Cell Wall Biosynthesis


Plant cell walls encase the plant cells and provide many structural and functional roles. Cell walls are complex structures consisting mainly of polysaccharides (cellulose, hemicellulose and pectin), glycoproteins and phenolic compounds (lignin). Most plant cell wall polymers are synthesised from ‘activated’ precursors by the action of transferases. Cellulose is synthesised at the plasma membrane (PM) by a large complex moving in the plane of PM and extruding cellulose chains into the wall. The hemicelluloses are synthesised in the endomembrane system by resident glycosyl transferase enzymes and transported to the wall. These individual components are then cross‐linked and assembled into the complex matrix structure that comprises functional cell walls. Woody cell walls also contain lignin. Lignin is synthesised in the wall by oxidative coupling of individual monomeric subunits.

Key Concepts

  • All plant cells are encased in a multilayered cell wall.
  • Plant cell walls are an important renewable resource.
  • Plant cell walls are composed of polysaccharides (cellulose, hemicellulose and pectin), cell wall proteins and phenolic compounds.
  • Most of the polysaccharides are synthesised from activated precursors, NDP sugars and incorporated into the growing polymer catalysed by the action of glycosyl transferases (GTs).
  • Cellulose is synthesised by the cellulose synthase complex (CSC) moving in the plane of plasma membrane, extruding cellulose chains into the cell wall.
  • Hemi‐celluloses and pectin are synthesised and modified in the endomembrane system before being deposited into the apoplast for assembly into the cell wall.
  • Lignin monomers are synthesised in the cytoplasm and transported to the wall.
  • Laccases have an essential role in lignin formation by catalysing the oxidative coupling of the lignin monomers in the apoplast.
  • Cell wall composition varies greatly between cell types and can change as the cell differentiates and with the developmental stages of cells and the plant.
  • The cell wall has an important function in regulating how plant cells achieve their final size and shape and consequently have an essential role in regulating plant growth.

Keywords: plant cell walls; primary cell wall; secondary cell wall; polysaccharides; cellulose; hemicellulose; pectin; lignin; glycosyl transferases; cell wall polymer assembly

Figure 1. Major pathways for the synthesis and interconversion of the NDP sugars used in the biosynthesis of plant cell wall polysaccharides. Some indication of the relative flux through the various pathways is given by arrow thickness. In addition, solid arrows indicate primary pathways for de novo synthesis of NDP sugars and inositol. Pecked arrows (‐‐‐‐‐‐) imply great variation between tissues. Dot‐dashed arrows (–⋅–⋅–⋅) indicate scavenger pathways involved in recycling monosaccharides, for example, released by polysaccharide turnover. The ‘box’ is the pool of hexose monophosphates mentioned in the text. Reactions marked * are isomerisations, with no other reactants. Numbered enzymes are 1, phosphoglucomutase; 2, glucose 6‐phosphate isomerase; 3, mannose 6‐phosphate isomerase; 4, phosphomannomutase; 5, UDP‐glucose pyrophosphorylase; 6, GDP‐mannose pyrophosphorylase; 7, myo‐inositol 1‐phosphate synthase; 8, myo‐inositol 1‐phosphatase; 9, myo‐inositol oxygenase; 10, glucuronokinase; 11, UDP‐glucuronate pyrophosphorylase; 12, UDP‐glucose dehydrogenase; 13, UDP‐glucuronate decarboxylase; 14, UDP‐glucose 4‐epimerase; 15, UDP‐glucuronate 4‐epimerase; 16, UDP‐xylose 4‐epimerase; 17, ‘GDP‐fucose synthase’ (three individual activities: (a) GDP‐d‐mannose 4,6‐dehydratase, (b) GDP‐4‐keto‐6‐deoxy‐d‐mannose 3,5‐epimerase, and (c) GDP‐4‐keto‐l‐fucose 4‐reductase); 18, GDP‐mannose 3,5‐epimerase; 19, ‘UDP‐rhamnose synthase’ (probably three activities, cf. 17); 20, UDP‐apiose synthase; 21, d‐galactokinase; 22, galacturonokinase; 23, arabinokinase; 24, fucokinase; 25, hexokinase or glucokinase; 26, fructokinase; 27, mannokinase; 28, UDP‐d‐galactose pyrophosphorylase; 29, UDP‐galacturonate pyrophosphorylase; 30, UDP‐arabinose pyrophosphorylase; 31, GDP‐fucose pyrophosphorylase.
Figure 2. (a–j) Polysaccharide structure. Individual monosaccharide units are depicted with their respective notations as described by Consortium of Functional Glycomics (CFG) (Berger et al., ). Monosaccharide units are joined by a solid line indicating the linkage with the type of linkage depicted above or below the joining line. The enzymes involved in formation of linkages at the numbered steps (1–22) are expanded upon in Table.


Alejandro S, Lee Y, Tohge T, et al. (2012) AtABCG29 Is a monolignol transporter involved in lignin biosynthesis. Current Biology 22: 1207–1212.

Anders N, Wilkinson MD, Lovegrove A, et al. (2012) Glycosyl transferases in family 61 mediate arabinofuranosyl transfer onto xylan in grasses. Proceedings of the National Academy of Sciences of the United States of America 109: 989–993.

Arioli T, Peng LC, Betzner AS, et al. (1998) Molecular analysis of cellulose biosynthesis in Arabidopsis. Science 279: 717–720.

Atmodjo MA, Sakuragi Y, ZHU X, et al. (2011) Galacturonosyltransferase (GAUT)1 and GAUT7 are the core of a plant cell wall pectin biosynthetic homogalacturonan: galacturonosyltransferase complex. Proceedings of the National Academy of Sciences of the United States of America 108: 20225–20230.

Berger, O, Mcbride, R, Razi, N and Paulson, J (2008) Symbol Notation Extension for Pathogen Polysaccharides. Consortium for Functional Glycomics. The Scripps Research Institute.

Boerjan W, Ralph J and Baucher M (2003a) Lignin biosynthesis. Annual Review of Plant Biology 54: 519–546.

Bouton S, Leboeuf E, Mouille G, et al. (2002) Quasimodo1 encodes a putative membrane‐bound glycosyltransferase required for normal pectin synthesis and cell adhesion in Arabidopsis. Plant Cell 14: 2577–2590.

Brown D, Wightman R, Zhang ZN, et al. (2011) Arabidopsis genes IRREGULAR XYLEM (IRX15) and IRX15L encode DUF579‐containing proteins that are essential for normal xylan deposition in the secondary cell wall. Plant Journal 66: 401–413.

Brown DM, Goubet F, Vicky WWA, et al. (2007) Comparison of five xylan synthesis mutants reveals new insight into the mechanisms of xylan synthesis. Plant Journal 52: 1154–1168.

Brown DM, Zhang ZN, Stephens E, Dupree P and Turner SR (2009) Characterization of IRX10 and IRX10‐like reveals an essential role in glucuronoxylan biosynthesis in Arabidopsis. Plant Journal 57: 732–746.

Chiniquy D, Sharma V, Schultink A, et al. (2012) XAX1 from glycosyltransferase family 61 mediates xylosyltransfer to rice xylan. Proceedings of the National Academy of Sciences of the United States of America 109: 17117–17122.

Davin LB, Wang HB, Crowell AL, et al. (1997) Stereoselective bimolecular phenoxy radical coupling by an auxiliary (dirigent) protein without an active center. Science 275: 362–366.

Dhugga KS, Barreiro R, Whitten B, et al. (2004) Guar seed beta‐mannan synthase is a member of the cellulose synthase super gene family. Science 303: 363–366.

Egelund J, Petersen BL, Motawia MS, et al. (2006) Arabidopsis thaliana RGXT1 and RGXT2 encode Golgi‐localized (1,3)‐alpha‐D‐xylosyltransferases involved in the synthesis of pectic rhamnogalacturonan‐II. Plant Cell 18: 2593–2607.

Gille S, De Souza A, Xiong GY, et al. (2011) O‐Acetylation of arabidopsis hemicellulose xyloglucan requires AXY4 or AXY4L, proteins with a TBL and DUF231 domain. Plant Cell 23: 4041–4053.

Harholt J, Jensen JK, Sorensen SO, et al. (2006) ARABINAN DEFICIENT 1 is a putative arabinosyltransferase involved in biosynthesis of Pectic Arabinan in Arabidopsis. Plant Physiology 140: 49–58.

Jensen JK, Johnson NR and Wilkerson CG (2014) Arabidopsis thaliana IRX10 and two related proteins from psyllium and Physcomitrella patens are xylan xylosyltransferases. Plant Journal 80: 207–215.

Jensen JK, Kim H, Cocuron JC, et al. (2011) The DUF579 domain containing proteins IRX15 and IRX15‐L affect xylan synthesis in Arabidopsis. Plant Journal 66: 387–400.

Kumar M and Turner S (2015) Plant cellulose synthesis: CESA proteins crossing kingdoms. Phytochemistry 112: 91–99.

Lee C, Teng Q, Zhong RQ, et al. (2012) Three Arabidopsis DUF579 domain‐containing GXM proteins are methyltransferases catalyzing 4‐O‐methylation of glucuronic acid on xylan. Plant and Cell Physiology 53: 1934–1949.

Liu C‐J, Miao Y‐C and Zhang K‐W (2011) Sequestration and transport of lignin monomeric precursors. Molecules 16: 710–727.

Liu S (2010) Woody biomass: niche position as a source of sustainable renewable chemicals and energy and kinetics of hot‐water extraction/hydrolysis. Biotechnology Advances 28: 563–582.

Liwanag AJM, Ebert B, Verhertbruggen Y, et al. (2012) Pectin biosynthesis: GALS1 in Arabidopsis thaliana Is a beta‐1,4‐galactan beta‐1,4‐galactosyltransferase. Plant Cell 24: 5024–5036.

Manabe Y, Nafisi M, Verhertbruggen Y, et al. (2011) Loss‐of‐function mutation of reduced wall acetylation2 in Arabidopsis leads to reduced cell wall acetylation and increased resistance to Botrytis cinerea. Plant Physiology 155: 1068–1078.

Manabe Y, Verhertbruggen Y, Gille S, et al. (2013) Reduced wall acetylation proteins play vital and distinct roles in cell wall O‐acetylation in Arabidopsis. Plant Physiology 163: 1107–1117.

Miao Y‐C and Liu C‐J (2010) ATP‐binding cassette‐like transporters are involved in the transport of lignin precursors across plasma and vacuolar membranes. Proceedings of the National Academy of Sciences of the United States of America 107: 22728–22733.

Mortimer JC, Miles GP, Brown DM, et al. (2010) Absence of branches from xylan in Arabidopsis gux mutants reveals potential for simplification of lignocellulosic biomass. Proceedings of the National Academy of Sciences of the United States of America 107: 17409–17414.

Paredez AR, Somerville CR and Ehrhardt DW (2006) Visualization of cellulose synthase demonstrates functional association with microtubules. Science 312: 1491–1495.

Pauly M, Gille S, Liu LF, et al. (2013) Hemicellulose biosynthesis. Planta 238: 627–642.

Perrin RM, Derocher AE, Bar‐Peled M, et al. (1999) Xyloglucan fucosyltransferase, an enzyme involved in plant cell wall biosynthesis. Science 284: 1976–1979.

Persson S, Paredez A, Carroll A, et al. (2007) Genetic evidence for three unique components in primary cell‐wall cellulose synthase complexes in Arabidopsis. Proceedings of the National Academy of Sciences of the United States of America 104: 15566–15571.

Roland JC, Vian B and Reis D (1977) Further observations on cell‐wall morphogenesis and polysaccharide arrangement during plant‐growth. Protoplasma 91: 125–141.

Scheller HV and Ulvskov P (2010) Hemicelluloses. Annual Review of Plant Biology 61: 263–289.

Schuetz M, Benske A, Smith RA, et al. (2014) Laccases direct lignification in the discrete secondary cell wall domains of protoxylem. Plant Physiology 166: 798–807.

Sibout R and Hoefte H (2012) Plant cell biology: the ABC of monolignol transport. Current Biology 22: R533–R535.

Sterling JD, Atmodjo MA, Inwood SE, et al. (2006) Functional identification of an Arabidopsis pectin biosynthetic homogalacturonan galacturonosyltransferase. Proceedings of the National Academy of Sciences of the United States of America 103: 5236–5241.

Taylor NG, Howells RM, Huttly AK, Vickers K and Turner SR (2003) Interactions among three distinct CesA proteins essential for cellulose synthesis. Proceedings of the National Academy of Sciences of the United States of America 100: 1450–1455.

Thompson JE, Smith RC and Fry SC (1997) Xyloglucan undergoes interpolymeric transglycosylation during binding to the plant cell wall in vivo: evidence from C‐13/H‐3 dual labelling and isopycnic centrifugation in caesium trifluoroacetate. Biochemical Journal 327: 699–708.

Urbanowicz BR, Pena MJ, Moniz HA, Moremen KW and York WS (2014) Two Arabidopsis proteins synthesize acetylated xylan in vitro. Plant Journal 80: 197–206.

Urbanowicz BR, Pena MJ, Ratnaparkhe S, et al. (2012) 4‐O‐methylation of glucuronic acid in Arabidopsis glucuronoxylan is catalyzed by a domain of unknown function family 579 protein. Proceedings of the National Academy of Sciences of the United States of America 109: 14253–14258.

Vain T, Crowell EF, Timpano H, et al. (2014) The cellulase KORRIGAN is part of the cellulose synthase complex. Plant Physiology 165: 1521–1532.

Vanholme R, Cesarino I, Rataj K, et al. (2013) Caffeoyl shikimate esterase (CSE) is an enzyme in the lignin biosynthetic pathway in Arabidopsis. Science 341: 1103–1106.

Vanholme R, Demedts B, Morreel K, Ralph J and Boerjan W (2010a) Lignin biosynthesis and structure. Plant Physiology 153: 895–905.

Wang Y, Chantreau M, Sibout R and Hawkins S (2013) Plant cell wall lignification and monolignol metabolism. Frontiers in Plant Science 4.

Wu AM, Hornblad E, Voxeur A, et al. (2010) Analysis of the Arabidopsis IRX9/IRX9‐L and IRX14/IRX14‐L pairs of glycosyltransferase genes reveals critical contributions to biosynthesis of the hemicellulose glucuronoxylan. Plant Physiology 153: 542–554.

Wu AM, Rihouey C, Seveno M, et al. (2009) The Arabidopsis IRX10 and IRX10‐LIKE glycosyltransferases are critical for glucuronoxylan biosynthesis during secondary cell wall formation. Plant Journal 57: 718–731.

Further Reading

Albersheim P, Darvill A, Roberts K, Sederoff R and Staehelin A (eds) (2011) Plant Cell Walls: From Chemistry to Biology. New York: Garland Science, Taylor & Francis.

Atmodjo MA, Hao Z and Mohnen D (2013) Evolving views of pectin biosynthesis. Annual Review of Plant Biology 64: 747–779.

Boerjan W, Ralph J and Baucher M (2003b) Lignin biosynthesis. Annual Review of Plant Biology 54: 519–546.

Cosgrove DJ (1997) Assembly and enlargement of the primary cell wall in plants. Annual Review of Cell and Developmental Biology 13: 171–201.

Delmer DP (1999) Cellulose biosynthesis: exciting times for a difficult field of study. Annual Review of Plant Physiology and Plant Molecular Biology 50: 245–276.

Hao ZY and Mohnen D (2014) A review of xylan and lignin biosynthesis: foundation for studying Arabidopsis irregular xylem mutants with pleiotropic phenotypes. Critical Reviews in Biochemistry and Molecular Biology 49: 212–241.

Jamet E, Canut H, Boudart G and Pont‐Lezica RF (2006) Cell wall proteins: a new insight through proteomics. Trends in Plant Science 11: 33–39.

Lerouxel O, Cavalier DM, Liepman AH and Keegstra K (2006) Biosynthesis of plant cell wall polysaccharides – a complex process. Current Opinion in Plant Biology 9: 621–630.

Liepman AH, Wightman R, Geshi N, Turner SR and Scheller HV (2010) Arabidopsis – a powerful model system for plant cell wall research. Plant Journal 61: 1107–1121.

Mohnen D (2008) Pectin structure and biosynthesis. Current Opinion in Plant Biology 11: 266–277.

Scheible WR and Pauly M (2004) Glycosyltransferases and cell wall biosynthesis: novel players and insights. Current Opinion in Plant Biology 7: 285–295.

Somerville C (2006) Cellulose synthesis in higher plants. Annual Review of Cell and Developmental Biology 22: 53–78.

Vanholme R, Demedts B, Morreel K, Ralph J and Boerjan W (2010b) Lignin biosynthesis and structure. Plant Physiology 153: 895–905.

York WS and O'Neill MA (2008) Biochemical control of xylan biosynthesis – which end is up? Current Opinion in Plant Biology 11: 258–265.

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Kumar, Manoj, and Turner, Simon(May 2015) Cell Wall Biosynthesis. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1002/9780470015902.a0001683.pub2]