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Abstract

Lipid droplets, also known as oil bodies or lipid bodies, are plant organelles that compartmentalize neutral lipids as a hydrophobic matrix covered by proteins embedded in a phospholipid monolayer. Some of these proteins have been known for decades, such as oleosins, caleosins, and steroleosins, whereas a host of others have been discovered more recently with various levels of abundance on lipid droplets, depending on the tissue and developmental stage. In addition to a growing inventory of lipid droplet proteins, the subcellular machinery that contributes to the biogenesis and degradation of lipid droplets is being identified and attention is turning to more mechanistic questions regarding lipid droplet dynamics. While lipid droplets are mostly regarded as storage deposits for carbon and energy in lipid-rich plant tissues such as seeds, these organelles are present in essentially all plant cells, where they display additional functions in signaling, membrane remodeling, and the compartmentalization of a variety of hydrophobic components. Remarkable metabolic engineering efforts have demonstrated the plasticity of vegetative tissues such as leaves to synthesize and package large amounts of storage lipids, which enable future applications in bioenergy and the engineering of high-value lipophilic compounds. Here, we review the growing body of knowledge about lipid droplets in plant cells, describe the evolutionary similarity and divergence in their associated subcellular machinery, and point to gaps that deserve future attention.

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2023-05-22
2024-04-28
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Literature Cited

  1. 1.
    Abell BM, Hahn M, Holbrook LA, Moloney MM. 2004. Membrane topology and sequence requirements for oil body targeting of oleosin. Plant J. 37:461–70
    [Google Scholar]
  2. 2.
    Andrianov V, Borisjuk N, Pogrebnyak N, Brinker A, Dixon J et al. 2010. Tobacco as a production platform for biofuel: overexpression of Arabidopsis DGAT and LEC2 genes increases accumulation and shifts the composition of lipids in green biomass. Plant Biotechnol. J. 8:277–87
    [Google Scholar]
  3. 3.
    Arlt H, Sui X, Folger B, Adams C, Chen X et al. 2022. Seipin forms a flexible cage at lipid droplet formation sites. Nat. Struct. Mol. Biol. 29:194–202
    [Google Scholar]
  4. 4.
    Aubert Y, Vile D, Pervent M, Aldon D, Ranty B et al. 2010. RD20, a stress-inducible caleosin, participates in stomatal control, transpiration and drought tolerance in Arabidopsis thaliana. Plant Cell Physiol. 51:1975–87
    [Google Scholar]
  5. 5.
    Austin JR II, Frost E, Vidi P-A, Kessler F, Staehelin LA 2006. Plastoglobules are lipoprotein subcompartments of the chloroplast that are permanently coupled to thylakoid membranes and contain biosynthetic enzymes. Plant Cell 18:1693–703
    [Google Scholar]
  6. 6.
    Bae SW, Jung S, Choi SC, Kim MY, Ryu SB. 2020. Lipid composition of latex and rubber particles in Hevea brasiliensis and Taraxacum kok-saghyz. Molecules 25:5110
    [Google Scholar]
  7. 7.
    Barajas-Lopez JD, Tiwari A, Zarza X, Shaw MW, Pascual J et al. 2021. EARLY RESPONSE TO DEHYDRATION 7 remodels cell membrane lipid composition during cold stress in Arabidopsis. Plant Cell Physiol. 62:80–91
    [Google Scholar]
  8. 8.
    Bates PD, Browse J. 2012. The significance of different diacylglycerol synthesis pathways on plant oil composition and bioengineering. Front. Plant Sci. 3:147
    [Google Scholar]
  9. 9.
    Bates PD, Ohlrogge JB, Pollard M. 2007. Incorporation of newly synthesized fatty acids into cytosolic glycerolipids in pea leaves occurs via acyl editing. J. Biol. Chem. 282:31206–16
    [Google Scholar]
  10. 10.
    Bates PD, Stymne S, Ohlrogge J. 2013. Biochemical pathways in seed oil synthesis. Curr. Opin. Plant Biol. 16:358–64
    [Google Scholar]
  11. 11.
    Baud S, Lepiniec L. 2009. Regulation of de novo fatty acid synthesis in maturing oilseeds of Arabidopsis. Plant Physiol. Biochem. 47:448–55
    [Google Scholar]
  12. 12.
    Berthelot K, Lecomte S, Estevez Y, Zhendre V, Henry S et al. 2014. Rubber particle proteins, HbREF and HbSRPP, show different interactions with model membranes. Biochim. Biophys. Acta Biomembr. 1838:287–99
    [Google Scholar]
  13. 13.
    Besagni C, Kessler F. 2013. A mechanism implicating plastoglobules in thylakoid disassembly during senescence and nitrogen starvation. Planta 237:463–70
    [Google Scholar]
  14. 14.
    Bohnert M. 2020. Tethering fat: tethers in lipid droplet contact sites. Contact 3: https://doi.org/10.1177/2515256420908142
    [Google Scholar]
  15. 15.
    Bouvier-Nave P, Berna A, Noiriel A, Compagnon V, Carlsson AS et al. 2010. Involvement of the Phospholipid Sterol Acyltransferase1 in plant sterol homeostasis and leaf senescence. Plant Physiol. 152:107–19
    [Google Scholar]
  16. 16.
    Boyle NR, Page MD, Liu B, Blaby IK, Casero D et al. 2012. Three acyltransferases and nitrogen-responsive regulator are implicated in nitrogen starvation-induced triacylglycerol accumulation in Chlamydomonas. J. Biol. Chem. 287:15811–25
    [Google Scholar]
  17. 17.
    Brocard L, Immel F, Coulon D, Esnay N, Tuphile K et al. 2017. Proteomic analysis of lipid droplets from Arabidopsis aging leaves brings new insight into their biogenesis and functions. Front. Plant Sci. 8:894
    [Google Scholar]
  18. 18.
    Busta L, Chapman KD, Cahoon EB. 2022. Better together: protein partnerships for lineage-specific oil accumulation. Curr. Opin. Plant Biol. 66:102191
    [Google Scholar]
  19. 19.
    Cai Y, Goodman JM, Pyc M, Mullen RT, Dyer JM, Chapman KD. 2015. Arabidopsis SEIPIN proteins modulate triacylglycerol accumulation and influence lipid droplet proliferation. Plant Cell 27:2616–36Identification of SEIPIN homologs in Arabidopsis that highlighted similarities and functional differences among three plant isoforms and the single human and yeast SEIPINs.
    [Google Scholar]
  20. 20.
    Cai Y, Whitehead P, Chappell J, Chapman KD. 2019. Mouse lipogenic proteins promote the co-accumulation of triacylglycerols and sesquiterpenes in plant cells. Planta 250:79–94
    [Google Scholar]
  21. 21.
    Cartwright BR, Goodman JM. 2012. Seipin: from human disease to molecular mechanism. J. Lipid Res. 53:1042–55
    [Google Scholar]
  22. 22.
    Chapman KD, Aziz M, Dyer JM, Mullen RT. 2019. Mechanisms of lipid droplet biogenesis. Biochem. J. 476:1929–42
    [Google Scholar]
  23. 23.
    Chapman KD, Dyer JM, Mullen RT. 2012. Biogenesis and functions of lipid droplets in plants: Thematic Review Series: lipid droplet synthesis and metabolism: from yeast to man. J. Lipid Res. 53:215–26
    [Google Scholar]
  24. 24.
    Chapman KD, Ohlrogge JB. 2012. Compartmentation of triacylglycerol accumulation in plants. J. Biol. Chem. 287:2288–94
    [Google Scholar]
  25. 25.
    Chen JC, Tsai CCY, Tzen JTC. 1999. Cloning and secondary structure analysis of caleosin, a unique calcium-binding protein in oil bodies of plant seeds. Plant Cell Physiol. 40:1079–86
    [Google Scholar]
  26. 26.
    Chen Q, Steinhauer L, Hammerlindl J, Keller W, Zou J. 2007. Biosynthesis of phytosterol esters: identification of a sterol O-acyltransferase in Arabidopsis. Plant Physiol. 145:974–84
    [Google Scholar]
  27. 27.
    Cherian S, Ryu SB, Cornish K. 2019. Natural rubber biosynthesis in plants, the rubber transferase complex, and metabolic engineering progress and prospects. Plant Biotechnol. J. 17:2041–61
    [Google Scholar]
  28. 28.
    Chernova A, Gubaev R, Mazin P, Goryunova S, Demurin Y et al. 2019. UPLC–MS triglyceride profiling in sunflower and rapeseed seeds. Biomolecules 9:9
    [Google Scholar]
  29. 29.
    Chung J, Wu X, Lambert TJ, Lai ZW, Walther TC, Farese RV Jr. 2019. LDAF1 and seipin form a lipid droplet assembly complex. Dev. Cell 51:551–63.e7
    [Google Scholar]
  30. 30.
    Clemente TE, Cahoon EB. 2009. Soybean oil: genetic approaches for modification of functionality and total content. Plant Physiol. 151:1030–40
    [Google Scholar]
  31. 31.
    Cook R, Lupette J, Benning C. 2021. The role of chloroplast membrane lipid metabolism in plant environmental responses. Cells 10:706
    [Google Scholar]
  32. 32.
    Coulon D, Brocard L, Tuphile K, Bréhélin C. 2020. Arabidopsis LDIP protein locates at a confined area within the lipid droplet surface and favors lipid droplet formation. Biochimie 169:29–40
    [Google Scholar]
  33. 33.
    da Silva Ramos LC, Tango JS, Savi A, Leal NR. 1984. Variability for oil and fatty acid composition in castorbean varieties. J. Am. Oil Chem. Soc. 61:1841–43
    [Google Scholar]
  34. 34.
    Davidi L, Katz A, Pick U. 2012. Characterization of major lipid droplet proteins from Dunaliella. Planta 236:19–33
    [Google Scholar]
  35. 35.
    De Domenico S, Bonsegna S, Lenucci MS, Poltronieri P, Di Sansebastiano GP, Santino A. 2011. Localization of seed oil body proteins in tobacco protoplasts reveals specific mechanisms of protein targeting to leaf lipid droplets. J. Integr. Plant Biol. 53:858–68
    [Google Scholar]
  36. 36.
    De Marcos Lousa C, van Roermund CW, Postis VLG, Dietrich D, Kerr ID et al. 2013. Intrinsic acyl-CoA thioesterase activity of a peroxisomal ATP binding cassette transporter is required for transport and metabolism of fatty acids. PNAS 110:1279–84
    [Google Scholar]
  37. 37.
    de Vries J, Ischebeck T. 2020. Ties between stress and lipid droplets pre-date seeds. Trends Plant Sci. 25:1203–14Intriguing conceptual overview of LD protein phylogenies in plant and algal species that identifies distant evolutionary relationships.
    [Google Scholar]
  38. 38.
    Deruyffelaere C, Bouchez I, Morin H, Guillot A, Miquel M et al. 2015. Ubiquitin-mediated proteasomal degradation of oleosins is involved in oil body mobilization during post-germinative seedling growth in Arabidopsis. Plant Cell Physiol. 56:1374–87
    [Google Scholar]
  39. 39.
    Deruyffelaere C, Purkrtova Z, Bouchez I, Collet B, Cacas J-L et al. 2018. PUX10 is a CDC48A adaptor protein that regulates the extraction of ubiquitinated oleosins from seed lipid droplets in Arabidopsis. Plant Cell 30:2116–36
    [Google Scholar]
  40. 40.
    Doner NM, Seay D, Mehling M, Sun S, Gidda SK et al. 2021. Arabidopsis thaliana EARLY RESPONSIVE TO DEHYDRATION 7 localizes to lipid droplets via its senescence domain. Front. Plant Sci. 12:658961
    [Google Scholar]
  41. 41.
    Dyer JM, Stymne S, Green AG, Carlsson AS. 2008. High-value oils from plants. Plant J. 54:640–55
    [Google Scholar]
  42. 42.
    Eastmond PJ. 2004. Cloning and characterization of the acid lipase from castor beans. J. Biol. Chem. 279:45540–45
    [Google Scholar]
  43. 43.
    Eastmond PJ. 2006. SUGAR-DEPENDENT1 encodes a patatin domain triacylglycerol lipase that initiates storage oil breakdown in germinating Arabidopsis seeds. Plant Cell 18:665–75
    [Google Scholar]
  44. 44.
    Esnay N, Dyer JM, Mullen RT, Chapman KD. 2020. Lipid droplet–peroxisome connections in plants. Contact 3: https://doi.org/10.1177/2515256420908765
    [Google Scholar]
  45. 45.
    Espinoza-Corral R, Schwenkert S, Lundquist PK. 2021. Molecular changes of Arabidopsis thaliana plastoglobules facilitate thylakoid membrane remodeling under high light stress. Plant J. 106:1571–87
    [Google Scholar]
  46. 46.
    Fan J, Yan C, Zhang X, Xu C. 2013. Dual role for phospholipid:diacylglycerol acyltransferase: enhancing fatty acid synthesis and diverting fatty acids from membrane lipids to triacylglycerol in Arabidopsis leaves. Plant Cell 25:3506–18
    [Google Scholar]
  47. 47.
    Fan J, Yu L, Xu C. 2017. A central role for triacylglycerol in membrane lipid breakdown, fatty acid β-oxidation, and plant survival under extended darkness. Plant Physiol. 174:1517–30
    [Google Scholar]
  48. 48.
    Fan J, Yu L, Xu C 2019. Dual role for autophagy in lipid metabolism in Arabidopsis. Plant Cell 31:1598–613Evidence is presented in this article for a role of lipophagy in plant cells; it includes a complex model for the interaction of LD turnover by lipophagy with TAG metabolism.
    [Google Scholar]
  49. 49.
    Fan J, Zhai Z, Yan C, Xu C 2015. Arabidopsis TRIGALACTOSYLDIACYLGLYCEROL5 interacts with TGD1, TGD2, and TGD4 to facilitate lipid transfer from the endoplasmic reticulum to plastids. Plant Cell 27:2941–55
    [Google Scholar]
  50. 50.
    Fatiha A 2019. Plant lipid metabolism. Advances in Lipid Metabolism RV Baez London: InTech Open https://doi.org/10.5772/intechopen.81355
    [Google Scholar]
  51. 51.
    Fernández-Santos R, Izquierdo Y, López A, Muñiz L, Martínez M et al. 2020. Protein profiles of lipid droplets during the hypersensitive defense response of Arabidopsis against Pseudomonas infection. Plant Cell Physiol. 61:1144–57
    [Google Scholar]
  52. 52.
    Feußner I, Kindl H. 1992. A lipoxygenase is the main lipid body protein in cucumber and soybean cotyledons during the stage of triglyceride mobilization. FEBS Lett. 298:223–25
    [Google Scholar]
  53. 53.
    Footitt S, Dietrich D, Fait A, Fernie AR, Holdsworth MJ et al. 2007. The COMATOSE ATP-binding cassette transporter is required for full fertility in Arabidopsis. Plant Physiol. 144:1467–80
    [Google Scholar]
  54. 54.
    Fulda M, Schnurr J, Abbadi A, Heinz E, Browse J 2004. Peroxisomal Acyl-CoA synthetase activity is essential for seedling development in Arabidopsis thaliana. Plant Cell 16:394–405
    [Google Scholar]
  55. 55.
    Gaydou EM, Ralaimanarivo A, Bianchini JP. 1993. Cyclopropanoic fatty acids of litchi (Litchi chinensis) seed oil. A reinvestigation. J. Agric. Food Chem. 41:886–90
    [Google Scholar]
  56. 56.
    Gidda SK, Park S, Pyc M, Yurchenko O, Cai Y et al. 2016. Lipid droplet-associated proteins (LDAPs) are required for the dynamic regulation of neutral lipid compartmentation in plant cells. Plant Physiol. 170:2052–71
    [Google Scholar]
  57. 57.
    Gidda SK, Shockey JM, Falcone M, Kim PK, Rothstein SJ et al. 2011. Hydrophobic-domain-dependent protein–protein interactions mediate the localization of GPAT enzymes to ER subdomains. Traffic 12:452–72
    [Google Scholar]
  58. 58.
    Gonzalez KL, Fleming WA, Kao Y-T, Wright ZJ, Venkova SV et al. 2017. Disparate peroxisome-related defects in Arabidopsis pex6 and pex26 mutants link peroxisomal retrotranslocation and oil body utilization. Plant J. 92:110–28
    [Google Scholar]
  59. 59.
    Goold H, Beisson F, Peltier G, Li-Beisson Y. 2015. Microalgal lipid droplets: composition, diversity, biogenesis and functions. Plant Cell Rep. 34:545–55
    [Google Scholar]
  60. 60.
    Greer MS, Cai Y, Gidda SK, Esnay N, Kretzschmar FK et al. 2020. SEIPIN isoforms interact with the membrane-tethering protein VAP27-1 for lipid droplet formation. Plant Cell 32:2932–50First evidence that a tethering complex—VAP27-1 in plants—might be involved in LD formation through its interaction with SEIPINs.
    [Google Scholar]
  61. 61.
    Guéguen N, Le Moigne D, Amato A, Salvaing J, Maréchal E 2021. Lipid droplets in unicellular photosynthetic stramenopiles. Front. Plant Sci. 12:639276
    [Google Scholar]
  62. 62.
    Hayashi H, De Bellis L, Hayashi Y, Nito K, Kato A et al. 2002. Molecular characterization of an Arabidopsis acyl-coenzyme A synthetase localized on glyoxysomal membranes. Plant Physiol. 130:2019–26
    [Google Scholar]
  63. 63.
    Hayashi Y, Hayashi M, Hayashi H, Hara-Nishimura I, Nishimura M. 2001. Direct interaction between glyoxysomes and lipid bodies in cotyledons of the Arabidopsis thaliana ped1 mutant. Protoplasma 218:83–94
    [Google Scholar]
  64. 64.
    Heneen WK, Karlsson G, Brismar K, Gummeson PO, Marttila S et al. 2008. Fusion of oil bodies in endosperm of oat grains. Planta 228:589–99
    [Google Scholar]
  65. 65.
    Herker E, Vieyres G, Beller M, Krahmer N, Bohnert M. 2021. Lipid droplet contact sites in health and disease. Trends Cell Biol. 31:345–58
    [Google Scholar]
  66. 66.
    Herman EM. 2008. Endoplasmic reticulum bodies: solving the insoluble. Curr. Opin. Plant Biol. 11:672–79
    [Google Scholar]
  67. 67.
    Hernandez ML, Lima-Cabello E, Alche JD, Martinez-Rivas JM, Castro AJ. 2020. Lipid composition and associated gene expression patterns during pollen germination and pollen tube growth in olive (Olea europaea L.). Plant Cell Physiol. 61:1348–64
    [Google Scholar]
  68. 68.
    Higashi Y, Okazaki Y, Myouga F, Shinozaki K, Saito K. 2015. Landscape of the lipidome and transcriptome under heat stress in Arabidopsis thaliana. Sci. Rep. 5:10533
    [Google Scholar]
  69. 69.
    Higashi Y, Okazaki Y, Takano K, Myouga F, Shinozaki K et al. 2018. HEAT INDUCIBLE LIPASE1 remodels chloroplastic monogalactosyldiacylglycerol by liberating α-linolenic acid in Arabidopsis leaves under heat stress. Plant Cell 30:1887–905
    [Google Scholar]
  70. 70.
    Horn PJ, James CN, Gidda SK, Kilaru A, Dyer JM et al. 2013. Identification of a new class of lipid droplet-associated proteins in plants. Plant Physiol. 162:1926–36First identification of LDAP as a major LD protein in TAG-containing LDs from nonseed tissues.
    [Google Scholar]
  71. 71.
    Hsiao ES, Tzen JTC. 2011. Ubiquitination of oleosin-H and caleosin in sesame oil bodies after seed germination. Plant Physiol. Biochem. 49:77–81
    [Google Scholar]
  72. 72.
    Hsieh K, Huang AH. 2004. Endoplasmic reticulum, oleosins, and oils in seeds and tapetum cells. Plant Physiol. 136:3427–34
    [Google Scholar]
  73. 73.
    Hsieh K, Huang AHC. 2005. Lipid-rich tapetosomes in Brassica tapetum are composed of oleosin-coated oil droplets and vesicles, both assembled in and then detached from the endoplasmic reticulum. Plant J. 43:889–99
    [Google Scholar]
  74. 74.
    Hu Q, Sommerfeld M, Jarvis E, Ghirardi M, Posewitz M et al. 2008. Microalgal triacylglycerols as feedstocks for biofuel production: perspectives and advances. Plant J. 54:621–39
    [Google Scholar]
  75. 75.
    Huang AHC. 2018. Plant lipid droplets and their associated proteins: potential for rapid advances. Plant Physiol. 176:1894–918
    [Google Scholar]
  76. 76.
    Huang CY, Huang AHC. 2017. Unique motifs and length of hairpin in oleosin target the cytosolic side of endoplasmic reticulum and budding lipid droplet. Plant Physiol. 174:2248–60
    [Google Scholar]
  77. 77.
    Hughes MA, Goad LJ. 1983. The hydrolysis of steryl esters during the germination of barley seed. Biochem. Soc. Trans. 11:588–99
    [Google Scholar]
  78. 78.
    Ischebeck T. 2016. Lipids in pollen—They are different. Biochim. Biophys. Acta Mol. Cell Biol. Lipids 1861:1315–28
    [Google Scholar]
  79. 79.
    Ischebeck T, Krawczyk HE, Mullen RT, Dyer JM, Chapman KD. 2020. Lipid droplets in plants and algae: distribution, formation, turnover and function. Semin. Cell Dev. Biol. 108:82–93
    [Google Scholar]
  80. 80.
    Ischebeck T, Zbierzak AM, Kanwischer M, Dörmann P. 2006. A salvage pathway for phytol metabolism in Arabidopsis. J. Biol. Chem. 281:2470–77
    [Google Scholar]
  81. 81.
    Iven T, Herrfurth C, Hornung E, Heilmann M, Hofvander P et al. 2013. Wax ester profiling of seed oil by nano-electrospray ionization tandem mass spectrometry. Plant Methods 9:24
    [Google Scholar]
  82. 82.
    Iven T, Hornung E, Heilmann M, Feussner I. 2016. Synthesis of oleyl oleate wax esters in Arabidopsis thaliana and Camelina sativa seed oil. Plant Biotechnol. J. 14:252–59
    [Google Scholar]
  83. 83.
    Jin K, Xia H, Liu Y, Li J, Du G et al. 2022. Compartmentalization and transporter engineering strategies for terpenoid synthesis. Microb. Cell Fact. 21:92
    [Google Scholar]
  84. 84.
    Jolivet P, Roux E, D'Andrea S, Davanture M, Negroni L et al. 2004. Protein composition of oil bodies in Arabidopsis thaliana ecotype WS. Plant Physiol. Biochem. 42:501–9
    [Google Scholar]
  85. 85.
    Kang BH, Anderson CT, Arimura SI, Bayer E, Bezanilla M et al. 2022. A glossary of plant cell structures: current insights and future questions. Plant Cell 34:10–52
    [Google Scholar]
  86. 86.
    Kawiński A, Miklaszewska M, Stelter S, Głab B, Banaś A. 2021. Lipases of germinating jojoba seeds efficiently hydrolyze triacylglycerols and wax esters and display wax ester-synthesizing activity. BMC Plant Biol. 21:50
    [Google Scholar]
  87. 87.
    Kelly AA, Quettier A-L, Shaw E, Eastmond PJ 2011. Seed storage oil mobilization is important but not essential for germination or seedling establishment in Arabidopsis. Plant Physiol. 157:866–75
    [Google Scholar]
  88. 88.
    Kelly AA, van Erp H, Quettier AL, Shaw E, Menard G et al. 2013. The SUGAR-DEPENDENT1 lipase limits triacylglycerol accumulation in vegetative tissues of Arabidopsis. Plant Physiol. 162:1282–89
    [Google Scholar]
  89. 89.
    Kim HU. 2020. Lipid metabolism in plants. Plants 9:871
    [Google Scholar]
  90. 90.
    Kim HU, Hsieh K, Ratnayake C, Huang AHC. 2002. A novel group of oleosins is present inside the pollen of Arabidopsis. J. Biol. Chem. 277:22677–84
    [Google Scholar]
  91. 91.
    Kim RJ, Kim HJ, Shim D, Suh MC. 2016. Molecular and biochemical characterizations of the monoacylglycerol lipase gene family of Arabidopsis thaliana. Plant J. 85:758–71
    [Google Scholar]
  92. 92.
    Kong F, Yamaoka Y, Ohama T, Lee Y, Li-Beisson Y. 2019. Molecular genetic tools and emerging synthetic biology strategies to increase cellular oil content in Chlamydomonas reinhardtii. Plant Cell Physiol. 60:1184–96
    [Google Scholar]
  93. 93.
    Krawczyk HE, Sun S, Doner NM, Yan Q, Lim MSS et al. 2022. SEED LIPID DROPLET PROTEIN1, SEED LIPID DROPLET PROTEIN2, and LIPID DROPLET PLASMA MEMBRANE ADAPTOR mediate lipid droplet–plasma membrane tethering. Plant Cell 34:2424–48
    [Google Scholar]
  94. 94.
    Kretzschmar FK, Doner NM, Krawczyk HE, Scholz P, Schmitt K et al. 2020. Identification of low-abundance lipid droplet proteins in seeds and seedlings. Plant Physiol. 182:1326–45
    [Google Scholar]
  95. 95.
    Kretzschmar FK, Mengel LA, Müller AO, Schmitt K, Blersch KF et al. 2018. PUX10 is a lipid droplet-localized scaffold protein that interacts with CELL DIVISION CYCLE48 and is involved in the degradation of lipid droplet proteins. Plant Cell 30:2137–60Identified a novel LD protein, PUX10, and provided evidence to support its role in the ubiquitin-mediated degradation of LD proteins as part of LD degradation during postgerminative seedling growth.
    [Google Scholar]
  96. 96.
    Kunz H-H, Scharnewski M, Feussner K, Feussner I, Flügge U-I et al. 2009. The ABC transporter PXA1 and peroxisomal β-oxidation are vital for metabolism in mature leaves of Arabidopsis during extended darkness. Plant Cell 21:2733–49
    [Google Scholar]
  97. 97.
    Laibach N, Hillebrand A, Twyman RM, Prufer D, Schulze Gronover C. 2015. Identification of a Taraxacum brevicorniculatum rubber elongation factor protein that is localized on rubber particles and promotes rubber biosynthesis. Plant J. 82:609–20
    [Google Scholar]
  98. 98.
    Lévesque-Lemay M, Chabot D, Hubbard K, Chan JK, Miller S, Robert LS. 2016. Tapetal oleosins play an essential role in tapetosome formation and protein relocation to the pollen coat. New Phytol. 209:691–704
    [Google Scholar]
  99. 99.
    Leyland B, Zarka A, Didi-Cohen S, Boussiba S, Khozin-Goldberg I. 2020. High resolution proteome of lipid droplets isolated from the pennate diatom Phaeodactylum tricornutum (Bacillariophyceae) strain pt4 provides mechanistic insights into complex intracellular coordination during nitrogen deprivation. J. Phycol. 56:1642–63
    [Google Scholar]
  100. 100.
    Li F, Han X, Guan H, Xu MC, Dong YX, Gao X-Q. 2022. PALD encoding a lipid droplet-associated protein is critical for the accumulation of lipid droplets and pollen longevity in Arabidopsis. New Phytol. 235:204–19
    [Google Scholar]
  101. 101.
    Li N, Meng H, Li S, Zhang Z, Zhao X et al. 2020. Two plastid fatty acid exporters contribute to seed oil accumulation in Arabidopsis. Plant Physiol. 182:1910–19
    [Google Scholar]
  102. 102.
    Li-Beisson Y, Kong F, Wang P, Lee Y, Kang B-H. 2021. The disassembly of lipid droplets in Chlamydomonas. New Phytol. 231:1359–64
    [Google Scholar]
  103. 103.
    Li-Beisson Y, Shorrosh B, Beisson F, Andersson MX, Arondel V et al. 2013. Acyl-lipid metabolism. Arabidopsis Book 11:e0161
    [Google Scholar]
  104. 104.
    Li-Beisson Y, Thelen JJ, Fedosejevs E, Harwood JL. 2019. The lipid biochemistry of eukaryotic algae. Prog. Lipid Res. 74:31–68
    [Google Scholar]
  105. 105.
    Lin LJ, Tai SS, Peng CC, Tzen JT. 2002. Steroleosin, a sterol-binding dehydrogenase in seed oil bodies. Plant Physiol. 128:1200–11
    [Google Scholar]
  106. 106.
    Lippold F, vom Dorp K, Abraham M, Holzl G, Wewer V et al. 2012. Fatty acid phytyl ester synthesis in chloroplasts of Arabidopsis. Plant Cell 24:2001–14
    [Google Scholar]
  107. 107.
    Listenberger LL, Han X, Lewis SE, Cases S, Farese RV Jr. et al. 2003. Triglyceride accumulation protects against fatty acid-induced lipotoxicity. PNAS 100:3077–82
    [Google Scholar]
  108. 108.
    Liu WX, Liu HL, Qu LQ. 2013. Embryo-specific expression of soybean oleosin altered oil body morphogenesis and increased lipid content in transgenic rice seeds. Theor. Appl. Genet. 126:2289–97
    [Google Scholar]
  109. 109.
    Liu Y, Li J. 2014. Endoplasmic reticulum-mediated protein quality control in Arabidopsis. Front. Plant Sci. 5:162
    [Google Scholar]
  110. 110.
    Lundquist PK, Poliakov A, Bhuiyan NH, Zybailov B, Sun Q, van Wijk KJ. 2012. The functional network of the Arabidopsis plastoglobule proteome based on quantitative proteomics and genome-wide coexpression analysis. Plant Physiol. 158:1172–92
    [Google Scholar]
  111. 111.
    Lundquist PK, Shivaiah KK, Espinoza-Corral R. 2020. Lipid droplets throughout the evolutionary tree. Prog. Lipid Res. 78:101029
    [Google Scholar]
  112. 112.
    Lunn D, Wallis JG, Browse J. 2018. Overexpression of Seipin1 increases oil in hydroxy fatty acid-accumulating seeds. Plant Cell Physiol. 59:205–14
    [Google Scholar]
  113. 113.
    Lunn D, Wallis JG, Browse J. 2022. A multigene approach secures hydroxy fatty acid production in Arabidopsis. J. Exp. Bot. 73:2875–88
    [Google Scholar]
  114. 114.
    Miquel M, Trigui G, d'Andréa S, Kelemen Z, Baud S et al. 2014. Specialization of oleosins in oil body dynamics during seed development in Arabidopsis seeds. Plant Physiol. 164:1866–78Higher-order oleosin mutants demonstrate their importance in the stability of LD structural organization during seed development.
    [Google Scholar]
  115. 115.
    Moellering ER, Benning C. 2010. RNA interference silencing of a major lipid droplet protein affects lipid droplet size in Chlamydomonas reinhardtii. Eukaryot. Cell 9:97–106
    [Google Scholar]
  116. 116.
    Morabito C, Bournaud C, Maës C, Schuler M, Aiese Cigliano R et al. 2019. The lipid metabolism in thraustochytrids. Prog. Lipid Res. 76:101007
    [Google Scholar]
  117. 117.
    Mueller SP, Krause DM, Mueller MJ, Fekete A. 2015. Accumulation of extra-chloroplastic triacylglycerols in Arabidopsis seedlings during heat acclimation. J. Exp. Bot. 66:4517–26
    [Google Scholar]
  118. 118.
    Müller AO, Ischebeck T. 2018. Characterization of the enzymatic activity and physiological function of the lipid droplet-associated triacylglycerol lipase AtOBL1. New Phytol. 217:1062–76
    [Google Scholar]
  119. 119.
    Murphy DJ. 2016. Plant storage lipids. eLS Hoboken, NJ: Wiley https://doi.org/10.1002/9780470015902.a0001918.pub3
    [Google Scholar]
  120. 120.
    Nguyen HM, Baudet M, Cuiné S, Adriano J-M, Barthe D et al. 2011. Proteomic profiling of oil bodies isolated from the unicellular green microalga Chlamydomonas reinhardtii: with focus on proteins involved in lipid metabolism. Proteomics 11:4266–73
    [Google Scholar]
  121. 121.
    Oh SK, Kang H, Shin DH, Yang J, Chow KS et al. 1999. Isolation, characterization, and functional analysis of a novel cDNA clone encoding a small rubber particle protein from Hevea brasiliensis. J. Biol. Chem. 274:17132–38
    [Google Scholar]
  122. 122.
    Ohlrogge J, Browse J. 1995. Lipid biosynthesis. Plant Cell 7:957–70
    [Google Scholar]
  123. 123.
    Ohlrogge J, Thrower N, Mhaske V, Stymne S, Baxter M et al. 2018. PlantFAdb: a resource for exploring hundreds of plant fatty acid structures synthesized by thousands of plants and their phylogenetic relationships. Plant J. 96:1299–308
    [Google Scholar]
  124. 124.
    Olzmann JA, Carvalho P. 2019. Dynamics and functions of lipid droplets. Nat. Rev. Mol. Cell Biol. 20:137–55
    [Google Scholar]
  125. 125.
    Ortiz R, Geleta M, Gustafsson C, Lager I, Hofvander P et al. 2020. Oil crops for the future. Curr. Opin. Plant Biol. 56:181–89
    [Google Scholar]
  126. 126.
    Parker ML, Murphy GJP. 1981. Oleosomes in flag leaves of wheat; their distribution, composition and fate during senesence and rust-infection. Planta 152:36–43
    [Google Scholar]
  127. 127.
    Partridge M, Murphy DJ. 2009. Roles of a membrane-bound caleosin and putative peroxygenase in biotic and abiotic stress responses in Arabidopsis. Plant Physiol. Biochem. 47:796–806
    [Google Scholar]
  128. 128.
    Peled E, Leu S, Zarka A, Weiss M, Pick U et al. 2011. Isolation of a novel oil globule protein from the green alga Haematococcus pluvialis (Chlorophyceae). Lipids 46:851–61
    [Google Scholar]
  129. 129.
    Peramuna A, Bae H, Quinonero Lopez C, Fromberg A, Petersen B, Simonsen HT 2020. Connecting moss lipid droplets to patchoulol biosynthesis. PLOS ONE 15:e0243620
    [Google Scholar]
  130. 130.
    Petan T, Jarc E, Jusović M. 2018. Lipid droplets in cancer: guardians of fat in a stressful world. Molecules 23:1941
    [Google Scholar]
  131. 131.
    Poxleitner M, Rogers SW, Samuels AL, Browse J, Rogers JC. 2006. A role for caleosin in degradation of oil-body storage lipid during seed germination. Plant J. 47:917–33
    [Google Scholar]
  132. 132.
    Prevost C, Sharp ME, Kory N, Lin Q, Voth GA et al. 2018. Mechanism and determinants of amphipathic helix-containing protein targeting to lipid droplets. Dev. Cell 44:73–86.e4
    [Google Scholar]
  133. 133.
    Price AM, Doner NM, Gidda SK, Jambunathan S, James CN et al. 2020. Mouse Fat-Specific Protein 27 (FSP27) expressed in plant cells localizes to lipid droplets and promotes lipid droplet accumulation and fusion. Biochimie 169:41–53
    [Google Scholar]
  134. 134.
    Pyc M, Cai Y, Gidda SK, Yurchenko O, Park S et al. 2017. Arabidopsis lipid droplet-associated protein (LDAP)–interacting protein (LDIP) influences lipid droplet size and neutral lipid homeostasis in both leaves and seeds. Plant J. 92:1182–201
    [Google Scholar]
  135. 135.
    Pyc M, Cai Y, Greer MS, Yurchenko O, Chapman KD et al. 2017. Turning over a new leaf in lipid droplet biology. Trends Plant Sci. 22:596–609
    [Google Scholar]
  136. 136.
    Pyc M, Gidda SK, Seay D, Esnay N, Kretzschmar FK et al. 2021. LDIP cooperates with SEIPIN and LDAP to facilitate lipid droplet biogenesis in Arabidopsis. Plant Cell 33:3076–103First demonstration of a functional interaction between SEIPIN and LDIP to support LD formation.
    [Google Scholar]
  137. 137.
    Qu R, Wang SM, Lin YH, Vance VB, Huang AH. 1986. Characteristics and biosynthesis of membrane proteins of lipid bodies in the scutella of maize (Zea mays L.). Biochem. J. 235:57–65
    [Google Scholar]
  138. 138.
    Rahman F, Hassan M, Rosli R, Almousally I, Hanano A, Murphy DJ. 2018. Evolutionary and genomic analysis of the caleosin/peroxygenase (CLO/PXG) gene/protein families in the Viridiplantae. PLOS ONE 13:e0196669
    [Google Scholar]
  139. 139.
    Rajangam AS, Gidda SK, Craddock C, Mullen RT, Dyer JM, Eastmond PJ. 2013. Molecular characterization of the fatty alcohol oxidation pathway for wax-ester mobilization in germinated jojoba seeds. Plant Physiol. 161:72–80
    [Google Scholar]
  140. 140.
    Reed J, Osbourn A 2018. Engineering terpenoid production through transient expression in Nicotiana benthamiana. Plant Cell Rep. 37:1431–41
    [Google Scholar]
  141. 141.
    Rowe ER, Mimmack ML, Barbosa AD, Haider A, Isaac I et al. 2016. Conserved amphipathic helices mediate lipid droplet targeting of perilipins 1–3. J. Biol. Chem. 291:6664–78
    [Google Scholar]
  142. 142.
    Sadre R, Kuo P, Chen J, Yang Y, Banerjee A et al. 2019. Cytosolic lipid droplets as engineered organelles for production and accumulation of terpenoid biomaterials in leaves. Nat. Commun. 10:853Biotechnology application highlighting LDs as locations to store high-value, engineered terpenes.
    [Google Scholar]
  143. 143.
    Schmidt MA, Herman EM. 2008. Suppression of soybean oleosin produces micro-oil bodies that aggregate into oil body/ER complexes. Mol. Plant 1:910–24
    [Google Scholar]
  144. 144.
    Scholz P, Chapman KD, Mullen RT, Ischebeck T. 2022. Finding new friends and revisiting old ones—how plant lipid droplets connect with other subcellular structures. New Phytol. 236:833–38
    [Google Scholar]
  145. 145.
    Selinski J, Scheibe R. 2014. Pollen tube growth: Where does the energy come from?. Plant Signal. Behav. 9:e977200
    [Google Scholar]
  146. 146.
    Shimada TL, Shimada T, Okazaki Y, Higashi Y, Saito K et al. 2019. HIGH STEROL ESTER 1 is a key factor in plant sterol homeostasis. Nat. Plants 5:1154–66
    [Google Scholar]
  147. 147.
    Shimada TL, Shimada T, Takahashi H, Fukao Y, Hara-Nishimura I. 2008. A novel role for oleosins in freezing tolerance of oilseeds in Arabidopsis thaliana. Plant J. 55:798–809
    [Google Scholar]
  148. 148.
    Shimada TL, Takano Y, Hara-Nishimura I. 2015. Oil body-mediated defense against fungi: from tissues to ecology. Plant Signal. Behav. 10:e989036
    [Google Scholar]
  149. 149.
    Shimada TL, Takano Y, Shimada T, Fujiwara M, Fukao Y et al. 2014. Leaf oil body functions as a subcellular factory for the production of a phytoalexin in Arabidopsis. Plant Physiol. 164:105–18
    [Google Scholar]
  150. 150.
    Sibi G, Shetty V, Mokashi K. 2016. Enhanced lipid productivity approaches in microalgae as an alternate for fossil fuels—a review. J. Energy Inst. 89:330–34
    [Google Scholar]
  151. 151.
    Siloto RMP, Findlay K, Lopez-Villalobos A, Yeung EC, Nykiforuk CL, Moloney MM. 2006. The accumulation of oleosins determines the size of seed oilbodies in Arabidopsis. Plant Cell 18:1961–74
    [Google Scholar]
  152. 152.
    Slee JA, Levine TP. 2019. Systematic prediction of FFAT motifs across eukaryote proteomes identifies nucleolar and eisosome proteins with the predicted capacity to form bridges to the endoplasmic reticulum. Contact 2. https://doi.org/10.1177/2515256419883136
  153. 153.
    Stark C, Breitkreutz B-J, Reguly T, Boucher L, Breitkreutz A, Tyers M. 2006. BioGRID: a general repository for interaction datasets. Nucleic Acids Res. 34:D535–39
    [Google Scholar]
  154. 154.
    Steinmuller D, Tevini M. 1985. Composition and function of plastoglobuli: I. Isolation and purification from chloroplasts and chromoplasts. Planta 163:201–7
    [Google Scholar]
  155. 155.
    Sturtevant D, Lu S, Zhou ZW, Shen Y, Wang S et al. 2020. The genome of jojoba (Simmondsia chinensis): a taxonomically isolated species that directs wax ester accumulation in its seeds. Sci. Adv. 6:eaay3240
    [Google Scholar]
  156. 156.
    Sui X, Arlt H, Brock KP, Lai ZW, DiMaio F et al. 2018. Cryo-electron microscopy structure of the lipid droplet-formation protein seipin. J. Cell Biol. 217:4080–91
    [Google Scholar]
  157. 157.
    Sun Y, Liu B, Xue J, Wang X, Cui H et al. 2022. Critical metabolic pathways and genes cooperate for epoxy fatty acid-enriched oil production in developing seeds of Vernonia galamensis, an industrial oleaginous plant. Biotechnol. Biofuels Bioprod. 15:21
    [Google Scholar]
  158. 158.
    Suzuki T, Tsunekawa S, Koizuka C, Yamamoto K, Imamura J et al. 2013. Development and disintegration of tapetum-specific lipid-accumulating organelles, elaioplasts and tapetosomes, in Arabidopsis thaliana and Brassica napus. Plant Sci. 207:25–36
    [Google Scholar]
  159. 159.
    Takagi M, Karseno, Yoshida T. 2006. Effect of salt concentration on intracellular accumulation of lipids and triacylglyceride in marine microalgae Dunaliella cells. J. Biosci. Bioeng. 101:223–26
    [Google Scholar]
  160. 160.
    Tan W-J, Yang Y-C, Zhou Y, Huang L-P, Xu L et al. 2018. DIACYLGLYCEROL ACYLTRANSFERASE and DIACYLGLYCEROL KINASE modulate triacylglycerol and phosphatidic acid production in the plant response to freezing stress. Plant Physiol. 177:1303–18
    [Google Scholar]
  161. 161.
    Tang G, Xu P, Ma W, Wang F, Liu Z et al. 2018. Seed-specific expression of AtLEC1 increased oil content and altered fatty acid composition in seeds of peanut (Arachis hypogaea L.). Front. Plant Sci. 9:260
    [Google Scholar]
  162. 162.
    Taurino M, Costantini S, De Domenico S, Stefanelli F, Ruano G et al. 2018. SEIPIN proteins mediate lipid droplet biogenesis to promote pollen transmission and reduce seed dormancy. Plant Physiol. 176:1531–46
    [Google Scholar]
  163. 163.
    Tevini M, Steinmuller D. 1985. Composition and function of plastoglobuli: II. Lipid composition of leaves and plastoglobuli during beech leaf senescence. Planta 163:91–96
    [Google Scholar]
  164. 164.
    Thazar-Poulot N, Miquel M, Fobis-Loisy I, Gaude T 2015. Peroxisome extensions deliver the Arabidopsis SDP1 lipase to oil bodies. PNAS 112:4158–63Impressive article that describes the extension of peroxisome membranes to deliver SDP1 from peroxisomes to LDs for lipid mobilization.
    [Google Scholar]
  165. 165.
    Thiam AR, Ikonen E. 2021. Lipid droplet nucleation. Trends Cell Biol. 31:108–18
    [Google Scholar]
  166. 166.
    Thompson J, Madey E, Nowack L, Froese C. 1998. Distinguishable populations of lipid particles of membrane origin in petals and stems. Advances in Plant Lipid Research: The Proceedings of the 13th International Symposium on Plant Lipids, Held at Sevilla, Spain, July 1998 J Sánchez, E Cerdá-Olmedo, E Martínez-Force 377–379. Sevilla, Spain: Univer. Sevilla
    [Google Scholar]
  167. 167.
    Thompson PA, Guo M-x, Harrison PJ, Whyte JNC. 1992. Effects of variation in temperature. II. On the fatty acid composition of eight species of marine phytoplankton. J. Phycol. 28:488–97
    [Google Scholar]
  168. 168.
    Trinh D-C, Lavenus J, Goh T, Boutte Y, Drogue Q et al. 2019. PUCHI regulates very long chain fatty acid biosynthesis during lateral root and callus formation. PNAS 116:14325–30
    [Google Scholar]
  169. 169.
    Tsai CH, Zienkiewicz K, Amstutz CL, Brink BG, Warakanont J et al. 2015. Dynamics of protein and polar lipid recruitment during lipid droplet assembly in Chlamydomonas reinhardtii. Plant J. 83:650–60
    [Google Scholar]
  170. 170.
    Tzen JTC 2012. Integral proteins in plant oil bodies. ISRN Bot. 2012:173954
    [Google Scholar]
  171. 171.
    Usher S, Han L, Haslam RP, Michaelson LV, Sturtevant D et al. 2017. Tailoring seed oil composition in the real world: optimising omega-3 long chain polyunsaturated fatty acid accumulation in transgenic Camelina sativa. Sci. Rep. 7:6570
    [Google Scholar]
  172. 172.
    van Erp H, Bryant FM, Martin-Moreno J, Michaelson LV, Eastmond PJ. 2021. Production of the infant formula ingredient 1,3-olein-2-palmitin in Arabidopsis thaliana seeds. Metab. Eng. 67:67–74
    [Google Scholar]
  173. 173.
    Van Vooren G, Le Grand F, Legrand J, Cuiné S, Peltier G, Pruvost J 2012. Investigation of fatty acids accumulation in Nannochloropsis oculata for biodiesel application. Bioresour. Technol. 124:421–32
    [Google Scholar]
  174. 174.
    van Zutphen T, Todde V, de Boer R, Kreim M, Hofbauer HF et al. 2014. Lipid droplet autophagy in the yeast Saccharomyces cerevisiae. Mol. Biol. Cell 25:290–301
    [Google Scholar]
  175. 175.
    Vanhercke T, Divi UK, El Tahchy A, Liu Q, Mitchell M et al. 2017. Step changes in leaf oil accumulation via iterative metabolic engineering. Metab. Eng. 39:237–46
    [Google Scholar]
  176. 176.
    Vanhercke T, Dyer JM, Mullen RT, Kilaru A, Rahman MM et al. 2019. Metabolic engineering for enhanced oil in biomass. Prog. Lipid Res. 74:103–29
    [Google Scholar]
  177. 177.
    Vanhercke T, El Tahchy A, Liu Q, Zhou XR, Shrestha P et al. 2014. Metabolic engineering of biomass for high energy density: oilseed-like triacylglycerol yields from plant leaves. Plant Biotechnol. J. 12:231–39Remarkable enhancement of LDs and TAGs in vegetative biomass; demonstrates the feasibility of producing large amounts of oils in plant tissues outside of seeds by the “push, pull and protect” strategy to package TAGs in cytoplasmic lipid droplets.
    [Google Scholar]
  178. 178.
    Vieler A, Brubaker SB, Vick B, Benning C. 2012. A lipid droplet protein of Nannochloropsis with functions partially analogous to plant oleosins. Plant Physiol. 158:1562–69
    [Google Scholar]
  179. 179.
    Vlahakis C, Hazebroek J. 2000. Phytosterol accumulation in canola, sunflower, and soybean oils: effects of genetics, planting location, and temperature. J. Am. Oil Chem. Soc. 77:49–53
    [Google Scholar]
  180. 180.
    Wadeesirisak K, Castano S, Berthelot K, Vaysse L, Bonfils F et al. 2017. Rubber particle proteins REF1 and SRPP1 interact differently with native lipids extracted from Hevea brasiliensis latex. Biochim. Biophys. Acta Biomembr. 1859:201–10
    [Google Scholar]
  181. 181.
    Walther TC, Chung J, Farese RV Jr. 2017. Lipid droplet biogenesis. Annu. Rev. Cell Dev. Biol. 33:491–510
    [Google Scholar]
  182. 182.
    Wang P, Richardson C, Hawkins TJ, Sparkes I, Hawes C, Hussey PJ. 2016. Plant VAP27 proteins: domain characterization, intracellular localization and role in plant development. New Phytol. 210:1311–26
    [Google Scholar]
  183. 183.
    Wang X, Hao T-B, Balamurugan S, Yang W-D, Liu J-S et al. 2017. A lipid droplet-associated protein involved in lipid droplet biogenesis and triacylglycerol accumulation in the oleaginous microalga Phaeodactylum tricornutum. Algal Res. 26:215–24
    [Google Scholar]
  184. 184.
    Wang X, Wei H, Mao X, Liu J. 2019. Proteomics analysis of lipid droplets from the oleaginous alga Chromochloris zofingiensis reveals novel proteins for lipid metabolism. Genom. Proteom. Bioinform. 17:260–72
    [Google Scholar]
  185. 185.
    Wang X, Wei W, Li N-J, Yuan W, Ding Y et al. 2018. Heterogeneous expression of human PNPLA3 triggers algal lipid accumulation and lipid droplet enlargement. Algal Res. 31:276–81
    [Google Scholar]
  186. 186.
    Wright ZJ, Bartel B. 2020. Peroxisomes form intralumenal vesicles with roles in fatty acid catabolism and protein compartmentalization in Arabidopsis. Nat. Commun. 11:6221
    [Google Scholar]
  187. 187.
    Yan R, Qian H, Lukmantara I, Gao M, Du X et al. 2018. Human SEIPIN binds anionic phospholipids. Dev. Cell 47:248–56.e4
    [Google Scholar]
  188. 188.
    Yoneda K, Yoshida M, Suzuki I, Watanabe MM. 2016. Identification of a major lipid droplet protein in a marine diatom Phaeodactylum tricornutum. Plant Cell Physiol. 57:397–406
    [Google Scholar]
  189. 189.
    You Z, Zhang Q, Peng Z, Miao X. 2019. Lipid droplets mediate salt stress tolerance in Parachlorella kessleri. Plant Physiol. 181:510–26
    [Google Scholar]
  190. 190.
    Yu D, Hornung E, Iven T, Feussner I. 2018. High-level accumulation of oleyl oleate in plant seed oil by abundant supply of oleic acid substrates to efficient wax ester synthesis enzymes. Biotechnol. Biofuels 11:53
    [Google Scholar]
  191. 191.
    Yu L, Fan J, Zhou C, Xu C. 2021. Sterols are required for the coordinated assembly of lipid droplets in developing seeds. Nat. Commun. 12:5598
    [Google Scholar]
  192. 192.
    Zbierzak AM, Kanwischer M, Wille C, Vidi P-A, Giavalisco P et al. 2009. Intersection of the tocopherol and plastoquinol metabolic pathways at the plastoglobule. Biochem. J. 425:389–99
    [Google Scholar]
  193. 193.
    Zhai Z, Liu H, Shanklin J. 2021. Ectopic expression of OLEOSIN 1 and inactivation of GBSS1 have a synergistic effect on oil accumulation in plant leaves. Plants 10:513
    [Google Scholar]
  194. 194.
    Zhang C, Qu Y, Lian Y, Chapman M, Chapman N et al. 2020. A new insight into the mechanism for cytosolic lipid droplet degradation in senescent leaves. Physiol. Plant. 168:835–44
    [Google Scholar]
  195. 195.
    Zhao P, Zhou XM, Zhao LL, Cheung AY, Sun MX. 2020. Autophagy-mediated compartmental cytoplasmic deletion is essential for tobacco pollen germination and male fertility. Autophagy 16:2180–92
    [Google Scholar]
  196. 196.
    Zhou X, Chen X, Du Z, Zhang Y, Zhang W et al. 2019. Terpenoid esters are the major constituents from leaf lipid droplets of Camellia sinensis. Front. Plant Sci. 10:179
    [Google Scholar]
  197. 197.
    Zhu L-H, Krens F, Smith MA, Li X, Qi W et al. 2016. Dedicated industrial oilseed crops as metabolic engineering platforms for sustainable industrial feedstock production. Sci. Rep. 6:22181
    [Google Scholar]
  198. 198.
    Zienkiewicz A, Zienkiewicz K, Rejon JD, Rodriguez-Garcia MI, Castro AJ. 2013. New insights into the early steps of oil body mobilization during pollen germination. J. Exp. Bot. 64:293–302
    [Google Scholar]
  199. 199.
    Zienkiewicz K, Castro AJ, de Dios Alché J, Zienkiewicz A, Suárez C, Rodriguez-Garcia MI. 2010. Identification and localization of a caleosin in olive (Olea europaea L.) pollen during in vitro germination. J. Exp. Bot. 61:1537–46
    [Google Scholar]
  200. 200.
    Zienkiewicz K, Zienkiewicz A. 2020. Degradation of lipid droplets in plants and algae—right time, many paths, one goal. Front. Plant Sci. 11:579019
    [Google Scholar]
  201. 201.
    Zienkiewicz K, Zienkiewicz A, Rodríguez-García MI, Castro AJ. 2011. Characterization of a caleosin expressed during olive (Olea europaea L.) pollen ontogeny. BMC Plant Biol. 11:122
    [Google Scholar]
  202. 202.
    Zolman BK, Silva ID, Bartel B 2001. The Arabidopsis pxa1 mutant is defective in an ATP-binding cassette transporter-like protein required for peroxisomal fatty acid β-oxidation. Plant Physiol. 127:1266–78
    [Google Scholar]
  203. 203.
    Zouiouich M, Di Mattia T, Martinet A, Eichler J, Wendling C et al. 2022. MOSPD2 is an endoplasmic reticulum-lipid droplet tether functioning in LD homeostasis. J. Cell Biol. 221:e202110044
    [Google Scholar]
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