Synchronization dynamics of phase oscillators on power grid models.

We investigate topological and spectral properties of models of European and US-American power grids and of paradigmatic network models as well as their implications for the synchronization dynamics of phase oscillators with heterogeneous natural frequencies. We employ the complex-valued order parameter-a widely used indicator for phase ordering-to assess the synchronization dynamics and observe the order parameter to exhibit either constant or periodic or non-periodic, possibly chaotic temporal evolutions for a given coupling strength but depending on initial conditions and the systems' disorder. Interestingly, both topological and spectral characteristics of the power grids point to a diminished capability of these networks to support a temporarily stable synchronization dynamics. We find non-trivial commonalities between the synchronization dynamics of oscillators on seemingly opposing topologies.

Impact of biological rhythms on the importance hierarchy of constituents in time-dependent functional brain networks.

Biological rhythms are natural, endogenous cycles with period lengths ranging from less than 24 h (ultradian rhythms) to more than 24 h (infradian rhythms). The impact of the circadian rhythm (approximately 24 h) and ultradian rhythms on spectral characteristics of electroencephalographic (EEG) signals has been investigated for more than half a century. Yet, only little is known on how biological rhythms influence the properties of EEG-derived evolving functional brain networks. Here, we derive such networks from multiday, multichannel EEG recordings and use different centrality concepts to assess the time-varying importance hierarchy of the networks' vertices and edges as well as the various aspects of their structural integration in the network. We observe strong circadian and ultradian influences that highlight distinct subnetworks in the evolving functional brain networks. Our findings indicate the existence of a vital and fundamental subnetwork that is rather generally involved in ongoing brain activities during wakefulness and sleep.

A perturbation-based approach to identifying potentially superfluous network constituents.

Constructing networks from empirical time-series data is often faced with the as yet unsolved issue of how to avoid potentially superfluous network constituents. Such constituents can result, e.g., from spatial and temporal oversampling of the system's dynamics, and neglecting them can lead to severe misinterpretations of network characteristics ranging from global to local scale. We derive a perturbation-based method to identify potentially superfluous network constituents that makes use of vertex and edge centrality concepts. We investigate the suitability of our approach through analyses of weighted small-world, scale-free, random, and complete networks.

Epileptic-network-based prediction and control of seizures in humans.

Epilepsy is now conceptualized as a network disease. The epileptic brain network comprises structurally and functionally connected cortical and subcortical brain regions - spanning lobes and hemispheres -, whose connections and dynamics evolve in time. With this concept, focal and generalized seizures as well as other related pathophysiological phenomena are thought to emerge from, spread via, and be terminated by network vertices and edges that also generate and sustain normal, physiological brain dynamics. Research over the last years has advanced concepts and techniques to identify and characterize the evolving epileptic brain network and its constituents on various spatial and temporal scales. Network-based approaches further our understanding of how seizures emerge from the evolving epileptic brain network, and they provide both novel insights into pre-seizure dynamics and important clues for success or failure of measures for network-based seizure control and prevention. In this review, we summarize the current state of knowledge and address several important challenges that would need to be addressed to move network-based prediction and control of seizures closer to clinical translation.

The time-evolving epileptic brain network: concepts, definitions, accomplishments, perspectives.

Epilepsy is now considered a network disease that affects the brain across multiple levels of spatial and temporal scales. The paradigm shift from an epileptic focus-a discrete cortical area from which seizures originate-to a widespread epileptic network-spanning lobes and hemispheres-considerably advanced our understanding of epilepsy and continues to influence both research and clinical treatment of this multi-faceted high-impact neurological disorder. The epileptic network, however, is not static but evolves in time which requires novel approaches for an in-depth characterization. In this review, we discuss conceptual basics of network theory and critically examine state-of-the-art recording techniques and analysis tools used to assess and characterize a time-evolving human epileptic brain network. We give an account on current shortcomings and highlight potential developments towards an improved clinical management of epilepsy.

Network structure from a characterization of interactions in complex systems.

Many natural and man-made complex dynamical systems can be represented by networks with vertices representing system units and edges the coupling between vertices. If edges of such a structural network are inaccessible, a widely used approach is to identify them with interactions between vertices, thereby setting up a functional network. However, it is an unsolved issue if and to what extent important properties of a functional network on the global and the local scale match those of the corresponding structural network. We address this issue by deriving functional networks from characterizing interactions in paradigmatic oscillator networks with widely-used time-series-analysis techniques for various factors that alter the collective network dynamics. Surprisingly, we find that particularly key constituents of functional networks-as identified with betweenness and eigenvector centrality-coincide with ground truth to a high degree, while global topological and spectral properties-clustering coefficient, average shortest path length, assortativity, and synchronizability-clearly deviate. We obtain similar concurrences for an empirical network. Our findings are of relevance for various scientific fields and call for conceptual and methodological refinements to further our understanding of the relationship between structure and function of complex dynamical systems.

Transcutaneous Auricular Vagus Nerve Stimulation Differently Modifies Functional Brain Networks of Subjects With Different Epilepsy Types.

Epilepsy types differ by pathophysiology and prognosis. Transcutaneous auricular vagus nerve stimulation (taVNS) is a non-invasive treatment option in epilepsy. Nevertheless, its mode of action and impact on different types of epilepsy are still unknown. We investigated whether short-term taVNS differently affects local and global characteristics of EEG-derived functional brain networks in different types of epilepsy. Thirty subjects (nine with focal epilepsy, 11 with generalized epilepsy, and 10 without epilepsy or seizures) underwent a 3-h continuous EEG-recording (1 h pre-stimulation, 1 h taVNS stimulation, 1 h post-stimulation) from which we derived evolving functional brain networks. We assessed-in a time-resolved manner-important global (topological, robustness, and stability properties) and local (centralities of vertices and edges) network characteristics. Compared to the subjects with focal epilepsies and without epilepsy, those with generalized epilepsies clearly presented with different topological properties of their functional brain network already at rest. Furthermore, subjects with focal and generalized epilepsies reacted differently to the stimulation, expressed as different taVNS-induced immediate and enduring reorganization of global network characteristics. On the local network scale, no discernible spatial pattern could be detected, which points to a rather unspecific and generalized modification of brain activity. Assessing functional brain network characteristics can provide additional information for differentiating between focal and generalized epilepsy. TaVNS-related modifications of global network characteristics clearly differ between epilepsy types. Impact of such a non-pharmaceutical intervention on clinical decision-making in the treatment of different epilepsy types needs to be assessed in future studies.

Modifications of Functional Human Brain Networks by Transcutaneous Auricular Vagus Nerve Stimulation: Impact of Time of Day.

Transcutaneous auricular vagus nerve stimulation (taVNS) is a novel non-invasive treatment option for different diseases and symptoms, such as epilepsy or depression. Its mechanism of action, however, is still not fully understood. We investigated short-term taVNS-induced changes of local and global properties of EEG-derived, evolving functional brain networks from eighteen subjects who underwent two 1 h stimulation phases (morning and afternoon) during continuous EEG-recording. In the majority of subjects, taVNS induced measurable modifications of network properties. Network alterations induced by stimulation in the afternoon were clearly more pronounced than those induced by stimulation in the morning. Alterations mostly affected the networks' topology and stability properties. On the local network scale, no clear-cut spatial stimulation-related patterns could be discerned. Our findings indicate that the possible impact of diurnal influences on taVNS-induced network modifications would need to be considered for future research and clinical studies of this non-pharmaceutical intervention approach.

Electrodermal Activity Biofeedback Alters Evolving Functional Brain Networks in People With Epilepsy, but in a Non-specific Manner.

There is evidence that biofeedback of electrodermal activity (EDA) can reduce seizure frequency in people with epilepsy. Prior studies have linked EDA biofeedback to a diffuse brain activation as a potential functional mechanism. Here, we investigated whether short-term EDA biofeedback alters EEG-derived large-scale functional brain networks in people with epilepsy. In this prospective controlled trial, thirty participants were quasi-randomly assigned to one of three biofeedback conditions (arousal, sham, or relaxation) and performed a single, 30-min biofeedback training while undergoing continuous EEG recordings. Based on the EEG, we derived evolving functional brain networks and examined their topological, robustness, and stability properties over time. Potential effects on attentional-executive functions and mood were monitored via a neuropsychological assessment and subjective self-ratings. Participants assigned to the relaxation group seemed to be most successful in meeting the task requirements for this specific control condition (i.e., decreasing EDA). Participants in the sham group were more successful in increasing EDA than participants in the arousal group. However, only the arousal biofeedback training was associated with a prolonged robustness-enhancing effect on networks. Effects on other network properties were mostly unspecific for the different groups. None of the biofeedback conditions affected attentional-executive functions or subjective behavioral measures. Our results suggest that global characteristics of evolving functional brain networks are modified by EDA biofeedback. Some alterations persisted after the single training session; however, the effects were largely unspecific across the different biofeedback protocols. Further research should address changes of local network characteristics and whether multiple training sessions will result in more specific network modifications.

A straightforward edge centrality concept derived from generalizing degree and strength.

Vertex degree-the number of edges that are incident to a vertex-is a fundamental concept in network theory. It is the historically first and conceptually simplest centrality concept to rate the importance of a vertex for a network's structure and dynamics. Unlike many other centrality concepts, for which joint metrics have been proposed for both vertices and edges, by now there is no concept for an edge centrality analogous to vertex degree. Here, we propose such a concept-termed nearest-neighbor edge centrality-and demonstrate its suitability for a non-redundant identification of central edges in paradigmatic network models as well as in real-world networks from various scientific domains.

Extreme events in a complex network: Interplay between degree distribution and repulsive interaction.

The role of topological heterogeneity in the origin of extreme events in a network is investigated here. The dynamics of the oscillators associated with the nodes are assumed to be identical and influenced by mean-field repulsive interactions. An interplay of topological heterogeneity and the repulsive interaction between the dynamical units of the network triggers extreme events in the nodes when each node succumbs to such events for discretely different ranges of repulsive coupling. A high degree node is vulnerable to weaker repulsive interactions, while a low degree node is susceptible to stronger interactions. As a result, the formation of extreme events changes position with increasing strength of repulsive interaction from high to low degree nodes. Extreme events at any node are identified with the appearance of occasional large-amplitude events (amplitude of the temporal dynamics) that are larger than a threshold height and rare in occurrence, which we confirm by estimating the probability distribution of all events. Extreme events appear at any oscillator near the boundary of transition from rotation to libration at a critical value of the repulsive coupling strength. To explore the phenomenon, a paradigmatic second-order phase model is used to represent the dynamics of the oscillator associated with each node. We make an annealed network approximation to reduce our original model and, thereby, confirm the dual role of the repulsive interaction and the degree of a node in the origin of extreme events in any oscillator associated with a node.

Impact of Transcutaneous Auricular Vagus Nerve Stimulation on Large-Scale Functional Brain Networks: From Local to Global.

Transcutaneous auricular vagus nerve stimulation (taVNS) is a novel non-invasive brain stimulation technique considered as a potential supplementary treatment option for a wide range of diseases. Although first promising findings were obtained so far, the exact mode of action of taVNS is not fully understood yet. We recently developed an examination schedule to probe for immediate taVNS-induced modifications of large-scale epileptic brain networks. With this schedule, we observed short-term taVNS to have a topology-modifying, robustness- and stability-enhancing immediate effect on large-scale functional brain networks from subjects with focal epilepsies. We here expand on this study and investigate the impact of short-term taVNS on various local and global characteristics of large-scale evolving functional brain networks from a group of 30 subjects with and without central nervous system diseases. Our findings point to differential, at first glance counterintuitive, taVNS-mediated alterations of local and global topological network characteristics that result in a reconfiguration of networks and a modification of their stability and robustness properties. We propose a model of a stimulation-related stretching and compression of evolving functional brain networks that may help to better understand the mode of action of taVNS.

Time in Brain: How Biological Rhythms Impact on EEG Signals and on EEG-Derived Brain Networks.

Electroencephalography (EEG) is a widely employed tool for exploring brain dynamics and is used extensively in various domains, ranging from clinical diagnosis via neuroscience, cognitive science, cognitive psychology, psychophysiology, neuromarketing, neurolinguistics, and pharmacology to research on brain computer interfaces. EEG is the only technique that enables the continuous recording of brain dynamics over periods of time that range from a few seconds to hours and days and beyond. When taking long-term recordings, various endogenous and exogenous biological rhythms may impinge on characteristics of EEG signals. While the impact of the circadian rhythm and of ultradian rhythms on spectral characteristics of EEG signals has been investigated for more than half a century, only little is known on how biological rhythms influence characteristics of brain dynamics assessed with modern EEG analysis techniques. At the example of multiday, multichannel non-invasive and invasive EEG recordings, we here discuss the impact of biological rhythms on temporal changes of various characteristics of human brain dynamics: higher-order statistical moments and interaction properties of multichannel EEG signals as well as local and global characteristics of EEG-derived evolving functional brain networks. Our findings emphasize the need to take into account the impact of biological rhythms in order to avoid erroneous statements about brain dynamics and about evolving functional brain networks.

Reconfiguration of human evolving large-scale epileptic brain networks prior to seizures: an evaluation with node centralities.

Previous research has indicated that temporal changes of centrality of specific nodes in human evolving large-scale epileptic brain networks carry information predictive of impending seizures. Centrality is a fundamental network-theoretical concept that allows one to assess the role a node plays in a network. This concept allows for various interpretations, which is reflected in a number of centrality indices. Here we aim to achieve a more general understanding of local and global network reconfigurations during the pre-seizure period as indicated by changes of different node centrality indices. To this end, we investigate-in a time-resolved manner-evolving large-scale epileptic brain networks that we derived from multi-day, multi-electrode intracranial electroencephalograpic recordings from a large but inhomogeneous group of subjects with pharmacoresistant epilepsies with different anatomical origins. We estimate multiple centrality indices to assess the various roles the nodes play while the networks transit from the seizure-free to the pre-seizure period. Our findings allow us to formulate several major scenarios for the reconfiguration of an evolving epileptic brain network prior to seizures, which indicate that there is likely not a single network mechanism underlying seizure generation. Rather, local and global aspects of the pre-seizure network reconfiguration affect virtually all network constituents, from the various brain regions to the functional connections between them.

Identifying edges that facilitate the generation of extreme events in networked dynamical systems.

The collective dynamics of complex networks of FitzHugh-Nagumo units exhibits rare and recurrent events of high amplitude (extreme events) that are preceded by so-called proto-events during which a certain fraction of the units become excited. Although it is well known that a sufficiently large fraction of excited units is required to turn a proto-event into an extreme event, it is not yet clear how the other units are being recruited into the final generation of an extreme event. Addressing this question and mimicking typical experimental situations, we investigate the centrality of edges in time-dependent interaction networks. We derived these networks from time series of the units' dynamics employing a widely used bivariate analysis technique. Using our recently proposed edge-centrality concepts together with an edge-based network decomposition technique, we observe that the recruitment is primarily facilitated by sets of certain edges that have no equivalent in the underlying topology. Our finding might aid to improve the understanding of generation of extreme events in natural networked dynamical systems.

The Human Organism as an Integrated Interaction Network: Recent Conceptual and Methodological Challenges.

The field of Network Physiology aims to advance our understanding of how physiological systems and sub-systems interact to generate a variety of behaviors and distinct physiological states, to optimize the organism's functioning, and to maintain health. Within this framework, which considers the human organism as an integrated network, vertices are associated with organs while edges represent time-varying interactions between vertices. Likewise, vertices may represent networks on smaller spatial scales leading to a complex mixture of interacting homogeneous and inhomogeneous networks of networks. Lacking adequate analytic tools and a theoretical framework to probe interactions within and among diverse physiological systems, current approaches focus on inferring properties of time-varying interactions-namely strength, direction, and functional form-from time-locked recordings of physiological observables. To this end, a variety of bivariate or, in general, multivariate time-series-analysis techniques, which are derived from diverse mathematical and physical concepts, are employed and the resulting time-dependent networks can then be further characterized with methods from network theory. Despite the many promising new developments, there are still problems that evade from a satisfactory solution. Here we address several important challenges that could aid in finding new perspectives and inspire the development of theoretic and analytical concepts to deal with these challenges and in studying the complex interactions between physiological systems.

Centrality-based identification of important edges in complex networks.

Centrality is one of the most fundamental metrics in network science. Despite an abundance of methods for measuring centrality of individual vertices, there are by now only a few metrics to measure centrality of individual edges. We modify various, widely used centrality concepts for vertices to those for edges, in order to find which edges in a network are important between other pairs of vertices. Focusing on the importance of edges, we propose an edge-centrality-based network decomposition technique to identify a hierarchy of sets of edges, where each set is associated with a different level of importance. We evaluate the efficiency of our methods using various paradigmatic network models and apply the novel concepts to identify important edges and important sets of edges in a commonly used benchmark model in social network analysis, as well as in evolving epileptic brain networks.