Phase transitions and critical behavior are crucial issues both in theoretical and experimental neuroscience. We report analytic and computational results about phase transitions and self-organized criticality (SOC) in networks with general stochastic neurons. The stochastic neuron has a firing probability given by a smooth monotonic function Φ(V) of the membrane potential V, rather than a sharp firing threshold. We find that such networks can operate in several dynamic regimes (phases) depending on the average synaptic weight and the shape of the firing function Φ. In particular, we encounter both continuous and discontinuous phase transitions to absorbing states. At the continuous transition critical boundary, neuronal avalanches occur whose distributions of size and duration are given by power laws, as observed in biological neural networks. We also propose and test a new mechanism to produce SOC: the use of dynamic neuronal gains – a form of short-term plasticity probably located at the axon initial segment (AIS) – instead of depressing synapses at the dendrites (as previously studied in the literature). The new self-organization mechanism produces a slightly supercritical state, that we called SOSC, in accord to some intuitions of Alan Turing.
Recent experiments suggested that homeostatic regulation of synaptic balance leads the visual system to recover and maintain a regime of power-law avalanches. Here we study an excitatory/inhibitory (E/I) mean-field neuronal network that has a critical point with power-law avalanches and synaptic balance. When short term depression in inhibitory synapses and firing threshold adaptation are added, the system hovers around the critical point. This homeostatically self-organized quasi-critical (SOqC) dynamics generates E/I synaptic current cancellation in fast time scales, causing fluctuation-driven asynchronous-irregular (AI) firing. We present the full phase diagram of the model without adaptation varying external input versus synaptic coupling.This system has a rich dynamical repertoire of spiking patterns: synchronous regular (SR), asynchronous regular (AR), synchronous irregular (SI), slow oscillations (SO) and AI. It also presents dynamic balance of synaptic currents, since inhibitory currents try and compensate excitatory currents over time, resulting in both of them scaling linearly with external input.Our model thus unifies two different perspectives on cortical spontaneous activity: both critical avalanches and fluctuation-driven AI firing arise from SOqC homeostatic adaptation, and are indeed two sides of the same coin.
Networks of stochastic spiking neurons are interesting models in the area of theoretical neuroscience, presenting both continuous and discontinuous phase transitions. Here, we study fully-connected networks analytically, numerically and by computational simulations. The neurons have dynamic gains that enable the network to converge to a stationary slightly supercritical state (self-organized supercriticality (SOSC)) in the presence of the continuous transition. We show that SOSC, which presents power laws for neuronal avalanches plus some large events, is robust as a function of the main parameter of the neuronal gain dynamics. We discuss the possible applications of the idea of SOSC to biological phenomena like epilepsy and Dragon-king avalanches. We also find that neuronal gains can produce collective oscillations that coexist with neuronal avalanches.
In the last decade, several models with network adaptive mechanisms (link deletion-creation, dynamic synapses, dynamic gains) have been proposed as examples of self-organized criticality (SOC) to explain neuronal avalanches. However, all these systems present stochastic oscillations hovering around the critical region that are incompatible with standard SOC. Here we make a linear stability analysis of the mean field fixed points of two self-organized quasi-critical systems: a fully connected network of discrete time stochastic spiking neurons with firing rate adaptation produced by dynamic neuronal gains and an excitable cellular automata with depressing synapses. We find that the fixed point corresponds to a stable focus that loses stability at criticality. We argue that when this focus is close to become indifferent, demographic noise can elicit stochastic oscillations that frequently fall into the absorbing state. This mechanism interrupts the oscillations, producing both power law avalanches and dragon king events, which appear as bands of synchronized firings in raster plots. Our approach differs from standard SOC models in that it predicts the coexistence of these different types of neuronal activity.
Burst firing is ubiquitous in nervous systems and has been intensively studied in central pattern generators (CPGs). Previous works have described subtle intraburst spike patterns (IBSPs) that, despite being traditionally neglected for their lack of relation to CPG motor function, were shown to be cell-type specific and sensitive to CPG connectivity. Here we address this matter by investigating how a bursting motor neuron expresses information about other neurons in the network. We performed experiments on the crustacean stomatogastric pyloric CPG, both in control conditions and interacting in real-time with computer model neurons. The sensitivity of postsynaptic to presynaptic IBSPs was inferred by computing their average mutual information along each neuron burst. We found that details of input patterns are nonlinearly and inhomogeneously coded through a single synapse into the fine IBSPs structure of the postsynaptic neuron following burst. In this way, motor neurons are able to use different time scales to convey two types of information simultaneously: muscle contraction (related to bursting rhythm) and the behavior of other CPG neurons (at a much shorter timescale by using IBSPs as information carriers). Moreover, the analysis revealed that the coding mechanism described takes part in a previously unsuspected information pathway from a CPG motor neuron to a nerve that projects to sensory brain areas, thus providing evidence of the general physiological role of information coding through IBSPs in the regulation of neuronal firing patterns in remote circuits by the CNS.
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