Human Sensorimotor Beta Event Characteristics and Aperiodic Signal Are Highly Heritable.

Individuals' phenotypes, including the brain's structure and function, are largely determined by genes and their interplay. The resting brain generates salient rhythmic patterns that can be characterized noninvasively using functional neuroimaging such as magnetoencephalography (MEG). One of these rhythms, the somatomotor (rolandic) beta rhythm, shows intermittent high amplitude "events" that predict behavior across tasks and species. Beta rhythm is altered in neurological disease. The aperiodic (1/f) signal present in electrophysiological recordings is also modulated by some neurological conditions and aging. Both sensorimotor beta and aperiodic signal could thus serve as biomarkers of sensorimotor function. Knowledge about the extent to which these brain functional measures are heritable could shed light on the mechanisms underlying their generation. We investigated the heritability and variability of human spontaneous sensorimotor beta rhythm events and aperiodic activity in 210 healthy male and female adult siblings' spontaneous MEG activity. The most heritable trait was the aperiodic 1/f signal, with a heritability of 0.87 in the right hemisphere. Time-resolved beta event amplitude parameters were also highly heritable, whereas the heritabilities for overall beta power, peak frequency, and measures of event duration remained nonsignificant. Human sensorimotor neural activity can thus be dissected into different components with variable heritability. We postulate that these differences partially reflect different underlying signal-generating mechanisms. The 1/f signal and beta event amplitude measures may depend more on fixed, anatomical parameters, whereas beta event duration and its modulation reflect dynamic characteristics, guiding their use as potential disease biomarkers.

Impaired Cortical Tracking of Speech in Children with Developmental Language Disorder.

In developmental language disorder (DLD), learning to comprehend and express oneself with spoken language is impaired, but the reason for this remains unknown. Using millisecond-scale magnetoencephalography recordings combined with machine learning models, we investigated whether the possible neural basis of this disruption lies in poor cortical tracking of speech. The stimuli were common spoken Finnish words (e.g., dog, car, hammer) and sounds with corresponding meanings (e.g., dog bark, car engine, hammering). In both children with DLD (10 boys and 7 girls) and typically developing (TD) control children (14 boys and 3 girls), aged 10-15 years, the cortical activation to spoken words was best modeled as time-locked to the unfolding speech input at ∼100 ms latency between sound and cortical activation. Amplitude envelope (amplitude changes) and spectrogram (detailed time-varying spectral content) of the spoken words, but not other sounds, were very successfully decoded based on time-locked brain responses in bilateral temporal areas; based on the cortical responses, the models could tell at ∼75-85% accuracy which of the two sounds had been presented to the participant. However, the cortical representation of the amplitude envelope information was poorer in children with DLD compared with TD children at longer latencies (at ∼200-300 ms lag). We interpret this effect as reflecting poorer retention of acoustic-phonetic information in short-term memory. This impaired tracking could potentially affect the processing and learning of words as well as continuous speech. The present results offer an explanation for the problems in language comprehension and acquisition in DLD.

Beta- and gamma-band cortico-cortical interactions support naturalistic reading of continuous text.

Large-scale integration of information across cortical structures, building on neural connectivity, has been proposed to be a key element in supporting human cognitive processing. In electrophysiological neuroimaging studies of reading, quantification of neural interactions has been limited to the level of isolated words or sentences due to artefacts induced by eye movements. Here, we combined magnetoencephalography recording with advanced artefact rejection tools to investigate both cortico-cortical coherence and directed neural interactions during naturalistic reading of full-page texts. Our results show that reading versus visual scanning of text was associated with wide-spread increases of cortico-cortical coherence in the beta and gamma bands. We further show that the reading task was linked to increased directed neural interactions compared to the scanning task across a sparse set of connections within a wide range of frequencies. Together, the results demonstrate that neural connectivity flexibly builds on different frequency bands to support continuous natural reading.

Bayesian reduced rank regression models generalizable neural fingerprints that differentiate between individuals in magnetoencephalography data.

Recent magnetoencephalography (MEG) studies have reported that functional connectivity (FC) and power spectra can be used as neural fingerprints in differentiating individuals. Such studies have mainly used correlations between measurement sessions to distinguish individuals from each other. However, it has remained unclear whether such correlations might reflect a more generalizable principle of individually distinctive brain patterns. Here, we evaluated a machine-learning based approach, termed latent-noise Bayesian reduced rank regression (BRRR) as a means of modelling individual differences in the resting-state MEG data of the Human Connectome Project (HCP), using FC and power spectra as neural features. First, we verified that BRRR could model and reproduce the differences between metrics that correlation-based fingerprinting yields. We trained BRRR models to distinguish individuals based on data from one measurement and used the models to identify subsequent measurement sessions of those same individuals. The best performing BRRR models, using only 20 spatiospectral components, were able to identify subjects across measurement sessions with over 90% accuracy, approaching the highest correlation-based accuracies. Using cross-validation, we then determined whether that BRRR model could generalize to unseen subjects, successfully classifying the measurement sessions of novel individuals with over 80% accuracy. The results demonstrate that individual neurofunctional differences can be reliably extracted from MEG data with a low-dimensional predictive model and that the model is able to classify novel subjects.

Semantic feature norms: a cross-method and cross-language comparison.

The ability to assign meaning to perceptual stimuli forms the basis of human behavior and the ability to use language. The meanings of things have primarily been probed using behavioral production norms and corpus-derived statistical methods. However, it is not known to what extent the collection method and the language being probed influence the resulting semantic feature vectors. In this study, we compare behavioral with corpus-based norms, across Finnish and English, using an all-to-all approach. To complete the set of norms required for this study, we present a new set of Finnish behavioral production norms, containing both abstract and concrete concepts. We found that all the norms provide largely similar information about the relationships of concrete objects and allow item-level mapping across norms sets. This validates the use of the corpus-derived norms which are easier to obtain than behavioral norms, which are labor-intensive to collect, for studies that do not depend on subtle differences in meaning between close semantic neighbors.

Picture naming yields highly consistent cortical activation patterns: Test-retest reliability of magnetoencephalography recordings.

Reliable paradigms and imaging measures of individual-level brain activity are paramount when reaching from group-level research studies to clinical assessment of individual patients. Magnetoencephalography (MEG) provides a direct, non-invasive measure of cortical processing with high spatiotemporal accuracy, and is thus well suited for assessment of functional brain damage in patients with language difficulties. This MEG study aimed to identify, in a delayed picture naming paradigm, source-localized evoked activity and modulations of cortical oscillations that show high test-retest reliability across measurement days in healthy individuals, demonstrating their applicability in clinical settings. For patients with a language disorder picture naming can be a challenging task. Therefore, we also determined whether a semantic judgment task ('Is this item living?') with a spoken response ("yes"/"no") would suffice to induce comparably consistent activity within brain regions related to language production. The MEG data was collected from 19 healthy participants on two separate days. In picture naming, evoked activity was consistent across measurement days (intraclass correlation coefficient (ICC)>0.4) in the left frontal (400-800 ms after image onset), sensorimotor (200-800 ms), parietal (200-600 ms), temporal (200-800 ms), occipital (400-800 ms) and cingulate (600-800 ms) regions, as well as the right temporal (600-800 ms) region. In the semantic judgment task, consistent evoked activity was spatially more limited, occurring in the left temporal (200-800 ms), sensorimotor (400-800 ms), occipital (400-600 ms) and subparietal (600-800 ms) regions, and the right supramarginal cortex (600-800 ms). The delayed naming task showed typical beta oscillatory suppression in premotor and sensorimotor regions (800-1200 ms) but other consistent modulations of oscillatory activity were mostly observed in posterior cortical regions that have not typically been associated with language processing. The high test-retest consistency of MEG evoked activity in the picture naming task testifies to its applicability in clinical evaluations of language function, as well as in longitudinal MEG studies of language production in clinical and healthy populations.

Selective auditory attention within naturalistic scenes modulates reactivity to speech sounds.

Rapid recognition and categorization of sounds are essential for humans and animals alike, both for understanding and reacting to our surroundings and for daily communication and social interaction. For humans, perception of speech sounds is of crucial importance. In real life, this task is complicated by the presence of a multitude of meaningful non-speech sounds. The present behavioural, magnetoencephalography (MEG) and functional magnetic resonance imaging (fMRI) study was set out to address how attention to speech versus attention to natural non-speech sounds within complex auditory scenes influences cortical processing. The stimuli were superimpositions of spoken words and environmental sounds, with parametric variation of the speech-to-environmental sound intensity ratio. The participants' task was to detect a repetition in either the speech or the environmental sound. We found that specifically when participants attended to speech within the superimposed stimuli, higher speech-to-environmental sound ratios resulted in shorter sustained MEG responses and stronger BOLD fMRI signals especially in the left supratemporal auditory cortex and in improved behavioural performance. No such effects of speech-to-environmental sound ratio were observed when participants attended to the environmental sound part within the exact same stimuli. These findings suggest stronger saliency of speech compared with other meaningful sounds during processing of natural auditory scenes, likely linked to speech-specific top-down and bottom-up mechanisms activated during speech perception that are needed for tracking speech in real-life-like auditory environments.

The neural representation of abstract words may arise through grounding word meaning in language itself.

In order to describe how humans represent meaning in the brain, one must be able to account for not just concrete words but, critically, also abstract words, which lack a physical referent. Hebbian formalism and optimization are basic principles of brain function, and they provide an appealing approach for modeling word meanings based on word co-occurrences. We provide proof of concept that a statistical model of the semantic space can account for neural representations of both concrete and abstract words, using MEG. Here, we built a statistical model using word embeddings extracted from a text corpus. This statistical model was used to train a machine learning algorithm to successfully decode the MEG signals evoked by written words. In the model, word abstractness emerged from the statistical regularities of the language environment. Representational similarity analysis further showed that this salient property of the model co-varies, at 280-420 ms after visual word presentation, with activity in regions that have been previously linked with processing of abstract words, namely the left-hemisphere frontal, anterior temporal and superior parietal cortex. In light of these results, we propose that the neural encoding of word meanings can arise through statistical regularities, that is, through grounding in language itself.

Task-Modulated Corticocortical Synchrony in the Cognitive-Motor Network Supporting Handwriting.

Both motor and cognitive aspects of behavior depend on dynamic, accurately timed neural processes in large-scale brain networks. Here, we studied synchronous interplay between cortical regions during production of cognitive-motor sequences in humans. Specifically, variants of handwriting that differed in motor variability, linguistic content, and memorization of movement cues were contrasted to unveil functional sensitivity of corticocortical connections. Data-driven magnetoencephalography mapping (n = 10) uncovered modulation of mostly left-hemispheric corticocortical interactions, as quantified by relative changes in phase synchronization. At low frequencies (~2-13 Hz), enhanced frontoparietal synchrony was related to regular handwriting, whereas premotor cortical regions synchronized for simple loop production and temporo-occipital areas for a writing task substituting normal script with loop patterns. At the beta-to-gamma band (~13-45 Hz), enhanced synchrony was observed for regular handwriting in the central and frontoparietal regions, including connections between the sensorimotor and supplementary motor cortices and between the parietal and dorsal premotor/precentral cortices. Interpreted within a modular framework, these modulations of synchrony mainly highlighted interactions of the putative pericentral subsystem of hand coordination and the frontoparietal subsystem mediating working memory operations. As part of cortical dynamics, interregional phase synchrony varies depending on task demands in production of cognitive-motor sequences.

Post-hoc modification of linear models: Combining machine learning with domain information to make solid inferences from noisy data.

Linear machine learning models "learn" a data transformation by being exposed to examples of input with the desired output, forming the basis for a variety of powerful techniques for analyzing neuroimaging data. However, their ability to learn the desired transformation is limited by the quality and size of the example dataset, which in neuroimaging studies is often notoriously noisy and small. In these cases, it is desirable to fine-tune the learned linear model using domain information beyond the example dataset. To this end, we present a framework that decomposes the weight matrix of a fitted linear model into three subcomponents: the data covariance, the identified signal of interest, and a normalizer. Inspecting these subcomponents in isolation provides an intuitive way to inspect the inner workings of the model and assess its strengths and weaknesses. Furthermore, the three subcomponents may be altered, which provides a straightforward way to inject prior information and impose additional constraints. We refer to this process as "post-hoc modification" of a model and demonstrate how it can be used to achieve precise control over which aspects of the model are fitted to the data through machine learning and which are determined through domain information. As an example use case, we decode the associative strength between words from electroencephalography (EEG) reading data. Our results show how the decoding accuracy of two example linear models (ridge regression and logistic regression) can be boosted by incorporating information about the spatio-temporal nature of the data, domain information about the N400 evoked potential and data from other participants.

Children show hemispheric differences in the basic auditory response properties.

Auditory cortex in each hemisphere shows preference to sounds from the opposite hemifield in the auditory space. Besides this contralateral dominance, the auditory cortex shows functional and structural lateralization, presumably influencing the features of subsequent auditory processing. Children have been shown to differ from adults in the hemispheric balance of activation in higher-order auditory based tasks. We studied, first, whether the contralateral dominance can be detected in 7- to 8-year-old children and, second, whether the response properties of auditory cortex in children differ between hemispheres. Magnetoencephalography (MEG) responses to simple tones revealed adult-like contralateral preference that was, however, extended in time in children. Moreover, we found stronger emphasis towards mature response properties in the right than left hemisphere, pointing to faster maturation of the right-hemisphere auditory cortex. The activation strength of the child-typical prolonged response was significantly decreased with age, within the narrow age-range of the studied child population. Our results demonstrate that although the spatial sensitivity to the opposite hemifield has emerged by 7 years of age, the population-level neurophysiological response shows salient immature features, manifested particularly in the left hemisphere. The observed functional differences between hemispheres may influence higher-level processing stages, for example, in language function.

Discovering heritable modes of MEG spectral power.

Brain structure and many brain functions are known to be genetically controlled, but direct links between neuroimaging measures and their underlying cellular-level determinants remain largely undiscovered. Here, we adopt a novel computational method for examining potential similarities in high-dimensional brain imaging data between siblings. We examine oscillatory brain activity measured with magnetoencephalography (MEG) in 201 healthy siblings and apply Bayesian reduced-rank regression to extract a low-dimensional representation of familial features in the participants' spectral power structure. Our results show that the structure of the overall spectral power at 1-90 Hz is a highly conspicuous feature that not only relates siblings to each other but also has very high consistency within participants' own data, irrespective of the exact experimental state of the participant. The analysis is extended by seeking genetic associations for low-dimensional descriptions of the oscillatory brain activity. The observed variability in the MEG spectral power structure was associated with SDK1 (sidekick cell adhesion molecule 1) and suggestively with several other genes that function, for example, in brain development. The current results highlight the potential of sophisticated computational methods in combining molecular and neuroimaging levels for exploring brain functions, even for high-dimensional data limited to a few hundred participants.

Statistical models of morphology predict eye-tracking measures during visual word recognition.

We studied how statistical models of morphology that are built on different kinds of representational units, i.e., models emphasizing either holistic units or decomposition, perform in predicting human word recognition. More specifically, we studied the predictive power of such models at early vs. late stages of word recognition by using eye-tracking during two tasks. The tasks included a standard lexical decision task and a word recognition task that assumedly places less emphasis on postlexical reanalysis and decision processes. The lexical decision results showed good performance of Morfessor models based on the Minimum Description Length optimization principle. Models which segment words at some morpheme boundaries and keep other boundaries unsegmented performed well both at early and late stages of word recognition, supporting dual- or multiple-route cognitive models of morphological processing. Statistical models based on full forms fared better in late than early measures. The results of the second, multi-word recognition task showed that early and late stages of processing often involve accessing morphological constituents, with the exception of short complex words. Late stages of word recognition additionally involve predicting upcoming morphemes on the basis of previous ones in multimorphemic words. The statistical models based fully on whole words did not fare well in this task. Thus, we assume that the good performance of such models in global measures such as gaze durations or reaction times in lexical decision largely stems from postlexical reanalysis or decision processes. This finding highlights the importance of considering task demands in the study of morphological processing.

Exploring the Organization of Semantic Memory through Unsupervised Analysis of Event-related Potentials.

Modern multivariate methods have enabled the application of unsupervised techniques to analyze neurophysiological data without strict adherence to predefined experimental conditions. We demonstrate a multivariate method that leverages priming effects on the evoked potential to perform hierarchical clustering on a set of word stimuli. The current study focuses on the semantic relationships that play a key role in the organization of our mental lexicon of words and concepts. The N400 component of the event-related potential is considered a reliable neurophysiological response that is indicative of whether accessing one concept facilitates subsequent access to another (i.e., one "primes" the other). To further our understanding of the organization of the human mental lexicon, we propose to utilize the N400 component to drive a clustering algorithm that can uncover, given a set of words, which particular subsets of words show mutual priming. Such a scheme requires a reliable measurement of the amplitude of the N400 component without averaging across many trials, which was here achieved using a recently developed multivariate analysis method based on beamforming. We validated our method by demonstrating that it can reliably detect, without any prior information about the nature of the stimuli, a well-known feature of the organization of our semantic memory: the distinction between animate and inanimate concepts. These results motivate further application of our method to data-driven exploration of disputed or unknown relationships between stimuli.

Cortical Tracking of Global and Local Variations of Speech Rhythm during Connected Natural Speech Perception.

During natural speech perception, listeners must track the global speaking rate, that is, the overall rate of incoming linguistic information, as well as transient, local speaking rate variations occurring within the global speaking rate. Here, we address the hypothesis that this tracking mechanism is achieved through coupling of cortical signals to the amplitude envelope of the perceived acoustic speech signals. Cortical signals were recorded with magnetoencephalography (MEG) while participants perceived spontaneously produced speech stimuli at three global speaking rates (slow, normal/habitual, and fast). Inherently to spontaneously produced speech, these stimuli also featured local variations in speaking rate. The coupling between cortical and acoustic speech signals was evaluated using audio-MEG coherence. Modulations in audio-MEG coherence spatially differentiated between tracking of global speaking rate, highlighting the temporal cortex bilaterally and the right parietal cortex, and sensitivity to local speaking rate variations, emphasizing the left parietal cortex. Cortical tuning to the temporal structure of natural connected speech thus seems to require the joint contribution of both auditory and parietal regions. These findings suggest that cortical tuning to speech rhythm operates on two functionally distinct levels: one encoding the global rhythmic structure of speech and the other associated with online, rapidly evolving temporal predictions. Thus, it may be proposed that speech perception is shaped by evolutionary tuning, a preference for certain speaking rates, and predictive tuning, associated with cortical tracking of the constantly changing-rate of linguistic information in a speech stream.

Information properties of morphologically complex words modulate brain activity during word reading.

Neuroimaging studies of the reading process point to functionally distinct stages in word recognition. Yet, current understanding of the operations linked to those various stages is mainly descriptive in nature. Approaches developed in the field of computational linguistics may offer a more quantitative approach for understanding brain dynamics. Our aim was to evaluate whether a statistical model of morphology, with well-defined computational principles, can capture the neural dynamics of reading, using the concept of surprisal from information theory as the common measure. The Morfessor model, created for unsupervised discovery of morphemes, is based on the minimum description length principle and attempts to find optimal units of representation for complex words. In a word recognition task, we correlated brain responses to word surprisal values derived from Morfessor and from other psycholinguistic variables that have been linked with various levels of linguistic abstraction. The magnetoencephalography data analysis focused on spatially, temporally and functionally distinct components of cortical activation observed in reading tasks. The early occipital and occipito-temporal responses were correlated with parameters relating to visual complexity and orthographic properties, whereas the later bilateral superior temporal activation was correlated with whole-word based and morphological models. The results show that the word processing costs estimated by the statistical Morfessor model are relevant for brain dynamics of reading during late processing stages.

The right hemisphere is highlighted in connected natural speech production and perception.

Current understanding of the cortical mechanisms of speech perception and production stems mostly from studies that focus on single words or sentences. However, it has been suggested that processing of real-life connected speech may rely on additional cortical mechanisms. In the present study, we examined the neural substrates of natural speech production and perception with magnetoencephalography by modulating three central features related to speech: amount of linguistic content, speaking rate and social relevance. The amount of linguistic content was modulated by contrasting natural speech production and perception to speech-like non-linguistic tasks. Meaningful speech was produced and perceived at three speaking rates: normal, slow and fast. Social relevance was probed by having participants attend to speech produced by themselves and an unknown person. These speech-related features were each associated with distinct spatiospectral modulation patterns that involved cortical regions in both hemispheres. Natural speech processing markedly engaged the right hemisphere in addition to the left. In particular, the right temporo-parietal junction, previously linked to attentional processes and social cognition, was highlighted in the task modulations. The present findings suggest that its functional role extends to active generation and perception of meaningful, socially relevant speech.

Evidence for genetic regulation of the human parieto-occipital 10-Hz rhythmic activity.

Several functional and morphological brain measures are partly under genetic control. The identification of direct links between neuroimaging signals and corresponding genetic factors can reveal cellular-level mechanisms behind the measured macroscopic signals and contribute to the use of imaging signals as probes of genetic function. To uncover possible genetic determinants of the most prominent brain signal oscillation, the parieto-occipital 10-Hz alpha rhythm, we measured spontaneous brain activity with magnetoencephalography in 210 healthy siblings while the subjects were resting, with eyes closed and open. The reactivity of the alpha rhythm was quantified from the difference spectra between the two conditions. We focused on three measures: peak frequency, peak amplitude and the width of the main spectral peak. In accordance with earlier electroencephalography studies, spectral peak amplitude was highly heritable (h(2)  > 0.75). Variance component-based analysis of 28 000 single-nucleotide polymorphism markers revealed linkage for both the width and the amplitude of the spectral peak. The strongest linkage was detected for the width of the spectral peak over the left parieto-occipital cortex on chromosome 10 (LOD = 2.814, nominal P < 0.03). This genomic region contains several functionally plausible genes, including GRID1 and ATAD1 that regulate glutamate receptor channels mediating synaptic transmission, NRG3 with functions in brain development and HRT7 involved in the serotonergic system and circadian rhythm. Our data suggest that the alpha oscillation is in part genetically regulated, and that it may be possible to identify its regulators by genetic analyses on a realistically modest number of samples.

A multimodal spectral approach to characterize rhythm in natural speech.

Human utterances demonstrate temporal patterning, also referred to as rhythm. While simple oromotor behaviors (e.g., chewing) feature a salient periodical structure, conversational speech displays a time-varying quasi-rhythmic pattern. Quantification of periodicity in speech is challenging. Unimodal spectral approaches have highlighted rhythmic aspects of speech. However, speech is a complex multimodal phenomenon that arises from the interplay of articulatory, respiratory, and vocal systems. The present study addressed the question of whether a multimodal spectral approach, in the form of coherence analysis between electromyographic (EMG) and acoustic signals, would allow one to characterize rhythm in natural speech more efficiently than a unimodal analysis. The main experimental task consisted of speech production at three speaking rates; a simple oromotor task served as control. The EMG-acoustic coherence emerged as a sensitive means of tracking speech rhythm, whereas spectral analysis of either EMG or acoustic amplitude envelope alone was less informative. Coherence metrics seem to distinguish and highlight rhythmic structure in natural speech.

Task- and stimulus-related cortical networks in language production: Exploring similarity of MEG- and fMRI-derived functional connectivity.

Large-scale networks support the dynamic integration of information across multiple functionally specialized brain regions. Network analyses of haemodynamic modulations have revealed such functional brain networks that show high consistency across subjects and different cognitive states. However, the relationship between the slowly fluctuating haemodynamic responses and the underlying neural mechanisms is not well understood. Resting state studies have revealed spatial similarities in the estimated network hub locations derived using haemodynamic and electrophysiological recordings, suggesting a direct neural basis for the widely described functional magnetic resonance imaging (fMRI) resting state networks. To truly understand the nature of the relationship between electrophysiology and haemodynamics it is important to move away from a task absent state and to establish if such networks are differentially modulated by cognitive processing. The present parallel fMRI and magnetoencephalography (MEG) experiment investigated the structural similarities between haemodynamic networks and their electrophysiological counterparts when either the stimulus or the task was varied. Connectivity patterns underlying action vs. object naming (task-driven modulations), and action vs. object images (stimulus-driven modulations) were identified in a data driven all-to-all connectivity analysis, with cross spectral coherence adopted as a metric of functional connectivity in both MEG and fMRI. We observed a striking difference in functional connectivity between conditions. The spectral profiles of the frequency-specific network similarity differed significantly for the task-driven vs. stimulus-driven connectivity modulations. While the greatest similarity between MEG and fMRI derived networks was observed at neural frequencies below 30 Hz, haemodynamic network interactions could not be attributed to a single frequency band. Instead, the entire spectral profile should be taken into account when assessing the correspondence between MEG and fMRI networks. Task-driven network hubs, evident in both MEG and fMRI, were found in cortical regions previously associated with language processing, including the posterior temporal cortex and the inferior frontal cortex. Network hubs related to stimulus-driven modulations, however, were found in regions related to object recognition and visual processing, including the lateral occipital cortex. Overall, the results depict a shift in network structure when moving from a task dependent modulation to a stimulus dependent modulation, revealing a reorganization of large-scale functional connectivity during task performance.