Biogenesis of the multifunctional lipid droplet: Lipids, proteins, and sites

Journal of Cell Biology - Tập 204 Số 5 - Trang 635-646 - 2014
Albert Pol1,2,3, Steven P. Gross4, Robert G. Parton5,6,7
1Departament de Biologia Cellular, Immunologia i Neurociències, Facultat de Medicina, Universitat de Barcelona, 08036 Barcelona, Spain 2
2Equip de Compartiments Cellulars i Senyalització, Institut d’Investigacions Biomèdiques August Pi i Sunyer (IDIBAPS), 08036 Barcelona, Spain 1
3Institució Catalana de Recerca i Estudis Avançats (ICREA), 08010 Barcelona, Spain 3
4Department of Developmental and Cell Biology, University of California, Irvine, Irvine, CA 92697 4
5Centre for Microscopy and Microanalysis, The University of Queensland, Brisbane, Queensland 4072, Australia 5 and 6
6The Institute for Molecular Bioscience 5 and 6
7The Institute for Molecular Bioscience and Centre for Microscopy and Microanalysis, The University of Queensland, Brisbane, Queensland 4072, Australia

Tóm tắt

Lipid droplets (LDs) are ubiquitous dynamic organelles that store and supply lipids in all eukaryotic and some prokaryotic cells for energy metabolism, membrane synthesis, and production of essential lipid-derived molecules. Interest in the organelle’s cell biology has exponentially increased over the last decade due to the link between LDs and prevalent human diseases and the discovery of new and unexpected functions of LDs. As a result, there has been significant recent progress toward understanding where and how LDs are formed, and the specific lipid pathways that coordinate LD biogenesis.

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Tài liệu tham khảo

Adeyo, 2011, The yeast lipin orthologue Pah1p is important for biogenesis of lipid droplets, J. Cell Biol., 192, 1043, 10.1083/jcb.201010111

Anand, 2012, A novel role for lipid droplets in the organismal antibacterial response, Elife., 1, e00003, 10.7554/eLife.00003

Andersson, 2006, PLD1 and ERK2 regulate cytosolic lipid droplet formation, J. Cell Sci., 119, 2246, 10.1242/jcs.02941

Athenstaedt, 1997, Biosynthesis of phosphatidic acid in lipid particles and endoplasmic reticulum of Saccharomyces cerevisiae, J. Bacteriol., 179, 7611, 10.1128/jb.179.24.7611-7616.1997

Ayciriex, 2012, YPR139c/LOA1 encodes a novel lysophosphatidic acid acyltransferase associated with lipid droplets and involved in TAG homeostasis, Mol. Biol. Cell., 23, 233, 10.1091/mbc.E11-07-0650

Bartz, 2007, Lipidomics reveals that adiposomes store ether lipids and mediate phospholipid traffic, J. Lipid Res., 48, 837, 10.1194/jlr.M600413-JLR200

Bi, 2012, Opposite and redundant roles of the two Drosophila perilipins in lipid mobilization, J. Cell Sci., 125, 3568, 10.1242/jcs.101329

Bickel, 2009, PAT proteins, an ancient family of lipid droplet proteins that regulate cellular lipid stores, Biochim. Biophys. Acta., 1791, 419, 10.1016/j.bbalip.2009.04.002

Blanchette-Mackie, 1995, Perilipin is located on the surface layer of intracellular lipid droplets in adipocytes, J. Lipid Res., 36, 1211, 10.1016/S0022-2275(20)41129-0

Borgese, 2007, How tails guide tail-anchored proteins to their destinations, Curr. Opin. Cell Biol., 19, 368, 10.1016/j.ceb.2007.04.019

Brasaemle, 2012, Packaging of fat: an evolving model of lipid droplet assembly and expansion, J. Biol. Chem., 287, 2273, 10.1074/jbc.R111.309088

Brasaemle, 2004, Proteomic analysis of proteins associated with lipid droplets of basal and lipolytically stimulated 3T3-L1 adipocytes, J. Biol. Chem., 279, 46835, 10.1074/jbc.M409340200

Bulankina, 2009, TIP47 functions in the biogenesis of lipid droplets, J. Cell Biol., 185, 641, 10.1083/jcb.200812042

Butler, 2010, Cross-talk between remodeling and de novo pathways maintains phospholipid balance through ubiquitination, J. Biol. Chem., 285, 6246, 10.1074/jbc.M109.017350

Cartwright, 2012, Seipin: from human disease to molecular mechanism, J. Lipid Res., 53, 1042, 10.1194/jlr.R023754

Chang, 2006, Protection against fatty liver but normal adipogenesis in mice lacking adipose differentiation-related protein, Mol. Cell. Biol., 26, 1063, 10.1128/MCB.26.3.1063-1076.2006

Chapman, 2012, Biogenesis and functions of lipid droplets in plants: Thematic review series: lipid droplet synthesis and metabolism: from yeast to man, J. Lipid Res., 53, 215, 10.1194/jlr.R021436

Chen, 2002, Oleic acid-induced PKC isozyme translocation in RAW 264.7 macrophages, J. Cell. Biochem., 86, 784, 10.1002/jcb.10266

Choudhary, 2011, The topology of the triacylglycerol synthesizing enzyme Lro1 indicates that neutral lipids can be produced within the luminal compartment of the endoplasmatic reticulum: Implications for the biogenesis of lipid droplets, Commun. Integr. Biol., 4, 781, 10.4161/cib.17830

Cornell, 2000, Regulation of CTP:phosphocholine cytidylyltransferase by amphitropism and relocalization, Trends Biochem. Sci., 25, 441, 10.1016/S0968-0004(00)01625-X

Cui, 2012, Overexpression of a short human seipin/BSCL2 isoform in mouse adipose tissue results in mild lipodystrophy, Am. J. Physiol. Endocrinol. Metab., 302, E705, 10.1152/ajpendo.00237.2011

Czabany, 2008, Structural and biochemical properties of lipid particles from the yeast Saccharomyces cerevisiae, J. Biol. Chem., 283, 17065, 10.1074/jbc.M800401200

Doege, 2006, Protein-mediated fatty acid uptake: novel insights from in vivo models, Physiology (Bethesda)., 21, 259, 10.1152/physiol.00014.2006

Du, 2009, Starving neurons show sex difference in autophagy, J. Biol. Chem., 284, 2383, 10.1074/jbc.M804396200

Eastman, 2009, A role for ubiquitin ligases and Spartin/SPG20 in lipid droplet turnover, J. Cell Biol., 184, 881, 10.1083/jcb.200808041

Ellis, 2010, Acyl-coenzyme A synthetases in metabolic control, Curr. Opin. Lipidol., 21, 212, 10.1097/MOL.0b013e32833884bb

Fei, 2008, Fld1p, a functional homologue of human seipin, regulates the size of lipid droplets in yeast, J. Cell Biol., 180, 473, 10.1083/jcb.200711136

Fei, 2011, Seipin, adipogenesis and lipid droplets, Trends Endocrinol. Metab., 22, 204, 10.1016/j.tem.2011.02.004

Fei, 2011, Molecular characterization of seipin and its mutants: implications for seipin in triacylglycerol synthesis, J. Lipid Res., 52, 2136, 10.1194/jlr.M017566

Fujimoto, 2011, Not just fat: the structure and function of the lipid droplet, Cold Spring Harb. Perspect. Biol., 3, a004838, 10.1101/cshperspect.a004838

Fujimoto, 2004, Identification of major proteins in the lipid droplet-enriched fraction isolated from the human hepatocyte cell line HuH7, Biochim. Biophys. Acta., 1644, 47, 10.1016/j.bbamcr.2003.10.018

Fujimoto, 2007, Involvement of ACSL in local synthesis of neutral lipids in cytoplasmic lipid droplets in human hepatocyte HuH7, J. Lipid Res., 48, 1280, 10.1194/jlr.M700050-JLR200

Füllekrug, 2012, Outlook: membrane junctions enable the metabolic trapping of fatty acids by intracellular acyl-CoA synthetases, Front Physiol., 3, 401, 10.3389/fphys.2012.00401

Fuller, 2001, The influence of lysolipids on the spontaneous curvature and bending elasticity of phospholipid membranes, Biophys. J., 81, 243, 10.1016/S0006-3495(01)75695-0

Garbarino, 2009, Sterol and diacylglycerol acyltransferase deficiency triggers fatty acid-mediated cell death, J. Biol. Chem., 284, 30994, 10.1074/jbc.M109.050443

Gorrissen, 1980, Deuterium magnetic resonance of selectively deuterated cholesteryl esters in phosphatidylcholine vesicles, Biochemistry., 19, 3422, 10.1021/bi00556a003

Gorrissen, 1982, Deuterium magnetic resonance of triacylglycerols in phospholipid bilayers, Chem. Phys. Lipids., 31, 245, 10.1016/0009-3084(82)90060-3

Gross, 2010, Structural insights into triglyceride storage mediated by fat storage-inducing transmembrane (FIT) protein 2, PLoS ONE., 5, e10796, 10.1371/journal.pone.0010796

Gross, 2011, Direct binding of triglyceride to fat storage-inducing transmembrane proteins 1 and 2 is important for lipid droplet formation, Proc. Natl. Acad. Sci. USA., 108, 19581, 10.1073/pnas.1110817108

Gubern, 2008, Group IVA phospholipase A2 is necessary for the biogenesis of lipid droplets, J. Biol. Chem., 283, 27369, 10.1074/jbc.M800696200

Ha, 2012, Regulation of the Golgi complex by phospholipid remodeling enzymes, Biochim. Biophys. Acta., 1821, 1078, 10.1016/j.bbalip.2012.04.004

Hapala, 2011, Is fat so bad? Modulation of endoplasmic reticulum stress by lipid droplet formation, Biol. Cell., 103, 271, 10.1042/BC20100144

Harris, 2011, DGAT enzymes are required for triacylglycerol synthesis and lipid droplets in adipocytes, J. Lipid Res., 52, 657, 10.1194/jlr.M013003

Herker, 2012, Emerging role of lipid droplets in host/pathogen interactions, J. Biol. Chem., 287, 2280, 10.1074/jbc.R111.300202

Herms, 2013, Cell-to-cell heterogeneity in lipid droplets suggests a mechanism to reduce lipotoxicity, Curr. Biol., 23, 1489, 10.1016/j.cub.2013.06.032

Hinson, 2009, The antiviral protein, viperin, localizes to lipid droplets via its N-terminal amphipathic alpha-helix, Proc. Natl. Acad. Sci. USA., 106, 20452, 10.1073/pnas.0911679106

Hörl, 2011, Sequential synthesis and methylation of phosphatidylethanolamine promote lipid droplet biosynthesis and stability in tissue culture and in vivo, J. Biol. Chem., 286, 17338, 10.1074/jbc.M111.234534

Igal, 1996, Acylglycerol recycling from triacylglycerol to phospholipid, not lipase activity, is defective in neutral lipid storage disease fibroblasts, J. Biol. Chem., 271, 16644, 10.1074/jbc.271.28.16644

Ingelmo-Torres, 2009, Hydrophobic and basic domains target proteins to lipid droplets, Traffic., 10, 1785, 10.1111/j.1600-0854.2009.00994.x

Jacquier, 2011, Lipid droplets are functionally connected to the endoplasmic reticulum in Saccharomyces cerevisiae, J. Cell Sci., 124, 2424, 10.1242/jcs.076836

Jacquier, 2013, Expression of oleosin and perilipins in yeast promotes formation of lipid droplets from the endoplasmic reticulum, J. Cell Sci., 126, 5198, 10.1242/jcs.131896

Jamil, 1990, Feedback regulation of CTP:phosphocholine cytidylyltransferase translocation between cytosol and endoplasmic reticulum by phosphatidylcholine, J. Biol. Chem., 265, 4332, 10.1016/S0021-9258(19)39568-7

Kadereit, 2008, Evolutionarily conserved gene family important for fat storage, Proc. Natl. Acad. Sci. USA., 105, 94, 10.1073/pnas.0708579105

Karanasios, 2010, A phosphorylation-regulated amphipathic helix controls the membrane translocation and function of the yeast phosphatidate phosphatase, Proc. Natl. Acad. Sci. USA., 107, 17539, 10.1073/pnas.1007974107

Karanasios, 2013, Regulation of lipid droplet and membrane biogenesis by the acidic tail of the phosphatidate phosphatase Pah1p, Mol. Biol. Cell., 24, 2124, 10.1091/mbc.E13-01-0021

Kassan, 2013, Acyl-CoA synthetase 3 promotes lipid droplet biogenesis in ER microdomains, J. Cell Biol., 203, 985, 10.1083/jcb.201305142

Khandelia, 2010, Triglyceride blisters in lipid bilayers: implications for lipid droplet biogenesis and the mobile lipid signal in cancer cell membranes, PLoS ONE., 5, e12811, 10.1371/journal.pone.0012811

Khelef, 1998, Immunolocalization of acyl-coenzyme A:cholesterol O-acyltransferase in macrophages, J. Biol. Chem., 273, 11218, 10.1074/jbc.273.18.11218

Kimmel, 2010, Adoption of PERILIPIN as a unifying nomenclature for the mammalian PAT-family of intracellular lipid storage droplet proteins, J. Lipid Res., 51, 468, 10.1194/jlr.R000034

Klemm, 2011, Dual role of ancient ubiquitous protein 1 (AUP1) in lipid droplet accumulation and endoplasmic reticulum (ER) protein quality control, J. Biol. Chem., 286, 37602, 10.1074/jbc.M111.284794

Kooijman, 2005, Spontaneous curvature of phosphatidic acid and lysophosphatidic acid, Biochemistry., 44, 2097, 10.1021/bi0478502

Krahmer, 2011, Phosphatidylcholine synthesis for lipid droplet expansion is mediated by localized activation of CTP:phosphocholine cytidylyltransferase, Cell Metab., 14, 504, 10.1016/j.cmet.2011.07.013

Krahmer, 2013, Balancing the fat: lipid droplets and human disease, EMBO Mol Med., 5, 905, 10.1002/emmm.201100671

Krause, 1984, Adipose tissue and cholesterol metabolism, J. Lipid Res., 25, 97, 10.1016/S0022-2275(20)37830-5

Kuerschner, 2008, Imaging of lipid biosynthesis: how a neutral lipid enters lipid droplets, Traffic., 9, 338, 10.1111/j.1600-0854.2007.00689.x

Kühnlein, 2012, Thematic review series: Lipid droplet synthesis and metabolism: from yeast to man. Lipid droplet-based storage fat metabolism in Drosophila, J. Lipid Res., 53, 1430, 10.1194/jlr.R024299

Lacey, 1999, The accumulation of triacylglycerols within the endoplasmic reticulum of developing seeds of Helianthus annuus, Plant J., 17, 397, 10.1046/j.1365-313X.1999.00387.x

Lands, 2000, Stories about acyl chains, Biochim. Biophys. Acta., 1483, 1, 10.1016/S1388-1981(99)00177-8

Liu, 2004, Chinese hamster ovary K2 cell lipid droplets appear to be metabolic organelles involved in membrane traffic, J. Biol. Chem., 279, 3787, 10.1074/jbc.M311945200

Long, 2012, Lipid droplet de novo formation and fission are linked to the cell cycle in fission yeast, Traffic., 13, 705, 10.1111/j.1600-0854.2012.01339.x

Mak, 2012, Lipid droplets as fat storage organelles in Caenorhabditis elegans: Thematic review series: Lipid droplet synthesis and metabolism: From yeast to man, J. Lipid Res., 53, 28, 10.1194/jlr.R021006

Malhotra, 2011, PKD regulates membrane fission to generate TGN to cell surface transport carriers, Cold Spring Harb. Perspect. Biol., 3, a005280, 10.1101/cshperspect.a005280

Marchesan, 2003, A phospholipase D-dependent process forms lipid droplets containing caveolin, adipocyte differentiation-related protein, and vimentin in a cell-free system, J. Biol. Chem., 278, 27293, 10.1074/jbc.M301430200

Martin, 2005, Regulated localization of Rab18 to lipid droplets: effects of lipolytic stimulation and inhibition of lipid droplet catabolism, J. Biol. Chem., 280, 42325, 10.1074/jbc.M506651200

McMahon, 2005, Membrane curvature and mechanisms of dynamic cell membrane remodelling, Nature., 438, 590, 10.1038/nature04396

Milewska, 2009, Identification of novel spartin-interactors shows spartin is a multifunctional protein, J. Neurochem., 111, 1022, 10.1111/j.1471-4159.2009.06382.x

Miranda, 2011, Re-patterning of skeletal muscle energy metabolism by fat storage-inducing transmembrane protein 2, J. Biol. Chem., 286, 42188, 10.1074/jbc.M111.297127

Moessinger, 2011, Human lysophosphatidylcholine acyltransferases 1 and 2 are located in lipid droplets where they catalyze the formation of phosphatidylcholine, J. Biol. Chem., 286, 21330, 10.1074/jbc.M110.202424

Mountford, 1988, Organization of lipids in the plasma membranes of malignant and stimulated cells: a new model, Trends Biochem. Sci., 13, 172, 10.1016/0968-0004(88)90145-4

Murphy, 1999, Mechanisms of lipid-body formation, Trends Biochem. Sci., 24, 109, 10.1016/S0968-0004(98)01349-8

Nakamura, 2003, Adipose differentiation-related protein has two independent domains for targeting to lipid droplets, Biochem. Biophys. Res. Commun., 306, 333, 10.1016/S0006-291X(03)00979-3

Novikoff, 1980, Organelle relationships in cultured 3T3-L1 preadipocytes, J. Cell Biol., 87, 180, 10.1083/jcb.87.1.180

Ohsaki, 2008, Lipid droplets are arrested in the ER membrane by tight binding of lipidated apolipoprotein B-100, J. Cell Sci., 121, 2415, 10.1242/jcs.025452

Ohsaki, 2009, Biogenesis of cytoplasmic lipid droplets: from the lipid ester globule in the membrane to the visible structure, Biochim. Biophys. Acta., 1791, 399, 10.1016/j.bbalip.2008.10.002

Ost, 2005, Triacylglycerol is synthesized in a specific subclass of caveolae in primary adipocytes, J. Biol. Chem., 280, 5, 10.1074/jbc.C400429200

Ozeki, 2005, Rab18 localizes to lipid droplets and induces their close apposition to the endoplasmic reticulum-derived membrane, J. Cell Sci., 118, 2601, 10.1242/jcs.02401

Park, 2010, Further assembly required: construction and dynamics of the endoplasmic reticulum network, EMBO Rep., 11, 515, 10.1038/embor.2010.92

Penno, 2013, Phospholipids and lipid droplets, Biochim. Biophys. Acta., 1831, 589, 10.1016/j.bbalip.2012.12.001

Ploegh, 2007, A lipid-based model for the creation of an escape hatch from the endoplasmic reticulum, Nature., 448, 435, 10.1038/nature06004

Pol, 2004, Dynamic and regulated association of caveolin with lipid bodies: modulation of lipid body motility and function by a dominant negative mutant, Mol. Biol. Cell., 15, 99, 10.1091/mbc.E03-06-0368

Poppelreuther, 2012, The N-terminal region of acyl-CoA synthetase 3 is essential for both the localization on lipid droplets and the function in fatty acid uptake, J. Lipid Res., 53, 888, 10.1194/jlr.M024562

Rajakumari, 2010, Janus-faced enzymes yeast Tgl3p and Tgl5p catalyze lipase and acyltransferase reactions, Mol. Biol. Cell., 21, 501, 10.1091/mbc.E09-09-0775

Rajakumari, 2010, Multiple functions as lipase, steryl ester hydrolase, phospholipase, and acyltransferase of Tgl4p from the yeast Saccharomyces cerevisiae, J. Biol. Chem., 285, 15769, 10.1074/jbc.M109.076331

Rao, 1966, Purification and properties of triglyceride synthetase from the intestinal mucosa, Biochim. Biophys. Acta., 125, 465, 10.1016/0005-2760(66)90035-X

Robenek, 2006, Adipophilin-enriched domains in the ER membrane are sites of lipid droplet biogenesis, J. Cell Sci., 119, 4215, 10.1242/jcs.03191

Roingeard, 2011, The birth and life of lipid droplets: learning from the hepatitis C virus, Biol. Cell., 103, 223, 10.1042/BC20100119

Saka, 2012, Emerging roles for lipid droplets in immunity and host-pathogen interactions, Annu. Rev. Cell Dev. Biol., 28, 411, 10.1146/annurev-cellbio-092910-153958

Sandoval, 2008, Fatty acid transport and activation and the expression patterns of genes involved in fatty acid trafficking, Arch. Biochem. Biophys., 477, 363, 10.1016/j.abb.2008.06.010

Sato, 2006, Proteomic profiling of lipid droplet proteins in hepatoma cell lines expressing hepatitis C virus core protein, J. Biochem., 139, 921, 10.1093/jb/mvj104

Sembongi, 2013, Distinct roles of the phosphatidate phosphatases lipin 1 and 2 during adipogenesis and lipid droplet biogenesis in 3T3-L1 cells, J. Biol. Chem., 288, 34502, 10.1074/jbc.M113.488445

Siniossoglou, 2013, Phospholipid metabolism and nuclear function: roles of the lipin family of phosphatidic acid phosphatases, Biochim. Biophys. Acta., 1831, 575, 10.1016/j.bbalip.2012.09.014

Skinner, 2009, Diacylglycerol enrichment of endoplasmic reticulum or lipid droplets recruits perilipin 3/TIP47 during lipid storage and mobilization, J. Biol. Chem., 284, 30941, 10.1074/jbc.M109.013995

Spandl, 2011, Ancient ubiquitous protein 1 (AUP1) localizes to lipid droplets and binds the E2 ubiquitin conjugase G2 (Ube2g2) via its G2 binding region, J. Biol. Chem., 286, 5599, 10.1074/jbc.M110.190785

Stehr, 2012, Cytosolic lipid inclusions formed during infection by viral and bacterial pathogens, Microbes Infect., 14, 1227, 10.1016/j.micinf.2012.08.001

Stevanovic, 2013, Monotopic topology is required for lipid droplet targeting of ancient ubiquitous protein 1, J. Lipid Res., 54, 503, 10.1194/jlr.M033852

Stone, 2009, The endoplasmic reticulum enzyme DGAT2 is found in mitochondria-associated membranes and has a mitochondrial targeting signal that promotes its association with mitochondria, J. Biol. Chem., 284, 5352, 10.1074/jbc.M805768200

Sturley, 2012, Lipid droplet formation on opposing sides of the endoplasmic reticulum, J. Lipid Res., 53, 1800, 10.1194/jlr.R028290

Suzuki, 2011, Lipid droplets: size matters, J. Electron Microsc. (Tokyo)., 60, S101, 10.1093/jmicro/dfr016

Suzuki, 2013, Protein kinase Cη is targeted to lipid droplets, Histochem. Cell Biol., 139, 505, 10.1007/s00418-013-1083-z

Suzuki, 2005, Molecular determinants for subcellular localization of hepatitis C virus core protein, J. Virol., 79, 1271, 10.1128/JVI.79.2.1271-1281.2005

Sztalryd, 2006, Functional compensation for adipose differentiation-related protein (ADFP) by Tip47 in an ADFP null embryonic cell line, J. Biol. Chem., 281, 34341, 10.1074/jbc.M602497200

Szymanski, 2007, The lipodystrophy protein seipin is found at endoplasmic reticulum lipid droplet junctions and is important for droplet morphology, Proc. Natl. Acad. Sci. USA., 104, 20890, 10.1073/pnas.0704154104

Tauchi-Sato, 2002, The surface of lipid droplets is a phospholipid monolayer with a unique Fatty Acid composition, J. Biol. Chem., 277, 44507, 10.1074/jbc.M207712200

Tomoda, 1987, Inhibition of acyl-CoA synthetase by triacsins, Biochim. Biophys. Acta., 921, 595, 10.1016/0005-2760(87)90088-9

Vallet-Pichard, 2012, Nonalcoholic fatty liver disease and HIV infection, Semin. Liver Dis., 32, 158, 10.1055/s-0032-1316471

van den Bosch, 1997, Alkyl-dihydroxyacetonephosphate synthase, Biochim. Biophys. Acta., 1348, 35, 10.1016/S0005-2760(97)00107-0

Vance, 2009, Physiological consequences of disruption of mammalian phospholipid biosynthetic genes, J. Lipid Res., 50, S132, 10.1194/jlr.R800048-JLR200

Vance, 2004, Phospholipid biosynthesis in mammalian cells, Biochem. Cell Biol., 82, 113, 10.1139/o03-073

Walther, 2009, The life of lipid droplets, Biochim. Biophys. Acta., 1791, 459, 10.1016/j.bbalip.2008.10.009

Walther, 2012, Lipid droplets and cellular lipid metabolism, Annu. Rev. Biochem., 81, 687, 10.1146/annurev-biochem-061009-102430

Wang, 2012, The ubiquitin-like (UBX)-domain-containing protein Ubx2/Ubxd8 regulates lipid droplet homeostasis, J. Cell Sci., 125, 2930, 10.1242/jcs.100230

Wang, 2011, Lipin-1γ isoform is a novel lipid droplet-associated protein highly expressed in the brain, FEBS Lett., 585, 1979, 10.1016/j.febslet.2011.05.035

Wanner, 1981, The ontogeny of lipid bodies (spherosomes) in plant cells: Ultrastructural evidence, Planta., 151, 109, 10.1007/BF00387812

Watkins, 2007, Evidence for 26 distinct acyl-coenzyme A synthetase genes in the human genome, J. Lipid Res., 48, 2736, 10.1194/jlr.M700378-JLR200

Welte, 2009, Fat on the move: intracellular motion of lipid droplets, Biochem. Soc. Trans., 37, 991, 10.1042/BST0370991

Wilfling, 2013, Triacylglycerol synthesis enzymes mediate lipid droplet growth by relocalizing from the ER to lipid droplets, Dev. Cell., 24, 384, 10.1016/j.devcel.2013.01.013

Wolins, 2006, A proposed model of fat packaging by exchangeable lipid droplet proteins, FEBS Lett., 580, 5484, 10.1016/j.febslet.2006.08.040

Xu, 2012, The FATP1-DGAT2 complex facilitates lipid droplet expansion at the ER-lipid droplet interface, J. Cell Biol., 198, 895, 10.1083/jcb.201201139

Yang, 2011, COPI acts in both vesicular and tubular transport, Nat. Cell Biol., 13, 996, 10.1038/ncb2273

Yang, 2012, The proteomics of lipid droplets: structure, dynamics, and functions of the organelle conserved from bacteria to humans, J. Lipid Res., 53, 1245, 10.1194/jlr.R024117

Zanghellini, 2010, Phospholipid demixing and the birth of a lipid droplet, J. Theor. Biol., 264, 952, 10.1016/j.jtbi.2010.02.025

Zechner, 2012, FAT SIGNALS—lipases and lipolysis in lipid metabolism and signaling, Cell Metab., 15, 279, 10.1016/j.cmet.2011.12.018

Zweytick, 2000, Contribution of Are1p and Are2p to steryl ester synthesis in the yeast Saccharomyces cerevisiae, Eur. J. Biochem., 267, 1075, 10.1046/j.1432-1327.2000.01103.x