found that the Stroop-evoked N2 potential (usually associated to conflict detection and localized in the anterior cingulate cortex) was modulated by both conflict detection and conscious perception processes. These results suggest that the relation between executive control and conscious perception lies in frontal lobe regions associated to conflict detection, supporting the gateway hypothesis over the cumulative influence hypothesis.

Semantic incongruity attracts attention at a pre-conscious level: Evidence from a TMS study.

Unpredicted objects, i.e., those that do not fit in a specific context, have been shown to quickly attract attention as a mean of extracting more information about potentially relevant items. Whether the required semantic processing triggering the attraction of attention can occur independently of participants' awareness of the object is still a highly debated topic. In the present study we make use of a change detection task in which we manipulate the semantic congruity between the to-be-detected object and the background scene. We applied inhibitory repetitive transcranial magnetic stimulation (rTMS) over the right temporo-parietal junction (right TPJ) and a control location (vertex) to test the causal role of the former in the processing of objects at a pre-conscious level. Our results clearly show that semantic congruity can impact detection and identification processes in opposite ways, even when low-level features are controlled for. Incongruent objects are quickly detected but poorly identified. rTMS over the right TPJ effectively diminishes semantic effects on object detection. These results suggest that at least some high order category processing takes place before conscious detection to direct attention towards the most informative regions of space. Moreover, rTMS over right TPJ also impacts object identification, which calls for a re-evaluation of right TPJ's role on object processing.

White matter microstructure of attentional networks predicts attention and consciousness functional interactions.

Attention is considered as one of the pre-requisites of conscious perception. Phasic alerting and exogenous orienting improve conscious perception of near-threshold information through segregated brain networks. Using a multimodal neuroimaging approach, combining data from functional MRI (fMRI) and diffusion-weighted imaging (DWI), we investigated the influence of white matter properties of the ventral branch of superior longitudinal fasciculus (SLF III) in functional interactions between attentional systems and conscious perception. Results revealed that (1) reduced integrity of the left hemisphere SLF III was predictive of the neural interactions observed between exogenous orienting and conscious perception, and (2) increased integrity of the left hemisphere SLF III was predictive of the neural interactions observed between phasic alerting and conscious perception. Our results combining fMRI and DWI data demonstrate that structural properties of the white matter organization determine attentional modulations over conscious perception.

Interference Control Modulations Over Conscious Perception.

The relation between attention and consciousness has been a controversial topic over the last decade. Although there seems to be an agreement on their distinction at the functional level, no consensus has been reached about attentional processes being or not necessary for conscious perception. Previous studies have explored the relation of alerting and orienting systems of attention and conscious perception, but the impact of the anterior executive attention system on conscious access remains unexplored. In the present study, we investigated the behavioral interaction between executive attention and conscious perception, testing control mechanisms both at stimulus-level representation and after error commission. We presented a classical Stroop task, manipulating the proportion of congruent and incongruent trials, and analyzed the effect of reactive and proactive control on the conscious perception of near-threshold stimuli. Reactive control elicited under high proportion congruent conditions influenced participants' decision criterion, whereas proactive control elicited under low proportion congruent conditions was ineffective in modulating conscious perception. In addition, error commission affected both perceptual sensitivity to detect near-threshold information and response criterion. These results suggest that reactivation of task goals through reactive control strategies in conflict situations impacts decision stages of conscious processing, whereas interference control elicited by error commission impacts both perceptual sensitivity and decision stages of conscious processing. We discuss the implications of our results for the gateway hypothesis about attention and consciousness, as they showed that interference control (both at stimulus-level representation and after error commission) can modulate the conscious access of near-threshold stimuli.