{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,5]],"date-time":"2026-11-05T13:31:56Z","timestamp":1793885516233,"version":"4.8.0"},"reference-count":122,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2024,10,3]],"date-time":"2024-10-03T00:00:00Z","timestamp":1727913600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Cell Dev. Biol."],"abstract":"<jats:p>Lipid droplets (LDs) are evolutionarily conserved dynamic organelles that play an important role in cellular physiology. Growing evidence suggests that LD biogenesis occurs at discrete endoplasmic reticulum (ER) subdomains demarcated by the lipodystrophy protein, Seipin, lack of which impairs adipogenesis. However, the mechanisms of how these domains are selected is not completely known. These ER sites undergo ordered assembly of proteins and lipids to initiate LD biogenesis and facilitate establishment of ER-LD contact sites, a prerequisite for proper growth and maturation of droplets. LDs retain both physical and functional association with the ER throughout their lifecycle to facilitate bi-directional communication, such as exchange of proteins and lipids between the two organelles at these ER-LD contact sites. In recent years several molecular tethers have been identified that bridge ER and LDs together including few proteins that are found exclusively at these ER-LD contact interface. Here, we discuss recent advances in understanding the role of factors that ensure functionality of ER-LD contact site machinery for LD homeostasis.<\/jats:p>","DOI":"10.3389\/fcell.2024.1483902","type":"journal-article","created":{"date-parts":[[2024,10,3]],"date-time":"2024-10-03T04:38:14Z","timestamp":1727930294000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":14,"title":["The evolving landscape of ER-LD contact sites"],"prefix":"10.3389","volume":"12","author":[{"given":"Akhil","family":"Kumar","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Surabhi","family":"Yadav","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Vineet","family":"Choudhary","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2024,10,3]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"194","DOI":"10.1038\/s41594-021-00718-y","article-title":"Seipin forms a flexible cage at lipid droplet formation sites","volume":"29","author":"Arlt","year":"2022","journal-title":"Nat. Struct. Mol. Biol."},{"key":"B2","doi-asserted-by":"publisher","first-page":"dev192997","DOI":"10.1242\/dev.192997","article-title":"Loss of the seipin gene perturbs eggshell formation in Caenorhabditiselegans","volume":"147","author":"Bai","year":"2020","journal-title":"Development"},{"key":"B3","doi-asserted-by":"publisher","first-page":"e202006111","DOI":"10.1083\/jcb.202006111","article-title":"FIT2 is an acyl-coenzyme A diphosphatase crucial for endoplasmic reticulum homeostasis","volume":"219","author":"Becuwe","year":"2020","journal-title":"J. Cell Biol."},{"key":"B4","doi-asserted-by":"publisher","first-page":"591","DOI":"10.1016\/j.devcel.2017.05.012","article-title":"ER membrane phospholipids and surface tension control cellular lipid droplet formation","volume":"41","author":"Ben M'barek","year":"2017","journal-title":"Dev. Cell"},{"key":"B5","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1016\/j.devcel.2017.11.020","article-title":"A proximity labeling strategy provides insights into the composition and dynamics of lipid droplet proteomes","volume":"44","author":"Bersuker","year":"2018","journal-title":"Dev. Cell"},{"key":"B6","doi-asserted-by":"publisher","first-page":"4794","DOI":"10.1242\/jcs.134254","article-title":"Targeting of the Arf-GEF GBF1 to lipid droplets and Golgi membranes","volume":"126","author":"Bouvet","year":"2013","journal-title":"J. Cell Sci."},{"key":"B7","doi-asserted-by":"publisher","first-page":"711","DOI":"10.1242\/dmm.015222","article-title":"A novel mouse model of Warburg Micro syndrome reveals roles for RAB18 in eye development and organisation of the neuronal cytoskeleton","volume":"7","author":"Carpanini","year":"2014","journal-title":"Dis. Model Mech."},{"key":"B8","doi-asserted-by":"publisher","first-page":"726","DOI":"10.1091\/mbc.E14-08-1303","article-title":"Seipin performs dissectible functions in promoting lipid droplet biogenesis and regulating droplet morphology","volume":"26","author":"Cartwright","year":"2015","journal-title":"Mol. Biol. Cell"},{"key":"B9","doi-asserted-by":"publisher","first-page":"1042","DOI":"10.1194\/jlr.R023754","article-title":"Seipin: from human disease to molecular mechanism","volume":"53","author":"Cartwright","year":"2012","journal-title":"J. Lipid Res."},{"key":"B10","doi-asserted-by":"publisher","first-page":"38870","DOI":"10.1074\/jbc.M106219200","article-title":"Cloning of DGAT2, a second mammalian diacylglycerol acyltransferase, and related family members","volume":"276","author":"Cases","year":"2001","journal-title":"J. Biol. Chem."},{"key":"B11","doi-asserted-by":"publisher","first-page":"268","DOI":"10.3390\/cells8030268","article-title":"Promethin is a conserved seipin partner protein","volume":"8","author":"Castro","year":"2019","journal-title":"Cells"},{"key":"B12","doi-asserted-by":"publisher","first-page":"e201907183","DOI":"10.1083\/jcb.201907183","article-title":"FIT2 organizes lipid droplet biogenesis with ER tubule-forming proteins and septins","volume":"220","author":"Chen","year":"2021","journal-title":"J. Cell Biol."},{"key":"B13","doi-asserted-by":"publisher","first-page":"25152564221125613","DOI":"10.1177\/25152564221125613","article-title":"VPS13A and VPS13C influence lipid droplet abundance","volume":"5","author":"Chen","year":"2022","journal-title":"Contact (Thousand Oaks)"},{"key":"B14","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1016\/j.devcel.2019.05.003","article-title":"Membrane asymmetry imposes directionality on lipid droplet emergence from the ER","volume":"50","author":"Chorlay","year":"2019","journal-title":"Dev. Cell"},{"key":"B15","doi-asserted-by":"publisher","first-page":"631","DOI":"10.1016\/j.bpj.2017.12.014","article-title":"An asymmetry in monolayer tension regulates lipid droplet budding direction","volume":"114","author":"Chorlay","year":"2018","journal-title":"Biophys. J."},{"key":"B16","doi-asserted-by":"publisher","first-page":"e201910177","DOI":"10.1083\/jcb.201910177","article-title":"Seipin and Nem1 establish discrete ER subdomains to initiate yeast lipid droplet biogenesis","volume":"219","author":"Choudhary","year":"2020","journal-title":"J. Cell Biol."},{"key":"B17","doi-asserted-by":"publisher","first-page":"915","DOI":"10.1016\/j.cub.2018.02.020","article-title":"Architecture of lipid droplets in endoplasmic reticulum is determined by phospholipid intrinsic curvature","volume":"28","author":"Choudhary","year":"2018","journal-title":"Curr. Biol."},{"key":"B18","doi-asserted-by":"publisher","first-page":"e1170276","DOI":"10.1080\/21624054.2016.1170276","article-title":"Keeping FIT, storing fat: lipid droplet biogenesis","volume":"5","author":"Choudhary","year":"2016","journal-title":"Worm"},{"key":"B19","doi-asserted-by":"publisher","first-page":"261","DOI":"10.1083\/jcb.201505067","article-title":"A conserved family of proteins facilitates nascent lipid droplet budding from the ER","volume":"211","author":"Choudhary","year":"2015","journal-title":"J. Cell Biol."},{"key":"B20","doi-asserted-by":"publisher","first-page":"218","DOI":"10.15698\/mic2020.08.727","article-title":"Lipid droplet biogenesis from specialized ER subdomains","volume":"7","author":"Choudhary","year":"2020","journal-title":"Microb. Cell"},{"key":"B21","doi-asserted-by":"publisher","first-page":"650186","DOI":"10.3389\/fcell.2021.650186","article-title":"A unique junctional interface at contact sites between the endoplasmic reticulum and lipid droplets","volume":"9","author":"Choudhary","year":"2021","journal-title":"Front. Cell Dev. Biol."},{"key":"B22","doi-asserted-by":"publisher","first-page":"551","DOI":"10.1016\/j.devcel.2019.10.006","article-title":"LDAF1 and seipin form a lipid droplet assembly complex","volume":"51","author":"Chung","year":"2019","journal-title":"Dev. Cell"},{"key":"B23","doi-asserted-by":"publisher","first-page":"jcs258819","DOI":"10.1242\/jcs.258819","article-title":"Lipid droplets form a network interconnected by the endoplasmic reticulum through which their proteins equilibrate","volume":"135","author":"Cottier","year":"2022","journal-title":"J. Cell Sci."},{"key":"B24","doi-asserted-by":"publisher","first-page":"1335","DOI":"10.1083\/jcb.201808133","article-title":"Cerebellar ataxia disease-associated Snx14 promotes lipid droplet growth at ER-droplet contacts","volume":"218","author":"Datta","year":"2019","journal-title":"J. Cell Biol."},{"key":"B25","doi-asserted-by":"publisher","first-page":"e45453","DOI":"10.15252\/embr.201745453","article-title":"Identification of MOSPD2, a novel scaffold for endoplasmic reticulum membrane contact sites","volume":"19","author":"Di Mattia","year":"2018","journal-title":"EMBO Rep."},{"key":"B26","doi-asserted-by":"publisher","first-page":"362","DOI":"10.1038\/nrm3117","article-title":"ARF family G proteins and their regulators: roles in membrane transport, development and disease","volume":"12","author":"Donaldson","year":"2011","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"B27","doi-asserted-by":"publisher","first-page":"915","DOI":"10.1038\/s41467-023-36375-6","article-title":"Cholesterol esters form supercooled lipid droplets whose nucleation is facilitated by triacylglycerols","volume":"14","author":"Dumesnil","year":"2023","journal-title":"Nat. Commun."},{"key":"B28","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1083\/jcb.201704122","article-title":"Identification of seipin-linked factors that act as determinants of a lipid droplet subpopulation","volume":"217","author":"Eisenberg-Bord","year":"2018","journal-title":"J. Cell Biol."},{"key":"B29","doi-asserted-by":"publisher","first-page":"1157","DOI":"10.1038\/s41556-023-01180-2","article-title":"Arf1 coordinates fatty acid metabolism and mitochondrial homeostasis","volume":"25","author":"Enkler","year":"2023","journal-title":"Nat. Cell Biol."},{"key":"B30","doi-asserted-by":"publisher","first-page":"473","DOI":"10.1083\/jcb.200711136","article-title":"Fld1p, a functional homologue of human seipin, regulates the size of lipid droplets in yeast","volume":"180","author":"Fei","year":"2008","journal-title":"J. Cell Biol."},{"key":"B31","doi-asserted-by":"publisher","first-page":"e202103176","DOI":"10.1083\/jcb.202103176","article-title":"Pex30-like proteins function as adaptors at distinct ER membrane contact sites","volume":"220","author":"Ferreira","year":"2021","journal-title":"J. Cell Biol."},{"key":"B32","doi-asserted-by":"publisher","first-page":"1492","DOI":"10.1002\/cbin.11199","article-title":"DFCP1 associates with lipid droplets","volume":"43","author":"Gao","year":"2019","journal-title":"Cell Biol. Int."},{"key":"B33","doi-asserted-by":"publisher","first-page":"25686","DOI":"10.1074\/jbc.M115.676700","article-title":"Postnatal deletion of fat storage-inducing transmembrane protein 2 (FIT2\/FITM2) causes lethal enteropathy","volume":"290","author":"Goh","year":"2015","journal-title":"J. Biol. Chem."},{"key":"B34","doi-asserted-by":"publisher","first-page":"1422032","DOI":"10.3389\/fcell.2024.1422032","article-title":"FIT2 proteins and lipid droplet emergence, an interplay between phospholipid synthesis, surface tension, and membrane curvature","volume":"12","author":"Graff","year":"2024","journal-title":"Front. Cell Dev. Biol."},{"key":"B35","doi-asserted-by":"publisher","first-page":"829","DOI":"10.1083\/jcb.201502070","article-title":"The seipin complex Fld1\/Ldb16 stabilizes ER-lipid droplet contact sites","volume":"211","author":"Grippa","year":"2015","journal-title":"J. Cell Biol."},{"key":"B36","doi-asserted-by":"publisher","first-page":"19581","DOI":"10.1073\/pnas.1110817108","article-title":"Direct binding of triglyceride to fat storage-inducing transmembrane proteins 1 and 2 is important for lipid droplet formation","volume":"108","author":"Gross","year":"2011","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B37","doi-asserted-by":"publisher","first-page":"1319","DOI":"10.1083\/jcb.201808119","article-title":"Mdm1 maintains endoplasmic reticulum homeostasis by spatially regulating lipid droplet biogenesis","volume":"218","author":"Hariri","year":"2019","journal-title":"J. Cell Biol."},{"key":"B38","doi-asserted-by":"publisher","first-page":"88","DOI":"10.15698\/mic2018.02.614","article-title":"Fat storage-inducing transmembrane (FIT or FITM) proteins are related to lipid phosphatase\/phosphotransferase enzymes","volume":"5","author":"Hayes","year":"2017","journal-title":"Microb. Cell"},{"key":"B39","doi-asserted-by":"publisher","first-page":"9247","DOI":"10.1021\/acs.langmuir.2c01014","article-title":"Lens nucleation and droplet budding in a membrane model for lipid droplet biogenesis","volume":"38","author":"Hegaard","year":"2022","journal-title":"Langmuir"},{"key":"B40","doi-asserted-by":"publisher","first-page":"1317","DOI":"10.1093\/oxfordjournals.jbchem.a124681","article-title":"Cloning and sequence of the SCS3 gene which is required for inositol prototrophy in Saccharomyces cerevisiae","volume":"116","author":"Hosaka","year":"1994","journal-title":"J. Biochem."},{"key":"B41","doi-asserted-by":"publisher","first-page":"118603","DOI":"10.1016\/j.bbamcr.2019.118603","article-title":"Come a little bit closer! Lipid droplet-ER contact sites are getting crowded","volume":"1867","author":"Hugenroth","year":"2020","journal-title":"Biochim. Biophys. Acta Mol. Cell Res."},{"key":"B42","doi-asserted-by":"publisher","first-page":"863907","DOI":"10.3389\/fcell.2022.863907","article-title":"Functions of stress-induced lipid droplets in the nervous system","volume":"10","author":"Islimye","year":"2022","journal-title":"Front. Cell Dev. Biol."},{"key":"B43","doi-asserted-by":"publisher","first-page":"2424","DOI":"10.1242\/jcs.076836","article-title":"Lipid droplets are functionally connected to the endoplasmic reticulum in Saccharomyces cerevisiae","volume":"124","author":"Jacquier","year":"2011","journal-title":"J. Cell Sci."},{"key":"B44","doi-asserted-by":"publisher","first-page":"515","DOI":"10.1083\/jcb.201602064","article-title":"A family of membrane-shaping proteins at ER subdomains regulates pre-peroxisomal vesicle biogenesis","volume":"215","author":"Joshi","year":"2016","journal-title":"J. Cell Biol."},{"key":"B45","doi-asserted-by":"publisher","first-page":"2940","DOI":"10.1038\/s41467-018-05277-3","article-title":"Lipid droplet and peroxisome biogenesis occur at the same ER subdomains","volume":"9","author":"Joshi","year":"2018","journal-title":"Nat. Commun."},{"key":"B46","doi-asserted-by":"publisher","first-page":"1147","DOI":"10.1091\/mbc.E20-09-0590","article-title":"Multiple C2 domain-containing transmembrane proteins promote lipid droplet biogenesis and growth at specialized endoplasmic reticulum subdomains","volume":"32","author":"Joshi","year":"2021","journal-title":"Mol. Biol. Cell"},{"key":"B47","doi-asserted-by":"publisher","first-page":"94","DOI":"10.1073\/pnas.0708579105","article-title":"Evolutionarily conserved gene family important for fat storage","volume":"105","author":"Kadereit","year":"2008","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B48","doi-asserted-by":"publisher","first-page":"985","DOI":"10.1083\/jcb.201305142","article-title":"Acyl-CoA synthetase 3 promotes lipid droplet biogenesis in ER microdomains","volume":"203","author":"Kassan","year":"2013","journal-title":"J. Cell Biol."},{"key":"B49","doi-asserted-by":"publisher","first-page":"jcs256206","DOI":"10.1242\/jcs.256206","article-title":"The surface of lipid droplets constitutes a barrier for endoplasmic reticulum-resident integral membrane proteins","volume":"135","author":"Khaddaj","year":"2022","journal-title":"J. Cell Sci."},{"key":"B50","doi-asserted-by":"publisher","first-page":"e12811","DOI":"10.1371\/journal.pone.0012811","article-title":"Triglyceride blisters in lipid bilayers: implications for lipid droplet biogenesis and the mobile lipid signal in cancer cell membranes","volume":"5","author":"Khandelia","year":"2010","journal-title":"PLoS One"},{"key":"B51","doi-asserted-by":"publisher","first-page":"1465","DOI":"10.1016\/j.celrep.2013.04.015","article-title":"A conserved role for atlastin GTPases in regulating lipid droplet size","volume":"3","author":"Klemm","year":"2013","journal-title":"Cell Rep."},{"key":"B52","doi-asserted-by":"publisher","first-page":"5892","DOI":"10.1038\/s41467-021-26162-6","article-title":"Mechanism of lipid droplet formation by the yeast Sei1\/Ldb16 Seipin complex","volume":"12","author":"Klug","year":"2021","journal-title":"Nat. Commun."},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.1101\/2023.08.14.553257","article-title":"ATG2A-mediated bridge-like lipid transport regulates lipid droplet accumulation","author":"Korfhage","year":"2023","journal-title":"bioRxiv"},{"key":"B54","doi-asserted-by":"publisher","first-page":"3625","DOI":"10.1083\/jcb.201807019","article-title":"VPS13A and VPS13C are lipid transport proteins differentially localized at ER contact sites","volume":"217","author":"Kumar","year":"2018","journal-title":"J. Cell Biol."},{"key":"B55","doi-asserted-by":"publisher","first-page":"151362","DOI":"10.1016\/j.ejcb.2023.151362","article-title":"Concept of lipid droplet biogenesis","volume":"102","author":"Kumari","year":"2023","journal-title":"Eur. J. Cell Biol."},{"key":"B56","doi-asserted-by":"publisher","first-page":"1421","DOI":"10.1002\/yea.320101105","article-title":"Characterization of lipid particles of the yeast, Saccharomyces cerevisiae","volume":"10","author":"Leber","year":"1994","journal-title":"Yeast"},{"key":"B57","doi-asserted-by":"publisher","first-page":"500","DOI":"10.1016\/j.ajhg.2016.01.014","article-title":"Loss of VPS13C function in autosomal-recessive parkinsonism causes mitochondrial dysfunction and increases PINK1\/parkin-dependent mitophagy","volume":"98","author":"Lesage","year":"2016","journal-title":"Am. J. Hum. Genet."},{"key":"B58","doi-asserted-by":"publisher","first-page":"441","DOI":"10.15252\/embj.201694866","article-title":"COPI-TRAPPII activates Rab18 and regulates its lipid droplet association","volume":"36","author":"Li","year":"2017","journal-title":"EMBO J."},{"key":"B59","doi-asserted-by":"publisher","first-page":"1026","DOI":"10.1002\/cbin.10650","article-title":"Rab proteins as regulators of lipid droplet formation and lipolysis","volume":"40","author":"Li","year":"2016","journal-title":"Cell Biol. Int."},{"key":"B60","doi-asserted-by":"publisher","first-page":"343","DOI":"10.1016\/j.celrep.2019.03.025","article-title":"The ER-localized protein DFCP1 modulates ER-lipid droplet contact formation","volume":"27","author":"Li","year":"2019","journal-title":"Cell Rep."},{"key":"B61","doi-asserted-by":"publisher","first-page":"e202001161","DOI":"10.1083\/jcb.202001161","article-title":"Cryo-EM reconstruction of a VPS13 fragment reveals a long groove to channel lipids between membranes","volume":"219","author":"Li","year":"2020","journal-title":"J. Cell Biol."},{"key":"B62","doi-asserted-by":"publisher","first-page":"e2321991121","DOI":"10.1073\/pnas.2321991121","article-title":"Different ER-plasma membrane tethers play opposing roles in autophagy of the cortical ER","volume":"121","author":"Liu","year":"2024","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B63","doi-asserted-by":"publisher","first-page":"ar131","DOI":"10.1091\/mbc.E22-07-0279","article-title":"PI(3)P and DFCP1 regulate the biogenesis of lipid droplets","volume":"33","author":"Lukmantara","year":"2022","journal-title":"Mol. Biol. Cell"},{"key":"B64","doi-asserted-by":"publisher","first-page":"365","DOI":"10.1038\/ng585","article-title":"Identification of the gene altered in Berardinelli-Seip congenital lipodystrophy on chromosome 11q13","volume":"28","author":"Magre","year":"2001","journal-title":"Nat. Genet."},{"key":"B65","doi-asserted-by":"publisher","first-page":"eade7753","DOI":"10.1126\/sciadv.ade7753","article-title":"Rab1b facilitates lipid droplet growth by ER-to-lipid droplet targeting of DGAT2","volume":"10","author":"Malis","year":"2024","journal-title":"Sci. Adv."},{"key":"B66","doi-asserted-by":"publisher","first-page":"42325","DOI":"10.1074\/jbc.M506651200","article-title":"Regulated localization of Rab18 to lipid droplets: effects of lipolytic stimulation and inhibition of lipid droplet catabolism","volume":"280","author":"Martin","year":"2005","journal-title":"J. Biol. Chem."},{"key":"B67","doi-asserted-by":"publisher","first-page":"7113","DOI":"10.1111\/febs.16280","article-title":"ATG2 and VPS13 proteins: molecular highways transporting lipids to drive membrane expansion and organelle communication","volume":"289","author":"McEwan","year":"2022","journal-title":"FEBS J."},{"key":"B68","doi-asserted-by":"publisher","first-page":"1318","DOI":"10.1016\/j.bbalip.2014.06.004","article-title":"Characterization of the interaction of diacylglycerol acyltransferase-2 with the endoplasmic reticulum and lipid droplets","volume":"1841","author":"McFie","year":"2014","journal-title":"Biochim. Biophys. Acta"},{"key":"B69","doi-asserted-by":"publisher","first-page":"9560","DOI":"10.1074\/jbc.M114.547687","article-title":"Fat storage-inducing transmembrane protein 2 is required for normal fat storage in adipose tissue","volume":"289","author":"Miranda","year":"2014","journal-title":"J. Biol. Chem."},{"key":"B70","doi-asserted-by":"publisher","first-page":"e1002890","DOI":"10.1371\/journal.pgen.1002890","article-title":"SCS3 and YFT2 link transcription of phospholipid biosynthetic genes to ER stress and the UPR","volume":"8","author":"Moir","year":"2012","journal-title":"PLoS Genet."},{"key":"B71","doi-asserted-by":"publisher","first-page":"117","DOI":"10.1016\/j.bbrc.2005.07.050","article-title":"Arf1-dependent PLD1 is localized to oleic acid-induced lipid droplets in NIH3T3 cells","volume":"335","author":"Nakamura","year":"2005","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"B72","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1038\/s41580-018-0085-z","article-title":"Dynamics and functions of lipid droplets","volume":"20","author":"Olzmann","year":"2019","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"B73","doi-asserted-by":"publisher","first-page":"2601","DOI":"10.1242\/jcs.02401","article-title":"Rab18 localizes to lipid droplets and induces their close apposition to the endoplasmic reticulum-derived membrane","volume":"118","author":"Ozeki","year":"2005","journal-title":"J. Cell Sci."},{"key":"B74","doi-asserted-by":"publisher","first-page":"1546","DOI":"10.1016\/j.celrep.2016.10.037","article-title":"SEIPIN regulates lipid droplet expansion and adipocyte development by modulating the activity of glycerol-3-phosphate acyltransferase","volume":"17","author":"Pagac","year":"2016","journal-title":"Cell Rep."},{"key":"B75","doi-asserted-by":"publisher","first-page":"e3000998","DOI":"10.1371\/journal.pbio.3000998","article-title":"Seipin traps triacylglycerols to facilitate their nanoscale clustering in the endoplasmic reticulum membrane","volume":"19","author":"Prasanna","year":"2021","journal-title":"PLoS Biol."},{"key":"B76","doi-asserted-by":"publisher","first-page":"e2106524","DOI":"10.1002\/smll.202106524","article-title":"Lipid droplets embedded in a model cell membrane create a phospholipid diffusion barrier","volume":"18","author":"Puza","year":"2022","journal-title":"Small"},{"key":"B77","doi-asserted-by":"publisher","first-page":"912","DOI":"10.1016\/j.tcb.2021.06.003","article-title":"Seipin: harvesting fat and keeping adipocytes healthy","volume":"31","author":"Rao","year":"2021","journal-title":"Trends Cell Biol."},{"key":"B78","doi-asserted-by":"publisher","first-page":"e202112068","DOI":"10.1083\/jcb.202112068","article-title":"Seipin concentrates distinct neutral lipids via interactions with their acyl chain carboxyl esters","volume":"221","author":"Renne","year":"2022","journal-title":"J. Cell Biol."},{"key":"B79","doi-asserted-by":"publisher","first-page":"726261","DOI":"10.3389\/fcell.2021.726261","article-title":"Lipid droplet-organelle contact sites as hubs for fatty acid metabolism, trafficking, and metabolic channeling","volume":"9","author":"Renne","year":"2021","journal-title":"Front. Cell Dev. Biol."},{"key":"B80","doi-asserted-by":"publisher","first-page":"5111","DOI":"10.1093\/hmg\/ddw315","article-title":"Reep1 null mice reveal a converging role for hereditary spastic paraplegia proteins in lipid droplet regulation","volume":"25","author":"Renvoise","year":"2016","journal-title":"Hum. Mol. Genet."},{"key":"B81","doi-asserted-by":"publisher","first-page":"1112954","DOI":"10.3389\/fcell.2023.1112954","article-title":"Seipin-still a mysterious protein?","volume":"11","author":"Salo","year":"2023","journal-title":"Front. Cell Dev. Biol."},{"key":"B82","doi-asserted-by":"publisher","first-page":"2699","DOI":"10.15252\/embj.201695170","article-title":"Seipin regulates ER-lipid droplet contacts and cargo delivery","volume":"35","author":"Salo","year":"2016","journal-title":"EMBO J."},{"key":"B83","doi-asserted-by":"publisher","first-page":"478","DOI":"10.1016\/j.devcel.2019.05.016","article-title":"Seipin facilitates triglyceride flow to lipid droplet and counteracts droplet ripening via endoplasmic reticulum contact","volume":"50","author":"Salo","year":"2019","journal-title":"Dev. Cell"},{"key":"B84","doi-asserted-by":"publisher","first-page":"2481","DOI":"10.1016\/j.cub.2020.04.066","article-title":"Membrane curvature catalyzes lipid droplet assembly","volume":"30","author":"Santinho","year":"2020","journal-title":"Curr. Biol."},{"key":"B85","doi-asserted-by":"publisher","first-page":"1896","DOI":"10.1002\/hep.27667","article-title":"The small GTPase Rab7 as a central regulator of hepatocellular lipophagy","volume":"61","author":"Schroeder","year":"2015","journal-title":"Hepatology"},{"key":"B86","doi-asserted-by":"publisher","first-page":"1188","DOI":"10.1016\/j.bbalip.2017.06.005","article-title":"A different kind of love - lipid droplet contact sites","volume":"1862","author":"Schuldiner","year":"2017","journal-title":"Biochim. Biophys. Acta Mol. Cell Biol. Lipids"},{"key":"B87","doi-asserted-by":"publisher","first-page":"1287","DOI":"10.1038\/s41467-019-09253-3","article-title":"Coming together to define membrane contact sites","volume":"10","author":"Scorrano","year":"2019","journal-title":"Nat. Commun."},{"key":"B88","doi-asserted-by":"publisher","first-page":"3259","DOI":"10.1038\/s41598-020-59982-5","article-title":"Oligomers of the lipodystrophy protein seipin may co-ordinate GPAT3 and AGPAT2 enzymes to facilitate adipocyte differentiation","volume":"10","author":"Sim","year":"2020","journal-title":"Sci. Rep."},{"key":"B89","doi-asserted-by":"publisher","first-page":"1834","DOI":"10.1242\/jcs.045849","article-title":"Coatomer-dependent protein delivery to lipid droplets","volume":"122","author":"Soni","year":"2009","journal-title":"J. Cell Sci."},{"key":"B90","doi-asserted-by":"publisher","first-page":"423","DOI":"10.1016\/s0955-0674(02)00346-0","article-title":"ARF1 regulatory factors and COPI vesicle formation","volume":"14","author":"Spang","year":"2002","journal-title":"Curr. Opin. Cell Biol."},{"key":"B91","doi-asserted-by":"publisher","first-page":"40273","DOI":"10.1074\/jbc.M607986200","article-title":"Membrane topology and identification of key functional amino acid residues of murine acyl-CoA:diacylglycerol acyltransferase-2","volume":"281","author":"Stone","year":"2006","journal-title":"J. Biol. Chem."},{"key":"B92","doi-asserted-by":"publisher","first-page":"5352","DOI":"10.1074\/jbc.M805768200","article-title":"The endoplasmic reticulum enzyme DGAT2 is found in mitochondria-associated membranes and has a mitochondrial targeting signal that promotes its association with mitochondria","volume":"284","author":"Stone","year":"2009","journal-title":"J. Biol. Chem."},{"key":"B93","doi-asserted-by":"publisher","first-page":"4080","DOI":"10.1083\/jcb.201809067","article-title":"Cryo-electron microscopy structure of the lipid droplet-formation protein seipin","volume":"217","author":"Sui","year":"2018","journal-title":"J. Cell Biol."},{"key":"B94","doi-asserted-by":"publisher","first-page":"20890","DOI":"10.1073\/pnas.0704154104","article-title":"The lipodystrophy protein seipin is found at endoplasmic reticulum lipid droplet junctions and is important for droplet morphology","volume":"104","author":"Szymanski","year":"2007","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B95","doi-asserted-by":"publisher","first-page":"3819","DOI":"10.1002\/1873-3468.12901","article-title":"Differential requirement for ATG2A domains for localization to autophagic membranes and lipid droplets","volume":"591","author":"Tamura","year":"2017","journal-title":"FEBS Lett."},{"key":"B96","doi-asserted-by":"publisher","first-page":"44507","DOI":"10.1074\/jbc.M207712200","article-title":"The surface of lipid droplets is a phospholipid monolayer with a unique Fatty Acid composition","volume":"277","author":"Tauchi-Sato","year":"2002","journal-title":"J. Biol. Chem."},{"key":"B97","doi-asserted-by":"publisher","first-page":"127","DOI":"10.1083\/jcb.201704115","article-title":"Regulation of lipid droplets by metabolically controlled Ldo isoforms","volume":"217","author":"Teixeira","year":"2018","journal-title":"J. Cell Biol."},{"key":"B98","doi-asserted-by":"publisher","first-page":"13244","DOI":"10.1073\/pnas.1307685110","article-title":"COPI buds 60-nm lipid droplets from reconstituted water-phospholipid-triacylglyceride interfaces, suggesting a tension clamp function","volume":"110","author":"Thiam","year":"2013","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B99","doi-asserted-by":"publisher","first-page":"108","DOI":"10.1016\/j.tcb.2020.11.006","article-title":"Lipid droplet nucleation","volume":"31","author":"Thiam","year":"2021","journal-title":"Trends Cell Biol."},{"key":"B100","doi-asserted-by":"publisher","first-page":"611","DOI":"10.1016\/j.ajhg.2014.10.007","article-title":"Mutations in SNX14 cause a distinctive autosomal-recessive cerebellar ataxia and intellectual disability syndrome","volume":"95","author":"Thomas","year":"2014","journal-title":"Am. J. Hum. Genet."},{"key":"B101","doi-asserted-by":"publisher","first-page":"121","DOI":"10.1038\/88825","article-title":"The gene encoding a newly discovered protein, chorein, is mutated in chorea-acanthocytosis","volume":"28","author":"Ueno","year":"2001","journal-title":"Nat. Genet."},{"key":"B102","doi-asserted-by":"publisher","first-page":"896","DOI":"10.1091\/mbc.E11-09-0785","article-title":"Mammalian Atg2 proteins are essential for autophagosome formation and important for regulation of size and distribution of lipid droplets","volume":"23","author":"Velikkakath","year":"2012","journal-title":"Mol. Biol. Cell"},{"key":"B103","doi-asserted-by":"publisher","first-page":"491","DOI":"10.1146\/annurev-cellbio-100616-060608","article-title":"Lipid droplet biogenesis","volume":"33","author":"Walther","year":"2017","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"B104","doi-asserted-by":"publisher","first-page":"1214","DOI":"10.1242\/jcs.137737","article-title":"Control of lipid droplet size in budding yeast requires the collaboration between Fld1 and Ldb16","volume":"127","author":"Wang","year":"2014","journal-title":"J. Cell Sci."},{"key":"B105","doi-asserted-by":"publisher","first-page":"e16582","DOI":"10.7554\/eLife.16582","article-title":"Seipin is required for converting nascent to mature lipid droplets","volume":"5","author":"Wang","year":"2016","journal-title":"Elife"},{"key":"B106","doi-asserted-by":"publisher","DOI":"10.1101\/2023.12.05.570183","article-title":"FITM2 deficiency results in ER lipid accumulation, ER stress, reduced apolipoprotein B lipidation, and VLDL triglyceride secretion in vitro and in mouse liver","author":"Wang","year":"2023","journal-title":"bioRxiv"},{"key":"B107","doi-asserted-by":"publisher","first-page":"1252","DOI":"10.1038\/s41467-021-21525-5","article-title":"An ESCRT-dependent step in fatty acid transfer from lipid droplets to mitochondria through VPS13D-TSG101 interactions","volume":"12","author":"Wang","year":"2021","journal-title":"Nat. Commun."},{"key":"B108","doi-asserted-by":"publisher","first-page":"2939","DOI":"10.1038\/s41467-018-05278-2","article-title":"Seipin and the membrane-shaping protein Pex30 cooperate in organelle budding from the endoplasmic reticulum","volume":"9","author":"Wang","year":"2018","journal-title":"Nat. Commun."},{"key":"B109","doi-asserted-by":"publisher","first-page":"1260","DOI":"10.1016\/j.bbalip.2017.07.006","article-title":"Lipid droplet functions beyond energy storage","volume":"1862","author":"Welte","year":"2017","journal-title":"Biochim. Biophys. Acta Mol. Cell Biol. Lipids"},{"key":"B110","doi-asserted-by":"publisher","first-page":"e01607","DOI":"10.7554\/eLife.01607","article-title":"Arf1\/COPI machinery acts directly on lipid droplets and enables their connection to the ER for protein targeting","volume":"3","author":"Wilfling","year":"2014","journal-title":"Elife"},{"key":"B111","doi-asserted-by":"publisher","first-page":"384","DOI":"10.1016\/j.devcel.2013.01.013","article-title":"Triacylglycerol synthesis enzymes mediate lipid droplet growth by relocalizing from the ER to lipid droplets","volume":"24","author":"Wilfling","year":"2013","journal-title":"Dev. Cell"},{"key":"B112","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1038\/ng1313","article-title":"Heterozygous missense mutations in BSCL2 are associated with distal hereditary motor neuropathy and Silver syndrome","volume":"36","author":"Windpassinger","year":"2004","journal-title":"Nat. Genet."},{"key":"B113","doi-asserted-by":"publisher","first-page":"378","DOI":"10.1016\/j.devcel.2014.07.005","article-title":"Rab8a-AS160-MSS4 regulatory circuit controls lipid droplet fusion and growth","volume":"30","author":"Wu","year":"2014","journal-title":"Dev. Cell"},{"key":"B114","doi-asserted-by":"publisher","first-page":"975","DOI":"10.1083\/jcb.201704184","article-title":"Rab18 promotes lipid droplet (LD) growth by tethering the ER to LDs through SNARE and NRZ interactions","volume":"217","author":"Xu","year":"2018","journal-title":"J. Cell Biol."},{"key":"B115","doi-asserted-by":"publisher","first-page":"895","DOI":"10.1083\/jcb.201201139","article-title":"The FATP1-DGAT2 complex facilitates lipid droplet expansion at the ER-lipid droplet interface","volume":"198","author":"Xu","year":"2012","journal-title":"J. Cell Biol."},{"key":"B116","doi-asserted-by":"publisher","first-page":"248","DOI":"10.1016\/j.devcel.2018.09.010","article-title":"Human SEIPIN binds anionic phospholipids","volume":"47","author":"Yan","year":"2018","journal-title":"Dev. Cell"},{"key":"B117","doi-asserted-by":"publisher","first-page":"443","DOI":"10.1038\/s41574-023-00845-0","article-title":"Lipid droplet biogenesis and functions in health and disease","volume":"19","author":"Zadoorian","year":"2023","journal-title":"Nat. Rev. Endocrinol."},{"key":"B118","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1242\/dmm.026476","article-title":"A homozygous FITM2 mutation causes a deafness-dystonia syndrome with motor regression and signs of ichthyosis and sensory neuropathy","volume":"10","author":"Zazo Seco","year":"2017","journal-title":"Dis. Model Mech."},{"key":"B119","doi-asserted-by":"publisher","first-page":"e2113074119","DOI":"10.1073\/pnas.2113074119","article-title":"Destabilization of \u03b2 Cell FIT2 by saturated fatty acids alter lipid droplet numbers and contribute to ER stress and diabetes","volume":"119","author":"Zheng","year":"2022","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B120","doi-asserted-by":"publisher","first-page":"e168594","DOI":"10.1172\/jci.insight.168594","article-title":"Altered lipid homeostasis is associated with cerebellar neurodegeneration in SNX14 deficiency","volume":"9","author":"Zhou","year":"2024","journal-title":"JCI Insight"},{"key":"B121","doi-asserted-by":"publisher","first-page":"e2017205118","DOI":"10.1073\/pnas.2017205118","article-title":"Seipin accumulates and traps diacylglycerols and triglycerides in its ring-like structure","volume":"118","author":"Zoni","year":"2021","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B122","doi-asserted-by":"publisher","first-page":"e202110044","DOI":"10.1083\/jcb.202110044","article-title":"MOSPD2 is an endoplasmic reticulum-lipid droplet tether functioning in LD homeostasis","volume":"221","author":"Zouiouich","year":"2022","journal-title":"J. Cell Biol."}],"container-title":["Frontiers in Cell and Developmental Biology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcell.2024.1483902\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,10,3]],"date-time":"2024-10-03T04:38:16Z","timestamp":1727930296000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcell.2024.1483902\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,10,3]]},"references-count":122,"alternative-id":["10.3389\/fcell.2024.1483902"],"URL":"https:\/\/doi.org\/10.3389\/fcell.2024.1483902","relation":{},"ISSN":["2296-634X"],"issn-type":[{"value":"2296-634X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2024,10,3]]},"article-number":"1483902"}}