This text is designed for a first course in biological mass transport, and the material in it is presented at a level that is appropriate to advanced undergraduates or early graduate level students. Its orientation is somewhat more physical and mathematical than a biology or standard physiology text, reflecting its origins in a transport course that I teach to undergraduate (and occasional graduate) biomedical engineering students in the Whiting School of Engineering at Johns Hopkins. The audience for my cours- and presumably for this text - also includes chemical engineering undergraduates concentrating in biotechnology, and graduate students in biophysics. The organization of this book differs from most texts that at tempt to present an engineering approach to biological transport. What distinguishes biological transport from other mass transfer processes is the fact that biological transport is biological. Thus, we do not start with the engineering principles of mass transport (which are well presented elsewhere) and then seek biological ap plications of these principles; rather, we begin with the biological processes themselves, and then develop the tools that are needed to describe them. As a result, more physiology is presented in this text than is often found in books dealing with engineering applica tions in the life sciences.
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Zielgruppe
Für höhere Schule und Studium
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ISBN-13
978-3-540-16370-1 (9783540163701)
DOI
10.1007/978-3-662-02467-6
Schweitzer Klassifikation
1 Equilibrium Thermodynamics.- 1.1 Chemical Potentials and Activities.- Thermodynamic Preliminaries; The Electrochemical Potential.- The Interphase Equilibrium Condition.- Electrochemical Potentials in Terms of Measurable Solution Variables: Introduction, and the Effect of Electrostatic Potential.- The Gibbs-Duhem Equation.- Dependence of the Chemical Potential on Pressure.- Dependence of the Chemical Potential on Composition.- Units.- Activity and Activity Coefficient.- 1.2 Ion Equilibrium Across Membranes.- The Nernst Equilibrium.- Origin of the Nernst Potential.- Specific Ion Electrodes.- Activity Coefficient Considerations.- The Donnan Equilibrium.- 1.3 Chemical Equilibrium.- 2 Free Diffusion.- 2.1 Free Diffusion of Nonelectrolytes.- The Teorell Equation.- Integration of the Teorell Equation; Fick's First Law.- Unstirred Layers.- A Few Remarks About Solute Permeability.- Applications of Solution Theory.- Fick's Second Law and Convective Diffusion.- 2.2 Free Diffusion of Electrolytes.- Differences Between Electrolyte and Nonelectrolyte Diffusion.- The Electrodiffusion Equation.- Integration of the Electrodiffusion Equation.- Some Special Cases.- Charged Membranes.- 3 Facilitated Diffusion.- 3.1 Mechanisms of Channels and Carriers.- Hallmarks of Facilitated Transport.- Ion Selectivity of Channels.- Energetics of Ion Selectivity, and Steric Effects.- Ion Selectivity of Channels: Summary.- The Structure of Ion Channels: Filters, Gates, and Energy Profiles.- Regulation of the Gating Process.- Classification of Channels.- Channel Transport of Anions and Divalent Cations.- Some Carrier Models.- Carriers and Channels: Convergences and Differences.- 3.2 Kinetics of Facilitated Transport.- The Simplest Carrier: Assumptions.- Physical Significance of the Rate Constants and Carrier Concentrations.- The Simplest Carrier: Equations.- More Complex Carrier Models.- Energy Barrier Models of Channel Transport.- Membrane Noise Analysis.- 3.3 Inhibition of Carrier Transport.- Competitive Inhibition.- Countertransport: The Other Side of Competitive Inhibition.- Noncompetitive Inhibition.- 4 Active Transport.- 4.1 Active Transport: General Considerations.- Metabolic Coupling and Affinity.- Metabolism in Brief.- Classification of Active Transport Processes.- Identification of Active Transport Processes.- 4.2 Mechanisms of Active Transport.- Scalar Active Transport.- Primary Scalar Transport.- Secondary Scalar Transport.- Vectorial Coupling and the Curie Theorem.- Mechanisms of Vectorial Active Transport; Substrate Activation.- Models of Sodium-Potassium Exchange.- Selectivity and Other Carrier Properties.- Endocytosis.- 4.3 Kinetics of Active Transport.- A Simple Secondary Scalar Transport Model: Assumptions.- A Simple Secondary Scalar Transport Model: Equations.- More Complex Symport Models.- Primary Scalar Transport.- Flux Equations for Primary Scalar Transport.- Relation Between the Coupling Parameter ? and the Affinity of the Metabolic Reaction.- Vectorial Active Transport.- Pumps and Leaks: Introduction.- Slippage.- Shunts.- 5 Nonequilibrium Thermodynamics.- 5.1 The Basic Phenomenological Equations.- Conjugate Forces and Fluxes.- Phenomenological Coefficients and Linear Thermodynamics.- Frictional Interpretation of the Phenomenological Equations.- A Cautionary Note Before Proceeding.- 5.2 Nonequilibrium Thermodynamic Description of Passive Transport.- Setting the Stage.- The Chemical Potential of the Solvent.- A New Set of Forces and Fluxes; Osmotic Pressure.- The Kedem-Katchalsky Equations.- Physical Significance of the Reflection Coefficient.- Osmotic Pressure of Electrolyte Solutions; Donnan Osmotic Pressure.- Passive Transport of Multiple Nonelectrolytes.- Passive Transport of Electrolytes; Electrokinetic Phenomena.- 5.3 Nonequilibrium Thermodynamic Description of Active Transport.- Definition of Active Transport.- Coupling Between Nonconjugate Forces and Fluxes.- 5.4 Limitations of Nonequilibrium Thermodynamics.- Closeness to Equilibrium: A Limitation Intrinsic to Linearized Nonequilibrium Thermodynamics.- The Concentration Dependence of the Phenomenological Coefficients.- Closeness to Equilibrium in Biological Systems.- The Information Content of Nonequilibrium Thermodynamics.- Approximations in the Derivation of the Kedem-Katchalsky Equations.- 6 Models of Transport Across Cell Membranes.- 6.1 Composition and Structure of Cell Membranes.- Heterogeneity of Cell Membranes.- The Mosaic Model of Cell Membranes.- 6.2 Transport Across the Lipid Bilayer of Cell Membranes.- Evidence for Nonelectrolyte Diffusion Across the Lipid Bilayer.- A Simple Model of Transbilayer Diffusion.- Potential Barriers in the Bilayer (and Channels Too).- 6.3 Models of Transport Through Pores.- Classification of Pore Transport Models.- Hydraulic Conductivity of a Pore.- Solute Permeability as a Probe of Pore Radius.- Other Factors Affecting Estimated Pore Size; The Equivalent Pore.- The Reflection Coefficient as a Probe of Pore Radius.- Single-File Transport Through Pores.- The Permeability Ratio of Larger Pores.- 6.4 Electrical Analogs.- Equivalent Circuit for the Passive Flux of a Single Ion.- Equivalent Circuit for Multiple Ions.- The Electrical Analog of a Rheogenic Pump.- Some Final Remarks.- 7 Single Cells.- 7.1 Erythrocytes.- Monosaccharide Transport.- The Michaelis-Menten Equation.- Sodium-Potassium Exchange.- The Red Cell Calcium Pump.- Anion Transport: Exchange Diffusion.- Anion Transport: The Red Cell Transporter.- 7.2 Nerve.- The Resting Neuron.- The Action Potential.- The Membrane Action Potential.- Hodgkin and Huxley's Equations for the Dependence of Conductance on Membrane Potential.- Excitation of the Membrane Action Potential.- The Propagating Action Potential: Cable Theory.- The Role of Myelin.- A Channel Model to Explain the Dependence of Conductance on Membrane Potential.- More About Nerve Membrane Channels.- Synaptic Transmission.- Neurotransmitters and Neuromodulators.- 7.3 Muscle.- The Resting Muscle Fiber.- Excitation.- Excitation-Contraction Coupling.- The Association-Induction Hypothesis.- 8 Epithelial Transport.- 8.1 Organization of Epithelial Tissue and Some Consequences.- Epithelial Structure; Leaky and Tight Epithelia.- The Shunt Pathway.- The Transcellular Pathway.- Feedback Between the Pump Rate and the Conductance of the Apical Sodium Channel: The Role of Calcium.- Transport in a Parallel Path System.- The Solute Flux Error for a Membrane Containing Multiple Parallel Paths: Membrane Heteroreflectivity.- Coupling of Transepithelial Water Flow to Active Ion Transport: The Curran Model.- The Standing Gradient Model of Water Transport by Epithelia.- Effect of Unstirred Layers on Transepithelial Diffusion and Osmosis.- Electrical Analogs of Epithelia.- 8.2 Example Epithelia.- Transport in the Small Intestine.- Intestinal Absorption of Sugars.- Transport in the Kidney.- Glomerular Filtration.- The Proximal Tubule.- The Loop of Henle.- The Distal Tubule.- The Collecting Tubule.- 8.3 Regulation of Epithelial Transport.- The Cyclic-AMP System.- The Aldosterone Mechanism.- 9 Gas Transport.- 9.1 Overview of the Gas Transport Process.- Partial Pressure and the Equations for Gas Flux.- Oxygen Consumption.- Oxygen Transport in the Blood.- Transport Processes in the Capillaries.- 9.2 Models of Capillary Perfusion.- The Krogh Tissue Cylinder.- Modifications to the Basic Krogh Model.- Deficiencies of the Krogh Cylinder; Some Other Models of the Oxygenation Process.- References.