Boolean Dynamics with Random Couplings

This paper reviews a class of generic dissipative dynamical systems called N-K models. In these models, the dynamics of N elements, defined as Boolean variables, develop step by step, clocked by a discrete time variable. Each of the N Boolean elements at a given time is given a value which depends upon K elements in the previous time step. We review the work of many authors on the behavior of the models, looking particularly at the structure and lengths of their cycles, the sizes of their basins of attraction, and the flow of information through the systems. In the limit of infinite N, there is a phase transition between a chaotic and an ordered phase, with a critical phase in between. We argue that the behavior of this system depends significantly on the topology of the network connections. If the elements are placed upon a lattice with dimension d, the system shows correlations related to the standard percolation or directed percolation phase transition on such a lattice. On the other hand, a very different behavior is seen in the Kauffman net in which all spins are equally likely to be coupled to a given spin. In this situation, coupling loops are mostly suppressed, and the behavior of the system is much more like that of a mean field theory. We also describe possible applications of the models to, for example, genetic networks, cell differentiation, evolution, democracy in social systems and neural networks.Comment: 69 pages, 16 figures, Submitted to Springer Applied Mathematical Sciences Serie

Relative Stability of Network States in Boolean Network Models of Gene Regulation in Development

Progress in cell type reprogramming has revived the interest in Waddington's concept of the epigenetic landscape. Recently researchers developed the quasi-potential theory to represent the Waddington's landscape. The Quasi-potential U(x), derived from interactions in the gene regulatory network (GRN) of a cell, quantifies the relative stability of network states, which determine the effort required for state transitions in a multi-stable dynamical system. However, quasi-potential landscapes, originally developed for continuous systems, are not suitable for discrete-valued networks which are important tools to study complex systems. In this paper, we provide a framework to quantify the landscape for discrete Boolean networks (BNs). We apply our framework to study pancreas cell differentiation where an ensemble of BN models is considered based on the structure of a minimal GRN for pancreas development. We impose biologically motivated structural constraints (corresponding to specific type of Boolean functions) and dynamical constraints (corresponding to stable attractor states) to limit the space of BN models for pancreas development. In addition, we enforce a novel functional constraint corresponding to the relative ordering of attractor states in BN models to restrict the space of BN models to the biological relevant class. We find that BNs with canalyzing/sign-compatible Boolean functions best capture the dynamics of pancreas cell differentiation. This framework can also determine the genes' influence on cell state transitions, and thus can facilitate the rational design of cell reprogramming protocols.Comment: 24 pages, 6 figures, 1 tabl

Boolean networks with robust and reliable trajectories

We construct and investigate Boolean networks that follow a given reliable trajectory in state space, which is insensitive to fluctuations in the updating schedule, and which is also robust against noise. Robustness is quantified as the probability that the dynamics return to the reliable trajectory after a perturbation of the state of a single node. In order to achieve high robustness, we navigate through the space of possible update functions by using an evolutionary algorithm. We constrain the networks to having the minimum number of connections required to obtain the reliable trajectory. Surprisingly, we find that robustness always reaches values close to 100 percent during the evolutionary optimization process. The set of update functions can be evolved such that it differs only slightly from that of networks that were not optimized with respect to robustness. The state space of the optimized networks is dominated by the basin of attraction of the reliable trajectory.Comment: 12 pages, 9 figure

Numerical and Theoretical Studies of Noise Effects in the Kauffman Model

In this work we analyze the stochastic dynamics of the Kauffman model evolving under the influence of noise. By considering the average crossing time between two distinct trajectories, we show that different Kauffman models exhibit a similar kind of behavior, even when the structure of their basins of attraction is quite different. This can be considered as a robust property of these models. We present numerical results for the full range of noise level and obtain approximate analytic expressions for the above crossing time as a function of the noise in the limit cases of small and large noise levels.Comment: 24 pages, 9 figures, Submitted to the Journal of Statistical Physic

Stochastic neural network models for gene regulatory networks

Recent advances in gene-expression profiling technologies provide large amounts of gene expression data. This raises the possibility for a functional understanding of genome dynamics by means of mathematical modelling. As gene expression involves intrinsic noise, stochastic models are essential for better descriptions of gene regulatory networks. However, stochastic modelling for large scale gene expression data sets is still in the very early developmental stage. In this paper we present some stochastic models by introducing stochastic processes into neural network models that can describe intermediate regulation for large scale gene networks. Poisson random variables are used to represent chance events in the processes of synthesis and degradation. For expression data with normalized concentrations, exponential or normal random variables are used to realize fluctuations. Using a network with three genes, we show how to use stochastic simulations for studying robustness and stability properties of gene expression patterns under the influence of noise, and how to use stochastic models to predict statistical distributions of expression levels in population of cells. The discussion suggest that stochastic neural network models can give better description of gene regulatory networks and provide criteria for measuring the reasonableness o mathematical models

Scaling in a general class of critical random Boolean networks

We derive analytically the scaling behavior in the thermodynamic limit of the number of nonfrozen and relevant nodes in the most general class of critical Kauffman networks for any number of inputs per node, and for any choice of the probability distribution for the Boolean functions. By defining and analyzing a stochastic process that determines the frozen core we can prove that the mean number of nonfrozen nodes in any critical network with more than one input per node scales with the network size $N$ as $N^{2/3}$, with only $N^{1/3}$ nonfrozen nodes having two nonfrozen inputs and the number of nonfrozen nodes with more than two inputs being finite in the thermodynamic limit. Using these results we can conclude that the mean number of relevant nodes increases for large $N$ as $N^{1/3}$, with only a finite number of relevant nodes having two relevant inputs, and a vanishing fraction of nodes having more than three of them. It follows that all relevant components apart from a finite number are simple loops, and that the mean number and length of attractors increases faster than any power law with network size.Comment: 11 page

Topology of biological networks and reliability of information processing

Biological systems rely on robust internal information processing: Survival depends on highly reproducible dynamics of regulatory processes. Biological information processing elements, however, are intrinsically noisy (genetic switches, neurons, etc.). Such noise poses severe stability problems to system behavior as it tends to desynchronize system dynamics (e.g. via fluctuating response or transmission time of the elements). Synchronicity in parallel information processing is not readily sustained in the absence of a central clock. Here we analyze the influence of topology on synchronicity in networks of autonomous noisy elements. In numerical and analytical studies we find a clear distinction between non-reliable and reliable dynamical attractors, depending on the topology of the circuit. In the reliable cases, synchronicity is sustained, while in the unreliable scenario, fluctuating responses of single elements can gradually desynchronize the system, leading to non-reproducible behavior. We find that the fraction of reliable dynamical attractors strongly correlates with the underlying circuitry. Our model suggests that the observed motif structure of biological signaling networks is shaped by the biological requirement for reproducibility of attractors.Comment: 7 pages, 7 figure

A characterization of the Edge of Criticality in Binary Echo State Networks

Echo State Networks (ESNs) are simplified recurrent neural network models composed of a reservoir and a linear, trainable readout layer. The reservoir is tunable by some hyper-parameters that control the network behaviour. ESNs are known to be effective in solving tasks when configured on a region in (hyper-)parameter space called \emph{Edge of Criticality} (EoC), where the system is maximally sensitive to perturbations hence affecting its behaviour. In this paper, we propose binary ESNs, which are architecturally equivalent to standard ESNs but consider binary activation functions and binary recurrent weights. For these networks, we derive a closed-form expression for the EoC in the autonomous case and perform simulations in order to assess their behavior in the case of noisy neurons and in the presence of a signal. We propose a theoretical explanation for the fact that the variance of the input plays a major role in characterizing the EoC

Mutual information in random Boolean models of regulatory networks

The amount of mutual information contained in time series of two elements gives a measure of how well their activities are coordinated. In a large, complex network of interacting elements, such as a genetic regulatory network within a cell, the average of the mutual information over all pairs is a global measure of how well the system can coordinate its internal dynamics. We study this average pairwise mutual information in random Boolean networks (RBNs) as a function of the distribution of Boolean rules implemented at each element, assuming that the links in the network are randomly placed. Efficient numerical methods for calculating show that as the number of network nodes N approaches infinity, the quantity N exhibits a discontinuity at parameter values corresponding to critical RBNs. For finite systems it peaks near the critical value, but slightly in the disordered regime for typical parameter variations. The source of high values of N is the indirect correlations between pairs of elements from different long chains with a common starting point. The contribution from pairs that are directly linked approaches zero for critical networks and peaks deep in the disordered regime.Comment: 11 pages, 6 figures; Minor revisions for clarity and figure format, one reference adde