Characterization of physiological traits, yield and fiber quality in three upland cotton cultivars grown under cadmium stress

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1 JS (11):7- (1) ISSN:1-77 hrteriztion of physiologil trits, yiel n fier qulity in three upln otton ultivrs grown uner mium stress Ling Li 1, Jinhong hen 1, Qiuling He 1, Muhmm Khn u 1,, Shuijin Zhu *1 1 eprtment of gronomy, ollege of griulture n iotehnology, Zhejing University, Hngzhou, P.R. hin eprtment of iotehnology & Geneti ngineering, Koht University of Siene & Tehnology (KUST), Pkistn *orresponene uthor: shjzhu@zju.eu.n strt mium () y plnts from ontminte soils inues vrious physiologil n iohemil hnges resulting reution in the yiel n the qulity of the plnt prouts. pot experiment ws performe in orer to stuy inue ltertions in physiologil hrters, yiel omponents n fier qulity of two trnsgeni otton ultivrs (Z-9 n SGK) n n upln otton stnr genotype (TM-1). Z-9, erive from our lortory, is glyphoste-resistnt ultivr with the PSPS-G gene, n SGK is n inset-resistnt ultivr with the t gene. stress signifintly reue the see otton yiel, lint yiel, oll numers per plnt, oll weight, net photosyntheti rte, n wter use effiieny et., while inrese the tivity of severl ntioxint enzymes suh s peroxise (PO) n superoxie ismutse (SO) in the three otton ultivrs. The umultion in ifferent prts of otton plnt inrese in the following orer: fier < see < see shell < root < lef < shoot < oll shell < petiole. The lest ffete ultivr y tretments in see otton yiel n olls per plnt mong the three otton ultivrs ws SGK, suggesting tht it ws less sensitive to stress thn the other two ultivrs. From the present experiment, it n e well onfirme tht the non-hrvestle prts of otton plnt umulte more (F=1.~.9) thn the fier n see (F=.9~.), initing tht otton is suitle for ultivtion in inustrilly pollute regions n oul e use s potentil rop for lening up ontminte soils y phytoremeition tehnology. Keywors: ntioxints; mium stress; otton; Fier qulity; Photosynthesis; Yiel. revitions: PX-sorte peroxise; F-ioonentrtion ftors; -mium; PSP--enolpyruvylshikimte--phosphte synthse; Gs-stomtl onutne; M-mlonilehye; Pn-net photosyntheti rte; PO-peroxise; ROS-retive oxygen speies; SO-superoxie ismutse; T--thiorituri i; Tr-trnspirtion rte; WU-wter use effiieny. Introution mium () is one of the most toxi environmentl pollutnts, whih hs thretene our eosystems n thus hs een hzrous to men n nimls (Wgner, 199). It n e reily tken up y plnt roots, trnslote to their eril prts n resultntly n esily enter the foo hin (Strug et l., ). intke y humn eings through plnt foos is lmost 7% (Wgner, 199). In hin, more thn 1. 1 km of griulturl soils re ontminte y, n 1. 1 kg of griulturl prouts re pollute y every yer (Gu n Zhou, ). mium is highly phytotoxi, resulting in growth retrtion, lef hlorosis or nerosis, mge to ell strutures s well s isturne in wter lne, minerl nutrition, photosynthesis, respirtion n plnt evelopment (Prs, 199). n thus it hs signifintly influene the qulittive n quntittive trits of the rops n plnt y-prouts (Myśliw-Kurziel n Strzłk, ). ue to its toxi nture, n stimulte the formtion of retive oxygen speies (ROS) either y iret eletron trnsfer involving metl tions, or s onsequene of metl meite inhiition of metoli retions (Hlliwell n Gutterige, 19). To omt ginst these SOS, vrious ntioxints n ntioxitive enzymes eome tive oth t ellulr n su-ellulr levels (Foyer et l., 1997). Regring tolerne, uptke n umultion, wie ifferenes mong plnt speies n ultivrs hve een foun in rops suh s lettue (Ltu stiv L.; rews n vies, 19) n whet (Tritium estivum L.; Zhng et l., ). ue to the vnements in the phytoremeition tehnology in reent yers, more ttention hs een pi to look for hyperumultors eing hving the pility to reue hevy metl ontent of the ontminte soils (Krämer, ). However, the pplition of phytoremeition tehnology is still very limite ue to low iomss of hyperumultors. Thus, seletive reeing for these hyperumultors is the nee of the hour in orer to selet -tolernt ultivrs. otton (Gossypium hirsutum L.) is n importnt sh rop in the worl, eing use for nturl fier n eile oil. However, its proutivity is versely ffete y vrious forms of ioti n ioti stresses. How these stresses, prtiulrly iretly n iniretly influene the iometri trits of the otton plnt nee reserh. However, stuies regring umultion s well s its effets on fier qulity re very rre. In erlier reserh y Senglevith (1999) onsiere otton s suitle rop for inustrilly pollute regions. Keeping in view of the ner unvilility of the relte informtion out physiologil n iohemil responses of the otton rop uner stress, we esigne the present reserh stuies. The ojetives of the stuy were: (1) to fin out the possile toxi effets of on otton; () to etermine the quntity n plnt prts of umultion; () to stuy the mehnism of phytoremeition of 7

2 ontminte soils using otton ultivrs. Results n isussion Photosyntheti hrteristis mium toxiity inues vrious physiologil hnges y inhiiting numer of photosyntheti prmeters suh s net photosyntheti rte (Pn), stomtl onutne (Gs) n trnspirtion rte (Tr) in the three growth perios of otton ultivrs, were shown in Figure 1(-L). Higher Pn ws oserve t the flowering n oll-forming stge thn t the uing n oll-opening stges. In ontrst with their ontrols, Pn ws higher t µm, signifintly for TM-1 n insignifintly for Z-9, then eline to the lowest t µm level t the uing stge, lthough signifint ifferene ompre with their hek ws foun only in SGK. There ws liner erese in Gs s the growth progresse, ut no signifint ifferenes were foun in Gs vlues t ll levels of tretment in the three ultivrs. Higher Tr ws oserve t the uing stge thn t the other two stges. ompre to their ontrols, Tr inrese grully t n μm for the three ultivrs, n then erese t μm t the flowering n oll-forming stge. liner inrese in Tr ws oserve with the inrese of in the three ultivrs t the flowering n oll-opening stge. Wter use effiieny (WU) in Figure 1 showe hnges similr to those of Pn ut n inverse tren ws oserve with regrs to Tr. The results from this stuy hve lerly shown tht inhiite Pn n Gs t the three growth stges of the otton ultivrs, onfirming the finings y other reserhers uring their stuies on pe (Pisum stivum L.; Htt et l., 9) n mustr (rssi june L.; Moin n Khn, 7). The reution of Pn uner stress mye ue to rresting of the photosyntheti eletron flow, inhiiting the wter-splitting omplex of the oxiizing site of PSII or ompetitively ining to the essentil + site in PSII uring photo tivtion (Fller et l., ). There were no sttistil ifferenes for Tr n Gs mong the tretment in the three ultivrs, lthough some vritions on Tr n Gs were foun mong the ifferent tretments n ifferent genotypes t the ifferent plnt evelopment stges, shown in Figure 1(-I). nzyme ssy n estimtion of lipi peroxies mium n inrese the proution of retive oxygen speies (ROS) in ells, whih n e extremely hrmful to plnts t high onentrtions (Snlio et l., 1). To protet themselves ginst ROS, plnt ells hve evelope ntioxitive mehnisms. In this experiment, the tivities of ntioxint enzymes were ssye in orer to evlute the efensive pilities ginst stress, shown in Figure (-L). ompre with their respetive ontrols, the tivity of peroxise (PO) grully inrese with n inrese in levels in SGK. Higher PO tivity ws oserve in Z-9 thn other two ultivrs, espeilly t the oll-opening stge. The tivity of superoxie ismutse (SO) inrese t n μm for TM-1 n SGK, n then erese t μm t ll growth stges, while Z-9 showe n inverse tren t the uing stge ompre with its ontrol. Superoxie ismutse is the first mjor ntioxint enzyme in the etoxifying proess s it rpily onverts superoxie rils into H O (Tnyolç et l., 7). The results from this stuy were in greement with the finings of hm et l. (11), who reporte n inrese in SO tivity when mustr plnts were expose to toxi onentrtions. ompre with the ontrols, the tivity of sorte peroxise (PX) inrese t lower levels, n then erese t μm for Z-9 t three growth stges. erese in PX tivity ws foun for TM-1 t the flowering n oll-forming stge n SGK t the uing stge t ll levels. Higher PX tivity ws oserve t the uing stge thn t the other two stges. signifint inrese in lipi peroxition ws oserve t the oll-opening stge s ompre to other two stges t ll levels, shown in Figure (-). The mlonilehye (M) ontent inrese up to the highest t μm ompre with their ontrols, following minor erese t μm in oth Z-9 n SGK, while there ws liner inrese with elevte levels in TM-1 t oll-forming stge. The ontent of M ws higher t μm thn the ontrol t ll growth stges in the three ultivrs. liner inrese in M onentrtion ws foun in the three otton ultivrs with the inrese in up to moerte level s well s with the extension of the growth stge in this experiment. Similr results hve een foun in rie (Oryz stiv L.; Shh et l., 1) n mung en (Phseolus vulgris L.; Somshekrih et l., 199) uner stress. Fier qulity The fier qulity in the three otton ultivrs with ifferent tretments is shown in Figure (-). In omprison to their ontrols, SGK possesse etter fier t lower level n more elongtion uner vrious levels thn TM-1. Z-9 showe lesser hnges thn other ultivrs se on fier length n strength expose to. The rise in + onentrtion in plnt ells, whih my hve een use y environmentl stress. Qin n Zhu (11) reporte tht + influx to the fier tips ws require for sustining fst ell elongtion. The promotion of fier elongtion expose to lower mounts of in this stuy might e ue to n elevtion in + level, whih is require for fier tips growth. lium-meite signl trnsution plys ruil role in the growth of fier tips (Qin n Zhu, 11). Therefore, it n e eue tht + might hve een entere into the ells through lium hnnels ompetitively. ompre to their ontrols, fier uniformity ws foun to e the highest t μm n then erese t higher level in the three ultivrs. There were no signifint ifferenes in fier qulity in the three ultivrs expose to. Yiel n its omponents Reution in yiel n yiel-relte trits is typil inex of plnt sensitivity to vrious stresses. In vrious rop speies (e.g. pe, whet, foer veth, rpesee n mize), yiel s reution uner multiple hevy metl stresses hs een oserve (Shrm n grwl, ). The effet of on yiel n its omponents is presente in Figure (-F). The eleterious effet of on otton yiel ws oze-epenent. With the inrese in levels, hnges in yiel n its omponents lso eme evient. s ompre to ontrol plnts, see otton yiel n oll weight first inrese up to the highest t μm, n then signifintly eline t μm for SGK, while ontinuously eresing tren ws oserve for Z-9 n TM-1. Men vlues of lint yiel n the numer of olls signifintly eline in the three ultivrs. ompre to their ontrols, the men vlues of fuzz perent progressively eline with the inrese in levels only for SGK. There were no signifint ifferenes in lint inex t ll tretment levels in the three ultivrs. Moreover, the eleterious effet of on otton yiel ws lso

3 WU(mmolO mol -1 HO) Gs(mmolHOm - s -1 ) Tr(mmolHOm - s -1 ) Pn(μmolO m - s -1 ) Tle 1. io-onentrtion ftors from soil to ifferent plnt prts or orgns in the three otton ultivrs (TM-1, Z-9, SGK). Plnt prts μm μm μm μm μm μm μm μm μm See kernel Fier Lef Petiole oll shell See shell Shoot Root Reproutive orgns Vegetle orgns ovegroun prts The whole plnts otton ultivrs 1 otton ultivrs 1 otton ultivrs F otton ultivrs otton ultivrs otton ultivrs G..... H..... I otton ultivrs otton ultivrs otton ultivrs J K L 1 otton ultivrs otton ultivrs otton ultivrs Fig 1. ffets of stress on the physiologil prmeters uring ifferent stges in the three otton ultivrs (TM-1, Z-9, SGK). : net photosynthesis (Pn) t the uing stge; : Pn t the flowering n oll-forming stge; : Pn t the oll-opening stge; : trnspirtion rte (Tr) t the uing stge; : Tr t the flowering n oll-forming stge; F: Tr t the oll-opening stge; G: stomtl onutne (Gs) t the uing stge; H: Gs t the flowering n oll-forming stge; I: Gs t the oll-opening stge; J: wter use effiieny (WU) t the uing stge; K: WU t the flowering n oll-forming stge; L: WU t the oll-opening stge. h vlue is the men of three replitions n the vertil rs give the stnr error (S) of the men. In ll figures, the ifferent letters of the sme ultivr inite signifint ifferene mong the tretments n ontrol t the % level. signifintly ifferent mong genotypes. It ws foun tht SGK ws the lest, while TM-1 ws the most ffete mong the three ultivrs. ompre to the ontrols n other tretments, yiel erese ws more prominent t μm in ll the three ultivrs. These results were in greement with those of Zhng et l. (), who foun signifint reution of whet grin weight per plnt uner tretment. umultion mium umultion in ifferent prts of the experimentl ultivrs losely orrelte with onentrtion in the soils shown in Figure (-H). The results lerly emonstrte tht the umultion enhne with the inrese in onentrtion. There ws liner inrese in uptke in ifferent plnt prts. Its umultion ws in orer of fier < see < see shell < root < lef < shoot < oll shell < petiole. In generl, roots re the primry sites through whih mium gins ess into the plnt. mium is sore y the plnt root n trnsporte to the other prts vi xylem. In most ses, lrge frtion of is retine in the roots, n only omprtively smll mounts re trnsporte to the shoots n the seelings. The results otine in this experiment were not in greement with Lozno-Roríguez et l. (1997) who inite tht umulte more in roots thn in shoots of mize n pe. These iverge ifferenes might e ue to 9

4 sorte peroxise (nmol g -1 FW min -1 ) Superoxie ismutse (Unit g -1 FW) Peroxise (nmol g -1 FW min -1 ) Mlonilehye (mmol g -1 FW) otton ultivrs otton ultivrs otton ultivrs F otton ultivrs otton ultivrs otton ultivrs 1 G 1 H 1 9 I otton ultivrs otton ultivrs otton ultivrs 1 otton ultivrs J 1 Fig. ffet of stress on the tivity of ntioxint systems (SO, PO, PX) n mlonilehye ontent uring ifferent stges in the three otton ultivrs (TM-1, Z-9, SGK). : mlonilehye (M) t the uing stge; : M t the flowering n oll-forming stge; : M t the oll-opening stge; : peroxise (PO) t the uing stge; : PO t the flowering n oll-forming stge; F: PO t the oll-opening stge; G: superoxie ismutse (SO) t the uing stge; H: SO t the flowering n oll-forming stge; I: SO t the oll-opening stge; J: sorte peroxise (PX) t the uing stge; K: PX t the flowering n oll-forming stge; L: PX t the oll-opening stge. h vlue is the mens of three replitions n the vertil rs give the stnr error (S) of the men. In ll figures, the ifferent letters of the sme ultivr inite signifint ifferene mong the tretments n ontrol t the % level. otton ultivrs K 1 otton ultivrs L ifferent mterils use in our experiment, whih inite tht the umultion in otton is ifferent to other plnt gretly. For the plnt prts investigte, petiole h the highest umultion in ll the three ultivrs. The ontent in petiole ws lmost times more thn tht in fier. This might e ue to the presene of rrier proteins in vsulr tissues (Prs, 199), n the formtion of heltion n metllothioneins in these vsulr tissues. Moreover, there ws low ontent in otton fier, suggesting tht these otton ultivrs oul e grown in pollute res. Like other fier rops, otton plnts extrt hevy metls from the soil through their root system n umulte them in the ove groun prts. These finings were in line with those of ngelov et l. () who stte tht P ws omprtively evenly istriute in otton roots n stems, n offere high resistne to pollution with the presene of protetive mehnisms in otton. In this stuy, sorption ws retine in ovegroun plnt prts n the umultion ws higher thn 1 μg g -1 t μm, n the ioonentrtion ftors (F) of ws more thn 1. (Tle 1) in the three ultivrs. It ws suggeste tht high onentrtions of oul e trnslote y otton from roots to ovegroun prts. The greter uptke in ifferent plnt prts ws oserve in SGK n Z-9. It inite tht SGK h lower mge in yiel n fier qulity thn the other otton ultivrs, n h higher potentil for soring n removing from moertely ontminte soil. It oul thus e use for the phytoremeition of pollute soils. Mterils n methos Plnt mterils, growth onitions n -tretment levels Three otton ultivrs (TM-1, Z-9 n SGK) were use in the present experiment. Z-9 is trnsgeni glyphoste tolernt otton ultivr erive from our lortory with the PSP-G gene lone y Zhejing University. SGK is n inset-resistnt ultivr with t otine from the iotehnology enter of the hinese emy of griulturl Sienes. TM-1 is the upln otton geneti stnr line otine from US, RS, ollege Sttion, Texs, US. The experiment ws rrie out t the experimentl sttion of Zhejing University. The sees of eh otton genotype were sown in pots ( m imeter m epth) on My 9 n 1. One plnt in eh pot ws grown outoors in lrge n rinproof net house till mturity. h pot ws fille with kg unontminte soil with ph=.,.99% orgni mtter, 1.% totl N (1. mg/kg ville N),.1% totl P (. mg/kg ville P), n.% totl K (11.79 mg/kg ville K). mium in form of l.h O ws e,

5 longtion (%) Strength (N/tex) Mironire Length (m) Uniformity (%) otton ultivrs otton ultivrs 9 1 otton ultivrs otton ultivrs otton ultivrs Fig. ffets of stress on fier length (), uniformity (), strength (), mironire () n elongtion () in the three otton ultivrs (TM-1, Z-9, SGK). h vlue is the men of three replites n the vertil rs give the stnr error (S) of the men. In ll figures, the ifferent letters of the sme ultivr inite signifint ifferene mong the tretments n ontrol t the % level. keeping reltively stle onentrtion t (ontrol),, n μm for the whole growing perio. There were three replitions per tretment with 1 pots for eh tretment, whih were rrnge in ompletely rnomize experimentl esign. etermintion of physiologil prmeters Physiologil prmeters suh s net photosyntheti rte (Pn), trnspirtion rte (Tr) n stomtl onutne (Gs) were mesure on the fourth lef three times from 9: to 11: t eh growing stge y portle photosynthesis system Li- (Li-or, Linoln, US). These prmeters were etermine t O flow of L min -1 n photon flux ensity of 1 μmols -1 m -. The wter use effiieny (WU) ws lulte s the rtio of net photosyntheti rte n trnspirtion rte. ntioxitive enzyme ssy n estimtion of lipi peroxies Totl superoxie ismutse (SO) tivity ws mesure spetrophotometrilly t nm (eyer n Friovih, 197). Peroxise (PO) tivity ws se on the etermintion of guiol oxition t 7 nm y H O (Pütter, 197). sorte peroxise (PX) tivity ws mesure oring to the proeure of hen n s (199). The level of lipi peroxition prouts in lef smples ws expresse s mlonilehye (M) ontent n ws etermine s -thiorituri i (T) retive metolites (Heth n Pker, 19). Mesurements of yiel prmeters, yiel n fier qulity For eh tretment, yiel n its omponents were lulte fter hrvesting. Fier properties (stple length, uniformity, strength, elongtion, n mironire) were etermine y the Supervision, Inspetion n Testing enter for otton Qulity in hin. etermintion of ontent For the quntifition of in otton plnt prts, eh smple ws rie to onstnt weight t n groun into the fine powers. They were then wet igeste for - hours fter ing mixture of - ml HNO n. ml H O. h igeste smple ws trnsferre t onstnt volume to ml in flsk for etermintion. mium ws quntifie using n IP-Mss Spetrogrph (7, gilent). Moreover, the ioonentrtion ftors (F) in ifferent prts t vrious levels were lulte oring to it et l. () using the following formul; F Sttistil nlysis shoot or root soil One-wy NOV ws performe using SS v.9 softwre n Mirosoft Offie xel for sttistil signifine t 1

6 ontent in shoot ontent in root ontent in oll shell ontent in see shell ontent in lef ontent in petiole ontent in see kernel ontent in fier Fuzz perent (%) Lint perent (%) olls No per plnt oll weight (g) See otton yiel (g plnt -1 ) otton lint yiel (g plnt -1 ) otton ultivrs otton ultivrs 1 9 otton ultivrs 1 otton ultivrs 1 otton ultivrs Fig. Yiel hrters of the three otton ultivrs (TM-1, Z-9, SGK) t hrvest sujete to inresing onentrtions. : see otton yiel; : otton lint yiel; : olls numer per plnt; : oll weight; : fuzz perent; F: lint perent. h vlue is the men of three replites n the vertil rs give the stnr error (S) of the men. In ll figures, the ifferent letters of the sme ultivr inite signifint ifferene mong the tretments n ontrol t the % level. 1 F otton ultivrs otton ultivrs otton ultivrs otton ultivrs otton ultivrs 1 1 otton ultivrs otton ultivrs G Fig. mium ontent in ifferent plnt prts of the three otton ultivrs (TM-1, Z-9, SGK) t hrvest uner vrious tretments. : ontent in ottonsee kernel; : ontent in fier; : ontent in lef; : ontent in petiole; : ontent in oll shell; F: ontent in see shell; G: ontent in shoot; H: ontent in root. h vlue is the men of three replites n the vertil rs give the stnr error (S) of the men. In ll figures, the ifferent letters of the sme ultivr inite signifint ifferene mong the tretments n ontrol t the % level. 1 1 otton ultivrs F (μm) H

7 P=.. ll the results were expresse s men ± S for three replitions. The men vlues were seprte y Lest Signifint ifferene (LS) test t % level of signifine. onlusion The phytotoxi effet of on otton yiel ws genotype s well s onentrtion-epenent. The effet eme more notle t the highest onentrtion (μm) thn t other levels. SGK ws the lest ffete mong the three ultivrs, while TM-1 ws the most sensitive one. For fier qulity, Z-9 showe greter tolerne to in fier length n strength, n SGK ws more tolernt to in fier elongtion. Less umultion ourre in fier. Therefore, it is suitle for growing in inustrilly pollute regions, sine it n remove onsierle quntities of from the soil with its non-hrveste prts. knowlegements The projet ws finnilly supporte y 97 Projet of Ntionl Nturl Siene Fountion of hin (11) n the Importnt Progrm for Trnsgeni iologil New ultivr reeing (ZX-). Referenes hm P, Ni G, shrf M (11) mium-inue oxitive mge in mustr [rssi june (L.) zern.& oss.] plnts n e llevite y sliyli i. S fr J ot 77:-. it li N, ernl MP, ter M () Tolerne n ioumultion of opper in Phrgmites ustrlis n Zemys. Plnt Soil 9: ngelov V, Ivnov R, eliltov V, Ivnov K () io-umultion n istriution of hevy metls in fire rops (flx, otton n hemp). In rop Pro 19:197-. eyer WF Jr, Friovih I (197) ssying for superoxie ismutse tivity: some lrge onsequenes of minor hnges in onitions. nl iohem 11:9-. hen GX, s K (199) sorte peroxise in te leves: ourrene of two isozymes n the ifferenes in their enzymti n moleulr properties. Plnt ell Physiol : rews HM, vies (19) Hevy metl uptke from ontminte soils y six vrieties of lettue (Ltu stiv L.). J gr Si :91-9. Fller P, Kienzler K, Krieger-Liszky () Mehnism of + toxiity: + inhiits phototivtion of Photosystem II y ompetitive ining to the essentil + site. iohim iophys t 17:-1. Foyer H, López-elgo H, t JF, Sott IM (1997) Hyrogen peroxie n glutthione ssoite mehnisms of lmtory stress tolerne n signling. Physiol Plntrum 1:1-. Gu JG, Zhou QX () lening up through phytoremeition: review of ontminte soils (in hinese with nglish strt). ol Si 1:-. Hlliwell, Gutterige JM (19) Oxygen toxiity, oxygen rils, trnsition metls n isese. iohem J 19:1-1. Htt S, rii, hou L, Kheer M, ousett H (9) Photosynthesis n growth responses of pe (Pisum stivum L.) uner hevy metls stress. J nviron Si 1:-. Heth RL, Pker L (19) Photoperoxition in isolte hloroplsts: I. Kinetis n stoihiometry of ftty i peroxition. rh iohem iophys 1: Krämer U () Phytoremeition: novel pprohes to lening up pollute soils. urr Opin ioteh 1:1-11. Lozno-Roríguez, Hernánez L, ony P, rpen-ruiz RO (1997) istriution of mium in shoot n root tissues of mize n pe plnts: physiologil isturnes. J xp ot :1-1. Moin M, Khn N (7) Photosyntheti tivity, pigment omposition n ntioxitive response of two mustr (rssi june L.) ultivrs iffering in photosyntheti pity sujete to mium stress. J Plnt Physiol 1:1-1. Myśliw-Kurziel, Strzłk K () Influene of (II), r (VI) n Fe (III) on erly steps of eetioltion proess in whet: fluoresene spetrl hnges of protohlorophyllie n newly forme hlorophyllie. gr osyst nviron 1: Prs MNV (199) mium toxiity n tolerne in vsulr plnts. nviron xp ot :-. Pütter J (197) Peroxise. In: ergmeyer HU eitor. Methos of enzymti nlysis. New York: emi Press: -9. Qin YM, Zhu YX (11) How otton fiers elongte: tle of liner ell-growth moe. urr Opin Plnt iol 1: Snlio LM, lurzo H, Gόmez M, Romero-Puerts M, el Río LR (1) mium inue hnges in the growth n oxitive metolism of pe plnts. J xp ot :1-1. Strug S, ker JR, Urenjpol S, Hswell-lkins M, Reilly P, Willims J, Moore MR () glol perspetive on mium pollution n toxiity in non-ouptionlly expose popultion. Toxiol Lett 17:-. Senglevith G (1999) Prolems of hevy metls ontmintion n using the griulturl lns in the region of KM-. Ploviv Prilozenie z ekologi km juletin KM 1:1-1. Shh K, Kumr RG, Verm S, uey RS (1) ffet of mium on lipi peroxition, superoxie nion genertion n tivities of ntioxint enzymes in growing rie seelings. Plnt Si 11: Shrm RK, grwl M () iologil effets of hevy metls: n overview. J nviron iol :1-1. Somshekrih V, Pmj K, Prs RK (199) Phytotoxiity of mium ions on germinting seelings of mung en (Phseolus Vulgris): involvement of lipi peroxies in hlorophyll egrtion. Physiol Plntrum :-9. Tnyolç, kmekçi Y, Ünln S (7) hnges in photohemil n ntioxint enzyme tivities in mize (Ze mys L.) leves expose to exess opper. hemosphere 7:9-9. Wgner GJ (199) umultion of mium in rop plnts n its onsequenes to humn helth. v gron 1:17-1. Zhng GP, Fukmi M, Sekimoto H () Influene of mium on minerl onentrtions n yiel omponents in whet genotypes iffering in tolerne t seeling stge. Fiel rop Res 77:9-9.