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Register a new userAsynchronous and coherent dynamics in balanced excitatory-inhibitory populations
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Dynamic excitatory-inhibitory (E-I) balance is a paradigmatic mechanism invoked to explain the irregular low firing activity observed in the cortex. However, we will show that the E-I balance can be at the origin of other regimes observable in the brain. The analysis is performed by combining simulations of sparse E-I networks composed of N spiking neurons with analytical investigations of low dimensional neural mass models. The bifurcation diagrams, derived for the neural mass model, allow to classify the asynchronous and coherent behaviours emerging any finite in-degree K. In the limit N >> K >> 1 both supra and sub-threshold balanced asynchronous regimes can be observed. Due to structural heterogeneity the asynchronous states are characterized by the splitting of the neurons in three groups: silent, fluctuation and mean driven. The coherent rhythms are characterized by regular or irregular temporal fluctuations joined to spatial coherence similar to coherent fluctuations observed in the cortex over multiple spatial scales. Collective Oscillations (COs) can emerge due to two different mechanisms. A first mechanism similar to the pyramidal-interneuron gamma (PING) one. The second mechanism is intimately related to the presence of current fluctuations, which sustain COs characterized by an essentially simultaneous bursting of the two populations. We observe period-doubling cascades involving the PING-like COs finally leading to the appearance of coherent chaos. For sufficiently strong current fluctuations we report a novel mechanism of frequency locking among collective rhythms promoted by these intrinsic fluctuations. Our analysis suggest that despite PING-like or fluctuation driven COS are observable for any finite in-degree K, in the limit N >> K >> 1 these solutions result in two coexisting balanced regimes: an asynchronous and a fully synchronized one.
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It is widely appreciated that well-balanced excitation and inhibition are necessary for proper function in neural networks. However, in principle, such balance could be achieved by many possible configurations of excitatory and inhibitory strengths, and relative numbers of excitatory and inhibitory neurons. For instance, a given level of excitation could be balanced by either numerous inhibitory neurons with weak synapses, or few inhibitory neurons with strong synapses. Among the continuum of different but balanced configurations, why should any particular configuration be favored? Here we address this question in the context of the entropy of network dynamics by studying an analytically tractable network of binary neurons. We find that entropy is highest at the boundary between excitation-dominant and inhibition-dominant regimes. Entropy also varies along this boundary with a trade-off between high and robust entropy: weak synapse strengths yield high network entropy which is fragile to parameter variations, while strong synapse strengths yield a lower, but more robust, network entropy. In the case where inhibitory and excitatory synapses are constrained to have similar strength, we find that a small, but non-zero fraction of inhibitory neurons, like that seen in mammalian cortex, results in robust and relatively high entropy.
Collective oscillations and their suppression by external stimulation are analyzed in a large-scale neural network consisting of two interacting populations of excitatory and inhibitory quadratic integrate-and-fire neurons. In the limit of an infinite number of neurons, the microscopic model of this network can be reduced to an exact low-dimensional system of mean-field equations. Bifurcation analysis of these equations reveals three different dynamic modes in a free network: a stable resting state, a stable limit cycle, and bistability with a coexisting resting state and a limit cycle. We show that in the limit cycle mode, high-frequency stimulation of an inhibitory population can stabilize an unstable resting state and effectively suppress collective oscillations. We also show that in the bistable mode, the dynamics of the network can be switched from a stable limit cycle to a stable resting state by applying an inhibitory pulse to the excitatory population. The results obtained from the mean-field equations are confirmed by numerical simulation of the microscopic model.
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.
We investigate the collective dynamics of excitatory-inhibitory excitable networks in response to external stimuli. How to enhance dynamic range, which represents the ability of networks to encode external stimuli, is crucial to many applications. We regard the system as a two-layer network (E-Layer and I-Layer) and explore the criticality and dynamic range on diverse networks. Interestingly, we find that phase transition occurs when the dominant eigenvalue of E-layers weighted adjacency matrix is exactly one, which is only determined by the topology of E-Layer. Meanwhile, it is shown that dynamic range is maximized at critical state. Based on theoretical analysis, we propose an inhibitory factor for each excitatory node. We suggest that if nodes with high inhibitory factors are cut out from I-Layer, dynamic range could be further enhanced. However, because of the sparseness of networks and passive function of inhibitory nodes, the improvement is relatively small compared tooriginal dynamic range. Even so, this provides a strategy to enhance dynamic range.
A companion paper introduces a nonlinear network with Hebbian excitatory (E) neurons that are reciprocally coupled with anti-Hebbian inhibitory (I) neurons and also receive Hebbian feedforward excitation from sensory (S) afferents. The present paper derives the network from two normative principles that are mathematically equivalent but conceptually different. The first principle formulates unsupervised learning as a constrained optimization problem: maximization of S-E correlations subject to a copositivity constraint on E-E correlations. A combination of Legendre and Lagrangian duality yields a zero-sum continuous game between excitatory and inhibitory connections that is solved by the neural network. The second principle defines a zero-sum game between E and I cells. E cells want to maximize S-E correlations and minimize E-I correlations, while I cells want to maximize I-E correlations and minimize power. The conflict between I and E objectives effectively forces the E cells to decorrelate from each other, although only incompletely. Legendre duality yields the neural network.
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