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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

Imagery of a moving object: the role of occipital cortex and human MT/V5+.

Visual imagery--similar to visual perception--activates feature-specific and category-specific visual areas. This is frequently observed in experiments where the instruction is to imagine stimuli that have been shown immediately before the imagery task. Hence, feature-specific activation could be related to the short-term memory retrieval of previously presented sensory information. Here, we investigated mental imagery of stimuli that subjects had not seen before, eliminating the effects of short-term memory. We recorded brain activation using fMRI while subjects performed a behaviourally controlled guided imagery task in predefined retinotopic coordinates to optimize sensitivity in early visual areas. Whole brain analyses revealed activation in a parieto-frontal network and lateral-occipital cortex. Region of interest (ROI) based analyses showed activation in left hMT/V5+. Granger causality mapping taking left hMT/V5+ as source revealed an imagery-specific directed influence from the left inferior parietal lobule (IPL). Interestingly, we observed a negative BOLD response in V1-3 during imagery, modulated by the retinotopic location of the imagined motion trace. Our results indicate that rule-based motion imagery can activate higher-order visual areas involved in motion perception, with a role for top-down directed influences originating in IPL. Lower-order visual areas (V1, V2 and V3) were down-regulated during this type of imagery, possibly reflecting inhibition to avoid visual input from interfering with the imagery construction. This suggests that the activation in early visual areas observed in previous studies might be related to short- or long-term memory retrieval of specific sensory experiences.

The timing of feedback to early visual cortex in the perception of long-range apparent motion.

When 2 visual stimuli are presented one after another in different locations, they are often perceived as one, but moving object. Feedback from area human motion complex hMT/V5+ to V1 has been hypothesized to play an important role in this illusory perception of motion. We measured event-related responses to illusory motion stimuli of varying apparent motion (AM) content and retinal location using Electroencephalography. Detectable cortical stimulus processing started around 60-ms poststimulus in area V1. This component was insensitive to AM content and sequential stimulus presentation. Sensitivity to AM content was observed starting around 90 ms post the second stimulus of a sequence and most likely originated in area hMT/V5+. This AM sensitive response was insensitive to retinal stimulus position. The stimulus sequence related response started to be sensitive to retinal stimulus position at a longer latency of 110 ms. We interpret our findings as evidence for feedback from area hMT/V5+ or a related motion processing area to early visual cortices (V1, V2, V3).

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 cortical representation of objects rotating in depth.

The perception of motion provides valuable interpolations of the visual scene. This fundamental capacity of the visual system is evident in apparent rotation: by presenting only two images of an object rotated in space, a vivid illusion of a smooth apparent motion in three dimensions can be induced. The unseen interpolated rotation views are filled in by the visual system. In the present study, we identified the cortical network responsible for this filling-in process. We argue that cross talk between areas of the ventral and dorsal visual pathways promote the illusion of smooth apparent rotation. Most interestingly, the network represents the unseen object views. Using functional magnetic resonance adaptation, we are able to show that the cortical network selectively adapts to the illusory object views. Our findings provide strong evidence for cortical representations of three-dimensional rotating objects that are view invariant with respect to the rotation path. Furthermore, our results confirm psychophysical investigations that unseen interpolated rotation views can be primed by apparent motion. By applying functional magnetic resonance adaptation, we show for the first time cortical adaptation to unseen objects. Together, our neuroimaging study advances the understanding of the cortical mechanisms mediating the influence of motion on object processing.

Functional magnetic resonance adaptation in visual neuroscience.

Functional magnetic resonance imaging (fMRI) is a powerful non-invasive tool to investigate neuronal processing. In the last ten years a new methodological approach in the field of fMRI has been developed: fMRI adaptation. It has been found that the repetition of a stimulus leads to a decrease of the fMRI signal in the brain region that processes this stimulus. The phenomenon has been related to neuronal adaptation effects found in single-cell recordings. Since the first experiments that observed fMRI-adaptation effects, the method has been applied extensively to study various visual phenomena, such as the perception of motion, shape, objects, and orientation. The great advantage of fMRI adaptation is that it allows assessing the functional response profile of a brain region at a subvoxel level. The purpose of the current review is to evaluate the different experimental approaches used to elicit fMRI-adaptation effects. We discuss papers published in the domain of visual neuroscience that made use of fMRI-adaptation paradigms. In doing so, we focus on methodological considerations concerning experimental design, stimulus presentation and influencing factors such as awareness and attention. In the course of this review, we show that different fMRI-adaptation designs capture heterogeneous neuronal adaptation effects. As the picture of the mechanisms underlying neuronal adaptation changes from simple synaptic fatigue to complex network interactions, the concept of fMRI adaptation has to be redefined.

Primary visual cortex activity along the apparent-motion trace reflects illusory perception.

The illusion of apparent motion can be induced when visual stimuli are successively presented at different locations. It has been shown in previous studies that motion-sensitive regions in extrastriate cortex are relevant for the processing of apparent motion, but it is unclear whether primary visual cortex (V1) is also involved in the representation of the illusory motion path. We investigated, in human subjects, apparent-motion-related activity in patches of V1 representing locations along the path of illusory stimulus motion using functional magnetic resonance imaging. Here we show that apparent motion caused a blood-oxygenation-level-dependent response along the V1 representations of the apparent-motion path, including regions that were not directly activated by the apparent-motion-inducing stimuli. This response was unaltered when participants had to perform an attention-demanding task that diverted their attention away from the stimulus. With a bistable motion quartet, we confirmed that the activity was related to the conscious perception of movement. Our data suggest that V1 is part of the network that represents the illusory path of apparent motion. The activation in V1 can be explained either by lateral interactions within V1 or by feedback mechanisms from higher visual areas, especially the motion-sensitive human MT/V5 complex.