Cellulose Biosynthesis in Plants: from Genes to RosettesCellulose Biosynthesis in Plants: from Genes to RosettesCellulose Biosynthesis in Plants: from Genes to RosettesCellulose Biosynthesis in Plants: from Genes to RosettesCellulose Biosynthesis in Plants: from Genes to RosettesDoblin,M.S.Doblin,M.S.Doblin,M.S.Monika S.Kurek,I.Kurek,I.IsaacJacob-Wilk,D.Jacob-Wilk,D.DeborahDelmer,D.P.Delmer,D.P.Deborah P.University of California, Davis, CA 95616, U.S.A.University of California, Davis, CA 95616, U.S.A.University of California, Davis, CA 95616, U.S.A.Modern techniques of gene cloning have identified theKeywordsKeywordsAbbreviations: CalS, callose synthase enzyme complex; CD, cellodextrin; c-di-GMP, cyclic diguanyl...IntroductionIntroductionIntroductionIdentification of CesA GenesIdentification of CesA GenesDiscovery of the CesA/Csl superfamilyDiscovery of the CesA/Csl superfamilyOrigin and evolution of the plant CesA/Csl supergene familyOrigin and evolution of the plant CesA/Csl supergene familyEvidence for a specialized role for different CesA family membersEvidence for a specialized role for different CesA family members(a) CesA expression and regulation(a) CesA expression and regulation(b) Rosette assembly(b) Rosette assembly(c) Possible functional roles for CesA isoforms(c) Possible functional roles for CesA isoformsOther genes involved in cellulose biosynthesisOther genes involved in cellulose biosynthesisOpen questions for the futureOpen questions for the futureStatus of in vitro cellulose synthesis in plantsStatus of in vitro cellulose synthesis in plantsConclusionsConclusionsReferencesReferencesAlbersheimAlbersheimAlbersheimAlbersheimAlbersheimAmorAmorAmorArabidopsis Genome Initiative, The (ArioliArioliArioliArioliArioliArioliAusmeesAusmeesAusmeesAusmeesAusmeesAusmeesBaskinBaskinBaskinBeckerBeckerBeckerBlantonBlantonBlantonBrownBrownBrownBrownBrownBrownBuckeridgeBuckeridgeBuckeridgeBurnBurnBurnCampbellCampbellCampbellCantatoreCantatoreCantatoreCarpitaCarpitaCarpitaCharnockCharnockCharnockCharnockCharnockCharnockCronnCronnCronnCutlerCutlerCutlerDelmerDelmerDelmerDelmerDelmerDelmerDelmerDelmerDelmerDelmerDelmerDelmerDelmerDelmerDelmerDesprezDesprezDesprezDhuggaDhuggaDhuggaDoblinDoblinDoblinElbeinElbeinElbeinFagardFagardFagardGibeautGibeautGibeautHaHaHaHaiglerHaiglerHaiglerHaiglerHaiglerHaiglerHazenHazenHazenHenrissatHenrissatHenrissatHollandHollandHollandKawagoeKawagoeKawagoeKimuraKimuraKimuraKimuraKimuraKimuraKoyamaKoyamaKoyamaKudlickaKudlickaKudlickaKudlickaKudlickaKudlickaKurekKurekKurekLiuLiuLiuLukowitzLukowitzLukowitzMaMaMaMaretMaretMaretMaretMaretMaretMatthysseMatthysseMatthysseMatthysseMatthysseMatthysseMayerMayerMayerMizunoMizunoMizunoMølhøjMølhøjMølhøjMontezinosMontezinosMontezinosNagahashiNagahashiNagahashiNakagawaNakagawaNakagawaNakaiNakaiNakaiNicolNicolNicolNoblesNoblesNoblesOkudaOkudaOkudaPearPearPearPengPengPengPengPengPengPerrinPerrinPerrinPotikhaPotikhaPotikhaPotikhaPotikhaPotikhaReadReadReadRichmondRichmondRichmondRichmondRichmondRichmondRobertsRobertsRobertsRossRossRossRömlingRömlingRömlingRudolphRudolphRudolphRujanRujanRujanSatoSatoSatoSaxenaSaxenaSaxenaSaxenaSaxenaSaxenaSaxenaSaxenaSaxenaSaxenaSaxenaSaxenaSaxenaSaxenaSaxenaScheibleScheibleScheibleSchmidtSchmidtSchmidtSeagullSeagullSeagullStasinopoulosStasinopoulosStasinopoulosTaylorTaylorTaylorTaylorTaylorTaylorTaylorTaylorTaylorTsekosTsekosTsekosTurnerTurnerTurnerVasakVasakVasakVergaraVergaraVergaraWarneckeWarneckeWarneckeWendelWendelWendelWilliamsonWilliamsonWilliamsonWilliamsonWilliamsonWilliamsonWhittakerWhittakerWhittakerWuWuWuZogajZogajZogaj(Received April 22, 2002(Received April 22, 2002Accepted July 9, 2002)Plant Cell Physiol. 43(12): 1407–1420 (2002)JSPP © 20021407Cellulose Biosynthesis in Plants: from Genes to RosettesMonika S. Doblin1, 2, Isaac Kurek, Deborah Jacob-Wilk and Deborah P. DelmerUniversity of California, Davis, CA 95616, U.S.A.;Modern techniques of gene cloning have identified theCesA genes as encoding the probable catalytic subunits of theplant CelS, the cellulose synthase enzyme complex visualizedin the plasma membrane as rosettes. At least 10 CesA iso-forms exist in Arabidopsis and have been shown by mutantanalyses to play distinct role/s in the cellulose synthesis proc-ess. Functional specialization within this family includes dif-ferences in gene expression, regulation and, possibly, cata-lytic function. Current data points towards some CesAisoforms potentially being responsible for initiation or elon-gation of the recently identified sterol b-glucoside primerwithin different cell types, e.g. those undergoing either pri-mary or secondary wall cellulose synthesis. Different CesAisoforms may also play distinct roles within the rosette, andthere is some circumstantial evidence that CesA genes mayencode the catalytic subunit of the mixed linkage glucan syn-thase or callose synthase. Various other proteins such as theKorrigan endocellulase, sucrose synthase, cytoskeletal com-ponents, Rac13, redox proteins and a lipid transfer proteinhave been implicated to be involved in synthesizing cellulosebut, apart from CesAs, only Korrigan has been definitivelylinked with cellulose synthesis. These proteins should provevaluable in identifying additional CelS components.Keywords: Arabidopsis thaliana — Cellulose — CesA —Cotton (Gossypium hirsutum) — Plant polysaccharide biosyn-thesis.Abbreviations: CalS, callose synthase enzyme complex; CD, cel-lodextrin; c-di-GMP, cyclic diguanylic acid; Csl, cellulose synthase-like; CelS, cellulose synthase enzyme complex; CesA (formerlyCelA), cellulose synthase catalytic subunit; CGAbp, cellulose synthe-sis inhibitor CGA 325¢615 binding protein; CR-P, plant conservedregion; dpa, days post anthesis; DCB, 2,6-dichlorobenzonitrile; Glc,glucose; GT, glycosyltransferase; HVR, hypervariable region; Kor,Korrigan endocellulase; LTP, lipid transfer protein; Mt, metal-lothionein; MT, microtubule; SCD, sterol cellodextrin; SG, sitosterol-b-glucoside; SuSy, sucrose synthase; TC, terminal complex; TMH,transmembrane helix, UDP-Glc, uridine diphospho-glucose.IntroductionUnderstanding the biosynthesis of wall polysaccharidecomponents has attracted considerable interest in light of thefundamental importance of these molecules not just to plantfunction, but also to man. Unfortunately, very little is known ofthe mechanism(s) and regulation of the biosynthetic steps thatcontrol polysaccharide biosynthesis, deposition and assembly,or the interaction of