Lipids and membrane-associated proteins in autophagy
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Protein & Cell
REVIEW Lipids and membrane-associated proteins in autophagy Linsen Li1,2, Mindan Tong2, Yuhui Fu2, Fang Chen2, Shen Zhang2, Hanmo Chen2, Xi Ma1, Defa Li1&, Xiaoxia Liu2&, Qing Zhong2& State Key Laboratory of Animal Nutrition, Ministry of Agriculture Feed Industry Centre, College of Animal Science and Technology, China Agricultural University, Beijing 100193, China 2 Key Laboratory of Cell Differentiation and Apoptosis of Chinese Ministry of Education, Department of Pathophysiology, Shanghai Jiao Tong University School of Medicine, Shanghai 200025, China & Correspondence: [email protected] (D. Li), [email protected] (X. Liu), [email protected] (Q. Zhong) Received July 2, 2020 Accepted August 7, 2020
ABSTRACT Autophagy is essential for the maintenance of cellular homeostasis and its dysfunction has been linked to various diseases. Autophagy is a membrane driven process and tightly regulated by membrane-associated proteins. Here, we summarized membrane lipid composition, and membrane-associated proteins relevant to autophagy from a spatiotemporal perspective. In particular, we focused on three important membrane remodeling processes in autophagy, lipid transfer for phagophore elongation, membrane scission for phagophore closure, and autophagosome-lysosome membrane fusion. We discussed the significance of the discoveries in this field and possible avenues to follow for future studies. Finally, we summarized the membrane-associated biochemical techniques and assays used to study membrane properties, with a discussion of their applications in autophagy.
KEYWORDS autophagy, membrane-associated proteins, membrane-associated biochemistry assays, ATG2, ESCRT, lipid transfer, elongation, scission, fusion INTRODUCTION Autophagy is a process by which cells break down and recycle their components. Autophagy occurs at a basal level in all cells, but it can be upregulated during stress, starvation, or infection. Autophagy is essential for the maintenance of cellular homeostasis and its dysfunction has been linked to various diseases, including cancer, neurodegeneration, and immune diseases (Levine and Kroemer, 2019). Autophagy is © The Author(s) 2020
an intracellular degradation system that delivers cytoplasmic materials to the lysosome via the double-membraned autophagosome, which is highly conserved in the evolution of eukaryotes and includes the following steps: 1) autophagy initiation (signals activating autophagy) and nucleation of the phagophore/isolation membrane (IM); 2) phagophore elongation; 3) closure to form the autophagosome; 4) fusion between autophagosome and lysosome; 5) degradation of substrates in autolysosomes (Dikic and Elazar, 2018) (Fig. 1). Extensive studies have focused on the molecular mechanisms behind these processes, and our knowledge of them is constantly updated with discoveries. So far, there are many works reviewing autophagy from various angles. In the past four years, there are review papers covering topics including functions of autophagy in disease (Dikic and Elazar, 2018;
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