Expanding roles for pectins in plant development
|
摘要
Pectins are complex cell wall polysaccharides important for many aspects of plant development.Recent studies have discovered extensive physical interactions between pectins and other cell wall components,implicating pectins in new molecular functions.Pectins are often localized in spatially-restricted patterns,and some of these non-uniform pectin distributions contribute to multiple aspects of plant development,including the morphogenesis of cells and organs.Furthermore,a growing number of mutants affecting cell wall composition have begun to reveal the distinct contributions of different pectins to plant development.This review discusses the interactions of pectins with other cell wall components,the functions of pectins in controlling cellular morphology,and how non-uniform pectin composition can be an important determinant of developmental processes.
Pectins are complex cell wall polysaccharides important for many aspects of plant development.Recent studies have discovered extensive physical interactions between pectins and other cell wall components,implicating pectins in new molecular functions.Pectins are often localized in spatially-restricted patterns,and some of these non-uniform pectin distributions contribute to multiple aspects of plant development,including the morphogenesis of cells and organs.Furthermore,a growing number of mutants affecting cell wall composition have begun to reveal the distinct contributions of different pectins to plant development.This review discusses the interactions of pectins with other cell wall components,the functions of pectins in controlling cellular morphology,and how non-uniform pectin composition can be an important determinant of developmental processes.
Pectins are complex cell wall polysaccharides important for many aspects of plant development.Recent studies have discovered extensive physical interactions between pectins and other cell wall components,implicating pectins in new molecular functions.Pectins are often localized in spatially-restricted patterns,and some of these non-uniform pectin distributions contribute to multiple aspects of plant development,including the morphogenesis of cells and organs.Furthermore,a growing number of mutants affecting cell wall composition have begun to reveal the distinct contributions of different pectins to plant development.This review discusses the interactions of pectins with other cell wall components,the functions of pectins in controlling cellular morphology,and how non-uniform pectin composition can be an important determinant of developmental processes.
引文
Abasolo W,Eder M,Yamauchi K,Obel N,Reinecke A,Neumetzler L,Dunlop JWC,Mouille G,Pauly M,H€ofte H,Burgert I(2009)Pectin may hinder the unfolding of xyloglucan chains during cell deformation:Implications of the mechanical performance of Arabidopsis hypocotyls with pectin alterations.Mol Plant 2:990-999
Amsbury S,Hunt L,Elhaddad N,Baillie A,Lundgren M,Verhertbruggen Y,Scheller HV,Knox JP,Fleming AJ,Gray JE(2016)Stomatal function requires pectin demethyl-esterification of the guard cell wall.Curr Biol 26:2899-2906
Anderson CT,Wallace IS,Somerville CR(2012)Metabolic click-labeling with a fucose analog reveals pectin delivery,architecture,and dynamics in Arabidopsis cell walls.Proc Natl Acad Sci USA 109:1329-1334
Atmodjo MA,Hao Z,Mohnen D(2013)Evolving views of pectin biosynthesis.Annu Rev Plant Biol 64:747-779
Baron-Epel O,Gharyal PK,Schindler M(1988)Pectins as mediators of wall porosity in soybean cells.Planta 175:389-395
Bootten TJ,Harris PJ,Melton LD,Newman RH(2004)Solidstate13C-NMR spectroscopy shows that the xyloglucans in the primary cell walls of mung bean(Vigna radiata L.)occur in different domains:A new model for xyloglucancellulose interactions in the cell wall.J Exp Bot 55:571-583
Bouton S,Leboeuf E,Mouille G,Leydecker MT,Talbotec J,Granier F,Lahaye M,Hofte H,Truong HN(2002)QUASIMODO1 encodes a putative membrane-bound glycosyltransferase required for normal pectin synthesis and cell adhesion in Arabidopsis.Plant Cell 14:2577-2590
Braybrook SA,Peaucelle A(2013)Mechano-chemical aspects of organ formation in Arabidopsis thaliana:The relationship between auxin and pectin.PLoS ONE 8:e57813
Buffetto F,Cornuault V,Rydahl MG,Ropartz D,Alvarado C,Echasserieau V,Le Gall S,Bouchet B,Tranquet O,Verhertbruggen Y,Willats WGT,Knox JP,Ralet MC,Guillon F(2015)The deconstruction of pectic rhamnogalacturonan I unmasks the occurrence of a novel arabinogalactan oligosaccharide epitope.Plant Cell Physiol 56:2181-2196
Bush MS,Marry M,Huxham MI,Jarvis MC,McCann MC(2001)Developmental regulation of pectic epitopes during potato tuberisation.Planta 213:869-880
Carpita NC,Gibeaut DM(1993)Structural models of primary cell walls in flowering plants:Consistency of molecular structure with the physical properties of the walls during growth.Plant J 3:1-30
Carter R,Woolfenden H,Baillie A,Amsbury S,Carroll S,Healicon E,Sovatzoglou S,Braybrook S,Gray JE,Hobbs J,Morris RJ,Fleming AJ(2017)Stomatal opening involves polar,not radial,stiffening of guard cells.Curr Biol 27:2974-2983.e2
Cavalier DM,Lerouxel O,Neumetzler L,Yamauchi K,Reinecke A,Freshour G,Zabotina OA,Hahn MG,Burgert I,Pauly M,Raikhel NV,Keegstra K(2008)Disrupting two Arabidopsis thaliana xylosyltransferase genes results in plants deficient in xyloglucan,a major primary cell wall component.Plant Cell 20:1519-1537
Coenen GJ,Bakx EJ,Verhoef RP,Schols HA,Voragen AGJ(2007)Identification of the connecting linkage between homo-or xylogalacturonan and rhamnogalacturonan type I.Carbohydr Polym 70:224-235
Cornuault V,Buffetto F,Rydahl MG,Marcus SE,Torode TA,Xue J,Crepeau MJ,Faria-Blanc N,Willats WGT,Dupree P,Ralet MC,Knox JP(2015)Monoclonal antibodies indicate low-abundance links between heteroxylan and other glycans of plant cell walls.Planta 242:1321-1334
Daher FB,Braybrook SA(2015)How to let go:Pectin and plant cell adhesion.Front Plant Sci 6:523
de Souza A,Hull PA,Gille S,Pauly M(2014)Identification and functional characterization of the distinct plant pectin esterases PAE8 and PAE9 and their deletion mutants.Planta 240:1123-1138
Derbyshire P,McCann MC,Roberts K(2007)Restricted cell elongation in Arabidopsis hypocotyls is associated with a reduced average pectin esterification level.BMC Plant Biol7:31
Dick-Perez M,Wang T,Salazar A,Zabotina OA,Hong M(2012)Multidimensional solid-state NMR studies of the structure and dynamics of pectic polysaccharides in uniformly13C-labeled Arabidopsis primary cell walls.Magn Reson Chem50:539-550
Dick-Perez M,Zhang Y,Hayes J,Salazar A,Zabotina OA,Hong M(2011)Structure and interactions of plant cell-wall polysaccharides by two-and three-dimensional magic-anglespinning solid-state NMR.Biochemistry 50:989-1000
Dumont M,Lehner A,Bouton S,Kiefer-Meyer MC,Voxeur A,Pelloux J,Lerouge P,Mollet JC(2014)The cell wall pectic polymer rhamnogalacturonan-II is required for proper pollen tube elongation:Implications of a putative sialyltransferase-like protein.Ann Bot 114:1177-1188
Ermel FF,Follet-Gueye ML,Cibert C,Vian B,Morvan C,Catesson AM,Goldberg R(2000)Differential localization of arabinan and galactan side chains of rhamnogalacturonan 1 in cambial derivatives.Planta 210:732-740
Goldberg R,Morvan C,Roland JC(1986)Composition,properties and localisation of pectins in young and mature cells of the mung bean hypocotyl.Plant Cell Physiol 27:417-429
Gou JY,Miller LM,Hou G,Yu XH,Chen XY,Liu CJ(2012)Acetylesterase-mediated deacetylation of pectin impairs cell elongation,pollen germination,and plant reproduction.Plant Cell 24:50-65
Grant GT,Morris ER,Rees DA,Smith PJC,Thom D(1973)Biological interactions between polysaccharides and divalent cations:The egg-box model.FEBS Lett 32:195-198
Guillemin F,Guillon F,Bonnin E,Devaux MF,Chevalier T,Knox JP,Liners F,Thibault JF(2005)Distribution of pectic epitopes in cell walls of the sugar beet root.Planta 222:355-371
Hall HC,Cheung J,Ellis BE(2013)Immunoprofiling reveals unique cell-specific patterns of wall epitopes in the expanding Arabidopsis stem.Plant J 74:134-147
Harholt J(2005)ARABINAN DEFICIENT 1 is a putative arabinosyltransferase involved in biosynthesis of pectic arabinan in Arabidopsis.Plant Physiol 140:49-58
Hongo S,Sato K,Yokoyama R,Nishitani K(2012)Demethylesterification of the primary wall by PECTIN METHYL-ESTERASE35 provides mechanical support to the Arabidopsis stem.Plant Cell 24:2624-2634
Ishii T(1997)O-Acetylated oligosaccharides from pectins of potato tuber cell walls.Plant Physiol 113:1265-1272
Iwai H,Masaoka N,Ishii T,Satoh S(2002)A pectin glucuronyltransferase gene is essential for intercellular attachment in the plant meristem.Proc Natl Acad Sci USA99:16319-16324
Jensen JK,Sorensen SO,Harholt J,Geshi N,Sakuragi Y,Moller I,Zandleven J,Bernal AJ,Jensen NB,Sorensen C,Pauly M,Beldman G,Willats WGT,Scheller HV(2008)Identification of a xylogalacturonan xylosyltransferase involved in pectin biosynthesis in Arabidopsis.Plant Cell 20:1289-1302
Jones L,Milne JL,Ashford D,McQueen-Mason SJ(2003)Cell wall arabinan is essential for guard cell function.Proc Natl Acad Sci USA 100:11783-11788
Kim S-J,Held MA,Zemelis S,Wilkerson C,Brandizzi F(2015)CGR2 and CGR3 have critical overlapping roles in pectin methylesterification and plant growth in Arabidopsis thaliana.Plant J 82:208-220
Knox JP,Linstead PJ,King J,Cooper C,Roberts K(1990)Pectin esterification is spatial y regulated both within cel walls and between developing tissues of root apices.Planta 181:512-521
Kong Y,Pe~na MJ,Renna L,Avci U,Pattathil S,Tuomivaara ST,Li X,Reiter WD,Brandizzi F,Hahn MG,Darvill AG,York WS,O’Neill MA(2015)Galactose-depleted xyloglucan is dysfunctional and leads to dwarfism in Arabidopsis.Plant Physiol 167:1296-1306
Krupkova E,Immerzeel P,Pauly M,Schm€ulling T(2007)The TUMOROUS SHOOT DEVELOPMENT2 gene of Arabidopsis encoding a putative methyltransferase is required for cell adhesion and co-ordinated plant development.Plant J 50:735-750
Lee KJD,Cornuault V,Manfield IW,Ralet M-C,Paul Knox J(2013)Multi-scale spatial heterogeneity of pectic rhamnogalacturonan I(RG-I)structural features in tobacco seed endosperm cell walls.Plant J 75:1018-1027
Lin D,Lopez-Sanchez P,Gidley MJ(2015)Binding of arabinan or galactan during cellulose synthesis is extensive and reversible.Carbohydr Polym 126:108-121
Lin D,Lopez-Sanchez P,Gidley MJ(2016)Interactions of pectins with cellulose during its synthesis in the absence of calcium.Food Hydrocoll 52:57-68
Liners F,Letesson JJ,Didembourg C,Van Cutsem P(1989)Monoclonal antibodies against pectin:Recognition of a conformation induced by calcium.Plant Physiol 91:1419-1424
Liners F,Thibault JF,Van Cutsem P(1992)Influence of the degree of polymerization of oligogalacturonates and of esterification pattern of pectin on their recognition by monoclonal antibodies.Plant Physiol 99:1099-1104
Liners F,Van Cutsem P(1992)Distribution of pectic polysaccharides throughout walls of suspension-cultured carrot cells.Protoplasma 170:10-21
Liu XL,Liu L,Niu QK,Xia C,Yang KZ,Li R,Chen LQ,Zhang XQ,Zhou Y,Ye D(2011)MALE GAMETOPHYTE DEFECTIVE 4encodes a rhamnogalacturonan II xylosyltransferase and is important for growth of pollen tubes and roots in Arabidopsis.Plant J 65:647-660
Liwanag AJM,Ebert B,Verhertbruggen Y,Rennie EA,Rautengarten C,Oikawa A,Andersen MCF,Clausen MH,Scheller HV(2012)Pectin biosynthesis:GALS1 in Arabidopsis thaliana is a b-1,4-galactan b-1,4-galactosyltransferase.Plant Cell 24:5024-5036
Lopez-Sanchez P,Martinez-Sanz M,Bonilla MR,Wang D,Walsh CT,Gilbert EP,Stokes JR,Gidley MJ(2016)Pectin impacts cellulose fibre architecture and hydrogel mechanics in the absence of calcium.Carbohydr Polym 153:236-245
Manabe Y,Nafisi M,Verhertbruggen Y,Orfila C,Gille S,Rautengarten C,Cherk C,Marcus SE,Somerville S,Pauly M,Knox JP,Sakuragi Y,Scheller HV(2011)Loss-offunction mutation of REDUCED WALL ACETYLATION2 in Arabidopsis leads to reduced cell wall acetylation and increased resistance to Botrytis cinerea.Plant Physiol155:1068-1078
Marcus SE,Verhertbruggen Y,Herve C,Ordaz-Ortiz JJ,Farkas V,Pedersen HL,Willats WG,Knox JP(2008)Pectic homogalacturonan masks abundant sets of xyloglucan epitopes in plant cell walls.BMC Plant Biol 8:60
Matoh T,Takasaki M,Takabe K,Kobayashi M(1998)Immunocytochemistry of rhamnogalacturonan II in cell walls of higher plants.Plant Cell Physiol 39:483-491
McCann MC,Stacey NJ,Wilson R,Roberts K(1993)Orientation of macromolecules in the walls of elongating carrot cells.J Cell Sci 106:1347-1356
McCartney L,Ormerod AP,Gidley MJ,Knox JP(2000)Temporal and spatial regulation of pectic(1!4)-b-D-galactan in cell walls of developing pea cotyledons:Implications for mechanical properties.Plant J 22:105-113
McCartney L,Steele-King CG,Jordan E,Knox JP(2003)Cell wall pectic(1!4)-b-D-galactan marks the acceleration of cell elongation in the Arabidopsis seedling root meristem.Plant J 33:447-454
Morris ER,Powell DA,Gidley MJ,Rees DA(1982)Conformations and interactions of pectins.I.Polymorphism between gel and solid states of calcium polygalacturonate.J Mol Biol 155:507-516
Mouille G,Ralet MC,Cavelier C,Eland C,Effroy D,Hematy K,McCartney L,Truong HN,Gaudon V,Thibault JF,Marchant A,H€ofte H(2007)Homogalacturonan synthesis in Arabidopsis thaliana requires a Golgi-localized protein with a putative methyltransferase domain.Plant J 50:605-614
Nakamura A,Furuta H,Maeda H,Takao T,Nagamatsu Y(2002)Structural studies by stepwise enzymatic degradation of the main backbone of soybean soluble polysaccharides consisting of galacturonan and rhamnogalacturonan.Biosci Biotechnol Biochem 66:1301-1313
Naran R,Chen G,Carpita NC(2008)Novel rhamnogalacturonan I and arabinoxylan polysaccharides of flax seed mucilage.Plant Physiol 148:132-141
Ndeh D,Rogowski A,Cartmell A,Luis AS,Basle A,Gray J,Venditto I,Briggs J,Zhang X,Labourel A,Terrapon N,Buffetto F,Nepogodiev S,Xiao Y,Field RA,Zhu Y,O’Neil MA,Urbanowicz BR,York WS,Davies GJ,Abbott DW,Ralet MC,Martens EC,Henrissat B,Gilbert HJ(2017)Complex pectin metabolism by gut bacteria reveals novel catalytic functions.Nature 544:65-70
O’Donoghue EM,Somerfield SD,Deroles SC,Sutherland PW,Hallett IC,Erridge ZA,Brummell DA,Hunter DA(2017)Simultaneous knock-down of six b-galactosidase genes in petunia petals prevents loss of pectic galactan but decreases petal strength.Plant Physiol Biochem 113:208-221
O’Neil MA,Eberhard S,Albersheim P,Darvill AG(2001)Requirement of borate cross-linking of cel wal rhamnogalacturonan II for Arabidopsis growth.Science 294:846-849
O’Neill MA,Ishii T,Albersheim P,Darvill AG(2004)Rhamnogalacturonan II:Structure and function of a borate crosslinked cell wall pectic polysaccharide.Annu Rev Plant Biol55:109-139
Obro J,Borkhardt B,Harholt J,Skj?t M,Willats WGT,Ulvskov P(2009)Simultaneous in vivo truncation of pectic side chains.Transgenic Res 18:961-969
Orfila C,Knox JP(2000)Spatial regulation of pectic polysaccharides in relation to pit fields in cell walls of tomato fruit pericarp.Plant Physiol 122:775-782
Pabst M,Fischl RM,Brecker L,Morelle W,Fauland A,K€ofeler H,Altmann F,Leonard R(2013)Rhamnogalacturonan IIstructure shows variation in the side chains monosaccharide composition and methylation status within and across different plant species.Plant J 76:61-72
Palin R,Geitmann A(2012)The role of pectin in plant morphogenesis.Biosystems 109:397-402
Park YB,Cosgrove DJ(2012a)A revised architecture of primary cell walls based on biomechanical changes induced by substrate-specific endoglucanases.Plant Physiol 158:1933-1943
Park YB,Cosgrove DJ(2012b)Changes in cell wall biomechanical properties in the xyloglucan-deficient xxt1/xxt2 mutant of Arabidopsis.Plant Physiol 158:465-475
Peaucelle A,Braybrook SA,Le Guillou L,Bron E,Kuhlemeier C,H€ofte H(2011a)Pectin-induced changes in cell wall mechanics underlie organ initiation in Arabidopsis.Curr Biol 21:1720-1726
Peaucelle A,Louvet R,Johansen JN,H€ofte H,Laufs P,Pelloux J,Mouille G(2008)Arabidopsis phyllotaxis is controlled by the methyl-esterification status of cell-wall pectins.Curr Biol 18:1943-1948
Peaucelle A,Louvet R,Johansen JN,Salsac F,Morin H,Fournet F,Belcram K,Gillet F,Hofte H,Laufs P,Mouille G,Pelloux J(2011b)The transcription factor BELLRINGER modulates phyllotaxis by regulating the expression of a pectin methylesterase in Arabidopsis.Development 138:4733-4741
Peaucelle A,Wightman R,H€ofte H(2015)The control of growth symmetry breaking in the Arabidopsis hypocotyl.Curr Biol 25:1746-1752
Phyo P,Wang T,Kiemle SN,O’Neill H,Pingali SV,Hong M,Cosgrove DJ(2017a)Gradients in wall mechanics and polysaccharides along growing inflorescence stems.Plant Physiol 175:1593-1607
Phyo P,Wang T,Xiao C,Anderson CT,Hong M(2017b)Effects of pectin molecular weight changes on the structure,dynamics,and polysaccharide interactions of primary cell walls of Arabidopsis thaliana:Insights from solid-state NMR.Biomacromolecules 18:2937-2950
Popper ZA,Fry SC(2008)Xyloglucan-pectin linkages are formed intra-protoplasmically,contribute to wallassembly,and remain stable in the cell wall.Planta 227:781-794
Qi J,Wu B,Feng S,L€u S,Guan C,Zhang X,Qiu D,Hu Y,Zhou Y,Li C,Long M,Jiao Y(2017)Mechanical regulation of organ asymmetry in leaves.Nat Plants 3:724-733
Ralet MC,Cabrera JC,Bonnin E,Quemener B,Hell?n P,Thibault JF(2005)Mapping sugar beet pectin acetylation pattern.Phytochemistry 66:1832-1843
Ralet MC,Crepeau MJ,Buchholt HC,Thibault JF(2003)Polyelectrolyte behaviour and calcium binding properties of sugar beet pectins differing in their degrees of methylation and acetylation.Biochem Eng J 16:191-201
Ralet MC,Crepeau MJ,Vigouroux J,Tran J,Berger A,Salle C,Granier F,Botran L,North HM(2016)Xylans provide the structural driving force for mucilage adhesion to the Arabidopsis seed coat.Plant Physiol 171:165-178
Reiter WD,Chapple CC,Somerville CR(1993)Altered growth and cell walls in a fucose-deficient mutant of Arabidopsis.Science 261:1032-1035
Rui Y,Xiao C,Yi H,Kandemir B,Wang JZ,Puri VM,Anderson CT(2017)POLYGALACTURONASE INVOLVED IN EXPANSION3functions in seedling development,rosette growth,and stomatal dynamics in Arabidopsis thaliana.Plant Cell 29:2413-2432
Ryden P,Sugimoto-Shirasu K,Smith AC,Findlay K,Reiter W-D,McCann MC(2003)Tensile properties of Arabidopsis cell walls depend on both a xyloglucan cross-linked microfibrillar network and rhamnogalacturonan II-borate complexes.Plant Physiol 132:1033-1040
Saffer AM,Carpita NC,Irish VF(2017)Rhamnose-containing cell wall polymers suppress helical plant growth independently of microtubule orientation.Curr Biol 27:2248-2259.e4
Skj?t M,Pauly M,Bush MS,Borkhardt B,McCann MC,Ulvskov P(2002)Direct interference with rhamnogalacturonan Ibiosynthesis in Golgi vesicles.Plant Physiol 129:95-102
S?rensen SO,Pauly M,Bush M,Skj?t M,McCann MC,Borkhardt B,Ulvskov P(2000)Pectin engineering:Modification of potato pectin by in vivo expression of an endo-1,4-beta-D-galactanase.Proc Natl Acad Sci USA97:7639-7644
Stonebloom S,Ebert B,Xiong G,Pattathil S,Birdseye D,Lao J,Pauly M,Hahn MG,Heazlewood JL,Scheller HV(2016)A DUF-246 family glycosyltransferase-like gene affects male fertility and the biosynthesis of pectic arabinogalactans.BMC Plant Biol 16:747
Tan L,Eberhard S,Pattathil S,Warder C,Glushka J,Yuan C,Hao Z,Zhu X,Avci U,Miller JS,Baldwin D,Pham C,Orlando R,Darvill A,Hahn MG,Kieliszewski MJ,Mohnen D(2013)An Arabidopsis cell wall proteoglycan consists of pectin and arabinoxylan covalently linked to an arabinogalactan protein.Plant Cell 25:270-287
Thimm JC,Burritt DJ,Ducker WA,Melton LD(2009)Pectins influence microfibril aggregation in celery cell walls:An atomic force microscopy study.J Struct Biol 168:337-344
Ulvskov P,Wium H,Bruce D,J?rgensen B,Qvist KB,Skj?t M,Hepworth D,Borkhardt B,S?rensen SO(2005)Biophysical consequences of remodeling the neutral side chains of rhamnogalacturonan I in tubers of transgenic potatoes.Planta 220:609-620
Verger S,Chabout S,Gineau E,Mouille G(2016)Cell adhesion in plants is under the control of putative O-fucosyltransferases.Development 143:2536-2540
Verhertbruggen Y,Marcus SE,Chen J,Knox JP(2013)Cell wall pectic arabinans influence the mechanical properties of Arabidopsis thaliana inflorescence stems and their response to mechanical stress.Plant Cell Physiol 54:1278-1288
Verhertbruggen Y,Marcus SE,Haeger A,Verhoef R,Schols HA,McCleary BV,McKee L,Gilbert HJ,Paul Knox J(2009)Developmental complexity of arabinan polysaccharides and their processing in plant cell walls.Plant J 59:413-425
Vogel J(2008)Unique aspects of the grass cell wall.Curr Opin Plant Biol 11:301-307
Voiniciuc C,G€unl M,Schmidt MHW,Usadel B(2015)Highly branched xylan made by IRREGULAR XYLEM14 and MUCILAGE-RELATED21 links mucilage to Arabidopsis seeds.Plant Physiol 169:2481-2495
Wang T,Park YB,Caporini MA,Rosay M,Zhong L,Cosgrove DJ,Hong M(2013)Sensitivity-enhanced solid-state NMRdetection of expansin’s target in plant cell walls.Proc Natl Acad Sci USA 110:16444-16449
Wang T,Park YB,Cosgrove DJ,Hong M(2015)Cellulose-pectin spatial contacts are inherent to never-dried Arabidopsis primary cell walls:Evidence from solid-state nuclear magnetic resonance.Plant Physiol 168:871-884
Wang T,Zabotina O,Hong M(2012)Pectin-cellulose interactions in the Arabidopsis primary cell wall from two-dimensional magic-angle-spinning solid-state nuclear magnetic resonance.Biochemistry 51:9846-9856
Willats WGT,Steele-King CG,Marcus SE,Knox JP(1999)Side chains of pectic polysaccharides are regulated in relation to cell proliferation and cell differentiation.Plant J 20:619-628
Wolf S,Mouille G,Pelloux J(2009)Homogalacturonan methylesterification and plant development.Mol Plant 2:851-860
Xiao C,Barnes WJ,Zamil MS,Yi H,Puri VM,Anderson CT(2017)Activation tagging of Arabidopsis POLYGALACTURONASEINVOLVED IN EXPANSION2 promotes hypocotyl elongation,leaf expansion,stem lignification,mechanical stiffening,and lodging.Plant J 89:1159-1173
Xiao C,Somerville C,Anderson CT(2014)POLYGALACTURONASEINVOLVED IN EXPANSION1 functions in cell elongation and flower development in Arabidopsis.Plant Cell 26:1018-1035
Yang W,Schuster C,Beahan CT,Charoensawan V,Peaucelle A,Bacic A,Doblin MS,Wightman R,Meyerowitz EM(2016)Regulation of meristem morphogenesis by cell wall synthases in Arabidopsis.Curr Biol 26:1404-1415
Yoneda A,Higaki T,Kutsuna N,Kondo Y,Osada H,Hasezawa S,Matsui M(2007)Chemical genetic screening identifies a novel inhibitor of parallel alignment of cortical microtubules and cellulose microfibrils.Plant Cell Physiol 48:1393-1403
Yoneda A,Ito T,Higaki T,Kutsuna N,Saito T,Ishimizu T,Osada H,Hasezawa S,Matsui M,Demura T(2010)Cobtorin target analysis reveals that pectin functions in the deposition of cellulose microfibrils in parallel with cortical microtubules.Plant J 64:657-667
Zablackis E,Huang J,M€uller B,Darvill AG,Albersheim P(1995)Characterization of the cell-wall polysaccharides of Arabidopsis thaliana leaves.Plant Physiol 107:1129-1138
Zabotina OA,Avci U,Cavalier D,Pattathil S,Chou YH,Eberhard S,Danhof L,Keegstra K,Hahn MG(2012)Mutations in multiple XXT genes of Arabidopsis reveal the complexity of xyloglucan biosynthesis.Plant Physiol159:1367-1384
Zandleven J,S?rensen SO,Harholt J,Beldman G,Schols HA,Scheller HV,Voragen AJ(2007)Xylogalacturonan exists in cell walls from various tissues of Arabidopsis thaliana.Phytochemistry 68:1219-1226
Zhao Q,Yuan S,Wang X,Zhang Y,Zhu H,Lu C(2008)Restoration of mature etiolated cucumber hypocotyl cell wall susceptibility to expansin by pretreatment with fungal pectinases and EGTA in vitro.Plant Physiol 147:1874-1885
Zykwinska A,Thibault JF,Ralet MC(2008)Competitive binding of pectin and xyloglucan with primary cell wall cellulose.Carbohydr Polym 74:957-961
Zykwinska A,Thibault JF,Ralet MC(2007)Organization of pectic arabinan and galactan side chains in association with cellulose microfibrils in primary cell walls and related models envisaged.J Exp Bot 58:1795-1802
Zykwinska AW,Ralet MCJ,Garnier CD,Thibault JFJ(2005)Evidence for in vitro binding of pectin side chains to cellulose.Plant Physiol 139:397-407
Amsbury S,Hunt L,Elhaddad N,Baillie A,Lundgren M,Verhertbruggen Y,Scheller HV,Knox JP,Fleming AJ,Gray JE(2016)Stomatal function requires pectin demethyl-esterification of the guard cell wall.Curr Biol 26:2899-2906
Anderson CT,Wallace IS,Somerville CR(2012)Metabolic click-labeling with a fucose analog reveals pectin delivery,architecture,and dynamics in Arabidopsis cell walls.Proc Natl Acad Sci USA 109:1329-1334
Atmodjo MA,Hao Z,Mohnen D(2013)Evolving views of pectin biosynthesis.Annu Rev Plant Biol 64:747-779
Baron-Epel O,Gharyal PK,Schindler M(1988)Pectins as mediators of wall porosity in soybean cells.Planta 175:389-395
Bootten TJ,Harris PJ,Melton LD,Newman RH(2004)Solidstate13C-NMR spectroscopy shows that the xyloglucans in the primary cell walls of mung bean(Vigna radiata L.)occur in different domains:A new model for xyloglucancellulose interactions in the cell wall.J Exp Bot 55:571-583
Bouton S,Leboeuf E,Mouille G,Leydecker MT,Talbotec J,Granier F,Lahaye M,Hofte H,Truong HN(2002)QUASIMODO1 encodes a putative membrane-bound glycosyltransferase required for normal pectin synthesis and cell adhesion in Arabidopsis.Plant Cell 14:2577-2590
Braybrook SA,Peaucelle A(2013)Mechano-chemical aspects of organ formation in Arabidopsis thaliana:The relationship between auxin and pectin.PLoS ONE 8:e57813
Buffetto F,Cornuault V,Rydahl MG,Ropartz D,Alvarado C,Echasserieau V,Le Gall S,Bouchet B,Tranquet O,Verhertbruggen Y,Willats WGT,Knox JP,Ralet MC,Guillon F(2015)The deconstruction of pectic rhamnogalacturonan I unmasks the occurrence of a novel arabinogalactan oligosaccharide epitope.Plant Cell Physiol 56:2181-2196
Bush MS,Marry M,Huxham MI,Jarvis MC,McCann MC(2001)Developmental regulation of pectic epitopes during potato tuberisation.Planta 213:869-880
Carpita NC,Gibeaut DM(1993)Structural models of primary cell walls in flowering plants:Consistency of molecular structure with the physical properties of the walls during growth.Plant J 3:1-30
Carter R,Woolfenden H,Baillie A,Amsbury S,Carroll S,Healicon E,Sovatzoglou S,Braybrook S,Gray JE,Hobbs J,Morris RJ,Fleming AJ(2017)Stomatal opening involves polar,not radial,stiffening of guard cells.Curr Biol 27:2974-2983.e2
Cavalier DM,Lerouxel O,Neumetzler L,Yamauchi K,Reinecke A,Freshour G,Zabotina OA,Hahn MG,Burgert I,Pauly M,Raikhel NV,Keegstra K(2008)Disrupting two Arabidopsis thaliana xylosyltransferase genes results in plants deficient in xyloglucan,a major primary cell wall component.Plant Cell 20:1519-1537
Coenen GJ,Bakx EJ,Verhoef RP,Schols HA,Voragen AGJ(2007)Identification of the connecting linkage between homo-or xylogalacturonan and rhamnogalacturonan type I.Carbohydr Polym 70:224-235
Cornuault V,Buffetto F,Rydahl MG,Marcus SE,Torode TA,Xue J,Crepeau MJ,Faria-Blanc N,Willats WGT,Dupree P,Ralet MC,Knox JP(2015)Monoclonal antibodies indicate low-abundance links between heteroxylan and other glycans of plant cell walls.Planta 242:1321-1334
Daher FB,Braybrook SA(2015)How to let go:Pectin and plant cell adhesion.Front Plant Sci 6:523
de Souza A,Hull PA,Gille S,Pauly M(2014)Identification and functional characterization of the distinct plant pectin esterases PAE8 and PAE9 and their deletion mutants.Planta 240:1123-1138
Derbyshire P,McCann MC,Roberts K(2007)Restricted cell elongation in Arabidopsis hypocotyls is associated with a reduced average pectin esterification level.BMC Plant Biol7:31
Dick-Perez M,Wang T,Salazar A,Zabotina OA,Hong M(2012)Multidimensional solid-state NMR studies of the structure and dynamics of pectic polysaccharides in uniformly13C-labeled Arabidopsis primary cell walls.Magn Reson Chem50:539-550
Dick-Perez M,Zhang Y,Hayes J,Salazar A,Zabotina OA,Hong M(2011)Structure and interactions of plant cell-wall polysaccharides by two-and three-dimensional magic-anglespinning solid-state NMR.Biochemistry 50:989-1000
Dumont M,Lehner A,Bouton S,Kiefer-Meyer MC,Voxeur A,Pelloux J,Lerouge P,Mollet JC(2014)The cell wall pectic polymer rhamnogalacturonan-II is required for proper pollen tube elongation:Implications of a putative sialyltransferase-like protein.Ann Bot 114:1177-1188
Ermel FF,Follet-Gueye ML,Cibert C,Vian B,Morvan C,Catesson AM,Goldberg R(2000)Differential localization of arabinan and galactan side chains of rhamnogalacturonan 1 in cambial derivatives.Planta 210:732-740
Goldberg R,Morvan C,Roland JC(1986)Composition,properties and localisation of pectins in young and mature cells of the mung bean hypocotyl.Plant Cell Physiol 27:417-429
Gou JY,Miller LM,Hou G,Yu XH,Chen XY,Liu CJ(2012)Acetylesterase-mediated deacetylation of pectin impairs cell elongation,pollen germination,and plant reproduction.Plant Cell 24:50-65
Grant GT,Morris ER,Rees DA,Smith PJC,Thom D(1973)Biological interactions between polysaccharides and divalent cations:The egg-box model.FEBS Lett 32:195-198
Guillemin F,Guillon F,Bonnin E,Devaux MF,Chevalier T,Knox JP,Liners F,Thibault JF(2005)Distribution of pectic epitopes in cell walls of the sugar beet root.Planta 222:355-371
Hall HC,Cheung J,Ellis BE(2013)Immunoprofiling reveals unique cell-specific patterns of wall epitopes in the expanding Arabidopsis stem.Plant J 74:134-147
Harholt J(2005)ARABINAN DEFICIENT 1 is a putative arabinosyltransferase involved in biosynthesis of pectic arabinan in Arabidopsis.Plant Physiol 140:49-58
Hongo S,Sato K,Yokoyama R,Nishitani K(2012)Demethylesterification of the primary wall by PECTIN METHYL-ESTERASE35 provides mechanical support to the Arabidopsis stem.Plant Cell 24:2624-2634
Ishii T(1997)O-Acetylated oligosaccharides from pectins of potato tuber cell walls.Plant Physiol 113:1265-1272
Iwai H,Masaoka N,Ishii T,Satoh S(2002)A pectin glucuronyltransferase gene is essential for intercellular attachment in the plant meristem.Proc Natl Acad Sci USA99:16319-16324
Jensen JK,Sorensen SO,Harholt J,Geshi N,Sakuragi Y,Moller I,Zandleven J,Bernal AJ,Jensen NB,Sorensen C,Pauly M,Beldman G,Willats WGT,Scheller HV(2008)Identification of a xylogalacturonan xylosyltransferase involved in pectin biosynthesis in Arabidopsis.Plant Cell 20:1289-1302
Jones L,Milne JL,Ashford D,McQueen-Mason SJ(2003)Cell wall arabinan is essential for guard cell function.Proc Natl Acad Sci USA 100:11783-11788
Kim S-J,Held MA,Zemelis S,Wilkerson C,Brandizzi F(2015)CGR2 and CGR3 have critical overlapping roles in pectin methylesterification and plant growth in Arabidopsis thaliana.Plant J 82:208-220
Knox JP,Linstead PJ,King J,Cooper C,Roberts K(1990)Pectin esterification is spatial y regulated both within cel walls and between developing tissues of root apices.Planta 181:512-521
Kong Y,Pe~na MJ,Renna L,Avci U,Pattathil S,Tuomivaara ST,Li X,Reiter WD,Brandizzi F,Hahn MG,Darvill AG,York WS,O’Neill MA(2015)Galactose-depleted xyloglucan is dysfunctional and leads to dwarfism in Arabidopsis.Plant Physiol 167:1296-1306
Krupkova E,Immerzeel P,Pauly M,Schm€ulling T(2007)The TUMOROUS SHOOT DEVELOPMENT2 gene of Arabidopsis encoding a putative methyltransferase is required for cell adhesion and co-ordinated plant development.Plant J 50:735-750
Lee KJD,Cornuault V,Manfield IW,Ralet M-C,Paul Knox J(2013)Multi-scale spatial heterogeneity of pectic rhamnogalacturonan I(RG-I)structural features in tobacco seed endosperm cell walls.Plant J 75:1018-1027
Lin D,Lopez-Sanchez P,Gidley MJ(2015)Binding of arabinan or galactan during cellulose synthesis is extensive and reversible.Carbohydr Polym 126:108-121
Lin D,Lopez-Sanchez P,Gidley MJ(2016)Interactions of pectins with cellulose during its synthesis in the absence of calcium.Food Hydrocoll 52:57-68
Liners F,Letesson JJ,Didembourg C,Van Cutsem P(1989)Monoclonal antibodies against pectin:Recognition of a conformation induced by calcium.Plant Physiol 91:1419-1424
Liners F,Thibault JF,Van Cutsem P(1992)Influence of the degree of polymerization of oligogalacturonates and of esterification pattern of pectin on their recognition by monoclonal antibodies.Plant Physiol 99:1099-1104
Liners F,Van Cutsem P(1992)Distribution of pectic polysaccharides throughout walls of suspension-cultured carrot cells.Protoplasma 170:10-21
Liu XL,Liu L,Niu QK,Xia C,Yang KZ,Li R,Chen LQ,Zhang XQ,Zhou Y,Ye D(2011)MALE GAMETOPHYTE DEFECTIVE 4encodes a rhamnogalacturonan II xylosyltransferase and is important for growth of pollen tubes and roots in Arabidopsis.Plant J 65:647-660
Liwanag AJM,Ebert B,Verhertbruggen Y,Rennie EA,Rautengarten C,Oikawa A,Andersen MCF,Clausen MH,Scheller HV(2012)Pectin biosynthesis:GALS1 in Arabidopsis thaliana is a b-1,4-galactan b-1,4-galactosyltransferase.Plant Cell 24:5024-5036
Lopez-Sanchez P,Martinez-Sanz M,Bonilla MR,Wang D,Walsh CT,Gilbert EP,Stokes JR,Gidley MJ(2016)Pectin impacts cellulose fibre architecture and hydrogel mechanics in the absence of calcium.Carbohydr Polym 153:236-245
Manabe Y,Nafisi M,Verhertbruggen Y,Orfila C,Gille S,Rautengarten C,Cherk C,Marcus SE,Somerville S,Pauly M,Knox JP,Sakuragi Y,Scheller HV(2011)Loss-offunction mutation of REDUCED WALL ACETYLATION2 in Arabidopsis leads to reduced cell wall acetylation and increased resistance to Botrytis cinerea.Plant Physiol155:1068-1078
Marcus SE,Verhertbruggen Y,Herve C,Ordaz-Ortiz JJ,Farkas V,Pedersen HL,Willats WG,Knox JP(2008)Pectic homogalacturonan masks abundant sets of xyloglucan epitopes in plant cell walls.BMC Plant Biol 8:60
Matoh T,Takasaki M,Takabe K,Kobayashi M(1998)Immunocytochemistry of rhamnogalacturonan II in cell walls of higher plants.Plant Cell Physiol 39:483-491
McCann MC,Stacey NJ,Wilson R,Roberts K(1993)Orientation of macromolecules in the walls of elongating carrot cells.J Cell Sci 106:1347-1356
McCartney L,Ormerod AP,Gidley MJ,Knox JP(2000)Temporal and spatial regulation of pectic(1!4)-b-D-galactan in cell walls of developing pea cotyledons:Implications for mechanical properties.Plant J 22:105-113
McCartney L,Steele-King CG,Jordan E,Knox JP(2003)Cell wall pectic(1!4)-b-D-galactan marks the acceleration of cell elongation in the Arabidopsis seedling root meristem.Plant J 33:447-454
Morris ER,Powell DA,Gidley MJ,Rees DA(1982)Conformations and interactions of pectins.I.Polymorphism between gel and solid states of calcium polygalacturonate.J Mol Biol 155:507-516
Mouille G,Ralet MC,Cavelier C,Eland C,Effroy D,Hematy K,McCartney L,Truong HN,Gaudon V,Thibault JF,Marchant A,H€ofte H(2007)Homogalacturonan synthesis in Arabidopsis thaliana requires a Golgi-localized protein with a putative methyltransferase domain.Plant J 50:605-614
Nakamura A,Furuta H,Maeda H,Takao T,Nagamatsu Y(2002)Structural studies by stepwise enzymatic degradation of the main backbone of soybean soluble polysaccharides consisting of galacturonan and rhamnogalacturonan.Biosci Biotechnol Biochem 66:1301-1313
Naran R,Chen G,Carpita NC(2008)Novel rhamnogalacturonan I and arabinoxylan polysaccharides of flax seed mucilage.Plant Physiol 148:132-141
Ndeh D,Rogowski A,Cartmell A,Luis AS,Basle A,Gray J,Venditto I,Briggs J,Zhang X,Labourel A,Terrapon N,Buffetto F,Nepogodiev S,Xiao Y,Field RA,Zhu Y,O’Neil MA,Urbanowicz BR,York WS,Davies GJ,Abbott DW,Ralet MC,Martens EC,Henrissat B,Gilbert HJ(2017)Complex pectin metabolism by gut bacteria reveals novel catalytic functions.Nature 544:65-70
O’Donoghue EM,Somerfield SD,Deroles SC,Sutherland PW,Hallett IC,Erridge ZA,Brummell DA,Hunter DA(2017)Simultaneous knock-down of six b-galactosidase genes in petunia petals prevents loss of pectic galactan but decreases petal strength.Plant Physiol Biochem 113:208-221
O’Neil MA,Eberhard S,Albersheim P,Darvill AG(2001)Requirement of borate cross-linking of cel wal rhamnogalacturonan II for Arabidopsis growth.Science 294:846-849
O’Neill MA,Ishii T,Albersheim P,Darvill AG(2004)Rhamnogalacturonan II:Structure and function of a borate crosslinked cell wall pectic polysaccharide.Annu Rev Plant Biol55:109-139
Obro J,Borkhardt B,Harholt J,Skj?t M,Willats WGT,Ulvskov P(2009)Simultaneous in vivo truncation of pectic side chains.Transgenic Res 18:961-969
Orfila C,Knox JP(2000)Spatial regulation of pectic polysaccharides in relation to pit fields in cell walls of tomato fruit pericarp.Plant Physiol 122:775-782
Pabst M,Fischl RM,Brecker L,Morelle W,Fauland A,K€ofeler H,Altmann F,Leonard R(2013)Rhamnogalacturonan IIstructure shows variation in the side chains monosaccharide composition and methylation status within and across different plant species.Plant J 76:61-72
Palin R,Geitmann A(2012)The role of pectin in plant morphogenesis.Biosystems 109:397-402
Park YB,Cosgrove DJ(2012a)A revised architecture of primary cell walls based on biomechanical changes induced by substrate-specific endoglucanases.Plant Physiol 158:1933-1943
Park YB,Cosgrove DJ(2012b)Changes in cell wall biomechanical properties in the xyloglucan-deficient xxt1/xxt2 mutant of Arabidopsis.Plant Physiol 158:465-475
Peaucelle A,Braybrook SA,Le Guillou L,Bron E,Kuhlemeier C,H€ofte H(2011a)Pectin-induced changes in cell wall mechanics underlie organ initiation in Arabidopsis.Curr Biol 21:1720-1726
Peaucelle A,Louvet R,Johansen JN,H€ofte H,Laufs P,Pelloux J,Mouille G(2008)Arabidopsis phyllotaxis is controlled by the methyl-esterification status of cell-wall pectins.Curr Biol 18:1943-1948
Peaucelle A,Louvet R,Johansen JN,Salsac F,Morin H,Fournet F,Belcram K,Gillet F,Hofte H,Laufs P,Mouille G,Pelloux J(2011b)The transcription factor BELLRINGER modulates phyllotaxis by regulating the expression of a pectin methylesterase in Arabidopsis.Development 138:4733-4741
Peaucelle A,Wightman R,H€ofte H(2015)The control of growth symmetry breaking in the Arabidopsis hypocotyl.Curr Biol 25:1746-1752
Phyo P,Wang T,Kiemle SN,O’Neill H,Pingali SV,Hong M,Cosgrove DJ(2017a)Gradients in wall mechanics and polysaccharides along growing inflorescence stems.Plant Physiol 175:1593-1607
Phyo P,Wang T,Xiao C,Anderson CT,Hong M(2017b)Effects of pectin molecular weight changes on the structure,dynamics,and polysaccharide interactions of primary cell walls of Arabidopsis thaliana:Insights from solid-state NMR.Biomacromolecules 18:2937-2950
Popper ZA,Fry SC(2008)Xyloglucan-pectin linkages are formed intra-protoplasmically,contribute to wallassembly,and remain stable in the cell wall.Planta 227:781-794
Qi J,Wu B,Feng S,L€u S,Guan C,Zhang X,Qiu D,Hu Y,Zhou Y,Li C,Long M,Jiao Y(2017)Mechanical regulation of organ asymmetry in leaves.Nat Plants 3:724-733
Ralet MC,Cabrera JC,Bonnin E,Quemener B,Hell?n P,Thibault JF(2005)Mapping sugar beet pectin acetylation pattern.Phytochemistry 66:1832-1843
Ralet MC,Crepeau MJ,Buchholt HC,Thibault JF(2003)Polyelectrolyte behaviour and calcium binding properties of sugar beet pectins differing in their degrees of methylation and acetylation.Biochem Eng J 16:191-201
Ralet MC,Crepeau MJ,Vigouroux J,Tran J,Berger A,Salle C,Granier F,Botran L,North HM(2016)Xylans provide the structural driving force for mucilage adhesion to the Arabidopsis seed coat.Plant Physiol 171:165-178
Reiter WD,Chapple CC,Somerville CR(1993)Altered growth and cell walls in a fucose-deficient mutant of Arabidopsis.Science 261:1032-1035
Rui Y,Xiao C,Yi H,Kandemir B,Wang JZ,Puri VM,Anderson CT(2017)POLYGALACTURONASE INVOLVED IN EXPANSION3functions in seedling development,rosette growth,and stomatal dynamics in Arabidopsis thaliana.Plant Cell 29:2413-2432
Ryden P,Sugimoto-Shirasu K,Smith AC,Findlay K,Reiter W-D,McCann MC(2003)Tensile properties of Arabidopsis cell walls depend on both a xyloglucan cross-linked microfibrillar network and rhamnogalacturonan II-borate complexes.Plant Physiol 132:1033-1040
Saffer AM,Carpita NC,Irish VF(2017)Rhamnose-containing cell wall polymers suppress helical plant growth independently of microtubule orientation.Curr Biol 27:2248-2259.e4
Skj?t M,Pauly M,Bush MS,Borkhardt B,McCann MC,Ulvskov P(2002)Direct interference with rhamnogalacturonan Ibiosynthesis in Golgi vesicles.Plant Physiol 129:95-102
S?rensen SO,Pauly M,Bush M,Skj?t M,McCann MC,Borkhardt B,Ulvskov P(2000)Pectin engineering:Modification of potato pectin by in vivo expression of an endo-1,4-beta-D-galactanase.Proc Natl Acad Sci USA97:7639-7644
Stonebloom S,Ebert B,Xiong G,Pattathil S,Birdseye D,Lao J,Pauly M,Hahn MG,Heazlewood JL,Scheller HV(2016)A DUF-246 family glycosyltransferase-like gene affects male fertility and the biosynthesis of pectic arabinogalactans.BMC Plant Biol 16:747
Tan L,Eberhard S,Pattathil S,Warder C,Glushka J,Yuan C,Hao Z,Zhu X,Avci U,Miller JS,Baldwin D,Pham C,Orlando R,Darvill A,Hahn MG,Kieliszewski MJ,Mohnen D(2013)An Arabidopsis cell wall proteoglycan consists of pectin and arabinoxylan covalently linked to an arabinogalactan protein.Plant Cell 25:270-287
Thimm JC,Burritt DJ,Ducker WA,Melton LD(2009)Pectins influence microfibril aggregation in celery cell walls:An atomic force microscopy study.J Struct Biol 168:337-344
Ulvskov P,Wium H,Bruce D,J?rgensen B,Qvist KB,Skj?t M,Hepworth D,Borkhardt B,S?rensen SO(2005)Biophysical consequences of remodeling the neutral side chains of rhamnogalacturonan I in tubers of transgenic potatoes.Planta 220:609-620
Verger S,Chabout S,Gineau E,Mouille G(2016)Cell adhesion in plants is under the control of putative O-fucosyltransferases.Development 143:2536-2540
Verhertbruggen Y,Marcus SE,Chen J,Knox JP(2013)Cell wall pectic arabinans influence the mechanical properties of Arabidopsis thaliana inflorescence stems and their response to mechanical stress.Plant Cell Physiol 54:1278-1288
Verhertbruggen Y,Marcus SE,Haeger A,Verhoef R,Schols HA,McCleary BV,McKee L,Gilbert HJ,Paul Knox J(2009)Developmental complexity of arabinan polysaccharides and their processing in plant cell walls.Plant J 59:413-425
Vogel J(2008)Unique aspects of the grass cell wall.Curr Opin Plant Biol 11:301-307
Voiniciuc C,G€unl M,Schmidt MHW,Usadel B(2015)Highly branched xylan made by IRREGULAR XYLEM14 and MUCILAGE-RELATED21 links mucilage to Arabidopsis seeds.Plant Physiol 169:2481-2495
Wang T,Park YB,Caporini MA,Rosay M,Zhong L,Cosgrove DJ,Hong M(2013)Sensitivity-enhanced solid-state NMRdetection of expansin’s target in plant cell walls.Proc Natl Acad Sci USA 110:16444-16449
Wang T,Park YB,Cosgrove DJ,Hong M(2015)Cellulose-pectin spatial contacts are inherent to never-dried Arabidopsis primary cell walls:Evidence from solid-state nuclear magnetic resonance.Plant Physiol 168:871-884
Wang T,Zabotina O,Hong M(2012)Pectin-cellulose interactions in the Arabidopsis primary cell wall from two-dimensional magic-angle-spinning solid-state nuclear magnetic resonance.Biochemistry 51:9846-9856
Willats WGT,Steele-King CG,Marcus SE,Knox JP(1999)Side chains of pectic polysaccharides are regulated in relation to cell proliferation and cell differentiation.Plant J 20:619-628
Wolf S,Mouille G,Pelloux J(2009)Homogalacturonan methylesterification and plant development.Mol Plant 2:851-860
Xiao C,Barnes WJ,Zamil MS,Yi H,Puri VM,Anderson CT(2017)Activation tagging of Arabidopsis POLYGALACTURONASEINVOLVED IN EXPANSION2 promotes hypocotyl elongation,leaf expansion,stem lignification,mechanical stiffening,and lodging.Plant J 89:1159-1173
Xiao C,Somerville C,Anderson CT(2014)POLYGALACTURONASEINVOLVED IN EXPANSION1 functions in cell elongation and flower development in Arabidopsis.Plant Cell 26:1018-1035
Yang W,Schuster C,Beahan CT,Charoensawan V,Peaucelle A,Bacic A,Doblin MS,Wightman R,Meyerowitz EM(2016)Regulation of meristem morphogenesis by cell wall synthases in Arabidopsis.Curr Biol 26:1404-1415
Yoneda A,Higaki T,Kutsuna N,Kondo Y,Osada H,Hasezawa S,Matsui M(2007)Chemical genetic screening identifies a novel inhibitor of parallel alignment of cortical microtubules and cellulose microfibrils.Plant Cell Physiol 48:1393-1403
Yoneda A,Ito T,Higaki T,Kutsuna N,Saito T,Ishimizu T,Osada H,Hasezawa S,Matsui M,Demura T(2010)Cobtorin target analysis reveals that pectin functions in the deposition of cellulose microfibrils in parallel with cortical microtubules.Plant J 64:657-667
Zablackis E,Huang J,M€uller B,Darvill AG,Albersheim P(1995)Characterization of the cell-wall polysaccharides of Arabidopsis thaliana leaves.Plant Physiol 107:1129-1138
Zabotina OA,Avci U,Cavalier D,Pattathil S,Chou YH,Eberhard S,Danhof L,Keegstra K,Hahn MG(2012)Mutations in multiple XXT genes of Arabidopsis reveal the complexity of xyloglucan biosynthesis.Plant Physiol159:1367-1384
Zandleven J,S?rensen SO,Harholt J,Beldman G,Schols HA,Scheller HV,Voragen AJ(2007)Xylogalacturonan exists in cell walls from various tissues of Arabidopsis thaliana.Phytochemistry 68:1219-1226
Zhao Q,Yuan S,Wang X,Zhang Y,Zhu H,Lu C(2008)Restoration of mature etiolated cucumber hypocotyl cell wall susceptibility to expansin by pretreatment with fungal pectinases and EGTA in vitro.Plant Physiol 147:1874-1885
Zykwinska A,Thibault JF,Ralet MC(2008)Competitive binding of pectin and xyloglucan with primary cell wall cellulose.Carbohydr Polym 74:957-961
Zykwinska A,Thibault JF,Ralet MC(2007)Organization of pectic arabinan and galactan side chains in association with cellulose microfibrils in primary cell walls and related models envisaged.J Exp Bot 58:1795-1802
Zykwinska AW,Ralet MCJ,Garnier CD,Thibault JFJ(2005)Evidence for in vitro binding of pectin side chains to cellulose.Plant Physiol 139:397-407