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

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1 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 Miz (Z myz L.) Plnts in Hyroponi Cultur 1 Hmi Noorni Az, Mohmm Rz Hjighri, 1 Frshi Kfilzh n Mji Shni 1 Dprtmnt of Biology, Jhrom Brnh, Islmi Az Univrsity, Jhrom, Irn Dprtmnt of Agriultur, Esthn Brnh, Islmi Az Univrsity, Esthn, Irn Astrt: Th ffts of NCl slinity on iomss, Mlonilhy (MDA), proxis (POD), Ctls (CAT), N +, K +, C + n prolin in Z mys L. slings wr invstigt unr hyroponi onition. Slings wr sujt to NCl strss (,, 1, 1 n mm) for 1 ys. A ompltly rnomiz sign with four rplits for h trtmnt ws us. Slinity strss fft on th growth n us rution in root n shoot iomss. NCl trtmnt us signifint inrs in root MDA ontnt. NCl t 1 mm n highr inrs lso th shoot MDA ontnt signifintly. Ctls tivity of lf ws signifintly inrs t 1, 1 n mm NCl in omprison with th ontrol. Proxis tivity in lf strt to signifint inrs with th ris of NCl ontnt t 1 n mm. Th lf N + ontnt, root n shoot prolin onntrtions inrs with th inrs in slinity strss. Th lf K + n C + mounts wr signifintly rs with th ris of slinity strss in omprison with ontrol. Ky wors: Antioxitiv nzyms, miz, MDA, prolin, slt strss INTRODUCTION Slt strss in soil or wtr is on of th mjor ioti strsss spilly in ri n smi-ri rgions n n svrly limit plnt growth n yil (Prviz n Stywti, ). Dmgs us y high slinity r oftn ssoit with thr iffrnt mhnisms (Lvinsh, ). First, ion toxiity is us y xssiv umultion of N + n ClG in th ytoplsm ling to n ioni imln. This n ountrt y n inrs trnsport intnsity of th ions to th vuol. Son, vn if mssiv ion omprtmnttion ourr in th vuol, th ytosol wtr potntil must lowr to ln low-xtrnl wtr potntil, thus llowing wtr intk in th plnt ll n prvnting mromolul mgs. Thir, high llulr NCl onntrtion uss n inrs formtion of Rtiv Oxygn Spis (ROS) (Hrnnz n Almns, ). Slt strss n l to stomtl losur, whih rus CO vilility in th lvs n inhiits ron fixtion, xposing hloroplsts to xssiv xittion nrgy, whih in turn oul inrs th gnrtion of ROS n inu oxitiv strss (Prviz n Stywti, ). Ths spis of oxygn r highly ytotoxi n n sriously rt with vitl iomoluls suh s lipis, protins, nuli i, t, using lipi proxition, protin nturing n DNA muttion, rsptivly (Quils n Lópz, ). Plnts hv vlop sris of nzymti n non-nzymti toxifition systms to ountrt ROS, n prott lls from oxitiv mg (Sirm n Tygi, ). Th Antioxint nzyms suh s CAT, n POD funtion in toxifition of supr oxi n H O (Mittlr, ). Bsi, Munns n Jms (3) suggst tht th ssssmnt of ll mmrn stility is n pproprit thniqu to srn plnts unr slin onition. Bnoglu t l. () foun tht slt strss inrs MDA proution n ll mmrn mg in lf of ri sling. Rution in ll mmrn mg n lipi istriution (MDA proution) improvmnt with ntioxint nzym tivity. Whi n Clos (7) Rport tht Prolin umults in lrg quntitis in rspons to nvironmntl strsss ts s omptil solut, n uffrs llulr rox potntil. ROS srv s intrmit moluls in inuing slt rsistn in th llus of Populus uphrti, y inrsing th K + /N + rtio, whih ws pnnt on th inrs plsm mmrn H + -ATPs tivity (Zhng t l., 7). Slinity is known to inu K + fiiny in plnts us of th omptition of N + n K + for ining sits in trnsport systms tht mit K + uptk (Shl, 3). Miz (Z mys L.) is on of th most importnt rops in Irn, whih plys spil rol in popl s nutrition. But unfortuntly ioti strsss, suh s slinity, rs miz growth n proutivity y ruing wtr uptk n us nutrint isorrs n ion toxiity in this rgion. In th prsnt stuy, th ffts Corrsponing Author: Hmi Noorni Az, Dprtmnt of Biology, Jhrom Brnh, Islmi Az Univrsity, Jhrom, Irn 17

2 Curr. Rs. J. Biol. Si., (1): 17-, 1 of inrsing NCl onntrtions on iomss, MDA, POD, CAT, tion onntrtions n prolin ontnt in Z mys L. slings wr invstigt. MATERIALS AND METHODS Plnt ultur n trtmnts: Th xprimnt ws prform in glsshous of Islmi Az Univrsity, Jhrom Brnh, Irn, in 1, unr th following ontroll onitions: photoprio of 1/ h (light/rknss), ily ir tmprtur n rltiv humiity rnging -ºC n -7%, rsptivly, n photosynthtilly tiv ritions of :mol/m.s. Mtur ss of miz (Z mys L.) ultivrs, singl ross (No -7) otin from vgtl rsrh ntr, Frs, Irn. S wr surf striliz with solution of mruri hlori ).1% for 3 S ( n wr wsh immitly with lrg volum of stril istill wtr, n grmint in plsti pots (1 m imtr n m pth) fill with stril sn n wtr ily with istill wtr until grmintion. Thn, slings wr irrigt for wks with Hogln, s nutrint solution (1/ strngth stril n ph = 7). Aftrwr, th slings wr trnsfrr to Hogln solution ontinrs n trt with ing iffrnt onntrtions of NCl: (ontrol),, 1, 1 n mm for 1. A ompltly rnomiz sign with four rplits for h trtmnt ws us. Aftr trtmnt for 1 ys, th miz slings wr smpl n trnsfrr to liqui nitrogn n mintin t -7ºC until th msurmnt of vrils unr stuy. Biomss ssy: Th ry wight (DW) of shoot n root wr msur t h smpling. Th DW is otin ftr ovn-rying th shoots n roots t ºC for 7 h. Lipi proxition in lf n root: Lipi proxition ws trmin y msuring th mount of mlonilhy (MDA) formtion, using th thiorituri i mtho sri y Vlntovi t l. (). Brifly, mg of frsh tissus lf n root wr homogniz in ml of.1% (w/v) trihloroti i (TCA) solution on i. Th homognt ws ntrifug t 1, rpm for 1 min, n. ml of th suprntnt ws to. ml of.% (w/v) TBA in % TCA. Th mixtur ws inut in oiling wtr for min n th rtion ws stopp y inution in i. Smpls wr thn ntrifug for min t 1, rpm, n th sorn of th suprntnt ws msur t 3 nm, sutrting th vlu for nonspifi sorption t nm (Vlntovi t l., ). Antioxint nzymti tivitis in lf: Lvs (. g) wr homogniz in ml of i-ol potssium phospht uffr ( mm, ph = 7.) ontining. mm EDTA, 1.% (w/v) polyvinylpyrrolion n.1% triton. Th homognt ws ntrifug t 1g t ºC for min. Protins wr quntifi using ovin srum lumin s stnr (Brfor, 197). Ctls (CAT) tivity ws trmin y onsumption of H O in sorn t nm oring to mtho of Ai (19). Th msurmnts of proxis (POD) tivity wr on oring to Filing n Hll (197). Enzym xtrtion ws to rtion mium ontining mm potssium phospht uffr (ph = 7),.1 mm N EDTA, mm H O n 3 mm gyol. Th inrs in sorption ws ror t 7 nm. N +, K + n C + ontnts in lf: Ctions wr xtrt from th sit lf tissu with.% (v/v) nitri i. Th onntrtion of N + n K + ws ssy y flm mission photomtry. Th onntrtions of C + wr msur y tomi sorption photomtry (Hsn t l., 11). Root n shoot prolin ontnts: Extrtion n trmintion of prolin ws prform oring to th mtho of Bts t l. (1973). Root n shoot smpls (. g) wr xtrt with 3% sulfosliyli i. Extrts ( ml) wr hl for 1 h in oiling wtr y ing ml ninhyrin n ml glil ti i, ftr whih ol tolun ( ml) ws. Prolin ontnt ws msur y sptrophotomtr (Shimzu-UV- 11-Jpn) t nm n lult s mmol/g FW ginst stnr prolin. Th sttistis mtho: Sttistil nlyss wr rri out y nlysis of vrin (ANOVA) using SAS 9 softwr. Diffrns twn trtmnts wr nlyz y th Dunn s multipl rng tst. RESULTS Effts of slt strss on iomss: Th iomss of shoots n roots, stimt on th sis of thir ry wight, rs signifintly (**p<.1 ) in slt-trt plnts, whn ompr with ontrol plnts (Fig. 1, ). Thrfor, slinity in miz slings us inhiition of shoots n roots growth n rution in th iomss of orgns. Efft on th MDA ontnts: A ris in MDA, n initor of mmrn mg ws osrv unr NCl slinity. Slt strss for 1 inrs MDA ontnts in root n lf of Z mys L. slings. Howvr, th iffrn twn ontrol n mm NCl ws not signifint in lf. Trtmnts with NCl slinity inu signifint (**p<.1) inrss in root MDA ontnt (Fig. 1, ). Th inrs of MDA lvl in root ws grtr thn in lf.

3 Curr. Rs. J. Biol. Si., (1): 17-, 1 Shoot ry wight (mg/plnt) () 1 1 NCl onntrtions (mm) Root ry wight (mg/plnt) () NCl onntrtions (mm) Root MDA ontnts (nmol/g FW ) () NCl onntrtions (mm) Lf MDA ontnts (nmol/g FW) () NCl onntrtions (mm) Lf POD tivity (U/mg protin) () NCl onntrtions(mm) Lf PCAT tivity (U/mg) protin) (f) NCl onntrtions (mm) Fig. 1: Effts of NCl on shoot ry wight, () root ry wight, () root MDA ontnts, () lf MDA ontnts, () lf POD tivity, () lf CAT tivity, (f) in miz Chngs in ntioxint nzyms tivitis: In th lf of Z mys L. slings ntioxint nzym tivitis inrs t th slt strss. Proxis tivity in lf strt to signifint inrs with th ris of NCl ontnt t 1 n mm in omprison with th ontrol (**p<.1) (Fig. 1). Ctls tivity ws signifintly (**p<.1) inrs in lf t 1, 1 n mm NCl in omprison with th ontrol (Fig. 1f). Effts of slt strss on tion ontnts: Figurs., n C illustrt N +, K + n C + hngs in lvs of miz slings y inrs in NCl slinity. Th onntrtions of N + inrs in slt-trt plnts of Z mys L. whn ompr with ontrol plnts (**p<.1). K + n C + ontnts in lf wr signifintly rs with th ris of slinity strss in omprison with ontrol onitions (**p<.1). Drs in K+ onntrtion ws signifint in ll trtmnts xpt mm NCl. Root n shoot prolin ontnts: As n sn in Fig.,, Prolin umultion in root n shoot inrs in NCl trtmnts. Howvr, In s of root it ws signifint for ll trtmnts (**p<.1), xpt mm NCl, ut in shoot thr ws signifint inrs only in 1 n mm trtmnts. 19

4 Curr. Rs. J. Biol. Si., (1): 17-, 1 Lf N ontnts (mg/g DW) + () NCl onntrtions (mm) Lf K ontnts (mg/g DW) + () 1 1 NCl onntrtions (mm) Lf C + ontnts (mg/g) () NCl onntrtions (mm) Root prolin ontnts (nmol/g FW) () NCl onntrtions (mm) Shoot prolin ontnts (mmol/g FW) () 1 1 NCl onntrtions (mm) Fig. : Effts of NCl on lf N + ontnts, (), lf K + ontnts, (), lf C + ontnts, (), root prolin ontnts, () shoot prolin ontnts, () in miz DISCUSSION Slinity strss uss ion toxiity n osmoti imlns, ling to oxitiv strss in plnts (Frouk, 11). Soil slinity is urrnt ioti strss for plnts. Growth inhiition is ommon rspons to slinity n plnt growth is on of th most importnt griulturl inis of slt strss tolrn s init y iffrnt stuis (Pri n Ds, ). Our rsults show tht th growth of Z mys L. slings strt to rs signifintly in 1 slinity trtmnt. Rution in shoot n root growth rts, s wll s iomss osrv in slt strss slings r in onformity with provn ffts of slinity strss. Slinity ffts vlopmnt prosss suh s s grmintion, sling growth n vigor, vgttiv growth, flowring n fruit st (Sirm n Tygi, ). Hsn t l. (11) rport tht rution in iomss n growth n u to highr N + onntrtion in th shoots of Ariopsis thlin n ioni toxiity n rss osmoti potntil. It hs n shown slinity inrs llulr umultion of ROS (Hrnnz n Almns, ). Unlss ontroll, ROS my oxiiz n vntully us mg to protins, lipis n nuli is. Howvr, som of thm n lso srv s signling moluls, if thy r prou trnsintly ut o not umult (Foyr n Notor, ). Thir funtion s signling moluls or th gr of mg us y ROS ultimtly pns on th ln twn thir formtion n rmovl y th ntioxitiv svnging systms (Hrnnz n Almns, ). In this stuy, w prsnt th vin tht slt strss is l to xssiv

5 Curr. Rs. J. Biol. Si., (1): 17-, 1 gnrtion of MDA in th root n lf of Z mys L. slings. Inrs in th lvl of MDA, prou uring proxition of mmrn lipis, is oftn us s n initor of oxitiv mg. Fr ril-inu proxition of lipi mmrns is rfltion of strssinu mg t th llulr lvl (Ngsh n Dvrj, ). As squl, lipi proxition prouts suh s MDA, will umult n svr mmrn mg will invitly our. Similr rsults wr osrv in sltsnsitiv plnts suh s otton (Gosstt t l., 199) n miz (Aror t l., ) unr slinity. MDA onntrtion n ll mmrn mg inrs unr slt strss onition us of lvting of ROS proution. Enzymti ntioxint fns systm n prott plnt lls from oxitiv injury. POD n CAT r th most importnt prottiv nzyms to rmov rtiv oxygn spis. Our rsults show tht in th lvs of miz slings ntioxint nzym (POD n CAT) tivitis inrs t th slt strss. Th highst ntioxint tivity hlp plnts unr nvironmntl strss lik slt strss (Munns n Jms, 3). A orrltion twn th ntioxint nzym tivitis n slinity tolrn wr rport in mny plnts (Ashrf n Ali, ). Bnoglu t l. () rport tht ntioxint nzym tivitis lik CAT n POD hlps plnt to rs MDA onntrtion n ll mg unr slinity strss. In our stuy, th inrs tivitis of ths nzyms wr suggst to insuffiint to ovrom lipi proxition n thus slt-inu oxitiv strss in Z mys L. slings.ymzki t l. (3) show tht High tivity of CAT n POD rs H O lvl in ll n inrs th stility of mmrns n CO fixtion us svrl nzyms of th Clvin yl within hloroplsts r xtrmly snsitiv to H O. A high lvl of H O irtly inhiits CO fixtion. In this stuy rution of iomss n growth in miz slings oul ttriut to oxitiv strssss n inhiition in photosynthti CO fixtion. Morovr, slt strss inu grtr inrs of MDA in roots thn in lvs of Z mys L., suggsting som gr of oxitiv strss, proly u to th high NCl onntrtion n highr lipi proxition in roots. Thrfor, in Z mys L. slt toxiity ws minly onntrt in roots, strong rs of plnt growth ws osrv, ssoit with oxitiv strss in th roots t th highst NCl supply. Also, highr iomss rution in roots oul rlt to th svr inrs of th oxitiv strss in roots in rspons to NCl. It this xprimnt, N + onntrtion inrs, whil K + n C + ontnts in lf wr signifintly rs with th ris of slinity strss. Slt strss is known to inu K + fiiny in plnts us of th omptition of N + n K + for ining sits in trnsport systms tht mit K + uptk (Shl, 3). Mthius n Amtmnn (1999) rport tht for most othr spis, N + is not nssry for growth. Th vilility of som N + is nfiil to th plnt, ut too muh will s mg. Volkmr t l. (199) show tht N+ ompting for K + ining sits in th ytoplsm inhiit mtoli prosss tht pn on K +. It is usully ssum tht N + n ngtivly fft plnt C + rltions. N + n rpl ltrosttilly oun C + in ll wlls n ll mmrns. In ition, C + nutrition n rgult us th osmoti ffts of slinity strss l to rution in trnspirtion, prtiulrly ffting rltivly immoil ions suh s C + in thir root-shoot trnslotion (Mthuis, ). W show in our stuy tht slinity strss inrs prolin umultion in root n shoot of Z mys L. slings. Prolin umultion ours normlly in ytosol in rspons to rought n slinity strss. Aprt from its rol s osmolyt, prolin ontriuts to stiliztion of su-llulr struturs, svnging ROS, n uffring llulr rox potntil unr strss (Ashrf n Fool, ). Rnt stuis suggst tht prolin my ply s n nzym stilizing rol in nol (Bhtthrj n Mukhrj, ) n ru lipi proxition in ri ultivrs (Frhoui t l., 7) unr slt strss. Thrfor, th inrs in th prolin onntrtion of Z mys L. slings spilly in roots, n mintin osmorgultion n prvnt nzym strution unr slt strss. CONCLUSION In onlusion, this stuy show tht slt strss rs iomss n growth in miz plnts u to ioni toxiity n rs osmoti potntil n oxitiv strss. Inrs in th lvl of MDA, prou uring proxition of mmrn lipis, is s n initor of oxitiv mg. Enzymti ntioxint fns systm n prott plnt lls from oxitiv injury. Slt strss is inu K + fiiny in miz plnts us of th omptition of N + n K + for ining sits in trnsport systms tht mit K + uptk. Th inrs in th prolin onntrtion of. slings spilly in roots, n mintin osmorgultion n prvnt nzym strution unr slt strss. REFERENCES Ai, H., 19. Ctls in vitro. Mtho Enzymol., 1: Aror, N., R. Bhrwj, P. Shrm n H.K. Aror,. -Homorssinoli llvits oxitiv strss in slttrt miz (Z mys L.) plnts. Brz. J. Plnt Physiol., (): Ashrf, M. n M.R. Fool,. Rols of glyin tin n prolin in improving plnt ioti strss rsistn. Environ. Exptl. Bot., 9: -1. 1

6 Curr. Rs. J. Biol. Si., (1): 17-, 1 Ashrf, M. n Q. Ali,. Rltiv mmrn prmility n tivitis of som ntioxint nzyms s th ky trminnts of slt tolrn in nol (Brssi npus L.)Environ. Exp. Bot., 3: -73. Bnoglu, E., F. Eyiogn, M. Yul n H.A. Oktm,. Antioxint rspons of shoots n roots of lntil to NACL Slinity strss. Plnt Grth. Rgu., : Bts, L.S., S.P. Wlrn n I.D. Tr, Rpi trmintion of fr prolin for wtr-strss stuis. Plnt Soil, 39: -. Bhtthrj, S. n A.K. Mukhrj,. Slt strss inu ytosolut umultion, ntioxint rspons n mmrn triortion in thr ri ultivrs uring rly grmintion. S Si. Th., 3: Brfor, M.M., 197. A rpi snsitiv mtho for th quntittion of mirogrm quntitis of protin utilizing th prinipl of protin-y ining. Anl. Biohm., 7: -. Frhoui, R., F. Shrifzh, K. Poustini, M.T. Mkkizh n M. Kohkpor, 7. Th ffts of NACL priming on slt tolrn in nol (Brssi npus) slings grown unr slin onitions. S Si. Th., 3: Frouk, S., 11. Osmoti justmnt in wht flg lf in rltion to flg lf r n grin yil pr plnt. J. Plnt Physiol. Biohm., 7(): Filing, J.L. n J. Hll, 197. A iohmil n ytohmil stuy of proxis tivity in roots of Pisum stivum. J. Exp. Bot., 9: Foyr, C.H. n G. Notor,. Oxint n ntioxint signl ling in plnts: A r-vlution of th onpt of oxitiv strss in physiologil ontxt. Plnt Cll Environ., : Gosstt, D.R., E.P. Millhollon n M.C. Lus, 199. Antioxint rspons to NACL strss in slt-tolrnt n slt-snsitiv ultivrs of otton. Crop Si., 3: Hsn, E., B.H. Krim, C. Jn, M.B. Srgi n A. Chly, 11. Erly ffts of slt strss on th physiologil n oxitiv sttus of Ckil mritim (hlophyt) n Ariopsis thlin (glyophyt). Physiol. Plntrum., 1: -13. Hrnnz, J.A. n M.S. Almns,. Short-trm ffts of slt strss on ntioxint systms n lf wtr rltions of p lvs. Physiol. Plnt, 11: 1-7. Lvinsh, G.,. Biologil sis of iologil ivrsity: Physiologil pttions of plnts to htrognous hitts long s ost. At. Univ. Lt., 71: Mthius, F.J.M. n A. Amtmnn, K + nutrition n N + toxiity: Th sis of llulr K + /N + rtios. Ann. Bot., : Mthuis, F.J.,. Th rol of monovlnt tion trnsportrs in plnt rspons to slinity. J. Exp. Bot., 7: Mittlr, R.,. Oxitiv strss; ntioxints n strss tolrn. Trn. Plnt Si., 7: -1. Munns, R. n R.A. Jms, 3. Srning mthos for slinity tolrn: As stuy with ttrploi wht. Plnt Soil, 3: 1-. Ngsh, B.R. n V.R. Dvrj,. High tmprtur n slt strss rspons in Frnh n (Phsolus vulgris). Aust. J. Crop Si., (): -. Pri, A.K. n A.B. Ds,. Slt tolrn n slinity fft on plnts: A rviw. Eotoxiol. Environ. Sf., : Prviz, A. n S. Stywti,. Slt strss n phytoiohmil rsponss of plnts: A rviw. Plnt Soil Environ., : Quils, M.J. n N.I. Lópz,. Photoinhiition of photosystms I n II inu y xposur to high light intnsity uring ot plnt grown ffts on th hloroplsti NADH hyrogns omplx. Plnt Si., 1: 1-3. Sirm R.K. n A. Tygi,. Physiologil n molulr iology of slinity strss tolrn in plnts. Curr. Si., : 7-. Shl, S., 3. Rgultion of potssium trnsport in lvs: From molulr to tissu lvl. Ann. Bot., 9: 7-3. Vlntovi, P., M. Luxov, L. Kolrovi n O. Gsprikor,. Efft of osmoti strss on omptil soluts ontnt, mmrn stility n wtr rltion in two miz. Plnt Soil Environ., (): Volkmr, K.M., Y. Hu n H. Stppuhn, 199. Physiologil rsponss of plnts to slinity: A rviw. Cn. J. Plnt Si., 7: Whi, A. n T.J. Clos, 7. Exprssion of hyrins unr ht strss n thir rltionship with wtr rltions of sugrn lvs. Biol. Plnt., 1: Ymzki, J., A. Ohshi, Y. Hshimoto, E. Ngishi, S. Kumgi, T. Kuo, T. Oikw, E. Mrut n Y. Kmimur, 3. Effts of high light n low tmprtur uring hrsh wintr on nl photomg of Ais mrisii growing t th forst limit on Mt. Norikur in Cntrl Jpn. Plnt Si., 1: 7-. Zhng, F., Y. Wng, Y. Yng, H. Wu, D. Wng n J. Liu, 7. Involvmnt of hyrogn proxi n nitri oxi in slt rsistn in th lluss from Populus uphrti. Plnt Cll Environ., 3: 77-7.

Antioxidant enzymes and physiological traits of Vicia faba L. as affected by salicylic acid under salt stress

Antioxidant enzymes and physiological traits of Vicia faba L. as affected by salicylic acid under salt stress Journl of Mtrils n Environmntl Sins ISSN : 8-58 Copyright 17, Univrsity of Mohmm 1r Ouj Moroo JMES, 17 Volum 8, Issu 7, Pg 59-563 http://www.jmtrnvironsi.om / Antioxint nzyms n physiologil trits of Vii