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.

The neural correlates of trustworthiness evaluations of faces and brands: Implications for behavioral and consumer neuroscience.

When we buy a product of a brand, we trust the brand to provide good quality and reliability. Therefore, trust plays a major role in consumer behavior. It is unclear, however, how trust in brands is processed in the brain and whether it is processed differently from interpersonal trust. In this study, we used fMRI to investigate the neural correlates of interpersonal and brand trust by comparing the brain activation patterns during explicit trustworthiness judgments of faces and brands. Our results showed that while there were several brain areas known to be linked to trustworthiness evaluations, such as the amygdalae, more active in trustworthiness judgments when compared to a control task (familiarity judgment) for faces, no such difference was found for brands. Complementary ROI analysis revealed that the activation of both amygdalae was strongest for faces in the trustworthiness judgments. The direct comparison of the brain activation patterns during the trustworthiness evaluations between faces and brands in this analysis showed that trustworthiness judgments of faces activated the orbitofrontal cortex, another region that was previously linked to interpersonal trust, more strongly than trustworthiness judgments of brands. Further, trustworthiness ratings of faces, but not brands, correlated with activation in the orbitofrontal cortex. Our results indicate that the amygdalae, as well as the orbitofrontal cortex, play a prominent role in interpersonal trust (faces), but not in trust for brands. It is possible that this difference is due to brands being processed as cultural objects rather than as having human-like personality characteristics.

The effects of transcranial alternating current stimulation (tACS) at individual alpha peak frequency (iAPF) on motor cortex excitability in young and elderly adults.

Transcranial alternating current stimulation (tACS) can modulate brain oscillations, cortical excitability and behaviour. In aging, the decrease in EEG alpha activity (8-12 Hz) in the parieto-occipital and mu rhythm in the motor cortex are correlated with the decline in cognitive and motor functions, respectively. Increasing alpha activity using tACS might therefore improve cognitive and motor function in the elderly. The present study explored the influence of tACS on cortical excitability in young and old healthy adults. We applied tACS at individual alpha peak frequency for 10 min (1.5 mA) to the left motor cortex. Transcranial magnetic stimulation was used to assess the changes in cortical excitability as measured by motor-evoked potentials at rest, before and after stimulation. TACS increased cortical excitability in both groups. However, our results also suggest that the mechanism behind the effects was different, as we observed an increase and decrease in intracortical inhibition in the old group and young group, respectively. Our results indicate that both groups profited similarly from the stimulation. There was no indication that tACS was more effective in conditions of low alpha power, that is, in the elderly.

Action affordances and visuo-spatial complexity in motor imagery: An fMRI study.

Imagining a complex action requires not only motor-related processing but also visuo-spatial imagery. In the current study, we examined visuo-spatial complexity and action affordances in motor imagery (MI). Using functional magnetic resonance imaging, we investigated the neural activity in MI of reach-to-grasp movements of the right hand in five conditions. Thirty participants were scanned while imagining grasping an everyday object, grasping a geometrical shape, grasping next to an everyday object, grasping next to a geometrical shape, and grasping at nothing (no object involved). We found that MI of grasping next to an object recruited the visuo-spatial cognition network including posterior parietal and premotor regions more strongly than MI of grasping an object. This indicates that grasping next to an object requires additional processing resources rendering MI more complex. MI of a grasping movement involving a familiar everyday object compared to a geometrical shape yielded stronger activation in motor-related regions, including the bilateral supplementary motor area. This activation might be due to inhibitory processes preventing motor execution of motor scripts evoked by everyday objects (action affordances). Our results indicate that visuo-spatial cognition plays a significant role in MI.

Neuronal interactions in areas of spatial attention reflect avoidance of disgust, but orienting to danger.

For survival, it is necessary to attend quickly towards dangerous objects, but to turn away from something that is disgusting. We tested whether fear and disgust sounds direct spatial attention differently. Using fMRI, a sound cue (disgust, fear or neutral) was presented to the left or right ear. The cue was followed by a visual target (a small arrow) which was located on the same (valid) or opposite (invalid) side as the cue. Participants were required to decide whether the arrow pointed up- or downwards while ignoring the sound cue. Behaviorally, responses were faster for invalid compared to valid targets when cued by disgust, whereas the opposite pattern was observed for targets after fearful and neutral sound cues. During target presentation, activity in the visual cortex and IPL increased for targets invalidly cued with disgust, but for targets validly cued with fear which indicated a general modulation of activation due to attention. For the TPJ, an interaction in the opposite direction was observed, consistent with its role in detecting targets at unattended positions and in relocating attention. As a whole our results indicate that a disgusting sound directs spatial attention away from its location, in contrast to fearful and neutral sounds.

Neural efficiency as a function of task demands.

The neural efficiency hypothesis describes the phenomenon that brighter individuals show lower brain activation than less bright individuals when working on the same cognitive tasks. The present study investigated whether the brain activation-intelligence relationship still applies when more versus less intelligent individuals perform tasks with a comparable person-specific task difficulty. In an fMRI-study, 58 persons with lower (n = 28) or respectively higher (n = 30) intelligence worked on simple and difficult inductive reasoning tasks having the same person-specific task difficulty. Consequently, less bright individuals received sample-based easy and medium tasks, whereas bright subjects received sample-based medium and difficult tasks. This design also allowed a comparison of lower versus higher intelligent individuals when working on the same tasks (i.e. sample-based medium task difficulty). In line with expectations, differences in task performance and in brain activation were only found for the subset of tasks with the same sample-based task difficulty, but not when comparing tasks with the same person-specific task difficulty. These results suggest that neural efficiency reflects an (ability-dependent) adaption of brain activation to the respective task demands.

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.

Mechanical flutter stimulation induces a lasting response in the sensorimotor cortex as revealed with BOLD fMRI.

It has been recently shown that 20 min of mechanical flutter stimulation induces lasting motor cortical excitability changes, as assessed by transcranial magnetic stimulation in relaxed hand muscles. The present functional magnetic resonance imaging (fMRI) study aims to examine if such neuromodulatory changes are reflected in the BOLD signal during a motor test. Therefore, two groups were recruited: one group receiving whole-hand flutter stimulation with a frequency of 25 Hz (FSTIM group, n = 22) and a second group receiving no stimulation (NOSTIM group, n = 22). As motor test finger-to-thumb tapping was performed to activate a wide sensorimotor network during the fMRI measurements. Three fMRI measurements were obtained with this test: before stimulation (PRE), after stimulation (POST1), and 1 h after stimulation (POST2). Three regions of interest (ROIs) were defined: primary motor area (M1), primary somatosensory area (S1), and supplementary motor area. In the absence of baseline differences between both groups, the FSTIM group showed increased movement-related brain activations compared with the NOSTIM group, both at POST1 and POST2. ROI analysis revealed increased blood-oxygenation-level-dependent (BOLD) responses within contralateral S1 (+20%) and M1 (+25%) at POST1, which lasted until POST2. These poststimulatory effects within S1 and M1 obviously reflect neuroplastic changes associated with augmented cortical excitability. These findings are of high clinical relevance, for example, to improve the treatment of stroke patients.

Individual differences in solving arithmetic word problems.

BACKGROUND: With the present functional magnetic resonance imaging (fMRI) study at 3 T, we investigated the neural correlates of visualization and verbalization during arithmetic word problem solving. In the domain of arithmetic, visualization might mean to visualize numbers and (intermediate) results while calculating, and verbalization might mean that numbers and (intermediate) results are verbally repeated during calculation. If the brain areas involved in number processing are domain-specific as assumed, that is, that the left angular gyrus (AG) shows an affinity to the verbal domain, and that the left and right intraparietal sulcus (IPS) shows an affinity to the visual domain, the activation of these areas should show a dependency on an individual's cognitive style. METHODS: 36 healthy young adults participated in the fMRI study. The participants habitual use of visualization and verbalization during solving arithmetic word problems was assessed with a short self-report assessment. During the fMRI measurement, arithmetic word problems that had to be solved by the participants were presented in an event-related design. RESULTS: We found that visualizers showed greater brain activation in brain areas involved in visual processing, and that verbalizers showed greater brain activation within the left angular gyrus. CONCLUSIONS: Our results indicate that cognitive styles or preferences play an important role in understanding brain activation. Our results confirm, that strong visualizers use mental imagery more strongly than weak visualizers during calculation. Moreover, our results suggest that the left AG shows a specific affinity to the verbal domain and subserves number processing in a modality-specific way.

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.

Stimulating creativity via the exposure to other people's ideas.

As it is the case in brainstorming, each single idea a person generates to a specific problem may stimulate new ideas or solutions in others. In this fMRI study, we investigate the effects of cognitive stimulation via the exposure to other people's ideas on the originality of generated ideas. Participants are requested to generate alternative uses of conventional everyday objects subsequent to a short cognitive stimulation intervention in which they are exposed to other ideas, which were either common or highly original. In a control condition, meaningless pseudowords are shown. Results suggest that cognitive stimulation via common or moderately creative ideas was effective in improving creativity. At the neurophysiological level, temporo-parietal brain regions (primarily right-hemispheric) turned out to be particularly sensitive to cognitive stimulation, possibly indicating that cognitive stimulation via relevant memory cues results in a state of heightened focused attention to memory that facilitates efficient retrieval and recombination of existing knowledge.

The Influence of verbalization on the pattern of cortical activation during mental arithmetic.

BACKGROUND: The aim of the present functional magnetic resonance imaging (fMRI) study at 3 T was to investigate the influence of the verbal-visual cognitive style on cerebral activation patterns during mental arithmetic. In the domain of arithmetic, a visual style might for example mean to visualize numbers and (intermediate) results, and a verbal style might mean, that numbers and (intermediate) results are verbally repeated. In this study, we investigated, first, whether verbalizers show activations in areas for language processing, and whether visualizers show activations in areas for visual processing during mental arithmetic. Some researchers have proposed that the left and right intraparietal sulcus (IPS), and the left angular gyrus (AG), two areas involved in number processing, show some domain or modality specificity. That is, verbal for the left AG, and visual for the left and right IPS. We investigated, second, whether the activation in these areas implied in number processing depended on an individual's cognitive style. METHODS: 42 young healthy adults participated in the fMRI study. The study comprised two functional sessions. In the first session, subtraction and multiplication problems were presented in an event-related design, and in the second functional session, multiplications were presented in two formats, as Arabic numerals and as written number words, in an event-related design. The individual's habitual use of visualization and verbalization during mental arithmetic was assessed by a short self-report assessment. RESULTS: We observed in both functional sessions that the use of verbalization predicts activation in brain areas associated with language (supramarginal gyrus) and auditory processing (Heschl's gyrus, Rolandic operculum). However, we found no modulation of activation in the left AG as a function of verbalization. CONCLUSIONS: Our results confirm that strong verbalizers use mental speech as a form of mental imagination more strongly than weak verbalizers. Moreover, our results suggest that the left AG has no specific affinity to the verbal domain and subserves number processing in a modality-general way.

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.

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.

Automated macrovessel artifact correction in dynamic susceptibility contrast magnetic resonance imaging using independent component analysis.

Dynamic susceptibility contrast-MRI is the most commonly used functional MRI-based method for studying changes in cerebral perfusion. However, several studies indicated a systematic overestimation of perfusion parameters compared with other imaging modalities related to the high sensitivity of dynamic susceptibility contrast-MRI for blood flow in large vessels. In this study, we therefore suggest an improved, automated, robust, and efficient method allowing for generating hemodynamic parameter maps where signal influence from large vessels is minimized. Based on independent component analysis, this fully automated approach corrects dynamic susceptibility contrast-MRI data without any user interaction, thus making a clinical applicability possible. The accuracy of the proposed method was tested in 10 patients with cerebrovascular disease. Application of our correction algorithm resulted in a significant reduction of the effect of macrovessel signal on hemodynamic parameters like the cerebral blood flow and the cerebral blood volume compared with uncorrected data. As desired, our method specifically corrected for macrovessel artifacts in cortical grey matter tissue, leaving white matter tissue parameters largely unaffected. This may increase sensitivity and reliability of detecting perfusion abnormalities in patient groups, in particular with regard to stroke and other cerebrovascular disorders.

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.