Regulation of Irregular Neuronal Firing by Autaptic Transmission

The importance of self-feedback autaptic transmission in modulating spike-time irregularity is still poorly understood. By using a biophysical model that incorporates autaptic coupling, we here show that self-innervation of neurons participates in the modulation of irregular neuronal firing, primarily by regulating the occurrence frequency of burst firing. In particular, we find that both excitatory and electrical autapses increase the occurrence of burst firing, thus reducing neuronal firing regularity. In contrast, inhibitory autapses suppress burst firing and therefore tend to improve the regularity of neuronal firing. Importantly, we show that these findings are independent of the firing properties of individual neurons, and as such can be observed for neurons operating in different modes. Our results provide an insightful mechanistic understanding of how different types of autapses shape irregular firing at the single-neuron level, and they highlight the functional importance of autaptic self-innervation in taming and modulating neurodynamics.Comment: 27 pages, 8 figure

A neuronal network model of interictal and recurrent ictal activity

We propose a neuronal network model which undergoes a saddle-node bifurcation on an invariant circle as the mechanism of the transition from the interictal to the ictal (seizure) state. In the vicinity of this transition, the model captures important dynamical features of both interictal and ictal states. We study the nature of interictal spikes and early warnings of the transition predicted by this model. We further demonstrate that recurrent seizures emerge due to the interaction between two networks.Comment: 9 pages, 7 figure

Sustained Activity in Hierarchical Modular Neural Networks: Self-Organized Criticality and Oscillations

Cerebral cortical brain networks possess a number of conspicuous features of structure and dynamics. First, these networks have an intricate, non-random organization. In particular, they are structured in a hierarchical modular fashion, from large-scale regions of the whole brain, via cortical areas and area subcompartments organized as structural and functional maps to cortical columns, and finally circuits made up of individual neurons. Second, the networks display self-organized sustained activity, which is persistent in the absence of external stimuli. At the systems level, such activity is characterized by complex rhythmical oscillations over a broadband background, while at the cellular level, neuronal discharges have been observed to display avalanches, indicating that cortical networks are at the state of self-organized criticality (SOC). We explored the relationship between hierarchical neural network organization and sustained dynamics using large-scale network modeling. Previously, it was shown that sparse random networks with balanced excitation and inhibition can sustain neural activity without external stimulation. We found that a hierarchical modular architecture can generate sustained activity better than random networks. Moreover, the system can simultaneously support rhythmical oscillations and SOC, which are not present in the respective random networks. The mechanism underlying the sustained activity is that each dense module cannot sustain activity on its own, but displays SOC in the presence of weak perturbations. Therefore, the hierarchical modular networks provide the coupling among subsystems with SOC. These results imply that the hierarchical modular architecture of cortical networks plays an important role in shaping the ongoing spontaneous activity of the brain, potentially allowing the system to take advantage of both the sensitivity of critical states and the predictability and timing of oscillations for efficient information processing

The Impact Of Spike-Frequency Adaptation On Balanced Network Dynamics

A dynamic balance between strong excitatory and inhibitory neuronal inputs is hypothesized to play a pivotal role in information processing in the brain. While there is evidence of the existence of a balanced operating regime in several cortical areas and idealized neuronal network models, it is important for the theory of balanced networks to be reconciled with more physiological neuronal modeling assumptions. In this work, we examine the impact of spike-frequency adaptation, observed widely across neurons in the brain, on balanced dynamics. We incorporate adaptation into binary and integrate-and-fire neuronal network models, analyzing the theoretical effect of adaptation in the large network limit and performing an extensive numerical investigation of the model adaptation parameter space. Our analysis demonstrates that balance is well preserved for moderate adaptation strength even if the entire network exhibits adaptation. In the common physiological case in which only excitatory neurons undergo adaptation, we show that the balanced operating regime in fact widens relative to the non-adaptive case. We hypothesize that spike-frequency adaptation may have been selected through evolution to robustly facilitate balanced dynamics across diverse cognitive operating states

Characterization and Compensation of Network-Level Anomalies in Mixed-Signal Neuromorphic Modeling Platforms

Advancing the size and complexity of neural network models leads to an ever increasing demand for computational resources for their simulation. Neuromorphic devices offer a number of advantages over conventional computing architectures, such as high emulation speed or low power consumption, but this usually comes at the price of reduced configurability and precision. In this article, we investigate the consequences of several such factors that are common to neuromorphic devices, more specifically limited hardware resources, limited parameter configurability and parameter variations. Our final aim is to provide an array of methods for coping with such inevitable distortion mechanisms. As a platform for testing our proposed strategies, we use an executable system specification (ESS) of the BrainScaleS neuromorphic system, which has been designed as a universal emulation back-end for neuroscientific modeling. We address the most essential limitations of this device in detail and study their effects on three prototypical benchmark network models within a well-defined, systematic workflow. For each network model, we start by defining quantifiable functionality measures by which we then assess the effects of typical hardware-specific distortion mechanisms, both in idealized software simulations and on the ESS. For those effects that cause unacceptable deviations from the original network dynamics, we suggest generic compensation mechanisms and demonstrate their effectiveness. Both the suggested workflow and the investigated compensation mechanisms are largely back-end independent and do not require additional hardware configurability beyond the one required to emulate the benchmark networks in the first place. We hereby provide a generic methodological environment for configurable neuromorphic devices that are targeted at emulating large-scale, functional neural networks

Sustained oscillations, irregular firing, and chaotic dynamics in hierarchical modular networks with mixtures of electrophysiological cell types

The cerebral cortex exhibits neural activity even in the absence of externalstimuli. This self-sustained activity is characterized by irregular firing ofindividual neurons and population oscillations with a broad frequency range.Questions that arise in this context, are: What are the mechanismsresponsible for the existence of neuronal spiking activity in the cortexwithout external input? Do these mechanisms depend on the structural organization of the cortical connections? Do they depend onintrinsic characteristics of the cortical neurons? To approach the answers to these questions, we have used computer simulations of cortical network models. Our networks have hierarchical modular architecture and are composedof combinations of neuron models that reproduce the firing behavior of the five main cortical electrophysiological cell classes: regular spiking (RS), chattering (CH), intrinsically bursting (IB), low threshold spiking (LTS) and fast spiking (FS). The population of excitatory neurons is built of RS cells(always present) and either CH or IB cells. Inhibitoryneurons belong to the same class, either LTS or FS. Long-lived self-sustained activity states in our networksimulations display irregular single neuron firing and oscillatoryactivity similar to experimentally measured ones. The duration of self-sustained activity strongly depends on the initial conditions,suggesting a transient chaotic regime. Extensive analysis of the self-sustainedactivity states showed that their lifetime expectancy increases with the numberof network modules and is favored when the network is composed of excitatory neurons of the RS and CH classes combined with inhibitory neurons of the LTS class. These results indicate that the existence and properties of the self-sustained cortical activity states depend on both the topology of the network and the neuronal mixture that comprises the network

Membrane resonance enables stable and robust gamma oscillations

Neuronal mechanisms underlying beta/gamma oscillations (20-80 Hz) are not completely understood. Here, we show that in vivo beta/gamma oscillations in the cat visual cortex sometimes exhibit remarkably stable frequency even when inputs fluctuate dramatically. Enhanced frequency stability is associated with stronger oscillations measured in individual units and larger power in the local field potential. Simulations of neuronal circuitry demonstrate that membrane properties of inhibitory interneurons strongly determine the characteristics of emergent oscillations. Exploration of networks containing either integrator or resonator inhibitory interneurons revealed that: (i) Resonance, as opposed to integration, promotes robust oscillations with large power and stable frequency via a mechanism called RING (Resonance INduced Gamma); resonance favors synchronization by reducing phase delays between interneurons and imposes bounds on oscillation cycle duration; (ii) Stability of frequency and robustness of the oscillation also depend on the relative timing of excitatory and inhibitory volleys within the oscillation cycle; (iii) RING can reproduce characteristics of both Pyramidal INterneuron Gamma (PING) and INterneuron Gamma (ING), transcending such classifications; (iv) In RING, robust gamma oscillations are promoted by slow but are impaired by fast inputs. Results suggest that interneuronal membrane resonance can be an important ingredient for generation of robust gamma oscillations having stable frequency