Thermal Biophysics of Membranes (Tutorials in Biophysics)

Thermal Biophysics of Membranes......Page 5
Contents......Page 7
Preface......Page 15
1.1 Overton (1895)......Page 19
1.3 Danielli and Davson (1935)......Page 20
1.4 Robertson (1958)......Page 22
1.5 The Fluid Mosaic Model of Singer and Nicolson (1972)......Page 23
1.6 The Mattress Model by Mouritsen and Bloom (1984)......Page 25
1.7 Domain Formation and Protein Clusters......Page 26
1.8 Perspectives of this Book......Page 28
1.9 Summary: Key Ideas of Chapter 1......Page 31
2.1 Lipid Membrane Structure......Page 33
2.2 X-Ray Diffraction......Page 36
2.3.1 Inverse Hexagonal Phase (H(II)-Phase)......Page 41
2.3.2 Cubic Phases......Page 42
2.3.3 Sponge Phases......Page 43
2.4 Summary: Key Ideas of Chapter 2......Page 45
3.1 Composition of Membranes......Page 47
3.2 Head Group Composition......Page 48
3.3 Hydrocarbon Chain Composition......Page 51
3.4 Asymmetry Across Membranes......Page 52
3.5 Dependence of Lipid Composition on Growth Temperature......Page 53
3.6 Dependence of Lipid Composition on Pressure......Page 55
3.7 Dependence of Lipid Composition on Changes in Other Thermodynamic Variables......Page 57
3.8 Summary: Key Ideas of Chapter 3......Page 58
4.1 Functions of State......Page 59
4.2 First Law of Thermodynamics......Page 60
4.3 Second Law of Thermodynamics......Page 62
4.4 Other Functions of State......Page 63
4.5 The Chemical Potential......Page 65
4.7 Chemical Equilibrium in Solutions......Page 66
4.8 Statistical Interpretation of Entropy......Page 68
4.9 Statistical Averages......Page 70
4.10.1 Heat Capacity......Page 71
4.10.2 Isothermal Volume and Area Compressibility......Page 72
4.11 Maxwell Relations......Page 73
4.12 Adiabatic Compressibility......Page 74
4.13 Thermodynamic Forces and Fluxes......Page 76
4.14 Summary: Key Ideas of Chapter 4......Page 78
5 Water......Page 81
5.1 The Electrostatic Potential......Page 82
5.2 The Electrostatic Potential in Electrolytes......Page 83
5.3 The Hydrophobic Effect......Page 84
5.3.1 Temperature Dependence of the Hydrophobic Effect......Page 85
5.4 The Wimley–White Hydrophobicity Scale......Page 88
5.5 Hydrophobic Matching......Page 89
5.6 Hofmeister Series......Page 90
5.7 Summary: Key Ideas of Chapter 5......Page 92
6.1.1 Calorimetry and Heat Capacity......Page 93
6.1.2 Melting of Membranes Consisting of One Lipid Species......Page 96
6.1.3 Some Simple Considerations on the Melting Enthalpies and Entropies......Page 101
6.2 Cooperativity and Cooperative Unit Size......Page 103
6.3 Influence of Pressure......Page 105
6.4 Metastable States......Page 108
6.5 Melting of Membranes Consisting of Lipid Mixtures......Page 109
6.6 Melting in Biological Membranes......Page 110
6.7 Lipid Monolayers......Page 111
6.8 Summary: Key Ideas of Chapter 6......Page 115
7.1 Ideal Mixture......Page 117
7.2 On the Number of Coexisting Phases......Page 122
7.2.1 Gibbs’ Phase Rule......Page 123
7.2.2 The Role of Water as a Component......Page 124
7.2.3 What Exactly is a Phase?......Page 125
7.3 Regular Solution......Page 127
7.3.1 Simple Eutectic Phase Diagram......Page 128
7.3.2 Melting Point Depression and the Effect of Anesthetics......Page 129
7.3.3 Regular Solution Theory......Page 131
7.4.1 Mixtures of Phospholipids......Page 134
7.4.2 Mixtures of Phospholipids with Other Lipids......Page 136
7.4.3 Ternary Phase Diagrams: Mixtures of Three Lipids......Page 138
7.5 Conclusions......Page 139
7.6 Summary: Key Ideas of Chapter 7......Page 140
8 Statistical Models for Lipid Melting......Page 141
8.1.1 Simple Monte Carlo Procedure......Page 142
8.1.2 Metropolis Algorithm......Page 143
8.2 Magnitude of Fluctuations......Page 144
8.3 Simple Statistical Thermodynamics Model......Page 146
8.4 Monte Carlo Simulations......Page 147
8.5 Derivation of the Partition Function for a Known Distribution of all States: The Ferrenberg–Swendsen Method......Page 150
8.6 Two-Component Membranes......Page 153
8.7 Local Fluctuations at Domain Boundaries......Page 155
8.8 The 10-State Pink Model......Page 156
8.9 Molecular Dynamics......Page 157
8.10 Summary: Key Ideas of Chapter 8......Page 158
9.1 Hydrophobic Matching......Page 159
9.2.1 Ising Model for the Interaction with Integral Proteins......Page 161
9.2.2 Lipid Fluctuations at Protein Interfaces......Page 168
9.2.3 Pore Formation......Page 169
9.3.1 Ising Model for the Interaction with Peripheral Proteins......Page 170
9.3.2 Monte Carlo Simulations on the Binding of Proteins to Membranes......Page 173
9.4 Action of Phospholipases on Membrane Domains......Page 175
9.5 Domains and “Rafts” in Biological Membranes......Page 176
9.6 Summary: Key Ideas of Chapter 9......Page 180
10 Diffusion......Page 183
10.1 Percolation......Page 184
10.2 Diffusion Models......Page 186
10.3 Diffusion of Lipids and Proteins......Page 187
10.4 Summary: Key Ideas of Chapter 10......Page 190
11.1 Diffuse Double Layer—Gouy–Chapman Theory......Page 191
11.2 Potential and Free Energy of Membranes......Page 198
11.3 Influence of Electrostatics on Melting Temperatures of Membranes......Page 199
11.4 Titration of Charged Lipid Membranes with Protons......Page 201
11.5 Binding of Charged Proteins......Page 202
11.6 Lateral Pressure Induced by Charges......Page 203
11.7 Summary: Key Ideas of Chapter 11......Page 205
12.1 The Langmuir Isotherm......Page 207
12.2 The Adsorption to a Continuous Surface......Page 210
12.2.1 The Protein Adsorbate as a Two-Dimensional Gas......Page 211
12.2.2 The van der Waals Adsorption Isotherm......Page 212
12.2.3 Scaled Particle Theory......Page 213
12.3 Aggregation Equilibria of Adsorbed Proteins......Page 214
12.4 Binding of Asymmetric Proteins......Page 215
12.5 Binding in the Presence of Electrostatic Interactions......Page 217
12.6 Lateral Pressure Changes Induced by Protein Binding......Page 219
12.7 Protein Insertion and Pore Formation......Page 220
12.7.1 Insertion Triggered by the Binding of Secondary Proteins......Page 223
12.8 Binding to Mixed Lipid Membranes......Page 225
12.9 Summary: Key Ideas of Chapter 12......Page 228
13.1 Liquid Crystalline Phases......Page 229
13.2 Elastic Theory of Incompressible Liquid Crystalline Phases......Page 230
13.2.1 Using Symmetries......Page 232
13.2.3 Achiral and Apolar Molecules......Page 233
13.2.4 Spontaneous Curvature......Page 234
13.3.1 Membranes Without Twist......Page 235
13.3.2 Radii of Curvature......Page 236
13.3.3 Topology of Vesicles and Saddle Points......Page 238
13.3.5 Fusion Pores......Page 239
13.4 Summary: Key Ideas of Chapter 13......Page 241
14 Thermodynamics of the Elastic Constants......Page 243
14.1 Heat Capacity......Page 244
14.2.1 Volume Compressibility......Page 245
14.2.2 Area Compressibility......Page 246
14.3 The Coupling Between Area Compressibility and Curvature Elasticity......Page 248
14.4 The Temperature Dependence of the Elastic Constants......Page 249
14.4.1 Proportionality Between Enthalpy and Volume Changes......Page 250
14.4.2 Proportionality Between Enthalpy and Volume Changes by Pressure Calorimetry......Page 252
14.4.5 Proportionality Between Isothermal Compressibility, Bending Elasticity and Heat Capacity Changes......Page 255
14.4.6 The Relations Between Heat Capacity and the Elastic Constants......Page 256
14.4.7 Proportionality Between Bending Elasticity and Heat Capacity......Page 257
14.5 Adiabatic Volume Compressibility......Page 259
14.6 Sound Propagation in Vesicle Dispersions......Page 261
14.7.1 Curvature Fluctuations......Page 263
14.7.2 Critical Swelling of Multilayered Membranes......Page 264
14.8 Local Fluctuations at Domain Interfaces......Page 266
14.9 Summary: Key Ideas of Chapter 14......Page 268
15.1 Coupling of Curvature and Domain Distribution......Page 269
15.2 Secretion, Endo- and Exocytosis in the Chain Melting Regime......Page 270
15.3 Curvature and the Broadening of the Melting Transition......Page 271
15.4 Structural Transitions of Vesicles in the Melting Regime......Page 273
15.5 Charged Lipid Membranes......Page 276
15.5.1 DMPG......Page 277
15.5.2 Geometry Changes Introduced by Charged Drugs......Page 278
15.6 The Ripple Phase......Page 280
15.6.1 Geometrical Considerations......Page 282
15.6.2 Modeling the Pretransition......Page 284
15.7 Peculiarities in the Melting of Zwitterionic Lipids......Page 288
15.7.1 Short Chains......Page 289
15.7.2 The Sub-Main Transition......Page 290
15.8 Summary: Key Ideas of Chapter 15......Page 293
16 Relaxation Processes in Membranes......Page 295
16.1 Thermodynamic Forces and Fluxes and Their Relation to Relaxation......Page 296
16.2 Relaxation Times of Domain Formation Processes......Page 298
16.2.1 Coupling Between Relaxation Times and Excess Heat Capacity......Page 299
16.2.2 Relaxation Experiments......Page 302
16.2.3 Relaxation Times of Biomembranes......Page 303
16.3 Summary: Key Ideas of Chapter 16......Page 306
17.1 Permeability of Lipid Membranes in the Melting Transition......Page 307
17.2 Lipid Pores......Page 309
17.3 Quantized Currents in Pure Lipid Membranes and Their Dependence on Thermodynamic Variables......Page 310
17.3.1 Temperature......Page 311
17.3.2 Calcium and Protons......Page 312
17.3.3 Lateral Pressure and Mechanosensitivity......Page 314
17.4 The Coupling of Lipid Phase Behavior and Ion Channels Proteins......Page 315
17.5 Summary: Key Ideas of Chapter 17......Page 318
18.1 The Hodgkin–Huxley Model......Page 319
18.2 Thermodynamics of the Nerve Pulse......Page 323
18.2.1 Heat Changes During the Action Potential......Page 324
18.2.2 Mechanical Changes During the Action Potential......Page 327
18.2.3 Are There Phase Transitions During the Action Potential?......Page 328
18.3 Isentropic Pulse Propagation......Page 329
18.3.1 Solitons......Page 330
18.3.2 Energy of the Solitons......Page 334
18.3.3 Thickness Changes......Page 336
18.4 Consequences of the Isentropic Theory......Page 337
18.5 Summary: Key Ideas of Chapter 18......Page 339
19 Anesthesia......Page 341
19.1 The Meyer–Overton Rule......Page 342
19.2 The Effect of Anesthetics on the Lipid Melting Points......Page 345
19.3 The Lateral Pressure Profile......Page 348
19.4 Dependence of Anesthesia on Hydrostatic Pressure......Page 350
19.5 pH Dependence of Anesthesia......Page 351
19.6 Neurotransmitters......Page 352
19.7 Summary: Key Ideas of Chapter 19......Page 353
Appendix A Abbreviations Used in this Book......Page 355
References......Page 357
Index......Page 371
Acknowledgments......Page 381