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  • 1
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    Bulletin of mathematical biology 15 (1953), S. 311-338 
    ISSN: 1522-9602
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Mathematics
    Notes: Abstract The partial differential equation of the random walk problem with persistence of direction and external bias is derived. By persistence of direction or internal bias we mean that the probability a particle will travel in a given direction need not be the same for all directions, but depends solely upon the particle's previous direction of motion. The external bias arises from an anisotropy of the medium or an external force on the particle. The problem is treated by considering that the net displacement of a particle arises from two factors, namely, that neither the probability of the particle traveling in any direction after turning nor the distance the particle travels in a given direction need be the same for all directions. A modified Fokker-Planck equation is first obtained using the assumptions that the particles have a distribution of travel times and speeds and that the average time of travel between turns need not be zero. The fional equation incopporating the assumption of a persistence of direction and an external bias is then derived. Applications to the study of diffusion and to long-chain polymers are then made.
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  • 2
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    Bulletin of mathematical biology 15 (1953), S. 383-383 
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  • 3
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    Bulletin of mathematical biology 15 (1953), S. 385-385 
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  • 4
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    Bulletin of mathematical biology 15 (1953), S. 367-381 
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    Notes: Abstract The transmission of some information or behavior pattern is treated as a flow of “particles” which execute random motions over a population of individuals and which may multiply or disappear. Equations are derived for the number density of these “particles” and from this is calculated the number of individuals through which the “particles” have passed. The results are applied to a number of situations such as 1) uniform spatial distribution with multiplication factor decreasing with time because of loss of interest or confusion of the information, 2) multiplication factor constant but the rate of spreal decreasing with multiple hearings, 3) one-dimensional region with a small starting region with or without an absorbing barrier 4) two-dimensional region with absorbing barrier, 5) continous sources of information within a small region in one dimension, 6) uniform spatial distribution in which individuals do not respond to more than one hearing.
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  • 5
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    Bulletin of mathematical biology 15 (1953), S. 387-394 
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    Notes: Abstract A situation is considered in which a fluid containing a substance flows through a vessel at a constant rate, the substance being permeable to the vessel wall. In the region outside the vessel there is supposed to be rapid mixing in the direction perpendicular to the axis of the vessel but no mixing longitudinally. The solution for the spatial distribution at any time is given for the case of an arbitrary initial distribution along the vessel length in the absence of an input. The solution is also given for the case of a single impulsive input, the concentration being initially zero everywhere.
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    Bulletin of mathematical biology 15 (1953), S. 431-476 
    ISSN: 1522-9602
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    Notes: Abstract Organisms orient themselves to a stimulus by two general methods. One method is by directed orientation (taxis); the other is by undirected locomotory reaction (kinesis). An equation, and the methods for finding the necessary parameters of this equation, is derived for the distribution of organisms within a container, with the following limitations: (1) the organisms have no accommodation, (2) they are always active, and (3) the stimulus changes slowly with position. Necessary modifications of the equation are then derived, so that the last two limitations may be eliminated. The equation cannot be solved excatly because of its complexity; hence an approximation method must be used. This method is discussed, an approximate solution is found, and a time constant for equilibrium to be established is derived. Applications tovarious experiments in the literature are then made with fairly satisfactory results. A new interpretation of the theory of klino-kinesis with accommodation is found upon application of the equations developed to experimental work. Further limitations and uses of these equations are then discussed.
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  • 7
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    Bulletin of mathematical biology 15 (1953), S. 501-507 
    ISSN: 1522-9602
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    Notes: Abstract Certain parameters are defined which roughly characterize the internal structure of networks. A given network structure uniquely determines the values of the parameters, but the reverse is not true. The parameters therefore define certain classes of networks. One of the parameters, thedispersion D(S) gives an indication of the “compactness” of the internal structure. Addition theorems and inequalities are derived relating the dispersions of sub-systems to the dispersion of the complete structure.
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  • 8
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    Bulletin of mathematical biology 15 (1953), S. 489-500 
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    Notes: Abstract A mathematical theory is developed which permits the determination of certain parameters of an inhomogenous tissue, such as a nerve trunk without its epineurium. The parameters are the permeability coefficients for entrance into an exit of a substance from the nerve fibers, and the diffusion coefficient of the interstitial material. The experimental data required are the dimensions of the cross-section, the average diameter of the fibers, and the ratio of the cross-sectional are of the fibers to the total cross-section, as well as the time course of the decrease of the fraction of the substance left in the nerve trunk, when the trunk is immersed in a bathing solution containing none of it.
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  • 9
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    Bulletin of mathematical biology 15 (1953), S. 509-522 
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    Notes: Abstract A model based on enzyme localization is developed which gives rise to an apparent active transport of a metabolite into or out of cells. The model is applied to three simple situations, using Fick's equation and the Rashevsky approximation. It is shown that the apparent efficiency can be made as large as desired if, for constant reaction, the outer cell region is made sufficiently small, or, for autocatalytic reaction, if the metabolite concentration in the outer region is sufficiently small. The physical limitations imposed by this mechanism are developed for all three situations.
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    Bulletin of mathematical biology 15 (1953), S. 523-533 
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    Notes: Abstract A previously derived iteration formula for a random net was applied to some data on the spread of information through a population. It was found that if the axon density (the only free parameter in the formula) is determined by the first pair of experimental values, the predicted spread is much more rapid than the observed one. If the successive values of the “apparent axon density” are calculated from the successive experimental values, it is noticed that this quantity at first suffers a sharp drop from an initial high value to its lowest value and then gradually “recovers”. An attempt is made to account for this behavior of the apparent axon density in terms of the “assumption of transitivity”, based on a certain socio-structural bias, namely, that the likely contacts of two individuals who themselves have been in contact are expected to be strongly overlapping. The assumption of transitivity leads to a drop in the apparent axon density from an arbitrary initial value to the vicinity of unity (if the actual axon density is not too small). However, the “recovery” is not accounted for, and thus the predicted spread turns out to beslower than the observed.
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    Bulletin of mathematical biology 15 (1953), S. 535-546 
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    Notes: Abstract The assumption of transitivity treated in part I is modified in various ways to describe an information-diffusion process, in which a certain amount of randomness of contact does occur. In one model a parameter is introduced which is indicative of a tendency to go beyond one's immediate vicinity to spread the information as the vicinity becomes saturated with knowers. In another model the randomness appears in the assumption that new knowers are uniformly distributed among the knowers. Two of the equations thus derived, each with two free parameters are in good agreement with experimental results.
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  • 12
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    Bulletin of mathematical biology 29 (1967), S. 1-16 
    ISSN: 1522-9602
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    Topics: Biology , Mathematics
    Notes: Abstract A vast number of biologically important processes are based upon bimolecular systems. In these systems intermediate complexes are formed. Bimolecular systems in which no complex-complex interactions occur are called linear systems of complexes. A definition and some characteristic properties of these systems are given here. There may exist a contradiction of Onsager's principle of detailed balancing in these systems; however, no principal differences are found between the steady state behavior of an open system and that of a closed system. It is shown that the steady state behavior of a linear system of complexes of arbitrary complexity has some similarities with the steady state behavior of a simple bimolecular system, e.g., Michaelis-Menten enzymatic reaction. Multiplicity of action of the substances participating in biomolecular processes may produce some qualitative differences in the steady state behavior of the system.
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  • 13
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    Bulletin of mathematical biology 29 (1967), S. 17-32 
    ISSN: 1522-9602
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    Topics: Biology , Mathematics
    Notes: Abstract A time-dependent DNA histogram is calculated for an irradiated population of cells under the limiting assumption that the cells cannot pass through prophase due to the effects of the radiation. The population is assumed to increase exponentially prior to irradiation, but after irradiation to neither gain nor lose cells. Chromosome-number dispersion is taken into account in the calculation. The qualitative behavior of the calculated and experimental histograms are in reasonable agreement. The quantitative agreement between the two is relatively good at short post-irradiation times but is poor at long post-irradiation times (say, greater than half the doubling time). This suggests that recovery phenomena cannot be neglected at long post-irradiation times.
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  • 14
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    Bulletin of mathematical biology 29 (1967), S. 187-188 
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    Notes: Abstract It is pointed out that the three different stimuli for a corrective turn, namely the distance from the edge of the lane, the rate of approach to the edge, and the angle between the direction of the car and the direction of the lane (Bull. Math. Biophysics,28, 645–654, 1966,29, 181–186, 1967) may act all three simultaneously. It is found that in that case the tracking curve of the car is stable below a critical speed and becomes unstable above it.
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  • 15
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    Bulletin of mathematical biology 29 (1967), S. 181-186 
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    Notes: Abstract Continuing a previous study (Bull. Math. Biophysics, 28, 645–654, 1966), the biophysical mechanism of a corrective turn is investigated for the case where the stimulus for the corrective turn is produced not only by the perception of the nearness of an edge of the lane, but also by the rate of approach of the car towards the edge. In that case it is found that the tracking curve of the car may consist of a series of damped sinusoids and safe driving would be possible at any speed if it were not for the endogenous fluctuation in the driver's central nervous system. If the effect of the rate of approach increases sufficiently rapidly as the distance to the edge of the lane decreases, then a stable undamped oscillating tracking curve is possible. The case is also studied where the driver makes a corrective turn in response to a direct perception of the angle between the direction of the lane and the longitudinal axis of the car.
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  • 16
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    Bulletin of mathematical biology 29 (1967), S. 245-259 
    ISSN: 1522-9602
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    Notes: Abstract The principle of minimal work requires that the conducting airways of the human lung should have a maximum radius for minimal resistance to gas flow. At the same time there is a requirement that the airways should have a minimal volume for economy of space. These two opposing requirements have been investigated mathematically, and a method for calculating the angle of branching which produces minimal volume has been derived. The relationship of the radii of the parent and daughter branches to produce minimal resistance has been similarly defined. By measurement of a bronchial cast from a human lung the extent to which the predicted optimum structure is realized in practice has been shown. The change in structure associated with change of function at the transition from conducting airway to diffusion zone has been demonstrated.
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    Bulletin of mathematical biology 29 (1967), S. 191-206 
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    Notes: Abstract This paper considers a class of set-theoretical entities, calledn-rank Linnaean structures, which are intended as abstract models of the taxonomic classificatory systems of biology. In the first part, devoted to formalism, finite Linnaean structures are discussed in complete generality; but, in addition, eight distinct subclasses are noted and some of the properties of their elements are explored. In the second part, concerned with applications, it is shown that taxonomic systems may be recast in the form of finite Linnaean structures, and an effort is made to show that some undesirable features of earlier models are avoided without artificiality and without abandoning extensional mathematics.
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    Bulletin of mathematical biology 29 (1967), S. 207-216 
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    Notes: Abstract Generalizing an idea of M. Richardson (Fundamentals of Mathematics, New York: Macmillan Co., 1958), an APS on a given populationP is a non-empty collection of non-empty subsets ofP such that ifA is in the collection andA⊆B, thenB is in the collection. From a structure of this kind a partial ordering ofP, called therelated bumping order, is derived. The question is raised as to what kinds of partial orderings can be so obtained. For structures determined by voting weights of the members of the population, a complete characterization of all possible bumping orders is obtained.
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    Bulletin of mathematical biology 29 (1967), S. 217-226 
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    Notes: Abstract The “second method” of Liapunov is used to perform a stability analysis of a mathematical model of the neuron. This analysis is based on the hypothesis that the firing of the neuron coincides with a temporary state of instability of the system, and that the initiation of all-or-none process depends on the magnitude of membrane depolarization and its first time derivative. It is found that the stability (and hence the possibility of a second firing) is restored approximately when the rate of membrane repolarization is at a maximum. This result predicts that the duration of the period of absolute refractoriness in neurons would be about 75 per cent of the spike duration, and thus shorter than the value usually obtained from experimental measurements.
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    Bulletin of mathematical biology 29 (1967), S. 227-232 
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    Notes: Abstract Some aspects of masking phenomena are considered in terms of the simplest possible model of two-factor neural elements. The effect of a number of variables can be accounted for, but the introduction of an internuncial element results in a masking function which need not be symmetric about zero delay interval. As an illustration, the results for a special case are compared with available data. In general, such a model results in a masking function which depends on the intensity, area, and duration of the stimuli, as well as on the temporal and spatial separation between them.
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    Bulletin of mathematical biology 29 (1967), S. 377-388 
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    Notes: Abstract The general equations are discussed describing two species in competition or in symbiosis or feeding one on the other.
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    Bulletin of mathematical biology 29 (1967), S. 403-404 
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    Bulletin of mathematical biology 29 (1967), S. 389-393 
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    Notes: Abstract It is shown that the principle of biological epimorphism (Rashevsky,Mathematical Principles in Biology and Their Applications, Springfield, Ill.: Charles Thomas, 1960) is contained in the theory of organismic sets (Bull. Math. Biophysics,29, 139–152, 1967) if an additional postulate not directly connected to mappings is made.
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    Bulletin of mathematical biology 29 (1967), S. 407-407 
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    Bulletin of mathematical biology 29 (1967), S. 409-409 
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    Bulletin of mathematical biology 43 (1981), S. 1-19 
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    Notes: Abstract By studying the behavior of various tracer species in the lungs, one can assess many important characteristics which distinguish normal and abnormal function. Quantitative evaluation of function depends on the use of an appropriate model in conjunction with experimental data. A multi-compartment model is derived from mass balances to describe dynamic as well as (breath-averaged) steady-state transport processes between the environment and pulmonary capillary blood. The breathing cycle is divided into three time periods (inspiration, expiration, and pause) so that the model equations are discrete in time. No other model of tracer species transport in the lungs deals simultaneously with species dynamics, variable breathing pattern, distribution inhomogeneities, and non-equilibrium between alveolar gas and capillary blood. Models currently in the literature are shown to be special cases of the model presented here.
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    Bulletin of mathematical biology 43 (1981), S. 47-58 
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    Notes: Abstract Local stability seems to imply global stability for population models. To investigate this claim, we formally define apopulation model. This definition seems to include the one-dimensional discrete models now in use. We derive a necessary and sufficient condition for the global stability of our defined class of models. We derive an easily testable sufficient condition for local stability to imply global stability. We also show that if a discrete model is majorized by one of these stable population models, then the discrete model is globally stable. We demonstrate the utility of these theorems by using them to prove that the regions of local and global stability coincide for six models from the literature. We close by arguing that these theorems give a method for demonstrating global stability that is simpler and easier to apply than the usual method of Liapunov functions.
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    Bulletin of mathematical biology 43 (1981), S. 125-140 
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    Notes: Abstract The asymptotic behaviour of a logistic equation with diffusion on a bounded region and a diffusionally coupled delay is investigated. An equivelent parabolic system is derived for certain types of delays. Using a Layapunov functional, sufficient conditions for the global asymptotic stability of the constant steady state are obtained. When the global stability is lost, using Hopf's bifurcation theory, existence of travelling waves is shown for ring-like and periodic one dimensional habitats.
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    Bulletin of mathematical biology 43 (1981), S. 141-149 
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    Notes: Abstract It was hypothesized in an earlier work that sensory perception can occur only when the perceiving system is uncertain about the nature of the event being perceived. In the absence of any uncertainty, perception will not take place. The response of the sensory afferent neuron (impulse transmission rate) was calculated using Shannon's measure of uncertainty or entropy. It will now be shown that when the event being perceived is the position and momentum of a particle, Shannon's measure of uncertainty leads to the Heisenberg Uncertainty relationship.
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    Bulletin of mathematical biology 29 (1967), S. 605-613 
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    Notes: Abstract This paper deals with bimolecular systems in which also complex-complex interactions occur. Because of the complexity of the problem, an approximation in a form of coupled linear systems of complexes (Bull. Math. Biophysics,29, 1–16, 1967) is considered. Two types of couplings, serial and parallel, are studied. In the serial coupling the nonlinear system of complexes has the same behavior as its subsystems. An entity, initial sensitivity, has interesting properties: in serial coupling it is at most equal to the product and in parallel coupling, at most equal to the sum of partial initial sensitivities.
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    Bulletin of mathematical biology 29 (1967), S. 615-623 
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    Notes: Abstract Amplification effect in the catalytic bimolecular systems is a consequence of the kinetic characteristic of the catalyst. Two types of the coefficient of amplification are defined. The applicability of these definitions is given by the type of the bimolecular system. In a simple example it is shown that the concept of amplification is meaningful in these systems. Furthermore, two rules, analogous to those for a coupling of amplifiers, are derived for the two basic modes of coupling of catalytic systems. Thus, in biological systems the catalytic reactions may be regarded as biologically effective amplifiers.
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    Bulletin of mathematical biology 29 (1967), S. 583-596 
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    Notes: Abstract It is postulated that cell hydration is governed by adsorption of water on cell proteins in accord with the Bradley adsorption isotherm, and that the action of a solute in the surrounding solution is to lower the vapor pressure of the solution so that cell water adsorption is decreased by moving down the Bradley isotherm. From these concepts, it is derived that cell volume (V) should be related to solute concentration (x) by the equationV=−E log10 x+F whereE andF are constants which are independent of type of solute. For a non-adsorbed solute this agrees well with experimental data. For solutes which are adsorbed by cell proteins, a correction in the above equation may be necessary at higher solute concentrations, which is shown to be compatible with various experimental data. The types of experiments which are generally used to support the osmotic pressure theory of cell hydration agree equally well with the adsorption theory. The virtue of the adsorption theory is that, unlike the osmotic pressure theory of cell swelling, it is compatible with permeability of the cell membrane to solutes, which has been experimentally observed for various solutes.
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    Bulletin of mathematical biology 43 (1981), S. 239-244 
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    Notes: Abstract It is not unusual for several classifications to be given for the same collection of objects. We present a method, called majority rule, which can be used to define a consensus of these classifications. We also discuss some mathematical properties of this consensus tree.
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    Bulletin of mathematical biology 43 (1981), S. 259-270 
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    Notes: Abstract The dependence of the spatial concentration profiles of morphogens on a characteristic dimension is obtained by continuation techniques for Gierer and Meinhardt's activator-inhibitor model of morphogenesis. The study of the behaviour of the system during growth, where the linear and exponential increase of the characteristic dimension is considered, revealed that more complex patterns of morphogen spatial concentrations appear regularly in a reproducible way.
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    Bulletin of mathematical biology 43 (1981), S. 271-278 
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    Notes: Abstract Computer models have been used by various authors to simulate both the growth of normal cellular tissue and the development of cancerous cells within normal tissue. As these models were the result of considerable idealization, the purpose of the present paper is to propose a model in which the degree of simplification is relaxed: the features of simultaneous growth, and cell growth whose rate depends on the free absorbing periphery of the cell are introduced. Simulation experiments have been conducted using the model, and the results are presented.
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    Bulletin of mathematical biology 43 (1981), S. 341-346 
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    Notes: Abstract The theory of complementary variational principles is used to obtain maximum and minimum principles for a nonlinear model of heat conduction in the human head. Accurate variational solutions are obtained in illustrative calculations. The effect of nonlinearity is seen to be significant from a comparison with the linearized model.
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    Bulletin of mathematical biology 43 (1981), S. 279-325 
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    Notes: Abstract A model for the nerve impulse due to Zeeman (1972) and based on catastrophe theory is compared with alternative models and criticisms of Zeeman's model by Sussmann and Zahler (1977, 1978) are assessed. The criticisms of Zeeman's motivation for his model are found to carry some weight. Sussmann and Zahler (1977, 1978) list numerous features of Zeeman's model which, they state, are not in agreement with experiment. These statements as they stand are largely erroneous, and the model still remains to be tested by a critical series of experiments. However, a detailed analysis reveals defects in Zeeman's model, not among those claimed by Sussmann and Zahler, showing that the explicit equations of the model cannot be correct. The possibility of a modified approach along similar lines and its ultimate adoption remains open.
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    Bulletin of mathematical biology 29 (1967), S. 657-664 
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    Notes: Abstract Conditions under which a time varying electromagnetic field problem (such as arises in electrophysiology, electrocardiography, etc.) can be reduced to the conventional quasistatic problem are summarized. These conditions are discussed for typical physiological parameters.
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    Bulletin of mathematical biology 29 (1967), S. 711-718 
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    Notes: Abstract A compartmental lung model with any number of synchronously filling and emptying functional chambers and a common dead space or conducting region is considered. It is shown that the model gives rise to an output, in an open circuit washout determination, which is a weighted sum of exponentials. From estimates of these weights and exponential components, estimates of the model parameters can be recovered. Relations giving the unique correspondence between the output parameters and the model parameters are derived and the existence and uniqueness of solutions established.
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    Bulletin of mathematical biology 29 (1967), S. 677-690 
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    Notes: Abstract A physical model that incorporates all the experimental information on the formation of the visual pigment rhodopsin is presented. The visual pigments consist of a chromophore bound to an appropriate protein. Thus rhodopsin (λm 505 mμ) is formed by a Schiff’s base linkage C19H27CH=NH+-opsin (λm 440 mμ) between 11-cis retinal (λm 380 mμ) and the protein opsin (λm 280 mμ). It is found that there exists a red shift in the spectrum of rhodopsin from the Schiff’s base. The model brings an explanation for this red shift. It is shown that such a shift may be due to a charge transfer process (R. S. Mulliken,J. Am. Chem. Soc.,74, 811–824, 1952) between an electron at the double bond of carbons C11−C12 and an atomic orbital of the sulphur present in cysteine. This provides an explanation of the presence of SH-groups in the protein after the absorption of light. A one-electron approximation is used and the dipole momentμ NV ; hence, the oscillator strengthf of the transitionNV is estimated and compared with the experimentally determined extinction coefficient ∈m by mixing 3.5×10−3 M of 11-cis retinal with 8.3×10−5 M of cysteine at pH ranges 6 through 8. Reasonable agreement is found. Solvent, concentration and temperature dependence are shown also.
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    Bulletin of mathematical biology 43 (1981), S. 375-388 
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    Notes: Abstract The irreversible Michaelis-Menten reaction is studied by the use of the method of multiple scales. Three stages of the reaction are identified, one of which is studied in detail. The results are compared with those of two earlier analyses.
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    Bulletin of mathematical biology 43 (1981), S. 389-400 
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    Notes: Abstract A numerical study of the coupled nerve fibre problem is given which verifies and extends the perturbation theory of Luzader. Pulses on adjacent fibres can couple together with two possible stable pulse separations.
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    Bulletin of mathematical biology 43 (1981), S. 401-413 
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    Notes: Abstract A possible mechanism for effects of microwave radiation on the auditory system is the generation of field-induced forces at interfaces that divide materials of dissimilar electrical properties. A general expression for these “Maxwell stresses” is derived and then used to calculate the approximate magnitude of field-induced force within the organ of Corti during microwave exposure. Comparison of the results with data on the force needed to excite cochlear hair cells indicates auditory responses could be evoked by this mechanism at power densities near the threshold of rf hearing sensations.
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    Bulletin of mathematical biology 43 (1981), S. 415-426 
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    Notes: Abstract A definition of homogeneity for neural networks is given which permits their construction as group quotients. The significance of this for neural dynamics is discussed.
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    Bulletin of mathematical biology 43 (1981), S. 447-461 
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    Notes: Abstract The left ventricle is represented as a cylinder contracting both radially and longitudinally. A simple method is indicated to derive an expression for the rate of change of the kinetic energy of this three-dimensional model, which quantity can be used as an index for the study of the contractile behaviour of the myocardium. An application to the study of muscle mechanics is also indicated.
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    Bulletin of mathematical biology 43 (1981), S. 463-485 
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    Notes: Abstract A perturbation method is proposed to calculate approximately the limit cycle type nonequilibrium steady-state resulting from periodic perturbation of coefficients of stable population systems; the two species Lotka-Volterra competition system is explicity studied and the results are formulated for general multi-species population systems. Avoidance of competitive or other types of exclusion of species in a periodic environment is indicated.
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    Bulletin of mathematical biology 43 (1981), S. 513-516 
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    Bulletin of mathematical biology 29 (1967), S. 841-862 
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    Notes: Abstract By assigning coordinates to the environmental function space comprising all physical and mental stimuli, mathematical interpretations can be based on such terms as adaptability, and reactivity which relate to individuals interacting with their environment within a society. These psychometric concepts are incorporated into a framework of functional analysis, which permits the optimization of social change by maximizing the satisfaction integral through the use of variational or dynamic programming methods in conjunction with some optimal social policy. The approach provides a mathematical connection between psychology and sociology, and further demonstrates that existing forms of government are simulated by differential equations belonging to the same general class. The synthesis of new classes of functional equations describing social progress is visualized as a legitimate objective for abstract mathematical sociology.
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    Bulletin of mathematical biology 29 (1967), S. 863-877 
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    Notes: Abstract The theory of imitative behavior as developed hitherto by the author was based on the assumption that each individual has a natural preference for one of the two mutually exclusive behaviors. The endogenous fluctuations in the central nervous system then result in the individual’s exhibiting the two behaviors alternately with a relative frequency determined by the natural preference. Imitation shifts the natural preference towards one or the other of the two mutually exclusive behaviors. In the present approach it is suggested that the relative frequency of the two mutually exclusive behaviors exhibited alternately is determined by maximizing the “satisfaction function” of the individual, that is by hedonistic factors rather than by purely random fluctuations. Corresponding equations are developed. It is shown that in certain cases, even when the imitation effect is absent, a sort of “pseudoimitation” may occur. Another situation leads, in the case of two individuals only, to a complete “division of labor” between them, with respect to the two behaviors. Each one exhibits only one behavior. After that imitation is introduced explicitly by assuming that imitation by one individual or another increases the satisfaction function of the imitating individual. Results thus obtained show similarities to the results of the old theory.
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    Bulletin of mathematical biology 15 (1953), S. 1-13 
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    Notes: Abstract The impressed field, “Window Field” (WF), due to a half-wave action potential on a muscle fiber, has been calculated on the basis of potential theory. It has been shown that in spite of the small intensity of the field, its integrated action can transfer the energy needed to induce, contraction from the membrane to the interior of the fiber. The energy of polarization has been found to be sufficient to exceed the energy of, thermal agitation on that length of fiber, which can be identified as the length of a sarcomere. The changes of ion concentration, caused by the WF, if calculated on the assumption of the semipermeability of theZ membranes, was found to be equal to the changes necessary to induce contraction of actomyosinin vitro.
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    Bulletin of mathematical biology 15 (1953), S. 15-21 
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    Notes: Abstract Some general properties of the solution of the diffusion equation are deduced for the steady-state, spherically symmetric system. On the basis of these developments some results of N. Rashevsky (Bull. Math. Biophysics,11, 15, 1949) are discussed and the results of a previous investigation (Hearon,Bull. Math. Biophysics,12, 135, 1950b) are extended to more general conditions. In particular these extensions apply to the flow of a soluteagainst its concentration gradient, the nonzero gradient of an inert metabolite, and theaccumulation or exclusion of an inert metabolite in a metabolic system.
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    Bulletin of mathematical biology 15 (1953), S. 23-31 
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    Notes: Abstract The approximation method of N. Rashevsky is discussed and reviewed. It is shown that in addition to theexplicit assumptions and approximations there is involved the assumption that the rate of metabolism is the same at every point in the cell and that theaverage rate of metabolism is different from zero. An expression is given for the error in the approximate method when the rate of metabolism is any function of the concentration. It is also shown that a solution in theform of that obtained by the approximate method is not possible if the generalized laws of diffusion are assumed to apply.
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    Bulletin of mathematical biology 15 (1953), S. 33-42 
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    Notes: Abstract Rate equations for the enzymatic oxidation of succinic acid are derived on the assumption that when a single molecule of substrate combines with an enzyme molecule, it can do so with either one or two sites on the enzyme, and that oxidation occurs only in the second case. In addition it is assumed that the product of the reaction, fumaric acid, combines reversibly with the enzyme. With certain enzyme preparations the data fitted such an equation satisfactorily. In others the rate was that of a first-order reaction, but addition of cytochrome changed it to the former type. It was concluded that the transfer of hydrogen to oxygen was a first-order reaction and dominated the whole rate when enzyme preparations were used which had been washed relatively free of cytochrome. When the limiting factor was succino-dehydrogenase the rates followed the new equation. Criteria for recognizing noncompetitive inhibition are given, and inhibition by di-tertiary butyl peroxide was shown to be of this type.
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    Bulletin of mathematical biology 15 (1953), S. 43-47 
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    Notes: Abstract This paper deals with the following question: Which distributions of radiosensitivity in a population can lead to an exponential survival curve? The problem is solved exactly, with statistical fluctuations in dose fully accounted for. It is shown that only an exponential distribution of sensitivities can give rise to an exponential survival curve.
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    Bulletin of mathematical biology 15 (1953), S. 49-61 
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    Notes: Abstract An approximation method is introduced which enables a number of diffusion-type problems to be solved in an approximate but simple manner. Many cases require only the solution of a simple first-order differential equation. The method is applied to a number of cases in which the exact solutions are available. A comparison shows that the method is quite satisfactory in these cases. The method is applied to diffusion problems with rate of consumption proportional to concentration or to the square of the concentration. In the latter case, the result obtained is essentially the same as that found by H. G. Landau (1950) after elaborate calculations.
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    Bulletin of mathematical biology 15 (1953), S. 83-91 
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    Notes: Abstract It is shown that a slight modification of a model of excitatory phenomena in irritable tissues, which has been treated before, exhibits spontaneous oscillations. The frequency of these oscillations and the time-course of the potential across the model membrane have been determined, together with the dependence of some of their characteristics on some important parameters, particularly (Ca++).
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    Bulletin of mathematical biology 15 (1953), S. 73-81 
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    Notes: Abstract It is demonstrated that, if the variations of viscosity throughout a cell are considered, swelling stresses may produce elongation and division. To do this it is necessary to generalize Betti's theorem to cover systems containing viscosity gradients and such a generalization is presented. On the basis of two special assumptions it is shown that most of the results of the diffusion drag theory of cell division may be duplicated by the present theory.
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    Bulletin of mathematical biology 15 (1953), S. 63-71 
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    Notes: Abstract The theory of imitative behavior, developed previously, is applied to the case of two social groups which are separated spatially. If the information of each group as to the behavior of the other is complete, the case reduces to that of a single group. When any information is lacking at all, the two groups are independent. If we have two mutually exclusive behaviorsA andB, all four combinationsAA, AB, BA, andBB are possible. If the mutual information gradually increases from zero, then for a certain value of it, the group which is more informed about the behavior of the other will change to that behavior if it did not already exhibit it. If for constant information the size of the group increases, then above a certain threshold value, the larger group imposes its behavior on the smaller.
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    Bulletin of mathematical biology 15 (1953), S. 103-104 
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    Bulletin of mathematical biology 15 (1953), S. 107-107 
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    Bulletin of mathematical biology 15 (1953), S. 105-106 
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    Bulletin of mathematical biology 15 (1953), S. 93-101 
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    Notes: Abstract Local regulation of blood flow as determined by capillary diameter and the number of open capillaries in a region is considered. The local changes in capillary diameter and in the number of open capillaries are assumed to be due to concentration changes of a diffusible, nonspecified metabolite. This metabolite is produced in the tissue and carried away by the blood stream. Using these assumptions and applying pertinent data on capillaries, deductions are made concerning: (a) the law of blood flow as a function of temperature and capillary radius for the hyperemia of high temperature, (b) high flow as it depends on metabolism during strenuous exercise of muscle, and (c) a first approximation to the time duration of occlusion hyperemia.
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    Bulletin of mathematical biology 15 (1953), S. 109-109 
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    Bulletin of mathematical biology 15 (1953), S. 143-148 
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    Notes: Abstract The necessary and sufficient condition is given forn integers to be the score structure of a society with a dominance relation. A proof is also given for a theorem showing that there are members who dominate every other member either directly or indirectly through a single intermediate member.
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    Bulletin of mathematical biology 15 (1953), S. 111-119 
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    Notes: Abstract On the basis of a previous general formulation (Bull. Math. Biophysics,15, 21–29, 1953a) a discussion is given of the error in the approximation method of N. Rashevsky. This error, inherent in the method when the metabolic rate is different at each point in the cell, is discussed in detail and numerical values are presented for two particular cases: the rate proportional to the concentration and the rate a prescribed function of the spatial coordinates. It is shown that the formulation for the first case also applies to several other cases, that the error is negligible provided the rate is sufficiently small, and that the error is fairly sensitive to the cell size. If the rate depends upon the coordinatesalone a small rate is not sufficient to insure a negligible error. The relations between the exact method, the standard approximate method, an earlier approximate method (Physics,7 260, 1936), and a more recent refinement (Bull. Math. Biophysics,10, 201, 1948) of the standard method are discussed.
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    Bulletin of mathematical biology 15 (1953), S. 121-141 
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    Notes: Abstract It is shown on the basis of (1) conservation of mass, (2) positive concentrations, and (3) the principle of detail balancing that periodic reactions cannot occur in a closed system described bylinear differential equations. The matrix,A, of the rate equations must be such that |A|=0,a ij〉0 fori≠j,a ii〈0, andVAV −1=B, whereV is diagonal andB is symmetric. These properties ofA imply that the latent roots are real and non-positive and that neither catalysis nor inhibition can be described bylinear equations. It is further shown that periodic reactions cannot occur in anopen system for which the matrix associated with the chemical reactions has the above properties and in which thesimple law of diffusion is obeyed. The relation of these results to Onsager's reciprocal relations and to previous work on periodic and cyclic chemical reactions is discussed. The utility of certain of these results for the treatment of isotope kinetics is indicated.
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    Bulletin of mathematical biology 15 (1953), S. 149-152 
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    Notes: Abstract It is demonstrated that an explanation of the small radius effect or the so-called sigma phenomenon may be obtained by noting that one of the effects of the presence of suspended particles in a flowing fluid is to increase the velocity of flow near the wall over that existing in the absence of particles. This effect may be considered equivalent to relaxing the boundary conditions at the wall. An expression for the viscosity is compared with data and fit is found to be good.
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    Bulletin of mathematical biology 15 (1953), S. 153-159 
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    Notes: Abstract The solution for the spatial distribution of ions in a Donnan equilibrium has been given by J. H. Bartlett and R. A. Kromhout (1952). The present note gives an explicit solution for the case in which the length of the region containing the membrane is large; in biological situations this requires only that the length considered should be greater than a few hundred Ångstrom units. The Donnan equilibrium may be considered to be a special case of a situation in which forces other than electrical act upon the ions; in particular, it represents the case in which only one ion is acted upon and the energy difference on the two sides of the membrane is infinite. An expression is given for the difference in energy of theith in terms of the electrical potential and of the ion concentrations. As an illustration, the results are applied to nerve membrane potentials.
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    Bulletin of mathematical biology 15 (1953), S. 161-165 
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    Notes: Abstract A mechanism is described which accounts for the active transport of Na+ ions through a membrane. It is assumed that at one side of the membrane the ion combines with a carrier ion, the resulting carrier compound then diffuses through the membrane and decomposes at the other side of the membrane. The free diffusion of the ions is also taken into account. The time rate of accumulation of the ion in question at the latter side of the membrane is calculated in terms of the concentrations of the ion at both sides of the membrane.
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    Bulletin of mathematical biology 15 (1953), S. 167-171 
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    Notes: Abstract The recent extension of the approximation method is applied to enable us to arrive at the time course of the concentrations at both sides of a membrane. From the differential equations which govern these, the steady-state solution is obtained in terms of the parameters, which are determined by the thickness of the diffusion layers, the chemical composition and reactions, and the diffusion constant of the membrane.
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    Bulletin of mathematical biology 15 (1953), S. 173-183 
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    Notes: Abstract An equation is derived from the spread of a “state” by contact through a thoroughly mixed population, in which the probability of transmission depends both on the over-all duration of the process and on the time an individual has been in the “state.” Cases in which this probability is a function of only one or the other of the two “times” are worked out. It is shown that in the case of dependence on “private time” alone the asymptotic value of the fraction of the population effected is the same as that derived by the random net approach.
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    Bulletin of mathematical biology 15 (1953), S. 235-235 
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    Bulletin of mathematical biology 15 (1953), S. 185-195 
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    Notes: Abstract The reflection of pressure waves in a fluid enclosed within a tube with an elastic wall is studied for the case of a localized change in diameter of the tube. The concept of impedance is introduced. The relation of the reflection characteristics of the parts of the tube at either side of the change is derived on the basis of the continuity of pressure and mass flow at the site of the change. This relations is used to derive the expression for the ratio of the pressure oscillations measured in front of, and behind, the constriction in terms of the constants of the system. As a result, a method is indicated to locate the coarctation from measurements of the pressures in front of, and behind it.
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    Notes: Abstract The question is raised concerning the possible causes of abnormally small standard deviations found in counting samples in which particles are distributed at random (e.g., blood cells, fat globules in milk, etc.). The effect of discarding abnormal samples is discounted inasmuch as small standard deviations occur even when all samples are counted. An approximation method is used to calculate the effect of finite particle size, of known repulsive forces between particles and of convection currents. This calculation shows that neither finite size nor the known repulsive forces are sufficient to account for the observed abnormality of standard deviation, but that convection currents can possibly account for it. The possible presence of long-range repulsive forces cannot, however, be excluded.
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    Bulletin of mathematical biology 15 (1953), S. 245-250 
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    Notes: Abstract The effect of finite particle size on the standard deviation in sample counts is computed for the one-dimensional case. To a first order of approximation the correction is found to be identifical with that found by H. de Vries (1953) using a general approximation method.
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    Bulletin of mathematical biology 15 (1953), S. 251-260 
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    Notes: Abstract A critical examination of the “classical” theories of photoreception in view of more recent experimental findings yields the result that these theories do not possess the property to describe all the more significant phenomena of photoreception correctly, and to some extent suffer the lack of more general applicability. The basis for a new and presumably more general theory of photoreception based on dynamical aspects is laid out. Emphasis is put on the time course of afferent and efferent excitation in the photoreception model, consisting of a receptor element, an afferent and an efferent neuron of the one-factor Rashevsky-type, and an effector organ.
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    Bulletin of mathematical biology 15 (1953), S. 197-234 
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    Notes: Abstract A mathematical model for the development of human society, beginning with the earliest stages of urban cultures, is outlined. In the early stages of history, behavior was characterized largely by adherence to a number of beliefs and prejudices of diffeirent kinds, which were accepted on faith and not subject to critical rational analysis. Due to psychobiological variability a very small number of individuals spontaneously appear at all times who challenge the accepted beliefs and prejudices and do not follow the accepted patterns of social behavior. The effect of these individuals upon the rest of the society, especially upon the younger generation, depends on the facilities with which information spreads in society. In earliest societies, when modern methods of mass communication were unknown, the channels of communication were practically identical with the channels of economic transport. The latter in its turn depended on the nature of the roads, and especially on the presence of waterway, which facilitated transportation. The sizes of the earliest cities and the distances between them were largely determined by relative ease of transporation. Expressions are derived for the average size of the earliest cities and for the average distance between them. The calculated average populations of the earliest cities are of the order of 103; the distance of the order of 102 km. Both are in agreement with some archaeological findings. An expression for the time spaon required for the development from the earliest stages of urban cultures to the present time is derived and shown to depend on the specific shoreline of the country, that is, the length of the shorline divided by the area of the country. It is pointed out that western Europe's specific shoreline, including land bordering both seas and rivers, is ten times as large as the shoreline area of other parts of the world. It is shown that this greater specific shoreline may account quantitatively for the faster social and technological development of western Europe in the last few centuries. The calculated total span of time of development from earliest urban cultures to our days is found to be of the order of magnitude of ten thousand years. It is shown that the model accounts for the existence at the present time of primitive cultures. A number of suggestions is made in regard to other possible applications of mathematics to history.
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    Bulletin of mathematical biology 15 (1953), S. 269-276 
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    Notes: Abstract The assumptions latent in the derivation of the integral equation of Branson are rendered explicit and discussed. It is shown that the equation is valid only for systems in which the substance disappears according to a linear rate law.
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    Bulletin of mathematical biology 15 (1953), S. 261-268 
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    Notes: Abstract It is shown that the validity of Branson's integral description of metabolizing systems is subject to severe limitations. The validity is insured only in cases where the reaction is of first order, or quasi of first order. In all other cases Branson's equation has to be modified to insure general applicability. The consequences of a different definition of the metabolizing functionF have also been investigated. With the new definitionF describes the pure effect of metabolization. It is found that in this case the integral equation is only capable of describing first-order reactions. With a slight modification of the integral equation it is possible to describe metabolites “with age”, which do not have reactions of definite order, but which satisfy the superposition principle.
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    Bulletin of mathematical biology 15 (1953), S. 277-292 
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    Notes: Abstract A problem in probability is stated with included the problem of the distribution of bacterial mutants as a special case. This problem is solved exactly but since the resulting expressions are too complicated for practical use, various approximate expressions for the distribution are considered, especially for the bacterial mutation case.
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    Bulletin of mathematical biology 15 (1953), S. 293-300 
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    Notes: Abstract Simple reaction and discrimination reaction, under the influence of imitation, are considered for the situation in which the stimulus or the stimuli vary slowly with time. The result is analogous to hysteresis under certain conditions. The calculations are facilitated by the solution of $$x = \int_{ - \infty }^{a + \beta x} {g\left( \xi \right)d} \xi ,$$ g(ξ) being the normal error function. Values ofx(α, β) are given in a table.
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    Bulletin of mathematical biology 15 (1953), S. 301-309 
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    Notes: Abstract On the basis of simple physical considerations the blood flow in a branching circulatory system is studied. The case of two groups of parallel vessels is treated. The vessels of the same group are supposed to be identical. The resistance of each group is determined by the resistance of each vessel in the group and by the number of vessels in the group. From the dependence of the resistance of each vessel on its radius an expression is obtained for the blood flow through each group of vessels in terms of the numbers and sizes of the vessels in each group. The number of open vessels in an organ and the radius of each of those vessels are assumed to depend on the metabolic rate of that organ. The relations so obtained, together with the expression above, are applied to derive the blood flow through an organ as a function of the metabolic rate of that organ. It is indicated that the relations obtained might describe the shifting of blood from one organ to another if the activity of one of them changes. A way is pointed out to treat neural regulation of this phenomenon.
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    Bulletin of mathematical biology 15 (1953), S. 361-365 
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    Notes: Abstract A stochastic model of population growth is treated using the Bellman-Harris theory of agedependent stochastic branching processes. The probability distribution for the population size at any time and the expectation are obtained when it is assumed that there is probability (1−σ), 0≤σ〈1, of the organism dividing into two at the end of its lifetime, and probability σ that division will not take place.
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    Bulletin of mathematical biology 15 (1953), S. 339-359 
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    Notes: Abstract In a previous paper, in which a possible mathematical approach to history was outlined, it was shown that urbanization plays an important part in the propagation of new ideas. The rate of such propagation influences the rate of historical developments. The present paper deals in more detail with possible mechanisms of formation of earliest cities. Equations are derived which give the limiting size of such cities and their rate of growth. Of particular importance for the spread of new ideas is the spread of information. The latter largely depends on the fraction of individuals who travel between city and country. Expressions for this quantity are derived. An approach is outlined to the mathematical study of the earliest social classes, which may have been formed as a result of military, religious, or economic stratifications.
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    Bulletin of mathematical biology 15 (1953), S. 395-409 
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    Notes: Abstract The velocity of propagation of a disturbance wave in a liquid flowing in a distensible tube is computed. The mathematical model is more general than those used in previous analyses: the tube wall properties are realistic; the convective part of the axial inertia forces is taken into account; radial inertia forces of both the fluid and tube wall are present; viscous stresses are present. Four parameters influencing the velocity of propagation are obtained and discussed. Curves are plotted illustrating the effects of the parameters. Contrary to the results of previous analyses, viscous effects are shown to be appreciable in blood flow. It is also shown that radial inertia effects can be important in laboratory set-ups.
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    Bulletin of mathematical biology 15 (1953), S. 411-429 
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    Notes: Abstract A general theory of the drying of frozen tissue is developed and applied to the measurement of the drying rate of frozen guinea pig liver. It is shown that for a given temperature of the subliming ice crystals the mininum drying time of a piece of guinea pig liver is greater than the minimum sublimation time of a piece of ice of the same size and shape by a factor of the order of one thousand. This fact has many implications in the design of freeze-dry apparatus which will be developed in a following paper.
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    Bulletin of mathematical biology 15 (1953), S. 477-488 
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    Notes: Abstract The equations governing the time course of the exchange of substances between the blood in the capillaries and the extracellular space are solved for the case of substances which do not penetrate the cells. The equations given relate the time course of the exchange process to the various tissue and circulation parameters such as the specific capillary wall area, the pore area, the inter-capillary distance, the size of the extra-vascular, extra-cellular space, the diffusion coefficient in this space, and the velocity of blood in the capillaries. Some experimental work on capillary exchange is discussed in relation to the theory and estimates are made of the relative importance of the various tissue and circulation parameters in the exchange of substances in different tissues.
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    Bulletin of mathematical biology 29 (1967), S. 233-243 
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    Notes: Abstract A neuron is assumed to receive synaptic input of both excitatory and inhibitory natures from a large number of neighboring neurons; it is also assumed that a large number of such impulses are required to raise the neuron’s transmembrane potential to its threshold potential, at which it “fires” or “spikes”. The model is similar to one of Gerstein and Mandelbrot, except that in the absence of input an exponential decay of potential toward a resting level is introduced. Computational methods of determining the spike timeinterval distribution are discussed, along with the inverse problem of estimating the parameters of the system from observed spike time-interval data.
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    Bulletin of mathematical biology 29 (1967), S. 311-318 
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    Notes: Abstract Moens-Korteweg relations are developed for the velocity of wave propagation through an orthotropic elastic tube based on the three dimensional equations of elasticity. Numerical examples are presented for the femoral artery of a dog and several other orthotropic materials. These results are compared with those obtained from the equations of motion for an orthotropic elastic medium.
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    Bulletin of mathematical biology 29 (1967), S. 335-341 
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    Notes: Abstract Up to the present time, the following property of the product component in the reversible one substrate-one intermediate-one product enzymic mechanism has been taken only as anassumption, viz., during the course of the reaction, the time-rate of change of product concentration is never negative and the product concentration never exceeds its equilibrium value. Applying the methods of the geometric theory of ordinary differential equations it is shown that this result follows as a direct deduction from the differential equations governing the mechanism together with the initial conditions. Further, the nature of the equilibrium point as a stable node is established.
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    Bulletin of mathematical biology 29 (1967), S. 319-333 
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    Notes: Abstract An integro-differential equation treatment of multi-compartment systems is developed which permits formal analysis of the incomplete data which is available from partly accessible, partly injectable systems. New transport functions are defined which can be obtained directly from the experimental data. These functions serve to characterize the communication and topology between different accessible compartments and also the reentrant contributions from inaccessible sites. The method gives solutions consistent with those of the differential equation approach when the system is uniformly contiguous and accessible, more complete solutions than those of the integral equation approach when all measured compartments are injectable, and in addition provides complete or partial solutions for certain otherwise analytically intractable systems. Detailed numerical illustrations of the method are given.
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    Bulletin of mathematical biology 29 (1967), S. 405-405 
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    Bulletin of mathematical biology 29 (1967), S. 395-401 
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    Notes: Abstract In continuation of previous work (Rashevsky,Some Medical Aspects of Mathematical Biology, Springfield, Ill.: Charles C. Thomas, 1964, Chap. 23 and Appendix 14), the study of the effects of the physical parameters of the cells of endocrine glands on the onset of sustained periodical oscillations in the interaction between the anterior pituitary and the thyroid hormones is generalized to include the possible effect of the intercellular fluid and of the degree of vascularization. Some conclusions of the previous study remain valid although some modifications must be made. A decreased relative volume of the intercellular fluid and an increased vascularization favor the conditions for sustained oscillations. The permeability of the cells and the permeability of the capillaries appear explicitly in the expressions which show the conditions for sustained periodicities.
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    Bulletin of mathematical biology 43 (1981), S. 59-67 
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    Notes: Abstract The theory of computational complexity and certain explicitly-stated hypotheses imply limitations on the information processing power of biological systems. Parallelism, special purpose organization, and analog mechanisms may provide speedup critical for life processes, but have little power in the face of exponential growth. We show that “polynomially simulatable” biological systems cannot exhibit dynamic behavior which produces the solution of an intractable problem. The argument implies that parallelism does not allow biological systems to defeat the exponential explosion, but rather is important because it allows polynomial time algorithms to be used more efficiently.
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    Bulletin of mathematical biology 29 (1967), S. 437-449 
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    Notes: Abstract A general model of a large 2m-ploid breeding population, withr loci ands h alleles at the h th locus is considered. It is assumed that the population is bisexual, non-overlapping and breeds by random mating. The genotypic structure of the population is presented as a bilinear form in the gametic output vectors where the genotype distribution is in the matrix form. Using the concept of the segregation distribution, the genotype proportions in the (n+1)st generation are given. An equilibrium condition for random chromosome segregation is obtained in terms of gene frequencies.
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    Bulletin of mathematical biology 43 (1981), S. 81-88 
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    Notes: Abstract A correlation matrix analysis is applied to the base sequence of MS2 and ϕX174 in comparison with sets of simulated sequences with different degrees of constaint Significant differences between a codified sequence, and a statistical one in terms of the “correlation matrix” for sets of different length cannot be found. This result is analysed in terms of nucleotide sequences with different levels of informational content.
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    Bulletin of mathematical biology 43 (1981), S. 101-109 
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    Notes: Abstract A method of calculating the volume of a tree distal to a cut at the origin of a branch, using branching, diameter and length ratios, has been developed. The method was applied to bronchial tree casts from human, dog, sheep, hamster, and rat lungs. It was found that the exponenta in the equation weight=k×diameter a is approximately equal to 3.0 in sheep lung casts, as found by Hooper (1977), but it is greater than 3.0 in casts from the other four species.
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    Bulletin of mathematical biology 43 (1981), S. 111-116 
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    Notes: Abstract In this note we examine a continuous time version of a compartmental model introduced in a discrete time setting by S. R. Bernard. The model allows for more than one particle to leave the system at any time. This introduces additional randomness into the system, over the pure death system and this is reflected in the variance function.
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    Bulletin of mathematical biology 43 (1981), S. 89-99 
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    Notes: Abstract The mean first passage time for free diffusion can be derived directly by solving a simple analogue steady state problem. In this problem the diffusion starting region is considered as a time independent source of diffusing particles and the diffusion target assumes the behaviour of a perfectly absorbing sink. It is shown here that the transit time between the source and the sink, which in this particular problem is equal to the ratio between the holdup of the system and the total flux, is identical to the Brownian movement concept of the mean first passage time for free diffusion. This established identity considerably facilitates the derivation and investigation of the timing of diffusion in complicated structures such as those commonly found in living organisms.
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    Bulletin of mathematical biology 43 (1981), S. 121-123 
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