Effects of aluminum toxicity on plant height, total chlorophyll (Chl a+b), potassium and calcium contents in spinach (Spinacia oleracea L.

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1 Interntionl Journl of Frming nd Allied Sciences Aville online t IJFAS Journl /76-82/ 29 Ferury, 2016 ISSN IJFAS Effects of luminum toxicity on plnt height, totl chlorophyll (Chl +), potssium nd clcium contents in spinch (Spinci olerce L.) Melik Krimei 1, Vhid Poozesh 1* 1 Biology Fculty, Dmghn University, Dmghn, Irn Corresponding uthor: Vhid Poozesh ABSTRACT: Toxicity of hevy metls such s luminum nd their ccumultions in the food chin is one of the min environmentl issues in griculture nd nturl ecosystems. Aluminum is one of the most importnt fctors limiting plnt growth nd reduces the efficiency of griculturl production in cid soils. Effects of the luminum chloride (AlCl3) were studied on morphologicl nd physiologicl trits in spinch (Spinci olerce L. vr. Virofly). For this purpose, ttriutes tht contins plnt height (root nd shoot), totl chlorophyll (Chl +) nd concentrtions of potssium nd clcium were studied in greenhouse. The luminum toxicity reduced the plnt growth, totl chlorophyll nd clcium concentrtions drsticlly. In the presence of the luminum, the potssium concentrtion incresed. Due to significnt reduction in plnt growth nd C sorption which is ssocited with incresed concentrtions of luminum, it cn e concluded tht Virofly vriety of spinch is not resistnt to luminum toxicity. Keywords: hevy metls; morphologicl; physiologicl; resistnt; spinch INTRODUCTION Aluminum is the second most undnt element fter oxygen nd silicon tht ility to pollute wter, soil nd the food chin (Bhlero nd Prhu., 2013). It is estimted tht out 40% of the potentilly rle lnds worldwide re cidic (Kochin et l., 2004). Acid soils (with ph of 5.5 or lower) re mong the most importnt limittions to griculturl production (Ryn nd Delhize., 2010). There re vrious Al forms in soils in monomer, polymer or solid phse. Their concentrtion depends on the hydrolysis rte of the Al compounds. Al species s Al 3+ nd Al13, re the most toxic to plnts (Merino et l., 2010). High concentrtions of hevy metls re toxic to plnts nd cuse oxidtive stress. In such circumstnces, these metl my replcing essentil elements nd disrupt metolic functions of plnts such s the formtion of pigments (Ghosh nd Singh., 2005). In ddition to cidity, luminum concentrtion, temperture nd concentrtions of ctions nd nions re other fctors tht cn ffect the mount of luminum toxicity (Wng et l., 2006). Fctors such s mmonium fertilizers, orgnic mtter, cid rin cn lso e cidifying tht is reson for soluiliztion of toxic ions of metls such s luminum, which eventully cuses prolems in griculture nd nturl ecosystems (Vitorello et l., 2005). Root growth is most sensitive indictor of luminum toxicity. The first sign of luminum toxicity is severe stunting nd thickening of the roots (Nosko et l., 1988). Numerous toxicity symptoms re lso mnifested in shoots. Some of the common responses in shoots to Al toxicity re reduction in shoot length, cellulr nd ultr-structurl modifictions in leves, symptoms resemle to phosphorus deficiency, turning mture leves drk green, stems purple nd killing lef pexes, due to reduced clcium (C) trnsport young leves curl, decresed photosynthetic ctivity, chlorosis nd folir necrosis (Wng et l., 2006). Al ppers to preferentilly impir thylkoids nd photosynthetic electron trnsport chin in most plnts (Chen et l., 2010).

2 Height of Shoot (cm) Intl J Frm & Alli Sci. Vol., 5 (2): 76-82, 2016 So, with regrd to the issue of hevy metls pollution such s luminum nd lso due to the lck of informtion out the rection of spinch to luminum toxicity; this study done to investigte the effects of vrious concentrtions of luminum on sorption of potssium nd clcium, chlorophyll content nd plnt height in oth root nd shoot orgns of Virofly spinch. Mteril nd method A fctoril experiment, in completely rndomized design with three replictions, ws done in greenhouse of Dmghn University. The photoperiod ws under 14 hour dy nd 10 hour night with temperture of 28 ± 3 C in dy nd 20 ± 3 C in night. Uniform nd homogeneous of spinch (Spinci olerce L. vr. Virofly) seeds were plnted in trys contining vermicompost nd pet moss. The seedlings were wtered dily. At four lef stge, seedlings were grown in plstic trys contining hlf strength Hoglnd's nutrient solution. Aluminum tretment ws pplied fter week. Aluminum chloride (AlCl3) is the min compound of luminum nd dded to growth medium in three levels of 0, 100 nd 200 mm. To mesure chlorophyll, some lef tissue ws homogenized with 80% cetone. After centrifugtion, extrct sorption red with spectrophotometer (UV.2100 pc) in wvelength of 646 nd 663 nm (Lichtenthler., 1989). To mesure the mount of potssium nd clcium of fresh sh ws used (Donohue nd Aho., 1992). The solution sorption red with ICP device (CBC Integr). Sttisticl nlysis Experiments were rrnged in rndomized complete lock design nd dt were sttisticlly evluted y Duncn s multiple rnge test. All sttistic nlyses were crried out using the SPSS 23 sttisticl softwre. Results nd discussion Plnt Height (roots nd shoots) Anlysis of vrince (ANOVA) showed significnt difference in height of oth shoot nd root etween different tretment of luminum chloride (Figure 1 nd 2). Al reduces sorption of wter nd nutrients such s phosphorus nd clcium nd chnge Donnn free spce. So, the movement of wter nd nutrients through plnt - from its roots to its shoots reduce tht cusing decrese in growth rte nd iomss of plnt (Gupt et l., 2013). One of the first effects of luminum toxicity is its negtive effect on plnt growth. Butler et l (2001) reported tht luminum tretments, decrese shoot height. The first effect of luminum toxicity is its negtive effect on root growth nd meiosis. Inhiition of root division is relting to decline mitotic ctivity in the meristem of root tip (Horst et l., 1982) AlCl 3 (µm) Figure 1 The effect of different concentrtions of AlCl3 on shoot height (cm). The sme nd different letters show no significnt nd significnt difference, respectively (P<5). (Vlues re men of three replictes ± SE). 77

3 Totl chlorophyll content (mg. g -1 FW) Height of Root (cm) Intl J Frm & Alli Sci. Vol., 5 (2): 76-82, AlCl 3 (µm) Figure 2 The effect of different concentrtions of AlCl3 on root height (cm). The sme nd different letters show no significnt nd significnt difference, respectively (P<5). (Vlues re men of three replictes ± SE). It hs een reported tht luminum toxicity inhiits root growth on wild grmine nd grsses (Poozesh et l., 2007; Poozesh et l., 2010). Studies on soyen plnt showed tht luminum toxicity decrese the height of the root nd shoot (Hider et l., 2007). The results of these reports re corresponding with our experimentl dt. Totl chlorophyll (Chl +) The totl chlorophyll (Chl + ) in spinch (Spinci olerce L. vr. Virofly) significntly decresed under Al stress, while it did not exhiit significnt reduction etween the different tretments of luminum in terms of impct on the mount of photosynthetic pigments (Figure 3). Considerle studies hve showed tht, hevy metls including luminum reduce photosynthesis reduction in mny plnt species (Jing et l., 2008). It hs een shown tht Al decrese in Chl synthesis y restrins the ctivity of minolevulinic cid dehydrtse responsile for the formtion of monopyrrole porphoilonogen (Pereir et l., 2006). In this experiment with incresing Al tretments, the mount of chlorophyll reduced Figure 3. Effects of luminum toxicity on the content of totl chlorophyll (Chl +). The sme nd different letters show no significnt nd significnt difference, respectively (P<5). (Vlues re men of three replictes ± SE). 78

4 Potssium content of Shoot Intl J Frm & Alli Sci. Vol., 5 (2): 76-82, 2016 This result ws consistent with the results of Zhng et l (2007) nd Rdic et l. (2010), who oserved tht Al decresed totl chlorophyll content in soyen (Glycine mx L.) nd duckweed (lemn minor L.), respectively. The te (Cmelli sinensis L.) plnt ws shown tht physiologicl prmeters such s pigments nd photosynthetic rte hve declined ecuse of the toxicity (Mukhopdyy et l., 2012). Al 3+ ion disle mny enzymes of chloroplsts. Enzyme of Delt-minolevulinic cid dehydrtse is needed y plnts to synthesis chlorophyll. Mg 2+ compete with Al 3+ for connect to ctive site of enzyme, which leds to inhiiting of chlorophyll synthesis (Pereir et l., 2006 ; Zheng et l., 2005). Hider et l. (2007) found tht Al decresed Chlorophyll tht coincided with decline of Mg nd Fe sorption. Potssium content The potssium content ws influenced y Al stress, while it incresed significntly under Al stress in oth root nd shoot (Figure 4 nd 5). Potssium is one of the most undnt ions in plnt cells tht forms out 1 10% of dry weight of plnts (Epstein nd Bloom., 2005). Potssium cn mintin electricl potentil grdient memrne (Cheesemn nd Hnson., 1979), cell development (Mouline et l., 2002), opening nd closing of stomt (Dietrich et l., 2001). Activities of numer of enzymes, nitrte nd sucrose trnsition nd stiliztion of intrcellulr nion is one of the potssium iologicl functions (Brito nd Kronzucker., 2008). Severl studies hve demonstrted tht incresing the Al supply could increse the potssium sorption in plnts (Lin., 2010; jn., 1991 nd Mcedo nd Jn., 2008). These reports re ppers to e in greement with re oservtions. Also, Thornton et l. (1986) found tht low concentrtions of luminum cn incresing potssium content while with the rise of luminum concentrtions, the potssium levels slightly reduce. It hs een reported tht Al resistnt genotypes showed higher uptke, influx, nd trnsport of potssium thn did the Al sensitive Genotypes (Mrino nd Keltjens., 2005). Uptke of the monovlent ction K hs een shown to e less inhiited y Al thn the divlent ctions C nd Mg (Rengel nd Roinson, 1989). Also, it hs een showed tht in certin cses the K uptke t low concentrtions of Al my e even higher thn in solutions without Al (Rengel nd Roinson, 1989) Figure 4. Effects of luminum toxicity on the potssium content in shoot. The sme nd different letters show no significnt nd significnt difference, respectively (P<5). (Vlues re men of three replictes ± SE). 79

5 Clcium content of Shoot potssium content of Root Intl J Frm & Alli Sci. Vol., 5 (2): 76-82, c 0.5 Figure 5. Effects of luminum toxicity on the potssium content in root. The sme nd different letters show no significnt nd significnt difference, respectively (P<5). (Vlues re men of three replictes ± SE). Clcium content The clcium content ws influenced y Al stress, while it decresed significntly under Al stress in oth root nd shoot (Figure 6 nd 7). Clcium is one of the essentil elements in plnts (White nd Brodley., 2003). Clcium is regultor of growth nd development tht is essentil for mny metolic pthwys (Hepler., 2005). Al 3+ cn lock the C 2+ chnnels nd ffect cell memrne structure nd permeility (Plieth, 2005). Clcium enhnces root nd shoot growth y effect on cell elongtion (White nd Brodley., 2003). Ryn et l. (1997) demonstrted tht high quntity of C 2+ is present in the poplst, where it mintins cell nd tissue viility. M et l. (2007) indicted tht out 85-99% of totl Al present in poplstic content, nd C 2+ is displced from negtive inding sites y Al 3+ in the poplsm (Kinride, 1998). Al 3+ my inhiit the influx of divlent ctions s C 2+ into cells more thn monovlent ctions, ut it stimultes the nion cell influxes (Merino et l., 2010). Disruption of clcium (C) uptke y AI hs long een considered possile cuse of toxicity, nd recent work with whet (Triticum estivum L.) hs demonstrted tht C uptke t the root pex in n AI-sensitive cultivr ws inhiited more thn in tolernt cultivr (Ryn nd Kochin., 1993). These results gree with our finding c Figure 6. Effects of luminum toxicity on the clcium content in shoot. The sme nd different letters show no significnt nd significnt difference, respectively (P<5). (Vlues re men of three replictes ± SE). 80

6 Clcium content of Root Intl J Frm & Alli Sci. Vol., 5 (2): 76-82, c 0.2 Figure 7. Effects of luminum toxicity on the clcium content in root. The sme nd different letters show no significnt nd significnt difference, respectively (P<5). (Vlues re men of three replictes ± SE). In conclusion, our results demonstrte tht clcium my ply role Al tolernce in Virofly spinch. Essentil elements my enhnce plnt tolernce to Al toxicity. Due to significnt reduction in plnt growth nd C sorption which is ssocited with incresed concentrtions of luminum, it cn e concluded tht Virofly vriety of spinch is not resistnt to luminum toxicity. REFERENCES Bhlero SA nd Bhlero DV Aluminium Toxicity in Plnts A Review. J. Applicle Chem. 2 (3), Cheesemn JM nd Hnson JB Mthemticl nlysis of the dependence of cell potentil on externl potssium in corn roots. Plnt Physiol. 63, 1 4. Chen LS, Qi YP, Jing HX, Yng LT nd Yng GH Photosynthesis nd photoprotective systems of plnts in respons to luminium toxicity. Afr. J. Biotechnol. 9 (54), Dietrich P, Snders D nd Hedrich R The role of ion chnnels in light-dependent stomtl opening. J. Exp. Bot. 52, Doln L nd Dvies J Cell expnsion in roots. Curr. Opin. Plnt Biol. 7, Echrt CL nd Cvlli-Molin S Aluminum phytotoxicity: Effects, tolernce mechnisms nd its genetic control. Cncer informtion needs in rurl res. Cienc. Rurl 31, Epstein E nd Bloom AJ Minerl Nutrition of Plnts: Principles nd Perspectives, 2nd Edn. Sinuer Assocites Inc., Sunderlnd MA. Hider SI, Kng W, Ghulm J nd Guo-ping Z Interctions of cdmium nd luminum toxicity in their effect on growth nd physiologicl prmeters in soyen. J. Zhejing Univ. Sci. B. 8(3), Kinride TB Three mechnisms for the clcium llevition of minerl toxicities. Plnt Physiol. 118, Kochin LV, Hoekeng OA nd Pineros MA How do crop plnts tolerte cid soils? Mechnisms of luminum tolernce nd phosphorous efficiency. Annul Reviews. Plnt Biol. 55, Jing HX, Chen LS, Zheng JG, Hn S, Tng N nd Smith BR Aluminium induced effect on photosystem Π photochemistry in citrus leve ssessed y the chlorophyll fluorescence trnsient. Tree Physiol. 28, M B Wn J nd Shen Z H2O2 production nd ntioxidnt responses in seeds nd erly seedlings of two different rice vrieties exposed to luminum. Plnt Growth Regul. 52, Mrino ED nd Keltjens WG Long-Term Effects of Aluminum Exposure on Nutrient Uptke y Mize Genotypes Differing in Aluminum Resistnce. J. Plnt Nutr. 28, Merino GC, Alerdi M, Ivnov AG nd Reyes DM Al3+ - C2+ interction plnts growing in cid soils: AL-Phytotoxicity response to clcreous mendments. J. soil. sci. plnt nutr. 10 (3), Morn N Osmoregultion of lef motor cells. FEBS Lett. 581, Mouline K, Very AA, Gymrd F, Boucherez J, Pilot G, Devic M, Bouchez D, Thiud JB nd Sentenc H Pollen tue development nd competitive ility re impired y disruption of Shker K1 chnnel in Aridopsis. Genes Dev. 16, Pereir LB, Tldi LA, Gonclves JF, Jucoski GO, Puletto MM, Weis SN, Nicoloso FT, Borher D, Roch JBT nd Schetinger MRC Effect of luminium on mino levulinic cid dehydrotse (ALA- D) nd the development of cucumer (Cuccumis Stivus). Environ. Exp. Bot. 57, Plieth C Clcium: Just nother regultor in the mchinery of life? Ann. Bot. 96, 1-8. Poozesh V, Cruz P, Choler P nd Bertoni G Reltionship etween the Al Resistnce of Grsses nd their Adpttion to n Infertile Hitt. Ann. Bot. 99,

7 Intl J Frm & Alli Sci. Vol., 5 (2): 76-82, 2016 Poozesh V, Cruz P nd Bertoni G Evlution of resistnce to Al toxicity in wild grmine of cid medows. J. Agroecol. (In Persin). 1 (2), Rengel Z nd Roinson DL Competitive Al3+ inhiition of net Mg2+ uptke y intct Lolium multiflorum roots. I. Kinetic. Plnt Physiol. 91(1), Ryn PR nd Delhize E The convergent evolution of luminium resistnce in plnts exploits convenient currency. Funct. J. Plnt Biol. 37, Ryn PR nd Kochin LV Interction etween luminum toxicity nd clcium uptke t the root pex in ner-lsogenic lines of whet (Triticum estivum L.) differing in luminum tolernce. Plnt Physiol. 102, Ryn PR, Reid RJ nd Smith FA Direct evlution of the C 2+ - displcement hypothesis for Al-toxicity. Plnt Physiol. 113, Thornton FC, Schedle M nd Rynl DJ Effects of luminum on growth, development, nd nutrient composition of honeylocust (Gleditsi tricnthos L.) seedlings. Tree Physiol. 2, Wng JP, Hrsh R, Guo-Ping Z, Neville M nd Zhou X Aluminium tolernce in rley (Hordeumvulgre L.): physiologicl mechnisms, genetics nd screening methods. J. Zhejing Univ. Sci. B. 7, White P nd Brodley MR Clcium in plnts. Ann. Bot. 92, Yocum CF The clcium nd chloride requirements of the O2 evolving complex. Coordin. Chem. Rev. 252, Zhou G, Delhize E, Zhou M nd Ryn PR Biotechnologicl solutions for enhncing the luminium resistnce of crop plnts. In: Shnker A, Venkteswrlu B, (eds) Aiotic stress in plnts mechnisms nd dpttions. Intech-Open Access Pulisher, pp