Lipids

, Volume 50, Issue 11, pp 1057–1068 | Cite as

Acyl-Trafficking During Plant Oil Accumulation

Review

Abstract

Vegetable oils are an extremely important agricultural commodity. Their use has risen inexorably for the last 50 years and will undoubtedly be even more prevalent in the future. They have a role not only in foodstuffs but also as renewable chemicals. However, our understanding of their metabolism, and particularly its control, is incomplete. In this article we highlight current knowledge and its deficiencies. In particular, we focus on the important role that phosphatidylcholine plays in lipid accumulation and in influencing the quality of the vegetable oils produced.

Keywords

Oil crops Triacylglycerol biosynthesis Kennedy pathway Phosphatidylcholine Metabolic control 

Abbreviations

ACCase

Acetyl-CoA carboxylase

ACP

Acyl carrier protein

CDP-choline

Cytidine diphosphate-choline

CPT

Cholinephosphotransferase

DAG

Diacylglycerol

DGAT

Diacylglycerol acyltransferase

ER

Endoplasmic reticulum

FAD

Fatty acid desaturase

FAS

Fatty acid synthase

FAT

Fatty acyl-ACP thioesterase

FAX1

Fatty acid export 1

G3P

sn-Glycerol 3-phosphate

GPAT

Glycerol 3-phosphate acyltransferase

GPC

Glycerophosphocholine

GPCAT

Glycerophosphocholine acyltransferase

KAS

β-Ketoacyl-ACP synthase

LACS

Long-chain acyl-CoA synthetase

LPA

Lysophosphatidic acid

LPAAT

Lysophosphatidate acyltransferase

LPC

Lysophosphatidylcholine

LPCAT

Lysophosphatidylcholine acyltransferase

LPCT

Lysophosphatidylcholine transacylase

MGDG

Monogalactosyldiacylglycerol

NAD(P)H

Nicotinamide adenine dinucleotide (phosphate)

NEFA

Nonesterified fatty acid (free fatty acid)

PA

Phosphatidic acid (PtdOH)

PAPase

Phosphatidate phosphohydrolase

PDAT

Phospholipid:diacylglycerol acyltransferase

PDCT

Phosphatidylcholine:diacylglycerol cholinephosphotransferase

PLA

Phospholipase A

PLC

Phospholipase C

PLD

Phospholipase D

PtdCho

Phosphatidylcholine

PtdGro

Phosphatidylglycerol

PtdIns

Phosphatidylinositol

PUFA

Polyunsaturated fatty acid

ROD1

Reduced oleate desaturation 1

TAG

Triacylglycerol

References

  1. 1.
    Oils and fats in the market place—prices of commodity oils. http://lipidlibrary.aocs.org/market/prices.htm. Accessed Mar 2015
  2. 2.
    Gunstone FD, Harwood JL, Dijkstra AJ (eds) (2007) The lipid handbook, 3rd edn. CRC Press, Boca RatonGoogle Scholar
  3. 3.
    Carlsson AS, Yilmaz JL, Green AG, Stymne S, Hofvander P (2011) Replacing fossil oil with fresh oil—with what and for what? Eur J Lipid Sci Technol 113:812–831PubMedCentralPubMedCrossRefGoogle Scholar
  4. 4.
    Vanhercke T, El Tahchy A, Liu Q, Zhou X-R, Shrestha P, Divi UK, Ral J-P, Mansour MP, Nichols PD, James CN, Horn PJ, Chapman KD, Beaudoin F, Ruiz-López N, Larkin PJ, de Feyter RC, Singh SP, Petrie JR (2014) Metabolic engineering of biomass for high energy density: oilseed-like triacylglycerol yields from plant leaves. Plant Biotech J 12:231–239CrossRefGoogle Scholar
  5. 5.
    Murphy DJ (ed) (2005) Plant lipids: biology, utilisation and manipulation. CRC Press, Boca RatonGoogle Scholar
  6. 6.
    Harwood JL, Ramli US, Tang M, Quant PA, Weselake RJ, Fawcett T, Guschina IA (2013) Regulation and enhancement of lipid accumulation in oil crops: the use of metabolic control analysis for informed genetic manipulation. Eur J Lipid Sci Technol 115:1239–1246CrossRefGoogle Scholar
  7. 7.
    Murphy DJ (2014) The future of oil palm as a major global crop: opportunities and challenges. J Oil Palm Res 26:1–24Google Scholar
  8. 8.
    Salas JJ, Martinez-Force E, Harwood JL, Venegas-Caleron M, Aznar-Moreno JA, Moreno-Perez AJ, Ruiz-Lopez N, Serrano-Vega MJ, Graham IA, Mullen RT, Garces R (2014) Biochemistry of high stearic sunflower, a new source of saturated fats. Prog Lipid Res 55:30–42PubMedCrossRefGoogle Scholar
  9. 9.
    Murphy DJ (ed) (1994) Designer oil crops—breeding, processing and biotechnology. VCH Weinheim, New YorkGoogle Scholar
  10. 10.
    Voelker TA, Worrell AC, Anderson L, Bleibaum J, Fan C, Hawkins DJ, Radke SE, Davies HM (1992) Fatty acid biosynthesis redirected to medium chains in transgenic oilseed plants. Science 257:72–74PubMedCrossRefGoogle Scholar
  11. 11.
    Weselake RJ, Taylor DC, Rahman MH, Shah S, Laroche A, McVetty PBE, Harwood JL (2009) Increasing the flow of carbon into seed oil. Biotechnol Adv 27:866–878PubMedCrossRefGoogle Scholar
  12. 12.
    Rahman H, Harwood J, Weselake R (2013) Increasing seed oil content in Brassica species through breeding and biotechnology. Lipid Tech 25:182–185CrossRefGoogle Scholar
  13. 13.
    Napier JA (2007) The production of unusual fatty acids in transgenic plants. Annu Rev Plant Physiol 58:295–319Google Scholar
  14. 14.
    Haslam RP, Ruiz-Lopez N, Eastmond P, Moloney M, Sayanova O, Napier JA (2013) The modification of plant oil composition via metabolic engineering-better nutrition by design. Plant Biotech J 11:157–168CrossRefGoogle Scholar
  15. 15.
    Bates PD, Stymne S, Ohlrogge J (2013) Biochemical pathways in seed oil synthesis. Curr Opin Plant Biol 16:358–364PubMedCrossRefGoogle Scholar
  16. 16.
    Chapman KD, Ohlrogge JB (2012) Compartmentation of triacylglycerol accumulation in plants. J Biol Chem 287:2288–2294PubMedCentralPubMedCrossRefGoogle Scholar
  17. 17.
    Sanjaya Durrett TP, Weise SE, Benning C (2011) Increasing the energy density of vegetative tissues by diverting carbon from starch to oil biosynthesis in transgenic Arabidopsis. Plant Biotechnol J 9:874–883PubMedCrossRefGoogle Scholar
  18. 18.
    Liedvogel B (1987) Lipid precursors in plant cells: the problem of acetyl CoA generation for plastid fatty acid synthesis. In: Stumpf P, Mudd JB, Nes WD (eds) The metabolism, structure, and function of plant lipids. Springer, New YorkGoogle Scholar
  19. 19.
    Allen DK, Bates PD, Tjellström H (2015) Tracking the metabolic pulse of plant lipid production with isotopic labeling and flux analyses: past, present and future. Prog Lipid Res 58:97–120PubMedCrossRefGoogle Scholar
  20. 20.
    Harwood J (2005) Fatty acid biosynthesis. In: Murphy DJ (ed) Plant lipids: biology, utilisation and manipulation. CRC Press, Boca RatonGoogle Scholar
  21. 21.
    Walker A, Ridley S, Lewis T, Harwood J (1989) Action of aryloxy-phenoxy carboxylic acids on lipid metabolism. Rev Weed Sci 4:71–84Google Scholar
  22. 22.
    Harwood J (1991) Herbicides affecting chloroplast lipid synthesis. In: Baker N, Percival M (eds) Topics in photosynthesis, vol 10. Elsevier, AmsterdamGoogle Scholar
  23. 23.
    Page RA, Okada S, Harwood JL (1994) Acetyl-CoA carboxylase exerts strong flux control over lipid synthesis in plants. Biochim Biophys Acta 1210:369–372PubMedCrossRefGoogle Scholar
  24. 24.
    Turnham E, Northcote DH (1983) Changes in the activity of acetyl-CoA carboxylase during rape-seed formation. Biochem J 212:223–229PubMedCentralPubMedCrossRefGoogle Scholar
  25. 25.
    Kang F, Ridout C, Morgan C, Rawsthorne S (1994) The activity of acetyl-CoA carboxylase is not correlated with the rate of lipid synthesis during development of oilseed rape (Brassica napus L.) embryos. Planta 193:320–325CrossRefGoogle Scholar
  26. 26.
    Roesler K, Shintani D, Savage L, Boddupalli S, Ohlrogge J (1997) Targeting of the Arabidopsis homomeric acetyl-coenzyme A carboxylase to plastids of rapeseeds. Plant Physiol 113:75–81PubMedCentralPubMedCrossRefGoogle Scholar
  27. 27.
    Andre C, Haslam RP, Shanklin J (2012) Feedback regulation of plastidic acetyl-CoA carboxylase by 18:1-acyl carrier protein in Brassica napus. Proc Natl Acad Sci USA 109:10107–10112PubMedCentralPubMedCrossRefGoogle Scholar
  28. 28.
    Ramli US, Salas JJ, Quant PA, Harwood JL (2009) Use of metabolic control analysis to give quantitative information on control of lipid biosynthesis in the important oil crop, Elaeis guineensis (oilpalm). New Phytol 184:330–339PubMedCrossRefGoogle Scholar
  29. 29.
    Ramli US. Biochemical studies of lipid biosynthesis in oil palm and olive callus cultures, 1999, PhD thesis, Cardiff University, Wales, UKGoogle Scholar
  30. 30.
    Bates PD, Johnson SR, Cao X, Li J, Nam JW, Jaworski JG, Ohlrogge JB, Browse J (2014) Fatty acid synthesis is inhibited by inefficient utilization of unusual fatty acids for glycerolipid assembly. Proc Natl Acad Sci USA 111:1204–1209PubMedCentralPubMedCrossRefGoogle Scholar
  31. 31.
    Harwood JL (1988) Fatty acid metabolism. Annu Rev Plant Physiol Plant Mol Biol 39:101–138CrossRefGoogle Scholar
  32. 32.
    Pollard MR, Anderson L, Fan C, Hawkins DJ, Davies HM (1991) A specific acyl-ACP thioesterase implicated in medium-chain fatty acid production in immature cotyledons of Umbellularia californica. Arch Biochem Biophys 284:306–312PubMedCrossRefGoogle Scholar
  33. 33.
    Wu Y, Li R, Hildebrand DF (2012) Biosynthesis and metabolic engineering of palmitoleate production, an important contributor to human health and sustainable industry. Prog Lipid Res 51:340–349PubMedCrossRefGoogle Scholar
  34. 34.
    Cahoon EB, Shah S, Shanklin J, Browse J (1998) A determinant of substrate specificity predicted from the acyl-acyl carrier protein desaturase of developing cat’s claw seed. Plant Physiol 117:593–598PubMedCentralPubMedCrossRefGoogle Scholar
  35. 35.
    Lindqvist Y, Huang W, Schneider G, Shanklin J (1996) Crystal structure of delta9 stearoyl-acyl carrier protein desaturase from castor seed and its relationship to other di-iron proteins. EMBO J 15:4081–4092PubMedCentralPubMedGoogle Scholar
  36. 36.
    Cahoon EB, Lindqvist Y, Schneider G, Shanklin J (1997) Redesign of soluble fatty acid desaturases from plants for altered substrate specificity and double bond position. Proc Natl Acad Sci USA 94:4872–4877PubMedCentralPubMedCrossRefGoogle Scholar
  37. 37.
    Shukla VKS (1995) Cocoa butter properties and quality. Lipid Tech 7:54–57Google Scholar
  38. 38.
    Ohlrogge JB, Jaworski JG (1997) Regulation of fatty acid synthesis. Annu Rev Plant Physiol Plant Mol Biol 48:109–136PubMedCrossRefGoogle Scholar
  39. 39.
    Li N, Gügel IL, Giavalisco P, Zeisler V, Schreiber L, Soll J, Philippar K (2015) FAX1, a novel membrane protein mediating plastid fatty acid export. PLoS Biol 13:e1002053PubMedCentralPubMedCrossRefGoogle Scholar
  40. 40.
    Joyard J, Stumpf PK (1981) Synthesis of long-chain acyl-CoA in chloroplast envelope membranes. Plant Physiol 67:250–256PubMedCentralPubMedCrossRefGoogle Scholar
  41. 41.
    Koo AJ, Ohlrogge JB, Pollard M (2004) On the export of fatty acids from the chloroplast. J Biol Chem 279:16101–16110PubMedCrossRefGoogle Scholar
  42. 42.
    Schnurr JA, Shockey JM, de Boer GJ, Browse JA (2002) Fatty acid export from the chloroplast. Molecular characterization of a major plastidial acyl-coenzyme A synthetase from Arabidopsis. Plant Physiol 129:1700–1709PubMedCentralPubMedCrossRefGoogle Scholar
  43. 43.
    Zhao L, Katavic V, Li F, Haughn GW, Kunst L (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 64:1048–1058PubMedCrossRefGoogle Scholar
  44. 44.
    Xiao S, Chye ML (2011) New roles for acyl-CoA-binding proteins (ACBPs) in plant development, stress responses and lipid metabolism. Prog Lipid Res 50:141–151PubMedCrossRefGoogle Scholar
  45. 45.
    Cassagne C, Lessire R, Bessoule JJ, Moreau P, Creach A, Schneider F, Sturbois B (1994) Biosynthesis of very long chain fatty acids in higher plants. Prog Lipid Res 33:55–69PubMedCrossRefGoogle Scholar
  46. 46.
    Jessen D, Roth C, Wiermer M, Fulda M (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 167:351–366PubMedCrossRefGoogle Scholar
  47. 47.
    Kennedy EP (1961) Biosynthesis of complex lipids. Fed Proc 20:934–940PubMedGoogle Scholar
  48. 48.
    Kornberg A, Pricer WE Jr (1953) Enzymatic esterification of alpha-glycerophosphate by long chain fatty acids. J Biol Chem 204:345–357PubMedGoogle Scholar
  49. 49.
    Weiss SB, Kennedy EP, Kiyasu JY (1960) The enzymatic synthesis of triglycerides. J Biol Chem 235:40–44PubMedGoogle Scholar
  50. 50.
    Chapman KD, Dyer JM, Mullen RT (2013) Commentary: why don’t plant leaves get fat? Plant Sci 207:128–134PubMedCrossRefGoogle Scholar
  51. 51.
    Heinz E, Roughan PG (1983) Similarities and differences in lipid metabolism of chloroplasts isolated from 18:3 and 16:3 plants. Plant Physiol 72:273–279PubMedCentralPubMedCrossRefGoogle Scholar
  52. 52.
    Wallis JG, Browse J (2002) Mutants of Arabidopsis reveal many roles for membrane lipids. Prog Lipid Res 41:254–278PubMedCrossRefGoogle Scholar
  53. 53.
    Nichols BW, James AT, Breuer J (1967) Interrelationships between fatty acid biosynthesis and acyl-lipid synthesis in Chlorella vulgaris. Biochem J 104:486–496PubMedCentralPubMedCrossRefGoogle Scholar
  54. 54.
    Gurr MI, Robinson MP, James AT (1969) The mechanism of formation of polyunsaturated fatty acids by photosynthetic tissue. The tight coupling of oleate desaturation with phospholipid synthesis in Chlorella vulgaris. Eur J Biochem 9:70–78PubMedCrossRefGoogle Scholar
  55. 55.
    Roughan PG (1970) Turnover of the glycerolipids of pumpkin leaves. The importance of phosphatidylcholine. Biochem J 117:1–8PubMedCentralPubMedCrossRefGoogle Scholar
  56. 56.
    Chung AE, Law JH (1964) Cyclopropane fatty acid synthetase: partial purification and properties. Biochemistry (Mosc) 3:967–974CrossRefGoogle Scholar
  57. 57.
    Citharel B, Oursel A, Mazliak P (1983) Desaturation of oleoyl and linoleoyl residues linked to phospholipids in growing roots of yellow lupin. FEBS Lett 161:251–256CrossRefGoogle Scholar
  58. 58.
    Demandre C, Trémolières A, Justin AM, Mazliak P (1986) Oleate desaturation in six phosphatidylcholine molecular species from potato tuber microsomes. Biochim Biophys Acta 877:380–386CrossRefGoogle Scholar
  59. 59.
    Murphy DJ, Woodrow IE, Mukherjee KD (1985) Substrate specificities of the enzymes of the oleate desaturase system from photosynthetic tissue. Biochem J 225:267–270PubMedCentralPubMedCrossRefGoogle Scholar
  60. 60.
    Slack CR, Roughan PG, Browse J (1979) Evidence for an oleoyl phosphatidylcholine desaturase in microsomal preparations from cotyledons of safflower (Carthamus tinctorius) seed. Biochem J 179:649–656PubMedCentralPubMedCrossRefGoogle Scholar
  61. 61.
    Stymne S, Stobart AK, Glad G (1983) The role of the acyl-CoA pool in the synthesis of polyunsaturated 18-carbon fatty acids and triacylglycerol production in the microsomes of developing safflower seeds. Biochim Biophys Acta 752:198–208PubMedCrossRefGoogle Scholar
  62. 62.
    Jones AV, Harwood JL (1980) Desaturation of linoleic acid from exogenous lipids by isolated chloroplasts. Biochem J 190:851–854PubMedCentralPubMedCrossRefGoogle Scholar
  63. 63.
    Wharfe J, Harwood JL (1978) Fatty acid biosynthesis in the leaves of barley, wheat and pea. Biochem J 174:163–169PubMedCentralPubMedCrossRefGoogle Scholar
  64. 64.
    Ohnishi J, Yamada M (1982) Glycerolipid synthesis in avena leaves during greening of etiolated seedlings III. Synthesis of α-linolenoyl-monogalactosyl diacylglycerol from liposomal linoleoyl-phosphatidylcholine by avena plastids in the presence of phosphatidylcholine-exchange protein. Plant Cell Physiol 23:767–773Google Scholar
  65. 65.
    Pugh EL, Kates M (1973) Desaturation of phosphatidylcholine and phosphatidylethanolamine by a microsomal enzyme system from Candida lipolytica. Biochim Biophys Acta 316:305–316PubMedCrossRefGoogle Scholar
  66. 66.
    Pugh EL, Kates M, Szabo AG (1980) Fluorescence polarization studies of rat liver microsomes with altered phospholipid desaturase activities. Can J Biochem 58:952–958PubMedCrossRefGoogle Scholar
  67. 67.
    Avery SV, Lloyd D, Harwood JL (1995) Temperature-dependent changes in plasma-membrane lipid order and the phagocytotic activity of the amoeba Acanthamoeba castellanii are closely correlated. Biochem J 312:811–816PubMedCentralPubMedCrossRefGoogle Scholar
  68. 68.
    Jones AL, Lloyd D, Harwood JL (1993) Rapid induction of microsomal delta 12 (omega 6)-desaturase activity in chilled Acanthamoeba castellanii. Biochem J 296:183–188PubMedCentralPubMedCrossRefGoogle Scholar
  69. 69.
    Sayanova O, Haslam R, Guschina I, Lloyd D, Christie WW, Harwood JL, Napier JA (2006) A bifunctional Delta12, Delta15-desaturase from Acanthamoeba castellanii directs the synthesis of highly unusual n-1 series unsaturated fatty acids. J Biol Chem 281:36533–36541PubMedCrossRefGoogle Scholar
  70. 70.
    Lager I, Glab B, Eriksson L, Chen G, Banas A, Stymne S (2015) Novel reactions in acyl editing of phosphatidylcholine by lysophosphatidylcholine transacylase (LPCT) and acyl-CoA:glycerophosphocholine acyltransferase (GPCAT) activities in microsomal preparations of plant tissues. Planta 241:347–358PubMedCentralPubMedCrossRefGoogle Scholar
  71. 71.
    Chen G, Snyder CL, Greer MS, Weselake RJ (2011) Biology and biochemistry of plant phospholipases. Crit Rev Plant Sci 30:239–258CrossRefGoogle Scholar
  72. 72.
    Li M, Wei F, Tawfall A, Tang M, Saettele A, Wang X (2015) Overexpression of patatin-related phospholipase AIIIδ altered plant growth and increased seed oil content in camelina. Plant Biotechnol J 13:766–778PubMedCrossRefGoogle Scholar
  73. 73.
    Li M, Bahn SC, Fan C, Li J, Phan T, Ortiz M, Roth M, Welti R, Jaworski J, Wang X (2013) Patatin-related phospholipase pPLAIIIδ increases seed oil content with long chain fatty acids in Arabidopsis. Plant Physiol 162:39–51PubMedCentralPubMedCrossRefGoogle Scholar
  74. 74.
    Wang G, Ryu S, Wang X (2012) Plant phospholipases: an overview. Methods Mol Biol 861:123–137PubMedCrossRefGoogle Scholar
  75. 75.
    Lu CF, Xin ZG, Ren ZH, Miquel M, Browse J (2009) An enzyme regulating triacylglycerol composition is encoded by the ROD1 gene of Arabidopsis. Proc Natl Acad Sci USA 106:18837–18842PubMedCentralPubMedCrossRefGoogle Scholar
  76. 76.
    Slack CR, Roughan PG, Browse JA, Gardiner SE (1985) Some properties of cholinephosphotransferase from developing safflower cotyledons. Biochim Biophys Acta 833:438–448CrossRefGoogle Scholar
  77. 77.
    Dahlqvist A, Stahl U, Lenman M, Banas A, Lee M, Sandager L, Ronne H, Stymne H (2000) Phospholipid:diacylglycerol acyltransferase: an enzyme that catalyzes the acyl-CoA-independent formation of triacylglycerol in yeast and plants. Proc Natl Acad Sci USA 97:6487–6492PubMedCentralPubMedCrossRefGoogle Scholar
  78. 78.
    Bayon S, Chen G, Weselake R, Browse J (2015) A small phospholipase A2-α from castor catalyzes the removal of hydroxy fatty acids from phosphatidylcholine in transgenic Arabidopsis seeds. Plant Physiol 167:1259–1270PubMedCentralPubMedCrossRefGoogle Scholar
  79. 79.
    Lager I, Yilmaz JL, Zhou X-R, Jasieniecka K, Kazachkov M, Wang P, Zou J, Weselake R, Smith MA, Bayon S, Dyer JM, Shockey JM, Heinz E, Green A, Banas A, Stymne S (2013) Plant acyl-CoA:lysophosphatidylcholine acyltransferases (LPCATs) have different specificities in their forward and reverse reactions. J Biol Chem 288:36902–36914PubMedCentralPubMedCrossRefGoogle Scholar
  80. 80.
    Wang L, Kazachkov M, Shen W, Bai M, Wu H, Zou J (2014) Deciphering the roles of Arabidopsis LPCAT and PAH in phosphatidylcholine homeostasis and pathway coordination for chloroplast lipid synthesis. Plant J 80:965–976PubMedCrossRefGoogle Scholar
  81. 81.
    Wang LP, Shen WY, Kazachkov M, Chen GQ, Chen QL, Carlsson AS, Stymne S, Weselake RJ, Zou JT (2012) Metabolic interactions between the Lands cycle and the Kennedy pathway of glycerolipid synthesis in Arabidopsis developing seeds. Plant Cell 24:4652–4669PubMedCentralPubMedCrossRefGoogle Scholar
  82. 82.
    Williams JP, Imperial V, Khan MU, Hodson JN (2000) The role of phosphatidylcholine in fatty acid exchange and desaturation in Brassica napus L. leaves. Biochem J 349:127–133PubMedCentralPubMedCrossRefGoogle Scholar
  83. 83.
    Bates PD, Durrett TP, Ohlrogge JB, Pollard M (2009) Analysis of acyl fluxes through multiple pathways of triacylglycerol synthesis in developing soybean embryos. Plant Physiol 150:55–72PubMedCentralPubMedCrossRefGoogle Scholar
  84. 84.
    Bates PD, Fatihi A, Snapp AR, Carlsson AS, Browse J, Lu CF (2012) Acyl editing and headgroup exchange are the major mechanisms that direct polyunsaturated fatty acid flux into triacylglycerols. Plant Physiol 160:1530–1539PubMedCentralPubMedCrossRefGoogle Scholar
  85. 85.
    Snyder CL, Yurchenko OP, Siloto RMP, Chen X, Liu Q, Mietkiewska E, Weselake RJ (2009) Acyltransferase action in the modification of seed oil biosynthesis. New Biotechnol 26:11–16CrossRefGoogle Scholar
  86. 86.
    Lands WEM (1960) Metabolism of glycerolipids: II. The enzymatic acylation of lysolecithin. J Biol Chem 235:2233–2237PubMedGoogle Scholar
  87. 87.
    Kennedy EP, Weiss SB (1955) Cytidine diphosphate choline: a new intermediate in lecithin biosynthesis. J Am Chem Soc 77:250–251CrossRefGoogle Scholar
  88. 88.
    Kennedy EP, Weiss SB (1956) The function of cytidine coenzymes in the biosynthesis of phospholipides. J Biol Chem 222:193–214PubMedGoogle Scholar
  89. 89.
    Weiss SB, Smith SW, Kennedy EP (1958) The enzymatic formation of lecithin from cytidine diphosphate choline and D-1,2-diglyceride. J Biol Chem 231:53–64PubMedGoogle Scholar
  90. 90.
    Wilgram GF, Kennedy EP (1963) Intracellular distribution of some enzymes catalyzing reactions in the biosynthesis of complex lipids. J Biol Chem 238:2615–2619PubMedGoogle Scholar
  91. 91.
    Devor KA, Mudd JB (1971) Biosynthesis of phosphatidylcholine by enzyme preparations from spinach leaves. J Lipid Res 12:403–411PubMedGoogle Scholar
  92. 92.
    Marshall MO, Kates M (1972) Biosynthesis of phosphatidylglycerol by cell-free preparations from spinach leaves. Biochim Biophys Acta 260:558–570PubMedCrossRefGoogle Scholar
  93. 93.
    Marshall MO, Kates M (1974) Biosynthesis of nitrogenous phospholipids in spinach leaves. Can J Biochem 52:469–482PubMedCrossRefGoogle Scholar
  94. 94.
    Jolliot A, Justin AM, Bimont E, Mazliak P (1982) Regulation by lipids of plant microsomal enzymes: III. Phospholipid dependence of the cytidine-diphospho-choline phosphotransferase of potato microsomes. Plant Physiol 70:206–210PubMedCentralPubMedCrossRefGoogle Scholar
  95. 95.
    Slack CR, Campbell LC, Browse JA, Roughan PG (1983) Some evidence for the reversibility of the cholinephosphotransferasecatalysed reaction in developing linseed cotyledons in vivo. Biochim Biophys Acta 754:10–20CrossRefGoogle Scholar
  96. 96.
    Bafor M, Jonsson L, Stobart AK, Stymne S (1990) Regulation of triacylglycerol biosynthesis in embryos and microsomal preparations from the developing seeds of Cuphea lanceolata. Biochem J 272:31–38PubMedCentralPubMedCrossRefGoogle Scholar
  97. 97.
    Vogel G, Browse J (1996) Cholinephosphotransferase and diacylglycerol acyltransferase: substrate specificities at a key branch point in seed lipid metabolism. Plant Physiol 110:923–931PubMedCentralPubMedGoogle Scholar
  98. 98.
    Hu ZH, Ren ZH, Lu CF (2012) The phosphatidylcholine diacylglycerol cholinephosphotransferase is required for efficient hydroxy fatty acid accumulation in transgenic Arabidopsis. Plant Physiol 158:1944–1954PubMedCentralPubMedCrossRefGoogle Scholar
  99. 99.
    Bates PD, Browse J (2012) The significance of different diacylgycerol synthesis pathways on plant oil composition and bioengineering. Front Plant Sci 3:147PubMedCentralPubMedCrossRefGoogle Scholar
  100. 100.
    Wickramarathna AD, Siloto RMP, Mietkiewska E, Singer SD, Pan X, Weselake RJ (2015) Heterologous expression of flax PHOSPHOLIPID:DIACYLGLYCEROL CHOLINE-PHOSPHOTRANSFERASE (PDCT) increases polyunsaturated fatty acid content in yeast and Arabidopsis seeds. BMC Biotechnol 15:63PubMedCentralPubMedCrossRefGoogle Scholar
  101. 101.
    Oelkers P, Tinkelenberg A, Erdeniz N, Cromley D, Billheimer JT, Sturley SL (2000) A lecithin cholesterol acyltransferase-like gene mediates diacylglycerol esterification in yeast. J Biol Chem 275:15609–15612PubMedCrossRefGoogle Scholar
  102. 102.
    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 25:3506–3518PubMedCentralPubMedCrossRefGoogle Scholar
  103. 103.
    Pan X, Peng FY, Weselake R (2015) Genome-wide analysis of PHOSPHOLIPID:DIACYLGLYCEROL ACYLTRANSFERASE (PDAT) genes in plants reveals the eudicot-wide PDAT gene expansion and altered selective pressures acting on the core eudicot PDAT paralogs. Plant Physiol 167:887–904PubMedCentralPubMedCrossRefGoogle Scholar
  104. 104.
    Arabolaza A, Rodriguez E, Altabe S, Alvarez H, Gramajo H (2008) Multiple pathways for triacylglycerol biosynthesis in Streptomyces coelicolor. Appl Environ Microbiol 74:2573–2582PubMedCentralPubMedCrossRefGoogle Scholar
  105. 105.
    Mhaske V, Beldjilali K, Ohlrogge J, Pollard M (2005) Isolation and characterization of an Arabidopsis thaliana knockout line for phospholipid:diacylglycerol transacylase gene (At5g13640). Plant Physiol Biochem 43:413–417PubMedCrossRefGoogle Scholar
  106. 106.
    Fan J, Yan C, Roston R, Shanklin J, Xu C (2014) Arabidopsis lipins, PDAT1 acyltransferase, and SDP1 triacylglycerol lipase synergistically direct fatty acids toward β-oxidation, thereby maintaining membrane lipid homeostasis. Plant Cell 26:4119–4134PubMedCentralPubMedCrossRefGoogle Scholar
  107. 107.
    Zhang M, Fan J, Taylor DC, Ohlrogge JB (2009) DGAT1 and PDAT1 acyltransferases have overlapping functions in Arabidopsis triacylglycerol biosynthesis and are essential for normal pollen and seed development. Plant Cell 21:3885–3901PubMedCentralPubMedCrossRefGoogle Scholar
  108. 108.
    Tjellstrom H, Strawsine M, Ohlrogge JB (2015) Tracking synthesis and turnover of triacylglycerol in leaves. J Exp Bot 66:1453–1461PubMedCentralPubMedCrossRefGoogle Scholar
  109. 109.
    Kim HU, Lee KR, Go YS, Jung JH, Suh MC, Kim JB (2011) Endoplasmic reticulum-located PDAT1-2 from castor bean enhances hydroxy fatty acid accumulation in transgenic plants. Plant Cell Physiol 52:983–993PubMedCrossRefGoogle Scholar
  110. 110.
    van Erp H, Bates PD, Burgal J, Shockey J, Browse J (2011) Castor phospholipid:diacylglycerol acyltransferase facilitates efficient metabolism of hydroxy fatty acids in transgenic Arabidopsis. Plant Physiol 155:683–693PubMedCentralPubMedCrossRefGoogle Scholar
  111. 111.
    Pan X, Siloto RM, Wickramarathna AD, Mietkiewska E, Weselake RJ (2013) Identification of a pair of phospholipid:diacylglycerol acyltransferases from developing flax (Linum usitatissimum L.) seed catalyzing the selective production of trilinolenin. J Biol Chem 288:24173–24188PubMedCentralPubMedCrossRefGoogle Scholar
  112. 112.
    Yoon K, Han D, Li Y, Sommerfeld M, Hu Q (2012) Phospholipid:diacylglycerol acyltransferase is a multifunctional enzyme involved in membrane lipid turnover and degradation while synthesizing triacylglycerol in the unicellular green microalga Chlamydomonas reinhardtii. Plant Cell 24:3708–3724PubMedCentralPubMedCrossRefGoogle Scholar
  113. 113.
    Stahl U, Carlsson AS, Lenman M, Dahlqvist A, Huang BQ, Banas W, Banas A, Stymne S (2004) Cloning and functional characterization of a phospholipid:diacylglycerol acyltransferase from Arabidopsis. Plant Physiol 135:1324–1335PubMedCentralPubMedCrossRefGoogle Scholar

Copyright information

© AOCS 2015

Authors and Affiliations

  1. 1.Alberta Innovates Phytola Centre, Department of Agricultural, Food and Nutritional ScienceUniversity of AlbertaEdmontonCanada
  2. 2.School of BiosciencesCardiff UniversityCardiffWales, UK
  3. 3.Department of Biological SciencesUniversity of ManitobaWinnipegCanada

Personalised recommendations