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  • Springer  (79,854)
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  • 1995-1999
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  • 1955-1959  (15,182)
  • 1993  (69,342)
  • 1959  (15,182)
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  • 1995-1999
  • 1990-1994  (69,342)
  • 1965-1969
  • 1955-1959  (15,182)
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  • 1
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    Bulletin of mathematical biology 21 (1959), S. 1-11 
    ISSN: 1522-9602
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Mathematics
    Notes: Abstract By means of the Laplace transform, the behavior of a simplified model of the cardiovascular system is mathematically formulated. This formulation allows mathematical expression of the periodicity of the cardiac output and the systemic response. With the cardiac output represented as half of a sine function cycle, the systolic aortic pressure becomes the sum of a sine term and exponential terms, while the sum of the exponential terms alone represents the diastolic pressure. The characteristics of the mathematical expressions for systole and diastole are analyzed, and some relationships of potentially practical value are derived. Variation in the parameters of the system yields mathematical results consistent with the expected physical ones.
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  • 2
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    Bulletin of mathematical biology 21 (1959), S. 19-32 
    ISSN: 1522-9602
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    Notes: Abstract A generalization of Landahl's approximation method (H. D. Landahl,Bull. Math. Biophysics,15, 49–61, 1953) for non-linear diffusion problems is suggested. The method is applied to sorption, desorption, and free diffusion problems involving concentration-dependent diffusion coefficients. With some limitations, the results compare favorably with those obtained by numerical methods.
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  • 3
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    Bulletin of mathematical biology 21 (1959), S. 33-60 
    ISSN: 1522-9602
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    Notes: Abstract Recently a theorem for representing current generators in a volume conductor by the superposition of a central dipole, quadrupole, octopole, etc., has been established by G. C. K. Yeh, J. Martinek, and H. de Beaumont (Bull. Math. Biophysics,20, 203–16, 1958). This theorem makes possible the representation of any discrete or line, surface- or volume-distributed current source by a unique model which can be determined for each given case by surface potential measurements and closed form analysis. In this paper the multipole representations of an eccentric dipole and an eccentric double-layer are obtained in terms of the various parameters of the assumed singularities, and the contributions to surface potentials due to each of the multipoles are compared. Certain numerical results corresponding to those of E. Frank (Amer. Heart J.,46, 364–78, 1953) are carried out and compared. Furthermore, the multipole representation of a partially damaged double-layer is also determined and compared with that of an undamaged one. It is concluded that within the range of parameters corresponding to human subjects the higher-order multipoles can contribute significantly to the surface potentials compared with the dipole.
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  • 4
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    Bulletin of mathematical biology 21 (1959), S. 97-100 
    ISSN: 1522-9602
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    Notes: Abstract In line with a recent suggestion by the author (Bull. Math. Biophysics,20, 267–73, September, 1958) that not only does the organism as a whole map on the primordial, but that each organ can also be thus mapped, it is shown that the previously introduced abstract spaces, which represent an organism, contain subspaces which map continuously on the space of the primordial. Several theorems about those subspaces are proven.
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  • 5
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    Bulletin of mathematical biology 21 (1959), S. 71-95 
    ISSN: 1522-9602
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    Notes: Abstract The DNA-protein coding problem is given a general algebraic formulation, the consequences of which are then explored by standard mathematical methods. To keep the treatment self-contained, the mathematical techniques to be used are explained in detail. It is demonstrated that there exista priori a countably infinite number of different abstract DNA-protein codes, thereby showing that inductive attempts to construct such a code will most likely be fruitless. A notion of ergodicity is then introduced, which imposes a number of restrictions on the admissible codes, and, in fact, these considerations enable us toderive a small portion of a code which, if our hypothesis of ergodicity is correct, must occur in nature. Finally, we discuss briefly the problem as to whether there can exist more than one DNA-protein code in nature.
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  • 6
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    Notes: Abstract The present-day practices of electrocardiography and vectorardiography are based upon the theory that the surface potential differences can be assumed to be due to a single dipole inside the body. It is shown in this paper that a dipole cannot account for all the surface potentials due to realistic current generators, and hence the determination of the current generator from surface potential measurements based upon such a theory will lead to inconsistent representations of the heart for one and the same subject. To demonstrate this point two eccentric dipoles of different strengths and locations representing two muscle fibers are taken to be the current generator in a homogeneous spherical conductor. The exact surface potentials are then expressed by means of the “interior sphere theorem” of the authors. With these expressions the magnitude, direction, and location of the resultant dipole are determined by the method of D. Gabor and C. V. Nelson (J. App. Physics,25, 413–16, 1954). The surface potentials due to this resultant dipole are again exactly expressed by means of the “interior sphere theorem” and compared with those due to the eccentric dipoles assumed. It can be seen that the differences can be considerable. It is suggested that the multipole model of the authors (Bull. Math. Biophysics,20, 203–16, 1958) be used as a more accurate and the only unique representation of the heart.
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  • 7
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    Bulletin of mathematical biology 21 (1959), S. 101-106 
    ISSN: 1522-9602
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    Notes: Abstract In a preceding paper (Bull. Math. Biophysics 20, 71–93, 1958) the principle of biotopological mapping was formulated in terms of a continuous mapping of an abstract space, made from the set of biological properties which characterize the organism, by an appropriate definition of neighborhoods. In this paper it is shown that we may consider directly the mappings of the different sets of properties which characterize different organisms without taking recourse to abstract spaces. All the verificable conclusions made in the preceding paper remain valid. A serious difficulty mentioned previously is, however, avoided and the possibility of more general predictions is established.
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  • 8
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    Bulletin of mathematical biology 21 (1959), S. 107-107 
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  • 9
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    Bulletin of mathematical biology 21 (1959), S. 109-128 
    ISSN: 1522-9602
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    Notes: Abstract The general Theory of Categories is applied to the study of the (M, R)-systems previously defined. A set of axioms is provided which characterize “abstract (M, R)-systems”, defined in terms of the Theory of Categories. It is shown that the replication of the repair components of these systems may be accounted for in a natural way within this framework, thereby obviating the need for anad hoc postulation of a replication mechanism. A time-lag structure is introduced into these abstract (M, R)-systems. In order to apply this structure to a discussion of the “morphology” of these systems, it is necessary to make certain assumptions which relate the morphology to the time lags. By so doing, a system of abstract biology is in effect constructed. In particular, a formulation of a general Principle of Optimal Design is proposed for these systems. It is shown under what conditions the repair mechanism of the system will be localized into a spherical region, suggestive of the nuclear arrangements in cells. The possibility of placing an abstract (M, R)-system into optimal form in more than one way is then investigated, and a necessary and sufficient condition for this occurrence is obtained. Some further implications of the above assumptions are then discussed.
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  • 10
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    Bulletin of mathematical biology 21 (1959), S. 141-151 
    ISSN: 1522-9602
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    Notes: Abstract The transient stage of the random dispersal of logistic populations is investigated, using a Sturm-Liouville series leading to an infinite system of non-linear integral equations. These equations are then solved via a successive approximation scheme. R. A. Fisher's (steady-state) velocity of advance paradox is discussed. An illustrative example is worked to the second order of approximation.
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    Bulletin of mathematical biology 21 (1959), S. 153-159 
    ISSN: 1522-9602
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    Notes: Abstract An approximation method using a sine function is used to solve the second degree growth equation for the case in which an organism may simultaneously become dispersed throughout a uniform region. The resulting expression for a special case is compared with the expression obtained by R. Barakat (1959,Bull. Math. Biophysics,21, 141–51), giving the first two terms, by an iterative, procedure. The agreement is satisfactory.
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  • 12
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    Bulletin of mathematical biology 21 (1959), S. 129-140 
    ISSN: 1522-9602
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    Notes: Abstract Diffusion through a flat pore into a large open region is proportional to the linear dimension of the pore and not to its area. This was first explained by Brown and Escombe (1900) for a circular pore and is here generalized, by means of a dimensional argument, to include any type of regular opening. The problem is further generalized to include diffusion through pores of finite thickness, finite distance apart, and into finite regions. Since this problem cannot be solved exactly, an approximation method is introduced. Reasons for the credibility of the approximation are presented. It is then shown, by means of the approximation method, that the diffusive flow through a pore is equal to the total concentration difference divided by the resistance of the system. The resistance, in turn, is the sum of the resistances of all portions of the system, each of which is calculated. The result is compared with results which have been calculated exactly for limiting cases and found to agree very well. The results are then applied to a standard method of computing pore size in membranes, and it is shown that the correction factor is negligible.
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  • 13
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    Bulletin of mathematical biology 21 (1959), S. 161-183 
    ISSN: 1522-9602
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    Topics: Biology , Mathematics
    Notes: Abstract V. S. Ivlev [Experimental Ecology of Nutrition of Fishes, 1955, Moscow (in Russia)] has shown that the food uptake by fishes during a fixed interval of time is an exponential function of the concentration of food. Ivlev's equation is derived here, and it is shown that it can hold only for non-stationary conditions, such as prevailed in Ivlev's experiments. For a stationary state, the rate of food uptake should tend asymptotically to a limiting value as the concentration increases, but the variation is not exponential. Different other aspects of the problem are investigated, and definite new experimental procedures suggested. The implications of Ivlev's findings on the effect of non-uniformity of food distribution upon the rate of food consumption are studied from a mathematical point of view. The conclusion is reached that whereas a fish does not, in the process of eating, move directly to an individual food particle which it perceives, it does move more or less directly to large aggregates of particles, if the latter are distributed nonuniformly.
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  • 14
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    Bulletin of mathematical biology 21 (1959), S. 185-193 
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    Notes: Abstract Some relational aspects of the property of self-reproduction of biological systems are studied. If in addition to the requirement of the property of self-reproduction we add also the requirement of adaptability of the organism to changing environment, this imposes certain conditions on the topology of the graphs which represent such systems. A further study of the relational properties of such systems seems to offer the possibility of deriving the principle of biological mapping from the requirement of self-reproduction and adaptability. An examination of the problem of the original formation of such self-reproducing systems in connection with the established fact of impossibility of spontaneous generation leads to the conclusion that an organism must inhibit such processes which, in the absence of organisms, would lead to spontaneous generation.
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    Bulletin of mathematical biology 21 (1959), S. 195-216 
    ISSN: 1522-9602
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    Notes: Abstract In the human, the antagonistic, extensor-flexor system of the leg is an example of a common type of neurophysiological feedback system. After a brief introduction to the neuroanatomy and physiology of this feedback system, the paper formulates transfer functions from temporal response data available in the literature. A feedback stability analysis, based on the extension of Nyquist's stability criteria to multiple-loop systems and utilizing flow-graph techniques, demonstrates the stable behavior of the system. Expressions are given relating the sensitivity of the system to variations in muscle response and Golgi tendon organ (tension receptor) response. By considering the events taking place at synapses and end-plates during “isometric tension-small knee angle excursion” conditions as stationary stochastic processes, an external “noise” input to the system is given, whose spectrum is derived from the statistics of a shot-process representation of these events. The paper concludes with some correlations between the analytical results and clinical syndromes.
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    Bulletin of mathematical biology 21 (1959), S. 217-255 
    ISSN: 1522-9602
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    Notes: Abstract In this continuation of a previous report it is shown how the Volterra population dynamics, which underlies the statistical theory, can be based on a variational principle; how the dynamics can be generalized as regards both the behavior of total populations and migration phenomena; and how many directly observable data, such as amplitudes and frequencies of oscillation of a population, fit into the statistical theory and can test it. Such a test is carried out in some detail using the fox-catch data of Elton, with a clear indication that the theory is capable of comprehending the major statistical properties of population-time curves. A final section sketches an extension of the theory to cover secular variations of external conditions such as temperature of the environment.
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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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  • 18
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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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    Circuits, systems and signal processing 12 (1993), S. 153-154 
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    Notes: Abstract Once designed, implementation of an optimal mean-square binary morphological filter is extremely fast, especially when the erosions are implemented on a suitable parallel processor. On the other hand, optimal filter design involes a computationally burdensome search procedure that can, in practice, be intractable. The present paper provides an algorithm for filter design that is based on the relationship between the optimal morphological filter and the conditional expectation. The algorithm proceeds by changing the conditional expectation into a morphological filter while at the same time increasing the mean-square error by a minimal amount. It does so by switching observations between the 1-set and the 0-set of the conditional expectation. The switching algorithm is extremely efficient in many noise environments, and therefore provides a filter design that can be useful for online structuring-element updating. Owing to the relationship between stack and morphological filters, the algorithm is at once useful for finding optimal binary stack filters.
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    Circuits, systems and signal processing 12 (1993), S. 263-278 
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    Notes: Abstract New characteristics of feedback neural networks are studied. We discuss in detail the question of updating of neurons given incomplete information about the state of the neural network. We show how the mechanism of self-indexing for such updating provides better results than assigning ‘don't know’ values to the missing parts of the state vector. Issues related to the choice of the neural model for a feedback network are also considered. Properties of a new complex valued neuron model that generalizes McCulloch-Pitts neurons are examined.
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    Circuits, systems and signal processing 12 (1993), S. 391-407 
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    Notes: Abstract This paper deals with chained eigenstructure assignment for strongly controllable singular systems of the form Êx (t)=Â x(t)+B u(t) with state feedback control of the formu(t)=Kx(t)+w(t). The development of our method depends crucially on the properties of standard form singular systems. The closed-loop system will satisfy the following requirements: regularity, impulse-free response and rankÊ arbitrary eigenvalues assignment. This parametric characterization conveniently organizes the nonunique gain matrixK to modify the dynamic response of the systems. The result can be used for discrete-time descriptor systems, in which a zero-value eigenvalue may well be a desired closed-loop eigenvalue. One illustrative example is included.
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    Circuits, systems and signal processing 12 (1993), S. 453-464 
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    Notes: Abstract An efficient bi-state stochastic gradient is proposed for spontaneous constrained time delay estimation. The quantized stochastic gradient is an approximation of the polarity of the instantaneous delay estimation error. It is adjusted in such a way that it has a much higher probability to move in the correct direction at each iteration so as to enable a speed-up in the delay estimate to converge to global minimum in steady state. The performance of the delay estimator is evaluated statistically and an analytical solution for its convergence behavior is established. It is demonstrated that the proposed algorithm has at least a two-fold improvement in convergence speed when compared with the conventional approach, and this is verified by extensive simulation results.
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    Circuits, systems and signal processing 12 (1993), S. 503-531 
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    Notes: Abstract In this work, we extend the coding theory approach to error control in redundant residue number systems (RRNS). The concept of erasure correction capability in RRNS is introduced. We derive the relationship between the minimum distance and the error detection and error/erasure correction capability. New computationally efficient algorithms are derived for simultaneously correcting single errors and multiple erasures and detecting multiple errors. These algorithms reduce the computational complexity of the previously known algorithms by at least an order of magnitude. Another attractive feature of the algorithms is that all the arithmetic operations are modulo operations. Consequently, the need to process large valued integers is avoided.
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    Circuits, systems and signal processing 12 (1993), S. 579-587 
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    Notes: Abstract This paper presents a general expression relating the complex-normalized scattering matrix of ann-port network to that of its augmentedn-port network normalizing to then 1 −Ω resistances, where the Darlington equivalent network may be either reciprocal or nonreciprocal.
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    Circuits, systems and signal processing 12 (1993), S. 211-221 
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    Notes: Abstract Recent research has shown that multilayer feedforward networks with sigmoidal activation functions are universal approximators, and that this holds for more general activations as well. The mathematical underpinning for these results has been various: Kolmogorov's resolution of Hilbert's thirteenth problem; the Stone-Weierstrass theorem; approximation of Fourier and Radon integral representations; and convergence of probability measures. This paper • Rigorously establishes the robustness of feedforward network realizations. • Uses a theorem of Wiener and ideas of translation invariant subspaces to provide conditions for universal approximations toL 1 andL 2 functions by networks, for quite general activation functions. The second result extends and simplifies some of the recent results of Stinchcombe and White, at least for the special cases ofL 1 andL 2 functions.
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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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    Circuits, systems and signal processing 12 (1993), S. 489-492 
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    Notes: Abstract In this short note, we establish a simple, yet precise, necessary and sufficient condition for the “right coprime factorization” of a nonlinear feedback control system. As a consequence, we also obtain similar conditions for the “stable right coprime factorizations ” of the nonlinear feedback control system.
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    Circuits, systems and signal processing 12 (1993), S. 557-566 
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    Notes: Abstract The pseudorandom sequence of arrays (PRSA) and a method to generate it was reported earlier by the authors. This paper presents another method to generate a PRSA. The mathematical recursion describing the PRSA and some of its properties are discussed.
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    Circuits, systems and signal processing 12 (1993), S. 37-49 
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    Notes: Abstract This paper proposes an effective method to improve the digital redesign method via the block-pulse function approach. The coefficients of the block-pulse function expansion are exactly evaluated such that the desired digitally redesigned feedback gain and forward gain will be obtained. A numerical example is given to demonstrate the effectiveness of the proposed method.
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    Circuits, systems and signal processing 12 (1993), S. 51-60 
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    Notes: Abstract In this paper we apply the technique of interval analysis to get bounds on the initial value response of a linearized single machine infinite bus problem when a parameter is varied. It is generally believed that responses for parameter variations in an interval should lie within the responses for the extremums of the parameter variations. This is not generally true and our example demonstrates this. The interval-analysis technique permits getting the overall bound on the response. Further experimentation also revealed that the method has some limitations particularly involving lightly damped long-term dynamics. The technique is useful in finding the robustness of a particular design such as the power system stabilizer for parameter variations.
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    Circuits, systems and signal processing 12 (1993), S. 105-117 
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    Notes: Abstract The high-order Yule-Walker (HOYW) method of sinusoidal frequency estimation based on a singular value decomposition (SVD) is known to have excellent statistical performance. Here, we show that the SVD-based step of the HOYW method can be replaced by a computationally more convenient QR decomposition (QRD)-based step, without affecting the asymptotic properties of the frequency estimates.
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    Circuits, systems and signal processing 12 (1993), S. 85-103 
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    Notes: Abstract For the given observations set of the ARMA (autoregressive moving average) process, the likelihood function depends, not only on model parameters, but on the starting values of the input and output. Therefore, it is called theconditional likelihood function. Theunconditional likelihood function can be obtained in two ways. The first is to set the starting values to zero, as is often done, and the second is to set them to the properly estimated values. The difference between these two types of likelihood functions is significant when the given data sequence is short, and any of the zeros of the moving average part is close to the boundary of the unit circle. In this paper the direct method of starting value estimation and its application to two off-line ARMA estimation algorithms, the maximum likelihood (ML) algorithm and the iterative inverse filtering (ITIF) algorithm, is proposed. Experimental results prove both increased efficiency and stability of these algorithms. The importance of setting the starting values properly is also significant when the recursive algorithm, with previously estimated parameters, has to be restarted. The advantage of the proposed reinitialization method is shown on the recursive lattice algorithm working in the block mode.
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    Circuits, systems and signal processing 12 (1993), S. 151-151 
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    Circuits, systems and signal processing 12 (1993), S. 133-149 
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    Notes: Abstract In this paper we consider nonlinear, infinite networks of purely resistive type where the voltage across a branch of the network is proportional to a fixed power of the current flowing in the branch. It is known that the study of currents in such networks amounts to studying the space of the functions on a network which have finite Dirichlet sums of orderp. Such a study was carried out in [7], [9], and [11]–[14] under the assumption that every node is connected to only finitely many different nodes of the network. In this paper we drop this assumption and work with general countable networks. We prove that most results of the locally finite case, and especially the classification theory, hold true in a more general context. Moreover, we give necessary and sufficient conditions for a network to have only constant Dirichlet finitep-harmonic functions. The relationship with discrete Markov processes is also pointed out.
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    Circuits, systems and signal processing 12 (1993), S. 155-175 
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    Notes: Abstract One of the major drawbacks of the backpropagation algorithm is its slow rate of convergence. Researchers have tried several different approaches to speed up the convergence of backpropagation learning. In this paper, we present those rapid learning methods as three categories, and implement the representative methods of each category: (1) for the numerical method based approach, the Aitken's Δ2 process, (2) for the heuristics based approach, the dynamic adaptation of learning rate, and (3) for the learning strategy based approach, the selective presentation of learning samples. Based on these implementations, the performance is evaluated with experiments and the merits and demerits are briefly discussed.
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    Circuits, systems and signal processing 12 (1993), S. 177-210 
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    Notes: Abstract The area of artificial neural networks has recently seen an explosion of theoretical and practical results. In this paper, we present an artificial neural network that is algebraically distinct from the classical artificial neural networks, and several applications which are different from the typical ones. In fact, this new class of networks, calledmorphology neural networks, is a special case of a general theory of artificial neural nets, which includes the classical neural nets. The main difference between a classical neural net and a morphology neural net lies in the way each node algebraically combines the numerical information. Each node in a classical neural net combines information by multiplying output values and corresponding weights and summing, while in a morphology neural net, the combining operation consists of adding values and corresponding weights, and taking the maximum value. We lay a theoretical foundation for morphology neural nets, describe their roots, and give several applications in image processing. In addition, theoretical results on the convergence issues for two networks are presented.
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    Circuits, systems and signal processing 12 (1993), S. 309-329 
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    Topics: Electrical Engineering, Measurement and Control Technology
    Notes: Abstract Higher order moment computations are generally necessary wherever the recognition task exceeds the confines of linearity. This paper provides an overview of recent progress on a specific neural network design, which explicitly uses higher order moment information. Attention is focused on the training algorithms used in the design and on network performance in prototype applications.
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    Circuits, systems and signal processing 12 (1993), S. 375-390 
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    Notes: Abstract In the present paper we study the problem of the existence and uniqueness of solutions of implicit systems considered on a finite interval of time. We consider two kinds of existence problems: input-acceptance and input-acceptance when the boundary conditions of a corresponding trajectory are set to zero, and two kinds of uniqueness problems: output uniqueness and output uniqueness when the boundary conditions of the corresponding trajectory are unknown. Geometric conditions for all of these notions are given, and the duality of these notions is studied.
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    The Geneva risk and insurance review 18 (1993), S. 103-105 
    ISSN: 1554-9658
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    Topics: Economics
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    Bulletin of mathematical biology 21 (1959), S. 13-17 
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    Notes: Abstract This paper extends Leslie's vector and matrix treatment of populations. A simple matrix is given for species in which adult mortality and fertility are independent of age, but in which the juvenile mortality rate differs from the adult. The population vector can be changed into a population matrix. This should allow equations using functions of the size of the population to be developed. Genetic variables such as sex or other polymorphisms can be introduced, and the notation allows different systems of selection or non-random mating to be specified.
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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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    Bulletin of mathematical biology 55 (1993), S. 491-502 
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    Bulletin of mathematical biology 55 (1993), S. 543-560 
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    Notes: Abstract We investigated the structural correlates of stability and resilience in strong hierarchies, that is, systems that can be represented by a rooted tree. A simple exponential model that incorporates three variables (the total number of nodesN; the number of basal nodesn B; and the number of single links among nodesN 1) accounts for 95% of the observed variability in stability among trees in our sample population. For resilience the situation is even simpler, with about 89% of the population variation being accounted for by tree size (N). For strong hierarchies, size and shape are the principal correlates of stability, while size alone explains the major proportion of the variability in resilience among stable trees. These results suggest that reasonably accurate statistical predictions about the stability and resilience of strong hierarchies can be made from a small set of (relatively) easily measured variables, without detailed knowledge of system topology.
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    Bulletin of mathematical biology 55 (1993), S. 525-541 
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    Notes: Abstract Numerous regulatory mechanisms in motor control involve the presence of time delays in the controlled behavior of the system. Experimentally, we have shown that an increase of the time delay in visual feedback induces different oscillations in control subjects and in patients with neurological diseases during the performance of a simple compensatory tracking task. A preliminary model is proposed to describe the oscillations observed in control subjects and in patients with neurological diseases. The influence of delays in two feedback loops are the main components of the motor control circuitry involved in this task and are studied from an analytical and physiological perspective. We analytically determine the influence in the model of each of these delays on the stability of the finger position. In addition, the influence of stochastic elements (“noise”) in the modeling equation is seen to contribute qualitatively to a more accurate reproduction of experimental traces in patients with Parkinson's disease but not in patients with cerebellar disease.
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    Bulletin of mathematical biology 55 (1993), S. 609-635 
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    Notes: Abstract A model of embryo energetics was fitted to data from the literature for species as different as snails and mammals. The model is based on assumptions about energy uptake, storage and utilization. It describes the animal by two state variables: volume and energy storage. Embryo weight is taken to be proportional to volume, yolk weight to energy storage, and respiration rate to storage utilization rate. The fits were good, with minor deviations occurring only in the early phases of development. For altricial birds, good model fits were obtained, but the parameter values markedly differed from those of other species. We hypothesized that, due to an increase in energy utilization towards hatching, the temperature of the embryo increases. As a result, metabolic processes are accelerated. When this was taken into account, parameter values were obtained that correspond better with those of other animals.
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    Bulletin of mathematical biology 55 (1993), S. 637-654 
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    Notes: Abstract Experimental evidence suggests that anisotropic stress induces alignment of intracellular actin filaments. We develop a model for this phenomenon, which includes a parameter reflecting the sensitivity of the microfilament network to changes in the stress field. When applied to a uniform cell sheet at rest, the model predicts that for sufficiently large values of the sensitivity parameter, all the actin filaments will spontaneously align in a single direction. Stress alignment can also be caused by a change in external conditions, and as an example of this we apply our model to the initial response of embryonic epidermis to wounding. Our solutions in this case are able to reflect the actin cable that has been found at the wound edge in recent experiments; the cable consists of microfilaments aligned with stress at the wound boundary of the epithelium. These applications suggest that stress-induced alignment of actin filaments could play a key role in some biological systems. This is the first attempt to include the alignment phenomenon in a mechanical model of cytogel.
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    Bulletin of mathematical biology 55 (1993), S. 715-730 
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    Notes: Abstract The problem of optimal dosage is studied for the administration of ddCyd using erythrocytes as carriers and bioreactors. The volume of erythrocytes and the initial amount of drug to be loaded have to be determined in such a way that the duration of the therapeutic effect is maximized without exceeding the toxic threshold. It is found that the optimal control is unique and it is at the upper vertex of the set of the admissible controls. A more general case is also briefly discussed.
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    Notes: Abstract To explore the biological similarities and differences between the HIV-1 and HIV-2 viruses, we model the probability of male-to-female transmission of either HIV virus as a function of the number of sexual partners, the prevalence of the viruses and the infectivity per contact. Using maximum likelihood estimation theory and data from a prospective study of registered female prostitutes in Dakar, Senegal, we estimate and compare the infectivities of HIV-1 and HIV-2. Graphical goodness-of-fit methods are used to show that our model fits the data well. We find that in male-to-female transmission HIV-1 is significantly more infectious than HIV-2. This findings is consistent with other data from laboratory and epidemiologic studies comparing the biology of HIV-1 and HIV-2.
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    Bulletin of mathematical biology 55 (1993), S. 817-827 
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    Notes: Abstract Predator-prey models where one or more terms involve ratios of the predator and prey populations may not be valid mathematically unless it can be shown that solutions with positive initial conditions never get arbitrarily close to the axis in question, i.e. that persistence holds. By means of a transformation of variables, criteria for persistence are derived for two classes of such models, thereby leading to their validity. Although local extinction certainly is a common occurrence in nature, it cannot be modeled by systems which are ratio-dependent near the axes.
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    Bulletin of mathematical biology 55 (1993), S. 847-864 
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    Notes: Abstract The apple twig borer (Amphicerus bicaudatus) is an insect pest of the grape vine, causing considerable damage to the grape vine in early spring. A simple difference equation model is formulated and analysed for this plant-herbivore system based on two control strategies, cane removal and pesticide application. The system has two equilibria, one where the pest is present and one where the pest is absent. Regions are found in parameter space for global stability of the equilibria and in the absence of global stability it is shown that there exist periodic or quasiperiodic solutions.
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    Bulletin of mathematical biology 55 (1993), S. 829-845 
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    Notes: Abstract A relative phase model of four coupled oscillators is used to interpret experiments on the coordination between rhythmically moving human limbs. The pairwise coupling functions in the model are motivated by experiments on two-limb coordination. Stable patterns of coordination between the limbs are represented by fixed points in relative phase coordinates. Four invariant circles exist in the model, each containing two patterns of coordination seen experimentally. The direction of switches between two four-limb patterns on the same circle can be understood in terms of two-limb coordination. Transitions between patterns in the human four-limb system are theoretically interpreted as bifurcations in a nonlinear dynamical system.
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    Bulletin of mathematical biology 55 (1993), S. 1063-1090 
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    Notes: Abstract A population dynamics approach based on a system of differential equations allows us to establish conditions for the emergence of mutualism for cases such as coelenterates-algae symbionts. A central assumption of the model is that a host organism is able to discriminate, via some molecular recognition mechanisms, among different invading organisms and preferentially rejectparasites rather thanbona fide symbionts. Large differential rejection rates allow the emergence of mutualism. Different attractors of the population dynamics correspond to the emergence of mutualism, predominance of “selfish” species, or coexistence of many species.
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    Bulletin of mathematical biology 55 (1993), S. 1091-1131 
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    Topics: Biology , Mathematics
    Notes: Abstract A Cayley tree model of idiotypic networks that includes both B cell and antibody dynamics is formulated and analysed. As in models with B cells only, localized states exist in the network with limited numbers of activated clones surrounded by virgin or near-virgin clones. The existence and stability of these localized network states are explored as a function of model parameters. As in previous models that have included antibody, the stability of immune and tolerant localized states are shown to depend on the ratio of antibody to B cell lifetimes as well as the rate of antibody complex removal. As model parameters are varied, localized steady-states can break down via two routes: dynamically, into chaotic attractors, or structurally into percolation attractors. For a given set of parameters percolation and chaotic attractors can coexist with localized attractors, and thus there do not exist clear cut boundaries in parameter space that separate regions of localized attractors from regions of percolation and chaotic attractors. Stable limit cycles, which are frequent in the two-clone antibody B cell (AB) model, are only observed in highly connected networks. Also found in highly connected networks are localized chaotic attractors. As in experiments by Lundkvistet al. (1989.Proc. natn. Acad. Sci. U.S.A. 86, 5074–5078), injection ofAb 1 antibodies into a system operating in the chaotic regime can cause a cessation of fluctuations ofAb 1 andAb 2 antibodies, a phenomenon already observed in the two-clone AB model. Interestingly, chaotic fluctuations continue at higher levels of the tree, a phenomenon observed by Lundkvistet al. but not accounted for previously.
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  • 98
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    Bulletin of mathematical biology 55 (1993), S. 1183-1198 
    ISSN: 1522-9602
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Mathematics
    Notes: Abstract The protein folding problem and the notion of NP-completeness and NP-hardness are discussed. A lattice model is suggested to capture the essece of protein folding. For this model we present a proof that finding the lowest free energy conformation belongs to the class of NP-hard problems. The implications of the proof are discussed and we suggest that the natural folding process cannot be considered as a search for the global free energy minimum. However, we suggest an explanation as to why, for many proteins, the native functional conformation maycoincide with the lowest free energy conformation.
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  • 99
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    Circuits, systems and signal processing 12 (1993), S. 3-35 
    ISSN: 1531-5878
    Source: Springer Online Journal Archives 1860-2000
    Topics: Electrical Engineering, Measurement and Control Technology
    Notes: Abstract In this paper we study the performance of direction-finding systems which attempt to estimate unknown array parameters (such as sensor gains and phases) in addition to the estimation of the directions of arrival (DOA), as a way of performing array self-calibration. We develop compact closed form expressions for the Cramér-Rao bound associated with the joint estimation of DOAs, gains, phases, and the signal covariance matrix. By evaluating the Cramér-Rao bound for selected cases we gain some insight into the performance of direction-finding systems in the presence of gain and phase uncertainties.
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    Circuits, systems and signal processing 12 (1993), S. 61-83 
    ISSN: 1531-5878
    Source: Springer Online Journal Archives 1860-2000
    Topics: Electrical Engineering, Measurement and Control Technology
    Notes: Abstract We consider input-output systems (not necessarily of feedback type) on the time domain [0, ∞) which are governed by nonlinear vector integral or differential equations that relate the input and the output. Assuming that these equations depend on a parameterA, describing perturbations, which is allowed to vary in a vicinity of a nominal valueA 0 in a linear space, we study how strongly the output is affected by changes of (a)A 0 when the input is fixed (insensitivity), and (b) the input whenA is fixed withA A 0 being not too large (robust stability). The results are based on the theory of input-output systems over abstract extended spaces given in Parts I and II of [3]. We discuss nominal systems described by nonlinear Volterra and differential equations, and consider two types of possible perturbations. We also prove a simple result on systems governed by singular differential equations whose perturbations can change the order of the system.
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