Chloroplast thylakoid membranes contain virtually all components of the energy-converting photosynthetic machinery. Their energized state, driving ATP synthesis, is enabled by the bilayer organization of the membrane. However, their most abundant lipid species is a non-bilayer-forming lipid, monogalactosyl-diacylglycerol; the role of lipid polymorphism in these membranes is poorly understood. Earlier 31P-NMR experiments revealed the coexistence of a bilayer and a non-bilayer, isotropic lipid phase in spinach thylakoids. Packing of lipid molecules, tested by fluorescence spectroscopy of the lipophilic dye, merocyanine-540 (MC540), also displayed heterogeneity. Now, our 31P-NMR experiments on spinach thylakoids uncover the presence of a bilayer and three non-bilayer lipid phases; time-resolved fluorescence spectroscopy of MC540 also reveals the presence of multiple lipidic environments. It is also shown by 31P-NMR that: (i) some lipid phases are sensitive to the osmolarity and ionic strength of the medium, (ii) a lipid phase can be modulated by catalytic hydrogenation of fatty acids and (iii) a marked increase of one of the non-bilayer phases upon lowering the pH of the medium is observed. These data provide additional experimental evidence for the polymorphism of lipid phases in thylakoids and suggest that non-bilayer phases play an active role in the structural dynamics of thylakoid membranes.

Department of Physics Faculty of Science University of Ostrava Chittussiho 10 CZ 710 00 Ostrava Czech Republic

EN FIST Center of Excellence Trg OF 13 Ljubljana Slovenia

Faculty of Chemistry and Chemical Technology Večna pot 113 Ljubljana Slovenia

Institute of Biochemistry Biological Research Centre Hungarian Academy of Sciences Temesvári körút 62 H 6726 Szeged Hungary

Institute of Plant Biology Biological Research Centre Hungarian Academy of Sciences Temesvári körút 62 H 6726 Szeged Hungary

Laboratory of BioNano Technology Wageningen University and Research Bornse Weilanden 9 6708 WG Wageningen The Netherlands

Laboratory of Biophysics Wageningen University and Research Stippeneng 4 6708 WE Wageningen The Netherlands

MAGNEFY Magnetic Resonance Research Facility Wageningen University and Research Stippeneng 4 6708 WE Wageningen The Netherlands

MicroSpectroscopy Centre Wageningen University and Research Stippeneng 4 6708 WE Wageningen The Netherlands

Slovenian NMR Center National Institute of Chemistry Hajdrihova 19 Ljubljana Slovenia

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Role of isotropic lipid phase in the fusion of photosystem II membranes

Editorial

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Modulation of non-bilayer lipid phases and the structure and functions of thylakoid membranes: effects on the water-soluble enzyme violaxanthin de-epoxidase