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Global neural dynamics emerge from multi-scale brain structures, with neurons communicating through synapses to form transiently communicating networks. Network activity arises from intercellular communication that depends on the structure of connectome tracts and local connection, intracellular signalling cascades, and the extracellular molecular milieu that regulate cellular properties. Multi-scale models of brain function have begun to directly link the emergence of global brain dynamics in conscious and unconscious brain states to microscopic changes at the level of cells. In particular, AdEx mean-field models representing statistical properties of local populations of neurons have been connected following human tractography data to represent multi-scale neural phenomena in simulations using The Virtual Brain (TVB). While mean-field models can be run on personal computers for short simulations, or in parallel on high-performance computing (HPC) architectures for longer simulations and parameter scans, the computational burden remains high and vast areas of the parameter space remain unexplored. In this work, we report that our TVB-HPC framework, a modular set of methods used here to implement the TVB-AdEx model for GPU and analyze emergent dynamics, notably accelerates simulations and substantially reduces computational resource requirements. The framework preserves the stability and robustness of the TVB-AdEx model, thus facilitating finer resolution exploration of vast parameter spaces as well as longer simulations previously near impossible to perform. Given that simulation and analysis toolkits are made public as open-source packages, our framework serves as a template onto which other models can be easily scripted and personalized datasets can be used for studies of inter-individual variability of parameters related to functional brain dynamics., Comment: 21 pages, 9 figures

Neuronal cable theory is usually derived from an electric analogue of the membrane, which contrasts with the slow movement of ions in aqueous media. We show here that it is possible to derive neuronal cable equations from a different perspective, based on the laws of hydrodynamic motion of charged particles (Navier-Stokes equations). This results in similar cable equations, but with additional contributions arising from nonlinear interactions inherent to fluid dynamics, and which may shape the integrative properties of the neurons.

Brain activity can be measured using magnetic fields located at some distance from the brain, a technique called magneto-encephalography (MEG). The origin of such magnetic fields are the ionic currents involved in neuronal activity. While these processes are well known at the microscopic scale, it is less clear how large populations of neurons generate magnetic fields. Here, we attempt to make such a link by deriving a mean-field theory of magnetic field generation by populations of neurons. We use the concept of current dipoles to build a theory that can be applied to populations of neurons. We also evaluate the contributions of neuronal current sources such as the synaptic currents or action potential currents. This theory should be useful to calculate the magnetic field located at long distances from neuronal populations, such as in MEG.

The brain is characterized by a strong heterogeneity of inhibitory neurons. We report that spiking neural networks display a resonance to the heterogeneity of inhibitory neurons, with optimal input/output responsiveness occurring for levels of heterogeneity similar to that found experimentally in cerebral cortex. A heterogeneous mean-field model predicts such optimal responsiveness. Moreover, we show that new dynamical regimes emerge from heterogeneity that were not present in the equivalent homogeneous system, such as sparsely synchronous collective oscillations.

Many studies have found evidence that the brain operates at a critical point, a processus known as self-organized criticality. A recent paper found remarkable scalings suggestive of criticality in systems as different as neural cultures, anesthetized or awake brains. We point out here that the diversity of these states would question any claimed role of criticality in information processing. Furthermore, we show that two non-critical systems pass all the tests for criticality, a control that was not provided in the original article. We conclude that such false positives demonstrate that the presence of criticality in the brain is still not proven and that we need better methods that scaling analyses.

A key element of the European Union's Human Brain Project (HBP) and other large-scale brain research projects is simulation of large-scale model networks of neurons. Here we argue why such simulations will likely be indispensable for bridging the scales between the neuron and system levels in the brain, and a set of brain simulators based on neuron models at different levels of biological detail should thus be developed. To allow for systematic refinement of candidate network models by comparison with experiments, the simulations should be multimodal in the sense that they should not only predict action potentials, but also electric, magnetic, and optical signals measured at the population and system levels.

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 to a different gain on inhibitory vs excitatory cells and that different levels of spike-frequency adaptation in excitatory cells can account for dynamical distinctions between the two types. This prediction was tested in vitro by varying adaptation strength using an agonist of acetylcholine receptors, which demonstrated a neuromodulatory switch between the two types of slow waves. Finally, we show that the first type of slow-wave dynamics is more sensitive to external stimuli, which can explain how slow waves in sleep and anesthesia differentially affect memory consolidation, as well as provide a link between slow-wave dynamics and memory diseases., Comment: 37 pages, 5 main figures, 4 supplementary figures

Neural electromagnetic (EM) signals recorded non-invasively from individual human subjects vary in complexity and magnitude. Nonetheless, variation in neural activity has been difficult to quantify and interpret, due to complex, broad-band features in the frequency domain. Studying signals recorded with magnetoencephalography (MEG) from healthy young adult subjects while in resting and active states, a systematic framework inspired by thermodynamics is applied to neural EM signals. Despite considerable inter-subject variation in terms of spectral entropy and energy across time epochs, data support the existence of a robust and linear relationship defining an effective state equation, with higher energy and lower entropy in the resting state compared to active, consistently across subjects. Mechanisms underlying the emergence of relationships between empirically measured effective state functions are further investigated using a model network of coupled oscillators, suggesting an interplay between noise and coupling strength can account for coherent variation of empirically observed quantities. Taken together, the results show macroscopic neural observables follow a robust, non-trivial conservation rule for energy modulation and information generation., Comment: 6 pages, 4 figures

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 Entropy models can also unveil novel features of neuronal interactions, which are found to be dominated by pairwise interactions during wakefulness, but are population-wide during deep sleep. In particular, inhibitory neurons are observed to be strongly tuned to the inhibitory population. Overall, we demonstrate Maximum Entropy models can be useful to analyze data-sets with classified neuron types, and to reveal the respective roles of excitatory and inhibitory neurons in organizing coherent dynamics in the cerebral cortex., Comment: 17 pages, 11 figures (including 5 supplementary)

The macroscopic electric permittivity of a given medium may depend on frequency, but this frequency dependence cannot be arbitrary, its real and imaginary parts are related by the well-known Kramers-Kronig relations. Here, we show that an analogous paradigm applies to the macroscopic electric conductivity. If the causality principle is taken into account, there exists Kramers-Kronig relations for conductivity, which are mathematically equivalent to the Hilbert transform. These relations impose strong constraints that models of heterogeneous media should satisfy to have a physically plausible frequency dependence of the conductivity and permittivity. We illustrate these relations and constraints by a few examples of known physical media. These extended relations constitute important constraints to test the consistency of past and future experimental measurements of the electric properties of heterogeneous media., Comment: 17 pages, 2 figures

During wakefulness and deep sleep brain states, cortical neural networks show a different behavior, with the second characterized by transients of high network activity. To investigate their impact on neuronal behavior, we apply a pairwise Ising model analysis by inferring the maximum entropy model that reproduces single and pairwise moments of the neuron's spiking activity. In this work we first review the inference algorithm introduced in Ferrari,Phys. Rev. E (2016). We then succeed in applying the algorithm to infer the model from a large ensemble of neurons recorded by multi-electrode array in human temporal cortex. We compare the Ising model performance in capturing the statistical properties of the network activity during wakefulness and deep sleep. For the latter, the pairwise model misses relevant transients of high network activity, suggesting that additional constraints are necessary to accurately model the data., Comment: 8 pages, 3 figures, Geometric Science of Information 2017 conference