The Cortex and the Critical Point

How the cerebral cortex operates near a critical phase transition point for optimum performance. Individual neurons have limited computational powers, but when they work together, it is almost like magic. Firing synchronously and then breaking off to improvise by themselves, they can be paradoxically both independent and interdependent. This happens near the critical point: when neurons are poised between a phase where activity is damped and a phase where it is amplified, where information processing is optimized, and complex emergent activity patterns arise. The claim that neurons in the cortex work best when they operate near the critical point is known as the criticality hypothesis. In this book John Beggs—one of the pioneers of this hypothesis—offers an introduction to the critical point and its relevance to the brain. Drawing on recent experimental evidence, Beggs first explains the main ideas underlying the criticality hypotheses and emergent phenomena. He then discusses the critical point and its two main consequences—first, scale-free properties that confer optimum information processing; and second, universality, or the idea that complex emergent phenomena, like that seen near the critical point, can be explained by relatively simple models that are applicable across species and scale. Finally, Beggs considers future directions for the field, including research on homeostatic regulation, quasicriticality, and the expansion of the cortex and intelligence. An appendix provides technical material; many chapters include exercises that use freely available code and data sets

A few strong connections: optimizing information retention in neuronal avalanches

<p>Abstract</p> <p>Background</p> <p>How living neural networks retain information is still incompletely understood. Two prominent ideas on this topic have developed in parallel, but have remained somewhat unconnected. The first of these, the "synaptic hypothesis," holds that information can be retained in synaptic connection strengths, or weights, between neurons. Recent work inspired by statistical mechanics has suggested that networks will retain the most information when their weights are distributed in a skewed manner, with many weak weights and only a few strong ones. The second of these ideas is that information can be represented by stable activity patterns. Multineuron recordings have shown that sequences of neural activity distributed over many neurons are repeated above chance levels when animals perform well-learned tasks. Although these two ideas are compelling, no one to our knowledge has yet linked the predicted optimum distribution of weights to stable activity patterns actually observed in living neural networks.</p> <p>Results</p> <p>Here, we explore this link by comparing stable activity patterns from cortical slice networks recorded with multielectrode arrays to stable patterns produced by a model with a tunable weight distribution. This model was previously shown to capture central features of the dynamics in these slice networks, including neuronal avalanche cascades. We find that when the model weight distribution is appropriately skewed, it correctly matches the distribution of repeating patterns observed in the data. In addition, this same distribution of weights maximizes the capacity of the network model to retain stable activity patterns. Thus, the distribution that best fits the data is also the distribution that maximizes the number of stable patterns.</p> <p>Conclusions</p> <p>We conclude that local cortical networks are very likely to use a highly skewed weight distribution to optimize information retention, as predicted by theory. Fixed distributions impose constraints on learning, however. The network must have mechanisms for preserving the overall weight distribution while allowing individual connection strengths to change with learning.</p

Stability of polarization singularities in disordered photonic crystal waveguides

The effects of short-range disorder on the polarization characteristics of light in photonic crystal waveguides were investigated using finite-difference time-domain simulations with a view to investigating the stability of polarization singularities. It was found that points of local circular polarization (C points) and contours of linear polarization (L lines) continued to appear even in the presence of high levels of disorder, and that they remained close to their positions in the ordered crystal. These results are a promising indication that devices exploiting polarization in these structures are viable given current fabrication standards

Extending Transfer Entropy Improves Identification of Effective Connectivity in a Spiking Cortical Network Model

Transfer entropy (TE) is an information-theoretic measure which has received recent attention in neuroscience for its potential to identify effective connectivity between neurons. Calculating TE for large ensembles of spiking neurons is computationally intensive, and has caused most investigators to probe neural interactions at only a single time delay and at a message length of only a single time bin. This is problematic, as synaptic delays between cortical neurons, for example, range from one to tens of milliseconds. In addition, neurons produce bursts of spikes spanning multiple time bins. To address these issues, here we introduce a free software package that allows TE to be measured at multiple delays and message lengths. To assess performance, we applied these extensions of TE to a spiking cortical network model (Izhikevich, 2006) with known connectivity and a range of synaptic delays. For comparison, we also investigated single-delay TE, at a message length of one bin (D1TE), and cross-correlation (CC) methods. We found that D1TE could identify 36% of true connections when evaluated at a false positive rate of 1%. For extended versions of TE, this dramatically improved to 73% of true connections. In addition, the connections correctly identified by extended versions of TE accounted for 85% of the total synaptic weight in the network. Cross correlation methods generally performed more poorly than extended TE, but were useful when data length was short. A computational performance analysis demonstrated that the algorithm for extended TE, when used on currently available desktop computers, could extract effective connectivity from 1 hr recordings containing 200 neurons in ∼5 min. We conclude that extending TE to multiple delays and message lengths improves its ability to assess effective connectivity between spiking neurons. These extensions to TE soon could become practical tools for experimentalists who record hundreds of spiking neurons

A simple spontaneously active Hebbian learning model: homeostasis of activity and connectivity, and consequences for learning and epileptogenesis

A spontaneously active neural system that is capable of continual learning should also be capable of homeostasis of both firing rate and connectivity. Experimental evidence suggests that both types of homeostasis exist, and that connectivity is maintained at a state that is optimal for information transmission and storage. This state is referred to as the critical state. We present a simple stochastic computational Hebbian learning model that incorporates both firing rate and critical homeostasis, and we explore its stability and connectivity properties. We also examine the behavior of our model with a simulated seizure and with simulated acute deafferentation. We argue that a neural system that is more highly connected than the critical state (i.e., one that is "supercritical") is epileptogenic. Based on our simulations, we predict that the post-seizural and post-deafferentation states should be supercritical and epileptogenic. Furthermore, interventions that boost spontaneous activity should be protective against epileptogenesis.Comment: 37 pages, 1 table, 7 figure

Rich-Club Organization in Effective Connectivity among Cortical Neurons.

The performance of complex networks, like the brain, depends on how effectively their elements communicate. Despite the importance of communication, it is virtually unknown how information is transferred in local cortical networks, consisting of hundreds of closely spaced neurons. To address this, it is important to record simultaneously from hundreds of neurons at a spacing that matches typical axonal connection distances, and at a temporal resolution that matches synaptic delays. We used a 512-electrode array (60 μm spacing) to record spontaneous activity at 20 kHz from up to 500 neurons simultaneously in slice cultures of mouse somatosensory cortex for 1 h at a time. We applied a previously validated version of transfer entropy to quantify information transfer. Similar to in vivo reports, we found an approximately lognormal distribution of firing rates. Pairwise information transfer strengths also were nearly lognormally distributed, similar to reports of synaptic strengths. Some neurons transferred and received much more information than others, which is consistent with previous predictions. Neurons with the highest outgoing and incoming information transfer were more strongly connected to each other than chance, thus forming a “rich club.” We found similar results in networks recorded in vivo from rodent cortex, suggesting the generality of these findings. A rich-club structure has been found previously in large-scale human brain networks and is thought to facilitate communication between cortical regions. The discovery of a small, but information-rich, subset of neurons within cortical regions suggests that this population will play a vital role in communication, learning, and memory.SIGNIFICANCE STATEMENT Many studies have focused on communication networks between cortical brain regions. In contrast, very few studies have examined communication networks within a cortical region. This is the first study to combine such a large number of neurons (several hundred at a time) with such high temporal resolution (so we can know the direction of communication between neurons) for mapping networks within cortex. We found that information was not transferred equally through all neurons. Instead, ∼70% of the information passed through only 20% of the neurons. Network models suggest that this highly concentrated pattern of information transfer would be both efficient and robust to damage. Therefore, this work may help in understanding how the cortex processes information and responds to neurodegenerative diseases