Diatom Morphogenesis. Группа авторов

Чтение книги онлайн.

Читать онлайн книгу Diatom Morphogenesis - Группа авторов страница 25

Diatom Morphogenesis - Группа авторов

Скачать книгу

surface. More generally, changes in this valve formation system exhibit ergodic behavior, and the stability of such behavior in a dynamical system is measurable as Lyapunov exponents.

      In a dynamical system, different states occupy different spaces that exist at different times. Changes from state to state can be delineated using a system of differential equations derived from the original relation among the variables of interest. A Lyapunov function of the general form x i = fi (x) has first derivatives that form the elements of the Jacobian matrix (i.e., Jacobian) image where x(t) is x0 at equilibrium [2.78]. The Jacobian represents the change in tangent vectors over time t as a linearization of the tangent space system [2.39].

      To determine the behavior characteristics of symmetry states, solution to the Lyapunov function of the form ATS + SA = -Q is evaluated, where Q is the resultant matrix. If the left side of the equation is determined to be positive definite when the right side of the equation is negative definite, then the symmetry states system is stable; otherwise, the system is unstable [2.23].

      Specific characteristics of instability are evaluated via A. At equilibrium, (A - 9I) = 0 and for det(A - 9I) = 0, 9 are eigenvalues of A, and tr image. For all symmetry states, ln image [2.23]. Transformation of produces image of A, and diagonalization and a change of basis leads to a decoupled system with the general solution of the time evolution matrix image [2.22].

      Lyapunov exponents [2.88, 2.101] are meaningfully characterized by the real parts of the complex valued eigenvalues of A [2.29] and are image where log is any base. Lyapunov exponents measure fluctuation as average rates of divergence (expansion and instability) or convergence (contraction and stability) and represent expected values [2.29]. Initial conditions influence the outcome, and changes in initial conditions are measurable as indicators of the behavior of fluctuations. Boltzmann entropy production is necessarily sensitive to exponentially changing initial conditions in tangent space [2.39]. However, slight perturbations in initial conditions will yield measurement of the degree and characterization of symmetry behavior at any point in the dynamical system.

      Lyapunov exponents comprise a multidimensional spectrum indicative of the rate of (exponential) divergence of multiple symmetry states with respect to direction of changes from one symmetry state to the next. Lyapunov exponents give bounds on information production of a dynamical system. They measure local behavior of the collective properties of a dynamical system yielding a global characterization of the system [2.8, 2.133]. A Lyapunov function close to equilibrium ensures that entropy tends to move toward a stable maximum in a dynamical system [2.29].

      By quantifying the relation between symmetry states and stability via Lyapunov exponents, instability is characterizable as a specific behavior, either deterministically chaotic or not. However, the picture may be more complicated. Slight changes in initial conditions can precipitate chaotic behavior resulting in instability [2.29] even though the initial conditions may be random. Chaotic systems may appear to be random when considering fewer than all possible states in the dynamical system [2.97]. High-dimensional systems may appear to be ordered despite non-convergent random behavior [2.35]. Different kinds of randomness as “noise” may appear to be chaotic in a given dynamical system [2.118]. Care must be taken to discern the behavior of a dynamical system in terms of randomness or chaoticity.

      2.2.8 Randomness and Instability

      Lyapunov exponents enable the assessment of randomness with respect to instability in a dynamical system [2.74, 2.118, 2.163]. While local unstable behavior on a state-by-state basis may be chaotic, the macro-state instability of valve formation may be inherently random at a global scale. Alternatively, there may be intermittent chaotic or random unstable states [2.74] during the morphogenetic process. Because of this, a test for randomness [2.34] is necessary.

      Thus far, our dynamical system model has been analyzed at equilibrium. However, to detect possible random instability, we need to consider the possible non-equilibrium consequences of such behavior in this system. Non-equilibrium may be a long-term phenomenon in contrast to short-term steady-states present in a dynamical system, for example, as time oscillations create Turing effects of diffusion driven instability [2.6]. Toy models have been used to link time intervals over multiple levels enabling the application of equilibrium dynamics to non-equilibrium behavior [2.96]. They have great potential in modeling complicated biological processes [2.2]. Non-equilibrium dynamics can be assessed in terms of constrained information loss or an increase in entropy [2.65].

      As entropy increases over longer and longer sequences for a given dynamical system, entropy blocks or sections that define the structure of these sequences do not characterize the entire system. What remains is randomness that is not taken into consideration and given as the source entropy rate image where L is the number of sequence steps [2.26]. One way to test for randomness is to consider the non-deterministic behavior of the valve formation simulated system as a simple renewal process [2.35, 2.74, 2.115]. In a study of morphology and inheritance, Cyclotella meneghiniana offspring cell structure was analyzed for partial renewal with regard to the mother cell and complete renewal in terms of genetic or epigenetic inheritance [2.131], so that valve formation was considered to be a renewal process.

      Evaluating entropy S using the PDF of associated probabilities involves first derivatives of S as entropy rates [2.26, 2.74] corresponding to bandwidth in the histogram used to determine the PDF [2.137]. For the expected value of the probabilities associated to image the associated Lyapunov exponents are found via Lagrange

Скачать книгу