摘要
根毛是植物从土壤中吸收养分及水分的重要器官,并提供一个根系与微生物互作的环境。根毛发育可分为三个阶段:细胞命运决定,根毛起始,及根毛伸长。植物根毛的机制研究目前主要集中在拟南芥中,水稻中报道较少。在拟南芥的报道中发现AtEXPA7与AtEXPA18与根毛的发育有关,但是AtEXPA7的T-DNA插入纯合突变体没有发现任何根毛突变表型。推测可能是同源基因AtEXPA18的功能互补,也可能由于EXPANSIN家族其他功能冗余的基因。
在水稻中,OsEXPA17基因发生点突变后,出现了短根毛表型。采用RNAi技术,抑制该基因表达也出现了短根毛表型。由此证实,点突变导致了OsEXPA17蛋白质失去了功能。点突变使第104位的氨基酸序列发生改变,由Gly变成了Arg。根据已报道的ZmEXPB1晶体结构分析推知,OsEXPA17第103位及第106位处的两个半胱氨酸,它们分别与第74位及第180位半胱氨酸形成二硫键。因此推测可能由于G104R的改变而使这两个二硫键形成受到了影响,而导致OsEXPA17蛋白功能丧失。
同源基因OsEXPA30, AtEXPA7互补实验发现,OsEXPA30能够部分回复OsexpA17短根毛表型,AtEXPA7也同样能够部分回复OsexpA17短根毛表型。说明EXPA7亚家族对根毛伸长功能是保守的。
通过质壁分离发现,OsEXPA17、OsEXPA30及OsexpA17是个一个膜蛋白。
Root hair is an important organ for plants to uptake nutrients and water from the rhizosphere and serves as a site of interaction with soil microorganisms. Root hair development can be divided into three stages:cell specification, initiation, and elongation. Extensive studies have been reported on root hair development in Arabidopsis (the dicot model plant). In rice (the monocot model plant) the knowledge on root hair development is still limited.It has been reported in Arabidopsis that AtEXPA7 and AtEXPA18 are tightly linked to root hair initiation and development. But the homozygous T-DNA insert mutant of AtEXPA7 has no obvious compaired phenotype with the wild type in root hair, for which the functional redundancy by AtEXPA18 and perhaps other expansin genes.
A short root hair mutant caused by mutation of OsEXPA17, is a orthology gene of AtEXPA7, has been founded. RNA interference of OsEXPA17 gene also results in short root hair phenotype, which conforms that the point mutant of OsEXPA17 makes the gene function lost. The point mutation results in a change of 104th site amino acid Gly to Arg. There are two cysteines in 103th site and 106th site near 104th site which can form two disulfide linkage with 74th site and 180th site cysteines alternatively. So we presume the amino acid change in 104th site may influence the disulfide linkage structure.
Overexpressing OsEXPA30 and AtEXPA7 can partially restore OsexpA17 short root hair mutant.
Through transient expression of OsEXPA17, OsEXPA30 and OsexpA17, we discover that they are plasma membrane proteins through plasmolysis.
引文
Allen-Baume V, Segui B, Cockcroft S.2002. Current thoughts on the phosphatidylinositol transfer protein family. FEBS Lett 531:74-80
Allwood EG, Anthony RG, Smertenko AP, Reichelt S, Drobak BK, et al.2002. Regulation of the pollen-specific actin-depolymerizing factor LlADF1. Plant Cell 14:2915-2927
Anthony RG, Henriques R, Helfer A, Meeszaros T, Rios G, et al.2004. A protein kinase target of a PDK1 signalling pathway is involved in root hair growth in Arabidopsis. EMBO J 23:572-581
Baluska F, Salaj J, Mathur J, Braun M, Jasper F, et al.2000. Root hair formation: F-actin-dependent tip growth is initiated by local assembly of proilin-supported F-actin meshworks accumulated within expansin-enriched bulges. Dev Biol 227:618-632
Bao Y, Kost B, Chua NH.2001. Reduced expression of α-tubulin genes in Arabidopsis thaliana specifically affects root growth and morphology, root hair development and root gravitropism. Plant J 28:145-157
Baumberger N, Ringli C, Keller B.2001. The chimeric leucine-rich repeat/extensin cell wall protein LRX1 is required for root hair morphogenesis in Arabidopsis thaliana. Genes Dev 15:1128-1139
Baumberger N, Steiner M, Ryser U, Keller B, Ringli C.2003. Synergistic interaction of the two paralogous Arabidopsis genes LRX1 and LRX2 in cell wall formation during root hair development. Plant J 35:71-81
Berger F, Haseloff J, Schiefelbein J, Dolan L.1998. Positional information in root epidermis is defined during embryogenesis and acts in domains with strict boundaries. Curr Biol 8:421-443
Bernhardt C, Lee MM, Gonzalez A, Zhang F, Lloyd A, Schiefelbein J.2003. The bHLH genes GLABRA3 (GL3) and ENHANCER OF GLABRA3 (EGL3) specify epidermal cell fate in the Arabidopsis root. Development 130:6431-6439
Bibikova TN, Blancaflor EB, Gilroy S.1999. Microtubules regulate tip growth and orientation in root hairs of Arabidopsis thaliana. Plant J 17:657-665
Bohme K, Li Y, Charlot F, Grierson C, Marrocco K, et al.2004. The Arabidopsis COW1 gene encodes a phosphatidylinositol transfer protein essential for root hair tip growth.Plant J 40:686-698
Bouzarelou D, Billini M, Roumelioti K, Sophianopoulou V.2008. EglD, a putative endoglucanase, with an expansin like domain is localized in the conidial cell wall of Aspergillus nidulans. Fungal Genet Biol 45:839-850
Brotman Y, Briff E, Viterbo A, Chet I.2008. Abstract Role of swollenin, an expansin-like protein from Trichoderma, in plant root colonization. Plant Physiol 147:779-789.
Brummell DA, Harpster MH, Civello PM, Palys JM, Bennett AB, and Dunsmuir P.1999. Modification of expansin protein abundance in tomato fruit alters softening and cell wall polymer metabolism during ripening. Plant Cell 11:2203-2216
Brummell, DA, Harpster, MH, Dunsmuir, P.1999. Differential expression of expansin gene family members during growth and ripening of tomato fruit. Plant Mol Biol 39:161-169
Brummell DA, Harpster MH.2001. Cell wall metabolism in fruit softening and quality and its manipulation in transgenic plants. Plant Mol Biol 47:311-340
Brummell,D.A., Howie,W.J., Ma,C., and Dunsmuir,P.2002. Postharvest fruit quality of transgenic tomatoes suppressed in expression of a ripening-related expansin. Postharvest Biology and Technology 25:209-220
Carey RE, Cosgrove DJ.2007. Portrait of the expansin superfamily in Physcomitrella patens: comparisons with angiosperm expansins. Ann Bot (Lond) 99:1131-1141
Carol RJ, Takeda S, Linstead P, Durrant MC, Kakesova H, et al.2005. A RhoGDP dissociation inhibitor spatially regulates growth in root hair cells. Nature 438:1013-1016
Catala C, Rose JK, Bennett AB.2000. Auxin-regulated genes encoding cell wall-modifying proteins are expressed during early tomato fruit growth. Plant Physiol 122:527-534
Cleves AE, McGee TP, Whitters EA, Champion KM, Aitken JR, et al.1991. Mutations in the CDP-choline pathway for phospholipid biosynthesis bypass the requirement for an essential phospholipid transfer protein. Cell 64:789-800
Clowes FAL.2000. Pattern in root meristem development in angiosperms. New Phytol 146:83-94
Chen F, Bradford KJ.2000. Expression of an expansin is associated with endosperm weakening during tomato seed germination. Plant Physiol 124:1265-1274
Chen F, Dahal P, Bradford KJ.2001. Two Tomato Expansin Genes Show Divergent Expression and Localization in Embryos during Seed Development and Germination. Plant Physiol 127:928-936
Cho HT, Cosgrove DJ.2002. Regulation of root hair initiation and expansin gene expression in Arabidopsis. Plant Cell 14:3237-3253
Cho HT, Cosgrove DJ.2002. Regulation of root hair initiation and expansin gene expression in Arabidopsis. Plant Cell 14:3237-3253
Cho HT, Kende H.1997a. Expressiongn of expansin gene is correlated with growth in deepwater rice. Plant Cell 9:1661-1671
Cho HT, Kende H.1997b. Expansins and internodalgrowth of deepwater rice. Plant Physiol 113:1145-1151
Cho HT, Kende H.1998. Tissue localization of expansins in deepwater rice. Plant J 15:805-812
Choi DS, Lee Y, Cho HT, KendeH.2003. Regulation of expansin gene expression affects growth and development in transgenic rice plants. Plant Cell 15:1386-1398
Civello PM, Powell ALT, Sabehat A, Bennett AB.1999. An expansin gene expressed in ripening strawberry fruit. Plant Physiol 121:1273-1279
Cosgrove DJ.1989. Characterization of long-term extension of isolated cell walls from growing cucumber hypocotyls. Planta 177:121-130
Cosgrove DJ.2000. Expansive growth of plant cells. Plant Physiolo Biochem 38:109-124
Cosgrove DJ, Bedinger P, Durchko DM.1997.Group I allergens of grass pollen as cell wall-loosening agents. Proc Natl Acad Sci 94:6559-6564
Cosgrove DJ, Durachko DM.1994. Autolysis and extension of isolated walls from growing cucumber hypocotyls. J Exp Bot 45:1711-1719
Cosgrove DJ, Li LC, Cho HT, Hoffmann-Benning, S Moore, RC, et al.2002. The growing world of expansins. Plant Cell Physiol 43:1436-1444
Costa S, Shaw P.2006. Chromatin organization and cell fate switch respond to positional information in Arabidopsis. Nature 439:493:496
Darby NJ, Morin PE, Talbo G, et al.1995. Refolding of bovine pancreatic trypsin inhibitor via non-native disulphide intermediates. JMol Biol 249:463-477
Desbrosses G, Josefsson C, Rigas S, Hatzopoulos P, Dolan L.2003. AKT1 and TRH1 are required during root hair elongation in Arabidopsis. J Exp Bot 54:781-88
Desbrosses G, Josefsson C, Rigas S, Hatzopoulos P, Dolan L.2003. AKT1 and TRH1 are required during root hair elongation in Arabidopsis. JExp Bot 54:781-788
Ding WN, Yu ZM, Tong YL, Huang W, Chen HM, Wu P.2009. A transcription factor with a bHLH domain regulates root hair development in rice. Cell Res 19:1309-1311
Dolan L, Duckett C, Grierson C, Linstead P, Schneider K, Lawson E, Dean C, Poethig RS, Roberts K.1994. Clonal relations and patterning in the root epidermis of Arabidopsis. Development 120:2465-2474
Downes BP, Steinbaker CR, Crowell DN.2001. Expression and Processing of a Hormonally Regulated beta-Expansin from Soybean. Plant Physiol 126:244-252
Durachko DM, Cosgrove DJ.2009. Measuring plant cell wall extension (creep) induced by acidic pH and by alpha-expansin. JOVE 25:1-4
Esch JJ, Chen M, Sanders M, Hillestad M, Ndkium S, et al.2003. A contradictory GLABRA3 allele helps define gene interactions controlling trichome development in Arabidopsis. Development 130:5885-5894
Favery B, Ryan E, Foreman J, Linstead P, Boudonck K, Steer M, Shaw P, Dolan L.2001. KOJAKencodes a cellulose synthase-like protein required or root haircell morphogenesis in Arabidopsis.Genes Dev 15:79-89
Fleming AJ, McQueen-Mason S, Mandel T, Kuhlemeier C.1997. Induction of leaf primordia by the cell wall protein expansin. Science 276:1415-1418
Foreman J, Demidchik V, Bothwell JH, Mylona P, Miedema H,et al.2003.Reactive oxygen species produced by NADPH oxidase regulate plant cell growth. Nature 422:442-46
Gal TZ, Aussenberg ER, Burdman S, Kapulnik Y, Koltai H.2006. Expression of a plant expansin is involved in the establishment of root knot nematode parasitism in tomato. Planta 224:155-162
Galway ME, Masucci JD, Lloyd AM, Walbot V, Davis RW, Schiefelbein JW.1994. The TTG gene is required to specify epidermal cell fate and cell patterning in the Arabidopsis root. Dev Biol 166:740-754
Gao Q, Guo QF, Xing SC, Zhao MR, Li F, Wang W.2007. The characteristics of expansins in wheat coleoptiles and their responses to water stress. Zhi Wu Sheng Li Yu Fen Zi Sheng Wu Xue Xue Bao 33:402-410
Grebe M, Friml J, Swarup R, Ljung K, Sandberg G, et al.2002. Cell polarity signaling in Arabidopsis involves a BFA-sensitive auxin influx pathway. Curr Biol 12:329-334
Greenwood MS, Xu FY, Hutchison KW.2006. The role of auxin-induced peaks of a-expansin expression during lateral root primordium formation in Pinus taeda. Physiologia Plantarum 126:279-288.
Grierson CS, Roberts K, Feldmann KA,Dolan L.1997.The COW1 locus of Arabidopsis acts after RHD2, and in parallel with RHD3 and TIP1, to determine the shape, rate of elongation, and number of root hairs produced from each site of hair formation. Plant Physiol 115:981-90
Guilhem D, Josefsson C, Rigas S, Hatzopoulos P, Dolan L.2003. AKT1 and TRH1 are required during root hair elongation in Arabidopsis. J Exp Bot 54:781-788
Harmer SE, Orford SJ, and Timmis JN.2002. Characterisation of six alpha-expansin genes in Gossypium hirsutum (upland cotton). Mol Genet Genomics 268:1-9
Harrison EP, McQueen-Mason SJ, Manning K.2001. Expression of six expansin genes in relation to extension activity in developing strawberry fruit. J Exp Bot 52:1437-46
Hochholdinger F, Wen TJ, Zimmermann R, Chimot-Marolle P, Silva OC, Bruce W, Lamkey KR, Wienand U, Schnable PS.2008. The maize (Zea mays L.) roothairless3 gene encodes a putative GPI-anchored, monocot-specific, COBRA-like protein that significantly affects grain yield. Plant J 54:888-898
Hoffmann-Benning S, Klomparens KL, and Kende H.1994. Characterization of growth-related osmiophilic particles in corn coleoptiles and deepwater rice internodes. Ann Bot 74:563-572
Huang J, Takano T, Akita S.2000. Expression of alpha-expansin genes in young seedlings of rice (Oryza sativa L.). Planta 211:467-473
Initiative AG.2000. Analysis of the genome sequence of the flowering plant Arabidopsis thaliana. Nature 408:796-815
Ishida T, Kurata T, Okada K, Wada T.2008. A genetic regulatory network in the development of trichomes and root hairs. Annu Rev Plant Biol 59:365-386
Jin Y, Tashpulatov AS, Katholnigg H, Heberle-Bors E, Touraev A.2006. Isolation and characterisation of two wheat beta-expansin genes expressed during male gametophyte development. Protoplasma 228:13-9
Jones MA,Raymond MJ, Yang Z, Smirnoff N.2007. NADPH oxidase-dependent reactive oxygen species formation required for root hair growth depends on ROP GTPase. J Exp Bot 58:1261-70
Jones MA, Shen JJ, Fu Y, LiH, Yang Z, GriersonCS.2002.The Arabidopsis Rop2GTPase is a positive regulator of both root hair initiation and tip growth. Plant Cell 14:763-76
Kende H, Bradford K, Brummell D, Cho HT, Cosgrove DJ, Fleming A, Gehring C, Lee Y, McQueen-Mason SM, Rose J, Voesenek LA.2004 Nomenclature for members of the expansin superfamily of genes and proteins. Plant Mol.Biol 55:311-314
Kerff F, Amoroso A, Herman R, Sauvage E, Petrella S, Filee P, Charlier P, Joris B, Tabuchi A, Nikolaidis N, Cosgrove DJ.2008. Crystal structure and activity of Bacillus subtilis YoaJ (EXLX1), a bacterial expansin that promotes root colonization. Proc Natl Acad Sci 105:16876-81
Ketelaar T, Allwood EG, Anthony R, Voigt B, Menzel D, Hussey PJ.2004. The actin-interacting protein AIP1 is essential for actin organization and plant development. Curr Biol 14:145-49
Ketelaar T, Faivre-Moskalenko C, Esseling JJ, de Ruijter NC, Grierson CS, et al. 2002.Positioning of nuclei in Arabidopsis root hairs:an actin-regulated process of tip growth. Plant Cell 14:2941-55
Kim CM, Park SH, Je BI, Park SH, Park SJ, et al.2007. OsCLSDl,a cellulose synthase-like D1 gene, is required for root hair morphogenesis in rice. Plant Physiol 143:1220-1230
Kim ES, Lee HJ, Bang WG, Choi IG, Kim KH.2009. Functional Characterization of a bacterial expansin from Bacillus subtilis for enhanced enzymatic hydrolysis of cellulos. Biotechnol Bioeng 102:1342-1353
Kim JH, Cho HT, Kende H.2000. Alpha-expansins in the semiaquatic ferns Marsilea quadrifolia and Regnellidium diphyllum:evolutionary aspects and physiological role in rachis elongation. Planta 212:85-92
Kirik V, Simon M, Huelskamp M, Schiefelbein J.2004. The ENHANCER OF TRY AND CPC1 gene acts redundantly with TRIPTYCHON and CAPRICE in trichome and root hair cell patterning in Arabidopsis. Dev Biol 268:506-513
Koshino-Kimura Y, Wada T, Tachibana T, Tsugeki R, Ishiguro S, Okada K.2005. Regulation of CAPRICE transcription by MYB proteins for root epidermis differentiation in Arabidopsis. Plant Cell Physiol 46:817-826
Kurata T, Ishida T, Kawabata-Awai C, Noguchi M, Hattori S, et al.2005. Cell-to- cell movement of the CAPRICE protein in Arabidopsis root epidermal cell differentiation. Development 132:5387-98
Kwak SH, Schiefelbein J.2006. The role of the SCRAMBLED receptor-like kinase in patterning the Arabidopsis root epidermis. Dev Biol 302:118-131
Kwak SH, Shen R, Schiefelbein J.2005. Positional signalling mediated by a receptor-like kinase in Arabidopsis. Science 307:1111-1113
Kwasniewski M, Szarejko I.2006. Molecular cloning and characterization of beta-expansin gene related to root hair formation in barley. Plant Physiol 141:1149-1158
Kwon YR, Lee HJ, Kim KH, Hong SW, Lee SJ, Lee H.2008. Ectopic expression of Expansin3 or Expansinbetal causes enhanced hormone and salt stress sensitivity in Arabidopsis. Biotechnol Lett 30:1281-1288
Lee DK, Ahn JH, Song SK, Choi YD, Lee JS.2003. Expression of an expansin gene is correlated with root elongation in soybean. Plant Physiol 131:985-997
Lee MM, Schiefelbein J.1999. WEREWOLF, a MYB-related protein in Arabidopsis, is a position-dependent regulator of epidermal cell patterning. Cell 99:473-483
Lee MM, Schiefelbein J.2002. Cell pattern in the Arabidopsis root epidermis determined by lateral inhibition with feedback. Plant Cell 14:611-618
Lee Y, Choi D.2005. Biochemical properties and localization of the beta-expansin OsEXPB3 in Rice(Oryza Sativa L.). Mol Cells 20:119-126
Lee Y, Kende H.2001. Expression of beta-Expansins Is Correlated with Internodal Elongation in Deepwater Rice. Plant Physiol 127:645-655
Li LC,Bedinger PA, Volk C, Jones AD, Cosgrove DJ.2003. Purification and characterization of four beta-expansins (Zea m 1 isoforms) from maize pollen. Plant Physiol 132:2073-2085
Li XX, Duan XP, Jiang HX, et al.2006. Genome-wide analysis of basic/helix-loop-helix transcription factor family in rice and Arabidopsis. Plant Physiol 141:1167-1184
Li Y, Darley CP, Ongaro V, et al.2002. Plant expansins are a complex multigen familiy with an ancient evolutionary origin. Plant Physiol 128:854-864
Lin Z, Ni Z, Zhang Y, Yao Y, Wu H, Sun Q.2005. Isolation and characterization of 18 genes encoding alpha- and beta-expansins in wheat (Triticum aestivum L.). Mol Genet Genomics 274:548-556
Link BM, Cosgrove DJ.1998. Acid-Growth Response and alpha-Expansins in Suspension Cultures of Bright Yellow 2 Tobacco. Plant Physiol 118:907-916
Ludidi NN, Heazlewood JL, Seoighe C, Irving HR, Gehring CA.2002. Expansin-like molecules: Novel functions derived from common domains. J Mol Evol 54:587-594
Masucci JD, Schiefelbein JW.1994.The rhd6mutation of Arabidopsis thaliana alters root-hair initiation through an auxin- and ethylene-associated process. Plant Physiol 106:1335-1346
Masucci JD, Schiefelbein JW.1996. Hormones act downstream of TTG and GL2 to promote root hair outgrowth during epidermis development in the Arabidopsis root. Plant Cell 8:1505-1517
McQueen-Mason S, Durachko DM, Cosgrove DJ.1992. Two endogenous proteins that induce cell wall expansion in plants. Plant Cell 4:1425-1433
McQueen-Mason SJ, Fry SC, Durachko DM, Cosgrove DJ.1993. The relationship between xyloglucan endotrans-glycosylase and in vitro cell wall extension in cucumber hypocotgyls. Planta 190:327-331
Meijer HJ, Munnik T.2003. Phospholipid-based signaling in plants. Annu Rev Plant Biol 54:265-306
Menand B, Yi K, Jouannic S, Hoffmann L, Ryan E, et al.2007. An ancient mechanism controls the development of cells with a rooting function in land plants. Science 316:1477-1480
Molendijk AJ, Bischoff F, Rajendrakumar CS, Friml J, Braun M, et al.2001. Arabidopsis thaliana Rop GTPases are localized to tips of root hairs and control polar growth. EMBO J 20:2779-27788
Nishitani K, Tominaga R.1992. Endo-xyloglucan transferase, a novel class of glycosyltransferase that catalyzes transfer of a segment of xyloglucan molecule to another xyloglucan molecule. JBiol Chem 267:21058-21064
Nishimura T, Yokota E, Wada T, Shimmen T, Okada K.2003. An Arabidopsis ACT2 dominant-negative mutation, which disturbs F-actin polymerization, reveals its distinctive function in root development. Plant Cell Physiol 44:1131-1140
O'Malley RC, Lynn DG.2000. Expansin message regulation in parasitic angiosperms:Marking time in development. Plant Cell 12:1455-1466
Ohashi Y, Oka A, Rodrigues-Pousada R, Possenti M, Ruberti I, et al.2003.Modulation of phospholipid signaling by GLABRA2 in root-hair pattern formation. Science 300:1427-1430
Ono S.2003.Regulation of actin filament dynamics by actin depolymerizing factor/cofilin and actin-interacting protein 1:new blades for twisted filaments. Biochemistry 42:13363-13370
Orford SJ, Timmis JN.1998. Specific expression of an expansin gene during elongation of cotton fibres. Biochim Biophys Acta 1398:342-346
Pesch M, Hulskamp M.2004. Creating a two-dimensional pattern de novo during Arabidopsis trichome and root hair initiation. Curr Opin Genet Dev 14:422-427
Pien S, Wyrzykowska J, McQueen-Mason S, Smart C, Fleming A.2001. Local expression of expansin induces the entire process of leaf development and modifies leaf shape. Proc Nat Acad Sci 98:11812-11817
Qin L, Kudla U, Roze EH, Goverse A, Popeijus H, Nieuwland J, Overmars H, Jones JT, et al. 2004. Expansin proteins, which have so far been identified only in plants, rapidly induce extension of plant cell walls by weakening the non-covalent interactions that help to maintain their integrity. Nature 427:30
Ramachandran S, Christensen HE, Ishimaru Y, Dong CH, Chao-Ming W, et al.2000.Profilin plays a role in cell elongation, cell shapemaintenance, and flowering in Arabidopsis. Plant Physiol 124:1637-1647
Rentel MC, Lecourieux D, Ouaked F, Usher SL, Petersen L, et al.2004. OXI1 kinase is necessary for oxidative burst-mediated signalling in Arabidopsis. Nature 427:858-861
Rerie WG, Feldmann KA, Marks MD.1994. The GLABRA2 gene encodes a homeo domain protein required for normal trichome development in Arabidopsis. Genes Dev 8:1388-1399
Ridge RW, Uozumi Y, Plazinski J, Hurley UA, Williamson RE.1999. Developmental transitions and dynamics of the cortical ER of Arabidopsis cells seen with green fluorescentprotein. Plant Cell Physiol 40:1253-1261
Rigas S, Debrosses G, Haralampidis K, Vicente-Agullo F, Feldmann KA, et al.2001. TRH1 encodes a potassium transporter required for tip growth in Arabidopsis root hairs. Plant Cell 13:139-151
Rochange SF, Wenzel CL, McQueen-Mason SJ.2001. Impaired growth in transgenic plants over-expressing an expansin isoform. Plant Mol Biol 46:581-589
Rose JKC, Lee HH, Bennett AB.1997. Expression of a divergent expansin gene is fruit-specific and ripening-regulated. Proc Nat Acad Sci 94:5955-5960
Row HC, Reeder JR.1957. Root-hair development as evidence of relationships among genera of gramineae. Am J Bot 44:596-601
Ruan Y, Llewellyn D, Furbank R.2001. The control of single-celled cotton fiber elongation by developmentally reversible gating of plasmodesmata and coordinated expression of sucrose and K+ transporters and expansin. Plant Cell 13:47-60
Ryu KH, Kang YH, Park YH, Hwang I, Schiefelbein J, Lee MM.2005. The WEREWOLF MYB protein directly regulates CAPRICE transcription during cell fate specification in the Arabidopsis root epidermis. Development 132:4765-4775
Sabirzhanova IB, Sabirzhanov BE, Chemeris AV, Veselov DS, Kudoyarova GR.2005. Fast changes in expression of expansin gene and leaf extensibility in osmotically stressed maize plants. Plant Physiol Biochem 43:419-422
Saloheimo M, Paloheimo M, Hakola S, Pere J, Swanson B, Nyyssonen E, Bhatia A, Ward M, Penttila M. 2002. Swollenin, a Trichoderma reesei protein with sequence similarity to the plant expansins, exhibits disruption activity on cellulosic materials. Eur J Biochem 269:4202-4211
Samaj J, Baluska F, Menzel D.2004. New signalling molecules regulating root hair tip growth. Trends Plant Sci 9:217-20
Samaj J, Ovecka M, Hlavacka A, Lecourieux F, Meskiene I, et al.2002. Involvement the mitogen-activated protein kinase SIMK in regulation of root hair tip growth. EMBO J 21:3296-306
Sampedro J, Cosgrove DJ. 2005. Genome biology the expansin superfamily. Genome Biol 6:242
Sampedro J, Carey RE, and Cosgrove DJ. 2006. Genome histories clarify evolution of the expansin superfamily:new insights from the poplar genome and pine ESTs. J Plant Res 119:11-21
Saloheimo M, Paloheimo M, Hakola S, Pere J, Swanson B, Nyyssonen E, Bhatia A, Ward M, Penttila M. 2002. Swollenin, a Trichoderma reesei protein with sequence similarity to the plant expansins, exhibits disruption activity on cellulosic materials. Eur J Biochem 269:4202-4211
Schellmann S, Schnittger A, Kirik V, Wada T, Okada K, Beermann A, Thumfahrt J, Jurgens G, Hulskamp M. 2002. TRIPTYCHON and CAPRICE mediate lateral inhibition during trichome and root hair patterning in Arabidopsis. EMBO J 21:5036-5046.
Schiefelbein JW.1992. Calcium influx at the tip of growing root-hair cells of Arabidopsis. Planta 187:455-59
Schiefelbein J.2003. Cell-fate specification in the epidermis:a common patterning mechanism in the root and shoot. Curr Opin Plant Biol 6:74-78
Schiefelbein J, Kwak SH, Wieckowski Y, Barron C, Bruex A.2009. The gene regulatory network for root epidermal cell-type pattern formation in Arabidopsis. J Exp Bot 60:151-1521
Schiefelbein JW, Somerville C.1990. Genetic control of root hair development in Arabidopsis thaliana. Plant Cell 2:235-243
Schipper O, Schaefer D, Reski R, Fleming A.2002. Expansins in the bryophyte Physcomitrella patens. Plant Mol Biol 50:789-802
Schultz SC, Dalbadie MG, Neitzel JJ, et al.1987. Stability of wild type and mutant RTEM-1 beta-lactamases:effect of the disulfide bond. Protein 2:290-297
Seifert GJ, Barber C, Wells B, Dolan L, Roberts K.2002. Galactose biosynthesis in Arabidopsis: genetic evidence for substrate channeling from UDP-D-galactose into cell wall polymers. Curr Biol 12:1840-1845
Shi YH, Zhu SW, Mao XZ, Feng JX, Qin YM, Zhang L, Cheng J, Wei LP, Wang ZY, Zhu YX. 2006. Transcriptome profiling, molecular biological, and physiological studies reveal a major role for ethylene in cotton fiber cell elongation. Plant Cell 18:651-664
Shin JH, Jeong DH, Park MC, An G.2005. Characterization and transcriptional expression of the alpha-expansin gene family in rice. Mol Cells 20:210-218
Simon M, Lee MM, Lin Y, Gish L, Schiefelbein J.2007. Distinct and overlapping roles of single-repeat MYB genes in root epidermal patterning. Dev Biol 311:566-578
Shcherban TY, Shi J, Durachko DM, Guiltinan MJ, McQueen-Mason S, Shieh M, Cosgrove DJ. 1995. Molecular cloning and sequence analysis of expansins-A highly conserved, multigene family of proteins that mediate cell wall extension in plants. Proc Natl Acad Sci 92:9245-9249
Sohn RH, Goldschmidt-Clermont PJ.1994. Profilin:at the crossroads of signal transduction and the actin cytoskeleton. BioEssays 16:465-472
Tang HB, Bowers JE, Wang XY, Ming R, Alam M, Peterson AH.2008. Synteny and Collinearity in Plant Genomes. Science 320:486-488
Thompson JE, Fry SC.2001. Restructuring of wall-bound xyloglucan by transglycosy-lation in living plant cells. Plant J 26:23-34
Tominaga R, Iwata M, Okada K, Wada T.2007. Functional analysis of the epidermal-specific MYB genes CAPRICE and WEREWOLF in Arabidopsis. Plant Cell 19:2264-2277
Torres MA, Onouchi H, Hamada S, Machida C, Hammond-Kosack KE, Jones JD.1998. Six Arabidopsis thaliana homologues of the human respiratory burst oxidase (gp91phox). Plant J 14:365-370
Torii KU.2004. Leucine-rich repeat receptor kinases in plants:structure, function, and signal transduction pathways. Int Rev Cytol 234:1-46
Ueda M, Koshino-Kimura Y, Okada K.2005. Stepwise under-standing of root development. Curr Opin Plant Biol 8:71-76
Valdivia ER, Sampedro J, Lamb JC, Chopra S, and Cosgrove DJ.2007. Recent Proliferation and Translocation of Pollen Group 1 Allergen Genes in the Maize Genome. Plant Physiol 143:1269-1281
Valdivia ER, Wu Y, Li LC, Cosgrove DJ, Stephenson AG.2007. A Group-1 Grass Pollen Allergen Influences the Outcome of Pollen Competition in Maize. PLoS ONE 2:e154
Van BN, Joss G, Van OP.2004. Reorganization and in vivo dynamics of microtubules during Arabidopsis root hair development. Plant Physiol 136:3905-3919
Vincent P, Chua M, Nogue F, Fairbrother A, Mekeel H, et al.2005. A Sec14p-nodulin domain phosphatidylinositol transfer protein polarizes membrane growth of Arabidopsis thaliana root hairs. J Cell Biol 168:801-812
Vissenberg K, Fry SC, Verbelen JP.2001. Root hair initiation is coupled to a highly localized increase of xyloglucan endotransglycosylase action in Arabidopsis roots. Plant Physiol 127:1125-1135
Wada T, Tachibana T, Shimura Y, Okada K.1997. Epidermal cell differentiation in Arabidopsis determined by a Myb homolog, CPC. Science 277:1113-1116
Wada T, Kurata T, Tominaga R, Koshino-Kimura Y, Tachibana T, Goto K, Marks MD, Shimura Y, Okada K.2002. Role of a positive regulator of root hair development, CAPRICE, in Arabidopsis root epidermal cell differentiation. Development 129:5409-5419
Walker AR, Davison PA, Bolognesi-Winfield AC, James CM, Srinivasan N, Blundell TL, Esch JJ, Marks MD, Gray JC.1999. The TRANSPARENT TESTA GLABRA1 locus, which regulates trichome differentiation and anthocyanin biosynthesis in Arabidopsis,encodes a WD40 repeat protein. Plant Cell 11:1337-1350
Webb M, Jouannic S, Foreman J, Linstead P, Dolan L.2002. Cell specification in the Arabidopsis root epidermis requires the activity of ECTOPIC ROOT HAIR 3-a katanin p60 protein. Development 129:123-131
Wen TJ, Hochholdinger F, Sauer M, Bruce W, Schnable PS.2005. The root hairless1 gene of maize encodes a homolog of sec3, which is involved in polar exocytosis. Plant Physiol 138:1637-1643
Wieczorek K, Golecki B, Gerdes L, Heinen P, Szakasits D, Durachko DM, Cosgrove DJ, Kreil DP, Puzio PS, Bohlmann H, Grundler FM.2006. Expansins are involved in the formation of nematode-induced syncytia in roots of Arabidopsis thaliana. Plant J 48:98-112
Wu YJ, Thorne ET, Sharp RE. 2001. Modification of expansin transcript levels in the maize primary root at low water potentials. Plant Physiol 126:1471-1479
Wymer CL, Bibikova TN, Gilroy S.1997. Cytoplasmic free calciumdistributions during the development of root hairs of Arabidopsis thaliana. Plant J 12:427-439
Xu CR, Liu C, Wang YL, Li LC, Chen WQ, Xu ZH, Bai SN.2005.Histone acetylation affects expression of cellular patterning genes in the Arabidopsis root epidermis. Proc Natl Acad Sci 102:14469-14474
Xu J, Scheres B.2005. Dissection of Arabidopsis ADP-RIBOSYLATION FACTOR 1 function in epidermal cell polarity. Plant Cell 17:525-536
Yang Z. 2002. Small GTPases:versatile signaling switches in plants. Plant Cell 14:375-388
Yennawar NH, Li LC, Dudsinski DM, Tabuchi A, Cosgrove DJ. 2006. Crystal structure and activities of EXPB1 (Zea m l), a-expansin and group-1 pollen allergen from maize. Proc Natl Acad Sci 103:14664-14671
Yi L, Jones L, McQueen-Mason S.2003. Expansins and cell growth. Curr Opin Plant Biol 6:603-610
Yoshida S, Forno DA, Cock JH, Gomez KA.1976. Laboratory Manual for Physiological Studies of Rice, Ed 3. The International Rice Research Institute, Manila, The Philippines
Yuo T, Toyota M, Ichii M, Taketa S.2009. Molecular cloning of a root hairless gene rthl in rice. Breeding science.59:13-20
Zenoni S, Reale L, Tornielli GB, Lanfaloni L, Porceddu A, Ferrarinin A, Moretti C, Zamboni A, Speghini A, Ferranti F, Pezzotti M.2004. Downregulation of the petunia hybrida alpha-expansin gene PhEXPl reduces the amount of crystalline cellulose in cell walls and leads to phenotypic changes in petal limbs. Plant Cell 16:295-308
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