Construct Validity and Applicability of a Team-Sport-Specific Change of Direction Test.

Cuts and changes of direction (COD) are frequent movements during games in team sports. Since those movements are seen as a key performance variable, COD assessments are included in performance diagnostics. However, some tests are criticized as they seem to be confounded by variables such as linear sprinting. Therefore, it is suggested that not only total COD time should be assessed, but also the athletes' COD movements should be examined more closely. For example, split times could be analyzed in tests with more than one COD like the Team-Sport-Specific COD (TSS-COD) test. We aimed to investigate the construct validity of the TSS-COD test, focusing on the homogeneity of the different test parts. We also tested how far sprint performance mapped onto COD performance. Test data were analyzed from 154 elite male and female volleyball and basketball athletes. A Fitlight© System was used to assess duration of the TSS-COD test. For the sprint tests, magnetic gates (Humotion GmbH) were used to measure sprint time. Explorative principal component analysis (PCA) was conducted including the test interval duration and the athletes' 5, 10, and 20 m sprint performance, to test the validity of the TSS-COD test. PCA results showed that the start interval formed a factor separate from the other COD sub-intervals. In addition, sprint performance was separated from all COD interval measures. The findings of the PCA were confirmed by split-half validation. Since sprint and COD performance represent independent performance domains within this analysis, we suggest the TSS-COD test to be a valid test to assess COD performance.

Surmising synchrony of sound and sight: Factors explaining variance of audiovisual integration in hurdling, tap dancing and drumming.

Auditory and visual percepts are integrated even when they are not perfectly temporally aligned with each other, especially when the visual signal precedes the auditory signal. This window of temporal integration for asynchronous audiovisual stimuli is relatively well examined in the case of speech, while other natural action-induced sounds have been widely neglected. Here, we studied the detection of audiovisual asynchrony in three different whole-body actions with natural action-induced sounds-hurdling, tap dancing and drumming. In Study 1, we examined whether audiovisual asynchrony detection, assessed by a simultaneity judgment task, differs as a function of sound production intentionality. Based on previous findings, we expected that auditory and visual signals should be integrated over a wider temporal window for actions creating sounds intentionally (tap dancing), compared to actions creating sounds incidentally (hurdling). While percentages of perceived synchrony differed in the expected way, we identified two further factors, namely high event density and low rhythmicity, to induce higher synchrony ratings as well. Therefore, we systematically varied event density and rhythmicity in Study 2, this time using drumming stimuli to exert full control over these variables, and the same simultaneity judgment tasks. Results suggest that high event density leads to a bias to integrate rather than segregate auditory and visual signals, even at relatively large asynchronies. Rhythmicity had a similar, albeit weaker effect, when event density was low. Our findings demonstrate that shorter asynchronies and visual-first asynchronies lead to higher synchrony ratings of whole-body action, pointing to clear parallels with audiovisual integration in speech perception. Overconfidence in the naturally expected, that is, synchrony of sound and sight, was stronger for intentional (vs. incidental) sound production and for movements with high (vs. low) rhythmicity, presumably because both encourage predictive processes. In contrast, high event density appears to increase synchronicity judgments simply because it makes the detection of audiovisual asynchrony more difficult. More studies using real-life audiovisual stimuli with varying event densities and rhythmicities are needed to fully uncover the general mechanisms of audiovisual integration.

Being right matters: Model-compliant events in predictive processing.

While prediction errors (PE) have been established to drive learning through adaptation of internal models, the role of model-compliant events in predictive processing is less clear. Checkpoints (CP) were recently introduced as points in time where expected sensory input resolved ambiguity regarding the validity of the internal model. Conceivably, these events serve as on-line reference points for model evaluation, particularly in uncertain contexts. Evidence from fMRI has shown functional similarities of CP and PE to be independent of event-related surprise, raising the important question of how these event classes relate to one another. Consequently, the aim of the present study was to characterise the functional relationship of checkpoints and prediction errors in a serial pattern detection task using electroencephalography (EEG). Specifically, we first hypothesised a joint P3b component of both event classes to index recourse to the internal model (compared to non-informative standards, STD). Second, we assumed the mismatch signal of PE to be reflected in an N400 component when compared to CP. Event-related findings supported these hypotheses. We suggest that while model adaptation is instigated by prediction errors, checkpoints are similarly used for model evaluation. Intriguingly, behavioural subgroup analyses showed that the exploitation of potentially informative reference points may depend on initial cue learning: Strict reliance on cue-based predictions may result in less attentive processing of these reference points, thus impeding upregulation of response gain that would prompt flexible model adaptation. Overall, present results highlight the role of checkpoints as model-compliant, informative reference points and stimulate important research questions about their processing as function of learning und uncertainty.

The Relationship between Expertise in Sports, Visuospatial, and Basic Cognitive Skills.

Team sports place high demands on visuospatial and other cognitive skills. However, there is a lack of research on visuospatial skills of elite athletes and there are heterogeneous results on basic cognitive skills of this population. Therefore, this series of studies tested different cognitive skills in elite team sports athletes. In Experiment 1, elite athletes were compared to recreational athletes, but no differences were observed between the groups in choice response time (CRT) and mental rotation (MR). To see if differences could be observed when the tested groups had a greater difference in expertise and more representative stimuli, in Experiment 2, we tested CRT and MR of elite athletes who had higher level of expertise, and we also used three-dimensional human stimuli. Overall, we still found no differences in MR; however, elite athletes did have shorter CRTs. In Experiment 3, instead of testing MR, we compared elite athletes' and recreational athletes' basic cognitive skills, such as processing speed, letter readout speed, memory span, and sustained attention. We found that elite athletes only performed better in sustained attention. Building on this data, in a supplementary analysis (Experiment 4) we tested whether MR and CRTs are correlated with basic cognitive skills. Results show that processing speed is the best predictor for MR, whereas letter readout speed explains most of the variance in CRTs. Finally, we discuss these findings against the backdrop of expertise and offer implications for future studies on mental rotation.

Neural Anatomy of Primary Visual Cortex Limits Visual Working Memory.

Despite the immense processing power of the human brain, working memory storage is severely limited, and the neuroanatomical basis of these limitations has remained elusive. Here, we show that the stable storage limits of visual working memory for over 9 s are bound by the precise gray matter volume of primary visual cortex (V1), defined by fMRI retinotopic mapping. Individuals with a bigger V1 tended to have greater visual working memory storage. This relationship was present independently for both surface size and thickness of V1 but absent in V2, V3 and for non-visual working memory measures. Additional whole-brain analyses confirmed the specificity of the relationship to V1. Our findings indicate that the size of primary visual cortex plays a critical role in limiting what we can hold in mind, acting like a gatekeeper in constraining the richness of working mental function.

V1 surface size predicts GABA concentration in medial occipital cortex.

A number of recent studies have established a link between behavior and the anatomy of the primary visual cortex (V1). However, one often-raised criticism has been that these studies provide little insight into the mechanisms of the observed relationships. As inhibitory neural interactions have been postulated as an important mechanism for those behaviors related to V1 anatomy, we measured the concentration of inhibitory gamma-amino butyric acid (GABA) in the medial occipital cortex where V1 is located using magnetic resonance spectroscopy (MRS) and estimated the surface area of V1 using fMRI retinotopic mapping. We found a significant positive relationship between GABA concentration and V1 surface area. This relationship was present irrespective of whether the MRS voxel had a fixed size across participants or was proportionally sized to each individual's V1 surface area. Hence, individuals with a larger V1 had a higher GABA concentration in the medial occipital cortex. By tying together V1 size and GABA concentration, our findings point towards individual differences in the level of neural inhibition that might partially mediate the relationships between behavior and V1 neuroanatomy. In addition, they illustrate how stable microscopic properties of neural activity and function are reflected in macro-measures of V1 structure.

Smaller Primary Visual Cortex Is Associated with Stronger, but Less Precise Mental Imagery.

Despite mental imagery's ubiquitous role in human perception, cognition and behavior, one standout question remains unanswered: Why does imagery vary so much from one individual to the next? Here, we used a behavioral paradigm that measures the functional impact of a mental image on subsequent conscious perception and related these measures to the anatomy of the early visual cortex estimated by fMRI retinotopic mapping. We observed a negative relationship between primary visual cortex (V1) surface area and sensory imagery strength, but found positive relationships between V1 and imagery precision (spatial location and orientation). Hence, individuals with a smaller V1 tended to have stronger, but less precise imagery. In addition, subjective vividness of imagery was positively related to prefrontal cortex volume, but unrelated to V1 anatomy. Our findings present the first evidence for the importance of the V1 layout in shaping the strength of human imagination.

Functional Connectivity Patterns of Visual Cortex Reflect its Anatomical Organization.

The brain is continuously active, even without external input or task demands. This so-called resting-state activity exhibits a highly specific spatio-temporal organization. However, how exactly these activity patterns map onto the anatomical and functional architecture of the brain is still unclear. We addressed this question in the human visual cortex. We determined the representation of the visual field in visual cortical areas of 44 subjects using fMRI and examined resting-state correlations between these areas along the visual hierarchy, their dorsal and ventral segments, and between subregions representing foveal versus peripheral parts of the visual field. We found that retinotopically corresponding regions, particularly those representing peripheral visual fields, exhibit strong correlations. V1 displayed strong internal correlations between its dorsal and ventral segments and the highest correlation with LGN compared with other visual areas. In contrast, V2 and V3 showed weaker correlations with LGN and stronger between-area correlations, as well as with V4 and hMT+. Interhemispheric correlations between homologous areas were especially strong. These correlation patterns were robust over time and only marginally altered under task conditions. These results indicate that resting-state fMRI activity closely reflects the anatomical organization of the visual cortex both with respect to retinotopy and hierarchy.

Population inversion in monolayer and bilayer graphene.

The recent demonstration of saturable absorption and negative optical conductivity in the Terahertz range in graphene has opened up new opportunities for optoelectronic applications based on this and other low dimensional materials. Recently, population inversion across the Dirac point has been observed directly by time- and angle-resolved photoemission spectroscopy (tr-ARPES), revealing a relaxation time of only ∼130 femtoseconds. This severely limits the applicability of single layer graphene to, for example, Terahertz light amplification. Here we use tr-ARPES to demonstrate long-lived population inversion in bilayer graphene. The effect is attributed to the small band gap found in this compound. We propose a microscopic model for these observations and speculate that an enhancement of both the pump photon energy and the pump fluence may further increase this lifetime.

Surface area of early visual cortex predicts individual speed of traveling waves during binocular rivalry.

Binocular rivalry ensues when different images are presented to the 2 eyes with conscious perception alternating between the possible interpretations. For large rivalry displays, perceptual transitions are initiated at one location and spread to other parts of the visual field, a phenomenon termed "traveling wave." Previous studies investigated the underlying neural mechanisms of the traveling wave and surmised that primary visual cortex might play an important role. We used magnetic resonance imaging and behavioral measures in humans to explore how interindividual differences in observers' subjective experience of the wave are related to anatomical characteristics of cortical regions. We measured wave speed in participants and confirmed the long-term stability of the individual values. Retinotopic mapping was employed to delineate borders of visual areas V1-V3 in order to determine surface area and cortical thickness in those regions. Only the surface areas of V1 and V2, but not V3 showed a correlation with wave speed. For individuals with larger V1/V2 area, the traveling wave needed longer to spread across the same distance in visual space. Our results highlight the role of early visual areas in mediating binocular rivalry and suggest possible mechanisms for the correlation between surface area and the traveling waves.

Snapshots of non-equilibrium Dirac carrier distributions in graphene.

The optical properties of graphene are made unique by the linear band structure and the vanishing density of states at the Dirac point. It has been proposed that even in the absence of a bandgap, a relaxation bottleneck at the Dirac point may allow for population inversion and lasing at arbitrarily long wavelengths. Furthermore, efficient carrier multiplication by impact ionization has been discussed in the context of light harvesting applications. However, all of these effects are difficult to test quantitatively by measuring the transient optical properties alone, as these only indirectly reflect the energy- and momentum-dependent carrier distributions. Here, we use time- and angle-resolved photoemission spectroscopy with femtosecond extreme-ultraviolet pulses to directly probe the non-equilibrium response of Dirac electrons near the K-point of the Brillouin zone. In lightly hole-doped epitaxial graphene samples, we explore excitation in the mid- and near-infrared, both below and above the minimum photon energy for direct interband transitions. Whereas excitation in the mid-infrared results only in heating of the equilibrium carrier distribution, interband excitations give rise to population inversion, suggesting that terahertz lasing may be possible. However, in neither excitation regime do we find any indication of carrier multiplication, questioning the applicability of graphene for light harvesting.

Feature-based attention affects direction-selective fMRI adaptation in hMT+.

Functional magnetic resonance adaptation has been successfully used to reveal direction-selective responses in the human motion complex (hMT+). Here, we aimed at further investigating direction-selective as well as position-selective responses of hMT+ by looking at how these responses are affected by feature-based attention. We varied motion direction and position of 2 consecutive random-dot stimuli. Participants had to either attend to the direction or the position of the stimuli in separate runs. We show that direction selectivity in hMT+ as measured by functional magnetic resonance imaging (fMRI) adaptation was strongly influenced by task set. Attending to the motion direction of the stimuli lead to stronger direction-selective fMRI adaptation than attending to their position. Position selectivity, on the other hand, was largely unaffected by attentional focus. Interestingly, the change in the direction-selective adaptation profile across tasks could not be explained by inheritance from earlier areas. The response pattern in the early retinotopic cortex was stable across conditions. In conclusion, our results provide further evidence for the flexible coding of direction information in hMT+ depending on task demands.

Auditory motion capturing ambiguous visual motion.

In this study, it is demonstrated that moving sounds have an effect on the direction in which one sees visual stimuli move. During the main experiment sounds were presented consecutively at four speaker locations inducing left or rightward auditory apparent motion. On the path of auditory apparent motion, visual apparent motion stimuli were presented with a high degree of directional ambiguity. The main outcome of this experiment is that our participants perceived visual apparent motion stimuli that were ambiguous (equally likely to be perceived as moving left or rightward) more often as moving in the same direction than in the opposite direction of auditory apparent motion. During the control experiment we replicated this finding and found no effect of sound motion direction on eye movements. This indicates that auditory motion can capture our visual motion percept when visual motion direction is insufficiently determinate without affecting eye movements.

Auditory motion direction encoding in auditory cortex and high-level visual cortex.

The aim of this functional magnetic resonance imaging (fMRI) study was to identify human brain areas that are sensitive to the direction of auditory motion. Such directional sensitivity was assessed in a hypothesis-free manner by analyzing fMRI response patterns across the entire brain volume using a spherical-searchlight approach. In addition, we assessed directional sensitivity in three predefined brain areas that have been associated with auditory motion perception in previous neuroimaging studies. These were the primary auditory cortex, the planum temporale and the visual motion complex (hMT/V5+). Our whole-brain analysis revealed that the direction of sound-source movement could be decoded from fMRI response patterns in the right auditory cortex and in a high-level visual area located in the right lateral occipital cortex. Our region-of-interest-based analysis showed that the decoding of the direction of auditory motion was most reliable with activation patterns of the left and right planum temporale. Auditory motion direction could not be decoded from activation patterns in hMT/V5+. These findings provide further evidence for the planum temporale playing a central role in supporting auditory motion perception. In addition, our findings suggest a cross-modal transfer of directional information to high-level visual cortex in healthy humans.

Orientation-selective functional magnetic resonance imaging adaptation in primary visual cortex revisited.

The processing of orientations is at the core of our visual experience. Orientation selectivity in human visual cortex has been inferred from psychophysical experiments and more recently demonstrated with functional magnetic resonance imaging (fMRI). One method to identify orientation-selective responses is fMRI adaptation, in which two stimuli-either with the same or with different orientations-are presented successively. A region containing orientation-selective neurons should demonstrate an adapted response to the "same orientation" condition in contrast to the "different orientation" condition. So far, human primary visual cortex (V1) showed orientation-selective fMRI adaptation only in experimental designs using prolonged pre-adaptation periods (∼40 s) in combination with top-up stimuli that are thought to maintain the adapted level. This finding has led to the notion that orientation-selective short-term adaptation in V1 (but not V2 or V3) cannot be demonstrated using fMRI. The present study aimed at re-evaluating this question by testing three differently timed adaptation designs. With the use of a more sensitive analysis technique, we show robust orientation-selective fMRI adaptation in V1 evoked by a short-term adaptation design.

Callosal connections of primary visual cortex predict the spatial spreading of binocular rivalry across the visual hemifields.

In binocular rivalry, presentation of different images to the separate eyes leads to conscious perception alternating between the two possible interpretations every few seconds. During perceptual transitions, a stimulus emerging into dominance can spread in a wave-like manner across the visual field. These traveling waves of rivalry dominance have been successfully related to the cortical magnification properties and functional activity of early visual areas, including the primary visual cortex (V1). Curiously however, these traveling waves undergo a delay when passing from one hemifield to another. In the current study, we used diffusion tensor imaging (DTI) to investigate whether the strength of interhemispheric connections between the left and right visual cortex might be related to the delay of traveling waves across hemifields. We measured the delay in traveling wave times (ΔTWT) in 19 participants and repeated this test 6 weeks later to evaluate the reliability of our behavioral measures. We found large interindividual variability but also good test-retest reliability for individual measures of ΔTWT. Using DTI in connection with fiber tractography, we identified parts of the corpus callosum connecting functionally defined visual areas V1-V3. We found that individual differences in ΔTWT was reliably predicted by the diffusion properties of transcallosal fibers connecting left and right V1, but observed no such effect for neighboring transcallosal visual fibers connecting V2 and V3. Our results demonstrate that the anatomical characteristics of topographically specific transcallosal connections predict the individual delay of interhemispheric traveling waves, providing further evidence that V1 is an important site for neural processes underlying binocular rivalry.

Interhemispheric connections shape subjective experience of bistable motion.

The right and left visual hemifields are represented in different cerebral hemispheres and are bound together by connections through the corpus callosum. Much has been learned on the functions of these connections from split-brain patients [1-4], but little is known about their contribution to conscious visual perception in healthy humans. We used diffusion tensor imaging and functional magnetic resonance imaging to investigate which callosal connections contribute to the subjective experience of a visual motion stimulus that requires interhemispheric integration. The "motion quartet" is an ambiguous version of apparent motion that leads to perceptions of either horizontal or vertical motion [5]. Interestingly, observers are more likely to perceive vertical than horizontal motion when the stimulus is presented centrally in the visual field [6]. This asymmetry has been attributed to the fact that, with central fixation, perception of horizontal motion requires integration across hemispheres whereas perception of vertical motion requires only intrahemispheric processing [7]. We are able to show that the microstructure of individually tracked callosal segments connecting motion-sensitive areas of the human MT/V5 complex (hMT/V5+; [8]) can predict the conscious perception of observers. Neither connections between primary visual cortex (V1) nor other surrounding callosal regions exhibit a similar relationship.

Investigating human audio-visual object perception with a combination of hypothesis-generating and hypothesis-testing fMRI analysis tools.

Primate multisensory object perception involves distributed brain regions. To investigate the network character of these regions of the human brain, we applied data-driven group spatial independent component analysis (ICA) to a functional magnetic resonance imaging (fMRI) data set acquired during a passive audio-visual (AV) experiment with common object stimuli. We labeled three group-level independent component (IC) maps as auditory (A), visual (V), and AV, based on their spatial layouts and activation time courses. The overlap between these IC maps served as definition of a distributed network of multisensory candidate regions including superior temporal, ventral occipito-temporal, posterior parietal and prefrontal regions. During an independent second fMRI experiment, we explicitly tested their involvement in AV integration. Activations in nine out of these twelve regions met the max-criterion (A < AV > V) for multisensory integration. Comparison of this approach with a general linear model-based region-of-interest definition revealed its complementary value for multisensory neuroimaging. In conclusion, we estimated functional networks of uni- and multisensory functional connectivity from one dataset and validated their functional roles in an independent dataset. These findings demonstrate the particular value of ICA for multisensory neuroimaging research and using independent datasets to test hypotheses generated from a data-driven analysis.

Neuroelectromagnetic correlates of perceptual closure processes.

Perceptual closure refers to the coherent perception of an object under circumstances when the visual information is incomplete. Although the perceptual closure index observed in electroencephalography reflects that an object has been recognized, the full spatiotemporal dynamics of cortical source activity underlying perceptual closure processing remain unknown so far. To address this question, we recorded magnetoencephalographic activity in 15 subjects (11 females) during a visual closure task and performed beamforming over a sequence of successive short time windows to localize high-frequency gamma-band activity (60-100 Hz). Two-tone images of human faces (Mooney faces) were used to examine perceptual closure. Event-related fields exhibited a magnetic closure index between 250 and 325 ms. Time-frequency analyses revealed sustained high-frequency gamma-band activity associated with the processing of Mooney stimuli; closure-related gamma-band activity was observed between 200 and 300 ms over occipitotemporal channels. Time-resolved source reconstruction revealed an early (0-200 ms) coactivation of caudal inferior temporal gyrus (cITG) and regions in posterior parietal cortex (PPC). At the time of perceptual closure (200-400 ms), the activation in cITG extended to the fusiform gyrus, if a face was perceived. Our data provide the first electrophysiological evidence that perceptual closure for Mooney faces starts with an interaction between areas related to processing of three-dimensional structure from shading cues (cITG) and areas associated with the activation of long-term memory templates (PPC). Later, at the moment of perceptual closure, inferior temporal cortex areas specialized for the perceived object are activated, i.e., the fusiform gyrus related to face processing for Mooney stimuli.

Stimulus predictability reduces responses in primary visual cortex.

In this functional magnetic resonance imaging study we tested whether the predictability of stimuli affects responses in primary visual cortex (V1). The results of this study indicate that visual stimuli evoke smaller responses in V1 when their onset or motion direction can be predicted from the dynamics of surrounding illusory motion. We conclude from this finding that the human brain anticipates forthcoming sensory input that allows predictable visual stimuli to be processed with less neural activation at early stages of cortical processing.