Oils in the form of triacylglycerols are the most abundant energy-dense storage compounds in eukaryotes, and their metabolism plays a key role in cellular energy balance, lipid homeostasis, growth, and maintenance. Plants accumulate oils primarily in seeds and fruits. Plant oils are used for food and feed and, increasingly, as feedstocks for biodiesel and industrial chemicals. Although plant vegetative tissues do not accumulate significant levels of triacylglycerols, they possess a high capacity for their synthesis, storage, and metabolism. The development of plants that accumulate oil in vegetative tissues presents an opportunity for expanded production of triacylglycerols as a renewable and sustainable bioenergy source. Here, we review recent progress in the understanding of triacylglycerol synthesis, turnover, storage, and function in leaves and discuss emerging genetic engineering strategies targeted at enhancing triacylglycerol accumulation in biomass crops. Such plants could potentially be modified to produce oleochemical feedstocks or nutraceuticals.


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Literature Cited

  1. Adeyo O, Horn PJ, Lee SK, Binns DD, Chandrahas A. 1.  et al. 2011. The yeast lipin orthologue Pah1p is important for biogenesis of lipid droplets. J. Cell Biol. 192:1043–55 [Google Scholar]
  2. Andre C, Haslam RP, Shanklin J. 2.  2012. Feedback regulation of plastidic acetyl-CoA carboxylase by 18:1-acyl carrier protein in Brassica napus. PNAS 109:10107–12Identified biochemical feedback for plastidial ACCase. [Google Scholar]
  3. Andrianov V, Borisjuk N, Pogrebnyak N, Brinker A, Dixon J. 3.  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. 8:277–87Demonstrates the synergistic effects that expressing DGAT and LEC2 has on oil accumulation in tobacco leaves. [Google Scholar]
  4. Athenstaedt K, Daum G. 4.  2003. YMR313c/TGL3 encodes a novel triacylglycerol lipase located in lipid particles of Saccharomyces cerevisiae. J. Biol. Chem. 278:23317–23 [Google Scholar]
  5. Athenstaedt K, Daum G. 5.  2005. Tgl4p and Tgl5p, two triacylglycerol lipases of the yeast Saccharomyces cerevisiae are localized to lipid particles. J. Biol. Chem. 280:37301–9 [Google Scholar]
  6. Aubert Y, Vile D, Pervent M, Aldon D, Ranty B. 6.  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]
  7. Barbosa AD, Savage DB, Siniossoglou S. 7.  2015. Lipid droplet-organelle interactions: emerging roles in lipid metabolism. Curr. Opin. Cell Biol. 35:91–97 [Google Scholar]
  8. Barthole G, Lepiniec L, Rogowsky PM, Baud S. 8.  2012. Controlling lipid accumulation in cereal grains. Plant Sci. 185–186:33–39 [Google Scholar]
  9. Bates PD, Browse J. 9.  2011. The pathway of triacylglycerol synthesis through phosphatidylcholine in Arabidopsis produces a bottleneck for the accumulation of unusual fatty acids in transgenic seeds. Plant J. 68:387–99 [Google Scholar]
  10. Bates PD, Durrett TP, Ohlrogge JB, Pollard M. 10.  2009. Analysis of acyl fluxes through multiple pathways of triacylglycerol synthesis in developing soybean embryos. Plant Physiol. 150:55–72 [Google Scholar]
  11. Bates PD, Fatihi A, Snapp AR, Carlsson AS, Browse J, Lu CF. 11.  2012. Acyl editing and headgroup exchange are the major mechanisms that direct polyunsaturated fatty acid flux into triacylglycerols. Plant Physiol. 160:1530–39 [Google Scholar]
  12. Bates PD, Ohlrogge JB, Pollard M. 12.  2007. Incorporation of newly synthesized fatty acids into cytosolic glycerolipids in pea leaves occurs via acyl editing. J. Biol. Chem. 282:31206–16Identified PC acyl editing as an important component of glycerolipid metabolism. [Google Scholar]
  13. Bates PD, Stymne S, Ohlrogge J. 13.  2013. Biochemical pathways in seed oil synthesis. Curr. Opin. Plant Biol. 16:358–64 [Google Scholar]
  14. Baud S, Bourrellier ABF, Azzopardi M, Berger A, Dechorgnat J. 14.  et al. 2010. PII is induced by WRINKLED1 and fine-tunes fatty acid composition in seeds of Arabidopsis thaliana. Plant J. 64:291–303 [Google Scholar]
  15. Baud S, Lepiniec L. 15.  2009. Regulation of de novo fatty acid synthesis in maturing oilseeds of Arabidopsis. Plant Physiol. Biochem. 47:448–55 [Google Scholar]
  16. Baud S, Lepiniec L. 16.  2010. Physiological and developmental regulation of seed oil production. Prog. Lipid Res. 49:235–49 [Google Scholar]
  17. Baud S, Mendoza MS, To A, Harscoet E, Lepiniec L, Dubreucq B. 17.  2007. WRINKLED1 specifies the regulatory action of LEAFY COTYLEDON2 towards fatty acid metabolism during seed maturation in Arabidopsis. Plant J. 50:825–38 [Google Scholar]
  18. Beller M, Sztalryd C, Southall N, Bell M, Jackle H. 18.  et al. 2008. COPI complex is a regulator of lipid homeostasis. PLOS Biol. 6e292
  19. Bessoule JJ, Testet E, Cassagne C. 19.  1995. Synthesis of phosphatidylcholine in the chloroplast envelope after import of lysophosphatidylcholine from endoplasmic-reticulum membranes. Eur. J. Biochem. 228:490–97 [Google Scholar]
  20. Block MA, Jouhet J. 20.  2015. Lipid trafficking at endoplasmic reticulum-chloroplast membrane contact sites. Curr. Opin. Cell Biol. 35:21–29 [Google Scholar]
  21. Bortesi L, Fischer R. 21.  2015. The CRISPR/Cas9 system for plant genome editing and beyond. Biotechnol. Adv. 33:41–52 [Google Scholar]
  22. Bourrellier ABF, Valot B, Guillot A, Ambard-Bretteville F, Vidal J, Hodges M. 22.  2010. Chloroplast acetyl-CoA carboxylase activity is 2-oxoglutarate-regulated by interaction of PII with the biotin carboxyl carrier subunit. PNAS 107:502–7 [Google Scholar]
  23. Bouvier-Nave P, Benveniste P, Noiriel A, Schaller H. 23.  2000. Expression in yeast of an acyl-CoA:diacylglycerol acyltransferase cDNA from Caenorhabditis elegans. Biochem. Soc. Trans. 28:692–95 [Google Scholar]
  24. Buchanan-Wollaston V, Page T, Harrison E, Breeze E, Lim PO. 24.  et al. 2005. Comparative transcriptome analysis reveals significant differences in gene expression and signalling pathways between developmental and dark/starvation-induced senescence in Arabidopsis. Plant J. 42:567–85 [Google Scholar]
  25. Cai Y, Goodman JM, Pyc M, Mullen PT, Dyer JM, Chapman KD. 25.  2015. Arabidopsis SEIPIN proteins modulate triacylglycerol accumulation and influence lipid droplet proliferation. Plant Cell 27:2616–36Demonstrates a role for SEIPIN in TAG accumulation in plants. [Google Scholar]
  26. Carlsson AS, Yilmaz JL, Green AG, Stymne S, Hofvander P. 26.  2011. Replacing fossil oil with fresh oil—with what and for what?. Eur. J. Lipid Sci. Technol. 113:812–31 [Google Scholar]
  27. Carrasco S, Merida I. 27.  2007. Diacylglycerol, when simplicity becomes complex. Trends Biochem. Sci. 32:27–36 [Google Scholar]
  28. Cernac A, Benning C. 28.  2004. WRINKLED1 encodes an AP2/EREB domain protein involved in the control of storage compound biosynthesis in Arabidopsis. Plant J. 40:575–85 [Google Scholar]
  29. Chapman KD, Dyer JM, Mullen RT. 29.  2012. Biogenesis and functions of lipid droplets in plants. J. Lipid Res. 53:215–26 [Google Scholar]
  30. Chapman KD, Dyer JM, Mullen RT. 30.  2013. Commentary: Why don't plant leaves get fat?. Plant Sci. 207:128–34 [Google Scholar]
  31. Chapman KD, Ohlrogge JB. 31.  2012. Compartmentation of triacylglycerol accumulation in plants. J. Biol. Chem. 287:2288–94 [Google Scholar]
  32. Chen M, Mooney BP, Hajduch M, Joshi T, Zhou M. 32.  et al. 2009. System analysis of an Arabidopsis mutant altered in de novo fatty acid synthesis reveals diverse changes in seed composition and metabolism. Plant Physiol. 150:27–41 [Google Scholar]
  33. Craddock CP, Adams N, Bryant FM, Kurup S, Eastmond PJ. 33.  2015. PHOSPHATIDIC ACID PHOSPHOHYDROLASE regulates phosphatidylcholine biosynthesis in Arabidopsis by phosphatidic acid-mediated activation of CTP:PHOSPHOCHOLINE CYTIDYLYLTRANSFERASE activity. Plant Cell 27:1251–64 [Google Scholar]
  34. Dahlqvist A, Stahl U, Lenman M, Banas A, Lee M. 34.  et al. 2000. Phospholipid:diacylglycerol acyltransferase: an enzyme that catalyzes the acyl-CoA-independent formation of triacylglycerol in yeast and plants. PNAS 97:6487–92 [Google Scholar]
  35. De Domenico S, Bonsegna S, Lenucci MS, Poltronieri P, Di Sansebastiano GP, Santino A. 35.  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. De Marcos Lousa C, van Roermund CW, Postis VL, Dietrich D, Kerr ID. 36.  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. Dehesh K, Tai H, Edwards P, Byrne J, Jaworski JG. 37.  2001. Overexpression of 3-ketoacyl-acyl-carrier protein synthase IIIs in plants reduces the rate of lipid synthesis. Plant Physiol. 125:1103–14 [Google Scholar]
  38. Durrett TP, Benning C, Ohlrogge J. 38.  2008. Plant triacylglycerols as feedstocks for the production of biofuels. Plant J. 54:593–607 [Google Scholar]
  39. Dwyer JR, Donkor J, Zhang P, Csaki LS, Vergnes L. 39.  et al. 2012. Mouse lipin-1 and lipin-2 cooperate to maintain glycerolipid homeostasis in liver and aging cerebellum. PNAS 109:E2486–95 [Google Scholar]
  40. Eastmond PJ. 40.  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]
  41. Eastmond PJ, Quettier AL, Kroon JTM, Craddock C, Adams N, Slabas AR. 41.  2010. PHOSPHATIDIC ACID PHOSPHOHYDROLASE1 and 2 regulate phospholipid synthesis at the endoplasmic reticulum in Arabidopsis. Plant Cell 22:2796–811 [Google Scholar]
  42. Eccleston VS, Ohlrogge JB. 42.  1998. Expression of lauroyl-acyl carrier protein thioesterase in Brassica napus seeds induces pathways for both fatty acid oxidation and biosynthesis and implies a set point for triacylglycerol accumulation. Plant Cell 10:613–21 [Google Scholar]
  43. El-Kouhen K, Blangy S, Ortiz E, Gardies AM, Ferté N, Arondel V. 43.  2005. Identification and characterization of a triacylglycerol lipase in Arabidopsis homologous to mammalian acid lipases. FEBS Lett. 579:6067–73 [Google Scholar]
  44. Fan J, Yan C, Roston R, Shanklin J, Xu C. 44.  2014. Arabidopsis lipins, PDAT1 acyltransferase, and SDP1 triacylglycerol lipase synergistically direct fatty acids toward β-oxidation, thereby maintaining membrane lipid homeostasis. Plant Cell 26:4119–34Identified TAG as an intermediate in FA breakdown in leaves. [Google Scholar]
  45. Fan J, Yan C, Xu C. 45.  2013. Phospholipid:diacylglycerol acyltransferase-mediated triacylglycerol biosynthesis is crucial for protection against fatty acid-induced cell death in growing tissues of Arabidopsis. Plant J. 76:930–42 [Google Scholar]
  46. Fan J, Yan C, Zhang X, Xu C. 46.  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. Flugge UI, Hausler RE, Ludewig F, Gierth M. 47.  2011. The role of transporters in supplying energy to plant plastids. J. Exp. Bot. 62:2381–92 [Google Scholar]
  48. Focks N, Benning C. 48.  1998. wrinkled1: a novel, low-seed-oil mutant of Arabidopsis with a deficiency in the seed-specific regulation of carbohydrate metabolism. Plant Physiol. 118:91–101 [Google Scholar]
  49. Footitt S, Slocombe SP, Larner V, Kurup S, Wu Y. 49.  et al. 2002. Control of germination and lipid mobilization by COMATOSE, the Arabidopsis homologue of human ALDP. EMBO J. 21:2912–22 [Google Scholar]
  50. Fukuda N, Ikawa Y, Aoyagi T, Kozaki A. 50.  2013. Expression of the genes coding for plastidic acetyl-CoA carboxylase subunits is regulated by a location-sensitive transcription factor binding site. Plant Mol. Biol. 82:473–83 [Google Scholar]
  51. Fulda M, Shockey J, Werber M, Wolter FP, Heinz E. 51.  2002. Two long-chain acyl-CoA synthetases from Arabidopsis thaliana involved in peroxisomal fatty acid β-oxidation. Plant J. 32:93–103 [Google Scholar]
  52. Furumoto T, Yamaguchi T, Ohshima-Ichie Y, Nakamura M, Tsuchida-Iwata Y. 52.  et al. 2011. A plastidial sodium-dependent pyruvate transporter. Nature 476:472–75 [Google Scholar]
  53. Geigenberger P. 53.  2011. Regulation of starch biosynthesis in response to a fluctuating environment. Plant Physiol. 155:1566–77 [Google Scholar]
  54. Geigenberger P, Kolbe A, Tiessen A. 54.  2005. Redox regulation of carbon storage and partitioning in response to light and sugars. J. Exp. Bot. 56:1469–79 [Google Scholar]
  55. Germain V, Rylott EL, Larson TR, Sherson SM, Bechtold N. 55.  et al. 2001. Requirement for 3-ketoacyl-CoA thiolase-2 in peroxisome development, fatty acid β-oxidation and breakdown of triacylglycerol in lipid bodies of Arabidopsis seedlings. Plant J. 28:1–12 [Google Scholar]
  56. Ghosh AK, Chauhan N, Rajakumari S, Daum G, Rajasekharan R. 56.  2009. At4g24160, a soluble acyl-coenzyme A-dependent lysophosphatidic acid acyltransferase. Plant Physiol. 151:869–81 [Google Scholar]
  57. Graham IA. 57.  2008. Seed storage oil mobilization. Annu. Rev. Plant Biol. 59:115–42 [Google Scholar]
  58. Graham IA, Leaver CJ, Smith SM. 58.  1992. Induction of malate synthase gene expression in senescent and detached organs of cucumber. Plant Cell 4:349–57 [Google Scholar]
  59. Greer MS, Truksa M, Deng W, Lung SC, Chen G, Weselake RJ. 59.  2015. Engineering increased triacylglycerol accumulation in Saccharomyces cerevisiae using a modified type 1 plant diacylglycerol acyltransferase. Appl. Microbiol. Biotechnol. 99:2243–53 [Google Scholar]
  60. Grennan AK. 60.  2006. Regulation of starch metabolism in Arabidopsis leaves. Plant Physiol. 142:1343–45 [Google Scholar]
  61. Grimberg A, Carlsson AS, Marttila S, Bhalerao R, Hofvander P. 61.  2015. Transcriptional transitions in Nicotiana benthamiana leaves upon induction of oil synthesis by WRINKLED1 homologs from diverse species and tissues. BMC Plant Biol. 15:192Identified WRI1-induced transcripts in leaf tissue. [Google Scholar]
  62. Gronke S, Mildner A, Fellert S, Tennagels N, Petry S. 62.  et al. 2005. Brummer lipase is an evolutionary conserved fat storage regulator in Drosophila. Cell Metab. 1:323–30 [Google Scholar]
  63. Guo Y, Walther TC, Rao M, Stuurman N, Goshima G. 63.  et al. 2008. Functional genomic screen reveals genes involved in lipid-droplet formation and utilization. Nature 453:657–61 [Google Scholar]
  64. Gut H, Matile P. 64.  1988. Apparent induction of key enzymes of the glyoxylic-acid cycle in senescent barley leaves. Planta 176:548–50 [Google Scholar]
  65. Han GS, Siniossoglou S, Carman GM. 65.  2007. The cellular functions of the yeast lipin homolog PAH1p are dependent on its phosphatidate phosphatase activity. J. Biol. Chem. 282:37026–35 [Google Scholar]
  66. Han GS, Wu WI, Carman GM. 66.  2006. The Saccharomyces cerevisiae lipin homolog is a Mg2+-dependent phosphatidate phosphatase enzyme. J. Biol. Chem. 281:9210–18 [Google Scholar]
  67. Harris TE, Finck BN. 67.  2011. Dual function lipin proteins and glycerolipid metabolism. Trends Endocrinol. Metab. 22:226–33 [Google Scholar]
  68. Hayashi M, Toriyama K, Kondo M, Nishimura M. 68.  1998. 2,4-Dichlorophenoxybutyric acid–resistant mutants of Arabidopsis have defects in glyoxysomal fatty acid β-oxidation. Plant Cell 10:183–95 [Google Scholar]
  69. He YH, Gan SS. 69.  2002. A gene encoding an acyl hydrolase is involved in leaf senescence in Arabidopsis. Plant Cell 14:805–15 [Google Scholar]
  70. Henry SA, Kohlwein SD, Carman GM. 70.  2012. Metabolism and regulation of glycerolipids in the yeast Saccharomyces cerevisiae. Genetics 190:317–49 [Google Scholar]
  71. Hernandez ML, Whitehead L, He ZS, Gazda V, Gilday A. 71.  et al. 2012. A cytosolic acyltransferase contributes to triacylglycerol synthesis in sucrose-rescued Arabidopsis seed oil catabolism mutants. Plant Physiol. 160:215–25 [Google Scholar]
  72. Horn PJ, James CN, Gidda SK, Kilaru A, Dyer JM. 72.  et al. 2013. Identification of a new class of lipid droplet-associated proteins in plants. Plant Physiol. 162:1926–36 [Google Scholar]
  73. Jako C, Kumar A, Wei Y, Zou J, Barton DL. 73.  et al. 2001. Seed-specific over-expression of an Arabidopsis cDNA encoding a diacylglycerol acyltransferase enhances seed oil content and seed weight. Plant Physiol. 126:861–74 [Google Scholar]
  74. James CN, Horn PJ, Case CR, Gidda SK, Zhang D. 74.  et al. 2010. Disruption of the Arabidopsis CGI-58 homologue produces Chanarin-Dorfman-like lipid droplet accumulation in plants. PNAS 107:17833–38 [Google Scholar]
  75. Jefferson T. 75.  2005 (1800). Summary of public service [after 2 September 1800]. The Papers of Thomas Jefferson 32 1 June 1800–16 February 1801 BB Oberg 122–25 Princeton, NJ: Princeton Univ. Press http://founders.archives.gov/documents/Jefferson/01-32-02-0080 [Google Scholar]
  76. Jessen D, Roth C, Wiermer M, Fulda M. 76.  2015. Two activities of long-chain acyl-coenzyme A synthetase are involved in lipid trafficking between the endoplasmic reticulum and the plastid in Arabidopsis. Plant Physiol. 167:351–U575 [Google Scholar]
  77. Kachroo A, Shanklin J, Whittle E, Lapchyk L, Hildebrand D, Kachroo P. 77.  2007. The Arabidopsis stearoyl-acyl carrier protein-desaturase family and the contribution of leaf isoforms to oleic acid synthesis. Plant Mol. Biol. 63:257–71 [Google Scholar]
  78. Kalt-Torres W, Kerr PS, Usuda H, Huber SC. 78.  1987. Diurnal changes in maize leaf photosynthesis: I. Carbon exchange rate, assimilate export rate, and enzyme activities. Plant Physiol. 83:283–88 [Google Scholar]
  79. Kammerer B, Fischer K, Hilpert B, Schubert S, Gutensohn M. 79.  et al. 1998. Molecular characterization of a carbon transporter in plastids from heterotrophic tissues: the glucose 6-phosphate/phosphate antiporter. Plant Cell 10:105–17 [Google Scholar]
  80. Katavic V, Reed DW, Taylor DC, Giblin EM, Barton DL. 80.  et al. 1995. Alteration of seed fatty acid composition by an ethyl methanesulfonate-induced mutation in Arabidopsis thaliana affecting diacylglycerol acyltransferase activity. Plant Physiol. 108:399–409 [Google Scholar]
  81. Ke J, Wen TN, Nikolau BJ, Wurtele ES. 81.  2000. Coordinate regulation of the nuclear and plastidic genes coding for the subunits of the heteromeric acetyl-coenzyme A carboxylase. Plant Physiol. 122:1057–71 [Google Scholar]
  82. Kelly AA, Quettier AL, Shaw E, Eastmond PJ. 82.  2011. Seed storage oil mobilization is important but not essential for germination or seedling establishment in Arabidopsis. Plant Physiol. 157:866–75 [Google Scholar]
  83. Kelly AA, Shaw E, Powers SJ, Kurup S, Eastmond PJ. 83.  2012. Suppression of the SUGAR-DEPENDENT1 triacylglycerol lipase family during seed development enhances oil yield in oilseed rape (Brassica napus L.). Plant Biotechnol. J. 11:355–61 [Google Scholar]
  84. Kelly AA, van Erp H, Quettier AL, Shaw E, Menard G. 84.  et al. 2013. The SUGAR-DEPENDENT1 lipase limits triacylglycerol accumulation in vegetative tissues of Arabidopsis. Plant Physiol. 162:1282–89Identified SDP1 as the predominant TAG lipase in vegetative tissues. [Google Scholar]
  85. Kim EY, Seo YS, Kim WT. 85.  2011. AtDSEL, an Arabidopsis cytosolic DAD1-like acylhydrolase, is involved in negative regulation of storage oil mobilization during seedling establishment. J. Plant Physiol. 168:1705–9 [Google Scholar]
  86. Kim HU, Jung SJ, Lee KR, Kim EH, Lee SM. 86.  et al. 2013. Ectopic overexpression of castor bean LEAFY COTYLEDON2 (LEC2) in Arabidopsis triggers the expression of genes that encode regulators of seed maturation and oil body proteins in vegetative tissues. FEBS Open Biol. 4:25–32 [Google Scholar]
  87. Kim HU, Lee KR, Jung SJ, Shin HA, Go YS. 87.  et al. 2015. Senescence-inducible LEC2 enhances triacylglycerol accumulation in leaves without negatively affecting plant growth. Plant Biotechnol. J. 13:1346–59 [Google Scholar]
  88. Kim YY, Jung KW, Yoo KS, Jeung JU, Shin JS. 88.  2011. A stress-responsive caleosin-like protein, AtCLO4, acts as a negative regulator of ABA responses in Arabidopsis. Plant Cell Physiol. 52:874–84 [Google Scholar]
  89. Kjellberg JM, Trimborn M, Andersson M, Sandelius AS. 89.  2000. Acyl-CoA dependent acylation of phospholipids in the chloroplast envelope. Biochim. Biophys. Acta 1485:100–10 [Google Scholar]
  90. Kohlwein SD. 90.  2010. Triacylglycerol homeostasis: insights from yeast. J. Biol. Chem. 285:15663–67 [Google Scholar]
  91. Kohlwein SD, Veenhuis M, van der Klei IJ. 91.  2013. Lipid droplets and peroxisomes: key players in cellular lipid homeostasis or a matter of fat—store 'em up or burn 'em down. Genetics 193:1–50 [Google Scholar]
  92. Kolling K, Thalmann M, Muller A, Jenny C, Zeeman SC. 92.  2015. Carbon partitioning in Arabidopsis thaliana is a dynamic process controlled by the plants metabolic status and its circadian clock. Plant Cell Environ. 38:1965–79 [Google Scholar]
  93. Koo AJ, Fulda M, Browse J, Ohlrogge JB. 93.  2005. Identification of a plastid acyl-acyl carrier protein synthetase in Arabidopsis and its role in the activation and elongation of exogenous fatty acids. Plant J. 44:620–32 [Google Scholar]
  94. Kunz HH, Scharnewski M, Feussner K, Feussner I, Flugge UI. 94.  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]
  95. Lager I, Yilmaz JL, Zhou XR, Jasieniecka K, Kazachkov M. 95.  et al. 2013. Plant acyl-CoA:lysophosphatidylcholine acyltransferases (LPCATs) have different specificities in their forward and reverse reactions. J. Biol. Chem. 288:36902–14 [Google Scholar]
  96. Landolt R, Matile P. 96.  1990. Glyoxisome-like microbodies in senescent spinach leaves. Plant Sci. 72:159–63 [Google Scholar]
  97. Lass A, Zimmermann R, Haemmerle G, Riederer M, Schoiswohl G. 97.  et al. 2006. Adipose triglyceridelipase-mediated lipolysis of cellular fat stores is activated by CGI-58 and defective in Chanarin-Dorfman syndrome. Cell Metab. 3:309–19 [Google Scholar]
  98. Lersten NR, Czlapinski AR, Curtis JD, Freckmann R, Horner HT. 98.  2006. Oil bodies in leaf mesophyll cells of angiosperms: overview and a selected survey. Am. J. Bot. 93:1731–39 [Google Scholar]
  99. Li FL, Asami T, Wu XZ, Tsang EWT, Cutler AJ. 99.  2007. A putative hydroxysteroid dehydrogenase involved in regulating plant growth and development. Plant Physiol. 145:87–97 [Google Scholar]
  100. Li H, Culligan K, Dixon RA, Chory J. 100.  1995. CUE1: a mesophyll cell-specific positive regulator of light-controlled gene expression in Arabidopsis. Plant Cell 7:1599–610 [Google Scholar]
  101. Li N, Gugel IL, Giavalisco P, Zeisler V, Schreiber L. 101.  et al. 2015. FAX1, a novel membrane protein mediating plastid fatty acid export. PLOS Biol. 13e1002053
  102. Li R, Yu K, Hildebrand DF. 102.  2010. DGAT1, DGAT2 and PDAT expression in seeds and other tissues of epoxy and hydroxy fatty acid accumulating plants. Lipids 45:145–57 [Google Scholar]
  103. Li Y, Beisson F, Pollard M, Ohlrogge J. 103.  2006. Oil content of Arabidopsis seeds: the influence of seed anatomy, light and plant-to-plant variation. Phytochemistry 67:904–15 [Google Scholar]
  104. Linka N, Theodoulou FL, Haslam RP, Linka M, Napier JA. 104.  et al. 2008. Peroxisomal ATP import is essential for seedling development in Arabidopsis thaliana. Plant Cell 20:3241–57 [Google Scholar]
  105. Lippold F, vom Dorp K, Abraham M, Holzl G, Wewer V. 105.  et al. 2012. Fatty acid phytyl ester synthesis in chloroplasts of Arabidopsis. Plant Cell 24:2001–14 [Google Scholar]
  106. Liu Q, Siloto RM, Lehner R, Stone SJ, Weselake RJ. 106.  2012. Acyl-CoA:diacylglycerol acyltransferase: molecular biology, biochemistry and biotechnology. Prog. Lipid Res. 51:350–77 [Google Scholar]
  107. Liu X, Sheng J, Curtiss R III. 107.  2011. Fatty acid production in genetically modified cyanobacteria. PNAS 108:6899–904 [Google Scholar]
  108. Lock YY, Snyder CL, Zhu W, Siloto RM, Weselake RJ, Shah S. 108.  2009. Antisense suppression of type 1 diacylglycerol acyltransferase adversely affects plant development in Brassica napus. Physiol. Plant. 137:61–71 [Google Scholar]
  109. Loewen CJ. 109.  2012. Lipids as conductors in the orchestra of life. F1000 Biol. Rep. 4:4 [Google Scholar]
  110. Lonien J, Schwender J. 110.  2009. Analysis of metabolic flux phenotypes for two Arabidopsis mutants with severe impairment in seed storage lipid synthesis. Plant Physiol. 151:1617–34 [Google Scholar]
  111. Lu CFL, de Noyer SB, Hobbs DH, Kang JL, Wen YC. 111.  et al. 2003. Expression pattern of diacylglycerol acyltransferase-1, an enzyme involved in triacylglycerol biosynthesis, in Arabidopsis thaliana. Plant Mol. Biol. 52:31–41 [Google Scholar]
  112. Lu SY, Song T, Kosma DK, Parsons EP, Rowland O, Jenks MA. 112.  2009. Arabidopsis CER8 encodes LONG-CHAIN ACYL-COA SYNTHETASE 1 (LACS1) that has overlapping functions with LACS2 in plant wax and cutin synthesis. Plant J. 59:553–64 [Google Scholar]
  113. Ma W, Kong Q, Grix M, Mantyla JJ, Yang Y. 113.  et al. 2015. Deletion of a C-terminal intrinsically disordered region of WRINKLED1 affects its stability and enhances oil accumulation in Arabidopsis. Plant J. 83:864–74 [Google Scholar]
  114. Maeo K, Tokuda T, Ayame A, Mitsui N, Kawai T. 114.  et al. 2009. An AP2-type transcription factor, WRINKLED1, of Arabidopsis thaliana binds to the AW-box sequence conserved among proximal upstream regions of genes involved in fatty acid synthesis. Plant J. 60:476–87 [Google Scholar]
  115. Martin S, Parton RG. 115.  2006. Lipid droplets: a unified view of a dynamic organelle. Nat. Rev. Mol. Cell Biol. 7:373–78 [Google Scholar]
  116. Masaki T, Mitsui N, Tsukagoshi H, Nishii T, Morikami A, Nakamura K. 116.  2005. ACTIVATOR of Spomin::LUC1/WRINKLED1 of Arabidopsis thaliana transactivates sugar-inducible promoters. Plant Cell Physiol. 46:547–56 [Google Scholar]
  117. Mitra MS, Chen Z, Ren H, Harris TE, Chambers KT. 117.  et al. 2013. Mice with an adipocyte-specific lipin 1 separation-of-function allele reveal unexpected roles for phosphatidic acid in metabolic regulation. PNAS 110:642–47 [Google Scholar]
  118. Morandini P. 118.  2013. Control limits for accumulation of plant metabolites: Brute force is no substitute for understanding. Plant Biotechnol. J. 11:253–67 [Google Scholar]
  119. Murphy DJ. 119.  2001. The biogenesis and functions of lipid bodies in animals, plants and microorganisms. Prog. Lipid Res. 40:325–438 [Google Scholar]
  120. Nakamura Y, Koizumi R, Shui G, Shimojima M, Wenk MR. 120.  et al. 2009. Arabidopsis lipins mediate eukaryotic pathway of lipid metabolism and cope critically with phosphate starvation. PNAS 106:20978–83 [Google Scholar]
  121. Napier JA. 121.  2007. The production of unusual fatty acids in transgenic plants. Annu. Rev. Plant Biol. 58:295–319 [Google Scholar]
  122. Napier JA, Haslam RP, Beaudoin F, Cahoon EB. 122.  2014. Understanding and manipulating plant lipid composition: Metabolic engineering leads the way. Curr. Opin. Plant Biol. 19:68–75 [Google Scholar]
  123. Niewiadomski P, Knappe S, Geimer S, Fischer K, Schulz B. 123.  et al. 2005. The Arabidopsis plastidic glucose 6-phosphate/phosphate translocator GPT1 is essential for pollen maturation and embryo sac development. Plant Cell 17:760–75 [Google Scholar]
  124. O'Grady J, Schwender J, Shachar-Hill Y, Morgan JA. 124.  2012. Metabolic cartography: experimental quantification of metabolic fluxes from isotopic labelling studies. J. Exp. Bot. 63:2293–308 [Google Scholar]
  125. Ogas J, Kaufmann S, Henderson J, Somerville C. 125.  1999. PICKLE is a CHD3 chromatin-remodeling factor that regulates the transition from embryonic to vegetative development in Arabidopsis. PNAS 96:13839–44 [Google Scholar]
  126. Ohlrogge J, Allen D, Berguson B, DellaPenna D, Shachar-Hill Y, Stymne S. 126.  2009. Driving on biomass. Science 324:1019–20 [Google Scholar]
  127. Ohlrogge J, Browse J. 127.  1995. Lipid biosynthesis. Plant Cell 7:957–70 [Google Scholar]
  128. Ohlrogge J, Chapman KD. 128.  2011. The seeds of green energy—expanding the contribution of plant oils as biofuels. Biochemist 33:34–38 [Google Scholar]
  129. Ohlrogge JB, Jaworski JG. 129.  1997. Regulation of fatty acid synthesis. Annu. Rev. Plant Physiol. Plant Mol. Biol. 48:109–36 [Google Scholar]
  130. Olofsson SO, Bostrom P, Andersson L, Rutberg M, Perman J, Boren J. 130.  2009. Lipid droplets as dynamic organelles connecting storage and efflux of lipids. Biochim. Biophys. Acta 1791:448–58 [Google Scholar]
  131. Padham AK, Hopkins MT, Wang TW, McNamara LM, Lo M. 131.  et al. 2007. Characterization of a plastid triacylglycerol lipase from Arabidopsis. Plant Physiol. 143:1372–84 [Google Scholar]
  132. Park S, Gidda SK, James CN, Horn PJ, Khuu N. 132.  et al. 2013. The α/β hydrolase CGI-58 and peroxisomal transport protein PXA1 coregulate lipid homeostasis and signaling in Arabidopsis. Plant Cell 25:1726–39 [Google Scholar]
  133. Pascual F, Carman GM. 133.  2013. Phosphatidate phosphatase, a key regulator of lipid homeostasis. Biochim. Biophys. Acta 1831:514–22 [Google Scholar]
  134. Petrie JR, Vanhercke T, Shrestha P, El Tahchy A, White A. 134.  et al. 2012. Recruiting a new substrate for triacylglycerol synthesis in plants: the monoacylglycerol acyltransferase pathway. PLOS ONE 7:e35214 [Google Scholar]
  135. Prabhakar V, Lottgert T, Geimer S, Dormann P, Kruger S. 135.  et al. 2010. Phosphoenolpyruvate provision to plastids is essential for gametophyte and sporophyte development in Arabidopsis thaliana. Plant Cell 22:2594–617 [Google Scholar]
  136. Prabhakar V, Lottgert T, Gigolashvili T, Bell K, Flugge UI, Hausler RE. 136.  2009. Molecular and functional characterization of the plastid-localized phosphoenolpyruvate enolase (ENO1) from Arabidopsis thaliana. FEBS Lett. 583:983–91 [Google Scholar]
  137. Rambold AS, Cohen S, Lippincott-Schwartz J. 137.  2015. Fatty acid trafficking in starved cells: regulation by lipid droplet lipolysis, autophagy, and mitochondrial fusion dynamics. Dev. Cell 32:678–92 [Google Scholar]
  138. Rani SH, Krishna TH, Saha S, Negi AS, Rajasekharan R. 138.  2010. Defective in cuticular ridges (DCR) of Arabidopsis thaliana, a gene associated with surface cutin formation, encodes a soluble diacylglycerol acyltransferase. J. Biol. Chem. 285:38337–47 [Google Scholar]
  139. Reynolds KB, Taylor MC, Zhou XR, Vanhercke T, Wood CC. 139.  et al. 2015. Metabolic engineering of medium-chain fatty acid biosynthesis in Nicotiana benthamiana plant leaf lipids. Front. Plant Sci. 6:164 [Google Scholar]
  140. Roesler K, Shintani D, Savage L, Boddupalli S, Ohlrogge J. 140.  1997. Targeting of the Arabidopsis homomeric acetyl-coenzyme A carboxylase to plastids of rapeseeds. Plant Physiol. 113:75–81 [Google Scholar]
  141. Rolland F, Sheen J. 141.  2005. Sugar sensing and signalling networks in plants. Biochem. Soc. Trans. 33:269–71 [Google Scholar]
  142. Routaboul JM, Benning C, Bechtold N, Caboche M, Lepiniec L. 142.  1999. The TAG1 locus of Arabidopsis encodes for a diacylglycerol acyltransferase. Plant Physiol. Biochem. 37:831–40 [Google Scholar]
  143. Ruiz-Lopez N, Haslam RP, Usher SL, Napier JA, Sayanova O. 143.  2013. Reconstitution of EPA and DHA biosynthesis in Arabidopsis: iterative metabolic engineering for the synthesis of n-3 LC-PUFAs in transgenic plants. Metab. Eng. 17:30–41 [Google Scholar]
  144. Ruuska SA, Girke T, Benning C, Ohlrogge JB. 144.  2002. Contrapuntal networks of gene expression during Arabidopsis seed filling. Plant Cell 14:1191–206 [Google Scholar]
  145. Sanjaya, Durrett TP, Weise SE, Benning C. 145.  2011. Increasing the energy density of vegetative tissues by diverting carbon from starch to oil biosynthesis in transgenic Arabidopsis. Plant Biotechnol. J. 9:874–83Demonstrated increased TAG accumulation by inhibition of starch synthesis. [Google Scholar]
  146. Sanjaya, Miller R, Durrett TP, Kosma DK, Lydic TA. 146.  et al. 2013. Altered lipid composition and enhanced nutritional value of Arabidopsis leaves following introduction of an algal diacylglycerol acyltransferase 2. Plant Cell 25:677–93 [Google Scholar]
  147. Santos-Mendoza M, Dubreucq B, Baud S, Parcy F, Caboche M, Lepiniec L. 147.  2008. Deciphering gene regulatory networks that control seed development and maturation in Arabidopsis. Plant J. 54:608–20 [Google Scholar]
  148. Santos-Mendoza M, Dubreucq B, Miquel M, Caboche M, Lepiniec L. 148.  2005. LEAFY COTYLEDON 2 activation is sufficient to trigger the accumulation of oil and seed specific mRNAs in Arabidopsis leaves. FEBS Lett. 579:4666–70 [Google Scholar]
  149. Santos-Rosa H, Leung J, Grimsey N, Peak-Chew S, Siniossoglou S. 149.  2005. The yeast lipin Smp2 couples phospholipid biosynthesis to nuclear membrane growth. EMBO J. 24:1931–41 [Google Scholar]
  150. Sasaki Y, Kozaki A, Hatano M. 150.  1997. Link between light and fatty acid synthesis: thioredoxin-linked reductive activation of plastidic acetyl-CoA carboxylase. PNAS 94:11096–101 [Google Scholar]
  151. Schnurr J, Shockey J, Browse J. 151.  2004. The acyl-CoA synthetase encoded by LACS2 is essential for normal cuticle development in Arabidopsis. Plant Cell 16:629–42 [Google Scholar]
  152. Schwender J, Hay JO. 152.  2012. Predictive modeling of biomass component tradeoffs in Brassica napus developing oilseeds based on in silico manipulation of storage metabolism. Plant Physiol. 160:1218–36 [Google Scholar]
  153. Schwender J, Hebbelmann I, Heinzel N, Hildebrandt TM, Rogers A. 153.  et al. 2015. Quantitative multilevel analysis of central metabolism in developing oilseeds of Brassica napus during in vitro culture. Plant Physiol 168:828–48Described a mathematical approach to understanding the basis of oil accumulation in seeds. [Google Scholar]
  154. Scott RW, Winichayakul S, Roldan M, Cookson R, Willingham M. 154.  et al. 2010. Elevation of oil body integrity and emulsion stability by polyoleosins, multiple oleosin units joined in tandem head-to-tail fusions. Plant Biotechnol. J. 8:912–27 [Google Scholar]
  155. Shi SB, Chen Y, Siewers V, Nielsen J. 155.  2014. Improving production of malonyl coenzyme A-derived metabolites by abolishing Snf1-dependent regulation of Acc1. mBio 5e01130–14
  156. Shimada TL, Hara-Nishimura I. 156.  2010. Oil-body-membrane proteins and their physiological functions in plants. Biol. Pharm. Bull. 33:360–63 [Google Scholar]
  157. Shimada TL, Shimada T, Takahashi H, Fukao Y, Hara-Nishimura I. 157.  2008. A novel role for oleosins in freezing tolerance of oilseeds in Arabidopsis thaliana. Plant J. 55:798–809 [Google Scholar]
  158. Shimada TL, Takano Y, Shimada T, Fujiwara M, Fukao Y. 158.  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]
  159. Shintani DK, Ohlrogge JB. 159.  1995. Feedback inhibition of fatty-acid synthesis in tobacco suspension cells. Plant J. 7:577–87 [Google Scholar]
  160. Shockey JM, Fulda MS, Browse JA. 160.  2002. Arabidopsis contains nine long-chain acyl-coenzyme A synthetase genes that participate in fatty acid and glycerolipid metabolism. Plant Physiol. 129:1710–22 [Google Scholar]
  161. Siloto RMP, Findlay K, Lopez-Villalobos A, Yeung EC, Nykiforuk CL, Moloney MM. 161.  2006. The accumulation of oleosins determines the size of seed oilbodies in Arabidopsis. Plant Cell 18:1961–74 [Google Scholar]
  162. Singh R, Kaushik S, Wang Y, Xiang Y, Novak I. 162.  et al. 2009. Autophagy regulates lipid metabolism. Nature 458:1131–35 [Google Scholar]
  163. Slocombe SP, Cornah J, Pinfield-Wells H, Soady K, Zhang Q. 163.  et al. 2009. Oil accumulation in leaves directed by modification of fatty acid breakdown and lipid synthesis pathways. Plant Biotechnol. J. 7:694–703 [Google Scholar]
  164. Sorokin HP. 164.  1955. Mitochondria and spherosomes in the living epidermal cell. Am. J. Bot. 42:225–31 [Google Scholar]
  165. Stahl U, Carlsson AS, Lenman M, Dahlqvist A, Huang B. 165.  et al. 2004. Cloning and functional characterization of a phospholipid:diacylglycerol acyltransferase from Arabidopsis. Plant Physiol. 135:1324–35 [Google Scholar]
  166. Stitt M, Zeeman SC. 166.  2012. Starch turnover: pathways, regulation and role in growth. Curr. Opin. Plant Biol. 15:282–92 [Google Scholar]
  167. Stone SL, Kwong LW, Yee KM, Pelletier J, Lepiniec L. 167.  et al. 2001. LEAFY COTYLEDON2 encodes a B3 domain transcription factor that induces embryo development. PNAS 98:11806–11 [Google Scholar]
  168. Thazar-Poulot N, Miquel M, Fobis-Loisy I, Gaude T. 168.  2015. Peroxisome extensions deliver the Arabidopsis SDP1 lipase to oil bodies. PNAS 112:4158–63 [Google Scholar]
  169. Thelen JJ, Ohlrogge JB. 169.  2002. Both antisense and sense expression of biotin carboxyl carrier protein isoform 2 inactivates the plastid acetyl-coenzyme A carboxylase in Arabidopsis thaliana. Plant J. 32:419–31 [Google Scholar]
  170. Theodoulou FL, Eastmond PJ. 170.  2012. Seed storage oil catabolism: a story of give and take. Curr. Opin. Plant Biol. 15:322–28 [Google Scholar]
  171. Theodoulou FL, Holdsworth M, Baker A. 171.  2006. Peroxisomal ABC transporters. FEBS Lett. 580:1139–55 [Google Scholar]
  172. Tjellstrom H, Strawsine M, Ohlrogge JB. 172.  2015. Tracking synthesis and turnover of triacylglycerol in leaves. J. Exp. Bot. 66:1453–61 [Google Scholar]
  173. Tjellstrom H, Strawsine M, Silva J, Cahoon EB, Ohlrogge JB. 173.  2013. Disruption of plastid acyl:acyl carrier protein synthetases increases medium chain fatty acid accumulation in seeds of transgenic Arabidopsis. FEBS Lett. 587:936–42 [Google Scholar]
  174. Tjellstrom H, Yang Z, Allen DK, Ohlrogge JB. 174.  2012. Rapid kinetic labeling of Arabidopsis cell suspension cultures: implications for models of lipid export from plastids. Plant Physiol. 158:601–11 [Google Scholar]
  175. To A, Joubes J, Barthole G, Lecureuil A, Scagnelli A. 175.  et al. 2012. WRINKLED transcription factors orchestrate tissue-specific regulation of fatty acid biosynthesis in Arabidopsis. Plant Cell 24:5007–23 [Google Scholar]
  176. Troncoso-Ponce MA, Cao X, Yang ZL, Ohlrogge JB. 176.  2013. Lipid turnover during senescence. Plant Sci. 205:13–19 [Google Scholar]
  177. Troncoso-Ponce MA, Nikovics K, Marchive C, Lepiniec L, Baud S. 177.  2016. New insights on the organization and regulation of the fatty acid biosynthetic network in the model higher plant Arabidopsis thaliana. Biochimie 120:3–8 [Google Scholar]
  178. Ugrankar R, Liu YL, Provaznik J, Schmitt S, Lehmann M. 178.  2011. Lipin is a central regulator of adipose tissue development and function in Drosophila melanogaster. Mol. Cell. Biol. 31:1646–56 [Google Scholar]
  179. Valdearcos M, Esquinas E, Meana C, Pena L, Gil-de-Gomez L. 179.  et al. 2012. Lipin-2 reduces proinflammatory signaling induced by saturated fatty acids in macrophages. J. Biol. Chem. 287:10894–904 [Google Scholar]
  180. Van der Graaff E, Schwacke R, Schneider A, Desimone M, Flugge UI, Kunze R. 180.  2006. Transcription analysis of Arabidopsis membrane transporters and hormone pathways during developmental and induced leaf senescence. Plant Physiol. 141:776–92 [Google Scholar]
  181. Vanhercke T, El Tahchy A, Liu Q, Zhou XR, Shrestha P. 181.  et al. 2014. Metabolic engineering of biomass for high energy density: oilseed-like triacylglycerol yields from plant leaves. Plant Biotechnol. J. 12:231–39Described a three-gene approach for optimizing oil accumulation in tobacco leaves. [Google Scholar]
  182. Vanhercke T, El Tahchy A, Shrestha P, Zhou XR, Singh SP, Petrie JR. 182.  2013. Synergistic effect of WRI1 and DGAT1 coexpression on triacylglycerol biosynthesis in plants. FEBS Lett. 587:364–69 [Google Scholar]
  183. Vanhercke T, Petrie JR, Singh S. 183.  2014. Energy densification in vegetative biomass through metabolic engineering. Biocatal. Agric. Biotechnol. 3:75–80 [Google Scholar]
  184. Voelker T. 184.  1996. Plant acyl-ACP thioesterases: chain-length determining enzymes in plant fatty acid biosynthesis. Genet. Eng. 18:111–33 [Google Scholar]
  185. Wahlroos T, Soukka J, Denesyuk A, Wahlroos R, Korpela T, Kilby NJ. 185.  2003. Oleosin expression and trafficking during oil body biogenesis in tobacco leaf cells. Genesis 35:125–32 [Google Scholar]
  186. Wang LP, Shen WY, Kazachkov M, Chen GQ, Chen QL. 186.  et al. 2012. Metabolic interactions between the Lands cycle and the Kennedy pathway of glycerolipid synthesis in Arabidopsis developing seeds. Plant Cell 24:4652–69 [Google Scholar]
  187. Wang Z, Benning C. 187.  2012. Chloroplast lipid synthesis and lipid trafficking through ER-plastid membrane contact sites. Biochem. Soc. Trans. 40:457–63 [Google Scholar]
  188. Weber APM, Linka N. 188.  2011. Connecting the plastid: transporters of the plastid envelope and their role in linking plastidial with cytosolic metabolism. Annu. Rev. Plant Biol. 62:53–77 [Google Scholar]
  189. Wilfling F, Haas JT, Walther TC, Farese RV. 189.  2014. Lipid droplet biogenesis. Curr. Opin. Cell Biol. 29:39–45 [Google Scholar]
  190. Winichayakul S, Scott RW, Roldan M, Hatier JH, Livingston S. 190.  et al. 2013. In vivo packaging of triacylglycerols enhances Arabidopsis leaf biomass and energy density. Plant Physiol. 162:626–39 [Google Scholar]
  191. Wu HY, Liu C, Li MC, Zhao MM, Gu D, Xu YN. 191.  2013. Effects of monogalactoglycerolipid deficiency and diacylglycerol acyltransferase overexpression on oil accumulation in transgenic tobacco. Plant Mol. Biol. Rep. 31:1077–88 [Google Scholar]
  192. Xu C, Fan J, Froehlich JE, Awai K, Benning C. 192.  2005. Mutation of the TGD1 chloroplast envelope protein affects phosphatidate metabolism in Arabidopsis. Plant Cell 17:3094–110 [Google Scholar]
  193. Yamaguchi T, Osumi T. 193.  2009. Chanarin-Dorfman syndrome: deficiency in CGI-58, a lipid droplet-bound coactivator of lipase. Biochim. Biophys. Acta 1791:519–23 [Google Scholar]
  194. Yang Y, Munz J, Cass C, Zienkiewicz A, Kong Q. 194.  et al. 2015. Ectopic expression of WRINKLED1 affects fatty acid homeostasis in Brachypodium distachyon vegetative tissues. Plant Physiol. 169:1836–47 [Google Scholar]
  195. Yang ZL, Ohlrogge JB. 195.  2009. Turnover of fatty acids during natural senescence of Arabidopsis, Brachypodium, and switchgrass and in Arabidopsis β-oxidation mutants. Plant Physiol. 150:1981–89 [Google Scholar]
  196. Yazdanbakhsh N, Sulpice R, Graf A, Stitt M, Fisahn J. 196.  2011. Circadian control of root elongation and C partitioning in Arabidopsis thaliana. Plant Cell Environ. 34:877–94 [Google Scholar]
  197. Zale J, Jung JH, Kim JY, Pathak B, Karan R. 197.  et al. 2016. Metabolic engineering of sugarcane to accumulate energy-dense triacylglycerols in vegetative biomass. Plant Biotechnol. J. 14661–69Demonstrated the feasibility of a strategy based on increasing vegetative TAG accumulation in monocot crop plants.
  198. Zhang M, Fan J, Taylor DC, Ohlrogge JB. 198.  2009. DGAT1 and PDAT1 acyltransferases have overlapping functions in Arabidopsis triacylglycerol biosynthesis and are essential for normal pollen and seed development. Plant Cell 21:3885–901 [Google Scholar]
  199. Zhang P, Takeuchi K, Csaki LS, Reue K. 199.  2012. Lipin-1 phosphatidic phosphatase activity modulates phosphatidate levels to promote peroxisome proliferator-activated receptor gamma (PPARγ) gene expression during adipogenesis. J. Biol. Chem. 287:3485–94 [Google Scholar]
  200. Zhao LF, Katavic V, Li FL, Haughn GW, Kunst L. 200.  2010. Insertional mutant analysis reveals that long-chain acyl-CoA synthetase 1 (LACS1), but not LACS8, functionally overlaps with LACS9 in Arabidopsis seed oil biosynthesis. Plant J. 64:1048–58 [Google Scholar]
  201. Zimmermann R, Strauss JG, Haemmerle G, Schoiswohl G, Birner-Gruenberger R. 201.  et al. 2004. Fat mobilization in adipose tissue is promoted by adipose triglyceride lipase. Science 306:1383–86 [Google Scholar]
  202. Zolman BK, Silva ID, Bartel B. 202.  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]

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