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  • 1
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    Bulletin of mathematical biology 44 (1982), S. 1-15 
    ISSN: 1522-9602
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Mathematics
    Notes: Abstract A theorem is proved, concerning expected values of a multitype branching process in a varying environment. The consequence of the theorem is that the branching process can be treated (in the sense of expected values) as a dynamical system with control terms. This is of importance in situations where the process serves as an abstract model of the dynamics of malignant cells for use in chemotherapy. A simple example of this kind is given.
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  • 2
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    Bulletin of mathematical biology 44 (1982), S. 29-42 
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    Notes: Abstract A method is developed for a full nonlinear evaluation of all velocities and stresses represented in the Navier-Stokes equations and in the general stress tensor. The information required is essentially that for solution of linearized forms. The solution is analytical except for the calculation of the axial velocity, which requires computer assistance to step through time and space. The treatment of the problem, although directed towards solutions involving fluid flow in elastic vessels, is also adaptable to solid deformations (strain vs rate of strain) where the general stress tensor applies. A special case for the distorting ellipse is presented as well as a limited, spatially analytic, solution.
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  • 3
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    Bulletin of mathematical biology 44 (1982), S. 43-56 
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    Notes: Abstract The stability characteristics and dynamical behavior of a system of mutually excitatory neurons in close spatial proximity are investigated with a mathematical model. The model predicts the existence of uniform, intermediate levels of activity other than those of no activity and maximal activity. The model also, yeilds a good explanation of data obtained from periglomerular neurons in the olfactory bulb of the cat.
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    Bulletin of mathematical biology 44 (1982), S. 75-86 
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    Notes: Abstract A multicompartmental model in which particles enter the system from the environment and reproduce according to a Markov branching process has been considered. Explicit expressions have been obtained for the mean vector and the correlation structure for the numbers of particles in different compartments in different time points of the system. Growth rates of the mean vector and some special cases of the system are also discussed.
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    Bulletin of mathematical biology 44 (1982), S. 57-74 
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    Notes: Abstract The study of bi-directionally coupled oscillators is relevant in biological modelling of such systems as gastro-intestinal electrical activity, cardiac pacemarkers, cardiovascular and respiratory interactions and circadian rhythms. Interconnecting pathways in biological systems often exhibit pure time-delay characteristics. In this paper the multiple-mode limit-cycle behaviour of such systems is analysed using the method of harmonic blance. It is shown that the coupling time delay radically affects the number, frequency and amplitudes of entrained limit-cycles.
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    Bulletin of mathematical biology 44 (1982), S. 87-102 
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    Notes: Abstract The effect of keeping all the parameters constant, except the diffusion coefficients, in a pair of reaction-diffusion equations is studied. It is shown that the stability of the constant solution and the bifurcation points can be easily established by constructing a simple stability diagram. The possible qualitatively different diagrams are enumerated.
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    Bulletin of mathematical biology 44 (1982), S. 103-117 
    ISSN: 1522-9602
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    Notes: Abstract We have numerically examined more than one million Large Complex Systems (LCS) of interacting variables (interpretable as interacting populations) governed by Generalized Lotka-Volterra Equations (GLV), with self-regulation term. The scope was to have some insight on the stability-complexity relationship. We considered systems of prey-predator type, and we gave appropriate rules for constructing the model systems, rules that specify the behaviour of model systems in order to put them near the biological reality. The results show, among other things, a strict correlation between the stability and the prey-predator ratio (which, in our model, uniquely determines the connectedness of the system).
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  • 8
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    Bulletin of mathematical biology 44 (1982), S. 149-150 
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  • 9
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    Bulletin of mathematical biology 44 (1982), S. 119-134 
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    Notes: Abstract The rate-controlling process in the oxygenation of red blood cells is investigated using a Roughton-like model for oxygen diffusion and reaction with hemoglobin. The mathematical equations describing the model are solved using two independent techniques, numerical inversions of the Laplace transform of the equations and numerical solutions via an implicit-explicit finite difference form of the equations. The model is used to re-examine previous theoretical models that incorporate either a red cell membrane that is resistive to oxygen diffusion or an unstirred layer of water surrounding the cell. Although both models have been postulated to be equivalent, the results of the computer simulations demonstrate significant differences between the two models in the rate of oxygenation of the red cells, depending upon the values chosen for the diffusion coefficient for O2 in the membrane and the thickness of the water layer. The difference is apparently due to differences in the induction and transient periods of the water layer model relative to the membrane model.
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    Bulletin of mathematical biology 44 (1982), S. 537-547 
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    Notes: Abstract We examine certain mathematical structures presented in Part I. The most important of these are the energy structures determined by the couple (ω×E, ψ) the space of causality defined by ψ-1(0) and the notion of collapsibility, i.e., the descent of a species from a higher to a lower equilibrium configuration as a result of energy loss.
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    Bulletin of mathematical biology 44 (1982), S. 557-570 
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    Notes: Abstract This paper discusses a general stochastic model for a two-compartment reversible system with non-homogeneous Poisson inputs, arbitrary residence times at each of the compartments and time-dependent transition probabilities. The probability distributions of the number of particles in each compartment and in the system are obtained together with the number of particles which depart from the system. In addition, various covariance functions with a time lag are obtained. Some of the above obtained results are deduced for time-independent arrivals, exponential residence times and time-independent transition probabilities. Fluctuations of the particles present in the system are also analysed. Similar analysis is provided for the model into which some particles are initially introduced at the system. Some possible applications are discussed at the end.
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    Bulletin of mathematical biology 44 (1982), S. 571-578 
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    Notes: Abstract The general multispecies prey-predator system with Gompertz's antisymmetric interactions is nonlinearly stable in the absence of dispersion and continues to remain stable with dispersion under both homogeneous reservoir and zero flux boundary conditions in a region containing the equilibrium state. It is proved that a general multispecies food-web model without antisymmetric interactions is stable in the absence of dispersion and remains stable with dispersion in the above-mentioned region.
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    Bulletin of mathematical biology 44 (1982), S. 593-593 
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    Bulletin of mathematical biology 44 (1982), S. 579-585 
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    Notes: Abstract We introduce a graphical approach in the study of the qualitative behavior ofm species predator-prey systems. We prove that tree graphs imply global stability for Volterra models and local stability for general models; furthermore, we derive sufficient conditions so that loop graphs imply stability and boundedness of the solutions.
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    Bulletin of mathematical biology 44 (1982), S. 594-595 
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    Bulletin of mathematical biology 44 (1982), S. 731-739 
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    Notes: Abstract A system of integro-differential equations is derived to describe epizootics of a fungal pathogen in an insect population. Because of piecewise continuous behavior under some parametric conditions, it is concluded that standard phase orbits can be misleading. Using a different analytic approach yields a simple system of finite difference equations. Both the continuous and discrete versions are compared to classical forms. The continuous version differs from a classical one in possessing a second derivative dependent on population density. The discrete version differs in maintaining positive, non-zero populations of both infectives and susceptibles in finite time.
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    Bulletin of mathematical biology 44 (1982), S. 741-748 
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    Notes: Abstract This note is an attempt to demonstrate that hypothalamic pulsatile GnRH secretion is not the result of a short-term, negative steroid hormone feedback. Clarification of this point is of importance for further modelling the control of gonads.
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    Bulletin of mathematical biology 44 (1982), S. 749-760 
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    Notes: Abstract A modern theory of the calculus of variations is used to form necessary and sufficient conditions for the existence of a Lagrangian representation of a system of first-order ordinary differential equations. There exists a theorem to the effect that when a system of ordinary differential equations is variationally self-adjoint, the fulfillment of such conditions is guaranteed. In addition, self-adjointness, allows establishement of an algorithm by which a Lagrangian for the system may be explicitly constructed. Examples in mathematical biology are given to illustrate the use of the stated theorem.
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    Bulletin of mathematical biology 44 (1982), S. 793-808 
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    Notes: Abstract Engineering optimal control theory is applied to equations describing insulin and glucose interactions. The nature of the optimal controller is established. It is shown how the results can be utilized in a closed loop feedback control system.
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    Bulletin of mathematical biology 44 (1982), S. 777-791 
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    Notes: Abstract An attempt is made to compare the conditions for the general error-optimality of linear systems developed by Kalman with the conditions for feasibility of linear models of neuromuscular and physiological control systems. Models of three actual physiological systems are tested for both the above criteria. Theoretical analysis presented here shows that there are no simple relationships between the two sets of conditions. Analysis carried out on the physiological systems models suggests the need for a general set of conditions for other optimality criteria, such as time and energy minimization, similar to Kalman's condition for error minimization.
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    Bulletin of mathematical biology 44 (1982), S. 851-877 
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    Notes: Abstract Following arteriolar occlusion, tissue oxygen concentration decreases and anoxic tissue eventually develops. Although anoxia first appears in the region most distal to the capillary at the venous end, it eventually spreads throughout the entire region of supply. In this paper the changing oxygen concentration, from the time of occlusion until the tissue is entirely anoxic, is examined mathematically. The equations governing oxygen transport to tissue are solved by iterating a nonlinear integral equation. This solution is valid until anoxia first appears. After anoxia develops it is necessary to solve a moving boundary problem. This is done using the method of matched asymptotic expansions, and accurate solutions are obtained for a wide range of physiological conditions.
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    Bulletin of mathematical biology 44 (1982), S. 899-900 
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    Bulletin of mathematical biology 55 (1993), S. 1-13 
    ISSN: 1522-9602
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    Notes: Abstract A simple one-dimensional model of single-species populations is studied by means of computer simulations. Although the model has a rich spectrum of dynamics including chaotic behavior, the introduction of survival thresholds makes the chaotic region so small that it can be hardly observed. Stochastic fluctuations further reduce the chaotic region because they accidentally lead populations to extinction. The model thus naturally explains the observation that the majority of natural populations do not show chaotic behavior but a monotonic return to a stable equilibrium point following a disturbance.
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    Notes: Abstract Current understanding of the pattern of proliferation within intestinal crypts involves the notion of a cutoff region introduced by Cairnieet al. (Exp. Cell. Res. 39, 539–553, 1965b). (Cells produced above the cutoff are non-cycling, whereas cells produced below the cutoff are cycling.) They contrasted the predicted distribution of proliferation in the extreme cases of a cutoff of width 0 (a sharp cutoff) with one eight cells wide (a slow cutoff) and concluded that the data were better explained by the latter. We have shown that crypt size variation artificially broadens the apparent distribution of proliferating cells in the crypt (Totafurnoet al., Biophys. J. 54, 845–858, 1988). Here we show that the measurement and analysis of crypts of a specified height reduces this artifact. This work introduces the use of distance from the crypt base (in microns) to specify the location of cells within the crypt as an improvement over the cell position ordering traditionally used in the determination of the distribution of proliferating cells. We also show how to explicitly correct for several artifacts in the measurement of the labelling index. We conclude that cell proliferation within the crypt is more localized than previously realized; in fact, a cutoff as slow as eight cells wide is rejected.
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    Bulletin of mathematical biology 55 (1993), S. 141-154 
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    Notes: Abstract Multiple string (sequence) alignment is a difficult and important problem in computational biology, where it is central in two related tasks: finding highly conserved subregions or embedded patterns of a set of biological sequences (strings of DNA, RNA or amino acids), and inferring the evolutionary history of a set of taxa from their associated biological sequences. Several precise measures have been proposed for evaluating the goodness of a multiple alignment, but no efficient methods are known which compute the optimal alignment for any of these measures in any but small cases. In this paper, we consider two previously proposed measures, and given two computationaly efficient multiple alignment methods (one for each measure) whose deviation from the optimal value isguaranteed to be less than a factor of two. This is the novel feature of these methods, but the methods have additional virtues as well. For both methods, the guaranteed bounds are much smaller than two when the number of strings is small (1.33 for three strings of any length); for one of the methods we give a related randomized method which is much faster and which gives, with high probability, multiple alignments with fairly small error bounds; and for the other measure, the method given yields a non-obviouslower bound on the value of the optimal alignment.
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    Bulletin of mathematical biology 55 (1993), S. 197-212 
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    Notes: Abstract The kinematics of helical motion are descirbed for an organism treated as a rigid body with six degrees of freedom relative to the organism's frame of reference, i.e. the organism can translate in the direction of, or rotate around any of, three orthogonal axes fixed to its body. Equations are derived that express the unit vectors of the Frenet trihedron and the torsion and curvature of the trajectory in terms of the organism's translational and rotational velocities. These equations permit description of the radius, pitch, angular velocity and axis of a helical trajectory in terms of the translational and rotational velocities of the organism swimming along that trajectory. The results of this analysis are then used in two later papers that describe how organisms can orient to an external stimulus.
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    Bulletin of mathematical biology 55 (1993), S. 257-257 
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    Bulletin of mathematical biology 55 (1993), S. 231-255 
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    Notes: Abstract Organisms that move along helical trajectories change their net direction of motion largely by changing the direction, with respect to the body of the organism, of their rotational velocity (Crenshaw and Edelstein-Keshet, 1993,Bull. math. Biol. 55, 213–230). This paper demonstrates that an organism orients to a stimulus field, such as a chemical concentration gradient or a ray of light, if the components of its rotational velocity, with respect to the, body of the organism, are simple functions of the stimulus intensity encountered by the organism. For example, an organism can orient to a chemical concentration gradient if the rate at which it rotates around its anterior-posterior axis is proportional to the chemical concentration it encounters. Such an orientation can be either positive or negative. Furthermore, it is true taxis—orientation of the axis of helical motion is direct. It is neither a kinesis nor a phobic response—there is no random component to this mechanism of orientation.
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    Bulletin of mathematical biology 55 (1993), S. 277-294 
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    Notes: Abstract A basic but neglected property of neuronal trees is their finite length. This finite length restricts the length of a segment to a certain maximum. The implications of the finite length of the tree with respect to the segment length distributions of terminal and intermediate segments are shown by means of a stochastic model. In the model it is assumed that branching is governed by a Poisson process. The model shows that terminal segments are expected to be longer than intermediate segments. Terminal and intermediate segments are expected to decrease in length with incrasing centrifugal order. The results are compared with data fromin vivo pyramidal cells from rat brain and tissue cultured ganglion cells from chicken. A good agreement between data and model was found.
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    Bulletin of mathematical biology 55 (1993), S. 345-364 
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    Notes: Abstract Shape and size of elongating cells were examined in three plant tissues: the adaxial epidermis of the petiole ofZebrina pendula L., the abaxial epidermis ofAnacharis densa L. leaves and the abaxial epidermis of the scale leaf ofAllium cepa L. Based on a few simple assumptions, the expected probability distribution frequencies (pdf) for cell length and number of adjacent walls were calculated. Actual data of cell lengths closely approximated those expected with the pdfs being asymmetrical since there are more younger, shorter cells than older, longer cells. Data for number of lateral walls of real cells were similar to that expected and these walls increase in compensating mechanism exists to maintain a constant range of cell lengths through many cell generations. It is expressed by longer than average new daughter cells dividing relatively soon while shorter than average new daughter cells divide after a relatively long cycle.
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    Bulletin of mathematical biology 55 (1993), S. 365-384 
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    Notes: Abstract Diffusion driven instability in reaction-diffusion systems has been proposed as a mechanism for pattern formation in numerous embryological and ecological contexts. However, the possible effects of environmental inhomogeneities has received relatively little attention. We consider a general two species reaction-diffusion model in one space dimension, with one diffusion coefficient a step function of the spatial coordinate. We derive the dispersion relation and the solution of the linearized system. We apply our results to Turing-type models for both embryogenesis and predator-prey interactions. In the former case we derive conditions for pattern to be isolated in one part of the domain, and in the latter we introduce the concept of “environmental instability”. Our results suggest that environmental inhomogeneity could be an important regulator of biological pattern formation.
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    Notes: Abstract The particular dynamics of the previously proposed model of a catalytic network formed byn error-prone self-replicative species without and with superimposed competition is analysed. In the first case, two situations are studied in detail: a uniform network in which all the species are inter-coordinated in the same way, and a network with a species differentiated in its catalytic relation with the remaining elements. In the second case, the superimposed competition is introduced at two levels: first, as an asymmetry in one of the network species amplification factor considering a null self-catalytic vector, and secondly, as a non-null self-catalytic vector with no asymmetry in the other propertics of the species. This kind of system does not present complex behaviour and can be adequately deseribed by performing a standard linear analysis, which gives direct information on the asymptotic behaviour of the sytem. Finally, the biological implications of this analysis within the framework of biological evolution are discussed.
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    Bulletin of mathematical biology 55 (1993), S. 451-464 
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    Notes: Abstract A theoretical model is proposed for the formation of cell distribution patterns in the slug stage of the cellular slime moldDictyostelium discoideum. The equilibrium distribution of two types of cells, prestalk and prespore, is obtained by minimizing the free energy, which is defined in terms of differential chemotaxis, differential cell adhesion and randomness of cell movement. Resulting distributions show various segregation patterns of cell types. The condition for cell sorting is obtained from stability analysis of the set of diffusion equations governing the evolution of cell type distribution and the concentration of chemoattractant. The intensities of differential chemotaxis and random cell movement are quantitatively evaluated from experimental data to show that two cell types can sort themselves completely by these forces.
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    Bulletin of mathematical biology 55 (1993), S. 655-674 
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    Notes: Abstract Multicell spheroids, small spherical clusters of cancer cells, have become an importantin vitro model for studying tumour development given the diffusion limited geometry associated with many solid tumour growths. Spheroids expand until they reach a dormant state where they exhibit a grossly static three-layered structure. However, at a cellular level, the spheroid is demonstrably dynamic with constituent cells migrating from the outer well-nourished region of the spheroid toward the necrotic central core. The mechanism that drives the migrating cells in the spheroid is not well understood. In this paper we demonstrate that recent experiments on internationalization can be adequately described by implicating pressure gradients caused by differential cell proliferation and cell death as the primary mechanism. Although chemotaxis plays a role in cell movement, we argue that it acts against the passive movement caused by pressure differences.
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    Bulletin of mathematical biology 55 (1993), S. 675-691 
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    Bulletin of mathematical biology 55 (1993), S. 693-693 
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    Bulletin of mathematical biology 55 (1993), S. 695-713 
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    Notes: Abstract In recent years, methods of consensus, developed for the solution of problems in the social sciences, have become widely used in molecular biology. Westudy a method of consensus originally due to Watermanet al. (Waterman, Galas and Arratia. 1984. Pattern recognition in several sequences: consensus and alignment.Bull. math. Biol. 46, 515–527) which is used to identify patterns or features in a molecular sequence where a pattern can vary in position within a given window. We show that some well-known consensus methods of the social sciences, the median and the mean, are special cases of this method for certain choices of the parameters used in it and give a precise account of the parameters for which these special cases arise. We also show that the specific parameters used in the method of Watermanet al. make their method equivalent to the median procedure which is widely used in the social sciences.
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    Bulletin of mathematical biology 55 (1993), S. 745-780 
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    Notes: Abstract We develop a model for the idiotypic interaction between two B cell clones. This model takes into account B cell proliferation, B cell maturation, antibody production, the formation and subsequent elimination of antibody-antibody complexes and recirculation of antibodies between the spleen and the blood. Here we investigate, by means of stability and bifurcation analysis, how each of the processes influences the model's behavior. After appropriate nondimensinalization, the model consists of eight ordinary differential equations and a number of parameters. We estimate the parameters from experimental sources. Using a coordinate system that exploits the pairwise symmetry of the interactions between two clones, we analyse two simplified forms of the model and obtain bifurcation diagrams showing how their five equilibrium states are related. We show that the so-called immune states lose stability if B cell and antibody concentrations change on different time scales. Additionally, we derive the structure of stable and unstable manifolds of saddle-tye equilibria, pinpoint their (global) bifurcations and show that these bifurcations play a crucial role in determining the parameter regimes in which the model exhibits oscillatory behavior.
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    Bulletin of mathematical biology 55 (1993), S. 781-816 
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    Notes: Abstract Two types of behavior have been previously reported in models of immune networks. The typical behavior of simple models, which involve B cells only, is stationary behavior involving several steady states. Finite amplitude perturbations may cause the model to switch between different equilibria. The typical behavior of more realistic models, which involve both B cells and antibody, consists of autonomous oscillations and/or chaos. While stationary behavior leads to easy interpretations in terms of idiotypic memory, oscillatory behavior seems to be in better agreement with experimental data obtained in unimmunized animals. Here we study a series of models of the idiotypic interaction between two B cell clones. The models differ with respect to the incorporation of antibodies, B cell maturation and compartmentalization. The most complicated model in the series has two realistic parameter regimes in which the behavior is respectively stationary and chaotic. The stability of the equilibrium states and the structure and interactions of the stable and unstable manifolds of the saddle-type equilibria turn out to be factors influencing the model's behavior. Whether or not the model is able to attain any form of sustained oscillatory behavior, i.e. limit cycles or chaos, seems to be determined by (global) bifurcations involving the stable and unstable manifolds of the equilibrium states. We attempt to determine whether such behavior should be expected to be attained from reasonable initial conditions by incorporating an immune response to an antigen in the model. A comparison of the behavior of the model with experimental data from the literature provides suggestions for the parameter regime in which the immune system is operating.
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    Bulletin of mathematical biology 55 (1993), S. 865-867 
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    Bulletin of mathematical biology 55 (1993), S. 869-889 
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    Notes: Abstract We show that the existence of diffusional resistance due to the presence of a solid phase can have a positive effect on the metabolic reactions of plant cells. In this case the efficiency of metabolic reactions, defined as the ratio of rate of production of biomass of aggregated cells/rate of production of biomass of dispersed cells, can be greater than unity for a certain range of aggregate sizes for both solid spheres (common plant cell aggregates) and hollow spheres (e.g.Volvox aggregates). This means that, under appropriate conditions, plant cells tend to stay in the aggregated form to improve the efficiency of their metabolic reactions. The result of the present analysis provides an explanation as to why aggregates of plant cells are observed under typical culture conditions.
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    Bulletin of mathematical biology 55 (1993), S. 937-952 
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    Notes: Abstract The Hodgkin and Huxley equations model action potentials in squid giant axons. Variants of these equations are used in most models for electrial activity of excitable membranes. Computational tools based upon the theory of nonlinear dynamical systems are used here to illustrate how the dynamical behavior of the Hodgkin Huxley model changes as functions of two of the system parameters.
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    Bulletin of mathematical biology 55 (1993), S. 919-936 
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    Notes: Abstract The description of the “microbial loop” has led to some major changes in our understanding of nutrient cycling within aquatic ecosystems. It now appears that in many settings it is not uncommon for some 50% of phytoplankton production to be diverted into microbial pathways rather than passing up to higher trophic levels. As a result the microbial loop is responsible for enhanced and rapid nutrient cycling at the very base of the food web. Since tight recycling is often associated with unstable positive feedback, we use a model to examine the possible repercussions in more detail. The model simulates the dynamics of the microbial loop and finds it to greatly affect the way in which aquatic primary production responds to nutrient pulses.
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    Bulletin of mathematical biology 55 (1993), S. 953-971 
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    Notes: Abstract The maintenance activity of plants is investigated in terms of a simple model. Maximization of a certain biomass fraction we refer to asnonactive biomass is postulated. Optimal behaviour of plants according to this principle is explicitly derived and expressed depending on environmental conditions. Several interesting hypotheses result, e.g. a quadratic law relating specific growth rate and gross rate of photosynthesis. A qualitative comparison with data from the literature is performed, with a special emphasis on the question whether plants stressed by air pollutants repair optimally. Regarding long-term constant environmental conditions, no data were found that contradict optimal behaviour. Exact quantitative testing of the theory is desirable, appropriate experiments are suggested.
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    Bulletin of mathematical biology 55 (1993), S. 993-1011 
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    Notes: Abstract In an earlier work a model of the autocrine and paracrine pathways of tumor growth control was developed (Michelson and Leith. 1991. Autocrine and paracrine growth factors in tumor growth.Bull. math. Biol. 53, 639–656). The target population, a generic tumor, was modeled as a single, homogeneous population using the standard Verhulst equation of logistic growth. Mitogenic signals were represented by modifications to the Malthusian growth parameter and adaptational signals were represented by modifications to the carrying capacity. Three growth scenarios were described: (1) normal tissue wound healing, (2) unperturbed tumor growth, and (3) tumor growth in a radiation damaged environment, a phenomenon termed the Tumor Bed Effect (TBE). In this paper, we extend those results to include a “triad” of growth factor controls (autocrine, paracrine and endocrine) and heterogeneity of the target population. The heterogeneous factors in the model represent either intrinsic, epigenetic or environmental differences in both normally differentiating tissues and tumors. Three types of growth are modeled: (1) normal tissue differentiation or wound healing, assuming no communication between differentiated and undifferentiated cell compartments; (2) normal wound healing with feedback inhibition, due to signalling from the differentiated compartment; and (3) the development of hypoxia in a spherical tumor. The signal processing within the triad is discussed for each model and biologically reasonable constraints are defined for limits on growth control.
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    Bulletin of mathematical biology 55 (1993), S. 1039-1061 
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    Notes: Abstract A transient multispecies model for quantifying microbial space competition in biofilm is derived from existing models, introducing a new approach to biomass detachment modelling. This model includes inert biomass, substrate diffusion and utilization rate within the biofilm and diffusional layers. It predicts the evolution of biofilm thickness, bulk substrate concentration, species distribution and substrate concentration within the biofilm. A zero-dimensional transient model is described. Its steady-state solution is used to set up initial conditions of the one-dimensional model and case computation towards steady-state solution. Some numerical tools have been developed, enabling fast computation on microcomputers. Simulations show the validity of a zero-dimensional model and perturbated systems are also simulated. Simulations with experimental data give acceptable results.
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    Bulletin of mathematical biology 55 (1993), S. 1025-1038 
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    Notes: Abstract Recently, we proposed a new model of DNA sequence evolution (Arquès and Michel. 1990b.Bull. math. Biol. 52, 741–772) according to which actual genes on the purine/pyrimidine (R/Y) alphabet (R=purine=adenine or guanine, Y=pyrimidine=cytosine or thymine) are the result of two successive evolutionary genetic processes: (i) a mixing (independent) process of non-random oligonucleotides (words of base length less than 10: YRY(N)6, YRYRYR and YRYYRY are so far identified; N=R or Y) leading to primitive genes (words of several hundreds of base length) and followed by (ii) a random mutation process, i.e. transformations of a base R (respectively Y) into the base Y (respectively R) at random sites in these primitive genes. Following this model the problem investigated here is the study of the variation of the 8 R/Y codon probabilities RRR,..., YYY under random mutations. Two analytical expressions solved here allow analysis of this variation in the classical evolutionary sense (from the past to the present, i.e. after random mutations), but also in the inverted evolutionary sense (from the present to the past, i.e. before random mutations). Different properties are also derived from these formulae. Finally, a few applications of these formulae are presented. They prove the proposition in Arquès and Michel (1990b.Bull. math. Biol. 52, 741–772), Section 3.3.2, with the existence of a miximal mean number of random mutations per base of the order 0.3 in the protein coding genes. They also confirm the mixing process of oligonucleotides by excluding the purine/pyrimidine contiguous and alternating tracts from the formation process of primitive genes.
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    Bulletin of mathematical biology 55 (1993), S. 1199-1210 
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    Notes: Abstract It is believed that the native folded three-dimensional conformation of a protein is its lowest free energy state, or one of its lowest. It is shown here that both a two-and three-dimensional mathematical model describing the folding process as a free energy minimization problems is NP-hard. This means that the problem belongs to a large set of computational problems, assumed to be very hard (“conditionally intractable”). Some of the possible ramifications of this results are speculated upon.
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    Bulletin of mathematical biology 55 (1993), S. 1133-1182 
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    Notes: Abstract A model employing separate dose-dependent response functions for proliferation and differentiation of idiotypically interacting B cell clones is presented. For each clone the population dynamics of proliferating B cells, non-proliferating B cells and free antibodies are considered. An effective response function, which contains the total impact of proliferation and differentiation at the fixed points, is defined in order to enable an exact analysis. The analysis of the memory states is restricted in this paper to a two-species system. The conditions for the existence of locally stable steady states with expanded B cell and antibody populations are established for various combinations of different field-response functions (e.g. linear, saturation, log-bell functions). The stable fixed points are interpreted as memory states in terms of immunity and tolerance. It is proven that a combination of linear response functions for both proliferation and differentiation does not give rise to stable fixed points. However, due to competition between proliferation and differentiation saturation response functions are sufficient to obtain two memory states, provided proliferation preceeds differentiation and also saturates earlier. The use of log-bell-shaped response functions for both proliferation and differentiation gives rise to a “mexican-hat” effective response function and allows for multiple (four to six) memory states. Both a primary response and a much more pronounced secondary response are observed. The stability of the memory states is studied as a function of the parameters of the model. The attractors lose their stability when the mean residence time of antibodies in the system is much longer than the B cells' lifetime. Neither the stability results nor the dynamics are qualitatively chanbed by the existence of non-proliferating B cells: memory states can exist and be stable without non-proliferating B cells. Nevertheless, the activation of non-proliferating B cells and the competition between proliferation and differentiation enlarge the parameter regime for which stable attractors are found. In addition, it is shown that a separate activation step from virgin to active B cells renders the virgin state stable for any choice of biologically reasonable parameters.
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    Bulletin of mathematical biology 44 (1982), S. 17-28 
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    Notes: Abstract The concept of natural selection is examined, which is one of basic principles for the Darwinian interpretation of evolution. In this model selection is defined as a solution of the deterministic Eigen equation. Next, the random effect is introduced through the mutation term. However, the probability of finding the solution expressing the selection is shown to be smallest. The validity of the model and its applicability to polynucleotide replication are discussed.
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    Bulletin of mathematical biology 44 (1982), S. 135-147 
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    Notes: Abstract Using numerical methods, the initial rates of oxygen uptake by the red blood cell have been computed. The methods accommodate both a water layer and membrane which may act as diffusive impedance to gas influx. The differential solubilities of the gas in these two layers have also been incorporated in the methods. The presence of a 0.50–0.65 μm deoxygenated water layer has been calculated to simulate the experimental results by Roughton (1959). Experimental studies of CO and NO uptake by the red cell could also be simulated. Although a membrane-only model with given parameters can also account for the observed rates of oxygenation of the red cell (Weingardenet al., submitted for publication), the additional incorporation of differential solubilities of oxygen in the different layers of the RBC yields results that indicate a three layer model to be more plausible. Using a thin layer-red cell oxygenation system, the rates of oxygenation were determined for red cells surrounded by a 4.2 μm deoxygenated water layer. The rates were found to compare favorably to the results of the theoretical model.
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    Bulletin of mathematical biology 44 (1982), S. 151-151 
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    Bulletin of mathematical biology 44 (1982), S. 152-152 
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    Bulletin of mathematical biology 44 (1982), S. 175-192 
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    Notes: Abstract Due to the complicated physiological structure of soft biological tissues, stresses can only be measured after the specimen has been stretched to many times of its related length. Therefore, the classical constitutive equations of finite elasticity developed for vulcanized, rubbery materials and the linear theories developed for most engineering materials cannot be applied to soft tissues which are highly elastic in nature. In this article, utilizing a mechanical model developed by Demiray for soft tissues, a class of finite deformations of some tissues is studied and the results are compared with experiment and the existing literature. These problems are the simultaneous extension and twisting of a circular cylindrical bar, the bending of a rectangular block, and the pure shear of a rectangular prism. It is believed that solutions to these problems may find some applications in plastic surgery.
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    Notes: Abstract In this paper, effects of convective and dispersive migration on the linear stability of the equilibrium state of a two species system with mutualistic interactions and functional response have been investigated. In both finite and semi-infinite habitats, it has been shown that the otherwise stable equilibrium state without dispersal remains so with dispersal also, both under flux and reservior conditions. In the case of finite habitat, the degree of stability increases as dispersal coefficients of the two species increase. The effect of convective migration also is to stabilize the equilibrium state in this case.
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    Bulletin of mathematical biology 44 (1982), S. 307-307 
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    Bulletin of mathematical biology 44 (1982), S. 283-305 
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    Notes: Abstract A linear spatially distributed model of a chain of neurons and interneurons was investigated in relation to the generation of propagated alpha rhythmic activity. It was assumed that the elements of the chain were interconnected by means of recurrent collaterals and inhibitory fibres in such a way that the connectivity functions were assumed to be homogeneous and their strength was an exponentially decreasing function of distance. It was found that such a neuronal chain shows propagation properties for frequencies in the alpha band. The results obtained with the model are in agreement with the phase velocities encountered experimentally. In this way, it was possible to estimate the length of the neural fibres responsible for the phenomenon of propagated activity. The estimates obtained are in good agreement with recent quantitative neuroanatomical data on the circuitry of the neocortex.
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    Bulletin of mathematical biology 44 (1982), S. 309-320 
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    Notes: Abstract This paper characterizes the cycle structure of a completely random net. Variables such as number of cycles of a specified length, number of cycles, number of cyclic states and length of cycle are studied. A square array of indicator variables enables conveninent study of moment structure. Additionally, exact and asymptotic distributional results are presented.
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    Bulletin of mathematical biology 44 (1982), S. 321-337 
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    Notes: Abstract A class of nonlinear equations describing the steady propagation of a disturbance on the infinite interval in one dimensional space are shown under certain conditions to admit solution with a unique velocity of propagation. The class of equations describe both initial and final homogeneous steady states which are asymptotically stable with respect to uniform perturbations, in contrast to the Fisher equation, which does not.
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    Bulletin of mathematical biology 44 (1982), S. 399-409 
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    Notes: Abstract A model for insulin secretion with a storage and a labile compartment, as well as a provisionary factor, is combined with a signal model in which the signal can be the difference between an excitation and an inhibition, or the difference in concentrations inside and outside some cell components. The model, using a single set of values for the parameters, accounts in a semiquantitative manner for all of the regularly appearing features of the insulin secretion from thein vitro perfused pancreas to a wide range of patterns of glucose and tolbutamide stimulation. Among the features which can be accounted for are: early and late secretion of insulin as a function of glucose in terms of a single parameter; the apparent depletion and recovery during a pulsed pattern of stimulation by tolbutamide; the hypersecretion following a short period of rest during a prolonged stimulation by glucose; the negative spike which occurs when the concentration of glucose, which has been maintained for a period of time, is suddenly reduced to a lower level; and the appropriate responses to slow and fast ramp functions of glucose concentration.
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    Bulletin of mathematical biology 44 (1982), S. 443-447 
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    Notes: Abstract The incorporation of a chaotic component in a computing system is incompatible with its being effectively programmable. The example presented shows that concepts of programming suitable for biological systems may differ from those which have grown out of our experience with present day digital computers.
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    Bulletin of mathematical biology 44 (1982), S. 411-423 
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    Notes: Abstract An analysis of the interaction between stimulus molecules and the olfactory receptor cell membrane is presented. The model is based upon a sequence of events, i.e. stimulus delivery at the olfactory epithelium, absorption of molecules in the mucus layer, diffusion of the molecules towards the receptor cells and molecule-receptor cell membrane interaction. The mathematical analysis considers the situation during electrophysiological experiments, where an odour puff is delivered at an exposed olfactory mucosa. Such a situation resembles sniffing of odour samples. The analysis is discussed in relation to experimental evidence.
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    Bulletin of mathematical biology 44 (1982), S. 425-442 
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    Notes: Abstract A model is developed to calculate the deposition of hygroscopic aerosols in the human tracheobronchial (TB) tree. The TB airflow pattern assumed is consistent with experimental observations and accounts for anatomical features such as the larynx and cartilaginous rings in large airways. Some original deposition efficiency formulae are presented for laminar and turbulent airstreams. Stepwise growth is simulated by changes in particle size and density at each TB generation. The dose distribution of NaCl aerosols is studied as a function of inhaled particle size and flow rate. Two NaCl growth rate curves are used which differ in the mode of aerosol-air mixing in the trachea. The initial rate of aerosol mixing in the human due to the laryngeal jet is shown to be an important factor affecting the deposition of hygroscopic aerosols. Total TB deposition of NaCl exceeds that for nonhygroscopic particles of the same inhaled aerodynamic size. Hygroscopic growth can also influence the regional TB distribution of dose when submicron NaCl particles grow rapidly enough to deposit by impaction and sedimentation.
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    Bulletin of mathematical biology 44 (1982), S. 449-449 
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    Bulletin of mathematical biology 44 (1982), S. 451-452 
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    Bulletin of mathematical biology 44 (1982), S. 491-500 
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    Notes: Abstract Granted that a single or complex gene is responsible for inbreeding depression, theoretical expressions for fertility and viability are obtained in different diploid populations: brother-sister, half-brother-sister, cousins and double-cousins. The conclusions of the study of viability variations according to the coefficient of parentage are proved by the results of experiments and lead to a new view of genetic load.
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    Bulletin of mathematical biology 44 (1982), S. 501-535 
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    Notes: Abstract The Thom gradient model of morphogenesis poses the followinga posteriori problem: “From the observed morphology of a given natural process (effect) determine the dynamics of the process (cause)”. In this paper we consider the classicala priori problem: “Given the cause (dynamics) determine the effect (resultant morphology)”. We find that in biochemical processes the mechanisms for energy activation, energy-matter interaction and energy dissipation determine the dynamics. Furthermore there exists basic energy mechanisms which drive the equilibrium states through the elementary catastrophes of Thom. A comparison with current theories shows that our models describe open ecological food chains and their dynamical systems generalize the equations of organisation posed by M. Eigen.
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    Bulletin of mathematical biology 44 (1982), S. 549-555 
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    Notes: Abstract We show that the elementary models of biochemical evolutionary processes are the bases for the study of open ecological systems and macromolecular self-organisation. We deduce a biochemical analogue for the basic closed phytoplankton-zooplankton food chain. The perturbation of the Michaelis-Menten mechanism which determines the nutrient-dependent growth rate of the phytoplankton species leads to higher catastrophes for the nutrient equilibrium manifold. The umbilic models are generalizations of the equations of organization of Eigen.
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    Bulletin of mathematical biology 44 (1982), S. 761-775 
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    Notes: Abstract The transport equation of Kedem and Katchalsky for the flux of ions through a membrane is generalized to demonstrate explicitly the role of impermeant ions in determining its mathematical form. Whereas the Kedem-Katchalsky equation is linear in the salt concentrations in the bathing solutions, the more general equation is bilinear (and symmetric) in the ionic concentrations of the permeant species. The Kedem-Katchalsky flux equation is further generalized to include explicitly a term for ion-exchange in systems having more than a single permeant salt. This additional term is also bilinear (and antisymmetric) in the concentrations of the exchanging ionic species. Flux equations are derived for systems having (1) a single mono-monovalent salt, (2) two mono-monovalent salts and (3) an arbitrary number of salts with no restriction upon the valencies of the ionic components. Since it has no effect upon the form of concentration-dependent terms in the flux equations, coupling to volume flow is neglected.
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    Bulletin of mathematical biology 44 (1982), S. 819-840 
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    Notes: Abstract A mathematical theory of the dynamics of the myocardium based on the concept of body forces is applied to account for two different experimental results. The first result consists of a relation between oxygen consumption and the kinetic energy of the myocardium. The second set of results provides a consistent interpretation of the relationship between different indices used to assess the ventricular performance.
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    Bulletin of mathematical biology 44 (1982), S. 809-817 
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    Notes: Abstract In this paper a semi-Markov process approach is developed to analyse stochastic compartmental systems using straightforward probabilistic arguments. Explicit expressions for several characteristics of thek-compartmental systems with a Poisson process input are derived and various models found in the literature arising from biological applications are generalised here using the semi-Markov process technique.
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    Bulletin of mathematical biology 44 (1982), S. 841-849 
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    Notes: Abstract A mathematical model for oxygen diffusion in a spherical cell with Michaelis-Menten oxygen uptake kinetics is analyzed by means of an intergral equation method. It is shown that an integral equation formulation can be used to obtain a numerical solution associated with this boundary and initial value problem. Through an illustrative numerical calculation we are able to obtain an accurate solution for both the steady and transient problems. Finally, a comparison is made with the numerical solution of McElwain and the variational solution of Anderson and Arthurs for the steady state and Lin's result concerning the unsteady state.
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    Bulletin of mathematical biology 44 (1982), S. 893-897 
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    Notes: Abstract Exact solutions are obtained and discussed for classes of Lotka-Volterra and Leslie-Gower systems governing the interaction of two species. The classes are defined by certain constraints which are imposed on the time-dependent parameters of the equations. A general result for such systems is that each species is characterised by two time-scales: one representing natural growth and the other, the interdependence of the species.
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    Bulletin of mathematical biology 44 (1982), S. 879-891 
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    Notes: Abstract We present an analysis of the diffusion of a tracer in a model of a cell-intercellular space system. The problem reduces to the resolution of a system of a linear partial differential equation and of a linear integral differential equation. The mathematical results have been obtained in terms of their Laplace transforms, which have been inverted by a numerical procedure for some parameter values. The importance of considering gradients of concentrations in intercellular spaces instead of lumping them with the external mediums has been discussed together with the possibility of extending Ussing's relation to transient cases, in order to detect active transports. Some possible implementations of the model to take into account more general situations have been considered.
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    Bulletin of mathematical biology 55 (1993), S. 891-918 
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    Notes: Abstract We present an algorithm for allocating individual ants to tasks that relies solely on task change being caused by the unavailability of work. We prove that such an algorithm will allocate the correct number of individuals to each job. Furthermore, we can demonstrate that if such an algorithm is used then an age structure emerges over the ants performing the various tasks. This matches closely with the weak temporal structure over tasks that is observed in Sendova-Franks and Franks (1993. Division of labour in ants nests within highly variable environments. (A study of temporal polyethism: experimental).Bull. math. Biol. 55, 75–96).
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    Bulletin of mathematical biology 55 (1993), S. 1013-1024 
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    Bulletin of mathematical biology 55 (1993), S. 973-991 
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    Notes: Abstract Biological regulatory systems can be described in terms of non-linear differential equations or in logical terms (using an “infinitely non-linear” approximation). Until recently, only part of the steady states of a system could be identified on logical grounds. The reason was that steady states frequently have one or more variable located on a threshold (see below); those steady states were not detected because so far no logical status was assigned to threshold values. This is why we introduced logical scales with values 0,1θ, 12θ, 2, ..., in which1θ,2θ, ... are the logical values assigned to the successive thresholds of the scale. We thus have, in addition to the regular logical states,singular states in which one or more variables is located on a threshold. This permits identifyingall the steady states on logical grounds. It was noticed that each feedback loop (or reunion of disjointed loops) can be characterized by a logical state located at the thresholds at which the variables of the loop operate. This led to the concept ofloop-characteristic state, which, as we will see, enormously simplifies the analysis.The core of this paper is a formal demonstration that among the singular states of a system, only loop-characteristic states can be steady. Reciprocally, given a loop-characteristic state, there are parameter values for which this state is steady; in this case, the loop is effective (i.e. it generates multistationarity if it is a positive loop, homeostasis if it is a negative loop). This not only results in the above-mentioned radical simplification of the identification of the steady states, but in an entirely new view of the relation between feedback loops and steady states.
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    Bulletin of mathematical biology 44 (1982), S. 259-269 
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    Notes: Abstract A discrete time stochastic model is formulated for the spread of a disease which is transmitted to an uninfected but susceptible individual through an environmental source and not through contact (either direct or indirect) with infected individuals. The model incorporates both exposure and infection components. The exposure component includes consideration of the introduction of an infectious agent into the environment and the subsequent diffusion of the agent. It also includes time and location patterns for visits by individuals in the target population to the affected environment. The infection component incorporates physiological responses of exposed individuals to the infectious agent. The goal of the model is to provide a method for developing a predicted epidemic curve. Comments are given on an application of the model to the study of an outbreak of toxoplasmosis in Atlanta, Georgia, in 1977.
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    Bulletin of mathematical biology 44 (1982), S. 450-450 
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    Bulletin of mathematical biology 44 (1982), S. 453-476 
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    Notes: Abstract The second order nonlinear differential equation $$\ddot x + \left( {\alpha _0 x + \alpha _1 } \right)\dot x + \alpha _2 x + \left( {x + \alpha _3 } \right) = 0$$ arises from a kinetics model of abrin binding in an Epstein-Barr virus-transformed lymphocyte culture. Some results on the dynamical behavior of this equation are given. These results are then discussed in relation to the known kinetics behavior of abrin in an EBV-lymphocyte cell culture.
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    Bulletin of mathematical biology 44 (1982), S. 477-489 
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    Notes: Abstract Measurements of the binding of ligand to receptors that are macromolecules, either free or components of biomembranes, often show deviation from what is expected of a simple reaction described by an association and a dissociation rate constant. A more versatile model and more discriminating experiments are required for a satisfactory explanation. This paper is based on a general model of the binding reaction in which the rate constants and equilibrium constant are dependent upon occupancy of receptors. The analysis of the model leads to three kinds of experiments: (1) equilibrium measurements which permit quantitative determination of a dissociation equilibrium parameter as a function of receptor occupancy; (2) measurements prior to equilibrium which yield the same information; and (3) measurements prior to equilibrium which reveal quantitatively the dependence of both association and dissociation rate parameters separately, on occupancy.
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    Bulletin of mathematical biology 44 (1982), S. 705-713 
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    Notes: Abstract Macromolecules and their aggregates (such as protein bundles in biomembranes) possess polar modes which, when excited, tend to deform the system and call into play elastic restoring forces. A model of such systems, characterised typically by electric polarisation modes stabilised on the one hand by quartic self-interactions and on the other through coupling to the elastic deformations, admits the possibility of localised excitations (solitary waves) propagating with subsonic velocities, possessing the features of relative stability and efficient transport characteristics (associated with the collective nature of the phenomena), and at the same time provides a mechanism of control and variability which could be of considerable interest in biology.
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    Bulletin of mathematical biology 44 (1982), S. 715-730 
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    Notes: Abstract Random Boolean networks have striking properties of self-organization. In this paper we propose an algorithm based on the different roles of Boolean mappings and on the connection structure to analyze the organization of the network. For a few cases— transfer mappings, AND/OR, equivalence/XOR—rigorous results are obtained about the dynamics of homogeneous networks. Conclusions are then drawn concerning the non-homogeneous networks.
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    Notes: Abstract Based on experimental work on the ontogeny of the electroretinogram circadian rhythm in crayfish, we present a mathematical model simulating changes in both frequency and amplitude of the electroretinogram oscillation during several developmental stages until shortly before the adult age. Simultaneously, we propose a hypothetical oscillation in the hormonal release whose frequency is imposed on the electroretinogram oscillation. The model consists of two coupled nonlinear oscillators in which a dynamical response is obtained mainly through an Andronov-Hopf bifurcation. Through the construction of the model, a biological hypothesis about the essential elements underlying the ERG circadian rhythm and their interrelations is formulated and discussed.
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    Bulletin of mathematical biology 55 (1993), S. 111-129 
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    Notes: Abstract Several of the known scaling laws in the animal kingdom are based on a so-called allometric correlation in which some physical quantity is presumed to scale as some power of the mass of the animal. Such a simple correlation, when deduced purely as an empirical result, often hides the physical balances that fix the relevant scaling law. In particular, the emphasis on a simple allometric scaling has often masked the fundamental role played by time scales associated with the physical balances being struck. In this paper I have concentrated on three different attributes to which the use of dimensional analysis, scaling arguments and some judicious guesswork have led to new results and an understanding of some balances that occur in the animal kingdom. The running speed of animals is examined and a rationale deduced for the resolution of a conundrum first posed by A.V. Hill of why it is that many animals appear to have approximately the same maximum speed. A complete dimensional analysis for scaling the basal metabolic rate for a class of animals suggests that a detailed understanding of the physical balances that fix the metabolic rate could be quite subtle. However, the use of such an analysis has led to the discovery of a new correlation for mammals, relating the metabolic rate to the mass and the pulse rate of the animal. At the heart of many scaling laws for animal motion is the provision of an estimate of how the skeletal structure depends on the mass of the animal. It has been known for some time that the assumption of isometry between the builds of animals is too constrictive to describe the observed scaling laws. It is shown here how to relax the isometric assumption and deduce scaling laws in good agreement with observation. Thus, it appears that the skeletal dimensions of many animals with exoskeletons are fixed by the need to support static rather than dynamical loads. The scaling laws associated with endoskeletons are more complex, apparently, though the analysis does suggest that it is dynamical loading which is decisive for the skeletal design of land mammals.
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    Bulletin of mathematical biology 55 (1993), S. 465-486 
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    Notes: Abstract This article presents a new method for the comparison of multiple macromolecular sequences. It is based on a hierarchical sequence synthesis procedure that does not require anya priori knowledge of the molecular structure of the sequences or the phylogenetic relations among the sequences. It differs from the existing methods as it has the capability of: (i) generating a statistical-structural model of the sequences through a synthesis process that detects homologous groups of the sequences, and (ii) aligning the sequences while the taxonomic tree of the sequences is being constructed in one single phase. It produces superior results when compared with some existing methods.
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    Bulletin of mathematical biology 55 (1993), S. 503-524 
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    Notes: Abstract This paper presents a dynamic programming algorithm for aligning two sequeces when the alignment is constrained to lie between two arbitrary boundary lines in the dynamic programming matrix. For affine gap penalties, the algorithm requires onlyO(F) computation time andO(M+N) space, whereF is the area of the feasible region andM andN are the sequence lengths. The result extends to concave gap penalties, with somewhat increased time and space bounds.
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    Bulletin of mathematical biology 55 (1993), S. 561-583 
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    Notes: Abstract A general model of zymogen activation is proposed and explicit kinetic equations for the time courses of the various species and products involved are given. These equations are valid for the whole course of the reaction and therefore for both the transient phase and the steady state. This model is sufficiently general to include mechanisms possessing one or more steps of zymogen activation besides possible steps of inhibition (reversible or irreversible) or inactivation.
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    Bulletin of mathematical biology 55 (1993), S. 585-608 
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    Notes: Abstract Cell migration can be characterized by two independent variables: the speed,v, and the migration angle, ϕ. Each variable can be described by a stochastic differential equation—a Langevin equation. The migration behaviour of an ensemble of cells can be predicted due to the stochastic processes involved in the signal transduction/response system of each cell. Distribution functions, correlation functions, etc. are determined by using the corresponding Fokker-Planck equation. The model assumptions are verified by experimental results. The theoretical predictions are mainly compared with the galvanotactic response of human granulocytes. The coefficient characterizing the mean effect of the signal transduction/response system of the cell is experimentally determined to 0.08 mm/V sec (galvanotaxis) or 0.7 mm/sec (chemotaxis) and the characteristic time characterizing stochastic effects in the signal transduction/response system is experimentally determined as 30 sec. The temporal directed response induced by electric field pulses is investigated: the experimental cells react slower but are more sensitive than predicted by theory.
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    Bulletin of mathematical biology 55 (1993), S. 15-35 
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    Notes: Abstract A classical predator-prey model is considered in this paper with reference to the case of periodically varying parameters. Six elementary seasonality mechanisms are identified and analysed in detail by means of a continuation technique producing complete bifurcation diagrams. The results show that each elementary mechanism can give rise to multiple attractors and that catastrophic transitions can occur when suitable parameters are slightly changed. Moreover, the two classical routes to chaos, namely, torus destruction and cascade of period doublings, are numerically detected. Since in the case of constant parameters the model cannot have multiple attractors, catastrophes and chaos, the results support the conjecture that seasons can very easily give rise to complex populations dynamics.
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    Bulletin of mathematical biology 55 (1993), S. 75-96 
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    Notes: Abstract We briefly review the literature on the division of labour in ant colonies with monomorphic worker populations, and show that there are anomalies in current theories and in the interpretation of existing data sets. Most ant colonies are likely to be in unstable situations and therefore we doubt if an age-based division of labour can be sufficiently flexible. We present data for a type of small ant colony in a highly seasonal environment, concentrating on individually marked older workers. We show that contrary to expectation such workers undertake a wide variety of tasks and can even retain their ability to reproduce, even whilst younger workers are actively foraging. Our analysis shows that old workers occupy four distinct spatial stations within the nest and that these are related to the tasks they perform. We suggest that correlations between age and task in many ant colonies might simply be based on ants foraging for work, i.e. actively seeking tasks to perform and remaining faithful to these as long as they are profitably employed. For this reason, employed older workers effectively displace unemployed younger workers into other tasks. In a companion paper, Tofts 1993,Bull. math. Biol. develops an algorithm that shows how foraging for work can be an efficient and flexible mechanism for the division of labour in social insects. The algorithm creates a correlation between age and task purely as a by-product of itsmodus operandi.
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    Bulletin of mathematical biology 55 (1993), S. 131-140 
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    Notes: Abstract A class of deformations of polyhedra which preserve both the combinatorial type and the location of all but one vertex are examined, leading to the concepts of unconditional and conditional vertex mobility. A simple criterion for vertex mobility is given, and equimobility classes of polyhedra are introduced. The polyhedral mobility characterization is suggested for applications in dynamic molecular modeling, shape analysis of protein folding, and the study of rearrangements of atomic clusters.
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    Bulletin of mathematical biology 55 (1993), S. 155-174 
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    Notes: Abstract The optimal shape of the front profile of the thick lens in the eye of the scallop,Pecten is theoretically, geometric optically investigated as a function of the refractive index of the lens and the retina, as well as of the geometrical parameters of the eye. The shape of the theoretical front surfaces is compared with that of the real, experimentally determined front face of the lens. The degree of correction of the lens for spherical aberration of the reflecting spherical mirror in thePecten eye is examined. The optimal shape of the front profile of the lens depends strongly on a set of parameters, such that a certain fine tuning is required among them to assure a full correction for spherical aberration. The extreme variability of the eye parameters and the shape of the front face of the lens in the scallop is inconsistent with this fine tuning requirement. The degree of correction of thePecten lens for spherical aberration might not be as good as it could be, a possible biooptical reason for which is discussed.
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    Bulletin of mathematical biology 55 (1993), S. 175-195 
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    Notes: Abstract A population of cells suspended in a liquid nutrient medium is considered. The process of growth, division and death of a cell is interpreted mathematically as the Bellman-Harris stochastic process governed by random meetings between the cell and nutrient particles. Growth of a cell is considered to be a result of two processes: mass inflow into and mass outflow from the cell. It is found that, in the absence of food limitations and inhibitors, population growth is not exponential. However, the exponential increase is approached asymptotically over time. Population net growth rate is a variable rather than a constant, but tends over time to a constant value which is the rate of exponential growth. The rate of exponential growth, the probabilities of cell division and death, and the life expectancy of a cell are expressed analytically via average rate of meetings between a cell and nutrient particles. The paper presents an independent phase in calculating mathematical relations between the rate of exponential growth and the concentration of food in a substrate.
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    Bulletin of mathematical biology 55 (1993), S. 213-230 
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    Notes: Abstract We analyse the helical motion of organisms, concentrating on the means by which organisms change the direction in space of the axis of the helical trajectory, which is the net direction of motion. We demonstrate that the direction of the axis is determined largely by the direction of the organism's rotational velocity. Changes in direction of the rotational velocity, with respect to the organism's body, change the direction in space of the axis of the helical trajectory. Conversely, changes in direction of the translational velocity, with respect to the body of the organism, have little effect on the direction in space of the axis of the trajectory. Because the axis of helical motion is the net direction of motion, it is likely that organisms that move in helices change direction by pointing their rotational velocity, not their translational velocity, in a new direction.
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    Bulletin of mathematical biology 55 (1993), S. 259-275 
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    Notes: Abstract The mechanistic model of the phytoplankton photosynthesis-light intensity relationship by Eilers and Peeters (1988.Ecol. Modelling 42, 199–215) is investigated mathematically. The model is based on the physiological idealization of transition probabilities between states of the photosynthetic factories,PSF. The model was found to have under constant light condition a globally stable unique positive equilibrium, while under periodically varying light (e.g. daily periodicity) there exists a unique globally asymptotically stable periodic solution. Based on this, the adaptation to a change of light intensity is defined as a process by which the state ofPSF converges to an equilibrium. Assuming that phytoplankton regulates its photosynthetic production rate with a certain strategy which maximizes production, two such possible strategies were examined. Both the instantaneous and the integral maximal photosynthetic production were shown to have the same result. With realistic qualitative assumptions of the shape of the dependence of the four model parameters on the light intensity to which phytoplankton is adapted, the numerical values of parameters under both constant and periodically varying conditions are determined by applying Pontryagin's maximum principle.
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    Bulletin of mathematical biology 55 (1993), S. 295-313 
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    Notes: Abstract Although the isobologram is presently the most widely used method of analysis for combined effects of agents, there are several different interpretations of the linear isobole isobole in regard to its use as a criterion of interaction. An investigation of the differential aspects of the linear isobole relation may cast some light in this regard. By conceptual extension of the present single effect level (i.e. effect-point) relation of the linear isobole to an effect-neighbourhood relation in which the linear isobole holds over a small continuous range of effect levels, the mathematical differential of the linear isobole can be developed and investigated. This differential aspect provides some useful insights into the implication and interpretation of the linear isobole relation when used as a general criterion in agent interaction studies. it can also serve as the mathematical basis for the formulation of analytic schemes in which the linear isobole relation is applicable over a continuous range of effect levels.
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    Bulletin of mathematical biology 55 (1993), S. 315-344 
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    Notes: Abstract We discuss in detail the behaviour of a model, proposed by Goldbeteret al. (1990.Proc. natn. Acad. Sci. 87, 1461–1465), for intracellular calcium wave propagation by calcium-induced calcium release, focusing our attention on excitability and the propagation of waves in one spatial dimension. The model with no diffusion behaves like a generic excitable system, and threshold behaviour, excitability and oscillations can be understood within this general framework. However, when diffusion is included, the model no longer behaves like a generic excitable system; the fast and slow variables are not distinct and previous results on excitable systems do not necessarily apply. We consider a piecewise linear simplification of the model, and construct travelling pulse and periodic plane wave solutions to the simplified model. The analogous behaviour in the full model is studied numerically. Goldbeter's model for calciuminduced calcium release is an excitable system of a type not previously studied in detail.
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    Bulletin of mathematical biology 55 (1993), S. 385-415 
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    Notes: Abstract A generalized model ofn catalytically-coupled self-replicative molecules witherror-prone replication is presented. A generalized mathematical formulation of this model and the outline of its asymptotic behaviour have been developed. Due to the complexity of the model, only in simple situations is it possible to draw general conclusions from the standard analysis. Some complex situations are illustrated by means of numerical integration of particular examples.
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    Bulletin of mathematical biology 55 (1993), S. 487-489 
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