207 Bulrin Journl o Ariulturl Sin, 18 (No 2) 2012, 207-218 Ariulturl Amy Slt tolrn urin vttiv rowth in ross o tomto n t o ytoplsm in rspons to slt tolrn A. Turhn 1 * n V. Sniz 2 1 Univrsity o Ulu, Dprtmnt o Ariultur n Animl Sin, Mustkmlps Votionl Shool, 16500 Burs, Turky 2 Univrsity o Ulu, Dprtmnt o Hortiultur, Fulty o Ariultur, 16059 Burs, Turky Astrt Turhn, A. n V. Sniz, 2012. Slt tolrn urin vttiv rowth in ross o tomto n t o ytoplsm in rspons to slt tolrn. Bul. J. Ari. Si., 18: 207-218 Th im o th prsnt stuy ws th trmintion o slt tolrn irns mon tomto rosss y usin morpholoil, physioloil prmtrs n th intiition ytoplsm rlt to slt tolrn mhnisms urin th vttiv sts o tomto. In th stuy, slt tolrnt (,, ) n slt snsitiv (, ) tomto notyps wr us s plnt mtril. Riprol rosss wr m twn slt tolrnt n slt snsitiv tomto notyps. Cross omintions wr xpos to 0 (ontrol), 8 n 12 S m -1 NCl or 40 ys. At th n o th xprimnt, th lvls o l, stm n ry root wihts n C 2+, K + n N + onntrtions wr trmin. Th rltionships twn th lvls o slinity n root, stm, l umultion; n twn K +, C 2+, N + n root, stm n ry l wihts wr invstit. C 2+ /N + n K + /N + rtios wr lso lult. A tolrn inx ws lult or vry sinl notyp in root, stm n l ry wihts n in th K + /N + n C 2+ /N + prmtrs o ths orns. In this stuy, tolrn inx wr us s slt slt tolrnt tomto rosss t irnt slt onntrtions. Thr ws lr vrition in root, stm n l ry wiht n K + /N +, C 2+ /N + TI mon 12 ross omintions unr inrsin slinity. Th hihst tolrn inx vlus wr nrlly otin rom th omintion o tolrnt notyps (, n ) n snsitiv notyp. Cytoplsm is nothr tor tht plys n importnt rol in th slt tolrn. Gnrlly, in riprol rosss wr m twn slt tolrnt n slt snsitiv tomto notyps;,, n ytoplsm positivly t tolrn inx vlus n mns whih thy wr lss t y slinity. Ky wors: tomto, notyps, rossin, slt tolrn, ytoplsm Introution Slinity ts lmost vry spt o th physioloy n iohmistry o plnts n siniintly rus yil. As slin soils n slin wtrs r ommon roun th worl, rt ort hs n vot to unrstnin physioloil spts o tolrn to slinity in plnts, s sis or plnt *Corrsponin uthor: E-mil: turhn@ulu.u.tr
208 A. Turhn n V. Sniz rrs to vlop slinity-tolrnt notyps. In spit o this rt ort, only smll numr o ultivrs, prtilly tolrnt to slinity, hv n vlop (Curtro t l., 2006). Ovr th yrs, plnt rrs hv vlop vry proutiv ultivrs o hih qulity, rsistnt to isss n pt to th mns o th mrkt n o inustry (Grnillo t l., 1999). Nw ultivrs r or slt tolrn hs to not only slt tolrnt, ut lso hiv th sm ttriuts o proutivity n qulity sn in morn ultivrs (Curtro t l., 2006). Th ultivt tomto (Lyoprsion sulntum Mill.) is in th mily Soln n is wily istriut nnul vtl rop. Tomto is lso vlul or rin stuis, us mkin rosss twn wil n ultivt tomto plnts is simpl n its wil rltivs provi rih rmplsm pool. Currntly, th tomto rop is rown in wi vrity o limts rnin rom th tropis to within w rs o th Arti Cirl (Fool, 2004). Tomto proution hs n limit y hih lvl o slinity in th soil or irrition wtr. Tomto is snsitiv to mort lvls o slinity lik most rop plnts. All sts o plnt vlopmnt inluin s rmintion, vttiv rowth n rproution show snsitivity to slt strss n onomi yil is ru unr slt strss (Jons t l., 1986; Ms, 1986; Bolrin t l., 1993). Most rsrh on tomto slt tolrn urin vttiv st ws ous minly on th physioloil rsponss to slt strss (Fool, 2004). Thr is positiv orrltion twn tomto yil n plnt siz urin vttiv rowth unr slt strss (Bolrin t l., 1993; Pstrnk t l., 1979), initin th importn o slt tolrn urin th vttiv st. Slinity strss rsults in lr stuntin o plnts (Hrnnz t l., 1995; Chrin t l., 1999; Tkmur t l., 2002). Slt strss lso rsults in onsirl rs in th rsh n ry wihts o lvs, stms, n roots (Hrnnz t l., 1995; AliDinr t l., 1999; Chrtzoulkis n Klpki, 2000). Inrsin slinity is ompni y siniint rutions in shoot wiht, plnt hiht, n numr o lvs pr plnt, root lnth, n root sur r pr plnt in tomto. Hih slinity in th root zon svrly imps norml plnt rowth n vlopmnt, rsultin in ru rop proutivity or rop ilur (Fool, 2004). Th physil rowth prmtrs suh s shoot rsh n ry wiht, root rsh n ry wiht r mor orrlt with rop slt tolrn t rly rowth sts n n us s srnin/sltion ritri (Irhim, 2003). Bolrin t l. (1991), Fool (1996) hv init tht shoot rowth unr slinity rltiv to ontrol is th st prtil initor o slt tolrn in tomto. Th lttr stuy sust tht tissu ion ontnt n ion sltivity wr oo sltion ritri or rin or slt tolrn in tomto (Ahsn t l., 2000). Potssium sltivity ovr N + ws lso rport s oo initor o slt tolrn in stuy o svrl notyps o th ultivt n wil spis o tomto (Curtro t l., 1992). Slinity tolrn is ntilly ontroll n trnsmittl mon notyps. Whol plnt prormn miht mor usul sltion ritri or slinity tolrn thn usin sinl trit hih K + /N + rtio, Low N +. Aorin to Fool (1997) nlysis o th prntl, ilil n kross popultions o ross twn slt-snsitiv tomto lin (UCT5) n slt-tolrnt primitiv ultivr (PI174263) init tht rowth unr slt strss ws positivly orrlt with l C 2+ ontnt n ntivly orrlt with l N + ontnt (Fool,, 1997). Gnrtion mns nlysis o ths popultions init tht unr slt strss, umultion o oth N + n C 2+ in th l ws ntilly ontroll with itivly in th mjor nti omponnt. Th rltionship twn slt tolrn n l ion omposition in th ultivt n thr wil spis o tomto prompt Srn t l. (1993) to onlu tht ry mttr proution
Slt Tolrn Durin Vttiv Growth in Cross o Tomto n Et o Cytoplsm... 209 unr slt strss ws positivly orrlt with K + / N + rtio. Th ntis o physioloil hrtrs tothr with othr tolrn omponnts rlt to mtoli nss inst slinity hv to stui in orr to vn th rin o notyps tolrnt to slinity (Curtro t l., 2006). Two mjor pprohs hv n propos n mploy to minimiz th ltrious ts o hih soil/wtr slinity in riultur (Epstin t l., 1980). First, thnoloil pproh o implmntin lr ninrin shms or rlmtion, rin n irrition with hih-qulity wtr. Althouh this pproh hs n tiv in som rs, th ssoit osts r hih n it otn provis only tmporry solution to th prolm. Th son pproh, ntilly moiyin omstit rops y rin n sltion or improv slt tolrn. Brin or slttolrnt notyps tht n row mor iintly thn th onvntionl vritis unr slin onitions (Fool, 2004). Th purpos o this stuy ws th trmintion o slt tolrn irns mon tomto rosss y usin morpholoil, physioloil prmtrs n th intiition ytoplsm rlt to slt tolrn mhnisms urin th vttiv sts o tomto. Mtrils n Mthos Th invstitions wr onut unr rnhous onitions t th prtmnt o Hortiultur n Soil Sins t Ulu Univrsity. Two xprimnts (2005 2006) hv n onut with tomto notyps. Gnti mtrils: As nti mtril, iv tomto (Lyoprsion sulntum Mill.) notyps wr us. Tomto notyps ollt rom irnt prts o Turky n mintin y An Ariultur Rsrh Institut, Izmir. Crosss: Riprol rosss wr m twn slt tolrnt (,, ) n slt snsitiv (, ) tomto notyps, Lyoprsion sulntum Mill. Plnts rom thos notyps hv n rown unr rnhous onitions n kpt until lowrin n hrvstin unr optiml nutritionl onitions. Mnul rosss hv n rri out usin slt tolrnt s polln onor n slt snsitiv (, ) tomto notyps s polln rptor. Polln rom h notyp ws ollt y rkin n mixin t lst tn mtur nthr ons in Ptri ish. Th lowr o th ml prtnr ws msult just or nthsis. Pollintion ws m y ruin th polln otin on th sur o th stim o th msult lowr (Yornov t l., 1983; Shinohr, 1989). Growth vlution n slt strss trtmnt: twlv rosss s wr initilly rmint in ornilly nrih pt with vrmiulit ovr to ilitt rtions, in opn plsti trys. Th vr rnhous tmprturs wr 17 n 25 o C t niht n y, rsptivly, ut th tmosphri humiity ws mintin t 70%. 35 ys tr rmintion, slins (3 4 tru l) wr trnsplnt into 14m plsti pots ill with pt: prlit (1:1 on volum sis) homonous mixtur. Th trnsplnt plnts wr trnsrr to th rnhous. Th slins wr rown in pt/prlit pproximtly or 40 ys. Whn th plnts h vlop 3 4 tru lvs (35 ys tr trnsplntin), pplitions o Holn solution ontinin 0 (ontrol), 8 n 12 S m -1 NCl wr strt vi rip irrition. Plnts wr irrit with thir rsptiv solution 1 2 tims pr y. It ws ttmpt to kp th quntity o rin wtr t 30 40% o th mount o nutrint solution ppli. Th ltril onutivity in th nvironmnt (pot, wtr, soil) ws rsptivly 1.5 S m -1 (ontrol), 8±0.5 S m -1 n 12±0.6 S m -1. Th slt lvl ws rully inrs ovr 4 ys to voi osmoti shok. Plnts wr rown in ontroll rnhous with y/niht vr tmprtur o 28.3 o C, vr rltiv humiity o 70.4%, n vr
210 A. Turhn n V. Sniz photoprio o 16 h. Th xprimnts wr onut with 3 rplits. Thr ws on plnt in h pot (1 l), with 4 pots in h rplit. Growth msurmnt: t th n o th xprimnts (orty ys tr trnsplntin), iniviul plnts rom oth slt strss n non-strss trtmnts wr hrvst n plnts wr sprt into root, stm n l. Plnt mtrils wr ovnri t 70 o C or two ys n ry wihts (DW) wr trmin (). Tissu ion nlysis: C 2+, K + n N + onntrtions wr lso msur on nitri-prhlori i ists o root, stm n l tissu y Eppnor Elx mol Flymotomtry. A C 2+ /N + n K + /N + rtio ws lult. In this stuy, Tolrn Inx (TI), vlop y LRos t l. (1989) is us to show nrl ttitu o notyps towrs NCl onntrtions, n, tr limintin irns, whih my rsult o notypi turs, only to ompr thir prormns inst slt onntrtions, whih r ppli. Tolrn Inx is lult or vry notyp on y on in root (DWTI), stm (DWTI) n l (DWTI) ry wihts n in C 2+ / N + n K + /N + prmtrs o ths orns. Tolrn Inx (TI) = 100 + n [X (Tx /T 0 )100] Whr n= numr o pplitions Tx= vlu o th prmtr s trmin on strss plnts T 0 = vlu o th prmtr s trmin on plnts without strss (ontrol) Sttistil nlysis: t wr nlyz usin MSTAT-C (vrsion 2.1, Mihin Stt Univrsity, 1991) n Minit 14.0 sotwr. Anlysis o vrin (ANOVA) ws onut n siniin o irns mon trtmnt ws tst usin th lst siniint irn (LSD). Dirns wr lr siniint t P<0.05 proility lvls y th F tst. Th F-prott LSD lult t 0.05 proility lvls. Rsults n Disussion Growth rspons In this stuy, th vlus o th ry wiht o th 12 tst tomto rosss wr oun to lowr with th ris o slinity lvl. Dt prsnt in Tl 1 shows tht th ry wiht oviously rs in ll ross omintions with slinity on omprin with ontrol. This similr to th rsult otin y Hjr t l. (2006), Mio t l. (2006), Li n Stnhllini (2001), Curtro n Frnnz-Munoz (1999). Furthrmor, thr is positiv orrltion twn tomto yil n plnt siz urin vttiv rowth unr slt strss (Bolrin t l., 1993; Pstrnk t l., 1979), initin th importn o slt tolrn urin th vttiv st. Th physil rowth prmtrs suh s rsh n ry shoot wiht, rsh n ry root wiht r mor orrlt with rop slt tolrn t rly rowth sts n n us s srnin/sltion ritri (Irhim, 2003). In xprimnt, tolrn Inx ws lult or vry sinl ross in root (DWTI), stm (DWTI) n l (DWTI) ry wihts prmtrs o ths orns. Anlysis o vrin rvl tht siniint irns mon th tst tomto rosss isply lr vrition in tolrn to trtmnts s on th Tolrn Inx (TI). Thr ws lr vrition in root, stm n l ry wiht TI twn 12 notyps unr inrsin slinity. Th rosss [ ] n [ x ] h th hihst root, stm DWTI n l DWTI t ll onntrtion lvls, rsptivly. It ws trmin tht th rosss, th most tolrnt towrs slt onntrtions, thus h th hihst TI. On ontrry, th lowst root, l DWTI n stm DWTI wr oun in [ x ] n [ ], rsptivly. Th [ x ] n [ ], howvr, wr lss unr slin onition ompr with th othr tn rosss (Tl 1, Fiur 1A, 1B, 1C).
Slt Tolrn Durin Vttiv Growth in Cross o Tomto n Et o Cytoplsm... 211 Ion umultion Slt strss uss xtnsiv physioloil n iohmil hns in plnt. Btwn ths hns, istriution in irnt orns o th plnt s intk o ions lik K +, C +2 n N + is mphsiz (Aziz t l., 1999). In nrl, C +2 n K + onntrtions rs with slinistions (Cruz, 1990). In onitions whih slt onntrtion is hih, th plnt ts mor N + ion thn it ns (Lvitt, 1980). Dspit N + ion hs no t on physioloil inints in th plnt, K + n C +2 ions ply ky rol. K + n N + ion umultion in tomto n plnt tissus is importnt in slt tolrn n N + ion inrss with slinity inrs, spit K + ion mount rss (Cruz t l., 2002; Dsn t l., 2002; Ysr, 2003). Also in rsrhs prorm on th sm sujt it is trmin tht, th tomto kins whih kp N + umultion limit n o whih K + /N + n C +2 /N + rtios r hih r slt tolrnt n this prmtr is rlil in slt tolrn (Rnl, 1992; Munns n Jms, 2003; Jun t l., 2005; Turhn t l., 2009). In ition tht, Physioloil ritri whih hv n sust or us s potntil initors o slt tolrn n in tomto inlu tissu wtr potntil, tissu ion ontnt, K + /N + rtio, (Fool, 2004; Ahsn t l., 2000). In this stuy, nlysis o vrin rvl tht siniint irns mon th tomto rosss or tissus (root, stm n l) K + /N + n C +2 /N +. In Tl 1 Dry wiht (DW, ) o vlus n thir Tolrn inx (TI) in root, stm, l tissu o tomto rosss, unr irnt NCl onntrtions Root DW Stm DW L DW Crosss Control 8 S 12 S m -1 m -1 TI Control 8 S 12 S m -1 m -1 TI Control 8 S 12 S m -1 m -1 TI x 4.05 h 2.69 1.83 1323.15 6.54 4.66 2.90 1355.25-12.31-7.19 6.72 1380.09 x 4.43-2.83-1.62 1201.76 6.00 4.05 2.22 1233.52 12.75 7.01 5.87 1242.59 4.29-3.60 1.74-1407.43 5.99 4.75 2.58 1403.65 12.73 7.52 6.16-1317.87-4.10 h 2.95-1.87 1370.98 6.36 3.99 2.61 1243.48 13.36 6.79 6.41-1233.55 4.06 h 2.97-1.63 1316.34 6.35 4.74 2.62 1347.15-11.67 6.94 5.98-1366.47 4.62 2.81 1.65 1169.82 5.88 4.40 2.08 1274.32 12.60 7.05 6.04-1274.01-4.72 3.57 1.83 1321.27 7.15 5.61 2.83 1369.24 12.21-7.62 6.33-1373.30 4.36-3.02 1.78-1296.31 6.75 4.78 2.77 1309.26-11.92 7.26 5.94 1342.09-4.51 3.14 1.88 1308.87 6.57 4.73 1.71 1139.10 11.78 8.38 5.37 1373.53 x 4.01 h 2.89-1.26 1210.33 6.32 4.58 2.38 1283.45 14.23 7.36 6.78 1240.73 x 4.94 2.89-1.70-1133.19 6.66 4.44 2.81 1290.25-14.30 7.68 6.20-1199.97 x 4.21 -h 2.80 1.58 1234.34 7.07 5.25 2.01 1185.31 12.71 7.51 5.69 1259.78 - Mns ollow y th sm lttr r sttistilly not siniint (Dunn s multipl rn tst, P=0.05)
212 A. Turhn n V. Sniz A Tolrn inx (TI) B Tolrn inx (TI) C Tolrn inx (TI) 1500 1400 1300 1200 1100 1000 1500 1400 1300 1200 1100 1000 1500 1400 1300 1200 1100 1000 1 2 3 4 5 6 7 8 9 10 11 12 Tomto rosss R oot 1 2 3 4 5 6 7 8 9 10 11 12 Tomto rosss S tm 1 2 3 4 5 6 7 8 9 10 11 12 Tomto rosss L Fi. 1. Dry wihts Tolrn inx (TI) rspons o th tomto rosss on ontrol n 8, 12 S m -1 NCl onntrtions (A) root ry wiht tolrn inx, (B) stm ry wiht tolrn inx, (C) l ry wiht tolrn inx. Tomto ross omintions wr s ollows: 1 ( x ), 2 ( x ), 3 ( ), 4 ( ), 5 ( ), 6 ( ), 7 ( ), 8 ( ), 9 ( ), 10 ( x ), 11 ( x ), 12 ( x ) omprison with th ontrol plnts, tissus o slttrt plnts umult mor N + ut lss K + n C +2, rsultin in lowr K + /N + n C +2 /N + rtios. K + /N + n C +2 /N + rtios r trmin or th plnt s root, stm n l prts sprtly. Tomto rosss with hihr K + /N + n C +2 /N + rtios tn to rt in th hihr TI showin lss m n thos with lowr K + /N + n C +2 / N + in lss TI. K + /N + TI n C +2 /N + TI wr us to hrtriz th ovrll tolrnt o plnt to slinity. In root tissu, ross [ ] show th hihst root K + /N + TI n C +2 /N + TI. In ition this rsult, ross [ ], rw ttntion with root C +2 /N + TI tht is hihr thn othr rosss. It is ovious tht th [ ] n [ ] rosss show hih r o rspons to slinity onitions muh mor thn th othr tn rosss. On th ontrry, th lowst root K + /N + TI n C +2 /N + TI vlus wr trmin in [ x ] n [ x ] rosss, rsptivly (Tl 2 n Fiur 2A) n this omintions o [ X ] n [ X ] show low tolrn. Anothr plnt tissu on whih th t o slt strss wr osrv ws th stm. Similr rsults wr otin in stm K + /N + TI n C +2 /N + TI. As shown in Tl 3; ll o slt trtmnt, stm s hihst K + /N + TI n C +2 /N + TI vlus wr prsnt y th [ ] ross, th lowst in [ x ] (K + /N + TI) n [ x ] (C +2 /N +1 TI) (Fiur 2B). Th rltionship o TI with th onntrtions o NCl n us s n initor o slt tolrn. Aorin to Turhn n Sniz (2010), TI oul us s simpl n tiv mtho to slt slt tomto ultivrs. Ahsn t l. (2000) sust tht ion ontnt oul us s rin tool or sltin slttolrnt notyps. Thr or it is vry importnt to know th xtnt o vrition in N +, K + in lvs. In ition tht, L K + onntrtion n K + / N + rtio hv n th most stui prmtrs rlt to slt tolrn in tomto (Curtro t l.,
Slt Tolrn Durin Vttiv Growth in Cross o Tomto n Et o Cytoplsm... 213 1992; Curtro n Frnnz-Munoz, 1999; Fool, 2004). In irnt tomto rosss stui in work, th hihst l K + /N + TI n C +2 /N + TI ppr in th [ ] ross, whrs th lowst wr prsnt y [ x ] n [ ], rsptivly. [ ] tomto ross v oo prormn whn it ws trt y NCl onntrtion (Tl 4 n Fiur 2C). Th t o ytoplsm on slt-tolrn In orr to osrv whthr thr is ytoplsmi t on slt tolrn or not, riprol rosss wr prpr with tolrnt, 40442, n snsitiv, tomto notyps. Cross omintions wr ppli with irnt slt onntrtion (ontrol, 8 n 12 S m -1 ). Atr th prti, th rt o K + /N + n C +2 /N +, ry wiht ws tt in root, stm n l tissu o omintions n Tolrn inx vlus o ths prmtrs wr lult. In this stuy, it ws tt tht ytoplsm hs irnt ts on irnt inx vlus. For xmpl, it ws tt tht ytoplsm os not hv n importnt t on ry wiht n root n l K + /N + TI - C +2 /N + TI, stm C +2 /N + TI prmtrs o tolrnt n snsitiv, whrs it hs importnt t on stm K + /N + TI prmtrs. In othr wors, it ws tt tht ytoplsm o notyp h positiv t on stm C +2 / N + TI (Tl 5). It ws oun tht ytoplsm hs no t on ry wiht, root n stm K + /N + TI prmtrs o [ x ] n [ x ] ross omintions otin rom notyp n snsitiv riprol mtril. Howvr, it ws tt tht ytoplsm hs quit importnt t on root, stm C +2 /N + TI, l K + /N + TI n C +2 / N + TI; n this positiv t is us y notyp in l K + /N + TI n C +2 /N + TI, stm C +2 /N + TI; n y notyp in root C +2 / N + TI (Tl 5). A B C Tolrn inx (TI) Tolrn inx (TI) Tolrn inx (TI) 700 650 600 550 500 450 400 350 300 900 800 700 600 500 400 300 1000 900 800 700 600 500 400 1 2 3 4 5 6 7 8 9 10 11 12 Tomto rosss K /N C /N 1 2 3 4 5 6 7 8 9 10 11 12 Tomto rosss K /N C /N 1 2 3 4 5 6 7 8 9 10 11 12 Tomto rosss K /N C /N Fi. 2. K + /N + n C 2+ /N + rtios Tolrn inx (TI) rspons o th tomto rosss on ontrol n 8, 12 S m -1 NCl onntrtions. (A) root K + /N + n C2 + /N + tolrn inx, (B) stm K + /N + n C2 + /N + tolrn inx, (C) l K + /N + n C2 + /N + tolrn inx. Tomto ross omintions wr s ollows: 1 ( x ), 2 ( x ), 3 (, 4 ( ), 5 ( ), 6 ( ), 7 ( ), 8 ( ), 9 ( ), 10 ( x ), 11 ( x ), 12 ( x )
214 A. Turhn n V. Sniz In nothr mtril, tolrnt notyp n snsitiv notyp wr riprol ross. It ws tt tht ytoplsm hs importnt ts on root n stm ry wiht vlus n root n l K + /N + n C +2 /N + TI vlus. It ws tt tht ths positiv ts wr spilly us rom th ytoplsm o notyp in root n stm ry wiht n K + /N + n C +2 / N + TI; n rom th ytoplsm o notyp in l K + /N + TI n C +2 /N + TI. Contrry to this, it ws tt tht ytoplsm hs no t on l ry wiht n stm K + /N + n C +2 /N + tolrn inx (Tl 5). Cytoplsm os not importnt t on root n stms ry wiht, K + /N + (root, stm, l) n C +2 /N + (l) tolrn inx; on rosss otin rom riprol mtril twn n tolrnt n snsitiv notyps. It n osrv rom Tl 5 tht ths positiv ts r us y ytoplsm in root, ry wiht n root K + /N + ; y notyp in stm; ry wiht n K + /N +, l; K + /N + n C +2 /N + tolrn inxs. With th riprol mtrils whih r tolrnt notyp n snsitiv notyps; omintions o [ ] n [ ] rosss wr otin. In ths omintions, it ws oun tht ytoplsm hs importnt ts on th prmtrs othr thn root n stm ry wiht n l K + /N + prmtrs. tol- Tl 2 Conntrtion o K +, C 2+, N +, K + /N +, C 2+ /N + n K + /N +, C 2+ /N + Tolrn inx (TI) in th root o th tomto rosss unr irnt NCl onntrtions Crosss x x x x x NCl Conntrtion (S m -1 ) Control 8 12 K + C +2 N + K + / N + C +2 / N + K + C +2 N + K + / N + C +2 / N + K + C 2+ N + K + / N + C 2+ / N + K + /N + C 2+ /N + 2.67 2.15 0.34 7.83 3.72 2.57 0.53 7.02 2.49 3.06 0.57 4.35 i 2.75 2.66 0.48 5.77 h 2.93 2.52 0.40 7.31 3.32 3.07 0.55 6.03 3.58 2.97 0.40 8.98 3.32 3.36 0.39 8.50 3.22 3.10 0.41 7.82 3.76 2.26 0.56 6.79 3.35 2.43 0.55 6.15 3.65 2.98 0.57 6.47 6.30 4.85 5.36 5.59 6.25 5.58 7.46 8.64 7.54 4.09 4.51 5.29 2.66 2.14 1.47 1.81 j 1.44 ij 2.55 1.96 1.79 1.42 k-m 1.09 j-l 1.68 1.96 1.09 1.24 1.80 2.00 2.00 2.62 1.24 1.35 1.60 1.66 2.68 2.08 1.61 2.21 2.11 1.97 1.72 2.71 1.24 2.58 1.84 1.99 1.82 1.56 2.66 1.29 1.76 2.05 1.54 kl 0.62 q-t 1.62 j-l 0.81 pq 1.66 jk 1.12 no 1.39 lm 1.29 mn 0.69 p-r 0.63 q-s 1.80 hi 0.90 l-o 2.12 h 0.96 k-n 1.29 j-k 1.79 1.59 2.74 1.43 1.31 2.96 1.21 1.66 2.36 1.11 1.04 2.56 1.46 1.63 2.11 1.10 0.76 2.53 2.02 1.78 2.21 0.65 q-s 0.48 r-u 0.51 r-u 0.44 s-u 0.69 p-r 0.43 s-u 0.92 op 1.07 j-m 2.00 1.99 2.15 0.93 op 2.19 0.92 k-o 0.59 n-s 0.86 l-p 1.53 1.54 2.38 1.46 1.61 2.63 1.01 1.49 3.07 1.19 0.61 3.09 0.64 q-s 0.55 r-u 1.61 u 0.38 tu 0.58 o-s Mns ollow y th sm lttr r sttistilly not siniint (Dunn s multipl rn tst, P=0.05) 535.52 TI 543.88 0.44 q-t 494.99 540.77 0.71m-r 675.29 676.10 0.41 r-t 426.27 466.29 0.78 l-r 541.70 678.68 0.30 st 444.63 452.85 0.81 l-q 521.63 0.93 k-o 519.72 487.09 478.12 0.64 n-s 490.42 585.77 0.61 ns 502.86 612.72 2.10 p-t 404.42 496.93 0.20 t 399.45 425.25
Slt Tolrn Durin Vttiv Growth in Cross o Tomto n Et o Cytoplsm... 215 rnt notyp ytoplsm spilly hs spilly mor positiv t on l ry wiht, root n stm K + /N + n C +2 /N +, l C +2 /N + tolrn inx whn ompr with snsitiv notyp (Tl 5). Dirnt slt onntrtions wr onut on [ ] n [ x ] ross omintions otin rom riprol mtril twn tolrnt notyp n othr snsitiv notyp. It ws tt tht ytoplsm hs positiv ts on stm ry wiht, root n l tissus K + /N + n C +2 /N + tolrn inxs n this t ws us y notyp. On th othr hn, s it n osrv rom Tl 5; thr is no t o ytoplsm on root n l ry wiht, stm K + /N + n C +2 /N + tolrn inxs. It ws sn on th ntir ross omintions tht whil ytoplsm hs importnt t on n inx vlu, it os not hv suh importnt t on nothr inx vlu in th sm ross omintion. Thror usin only on inx vlu tht is tolrnt towrs slt woul show th t o ytoplsm on slt tolrn ttr. Conlusions In this stuy, th ry wiht oviously rs in ll rosss with slinity on omprin with ontrol. Th N + onntrtion o th 12 Tl 3 Conntrtion o K +, C 2+, N +, K + /N +, C 2+ /N + n K + /N +, C 2+ /N + Tolrn inx (TI) in th stm o th tomto rosss unr irnt NCl onntrtions Crosss x x x x x NCl Conntrtion (S m -1 ) Control 8 12 TI K + C +2 N + K + / N + C +2 / N + K + C +2 N + K + / N + C +2 / N + K + C 2+ N + K + / N + C 2+ / N + K + / N + C 2+ / N + 3.10 2.48 0.51 6.07 4.86 2.04 1.81 1.71 1.19 1.06 1.99 1.20 1.82 1.10 0.66 623.96 588.61 3.37 2.44 0.46 7.35 5.32 1.31 0.83 1.31 1.01 0.63 1.25 0.63 2.70 3.47 3.02 0.43 8.13 7.06 2.37 2.62 1.23 1.93 2.13 1.34 1.49 2.01 3.77 2.47 0.56 6.76 4.43 h 1.74 0.87 1.30 1.34 0.67 1.05 0.72 2.60 0.46 0.67 0.40 4.15 2.49 0.47 8.94 5.37 2.60 2.11 0.85 3.20 2.59 1.92 1.54 1.73 1.11 4.50 2.61 0.52 8.70 5.03 1.57 0.95 1.16 1.35 0.82 1.01 0.88 2.98 3.58 2.54 0.46 7.89 5.60 0.34 0.23 435.51 0.74-539.06 0.28 480.56 0.89 2.26 1.86 1.67 1.36 1.12 1.85 1.69 1.78 1.04 0.95 3.33 2.87 0.42 7.88 6.80 2.51 2.84 1.30 1.93 2.19 1.57 1.39 2.31 4.07 2.53 0.42 9.82 6.10 2.85 2.44 0.96 2.98 2.55 1.98 1.50 2.51 3.51 2.50 0.43 8.19 5.81 1.51 0.70 1.38 1.09 4.80 2.71 0.64 7.46 4.21 h 3.09 0.91 1.50 2.05 0.51 1.22 0.61 2.48 0.60 1.16 0.81 2.79 4.36 2.75 0.54 8.05 5.08 1.39 1.10 1.39 1.00 0.79 1.13 0.73 2.56 Mns ollow y th sm lttr r sttistilly not siniint (Dunn s multipl rn tst, P=0.05) 0.68 0.79 0.49-0.41 0.44 685.64 0.29 421.93 546.78 0.60 550.29 0.60 590.32 0.25 429.23 0.29 537.10 0.28 415.51 397.60 617.92 446.09 835.72 452.41 614.74 614.04 702.38 371.09 448.01 441.70
216 A. Turhn n V. Sniz Tl 4 Conntrtion o K +, C 2+, N +, K + /N +, C 2+ /N + n K + /N +, C 2+ /N + Tolrn inx (TI) in th l o th tomto rosss unr irnt NCl onntrtions NCl Conntrtion (S m -1 ) Crosss Control 8 12 TI K + C +2 N + K + / N + C +2 / N + K + C +2 N + K + / N + C +2 / N + K + C 2+ N + K + / N + C 2+ / N + K + /N + C 2+ / N + x 4.30 3.24 0.54 7.93 5.97 3.29 3.00 1.12 2.95 2.69 1.71 2.71 1.61 1.06 1.68 708.79 949.59 x 4.13 3.49 0.63 6.62 5.58 4.03 2.22 1.85 2.19 1.20 1.29 2.07 2.04 0.63 1.02 629.32 641.94-4.77 3.82 0.70 6.84 5.46 3.27 3.00 1.55 2.12-1.95 2.20 2.28 2.78 0.79-0.82 637.28 715.22 4.86 4.33 0.73 6.63 5.91 3.83 2.58 2.20 1.74 1.17 1.82 1.44 3.01 0.60 0.48 569.42 505.91 h 4.25 3.14 0.51 8.31 6.15 3.22 2.98 0.93 3.47 3.21 2.11 2.75 2.28 0.92-1.21 717.75 903.47 4.60 3.99 0.59 7.78 6.75 3.74 2.91 1.66 2.25 1.75 1.52 2.09 2.22 0.68 0.94 588.59 626.42 4.73 3.49 0.62 7.63 5.63 3.49 3.33 1.20 2.91 2.30 1.74 2.77 1.75 1.00 1.59 712.33 915.80 4.45 3.30 0.66 6.82 5.05 h 3.76 3.18 1.30 2.89 2.44 2.35 2.75 1.82 1.30 1.51 817.14 998.06 4.26 3.41 0.54 7.85 6.28 4.07 2.89 1.71 2.38 1.69 2.89 2.85 1.90 1.53 1.50 727.06 753.08 x 4.65 3.72 0.61 7.66 6.13 3.48 2.12 1.55 2.25 1.37 1.25 2.29 2.36 0.53 0.97 568.43 619.01 x 3.93 3.12 0.67 5.87 4.66 ı 4.14 2.71 2.10 1.97-1.29 1.74 2.41 2.92 0.60 0.82 645.24 685.08 - x 5.43 3.78 0.60 9.05 6.31 3.49 3.57 1.83 1.90 1.95 1.60 2.37 2.37 0.68 1.00 508.15 688.0 Mns ollow y th sm lttr r sttistilly not siniint (Dunn s multipl rn tst, P=0.05) Tl 5 Et o notyps ytoplsm in rspons to slt tolrn (DW, Dry Wiht; TI, Tolrn Inx) Crosss Root Stm L Root Stm L DWTI DWTI DWTI K/NTI C/NTI K/NTI C/NTI K/NTI C/NTI 1) X 1323.15 1355.25 1380.09 535.52 543.88 623.96** 588.61 708.79 949.59 7) X 1321.27 1369.24 1373.30 521.63 519.72 546.78 614.74 712.33 915.80 2) X 1201.76 1233.52 1242.59 494.99 540.77** 435.51 397.60** 629.32** 641.94** 10) X 1210.33 1283.45 1240.73 502.86 612.72 429.23 371.09 568.43 619.01 3) X 1407.43** 1403.65** 1317.87 675.29** 676.10** 539.06 617.92 637.28** 715.22** 8) X 1296.31 1309.26 1342.09 487.09 478.12 550.29 614.04 817.14 998.06 4) X 1370.98** 1243.48** 1233.55 426.27** 466.29 480.56** 446.09 569.42** 505.91** 11) X 1133.19 1290.25 1199.97 404.42 496.93 537.10 448.01 645.24 685.08 5) X 1316.34 1347.15** 1366.47 541.70** 678.68** 685.64** 835.72** 717.75 903.47** 9) X 1308.87 1139.10 1373.53 490.42 585.77 590.32 702.38 727.06 753.08 6) X 1169.82 1274.32** 1274.01 444.63** 452.85** 421.93 452.41 588.59** 626.42** 12) X 1234.34 1185.31 1259.78 399.45 425.25 415.51 441.70 508.15 688.0 Siniint twn roups - ** - t P<0.01
Slt Tolrn Durin Vttiv Growth in Cross o Tomto n Et o Cytoplsm... 217 tst tomto rosss ws inrs, whrs K +, C +2, N + n K + /N +, C +2 /N + onntrtion rs. Tolrn inx vlus wr lult orin to ths inins. In this stuy, tolrn inx us s slt tolrnt slt tolrnt tomto rosss t irnt slt onntrtions. Thr ws lr vrition in root, stm n l ry wiht n K + /N +, C +2 /N + TI mon 12 ross omintions unr inrsin slinity. Th hihst tolrn inx vlus wr nrlly otin rom th omintion o tolrnt notyps (, n ) n snsitiv notyp. For xmpl, hihst root, stm n l K + /N + TI, C +2 /N + TI vlus wr xtrt rom th [ x ], [ ] n [ ] rosss omintion. Similr to rsults wr otin rom DWTI. Cytoplsm is nothr tor tht plys n importnt rol in th slt tolrn. It n onlu tht rossin, in twn tolrnt n snsitiv tomto notyps positivly or ntivly t notyp ytoplsm. Ths t wr hn y th irnt tolrn inx vlus in us. Gnrlly, in riprol rosss wr m twn slt tolrnt n slt snsitiv tomto notyps;,, n ytoplsm positivly t tolrn inx vlus (root, stm, l; ry wiht, K + /N +, C +2 /N + onntrtion) n mns whih thy wr lss t y slinity. Rrns Ahsn, M., I. Khli n S. S. Mhi, 2000. Phsionti n rin spt o slinity tolrn in r wht: rviw. Pkistn Journl o Bioloil Sin, 3(9): 1354 1363. AliDinr, H. M., G. Ert n P. Lurs, 1999. Growth, hlorophyll ontnt, photosynthsis n wtr rltions in uv unr slinity n irnt nitron supply. Grtnuwissnsht, 64: 54 59. Aziz, A., J. Mrtin-Tnuy n F. Lrhr, 1999. Slt strss- inu prolin umultion n hns in tyrmin n polymin lvls r link to ioni justmnt tomto ls iss. Plnt Sin, Elsvir Sin Irln Lt. P. 27 31. Bolrin, M. C., F. G. Frnnz, V. Cruz n J. Curtro, 1991. Slinity tolrn in our wil tomto spis usin vttiv yil slinity rspons urvs. J. Amr. So. Hort. Si., 116, 286-290. Bolrin, M. C., F. Prz-Alo, E. A. Cno, M T. Estn n M. Cro, 1993. Growth, ruit yil, n ion onntrtion in tomto notyps tr pr-mrn n post-mrn slt trtmnts. Journl o th Amrin Soity or Hortiulturl Sin, 118: 655 660. Chrtzoulkis, K. n G. Klpki, 2000. Rspons o two rn hous pppr hyris to NCl slinity urin irnt rowth sts. Si. Horti., 86: 247 260. Chrin, S., M. P. Ry n J. B. Pny, 1999. Stuis on slt tolrn in Avinni mrin (Forstk.) Virh.: t o NCl slinity on rowth, ion umultion n nzym tivity. Inin J. Plnt Physiol., 4: 266 270. Cruz, A. S., M M. Roriuz, F. P. Alo, R. R. Arn n M. C. Bolrin, 2002. Th rootstok t on th tomto slinity rspons pns on th shoot notyp.plnt Sin, 162: 825 831. Curtro, J., A. R. Yo n T. J. Flowrs, 1992. Sltion o onors or slt- tolrn in tomto usin physioloil trits. Nw Phytol., 121: 63-69. Curtro, J. n R. Frnnz-Munoz, 1999. Tomto n slinity. Si. Hortiultur, 78: 83 125. Curtro, J., M. C. Bolrin, M. J. Asins n V. Morno, 2006. Inrsin slt tolrn in th tomto. Journl o Exprimntl Botny, 5: 1045 1058. Dsn, H.Y., H. Akts, K. Ak, I. Ckmk, 2002. Dtrmintion o srnin thniqus to slt tolrn in tomtos n invstition o notyp rsponss. Plnt Sin, 163: 695 703. Epstin, E., J. D. Noryln, D. W. Rush, R. W. Kinury, D. B. Klly, G. A. Cunninhm n A. F. Wron, 1980. Slin ultur o rops: A Gnti Approh Sin, 210: 399 404. Fool, M. R., 1996. Anlysis o rspons n orr-
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