INFLUENCE OF CADMIUM ON ANTIOXIDATIVE DEFENCE SYSTEM, PHOTOSYNTHESIS, LEVEL OF OSMOLYTES AND IONS UPTAKE IN BRASSICA JUNCEA

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1 Interntionl Journl of Phrmy nd Phrmeutil Sienes ISSN Vol 8, Issue 10, 2016 Originl Artile INFLUENCE OF CADMIUM ON ANTIOXIDATIVE DEFENCE SYSTEM, PHOTOSYNTHESIS, LEVEL OF OSMOLYTES AND IONS UPTAKE IN BRASSICA JUNCEA ABSTRACT 1 DHRITI KAPOOR 1, AMANDEEP RATTAN 1, SATWINDERJEET KAUR 1, RENU BHARDWAJ 1 Deprtment of Botnil nd Environmentl Sienes, Guru Nnk Dev University, Amritsr-Punj (Indi) Emil: dhriti405@gmil.om Reeived: 05 Jul 2016 Revised nd Aepted: 12 Aug 2016 Ojetive: In the present study vrious physiologil nd iohemil spets of Brssi june were studied under dmium (Cd) stress onditions. Methods: Plnts of Brssi june were sujeted to different onentrtions of Cd (0, 0.2, 0.4 nd 0.6 mmol) metl. After 30 d of dmium exposure it ws found tht the level of ntioxidnts, osmolytes, photosyntheti prmeters, ions nd totl sugrs were ltered. To investigte the effets of metl in Brssi june plnts, level of sori id, toopherol, glutthione, ferri ion reduing ssy, molydte ion redution ssy, totl osmolytes ontent, nthoynins, xnthophylls, trnspirtion rte, stomtl ondutne, wter use effiieny, uptke of sodium nd potssium ions, level of ron, hydrogen, nitrogen, sulfur nd totl sugr ontent ws deteted. Results: Results from this study reveled the inrese in ntioxidnt potentil of Brssi june plnts under dmium metl stress. Photosyntheti prmeters nd uptke of sodium nd potssium ions were ffeted negtively due to metl exposure nd level of sugrs nd osmolytes were found to rise in the presene of dmium stress. Conlusion: Findings of present study suggested tht tretment of Cd tivted rnge of defene strtegies in Brssi june plnts. Keywords: Cd Stress, Osmoprotetnt, Antioxidtive defene system, Photosynthesis 2016 The Authors. Pulished y Innovre Ademi Sienes Pvt Ltd. This is n open ess rtile under the CC BY liense ( org/lienses/y/4. 0/) DOI: INTRODUCTION Hevy metl (HM) toxiity is reting troule for evolutionry, nutritionl, eologil, nd environmentl resons s these metls re mjor environmentl pollutnts [1]. Metl toxiity vries with plnt speies, speifi metl, onentrtion, hemil form nd ph of soil omposition in plnts [2]. Aumultion of higher doses of metl uses toxiity in different wys in plnts. Hevy metls my e divided into two groups: redox tive (Cu, Cr, Fe, Co) nd redox intive (Cd, Ni, Al, Zn, et.) metls. The redox tive metls re diretly involved in the redox retions of the ells nd trigger the formtion of O 2, H2O 2 nd OH susequently vi the Her-Weiss nd Fenton retion [3]. Exposure of plnts to redox intive metls results in oxidtive stress through indiret involvement in the mehnisms like disruption of the eletron trnsport hin, intertion with the ntioxidnt defene system nd indution of lipid peroxidtion. The most prevlent visul symptoms of HM toxiity is retrdtion in plnt growth[4] inluding lef hlorosis, nerosis, turgor loss, redution in the rte of seed germintion nd rippled photosyntheti pprtus, long with progressing senesene proesses nd finlly plnt deth [5]. Furthermore, HMs ffet homeostti events like wter uptke, trnsport, trnspirtion nd nutrient metolism [6] nd lso interfere with the uptke of Mg, K, C nd P [7]. Elevted levels of HMs usully inhiit photosynthesis due to their diret influene on the photosyntheti pprtus, inluding thylkoids. HM toxiity triggers the umultion of exess retive oxygen speies (ROS) inside the ell. Prodution of ROS depends upon the prtiulr HM element. Cd is redox-intive HM, whih produes ROS indiretly y intivtion of enzyme nd lso y induing the expression of lipoxygense (LOX) in plnt tissues nd onsequently leds to oxidtion of polyunsturted ftty ids [8]. Vrious stress protetive proteins, omptile solutes nd ntioxidnts re possessed y plnts in response to vrious stress onditions. They protet the plnts y inresing tolerne or voiding the toxiity of metls [9]. Brssi june elongs to fmily Cruifere (Brssiee). It is n oilseed rop, minly grown s food rop nd lso used for its mediinl purposes. It ontins ntioxidnts like rotenes, flvonoids, lutein, indoles, nd zexnthin [10], whih helps in the tivtion of ellulr defene system nd iologil system ginst oxidtive dmge. As Brssi june plnts re hyper umultor for hevy metls, thus present investigtion ws done to oserve the effets of Cd metl on the level of ntioxidnts, free rdil svenging pity, totl osmolytes, photosyntheti pigments, gseous exhnge prmeters, elementl nlysis nd sugrs in 30 d old Brssi june plnts. MATERIALS AND METHODS For experimenttion, ertified nd disese free seeds of Brssi june were proured from Punj Agriulturl University, Ludhin (Punj). Chemils nd regents CdCl2 (Sigm-Aldrih) is used for experimenttion. Antioxidnts Asori id ontent ws determined y following the method of Roe nd Kuether [11]. Toopherol ontent ws estimted y the method proposed y Mrtinek [12]. Glutthione ontent ws nlyzed y the method given y Sedlk nd Lindsy [13] Rdil svenging ssys Reduing power ssy (FRAP) ws performed y the method given y Oyizu [14] nd molydte ion redution ssy ws estimted y Prieto et l. [15]. Totl osmolyte ontent ws nlyzed y using vpor pressure osmometer (VPO) (Vpro 5600). Photosyntheti pigments Totl nthoynin ontent ws performed y the method of Minelli [16]. Xnthophylls ontent ws nlyzed y Lwrene [17].

2 Kpoor et l. Int J Phrm Phrm Si, Vol 8, Issue 10, Gseous exhnge of plnts like trnspirtion rte, stomtl ondutne nd wter use effiieny were mesured with the help of (IRGA) infr-red gs nlyzer (Li-COR 6400). Sodium nd potssium ion ontent ws mesured y flme emission photometer (systronis 128). Ions ontent ws nlyzed y method given y Allen et l. [18, 19]. The perentge of ron, hydrogen, nitrogen nd sulphur were determined with the help of CHNS nlyzer (Elementr Vrio ELIII). Totl sugrs were quntittively deteted y the method given y Sott nd Melvin [20]. Sugr ontent of 30 d old plnts were mesured y Metrohm Ion Chromtogrphy (Orion- 960). Sttistil nlysis All dt were sujeted to one-wy nlysis of vrine (ANOVA) for srutinizing the effet of Cd metl on vrious experiments nd expressed s the men±stndrd error of three replites. The Tukey post ho test (p 0.05) ws pplied for the omprisons ginst ontrol vlues using ssistt version 7.7 et. RESULTS In the present study, 1.96 folds inrese in the level of sori id ws reorded in 0.6 mmol Cd-treted seedlings (12.98 mg/g FW) in omprison to ontrol (6.61 mg/g FW). Inrese in vit-e level ws found from 3.96 to 5.29 mg/g FW i.e., from ontrol to 0.4 mmol Cd tretment. Further GSH ontent got enhned from 7.48 to 8.72 mg/g FW in Cd-stressed plnts (tle 1). Inhiition of molydte ion ws found mximum in 0.6 mmol Cdtreted plnts (83.29%) with respet to ontrol (68.66%). Inrese in the inhiition of FRAP ion ws noted from ontrol (46.45%) to 0.2 mmol Cd (49.19%). Mximum svenging of FRAP ws reported in 0.4 mmol Cd (67.11%) (tle 1). It ws found tht totl osmolyte ontent ws enhned with the metl tretment. It ws reorded mximum in 0.6 mmol Cd tretment (193.23m mol/kg), followed y 0.4 mmol Cd ( m mol/kg) nd 0.2 mmol Cd (179.17m mol/kg) tretments (tle 1). Tle 1: Effet of Cd on ntioxidnts nd ntioxidnt ssys in 30 d old B. june plnts Tretments Asori id Toopherol Glutthione FRAP Molydte ion Totl Osmolytes (µ mol/g FW) 0.0 mmol 6.61± ± ± ± ± ± mmol 8.96± ± ± ± ± ± mmol 11.63± ± ± ± ± ± mmol 12.98± ± ± ± ± ±2.47 Dt presented in men±se. Different letters (,, nd d) within vrious onentrtions of Cd (0, 0.2, 0.4 nd 0.6 mmol) re signifintly different (The Tukey post ho test, p 0.05) nd signify the effet of Cd metl on vrious ntioxidnts nd ntioxidnt ssys. It ws oserved tht ontinuous rise in nthoynin ontent from 7.45 to 9.65 mg/g FW our. A similr trend ws notied in the se of xnthophylls, where 0.4 mmol Cd tretment showed the highest xnthophylls level (9.92 mg/g DW) (tle 2). In the present study, trnspirtion rte deresed with the Cd toxiity from 1.97 to 1.11m mol H 2O m -2 s -1 In 30 d old plnts; results reveled deline in stomtl ondutne from ontrol (0.29 mol m -2 s -1 ) to 0.6 mmol Cd-stressed plnts (0.19 mol m -2 s -1 ). With inresing doses of Cd, wter use effiieny (WUE) ws found to derese from ontrol to 0.6 mm Cd-treted plnts. Its vlue deresed from 0.2 mmol (3.31 mgco 2/gH 2O) to 0.6 mmol Cd (3.02 mgco 2/gH 2O) treted plnts (tle 2). Tle 2: Effet of Cd on photosyntheti system in 30 d old B. june plnts Stomtl ondutne H Tretments Anthoynin Xnthophyll Trnspirtion rte 2O use effiieny (mg/g DW) (m mol H2O m -2 s -1 ) (mol m -2 s -1 ) (mg CO 2/gH 2O) 0.0 mmol 7.45± ± ± ± ± mmol 8.27± ± ± ± ± mmol 8.96± ± ± ± ± mmol 9.65± ± ± ± ±0.26 Dt presented in men±se. Different letters (,, nd d) within vrious onentrtions of Cd (0, 0.2, 0.4 nd 0.6 mmol) re signifintly different (The Tukey post ho test, p 0.05) nd signify the effet of Cd metl on the photosyntheti system. A steep deline ws oserved in the ions ontent due to Cd phytotoxiity from 7.63 to 6.87 ppm nd from 7.92 to 6.77 ppm respetively (tle 3). C ontent ws found to inrese from to 38.66%. Similrly, H ontent ws oserved mximum in plnts sujeted to 0.6 mmol Cd stress (5.84%), whih ws found to derese with 0.2 mmol Cd tretment (5.78%). A shrp inrese ws reorded in S ontent. The mximum vlue ws seen in the plnts exposed to 0.6 mmol Cd toxiity (0.35%). N ontent shrply delined with 0.4 mmol Cd tretment (1.13%), whih were lmost 4.07 folds less thn the vlue of ontrol (4.6%) (tle 3). Tle 3: Effet of Cd on Ion ontent in 30 d old B. june plnts Tretments Sodium ion (ppm) Potssium ion (ppm) Cron Hydrogen Nitrogen Sulphur Totl sugr (µ mol/g FW) 0.0 mmol 9.85± ± ± ± ± ± ± mmol 7.63± ± ±1 5.78± ± ± ± mmol 7.55± ± ± ± ± ± ± mmol 6.87± ± ± ± ± ± ±1.09 Dt presented in men±se. Different letters (,, nd d) within vrious onentrtions of Cd (0, 0.2, 0.4 nd 0.6 mmol) re signifintly different (The Tukey post ho test, p 0.05) nd signify the effet of Cd metl on vrious ions nd sugr ontent. 205

3 Kpoor et l. Int J Phrm Phrm Si, Vol 8, Issue 10, With inresing metl onentrtion, sugr ontent ws lso found to enhne from 0.2 mmol Cd to 0.6 mmol Cd onentrtion, i.e., from to 21.17µ mol/g FW s ompred to ontrol plnts (17.66µ mol/g FW) (tle 3). 30 d old plnts of B. june resulted in the umultion of soritol, mnnitol, gluose, frutose nd elloiose. Further, in 0.2 mmol Cd stress inrese in the mount of mnnitol from to ppm, gluose from to ppm nd frutose from to ppm ws reorded. Addition pek of sugr nmely surose ws expressed in 0.4 mmol Cd-treted plnts. Two dditionl sugrs, rinose, nd xylose were notied t the highest metl stress of 0.6 mmol Cd (fig. 1, tle 4). Tle 4: Conentrtions of sugrs in 30 d old Brssi june plnts treted with Cd stress S. No. Sugrs Conentrtions (ppm) Control 0.2 mmol Cd 0.4 mmol Cd 0.6 mmol Cd 1 Soritol _ _ 2. Mnnitol Gluose _ 4. Frutose _ 5. Celloiose _ Surose _ Arinose _ Xylose _ Fig. 1: Qulittive nlysis of sugrs y Ion hromtogrphy in 30-dys old Brssi june plnts treted with Cd DISCUSSION In the present study, ntioxidnt potentil of B. june plnts ws found to improve with inresing Cd metl stress. Improved ntioxidnt system ts s key plyer in the mitigtion of oxidtive stress produed y ROS [21]. The oservtions of the present study were supported y the findings of Choudhry et l. [22], where the level of ntioxidnts like sori id nd GSH were found to enhne in Rphnus stivus seedlings exposed to Cr stress. The report suggested tht level of GSH inresed under Cu nd Cd stress in Cleome gynndr [23]. Results reveled tht level of osmolytes ws enhned in B. june plnts sujeted to Cd stress. Osmolytes provide protetion ginst stress; they t s ntioxidnt y svenging ROS nd stilizing the memrnes [24]. Under stress onditions, Δ1pyrroline-5- roxylte synthse enzyme gets stimulted, whih further use n inrese in proline ontent. GB is lso umulted more during the stress onditions s it is formed y holine nd GB sustrtes. Two-step oxidtion of holine is stimulted vi the toxi intermedite etine ldehyde nd these retions re tlyzed y holine monooxygense (CMO) nd NAD+-dependent etine ldehyde dehydrogense (BADH), whih is tivted under stress onditions [25]. These results re in oherene with the oservtions of Choudhry et l. [22], where Cr toxiity used rise in the level of GB. Present investigtion showed the rise in the level of photosyntheti pigments with inresing Cd doses. This is due to the tivtion of ntioxidtive enzyme nmely glutthione-s-trnsferse tht uses iosynthesis of nthoynin pigment [26], ts s preursor for the synthesis of sisi id (ABA), whih is lso involved in the protetion of photosynthesis ginst oxidtive stress [27] nd thus lso ts s n ntioxidnt. The oservtions of present work re in oherene with the oservtions of Amiri et l. [28]. Wheres gseous exhnge prmeters suh s trnspirtion rte, stomtl ondutne nd wter use effiieny were found to deline with the Cd tretment. Regrding the toxiity of Cd metl to PSII tivities in plnts, it hs een investigted tht Cd inds in the sites of oth eptor nd donor sides of PSII [29]. On the donor side, the presene of Cd 2+ exhnges the C 2+ oftor in the C/Mn luster tht onstitutes the oxygen-evolving enter with high ffinity in slow retion tht uses redution of photosyntheti oxygen evolution [30]. Results were supported y the oservtions of Stnhev et l. [31], where wter use effiieny (WUE) ws deresed in Oimum silium nd inhiition of gs exhnge nd trnspirtion rtes were oserved in Orignum vulgre plnts exposed to hevy metl stress. The level of ions ws lso delined with Cd toxiity in B. june plnts. Metl tretment negtively ffets the uptke nd trnsport of minerl nutrients in the plnts [32]. A deline in the level of these ions often indites their efflux ross plsm memrne nd exess dose of metl dmge them y induing the lipid peroxidtion. Further, this dmge leds to loss of memrne 206

4 Kpoor et l. Int J Phrm Phrm Si, Vol 8, Issue 10, seletivity nd inrese in permeility [33]. Results of the present study were supported y Bouzizi et l. [34] where the level of K + ws negtively ffeted in Phseolus vulgris plnts with enhning Cu onentrtion. In Cuumis stivus plnt, Cu toxiity led to redution in N + nd K + ontent [35]. Perentge of ron, hydrogen nd sulphur ws found to enhne with inresing Cd doses. The high elementl ontent in the shoots of B. june might e relted to metl tolerne in umultors/hyperumultors. The level of elements ws rised y the enhned metl onentrtions in plnt shoots, suggesting tht B. june might hve speifi physiologil need for them if exposed to potentil metl toxiity [36]. In the present study, the level of sugrs ws notied to enhne in B. june plnts. Retive oxygen speies ret with sugrs nd oxidize them to relese formi id, whih is the min rekdown produt. Mnnitol ws found to umulte in the hloroplsts, showed inresed tolerne to oxidtive stress in trnsgeni too. As mnnitol uses the removl of HO nd therefore inresed mnnitol is the inditor of enhned stress protetion [37]. From the present investigtions, it is onluded tht Cd metl stress negtively ffets the physiology of B. june plnts in terms of inhiition of minerl nutrient uptke nd gseous exhnge prmeters. CONCLUSION In response to the dverse effets of metl, omptile solutes nd ntioxidtive defense system of the Brssi june plnt got tivted nd proved enefiil to ountert the effets of metl stress y svenging the ROS, generted during the stressed onditions. ACKNOWLEDGEMENT Authors re thnkful to Deprtment of Botnil nd Environmentl Sienes, Guru Nnk Dev University, Amritsr-Punj (Indi) for providing lortory filities for this work. ABBREVIATION ROS-retive oxygen speies, TCA-trihloro eti id, TPTZ-2,4,6- tripyridyl-s-trizine, GSH redued glutthione, DTNB-dinithionitroenzoi id, IRGA infrred gs nlyzer, VPO vpour pressure osmometer, N + -sodium ions, K + -potssium ions. 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