CONTENTS 3
Contents
1 Introduction 6 1.1 Biomembranes . . . 6 1.1.1 Natural membranes . . . 6 1.1.2 Membrane lipids . . . 81.1.3 Self-organization and lipid structures . . . 9
1.1.4 Phases and phase transitions . . . 11
1.1.5 Mechanical properties of membranes . . . 13
1.2 Carotenoids . . . 15
1.2.1 Chemical structure . . . 15
1.2.2 Functions of carotenoids . . . 16
1.2.3 Interaction with membranes . . . 19
1.3 Motivation and aims of this thesis . . . 22
1.3.1 Motivation . . . 22
1.3.2 Objectives . . . 23
2 Materials and methods 24 2.1 Chemicals . . . 24
2.2 Preparation of samples . . . 24
2.2.1 Preparation of multilamellar liposomes . . . 24
2.2.2 Preparation of samples for AFM measurement . . . 25
2.2.3 Isolation of violaxanthin . . . 25
2.3 Dierential scanning calorimetry . . . 28
2.3.1 Principles and conguration . . . 28
2.3.2 Details of the experiments . . . 30
2.4 Atomic force microscopy . . . 31
2.4.1 Introduction . . . 31
2.4.2 Principles and conguration . . . 31
2.4.3 Interaction forces . . . 33
CONTENTS 4
2.4.5 Force spectroscopy . . . 36
2.4.6 Adhesion . . . 37
2.4.7 Elasticity . . . 38
2.4.8 Details of the experiments . . . 40
3 Results 42 3.1 Phase behaviour of liposomes . . . 42
3.1.1 Pure DPPC liposomes . . . 42
3.1.2 Liposomes enriched in β-carotene . . . 44
3.1.3 Liposomes enriched in zeaxanthin . . . 44
3.2 Structural properties of liposomes . . . 46
3.2.1 Measurements performed in air . . . 46
3.2.2 Measurements performed in water . . . 48
3.3 Temperature dependence of the adhesion forces . . . 51
3.3.1 Measurements performed in air . . . 51
3.3.2 Measurements performed in water . . . 53
3.4 Inuence of loading rate on the adhesion forces . . . 59
3.4.1 Measurements performed in air . . . 59
3.4.2 Measurements performed in water . . . 61
4 Discussion 70 4.1 Phase behaviour of liposomes . . . 70
4.2 Structural properties of liposomes . . . 71
4.3 Temperature dependence of the adhesion forces . . . 72
4.4 Inuence of loading rate on the adhesion forces . . . 75
5 Conclusions 78
Acknowledgements 81
References 82
Abstract 99