Electrophysiological (EEG) microstates during dream-like bizarre experiences in a naturalistic scenario using immersive virtual reality.

Monitoring the reality status of conscious experience is essential for a human being to interact successfully with the external world. Despite its importance for everyday functioning, reality monitoring can systematically become erroneous, for example, while dreaming or during hallucinatory experiences. To investigate brain processes associated with reality monitoring occurring online during an experience, i.e., perceptual reality monitoring, we assessed EEG microstates in healthy, young participants. In a within-subjects design, we compared the experience of reality when being confronted with dream-like bizarre elements versus realistic elements in an otherwise highly naturalistic real-world scenario in immersive virtual reality. Dream-like bizarreness induced changes in the subjective experience of reality and bizarreness, and led to an increase in the contribution of a specific microstate labelled C'. Microstate C' was related to the suspension of disbelief, i.e. the suppression of bizarre mismatches. Together with the functional interpretation of microstate C' as reported by previous studies, the findings of this study point to the importance of prefrontal meta-conscious control processes in perceptual reality monitoring.

EEG Microstate Dynamics Associated with Dream-Like Experiences During the Transition to Sleep.

Consciousness always requires some representational content; that is, one can only be conscious about something. However, the presence of conscious experience (awareness) alone does not determine whether its content is in line with the external and physical world. Dreams, apart from certain forms of hallucinations, typically consist of non-veridical percepts, which are not recognized as false, but rather considered real. This type of experiences have been described as a state of dissociation between phenomenal and reflective awareness. Interestingly, during the transition to sleep, reflective awareness seems to break down before phenomenal awareness as conscious experience does not immediately fade with reduced wakefulness but is rather characterized by the occurrence of uncontrolled thinking and perceptual images, together with a reduced ability to recognize the internal origin of the experience. Relative deactivation of the frontoparietal and preserved activity in parieto-occipital networks has been suggested to account for dream-like experiences during the transition to sleep. We tested this hypothesis by investigating subjective reports of conscious experience and large-scale brain networks using EEG microstates in 45 healthy young subjects during the transition to sleep. We observed an inverse relationship between cognitive effects and physiological activation; dream-like experiences were associated with an increased presence of a microstate with sources in the superior and middle frontal gyrus and precuneus. Additionally, the presence of a microstate associated with higher-order visual areas was decreased. The observed inverse relationship might therefore indicate a disengagement of cognitive control systems that is mediated by specific, inhibitory EEG microstates.

BizarreVR: Dream-like bizarreness in immersive virtual reality induced changes in conscious experience of reality while leaving spatial presence intact.

Differences in conscious experience of reality occur between waking, dreaming, and psychotic states. Between these states, there are systematic differences in the judgment about the reality of the experience when being confronted with bizarre breaks. However, the mechanisms underlying experience of reality in these different states are still unknown. To investigate the effect of bizarre breaks on experience of reality during the wake state, we propose a new paradigm using dream-like bizarreness and immersive virtual reality. Results showed that the realistic non-bizarre virtual environment induced high levels of reality judgment and spatial presence, whereas the confrontation with bizarre breaks induced high levels of experienced bizarreness. Moreover, experienced bizarreness significantly reduced reality judgment in both the bizarre and the realistic condition. Further, there was no effect of bizarre breaks on spatial presence. These results provide proof of concept for the new method to elicit natural bizarre experience within a realistic scenario.

Pictorial low-level features in mental images: evidence from eye fixations.

It is known that eye movements during object imagery reflect areas visited during encoding. But will eye movements also reflect pictorial low-level features of imagined stimuli? In this paper, three experiments are reported in which we investigate whether low-level properties of mental images elicit specific eye movements. Based on the conceptualization of mental images as depictive representations, we expected low-level visual features to influence eye fixations during mental imagery, in the absence of a visual input. In a first experiment, twenty-five participants performed a visual imagery task with high vs. low spatial frequency and high vs. low contrast gratings. We found that both during visual perception and during mental imagery, first fixations were more often allocated to the low spatial frequency-high contrast grating, thus showing that eye fixations were influenced not only by physical properties of visual stimuli but also by its imagined counterpart. In a second experiment, twenty-two participants imagined high contrast and low contrast stimuli that they had not encoded before. Again, participants allocated more fixations to the high contrast mental images than to the low contrast mental images. In a third experiment, we ruled out task difficulty as confounding variable. Our results reveal that low-level visual features are represented in the mind's eye and thus, they contribute to the characterization of mental images in terms of how much perceptual information is re-instantiated during mental imagery.

Usability of Two New Interactive Game Sensor-Based Hand Training Devices in Parkinson's Disease.

This pilot cross-sectional study aimed to evaluate the usability of two new interactive game sensor-based hand devices (GripAble and Smart Sensor Egg) in both healthy adults as well as in persons with Parkinson's Disease (PD). Eight healthy adults and eight persons with PD participated in this study. Besides a standardised usability measure, the state of flow after one training session and the effect of cognitive abilities on flow were evaluated. High system usability scores (SUS) were obtained both in healthy participants (72.5, IQR = 64.375-90, GripAble) as well as persons with PD (77.5, IQR = 70-80.625, GripAble; 77.5, IQR = 75-82.5, Smart Sensor Egg). Similarly, high FSSOT scores were achieved after one training session (42.5, IQR = 39.75-50, GripAble; 50, IQR = 47-50, Smart Sensor Egg; maximum score 55). Across both groups, FSSOT scores correlated significantly with SUS scores (r = 0.52, p = 0.039). Finally, MoCA did not correlate significantly with FSSOT scores (r = 0.02, p = 0.9). The present study shows high usability for both interactive game sensor-based hand training devices, for persons with PD and healthy participants.

The prioritization of visuo-spatial associations during mental imagery.

While previous research has shown that during mental imagery participants look back to areas visited during encoding it is unclear what happens when information presented during encoding is incongruent. To investigate this research question, we presented 30 participants with incongruent audio-visual associations (e.g. the image of a car paired with the sound of a cat) and later asked them to create a congruent mental representation based on the auditory cue (e.g. to create a mental representation of a cat while hearing the sound of a cat). The results revealed that participants spent more time in the areas where they previously saw the object and that incongruent audio-visual information during encoding did not appear to interfere with the generation and maintenance of mental images. This finding suggests that eye movements can be flexibly employed during mental imagery depending on the demands of the task.

Anodal High-definition Transcranial Direct Current Stimulation over the Posterior Parietal Cortex Modulates Approximate Mental Arithmetic.

The representation and processing of numerosity is a crucial cognitive capacity. Converging evidence points to the posterior parietal cortex (PPC) as primary "number" region. However, the exact role of the left and right PPC for different types of numerical and arithmetic tasks remains controversial. In this study, we used high-definition transcranial direct current stimulation (HD-tDCS) to further investigate the causal involvement of the PPC during approximative, nonsymbolic mental arithmetic. Eighteen healthy participants received three sessions of anodal HD-tDCS at 1-week intervals in counterbalanced order: left PPC, right PPC, and sham stimulation. Results showed an improved performance during online parietal HD-tDCS (vs. sham) for subtraction problems. Specifically, the general tendency to underestimate the results of subtraction problems (i.e., the "operational momentum effect") was reduced during online parietal HD-tDCS. There was no difference between left and right stimulation. This study thus provides new evidence for a causal involvement of the left and right PPC for approximate nonsymbolic arithmetic and advances the promising use of noninvasive brain stimulation in increasing cognitive functions.

Vestibular Stimulation Modulates Neural Correlates of Own-body Mental Imagery.

There is growing evidence that vestibular information is not only involved in reflexive eye movements and the control of posture but it also plays an important role in higher order cognitive processes. Previous behavioral research has shown that concomitant vestibular stimuli influence performance in tasks that involve imagined self-rotations. These results suggest that imagined and perceived body rotations share common mechanisms. However, the nature and specificity of these effects remain largely unknown. Here, we investigated the neural mechanisms underlying this vestibulocognitive interaction. Participants (n = 20) solved an imagined self-rotation task during caloric vestibular stimulation. We found robust main effects of caloric vestibular stimulation in the core region of the vestibular network, including the rolandic operculum and insula bilaterally, and of the cognitive task in parietal and frontal regions. Interestingly, we found an interaction of stimulation and task in the left inferior parietal lobe, suggesting that this region represents the modulation of imagined body rotations by vestibular input. This result provides evidence that the inferior parietal lobe plays a crucial role in the neural integration of mental and physical body rotation.

Individual differences in basic numerical skills: The role of executive functions and motor skills.

The aim of the current study was to explore individual differences in basic numerical skills in a normative sample of 151 kindergarteners (mean age = 6.45 years). Whereas previous research claims a substantial link between executive functions and basic numerical skills, motor abilities have been put forward to explain variance in numerical skills. Regarding the current study, these two assumptions have been combined, revealing interesting results. Namely, executive functions (inhibition, switching, and visuospatial working memory) were found to relate to symbolic numerical skills, and motor skills (gross and fine motor skills) showed a significant correlation to nonsymbolic numerical skills. Suggesting that motor skills and executive functions are associated with basic numerical skills could lead to potential avenues for interventions in certain disorders or disabilities such as nonverbal learning disability, developmental dyscalculia, and developmental coordination disorder.

Reinterpretation in visual imagery is possible without visual cues: a validation of previous research.

Is visual reinterpretation of bistable figures (e.g., duck/rabbit figure) in visual imagery possible? Current consensus suggests that it is in principle possible because of converging evidence of quasi-pictorial functioning of visual imagery. Yet, studies that have directly tested and found evidence for reinterpretation in visual imagery, allow for the possibility that reinterpretation was already achieved during memorization of the figure(s). One study resolved this issue, providing evidence for reinterpretation in visual imagery (Mast and Kosslyn, Cognition 86:57-70, 2002). However, participants in that study performed reinterpretations with aid of visual cues. Hence, reinterpretation was not performed with mental imagery alone. Therefore, in this study we assessed the possibility of reinterpretation without visual support. We further explored the possible role of haptic cues to assess the multimodal nature of mental imagery. Fifty-three participants were consecutively presented three to be remembered bistable 2-D figures (reinterpretable when rotated 180°), two of which were visually inspected and one was explored hapticly. After memorization of the figures, a visually bistable exemplar figure was presented to ensure understanding of the concept of visual bistability. During recall, 11 participants (out of 36; 30.6%) who did not spot bistability during memorization successfully performed reinterpretations when instructed to mentally rotate their visual image, but additional haptic cues during mental imagery did not inflate reinterpretation ability. This study validates previous findings that reinterpretation in visual imagery is possible.

Motor response specificity in perceptual learning and its release by double training.

Perceptual learning is usually feature-specific. Recently, we showed that perceptual learning is even specific for the motor response type. In a three-line bisection task, participants indicated whether the central line was offset either to the left or right by pressing a left or a right button, respectively. We found no transfer when the same participants adjusted the offset by using a computer mouse. Here, we first show that perceptual learning with mouse adjustments transfers to the untrained hand, but only for the trained adjustment condition. There was no transfer to the button press conditions, neither for the trained nor the untrained hand. Second, we show that a double training procedure enables transfer from the mouse adjustment to the button press condition. Hence, the specificity of perceptual learning to the motor response type can be overcome by double training as it is the case for visual features. Our results suggest that during perceptual learning, perceptuo-decisional signals are encoded together with the corresponding actions.

Recurrence quantification analysis of eye movements during mental imagery.

Several studies demonstrated similarities of eye fixations during mental imagery and visual perception but-to our knowledge-the temporal characteristics of eye movements during imagery have not yet been considered in detail. To fill this gap, the same data is analyzed with conventional spatial techniques such as analysis of areas of interest (AOI), ScanMatch, and MultiMatch and with recurrence quantification analysis (RQA), a new way of analyzing gaze data by tracking re-fixations and their temporal dynamics. Participants viewed and afterwards imagined three different kinds of pictures (art, faces, and landscapes) while their eye movements were recorded. While fixation locations during imagery were related to those during perception, participants returned more often to areas they had previously looked at during imagery and their scan paths were more clustered and more repetitive when compared to visual perception. Furthermore, refixations of the same area occurred sooner after initial fixation during mental imagery. The results highlight not only content-driven spatial similarities between imagery and perception but also shed light on the processes of mental imagery maintenance and interindividual differences in these processes.

Subliminal encoding and flexible retrieval of objects in scenes.

Our episodic memory stores what happened when and where in life. Episodic memory requires the rapid formation and flexible retrieval of where things are located in space. Consciousness of the encoding scene is considered crucial for episodic memory formation. Here, we question the necessity of consciousness and hypothesize that humans can form unconscious episodic memories. Participants were presented with subliminal scenes, that is, scenes invisible to the conscious mind. The scenes displayed objects at certain locations for participants to form unconscious object-in-space memories. Later, the same scenes were presented supraliminally, that is, visibly, for retrieval testing. Scenes were presented absent the objects and rotated by 90°-270° in perspective to assess the representational flexibility of unconsciously formed memories. During the test phase, participants performed a forced-choice task that required them to place an object in one of two highlighted scene locations and their eye movements were recorded. Evaluation of the eye tracking data revealed that participants remembered object locations unconsciously, irrespective of changes in viewing perspective. This effect of gaze was related to correct placements of objects in scenes, and an intuitive decision style was necessary for unconscious memories to influence intentional behavior to a significant degree. We conclude that conscious perception is not mandatory for spatial episodic memory formation.

The Influence of Alertness on the Spatial Deployment of Visual Attention is Mediated by the Excitability of the Posterior Parietal Cortices.

With a reduced level of alertness, healthy individuals typically show a rightward shift when deploying visual attention in space. The impact of alertness on the neural networks governing visuospatial attention is, however, poorly understood. By using a transcranial magnetic stimulation twin-coil approach, the present study aimed at investigating the effects of an alertness manipulation on the excitability of the left and the right posterior parietal cortices (PPCs), crucial nodes of the visuospatial attentional network. Participants' visuospatial attentional deployment was assessed with a free visual exploration task and concurrent eye tracking. Their alertness level was manipulated through the time of the day, that is, by testing chronotypically defined evening types both during their circadian on- and off-peak times. The results revealed an increased excitability of the left compared with the right PPC during low alertness. On the horizontal dimension, these results were accompanied by a significant rightward shift in the center and a bilateral narrowing in the periphery of the visual exploration field, as well as a central upward shift on the vertical dimension. The findings show that the manipulation of non-spatial attentional aspects (i.e., alertness) can affect visuospatial attentional deployment and modulate the excitability of areas subtending spatial attentional control.

The Fantasy Questionnaire: A Measure to Assess Creative and Imaginative Fantasy.

This article reports the construction and validation of a comprehensive self-report measure of fantasy. Unlike previous measures of fantasy, which focus on psychopathology, we conceive fantasy as a trait with positive connotation. Principal component analysis (N = 318) and confirmatory factor analyses (N = 345) were conducted using 2 sociodemographically diverse samples. The results provided support for a 2-factor conceptualization of the construct, with the dimensions imaginative fantasy and creative fantasy. Imaginative fantasy refers to vivid imagination and absorption in these images and daydreams. Creative fantasy refers to the activity of using fantasy to create new ideas. The trait measure showed good internal consistency, test-retest reliability, discriminant and convergent construct validity, as well as incremental validity. Moreover, in 3 behavioral studies, we put fantasy scores in relationship with behavioral data to provide further proof of validity. A comprehensive measure of fantasy can contribute to our understanding of individual differences in inner experiences, creative processes, and problem solving.

Semantic incongruity influences response caution in audio-visual integration.

Multisensory stimulus combinations trigger shorter reaction times (RTs) than individual single-modality stimuli. It has been suggested that this inter-sensory facilitation effect is found exclusively for semantically congruent stimuli, because incongruity would prevent multisensory integration. Here we provide evidence that the effect of incongruity is due to a change in response caution rather than prevention of stimulus integration. In two experiments, participants performed two-alternative forced-choice decision tasks in which they categorized auditory stimuli, visual stimuli or audio-visual stimulus pairs. The pairs were either semantically congruent (e.g. ambulance image and horn sound) or incongruent (e.g. ambulance image and bell sound). Shorter RTs and violations of the race model inequality on congruent trials are in accordance with previous studies. However, Bayesian hierarchical drift diffusion analyses contradict former co-activation-based explanations of the effects of congruency. Instead, they show that longer RTs on incongruent compared to congruent trials are most likely the result of an incongruity caution effect-more cautious response behaviour in face of semantically incongruent sensory input. Further, they show that response caution can be adjusted on a trial-by-trial basis depending on incoming information. Finally, stimulus modality influenced non-cognitive components of the response. We suggest that the combined stimulus energy from simultaneously presented stimuli reduces encoding time.

Acute peripheral vestibular deficit increases redundancy in random number generation.

Unilateral peripheral vestibular deficit leads to broad cognitive difficulties and biases in spatial orientation. More specifically, vestibular patients typically show a spatial bias toward their affected ear in the subjective visual vertical, head and trunk orientation, fall tendency, and walking trajectory. By means of a random number generation task, we set out to investigate how an acute peripheral vestibular deficit affects the mental representation of numbers in space. Furthermore, the random number generation task allowed us to test if patients with peripheral vestibular deficit show evidence of impaired executive functions while keeping the head straight and while performing active head turns. Previous research using galvanic vestibular stimulation in healthy people has shown no effects on number space, but revealed increased redundancy of the generated numbers. Other studies reported a spatial bias in number representation during active and passive head turns. In this experiment, we tested 43 patients with acute vestibular neuritis (18 patients with left-sided and 25 with right-sided vestibular deficit) and 28 age-matched healthy controls. We found no bias in number space in patients with peripheral vestibular deficit but showed increased redundancy in patients during active head turns. Patients showed worse performance in generating sequences of random numbers, which indicates a deficit in the updating component of executive functions. We argue that RNG is a promising candidate for a time- and cost-effective assessment of executive functions in patients suffering from a peripheral vestibular deficit.

Using space to represent categories: insights from gaze position.

We investigated the boundaries among imagery, memory, and perception by measuring gaze during retrieved versus imagined visual information. Eye fixations during recall were bound to the location at which a specific stimulus was encoded. However, eye position information generalized to novel objects of the same category that had not been seen before. For example, encoding an image of a dog in a specific location enhanced the likelihood of looking at the same location during subsequent mental imagery of other mammals. The results suggest that eye movements can also be launched by abstract representations of categories and not exclusively by a single episode or a specific visual exemplar.

Time in the eye of the beholder: Gaze position reveals spatial-temporal associations during encoding and memory retrieval of future and past.

Time is grounded in various ways, and previous studies point to a "mental time line" with past associated with the left, and future with the right side. In this study, we investigated whether spontaneous eye movements on a blank screen would follow a mental timeline during encoding, free recall, and recognition of past and future items. In all three stages of processing, gaze position was more rightward during future items compared to past items. Moreover, horizontal gaze position during encoding predicted horizontal gaze position during free recall and recognition. We conclude that mental time line and the stored gaze position during encoding assist memory retrieval of past versus future items. Our findings highlight the spatial nature of temporal representations.

Perceptual learning is specific beyond vision and decision making.

Perceptual learning is usually assumed to occur within sensory areas or when sensory evidence is mapped onto decisions. Subsequent procedural and motor processes, involved in most perceptual learning experiments, are thought to play no role in the learning process. Here, we show that this is not the case. Observers trained with a standard three-line bisection task and indicated the offset direction of the central line by pressing either a left or right push button. Before and after training, observers adjusted the central line of the same bisection stimulus using a computer mouse. As expected, performance improved through training. Surprisingly, learning did not transfer to the untrained mouse adjustment condition. The same was true for the opposite, i.e., training with mouse adjustments did not transfer to the push button condition. We found partial transfer when observers adjusted the central line with two different adjustment procedures. We suggest that perceptual learning is specific to procedural motor aspects beyond visual processing. Our results support theories were visual stimuli are coded together with their corresponding actions.