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Biological Evolution and Statistical Physics

This review is an introduction to theoretical models and mathematical calculations for biological evolution, aimed at physicists. The methods in the field are naturally very similar to those used in statistical physics, although the majority of publications appeared in biology journals. The review has three parts, which can be read independently. The first part deals with evolution in fitness landscapes and includes Fisher's theorem, adaptive walks, quasispecies models, effects of finite population sizes, and neutral evolution. The second part studies models of coevolution, including evolutionary game theory, kin selection, group selection, sexual selection, speciation, and coevolution of hosts and parasites. The third part discusses models for networks of interacting species and their extinction avalanches. Throughout the review, attention is paid to giving the necessary biological information, and to pointing out the assumptions underlying the models, and their limits of validity.

cond-mat.stat-mech↗

Evolution of networks

We review the recent fast progress in statistical physics of evolving networks. Interest has focused mainly on the structural properties of random complex networks in communications, biology, social sciences and economics. A number of giant artificial networks of such a kind came into existence recently. This opens a wide field for the study of their topology, evolution, and complex processes occurring in them. Such networks possess a rich set of scaling properties. A number of them are scale-free and show striking resilience against random breakdowns. In spite of large sizes of these networks, the distances between most their vertices are short -- a feature known as the ``small-world'' effect. We discuss how growing networks self-organize into scale-free structures and the role of the mechanism of preferential linking. We consider the topological and structural properties of evolving networks, and percolation in these networks. We present a number of models demonstrating the main features of evolving networks and discuss current approaches for their simulation and analytical study. Applications of the general results to particular networks in Nature are discussed. We demonstrate the generic connections of the network growth processes with the general problems of non-equilibrium physics, econophysics, evolutionary biology, etc.

cond-mat.stat-mech↗

A Configurational Bias Monte Carlo Method for Linear and Cyclic Peptides

In this manuscript, we describe a new configurational bias Monte Carlo technique for the simulation of peptides. We focus on the biologically relevant cases of linear and cyclic peptides. Our approach leads to an efficient, Boltzmann-weighted sampling of the torsional degrees of freedom in these biological molecules, a feat not possible with previous Monte Carlo and molecular dynamics methods.

cond-mat.soft↗

Phase Transitions of Neural Networks

The cooperative behaviour of interacting neurons and synapses is studied using models and methods from statistical physics. The competition between training error and entropy may lead to discontinuous properties of the neural network. This is demonstrated for a few examples: Perceptron, associative memory, learning from examples, generalization, multilayer networks, structure recognition, Bayesian estimate, on-line training, noise estimation and time series generation.

cond-mat.dis-nn↗

Time ordering in the evolution of information processing and modulation systems

The ideas of optimization of learning algorithms in Artificial Neural Networks are reviewed emphasizing generic properties and the online implementations are interpreted from a biological perspective. A simple model of the relevant subsidiary variables needed to improve learning in artificial feedforward networks and the `time ordering' of the appearance of the respective information processing systems is proposed. We discuss the possibility that these results might be relevant in other contexts, not being restricted to the simple models from which they stem. The analysis of a few examples, which range from the lowest cellular scale to the macroscopic level, suggests that similar ideas could be applied to biological systems.

cond-mat.dis-nn↗

Microspectroscopy and Imaging in the THz Range Using Coherent CW Radiation

A novel THz near-field spectrometer is presented which allows to perform biological and medical studies with high spectral resolution combined with a spatial resolution down to l/100. In the setup an aperture much smaller than the used wavelength is placed in the beam very close to the sample. The sample is probed by the evanescent wave behind the aperture. The distance is measured extremely accurate by a confocal microscope. We use monochromatic sources which provide powerful coherent cw radiation tuneable from 50 GHz up to 1.5 THz. Transmission and reflection experiments can be performed which enable us to study solids and molecules in aqueous solution. Examples for spectroscopic investigations on biological tissues are presented.

physics.bio-ph↗

A Microscopic Mechanism for Muscle's Motion

The SIRM (Stochastic Inclined Rods Model) proposed by H. Matsuura and M. Nakano can explain the muscle's motion perfectly, but the intermolecular potential between myosin head and G-actin is too simple and only repulsive potential is considered. In this paper we study the SIRM with different complex potential and discuss the effect of the spring on the system. The calculation results show that the spring, the effective radius of the G-actin and the intermolecular potential play key roles in the motion. The sliding speed is about $4.7\times10^{-6}m/s$ calculated from the model which well agrees with the experimental data.

physics.bio-ph↗

Diffusion on a hypercubic lattice with pinning potential: exact results for the error-catastrophe problem in biological evolution

In the theoretical biology framework one fundamental problem is the so-called error catastrophe in Darwinian evolution models. We reexamine Eigen's fundamental equations by mapping them into a polymer depinning transition problem in a ``genotype'' space represented by a unitary hypercubic lattice. The exact solution of the model shows that error catastrophe arises as a direct consequence of the equations involved and confirms some previous qualitative results. The physically relevant consequence is that such equations are not adequate to properly describe evolution of complex life on the Earth.

cond-mat↗

Entropic Sampling and Natural Selection in Biological Evolution

With a view to connecting random mutation on the molecular level to punctuated equilibrium behavior on the phenotype level, we propose a new model for biological evolution, which incorporates random mutation and natural selection. In this scheme the system evolves continuously into new configurations, yielding non-stationary behavior of the total fitness. Further, both the waiting time distribution of species and the avalanche size distribution display power-law behaviors with exponents close to two, which are consistent with the fossil data. These features are rather robust, indicating the key role of entropy.

cond-mat↗

An exact learning algorithm for autoassociative neural networks with binary couplings

Exact solutions for the learning problem of autoassociative networks with binary couplings are determined by a new method. The use of a branch-and-bound algorithm leads to a substantial saving of computational time compared with complete enumeration. As a result, fully connected networks with up to 40 neurons could be investigated. The network capacity is found to be close to 0.83.

cond-mat.dis-nn↗

Error threshold in the evolution of diploid organisms

The effects of error propagation in the reproduction of diploid organisms are studied within the populational genetics framework of the quasispecies model. The dependence of the error threshold on the dominance parameter is fully investigated. In particular, it is shown that dominance can protect the wild-type alleles from the error catastrophe. The analysis is restricted to a diploid analogue of the single-peaked landscape.

cond-mat.dis-nn↗

Statics and dynamics of an Ashkin-Teller neural network with low loading

An Ashkin-Teller neural network, allowing for two types of neurons is considered in the case of low loading as a function of the strength of the respective couplings between these neurons. The storage and retrieval of embedded patterns built from the two types of neurons, with different degrees of (in)dependence is studied. In particular, thermodynamic properties including the existence and stability of Mattis states are discussed. Furthermore, the dynamic behaviour is examined by deriving flow equations for the macroscopic overlap. It is found that for linked patterns the model shows better retrieval properties than a corresponding Hopfield model.

cond-mat.dis-nn↗

Phase Diagram and Storage Capacity of Sequence Processing Neural Networks

We solve the dynamics of Hopfield-type neural networks which store sequences of patterns, close to saturation. The asymmetry of the interaction matrix in such models leads to violation of detailed balance, ruling out an equilibrium statistical mechanical analysis. Using generating functional methods we derive exact closed equations for dynamical order parameters, viz. the sequence overlap and correlation- and response functions, in the thermodynamic limit. We calculate the time translation invariant solutions of these equations, describing stationary limit-cycles, which leads to a phase diagram. The effective retarded self-interaction usually appearing in symmetric models is here found to vanish, which causes a significantly enlarged storage capacity of $α_c\sim 0.269$, compared to $α_\c\sim 0.139$ for Hopfield networks storing static patterns. Our results are tested against extensive computer simulations and excellent agreement is found.

cond-mat.dis-nn↗

Finite-size scaling of the error threshold transition in finite population

The error threshold transition in a stochastic (i.e. finite population) version of the quasispecies model of molecular evolution is studied using finite-size scaling. For the single-sharp-peak replication landscape, the deterministic model exhibits a first-order transition at $Q=Q_c=1/a$, where $% Q$ is the probability of exact replication of a molecule of length $L \to \infty$, and $a$ is the selective advantage of the master string. For sufficiently large population size, $N$, we show that in the critical region the characteristic time for the vanishing of the master strings from the population is described very well by the scaling assumption $τ= N^{1/2} f_a \left [ \left (Q - Q_c) N^{1/2} \right ] $, where $f_a$ is an $a$-dependent scaling function.

cond-mat.stat-mech↗

Dynamical properties of a randomly diluted neural network with variable activity

The subject of study is a neural network with binary neurons, randomly diluted synapses and variable pattern activity. We look at the system with parallel updating using a probabilistic approach to solve the one step dynamics with one condensed pattern. We derive restrictions on the storage capacity and the mutual information content occuring during the retrieval process. Special focus is on the constraints on the threshold for optimal performance. We also look at the effect of noisy updating, giving a dynamical version of the critical temperature, the corresponding threshold and an approximation for the time evolution for small temperatures. The description is applicable to the whole retrieval process in the limit of strong dilution. The analysis is carried out as exactly as possible and over the full parameter ranges, generalizing some former results.

cond-mat.dis-nn↗

A replica approach to the state-dependent synapses neural network

The replica method is applied to a neural network model with state-dependent synapses built from those patterns having a correlation with the state of the system greater than a certain threshold. Replica-symmetric and first-step replica-symmetry-breaking results are presented for the storage capacity at zero temperature as a function of this threshold value. A comparison is made with existing results based upon mean-field equations obtained by using a statistical method.

cond-mat.dis-nn↗

Oscillator neural network model with distributed native frequencies

We study associative memory of an oscillator neural network with distributed native frequencies. The model is based on the use of the Hebb learning rule with random patterns ($ξ_i^μ=\pm 1$), and the distribution function of native frequencies is assumed to be symmetric with respect to its average. Although the system with an extensive number of stored patterns is not allowed to get entirely synchronized, long time behaviors of the macroscopic order parameters describing partial synchronization phenomena can be obtained by discarding the contribution from the desynchronized part of the system. The oscillator network is shown to work as associative memory accompanied by synchronized oscillations. A phase diagram representing properties of memory retrieval is presented in terms of the parameters characterizing the native frequency distribution. Our analytical calculations based on the self-consistent signal-to-noise analysis are shown to be in excellent agreement with numerical simulations, confirming the validity of our theoretical treatment.

cond-mat.dis-nn↗

Delineation of the Native Basin in Continuum Models of Proteins

We propose two approaches for determining the native basins in off-lattice models of proteins. The first of them is based on exploring the saddle points on selected trajectories emerging from the native state. In the second approach, the basin size can be determined by monitoring random distortions in the shape of the protein around the native state. Both techniques yield the similar results. As a byproduct, a simple method to determine the folding temperature is obtained.

cond-mat.soft↗