We study living neural networks by measuring the neurons' response to a global electrical stimulation. Neural connectivity is lowered by reducing the synaptic strength, chemically blocking neurotransmitter receptors. We use a graph-theoretic approach to show that the connectivity undergoes a percolation transition. This occurs as the giant component disintegrates, characterized by a power law with critical exponent $β\simeq 0.65$ is independent of the balance between excitatory and inhibitory neurons and indicates that the degree distribution is gaussian rather than scale free
Cooperative effects in neural networks appear because a neuron fires only if a minimal number $m$ of its inputs are excited. The multiple inputs requirement leads to a percolation model termed {\it quorum percolation}. The connectivity undergoes a phase transition as $m$ grows, from a network--spanning cluster at low $m$ to a set of disconnected clusters above a critical $m$. Both numerical simulations and the model reproduce the experimental results well. This allows a robust quantification of biologically relevant quantities such as the average connectivity $\kbar$ and the distribution of co…
We study neural connectivity in cultures of rat hippocampal neurons. We measure the neurons' response to an electric stimulation for gradual lower connectivity, and characterize the size of the giant cluster in the network. The connectivity undergoes a percolation transition described by the critical exponent $β\simeq 0.65$. We use a theoretic approach based on bond.percolation on a graph to describe the process of disintegration of the network and extract its statistical properties. Together with numerical simulations we show that the connectivity in the neural culture is local, characterized…
Recent experimental studies of living neural networks reveal that their global activation induced by electrical stimulation can be explained using the concept of bootstrap percolation on a directed random network. The experiment consists in activating externally an initial random fraction of the neurons and observe the process of firing until its equilibrium. The final portion of neurons that are active depends in a non linear way on the initial fraction. The main result of this paper is a theorem which enables us to find the asymptotic of final proportion of the fired neurons in the case of r…
It is shown that, contrary to the claims in a recent letter by Haldeman and Beggs (PRL, 94, 058101, 2005), the branching ratio in epileptic cortical cultures is smaller than one. In addition, and also in contrast to claims made in that paper, the number of metastable states is not significantly different between cortical cultures in the critical state and cultures made epileptic using picrotoxin.
Improvements in technique in conjunction with an evolution of the theoretical and conceptual approach to neuronal networks provide a new perspective on living neurons in culture. Organization and connectivity are being measured quantitatively along with other physical quantities such as information, and are being related to function. In this review we first discuss some of these advances, which enable elucidation of structural aspects. We then discuss two recent experimental models that yield some conceptual simplicity. A one-dimensional network enables precise quantitative comparison to analy…
Shri Mata Vaishno Devi University（什里马塔瓦伊什诺德维大学）；Orthogonal Research and Education Lab（正交研究与教育实验室）；OpenWorm Foundation（OpenWorm基金会）
While Artificial Neural Networks (ANNs) have yielded impressive results in the realm of simulated intelligent behavior, it is important to remember that they are but sparse approximations of Biological Neural Networks (BNNs). We go beyond comparison of ANNs and BNNs to introduce principles from BNNs that might guide the further development of ANNs as embodied neural models. These principles include representational complexity, complex network structure/energetics, and robust function. We then consider these principles in ways that might be implemented in the future development of ANNs. In conc…
Organisms are nonequilibrium, stationary systems self-organized via spontaneous symmetry breaking and undergoing metabolic cycles with broken detailed balance in the environment. The thermodynamic free-energy principle describes an organism's homeostasis as the regulation of biochemical work constrained by the physical free-energy cost. In contrast, recent research in neuroscience and theoretical biology explains a higher organism's homeostasis and allostasis as Bayesian inference facilitated by the informational free energy. As an integrated approach to living systems, this study presents a f…
Neuronal networks in dissociated culture combined with cell engineering technology offer a pivotal platform to constructively explore the relationship between structure and function in living neuronal networks. Here, we fabricated defined neuronal networks possessing a modular architecture on high-density microelectrode arrays (HD-MEAs), a state-of-the-art electrophysiological tool for recording neural activity with high spatial and temporal resolutions. We first established a surface coating protocol using a cell-permissive hydrogel to stably attach polydimethylsiloxane microfluidic film on t…
To understand the learning process in brains, biologically plausible algorithms have been explored by modeling the detailed neuron properties and dynamics. On the other hand, simplified multi-layer models of neural networks have shown great success on computational tasks such as image classification and speech recognition. However, the computational models that can achieve good accuracy for these learning applications are very different from the bio-plausible models. This paper studies whether a bio-plausible model of a in vitro living neural network can be used to perform machine learning tas…
Eytan and Marom recently showed that the spontaneous burst activity of rat neuron cultures includes `first to fire' cells that consistently fire earlier than others. Here we analyze the behavior of these neurons in long term recordings of spontaneous activity of rat hippocampal and rat cortical neuron cultures from three different laboratories. We identify precursor events that may either subside (`small events') or can lead to a full-blown burst (`pre-bursts'). We find that the activation in the pre-burst typically has a first neuron (`leader'), followed by a localized response in its neighbo…
This paper contains an analysis of a simple neural network that exhibits self-organized criticality. Such criticality follows from the combination of a simple neural network with an excitatory feedback loop that generates bistability, in combination with an anti-Hebbian synapse in its input pathway. Using the methods of statistical field theory, we show how one can formulate the stochastic dynamics of such a network as the action of a path integral, which we then investigate using renormalization group methods. The results indicate that the network exhibits hysteresis in switching back and for…
The Wilson-Cowan model constitutes a paradigmatic approach to understanding the collective dynamics of networks of excitatory and inhibitory units. It has been profusely used in the literature to analyze the possible phases of neural networks at a mean-field level, e.g., assuming large fully-connected networks. Moreover, its stochastic counterpart allows one to study fluctuation-induced phenomena, such as avalanches. Here, we revisit the stochastic Wilson-Cowan model paying special attention to the possible phase transitions between quiescent and active phases. We unveil eight possible types o…
The dynamic range of stimulus processing in living organisms is much larger than a single neural network can explain. For a generic, tunable spiking network we derive that while the dynamic range is maximal at criticality, the interval of discriminable intensities is very similar for any network tuning due to coalescence. Compensating coalescence enables adaptation of discriminable intervals. Thus, we can tailor an ensemble of networks optimized to the distribution of stimulus intensities, e.g., extending the dynamic range arbitrarily. We discuss potential applications in machine learning.
We present a theoretical framework using quorum-percolation for describing the initiation of activity in a neural culture. The cultures are modeled as random graphs, whose nodes are excitatory neurons with kin inputs and kout outputs, and whose input degrees kin = k obey given distribution functions pk. We examine the firing activity of the population of neurons according to their input degree (k) classes and calculate for each class its firing probability Φ_k(t) as a function of t. The probability of a node to fire is found to be determined by its in-degree k, and the first-to-fire neurons ar…
Bridging integrated information theory and the free-energy principle in living neuronal networks
The University of Tokyo（东京大学）；International Research Center for Neurointelligence (WPI-IRCN), The University of Tokyo Institutes for Advanced Study (UTIAS)（东京大学高等研究院神经智能国际研究中心（WPI-IRCN））
Living organisms must actively maintain themselves in order to continue existing. Autopoiesis is a key concept in the study of living organisms, where the boundaries of the organism is not static by dynamically regulated by the system itself. To study the autonomous regulation of self-boundary, we focus on neural homeodynamic responses to environmental changes using both biological and artificial neural networks. Previous studies showed that embodied cultured neural networks and spiking neural networks with spike-timing dependent plasticity (STDP) learn an action as they avoid stimulation from…
The aim of this paper is to leverage the free-energy principle and its corollary process theory, active inference, to develop a generic, generalizable model of the representational capacities of living creatures; that is, a theory of phenotypic representation. Given their ubiquity, we are concerned with distributed forms of representation (e.g., population codes), whereby patterns of ensemble activity in living tissue come to represent the causes of sensory input or data. The active inference framework rests on the Markov blanket formalism, which allows us to partition systems of interest, suc…
Cyborg in the brain-machine interface field has attracted more attention in recent years. To control a creature via a machine called cyborg method, three stages are considerable: stimulation of neurons, neural response, and the behavioral reaction of the subject. Our main concern was to know how electrical stimulation induces neural activity and leads to a behavioral response. Additionally, we were interested to explore which type of electrical stimulation is optimal from different aspects such as maximum response with minimum induction stimulus field, minimum damage of the tissue and the elec…
The glow of a starry night sky, the smell of a freshly brewed cup of coffee or the sound of ocean waves breaking on the beach are representations of the physical world that have been created by the dynamic interactions of thousands of neurons in our brains. How the brain mediates perceptions, creates thoughts, stores memories and initiates actions remains one of the most profound puzzles in biology, if not all of science. A key to a mechanistic understanding of how the nervous system works is the ability to analyze the dynamics of neuronal networks in the living organism in the context of sens…
The dynamical mechanism underlying the processes of anesthesia-induced loss of consciousness and recovery is key to gaining insights into the working of the nervous system. Previous experiments revealed an asymmetry between neural signals during the anesthesia and recovery processes. Here we obtain experimental evidence for the hysteresis loop and articulate the dynamical mechanism based on percolation on multilayer complex networks with self-similarity. Model analysis reveals that, during anesthesia, the network is able to maintain its neural pathways despite the loss of a substantial fractio…
Despite the fact that the phenomenon of bursting activity is important for functioning of living neural networks, the mechanisms of its origin are still not clear. In this paper, we propose a new phenomenological model that can explain the mechanisms of the formation of bursting activity based on short-term synaptic plasticity, recurrent connections, and neuron-glial interactions. We show that neuron-glial interactions can induce bursting activity. The bifurcation scenarios of emergence of bursting activity are in the focus of the paper. Proposed study is important for understanding of the com…
One of the challenges of modern neuroscience is creating a "living computer" based on neural networks grown in vitro. Such an artificial device is supposed to perform neurocomputational tasks and interact with the environment when embodied in a robot. Recent studies have identified the most critical challenge, the search for a neural network architecture to implement associative learning. This work proposes a model of modular architecture with spiking neural networks connected by unidirectional couplings. We show that the model enables training a neuro-robot according to Pavlovian conditioning…
Centre National de la Recherche Scientifique（法国国家科学研究中心）；Strasbourg University（斯特拉斯堡大学）
Two universal functional principles of Adaptive Resonance Theory simulate the brain code of all biological learning and adaptive intelligence. Low level representations of multisensory stimuli in their immediate environmental context are formed on the basis of bottom up activation and under the control of top down matching rules that integrate high level long term traces of contextual configuration. These universal coding principles lead to the establishment of lasting brain signatures of perceptual experience in all living species, from aplysiae to primates. They are revisited in this paper h…
Carbon nanotubes are often seen as the only alternative technology to silicon transistors. While they are the most likely short-term one, other longer-term alternatives should be studied as well. While contemplating biological neurons as an alternative component may seem preposterous at first sight, significant recent progress in CMOS-neuron interface suggests this direction may not be unrealistic; moreover, biological neurons are known to self-assemble into very large networks capable of complex information processing tasks, something that has yet to be achieved with other emerging technologi…
Living neural networks emerge through a process of growth and self-organization that begins with a single cell and results in a brain, an organized and functional computational device. Artificial neural networks, however, rely on human-designed, hand-programmed architectures for their remarkable performance. Can we develop artificial computational devices that can grow and self-organize without human intervention? In this paper, we propose a biologically inspired developmental algorithm that can 'grow' a functional, layered neural network from a single initial cell. The algorithm organizes int…
