Many studies have shown that the excitation and inhibition received by cortical neurons remain roughly balanced across many conditions. A key question for understanding the dynamical regime of cortex is the nature of this balancing. Theorists have shown that network dynamics can yield systematic cancellation of most of a neuron's excitatory input by inhibition. We review a wide range of evidence pointing to this cancellation occurring in a regime in which the balance is loose, meaning that the net input remaining after cancellation of excitation and inhibition is comparable in size to the factors that cancel, rather than tight, meaning that the net input is very small relative to the cancelling factors. This choice of regime has important implications for cortical functional responses, as we describe: loose balance, but not tight balance, can yield many nonlinear population behaviors seen in sensory cortical neurons, allow the presence of correlated variability, and yield decrease of that variability with increasing external stimulus drive as observed across multiple cortical areas.
The electric activities of cortical pyramidal neurons are supported by structurally stable, morphologically complex axo-dendritic trees. Anatomical differences between axons and dendrites in regard to their length or caliber reflect the underlying functional specializations, for input or output of neural information, respectively. For a proper assessment of the computational capacity of pyramidal neurons, we have analyzed an extensive dataset of three-dimensional digital reconstructions from the NeuroMorpho.Org database, and quantified basic dendritic or axonal morphometric measures in differe…
In this paper the method and performance data of 'Boundary Element Method (BEM)'-based SMS-LORETA (Simultaneous Multiple Sources LORETA) are presented. According to these data the method is capable of locating efficiently multiple simultaneously active neural sources from scalp potential topographies automatically. BEM-based SMS-LORETA is a procedure to fully interpret sLORETA solutions, i.e., with a given scalp potential distribution it gives the number of identifiable sources as well as their strength and orientation. Performance data result from numerous analyses of simulated noise-free and…
Despite differences in brain sizes and cognitive niches among mammals, their cerebral cortices posses many common features and regularities. These regularities have been a subject of experimental investigation in neuroanatomy for the last 100 years. It is believed that such studies may provide clues about cortical design principles and perhaps function. However, on a theoretical side there has been little interest, until recently, in studying quantitatively these regularities. This article reviews some attempts in this direction with an emphasis on neuronal connectivity. It is suggested that t…
Neural correlations during a cognitive task are central to study brain information processing and computation. However, they have been poorly analyzed due to the difficulty of recording simultaneous single neurons during task performance. In the present work, we quantified neural directional correlations using spike trains that were simultaneously recorded in sensory, premotor, and motor cortical areas of two monkeys during a somatosensory discrimination task. Upon modeling spike trains as binary time series, we used a nonparametric Bayesian method to estimate pairwise directional correlations…
Mammalian brains span about 4 orders of magnitude in cortical volume and have to operate in different environments that require diverse behavioral skills. Despite these geometric and behavioral diversities, the examination of cerebral cortex across species reveals that it contains a substantial number of conserved characteristics that are associated with neuroanatomy and metabolism, i.e. with neuronal connectivity and function. Some of these cortical constants or invariants have been known for a long time but not sufficiently appreciated, and others were only recently discovered. The focus of …
We model the elasticity of the cerebral cortex as a layered material with bending energy along the layers and elastic energy between them in both planar and polar geometries. The cortex is also subjected to axons pulling from the underlying white matter. Above a critical threshold force, a "flat" cortex configuration becomes unstable and periodic unduluations emerge, i.e. a buckling instability occurs. These undulations may indeed initiate folds in the cortex. We identify analytically the critical force and the critical wavelength of the undulations. Both quantities are physiologically relevan…
We have studied effects of perturbations on the cat cerebral cortex. According to the literature, this cortex structure can be described by a clustered network. This way, we construct a clustered network with the same number of areas as in the cat matrix, where each area is described as a sub-network with small-world property. We focus on the suppression of neuronal phase synchronisation considering different kinds of perturbations. Among the various controlling interventions, we choose three methods: delayed feedback control, external time-periodic driving, and activation of selected neurons.…
The frequency-specific coupling mechanism of the functional human brain networks underpins its complex cognitive and behavioral functions. Nevertheless, it is not well unveiled what are the frequency-specific subdivisions and network topologies of the human brain. In this study, we estimated functional connectivity of the human cerebral cortex using spectral connection, and conducted frequency-specific parcellation using eigen-clustering and gradient-based methods, and then explored their topological structures. 7T fMRI data of 184 subjects in the HCP dataset were used for parcellation and exp…
Recent studies have pointed out the importance of transient synchronization between widely distributed neural assemblies to understand conscious perception. These neural assemblies form intricate networks of neurons and synapses whose detailed map for mammals is still unknown and far from our experimental capabilities. Only in a few cases, for example the C. elegans, we know the complete mapping of the neuronal tissue or its mesoscopic level of description provided by cortical areas. Here we study the process of transient and global synchronization using a simple model of phase-coupled oscilla…
Imperial College of Science, Technology and Medicine（帝国理工学院（科学、技术与医学））；Imperial College London（伦敦帝国学院）
In this thesis, we present robust and fully-automated methods for the subdivision of the entire human cerebral cortex based on connectivity information. Our contributions are four-fold: First, we propose a clustering approach to delineate a cortical parcellation that provides a reliable abstraction of the brain's functional organisation. Second, we cast the parcellation problem as a feature reduction problem and make use of manifold learning and image segmentation techniques to identify cortical regions with distinct structural connectivity patterns. Third, we present a multi-layer graphical m…
Cortical slow oscillations are an emergent property of the cortical network, a hallmark of low complexity brain states like sleep, and represent a default activity pattern. Here, we present a methodological approach for quantifying the spatial and temporal properties of this emergent activity. We improved and enriched a robust analysis procedure that has already been successfully applied to both in vitro and in vivo data acquisitions. We tested the new tools of the methodology by analyzing the electrocorticography (ECoG) traces recorded from a custom 32-channel multi-electrode array in wild-ty…
CytoNet: A Foundation Model for the Human Cerebral Cortex at Cellular Resolution
Institute of Neuroscience and Medicine (INM-1), Research Centre Jülich（神经科学与医学研究所（INM-1），贾格尔研究中心）；Helmholtz AI, Research Centre Jülich（海德堡人工智能，贾格尔研究中心）；Institute for Brain Research, University Hospital Düsseldorf（脑研究所在杜塞尔多夫大学医院）；Institute of Computational Biology, Computational Health Center, Helmholtz Munich（计算生物学研究所，计算健康中心，海德堡慕尼黑）；Institute for Stroke and Dementia Research (ISD), LMU University Hospital, LMU Munich（中风与痴呆研究所以（ISD），慕尼黑大学医院，慕尼黑大学）；Computer Vision, Institute for Computational Visualistics, University of Koblenz（计算机视觉，计算视觉研究所，科布伦兹大学）
Self-organized critical states are found in many natural systems, from earthquakes to forest fires, they have also been observed in neural systems, particularly, in neuronal cultures. However, the presence of critical states in the awake brain remains controversial. Here, we compared avalanche analyses performed on different in vivo preparations during wakefulness, slow-wave sleep and REM sleep, using high-density electrode arrays in cat motor cortex (96 electrodes), monkey motor cortex and premotor cortex and human temporal cortex (96 electrodes) in epileptic patients. In neuronal avalanches …
Sleep slow waves are known to participate in memory consolidation, yet slow waves occurring under anesthesia present no positive effects on memory. Here, we shed light onto this paradox, based on a combination of extracellular recordings in vivo, in vitro, and computational models. We find two types of slow waves, based on analyzing the temporal patterns of successive slow-wave events. The first type is consistently observed in natural slow-wave sleep, while the second is shown to be ubiquitous under anesthesia. Network models of spiking neurons predict that the two slow wave types emerge due …
The parcellation of the primate cerebral cortex into numbered regions, based on cytoarchitecture, began with the pioneering research of neuroanatomist Kobrinian Brodmann. While the borders between regions have changed somewhat, and in some cases continue to be disputed, the idea of dividing the cortex into distinct numbered areas has become central to the goal of understanding brain function. And indeed, areal maps do provide a good starting point for functional parcellation. It is obvious, for example, that visual area V1 has a different function than primary motor cortex. However, as with an…
It has been hypothesized that Gamma cortical oscillations play important roles in numerous cognitive processes and may involve psychiatric conditions including anxiety, schizophrenia, and autism. Gamma rhythms are commonly observed in many brain regions during both waking and sleep states, yet their functions and mechanisms remain a matter of debate. Spatiotemporal Gamma oscillations can explain neuronal representation, computation, and the shaping of communication among cortical neurons, even neurological and neuropsychiatric disorders in neo-cortex. In this study, the neural network dynamics…
One of the most well-established tools for modeling the brain as a complex system is the functional connectivity network, which examines the correlations between pairs of interacting brain regions. While powerful, the network model is limited by the restriction that only pairwise dependencies are visible and potentially higher-order structures are missed. In this work, we explore how multivariate information theory can reveal higher-order, synergistic dependencies in the human brain. Using the O-information, a measure of whether the structure of a system is redundancy- or synergy-dominated, we…
An Operating Principle of the Cerebral Cortex, and a Cellular Mechanism for Attentional Trial-and-Error Pattern Learning and Useful Classification Extraction
Functions of brain areas in complex animals are believed to rely on the dynamics of networks of neurons rather than on single neurons. On the other hand, the network dynamics reflect and arise from the integration and coordination of the activity of populations of single neurons. Understanding how single-neurons and neural-circuits dynamics complement each other to produce brain functions is thus of paramount importance. LFPs and EEGs are good indicators of the dynamics of mesoscopic and macroscopic populations of neurons, while microscopic-level activities can be documented by measuring the m…
In awake animals, the activity of the cerebral cortex is highly complex, with neurons firing irregularly with apparent Poisson statistics. One way to characterize this complexity is to take advantage of the high interconnectivity of cerebral cortex and use intracellular recordings of cortical neurons, which contain information about the activity of thousands of other cortical neurons. Identifying the membrane potential (Vm) to a stochastic process enables the extraction of important statistical signatures of this complex synaptic activity. Typically, one estimates the total synaptic conductanc…
Cortical network functioning critically depends on finely tuned interactions to afford neuronal activity propagation over long distances while avoiding runaway excitation. This importance is highlighted by the pathological consequences and impaired performance resulting from aberrant network excitability in psychiatric and neurological diseases, such as epilepsy. Theory and experiment suggest that the control of activity propagation by network interactions can be adequately described by a branching process. This hypothesis is partially supported by strong evidence for balanced spatiotemporal d…
The analysis of complex networks has revealed patterns of organization in a variety of natural and artificial systems, including neuronal networks of the brain at multiple scales. In this paper, we describe a novel analysis of the large-scale connectivity between regions of the mammalian cerebral cortex, utilizing a set of hierarchical measurements proposed recently. We examine previously identified functional clusters of brain regions in macaque visual cortex and cat cortex and find significant differences between such clusters in terms of several hierarchical measures, revealing differences …
An essential requirement for the representation of functional patterns in complex neural networks, such as the mammalian cerebral cortex, is the existence of stable network activations within a limited critical range. In this range, the activity of neural populations in the network persists between the extremes of quickly dying out, or activating the whole network. The nerve fiber network of the mammalian cerebral cortex possesses a modular organization extending across several levels of organization. Using a basic spreading model without inhibition, we investigated how functional activations …
Several studies suggest that cyclooxygenase (COX)-2 plays a pivotal role in the progression of ischaemic brain damage. In the present study, we investigated the effects of selective inhibition of COX-2 with nimesulide (12 mg/kg) and selective inhibition of COX-1 with valeryl salicylate (VAS, 12-120 mg/kg) on prostaglandin E2 (PGE2) levels, myeloperoxidase (MPO) activity, Evans blue (EB) extravasation and infarct volume in a standardized model of transient focal cerebral ischaemia in the rat. Post-ischaemic treatment with nimesulide markedly reduced the increase in PGE2 levels in the ischaemic …
UvA-Bosch Delta Lab（阿姆斯特丹大学-博世三角洲实验室）；University of Amsterdam（阿姆斯特丹大学）；ChemEngAI Lab（化学工程人工智能实验室）；Delft University of Technology（代尔夫特理工大学）
Category-selectivity in the brain describes the observation that certain spatially localized areas of the cerebral cortex tend to respond robustly and selectively to stimuli from specific limited categories. One of the most well known examples of category-selectivity is the Fusiform Face Area (FFA), an area of the inferior temporal cortex in primates which responds preferentially to images of faces when compared with objects or other generic stimuli. In this work, we leverage the newly introduced Topographic Variational Autoencoder to model the emergence of such localized category-selectivity …
Randomly-connected networks of integrate-and-fire (IF) neurons are known to display asynchronous irregular (AI) activity states, which resemble the discharge activity recorded in the cerebral cortex of awake animals. However, it is not clear whether such activity states are specific to simple IF models, or if they also exist in networks where neurons are endowed with complex intrinsic properties similar to electrophysiological measurements. Here, we investigate the occurrence of AI states in networks of nonlinear IF neurons, such as the adaptive exponential IF (Brette-Gerstner-Izhikevich) mode…
The traditional view of neural computation in the cerebral cortex holds that sensory neurons are specialized, i.e., selective for certain dimensions of sensory stimuli. This view was challenged by evidence of contextual interactions between stimulus dimensions in which a neuron's response to one dimension strongly depends on other dimensions. Here we use methods of mathematical modeling, psychophysics, and electrophysiology to address shortcomings of the traditional view. Using a model of a generic cortical circuit, we begin with the simple demonstration that cortical responses are always dist…
Recent experiments revealed that a certain class of inhibitory neurons in the cerebral cortex make synapses not onto cell bodies but at distal parts of dendrites of the target neurons, mediating highly nonlinear dendritic inhibition. We propose a novel form of competitive neural network model that realizes such dendritic inhibition. Contrary to the conventional lateral inhibition in neural networks, our dendritic inhibition models don't always show winner-take-all behaviors; instead, they converge to "I don't know" states when unknown input patterns are presented. We derive reduced two-dimensi…
Primary visual cortex (V1) provides crucial insights into the selectivity and emergence of specific output features such as orientation tuning. Tuning and selectivity of cortical neurons in mouse visual cortex is not equivocally resolved so far. While many in-vivo experimental studies found inhibitory neurons of all subtypes to be broadly tuned for orientation other studies report inhibitory neurons that are as sharply tuned as excitatory neurons. These diverging findings about the selectivity of excitatory and inhibitory cortical neurons prompted us to ask the following questions: (1) How dif…
We present a hybrid method that performs the complete parcellation of the cerebral cortex of an individual, based on the connectivity information of the white matter fibers from a whole-brain tractography dataset. The method consists of five steps, first intra-subject clustering is performed on the brain tractography. The fibers that make up each cluster are then intersected with the cortical mesh and then filtered to discard outliers. In addition, the method resolves the overlapping between the different intersection regions (sub-parcels) throughout the cortex efficiently. Finally, a post-pro…
Maximum Entropy models can be inferred from large data-sets to uncover how collective dynamics emerge from local interactions. Here, such models are employed to investigate neurons recorded by multielectrode arrays in the human and monkey cortex. Taking advantage of the separation of excitatory and inhibitory neuron types, we construct a model including this distinction. This approach allows to shed light upon differences between excitatory and inhibitory activity across different brain states such as wakefulness and deep sleep, in agreement with previous findings. Additionally, Maximum Entrop…
Experiments suggest that cerebral cortex gains several functional advantages by operating in a dynamical regime near the critical point of a phase transition. However, a long-standing criticism of this hypothesis is that critical dynamics are rather noisy, which might be detrimental to aspects of brain function that require precision. If the cortex does operate near criticality, how might it mitigate the noisy fluctuations? One possibility is that other parts of the brain may act to control the fluctuations and reduce cortical noise. To better understand this possibility, here we numerically a…
A wide range of evidence points toward the existence of a common algorithm underlying the processing of information throughout the cerebral cortex. Several hypothesized features of this cortical algorithm are reviewed, including sparse distributed representation, Bayesian inference, hierarchical organization composed of alternating template matching and pooling layers, temporal slowness and predictive coding. Hierarchical Temporal Memory (HTM) is a family of learning algorithms and corresponding theories of cortical function that embodies these principles. HTM has previously been applied mainl…
Partially inspired by features of computation in visual cortex, deep neural networks compute hierarchical representations of their inputs. While these networks have been highly successful in machine learning, it remains unclear to what extent they can aid our understanding of cortical function. Several groups have developed metrics that provide a quantitative comparison between representations computed by networks and representations measured in cortex. At the same time, neuroscience is well into an unprecedented phase of large-scale data collection, as evidenced by projects such as the Allen …
The cerebral cortex is composed of multiple cortical areas that exert a wide variety of brain functions. Although human brain neurons are genetically and areally mosaic, the three-dimensional structural differences between neurons in different brain areas or between the neurons of different individuals have not been delineated. Here, we report a nanometer-scale geometric analysis of brain tissues of the superior temporal gyrus of 4 schizophrenia and 4 control cases by using synchrotron radiation nanotomography. The results of the analysis and a comparison with results for the anterior cingulat…
The activity of neurons is correlated, and this correlation affects how the brain processes information. We study the neural circuit mechanisms of correlations by analyzing a network model characterized by strong and heterogeneous interactions: excitatory input drives the fluctuations of neural activity, which are counterbalanced by inhibitory feedback. In particular, excitatory input tends to correlate neurons, while inhibitory feedback reduces correlations. We demonstrate that heterogeneity of synaptic connections is necessary for this inhibition of correlations. We calculate statistical ave…
Columbia University（哥伦比亚大学）；Edmond and Lily Safra Center for Brain Sciences（埃德蒙和莉莉·萨弗拉脑科学中心）；Racah Institute of Physics（拉卡物理研究所）；The Hebrew University of Jerusalem（耶路撒冷希伯来大学）；Center for Brain Science（脑科学中心）；Harvard University（哈佛大学）
Neurons and networks in the cerebral cortex must operate reliably despite multiple sources of noise. To evaluate the impact of both input and output noise, we determine the robustness of single-neuron stimulus selective responses, as well as the robustness of attractor states of networks of neurons performing memory tasks. We find that robustness to output noise requires synaptic connections to be in a balanced regime in which excitation and inhibition are strong and largely cancel each other. We evaluate the conditions required for this regime to exist and determine the properties of networks…
