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Thermal Biophysics of Membranes

ISBN: 978-3-527-61160-7
378 pages
February 2008
Thermal Biophysics of Membranes (3527611606) cover image
An overview of recent experimental and theoretical developments in the field of the physics of membranes, including new insights from the past decade.
The author uses classical thermal physics and physical chemistry to explain our current understanding of the membrane. He looks at domain and 'raft' formation, and discusses it in the context of thermal fluctuations that express themselves in heat capacity and elastic constants. Further topics are lipid-protein interactions, protein binding, and the effect of sterols and anesthetics. Many seemingly unrelated properties of membranes are shown to be intimately intertwined, leading for instance to a coupling between membrane state, domain formation and vesicular shape. This also applies to non-equilibrium phenomena like the propagation of density pulses during nerve activity.
Also included is a discussion of the application of computer simulations on membranes.
For both students and researchers of biophysics, biochemistry, physical chemistry, and soft matter physics.
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1 Membranes—An Introduction.

1.1 Overton (1895).

1.2 Langmuir (1917) and Gorter and Grendel (1925).

1.3 Danielli and Davson (1935).

1.4 Robertson (1958).

1.5 The Fluid Mosaic Model of Singer and Nicolson (1972).

1.6 The Mattress Model by Mouritsen and Bloom (1984).

1.7 Domain Formation and Protein Clusters.

1.8 Perspectives of this Book.

1.9 Summary: Key Ideas of Chapter 1.

2 Membrane Structure.

2.1 Lipid Membrane Structure.

2.2 X-Ray Diffraction.

2.3 Nonlamellar Lipid Phases.

2.4 Summary: Key Ideas of Chapter 2.

3 The Composition of Biological Membranes.

3.1 Composition of Membranes.

3.2 Head Group Composition.

3.3 Hydrocarbon Chain Composition.

3.4 Asymmetry Across Membranes.

3.5 Dependence of Lipid Composition on Growth Temperature.

3.6 Dependence of Lipid Composition on Pressure.

3.7 Dependence of Lipid Composition on Changes in Other Thermodynamic Variables.

3.8 Summary: Key Ideas of Chapter 3.

4 Introduction Into Thermodynamics.

4.1 Functions of State.

4.2 First Law of Thermodynamics.

4.3 Second Law of Thermodynamics.

4.4 Other Functions of State.

4.5 The Chemical Potential.

4.6 The Gibbs–Duhem Equation.

4.7 Chemical Equilibrium in Solutions.

4.8 Statistical Interpretation of Entropy.

4.9 Statistical Averages.

4.10 Heat Capacity and Elastic Constants.

4.11 Maxwell Relations.

4.12 Adiabatic Compressibility.

4.13 Thermodynamic Forces and Fluxes.

4.14 Summary: Key Ideas of Chapter 4.

5 Water.

5.1 The Electrostatic Potential.

5.2 The Electrostatic Potential in Electrolytes.

5.3 The Hydrophobic Effect.

5.3.1 Temperature Dependence of the Hydrophobic Effect.

5.4 TheWimley–White Hydrophobicity Scale.

5.5 Hydrophobic Matching.

5.6 Hofmeister Series.

5.7 Summary: Key Ideas of Chapter 5.

6 Lipid Melting.

6.1 Lipid Melting.

6.2 Cooperativity and Cooperative Unit Size.

6.3 Influence of Pressure.

6.4 Metastable States.

6.5 Melting of Membranes Consisting of Lipid Mixtures.

6.6 Melting in Biological Membranes.

6.7 Lipid Monolayers.

6.8 Summary: Key Ideas of Chapter 6.

7 Phase Diagrams.

7.1 Ideal Mixture.

7.2 On the Number of Coexisting Phases.

7.3 Regular Solution.

7.4 Experimental Phase Diagrams.

7.5 Conclusions.

7.6 Summary: Key Ideas of Chapter 7.

8 Statistical Models for Lipid Melting.

8.1 Monte Carlo Simulations.

8.2 Magnitude of Fluctuations.

8.3 Simple Statistical Thermodynamics Model.

8.4 Monte Carlo Simulations.

8.5 Derivation of the Partition Function for a Known Distribution of all States: The Ferrenberg–Swendsen Method.

8.6 Two-Component Membranes.

8.7 Local Fluctuations at Domain Boundaries.

8.8 The 10-State Pink Model.

8.9 Molecular Dynamics.

8.10 Summary: Key Ideas of Chapter 8.

9 Lipid–Protein Interactions.

9.1 Hydrophobic Matching.

9.2 Integral Proteins.

9.3 Binding of Peripheral Proteins to One-ComponentMembranes.

9.4 Action of Phospholipases on Membrane Domains.

9.5 Domains and “Rafts” in Biological Membranes.

9.6 Summary: Key Ideas of Chapter 9.

10 Diffusion.

10.1 Percolation.

10.2 Diffusion Models.

10.3 Diffusion of Lipids and Proteins.

10.4 Summary: Key Ideas of Chapter 10.

11 Electrostatics.

11.1 Diffuse Double Layer—Gouy–Chapman Theory.

11.2 Potential and Free Energy of Membranes.

11.3 Influence of Electrostatics on Melting Temperatures of Membranes.

11.4 Titration of Charged Lipid Membranes with Protons.

11.5 Binding of Charged Proteins.

11.6 Lateral Pressure Induced by Charges.

11.7 Summary: Key Ideas of Chapter 11.

12 Adsorption, Binding, and Insertion of Proteins.

12.1 The Langmuir Isotherm.

12.2 The Adsorption to a Continuous Surface.

12.3 Aggregation Equilibria of Adsorbed Proteins.

12.4 Binding of Asymmetric Proteins.

12.5 Binding in the Presence of Electrostatic Interactions.

12.6 Lateral Pressure Changes Induced by Protein Binding.

12.7 Protein Insertion and Pore Formation.

12.8 Binding to Mixed Lipid Membranes.

12.9 Summary: Key Ideas of Chapter 12.

13 Elasticity and Curvature.

13.1 Liquid Crystalline Phases.

13.2 Elastic Theory of Incompressible Liquid Crystalline Phases.

13.3 Elastic Theory of Membrane Bending.

13.4 Summary: Key Ideas of Chapter 13.

14 Thermodynamics of the Elastic Constants.

14.1 Heat Capacity.

14.2 Volume and Area Compressibility.

14.3 The Coupling Between Area Compressibility and Curvature Elasticity.

14.4 The Temperature Dependence of the Elastic Constants.

14.5 Adiabatic Volume Compressibility.

14.6 Sound Propagation in Vesicle Dispersions.

14.7 Curvature Fluctuations and Critical Swelling of Multilayers.

14.8 Local Fluctuations at Domain Interfaces.

14.9 Summary: Key Ideas of Chapter 14.

15 Structural Transitions.

15.1 Coupling of Curvature and Domain Distribution.

15.2 Secretion, Endo- and Exocytosis in the Chain Melting Regime.

15.3 Curvature and the Broadening of the Melting Transition.

15.4 Structural Transitions of Vesicles in the Melting Regime.

15.5 Charged Lipid Membranes.

15.6 The Ripple Phase.

15.7 Peculiarities in the Melting of Zwitterionic Lipids.

15.8 Summary: Key Ideas of Chapter 15.

16 Relaxation Processes in Membranes.

16.1 Thermodynamic Forces and Fluxes and Their Relation to Relaxation.

16.2 Relaxation Times of Domain Formation Processes.

16.3 Summary: Key Ideas of Chapter 16.

17 Permeability.

17.1 Permeability of Lipid Membranes in the Melting Transition.

17.2 Lipid Pores.

17.3 Quantized Currents in Pure Lipid Membranes and Their Dependence on Thermodynamic Variables.

17.4 The Coupling of Lipid Phase Behavior and Ion Channels Proteins.

17.5 Summary: Key Ideas of Chapter 17.

18 Nerve Pulse Propagation.

18.1 The Hodgkin–Huxley Model.

18.2 Thermodynamics of the Nerve Pulse.

18.3 Isentropic Pulse Propagation.

18.4 Consequences of the Isentropic Theory.

18.5 Summary: Key Ideas of Chapter 18.

19 Anesthesia.

19.1 The Meyer–Overton Rule.

19.2 The Effect of Anesthetics on the Lipid Melting Points.

19.3 The Lateral Pressure Profile.

19.4 Dependence of Anesthesia on Hydrostatic Pressure.

19.5 pH Dependence of Anesthesia.

19.6 Neurotransmitters.

19.7 Summary: Key Ideas of Chapter 19.


A. Abbreviations Used in this Book.




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Thomas Heimburg received his Ph.D. in physics and his habilitation in biophysics both from the Physics Department of the University of G?ngen, Germany. He was a Heisenberg Fellow of the German Research Council (Deutsche Forschungsgemeinschaft) at the Max Planck Institute for Biophysical Chemistry in G?ngen and head of the independent research group "Membrane Biophysics & Thermodynamics". He was appointed associate professor in the Physics Department of the University of G?ngen. Now he is associate professor for biophysics at the Niels Bohr Institute of the University of Copenhagen and head of the Membrane Biophysics Group.
His primary research interests are experimental and theoretical thermodynamics and spectroscopy of artificial and biological membranes with a special focus on cooperative phenomena in biomembranes.
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