Protective effect of exogenousnitric oxide onalleviation ofoxidative damage induced by high salinity in rice (Oryza sativa L.

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1 Shirz Univrsity Irn Agriulturl Rsrh(215) 34(1) 63-7 Prottiv t o xognousnitri oxi onllvition ooxitiv mg inu y high slinity in ri (Oryz stiv L.) slings S. AsiSnm 1,M. Zvrh 1*, A. Hshmpour 2 1 Dprtmnt o Agronomy n Plnt Bring, Fulty o Agriulturl Sins, Univrsity o Guiln, Rsht, I. R. Irn 2 Dprtmnt o Hortiulturl Sin, Fulty o Agriulturl Sins, Univrsity o Guiln, Rsht, I. R. Irn * Corrsponing Author: mzvrh@guiln..ir ARTICLE INFO Artil history: Riv 2 Frury 214 Apt 13 April 215 Avill onlin3 Jun 215 Kywors: Antioxint nzyms Ri Soium nitroprussi Rtiv oxygn spis ABSTRACT-To in th prottiv rol o xognous nitri oxi (NO) on slinitystrss ri slings, CRD-s toril xprimnt with thr rplitions ws onut in Agronomy Lortory o th Fulty o Agriulturl Sins, Univrsity o Guiln, in 212. Th xprimntl sign onsist o hlthy n vigorous slings o two ri ultivrs, Khzr n Goohr, th lst lry known s promising SA13 lin, whih wr xpos to (Control), 5 mm NCl, 5 µm soium nitroprussi (SNP) s NO onor solution supplmnt with simultnous 5 mm NCl + 5 µm SNP or our ys. Atr 4 ys, ltrolyt lkg n mlonilhy (MDA) ontnt, tivitis o ntioxint nzym, strution o hlorophyll n solul protin ontnt in lvs wr msur in trt n ontrol plnts. Th rsults show tht simultnous trtmnt o ri lvs with SNP, supprss th ion lkg ontnt y 8.5% ompr with th rsults o NCl trtmnt. Furthrmor, SNPinrs th tivitis o suproxi ismuts (SOD), proxis (POD), sort proxis (APX) n tls (CAT). Exognous pplition o NO lso ru proxition o mmrn lipis, whrs inrs th solul protin ontnt n hlorophyll pigmnts in ri lvs unr slinity strss. Ths rsults suggst tht NO oul tivly prott ri slings rom slt strss mg y nhning tivitis o ntioxint nzyms to qunh th xssiv rtiv oxygn spis us y slt strss. INTRODUCTION Slinity hs long n intii s on o th most prvsiv nvironmntl hzrs, limiting rop proution mostly in ri rgions o th worl (Ashr n Hrris, 24). Slt strss n hv signiint impt on th yil o ll mjor rls inluing ri, whih is slt snsitiv spis (Koym t l., 21). It hs n unrstoo tht soil n wtr slinity hs long nintii s mjor issu or ri proution (Grnln, 1984). High slinity nvironmnt n us plnt th or rs proutivity tth whol-plnt lvl. Th loss in plnt proutivity is onsqun o imln inioni n nutrints onntrtion n osmoti ts (Ashr, 29) rsulting in th ovrproution o rtiv oxygn spis (ROS) suh s suproxi(o 2 ) n hyrogn proxi (H 2 O 2 ). In plnts, th ntioxint nzyms r importnt omponnts in svnging ROS (Ashr, 29). Ths nzyms hv n intii in plnts s suproxi ismuts (SOD), proxis (POD), sort proxis (APX), n tls (CAT). Prvious stuis hv monstrt tht th ntioxint systms ply importnt rols in protting plnts ginst oxitiv mg inu y slt strss. It hs n rport tht slt strss us signiint inrss o r ril proution n mmrn mg o ri rsulting rom rs in th tivity o ntioxint nzyms (Dionisio-Ss n Toit, 1998). NCl trtmnt lso uss supprssion o th growth o pursln slings u to rs ntioxint nzym tivity (Yziit l., 27). Thror, th us o thniqus n/or mthos to llvit vrs ts o slinity strss is xpt to rsult in sustinl rop proutivity. Howvr, th mngmnt o slinity strss hs n onsir iiult us o its multigni n quntittiv ntur.nitri oxi (NO) is n importnt signling molul xrting vrious physiologil untions in plnts (Zhngt l., 29). It is rport tht NO mits th plnts' rsponss to oth ioti n ioti strsss (Guo n Crwor, 25). For instn, it involv s grmintion (Bligni n Lmttin, 2), t th growth n vlopmnt o plnt tissu (Pgnusstt l., 23), inrs iron homostsis (Durnr n Klssig, 1999) n rgult plnt mturtion n snsn (Guo n Crwor, 25). A w stuis hv lry suggst tht NO n ply rol in protting plnts rom oxitiv strsss (Shit l., 25; Singh t l., 28). (Li t l. 28) oun tht SNP s n No-onor n tivly prott rly slings rom slt strss mg y nhning tivitis o ntioxint nzyms to qunh ROS y slt strss. Furthrmor, No-onor trtmnt hs shown to inrs th tivitis o SOD n CAT in th root o Lupinus lutus (Kopyr n Gwozz, 23), n to rs ontnts o MDA n H 2 O 2 in s Tritium stivum (Zhng t l., 29) unr slinity strss. Howvr, littl is known out th inluns o xognous NO trtmnt on oxitiv mg n

2 AsiSnm t l./ Irn Agriulturl Rsrh (215) 34(1) 63-7 ntioxint nzym tivitis in lvs o ri unr slt strss. Thror, this stuy intns to invstigt th t o soium nitroprussi (SNP), s n NO-onor, on prottion rsponss o two ri ultivrs unr slt strss y nhning th tivity o ntioxint nzyms n improving physiologil rsponss. MATERIALS AND METHODS Fil Exprimnt Ss o two ri (Oryz stiv L.) ultivrs, Khzr n Goohr, otin rom th Ri Rsrh Institut o Irn, wr sur-striliz with 2.5% soium hypohlorit or 1 min. Thn, thy wr wsh with stril istill wtr,n wr pl on stril iltr ppr moistn with ioniz wtr n inut t 27 C or 72 h in rk to grmint. Grmint ss wr sown in plsti pots ontining prlit n wr llow to grow in growth hmr with 26/22 C y/night tmprtur n 12/12 light/rk prio t 35 µmol m 2 s 1 light nsity. Th growth hmr ytim rltiv humiity ws mintin 65±5%. Eh pot ws irrigt with 2 ml o Yoshi nutrint solution just to ph 5.2 vry thr ys. NCl n Nitri Oxi Trtmnt Fitn-y-ol hlthy n vigorous slings wr slt or xprimntl trtmnts. Th xprimntl trtmnts onsist o (Control; no -SNP n NCl, 5 µm SNP, 5 mm NCl, 5 mm NCl + 5 µm SNP) n two ri ultivrs (Khzr; vry snsitiv n Goohr; th lst lry known s promising SA13 lin). Eh trtmnt ws rplit t lst thr tims with 1 slings. At 4 ys, ll hlthy lvs o uniorm ri slings wr ollt n immitly prsrv in liqui nitrogn n stor t 8 C or ssys iohmil hrtristis. Msur Prmtrs Eltrolyt Lkg: Th ltrolyt lkg ws trmin in th mnnr sri y Dionisio-Ss n Toit(1998). Frsh ri lvs wr ut into pis o 5 mm lngth n pl in tst tus ontining 1 ml istill ioniz wtr. Th tus wr inut in wtr th t 32 C or 2 h n th initil ltril onutivity o th mium (EG 1 ) ws msur. Th smpls wr utolv t 121 C or 2 min to rls ll ltrolyts; ool to 25 C n th inl ltril onutivity (EG 2 ) ws msur. Th rltiv ltrolyt lkg (REL) ws xprss s prntg n lult y th qution:rel= (EC 1/ EC 2 ) 1 Lipi Proxition: Th lvl o mmrn mg ws msur y trmining mlonilhy ontnt (MDA) s th n prout o mmrn lipi proxition (Hth & Pkr, 1968). Lvs wr wigh n homogniz in solution ontining 1% trihloroti i (TCA), n thn ntriug t 1, g or 1 min. To 1.5 ml o th suprntnt liquot, 1.5 ml o 2% (w/v) TCA ontining.5% (w/v) TBA, wr. Th mixtur ws ht t 95 C or 6 min n ool to room tmprtur n ntriug t 1, g or 1 min. Th sorn o th suprntnt ws r t 532 n 6 nm ginst TCA solution s rgnt lnk. Th ontnt o MDA ws trmin using th xtintion oiint o 1.55 M 1 m 1 or MDA, n xprss in nmol MDA g -1 o rsh wight (r wt). Enzym Ativity: Ri lv smpls (.5 g) wr homogniz in 1 ml o 5 mm potssium phospht ur, ph 7 ontining 1 mm o EDTA in prsn o polyvinylpolypyrrolion (PVPP). Th homognt ws ntriug t 15, g or 15 min t 4 C. Th suprntnt ws us to msur th tivitis o suproxi-ismuts (SOD), proxis (POD), sort proxis (APX), tls (CAT) n to trmin totl protin ontnt. Enzyms tivitis wr msur t 25 C using sptrophotomtr (P.G Instrumnt T.8, Engln). Th tivity o SOD (EC ) ws trmin y msuring its ility to inhiit th photorution o nitro lu ttrzolium (NBT) oring to Ginnopolitis n Ris's mtho. Th rtion mixtur ontin 5 mm phospht ur (ph 7.), 2 mm mthionin, mm NBT, 1.5 mm EDTA, 75 M riolvin, n -5 µl o nzym xtrt. Riolvin ws s th lst omponnt. Rtions wr rri out in tst tus t 25 C unr illumintion suppli y two luorsnt lmps (2 W). Th tus wr shkn or thy wr pl in th rtion ssmly. Th rtion ws initit y swithing on th light n ws llow to run or 15 min, n swithing th light o stopp th rtion. Th tus wr thn immitly ovr with luminum oil in orr to stop th rtion n sorn o th mixtur whih ws thn r t 56 nm. Intil tus with omplt rtion mixtur, ontining no nzym xtrt, n vloping mximum olor srv s ontrol. A non-illumint omplt rtion mixtur with no olor vlopmnt srv s lnk. Unr th xprimntl onitions, th initil rt o rtion, s msur y th irn in inrs in sorn t 56 nm in th prsn n sn o l xtrt, ws proportionl to th mount o nzym. On unit SOD tivity ws in s th mount o nzym rquir to rsult in 5% inhiition o th rt o NBT rution msur t 56 nm.th SOD tivity o th xtrt ws xprss s tivity unit/ g -1 r wt. POD (EC ) tivity in lvs ws ssy y th oxition o guiol in th prsn o H 2 O 2. Th inrs in sorn ws ror t 47 nm (Chn & Mhly, 1955). Th rtion mixtur ontin 5 µl o ru nzym xtrt, 5 µl o 5 mm H 2 O 2, 5 µl o 28 mm guiol n 195 µl o 5 mm potssium phospht ur (ph 7). POD tivity o th xtrt ws xprss s tivity unit/ g -1 r wt min

3 AsiSnm t l./ Irn Agriulturl Rsrh (215) 34(1) 63-7 APX (EC ) tivity ws msur oring to Nkno n As (1981). Th rtion mixtur in totl volum o 2 ml onsist o 25 mm (ph 7.) soium phospht ur,.1 mm EDTA,.25 mm sort, 1. mm H 2 O 2 n 5 ml nzyms xtrt. H 2 O 2 -pnnt oxition o sort ws ollow y rs in th sorn t 29 nm. Th APX tivity o th xtrt ws xprss s APX unit/ g -1 r wt. min -1. CAT (EC ) tivity ws ssy oring to Chn n Mhly's mtho (1955). Th omposition o H 2 O 2 ws monitor y th rs in sorn t 24 nm. Th ssy mixtur ontin 2.55 ml o 5 mm potssium phospht ur (ph 7.), 4 µl o 15 mm H 2 O 2 n 5 µl o nzym xtrt. Th CAT tivity o th xtrt ws xprss s tivity unit/ g -1 r wt. min -1. Chlorophyll n Protin Contnts: Chlorophyll (Chl) ontnt ws trmin using Arnon's mtho (1949). 1 mg o th rsh l tissus wr grounwithpstl n mortr in liqui nitrogn. Thn, in powr ws trnsrr into tus n 1 ml o solut mthnol ws susquntly to tus n stor in th rk or 3 min. Th homognt ws ntriug t 5 rpm or 15 min. Th suprntnt ws us to msur th Chl n Chl ontnt. 1 ml o th xtrt ws thn ilut with 9 ml o solut mthnol n th sorn o th hlorophyll solution ws msur using sptrophotomtr t 645 n 663 nm. Th Chl n Chl ontnt in µg wr stimt using Arnon's ormul. Solul protin ontnt in lvs ws trmin oring to using ovin srum lumin (BSA) s stnr. Sttistil Anlyss Th xprimnt ws rrng in ompltly rnomiz sign with thr rplitions. Dt wr sujt to nlysis o vrin (ANOVA) ollow y Dunn s multipl rng tst mn omprison t P<.5 using SAS sotwr (v. 9.1, SAS Institut, Cry, NC, USA). Th igurs wr on using SigmPlot sotwr (v. 12). RESULTS AND DISCUUSION Th t o NO Trtmnt onrltiv Eltrolyt Lkg n MDA Contnt: Thr ws no signiint irn in ltrolyt lkg ontnt twn th two ultivrs o ri (Tl 1). Howvr, slinity n SNPtrtmnts h signiint t on ltrolyt lkg prntg in th ri lvs (Tl 1). Slinity strss signiintly inu ri lvs REL y 44% ompr to th ontrol group (Fig. 1). In th prsn o NCl n SNP togthr, th ion lkg rs y 8% ompr with th rsults o NCl trtmnt (Fig. 1). NCl trtmnt rsult in th highst MDA ontnt in oth ultivrs in omprison to ontrol plnts (Fig. 2). Unr norml onitions, th pplition o SNP h ngtiv t on MDA ontnt ompr to ontrol trtmnt (Fig. 2). Th ition o SNP signiintly ru NCl-inu MDA umultion tr oury slt strss-xposur (Fig. 2). It n sn tht SNP hs prottiv t ginst NCl-inu lipi proxition in ri lvs. Also, ths rsults show high rt o lipi proxition in Goohr ultivr u to slt strss (Fig. 2). Tl 1. Mn squrs rom nlysis o vrin o t or irnt iohmil ttriuts o two ri ultivrs trt with or without SNP n NCl or our ys SOV.. EL MDA SOD POD APX CAT PRT Trtmnt (T) ** 4.6 ** ** 23.2 ** 16.6 **.21 ** ** Cultivr (C) ns 1.92 ** 42.7 ns.767 ns 18.9 **.31 ** 97.7 ** T C ns.46 ** 8.44 ns 1.16 ns 17.7 **.4 * 15.2 ** Error *, ** = signiint t.5 n.1 lvls, rsptivly; ns = non-signiint. EL- ltrolyt lkg; MDA- mlonilhy; SODsuproxi ismuts; POD- proxis; APX- sort proxis; CAT- tls; PRT- protin. 65

4 AsiSnm t l./ Irn Agriulturl Rsrh (215) 34(1) 63-7 Eltrolytlkg(%) th mor iint ntioxint systms in SNP-trt plnts. Shi t l. (25) oumnt tht NO rus ltrolyt lkg in n lvs. Also, (Ruo t l. 2) rport tht NO is potnt inhiitor o lipi proxition.a similr nti-oxitiv untion o xognous NO ws lso osrv in slt-strss rly (Ruo t l., 2) n wht (Zhng t l., 29) plnts. Thror, xognous SNP trtmnt oul n tiv prti to prott ri slings ginst oxitiv mg us y slt strss. Fig. 1.Iniviul n omin ts o NCl n SNP on th ltrolyt lkg prntg in th ri lvs. Mn (±S.E.) ws lult rom thr rplits or h trtmnt. Dirnt lttrs init signiint irns t P<.5 (Dunn s multipl rng tst). MDA (nmol/ g -1 r wt) h khzr SA13 g Fig. 2. Iniviul n omin ts o NCl n SNP on MDA ontnt in th ri lvs. Mn (±S.E.) ws lult rom thr rplits or h trtmnt. Dirnt lttrs init signiint irns t P<.5 (Dunn s multipl rng tst). In th prsnt stuy, w oun tht slt strssinu oxitiv mg ri n msur th prottiv t o xognous NO. Slt strss us inrss in ltrolyt lkg (Fig. 1) n MDA ontnt (Fig. 2) in ri lvs. By gnrting hngs in unsturt tty is tht t mmrn strutur n proprtis, this ltrolyt lkg n lipi proxition unr slt strss my hv lso rought out n inrs in mmrn prmility or loss o mmrn intgrity (Dionisio-Ss n Toit, 1998), n ruprtion (though prtil) upon NO supply inits prvntiv rol o NO ginst NCl-mg. In t, NO prvnts mmrn mg y rsing lipi proxition n ltrolyt lkg rom plnt tissus (Li t l., 28). Howvr, th highr lvl o prottion in plnts trt with SNP my rsult o Th t o NO Trtmnt on th Ativitis o Antioxint Enzyms: Th tivitis o ntioxint nzyms signiintly inrs unr NCl trtmnt. Th tivitis o SOD, POX, APX ncat nzyms wr shown in Fig. 3-6, rsptivly. Thr ws no signiint irn in SOD n POD tivity twn th two ultivrs o ri (Tl 1). Howvr, SOD n POD tivitis wr irnt mong trtmnts (Tl 1). In th prsn o NCl n SNP togthr, th SOD tivity ws 56% highr thn our-y NCl trtmnt tr xposur (Fig. 3); howvr, no signiint irn ws osrv twn ultivrs. Also, unr unslinity onitions, th pplition o xognous SNP i not grtly hng SOD tivity in ri lvs (Fig. 3). SNP h inu similr hng t th POD tivity n th mximl vlu o POD tivity ws units g -1 r wt. min t omin SNP + NCl trtmnt, whih ws 25.2% mor thn NCl trtmnt lon tr our-y trtmnts (Fig. 4). SOD(unit/g -1 rwt ) Fig. 3. Iniviul n omin ts o NCl n SNP on SOD tivity in th ri lvs. Mn (±S.E.) ws lult rom thr rplits or h trtmnt. Dirnt lttrs init signiint irns t p<.5 (Dunn s multipl rng tst). 66

5 AsiSnm t l./ Irn Agriulturl Rsrh (215) 34(1) 63-7 POD (unit/ g -1 rwt. min -1 ) CAT (unit/ g -1 r wt. min -1 ) Khzr SA13 g g Fig. 4. Iniviul n omin ts o NCl n SNP on POD tivity in th ri lvs. Mn (±S.E.) ws lult rom thr rplits or h trtmnt. Dirnt lttrs init signiint irns t p<.5 (Dunn s multipl rng tst). APX tivity in ri lvs ws signiintly inu y slt strss (p<.5), n pplition o SNP h n ovious t on APX tivity (Fig. 5). Inuil t o SNP on th tivity o APX unr strss onitions ws highr thn SOD n POD tivitis. Unlik SOD n POD, APX tivity signiintly inrs in th two ultivrs o ri (Fig. 5). Similr to APX, NCl strss signiintly inrs CAT tivity in lvs o Khzr ultivr in omprison to its tivity in ontrol plnts (Fig. 6). Unr norml onitions, CAT tivity ws signiintly promot y SNP trtmnt (Fig. 6). Th CAT tivity ws sttistilly irnt in th ontrol plnts n th plnts trt with SNP n thos trt with th omin trtmnt (NCl n SNP) n th plnts trt with NCl only (Fig. 6). APX (unit/ g -1 r wt. min -1 ) Khzr SA13 Fig. 5. Iniviul n omin ts o NCl n SNP on APX tivity in th ri lvs. Mn (±S.E.) ws lult rom thr rplits or h trtmnt. Dirnt lttrs init signiint irns t p<.5 (Dunn s multipl rng tst).. Fig. 6. Iniviul n omin ts o NCl n SNP on AT tivity in th ri lvs. Mn (±S.E.) ws lult Crom thr rplits or h trtmnt. Dirnt lttrs init signiint irns t p<.5 (Dunn s multipl rng tst). It is wll known tht th ntioxint nzyms suh s SOD, POD, APX n CAT ply signiint rol in svnging ROS in slt strss plnts (Ashr, 29; Tunt l., 28).Erlir stuis hv monstrt tht xognous NO protts lvs ginst oxitiv mg in r unr ht strss (Song t l., 26) n in wht unr rought strss (Tin n Li, 26), whih ws minly sri to th inrs tivitis o SOD, POD n CAT. Thus, th prottiv rol o NO my ttriut to its miting th xprssion o gns noing ths ROS-svnging nzyms unr slt strss (Uhi t l., 22; Li t l. 28; Zhng t l., 29). In orn with ths inings, w oun tht th tivitis o SOD (Fig. 3), POD (Fig. 4), APX (Fig. 5) n CAT (Fig. 6) in plnts xpos to slt strss wr signiintlyinrs whn NO ws, whih ws in grmnt with thrsults rport y Uhi t l. (22), Kopyr n Gwozz (23), Li t l. (28), n Zhng t l.(29). Ths rsults onirm tht NO-onor ition n prott plnts rom mg us y slt strss. Th t o NO Trtmnt on Chlorophyll Contnt: Chlorophyll, hlorophyll n totl hlorophyll ontnt signiintly rs in rspons to slinity strss (Tl 2). Th hlorophyll (38.1%) n (64.9%) lvls n lso totl hlorophyll ontnt inrs in rspons to 5 Mm NCl + 5 µm SNP (Tl 2). Chlorophyll loss hs n rport s ritrion o snsn (Hung & Ko, 23). Our rsults xhiit rs in hlorophyll ontnt unr NCl trtmnt in ri slings, whih hslso n prviously rport (Siringm t l., 29). Th rs o l hlorophyll unr slinity is u to th strution o hlorophyll pigmnts n th instility o pigmnt protin omplxs (Lvitt, 198). Aing SNP to th NCl strss slings mrkly inrs th hlorophyll ontnt (Tl 2). Th inrs hlorophyll ontnt with th SNP trtmnt my u to th ility o SNP to nhn ytokinin proution, whih stimults hlorophyll iosynthsis (Flthr t l., 21). 67

6 AsiSnm t l./ Irn Agriulturl Rsrh (215) 34(1) 63-7 Tl 2. Chlorophyll (Chl ), hlorophyll (Chl ) n totl hlorophyll (TC) in ri slings trt with or without SNP n NCl or our ys Trtmnt Chl Chl TC (µg g -1 r wt) (µg g -1 r wt) (µg g -1 r wt) ontrol 7.71± ± ± µμ SNP 8.76± ± ± mm NCl 5.96± ± ± mm NCl + 5 µμ SNP 8.19± ± ±.3 ANOVA ** * ** Mn (±S.E.) ws lult rom thr rplits or h trtmnt. Mns ollow y irnt lttrs r signiintly irnt s on Dunn s multipl rng tst (P<.5). *, ** = signiint t.5 n.1 lvls, rsptivly. Th t o NO Trtmnt on Protin Contnts: In this xprimnt, totl solul protin ontnt ru in th ri lvs unr NCl strss onition (Fig. 7). Howvr, omin SNP + NCl trtmnt inrs th totl solul protin ontnt in Khzr (53.1%) n Goohr (2/9%) ultivrs ompr with NCl strss plnts (Fig. 7). protin(mg/g -1 rwt) 35 khzr SA slt onition rsult in signiintly highr l solul protin umultion or oth ultivrs whih oul proly u to prouing n intiying spii protins (Ky t l., 1981; Homn t l., 212) tht irntilly umult in lvs. Among th irnt protin lsss, w vlut hngs in irnt ntioxint nzyms u to prvious rsrh showing tht slt strss oul llvit oxitiv strss y improving ROS svnging pity (Mittlr t l., 24; Slot n KhnnChopr, 21). In this stuy, NO s n ntioxint, irtly qunhs th ROS n moults vrious llulr physiologil prosss to limit oxitiv injury n prott ri slings rom slt strss mg (Uhi t l., 22). 5 Fig. 7. Iniviul n omin ts o NCl n SNP on protin ontnt in th ri lvs. Mn (±S.E.) ws lult rom thr rplits or h trtmnt. Dirnt lttrs init signiint irns t p<.5 (Dunn s multipl rng tst). Th rution in protin ontnt unr slinity oul u to th rs in protin synthsis, lrt protolysis, n rs vilility o mino is n nturtion o nzyms involv in protin synthsis (Shrm n Ditz, 29). SNP trtmnt in REFERENCES Arnon, D.I. (1949). Coppr nzyms in isolt hloroplsts, polyphnol oxis in Bt vulgris. Plnt Physiology, 24, Ashr, M. (29). Biothnologil pproh o improving plnt slt tolrn using ntioxints s mrkrs. Biothnology Avns, 27, Ashr, M., & Hrris, P.J.C. (24). Potntil iohmil initors o slinity tolrn in plnts. Plnt Sin, 166, Bligni, M.V., & Lmttin, L. (2). Nitri oxi stimults s grmintion n -tioltion, n inhiits hypootyl longtion, thr light-inuil rsponss in plnts. Plnt, 21, Bror, M.M. (1976). A rpi n snsitiv mtho or quntittion o mirogrm quntitis o protin utilizing th prinipl o protin-y ining. Anlytil Biohmistry, 72, CONCLUSIONS In onlusion, our stuy monstrt tht NO n tivly prott ri slings rom slt strss mg y nhning th tivity o ntioxint nzyms n ruing mmrn injury to prvnt mg ssoit with xssiv ROS. Howvr, th pplition os o NO onor ns urthr invstigtion oring to irnt plnt spis n irnt growth stgs. Chn, B., & Mhly, S.K. (1955). Assy o tls n proxiss. Mthos in Enzymology, 2, Dionisio-Ss, M.L., & Toit, S. (1998). Antioxint rsponss o ri slings to slinity strss. Plnt Sin. 135: 1-9. Durnr, J., & Klssig, D.F. (1999). Nitri oxi s signl in plnts. Currnt Opinion in Plnt Biology, 2, Flthr, R.A., Gilly, A., Snkhl, N., & Dvis, T.D. (21). Trizols s Plnt Growth Rgultors n Strss Prottnts. In "Hortiulturl Rviws", pp John Wily & Sons, In. Ginnopolitis, C.N., & Ris, S. K. (1977). Suproxi ismutss I. Ourrn in highr plnts. Plnt physiology, 59, Grnln, D.J. (1984). Exploit Plnts: Ri. Biologist, 31, Guo, F.Q., & Crwor, N.M. (25). Ariopsis nitri oxi synths 1 is trgt to mitohonri n protts ginst oxitiv mg n rk-inu snsn. Plnt Cll, 17,

7 AsiSnm t l./ Irn Agriulturl Rsrh (215) 34(1) 63-7 Hth, R.L., & Pkr, L. (1968). Photoproxition in isolt hloroplsts: I. Kintis n stoihiomtry o tty i proxition. Arhivs o Biohmistry n Biophysis, 125, Homn, L., DCost, M., Eon, J.S., & Zho, J. (212). Ets o rought pronitioning on rzing tolrn o prnnil rygrss. Environmntl n Exprimntl Botny, 79, Hung, K.T., & Ko, C.H. (23). Nitri oxi ountrts th snsn o ri lvs inu y sisi i. Journl o Plnt Physiology, 16, Ky, L.J., Lin, C.Y., & Chn, Y.M. (1981). Ht shok protins o highr plnts. Proings o th Ntionl Amy o Sins o th Unit Stts o Amri, 78, Kopyr, M., & Gwozz, E.A. (23). Nitri oxi stimults s grmintion n ountrts th inhiitory t o hvy mtls n slinity on root growth o Lupinus lutus. Plnt Physiology n Biohmistry,41, Koym, M.L., Lvsly, A., Konr, R.M.D., Flowrs, T J., & Yo, A.R. (21). Quntittiv trit loi or omponnt physiologil trits trmining slt tolrn in ri. Plnt Physiology. 125: Lvitt, J. (198). Rsponss o plnts to nvironmntl strsss. Ami Prss, Nw York. pp Li, Q.Y., Niu, H.B., Yin, J.. Wng, M.B., Sho, H.B., Dng, D.Z., Chn, X.X., Rn, J.P., & Li, Y.C. Li. (28). Prottiv rol o xognous nitri oxi ginst oxitiv-strss inu y slt strss in rly (Horum vulgr). Collois n Surs B: Biointrs,65, Mittlr, R., Vnruwr, S., Gollry, M., & Brusgm, F. (24). Rtiv oxygn gn ntwork o plnts. Trns in Plnt Sin, 9, Nkno, Y., & As K. (1981). H 2 O 2 is svng y sort-spii proxis in spinh hloroplsts. Plnt n Cll Physiology, 22, Pgnusst, G.C., Lntri, M.L., & Lmttin, L. (23). Nitri oxi n yli GMP r mssngrs in th inol ti i-inu vntitious rooting pross. Plnt Physiology, 132, Ruo, R., Ri, H.R., Anslmi, D., Kirk, M., Brns, S., Butlr, J., Eisrih, J.P. & Frmn, B.A. (2). Nitri oxi rtion with lipi proxyl rils sprs lphtoophrol uring lipi proxition. Grtr oxint prottion rom th pir nitri oxi/lph-toophrol thn lph-toophrol/sort. Th Journl o Biologil Chmistry, 275, Slot, D.S., & KhnnChopr, R. (21). Antioxint rspons o wht roots to rought limtion. Protoplsm, 245, Shrm, S.S., & Ditz, K.J. (29). Th rltionship twn mtl toxiity n llulr rox imln. Trns in Plnt Sin, 14, Shi, S.Y., Wng, G., Wng, Y.D., Zhng, L.A., & Zhng, L.X. (25). Prottiv t o nitri oxi ginst oxitiv strss unr ultrviolt-b rition. Nitri Oxi, 13, 1-9. Singh, H.P., Btish, D.R., Kur, G., Aror, K., & Kohli, R.K. (28). Nitri oxi (s soium nitroprussi) supplmnttion mliorts C toxiity in hyroponilly grown wht roots. Environmntl n Exprimntl Botny, 63, Siringm, K., Juntwong, N., Ch-um, S., & Kirmn, C. (29). Rltionships twn soium ion umultion n physiologil hrtristis in ri (Oryz stiv L.) slings grown unr iso -osmoti slinity strss. Pkistn Journl o Botny, 41, Song, L., Ding, W., Zho, M., Sun, B., & Zhng, L. (26). Nitri oxi protts ginst oxitiv strss unr ht strss in th lluss rom two otyps o r. Plnt Sin, 171, Tin, X., & Li, Y. (26). Nitri oxi trtmnt llvits rought strss in wht slings. Biologi plntrum, 5, Tun, A.L., Ky, C., Dikilits, M., & Higgs, D. (28). Th omin ts o girlli i n slinity on som ntioxint nzym tivitis, plnt growth prmtrs n nutritionl sttus in miz plnts. Environmntl n Exprimntl Botny, 62, 1-9. Uhi, A., Jgnor, A.T., Hiino, T. Tk, T., & Tk, T. (22). Ets o hyrogn proxi n nitri oxi on oth slt n ht strss tolrn in ri. Plnt Sin, 163, Yzii, I., Türkn, I., Skmn, A.H., & Dmirl, T. (27). Slinity tolrn o pursln (Portul olr L.) is hiv y nhn ntioxitiv systm, lowr lvl o lipi proxition n prolin umultion. Environmntl n Exprimntl Botny, 61, Yoshi, S., Forno, A.D., Cook, J.H., & Goms, K.A. (1976). Lortory Mnul or Physiologil Stuis o Ri, (3 r.). Th Intrntionl Ri Rsrh Institut, Los Bnos, Lgun, Philippins. Zhng, C., Jing, D., Liu, F., Di, T., Liu, W., Jing, Q., & Co, W. (29). Exognous nitri oxi improvs s grmintion in wht ginst mitohonril oxitiv mg inu y high slinity. Environmntl n Exprimntl Botny, 67,

7-63 (1)34 (1394) (Oryz stiv L.) 2 *1 1 1.. 2.. * (NO) -. 5 (NCl) 5 SA13 5 (SNP).

8 7-63 (1)34 (1394) (Oryz stiv L.) 2 * * (NO) -. 5 (NCl) 5 SA13 5 (SNP). ( ) SNP 5 (MDA). 8/5 SNP SNP. (APX) (POD) (SOD) NO. (CAT). NO. : 1391/11/25 : 1392/6/3 : 1394/4/9 : : 7

The Effects of NaCl Stress on the Physiological and Oxidative Situation of Maize (Zea mayz L.) Plants in Hydroponic Culture

The Effects of NaCl Stress on the Physiological and Oxidative Situation of Maize (Zea mayz L.) Plants in Hydroponic Culture Currnt Rsrh Journl of Biologil Sins (1): 17-, 1 ISSN: 1-77 Mxwll Sintifi Orgniztion, 1 Sumitt: Sptmr 1, 11 Apt: Otor 9, 11 Pulish: Jnury, 1 Th Effts of NCl Strss on th Physiologil n Oxitiv Sitution of