title: Subcellular localization of rice acyl-CoA-binding proteins (ACBPs) indicates that OsACBP6::GFP is targeted to the peroxisomes.
abstract: Acyl-CoA-binding proteins (ACBPs) show conservation at the acyl-CoA-binding (ACB) domain which facilitates binding to acyl-CoA esters. In Arabidopsis thaliana, six ACBPs participate in development and stress responses. Rice (Oryza sativa) also contains six genes encoding ACBPs. We investigated differences in subcellular localization between monocot rice and eudicot A. thaliana ACBPs. The subcellular localization of the six OsACBPs was achieved via transient expression of green fluorescence protein (GFP) fusions in tobacco (Nicotiana tabacum) epidermal cells, and stable transformation of A. thaliana. As plant ACBPs had not been reported in the peroxisomes, OsACBP6::GFP localization was confirmed by transient expression in rice sheath cells. The function of OsACBP6 was investigated by overexpressing 35S::OsACBP6 in the peroxisomal abc transporter1 (pxa1) mutant defective in peroxisomal fatty acid β-oxidation. As predicted, OsACBP1::GFP and OsACBP2::GFP were localized to the cytosol, and OsACBP4::GFP and OsACBP5::GFP to the endoplasmic reticulum (ER). However, OsACBP3::GFP displayed subcellular multi-localization while OsACBP6::GFP was localized to the peroxisomes. 35S::OsACBP6-OE/pxa1 lines showed recovery in indole-3-butyric acid (IBA) peroxisomal β-oxidation, wound-induced VEGETATIVE STORAGE PROTEIN1 (VSP1) expression and jasmonic acid (JA) accumulation. These findings indicate a role for OsACBP6 in peroxisomal β-oxidation, and suggest that rice ACBPs are involved in lipid degradation in addition to lipid biosynthesis.
authors.affiliation: School of Biological Sciences, The University of Hong Kong, Pokfulam, Hong Kong, China.|College of Life Science, Northeast Forestry University, Harbin, China.
authors.forename: Wei
authors.initials: W
authors.lastname: Meng
authors.affiliation: School of Biological Sciences, The University of Hong Kong, Pokfulam, Hong Kong, China.
authors.forename: An-Shan
authors.initials: AS
authors.lastname: Hsiao
authors.affiliation: School of Life Sciences, The Chinese University of Hong Kong, Shatin, New Territories, Hong Kong, China.
authors.forename: Caiji
authors.initials: C
authors.lastname: Gao
authors.affiliation: School of Life Sciences, The Chinese University of Hong Kong, Shatin, New Territories, Hong Kong, China.
authors.forename: Liwen
authors.initials: L
authors.lastname: Jiang
authors.affiliation: School of Biological Sciences, The University of Hong Kong, Pokfulam, Hong Kong, China.
authors.forename: Mee-Len
authors.initials: ML
authors.lastname: Chye
mesh_terms: D017360:Arabidopsis / Q000235:genetics / Q000378:metabolism; D029681:Arabidopsis Proteins / Q000235:genetics / Q000378:metabolism; D002352:Carrier Proteins / Q000235:genetics / Q000378:metabolism; D003517:Cyclopentanes / Q000378:metabolism; D003600:Cytosol / Q000378:metabolism; D049452:Green Fluorescent Proteins / Q000235:genetics / Q000378:metabolism; D007211:Indoles / Q000378:metabolism; D050356:Lipid Metabolism / Q000235:genetics; D012275:Oryza / Q000235:genetics / Q000378:metabolism*; D054883:Oxylipins / Q000378:metabolism; D020675:Peroxisomes / Q000235:genetics / Q000378:metabolism*; D010940:Plant Proteins / Q000235:genetics / Q000378:metabolism*; D030821:Plants, Genetically Modified; D021381:Protein Transport; D014026:Nicotiana / Q000235:genetics
chemical_list: D029681:Arabidopsis Proteins; D002352:Carrier Proteins; D003517:Cyclopentanes; D007211:Indoles; D054883:Oxylipins; D010940:Plant Proteins; C014612:indolebutyric acid; D049452:Green Fluorescent Proteins; C011006:jasmonic acid
keywords: Arabidopsis thaliana; Oryza sativa (rice); acyl-CoA-binding protein (ACBP); indole-3-butyric acid; jasmonic acid; peroxisomal β-oxidation; subcellular localization
references.citation: Awai K, Xu CC, Tamot B, Benning C. 2006. A phosphatidic acid-binding protein of the chloroplast inner envelope membrane involved in lipid trafficking. Proceedings of the National Academy of Sciences, USA 103: 10817-10822.
references.citation: Benning C. 2008. A role for lipid trafficking in chloroplast biogenesis. Progress in Lipid Research 47: 381-389.
references.citation: Benning C. 2009. Mechanisms of lipid transport involved in organelle biogenesis in plant cells. Annual Review of Cell and Developmental Biology 25: 71-91.
references.citation: Benning C, Xu CC, Awai K. 2006. Non-vesicular and vesicular lipid trafficking involving plastids. Current Opinion in Plant Biology 9: 241-247.
references.citation: Blée E. 1998. Phytooxylipins and plant defense reactions. Progress in Lipid Research 37: 33-72.
references.citation: Chaw SM, Chang CC, Chen HL, Li WH. 2004. Dating the monocot-dicot divergence and the origin of core eudicots using whole chloroplast genomes. Journal of Molecular Evolution 58: 424-441.
references.citation: Chen QF, Xiao S, Chye ML. 2008. Overexpression of the Arabidopsis 10-kilodalton acyl-CoA-binding protein ACBP6 enhances freezing tolerance. Plant Physiology 148: 304-315.
references.citation: Chen QF, Xiao S, Qi WQ, Mishra G, Ma JY, Wang MF, Chye ML. 2010. The Arabidopsis acbp1acbp2 double mutant lacking acyl-CoA-binding proteins ACBP1 and ACBP2 is embryo lethal. New Phytologist 186: 843-855.
references.citation: Chye ML. 1998. Arabidopsis cDNA encoding a membrane-associated protein with an acyl-CoA-binding domain. Plant Molecular Biology 38: 827-838.
references.citation: Chye ML, Huang BQ, Zee SY. 1999. Isolation of a gene encoding Arabidopsis membrane associated acyl-CoA-binding protein and immunolocalization of its gene product. Plant Journal 18: 205-214.
references.citation: Chye ML, Li HY, Yung MH. 2000. Single amino acid substitutions at the acyl-CoA-binding domain interrupt 14[C]palmitoyl-CoA binding of ACBP2, an Arabidopsis acyl-CoA-binding protein with ankyrin repeats. Plant Molecular Biology 44: 711-721.
references.citation: Clough SJ, Bent AF. 1998. Floral dip: a simplified method for Agrobacterium-mediated transformation of Arabidopsis thaliana. Plant Journal 16: 735-743.
references.citation: Cooper TG, Beevers H. 1969. β-oxidation in glyoxysomes from castor bean endosperm. Journal of Biological Chemistry 244: 3514-3520.
references.citation: Coutos-Thevenot P, Jouenne T, Maes O, Guerbette F, Grosbois M, Le Caer JP, Boulay M, Deloire A, Kader JC, Guern J. 1993. Four 9-kDa proteins excreted by somatic embryos of grapevine are isofoms of lipid-transfer proteins. European Journal of Biochemistry 217: 885-889.
references.citation: Dave A, Hernández ML, He ZS, Andriotis VME, Vaistij FE, Larson TR, Graham IA. 2011. 12-Oxo-phytodienoic acid accumulation during seed development represses seed germination in Arabidopsis. Plant Cell 23: 583-599.
references.citation: De Marcos Lousa C, van Roermund CW, Postis VL, Dietrich D, Kerr ID, Wanders RJ, Baldwin SA, Baker A, Theodoulou FL. 2013. Intrinsic acyl-CoA thioesterase activity of a peroxisomal ATP binding cassette transporter is required for transport and metabolism of fatty acids. Proceedings of the National Academy of Sciences, USA 110: 1279-1284.
references.citation: Du ZY, Chen MX, Chen QF, Xiao S, Chye ML. 2013a. Overexpression of Arabidopsis acyl-CoA-binding protein ACBP2 enhances drought tolerance. Plant, Cell & Environment 36: 300-314.
references.citation: Du ZY, Chen MX, Chen QF, Xiao S, Chye ML. 2013b. Arabidopsis acyl-CoA-binding protein ACBP1 participates in the regulation of seed germination and seedling development. Plant Journal 74: 294-309.
references.citation: Du ZY, Chye ML. 2013. Interactions between Arabidopsis acyl-CoA-binding proteins and protein partners. Planta 238: 239-245.
references.citation: Du ZY, Xiao S, Chen QF, Chye ML. 2010. Depletion of the membrane-associated acyl-coenzyme A-binding protein ACBP1 enhances the ability of cold acclimation in Arabidopsis. Plant Physiology 152: 1585-1597.
references.citation: Engeseth NJ, Pacovsky RS, Newman T, Ohlrogge JB. 1996. Characterization of an acyl-CoA-binding protein from Arabidopsis thaliana. Archives of Biochemistry and Biophysics 331: 55-62.
references.citation: Faergeman NJ, Knudsen J. 1997. Role of long-chain fatty acyl-CoA esters in the regulation of metabolism and in cell signaling. Journal of Biochemistry 323: 1-12.
references.citation: Faergeman NJ, Wadum M, Feddersen S, Burton M, Kragelund BB, Knudsen J. 2007. Acyl-CoA-binding proteins; structural and functional conservation over 2000 MYA. Molecular and Cellular Biochemistry 299: 55-65.
references.citation: Farese RV, Walther TC. 2009. Lipid droplets finally get a little R-E-S-P-E-C-T. Cell 139: 855-860.
references.citation: Finkemeier I, Laxa M, Miguet L, Howden AJM, Sweetlove LJ. 2011. Proteins of diverse function and subcellular location are lysine acetylated in Arabidopsis. Plant Physiology 155: 1779-1790.
references.citation: Flügge U, van Meer G. 2006. ABC Transporters. FEBS Letters 580: 997.
references.citation: Footitt S, Slocombe SP, Larner V, Kurup S, Wu YS, Larson T, Graham I, Baker A, Holdsworth M. 2002. Control of germination and lipid mobilization by COMATOSE, the Arabidopsis homologue of human ALDP. EMBO Journal 21: 2912-2922.
references.citation: Friedman JR, Voeltz GK. 2011. The ER in 3D: a multifunctional dynamic membrane network. Trends in Cell Biology 21: 709-711.
references.citation: Fyrst H, Knudsen J, Schott MA, Lubin BH, Kuypers FA. 1995. Detection of acyl-CoA-binding protein in human red blood cells and investigation of its role in membrane phospholipid renewal. Biochemical Journal 306: 793-799.
references.citation: Gao W, Li HY, Xiao S, Chye ML. 2010. Acyl-CoA-binding protein 2 binds lysophospholipase 2 and lysoPC to promote tolerance to cadmium-induced oxidative stress in transgenic Arabidopsis. Plant Journal 62: 989-1003.
references.citation: Gao W, Xiao S, Li HY, Tsao SW, Chye ML. 2009. Arabidopsis thaliana acyl-CoA-binding protein ACBP2 interacts with a heavy-metal-binding protein AtFP6. New Phytologist 181: 89-102.
references.citation: Hayashi M, Nito K, Takei-Hoshi R, Yagi M, Kondo M, Suenaga A, Yamaya T, Nishimura M. 2002. Ped3p is a peroxisomal ATP-Binding Cassette transporter that might supply substrates for fatty acid β-oxidation. Plant and Cell Physiology 43: 1-11.
references.citation: Hills MJ, Dann R, Lydiate D, Sharpe A. 1994. Molecular cloning of a cDNA from Brassica napus L. for a homologue of acyl-CoA-binding protein. Plant Molecular Biology 25: 917-920.
references.citation: Hu J, Baker A, Bartel B, Linka N, Mullen RT, Reumann S, Zolman BK. 2012. Plant peroxisomes: biogenesis and function. Plant Cell 24: 2279-2303.
references.citation: Huang H, Atshaves BP, Frolov A, Kier AB, Schroeder F. 2005. Acyl-coenzyme A binding protein expression alters liver fatty acyl-coenzyme A metabolism. Biochemistry 44: 10282-10297.
references.citation: Hunt MC, Tillander V, Alexson SE. 2014. Regulation of peroxisomal lipid metabolism: the role of acyl-CoA and coenzyme A metabolizing enzymes. Biochimie 98: 45-55.
references.citation: Jelitto-Van Dooren EPWM, Vidal S, Denecke J. 1999. Anticipating endoplasmic reticulum stress. A novel early response before pathogenesis-related gene induction. Plant Cell 11: 1935-1944.
references.citation: Kader JC. 1996. Lipid-transfer proteins in plants. Annual Review of Plant Physiology and Plant Molecular Biology 47: 627-654.
references.citation: Kaur N, Hu JP. 2011. Defining the plant peroxisomal proteome: from Arabidopsis to rice. Frontiers in Plant Science 2: 103.
references.citation: Knudsen J, Faergeman NJ, Skott H, Hummel R, Borsting C, Rose TM, Andersen JS, Hojrup P, Roepstorff P, Kristiansen K. 1994. Yeast acyl-CoA binding protein: acyl-CoA-binding affinity and effect on intracellular acyl-CoA pool size. Biochemical Journal 302: 479-485.
references.citation: Kroll D, Meierhoff K, Bechtold N, Kinoshita M, Westphal S, Vothknecht UC, Soll J, Westhoff P. 2001. VIPP1, a nuclear gene of Arabidopsis thaliana essential for thylakoid membrane formation. Proceedings of the National Academy of Sciences, USA 98: 4238-4242.
references.citation: Kuo MH, Allis CD. 1998. Roles of histone acetyltransferases and deacetylases in gene regulation. BioEssays 20: 615-626.
references.citation: Leung KC, Li HY, Mishra G, Chye ML. 2004. ACBP4 and ACBP5, novel Arabidopsis acyl-CoA-binding proteins with kelch motifs that bind oleoyl-CoA. Plant Molecular Biology 55: 297-309.
references.citation: Leung KC, Li HY, Xiao S, Tse MH, Chye ML. 2006. Arabidopsis ACBP3 is an extracellularly targeted acyl-CoA-binding protein. Planta 223: 871-881.
references.citation: Li HY, Chye ML. 2003. Membrane localization of Arabidopsis acyl-CoA-binding protein ACBP2. Plant Molecular Biology 51: 483-492.
references.citation: Li HY, Chye ML. 2004. Arabidopsis acyl-CoA-binding protein ACBP2 interacts with an ethylene responsive element-binding protein, AtEBP, via its ankyrin repeats. Plant Molecular Biology 54: 233-243.
references.citation: Li HY, Xiao S, Chye ML. 2008. Ethylene- and pathogen-inducible Arabidopsis acyl-CoA-binding protein 4 interacts with an ethylene-responsive element binding protein. Journal of Experimental Botany 59: 3997-4006.
references.citation: Li-Beisson Y, Shorrosh B, Beisson F, Andersson MX, Arondel V, Bates PD, Baud S, Bird D, DeBono A, Durrett TP et al. 2010. Acyl-lipid metabolism. The Arabidopsis Book 8: e0133.
references.citation: Linka N, Esser C. 2012. Transport proteins regulate the flux of metabolites and cofactors across the membrane of plant peroxisomes. Frontiers in Plant Science 3: 3.
references.citation: Lu B, Xu CC, Awai K, Jones AD, Benning C. 2007. A small ATPase protein of Arabidopsis, TGD3, involved in chloroplast lipid import. Journal of Biological Chemistry 282: 35945-35953.
references.citation: Mandrup S, Jepsen R, Skott H, Rosendal J, Hojrup P, Kristiansen K, Knudsen J. 1993. Effect of heterologous expression of acyl-CoA-binding protein on acyl-CoA level and composition in yeast. Biochemical Journal 290: 369-374.
references.citation: Mclntosh KB, Hulm JL, Young LW, Bonham-Smith PC. 2004. A rapid Agrobacterium-mediated Arabidopsis thaliana transient assay system. Plant Molecular Biology Reporter 22: 53-61.
references.citation: van Meer G, Voelker DR, Feigenson GW. 2008. Membrane lipids: where they are and how they behave. Nature Reviews Molecular Cell Biology 9: 112-124.
references.citation: Meng W. 2012. Characterization of acyl-coenzyme A-binding proteins from rice. PhD thesis, University of Hong Kong, Hong Kong, China.
references.citation: Meng W, Su YC, Saunders RM, Chye ML. 2011. The rice acyl-CoA-binding protein gene family: phylogeny, expression and functional analysis. New Phytologist 189: 1170-1184.
references.citation: Mikkelsen J, Knudsen J. 1987. Acyl-CoA-binding protein from cow. Binding characteristics and cellular and tissue distribution. Biochemical Journal 248: 709-714.
references.citation: Mundy J, Rogers JC. 1986. Selective expression of a probable amylase/protease inhibitor in barley aleurone cells: comparison to the barley amylase/subtilisin inhibitor. Planta 169: 51-63.
references.citation: Murashige T, Skoog F. 1962. A revised medium for rapid growth and bioassays with tobacco tissue cultures. Physiologia Plantarum 15: 473-497.
references.citation: Nyathi Y, Marcos Lousa C, van Roermund CW, Wanders RJA, Johnson B, Baldwin SA, Theodoulou FL, Baker A. 2010. The Arabidopsis peroxisomal ABC transporter, Comatose, complements the Saccharomyces cerevisiae pxa1 pxa2∆ mutant for metabolism of long-chain fatty acids and exhibits fatty acyl-coA-stimulated ATPase activity. Journal of Biological Chemistry 285: 29892-29902.
references.citation: Ohlrogge JB, Browse J. 1995. Lipid biosynthesis. Plant Cell 7: 957-970.
references.citation: Pan XQ, Welti R, Wang XM. 2010. Quantitative analysis of major plant hormones in crude plant extracts by high-performance liquid chromatography-mass spectrometry. Nature Protocols 5: 986-992.
references.citation: Park SJ, Gidda SK, James CN, Horn PJ, Khuu N, Seay DC, Keereetaweep J, Chapman KD, Mullen RT, Dyer JM. 2013. The α/β hydrolase CGI-58 and peroxisomal transport protein PXA1 coregulate lipid homeostasis and signaling in Arabidopsis. Plant Cell 25: 1726-1739.
references.citation: Poot M, Zhang YZ, Krämer JA, Wells KS, Jones LJ, Hanzel DK, Lugade AG, Singer VL, Haugland RP. 1996. Analysis of mitochondrial morphology and function with novel fixable fluorescent stains. Journal of Histochemistry & Cytochemistry 44: 1363-1372.
references.citation: Rasmussen JT, Rosendal J, Knudsen J. 1993. Interaction of acyl-CoA-binding protein (ACBP) on processes for which acyl-CoA is a substrate, product or inhibitor. Journal of Biochemistry 292: 907-913.
references.citation: Ritzenthaler C, Nebenführ A, Movafeghi A, Stussi-Garaud C, Behnia L, Pimpl P, Staehelin LA, Robinson DG. 2002. Reevaluation of the effects of brefeldin A on plant cells using tobacco Bright Yellow 2 cells expressing Golgi-targeted green fluorescent protein and COPI antisera. Plant Cell 14: 237-261.
references.citation: Ruzin SE. 1999. Plant microtechnique and microscopy. New York, NY, USA: Oxford University Press.
references.citation: Shi DQ, Liu J, Xiang YH, Ye D, Sundaresan V, Yang WC. 2005. SLOW WALKER1, essential for gametogenesis in Arabidopsis, encodes a WD40 protein involved in 18S ribosomal RNA biogenesis. Plant Cell 17: 2340-2354.
references.citation: Smith JA, Krauss MR, Borkird C, Sung ZR. 1988. A nuclear-protein associated with cell divisions in plants. Planta 174: 462-472.
references.citation: Soltis P, Soltis D, Edwards C. 2005. Angiosperms. Flowering plants. Tree of Life Web Project. [WWW document] URL http://tolweb.org/Angiosperms/20646/2005.06.03 [accessed 4 April 2012].
references.citation: Sparkes I, Hawes C, Frigerio L. 2011. FrontiERs: movers and shapers of the higher plant cortical endoplasmic reticulum. Current Opinion in Plant Biology 14: 658-665.
references.citation: Sterk P, Booij H, Scheleekens GA, Van Kammen A, De Vries SC. 1991. Cell specific expression of the carrot EP2 lipid transfer protein gene. Plant Cell 3: 907-921.
references.citation: Theodoulou FL, Holdsworthb M, Baker A. 2006. Peroxisomal ABC transporters. FEBS Letters 580: 1139-1155.
references.citation: Theodoulou FL, Job K, Slocombe SP, Footitt S, Holdsworth M, Baker A, Larson TR, Graham IA. 2005. Jasmonic acid levels are reduced in COMATOSE ATP-binding cassette transporter mutants. Implications for transport of jasmonate precursors into peroxisomes. Plant Physiology 137: 835-840.
references.citation: Toulmay A, Prinz WA. 2011. Lipid transfer and signaling at organelle contact sites: the tip of the iceberg. Current Opinion in Cell Biology 23: 458-463.
references.citation: Turner BM. 1991. Histone acetylation and control of gene expression. Journal of Cell Science 99: 13-20.
references.citation: Tusnády GE, Simon I. 2001. The HMMTOP transmembrane topology prediction server. Bioinformatics 17: 849-850.
references.citation: Vadaie N, Hulinsky RS, Jarvis DL. 2002. Identification and characterization of a Drosophila melanogaster ortholog of human beta 1,4-galactosyltransferase VII. Glycobiology 12: 589-597.
references.citation: Vick BA, Zimmerman DC. 1983. The biosynthesis of jasmonic acid: a physiological role for plant lipoxygenase. Biochemical and Biophysical Research Communications 111: 470-477.
references.citation: Wang YM, Wu JN, Kim SG, Kim ST, Kang KY. 2013. A transient gene expression protocol for subcellular protein localization and protein secretion analyses in rice. Protocol Exchange. doi: 10.1038/protex.2013.064.
references.citation: Wang Z, Xu CC, Benning C. 2012. TGD4 involved in endoplasmic reticulum-to-chloroplast lipid trafficking is a phosphatidic acid binding protein. Plant Journal 70: 614-623.
references.citation: Weber H, Vick BA, Farmer EE. 1997. Dinor-oxo-phytodienoic acid: a new hexadecanoid signal in the jasmonate family. Proceedings of the National Academy of Sciences, USA 94: 10473-10478.
references.citation: Xiao S, Chen QF, Chye ML. 2009. Light-regulated Arabidopsis ACBP4 and ACBP5 encode cytosolic acyl-CoA-binding proteins that bind phosphatidylcholine and oleoyl-CoA ester. Plant Physiology and Biochemistry 47: 926-933.
references.citation: Xiao S, Chye ML. 2009. An Arabidopsis family of six acyl-CoA-binding proteins has three cytosolic members. Plant Physiology and Biochemistry 47: 479-484.
references.citation: Xiao S, Chye ML. 2011a. New roles for acyl-CoA-binding proteins in plant development, stress responses and lipid metabolism. Progress in Lipid Research 50: 141-151.
references.citation: Xiao S, Chye ML. 2011b. Overexpression of Arabidopsis ACBP3 enhances NPR1-dependent plant resistance to Pseudomonas syringe pv tomato DC3000. Plant Physiology 156: 2069-2081.
references.citation: Xiao S, Gao W, Chen QF, Chan SW, Zheng SX, Ma JY, Wang MF, Welti R, Chye ML. 2010. Overexpression of Arabidopsis acyl-CoA-binding protein ACBP3 promotes starvation-induced and age-dependent leaf senescence. Plant Cell 22: 1463-1482.
references.citation: Xiao S, Gao W, Chen QF, Ramalingam S, Chye ML. 2008a. Overexpression of membrane-associated acyl-CoA-binding protein ACBP1 enhances lead tolerance in Arabidopsis. Plant Journal 54: 141-151.
references.citation: Xiao S, Li HY, Zhang JP, Chan SW, Chye ML. 2008b. Arabidopsis acyl-CoA-binding proteins ACBP4 and ACBP5 are subcellularly localized to the cytosol and ACBP4 depletion affects membrane lipid composition. Plant Molecular Biology 68: 571-583.
references.citation: Xu CC, Fan J, Cornish AJ, Benning C. 2008. Lipid trafficking between the endoplasmic reticulum and the plastid in Arabidopsis requires the extraplastidic TGD4 protein. Plant Cell 20: 2190-2204.
references.citation: Xu CC, Fan J, Froehlich J, Awai K, Benning C. 2005. Mutation of the TGD1 chloroplast envelope protein affects phosphatidate metabolism in Arabidopsis. Plant Cell 17: 3094-3110.
references.citation: Xu CC, Fan J, Riekhof W, Froehlich JE, Benning C. 2003. A permease-like protein involved in ER to thylakoid lipid transfer in Arabidopsis. EMBO Journal 22: 2370-2379.
references.citation: Yurchenko OP, Nykiforuk CL, Moloney MM, Ståhl U, Banaś A, Stymne S, Weselake RJ. 2009. A 10-kDa acyl-CoA-binding protein (ACBP) from Brassica napus enhances acyl exchange between acyl-CoA and phosphatidylcholine. Plant Biotechnology Journal 7: 602-610.
references.citation: 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 Physiology 127: 1266-1278.