The Chemical Basis of Morphogenesis
14 Aug 1952-Philosophical Transactions of the Royal Society B (The Royal Society)-Vol. 237, Iss: 641, pp 37-72
TL;DR: In this article, it is suggested that a system of chemical substances, called morphogens, reacting together and diffusing through a tissue, is adequate to account for the main phenomena of morphogenesis.
Abstract: It is suggested that a system of chemical substances, called morphogens, reacting together and diffusing through a tissue, is adequate to account for the main phenomena of morphogenesis. Such a system, although it may originally be quite homogeneous, may later develop a pattern or structure due to an instability of the homogeneous equilibrium, which is triggered off by random disturbances. Such reaction-diffusion systems are considered in some detail in the case of an isolated ring of cells, a mathematically convenient, though biologically unusual system. The investigation is chiefly concerned with the onset of instability. It is found that there are six essentially different forms which this may take. In the most interesting form stationary waves appear on the ring. It is suggested that this might account, for instance, for the tentacle patterns on Hydra and for whorled leaves. A system of reactions and diffusion on a sphere is also considered. Such a system appears to account for gastrulation. Another reaction system in two dimensions gives rise to patterns reminiscent of dappling. It is also suggested that stationary waves in two dimensions could account for the phenomena of phyllotaxis. The purpose of this paper is to discuss a possible mechanism by which the genes of a zygote may determine the anatomical structure of the resulting organism. The theory does not make any new hypotheses; it merely suggests that certain well-known physical laws are sufficient to account for many of the facts. The full understanding of the paper requires a good knowledge of mathematics, some biology, and some elementary chemistry. Since readers cannot be expected to be experts in all of these subjects, a number of elementary facts are explained, which can be found in text-books, but whose omission would make the paper difficult reading.
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TL;DR: A comprehensive review of spatiotemporal pattern formation in systems driven away from equilibrium is presented in this article, with emphasis on comparisons between theory and quantitative experiments, and a classification of patterns in terms of the characteristic wave vector q 0 and frequency ω 0 of the instability.
Abstract: A comprehensive review of spatiotemporal pattern formation in systems driven away from equilibrium is presented, with emphasis on comparisons between theory and quantitative experiments. Examples include patterns in hydrodynamic systems such as thermal convection in pure fluids and binary mixtures, Taylor-Couette flow, parametric-wave instabilities, as well as patterns in solidification fronts, nonlinear optics, oscillatory chemical reactions and excitable biological media. The theoretical starting point is usually a set of deterministic equations of motion, typically in the form of nonlinear partial differential equations. These are sometimes supplemented by stochastic terms representing thermal or instrumental noise, but for macroscopic systems and carefully designed experiments the stochastic forces are often negligible. An aim of theory is to describe solutions of the deterministic equations that are likely to be reached starting from typical initial conditions and to persist at long times. A unified description is developed, based on the linear instabilities of a homogeneous state, which leads naturally to a classification of patterns in terms of the characteristic wave vector q0 and frequency ω0 of the instability. Type Is systems (ω0=0, q0≠0) are stationary in time and periodic in space; type IIIo systems (ω0≠0, q0=0) are periodic in time and uniform in space; and type Io systems (ω0≠0, q0≠0) are periodic in both space and time. Near a continuous (or supercritical) instability, the dynamics may be accurately described via "amplitude equations," whose form is universal for each type of instability. The specifics of each system enter only through the nonuniversal coefficients. Far from the instability threshold a different universal description known as the "phase equation" may be derived, but it is restricted to slow distortions of an ideal pattern. For many systems appropriate starting equations are either not known or too complicated to analyze conveniently. It is thus useful to introduce phenomenological order-parameter models, which lead to the correct amplitude equations near threshold, and which may be solved analytically or numerically in the nonlinear regime away from the instability. The above theoretical methods are useful in analyzing "real pattern effects" such as the influence of external boundaries, or the formation and dynamics of defects in ideal structures. An important element in nonequilibrium systems is the appearance of deterministic chaos. A greal deal is known about systems with a small number of degrees of freedom displaying "temporal chaos," where the structure of the phase space can be analyzed in detail. For spatially extended systems with many degrees of freedom, on the other hand, one is dealing with spatiotemporal chaos and appropriate methods of analysis need to be developed. In addition to the general features of nonequilibrium pattern formation discussed above, detailed reviews of theoretical and experimental work on many specific systems are presented. These include Rayleigh-Benard convection in a pure fluid, convection in binary-fluid mixtures, electrohydrodynamic convection in nematic liquid crystals, Taylor-Couette flow between rotating cylinders, parametric surface waves, patterns in certain open flow systems, oscillatory chemical reactions, static and dynamic patterns in biological media, crystallization fronts, and patterns in nonlinear optics. A concluding section summarizes what has and has not been accomplished, and attempts to assess the prospects for the future.
6,145 citations
01 Aug 2000
TL;DR: This paper explores and evaluates the use of directed diffusion for a simple remote-surveillance sensor network and its implications for sensing, communication and computation.
Abstract: Advances in processor, memory and radio technology will enable small and cheap nodes capable of sensing, communication and computation. Networks of such nodes can coordinate to perform distributed sensing of environmental phenomena. In this paper, we explore the directed diffusion paradigm for such coordination. Directed diffusion is datacentric in that all communication is for named data. All nodes in a directed diffusion-based network are application-aware. This enables diffusion to achieve energy savings by selecting empirically good paths and by caching and processing data in-network. We explore and evaluate the use of directed diffusion for a simple remote-surveillance sensor network.
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TL;DR: The second volume in a series on terrestrial and marine comparisons focusing on the temporal complement of the earlier spatial analysis of patchiness and pattern was published by Levin et al..
Abstract: This book is the second of two volumes in a series on terrestrial and marine comparisons, focusing on the temporal complement of the earlier spatial analysis of patchiness and pattern (Levin et al. 1993). The issue of the relationships among pattern, scale, and patchiness has been framed forcefully in John Steele’s writings of two decades (e.g., Steele 1978). There is no pattern without an observational frame. In the words of Nietzsche, “There are no facts… only interpretations.”
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TL;DR: Work in different scientific fields is now suggesting the existence of generic early-warning signals that may indicate for a wide class of systems if a critical threshold is approaching.
Abstract: Complex dynamical systems, ranging from ecosystems to financial markets and the climate, can have tipping points at which a sudden shift to a contrasting dynamical regime may occur. Although predicting such critical points before they are reached is extremely difficult, work in different scientific fields is now suggesting the existence of generic early-warning signals that may indicate for a wide class of systems if a critical threshold is approaching.
3,450 citations
TL;DR: A mathematical formulation of the general interaction of amoebae, as mediated by acrasin is presented, and a detailed analysis of the aggregation process is provided.
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Cites background from "The Chemical Basis of Morphogenesis..."
...Instabilities of the kind which we shall consider have recently received some attention as possible mechanisms for certain kinds of structure or pattern formation in the biological world (Turing, 1952; Prigogine & Nicolis, 1967)....
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