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_Biological Physics of the Developing Embryo_ _During development, cells and tissues undergo dynamic changes in pattern and form that employ a wider range of physical mechanisms than at any other time during an organism’s life. Biological Physics of the Developing Embryo presents a framework within which physics can be used to analyze these biological phenomena._ _Written to be accessible to both biologists and physicists, major stages and components of biological development are introduced and then analyzed from the viewpoint of physics. The presentation of physical models requires no mathematics beyond basic calculus. Physical concepts introduced include diffusion, viscosity and elasticity, adhesion, dynamical systems, electrical potential, percolation, fractals, reaction–diffusion systems, and cellular automata._ _With full-color figures throughout, this comprehensive textbook teaches biophysics by application to developmental biology and is suitable for graduate and upper-undergraduate courses in physics and biology._ _Gábor Forgács is George H. Vineyard Professor of Biological Physics at the University of Missouri, Columbia. He received his Ph.D. in condensed matter physics from Roland Eötvös University in Budapest. He made contributions to the physics of phase transitions, surface and interfacial phenomena, and statistical mechanics before moving to biological physics, where he has studied the biomechanical properties of living materials and has modeled early developmental phenomena. His recent research on constructing models of living structures of prescribed geometry using automated printing technology has been the topic of numerous articles in the international press._ _Professor Forgács has held positions at the Central Research Institute for Physics, Budapest, at the French Atomic Energy Agency, Saclay, and at Clarkson University, Potsdam. He has been a Fulbright Fellow at the Institute of Biophysics of the Budapest Medical University and has organized several meetings on the frontiers between physics and biology at the Les Houches Center for Physics. He has also served as advisor to several federal agencies of the USA on the promotion of interdisciplinary research, in particular at the interface of physics and biology. He is a member of a number of professional associations such as The Biophysical Society, The American Society for Cell Biology, and The American Physical Society._ _Stuart A. Newman is Professor of Cell Biology and Anatomy at New York Medical College, Valhalla, New York. He received an A.B. from Columbia University and a Ph.D. in Chemical Physics from the University of Chicago. He has contributed to several scientific fields including developmental pattern formation and morphogenesis, cell differentiation, the theory of biochemical networks, protein folding and assembly, and mechanisms of morphological evolution. He has also written on the philosophy, cultural background, and social implications of biological research._ _Professor Newman has been an INSERM Fellow at the Pasteur Institute, Paris, and a Fogarty Senior International Fellow at Monash University, Australia. He is a co-editor (with Brian K. Hall) of _Cartilage: Molecular Aspects_ (CRC Press, 1991) and (with Gerd B. Müller) of _Origination of Organismal Form: Beyond the Gene in Developmental and Evolutionary Biology_ (MIT Press, 2003). He has testified before US Congressional committees on cloning, stem cells, and the patenting of organisms and has served as a consultant to the US National Institutes of Health on both technical and societal issues._ _Biological Physics of the Developing Embryo_ _Gábor Forgács_ _University of Missouri and_ _Stuart A. Newman_ _New York Medical College_ _Cambridge University Press, Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo_ _Published in the United States by Cambridge University Press, New York_ Www.cambridge.org © G. Forgács and S. A. Newman 2005 This publication is in copyright. Subject to statutory exception and to the provision of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published in print format 2005 ISBN-13: 978-0-521-78337-8 (hardback) ISBN-10: 0-521-78337-2 (hardback) ISBN-13: 978-0-521-78946-2 (paperback) ISBN-10: 0-521-78946-X (paperback) Cambridge University Press has no responsibility for the persistence or accuracy of URLs for external or third-party internet websites referred to in this publication and does not guarantee that any content on such websites is, or will remain, accurate or appropriate. Contents Acknowledgments Introduction: Biology and physics The cell: fundamental unit of developmental systems The eukaryotic cell Diffusion Osmosis Viscosity Elasticity and viscoelasticity Perspective Cleavage and blastula formation The cell biology of early cleavage and blastula formation Physical processes in the cleaving blastula Physical models of cleavage and blastula formation Perspective Cell states: stability, oscillation, differentiation Gene expression and biochemical state How physics describes the behavior of a complex system Oscillatory processes in early development Multistability in cell-type diversification Perspective Cell adhesion, compartmentalization, and lumen formation Adhesion and differential adhesion in development The cell surface Cell adhesion: specific and nonspecific aspects The kinetics of cell adhesion Differential adhesion of embryonic tissues The physics of cell sorting Perspective Epithelial morphogenesis: gastrulation and neurulation Physical properties of epithelia Gastrulation Convergence and extension Neurulation Perspective Appendix: Linear stability analysis Mesenchymal morphogenesis Development of the neural crest The extracellular matrix: networks and phase transformations Mesenchymal condensation Perspective Pattern formation: segmentation, axes, and asymmetry Basic mechanisms of cell pattern formation Segmentation Epithelial patterning by juxtacrine signaling Mesoderm induction by diffusion gradients Reaction–diffusion systems Control of axis formation and left–right asymmetry Perspective Organogenesis Development of the cardiovascular system Fractals and their biological significance Branching morphogenesis: development of the salivary gland Vertebrate limb development Perspective Fertilization: generating one living dynamical system from two Development of the egg and sperm Interaction of the egg and sperm Propagation of calcium waves: spatiotemporal encoding of postfertilization events Surface contraction waves and the initiation of development Perspective Kaneko–Yomo model juxtacrine signaling, see signaling Kaneko–Yomo model for cell differentiation52, 69–76 community effect in isologous diversification 70 Kartagener (immotile-cilia) syndrome Keller model for cell differentiation 65–69, 254 see also autoregulatory network Kerszberg–Changeux model for neurulation 125–126, 127, 156 see also neurulation compared with Drasdo–Forgacs model 126 compared with Mittenthal–Mazo model 126 kidney tubules 134, 152, 189 kinesin, see molecular motor and protein kinetic energy kinetochore tension Lagrange multiplier lamellipodia laminin ‘‘liquid” tissues mesenchymal condensation mesenchymal-to-epithelial transformation mesenchyme phase separation in primary (of sea urchin embryo) 110 mesoblast limb trilaminar embryo 131 mesoderm extraembryonic induction of 171–173 paraxial (presomitic, PSM) somitic 162, 163, 164 metaphase metazoa as excitable media body plans of origin of microarrays microenvironment (of cells) see also extracellular matrix and glycocalyx membrane channel composition cortex energy lipid bilayer in stretching of tether membrane potential membrane resting potential mesenchymal condensation mesenchymal-to-epithelial transformation mesoderm phase separation in primary (of sea urchin embryo) 110 mesoblast limb trilaminar embryo 131 mesoderm extraembryonic induction of 171–173 paraxial (presomitic, PSM) somitic 162, 163, 164 metaphase metazoa as excitable media body plans of origin of microarrays microenvironment (of cells) see also extracellular matrix and glycocalyx membrane channel composition cortex energy lipid bilayer in stretching of tether membrane potential membrane resting potential Ключевые слова: zona pellucida, tg, spinal cord, vitelline envelope, cleavage, region, net magnetization, developing embryo, adhesion, set, kon rtot, agellar motor, chromosome, cell, vice versa, local communication, rev, differentiation, mechanically excitable, egg, molecule, parameter, 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cycle, frozen zone, ceq, gap junction, negatively charged, organism, viscous drag, blastula, adult body, biology, liquid-like property, constant, development, burning log, division, wa, change, term, mechanical property, result, dynamical, ci x, johnston, biomechanical property, equation, ha, insulating matrix, direction, formation, function, thermal energy, soc lond, eq, discussed, evolutionary transition, thermodynamic spontaneity, forgacs, petri dish, extracellular matrix, differential adhesion, thermal uctuations, dnb, interfacial tension, cf eq, concentration, layer, intracellular store, pattern formation, chapter, sperm, xenopus laevis, eq ha, nucleic acid, biological, material, nature, tight junction, tension, intermediate laments, axis, salivary globule, epithelial, number, neural tube, radially symmetric, references, analysis, neural, sierpinski carpet, mothers body, point, alcian blue, exp, sci, basal lamina, adhesive differential, morphogenesis, model, membrane, case, early, small, pattern, chemical, sea urchin, form, adherens junction, fh bh, gene, schematic representation, body, random uctuations, solgel transition, shape, neural fold, endoplasmic reticulum, cortical tension, preexisting nonuniformity, factor, time, relative frequency, diffusion, physics developing, experimental ndings, agellar rod, imaginal disc, embryonic, vertical axis, gradient, xenopus, acad, stage, viscoelastic modulus, dev, energy