植物蔗糖转运蛋白及其生理功能的研究进展
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摘要
高等植物通过光合作用产生蔗糖,并以蔗糖形式将同化碳源分配到植物各组织器官中,蔗糖的跨膜转运需要蔗糖转运蛋白的参与。目前,已有研究表明蔗糖转运蛋白广泛存在于高等植物体内,主要在种子、花器、叶肉细胞、韧皮部和根等器官中发挥作用,而通过亚细胞定位可以发现,蔗糖转运蛋白主要定位在细胞的液泡膜及细胞质膜上。综述了国内外对蔗糖转运蛋白的发现、结构、分类、生理功能、功能验证及定位等方面的研究内容,分析了研究蔗糖转运蛋白的意义及目前存在的一些问题,旨为更好地理解蔗糖转运蛋白在植物体内的作用机制做铺垫。
Sucrose is the primary product of photosynthetic CO_2 fixation which is used for the distribution of assimilated carbon withinhigher plants. The transmembrane transport of sucrose requires the participation of sucrose transporters. Currently,studies have shown thatsucrose transporter are widely present in higher plants and played main role in seed,flower,leaf cells,phloem,root and other organs. Bythe way of subcellular localization,sucrose transporters have been found mostly to be located in the cell membrane and vacuole membrane.In this paper,we review the progresses on different aspects of SUTs,such as the protein structure,classification,physiological functions,functional verification and subcellular localization;then we discussed the research significance and remained problems in studying sucrosetransporter for better understanding the action mechanism of sucrose transporter proteins in plants.
Sucrose is the primary product of photosynthetic CO_2 fixation which is used for the distribution of assimilated carbon withinhigher plants. The transmembrane transport of sucrose requires the participation of sucrose transporters. Currently,studies have shown thatsucrose transporter are widely present in higher plants and played main role in seed,flower,leaf cells,phloem,root and other organs. Bythe way of subcellular localization,sucrose transporters have been found mostly to be located in the cell membrane and vacuole membrane.In this paper,we review the progresses on different aspects of SUTs,such as the protein structure,classification,physiological functions,functional verification and subcellular localization;then we discussed the research significance and remained problems in studying sucrosetransporter for better understanding the action mechanism of sucrose transporter proteins in plants.
引文
[1]Reinders A,Schulze W,Kühn C,et al.Protein-protein interactionsbetween sucrose transporters of different affinities colocalized inthe same enucleate sieve element[J].Plant Cell,2002,14(7):1567-1577.
[2]Truernit E.Plant physiology:the importance of sucrosetransporters[J].Curr Biol,2001,11(5):169-171.
[3]Viola R,Roberts AG,Haupt S,et al.Tuberization in potato involvesa switch from apoplastic to symplastic phloem unloading[J].PlantCell,2001,13(2):385-398.
[4]Patrick JW.Phloem unloading:sieve element unloading and postsieve element transport[J].Annu Rev Plant Physiol Plant MolBiol,1997,48(1):191-222.
[5]Vaughn MW,Harrington GN,Bush DR.Sucrose-mediatedtranscriptional regulation of sucrose symporter activity in thephloem[J].Proc Natl Acad Sci USA,2002,99(16):10876-10880.
[6]Riesmeier JW,Willmitzer L,Frommer WB.Isolation and characterization of a sucrose carrier c DNA from spinach by functionalexpression in yeast[J].EMBO J,1992,11(13):4705-4713.
[7]The Arabidopsis Genome Initiative.analysis of the genome sequenceof the flowering plant Arabidopsis thaliana[J].Nature,2000,408(6814):796-815.
[8]Srivastava AC,Ganesan S,Ismail IO,et al.Functional characterization of the Arabidopsis At SUC2 sucrose/H1 symporter by tissuespecific complementation reveals an essential role in phloem loadingbut not in long distance transport[J].Plant Physiol,2008,148(1):200-211.
[9]高蕾,肖文芳,李文燕,等.拟南芥蔗糖转运蛋白(SUTs)的功能研究进展[J].分子植物育种,2011,9(2):251-255.
[10]Aoki N,Hirose T,Scofield GN,et al.The sucrose transporter genefamily in rice[J].Plant Cell Physiol,2003,44(3):223-232.
[11]阳江华,黄德宝,刘术金,等.巴西橡胶树6个蔗糖转运蛋白基因的克隆与序列分析[J].热带作物学报,2007,28(4):32-38.
[12]Shakya R,Sturm A.Characterization of source and sink specificsucrose/H+symporters from carrot[J].Plant Physiol,1998,118(4):1473-1480.
[13]Noiraud N,Delrot S,Lemoine R.The sucrose transporter of celery.Identification and expression during salt stress[J].Plant Physiol,2000,122(4):1447-1456.
[14]Bush DR.Proton-coupled sugar and amino acid transporters inplants[J].Annu Rev Plant Physiol Plant Mol Bio,1993,44(44):513-542.
[15]Lemoine R.Sucrose transporters in plants:update on function andstructure[J].Biochim Biophys Acta,2000,1465(1-2):246-262.
[16]Williams LE,Lemoine R,Sauer N,et al.Sugar transporters inhigher plants:a diversity of roles and complex regulation[J].Trends Plant Sci,2000,5(7):283-290.
[17]Weise A,Barker L,Kühn C,et al.A new subfamily of sucrosetransporters,SUT4,with low affinity/high capacity localized inenucleate sieve elements of plants[J].Plant Cell,2000,12(8):1345-1355.
[18]Ward JM,Kühn C,Tegeder M,et al.Sucrose transport in higherplants[J].Int Rev Cytol,1998,178(335):41-71.
[19]Hackel A,Schauer N,Carrari F,et al.Sucrose transporter Le SUT1and Le SUT2 inhibition affects tomato fruit development in differentways[J].Plant J,2006,45(2):180-192.
[20]Endler A,Meyer S,Schelbert S,et al.Identification of a vacuolarsucrose transporter in barley and Arabidopsis mesophyll cells by atonoplast proteomic approach[J].Plant Physiol,2006,141(1):196-207.
[21]Stadler R,Brandner J,Schulz A,et al.Phloem loading by thePm SUC2 sucrose carrier from Plantago major occurs intocompanion cells[J].Plant Cell,1995,7(10):1545-1554.
[22]Stadler R,Sauer N.The Arabidopsis thaliana At SUC2 gene isspecifically expressed in companion cells[J].Bot Acta,1996,109(4):299-306.
[23]Riesmeier JW,Willmitzer L,Frommer WB.Evidence for anessential role of the sucrose transporter in phloem loading andassimilate partitioning[J].EMBO J,1994,13(1):1-7.
[24]Kühn C,Quick WP,Schulz A,et al.Companion cell-specificinhibition of the potato sucrose transporter SUT1[J].Plant CellEnviron,1996,19(10):1115-1123.
[25]Burkle L,Hibberd JM,Quick WP,et al.The H+-sucrosecotransporter Nt SUT1 is essential for sugar export from tobaccoleaves[J].Plant Physiol,1998,118(1):59-68.
[26]Truernit E and Sauer N.The promoter of the Arabidopsis thalianaSUC2 sucrose-H1 symporter gene directs expression of b-glucuronidase to the phloem:evidence for phloem loading and unloadingby SUC2[J].Planta,1995,196(1):564-570.
[27]Lemoine R,Kühn C,Thiele NS,et al.Antisense inhibition of thesucrose transporter in potato:effects on amount and activity[J].Plant Cell Environ,1996,19(10):1124-1131.
[28]Meyer S,Lauterbach C,Niedermeier M,et al.Wounding enhancesexpression of At SUC3,a sucrose transporter from Arabidopsis sieveelements and sink tissues[J].Plant Physiol,2004,134(2):684-693.
[29]Weber H,Borisjuk L,Heim U,et al.A role for sucrose transportersduring seed development:molecular characterization of a hexoseand a sucrose carrier in fava bean seeds[J].Plant Cell,1997,9(6):895-908.
[30]Bick JA,Neelam A,Smith E,et al.Expression analysis of a sucrosecarrier in the germinating seedling of Ricinus communis[J].Plant Mol Biol,1998,38(3):425-435.
[31]Stadler R,Truernit E,Gahrtz M,et al.The At SUC1 sucrose carriermay represent the osmotic driving force for anther dehiscence andpollen tube grow thin Arabidopsis[J].Plant J,1999,19(3):269-278.
[32]Lemoine R,Burkle L,Barker L,et al.Identification of apollen-specific sucrose transporter-like protein Nt SUT3 fromtobacco[J].Febs Lett,1999,454(3):325-330.
[33]Sivitz AB,Reinders A,Ward JM,et al.Arabdopsis sucrose transporter At SUC1 is important for pollen germination and sucrose induced anthocyanin in accumulation[J].Plant Physiol,2008,147(1):92-100.
[34]Flemetakis E,Dimou M,Cotzur D,et al.A sucrose transporter,Lj SUT4 is up regulated during lotus japonicas nodule development[J].J Exp Bot,2003,54(388):1789-1791.
[35]Smeekens S,Rook F.Sugar sensing and sugar-mediated signaltransduction in plants[J].Plant Physiol,1997,115(1):7-13.
[36]Matsukura C,Saitoh T,Hirose T,et al.Sugar uptake and transportin rice embryo.Expression of companion cellspecific sucrosetransporter(Os SUT1)induced by sugar and light[J].PlantPhysiol,2000,124(1):85-93.
[37]Chiou TJ,Bush DR.Sucrose is a signal molecule in assimilatepartitioning[J].Proc Natl Acad Sci USA,1998,95(8):4784-4788.
[38]Ransom-Hodgkins WD,Vaughn MW,et al.Protein phosphorylationplays a key role in sucrose-mediated transcriptional regulation of aphloem-specific proton-sucrose symporter[J].Planta,2003,217(3):483-489.
[39]Müller J,Aeschbacher RA,Sprenger N,et al.Disaccharidemediated regulation of sucrose fructan-6-fructosyl transferase,a keyenzyme of fructan synthesis in barley leaves[J].Plant Physiol,2000,123(1):265-273.
[40]Gabriel-Neumann E,Neumann G,Leggewie G,et al.Constitutiveoverexpression of the sucrose transporter So SUT1 in potato plantsincreases arbuscular mycorrhiza fungal root colonization underhigh,but not under low,soil phosphorus availability[J].J PlantPhysiol,2011,168(9):911-919.
[41]Zhang YL,Meng QY,Qu GQ,et al.The antisence expression of aputative grape sucrose transporter in tobacco and effects on growthof the transformants[J].Prog Biochem Biophys,2006,7(7):699-705.
[42]Bitterlich M,Krügel U,Boldt-Burisch K,et al.The sucrose transporter Sl SUT2 from tomato interacts with brassinosteroid functioningand affects arbuscular mycorrhiza formation[J].Plant J,2014,78(5):877-889.
[43]Weichert N,Saalbach I,Weichert H,et al.Increasing sucroseuptake capacity of wheat grains stimulates storage proteinsynthesis[J].Plant Physiol,2010,152(2):698-710.
[44]Leggewie G,Kolbe A,Lemoine R,et al.Overexpression of thesucrose transporter So SUT1 in potato results in alterations in leafcarbon partitioning and in tuber metabolism but has little impact ontuber morphology[J].Planta,2003,217(1):158-167.
[45]Rosche E,Blackmore D,Tegeder M,et al.Seed-specificoverexpression of a potato sucrose transporter increases sucroseuptake and growth rates of developing pea cotyledons[J].Plant J,2002,30(2):165-175.
[46]Li H,Zhou SY,Zhao WS,et al.A novel wall-associated receptorlike protein kinase gene,Os WAK1,plays important roles in riceblast disease resistance[J].Plant Mol Biol,2009,69(3):337-346.
[47]Brandizzi F,Frangne N,Marc-Martin S,et al.The destinationfor single-pass membrane proteins is influenced markedly by thelength of the hydrophobic domain[J].Plant Cell,2002,14(5):1077-1092.
[48]Friedrichsen DM,Joazeiro CA,Li J,et al.Brassinosteroidinsensitive-1 is a ubiquitously expressed leucine-rich repeatreceptor serine/threonine kinase[J].Plant Physiology,2000,123(4):1247-1256.
[49]邱礽,陶刚,李奇科,等.农杆菌渗入法介导的基因瞬时表达技术及应用[J].分子植物育种,2009,7(5):1032-1039.
[50]庞建周.小麦干旱胁迫应答基因Ta SUT1的克隆、表达与基因转化研究[D].河北:河北师范大学,2011.
[51]高志民,杨学文,彭镇华,等.绿竹Bo SUT2基因的分子特征与亚细胞定位[J].林业科学,2010,46(2):45-50.
[52]Fan RC,Peng CC,Xu YH,et al.Apple sucrose transporter SUT1and sorbitol transporter SOT6 interact with cytochrome b5 toregulate their affinity for substrate sugars[J].Plant Physiol,2009,150(4):1880-1901.
[53]Sivitz AB,Reinders A,Johnson ME,et al.Arabidopsis sucrosetransporter At SUC9,high-affinity transport activity,intrageniccontrol of expression,and Early Flowering Mutant Phenotype[J].Plant Physiol,2007,143(1):188-198.
[2]Truernit E.Plant physiology:the importance of sucrosetransporters[J].Curr Biol,2001,11(5):169-171.
[3]Viola R,Roberts AG,Haupt S,et al.Tuberization in potato involvesa switch from apoplastic to symplastic phloem unloading[J].PlantCell,2001,13(2):385-398.
[4]Patrick JW.Phloem unloading:sieve element unloading and postsieve element transport[J].Annu Rev Plant Physiol Plant MolBiol,1997,48(1):191-222.
[5]Vaughn MW,Harrington GN,Bush DR.Sucrose-mediatedtranscriptional regulation of sucrose symporter activity in thephloem[J].Proc Natl Acad Sci USA,2002,99(16):10876-10880.
[6]Riesmeier JW,Willmitzer L,Frommer WB.Isolation and characterization of a sucrose carrier c DNA from spinach by functionalexpression in yeast[J].EMBO J,1992,11(13):4705-4713.
[7]The Arabidopsis Genome Initiative.analysis of the genome sequenceof the flowering plant Arabidopsis thaliana[J].Nature,2000,408(6814):796-815.
[8]Srivastava AC,Ganesan S,Ismail IO,et al.Functional characterization of the Arabidopsis At SUC2 sucrose/H1 symporter by tissuespecific complementation reveals an essential role in phloem loadingbut not in long distance transport[J].Plant Physiol,2008,148(1):200-211.
[9]高蕾,肖文芳,李文燕,等.拟南芥蔗糖转运蛋白(SUTs)的功能研究进展[J].分子植物育种,2011,9(2):251-255.
[10]Aoki N,Hirose T,Scofield GN,et al.The sucrose transporter genefamily in rice[J].Plant Cell Physiol,2003,44(3):223-232.
[11]阳江华,黄德宝,刘术金,等.巴西橡胶树6个蔗糖转运蛋白基因的克隆与序列分析[J].热带作物学报,2007,28(4):32-38.
[12]Shakya R,Sturm A.Characterization of source and sink specificsucrose/H+symporters from carrot[J].Plant Physiol,1998,118(4):1473-1480.
[13]Noiraud N,Delrot S,Lemoine R.The sucrose transporter of celery.Identification and expression during salt stress[J].Plant Physiol,2000,122(4):1447-1456.
[14]Bush DR.Proton-coupled sugar and amino acid transporters inplants[J].Annu Rev Plant Physiol Plant Mol Bio,1993,44(44):513-542.
[15]Lemoine R.Sucrose transporters in plants:update on function andstructure[J].Biochim Biophys Acta,2000,1465(1-2):246-262.
[16]Williams LE,Lemoine R,Sauer N,et al.Sugar transporters inhigher plants:a diversity of roles and complex regulation[J].Trends Plant Sci,2000,5(7):283-290.
[17]Weise A,Barker L,Kühn C,et al.A new subfamily of sucrosetransporters,SUT4,with low affinity/high capacity localized inenucleate sieve elements of plants[J].Plant Cell,2000,12(8):1345-1355.
[18]Ward JM,Kühn C,Tegeder M,et al.Sucrose transport in higherplants[J].Int Rev Cytol,1998,178(335):41-71.
[19]Hackel A,Schauer N,Carrari F,et al.Sucrose transporter Le SUT1and Le SUT2 inhibition affects tomato fruit development in differentways[J].Plant J,2006,45(2):180-192.
[20]Endler A,Meyer S,Schelbert S,et al.Identification of a vacuolarsucrose transporter in barley and Arabidopsis mesophyll cells by atonoplast proteomic approach[J].Plant Physiol,2006,141(1):196-207.
[21]Stadler R,Brandner J,Schulz A,et al.Phloem loading by thePm SUC2 sucrose carrier from Plantago major occurs intocompanion cells[J].Plant Cell,1995,7(10):1545-1554.
[22]Stadler R,Sauer N.The Arabidopsis thaliana At SUC2 gene isspecifically expressed in companion cells[J].Bot Acta,1996,109(4):299-306.
[23]Riesmeier JW,Willmitzer L,Frommer WB.Evidence for anessential role of the sucrose transporter in phloem loading andassimilate partitioning[J].EMBO J,1994,13(1):1-7.
[24]Kühn C,Quick WP,Schulz A,et al.Companion cell-specificinhibition of the potato sucrose transporter SUT1[J].Plant CellEnviron,1996,19(10):1115-1123.
[25]Burkle L,Hibberd JM,Quick WP,et al.The H+-sucrosecotransporter Nt SUT1 is essential for sugar export from tobaccoleaves[J].Plant Physiol,1998,118(1):59-68.
[26]Truernit E and Sauer N.The promoter of the Arabidopsis thalianaSUC2 sucrose-H1 symporter gene directs expression of b-glucuronidase to the phloem:evidence for phloem loading and unloadingby SUC2[J].Planta,1995,196(1):564-570.
[27]Lemoine R,Kühn C,Thiele NS,et al.Antisense inhibition of thesucrose transporter in potato:effects on amount and activity[J].Plant Cell Environ,1996,19(10):1124-1131.
[28]Meyer S,Lauterbach C,Niedermeier M,et al.Wounding enhancesexpression of At SUC3,a sucrose transporter from Arabidopsis sieveelements and sink tissues[J].Plant Physiol,2004,134(2):684-693.
[29]Weber H,Borisjuk L,Heim U,et al.A role for sucrose transportersduring seed development:molecular characterization of a hexoseand a sucrose carrier in fava bean seeds[J].Plant Cell,1997,9(6):895-908.
[30]Bick JA,Neelam A,Smith E,et al.Expression analysis of a sucrosecarrier in the germinating seedling of Ricinus communis[J].Plant Mol Biol,1998,38(3):425-435.
[31]Stadler R,Truernit E,Gahrtz M,et al.The At SUC1 sucrose carriermay represent the osmotic driving force for anther dehiscence andpollen tube grow thin Arabidopsis[J].Plant J,1999,19(3):269-278.
[32]Lemoine R,Burkle L,Barker L,et al.Identification of apollen-specific sucrose transporter-like protein Nt SUT3 fromtobacco[J].Febs Lett,1999,454(3):325-330.
[33]Sivitz AB,Reinders A,Ward JM,et al.Arabdopsis sucrose transporter At SUC1 is important for pollen germination and sucrose induced anthocyanin in accumulation[J].Plant Physiol,2008,147(1):92-100.
[34]Flemetakis E,Dimou M,Cotzur D,et al.A sucrose transporter,Lj SUT4 is up regulated during lotus japonicas nodule development[J].J Exp Bot,2003,54(388):1789-1791.
[35]Smeekens S,Rook F.Sugar sensing and sugar-mediated signaltransduction in plants[J].Plant Physiol,1997,115(1):7-13.
[36]Matsukura C,Saitoh T,Hirose T,et al.Sugar uptake and transportin rice embryo.Expression of companion cellspecific sucrosetransporter(Os SUT1)induced by sugar and light[J].PlantPhysiol,2000,124(1):85-93.
[37]Chiou TJ,Bush DR.Sucrose is a signal molecule in assimilatepartitioning[J].Proc Natl Acad Sci USA,1998,95(8):4784-4788.
[38]Ransom-Hodgkins WD,Vaughn MW,et al.Protein phosphorylationplays a key role in sucrose-mediated transcriptional regulation of aphloem-specific proton-sucrose symporter[J].Planta,2003,217(3):483-489.
[39]Müller J,Aeschbacher RA,Sprenger N,et al.Disaccharidemediated regulation of sucrose fructan-6-fructosyl transferase,a keyenzyme of fructan synthesis in barley leaves[J].Plant Physiol,2000,123(1):265-273.
[40]Gabriel-Neumann E,Neumann G,Leggewie G,et al.Constitutiveoverexpression of the sucrose transporter So SUT1 in potato plantsincreases arbuscular mycorrhiza fungal root colonization underhigh,but not under low,soil phosphorus availability[J].J PlantPhysiol,2011,168(9):911-919.
[41]Zhang YL,Meng QY,Qu GQ,et al.The antisence expression of aputative grape sucrose transporter in tobacco and effects on growthof the transformants[J].Prog Biochem Biophys,2006,7(7):699-705.
[42]Bitterlich M,Krügel U,Boldt-Burisch K,et al.The sucrose transporter Sl SUT2 from tomato interacts with brassinosteroid functioningand affects arbuscular mycorrhiza formation[J].Plant J,2014,78(5):877-889.
[43]Weichert N,Saalbach I,Weichert H,et al.Increasing sucroseuptake capacity of wheat grains stimulates storage proteinsynthesis[J].Plant Physiol,2010,152(2):698-710.
[44]Leggewie G,Kolbe A,Lemoine R,et al.Overexpression of thesucrose transporter So SUT1 in potato results in alterations in leafcarbon partitioning and in tuber metabolism but has little impact ontuber morphology[J].Planta,2003,217(1):158-167.
[45]Rosche E,Blackmore D,Tegeder M,et al.Seed-specificoverexpression of a potato sucrose transporter increases sucroseuptake and growth rates of developing pea cotyledons[J].Plant J,2002,30(2):165-175.
[46]Li H,Zhou SY,Zhao WS,et al.A novel wall-associated receptorlike protein kinase gene,Os WAK1,plays important roles in riceblast disease resistance[J].Plant Mol Biol,2009,69(3):337-346.
[47]Brandizzi F,Frangne N,Marc-Martin S,et al.The destinationfor single-pass membrane proteins is influenced markedly by thelength of the hydrophobic domain[J].Plant Cell,2002,14(5):1077-1092.
[48]Friedrichsen DM,Joazeiro CA,Li J,et al.Brassinosteroidinsensitive-1 is a ubiquitously expressed leucine-rich repeatreceptor serine/threonine kinase[J].Plant Physiology,2000,123(4):1247-1256.
[49]邱礽,陶刚,李奇科,等.农杆菌渗入法介导的基因瞬时表达技术及应用[J].分子植物育种,2009,7(5):1032-1039.
[50]庞建周.小麦干旱胁迫应答基因Ta SUT1的克隆、表达与基因转化研究[D].河北:河北师范大学,2011.
[51]高志民,杨学文,彭镇华,等.绿竹Bo SUT2基因的分子特征与亚细胞定位[J].林业科学,2010,46(2):45-50.
[52]Fan RC,Peng CC,Xu YH,et al.Apple sucrose transporter SUT1and sorbitol transporter SOT6 interact with cytochrome b5 toregulate their affinity for substrate sugars[J].Plant Physiol,2009,150(4):1880-1901.
[53]Sivitz AB,Reinders A,Johnson ME,et al.Arabidopsis sucrosetransporter At SUC9,high-affinity transport activity,intrageniccontrol of expression,and Early Flowering Mutant Phenotype[J].Plant Physiol,2007,143(1):188-198.