Quasi-criticalidade auto-organizada
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Nested θ- and β/γ-oscillations organize in the form of neuronal avalanches. (A) Definition of neuronal avalanches formed by the nested θ- and β/γ-oscillations. (Top) Threshold detection (broken line) of nLFPs (filled circles) at a single electrode. (Middle) Corresponding time–amplitude raster plot of nLFPs on the MEA (color: nLFP amplitude). (Bottom) Spatiotemporal nLFP clusters occupy successive bins of width Δt avg (dotted rectangles). (B) Average cross-correlation function for nLFPs in vivo at P8 (red) and P13 (black; single experiments). (C) nLFP clusters from nested θ- and β/γ-oscillations organize in the form of neuronal avalanches, i.e., distribute in sizes according to a power law with slope close to α = −1.5 (broken line). Average cluster size distribution in vivo plotted in log–log coordinates for P8 (red open circles; n = 5) and P13 (black; n = 7). (D) Example of two simultaneous burst periods before (black) and after (red) phase-shuffling. (E) The power law in cluster sizes is established for cluster area and cluster intensity (G) in single in vivo experiments and in the average (n = 7; F; cp. also C; all P13), but is destroyed on phase-shuffling of the LFP (open red). (H) Average cluster size distribution in vitro follows a power law with slope α ≅ −1.5 (broken line; n = 15; ≥10 DIV). (Inset) Average nLFP cross-correlation function for single experiment. Published online before print May 22, 2008, doi:10.1073/pnas.0800537105
PNAS May 27, 2008 vol. 105no. 21 7576-7581Neuronal avalanches organize as nested theta- and beta/gamma-oscillations during development of cortical layer 2/3
+Author Affiliations
Edited by Nancy J. Kopell, Boston University, Boston, MA, and approved March 27, 2008 (received for review January 18, 2008)
Abstract
Maturation of the cerebral cortex involves the spontaneous emergence of distinct patterns of neuronal synchronization, which regulate neuronal differentiation, synapse formation, and serve as a substrate for information processing. The intrinsic activity patterns that characterize the maturation of cortical layer 2/3 are poorly understood. By using microelectrode array recordings in vivo and in vitro, we show that this development is marked by the emergence of nested θ- and β/γ-oscillations that require NMDA- and GABAA-mediated synaptic transmission. The oscillations organized as neuronal avalanches, i.e., they were synchronized across cortical sites forming diverse and millisecond-precise spatiotemporal patterns that distributed in sizes according to a power law with a slope of −1.5. The correspondence between nested oscillations and neuronal avalanches required activation of the dopamine D1 receptor. We suggest that the repetitive formation of neuronal avalanches provides an intrinsic template for the selective linking of external inputs to developing superficial layers.
Self-organized (quasi-)criticality: the extremal Feder and Feder model
A simple random-neighbor SOC model that combines properties of the Bak-Sneppen and the relaxation oscillators (slip-stick) models is introduced. The analysis in terms of branching processes is transparent and gives insight about the development of large but finite mean avalanche sizes in dissipative models. In the thermodynamic limit, the distribution of states has a simple analytical form and the mean avalanche size, as a function of the coupling parameter strength, is exactly calculable.
Self-organization without conservation: true or just apparent scale-invariance?
The existence of true scale-invariance in slowly driven models of self-organized criticality without a conservation law, as forest-fires or earthquake automata, is scrutinized in this paper. By using three different levels of description - (i) a simple mean field, (ii) a more detailed mean-field description in terms of a (self-organized) branching processes, and (iii) a full stochastic representation in terms of a Langevin equation-, it is shown on general grounds that non-conserving dynamics does not lead to bona fide criticality. Contrarily to conserving systems, a parameter, which we term "re-charging" rate (e.g. the tree-growth rate in forest-fire models), needs to be fine-tuned in non-conserving systems to obtain criticality. In the infinite size limit, such a fine-tuning of the loading rate is easy to achieve, as it emerges by imposing a second separation of time-scales but, for any finite size, a precise tuning is required to achieve criticality and a coherent finite-size scaling picture. Using the approaches above, we shed light on the common mechanisms by which "apparent criticality" is observed in non-conserving systems, and explain in detail (both qualitatively and quantitatively) the difference with respect to true criticality obtained in conserving systems. We propose to call this self-organized quasi-criticality (SOqC). Some of the reported results are already known and some of them are new. We hope the unified framework presented here helps to elucidate the confusing and contradictory literature in this field. In a second accompanying paper, we shall discuss the implications of the general results obtained here for models of neural avalanches in Neuroscience for which self-organized scale-invariance in the absence of conservation has been claimed.
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