The angular gyrus serves as an interface between the non-lexical reading network and the semantic system: evidence from dynamic causal modeling.

Understanding encoded language, such as written words, requires multiple cognitive processes that act in a parallel and interactive fashion. These processes and their interactions, however, are not fully understood. Various conceptual and methodical approaches including computational modeling and neuroimaging have been applied to better understand the neural underpinnings of these complex processes in the human brain. In this study, we tested different predictions of cortical interactions that derived from computational models for reading using dynamic causal modeling. Morse code was used as a model for non-lexical decoding followed by a lexical-decision during a functional magnetic resonance examination. Our results suggest that individual letters are first converted into phonemes within the left supramarginal gyrus, followed by a phoneme assembly to reconstruct word phonology, involving the left inferior frontal cortex. To allow the identification and comprehension of known words, the inferior frontal cortex then interacts with the semantic system via the left angular gyrus. As such, the left angular gyrus is likely to host phonological and semantic representations and serves as a bidirectional interface between the networks involved in language perception and word comprehension.

Temporal dynamics of oscillatory activity during nonlexical language decoding: Evidence from Morse code and magnetoencephalography.

Understanding encoded languages, such as written script or Morse code, requires nonlexical and lexical processing components that act in a parallel and interactive fashion. Decoding written script-as for example in reading-is typically very fast, making the investigation of the lexical and nonlexical components and their underlying neural mechanisms challenging. In the current study, we aimed to accomplish this problem by using Morse code as a model for language decoding. The decoding of Morse code is slower and thus allows a better and more fine-grained investigation of the lexical and nonlexical components of language decoding. In the current study, we investigated the impact of various components of nonlexical decoding of Morse code using magnetoencephalography. For this purpose, we reconstructed the time-frequency responses below 40 Hz in brain regions significantly involved in Morse code decoding and word comprehension that were identified in a previous study. Event-related reduction in beta- and alpha-band power were found in left inferior frontal cortex and angular gyrus, respectively, while event-related theta-band power increase was found at frontal midline. These induced oscillations reflect working-memory encoding, long-term memory retrieval as well as demanding cognitive control, respectively. In sum, by using Morse code and MEG, we were able to identify a cortical network underlying language decoding in a time- and frequency-resolved manner.

Saccade suppression of displacements, but not of contrast, depends on context.

Saccades let the visual scene sweep with high speed across the retina, thus producing a massive motion stimulus. Yet, in natural vision, we never perceive motion that is produced by saccades. The absence of perisaccadic motion perception might be caused by a transient reduction of visual sensitivity at the time of saccade initiation, so-called saccadic suppression. Saccade suppression occurs for contrast, displacement, and motion stimuli. Saccade suppression of displacements has been shown to be context sensitive. After performing saccades in sessions without perisaccadic stimulation, saccade suppression magnitude is drastically decreased (Zimmermann, 2020). Here, we aimed to test whether saccade suppression of contrast is similarly modulated by context. To this end, we projected stimuli on a homogeneously white wall such that we could establish a ganzfeld-like environment that, depending on the experimental session, did or did not contain any visible contrast stimuli. We first successfully replicated the context sensitivity of saccade suppression of displacements. Then, we tested context sensitivity of contrast suppression by asking subjects to perform several saccades either across the uniform white wall or across a background consisting of a sinusoidal grating. In contrast to perisaccadic context sensitivity for displacement suppression, we did not find context sensitivity for suppression of contrast.

Human movements do not look the same in a tilted world: Gravitational constraints influence the perception of biological motion.

We investigated whether gravitational constraints influence the interaction of visual, proprioceptive and vestibular cues for Biological Motion Perception (BMP). Participants were asked to distinguish between plausible and random point-light movements, while passively placed in either an upright or a tilted body orientation. Manipulating the body orientation with respect to gravity leads to different gravitational signals transmitted by the visual, proprioceptive, and vestibular systems. Participants were overall faster in distinguishing plausible point-light movements than random movements. Critically, response times for biologically plausible point-light movements - but not for random movements - were significantly prolonged in the tilted body orientation. Our results suggest that BMP depends not only on the spatial-temporal cues embedded in point-light movements but also rely on the congruency between current gravitational signals detected by the sensory systems and our previous knowledge of terrestrial gravity. STATEMENT OF RELEVANCE: As humankind is preparing for a new space age, understanding how gravity influences behaviour and cognition has never been more pressing. All living organisms have evolved to survive in a terrestrial gravitational field. Although we cannot consciously feel gravity, it has an impact in our life: it affects how we move and interact with the external environment. The sensory signals from the vestibular system are continuously combined with visual and proprioceptive cues to help us in maintaining a stable representation of the world. Here we placed participants in a tilted body orientation and were able to determine that a conflict between prior gravitational knowledge and what was actively sensed about gravity affected human Biological Movement Perception. Humans suffer changes in perception under non-terrestrial gravity conditions that may potentially compromise performance during space exploration.

10 Hz tACS Over Somatosensory Cortex Does Not Modulate Supra-Threshold Tactile Temporal Discrimination in Humans.

Perception of physical identical stimuli can differ over time depending on the brain state. One marker of this brain state can be neuronal oscillations in the alpha band (8-12 Hz). A previous study showed that the power of prestimulus alpha oscillations in the contralateral somatosensory area negatively correlate with the ability to temporally discriminate between two subsequent tactile suprathreshold stimuli. That is, with high alpha power subjects were impaired in discriminating two stimuli and more frequently reported to perceive only one stimulus. While this previous study found correlative evidence for a role of alpha oscillations on tactile temporal discrimination, here, we aimed to study the causal influence of alpha power on tactile temporal discrimination by using transcranial alternating current stimulation (tACS). We hypothesized that tACS in the alpha frequency should entrain alpha oscillations and thus modulate alpha power. This modulated alpha power should alter temporal discrimination ability compared to a control frequency or sham. To this end, 17 subjects received one or two electrical stimuli to their left index finger with different stimulus onset asynchronies (SOAs). They reported whether they perceived one or two stimuli. Subjects performed the paradigm before (pre), during (peri), and 25 min after tACS (post). tACS was applied to the contralateral somatosensory-parietal area with either 10, 5 Hz or sham on three different days. We found no significant difference in discrimination abilities between 10 Hz tACS and the control conditions, independent of SOAs. In addition to choosing all SOAs as the independent variable, we chose individually different SOAs, for which we expected the strongest effects of tACS. Again, we found no significant effects of 10 Hz tACS on temporal discrimination abilities. We discuss potential reasons for the inability to modulate tactile temporal discrimination abilities with tACS.

Impaired Tactile Temporal Discrimination in Patients With Hepatic Encephalopathy.

The sensory system constantly receives stimuli from the external world. To discriminate two stimuli correctly as two temporally distinct events, the temporal distance or stimulus onset asynchrony (SOA) between the two stimuli has to exceed a specific threshold. If the SOA between two stimuli is shorter than this specific threshold, the two stimuli will be perceptually fused and perceived as one single stimulus. Patients with hepatic encephalopathy (HE) are known to show manifold perceptual impairments, including slowed visual temporal discrimination abilities as measured by the critical flicker frequency (CFF). Here, we hypothesized that HE patients are also impaired in their tactile temporal discrimination abilities and, thus, require a longer SOA between two tactile stimuli to perceive the stimuli as two temporally distinct events. To test this hypothesis, patients with varying grades of HE and age-matched healthy individuals performed a tactile temporal discrimination task. All participants received two tactile stimuli with varying SOA applied to their left index finger and reported how many distinct stimuli they perceived ("1" vs. "2"). HE patients needed a significantly longer SOA (138.0 ± 11.3 ms) between two tactile stimuli to perceive the stimuli as two temporally distinct events than healthy controls (78.6 ± 13.1 ms; p < 0.01). In addition, we found that the temporal discrimination ability in the tactile modality correlated positively with the temporal discrimination ability in the visual domain across all participants (i.e., negative correlation between tactile SOA and visual CFF: r = -0.37, p = 0.033). Our findings provide evidence that temporal tactile perception is substantially impaired in HE patients. In addition, the results suggest that tactile and visual discrimination abilities are affected in HE in parallel. This finding might argue for a common underlying pathophysiological mechanism. We argue that the known global slowing of neuronal oscillations in HE might represent such a common mechanism.

Connecting occipital alpha band peak frequency, visual temporal resolution, and occipital GABA levels in healthy participants and hepatic encephalopathy patients.

Recent studies have proposed a connection between the individual alpha band peak frequency and the temporal resolution of visual perception in healthy human participants. This connection rests on animal studies describing oscillations in the alpha band as a mode of phasic thalamocortical information transfer for low-level visual stimuli, which critically relies on GABAergic interneurons. Here, we investigated the interplay of these parameters by measuring occipital alpha band peak frequency by means of magnetoencephalography, visual temporal resolution by means of behavioral testing, and occipital GABA levels by means of magnetic resonance spectroscopy. Importantly, we investigated a sample of healthy participants and patients with varying grades of hepatic encephalopathy, which are known to exhibit decreases in the investigated parameters, thus providing an increased parameter space. We found that occipital alpha band peak frequency and visual temporal resolution were positively correlated, i.e., higher occipital alpha band peak frequencies were on average related to a higher temporal resolution. Likewise, occipital alpha band peak frequency correlated positively with occipital GABA levels. However, correlations were significant only when both healthy participants and patients were included in the analysis, thereby indicating a connection of the measures on group level (instead of the individual level). These findings provide new insights into neurophysiological and neurochemical underpinnings of visual perception.

A new device for high-temperature in situ GISAXS measurements.

A heating stage originally designed for diffraction experiments is implemented into a Bruker NANOSTAR instrument for in situ grazing incidence small-angle x-ray scattering experiments. A controlled atmosphere is provided by a dome separating the sample environment from the evacuated scattering instrument. This dome is double shelled in order to enable cooling water to flow through it. A mesoporous silica film templated by a self-assembled block copolymer system is investigated in situ during step-wise heating in air. The GISAXS pattern shows the structural development of the ordered lattice of parallel cylindrical pores. The deformation of the elliptical pore-cross section perpendicular to the film surface was studied with increasing temperature. Moreover, the performance of the setup was tested by controlled in situ heating of a copper surface under controlled oxygen containing atmosphere.

Event-related desynchronization of mu and beta oscillations during the processing of novel tool names.

According to the embodied cognition framework, the formation of conceptual representations integrates the type of experience during learning. In this electroencephalographic study, we applied a linguistic variant of a training paradigm, in which participants learned to associate novel names to novel tools while either manipulating or visually exploring them. The analysis focused on event-related desynchronization (ERD) of oscillations in the mu and beta frequency range, which reflects activation of sensorimotor brain areas. After three training sessions, processing names of manipulated tools elicited a stronger ERD of the beta (18-25 Hz, 140-260 ms) and the lower mu rhythm (8-10 Hz, 320-440 ms) than processing names of visually explored tools, reflecting a possible reactivation of experiential sensorimotor information. Given the unexpected result that familiarized pseudo-words elicited an ERD comparable to names of manipulated tools, our findings could reflect a suppression of sensorimotor activity during the processing of objects with exclusively visual features.

U-shaped Relation between Prestimulus Alpha-band and Poststimulus Gamma-band Power in Temporal Tactile Perception in the Human Somatosensory Cortex.

Neuronal oscillations are a ubiquitous phenomenon in the human nervous system. Alpha-band oscillations (8-12 Hz) have been shown to correlate negatively with attention and performance, whereas gamma-band oscillations (40-150 Hz) correlate positively. Here, we studied the relation between prestimulus alpha-band power and poststimulus gamma-band power in a suprathreshold tactile discrimination task. Participants received two electrical stimuli to their left index finger with different SOAs (0 msec, 100 msec, intermediate SOA, intermediate SOA ± 10 msec). The intermediate SOA was individually determined so that stimulation was bistable, and participants perceived one stimulus in half of the trials and two stimuli in the other half. We measured neuronal activity with magnetoencephalography (MEG). In trials with intermediate SOAs, behavioral performance correlated inversely with prestimulus alpha-band power but did not correlate with poststimulus gamma-band power. Poststimulus gamma-band power was high in trials with low and high prestimulus alpha-band power and low for intermediate prestimulus alpha-band power (i.e., U-shaped). We suggest that prestimulus alpha activity modulates poststimulus gamma activity and subsequent perception: (1) low prestimulus alpha-band power leads to high poststimulus gamma-band power, biasing perception such that two stimuli were perceived; (2) intermediate prestimulus alpha-band power leads to low gamma-band power (interpreted as inefficient stimulus processing), consequently, perception was not biased in either direction; and (3) high prestimulus alpha-band power leads to high poststimulus gamma-band power, biasing perception such that only one stimulus was perceived.

Rapid temporal recalibration to visuo-tactile stimuli.

For a comprehensive understanding of the environment, the brain must constantly decide whether the incoming information originates from the same source and needs to be integrated into a coherent percept. This integration process is believed to be mediated by temporal integration windows. If presented with temporally asynchronous stimuli for a few minutes, the brain adapts to this new temporal relation by recalibrating the temporal integration windows. Such recalibration can occur even more rapidly after exposure to just a single trial of asynchronous stimulation. While rapid recalibration has been demonstrated for audio-visual stimuli, evidence for rapid recalibration of visuo-tactile stimuli is lacking. Here, we investigated rapid recalibration in the visuo-tactile domain. Subjects received visual and tactile stimuli with different stimulus onset asynchronies (SOA) and were asked to report whether the visuo-tactile stimuli were presented simultaneously. Our results demonstrate visuo-tactile rapid recalibration by revealing that subjects' simultaneity reports were modulated by the temporal order of stimulation in the preceding trial. This rapid recalibration effect, however, was only significant if the SOA in the preceding trial was smaller than 100 ms, while rapid recalibration could not be demonstrated for SOAs larger than 100 ms. Since rapid recalibration in the audio-visual domain has been demonstrated for SOAs larger than 100 ms, we propose that visuo-tactile recalibration works at shorter SOAs, and thus faster time scales than audio-visual rapid recalibration.

Beyond the Peak - Tactile Temporal Discrimination Does Not Correlate with Individual Peak Frequencies in Somatosensory Cortex.

The human sensory systems constantly receive input from different stimuli. Whether these stimuli are integrated into a coherent percept or segregated and perceived as separate events, is critically determined by the temporal distance of the stimuli. This temporal distance has prompted the concept of temporal integration windows or perceptual cycles. Although this concept has gained considerable support, the neuronal correlates are still discussed. Studies suggested that neuronal oscillations might provide a neuronal basis for such perceptual cycles, i.e., the cycle lengths of alpha oscillations in visual cortex and beta oscillations in somatosensory cortex might determine the length of perceptual cycles. Specifically, recent studies reported that the peak frequency (the frequency with the highest spectral power) of alpha oscillations in visual cortex correlates with subjects' ability to discriminate two visual stimuli. In the present study, we investigated whether peak frequencies in somatosensory cortex might serve as the correlate of perceptual cycles in tactile discrimination. Despite several different approaches, we were unable to find a significant correlation between individual peak frequencies in the alpha- and beta-band and individual discrimination abilities. In addition, analysis of Bayes factor provided evidence that peak frequencies and discrimination thresholds are unrelated. The results suggest that perceptual cycles in the somatosensory domain are not necessarily to be found in the peak frequency, but in other frequencies. We argue that studies based solely on analysis of peak frequencies might thus miss relevant information.

Subliminal stimuli modulate somatosensory perception rhythmically and provide evidence for discrete perception.

Despite being experienced as continuous, there is an ongoing debate if perception is an intrinsically discrete process, with incoming sensory information treated as a succession of single perceptual cycles. Here, we provide causal evidence that somatosensory perception is composed of discrete perceptual cycles. We used in humans an electrotactile temporal discrimination task preceded by a subliminal (i.e., below perceptual threshold) stimulus. Although not consciously perceived, subliminal stimuli are known to elicit neuronal activity in early sensory areas and modulate the phase of ongoing neuronal oscillations. We hypothesized that the subliminal stimulus indirectly, but systematically modulates the ongoing oscillatory phase in S1, thereby rhythmically shaping perception. The present results confirm that, without being consciously perceived, the subliminal stimulus critically influenced perception in the discrimination task. Importantly, perception was modulated rhythmically, in cycles corresponding to the beta-band (13-18 Hz). This can be compellingly explained by a model of discrete perceptual cycles.

Impaired perception of human movements in Parkinson's disease.

Interacting with other individuals in a social world requires fast and accurate perception of other individuals' identity, actions, or intentions. Humans are very efficient in these social tasks, as they can extract social information even if the actor is represented only by a handful of point-lights on an otherwise invisible body. Theories have argued that efficient visual perception of actions is based on intact motor system functioning. The motor system provides visuo-motor action representations shaped by the observer's own movements or motor repertoire. If the observer's motor repertoire is impaired, this should lead to impaired visuo-motor representations and ultimately to impaired visual perception of movements. Here we tested this hypothesis in a behavioral study with patients suffering from Parkinson's disease (PD). PD patients are typically impaired in movement execution. We tested these patients and a matched control group in a visual discrimination task on human movement perception. The results showed that PD patients were significantly impaired in the perception of human movements. This impairment was most prominent for transitive (object-related) movements. The results indicate that impaired movement execution critically influences movement perception. The results support the hypothesis that the motor system plays a causal role for the visual perception of human movements.

Beta Peak Frequencies at Rest Correlate with Endogenous GABA+/Cr Concentrations in Sensorimotor Cortex Areas.

Neuronal oscillatory activity in the beta band (15-30 Hz) is a prominent signal within the human sensorimotor cortex. Computational modeling and pharmacological modulation studies suggest an influence of GABAergic interneurons on the generation of beta band oscillations. Accordingly, studies in humans have demonstrated a correlation between GABA concentrations and power of beta band oscillations. It remains unclear, however, if GABA concentrations also influence beta peak frequencies and whether this influence is present in the sensorimotor cortex at rest and without pharmacological modulation. In the present study, we investigated the relation between endogenous GABA concentration (measured by magnetic resonance spectroscopy) and beta oscillations (measured by magnetoencephalography) at rest in humans. GABA concentrations and beta band oscillations were measured for left and right sensorimotor and occipital cortex areas. A significant positive linear correlation between GABA concentration and beta peak frequency was found for the left sensorimotor cortex, whereas no significant correlations were found for the right sensorimotor and the occipital cortex. The results show a novel connection between endogenous GABA concentration and beta peak frequency at rest. This finding supports previous results that demonstrated a connection between oscillatory beta activity and pharmacologically modulated GABA concentration in the sensorimotor cortex. Furthermore, the results demonstrate that for a predominantly right-handed sample, the correlation between beta band oscillations and endogenous GABA concentrations is evident only in the left sensorimotor cortex.

Prestimulus Alpha Power Influences Tactile Temporal Perceptual Discrimination and Confidence in Decisions.

Recent studies have demonstrated that prestimulus alpha-band activity substantially influences perception of near-threshold stimuli. Here, we studied the influence of prestimulus alpha power fluctuations on temporal perceptual discrimination of suprathreshold tactile stimuli and subjects' confidence regarding their perceptual decisions. We investigated how prestimulus alpha-band power influences poststimulus decision-making variables. We presented electrical stimuli with different stimulus onset asynchronies (SOAs) to human subjects, and determined the SOA for which temporal perceptual discrimination varied on a trial-by-trial basis between perceiving 1 or 2 stimuli, prior to recording brain activity with magnetoencephalography. We found that low prestimulus alpha power in contralateral somatosensory and occipital areas predicts the veridical temporal perceptual discrimination of 2 stimuli. Additionally, prestimulus alpha power was negatively correlated with confidence ratings in correctly perceived trials, but positively correlated for incorrectly perceived trials. Finally, poststimulus event-related fields (ERFs) were modulated by prestimulus alpha power and reflect the result of a decisional process rather than physical stimulus parameters around ∼150 ms. These findings provide new insights into the link between spontaneous prestimulus alpha power fluctuations, temporal perceptual discrimination, decision making, and decisional confidence. The results suggest that prestimulus alpha power modulates perception and decisions on a continuous scale, as reflected in confidence ratings.

Beta oscillations define discrete perceptual cycles in the somatosensory domain.

Whether seeing a movie, listening to a song, or feeling a breeze on the skin, we coherently experience these stimuli as continuous, seamless percepts. However, there are rare perceptual phenomena that argue against continuous perception but, instead, suggest discrete processing of sensory input. Empirical evidence supporting such a discrete mechanism, however, remains scarce and comes entirely from the visual domain. Here, we demonstrate compelling evidence for discrete perceptual sampling in the somatosensory domain. Using magnetoencephalography (MEG) and a tactile temporal discrimination task in humans, we find that oscillatory alpha- and low beta-band (8-20 Hz) cycles in primary somatosensory cortex represent neurophysiological correlates of discrete perceptual cycles. Our results agree with several theoretical concepts of discrete perceptual sampling and empirical evidence of perceptual cycles in the visual domain. Critically, these results show that discrete perceptual cycles are not domain-specific, and thus restricted to the visual domain, but extend to the somatosensory domain.

Lateralized modulation of beta-band power in sensorimotor areas during action observation.

The cortical network for action observation includes areas of the visual cortex and non-visual areas, including areas of the motoric system. Parts of this network are known for their contralateral organization during motion execution, i.e., they predominantly control and respond to movements of the contralateral body side. We were interested whether this lateralized organization was also present during action observation. Human participants viewed point-light displays of human actors, where the actor was facing and moving either to the right or to the left, while participants' neuromagnetic activity was recorded using magnetoencephalography (MEG). We found that right and left facing movements elicited different activity in left and right motoric areas. This lateralization effect was found in two distinct spatio-temporal-spectral clusters: An early lateralization effect in medial sensors at 12-16 Hz and ~276-675 ms after stimulus onset, and a second cluster in more lateral sensors at 22-28 Hz and ~1275-1775 ms. Our results demonstrate that in addition to the known somatotopic organization of parts of the human motoric system, these areas also show a lateralization effect during action observation. Thus, our results indicate that the hemispheric organization of one's own body map known for motion execution extends to the visual observation of others' bodily actions and movements.

The role of alpha oscillations for illusory perception.

Alpha oscillations are a prominent electrophysiological signal measured across a wide range of species and cortical and subcortical sites. Alpha oscillations have been viewed for a long time as an "idling" rhythm, purely reflecting inactive sites. Despite earlier evidence from neurophysiology, awareness that alpha oscillations can substantially influence perception and behavior has grown only recently in cognitive neuroscience. Evidence for an active role of alpha for perception comes mainly from several visual, near-threshold experiments. In the current review, we extend this view by summarizing studies showing how alpha-defined brain states relate to illusory perception, i.e. cases of perceptual reports that are not "objectively" verifiable by distinct stimuli or stimulus features. These studies demonstrate that ongoing or prestimulus alpha oscillations substantially influence the perception of auditory, visual or multisensory illusions.

Distinct spatio-temporal profiles of beta-oscillations within visual and sensorimotor areas during action recognition as revealed by MEG.

The neural correlates of action recognition have been widely studied in visual and sensorimotor areas of the human brain. However, the role of neuronal oscillations involved during the process of action recognition remains unclear. Here, we were interested in how the plausibility of an action modulates neuronal oscillations in visual and sensorimotor areas. Subjects viewed point-light displays (PLDs) of biomechanically plausible and implausible versions of the same actions. Using magnetoencephalography (MEG), we examined dynamic changes of oscillatory activity during these action recognition processes. While both actions elicited oscillatory activity in visual and sensorimotor areas in several frequency bands, a significant difference was confined to the beta-band (∼20 Hz). An increase of power for plausible actions was observed in left temporal, parieto-occipital and sensorimotor areas of the brain, in the beta-band in successive order between 1650 and 2650 msec. These distinct spatio-temporal beta-band profiles suggest that the action recognition process is modulated by the degree of biomechanical plausibility of the action, and that spectral power in the beta-band may provide a functional interaction between visual and sensorimotor areas in humans.