Expression pattern of two sugar transporter genes (SuT4 and SuT5) under salt stress in wheat

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1 POJ 3(6): (21) ISSN: Exprssion pttrn o two sugr trnsportr gns (SuT4 n SuT5) unr slt strss in wht Fhimh Chrkzi, S. Snz Rmznpour *, Hssn Soltnloo Dprtmnt o Plnt Bring n Biothnology, Gorgn Univrsity o Agriulturl Sins n Nturl Rsours, Gorgn, Irn *Corrsponing uthor: rmznpours@gu..ir Astrt Slinity strss is importnt ioti tor, limiting wht yil roun th worl n it is known to inu th umultion o wtr solul rohyrts. Ths hngs wr ompni y ltrtion in xprssion lvls o lrg numr o rohyrt mtoli gns. In prsnt stuy, rgultion o two wht rohyrt trnsportr gns, nmly SuT4 n SuT5 uring slinity strss t sling stg wr luits using quntittiv RT-PCR. Slinity strss ws inu using NCl. Msurmnt on grmintion n sling growth rvl Kvir s slt tolrnt n Flt s slt susptil ultivr. Exprssions o two sugr trnsportr gns wr irntilly rgult uring slt strss. Trnsripts lvls o oth gns wr lmost highr in tolrnt ultivr. Highst lvl o SuT4 n SuT5 trnsripts ws osrv in -.75 MP o NCl in slt tolrnt ultivr. Trnsript lvl o oth gns own rgult in slt susptil ultivr y inrsing NCl osmoti potntil up to -1 MP. Thror, it sms tht umultion o sugrs ws nssrily orrlt with umultion o sugr trnsportr gns trnsripts n slinity tolrn. Kywors: Slinity, wht, sugr trnsportr, xprssion, rohyrt Arvitions: SuT: Sugr Trnsportr, MP : Mg Psl, PCR: Polymrs Chin Rtion, QRT-PCR: Quntittiv Rl- Tim PCR, GAPDH: Glyrlhy-3-phospht hyrogns, REST: Rltiv Exprssion Sotwr Tools Introution Wht (Tritium stivum L.) is mjor rop or mor thn on thir o th worl popultion. It is origint in south wstrn Asi n hs n mjor griulturl ommoity sin pr histori tims. Wht proutivity pns upon th proution, trnslotion, storg n utiliztion o rohyrts. Crohyrts srv s sour o nrgy n lso t s signling molul in rgultion o mtoli pthwys unr norml n strss onitions (Gupt n Kur, 25). During grmintion, strh prsnt in th rl nosprm is hyrolyz to gluos y mylss n is thn onvrt to suros y th suros phospht synths. Suros, thus orm is thn trnsport to th growing mryoni xis, whr it is hyrolyz n th prouts so orm r us s nrgy sour or th growth o slings (Smith t l., 25). Sugrs intrt with th snsor protins n initit signl trnsution s tht rsults in llulr rsponss suh s ltr gn xprssion n nzymti tivity (Gupt n Kur, 25). Thror th sugr trnsportr protins ply n importnt rol in ioti strss us thy trnsport rohyrt within th lls. Th tolrn to slt strss is omplx pross tht involvs morphologil physiologil n iohmil moiitions. Survivl n growth unr slin nvironmnts r th rsult o ptiv prosss suh s ion trnsport n omprtmntlistion, synthsis n umultion o orgni soluts, ring to n osmoti justmnt (Fougr t l., 1991). Apttion o plnts to osmoti strss rings out th vlopmnt o low osmoti potntil similr to thos in plnt spis rom ri nvironmnts. As singl ll, w strtgis n onivl to surviv in suh onitions. On o ths mhnisms is th osmoti justmnt hiv y th storg o soluts within lls (Sntos-Diz n Oho-Aljo, 1994). Svrl ring strtgis, inluing mrkr-ssist sltion, r ing us to improv rop yils unr wtr strss. Ths hv n somwht sussul, ut thy r limit y th tim n othr rsours rquir. To improv th iiny o suh strtgis n to vlop nw options, ttr unrstning o th physiologil n molulr ss o slinity tolrn in plnts is rquir (Blum 1998; Trthown t l. 21). Exprssion hngs o som gns involv in rohyrt mtolism uring slt strss hv n rport in w stuis (Brtls n Sunkr 25; Chsmn, 1988; Rthrt, 1984; Zhu, 22). Th trmintion o xprssion pttrn o sugr trnsportr protins in rspons to slinity my ls to unrstning mor out mhnisms involv in suh strsss tolrn. In th othr hn, thr r so mny stuis on ltrntion rohyrt lvl unr ioti strss in physiologil lvl (Tmmm t l., 28; Hsnn t l., 29) ut molulr stuy on xprssion pttrns o nzym inlu in ioti strss ws so rr. In this stuy, w hv invstigt th hngs o xprssion pttrn o two sugr trnsportr gns in th growing slings unr slinity strss in susptil n tolrnt plnts. Ths gns wr isolt rom wht gnotyps unr ol strss y SSH mtho n show high homology sugr trnsportr gns in othr plnts lik rly (Wshk t l., 23) n ri (Bull t l., 23). 194

2 Tl 1. Squn n som hrtriztion o spii primrs. Nm Squn Tm GC% Prout lngth SuT4 For 5'-AGTGCCTTGACAACCTTGCT-3' Rv 5'-GGTTCCGACATCCAGAACAT-3' SuT5 For 5'-GAAGGCTGCAACAAAACCTC-3' Rv 5'- TTTGCCAAGGCTCTACTGTG-3' GAPDH For 5'-TCACCACCGACTACATGACC-3' Rv 5'-ACAGCAACCTCCTTCTCACC-3' 6 55 Mtril n mtho Plnt mtril n strss trtmnt This stuy ws rri out t th prtmnt o plnt ring n iothnology, ulty o rop sin, Gorgn Univrsity o Agriulturl Sin n Nturl Rsours, Irn. Slt strss ws inu y NCl. Six slt trtmnts with irnt osmoti potntils o -.25, -.5, -.75, -1, n -1.5 MP wr rrng s sri y Coons t l., 199. In this stuy istill wtr srv s ontrol. Wht (Tritium stivum L.) ss o v. Kvir (slinitytolrnt) n v. Flt (slinity-snsitiv) wr striliz with 5% trgnt n wsh svrl tims with istill wtr. Thr rplits o 2 ss o h ultivr wr grmint in 2 roll Whtmn iltr pprs with 1 ml o rsptiv tst solutions (Rhmn t l., 1996). In orr to prvnt vportion, h roll ppr ws put into sl plsti g. Ss wr llow to grmint t 2 ± 1 o C in th rk or 1 ys. Grmintion prntg, shoot lngth, root lngth n sling rsh wight wr trmin tr 1 ys. Anlysis o vrin Th xprimntl sign ws two toril, rrng in ompltly rnomiz sign with 3 rplitions n 2 ss pr rplit. Th irst tor ws ultivrs n th son ws NCl osmoti potntils (, -.25, -.5, -.75, -1, n -1.5 MP). Dt givn in prntgs wr sujt to rsin trnsormtion or sttistil nlysis. For ll invstigt prmtrs, nlysis o vrin ws prorm using th SAS vrsion 9.1 (SAS institut Cry, NC). Signiint irns mong th mn vlus wr ompr y LSD tst (P <.5 or.1). RNA xtrtion n DNA synthsis All slings or h trtmnt smpl tr 1 ys n put in luminum oil in irnt rplitions. Th smpls wr immitly rozn in liqui nitrogn n stor t - 8 o C until RNA xtrtion. Lithium hlori (Mrk, Grmny) ws us to xtrt totl RNA using moii mtho s sri y Nito t l., First strn DNA ws rvrs trnsri rom totl RNA y Frmnts protool using RvrtAi TM M-MuLV (Frmnts, Grmny) s mnutur sription n Tnol ilution ws us s tmplt or rl-tim PCR. Primr sign or RT-PCR Primrs wr sign y Primr 3 ( to otin p lngth, 59 n 61 o C mlting tmprtur n GC ontnt twn 55% n 65% voiing hirpins n omplmntrity twn primrs. Th primrs sign s on two sugr trnsportr nit gns nmly, SuT4 n SuT5. Ths gns wr show irntilly xprssion unr ol strss in wht y SSH mtho n show high homology with two irnt puttiv sugr trnsportr gns in rly n ri (ssion no. AJ n NM_ , rsptivly). Th hrtriztion o gns n spii primrs or SuT4, SuT5 n glyrlhy-3-phospht hyrogns (GAPDH) s rrn gn (ssion no. EF59218) ws prsnt in Tl 1. Th spiity o primrs ws hk y stnr PCR n ltrophorsis on gros gl s shown in Fig. 1. Quntittiv rl-tim PCR Rl-tim RT-PCR using SYBR Grn I thnology on iq5 Systm (BioR, USA) ws prorm. Th SYBR Grn PCR Mstr Mix (SYBR ioprs, GUASNR, Irn) ws us with inl onntrtion o 1x SYBR Grn PCR Mstr Mix, in totl volum o 25 ul, 3 nm o h spii sns n nti-sns primrs n 1 ng o DNA wr. Th ollowing mpliition progrm ws us: 94ºC uring 3 min, 35 yls t 15s t 94ºC ollow y 15s t 6ºC n 15s t 6ºC. All smpls wr mplii in triplit rom th sm RNA prprtion n th mn vlu ws onsir. Th I ws us s rrn gn. Two iologil rplitions wr us or h plt. Th rl-tim RT-PCR iiny ws trmin or h gn with th slop o linr rgrssion mol (Pl, 21). For this, ulks o h DNA smpl wr us s PCR tmplt in rng o 1 ol sril ilution. Th orrsponing rl-tim RT-PCR iinis wr lult oring to th qution: E = 1 (-1/slop) (Roni t l., 24). Rltiv xprssion ws omput using ollowing ormul whih prsnt y Pl, 21, s on its rl-tim PCR iiny (E) n th rossing point (CP) irn (Δ) o n trt smpl (unr strss) vrsus ontrol (ΔCP ontrol-smpl ) or oth trgt n rrn gns. ΔCP ΔCP trg ( ontrolsmpl) r( ontrolsmpl) ( E ) ( ) t Rtio= t rgt Er Th REST (Rltiv Exprssion Sotwr tools) sotwr (28) us this ormul to lult rtio twn th mount o trgt molul n rrn molul GAPDH within th sm smpl. In this mol th trgt gn xprssion ws normliz y GAPDH xprssion whih is non-rgult rrn gn. Th normliz vlu ws thn us to ompr irntil gn xprssion in irnt smpls. A mlt urv n gros gl sprtion ws us to tt whthr th PCR rtion yils non-spii rgmnts or primr imrs. Bus SYBR Grn y whih us in this stuy is non-spii y n in to ll typs o oul strn DNA. To vlop mlt urv th rl-tim PCR prouts ws ht t 94 C or 1 min ollow y 11 yls, 1 s pr yl, string t 4 C n inrsing.5 C tr h yl. Rsults Th t o NCl ws signiint (P<.1, =6) or ll invstigt trits xpt or grmintion prntg (Fig. 2). 195

2 p 1 p Fig 1. Agros gl ltrophorsis o spii primrs. Th lns r spii mplii rgmnt on two DNA smpl y SuT4 (1,2), SuT5 (3,4) n GAPDH (5,6) primrs. Grmintion % 12 1 8 6 4 2 -.25 -.5 -.75-1 -1.25-1.

3 2 p 1 p Fig 1. Agros gl ltrophorsis o spii primrs. Th lns r spii mplii rgmnt on two DNA smpl y SuT4 (1,2), SuT5 (3,4) n GAPDH (5,6) primrs. Grmintion % NCl osmoti potntil (MP) Fig 2. Chngs in grmintion prntg o wht ultivrs t irnt osmoti potntils o NCl. Vlus show th rl grmintion prntgs ut vrin nlysis ws prorm using rsin trnsorm vlus. Brs rprsnt on stnr vition. Mns ollow y th sm lttr(s) r not signiintly irnt t P =.1. Thr r no signiint irns twn irnt trtmnts in grmintion prntg. Sling root lngth (m) NCl osmoti potntil (MP) Fig 3. Root lngth o wht ultivrs t irnt osmoti potntils o NCl. Mns ollow y th sm lttr(s) r not signiintly irnt t P =.1. Brs rprsnt on stnr vition. Root lngth ws rs signiintly in ll NCl trtmnts. Sling shoot lngth (m) NCl osmoti potntil (MP) Fig 4. Shoot lngth o wht ultivrs t irnt osmoti potntils o NCl. Mns ollow y th sm lttr(s) r not signiintly irnt t P =.1. Brs rprsnt on stnr vition. NCl trtmnts h signiint t to rs Shoot lngth. g g Kvir Flt Kvir Flt Kvir Flt Th ontrol ( MP) i not show ny irns mong th ultivrs or ll trits. A non-signiint 2-wy intrtion (vrity strss) ws oun or grmintion prntg. Inrsing NCl rsult in rs root lngth in ll vrity. No ultivr ws l to grow roots t -1.5 MP o NCl (Fig. 3). Th irst lvl o NCl (-.25) i not rs root lngth signiintly n it lin onsirly in osmoti potntil mor thn NCl h stimulting t on th root growth o Kvir rom to -.25 MP o osmoti potntil. Root lngth i not hng signiintly twn slinity tolrnt (Kvir) n susptil (Flt) vrity. NCl show stimulting t on shoot growth in oth vritis t -.25 MP (Fig. 4). Shoot lngth i not show signiintly irns twn slt tolrnt n susptil vritis. 2-wy intrtion (vrity strss) ws non-signiint. Drsing wtr potntil y NCl us rution in sling rsh wight (Fig. 5). Dirns mong th vritis wr signiint. Drsing in sling rsh wight y NCl strt rom -.5 MP n NCl hs stimulting t on Kvir sling rsh wight rom to -.25 MP. NCl inlun sling rsh wight o vritis in irnt wys. Th sugr lvl in th slt-strss lvs is pnnt on th trnsporttion tivity o sugr trnsportrs. For this, w monitor th Sugr Trnsportr (SuT) xprssion pttrns o two gns with 96 98% nuloti intity to th Ri sugr trnsportrs gns (SuT4 n SuT5). This is th irst rport whih stui ths gns unr slt strss. Th SuT gns wr originlly isolt y SSH rom lirry onstrut rom low-tmprtur-trt wht rown tissu (unpulish t), us umultion o rohyrts our in ll wtr potntil strss inluing ol, rought n slinity. Anlysis o vrin or SuT4 n SuT5 trnsript umultion in sling show highly signiint ts o vrity, osmoti potntil n vrity osmoti potntil. Thr ws shrp inrs in SuT4 trnsripts in slt tolrnt vrity (Kvir) y inrsing osmoti potntil to -.25 Mp ollow y high sty stt to -.75 Mp o NCl osmoti potntil (Fig. 6). Its trnsripts rs 7% t -1 Mp n inrs 5% in -1.5 Mp in Kvir. In slt susptil vrity, SuT4 trnsripts grully lin to -1 Mp ollow y 15% inrs rom - 1 to -1.5 Mp. Th xprssion pttrn o SuT5 ws lmost th sm in slt tolrnt n susptil vrity t th irst NCl osmoti potntil ut its trnsript lvl is lmost highr in tolrnt ultivr thn in susptil ultivr (Fig. 7). It is own-rgult out (25 3%) in oth ultivr t -.25 Mp o NCl ut it n rovr 1% o its trnsripts lvl in -.75 Mp in Kvir n 8% in Flt. It shrply own rgult in slt susptil in -1 Mp n its trnsripts rs 75%, ollow y sun inrs t Mp n kp sty-stt lvls o trnsript to Kvir show rs in SuT5 xprssion rom to -.75 Mp to Mp ollow y sun inrs in trnsripts lvl t -1.5 Mp. Disussion NCl vrsly t sling growth o wht. Root n shoot lngth, n sling rsh wight wr rs y inrsing NCl onntrtions. Our rsults r onsistnt to stuis whih vlut ts o NCl on th grmintion n sling growth o p y Murillo-Amor t l., (22), owp y Oku t l., (25) n soyn y Khjh- Hossini t l. (23). Howvr, our stuy show tht NCl h grtr inhiitory ts on sling growth thn grmintion us no signiint rs in grmintion in th two ultivrs ws osrv. Ss lwys grmint 196

4 Sling rsh wight (mg pr plnt) NCl osmoti potntil (MP) Fig 5. Sling rsh wight o wht ultivrs t irnt osmoti potntils o NCl. Mns ollow y th sm lttr(s) r not signiintly irnt t P =.1. Brs rprsnt on stnr vition. Sling rsh wight ws rs signiintly in high NCl onntrtions SuT4-NCl Osmotp potntil (MP) Flt Kvir Fig 6. Quntittiv xprssion pttrns o SuT4 gn in sling o Kvir n Flt in irnt NCl osmoti potntil. Th highst xprssion lvl o this gn ws in mil NCl trtmnts (-.5 to -.75 Mp) in Kvir s tolrnt ultivr. Th xprssion lvl o this gn ws ru in susptil ultivr (Flt). % Exprssion Flt Kvir SuT5-NCl Osmoti potntil (Mp) Fig 7. Quntittiv xprssion pttrns o SuT5 gn in sling o Kvir n Flt in irnt NCl osmoti potntils. Th highst trnsript lvl o this gn ws umult in mil lvl o NCl (-.75 Mp) in tolrnt ultivr (Kvir) lik s SuT4 gn. proprly in NCl ompr to othr ioti strss in lin with rlir osrvtions m or soyn y Khjh-Hossini t l., (23). This my u to th uptk o N + n Cl ions y th s, mintining wtr potntil grint llowing wtr uptk uring s grmintion. In onlusion, rly sling growth o wht ws ru s wtr potntils inrs. Kvir ws mor tolrnt to slt onntrtion whil Flt ppr susptil. Howvr sling growth ws mor snsitiv to slt strss thn grmintion. Comptil soluts, low-molulr wight orgni moluls suh s th prviously isuss gluos, suros, rutos, Put n mny othrs, r known to umult unr strss onitions (Guy t l., 28). Thy r onsir to stiliz protins n mmrns n ontriut to ll osmoti Kvir Flt prssur (Kpln t l., 24). Th prsnt work ouss on gn xprssion upon NCl osmoti potntil in wht sling. Through QRT-PCR stuy w show irntil xprssion or sugr trnsportr gns involv in rohyrt umultion unr slinity strss. Although w oun ol strss n shr signl trnsution pthwy to inu rsponing gn to wtr iit strss (unpulish t). Slinity strss is known to ru photosynthsis rts n to inu th umultion o wtr solul rohyrts. Ths hngs wr ompni y ltrtion in xprssion lvls o lrg numr o rohyrt mtoli gns. With omprhnsiv mtoli-pthwy-s xprssion nlysis using urt n quntittiv RT-PCR, w monstrt signiint ltrtions in th xprssion lvls o two prviously unknown slinity rsponsiv gns involv in th mjor rohyrt mtoli pthwys in slt strss sling. Exprssion pttrn o SuT4 n SuT5 gns ws hng unr slt strss, whih mns prout o ths gns r importnt in signl trnsution pthwy unr osmoti strss us y irnt NCl onntrtions. Both gns signiintly up rgult in -1.5 Mp o NCl osmoti potntil. In this osmoti potntil thr ws no shoot growth ut root grw vry low. This mns th rohyrt nssry to growth shoot ws stor in root ut som inhiitory t likly ion toxi t inhiit growing shoot. In our stuy, th inrs o sugr trnsportr trnsripts in sling o Kvir ws highr thn Flt, thror it sms tht th umultion o sugrs ws nssrily orrlt with umultion o sugr trnsportr gns trnsripts n slinity tolrn. Inrsing in sugr ontnt long with inrsing NCl onntrtion ws rport y Chorro t l., 1993 in Phsolus vulgris. In nothr stuy, sorghum grown in highly slin soil show 28% inrs in solul sugr ontnt in lvs (Chvn n Krj, 1986). Sugr ontnt o tomto l sp ws t irntly y soium hlori strss (Bzrr, 1992). A omplx ssntil rol o solul sugrs in plnt mtolism is wll known s prouts o hyrolyti prosss, sustrts in iosynthsis prosss, nrgy proution ut lso in sugr snsing n signling systms. Rntly it hs n lim tht vn sugr lux my signl or mtoli rgultion (Gison, 25). Prottion ginst hyrtion y th ormr sugrs ws orrlt with th inrs in shoots n roots. Solul sugrs my lso untion s typil osmoprottnt, stilizing llulr mmrns n mintining turgor. Unr strss onitions, umultion o sugr trnsportr trnsripts my to ountr th osmoti strss. Aknowlgmnt This rsrh ws support y Gorgn Univrsity o Agriulturl Sin n Nturl Rsours (GUASNR), Irn. Rrns Brtls D, Sunkr R (25) Drought n slt tolrn in plnts. Crit Rv Plnt Si 24: Bzrr N E (1992) Slt tolrn in tomtos. Bngor, Univrsity Collg o North Wls. 161 p. Ts Doutoro. Blum A (1998) Plnt ring or strss nvironmnt. pp. 28. Flori: CRC Prss In. Bull CR, Wing RA, MComi WR, Mssing J, Yun Q (23) In-pth viw o strutur, tivity, n volution o ri hromosom 1. Sin 3 (5625):

5 Chorro P, Ortiz A, Cr A (1993) Growth, wtr rltions n solut omposition o Phsolus vulgris L. unr slin onitions. Plnt Si 95:23-29 Chvn PD, Krg BA (1986) Growth, minrl nutrition, orgni onstitunts n rt o photosynthsis in Ssni grnilor L. grown unr slin onitions. Plnt n Soil 93: Chsmn JM (1988) Mhnisms o Slinity Tolrn in Plnts. Plnt Physiol 117: Coons MJ, Kuhl RO, Simons NR (199) Tolrn o tn lttu ultivrs to high tmprtur omin with NCl uring grmintion. J Amri So Hort Si 115: Fougr F, Ruulir DL, Strtr JG (1991) Ets o slt strss on mino i, orgni i, n rohyrt omposition o roots, trois, n ytosol o l (Migo stiv L.). Plnt Physiol 96: Gison SI (25) Control o plnt vlopmnt n gn xprssion y sugr signling. Curr Opin Plnt Biol 8: Gupt AK, Kur N (25) Sugr signling n gn xprssion in rltion to rohyrt mtolism unr ioti strsss in plnts. J Biosi 3: Guy C, Kpln F, Kopk J, Slig J, Hinh DK (28) Mtolomis o tmprtur strss. Physiol Plnt 132: Hsnn MNA, Younis ME, Tourky SMN (29) Plnt growth, mtolism n pttion in rltion to strss onitions XXIII. Slinity-iortility intrtiv ts on growth, rohyrts n photosynthti iiny o ltu stiv. Plnt Omis J 2(2):6-69 Kpln F, Kopk J, Hskll DW, Zho W, Shillr C, Gtzk N, Sung DY, Guy CL (24) Exploring th tmprtur strss mtolom o Ariopsis. Plnt Physiol 136: Khjh-Hossini M, Powll AA, Binghm IJ (23) Th intrtion twn slinity strss n s vigour uring grmintion o soyn ss. S Si n Thnol 31: Murillo-Amor B, Lopz-Aguilr R, Ky C, Lrring- Myorl J, Flors-Hrnnz A (22) Comprtiv ts o NCl n polythyln glyol on grmintion, mrgn n sling growth o owp. J Agron Crop Si 188: Nito S, Hiri MY, Chino M, Kom Y (1994) Exprssion o soyn (Glyin mx (L.) Mrr.) s storg protin gn in trnsgni Ariopsis thlin n its rspons to rspons to nutritionl strss n to sisi i muttions. Plnt Physiol 14: Oku G, ky MD, Atk M (25) Ets o Slt n Drought Strsss on Grmintion n Sling Growth o P (Pisum stivum L.). Turk J Agri For 19: Pl MW (21) A nw mthmtil mol or rltiv quntiition in rl-tim RT-PCR. Nuli Ais Rs 29: Roni A, Thulk S, Mky IM, Lnt O, Sigrt W, Nitsh A (24) Guilin to rrn gn sltion or quntittiv rl-tim PCR. Biohm Biophys Rs Commun 313: Rthrt G (1984) Suros n Strh Contnt o Plnt Prts s Possil Initor or Slt Tolrn o Crops. Aust J Plnt Physiol 11: Rhmn S, Hrris PJC, Bourn WF, Wilkin J (1996) Th t o soium hlori on grmintion n th potssium n lium ontnt o Ais ss. S Si n Thnol. 25: Sntos-Diz MS, Oho-Aljo N (1994) PEG-tolrnt ll o hili pppr: growth, osmoti potntils n solut umultion. Plnt Cll, Tissu n Orgn Cultur, 37:1-8 Smith AM, Zmn SC, Smith SM (25) Strh grtion. Annu Rv Plnt Biol 56: Tmmm AA, Aou Alhm MF, Hm MM (28) Stuy o slt tolrn in wht (Tritium stivum L.) ultivr Bnysoi 1. Aust J Crop Si 1(3): Trthown RM, Cross J, Ginkl M, Vn Rjrm S (21) Rltionships mong r wht intrntionl yil tsting lotions in ry rs. Crop Si 41: Wshk W, Pnitz R, Gutz S, Wngy Q, Rhuk R, Wr H, Wous H (23) Th rol o invrtss n hxos trnsportrs in ontrolling sugr rtios in mtrnl n ilil tissus o rly ryopss uring rly vlopmnt. Th Plnt J (33): Zhu JK (22) Slt n rought strss signl trnsution in plnts. Annu Rv Plnt Biol 53:

WATER STRESS AFFECTS THE GERMINATION, EMERGENCE AND GROWTH OF DIFFERENT SORGHUM CULTIVARS

WATER STRESS AFFECTS THE GERMINATION, EMERGENCE AND GROWTH OF DIFFERENT SORGHUM CULTIVARS SINET: Ethiop. J. Si., 28(2):119 128, 25 Fulty of Sin, is Univrsity, 25 ISSN: 379 2897 WTER STRESS FFECTS THE GERMINTION, EMERGENCE ND GROWTH OF DIFFERENT SORGHUM CULTIVRS Wonimu yu 1, N.F.G. Rthmn 2,