Elucidating the difference between mind-wandering and day-dreaming terms.

Self-generated thoughts have been widely investigated in recent years, while the terms "mind-wandering" and "day-dreaming" are usually used interchangeably. But are these terms equivalent? To test this, online study participants were presented with situations of a protagonist engaged in self-generated thoughts. The scenarios differed with regard to type of situation, the activity in which the protagonist was engaged in, and the properties of the self-generated thoughts. Two different groups evaluated the same situations; one group evaluated the extent to which the protagonist mind-wandered and another the extent to which the protagonist day-dreamt. Our key findings were that the situations were perceived differently with regard to mind-wandering and day-dreaming, depending on whether self-generated thoughts occurred when the protagonist was busy with another activity and the type of self-generated thoughts. In particular, while planning, worrying, and ruminating thoughts were perceived more as mind-wandering in situations involving another activity/task, the situations without another activity/task involving recalling past events and fantasizing thoughts were perceived more as day-dreaming. In the additional experiment, we investigated laypeople's reasons for classifying the situation as mind-wandering or day-dreaming. Our results altogether indicate that mind-wandering and day-dreaming might not be fully equivalent terms.

Naturalistic auditory stimuli with fNIRS prefrontal cortex imaging: A potential paradigm for disorder of consciousness diagnostics (a study with healthy participants).

Disorder of consciousness (DOC) is a devastating condition due to brain damage. A patient in this condition is non-responsive, but nevertheless might be conscious at least at some level. Determining the conscious level of DOC patients is important for both medical and ethical reasons, but reliably achieving this has been a major challenge. Naturalistic stimuli in combination with neuroimaging have been proposed as a promising approach for DOC patient diagnosis. Capitalizing on and extending this proposal, the goal of the present study conducted with healthy participants was to develop a new paradigm with naturalistic auditory stimuli and functional near-infrared spectroscopy (fNIRS) - an approach that can be used at the bedside. Twenty-four healthy participants passively listened to 9 min of auditory story, scrambled auditory story, classical music, and scrambled classical music segments while their prefrontal cortex activity was recorded using fNIRS. We found much higher intersubject correlation (ISC) during story compared to scrambled story conditions both at the group level and in the majority of individual subjects, suggesting that fNIRS imaging of the prefrontal cortex might be a sensitive method to capture neural changes associated with narrative comprehension. In contrast, the ISC during the classical music segment did not differ reliably from scrambled classical music and was also much lower than the story condition. Our main result is that naturalistic auditory stories with fNIRS might be used in a clinical setup to identify high-level processing and potential consciousness in DOC patients.

Intracranial study in humans: Neural spectral changes during watching comedy movie of Charlie Chaplin.

Humor plays a prominent role in our lives. Thus, understanding the cognitive and neural mechanisms of humor is particularly important. Previous studies that investigated neural substrates of humor used functional MRI and to a lesser extent EEG. In the present study, we conducted intracranial recording in human patients, enabling us to obtain the signal with high temporal precision from within specific brain locations. Our analysis focused on the temporal lobe and the surrounding areas, the temporal lobe was most densely covered in our recording. Thirteen patients watched a fragment of a Charlie Chaplin movie. An independent group of healthy participants rated the same movie fragment, helping us to identify the most funny and the least funny frames of the movie. We compared neural activity occurring during the most funny and least funny frames across frequencies in the range of 1-170 Hz. The most funny compared to least funny parts of the movie were associated with activity modulation in the broadband high-gamma (70-170 Hz; mostly activation) and to a lesser extent gamma band (40-69Hz; activation) and low frequencies (1-12 Hz, delta, theta, alpha bands; mostly deactivation). With regard to regional specificity, we found three types of brain areas: (I) temporal pole, middle and inferior temporal gyrus (both anterior and posterior) in which there was both activation in the high-gamma/gamma bands and deactivation in low frequencies; (II) ventral part of the temporal lobe such as the fusiform gyrus, in which there was mostly deactivation the low frequencies; (III) posterior temporal cortex and its environment, such as the middle occipital and the temporo-parietal junction, in which there was activation in the high-gamma/gamma band. Overall, our results suggest that humor appreciation might be achieved by neural activity across the frequency spectrum.

Similarity in activity and laterality patterns in the angular gyrus during autobiographical memory retrieval and self-referential processing.

Long-term memory is arguably one of the key cognitive functions. At the neural level, the lateral parietal cortex and the angular gyrus, particularly in the left hemisphere, exhibit strong activations during autobiographical and episodic memory retrieval. In a separate sub-field, left-lateralized activations of the angular gyrus are also found during self-referential processing, defined as higher activity when a trait term is judged by participants as being related to them vs. related to someone else. The question is whether episodic/autobiographical memory retrieval and self-referential processing effects are related. In the present study, thirty participants participated in the fMRI study with two separate experiments: autobiographical memory retrieval (Experiment 1) and self-referential processing (Experiment 2). In a series of analyses, including the most critical spatial correlation analysis between experiments, we found neural similarity between autobiographical memory retrieval and self-referential processing. Given that self-referential processing was identified in a selective way, the most plausible interpretation of our findings is that self-referential processing might partly explain the activation of the left angular gyrus during autobiographical memory retrieval. Our results are in line with the seminal view of Endel Tulving that the sense of self is a fundamental attribute of long-term memory recollection. However, it should be emphasized that: a) our results do not imply that the left angular gyrus is not involved in the retrieval of episodic memory details; and b) given that our experiment included an autobiographical memory task, generalization of our results to the episodic memory laboratory tasks has yet to be tested.

Neural modulations in the auditory cortex during internal and external attention tasks: A single-patient intracranial recording study.

Brain sensory processing is not passive, but is rather modulated by our internal state. Different research methods such as non-invasive imaging methods and intracranial recording of the local field potential (LFP) have been used to study to what extent sensory processing and the auditory cortex in particular are modulated by selective attention. However, at the level of the single- or multi-units the selective attention in humans has not been tested. In addition, most previous research on selective attention has explored externally-oriented attention, but attention can be also directed inward (i.e., internal attention), like spontaneous self-generated thoughts and mind-wandering. In the present study we had a rare opportunity to record multi-unit activity (MUA) in the auditory cortex of a patient. To complement, we also analyzed the LFP signal of the macro-contact in the auditory cortex. Our experiment consisted of two conditions with periodic beeping sounds. The participants were asked either to count the beeps (i.e., an "external attention" condition) or to recall the events of the previous day (i.e., an "internal attention" condition). We found that the four out of seven recorded units in the auditory cortex showed increased firing rates in "external attention" compared to "internal attention" condition. The beginning of this attentional modulation varied across multi-units between 30-50 msec and 130-150 msec from stimulus onset, a result that is compatible with an early selection view. The LFP evoked potential and induced high gamma activity both showed attentional modulation starting at about 70-80 msec. As the control, for the same experiment we recorded MUA activity in the amygdala and hippocampus of two additional patients. No major attentional modulation was found in the control regions. Overall, we believe that our results provide new empirical information and support for existing theoretical views on selective attention and spontaneous self-generated cognition.

Functional near-infrared spectroscopy imaging of the prefrontal cortex during a naturalistic comedy movie.

Naturalistic stimulation (i.e., movies and auditory narratives of some minutes' length) has been a powerful approach to bringing more real-life experiences into laboratory experiments. Data-driven, intersubject correlation (ISC) analysis permits examining to what extent activity in a specific brain region correlates across participants during exposure to a naturalistic stimulus, as well as testing whether neural activity correlates with behavioral measures. Notably, most of the previous research with naturalistic stimuli was conducted using functional fMRI (fMRI). Here, we tested whether a naturalistic approach and the ISC are feasible using functional near-infrared spectroscopy (fNIRS) - the imaging method particularly suited for populations of patients and children. Fifty-three healthy adult participants watched twice a 3-min segment of a Charlie Chaplin movie while we recorded the brain activity on the surface of their prefrontal cortex using fNIRS. In addition, an independent group of 18 participants used a continuous scoring procedure to rate the extent to which they felt that different parts of the movie fragment were funny. Our two findings were as follows. First, we found higher-than-zero ISC in fNIRS signals in the prefrontal cortex lobes, a result that was particularly high in the oxygenated channels during the first repetition of the movie. Second, we found a significant negative correlation between oxygenated brain signals and ratings of the movie's humorousness. In a series of control analyses we demonstrated that this latter correlation could not be explained by various non-humor-related movie sensory properties (e.g., auditory volume and image brightness). The key overall outcome of the present study is that fNIRS in combination with the naturalistic paradigms and the ISC might be a sensitive and powerful research method to explore cognitive processing. Our results also suggest a potential role of the prefrontal cortex in humor appreciation.

Face-selective multi-unit activity in the proximity of the FFA modulated by facial expression stimuli.

When we see someone's face, our brain usually effortlessly extracts a variety of information such as facial identity, expression, or gaze direction. While it is widely accepted that dedicated subsystems are responsible for different aspects of face processing, how these subsystems work together is not yet fully understood. To this extent, one of the most explored questions is whether and if so, to what extent facial expression processing interacts with other stages of facial processing. In the present study, we report a rare case of a patient for whom we were able to record multi-unit activity (MUA) in the proximity of the fusiform face area (FFA) while two out of four recorded multi-units were face-selective. In our experiment, the human subject was shown images of neutral and fearful faces as well as everyday objects and frightening images of natural disaster. We found that activity of both face-selective units was modulated by facial expression stimuli, starting at about 150 ms from stimulus onset. For both facial conditions we observed abrupt increase in firing rate with a simultaneous peak, suggesting that this activity and the modulation by facial expression stimuli likely reflected feed-forward processing. Interestingly, while in one multi-unit, the firing rate for fearful faces was higher than for neutral faces, in the other multi-units the polarity was reversed. Finally, modulation in the face-selective units was specific to emotional facial stimuli, but not to emotional stimuli in general. The present multi-unit results, albeit obtained only for several multi-units, nevertheless are potentially valuable for understanding mechanisms of facial processing in humans.

Comparing stimulus-evoked and spontaneous response of the face-selective multi-units in the human posterior fusiform gyrus.

The stimulus-evoked neural response is a widely explored phenomenon. Conscious awareness is associated in many cases with the corresponding selective stimulus-evoked response. For example, conscious awareness of a face stimulus is associated with or accompanied by stimulus-evoked activity in the fusiform face area (FFA). In addition to the stimulus-evoked response, spontaneous (i.e. task-unrelated) activity in the brain is also abundant. Notably, spontaneous activity is considered unconscious. For example, spontaneous activity in the FFA is not associated with conscious awareness of a face. The question is: what is the difference at the neural level between stimulus-evoked activity in a case that this activity is associated with conscious awareness of some content (e.g. activity in the FFA in response to fully visible face stimuli) and spontaneous activity in that same region of the brain? To answer this question, in the present study, we had a rare opportunity to record two face-selective multi-units in the vicinity of the FFA in a human patient. We compared multi-unit face-selective task-evoked activity with spontaneous prestimulus and a resting-state activity. We found that when activity was examined over relatively long temporal windows (e.g. 100-200 ms), face-selective stimulus-evoked firing in the recorded multi-units was much higher than the spontaneous activity. In contrast, when activity was examined over relatively short windows, we found many cases of high firing rates within the spontaneous activity that were comparable to stimulus-evoked activity. Our results thus indicate that the sustained activity is what might differentiate between stimulus-evoked activity that is associated with conscious awareness and spontaneous activity.

Importance, limits and caveats of the use of "disorders of consciousness" to theorize consciousness.

The clinical and fundamental exploration of patients suffering from disorders of consciousness (DoC) is commonly used by researchers both to test some of their key theoretical predictions and to serve as a unique source of empirical knowledge about possible dissociations between consciousness and cognitive and/or neural processes. For instance, the existence of states of vigilance free of any self-reportable subjective experience [e.g. "vegetative state (VS)" and "complex partial epileptic seizure"] originated from DoC and acted as a cornerstone for all theories by dissociating two concepts that were commonly equated and confused: vigilance and conscious state. In the present article, we first expose briefly the major achievements in the exploration and understanding of DoC. We then propose a synthetic taxonomy of DoC, and we finally highlight some current limits, caveats and questions that have to be addressed when using DoC to theorize consciousness. In particular, we show (i) that a purely behavioral approach of DoC is insufficient to characterize the conscious state of patients; (ii) that the comparison between patients in a minimally conscious state (MCS) and patients in a VS [also coined as unresponsive wakefulness syndrome (UWS)] does not correspond to a pure and minimal contrast between unconscious and conscious states and (iii) we emphasize, in the light of original resting-state positron emission tomography data, that behavioral MCS captures an important but misnamed clinical condition that rather corresponds to a cortically mediated state and that MCS does not necessarily imply the preservation of a conscious state.

Spontaneous cognition and its relationship to human creativity: A functional connectivity study involving a chain free association task.

Resting-state functional connectivity (RSFC) between various brain regions is thought to be associated with creative abilities. Extensive research correlating RSFC with performance on creativity tasks has revealed some of the RSFC patterns characterizing 'the creative brain'. Yet, our understanding of the neurocognitive processes underlying creative thinking still remains limited. This limitation results, in part, from the fact that standard creativity tasks used in these studies do not distinguish between the different modes of cognitive processing that are critical in creative cognition (e.g., spontaneous cognition vs. controlled cognition). In the present fMRI research we address this limitation by using a chain free association task â€‹- â€‹a task that we have recently refined and validated for the purpose of isolating measures of spontaneous cognition that are relevant for creative thinking (referred to as associative fluency and associative flexibility). In our study, 27 female participants completed standardized creativity tasks, a chain free association task, and a fMRI scan in which RSFC was measured. Our results indicate that higher scores on associative fluency are associated with stronger positive RSFC within the default mode network (DMN; i.e., between DMN regions). Critically, we provide evidence that the previously-identified relationship between performance on creativity tasks and connectivity within the DMN is partially mediated by associative fluency. Thus, our observations suggest that the heightened DMN connectivity observed in 'the creative brain' can be explained, at least to some extent, by spontaneous cognition. Overall, our study identifies unique RSFC patterns that are related specifically to spontaneous cognitive processes involved in creative ideation, thus shedding new light on mechanisms of creative processing.

Transcranial stimulation of the frontal lobes increases propensity of mind-wandering without changing meta-awareness.

Mind-wandering is omnipresent in our lives. The benefits of mind-wandering are not yet clear, but given how much time we spend mind-wandering, this mental function is likely to be important. Accordingly, it is essential to understand the neural and cognitive mechanisms of mind-wandering. In a recent study by the leading author of the present paper it was demonstrated that by applying transcranial direct current stimulation (tDCS) of the frontal lobes, but not sham or occipital cortex stimulation, it was possible to increase propensity of mind-wandering. The goal of the present study has been to replicate these previous findings and to extend them by examining whether changes in mind-wandering as a result of stimulation are associated with a change of meta-awareness of the attentional focus. By using a larger sample size and by conducting the experiment in a different country and language, we fully replicated the key original findings by showing that stimulation of the prefrontal cortex increased the level of mind-wandering. We also show that stimulation had no major effect on the level of meta-awareness of the attentional focus. Taken together, our results indicate that mind-wandering - probably the most internal and self-related mental function - can be modulated externally, that at least in some cases mind-wandering might not be regulated by meta-awareness, and that the frontal lobes might play a causal role in mind-wandering.

Perceptual similarity and the neural correlates of geometrical illusions in human brain structure.

Geometrical visual illusions are an intriguing phenomenon, in which subjective perception consistently misjudges the objective, physical properties of the visual stimulus. Prominent theoretical proposals have been advanced attempting to find common mechanisms across illusions. But empirically testing the similarity between illusions has been notoriously difficult because illusions have very different visual appearances. Here we overcome this difficulty by capitalizing on the variability of the illusory magnitude across participants. Fifty-nine healthy volunteers participated in the study that included measurement of individual illusion magnitude and structural MRI scanning. We tested the Muller-Lyer, Ebbinghaus, Ponzo, and vertical-horizontal geometrical illusions as well as a non-geometrical, contrast illusion. We found some degree of similarity in behavioral judgments of all tested geometrical illusions, but not between geometrical illusions and non-geometrical, contrast illusion. The highest similarity was found between Ebbinghaus and Muller-Lyer geometrical illusions. Furthermore, the magnitude of all geometrical illusions, and particularly the Ebbinghaus and Muller-Lyer illusions, correlated with local gray matter density in the parahippocampal cortex, but not in other brain areas. Our findings suggest that visuospatial integration and scene construction processes might partly mediate individual differences in geometric illusory perception. Overall, these findings contribute to a better understanding of the mechanisms behind geometrical illusions.

The default network and the combination of cognitive processes that mediate self-generated thought.

Self-generated cognitions, such as recalling personal memories or empathizing with others, are ubiquitous and essential for our lives. Such internal mental processing is ascribed to the Default Mode Network, a large network of the human brain, though the underlying neural and cognitive mechanisms remain poorly understood. Here, we tested the hypothesis that our mental experience is mediated by a combination of activities of multiple cognitive processes. Our study included four functional MRI experiments with the same participants and a wide range of cognitive tasks, as well as an analytical approach that afforded the identification of cognitive processes during self-generated cognition. We showed that several cognitive processes functioned simultaneously during self-generated mental activity. The processes had specific and localized neural representations, suggesting that they support different aspects of internal processing. Overall, we demonstrate that internally directed experience may be achieved by pooling over multiple cognitive processes.

On the domain-specificity of the visual and non-visual face-selective regions.

What happens in our brains when we see a face? The neural mechanisms of face processing - namely, the face-selective regions - have been extensively explored. Research has traditionally focused on visual cortex face-regions; more recently, the role of face-regions outside the visual cortex (i.e., non-visual-cortex face-regions) has been acknowledged as well. The major quest today is to reveal the functional role of each this region in face processing. To make progress in this direction, it is essential to understand the extent to which the face-regions, and particularly the non-visual-cortex face-regions, process only faces (i.e., face-specific, domain-specific processing) or rather are involved in a more domain-general cognitive processing. In the current functional MRI study, we systematically examined the activity of the whole face-network during face-unrelated reading task (i.e., written meaningful sentences with content unrelated to faces/people and non-words). We found that the non-visual-cortex (i.e., right lateral prefrontal cortex and posterior superior temporal sulcus), but not the visual cortex face-regions, responded significantly stronger to sentences than to non-words. In general, some degree of sentence selectivity was found in all non-visual-cortex cortex. Present result highlights the possibility that the processing in the non-visual-cortex face-selective regions might not be exclusively face-specific, but rather more or even fully domain-general. In this paper, we illustrate how the knowledge about domain-general processing in face-regions can help to advance our general understanding of face processing mechanisms. Our results therefore suggest that the problem of face processing should be approached in the broader scope of cognition in general.

Increasing propensity to mind-wander with transcranial direct current stimulation.

Humans mind-wander quite intensely. Mind wandering is markedly different from other cognitive behaviors because it is spontaneous, self-generated, and inwardly directed (inner thoughts). However, can such an internal and intimate mental function also be modulated externally by means of brain stimulation? Addressing this question could also help identify the neural correlates of mind wandering in a causal manner, in contrast to the correlational methods used previously (primarily functional MRI). In our study, participants performed a monotonous task while we periodically sampled their thoughts to assess mind wandering. Concurrently, we applied transcranial direct current stimulation (tDCS). We found that stimulation of the frontal lobes [anode electrode at the left dorsolateral prefrontal cortex (DLPFC), cathode electrode at the right supraorbital area], but not of the occipital cortex or sham stimulation, increased the propensity to mind-wander. These results demonstrate for the first time, to our knowledge, that mind wandering can be enhanced externally using brain stimulation, and that the frontal lobes play a causal role in mind-wandering behavior. These results also suggest that the executive control network associated with the DLPFC might be an integral part of mind-wandering neural machinery.

Successful decoding of famous faces in the fusiform face area.

What are the neural mechanisms of face recognition? It is believed that the network of face-selective areas, which spans the occipital, temporal, and frontal cortices, is important in face recognition. A number of previous studies indeed reported that face identity could be discriminated based on patterns of multivoxel activity in the fusiform face area and the anterior temporal lobe. However, given the difficulty in localizing the face-selective area in the anterior temporal lobe, its role in face recognition is still unknown. Furthermore, previous studies limited their analysis to occipito-temporal regions without testing identity decoding in more anterior face-selective regions, such as the amygdala and prefrontal cortex. In the current high-resolution functional Magnetic Resonance Imaging study, we systematically examined the decoding of the identity of famous faces in the temporo-frontal network of face-selective and adjacent non-face-selective regions. A special focus has been put on the face-area in the anterior temporal lobe, which was reliably localized using an optimized scanning protocol. We found that face-identity could be discriminated above chance level only in the fusiform face area. Our results corroborate the role of the fusiform face area in face recognition. Future studies are needed to further explore the role of the more recently discovered anterior face-selective areas in face recognition.

Exploring the unconscious using faces.

Understanding the mechanisms of unconscious processing is one of the most substantial endeavors of cognitive science. While there are many different empirical ways to address this question, the use of faces in such research has proven exceptionally fruitful. We review here what has been learned about unconscious processing through the use of faces and face-selective neural correlates. A large number of cognitive systems can be explored with faces, including emotions, social cueing and evaluation, attention, multisensory integration, and various aspects of face processing.

Neural Correlates of Subliminal Language Processing.

Language is a high-level cognitive function, so exploring the neural correlates of unconscious language processing is essential for understanding the limits of unconscious processing in general. The results of several functional magnetic resonance imaging studies have suggested that unconscious lexical and semantic processing is confined to the posterior temporal lobe, without involvement of the frontal lobe-the regions that are indispensable for conscious language processing. However, previous studies employed a similarly designed masked priming paradigm with briefly presented single and contextually unrelated words. It is thus possible, that the stimulation level was insufficiently strong to be detected in the high-level frontal regions. Here, in a high-resolution fMRI and multivariate pattern analysis study we explored the neural correlates of subliminal language processing using a novel paradigm, where written meaningful sentences were suppressed from awareness for extended duration using continuous flash suppression. We found that subjectively and objectively invisible meaningful sentences and unpronounceable nonwords could be discriminated not only in the left posterior superior temporal sulcus (STS), but critically, also in the left middle frontal gyrus. We conclude that frontal lobes play a role in unconscious language processing and that activation of the frontal lobes per se might not be sufficient for achieving conscious awareness.