Abstract
Stochastic phenomena are central to biological modeling. Small numbers of molecules regulate and control genetics, epigenetics, and cellular metabolism, and small numbers of well-adapted individuals drive evolution. As far as processes are concerned, the relation between chemistry and biology is a tight one. Reproduction, the basis of all processes in evolution, is understood as autocatalysis with an exceedingly complex mechanism in the language of chemists, and replication of the genetic molecules, DNA and RNA, builds the bridge between chemistry and biology. The earliest stochastic models in biology applied branching processes in order to give answers to genealogical questions like, for example, the fate of family names in pedigrees. Branching processes, birth-and-death processes, and related stochastic models are frequently used in biology and they are defined, analyzed, and applied to typical problems. Although the master equation is not so dominant in biology as it is in chemistry, it is a very useful tool for deriving analytical solutions, and most birth-and-death processes can be analyzed successfully by means of master equations. Kimura’s neutral theory of evolution makes use of a Fokker–Planck equation and describes population dynamics in the absence of fitness differences. A section on coalescent theory demonstrates the applicability of backwards modeling to the problem of reconstruction of phylogenies. Unlike in the previous chapter we shall present and discuss numerical simulations here together with the analytical approaches. Simulations of stochastic reaction networks in systems biology are a rapidly growing field and several new monographs have come out during the last few years. Therefore only a brief account and a collection of references are given.
Nothing in biology makes sense except in the light of evolution.
Theodosius Dobzhansky 1972.
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Notes
- 1.
Throughout this monograph we use subspecies in the sense of molecular species or variant for the components of a population \(\boldsymbol{\Pi } =\{ \mathsf{X}_{1},\mathsf{X}_{2},\mathop{\ldots },\mathsf{X}_{n}\}\). We express its numbers by the random vector \(\mathcal{N} = \vert \boldsymbol{\Pi }\vert = (\mathcal{N}_{1},\mathcal{N}_{2},\mathop{\ldots },\mathcal{N}_{n})\) and indicate by using the notion biological species when we mean species in biology.
- 2.
Verhulst himself gave no biological interpretation of the logistic equation and its parameters in terms of a carrying capacity. For a brief historical account on the origin of this equation, see [282]. The Malthus parameter is commonly denoted by r. Since r is used as the flow rate in the CSTR, we choose f here in order to avoid confusion and to stress the close relationship between the Malthus parameter and fitness.
- 3.
In this section we shall use n for the number of molecules involved in the autocatalytic reaction, as well as for the numbers of stochastic variables.
- 4.
Termolecular and higher reaction steps are neglected in mass action kinetics, but they are nevertheless frequently used in models and simplified kinetic mechanisms. Examples are the Schlögl model aut]Schlögl, F. [479] and the Brusselator model [421] (Sect. 4.6.4). Thomas Wilhelm aut]Wilhelm, T. and Reinhart Heinrich aut]Heinrich, R. provided a rigorous proof that the smallest oscillating system with only mono- and bimolecular reaction steps has to be at least three-dimensional and must contain one bimolecular term [571]. A similar proof for the smallest system showing bistability can be found in [570].
- 5.
This relation is a result of mass conservation in the closed system.
- 6.
As for the time dependence in the closed system expressed by (5.5c), we make use of the uncommon implicit function \(r = f(\bar{x})\) rather than the direct relation \(\bar{x} = f(r)\).
- 7.
- 8.
Although confusion is highly improbable, we remark that the use of n as the exponent of the growth function in x n and as the number of particles in P n is ambiguous here.
- 9.
Reproduction is understood here as asexual reproduction, which under pseudo-first order conditions gives rise to linear reaction kinetics. Sexual reproduction requires two partners and gives rise to a special process of reaction order two (Table 4.2).
- 10.
Here we use the symbols commonly applied in biology: λ (n) for birth, μ (n) for death, ν for immigration, and ρ for emigration. Other notions and symbols are used in chemistry: f ≡ λ for birth corresponding to the production of a molecule and d ≡ μ for death understood as decomposition or degradation through a chemical reaction. Inflow and outflow are the equivalents of immigration and emigration. Pure immigration and emigration give rise to Poisson processes and continuous time random walks, which have been discussed extensively in Chap. 3 There we denoted the parameters by γ and ϑ, instead of ν and ρ.
- 11.
Here the conjugated Laplace variable is denoted by \(\hat{s}\) in order to avoid confusion with the dummy variable s in the generating function.
- 12.
The notion of allele was invented in genetics as a short form of allelomorph, which means other form, for the variants of a gene.
- 13.
Here we use 2N for the number of alleles in a population of size N, which refers to diploid organisms. For haploid organisms, 2N has to be replaced by N. In real populations, the population size is corrected for various other factors and taken to be 2N e or N e , respectively.
- 14.
The selection coefficient is denoted here by ϱ instead of s in order to avoid confusion with the auxiliary variable of the probability generation function. The definition here is the same as used by Kimura [96, 304]: ϱ > 0 implies greater fitness than the reference and an advantageous allele, ϱ < 0 reduced fitness and a deleterious allele. We remark that the conventional definition in population genetics uses the opposite sign: s = 1 means fitness zero, no progeny, and a lethal variant. In either case, selective neutrality occurs for s = 0 or ϱ = 0 (see also Sect. 5.3.3).
- 15.
The definition of Gegenbauer polynomials here is slightly different from the one given in Sect. 4.3.3: T n (β)(z) = (2β − 1)! ! C n (β+1∕2)(z).
- 16.
The definitions of the product terms of the ratios π k and ϱ k differ from those used in Sect. 3.2.3.
- 17.
The law of the mean expresses the difference in the values of a function f(x) in terms of the derivative at one particular point x = x 1 and the difference in the arguments
$$\displaystyle{ f(b) - f(a) = (b - a)\left.\frac{\partial f} {\partial x}\right \vert _{x=x_{1}}\;,\quad a < x < b. }$$The law of the mean is satisfied for at least one point x 1 on the arc between a and b.
- 18.
In real systems, we are always dealing with finite populations in finite time, and then expectation values do not diverge (but see, for example, the unrestricted birth-and-death process in Sect. 5.2.2).
- 19.
Situations may exist where it is for all practical purposes impossible to reach one population from another one through a chain of mutations in any reasonable time span. Then M is not irreducible in reality, and we are dealing with two independently mutating populations. In particular, when more involved mutation mechanisms comprising point mutations, deletions, and insertions are considered, it may be advantageous to deal with disjoint sets of subspecies.
- 20.
- 21.
The selection–mutation equation (5.46a) in the original formulation [130, 132] aut]Eigen, M. also contains a degradation term d j x j , and the corresponding definition of the value matrix reads W = (w ij = Q ij f j − d j ). If all individuals follow the same death law, i.e., d j = d, \(\forall \,j\), the parameter d can be absorbed into the population size conservation relation and need not be considered separately.
- 22.
The notation applied here is the conventional way of writing transitions in physics: W nm is the probability of the transition n ← m, whereas many mathematicians would write W mn , indicating m → n.
- 23.
When doing actual calculations, one has to use the convention 00 = 1 used in probability theory and combinatorics, but not usually in analysis, where 00 is an indefinite expression.
- 24.
A matrix W with this property is called a stochastic matrix.
- 25.
The second procedure can be visualized by a somewhat strange but nevertheless precise model assumption: after the replication event, the parent but not the offspring is put back into the pool from which the individual, which is doomed to die, is chosen in the second draw.
- 26.
In the non-degenerate case, stationary states do not depend on initial conditions, but this is no longer true for linear combinations of degenerate eigenvectors: α and β, and ϑ are functions of the initial state.
- 27.
In population genetics, the fitness parameter is conventionally denoted by s, but here we use ϱ in order to avoid confusion with the auxiliary variable s.
- 28.
The function3 F 2 belongs to the class of extended hypergeometric functions, referred to in Mathematica as HypergeometricPFQ.
- 29.
The result for ɛ is easily obtained by making use of the infinite series
$$\displaystyle{\sqrt{1 + x} = 1 + \frac{1} {2}x -\frac{1} {8}x^{2} + \frac{1} {16}x^{3} + \cdots \;,\quad 1/\sqrt{1 + x} = 1 -\frac{1} {2}x + \frac{3} {8}x^{2} - \frac{5} {16}x^{3} + \cdots \;,}$$for small x.
- 30.
Nucleic acids are linear polymers and have two different ends with the hydroxy group in the 5′ position or in the 3′ position, respectively.
- 31.
- 32.
Time is running backwards from the present τ = −t, with today as the origin (Fig. 5.29).
- 33.
The Y-chromosome in males is haploid and non-recombining.
- 34.
This means that reproduction lies in the domain of neutral evolution, i.e., all fitness values are assumed to be the same, or in other words no effects of selection are observable and the numbers of descendants of the individual alleles \(\mathcal{N}_{1},\mathcal{N}_{2},\mathop{\ldots },\mathcal{N}_{N}\), are entirely determined by random events.
- 35.
Assume that X i (n) has the ancestor X i (n + 1). The probability that X j (n) has the same ancestor is simply one out of N, i.e., 1∕N.
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Schuster, P. (2016). Applications in Biology. In: Stochasticity in Processes. Springer Series in Synergetics. Springer, Cham. https://doi.org/10.1007/978-3-319-39502-9_5
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