Based on the data of the NIH-funded Human Connectome Project, we have computed structural connectomes of 426 human subjects in five different resolutions of 83, 129, 234, 463 and 1015 nodes and several edge weights. The graphs are given in anatomically annotated GraphML format that facilitates better further processing and visualization. For 96 subjects, the anatomically classified sub-graphs can also be accessed, formed from the vertices corresponding to distinct lobes or even smaller regions of interests of the brain. For example, one can easily download and study the connectomes, restricted to the frontal lobes or just to the left precuneus of 96 subjects using the data. Partially directed connectomes of 423 subjects are also available for download. We also present a GitHub-deposited set of tools, called the Brain Graph Tools, for several processing tasks of the connectomes on the site \url{http://braingraph.org}.
The human brain is the most complex object of study we encounter today. Mapping the neuronal-level connections between the more than 80 billion neurons in the brain is a hopeless task for science. By the recent advancement of magnetic resonance imaging (MRI), we are able to map the macroscopic connections between about 1000 brain areas. The MRI data acquisition and the subsequent algorithmic workflow contain several complex steps, where errors can occur. In the present contribution, we describe and publish 1064 human connectomes, computed from the public release of the Human Connectome Project…
Here we show a method of directing the edges of the connectomes, prepared from diffusion tensor imaging (DTI) datasets from the human brain. Before the present work, no high-definition directed braingraphs (or connectomes) were published, because the tractography methods in use are not capable of assigning directions to the neural tracts discovered. Previous work on the functional connectomes applied low-resolution functional MRI-detected statistical causality for the assignment of directions of connectomes of typically several dozens of vertices. Our method is based on the phenomenon of the "…
Gaussian blurring is a well-established method for image data augmentation: it may generate a large set of images from a small set of pictures for training and testing purposes for Artificial Intelligence (AI) applications. When we apply AI for non-imagelike biological data, hardly any related method exists. Here we introduce the "Newtonian blurring" in human braingraph (or connectome) augmentation: Started from a dataset of 1053 subjects, we first repeat a probabilistic weighted braingraph construction algorithm 10 times for describing the connections of distinct cerebral areas, then take 7 r…
For more than a decade now, we can discover and study thousands of cerebral connections with the application of diffusion magnetic resonance imaging (dMRI) techniques and the accompanying algorithmic workflow. While numerous connectomical results were published enlightening the relation between the braingraph and certain biological, medical, and psychological properties, it is still a great challenge to identify a small number of brain connections, closely related to those conditions. In the present contribution, by applying the 1200 Subjects Release of the Human Connectome Project (HCP), we i…
In the applications of the graph theory it is unusual that one considers numerous, pairwise different graphs on the very same set of vertices. In the case of human braingraphs or connectomes, however, this is the standard situation: the nodes correspond to anatomically identified cerebral regions, and two vertices are connected by an edge if a diffusion MRI-based workflow identifies a fiber of axons, running between the two regions, corresponding to the two vertices. Therefore, if we examine the braingraphs of $n$ subjects, then we have $n$ graphs on the very same, anatomically identified vert…
Connections of the living human brain, on a macroscopic scale, can be mapped by a diffusion MR imaging based workflow. Since the same anatomic regions can be corresponded between distinct brains, one can compare the presence or the absence of the edges, connecting the very same two anatomic regions, among multiple cortices. Previously, we have constructed the consensus braingraphs on 1015 vertices first in five, then in 96 subjects in the Budapest Reference Connectome Server v1.0 and v2.0, respectively. Here we report the construction of the version 3.0 of the server, generating the common edg…
Determining important vertices in large graphs (e.g., Google's PageRank in the case of the graph of the World Wide Web) facilitated the construction of excellent web search engines, returning the most important hits corresponding to the submitted user queries. Interestingly, finding important edges -- instead of vertices -- in large graphs has received much less attention until now. Here we examine the human structural braingraph (or connectome), identified by diffusion magnetic resonance imaging (dMRI) methods, with edges connecting cortical and subcortical gray matter areas and weighted by f…
Graph theory in the last two decades penetrated sociology, molecular biology, genetics, chemistry, computer engineering, and numerous other fields of science. One of the more recent areas of its applications is the study of the connections of the human brain. By the development of diffusion magnetic resonance imaging (diffusion MRI), it is possible today to map the connections between the 1-1.5 cm$^2$ regions of the gray matter of the human brain. These connections can be viewed as a graph: the vertices are the anatomically identified regions of the gray matter, and two vertices are connected …
In our previous study we have shown that the female connectomes have significantly better, deep graph-theoretical parameters, related to superior "connectivity", than the connectome of the males. Since the average female brain is smaller than the average male brain, one cannot rule out that the significant advantages are due to the size- and not to the sex-differences in the data. To filter out the possible brain-volume related artifacts, we have chosen 36 small male and 36 large female brains such that all the brains in the female set are larger than all the brains in the male set. For the se…
While it is still not possible to describe the neural-level connections of the human brain, we can map the human connectome with several hundred vertices, by the application of diffusion-MRI based techniques. In these graphs, the nodes correspond to anatomically identified gray matter areas of the brain, while the edges correspond to the axonal fibers, connecting these areas. In our previous contributions, we have described numerous graph-theoretical phenomena of the human connectomes. Here we map the frequent complete subgraphs of the human brain networks: in these subgraphs, every pair of ve…
The human connectome has become the very frequent subject of study of brain-scientists, psychologists, and imaging experts in the last decade. With diffusion magnetic resonance imaging techniques, unified with advanced data processing algorithms, today we are able to compute braingraphs with several hundred, anatomically identified nodes and thousands of edges, corresponding to the anatomical connections of the brain. The analysis of these graphs without refined mathematical tools is hopeless. These tools need to address the high error rate of the MRI processing workflow, and need to find stru…
Deep, classical graph-theoretical parameters, like the size of the minimum vertex cover, the chromatic number, or the eigengap of the adjacency matrix of the graph were studied widely by mathematicians in the last century. Most researchers today study much simpler parameters of braingraphs or connectomes which were defined in the last twenty years for enormous networks -- like the graph of the World Wide Web -- with hundreds of millions of nodes. Since the connectomes, describing the connections of the human brain, typically contain several hundred vertices today, one can compute and analyze t…
The human connectome is the object of an intensive research today. In these graphs, the vertices correspond to the small areas of the gray matter, and two vertices are connected by an edge, if a diffusion-MRI based workflow finds connections between those areas. One main question of the field is discovering the directions of the edges. In a previous work we have reported the construction of the Budapest Reference Connectome Server http://connectome.pitgroup.org from the data recorded in the Human Connectome Project of the NIH. After the server had been published, we recognized a surprising and…
In the study of the human connectome, the vertices and the edges of the network of the human brain are analyzed: the vertices of the graphs are the anatomically identified gray matter areas of the subjects; this set is exactly the same for all the subjects. The edges of the graphs correspond to the axonal fibers, connecting these areas. In the biological applications of graph theory, it happens very rarely that scientists examine numerous large graphs on the very same, labeled vertex set. Exactly this is the case in the study of the connectomes. Because of the particularity of these sets of gr…
Investigations of the human connectome have elucidated core features of adult structural networks, particularly the crucial role of hub-regions. However, little is known regarding network organisation of the healthy elderly connectome, a crucial prelude to the systematic study of neurodegenerative disorders. Here, whole-brain probabilistic tractography was performed on high-angular diffusion-weighted images acquired from 115 healthy elderly subjects, whom were 76 to 94 years old. Structural networks were reconstructed between 512 cortical and subcortical brain regions. We sought to investigate…
The human connectome has been widely studied over the past decade. A principal finding is that it can be decomposed into communities of densely interconnected brain regions. This result, however, may be limited methodologically. Past studies have often used a flawed modularity measure in order to infer the connectome's community structure. Also, these studies relied on the intuition that community structure is best defined in terms of a network's static topology as opposed to a more dynamical definition. In this report we used the partition stability framework, which defines communities in ter…
The human braingraph, or connectome is a description of the connections of the brain: the nodes of the graph correspond to small areas of the gray matter, and two nodes are connected by an edge if a diffusion MRI-based workflow finds fibers between those brain areas. We have constructed 1015-vertex graphs from the diffusion MRI brain images of 395 human subjects and compared the individual graphs with respect to several different areas of the brain. The inter-individual variability of the graphs within different brain regions was discovered and described. We have found that the frontal and the…
The human connectome represents a network map of the brain's wiring diagram and the pattern into which its connections are organized is thought to play an important role in cognitive function. The generative rules that shape the topology of the human connectome remain incompletely understood. Earlier work in model organisms has suggested that wiring rules based on geometric relationships (distance) can account for many but likely not all topological features. Here we systematically explore a family of generative models of the human connectome that yield synthetic networks designed according to…
The connectomes of different human brains are pairwise distinct: we cannot talk about an abstract "graph of the brain". Two typical connectomes, however, have quite a few common graph edges that may describe the same connections between the same cortical areas. The Budapest Reference Connectome Server Ver. 2.0 (http://connectome.pitgroup.org) generates the common edges of the connectomes of 96 distinct cortexes, each with 1015 vertices, computed from 96 MRI data sets of the Human Connectome Project. The user may set numerous parameters for the identification and filtering of common edges, and …
Anatomical connectivity imposes strong constraints on brain function, but there is no general agreement about principles that govern its organization. Based on extensive quantitative data we tested the power of three models to predict connections of the primate cerebral cortex: architectonic similarity (structural model), spatial proximity (distance model) and thickness similarity (thickness model). Architectonic similarity showed the strongest and most consistent influence on connection features. This parameter was strongly associated with the presence or absence of inter-areal connections an…
Analyzing the brain in terms of organizational structures at intermediate scales provides an approach to negotiate the complexity arising from interactions between its large number of components. Focusing on a wiring diagram that spans the cortex, basal ganglia and thalamus of the Macaque brain, we provide a mesoscopic-level description of the topological architecture of one of the most well-studied mammalian connectomes. The robust modules we identify each comprise densely inter-connected cortical and sub-cortical areas that play complementary roles in executing specific cognitive functions. …
The connectome, or the entire connectivity of a neural system represented by network, ranges various scales from synaptic connections between individual neurons to fibre tract connections between brain regions. Although the modularity they commonly show has been extensively studied, it is unclear whether connection specificity of such networks can already be fully explained by the modularity alone. To answer this question, we study two networks, the neuronal network of C. elegans and the fibre tract network of human brains yielded through diffusion spectrum imaging (DSI). We compare them to th…
In order to understand the complex cognitive functions of the human brain, it is essential to study the structural connectome, i.e., the wiring of different brain regions to each other through axonal pathways. However, the high degree of plasticity and cross-population variability in human brains makes it difficult to relate structure to function, motivating a search for invariant patterns in the connectivity. At the same time, variability within a population can provide information about generative mechanisms. In this paper we analyze the connection topology and link-weight distribution of hu…
One of the crucial questions in neuroscience is how a rich functional repertoire of brain states relates to its underlying structural organization. How to study the associations between these structural and functional layers is an open problem that involves novel conceptual ways of tackling this question. We here propose an extension of the Connectivity Independent Component Analysis (connICA) framework, to identify joint structural-functional connectivity traits. Here, we extend connICA to integrate structural and functional connectomes by merging them into common hybrid connectivity patterns…
Databases of directed- and weighted- connectivity for mouse, macaque and marmoset monkeys, have recently become available and begun to be used to build dynamical models. A hierarchical organization can be defined based on laminar patterns of cortical connections, possibly improved by thalamocortical projections. A large-scale model of the macaque cortex endowed with a laminar structure accounts for layer-dependent and frequency-modulated interplays between bottom-up and top-down processes. Signal propagation in a version of the model with spiking neurons displays a threshold of stimulus amplit…
The increasing quantity and quality of the publicly available human cerebral diffusion MRI data make possible the study of the brain as it was unimaginable before. The Consensus Connectome Dynamics (CCD) is a remarkable phenomenon that was discovered by continuously decreasing the minimum confidence-parameter at the graphical interface of the Budapest Reference Connectome Server (\url{http://connectome.pitgroup.org}). The Budapest Reference Connectome Server depicts the cerebral connections of $n=418$ subjects with a frequency-parameter $k$: For any $k=1,2,...,n$ one can view the graph of the …
Overlooked weak structural connections support human cognition under nonlinear connectome scaling
Xi’an Jiaotong University（西安交通大学）；Hong Kong Baptist University（香港浸会大学）；Carl von Ossietzky Universität Oldenburg（奥尔登堡卡尔·冯·奥西茨基大学）；University of California San Francisco（加州大学旧金山分校）；Shanghai Artificial Intelligence Laboratory（上海人工智能实验室）；University of Roehampton（罗汉普顿大学）；Zhejiang University（浙江大学）
We have performed the comparative spectral analysis of structural connectomes for various organisms using open-access data. Our analysis indicates several new peculiar features of the human connectome. We found that the spectral density of human connectome has the maximal deviation from the spectral density of the randomized network compared to all other organisms. For many animals except human structural peculiarities of connectomes are well reproduced in the network evolution induced by the preference of 3-cycles formation. To get the reliable fit , we discovered the crucial role of the cons…
Autism spectrum disorder (ASD) is a neurodevelopmental condition impacting high-level cognitive processing and social behavior. Recognizing the distributed nature of brain function, neuroscientists are exploiting the connectome to aid with the characterization of this complex disease. The human connectome has demonstrated the brain to be a highly organized system with a centralized core vital for effective function. As such, many have used this topological principle to not only assess core regions, but have stratified the remaining graph into subnetworks depending on their relation to the core…
The relation between large-scale brain structure and function is an outstanding open problem in neuroscience. We approach this problem by studying the dynamical regime under which realistic spatio-temporal patterns of brain activity emerge from the empirically derived network of human brain neuroanatomical connections. The results show that critical dynamics unfolding on the structural connectivity of the human brain allow the recovery of many key experimental findings obtained with functional Magnetic Resonance Imaging (fMRI), such as divergence of the correlation length, anomalous scaling of…
Structural connectivity in the brain is typically studied by reducing its observation to a single spatial resolution. However, the brain possesses a rich architecture organized over multiple scales linked to one another. We explored the multiscale organization of human connectomes using datasets of healthy subjects reconstructed at five different resolutions. We found that the structure of the human brain remains self-similar when the resolution of observation is progressively decreased by hierarchical coarse-graining of the anatomical regions. Strikingly, a geometric network model, where dist…
Estimated connectomes by the means of neuroimaging techniques have enriched our knowledge of the organizational properties of the brain leading to the development of network-based clinical diagnostics. Unfortunately, to date, many of those network-based clinical diagnostics tools, based on the mere description of isolated instances of observed connectomes are noisy estimates of the true connectivity network. Modeling brain connectivity networks is therefore important to better explain the functional organization of the brain and allow inference of specific brain properties. In this report, we …
High-throughput methods for yielding the set of connections in a neural system, the connectome, are now being developed. This tutorial describes ways to analyze the topological and spatial organization of the connectome at the macroscopic level of connectivity between brain regions as well as the microscopic level of connectivity between neurons. We will describe topological features at three different levels: the local scale of individual nodes, the regional scale of sets of nodes, and the global scale of the complete set of nodes in a network. Such features can be used to characterize compon…
Yale University（耶鲁大学）；Yale School of Medicine（耶鲁医学院）
Functional connectomes derived from functional magnetic resonance imaging have long been used to understand the functional organization of the brain. Nevertheless, a connectome is intrinsically linked to the atlas used to create it. In other words, a connectome generated from one atlas is different in scale and resolution compared to a connectome generated from another atlas. Being able to map connectomes and derived results between different atlases without additional pre-processing is a crucial step in improving interpretation and generalization between studies that use different atlases. He…
The connectome is a wiring diagram mapping all the neural connections in the brain. At the cellular level, it provides a map of the neurons and synapses within a part or all of the brain of an organism. In recent years, significant advances have been made in the study of the connectome via network science and graph theory. This analysis is fundamental to understand neurotransmission (fast synaptic transmission) networks. However, neurons use other forms of communication as neuromodulation that, instead of conveying excitation or inhibition, change neuronal and synaptic properties. This additio…
The large-scale structural ingredients of the brain and neural connectomes have been identified in recent years. These are, similar to the features found in many other real networks: the arrangement of brain regions into modules and the presence of highly connected regions (hubs) forming rich-clubs. Here, we examine how modules and hubs shape the collective dynamics on networks and we find that both ingredients lead to the emergence of complex dynamics. Comparing the connectomes of C. elegans, cats, macaques and humans to surrogate networks in which either modules or hubs are destroyed, we fin…
The human connectome at the level of fiber tracts between brain regions has been shown to differ in patients with brain disorders compared to healthy control groups. Nonetheless, there is a potentially large number of different network organizations for individual patients that could lead to cognitive deficits prohibiting correct diagnosis. Therefore changes that can distinguish groups might not be sufficient to diagnose the disease that an individual patient suffers from and to indicate the best treatment option for that patient. We describe the challenges introduced by the large variability …
Neuropsychiatric disorders impact functional connectivity of the brain at the network level. The identification and statistical testing of disorder-related networks remains challenging. We propose novel methods to streamline the detection and testing of the hidden, disorder-related connectivity patterns as network-objects. We define an abnormal connectome subnetwork as a network-object that includes three classes: nodes of brain areas, edges representing brain connectomic features, and an organized graph topology formed by these nodes and edges. Comparing to the conventional statistical method…
The behavior of many complex systems is determined by a core of densely interconnected units. While many methods are available to identify the core of a network when connections between nodes are all of the same type, a principled approach to define the core when multiple types of connectivity are allowed is still lacking. Here we introduce a general framework to define and extract the core-periphery structure of multi-layer networks by explicitly taking into account the connectivity of the nodes at each layer. We show how our method works on synthetic networks with different size, density, an…
To meet ongoing cognitive demands, the human brain must seamlessly transition from one brain state to another, in the process drawing on different cognitive systems. How does the brain's network of anatomical connections help facilitate such transitions? Which features of this network contribute to making one transition easy and another transition difficult? Here, we address these questions using network control theory. We calculate the optimal input signals to drive the brain to and from states dominated by different cognitive systems. The input signals allow us to assess the contributions ma…
Human brain structural networks contain sets of centrally embedded hub regions that enable efficient information communication. However, it remains largely unknown about categories of structural brain hubs and their microstructural, functional and cognitive characteristics as well as contributions to individual identification. Here, we employed three multi-modal imaging data sets with structural MRI, diffusion MRI and resting-state functional MRI to construct individual structural brain networks, identify brain hubs based on eight commonly used graph-nodal metrics, and perform comprehensive va…
A major challenge in neuroscience is posed by the need for relating the emerging dynamical features of brain activity with the underlying modular structure of neural connections, hierarchically organized throughout several scales. The spontaneous emergence of coherence and synchronization across such scales is crucial to neural function, while its anomalies often relate to pathological conditions. Here we provide a numerical study of synchronization dynamics in the human connectome network. Our purpose is to provide a detailed characterization of the recently uncovered broad dynamic regime, in…
Examining task-free functional connectivity (FC) in the human brain offers insights on how spontaneous integration and segregation of information relate to human cognition, and how this organization may be altered in different conditions, and neurological disorders. This is particularly relevant for patients in disorders of consciousness (DOC) following severe acquired brain damage and coma, one of the most devastating conditions in modern medical care. We present a novel data-driven methodology, connICA, which implements Independent Component Analysis (ICA) for the extraction of robust indepe…
Mapping the connectivity of neurons in the brain (i.e., connectomics) is a challenging problem due to both the number of connections in even the smallest organisms and the nanometer resolution required to resolve them. Because of this, previous connectomes contain only hundreds of neurons, such as in the C.elegans connectome. Recent technological advances will unlock the mysteries of increasingly large connectomes (or partial connectomes). However, the value of these maps is limited by our ability to reason with this data and understand any underlying motifs. To aid connectome analysis, we int…
École Polytechnique Fédérale de Lausanne (EPFL)（洛桑联邦理工学院）；University of Geneva（日内瓦大学）
Spectral approaches of network analysis heavily rely upon the eigendecomposition of the graph Laplacian. For instance, in graph signal processing, the Laplacian eigendecomposition is used to define the graph Fourier transform and then transpose signal processing operations to graphs by implementing them in the spectral domain. Here, we build on recent work that generalized Slepian functions to the graph setting. In particular, graph Slepians are band-limited graph signals with maximal energy concentration in a given subgraph. We show how this approach can be used to guide network analysis; i.e…
Connectomes are spatially embedded networks whose architecture has been shaped by physical constraints and communication needs throughout evolution. Using a decentralized navigation protocol, we investigate the relationship between the structure of the connectomes of different species and their spatial layout. As a navigation strategy, we use greedy routing where nearest neighbors, in terms of geometric distance, are visited. We measure the fraction of successful greedy paths and their length as compared to shortest paths in the topology of connectomes. In Euclidean space, we find a striking d…
Understanding the modularity of fMRI-derived brain networks or connectomes can inform the study of brain function organization. However, fMRI connectomes additionally involve negative edges, which are not rigorously accounted for by existing approaches to modularity that either ignores or arbitrarily weight these connections. Furthermore, most Q maximization-based modularity algorithms yield variable results with suboptimal reproducibility. Here we present an alternative, reproducible approach that exploits how frequent the BOLD-signal correlation between two nodes is negative. We validated th…
Mapping the brain imaging data to networks, where each node represents a specific area of the brain, has enabled an objective graph-theoretic analysis of human connectome. However, the latent structure on higher-order connections remains unexplored, where many brain regions acting in synergy perform complex functions. Here we analyse this hidden structure using the simplicial complexes parametrisation where the shared faces of simplexes encode higher-order relationships between groups of nodes and emerging hyperbolic geometry. Based on data collected within the Human Connectome Project, we per…
The spontaneous emergence of coherent behavior through synchronization plays a key role in neural function, and its anomalies often lie at the basis of pathologies. Here we employ a parsimonious (mesoscopic) approach to study analytically and computationally the synchronization (Kuramoto) dynamics on the actual human-brain connectome network. We elucidate the existence of a so-far-uncovered intermediate phase, placed between the standard synchronous and asynchronous phases, i.e. between order and disorder. This novel phase stems from the hierarchical modular organization of the connectome. Whe…
The impact of developmental and aging processes on brain connectivity and the connectome has been widely studied. Network theoretical measures and certain topological principles are computed from the entire brain, however there is a need to separate and understand the underlying subnetworks which contribute towards these observed holistic connectomic alterations. One organizational principle is the rich-club - a core subnetwork of brain regions that are strongly connected, forming a high-cost, high-capacity backbone that is critical for effective communication in the network. Investigations pr…
Hierarchical Multiscale Structure-Function Coupling for Brain Connectome Integration
School of Medicine, University of Dundee, UK（邓迪大学医学院）；School of Science and Engineering, University of Dundee, UK（邓迪大学科学与工程学院）；College of Medicine and Biological Information Engineering, Northeastern University, China（东北大学医学院与生物信息工程学院）；College of Medical Informatics, Chongqing Medical University, China（重庆医科大学医学信息学院）；School of Airspace Science and Engineering, Shandong University, China（山东大学航空航天科学与工程学院）；Chinese Institute for Brain Research, China（中国脑科学研究院）；Department of Applied Mathematics and Theoretical Physics, University of Cambridge, UK（剑桥大学应用数学与理论物理系）；Department of Clinical Neurosciences, University of Cambridge, UK（剑桥大学临床神经科学系）
Extended numerical simulations of threshold models have been performed on a human brain network with N=836733 connected nodes available from the Open Connectome project. While in case of simple threshold models a sharp discontinuous phase transition without any critical dynamics arises, variable thresholds models exhibit extended power-law scaling regions. This is attributed to fact that Griffiths effects, stemming from the topological/interaction heterogeneity of the network, can become relevant if the input sensitivity of nodes is equalized. I have studied the effects effects of link directn…
