Triveni Enterprises, Lucknow (Indi) J. Environ. Biol. ISSN : 254-874 29(6), 959-964 (28) http : //www.geocities.com/j_environ_iol/ j_environ_iol@yhoo.com Effect of slinity nd temperture on the germintion of Spergulri mrin seeds nd meliorting effect of scoric nd slicylic cids Ysemin Ozdener* nd H. Gury Kuty Deprtment of Biology, Fculty of Science nd Art, University of Ondokuz Myis - 55139, Kurupelit, Smsun, Turkey (Received: June 11, 27; Revised received: Septemer 28, 27; Accepted: Octoer 1, 27 ) Astrct: Spergulri mrin (Cryophyllcee) is hlophytic species nd widely distriuted mong the se shores of Turkey. Its seeds my e unwinged or winged. Lortory experiments were conducted to determine the effects of slinity (, 5, 1 nd 5 mm NCl) nd temperture (1, 15, 25, 3, 35 o C) on seed germintion. S. mrin showed 73.3 % germintion in non-sline controls t 25 o C. No germintion occurred t 3 o C nd 35 o C. Seed germintion of Spergulri mrin ws significntly ffected y slinity levels, temperture nd their interctions. Mximum finl germintion percentge occured under the lowest slinity condition (5 mm) nd distilled wter t 25 o C. Seed germintion ws completely inhiited y 5 mm NCl, lthough seed germintion rte ws not ffected y slinity. Recovery germintion ws gretest in 5 mm nd t 25 o C.The rte of germintion ws significntly ffected y temperture (p<.1). The germintion percentge of the seeds pretreted with 4 mm nd 6 mm L-ASA in 5 mm nd 1 mm NCl ws improved compred with tht of untreted L-ASA. Addition of.5 mm slicylic cid (SA) improved the germintion in 5 mm NCl. Key words: Spergulri mrin, Slinity, Temperture, Germintion, Ascoric cid, Slicylic cid PDF of full length pper is ville with uthor (*ysemino@omu.edu.tr) Introduction Slinity stress is mjor limiting fctor for plnts germintion growing in costl hitts s it is one of the most criticl periods in life cycle of hlophytes (Gilles et l., 21; Ruio Csl et l., 23). Slt stress ffects germintion percentge, germintion rte nd seedling growth in different wys depending on the plnt species (Ungr, 1996; Gul nd Weer, 1999 ). It ws reported tht mximum germintion of the seeds of hlophytic plnts occurred in distilled wter or under reduced slinity (Gul nd Weer, 1999; Khn nd Gulzr, 23; Crter nd Ungr, 23), nd it hs een found tht germintion percentge is reduced with high NCl concentrtions (Toe et l., 21; Pujol 2; Ruio-Csl et l., 23). Germintion of hlophytes typiclly occurs fter rins which provide moisture nd leching the slt ions (Khn nd Ungr, 1997; Vincente, 24; El Kelwy, 24; El Kelwy nd Al Rwi, 25; Song et l., 26). Temperture is lso n importnt fctor tht determines germintion. The effects of slinity nd temperture on germintion of mny hlophytic plnts hve een studied y severl uthors nd response of the plnts to temperture nd slinity vries ccording to slinity response of the hlophytic species (Gul nd Weer, 1999; Khn nd Ungr, 21; Khn et l., 21; El Kelwy nd Al Rwi, 25; Al Khtee, 26; Gori et l., 26; Song et l., 26; Cicek nd Tilki, 27; Tilki nd Dirik, 27). Slinity my cuse disturnces in plnt metolism (El Tye, 25). Slinity- induced oxidtive stress could e reson for germintion inhiition (Amor et l., 25). The strtegy of osmotic djustment vried from one plnt to nother s well s tissues. Mny trils hve een mde to help the plnts to overcome these disturnces using vrious tretments in lortory. Slicylic cid (SA) plys n importnt role in iotic plnt stress tolernce incresing their resistnce to slinity, wter deficit, low temperture s well s detrimentl ction of hevy metls (Mishr nd Chudhri, 1997; Mishr nd Chudhri, 1999; Senrtr et l., 2; Bezrukov et l., 21; Shkirov et l., 23; Singh nd Ush, 23; Drzic nd Mihilovic, 25; El Tye, 25; Gunes et l., 27; Chndr et l., 27). Exogenously pplied slicylic cid my improve germintion of hlophytes. Similrily ppliction of L-scoric cid llevited the se slt effects in some hlophytes (Khn et l., 26). Also L-scoric cid (L-ASA) meliorted the toxic effect of hevy metls in nimls (Singh nd Rn, 27). Little informtion is ville on the effect of slicylic cid nd scoric cid on the germintion of the seeds under slt stress. Spergulri mrin (Cryophyllcee) is widely distriuted hlophytic species through the seshores of Turkey. The seeds of this species my e unwinged or winged. Previous experiments indicted tht there is n interction etween the effects of temperture nd slinity on germintion of S. mrin (Ungr, 1967). Germintion rte ws significntly reduced t higher slinity levels (Keiffer nd Ungr, 1997) nd high slt concentrtion with incresing temperture displyed n inhiitory effect on germintion (Ungr, 1967). Therefore, the ims of the present study were: 1) to investigte the effect of temperture nd slinity on the germintion of the seeds of S. mrin, 2) to determine other germintion chrcteristics (i.e. germintion rte, men dily germintion, recovery germintion) 3) to show improving effect of slicylic cid nd L- scoric cid on the germintion of the seeds under slt stress. Journl of Environmentl Biology Novemer, 28
96 Mterils nd Methods Seed collection: The seeds of S. mrin were collected in July 24 from pure stnd in sline soils in Smsun (41 o 42 17 nd 41 o 31 46 N nd 36 o 5 4 nd 36 o 3 1 E). Seeds were dried in lortory nd stored t room temperture efore eing used (seed weight fter drying : 44±2 mg 1-1 seeds). Effect of temperture nd slinity on seed germintion: Germintion trils were crried out in sterilized petri dishes on two lyers of filter pper moistened with 1 ml solution of 5, 1 or 5 mm NCl or distilled wter (control). Temperture ws kept constnt t 1, 15, 2, 25, 3 nd 35 o C. All petri dishes were kept in n incutor under continuous drkness for 15 dys. Ech of the three replictes of 1 seeds ech were used. The solution in ech petri dish ws renewed every three dys. Germinted seeds were counted, then these seeds were removed from petri dishes. Seeds were considered germinting with the emergence of the rdicle. Four prmeters of germintion were determined: 1. Finl germintion percentge 2. Germintion rte (GR): is mesure of rpidity of germintion, with lower vlues indicting fster germintion. It is clculted s follows: GR = (n 1 t 1 ) + (n 2 t 2 ) +..+(n x t x ) / x n Where n 1 is the numer. of germinnts t the first dy of germintion, t 1 is the dys from strt to first germintion, nd xn the totl numers of seeds germinted (Ruio-Csl et l., 23). 3. Men dily germintion (MDG): where MDG = Finl germintion percentge numer of dys to finl germintion percentge (Ruio-Csl et l., 23). 4. Recovery test: After 15 dys, ungerminted seeds from ech petri dish with 5, 1 nd 5 mm NCl were trnsferred to distilled wter nd the germintion ws recorded fter seven dys. The totl germintion ws determined y the following formul: ( + ) / c x 1 where is the totl numer of seeds germinted in sline solution, is the totl numer of seeds germinted in distilled wter fter trnsferred from NCl condition from distilled wter (Song et l., 26) nd c totl numer of seeds sowned. Recovery tests were mde t 1, 2 nd 25 o C. The seeds of S. mrin were exposed to NCl nd distilled wter pplictions nd kept t 12:12 light/drk photoperiods during the preliminry study which conducted to determine the effects of light on seed germintion. After 15 dys of the light ppliction it ws Ysemin Ozdener nd H. Gury Kuty determined tht there were no significnt differences etween drk nd 12:12 light/drk period pplictions. So tht the study ws continued over drk period. The effect of L-scoric cid nd slicylic cid on seed germintion in Ncl. L- Ascoric cid: The seeds of S. mrin were pretreted with L- scoric cid (2, 4 nd 6 mm) for 4 hr nd ir dried efore immersing in NCl solutions (, 5, 1 nd 5 mm). The experiments were crried out in n incutor t continuous drk nd 25±2 o C. Germinted seeds were counted nd finl germintion percentge ws determined. Slicylic cid: The seeds were imied with slicylic cid (.5, 1 nd 2 mm) for 24 hr nd ir dried, then were sown in petri dishes with NCl solution (, 5, 1 nd 5 mm). Finl germintion percentge ws determined. Sttisticl nlysis: Sttisticl nlysis ws performed using SPSS 1. version (Anonymous, 1999). Two-wy nlysis of vrince (ANOVA) ws performed to show the differences in seed germintion prmeters depending on temperture nd NCl concentrtions. Tukey s honestly significnt difference (HSD) test ws used to rnk mens following nlysis of vrince. Results nd Discussion The effects of temperture nd slinity on germintion: Finl germintion percentge of S. mrin seeds ws significntly ffected y the temperture, slinity nd the interctions of oth iotic fctors (p <.1) (Tle 1). Germintion did not occured t 3 o C nd 35 o C, in ll NCl concentrtions. Seed germintion ws highest in distilled wter t 25 o C second highest in 5 mm. Germintion decresed t lower temperture (1 o C) compred with tht of 25 o C. Compred with distilled wter, germintion percentge decresed little in 5 mm NCl, while it ws severely ffected y 1 mm NCl t 25 o C. Germintion ws completely inhiited in 5 mm t 1, 15 nd 25 o C (Fig. 1). The rte of germintion ws the highest t 25 o C nd the lowest t 1 o C (Fig. 2). Chnges in temperture significntly effected the rte of germintion (p<.1) while vrious slinity nd interction of temperture-slinity did not effect the germintion rte (Tle 1). There were significnt differences etween slinity nd temperture (p<.1) in terms of men dily germintion (MDG). The highest MDG ws otined t 25 o C in distilled wter while it decresed t lower temperture nd slinities (Fig. 3). Recovery germintion: Seeds showed recovery from slinity stress t ll tempertures. There ws reltively lower recovery t lower tempertures (1-15 o C) in high slinity tretments (1-5 mm). However t 25 o C, there ws sustntil recovery t higher slinities (Fig. 4). Journl of Environmentl Biology Novemer, 28
Germintion of Spergulri mrin Finl germintion (%) 9 8 7 6 5 4 3 2 1 1 C 15 C 25 C Temprture Fig. 1: Finl germintion percentge (%) with respect to temperture nd NCl concentrtions Germintion rte 4 35 3 25 2 15 1 5 1 C 15 C 25 C Temperture Fig. 2: Germintion rte with respect to temperture nd NCl concentrtions 961 Men dily germintion 14 12 1 8 6 4 2 c Recovery germintion 12 1 8 6 4 2 1 C 15 C 25 C Temprture 1 C 15 C 25 C Temperture Fig. 3: Men dily germintion (MDG) with respect to temperture nd NCl concentrtions Fig. 4: Recovery germintion with respect to temperture nd NCl concentrtions Germintion (%) 9 8 7 6 5 4 3 2 1 c 2 4 6 ASA (mm) Germintion (%) 9 8 7 6 5 4 3 2 1.5 1 2 SA (mm) Fig. 5: Germintion percentge with respect to L-scoric cid nd NCl concentrtions Fig. 6: Germintion percentge with respect to slicylic cid nd NCl concentrtions Control 5 NCl 1 NCl Journl of Environmentl Biology Novemer, 28
962 Ysemin Ozdener nd H. Gury Kuty Tle - 1: Results of two- wy ANOVA of chrcteristics of seed germintion when seeds were moistened with, 5, 1 nd 5 mm NCl t constnt 5, 1, 15, 25, 3 nd 35 o C in continuous drkness Independent vrile Temperture NCl Temp. & NCl Totl men ± SD Finl germintion percentge 5436.74** 1144.74** 561.815** 41.18 ± 24.76 Germintion rte 1548.98** 1.437 NS.78 NS 16,2 ± 1,95 Men dily germintion 18.1** 19.62** 12.41** 4.5 ± 3.44 Recovery germintion 6912.111 152.111** 383.389* 55.66 ± 25.62 Note: Numer represents men squre (*p <.5 ; **p <.1, NS : Not significnt), SD: Stndrd devition Tle - 2: Sttisticl nlysis of the effect of slicylic cid (SA) on the germintion of S. mrin seeds y two wy ANOVA Slicylic cid Independent vrile SA NCl SA & NCl Totl men ± SD Finl germintion percentge (%) 572.917** 3231.361** 56.472* 32.52 ± 29.61 Note: Numer represents men squre (*p <.5; **.1) SD: Stndrd devition Tle - 3: Sttisticl nlysis of the effect of scoric cid (A) on the germintion of S. mrin seeds y two wy ANOVA Ascoric cid Independent vrile L-ASA NCl L-ASA & NCl Totl men ± SD Finl germintion percentge (%) 413.731** 121.583** 497.62* 56.8 ± 15.36 Note: Numer represents men squre (**p <.5), SD: Stndrd devition Recovery of germintion ws significntly ffected y temperture (p<.1) nd the interctions of temperture with slinity (p <.5) (Tle 1). However there were no reltion etween recovery nd vrious slt concentrtions. Effect of L- scoric cid (L-ASA) nd slicylic cid (SA) on the seed germintion: L-scoric cid pretretments llevited the inhiitory effect of slt (5 mm nd 1 mm) on seed germintion of S. mrin (Fig. 5). The germintion percentge pretreted with 4 mm nd 6 mm L-ASA in 5 mm nd 1 mm NCl ws similr with tht of untreted seeds. The effect of L-ASA, NCl nd the interction of L-ASA nd NCl were sttisticlly significnt (p<.5) (Tle 3). Improved effect of L-ASA ws not oserved in 5 mm NCl. The germintion percentge of the seeds pretreted with L-ASA (2, 4 nd 6 mm) decresed in distilled wter (Fig. 5). Finl germintion percentge ws significntly improved y L- scoric cid pretretments t 5 nd 1 mm slt concentrtions. Effects of L- scoric cid pretretments on seed germintion of hlophytic species were found to e species- specific (Khn et l., 26). Finl germintion stge of S. mrin seeds emedded in 4 nd 5 mm scoric cid ws found very close to the control group when they were incuted in 5 mm NCl medi. A very slight improved effect ws oserved when they were incuted with 1 mm NCl. Improving effect of.5 mm slicylic cid on the germintion ws oserved in 5 mm NCl (Fig. 6). But germintion percentge of the seeds pretreted with 1 mm SA decresed in distilled wter nd ll slinity conditions tested. SA, NCl nd the interction of oth the fctors were significnt (Tle 2). The germintion of the seeds pretreted with 2 mm SA ws completely inhiited under ll conditions (Fig. 6). Spergulri mrin is n nnul plnt nd it hs cosmopolitn distriution nd occurred in costl hitts ll over the world. S. mrin hs occupied in sline res with soil totl slinity concentrtions out 4.% (Ungr, 1992). But seeds hve een reported to germinte in NCl concentrtions of 1.% ut not t 2.% under lortory conditions (Keiffer nd Ungr, 1997). Ungr (1967) ws determined tht the mximum germintion temperture ws 13-21 o C in non-sline nd low slt concentrtion (.5%). During the preliminry study it ws determined tht light hs no dominnt effect on the germintion of S. mrin seeds (dt not shown). However, Crter nd Ungr (24) explined tht light positively effects on the germintion of S. mrin seeds. This difference my e due to geogrphicl differences. The results of this study indicte tht the seeds of S. mrin were susceptile to slinity t germintion stge. Hlophyte seed germintion hs een reported to occur optimlly under reduced slinity stress (Ungr, 1967; Khn nd Ungr, 1997; Gul nd Weer, 1999; Aizzi et l., 22; Khn nd Ungr, 21; Khn et l., 21; Khn nd Gulzr, 23; Ruio-Csl et l., 23; El Kelwy nd Al Rwi, 25). However the effect of slinity nd temperture ws different on the germintion of some hlophyte seeds. For exmple, reports indicte tht some hlophyte seeds germinted t higher Journl of Environmentl Biology Novemer, 28
Germintion of Spergulri mrin 963 tempertures, nd mximum germintion occured under oth nonsline nd sline conditions (Khn et l., 2; Khn et l., 21; Song et l., 26). High slinity cuse germintion inhiition t higher tempertures, it suggests tht the mechnism of N + toxicity my not e present in some species (Aizzi et l., 22). In our study, the increse of temperture nd slt concentrtion effects germintion rte (GR). Exposure to lower or higher tempertures nd higher slt concentrtions not only provoke inhiition of germintion percentge ut lso decrese in germintion rte. In this reserch, germintion rte ws significntly reduced t 1 nd 15 o C, ut the effect of increse in slt concentrtion ws not sttisticlly significnt. In the seeds of Arthrocnemum mcrostchyum, GR ws relted to sliniy nd slowed in high slt concentrtion (Ruio-Csl et l., 23). Similr results were lso otined from Prosopis juliflor seeds (El Kelwy nd Al Rwi, 25) Slinity inhiits germintion of hlophyte seeds in one of two wys: (1) preventing germintion without loss of viility t higher slinities; nd (2) delying germintion of seeds t slinities tht cuse some stres to seeds ut not prevent germintion (Gulzr nd Khn, 21). The response of seeds of S. mrin trnsferred to distilled wter fter 15 d t vrious slinities vried depending on the temperture. Seeds exposed to higher slinity (1-5 mm NCl) recovered quickly t moderte temperture (25 o C). Under hypersline conditions, seed survivl rther thn germinility my e n pproprite criterion for success, since recovery germintion occur in seeds of hlophytes when hypersline conditions re llevited (Pujol et l., 2; Gulzr nd Khn 21; Hung et l., 23). However the recovery germintion of seeds tht were previously exposed to hypersline conditions ws ffected y temperture regime (Keiffer nd Ungr, 1997; Gulzr nd Khn, 21; Song et l., 26). Similr results were lso reported for Slicorni rmosissim nd Arthrocnemum mcrostchyum (Ruio Csl et l., 23) nd H. mmodendron (Hung et l., 23). Seed viility ws not ffected y exposure to high slt concentrtions nd the seeds of the species exposed to higher slinity showed greter germintion percentge in recovery germintion. At germintion level rective oxygen species (ROS) re generted minly during depletion of food reserves nd oxidtive phosphoryltion, while their quntittive level is conntrolled y seed s protective ntioxidnt system. However, it hs een reported tht seeds contin vrious ntioxidnts in smll mounts nd compounds like scoric cid re not present. An rtificil increse in cellulr level of n ntioxidnt such s scoric cid should e eneficil in improving stres tolernce t germintion level (Khn et l., 26). Detiled studies on components of ntioxidnt defense systems of seeds of hlophytes might provide stisfying evidence. Our results re consistent with those of Shkirov et l., (23) nd El Tye (25), who showed n encourgement in seed germintion with SA ppliction. Improved effect of slicylic cid on germintion of S. mrin seeds were oserved. It hs een found tht finl germintion rte ws higher when.5 M slicylic cid emedded seeds were incuted in 5 mm NCl contining medi nd control medi (without NCl) s compred to seeds without emedded in slicylic cid. The effect of NCl nd slicylic cid interction on finl germintion rte ws sttisticlly significnt. Since slicylic cid is n endogenous growth regultor, it ws importnt to study the influence of exogenous SA on seed germintion (El Tye, 25). Slicylic cid (SA) is n endogenous growth regultor of phenolic nture. Exogenous ppliction of SA my influence rnge of diverse processes in plnts, including seed germintion, stomtl closure, ion uptke nd trnsport, photosynthesis nd growth (El Tye, 25 ). 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