Probing Our Built-in Calculator: A Systematic Narrative Review of Noninvasive Brain Stimulation Studies on Arithmetic Operation-Related Brain Areas.

This systematic review presented a comprehensive survey of studies that applied transcranial magnetic stimulation and transcranial electrical stimulation to parietal and nonparietal areas to examine the neural basis of symbolic arithmetic processing. All findings were compiled with regard to the three assumptions of the triple-code model (TCM) of number processing. Thirty-seven eligible manuscripts were identified for review (33 with healthy participants and 4 with patients). Their results are broadly consistent with the first assumption of the TCM that intraparietal sulcus both hold a magnitude code and engage in operations requiring numerical manipulations such as subtraction. However, largely heterogeneous results conflicted with the second assumption of the TCM that the left angular gyrus subserves arithmetic fact retrieval, such as the retrieval of rote-learned multiplication results. Support is also limited for the third assumption of the TCM, namely, that the posterior superior parietal lobule engages in spatial operations on the mental number line. Furthermore, results from the stimulation of brain areas outside of those postulated by the TCM show that the bilateral supramarginal gyrus is involved in online calculation and retrieval, the left temporal cortex in retrieval, and the bilateral dorsolateral prefrontal cortex and cerebellum in online calculation of cognitively demanding arithmetic problems. The overall results indicate that multiple cortical areas subserve arithmetic skills.

The impact of different distractions on outdoor visual search and object memory.

We investigated whether and how different types of search distractions affect visual search behavior and target memory while participants searched in a real-world environment. They searched either undistracted (control condition), listened to a podcast (auditory distraction), counted down aloud at intervals of three while searching (executive working memory load), or were forced to stop the search on half of the trials (time pressure). In line with findings from laboratory settings, participants searched longer but made fewer errors when the target was absent than when it was present, regardless of distraction condition. Furthermore, compared to the auditory distraction condition, the executive working memory load led to higher error rates (but not longer search times). In a surprise memory test after the end of the search tasks, recognition was better for previously present targets than for absent targets. Again, this was regardless of the previous distraction condition, although significantly fewer targets were remembered by the participants in the executive working memory load condition than by those in the control condition. The findings suggest that executive working memory load, but likely not auditory distraction and time pressure affected visual search performance and target memory in a real-world environment.

Resting-state neural correlates of visual Gestalt experience.

Subjective perceptual experience is influenced not only by bottom-up sensory information and experience-based top-down processes, but also by an individual's current brain state. Specifically, a previous study found increased prestimulus insula and intraparietal sulcus (IPS) activity before participants perceived an illusory Gestalt (global) compared with the non-illusory (local) interpretation of a bistable stimulus. That study provided only a snapshot of the brain state that favors the illusory interpretation. In the current study, we tested whether areas that differentiate between the illusory and non-illusory perception, immediately before stimulus onset, are also associated with an individual's general tendency to perceive it, which remains stable over time. We examined individual differences in task-free functional connectivity of insula and IPS and related them to differences in the individuals' duration of the two stimulus interpretations. We found stronger connectivity of the IPS with areas of the default mode and visual networks to be associated with shorter local perceptual phases, i.e. a faster switch to an illusory percept, and an opposite effect for insula connectivity with the early visual cortex. Our findings suggest an important role of IPS and insula interactions with nodes of key intrinsic networks in forming a perceptual tendency toward illusory Gestalt perception.

Fast and functionally specific cortical thickness changes induced by visual stimulation.

Structural characteristics of the human brain serve as important markers of brain development, aging, disease progression, and neural plasticity. They are considered stable properties, changing slowly over time. Multiple recent studies reported that structural brain changes measured with magnetic resonance imaging (MRI) may occur much faster than previously thought, within hours or even minutes. The mechanisms behind such fast changes remain unclear, with hemodynamics as one possible explanation. Here we investigated the functional specificity of cortical thickness changes induced by a flickering checkerboard and compared them to blood oxygenation level-dependent (BOLD) functional MRI activity. We found that checkerboard stimulation led to a significant thickness increase, which was driven by an expansion at the gray-white matter boundary, functionally specific to V1, confined to the retinotopic representation of the checkerboard stimulus, and amounted to 1.3% or 0.022 mm. Although functional specificity and the effect size of these changes were comparable to those of the BOLD signal in V1, thickness effects were substantially weaker in V3. Furthermore, a comparison of predicted and measured thickness changes for different stimulus timings suggested a slow increase of thickness over time, speaking against a hemodynamic explanation. Altogether, our findings suggest that visual stimulation can induce structural gray matter enlargement measurable with MRI.

Dissociation between Attention-Dependent and Spatially Specific Illusory Shape Responses within the Topographic Areas of the Posterior Parietal Cortex.

The human visual system consists of multiple topographic maps that extend from the early visual cortex (EVC) along the dorsal and ventral processing streams. Responses to illusory shapes within these maps have been demonstrated in the ventral stream areas, in particular the lateral occipital complex (LOC). Recently, the intraparietal sulcus (IPS) of the dorsal stream has been linked to the processing of illusory shapes defined by motion. It remains unclear whether the topographically organized parietal areas also respond to stationary illusory shapes, which would suggest their generic role in representing illusory content. In the current study we measured brain responses using fMRI while 30 human participants (12 male) observed flickering inducers around the fixation task. The inducers either formed an illusory diamond in the center, a triangle in the left or right hemifield, or were inverted such that no illusory figure was formed. We compared responses of parietal regions IPS0-IPS5 and SPL1 to each illusory figure with the nonillusory condition. To determine the role of attentional modulation on illusory shape responses we manipulated the difficulty of the fixation task. Our results show that all IPS areas responded to illusory shapes. The more posterior areas IPS0-IPS3 additionally displayed a preference toward contralateral shapes, while the more anterior areas IPS4 and IPS5 showed response attenuation with increased task difficulty. We suggest that the IPS can represent illusory content generated not only by moving, but also by stationary stimuli, and that there is a functional dissociation between attention-dependent anterior and spatially specific posterior topographic maps.SIGNIFICANCE STATEMENT The traditional view of the ventral visual pathway being solely responsible for representation of objects has recently been challenged by demonstrating illusory shape representation within the dorsal visual pathway with moving bistable stimuli. Our results provide evidence for the dorsal stream contribution to representing not only moving, but also stationary illusory shapes. Our results also show a functional subdivision along the topographic maps, with spatially specific shape responses in the more posterior, and attention-dependent responses in the more anterior areas. These findings have implications for our understanding of the relationship between attention and grouping in healthy individuals and neuropsychological patients. Furthermore, IPS areas should be considered in theoretical accounts and models of how subjective content is generated in the brain.

Increased insula activity precedes the formation of subjective illusory Gestalt.

The constructive nature of human perception sometimes leads us to perceiving rather complex impressions from simple sensory input: for example, recognizing animal contours in cloud formations or seeing living creatures in shadows of objects. A special type of bistable stimuli gives us a rare opportunity to study the neural mechanisms behind this process. Such stimuli can be visually interpreted either as simple or as more complex illusory content on the basis of the same sensory input. Previous studies demonstrated increased activity in the superior parietal cortex during the perception of an illusory Gestalt impression compared to a simpler interpretation. Here, we examined the role of slow fluctuations of resting-state fMRI activity in shaping the subsequent illusory interpretation by investigating activity related to the illusory Gestalt not only during, but also prior to its perception. We presented 31 participants with a bistable motion stimulus, which can be perceived either as four moving dot pairs (local) or two moving illusory squares (global). fMRI was used to measure brain activity in a slow event-related design. We observed stronger IPS and putamen responses to the stimulus when participants perceived the global interpretation compared to the local, confirming the findings of previous studies. Most importantly, we also observed that the global stimulus interpretation was preceded by an increased activity of the bilateral dorsal insula, which is known to process saliency and gate information for conscious access. Our data suggest an important role of the dorsal insula in shaping complex illusory interpretations of the sensory input.

Modulation of proper name recall by transcranial direct current stimulation of the anterior temporal lobes.

We often fail to recall another person's name. Proper names might be more difficult to memorize and retrieve than other pieces of knowledge, such as one's profession because they are processed differently in the brain. Neuroimaging and neuropsychological studies associate the bilateral anterior temporal lobes (ATL) in the retrieval of proper names and other person-related knowledge. Specifically, recalling a person's name is thought to be supported by the left ATL, whereas recalling specific information such as a person's occupation is suggested to be subserved by the right ATL. To clarify and further explore the causal relationship between both ATLs and proper name retrieval, we stimulated these regions with anodal, cathodal and sham transcranial direct current stimulation (tDCS) while the participants memorized surnames (e.g., Mr. Baker) and professions (e.g., baker) presented with a person's face. The participants were then later asked to recall the surname and the profession. Left ATL anodal stimulation resulted in higher intrusion errors for surnames than sham, whereas right ATL anodal stimulation resulted in higher overall intrusion errors, both, surnames and professions, compared to cathodal stimulation. Cathodal stimulation of the left and right ATL had no significant effect on surname and profession recall. The results indicate that the left ATL plays a role in recalling proper names. On the other hand, the specific role of the right ATL remaines to be explored.

Driver drowsiness estimation using EEG signals with a dynamical encoder-decoder modeling framework.

Drowsiness is a leading cause of accidents on the road as it negatively affects the driver's ability to safely operate a vehicle. Neural activity recorded by EEG electrodes is a widely used physiological correlate of driver drowsiness. This paper presents a novel dynamical modeling solution to estimate the instantaneous level of the driver drowsiness using EEG signals, where the PERcentage of eyelid CLOSure (PERCLOS) is employed as the ground truth of driver drowsiness. Applying our proposed modeling framework, we find neural features present in EEG data that encode PERCLOS. In the decoding phase, we use a Bayesian filtering solution to estimate the PERCLOS level over time. A data set that comprises 18 driving tests, conducted by 13 drivers, has been used to investigate the performance of the proposed framework. The modeling performance in estimation of PERCLOS provides robust and repeatable results in tests with manual and automated driving modes by an average RMSE of 0.117 (at a PERCLOS range of 0 to 1) and average High Probability Density percentage of 62.5%. We further hypothesized that there are biomarkers that encode the PERCLOS across different driving tests and participants. Using this solution, we identified possible biomarkers such as Theta and Delta powers. Results show that about 73% and 66% of the Theta and Delta powers which are selected as biomarkers are increasing as PERCLOS grows during the driving test. We argue that the proposed method is a robust and reliable solution to estimate drowsiness in real-time which opens the door in utilizing EEG-based measures in driver drowsiness detection systems.

No common factor for illusory percepts, but a link between pareidolia and delusion tendency: A test of predictive coding theory.

Predictive coding theory is an influential view of perception and cognition. It proposes that subjective experience of the sensory information results from a comparison between the sensory input and the top-down prediction about this input, the latter being critical for shaping the final perceptual outcome. The theory is able to explain a wide range of phenomena extending from sensory experiences such as visual illusions to complex pathological states such as hallucinations and psychosis. In the current study we aimed at testing the proposed connection between different phenomena explained by the predictive coding theory by measuring the manifestation of top-down predictions at progressing levels of complexity, starting from bistable visual illusions (alternating subjective experience of the same sensory input) and pareidolias (alternative meaningful interpretation of the sensory input) to self-reports of hallucinations and delusional ideations in everyday life. Examining the correlation structure of these measures in 82 adult healthy subjects revealed a positive association between pareidolia proneness and a tendency for delusional ideations, yet without any relationship to bistable illusions. These results show that only a subset of the phenomena that are explained by the predictive coding theory can be attributed to one common underlying factor. Our findings thus support the hierarchical view of predictive processing with independent top-down effects at the sensory and cognitive levels.

Saccadic and manual response time data on inhibition of return during and after a visual search.

In the present paper we present a dataset that provides data of two experiments in which we investigated the presence of Inhibition of Return (IOR) during and after a visual search. Participants either had to saccade (Experiment 1 and 2) or make a manual response (Experiment 2) to a probe during a visual search task (searching for a target letter among a set of distractors) or immediately after its completion. The data consist of the unprocessed raw data and one csv-file of the processed eye tracking data on eight (Experiment 1) and 18 (Experiment 2) participants, respectively. In total, we obtained 5,116 trials in Experiment 1 and 18,424 in Experiment 2. The data set is stored at the repository DOOR hosted by the University of Krems (https://door.donau-uni.ac.at/view/o:1014). Detailed information about the experiments and the interpretation of the data can be found in the paper "Post-search IOR: Searching for inhibition of return after search" (Höfler et al., 2019) [1].

Target processing in overt serial visual search involves the dorsal attention network: A fixation-based event-related fMRI study.

In serial visual search we shift attention successively from location to location in search for the target. Although such search has been investigated using fMRI, overt attention (i.e., eye movements) was usually neglected or discouraged. As a result, it is unclear what happens in the instant when our gaze falls upon a target as compared to a distractor. In the present experiment, we used a multiple target search task that required eye movements and employed an analysis based on fixations as events of interest to investigate differences between target and distractor processing. Twenty young healthy adults indicated the number of targets (0-3) among distractors in a 20-item display. Compared to distractor fixations, we found that target fixations gave rise to wide-spread activation in the dorsal attention system, as well as in the visual cortex. Targets that were found later during the search activated the left inferior frontal gyrus and the left supramarginal gyrus more strongly than those that were found earlier. Finally, areas associated with visual and verbal working memory showed increased activation with a larger number of targets in the display.

Dissociating Arithmetic Operations in the Parietal Cortex Using 1 Hz Repetitive Transcranial Magnetic Stimulation: The Importance of Strategy Use.

The triple-code model (TCM) of number processing suggests the involvement of distinct parietal cortex areas in arithmetic operations: the bilateral horizontal segment of the intraparietal sulcus (hIPS) for arithmetic operations that require the manipulation of numerical quantities (e.g., subtraction) and the left angular gyrus (AG) for arithmetic operations that require the retrieval of answers from long-term memory (e.g., multiplication). Although neuropsychological, neuroimaging, and brain stimulation studies suggest the dissociation of these operations into distinct parietal cortex areas, the role of strategy (online calculation vs. retrieval) is not yet fully established. In the present study, we further explored the causal involvement of the left AG for multiplication and left hIPS for subtraction using a neuronavigated repetitive transcranial magnetic stimulation (rTMS) paradigm. Stimulation sites were determined based on an fMRI experiment using the same tasks. To account for the effect of strategy, participants were asked whether they used retrieval or calculation for each individual problem. We predicted that the stimulation of the left AG would selectively disrupt the retrieval of the solution to multiplication problems. On the other hand, stimulation of the left hIPS should selectively disrupt subtraction. Our results revealed that left AG stimulation was detrimental to the retrieval and online calculation of solutions for multiplication problems, as well as, the retrieval (but not online calculation) of the solutions to subtraction problems. In contrast, left hIPS stimulation had no detrimental effect on both operations regardless of strategy.

Age-Dependent Effect of Transcranial Alternating Current Stimulation on Motor Skill Consolidation.

Transcranial alternating current stimulation (tACS) is the application of subthreshold, sinusoidal current to modulate ongoing brain rhythms related to sensory, motor and cognitive processes. Electrophysiological studies suggested that the effect of tACS applied at an alpha frequency (8-12 Hz) was state-dependent. The effects of tACS, that is, an increase in parieto-occipital electroencephalography (EEG) alpha power and magnetoencephalography (MEG) phase coherence, was only observed when the eyes were open (low alpha power) and not when the eyes were closed (high alpha power). This state-dependency of the effects of alpha tACS might extend to the aging brain characterized by general slowing and decrease in spectral power of the alpha rhythm. We additionally hypothesized that tACS will influence the motor cortex, which is involved in motor skill learning and consolidation. A group of young and old healthy adults performed a serial reaction time task (SRTT) with their right hand before and after the tACS stimulation. Each participant underwent three sessions of stimulation: sham, stimulation applied at the individual participant's alpha peak frequency or individual alpha peak frequency (iAPF; α-tACS) and stimulation with iAPF plus 2 Hz (α2-tACS) to the left motor cortex for 10 min (1.5 mA). We measured the effect of stimulation on general motor skill (GMS) and sequence-specific skill (SS) consolidation. We found that α-tACS and α2-tACS improved GMS and SS consolidation in the old group. In contrast, α-tACS minimally improved GMS consolidation but impaired SS consolidation in the young group. On the other hand, α2-tACS was detrimental to the consolidation of both skills in the young group. Our results suggest that individuals with aberrant alpha rhythm such as the elderly could benefit more from tACS stimulation, whereas for young healthy individuals with intact alpha rhythm the stimulation could be detrimental.

Impact of Priming on Effectiveness of TMS in Detecting Language-eloquent Brain Areas in Tumor Patients.

BACKGROUND AND STUDY AIMS: Language is characteristically human, and preserving it is critical when resecting tumors in language-eloquent brain areas. Navigated repetitive transcranial magnetic stimulation (nrTMS) has been used in recent years as a noninvasive technique to identify preoperatively the language-eloquent cortical areas in tumor patients. An important objective is to increase the sensitivity and specificity of nrTMS in detecting language-related areas and increase the positive correlation of its results to that of intraoperative direct cortical stimulation (DCS). Although the technical aspects of the procedure have received enormous interest, factors related to the targeted cortical area such as previous cortical history or activity have been neglected. Therefore, the present study explores the impact of previous cortical history or activity on the effectiveness of a subsequent nrTMS mapping paradigm. MATERIALS AND METHODS: Twelve right-handed patients with a left hemispheric glioma underwent presurgical nrTMS language mapping and intraoperative language mapping with DCS. nrTMS was performed using a continuous theta burst stimulation paradigm to inhibit possible language relevant areas in the vicinity of the tumor, determined anatomically or based on functional magnetic resonance imaging hotspots. The nrTMS was applied in two separate sessions. One of the sessions randomly included a priming paradigm to precondition the targeted cortical areas. RESULTS: Priming stimulation decreased the error detection of the subsequent nrTMS mapping paradigm. This effect was more robust on major types of errors such as speech arrest and hesitation. CONCLUSION: Prior cortical activity as induced by the priming stimulation has a profound impact on the responsiveness to the nrTMS mapping paradigm. Our findings further showed that metaplasticity, a type of homeostatic plastic process, could be elicited even in cortical areas affected by a growing tumor.

Language Dominance in Patients With Malformations of Cortical Development and Epilepsy.

Background: Language function may be reorganized in patients with malformations of cortical development (MCD). This prospective cohort study aimed in assessing language dominance in a large group of patients with MCD and epilepsy using functional MRI (fMRI). Methods: Sixty-eight patients (40 women) aged 10-73 years (median, 28.0; interquartile range, 19) with MCD and epilepsy underwent 1.5 T MRI and fMRI (word generation task). Single-subject image analysis was performed with statistical parametric mapping (SPM12). Language lateralization indices (LIs) were defined for statistically significantly activated voxels in Broca's and Wernicke's areas using the formula: LI = (V L - V R)/(V L + V R) × 100, where V L and V R were sets of activated voxels on the left and on the right, respectively. Language laterality was considered typical if LI was between +20 and +100 or atypical if LI was between +19 and -100. Results: fMRI signal was elicited in 55 of 68 (81%) patients. In 18 of 55 (33%) patients, language dominance was typical, and in 37 of 55 (67%) patients, atypical (in 68%, right hemispheric; in 32%, bilateral). Language dominance was not influenced by handedness, electroclinical, and imaging features. Conclusions: In this prospective study on a large group of patients with MCD and epilepsy, about two-thirds had atypical language dominance. These results may contribute to assessing risks of postsurgical language deficits and could assist in planning of "cortical mapping" with intracranial electrodes in patients who undergo presurgical assessment.

Post-search IOR: Searching for inhibition of return after search.

Previous research has indicated that Inhibition of return (IOR) supports visual search by discouraging the re-inspection of recently inspected items during search. However, it is not clear whether IOR persists after a search is completed or whether this depends on the presence of a further search in the same display. To investigate this issue, we had participants search consecutively twice in the same display (Experiment 1). Immediately after the end of the first search and after the end of the second search we probed an item which had been recently inspected or not in the previous search. The results showed that IOR as measured by the saccadic latency to the probed items was absent after the end of each of the two successive searches. In Experiment 2, we measured both saccadic latencies and manual responses in a single-search paradigm. We found that IOR during and after the search was present for saccadic responses but absent for manual responses. This suggests that IOR during and after a visual search depends on the modality of the response and the number of required searches.

ERP-study on the time course of disgust-motivated spatial avoidance.

Recently, we showed that disgusting sound cues direct spatial attention away from the location of their origin to the opposite location indicating spatial disgust avoidance (Zimmer et al., 2015, Psychophysiology; 2016, Neuroimage). However, in these studies, we had solely used an inter-stimulus interval (ISI) of 200-300 ms, leaving unclear how disgust avoidance develops over time. Studies have shown that spatial attraction due to anger persists longer when induced by auditory rather than visual stimuli. In the present ERP-study, one of two laterally presented sounds (neutral/disgust) cued a laterally presented visual target. ISIs varied from short (50-150 ms) over middle (350-450 ms) to long (650-750 ms). For disgust stimuli, response times were longer for invalidly cued targets compared to validly cued targets, reflecting disgust avoidance. There was an increased P3 amplitude for validly versus invalidly cued targets consistent with disgust avoidance. In contrast, in the neutral condition, we found evidence of inhibition of return (IOR), as we observed a reversal of the usual validity effect from short to long ISI in the behavioral data and on the P1-component. These results indicate that spatial avoidance motivated by auditory disgust persists over time, presumably enforced by emotional rather than general attentional processes.

Depth of Encoding Through Observed Gestures in Foreign Language Word Learning.

Word learning is basic to foreign language acquisition, however time consuming and not always successful. Empirical studies have shown that traditional (visual) word learning can be enhanced by gestures. The gesture benefit has been attributed to depth of encoding. Gestures can lead to depth of encoding because they trigger semantic processing and sensorimotor enrichment of the novel word. However, the neural underpinning of depth of encoding is still unclear. Here, we combined an fMRI and a behavioral study to investigate word encoding online. In the scanner, participants encoded 30 novel words of an artificial language created for experimental purposes and their translation into the subjects' native language. Participants encoded the words three times: visually, audiovisually, and by additionally observing semantically related gestures performed by an actress. Hemodynamic activity during word encoding revealed the recruitment of cortical areas involved in stimulus processing. In this study, depth of encoding can be spelt out in terms of sensorimotor brain networks that grow larger the more sensory modalities are linked to the novel word. Word retention outside the scanner documented a positive effect of gestures in a free recall test in the short term.

So pretty! The neural correlates of self-other vs familiar-other attractiveness comparisons.

Previous research has demonstrated that comparing two persons activates a frontoparietal network associated with numbers and nonsocial magnitudes. However, it is unclear whether this network is also recruited by comparisons involving the self. Self-reflection engages self-serving motivations (e.g., the maintenance of a positive self-image) and is associated with specific brain structures, such as the medial prefrontal cortex (MPFC), the anterior insula (AI) and the anterior cingulate cortex (ACC). Self-other comparisons may thus rely on distinct neural activity. To clarify this question, we used fMRI and asked female participants to compare their own attractiveness (or the attractiveness of a familiar woman) to pictures of unknown women. Participants were slower for comparisons with targets whose attractiveness was similar to their own (or their familiar other). Yet although this behavioral result resembles the distance effect reported for nonsocial magnitudes, at the brain level, it was linked to the activity of the AI, the ACC and the MPFC. The effect of distance in these regions was stronger for self-other than familiar-other comparisons. We interpret these results in relation to previous literature in social psychology and social neuroscience.