Κ2005Κ25; 2ΓΠ413 418 A cta B oṫ B orea l. -O cciden ṫ S in. Π100024025; 2005Γ0220413206 Ξ ; 712100Γ Π Κ. ABA Κ Π Μ ΜABA ΠQ 945. 78 ΠA The Respon se M echan ism of Active Oxygen Spec ies Rem ov ing System to D rought Stress ZHAO L i2yingκd EN G X i2p ingκshan L un ; Institute of So il and W ater ConservationΚCh inese A cadem y of Sciences and M inistry of W ater ResourcesΚN o rthw est Sci2T ech U niversity of A griculture and Fo restryκyanglingκshaanxi 712100ΚChinaΓ AbstractΠD rough t stress is a bo ttleneck facto r fo r p lan t grow th and developm en ṫ W ater stress induce the increased generation of active oxygen speciesκw h ich cau se ox idative stress to p lan t cells. som e are h igh ly tox ic and rap idly detox ified th roughvariou s cellu lar enzym atic and nonenzym atic m echan ism sκhow everκthe changes of active oxygen species generation resu lted in the sign ifican t enhancem en t in the activities of an2 tiox idan t enzym eṡ T he generation of A ctive oxygen species is invo lved in the accum u lation of ABA Κp ro2 line and ch lo rophyll fluo rescence ex term inating p rocesṡ T herefo reκ it is necessary to understand the re2 spon se m echan ism of active oxygen rem oving system to drough t stress and that of function in som e physio2 logical and b iochem ical p rocesṡ Key wordsπdrough t stressμreactive oxygen species scavenging system ΜABA ; stress shockγκ [ 1Κ2 ] Κ ;O - 2 Γ H 2O 2 ;O 2 1 Γ ;OH Γ [ 3 Κ ] Ξ Π2004208220Μ Π2004210214 Π ; G1999011708ΓΜ Π ; 1972- ΓΚ Κ Κ
414 25 DNA Κ ; GR Γ Κ [ 1 ] Κ GR Κ Κ NAD P + gnad PH Κ Κ NAD P + NAD P + [ 1 6 Κ ] O 2 O - 2 Μ GSH ggssgκ 1 [ 6 ] A PX A sa 2GSH Κ A PX ; SOD s superox ide dism u taseγ Κ ; CA T ΚcatalaseΓ Κ A PX ; sa PXΓ ;A PXΚasco rbate perox idaseγ A PX; ta PXΓΜ ;DHA R Κdehydroasco rbate reductaseγ Κ ;M DHA R Κm onodehydroasco rbate re2 ductaseγ ; GR Κglu tath ione reduc2 taseγ ; POD Κnon2sp ecific perox idaseγ SOD s ; GR Γ H aber2w eiss ; 2 O - 2 + 2H + [ 2 H 2O 2+ O 2Γ ] ΚSOD s ΠCugZnSOD FeSOD M nsod Κ CugZnSOD 3 Κ PEG26000 Κ ΚM nsod ΚFeSOD Κ SOD CA T CA T H 2O 2Κ CA T H 2O 2 H alliw ell2a sada ΚSOD CA T Κ [ 8 ΚA PX GR Κ ] ΚSOD CA T Κ H 2O 2 CA T POD M DA Κ Κ Κ 35 S2M et ΚCA T Κ SOD CA T POD CA T POD Μ ΚO - 2 H 2O 2 CA T CA T Κ CA T ΚSOD CA T [ 6 ] POD ΚPOD Κ POD CA T [ 8 H 2O 2Κ Κ ΚSOD ] Μ Κ Κ POD Κ A sa Κ CO 2 Κ GR [ 7 A PX ] ;M DHA R Γ ; Γ ;DHA R Κ POD
2 Κ Π 415 2 Κ Μ ;A SA Γ Car ;CarΚcaro teno idsγ Α2 Β2 Κ ; ΚGSH Κglu tath ione; re2 ducedfo rm ΓΓ Κ Car ; Ch lκch lo rophyllγ ; Ch lg Κ CarΓ Car Ch lκ ; asco rbate ox idaseγ Car [ 10 ] ;M DHA ΓΚ H alli2 Κ Ch lg w ell2a sada A PX Κ Car Car Κ ΚCh lgcar Κ A SA O - 2 Κ Car ΚCh lgcar OH 1 O 2 H 2O 2 V E [ 9 ] A SA [ 27 OH ] H alliw ell2a sada M DA A SA E 1 O 2 [ 9 A SA ] Κ [ 7Κ9 ] GSH Κ ; GSH Γ ; GSSGΓ ΚCar Car Β2 3 ABA Π Μ Κ Μ Κ [ 1 ] ABA [ 12 ] [ 13 ] ABA Κ NAD PH H 2O 2Μ [ 14Κ15 ] ABA SO 2ΚGSH SO 2 NAD PH NAD PH Κ O 2Κ Κ 2SH H 2O 2 [ 16 ] GSH ABA Κ ΚPEG Κ GSH ggssg Μ Κ ABA Κ Κ ΚGSH 50 Κ Κ GSH J iang M [ 17 ] 102100 m o lgl ABA [ 9 ]
416 25 H 2O 2 SOD ΚCA T ΚA PX GR Κ OH 1 O 2 A SA ΚGSH ΚΑ2TOC CA R Κ 100 M [ 17 ] ABA Κ g OH 1 O 2 m o lgl ABA O - 2 H 2O 2 Κ Κ 100 m o lgl ABA Κ1 000 m o lgl ABA O - 2 H 2O 2 OH OH Κ Α2TOC CA R O 2 1 OH A SA GSH Κ OH ABA ; 10 100 M DA OH m o lgl Γ [ 3Κ27 Κ ; 1 000 m o lgl ΓABA RO S ] [ 28 ] ΜPei [ 18 ] H 2DCFDA Zhang [ 19 ] H 2DCFDA ABA 2 H 2O 2ΚH 2O 2 Ca 2+ Κ Κ Ca 2+ H 2O 2 ABA SOD Μ ABA 1 m o lgl Κ H 2O 2Κ ABA CuZn2SOD M n2 Κ [ 21 24 SOD Fe2SOD CA T ] Κ SOD A PX CA T GR Κ [ 5Κ23 ] ABA J iang 5 ABA RO S Κ ABA Κ Κ ABA Μ 4 Κ Κ Sm irnoff Cum bes; 1993Γ [ 5 ] II; PS IIΓ QA Κ ΚQA [ 27 ] Κ OH 1 O 2 Κ Κ Κ O - 2 H 2O 2 ΚO 2 O - 2 2 ΚO 2 PSg
2 Κ Π 417 Κ PSg Κ O - 2 H 2O 2 ABA Κ Κ ABA 1 O 2 Ν ;RO SΓ OH Κ O - 2 ΝRO S ΝABA Ν Π; 1Γ [ 29Κ30 ] Κ Μ; 2Γ Κ Μ; 3Γ 6 Κ Κ Κ Κ Κ Π [ 1 ] ZEN G Q P; ΓΚGUO Y; Γ. A dversity response and system atic resistance induction in p lants[j ]. Chem istry of L if e; ΓΚ1997Κ17; 3ΓΠ31-33; in Ch ineseγ. [ 2 ] SHAN L ; ΓΚCH EN P Y; Γeds. T he Physio logical and Eco logical Basis of D ryland Farm ing [M ]. BejingΠScience P ress. Π ΓΚ1998Π1-18; in ChineseΓ. [ 3 ] J IAN G M Y; ΓΚYAN G W Y; ΓΚXU J ; ΓΚCH EN G Q Y ; Γ. O smo tic stress2induced oxidative injury of rice seedlings[j ]. A cta A g ronom ica S inica. ; ΓΚ1994Κ20; 6ΓΠ733-738; in ChineseΓ. [ 4 ] SH I ZH J; ΓΚHU ZH S; ΓΚL I R SH ; Γ. W ater stress and active oxygen m etabo lism [J ]. J ournal of Guizhou U niversity A g ricu ltu ral and B iolog ical S cienceσ; ; ΓΓ. 2002Κ21; 2ΓΠ140-145; in ChineseΓ. [ 5 ] SM IRON FF N. T he ro le of active oxygen in the response of p lants to w ater deficit and desiccation [J ]. N ew P hy tolκ1993κ25; 1ΓΠ27-58. [6 ] A SADA K. T he w ater2w ater cycle in ch lo rop lastsπ scavenging of active oxygens and dissipation of excess pho tons. A nnual Review of P lant Physio logy and P lant M o lecular B io logy[j ]. 1999Κ50Π601-639. [ 7 ] P IGNOCCH I CΚFL ETCH ER J M. W IL K IN SON J E. T he function of asco rbate oxidase in tobacco [J ]. P lant P hy siology Κ 2003Κ132Π 1 631-1 641. [ 8 ] SUN C X; ΓΚL IU ZH G; ΓΚJ IN G Y D ; Γ. Effect of w ater stress on activity and isozym e of the m ajo r defense2defense in m aize leaves. [J ]. J ou rnal of M aiz e S cience; ΓΚ2003Κ11; 1ΓΠ63-66; in ChineseΓ. [ 9 ] YAN CH SH ; ΓΚL I D Q ; ΓΚZHAN G J H ; Γ. O xidative dam age and antioxidant responses during drought2induced w inter w heat flag leaf senescence[j ]. A cta B oṫ B oreal. O ccid enṫ S in. ; ΓΚ2000Κ20; 4ΓΠ568-576; in ChineseΓ. [ 10 ] M A J Y; ΓΚZHOU R ; ΓΚCH EN G B S; Γ. The relationship betw een caro teno id and activated oxygen m etabo lism [J ]. J ou rnal of S hand ong A g ricu ltu re U nivercity ; ΓΚ1997Κ28; 4ΓΠ518-522; in ChineseΓ. [ 11 ] BOHN ERT H JΚJEM SEN R G. Strategies fo r engineering w ater stress to lerance in p lants[j ]. T rend s in B iotechnology Κ1996Κ14Κ89-97. [ 12 ] DAV IES W JΚZHAN G J. Roo t signals and the regulation of grow th and developm ent of p lants in drying so il[j ]. A nnual R ev iew of P lant P hy siology and P lant M olecular B iology Κ1991Κ42Π55-76. [ 13 ] SH INO ZA K I KΚYAM A GU CH I2SH INO ZA K I K. Gene exp ression and signal transduction in w ater2stress response[j ]. P lant P hy siolo2 gy Κ1997Κ115Π327-334. [ 14 ] GRAN T J JΚLOA KE G J. Ro le of reactive oxygen interm ediates and cognate redox signaling in disease resistance[j ]. P lant P hy siolκ 2000Κ124Π21-29. [ 15 ] N E ILL S JΚD ES IKAN R ΚCLA RKE A ΚHAN COCK J T. N itric oxide is a novel component of abscisic acid signaling in stom atal guard cells[j ]. P lant P hy siolκ2002κ128π13-16. [ 16 ] YAN X; ΓΚL I F M ; ΓΚWAN G Y F; Γ. The response of endogenous ABA and enzym es activity of scavenging free ;
418 25 radical of w heat seedling to w ater stress [J ]. A cta B oṫ B oreal. O ccid enṫ S in. ; ΓΚ2000Κ20; 6ΓΠ1 003-1 009; in ChineseΓ. [ 17 ] J IAN G M ΚZHAN G J. Effect of abscisic acid on active oxygen speciesκantioxidative defence system and oxidative dam age in leaves of m aize seedlings[j ]. P lant and Cell P hy siology Κ2001Κ42Π1 265-1 273. [ 18 ] PE I ZM ΚM U RA TA YΚBENN IN G GΚTHOM IN E SΚKLU SEN ER BΚALL EN G JΚGR ILL EΚSCHRO ED ER J İ Calcium channels activated by hydrogen peroxide m ediate abscisic acid signalling in guard cells[j ]. N atureκ2000κ406π731-734. [ 19 ] M OR I I CΚSCHRO ED ER J İ Reactive oxygen species activation of p lant Ca 2+ channels. A signaling m echanism in po lar grow thκho r2 mone transductionκ stress signalingκand hypo thetically m echano transduction [J ]. P lant P hy siology Κ2004Κ135Π702-708. [ 20 ] CHN E ZΚGALL IE D R. T he asco rbic acid redox state contro ls guard cell signaling and stom atal movem ent[j ]. P lant Cell Κ2004Κ16Π 1 143-1 162. [ 21 ] ZHAN G XΚZHAN G L ΚDON G FΚGAO JΚGALBRA ITH D W ΚSON G C P. H ydrogen peroxide is invo lved in abscisic acid2induced stom atal clo sure in V icia faba[j ]. P lant P hy siology Κ2001Κ126Π1 438-1 448. [ 22 ] J IA W ΚZHAN G J. W ater stress2induced abscisic acid accum ulation in relation to reducing agents and sulphhydryl modifiers in m aize p lant[j ]. P lantψcell and E nv ironm entκ2000κ23π1 389-1 395. [ 23 ] GUAN L ΚSCANDAL IO S J G. Two structurally sim ilar m aize cyto so lic superoxide dism utase genesκsod4 and Sod4A Κrespond differen2 tially to abscisic acid and high o smo ticum [J ]. P lant P hy siology Κ1998Κ117Π217-224. [ 24 ] ZHU D ΚSCANDAL IO S J G. D ifferential accum ulation of m anganese2superoxide dism utase transcrip ts in m aize in response to abscisic acid and high o smo ticum [J ]. P lant P hy siology Κ1994Κ106Π173-178. [ 25 ] GUAN L ΚSCANDAL IO S J G. Effects of the p lant grow th regulato r abscisic acid and h igh o smo ticum on the developm ental exp ression of the m aize catalase genes[j ]. P hy siolog ia P lantarum Κ1998Κ104Π413-422. [ 26 ] GUAN L ΚZHAO JΚSCANDAL IO S J G. C is2elem ents and transfacto rs that regulate exp ression of the m aize Cat1 antioxidant gene in response to ABA and o smo tic stressπh 2O 2 is the likely interm ediary signalling mo lecule fo r the response[j ]. 22Π87-95. T he P lant J ournalκ2000κ [ 27 ] J IAN G M Y; ΓΚGUO SH CH ; ΓΚZHAN G X M ; Γ. P ro line accum ulation in rice seedlings expo sed to oxidative stress in relation to antioxidation [J ]. A cta P hy top gy siolog ica S inica; ΓΚ1997Κ23; 4ΓΠ347-352; in ChineseΓ. [ 28 ] WAN G J ; ΓΚL I D Q ; ΓΚGU L K; Γ. T he reponse to w ater stress of the antioxidant system in m aize seedling roo ts w ith different drough t resistance[j ]. A cta B oṫ B oreal. O ccid enṫ S in. ; ΓΚ2002Κ22; 6ΓΠ285-290; in ChineseΓ. [29 ] PEN G CH L ; ΓΚL IN G ZH ; Γ. The Feature of chlo rophyll F luo rescence Q uenching in Xanthophyll2deficient M utants of A rabidop sis thaliana[j ]. P rog. B iochem. B iop hy ṡ ; ΓΚ2003Κ30; 2ΓΠ251-256; in ChineseΓ. [ 30 ] XU ZH F; ΓΚLUO G H ; ΓΚKE D S; Γ et al. Chlo rophyll fluo rescence quenching induced by superoxide anion [J ]. P rog. B iochem. B iop hy ṡ ; ΓΚ2002Κ29; 1ΓΠ139-142; in ChineseΓ.