It is well established that plant thylakoid membranes (TMs), in addition to a bilayer, contain two isotropic lipid phases and an inverted hexagonal (HII) phase. To elucidate the origin of non-bilayer lipid phases, we recorded the 31P-NMR spectra of isolated spinach plastoglobuli and TMs and tested their susceptibilities to lipases and proteases; the structural and functional characteristics of TMs were monitored using biophysical techniques and CN-PAGE. Phospholipase-A1 gradually destroyed all 31P-NMR-detectable lipid phases of isolated TMs, but the weak signal of isolated plastoglobuli was not affected. Parallel with the destabilization of their lamellar phase, TMs lost their impermeability; other effects, mainly on Photosystem-II, lagged behind the destruction of the original phases. Wheat-germ lipase selectively eliminated the isotropic phases but exerted little or no effect on the structural and functional parameters of TMs-indicating that the isotropic phases are located outside the protein-rich regions and might be involved in membrane fusion. Trypsin and Proteinase K selectively suppressed the HII phase-suggesting that a large fraction of TM lipids encapsulate stroma-side proteins or polypeptides. We conclude that-in line with the Dynamic Exchange Model-the non-bilayer lipid phases of TMs are found in subdomains separated from but interconnected with the bilayer accommodating the main components of the photosynthetic machinery.

Belozersky Institute of Physico Chemical Biology M 5 Lomonosov Moscow State University 119991 Moscow Russia

Doctoral School of Biology University of Szeged 6726 Szeged Hungary

EN FIST Center of Excellence SI 1000 Ljubljana Slovenia

Faculty of Chemistry and Chemical Technology University of Ljubljana SI 1000 Ljubljana Slovenia

Group of Biophysics Department of Physics Faculty of Science University of Ostrava 710 00 Ostrava Czech Republic

Laboratory of Ecological Plant Physiology Global Change Research Institute of the Czech Academy of Sciences 603 00 Brno Czech Republic

Photosynthetic Membranes Group Institute of Plant Biology Biological Research Centre Eötvös Loránd Research Network 6726 Szeged Hungary

Slovenian NMR Center National Institute of Chemistry SI 1000 Ljubljana Slovenia

STEM Program Science Department Chaoyang KaiWen Academy Beijing 100018 China

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

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