Cutin and Suberin Polyesters

Abstract

Cutin and suberin are cell wall‐associated glycerolipid polymers that are specific to plants. Cutin forms the framework of the cuticle sealing the aerial epidermis, while suberin is present in the periderm of barks and underground organs. Suberised walls are also found in the root endodermis. Barriers based on cutin and suberin restrict the transport of water and solutes across cell walls and limit pathogen invasions. Chemical analysis shows that both polymers are polyesters composed mostly of fatty hydroxyacids, diacids and epoxyacids esterified to each other and to glycerol. Suberin, whose best‐known form is cork, usually differs from cutin (which has C16 and C18 fatty acids) by a higher content of C20–C24 aliphatics and aromatics. In the last 10 years, the identification of mutants of Arabidopsis or other model plants affected in cutin and/or suberin content has allowed the construction of a more complete picture of the polyester biosynthesis pathways, which currently include acyltransferases with unique specificities, fatty acid hydroxylases, acyl‐CoA synthetases, fatty acid elongases, fatty acyl‐CoA reductases, feruloyl transferases, ABC transporters and extracellular transacylases.

Key Concepts

  • The epidermal cells of plant aerial organs and periderm/endoderm cells synthesise the protective cell wall lipid polymers cutin and suberin respectively.
  • Cutin and suberin are both polyesters containing glycerol and oxygenated fatty acids.
  • Cutin structure is not completely understood and suberin structure remains controversial.
  • Oxygenated fatty acid monomers are produced by fatty acid oxidases of the cytochrome P450 superfamily.
  • Acylation of oxygenated fatty acids to glycerol is catalysed by special glycerol‐3‐phosphate acyltransferases.
  • Cutin acylglycerol building blocks are exported to the cell wall and polymerised by extracellular transacylases.
  • How suberin precursors are assembled is still unknown.

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

Figure 1. Localisation and ultrastructure of cutin and suberin layers. Top panel: Schematic representation of the cuticle (left) and suberised cell wall (right). Bottom panel: Observation of cutin and suberin using electron microscope. (a) Transmission electron microscopy (TEM) image of a cross section view of Arabidopsis stem epidermis. Scale bar: 500 nm. (b) A scanning electron microscopy (SEM) image of the epidermal surface of an Arabidopsis sepal. Scale bar: 5 μm. (c) A TEM image of a cross section view of Arabidopsis roots. Scale bar: 100 nm. Adapted from Molina et al., (2009) © American Society of Plant Biologists. Abbreviations: CW, cell wall; PC, peridermal cell.
Figure 2. Structure of the most common monomers of cutin and the major reactions/enzymes involved in their syntheses. In red: enzymes identified; 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). DH, dehydrogenase.
Figure 3. A possible structure of a polyester domain. Note: R: other fatty acid or glycerol molecules. Fatty acids and linkages represented are typical of cutins rich in hydroxyacids.
Figure 4. Major biochemical steps identified in cutin/suberin biosynthetic pathways. Abbreviations: ABC transporter, ATP binding cassette transporter; CW, cell wall; ER, endoplasmic reticulum; GPAT, glycerol‐3‐phosphate acyltransferase (with phosphatase activity for cutin‐related members); KCS, ketoacyl‐CoA synthetase; LACS, long chain acyl‐CoA synthetase; PM, plasma membrane.
Figure 5. The regiospecificity of glycerol‐3‐phosphate acyltransferases (GPATs) controls the flux of acyl chains to their final site of deposition. Numbering of sn2‐GPATs refers to Arabidopsis GPAT family. Abbreviations: G3P, glycerol‐3‐phosphate; MAG, monoacylglycerol; PA, phosphatidic acid; CW, cell wall; PM, plasma membrane.
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Further Reading

Andersen TG, Barberon M and Geldner N (2015) Suberization‐the second life of an endodermal cell. Current Opinion in Plant Biology 28: 9–15.

Beisson F, Li‐Beisson Y and Pollard M (2012) Solving the puzzles of cutin and suberin polymer biosynthesis. Current Opinion in Plant Biology 15: 329–337.

Graça J (2015) Suberin: the biopolyester at the frontier of plants. Frontiers in Chemistry 3: 62.

Nawrath C, Schreiber L, Franke RB, et al. (2013) Apoplastic diffusion barriers in Arabidopsis. Arabidopsis Book 11: e0167.

Pollard M, Beisson F, Li Y and Ohlrogge J (2008) Building lipid barriers: biosynthesis of cutin and suberin. Trends in Plant Sciences 13: 236–246.

Vishwanath SJ, Delude C, Domergue F and Rowland O (2015) Suberin: biosynthesis, regulation, and polymer assembly of a protective extracellular barrier. Plant Cell Reports 34: 573–586.

Yeats TH and Rose JK (2013) The formation and function of plant cuticles. Plant Physiology 163: 5–20.

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Li‐Beisson, Yonghua, Verdier, Gaëtan, Xu, Lin, and Beisson, Fred(May 2016) Cutin and Suberin Polyesters. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1002/9780470015902.a0001920.pub3]