沈元月;

• 1:北京农学院植物科学技术学院

摘要(Abstract)：

果实成熟调控是果树生产的核心问题。根据果实成熟的特点,果实可分为呼吸跃变及非呼吸跃变2个基本类型。葡萄属于典型的非呼吸跃变型果实。多年来,人们围绕跃变型果实开展了很多研究,但对非跃变型果实成熟调控的了解却相对贫乏。一般认为,植物激素脱落酸(abscisic acid,ABA)在非跃变果实成熟调控中起着关键的作用。近年来,越来越多的研究表明,葡萄果实中不仅存在着ABA受体,而且还存在着一系列ABA下游信号组分,如ABA应答的蛋白激酶,转录因子,糖载体和ASR蛋白(abscisic acid stress ripening protein)等,说明ABA的确在葡萄果实成熟调控中起着关键作用。综述了葡萄果实成熟与ABA信号传导相关研究的最新进展,以期促进和加快今后非跃变型果实成熟机理深入研究的步伐。

关键词(KeyWords)： 葡萄;果实成熟;脱落酸;信号传导

Abstract:

Keywords:

基金项目(Foundation): 国家自然科学基金(30971977);; 北京市自然科学基金(6082005);; 北京市教委联合资助重点项目(KZ200910020001);北京市教委平台建设项目(PXM2009);; 北京市属市管高校人才强教计划资助项目(PXM2007014207);;

作者(Author): 沈元月;

Email:

参考文献(References)：

• [1]COOMBEBG.Thedevelopmentoffleshyfruits[J].AnnuRevPlantPhys-iol,1976,27:207-228.
• [2]COOMBE B G,HALE C R.The hormone content of ripening grapeberries and the effects of growth substance treatments[J].Plant Phys-iol,1973,51:629-634.
• [3]WANG Xin-quan,ZHAO Min,SHANG Xiu-hong.The dynamicobservation of red earth-like grape berry growth and development[J].Deciduous fruit,2004(1):11-12.汪心泉,赵敏,尚秀红.红地球葡萄果实生长发育动态观察[J].落叶果树,2004(1):11-12.
• [4]COOMBE B G.Distribution of solutes within the developing grapeberry in relation to its morphology[J].American Journal of Enologyand Viticulture,1987,38:120-127.
• [5]COOMBE B G.Research on development and ripening of grape berry[J].Journal of Enology and Viticulture,1992,43(1):101-110.
• [6]DAVIES C,BOSS P K,ROBINSON S P.Treatment of grape berries,a nonclimacteric fruit with a synthetic auxin,retards ripening andalterstheexpressionofdevelopmentallyregulatedgenes[J].Plant Phys-iol,1997,115:1155-1161.
• [7]COOMBE B G.Relationship of growth and development to changes insugars,auxins and gibberellins in fruit of seeded and seedless vari-eties of Vitis vinifera[J].Plant Physiol,1960,35:241-250.
• [8]McGLASSON,W B.The ethylene factor.In:A.C.Hulme,ed.,TheBiochemistry of Fruits and Their Products[M].Vol.1.Academic Press,New York,1970:475-519.
• [9]CHEN Fa-he,CAI Hui-nong,FENG Zuo-shan,ZHANG Wei-yi,LIAO Kang.Changes in hormone levels in grape berries during de-velopment[J].Journal of Plant Physiology and Molecular Biology,2002,28(5):391-395.陈发河,蔡慧农,冯作山,张维一,廖康.葡萄浆果发育过程中激素水平的变化[J].植物生理与分子生物学学报,2002,28(5):391-395.
• [10]ZHANG You-lin,CHEN Jin-ping,WANG Ming-zhen,ZHANGBao-shan,HAN Jun-qi.Studies on abscisic acid(ABA)change ofgrape during storage[J].Acta Botacica Boreali-Occidentalia Sinica,2000,20(4):604-609.张有林,陈锦屏,王明珍,张宝善,韩军歧.葡萄贮期脱落酸(ABA)变化的研究[J].西北植物学报,2000,20(4):604-609.
• [11]ZHANG Da-peng,XU Xue-feng,ZHANG Zi-lian,JIA Wen-suo.Studies on the veraison mechanisms:Physiological and biochemicalresponses of grape berries to exogenous hormone treatment andgirdling during the lag phase of berry growth[J].Acta HorticulturaeSinica,1997,24(1):1-7.张大鹏,许雪峰,张子连,贾文锁.葡萄果实始熟机理的研究[J].园艺学报,1997,24(1):1-7.
• [12]XIA Guo-hai,ZHANG Da-peng,JIA Wen-suo.Effects of IAA,GA and ABA on 14C sucrose import and metabolism in grape berries[J].Acta Horticulturae Sinica,2000,27(1):6-10.夏国海,张大鹏,贾文锁.IAA、GA和ABA对葡萄果实14C蔗糖输入与代谢的调控[J].园艺学报,2000,27(1):6-10.
• [13]FINKESTTEIN R R,GAMPALA S S,ROCK C D.Abscisic acid sig-naling in seeds and seedlings[J].Plant Cell,2002,14:15-45.
• [14]XIONG L,SCHUMAKER K S,ZHU J K.Cell signaling during cold,drought and salt stress[J].Plant Cell,2002,14:165-183.
• [15]LEUNG J,GIRAUDAT J.Abscisic acid signal transduction[J].AnnuRev Plant Physiol Plant Mol Biol,1998,49:199-222.
• [16]HIRAYAMA T,SHINOZAKI K.Perception and transduction of ab-scisic acid signals:keys to the function of the versatile plant hormoneABA[J].Trends Plant Sci,2007,12(8):343-351.
• [17]WANG X F,ZHANG D P.Abscisic acid receptors:Multiple signal-perception sites[J].Annals of Botany,2008,101:311-317.
• [18]SHEN YY,WANG X F,WU F Q,DU S Y,CAO Z,SHANG Y,WANG X L,PENG C C,YU X C,FAN R C,XU Y H,ZHANG DP.The Mg-chelatase H subunit is an abscisic acid receptor[J].Nature,2006,443(7113):823-826.
• [19]PANDEY S,NELSON D C,ASSMANN S M.Two novel GPCR-typeG proteins are abscisic acid receptors in Arabidopsis[J].Cell,2009,136:136-148.
• [20]LIU X,YUE Y,LI B,NIE Y,LI W,WU W H,MA L A.G proteincoupled receptor is a plasma membrane receptor for the plant hor-mone abscisic acid[J].Science,2007,315:1712-1716.
• [21]NAMBARA E,MARION-POLL A.Abscisic acid biosynthesis andcatabolism[J].Annu Rev Plant Biol,2005,56:165-185.
• [22]BERTRAND S,BENHAMOU N,NADEAU P.Immunogold local-ization of free abscisic acid in tomato root cells[J].Canadian JournalBotany,1992,70(5):1001-1011.
• [23]GOLLAN T,SCHRRU U,SCHULZE E.Stomatal response to dryingsoil in relation to changes in the xylem sap composition of Helianthusannus.I.The concentration of cations,anions,amino acids and pHof the xylem sap[J].Plant Cell Environ,1992,15:551-559.
• [24]HUANG Cong-lin,ZHANG Da-peng,JIA Wen-suo.A study of thesources of abscisic acid in grape berry during its late developmentalphases[J].Acta Horticulturae Sinica,2001,28(5):385-391.黄丛林,张大鹏,贾文锁.葡萄果实发育后期脱落酸来源的研究[J].园艺学报,2001,28(5):385-391.
• [25]ZHANG Zi-lian,ZHANG Da-peng,CHEN Jia.The abscisic acidbinding proteins and their properties in grapevine fruits[J].ChineseBiochemical Journal,1996,12(4):404-408.张子连,张大鹏,陈珈.葡萄果实中脱落酸结合蛋白的存在和性质[J].生物化学杂志,1996,12(4):404-408.
• [26]ZHANG D P,ZHANG Z L,CHEN J,JIA W S.Specific abscisicacid-binding sites in mesocarp of grape berry:Properties and sub-cellular localization[J].Journal of Plant Physiology,1999,155:324-331.
• [27]ZHANG Da-peng,ZHANG Zi-lian,CHEN Jia.Changes in kineticcharacteristics of the ABA binding protein(s)of the grapevine fruit dur-ing its developmental process[J].Acta Botanica Sinica,1996,38(12):930-935.张大鹏,张子连,陈珈.葡萄果实发育过程中脱落酸结合蛋白动力学特性的变化[J].植物学报,1996,38(12):930-935.
• [28]STONE J M,WALKER J C.Plant protein kinase families and signaltransduction[J].Plant Physiol,1995,108:451-457.
• [29]MORI I C,MURATA Y,YANG Y,MUNERMAS A S,WANG Y F,ANDREOLI S,TIRIAC H,AIONSO J M,HARPER J F,ECKER JR,KWAK J M,SCHROEDER J I.CPK6 and CPK3 function in A-BA regulation of guard cell S-type anion-and Ca2+-permeablechannels and stomatal closure[J].PLoS Biology,2006,4:1749-1762.
• [30]FUJII H,VERSLUES P E,ZHU J K.Identification of two proteinkinases required for abscisic acid regulation of seed germination,root growth,and gene expression in Arabidopsis[J].Plant Cell,2007,19:485-494.
• [31]LU C,HAN M H,GUEVARA-GARCIA A,FEDOROFF N V.Mi-togen-activated protein kinase signaling in postgermination arrest ofdevelopment by abscisic acid[J].Proceeding of the National Academyof Sciences,2002,99:15812-15817.
• [32]OSAKABE Y,MARUYAMA K,SEKI M,SATOU M,SHINOZAKIK,YAMAGUCHI-SHINOZAKI K.Leucine-rich repeat receptor-like kinase1 is a key membrane-bound regulator of abscisic acidearly signaling in Arabidopsis[J].Plant Cell,2005,17:1105-1119.
• [33]FIKEELSTEIN R R,GAMPALA S S L,ROCK C D.Abscisic acidsignaling in seeds and seedlings[J].Plant Cell,2002,14:15-45.
• [34]HARONI A,O'Connell A P.Gene expression analysis of strawberryachene and receptacle maturation using DNA microarrays[J].Journalof Experimental Botany,2002,53(377):2073-2087.
• [35]SHEN Y Y,DUAN C Q,lIANG X E,ZHANG D P.Membrane-as-sociated protein kinase activities in the developing mesocarp of grapeberry[J].Journal of Plant Physiology,2004,16(1):15-23.
• [36]YU X C,LI M J,GAO G F,FENG H Z,GENG X Q,PENG C C,ZHU X Y,WANG X J,SHEN Y Y,ZHANG D P.Abscisic acidstimulates a calcium-dependent protein kinase in grape berry[J].Plant Physiol,2006,140(2):558-579.
• [37]YU X C,ZHU S Y,GAO G F,WANG X J,ZHAO R,ZOU K Q,WANG X F,ZHANG X Y,WU F Q,PENG C C ZHANG D P.Ex-pression of a grape calcium-dependent protein kinase ACPK1 inArabidopsis thaliana promotes plant growth and confers abscisicacid-hypersensitivity in germination,postgermination growth,andstomatal movement[J].Plant Mol Biol,2007,64(5):531-538.
• [38]LECOUTIEUX F,LECOURIEUX D,VIGBAULT C,DELROT S.Asugar inducible protein kinase,VvSK1,regulates hexose transportand sugar accumulation in grapevine cells[J].Plant Physiol,2010,152(2):1096-1106.
• [39]DAVIES C,ROBINSON S P.Differential screening indicates a dra-matic change in mRNA profiles during grape berry ripening.Cloningand characterization of cDNAs encoding putative cell wall and stressresponse proteins[J].Plant Physiol,2000,122(3):803-812.
• [40]SCHUSSLER J R,BRENNER M L,BRUN W A.Abscisic acid andits relationship to seed filling in soybeans[J].Plant Physiol,1984,76:301-307.
• [41]CLIFFORF P E,OFFLER C E,PATRICK J W.Growth regulatorshave rapid effects on photosynthetic unloading from seed coats ofPhaseolus rulgaris L[J].Plant Physiol,1986,80:635-641.
• [42]GIFFORD R M,THORNE J H.Phloem unloading in soybean seedcoats:Dynamics of stability of flux into attached empty ovules[J].Plant Physiol,1986,80:464-471.
• [43]BRADY C J.Fruit ripening[J].Annual Review of Plant Physiology,1987,38:155-178.
• [44]WAYNE H L,JOHN D E.Sugar alcohol metabolism in sinks andsources[M]//ZAMSKI E,SCHAFFER A A.Photoassimilate distribu-tion in plants and crops:source-sink relationships.Marcel Dekker,New York,1996.
• [45]ACKERSON R C.Invertase activity and abscisic acid in developingsoybean reproductive structures[J].Crop Science,1985,25:615-620.
• [46]ROCK C D,QUATRANO R S.The role of hormones during seed de-velopment[M].In Plant Hormones(Davies PJ).Kluwer AcademicPublishers,Dordrecht,The Netherlands,1995:671-697.
• [47]VENDRELL M,BUES A.Relationship between ABA content andripening of apples[J].Acta Horticulturae,1989,258:389-396.
• [48]FINDLAY N,OLIVER K J,NII N,COOMBE B G.Solute accumu-lation by grape pericarp cells.IV.Perfusion of pericarp apoplast viathe pedicel and evidence for xylem malfunction in ripening berries[J].J Exp Bot,1987,38:668-679.
• [49]FILLION L,AGEORGES A,PICAUD S,COUTOS T P,LEMOINER,ROMIEU C,DELROT S.Cloning and expression of a hexosetransporter gene expressed during the ripening of grape berry[J].Plant Physiol,1999,120(4):1083-1094.
• [50]CAKIR B,AGASSE A,GAILLARD C,SAUMINNEAU A,DEIROTS,ATANASSOVA R.A grape ASR protein involved in sugar and ab-scisic acid signaling[J].Plant Cell,2003,15(9):2165-2180.
• [51]IEASAKI T,SHINOZAKI K.Identification of a cis-regulatory regionof a gene in Arabidopsis thaliana whose induction by dehydration ismediated by abscisic acid and requires protein synthesis[J].Mol GenGenet,1995,247(4):391-398.
• [52]URAO T,SHINOZAKI K,URAO S.An Arabidopsis myb homolog isinduced by dehydration stress and its gene product binds to the con-served MYB recognition sequence[J].Plant Cell,1993,5:1528-1539.
• [53]WALKER AR,LEE E,BOGS J,McDAVID D A,THOMAS M R,ROBINSON S P.White grapes arose through the mutation of twosimilar and adjacent regulatory genes[J].Plant J,2007,49:772-785.
• [54]BOGS J,JAFFE F W,TAKOS A M,WALKER A R,ROBINSON SP.The grapevine transcription factor VvMYBPA1 regulates proan-thocyanidin synthesis during fruit development[J].Plant Physiol,2007,143(3):1347-1361.
• [55]MA Y,SZOSTKIEWICA I,KORTE A,MOES D,YANG Y,CHRISTMANN A,GRILL E.Regulators of PP2C phosphatase ac-tivity function as abscisic acid sensors[J].Science,2009,324:1064-1068.
• [56]PARK S Y,FUNG P,NISHIMURA N,JENSEN D R,FFUJII,ZHAO Y,LUMBA S,SANTIAGO J,RODRIGUES A,CHOW T F.Abscisic acid inhibits type 2C protein phosphatases via the PYR/PYL family of START proteins[J].Science,2009,324:1068-1071.
• [57]FUJII H,CHINNUSAMY V,RODRIGUES A,RUBIO S,AATONI R,PARK S Y,CUTLER S R,SHEEN J,ROTRIGUES P L,ZHU J K.Invitro reconstitution of an abscisic acid signalling pathway[J].Nature,2009,462:660-664.
• [58]GRIMPLET J,DELUC L G,TILLETT R L,WHEATLEY M D,SCHLAUCH K A,CRAMER G R,CUSHMAN J C.Tissue-specificmRNA expression profiling in grape berry tissues[J].BMC Genomics,2007,8:187.