Cutin and Suberin

Cutin and suberin are cell-wall associated glycerolipid polymers specific to plants. Cutin forms the framework of the cuticle sealing the aerial epidermis, whereas suberin is present in the periderm of barks and underground organs. Suberised walls are also found in root internal tissues. Barriers based on cutin and suberin restrict transport of water across cell walls and limit pathogen invasions. Chemical analysis shows that both polymers are polyesters composed mostly of C16-C18 hydroxyacids, diacids and epoxyacids esterified to each other and to glycerol. Suberin, whose best known form is cork, usually differs from cutin by a higher content in C20-C24 aliphatics and aromatics. In the last decade, the identification of Arabidopsis mutants affected in cutin or suberin content has allowed the identification of several proteins involved in polyester biosynthesis, including acyltransferases with unique specificities, fatty acid hydroxylases, acyl-CoA synthetases, fatty acid elongases and an ABC transporter.

Key Concepts:

  • The epidermal cells and suberising cells have specialised enzymes that convert the common cellular fatty acids into the unique components of cutin and suberin, respectively.
  • Oxygenated fatty acid monomers are produced by fatty acid oxidases of the cytochrome P450 superfamily.
  • Acylglycerol dimers can be synthesised by special glycerol-3-phosphate acyltransferases.
  • Whether polymerisation of dimers and monomers occurs in intracellular compartments or in the cell walls is still unknown.
  • How cutin and suberin polymers are linked to the cell walls remains to be determined.
  • Cutin contributes to the formation of surface nanostructures in epidermis.

Keywords: cutin; suberin; waxes; oxygenated fatty acids; glycerol-3-phosphate acyltransferase; P450 monooxygenase; cuticle; cork

Figure 1. Occurrence and morphology of cutin and suberin. Top panel: Schematic representation of the cuticle (left) and suberised cell wall (right). Bottom panel: Observation of cutin and suberin using electron microscope. (a) A TEM image of a cross section view of Arabidopsis stems (bar=5 m). (b) A scanning electron microscopy image of epidermal surface of Arabidopsis sepal (bar=1 m) (adapted from Li-Beisson et al. (2009), copyright (2009) National Academy of Sciences, USA). (c) A TEM image of a cross section view of Arabidopsis roots (bar=0.2 m) (adapted from Molina et al., 2009, copyright American Society of Plant Biologists, www.plantphysiol.org). Abbreviations: CW, cell wall and PC, peridermal cell.
Figure 2. Structure of the most common monomers of cutin and the tentative reactions/enzymes involved in their syntheses. In red: enzymes identified and in blue: unknown. Note: substrates are likely to be acyl-CoAs (R=CoA) but it cannot be ruled out that they are free fatty acids (R=H) or glycerolipids; DH, dehydrogenase.
Figure 3. A possible structure of polyesters cutin and suberin. Note: R, other domains of polyester.
Figure 4. Tentative biochemical steps identified in cutin/suberin biosynthetic pathways. Abbreviations: ABC transporter, ATP-binding cassette transporter; BDG, an / hydrolase family protein; CW, cell wall; DCR, defective in cuticular ridges, a member of the BAHD transferase family; ER, endoplasmic reticulum; FATB, fatty acid thioesterase; GPAT, glycerol 3-phosphate acyltransferase; KCS, ketoacyl-CoA synthetase; LACS, long chain acyl-CoA synthetase; PM, plasma membrane and WIN1: wax inducer 1.
Figure 5. The regiospecificity of glycerol 3-phosphate acyltransferase (GPAT) may control the flux of acyl chains to their final site of deposition. Abbreviations: G3P, glycerol 3-phosphate; MAG, monoacylglycerol; PA, phosphatidic acid; CW: cell wall and PM, plasma membrane. Courtesy of Dr Weili Yang and Professor John Ohlrogge, Michigan State University.
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 References
    Aharoni A, Dixit S, Jetter R et al. (2004) The SHINE clade of AP2 domain transcription factors activates wax biosynthesis, alters cuticle properties, and confers drought tolerance when overexpressed in Arabidopsis. Plant Cell 16: 2463–2480.
    Beisson F, Li Y, Bonaventure G, Pollard M and Ohlrogge JB (2007) The acyltransferase GPAT5 is required for the synthesis of suberin in seed coat and root of Arabidopsis. Plant Cell 19: 351–368.
    Bernards MA (2002) Demystifying suberin. Canadian Journal of Botany 80: 227–240.
    Bernards MA, Lopez ML, Zajicek J and Lewis NG (1995) Hydroxycinnamic acid-derived polymers constitute the polyaromatic domain of suberin. Journal of Biological Chemistry 270: 7382–7386.
    Bird D, Beisson F, Brigham A et al. (2007) Characterization of Arabidopsis ABCG11/WBC11, an ATP binding cassette (ABC) transporter that is required for cuticular lipid secretion. Plant Journal 52: 485–498.
    Bonaventure G, Beisson F, Ohlrogge J and Pollard M (2004) Analysis of the aliphatic monomer composition of polyesters associated with Arabidopsis epidermis: occurrence of octadeca-cis-6, cis-9-diene-1,18-dioate as the major component. Plant Journal 40: 920–930.
    Chaves I, Pinheiro C, Paiva JA et al. (2009) Proteomic evaluation of wound-healing processes in potato (Solanum tuberosum L.) tuber tissue. Proteomics 9: 4154–4175.
    Compagnon V, Diehl P, Benveniste I et al. (2009) CYP86B1 is required for very long chain omega-hydroxyacid and alpha, omega -dicarboxylic acid synthesis in root and seed suberin polyester. Plant Physiology 150: 1831–1843.
    Debono A, Yeats TH, Rose JK et al. (2009) Arabidopsis LTPG is a glycosylphosphatidylinositol-anchored lipid transfer protein required for export of lipids to the plant surface. Plant Cell 21: 1230–1238.
    Espelie KE, Dean BB and Kolattukudy PE (1979) Composition of lipid-derived polymers from different anatomical regions of several plant species. Plant Physiology 64: 1089–1093.
    Fang X, Qiu F, Yan B et al. (2001) NMR studies of molecular structure in fruit cuticle polyesters. Phytochemistry 57: 1035–1042.
    Franke R, Briesen I, Wojciechowski T et al. (2005) Apoplastic polyesters in Arabidopsis surface tissues – a typical suberin and a particular cutin. Phytochemistry 66: 2643–2658.
    Franke R, Höfer R, Briesen I et al. (2009) The DAISY gene from Arabidopsis encodes a fatty acid elongase condensing enzyme involved in the biosynthesis of aliphatic suberin in roots and the chalaza-micropyle region of seeds. Plant Journal 57: 80–95.
    Franke R and Schreiber L (2007) Suberin – a biopolyester forming apoplastic plant interfaces. Current Opinion in Plant Biology 10: 252–259.
    Graça J and Pereira H (2000) Diglycerol alkenedioates in suberin: building units of a poly(acylglycerol) polyester. Biomacromolecules 1: 519–522.
    Graça J and Santos S (2007) Suberin: a biopolyester of plants’ skin. Macromolecular Bioscience 7: 128–135.
    Graça J, Schreiber L, Rodrigues J and Pereira H (2002) Glycerol and glyceryl esters of omega-hydroxyacids in cutins. Phytochemistry 61: 205–215.
    Hima Rani S, Anantha Krishna TH, Saha S, Negi AS and Rajasekharan R (2010) Defective in cuticular ridges of Arabidopsis thaliana, a gene associated with surface cutin formation, encodes a soluble diacylglycerol acyltransferase. Journal of Biological Chemistry. In press.
    Höfer R, Briesen I, Beck M et al. (2008) The Arabidopsis cytochrome P450 CYP86A1 encodes a fatty acid omega-hydroxylase involved in suberin monomer biosynthesis. Journal of Experimental Botany 59: 2347–2360.
    Hoffmann-Benning S and Kende H (1994) Cuticle biosynthesis in rapidly growing internodes of deepwater rice. Plant Physiology 104: 719–723.
    book Jeffree CE (2006) "The fine structure of the plant cuticle". In: Riederer M and Müller C (eds) Biology of the Plant Cuticle, pp. 11–125. Oxford, UK: Blackwell Publishing Ltd.
    Kannangara R, Branigan C, Liu Y et al. (2007) The transcription factor WIN1/SHN1 regulates cutin biosynthesis in Arabidopsis thaliana. Plant Cell 19: 1278–1294.
    Kolattukudy PE (1981) Structure, biosynthesis and biodegradation of cutin and suberin. Annual Review of Plant Physiology 32: 539–567.
    Kolattukudy PE (2001) Polyesters in higher plants. Advances in Biochemical Engineering/Biotechnology 71: 1–49.
    Kurdyukov S, Faust A, Trenkamp S et al. (2006a) Genetic and biochemical evidence for involvement of HOTHEAD in the biosynthesis of long-chain -,-dicarboxylic fatty acids and formation of extracellular matrix. Planta 224: 315–329.
    Kurdyukov S, Faust A, Nawrath C et al. (2006b) The epidermis-specific extracellular BODYGUARD controls cuticle development and morphogenesis in Arabidopsis. Plant Cell 18: 321–339.
    Lee SB, Jung SJ, Go YS et al. (2009) Two Arabidopsis 3-ketoacyl CoA synthase genes, KCS20 and KCS2/DAISY, are functionally redundant in cuticular wax and root suberin biosynthesis, but differentially controlled by osmotic stress. Plant Journal 60: 462–475.
    Li Y and Beisson F (2009) The biosynthesis of cutin and suberin as an alternative source of enzymes for the production of bio-based chemicals and materials. Biochimie 91: 685–691.
    Li Y, Beisson F, Koo AJ et al. (2007) Identification of acyltransferases required for cutin biosynthesis and production of cutin with suberin-like monomers. Proceedings of the National Academy of Sciences of the USA 104: 18339–18344.
    Li-Beisson Y, Pollard M, Sauveplane V et al. (2009) Nanoridges that characterize the surface morphology of flowers require the synthesis of cutin polyester. Proceedings of the National Academy of Sciences of the USA 106: 22008–22013.
    book Li-Beisson Y, Shorrosh B, Beisson F et al. (2010) "Acyl lipid metabolism". In: Last R (ed.) The Arabidopsis Book. Rockville, MD: American Society of Plant Biologists.
    Lü S, Song T, Kosma DK et al. (2009) Arabidopsis CER8 encodes LONG-CHAIN ACYL-COA SYNTHETASE 1 (LACS1) that has overlapping functions with LACS2 in plant wax and cutin synthesis. Plant Journal 59: 553–564.
    Matas AJ, Agustí J, Tadeo FR, Talón M and Rose JK (2010) Tissue-specific transcriptome profiling of the citrus fruit epidermis and subepidermis using laser capture microdissection. Journal of Experimental Botany 61: 3321–3330.
    Moire L, Schmutz A, Buchala A et al. (1999) Glycerol is a suberin monomer. New experimental evidence for an old hypothesis. Plant Physiology 119: 1137–1146.
    Molina I, Bonaventure G, Ohlrogge J and Pollard M (2006) The lipid polyester composition of Arabidopsis thaliana and Brassica napus seeds. Phytochemistry 67: 2597–2610.
    Molina I, Li-Beisson Y, Beisson F, Ohlrogge JB and Pollard M (2009) Identification of an Arabidopsis feruloyl-coenzyme A transferase required for suberin synthesis. Plant Physiology 151: 1317–1328.
    Nawrath C (2006) Unraveling the complex network of cuticular structure and function. Current Opinion Plant Biology 9: 281–287.
    Panikashvili D, Savaldi-Goldstein S, Mandel T et al. (2007) The Arabidopsis DESPERADO/AtWBC11 transporter is required for cutin and wax secretion. Plant Physiology 145: 1345–1360.
    Panikashvili D, Shi JX, Schreiber L and Aharoni A (2009) The Arabidopsis DCR encoding a soluble BAHD acyltransferase is required for cutin polyester formation and seed hydration properties. Plant Physiology 151: 1773–1789.
    Pollard M, Beisson F, Li Y and Ohlrogge JB (2008) Building lipid barriers: biosynthesis of cutin and suberin. Trends in Plant Science 13: 236–246.
    Samuels L, Kunst L and Jetter R (2008) Sealing plant surfaces: cuticular wax formation by epidermal cells. Annual Review of Plant Biology 59: 683–707.
    Schnurr J, Shockey J and Browse J (2004) The acyl-CoA synthetase encoded by LACS2 is essential for normal cuticle development in Arabidopsis. Plant Cell 16: 629–642.
    Soler M, Serra O, Molinas M et al. (2007) A genomic approach to suberin biosynthesis and cork differentiation. Plant Physiology 144: 419–431.
    Suh MC, Samuels AL, Jetter R et al. (2005) Cuticular lipid composition, surface structure, and gene expression in Arabidopsis stem epidermis. Plant Physiology 139: 1649–1665.
    Villena JF, Domínguez E, Stewart D and Heredia A (1999) Characterization and biosynthesis of non-degradable polymers in plant cuticles. Planta 208: 181–187.
    Wellesen K, Durst F, Pinot F et al. (2001) Functional analysis of the LACERATA gene of Arabidopsis provides evidence for different roles of fatty acid omega -hydroxylation in development. Proceedings of the National Academy of Sciences of the USA 98: 9694–9699.
    Xiao F, Goodwin SM, Xiao Y et al. (2004) Arabidopsis CYP86A2 represses Pseudomonas syringae type III genes and is required for cuticle development. EMBO Journal 23: 2903–2913.
    Yang W, Pollard M, Li-Beisson Y et al. (2010) A distinct type of glycerol-3-phosphate acyltransferase with sn-2 preference and phosphatase activity producing 2-monoacylglycerol. Proceedings of the National Academy of Sciences of the USA 107: 12040–12045.
    Yeats TH, Howe KJ, Matas AJ et al. (2010) Mining the surface proteome of tomato (Solanum lycopersicum) fruit for proteins associated with cuticle biogenesis. Journal Experimental Botany 61: 3759–3771.
    Zheng Z, Xia Q, Dauk M et al. (2003) Arabidopsis AtGPAT1, a member of the membrane-bound glycerol-3-phosphate acyltransferase gene family, is essential for tapetum differentiation and male fertility. Plant Cell 15: 1872–1887.
    Zlotnik-Mazori T and Stark RE (1988) Nuclear magnetic resonance studies of cutin, an insoluble plant polyester. Macromolecules 21: 2412–2417.
 Further Reading
    book Kolattukudy PE (2001) "Polyesters in higher plants". In: Babel W and Steinbuschel A (eds) Advances in Biochemical Engineering/Biotechnology, vol. 71, Biopolyesters, pp. 1–49. Springer: Berlin.
    Kunst L and Samuels L (2009) Plant cuticles shine: advances in wax biosynthesis and export. Current Opinion in Plant Biology 12: 721–727.
    Schreiber L (2010) Transport barriers made of cutin, suberin and associated waxes. Trends in Plant Science 15: 546–553.
    book Stark RE and Tian S (2006) "The cutin biopolymer matrix". In: Riederer M (ed.) Biology of the Plant Cuticle, pp. 126–144. Oxford, UK: Blackwell Publishing Co.
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Li‐Beisson, Yonghua(Mar 2011) Cutin and Suberin. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1002/9780470015902.a0001920.pub2]