Mitigation of salt stress in radish (Raphanus Sativus l.) by plant growth promoting rhizobacteria

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1 Roumnin Biotechnologicl Letters Vol. 13, No. 5, 28, pp Copyright 28 Buchrest University Printed in Romni. All rights reserved Roumnin Society of Biologicl Sciences ORIGINAL PAPER Mitigtion of slt stress in rdish (Rphnus Stivus l.) y plnt growth promoting rhizocteri Astrct Received for puliction, August 2, 28 Accepted, Octoer 1, 28 ERTAN YILDIRIM*, METIN TURAN** MESUDE FIGEN DONMEZ*** *Ispir Hmz Polt Voctionl Trining School, Attürk University, 259, Ispir, Erzurum, Turkey **Deprtment of Soil Science, Fculty of Agriculture, Attürk University, Erzurum, Turkey ***Deprtment of Plnt Protection, Fculty of Agriculture, Attürk University, Erzurum, Turkey * Corresponding uthor: ertyil25@hotmil.com, ertyil25@yhoo.com The effects of Stphylococcus kloosii EY37 nd Kocuri erythromyx EY43 on some grophysicl properties such s shoot root fresh weight, shoot root dry weight, emergence percent (EP), chlorophyll content, lef numer per plnt (LNPP), lef reltive wter content (LRWC), electrolyte lekge (EL) nd ionic composition of leves of rdish were tested under sline conditions. Seeds were soked in the cteril suspension incuted t 27 C for 2 h. After incution the seeds were ir dried efore use. Slinity tretments were estlished y dding nd 8 mm of NCl to se complete nutrient solution. In control, slt tretment significntly decresed shoot nd root fresh dry weight, EP, LNPP, LRWC, chlorophyll nd minerl content, ut incresed EL. However, EY37 nd EY43 tretments under sline conditions significntly incresed shoot fresh weight (79.5%, 56.2%), root fresh weight (1.% 52.2%), shoot dry weight (18.1% 97.%), root dry weight (15.6% 157.4%), LNPP ( ), LRWC (37.% 2.6%), EP (17.2% 14.1%) nd chlorophyll content (21.2% 24.9%), ut decrese EL (19.5% 26.9%) compred to the control (without plnt growth promoting rhizoil cteri (PGPR)). Ionic compositions of the leves of rdish plnts were significntly ffected y slinity nd cteril inocultions. All nutrient element contents investigted were significntly decresed except for N nd Cl under slt stress, which were significntly incresed y slt tretment. Bcteril inocultion under slinity conditions usully incresed plnt nutrient element (PNE) contents of the leves. It cn e concluded from the study tht tretment with EY37 nd EY43 strins cn meliorte the deleterious effects of slt stress on nutrition nd on the growth prmeters of rdish plnts under slinity conditions; PGRB tretment could offer n economicl nd simple ppliction technology to llevite the modertely slt sensitive rdish production prolems in rid soil cused y high slinity. Keywords: Ameliortive effect, rdish, Stphylococcus kloosii, Kocuri erythromyx, slinity stress Introduction Cultivted soils worldwide re ecoming more sline from mrginl irrigtion wter, excessive fertiliztion, nd desertifiction processes. Impcted soils re mjor limiting production fctor worldwide for every mjor crop [1]. Furthermore, relloction of fresh wter from griculture to cities nd industries limits irrigted crop production in mny importnt food production regions. Under these conditions, production is dependent on the use of lterntive wter sources including sline or wste wter for irrigtion. Slinity is mjor iotic stress reducing the yield of wide vriety of crops ll over the world [2]. Glolly, more thn 77. km 2 of lnd is slt ffected y secondry sliniztion: 2% of irrigted lnd, nd out 2% of dry griculturl lnd [3]. The restriction of plnt growth nd productivity due to slinity is especilly cute in rid nd semi rid regions round the world. The slinity nd lklinity re lso prolem in some prt of Turkey. The glol estimte for 3933

2 ERTAN YILDIRIM, METIN TURAN, MESUDE FIGEN DONMEZ griculturl lnd thretened y or lredy lost to slinity exceeds 9 million h [4]. In Turkey, out 2 million h of cultivted lnd is ffected y slinity nd lklinity [5]. Slinity my occur when there is irregulr irrigtion, indequte dringe, wrong fertilizer ppliction, nd it extremely increses prticulrly in protected cultivtion [6]. Generlly plnts growing in sline medi come cross with mjor drwcks. The first is the increse in the osmotic stress due to high slt concentrtion of soil solution tht decreses wter potentil of soil. The second is the increses in concentrtion of N nd Cl, exhiiting tissue ccumultion of N nd Cl nd inhiition of minerl nutrients uptke, thus cusing ionic imlnce [7]. Mny ttempts hve een mde to reduce the drstic effect of slt stress on growth nd productivity, most focusing on chemicl meliortion, tht involve developing slt resistnt cultivrs, leching excess solule slts from upper to lower soil depths, flushing soils tht contin slt crusts t the surfce, reducing slt y hrvesting slt ccumulting eril plnt prts in res with negligile irrigtion wter or rinfll for leching, nd meliortion of sline soils under cropping nd leching [8]. Breeding for tolernce to slinity in crops hs usully een limited y lck of relile trits for selection. Multiple genes seem to ct in concert to increse slinity tolernce, nd certin proteins involved in slinity stress protection hve lso een recognized [9]. Therefore, the developments of methods nd strtegies to meliorte deleterious effects of slt stress on plnts hve received considerle ttention. Recently iologicl pproch using plnt growth promoting rhizocteri (PGPR) inocultion ws ttempted. The most pproprite solution in such conditions is to use slt tolernt cteril inoculnts tht my prove useful in developing strtegies to fcilitte plnt growth in sline soils [1]. GARCIA & HERNANDEZ [11] reported tht slinity negtively ffects iologicl ctivity y high osmotic strength (low wter potentil) which cn e ttriuted to the toxic effect on microil growth, except tolernt hlophytic cteri. Therefore, slt tolernt rootcolonizing cteri tht hve mnged to survive dverse environmentl fctors could gretly help in hrnessing them for their eneficil properties in such environments in which other microorgnisms hrdly survive [12]. These PGPR cn lso prevent the deleterious effects of one or more stressors from the environment. In ddition, the identifiction, selection nd ppliction of suitle eneficil microorgnisms cn increse the options to del with growing prolems, nd cn e lso environmentlly sound [13]. Although some studies hve een conducted on effects of PGPR on growth nd productivity in different plnts such s tomtoes [12] nd lettuce [14] no ttempts hve to dte een mde to study the effects of Stphylococcus kloosii EY37 nd Kocuri erythromyx EY43 on growth, productivity nd ionic compositions in rdish plnts under slinity conditions. Rdish is considered s modertely sensitive crop to slinity [15]. A review of literture revels tht no investigtions hve een crried out in reltion to slt cclimtion of rdish treted with Stphylococcus kloosii EY37 nd Kocuri erythromyx EY43. Therefore, the im of the present study ws to determine the effect of these cteri on emergence, plnt growth, some physiologicl properties nd resistnce of rdish grown under slt stress. Mterils nd methods Plnt mterils nd growing conditions The study ws conducted under greenhouse conditions t Attürk University, in Turkey, during 28. Rdish (Rphnus stivus L. cv Cherry Belle) plnts were mintined under nturl light conditions, dy/night temperture of pproximte 23/13 C nd 6% reltive humidity during the spn of the experiment Roum. Biotechnol. Lett., Vol. 13, No. 5, (28)

3 Mitigtion of slt stress in rdish (Rphnus Stivus l.) y plnt growth promoting rhizocteri Bcteril isoltion, culture nd inocultion Forty four cteril strins were originlly isolted from the rhizosphere of plnts nturlly grown on high slty soils in Upper Coruh Vlley (N+4 29 / L+41 1 / ) Erzurum, Turkey. The cteril strins were tested to determine their slt tolernce. One smll colony of ech strin ws plced in Petri dishes contining gr nd 2, 5 nd 1% NCl. Stphylococcus kloosii EY37, Kocuri erythromyx EY43 (with MIS similrity index (SIM) of.58,.6, respectively, sed on ftty cid methyl ester nlysis using MIDI system), which showed resistnce to the 1% NCl, were selected to test. Bcteri were grown on Nutrient Agr (NA) for routine use, nd mintined in Nutrient Broth (NB) with 3% glycerol t 8 C for long term storge. For this experiment, the cteril strins were grown on nutrient gr. A single colony ws trnsferred to 25 ml flsks contining NB, nd grown eroiclly in flsks on rotting shker (95 rpm) for 24 h t 27 C. The cteril suspension ws then diluted in sterile distilled wter to finl concentrtion of 1 8 CFU ml 1. For seed tretments, seeds were soked in the cteril suspension mended with sucrose to fcilitte the dherence of the cteri to the seeds nd incuted t 27 C for 2 h. After incution the seeds were ir dried efore use. Seeds soked in distilled wter mended with sucrose served s the control. The cteril strins were le to grow in N free sl medium indicting their N fixing potentil. In the present study, P soluilizing ctivities of the two new (EY 37 nd EY 43) were mesured ccording to the qulittive methods [16] (Tle 1). Tle 1. Some iochemicl chrcteristics of the cteril strins tested Bcteril strin Grm stin Ctlse Growth in YDC P soluiliztion Growth in N free Bsl medium Stphylococcus kloosii EY Kocuri erythromyx EY Plnt growth Rdish seeds were sown into plstic trys filled with mixture (1/1 v/v) of soil nd snd (ph: 7.5, EC:1.82 ds/cm, N:1.2 mg/kg, P:13.22 mg/kg, exchngele K 1.64 meq/1 g soil, orgnic mtter: 3.1 %). There were four rows in trys nd ech row contined 5 seeds. Trys were 8x8x2 cm (length x width x height). All trys were rndomized on the enches in the greenhouse. There were four replictes per tretment, 24 trys in totl. Slt (NCl) tretments Slinity tretments were estlished y dding nd 8 mm of NCl to se complete nutrient solution (SoFertig). The composition of the SoFertig (Elftochem Co., Pris, Frnce) ws (%): N, 17; P 2 O 5, 9; K 2 O, 31; Mg, 2; SO 4, 4; N,.1; Fe,.2; Zn,.2; Cu,.2; B,.1; Mn,.1; Mo,.1. The solution ws prepred y dding SoFertig to the distilled wter. The electricl conductivities of these solutions fter dding nd 8 mm of NCl were determined with conductivity meter, Model 47 (Jenwy Limited). Electricl conductivities (EC) of these solutions were 1.82 ds m 1 for mm NCl nd 1.21 ds m 1 for 8 mm NCl. Emergence percentge Norml seedlings were counted fter sowing nd emergence percentge (EP) determined. Chlorophyll mesurements A portle chlorophyll meter (SPAD 52, Konic Minolt Sensing, Inc., Jpn) ws used to mesure lef greenness of the rdish plnts t 2 dys efore hrvest. For ech plnt mesurements were tken t four loctions on ech lef, two on ech side of the midri on ll Roum. Biotechnol. Lett., Vol. 13, No. 5, (28) 3935

4 ERTAN YILDIRIM, METIN TURAN, MESUDE FIGEN DONMEZ fully expnded leves nd then verged. Reltive lef chlorophyll content ws expressed in SPAD units defined y mnufcturer s 1 equivlent to very ple green colortion (chlorotic) nd 5 equivlent to very drk green colortion Mesurement of electrolyte lekge (memrne permeility) For mesurement of electrolyte lekge, 2 lef discs (1 mm in dimeter) from the young fully expnded leves from four plnts per replicte were plced in 5 ml glss vils, rinsed with distilled wter to remove electrolytes relesed during lef disc excision. Vils were then filled with 3 ml of distilled wter nd llowed to stnd in the drk for 24 h t room temperture. Electricl conductivity (EC1) of the thing solution ws determined t the end of incution period. Vils were heted in temperture controlled wter th t 95 C for 2 min, nd then cooled to room temperture nd the electricl conductivity (EC2) ws mesured. Electrolyte lekge ws clculted s percentge of EC1/EC2. Lef Reltive Wter Content (LRWC) To determine LRWC of plnt, four leves were collected from the young fully expnded leves of fours plnts per replicte t 2 dys efore hrvest. Individul leves first detched from the stem nd then weighed to determine fresh weight (FW). In order to determine turgid weight (TW), leves were floted in distilled wter inside closed Petri dish. Lef smples were weighed periodiclly, fter gently wiping the wter from the lef surfce with the tissue pper until stedy stte chieved. At the end of imiitions period, lef smples were plced in pre heted oven t 7 C for 48 h, in order to determine dry weight (DW). Vlues of FW, TW, nd DW were used to clculte LRWC using the eqution elow [17]: LRWC (%)=[(FW DW)/(TW DW)]x1 Growth Prmeters Forty five dys fter sowing (DAS), twenty plnts from ech replicte were rndomly hrvested, nd dt on plnt growth vriles, such s shoot fresh weight, root fresh weight, shoot dry weight, root dry weight, nd lef numer were collected. Minerl Anlysis In order to determine the minerl contents of leves of rdish, plnt smples were collected from fully expnded leves t fourth five DAS thn oven dried t 68 o C for 48 h nd ground nd pssed 1 mm sieve size. The Kjeldhl method nd Vpodest 1 Rpid Kjeldhl Distilltion Unit (Gerhrdt, Konigswinter, Germny) were used to determine totl N. Phosphorus nd S contents were determined fter wet digestion using HNO 3 HClO 4 cid mixture (4:1 v/v) [18]. Phosphorus nd S in the extrction solution ws mesured spectrophotometriclly using the indophenol lue nd scoric cid method [18] nd UV/VIS Aqumt Spectrophotometer t 66 nm nd t 44 nm respectively (Thermo Electron Spectroscopy LTD, Cmridge, UK). Potssium, N, C, nd Mg, Fe, Mn, Zn, nd Cu were determined fter wet digestion using HNO 3 HClO 4 cid mixture (4:1 v/v). In the diluted digests, K, N, C, Mg, Fe, Mn, Zn, nd Cu nlysis were determined y tomic sorption spectrometry (Perkin Elmer 369) [18]. Sttisticl Anlysis Experimentl design ws hierrchicl with respect to two fctors rrnged in completely rndomized design with four replictions. The first fctor (NCl levels) hd two levels ( nd 8 mm), nd the second one (cteril strins) hd three levels (control, EY37, EY43). Dt were sujected to nlysis of vrince (ANOVA) to compre the effects of slt stress tretments nd cteril strins. Percentge dt were trnsformed using rcsine prior to sttisticl nlysis. The differences etween the mens were compred using lest significnt difference test (LSD, p <.5) Roum. Biotechnol. Lett., Vol. 13, No. 5, (28)

5 Mitigtion of slt stress in rdish (Rphnus Stivus l.) y plnt growth promoting rhizocteri Tle 2. Rdish lef minerl content s ffected y cteri tretments nd slt stress (men ± stndrd error, n = 4 ech tretment) Slt Bcteril N P K C Mg N Cu Mn Zn Fe Tretment Tretment (mm) g 1 g 1 dw mg kg 1 dw Control 2.37±.7c.46± ±.25.77±.4.53±.2 189±84 22.± ± ± ±7.32 EY37 3.3±.6.63± ±.3.84±.4.5±.2 1± ± ± ± ±2.65 EY43 2.9±.5.66± ±.3.81±.1.62±.4 113± ± ± ±1. 14.±3.61 LSD ns Control 2.12±.15.3±.1c 2.22±.17.61±.1.33± ± ± ± ± ±2.65c 8 EY ±.21.47± ±.3.74±.3.45±.4 267± ± ± ± ±5.51 EY43 19.± ±.8.42± ±.2.75±.3.42± ± ± ± ±2. LSD z : Numers with the sme letters in the sme column re not sttisticlly different (p <.5) n.s: non significnt Roum. Biotechnol. Lett., Vol. 13, No. 5, (28) 3937

6 ERTAN YILDIRIM, METIN TURAN, MESUDE FIGEN DONMEZ Results Effects of cteril ppliction on EP nd growth EP ws significntly influenced y NCl nd PGRB strins. There were no significnt differences etween tretments in terms of EP in slt stress sence wheres PGPR strins significntly (p <.5) incresed the EP vlues compred to the control (non inoculted plnts) under stress conditions. When evluted with nd without PGPR tretment in slt stress conditions, the lowest vlue of EP ws otined without PGPR, while the highest ws recorded in the EY37 tretment (Figure 1). Slt stress significntly (p <.5) decresed lef numer of rdish plnts regrdless of PGPR tretments compred to the non sline conditions. However, plnts treted with PGPR hd more lef numer thn the control under slt stress. E37 gve the highest lef numer in oth slt stress nd slt stress sence (Figure 1). The results of this study showed tht growth prmeters were significntly ffected y slinity nd cteril pplictions (Figure 1). Fresh dry shoot nd root weight significntly (p <.5) were decresed y 8 mm NCl tretment. An increse in NCl concentrtion from mm to 8 mm resulted in dry shoot nd root weight reduction of 77.9% nd 85.6%, respectively. Nevertheless, cteril tretments under slinity conditions hve positive effects on plnt growth. EY 37 inocultion gve the highest fresh shoot (1.1 g) nd root (5.1 g) weight nd dry shoot root (6.g 2.2 g) weight. EY43 cteril strins hd lso higher fresh nd dry shoot root weight thn the control under slt stress. Effects of cteril ppliction on chlorophyll content, LRWC nd EL Figure 1 shows the effect of slt stress nd PGRB tretments on chlorophyll content, LRWC nd electrolyte lekge of rdish plnts. Slinity nd cteril pplictions significntly (p <.5) ffected chlorophyll content, LRWC nd EL in this study. Chlorophyll contents of plnt leves were significntly (p <.5) decresed with the slt stress. Effects of PGRB ppliction on chlorophyll contents of leves ws not oserved in slt stress sence, ut significnt (p <.5) increse ws otined with PGRB ppliction under slt stress conditions (Figure 1). The lowest nd highest chlorophyll reding vlues were otined from without PGPR (32.1) nd EY43 (4.1) tretments in 8 mm NCl, respectively. Incresing the concentrtion of NCl from to 8 mm lowered LRWC of rdish plnts. The verge percentge of LRWC decrese ws 36.3% under slt stress. Bcteril pplictions under slinity condition elevted the LRWC level (61.93% for EY37, 54.5% for EY43) over to the control (Figure 1). Slinity induced significnt (p <.5) increses in EL compred with no slinity ( mm NCl) ut plnts treted with PGPR strins showed lower vlues thn non treted ones. There were no significnt (p <.5) differences etween tretments in regrd to electrolyte lekge under the slt sence (Figure 1). Effects of cteril pplictions on ionic composition of leves The results showed tht slinity nd cteril inocultions significntly ffected ionic compositions of leves (Tle 2). We hve found in this study tht compred with mm NCl tretment 8 mm NCl tretment decresed plnt nutrient element (PNE) contents of leves, except N nd Cl contents. On the other hnd, N nd Cl concentrtion of leves incresed with incresing NCl concentrtion in the nutrient solution. However, generlly cteril pplictions under slinity condition significntly incresed PNE content of leves except for N nd Cl concentrtion compred to the control. N nd Cl contents of leves significntly (p <.5) were decresed y cteril pplictions. Bcteril pplictions rought PNE contents of leves ner to tht of mm NCl tretment Roum. Biotechnol. Lett., Vol. 13, No. 5, (28)

7 Mitigtion of slt stress in rdish (Rphnus Stivus l.) y plnt growth promoting rhizocteri Shoot Fresh weight (g/plnt) c mm 8 mm Control EY37 EY43 Root fresh weight (g/plnt) c mm 8 mm NCl NCl Shoot dry weight (mg/plnt) mm 8 mm Root dry weight (mg/plnt) c mm 8 mm NCl NCl EP (%) mm 8 mm Lef numer per plnt c mm 8 mm NCl NCl LRWC (%) mm 8 mm c Electrolyte lekge (%) mm 8 mm NCl NCl Chlorophyll reding vlues mm 8 mm NCl Figure 1. Emergence percentge, shoot fresh weight, shoot dry weight, root fresh weight, root dry weight, lef numer per plnt, chlorophyll content, electrolyte lekge nd LRWC of rdish plnts in response to different PGRB tretments under slt stress. Roum. Biotechnol. Lett., Vol. 13, No. 5, (28) 3939

8 ERTAN YILDIRIM, METIN TURAN, MESUDE FIGEN DONMEZ Different letters on top of rs indicte significnt differences ccording to LSD test (p <.5) t ech slt level. Verticl rs indicte the men ± SE, n=4. Discussion PGPR re free living microorgnisms hving eneficil effects on plnts y colonizing in the rhizosphere of plnts. PGPR my improve plnt growth nd yield y indirect nd direct mechnism. In indirect mechnism, PGPR hve ntgonistic effect ginst phytopthogenic microorgnisms [19]. Direct mechnisms my ct on the plnt itself nd ffect growth y mens of plnt growth regultors such s uxin, cytokinins nd gierellins, soluilizing of inorgnic phosphte nd minerliztion of orgnic phosphte nd symiotic fixtion of tmospheric nitrogen [2]. In this study, evidence is provided tht PGPR improves emergence performnce, growth nd nutrient uptke of rdish nd to the est of our knowledge, this is the first report to suggest tht Stphylococcus kloosii EY37 nd Kocuri erythromyx EY43 inoculted onto seeds could induce tolernce to slt stress. Similr findings were reported in the previous studies showing tht ppliction of PGPR my stimulte yield, growth nd PNE uptke from soil in different crops such s tomtoes [12, 13] nd lettuce [14] under slinity conditions. However, Azospirillum sp., Achromocter sp., Serrti sp., Rhizoium sp., Aeromens sp. nd Bcillus spp. s PGPR ws used in these previous studies. Stphylococcus nd Kocuri were not used s PGPR under sline conditions. The present study demonstrted tht slt stress dversely ffected emergence nd growth of rdish (Figure 1). Slinity, s n iotic hzrd, induces numerous disorders in seeds nd propgules during germintion. Slinity either completely inhiits germintion t higher levels or induces stte of dormncy t lower levels [21]. Slinity cn lso ffect germintion y fcilitting the intke of toxic ions, which cn cuse chnge of certin enzymtic or hormonl ctivities of the seed. Slinity hs een reported to cuse significnt reductions in the rte nd finl percentge of germintion nd emergence of rdish, which in turn my led to uneven stnd estlishment nd reduced crop yields [22, 23, 24]. In the study, rdish seeds inoculted with PGPR strins showed greter EP thn non primed ones under slt stress. Similrly, PGPR inocultions improved germintion of seeds of lettuce [14] in sline conditions. This positive effect of the PGPR on seed germintion nd emergence could e ttriuted to the cteril ility to produce or modify plnt hormones including gierellins, which ply key role in germintion [14]. Slt stress lso significntly decresed lef numer, fresh nd dry weight of shoot nd root of rdish plnts (Figure 1). Similrly, MARCELIS & VAN HOOIJDONK [23] nd Jmil et l. [24] reported tht growth chrcteristics such s shoot nd root weights, nd lef re were negtively ffected when rdish plnts were grown under slt stress. The present study showed tht PGPR improved the growth prmeters of rdish plnts compred to the noninoculted controls under slt stress s well s non sline conditions (Figures 1). Our findings re concordnt with those of BOCHOW & l. [25] in eggplnt, MAYAK & l. [12] in tomto, YILDIRIM & TAYLOR [26] in en, BARASSI & l. [14] in lettuce, nd YILDIRIM & l. [27] in sqush, who found tht PGPR meliorted the deleterious effects of slt stress on plnt growth. This results lso confirmed tht the PGPR inocultions my effectively increse the surfce re of roots [28] nd the root weight [29]. Slt stress significntly decresed chlorophyll reding vlues compred to the nonsline conditions (Figure 1). Similrly, JAMIL & l. [24] indicted tht chlorophyll content ws reduced y slt stress. However, PGPR inocultion elevted the chlorophyll reding vlues compred to the controls under oth slt stress nd sence of slt stress. There were 394 Roum. Biotechnol. Lett., Vol. 13, No. 5, (28)

9 Mitigtion of slt stress in rdish (Rphnus Stivus l.) y plnt growth promoting rhizocteri no significnt (p <.5) differences etween tretments in regrd to LRWC in mm NCl while PGRB tretments cused to the increse in LRWC under slt stress. Electrolyte lekge ws higher in 8 mm NCl compred to the non sline conditions. PARIDA & DAS [3] reported tht slt stress led to significnt increse in the level of electrolyte lekge in mny crops. In the present study, PGRB inoculted plnts hd less electrolyte lekge compred to their respective non inoculted controls. PGRB cn trigger plnt medited induced systemic resistnce (ISR) response [31]. SASKIA & l. [32] reported tht PGPR could induce ccumultion of the signling molecules of slicylic cid nd jsmonte, which hve een shown to e importnt signl molecules for modulting plnt responses to environmentl stress [33]. Bcteril inocultions in rdish plnts could llevite deleterious effects of slt stress. This phenomenon cn e explined y couple of mechnisms: cteril inocultion cn restrict N nd Cl uptke nd enhnce the uptke of other PNE such s N, P, K nd C positively. We found tht cteri inoculted plnts under slt stress hve lower N nd Cl contents nd higher PNE contents compred with non inoculted plnts. Seed inocultion of rdish with the EY37 nd EY43 significntly incresed concentrtion of P nd N in the leves of rdish plnts. The higher totl N nd P uptke of rdish could e roused y N fixtion nd P soluiliztion, thus promoting the plnt growth (Tle 1). Phosphte soluilizing nd N2 fixing cteri cn improve the N nd P nutrition of plnts nd stimulte the plnt growth [34, 35]. Our dt showed tht higher nutrient uptke y PGPB inocultions significntly improved seedling growth. PGPR inoculnts could hold inorgnic ions, especilly N +, in roots, thus preventing their trnsfer to leves. It is known tht hlophoic plnt species hve this mechnism [36]. These cteri could contriute to increse to slt tolernce vi decresing N uptke. PGPR cn fcilitte rooting nd growth of plnts under sline conditions y enhncing wter use efficiency, y providing plnts with fixed nitrogen, iron nd solule phosphte [12, 37]. In these wys, plnt growth nd PNE uptke from soil could e incresed y stimultion of root growth. Thus, incresing root growth nd PNE uptke nd decresing N nd Cl uptke could supply the stiliztion of memrne permeility, nd increse chlorophyll reding vlue nd LRWC, stimulting the growth of rdish plnts under sline conditions. YILDIRIM & l. [27] reported tht some PGPR tretments reduced the N uptke of sqush plnts nd/or incresed the K nd C uptkes compred to the control tretment (non inoculted plnts) under slt stress. ASHRAF & l. [38] determined tht PGRB, exopolisccride producing cteri, could restrict N influx into roots. PGPB cn directly ffect plnt growth through the production of phytohormones. This direct growth promoting effect ppers to involve the plnt growth regultors indole 3 cetic cid nd cytokinins [39]. PGPR hve een lso reported to le to control ACC levels or lock ethylene iosynthesis in plnts, thus stimulting root growth nd helping to protect plnts from the deleterious effects of the slt stress (12). Conclusion Sline soils nd sline irrigtions constitute serious production prolem for vegetle crops s sline conditions re known to suppress plnt growth. The present study demonstrtes tht slinity stress induced lower plnt fresh dry weight, lower chlorophyll nd mcro nd micro element content in rdish plnts. The ssessment of the effect of slinity on the growth prmeters y different slinity levels enled us to conclude tht ll of the considered prmeters were ffected y slinity. To conclude the Roum. Biotechnol. Lett., Vol. 13, No. 5, (28) 3941

10 ERTAN YILDIRIM, METIN TURAN, MESUDE FIGEN DONMEZ present study, we suggest tht PGPR seed tretments cn meliorte the deleterious effects of slt stress. The ddition of PGPR could offer n economicl nd simple tretment to slt sensitive rdish plnts, helping to solve the production prolems cused y high slinity. The study does not provide evidence for induction of slt stress tolernce t plnt tissue, cell or moleculr level. Thus, further studies re required in order to determine the effect of PGPR tested t these levels nd the efficiency of these PGPR under nturl field condition nd for other plnt species. E37 tretment could show etter effect to llevite the slt stress in rdish thn the E43. Acknowledgments We re very grteful to The Attürk University, Scientific Reserch Projects Foundtion for generous finncil support (Project Numer 27/7). References [1] M.C. SHANNON, C.M. GRIEVE, Scienti Hort. 78, 5 38 (1999). [2] M. ASHRAF, M.A. FOOLAD. Env. Exp. Bot. 59, (27). [3] FAO, (2). [4] T.J. FLOWERS, M.A. HAJIBAGHERI, N.J.W. CLIPSON, Hlophytes. Qurterly Rev. Biol. 61, (1986), pp [5] Y. GUNGOR, Z. EREZOL, Drenj ve Arzi Islhi. Ders Kiti; Ankr University: Ankr, Turkey (1994). [6] R. GEORGE, D. MCFARLANE, B. NULSEN, Hydrogeology Journl 5, 6 21 (1997). [7] H. MARSCHNER, Minerl Nutrition of Higher Plnts (2nd. Edition) London, GB: Acdemic Press. Inc. (1995), p [8] M. QADIR, A. GHAFOOR, G. MURTAZA, Lnd Degrdtion Dev. 11, (2). [9] B. MURILLO AMADOR, H.G. JONES, C. KAYA, R.L. AGUILAR, J.L. GARCIA HERNANDEZ, E. TROYO DIEGUEZ, N.Y. AVILA SERRANO, E. RUEDA PUENTE, Environ. Exper. Botny 58, (26). [1] M. BACILIO, H. RODRIGEUEZ, M. MORENO, J. HARNENDEZ, Y. BASHAN, (24) Biol. Fertil Soils 4, [11] C. GARCIA, T. HERNANDEZ, Plnt Soil 178, (1996). [12] S. MAYAK, T. TIROSH, B.R. GLICK, Plnt Science 166, (24). [13] M. WOITKE, H. JUNGE, W.H. SCHNITZLER, Act Hort. 659, (24). [14] C.A. BARASSI, G. AYRAULT, C.M. CREUS, R.J. SUELDO, M.T. SOBRERO, Scienti Hort. 19, 8 14 (26). [15] E.V. MAAS, G.J. HOFFMAN, Crop slt tolernce Current ssessment. J Irr Drin Div Proc Am Soc Civ Eng. 13, (1977). [16] S. MEHTA, C.S. NAUTİYAL, Curr. Microiol. 43, (21). [17] C. KAYA, D. HIGGS, F. INCE, B.M. AMADOR, A. CAKIR, E. SAKAR, J. Plnt Nutrition 26, (23). [18] AOAC (Assocition of Officil Anlyticl Chemists Interntionl), Officil Methods of Anlysis, 18 th ed., Dr. Willim Hortwitz (Editor), Dr. George W Ltimer (Assist Edit), AOAC Int. Suite 5, 481 North Frederick Avenue, Githersurg, Mrylnd 25, pp , USA. [19] D. CUPPELS, F. SAHIN, S.A. MILLER, Phytopthology 89, (1999). [2] A.Z. ZAHIR, M. ARSHAD, W.T. FRANKENBERGER, Adv. Agron. 81, (24). [21] M.A. KHAN, I.A. UNGAR, Cn. J. Bot. 75, (1997). [22] E. YILDIRIM, A. DURSUN, I. GUVENC, A.M. KUMLAY, Act Agroot. 55, 75 8 (22). [23] L.F.M. MARCELIS, J. VAN HOOIJDONK, Plnt nd Soil 215, (1999). [24] M. JAMIL, S. REHMAN, K.J. LEE, J.M. KIM, H.S. KIM, E.S. RHA, Sci. Agric. 64, (27). [25] H. BOCHOW, S.F. EL SAYED, H. JUNGE, A. STAUROPOULOU, G. SCHMIEEKNECHT, J. Plnt Dis. nd Prot. 18, 21 3 (21). [26] E. YILDIRIM, A.G. TAYLOR, Annul Report of Ben Improvement Coopertive 48, (25). [27] E. YILDIRIM, A.G. TAYLOR, T.D. SPITTLER, Sci. Hortic. 111, 1 6. (26) 3942 Roum. Biotechnol. Lett., Vol. 13, No. 5, (28)

11 Mitigtion of slt stress in rdish (Rphnus Stivus l.) y plnt growth promoting rhizocteri [28] Y. BASHAN, G. HOLGUIN, L.E. DE BASHAN, Cn. J. Microiol. 5, (24). [29] H. BERTRAND, R. NALIN, R. BALLY, J.C. CLEYET MAREL, Biol. Fertil Soils 33, (21). [3] A.K. PARIDA, A.B. DAS, Ecotoxicology nd Environmentl Sfety 6, (25). [31 ] C.M.J. PIETERSE, J.A. VAN PELT, B.W.M. VERHAGEN, J. TON, S.C.M. VAN WEES, K.M. LEON KLOOSTERZIEL, L.C. VAN LOON, Symiosis 35, (23). [32] C.M.W. SASKIA, M. LUIJENDIJK, I. SMOORENBURG, L.C. VAN LOON, C.M.J. PIETERSE, Plnt Mol. Biol. 41, (1999). [33] F.V. BREUSEGEM, E. VRANOVA, J.F. DAT, D. INZE, Plnt Sci. 161, (21). [34] H. RODRIGUEZ, R. FRAGA, Biotechnol Adv. 17, (1999). [35] M.E. PUENTE, Y. BASHAN, C.Y. LI, V.K. LEBSKY, Plnt Biol. 6, (24). [36] A. GUREL, R. AVCIOGLU, Physiology of Resistnce to Stress in Plnts. In plnt Biotechnology Vol.2: genetic Engineering nd Its Applictions. Eds. Ozcn S, Gurel E nd Boğlu M. pp Pulishing of S.U. (In Turkish) (21) [37] E. SERGEEVA, S. SHAH, B.R. GLICK, World J. Micro. Biotech. 22, (26). [38] M. ASHRAF, S. HASNAIN, O. BERGE. T. MAHMOOD, Biol. Fert. Soils. 4, (24). [39] B. LIPPMANN, V. LEINHOS, H. BERGMANN, Angew Bot. 69, (1995). Roum. Biotechnol. Lett., Vol. 13, No. 5, (28) 3943

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