J. Elmntol. 2008, 13(4): 597-604 597 EFFECT OF COPPER DEFICIENCY ON GAS EXCHANGE PARAMETERS, LEAF GREENNESS (SPAD) AND YIELD OF PERENNIAL RYEGRASS (LOLIUM PERENNE L.) AND ORCHARD GRASS (DACTYLIS GLOMERATA L.) Mrznn Olszwsk, Stn Grzgorzyk, Jk Alrski, Ann B³uh-M³k, Am Kozikowski Chir o Grssln Mngmnt Univrsity o Wrmi n Mzury in Olsztyn Astrt Coppr is ssntil or th growth n vlopmnt o plnts. This mironutrint ts hlorophyll ontnt, photosynthsis pross n wtr rltions in th plnt. Th ojtiv o this stuy ws to trmin th t o oppr iiny in soil on th rt o photosynthsis n trnspirtion, wtr us iiny, l grnnss n th yil o slt ultivrs o prnnil rygrss (Lolium prnn L.) n orhr grss (Dtylis glomrt L.). During th growing sson, th rt o photosynthsis n trnspirtion ws msur using LI-COR 6400 gs nlyzr (Portl Photosynthsis Systm), n l grnnss ws stimt with Minolt SPAD-502 hlorophyll mtr. Wtr us iiny (WUE) ws lult s on instntnous vlus o photosynthsis n trnspirtion. Dry mttr yil ws trmin y grn mttr rying to onstnt wight t 105 o C. Th rsults o th stuy init tht oppr iiny signiintly rs th rt o photosynthsis n trnspirtion, hlorophyll onntrtion in lvs n th yil o ll invstigt ultivrs. Prnnil rygrss v. Mj ws oun to most rsistnt to oppr iiny it ws hrtriz y high rt o photosynthsis n trnspirtion, n y th highst hlorophyll ontnt. Th yil o v. Mj ttin unr oppr iit onitions ws omprl to tht o othr ultivrs grown unr ontrol onitions. Ky wors: oppr iiny, prnnil rygrss, orhr grss, photosynthsis, trnspirtion, wtr us iiny (WUE), l grnnss (SPAD), yil. r in. Mrznn Olszwsk, Chir o Grssln Mngmnt, Univrsity o Wrmi n Mzury, pl. ózki 1/8, 10-718 Olsztyn, Poln, phon: (089) 523 35 01, -mil: mrznn. olszwsk@uwm.u.pl
598 WP YW NIEDOBORU MIEDZI NA WSKAZNIKI WYMIANY GAZOWEJ, INDEKS ZIELONOŒCI LIŒCI (SPAD) ORAZ PLONOWANIE YCICY TRWA EJ (LOLIUM PERENNE L). I KUPKÓWKI POSPOLITEJ (DACTYLIS GLOMERATA L.) Astrkt MiŸ jst mikrolmntm nizênym o prwi³owgo wzrostu i rozwoju roœlin. Wp³yw n zwrtoœæ hloroilu, pros otosyntzy orz gosporkê won¹ roœlin. Clm pry y³ on wp³ywu niooru mizi w gli n intnsywnoœæ otosyntzy i trnspirji, wspó³zynnik wykorzystni woy, inks zilonoœi liœi orz plonowni wyrnyh omin yiy trw³j (Lolium prnn L.) i kupkówki pospolitj (Dtylis glomrt L.). W okrsi wgtji mirzono intnsywnoœæ otosyntzy i trnspirji z pomo¹ prznoœngo nliztor gzowgo LI-COR 6400 i inks zilonoœi liœi z pomo¹ optyzngo hloroilomtru Minolt SPAD-502. N postwi hwilowyh wrtoœi otosyntzy i trnspirji wylizono wspó³zynnik wykorzystni woy (WUE). Plon suhj msy okrœlono przz wysuszni zilonj msy w tmprturz 105 o C o st³j wgi. Wykzno, nioór mizi istotni ogrnizy³ intnsywnoœæ otosyntzy, trnspirji, poziom hloroilu w liœih orz plonowni wszystkih nyh omin. Omin¹ njrzij oporn¹ n nioór mizi y³ Mj, któr oznz³ siê wysok¹ intnsywnoœi¹ otosyntzy i trnspirji orz njwiêksz¹ zwrtoœi¹ hloroilu. W wrunkh niooru mizi plonow³ on n poziomi pozost³yh omin uprwinyh w wrunkh kontrolnyh. S³ow kluzow: nioór mizi, yi trw³, kupkówk pospolit, otosyntz, trnspirj, WUE, inks SPAD, plonowni. INTRODUCTION Coppr is ssntil or th growth n vlopmnt o plnts. This mironutrint inluns hlorophyll ontnt, prtiipts in th prosss o protin n rohyrt synthsis, n positivly ts th tivity o nitrt ruts (FALKOWSKI t l. 1990, KUDUK 1996, TERELAK t l. 1998, KABATA-PENDIAS, PENDIAS 1999, BARCZAK t l. 2006). Morovr, oppr hs n impt on th photosynthsis pross n wtr rltions in th plnt. Coppr iiny rss th rt o photosynthsis n inrss th rt o rspirtion (RUSZKOWSKA, WOJCIESKA-WYSKUPAJTYS 1996), n it inus hngs in th strutur o l pirmis n stm tissu. Unr oppr iit onitions, stomt, prnhyml n slrnhyml lls o th stm om smllr, whil xylm lls unrgo ormtions. Ths unsirl hngs limit wtr low in th soil-plnt systm, ling to plnt wilting (DYKI, BORKOWSKI 2000). Th oppr ontnt o plnts vris wily pning on th prt o plnt, growth stg, spis n vrity, s wll s on oppr onntrtions in soil n limt onitions (KABATA-PENDIAS 1996, RUSZKOWSKA t l. 1996). Th ojtiv o this stuy ws to trmin th t o oppr iiny in soil on th rt o photosynthsis n trnspirtion, wtr us iiny, l grnnss n th yil o slt ultivrs o prnnil rygrss n orhr grss.
599 MATERIALS AND METHODS Th xprimnt ws onut in grnhous o th Univrsity o Wrmi n Mzury in Olsztyn. Kik-Brukmnn pots wr ill with 10 kg o soil vlop rom loos sn, hrtriz y low ontnt o phosphorus (31 mg kg -1 ), potssium (42 mg kg -1 ), mgnsium (13 mg kg -1 ) n oppr (4.5 mg kg -1 ). Soil rtion ws slightly ii - ph 5.8 m -3 in 1 mol KCl. Th xprimnt ws prorm in our rplitions. In h pot, 2 to 3 ss o grss wr sown t 10 points, n th plnts wr thinn immitly tr mrgn, lving 7 plnts pr pot. Th xprimnt involv two ultivrs o prnnil rygrss (Lolium prnn L.): ttrploi Mj n iploi Argon, n two ultivrs o orhr grss (Dtylis glomrt L.): ttrploi Dl n iploi Ar. Control pots wr rtiliz with nutrint solution ontining 1.00 g N s CO(NH 2 ) 2, 0.25 g P s KH 2 PO 4, 1.00 g K s K 2 SO 4 n 0.50 g Mg s MgSO 4 7H 2 O. A mironutrint solution (30 mg pot -1 ), ompos o 2.65 mg F in EDTA, 0.09 mg MnCl 2 4H 2 O, 0.1 mg ZnCl 2, 0.03 mg CuCl 2 2H 2 O, 0.12 mg H 3 BO 3 n 0.01 mg (NH 4 ) 6 Mo 7 O 24 4H 2 O pr kg o soil, ws lso ppli. Th rmining pots wr not rtiliz with oppr. P, K n Mg wr ppli pr-sowing t singl rt, whil nitrogn n mirolmnts wr ppli t thr rts: pr-sowing, tr th irst n son utting. During th growing sson l grnnss ws stimt with Minolt SPAD-502 hlorophyll mtr, whil th rt o photosynthsis n trnspirtion ws msur using LI- COR 6400 gs nlyzr (Portl Photosynthsis Systm), t ir tmprtur o roun 25 o C, onstnt CO 2 onntrtion o 400 ppm, illumintion o 1000 µmol m -2 s -1 n soil moistur ontnt o roun 70% o il wtr pity. Msurmnts wr tkn on th youngst, ully vlop l o shoots slt rnomly in h trtmnt. Four msurmnts wr prorm or h rgrowth, n routs wr tkn t on-wk intrvls. Mn vlus or rgrowths r prsnt in th ppr. Wtr us iiny (WUE) ws lult s on instntnous vlus o photosynthsis n trnspirtion. Th ovgroun prts o plnts wr ut thr tims ovr th growing sson: ut I 9 July, ut II 30 August, ut III 25 Otor. Dry mttr yil ws trmin y grn mttr rying to onstnt wight t 105 o C. Th rsults wr pross sttistilly with th us o Sttisti sotwr. Th signiin o irns ws vrii y Tuky`s tst t p = 0.99.
600 RESULTS AND DISCUSSION Coppr iiny isturs th onvrsion o light nrgy into hmil nrgy, thus ruing th rt o photosynthsis n photosynthti tivity in plnts. Th vrg rt o photosynthsis ws low in th lvs o ll invstigt grss ultivrs, n it rng rom 5.59 to 8.88 µmol CO 2 m -2 s -1 (Tl 1). Cv. Mj ws mrk y th highst rt o photosynthsis. Th ours o this pross ws wily irntit in prtiulr rgrowths. Th lowst rt o photosynthsis ws osrv in th irst rgrowth, whil th highst in th son rgrowth. Coppr iiny in soil signiintly limit photosynthsis intnsity in th lvs o ll tst ultivrs. Compr to ontrol trtmnts, th rt o photosynthsis rs y 25%. Ttrploi ultivrs wr mor rsistnt to oppr iiny thn iploi ultivrs. Among th tst ultivrs, v. Dl ws oun to most rsistnt to oppr iiny. Th rt o photosynthsis rs to th slightst gr in this ultivr, i.. y 15% on vrg, ompr to 22-36% in th othr ultivrs. Orhr grss is known or its ility to umult oppr. FALKOWSKI t l. (1990) rport tht th onntrtion o this mironutrint ws sustntilly highr in orhr grss thn in prnnil rygrss unr oppr iit onitions. This proly xplins th wkr rspons o orhr grss to oppr iiny, in omprison with prnnil rygrss. In th prsnt stuy oppr iiny rs trnspirtion intnsity, on vrg y 31% (Tl 2). Th invstigt ultivrs ir in thir rsponss to oppr iit. Th wkst rspons ws ror in v. Mj, whil th strongst in v. Argon th rt o trnspirtion rs in ths ultivrs y pproximtly 16% n 42% rsptivly. DYKA n BORKOWSKI (2000) monstrt tht in tomto plnts oppr iiny us rution in th thiknss o th xylm lyr n ormtion o xylm lls s wll s th losing o stomt, thus isturing wtr trnsport n limiting trnspirtion. Prnnil rygrss ultivrs vport mor wtr thn orhr grss ultivrs. Prtiulr rgrowths ir with rspt to th rt o trnspirtion, whih ws th stst in th irst rgrowth n th slowst in th thir rgrowth. Wtr rltions in th plnt n trmin s on wtr us iiny (WUE). Th highst wtr us iiny ws osrv in th thir rgrowth, whih rsult rom low rt o trnspirtion (Tl 3). In orhr grss ultivrs, highr vlus o WUE wr ror in trtmnts not rtiliz with oppr, whrs in prnnil grss ultivrs in ontrol trtmnts. Among th tst ultivrs, th lowst WUE (mn vlu or rgrowths) ws rport or v. Argon, whil irns twn th rmining ultivrs wr sttistilly non-signiint. Coppr prtiipts in hlorophyll iosynthsis (GRZYŒ 2004) roun 70% o oppr n oun in hloroplsts. Coppr iiny in soil h signii-
601 Ar Dl Argon Mj Ar Dl Argon Mj Cultivrs Control ojt Coppr iiny Intnsity o photosynthsis (µmol CO2 m- 2 s - 1 ) ontrol ojt iiny ontrol ojt iiny ontrol ojt iiny ontrol ojt iiny Frtiliztion oppr oppr oppr oppr ut 5.98* 3.36 7.16 6.22 9.46 4.36 6.30 5.13 Mn or ultivrs 4.67 6.69 6.91 5.71 Mn or rtiliztion 7.22 4.77 2 n ut 9.17 8.29 8.39 6.98 10.25 6.82 10.43 8.96 8.73 7.68 8.54 9.69 9.56 7.76 3 r ut 7.38 5.12 5.92 5.02 6.93 5.87 9.01 5.91 6.25 5.47 6.40 7.46 7.31 5.48 Tl 1 Mn 7.51 5.59 7.16 6.07 8.88 5.69 8.58 6.66 6.55 6.61 7.28 7.62 8.03 6.00 Mn or uts 5.60 8.66 6.39 7.02 *homognous sttistil groups (vlus mrk with sm lttr i not ir sttistilly) Ar Dl Argon Mj Ar Dl Argon Mj Cultivrs Control ojt Coppr iiny Intnsity o trnspirtion (m mol H2O m -2 s - 1 ) ontrol ojt iiny ontrol ojt iiny ontrol ojt iiny ontrol ojt iiny Frtiliztion oppr oppr oppr oppr ut 0.90 0.64 1.04 0.92 2.03 0.87 1.21 1.10 Mn or ultivrs 0.77 0.98 1.45 1.15 Mn or rtiliztion 1.29 0.88 2 n ut 0.92 0.52 0.77 0.44 1.19 0.84 1.31 g 0.84 0.72 0.60 1.01 1.08 1.04 0.66 3 r ut 0.52 0.35 0.53 0.36 0.36 0.37 0.38 0.48 0.43 0.44 0.36 0.43 0.45 0.39 Tl 2 Mn 0.78 0.50 0.78 0.57 1.19 0.69 0.96 0.81 0.64 0.68 0.94 0.88 0.93 0.64 Mn or uts 1.09 0.85 0.42 0.79 Explntions, s Tl 1
602 Ar Dl Argon Mj Ar Dl Argon Mj Cultivrs Control ojt Coppr iiny Wtr us iiny (µmol CO2 m- 2 s - 1 m mol H2O m -2 s - 1 ) ontrol ojt iiny ontrol ojt iiny ontrol ojt iiny ontrol ojt iiny Frtiliztion oppr oppr oppr oppr ut 6.70 5.28 6.89 6.74 4.66 5.02 5.23 4.67 Mn or ultivrs 5.99 6.81 4.83 4.95 Mn or rtiliztion 5.87 5.43 2 n ut 10.02 16.03 10.93 15.95 8.63 8.17 7.97 10.63 13.02 13.44 8.40 9.30 9.39 13.70 3 r ut 14.26 14.84 11.28 13.96 19.11 16.09 23.89 12.38 14.55 12.62 17.60 18.14 17.14 14.32 Tl 3 Mn 10.33 12.05 9.70 12.23 10.80 9.80 12.36 9.23 11.19 10.96 10.28 10.80 10.80 10.81 Mn or uts 5.65 11.04 15.73 10.81 Explntions, s Tl 1 nt t on hlorophyll onntrtions in lvs. L grnnss (SPAD) vlus rs y pproximtly 7-8% in ll rgrowths unr oppr iit onitions (Tl 4). Cv. Mj show th strongst rspons to oppr iiny, whih ws mnist y n 11% rs in th hlorophyll ontnt o lvs. At th sm tim, this ultivr h th highst hlorophyll onntrtion (on vrg 42.73 SPAD). Th lowst hlorophyll ontnt o lvs ws ror in th irst rgrowth, rgrlss o ultivr. This is onsistnt with our rlir inings (OLSZEWSKA 2006, OLSZEWSKA t l. 2008). Prnnil rygrss v. Mj ws hrtriz y th highst yil o ll tst ultivrs, oth unr optimum rtiliztion onitions n unr oppr iit onitions (Tl 5). In this ultivr high yil orrspon with high rt o photosynthsis n trnspirtion, n with th highst hlorophyll ontnt o lvs. Diit in soil Cu us yil lin in ll th ultivrs, ut sttistilly signiint irns wr osrv only with rgr to v. Ar. Dry mttr yil ropp on vrg y roun 8%, n th highst lin ws not in v. Dl (pprox. 11%). In ll th nlyz ultivrs th strongst rspons to oppr iiny ws rport in th son rgrowth, whih ws rlt y th highst yil rs.
603 Ar Dl Argon Mj Ar Dl Argon Mj Cultivrs Control ojt Coppr iiny ontrol ojt iiny ontrol ojt iiny ontrol ojt iiny ontrol ojt iiny L grnnss inx Frtiliztion oppr oppr oppr oppr (SPAD) ut 36.13 33.48 35.43 34.40 39.95 36.88 41.98 37.85 Mn or ultivrs 34.80 34.91 34.41 39.91 Mn or rtiliztion 38.37 35.63 2 n ut 40.85 38.90 41.05 37.25 43.18 41.43 48.10 42.2 39.88 39.95 42.30 45.15 43.29 39.94 3 r ut 42.03 40.45 42.03 38.85 41.58 39.65 45.90 40.38 41.23 39.64 40.61 43.14 42.88 39.83 Tl 4 Mn 39.67 37.61 39.50 36.83 41.57 39.32 45.33 40.14 38.64 38.17 40.44 42.73 41.51 38.48 Mn or uts 37.01 41.82 41.16 39.99 Explntions, s Tl 1 Ar Dl Argon Mj Ar Dl Argon Mj Cultivrs Control ojt Coppr iiny ontrol ojt iiny ontrol ojt iiny ontrol ojt iiny ontrol ojt iiny D ry mttr yil (g pot - 1 ) Frtiliztion oppr oppr oppr oppr ut 7.38 7.23 8.15 6.98 7.50 7.68 8.73 7.93 Mn or ultivrs 7.30 7.56 7.59 8.33 Mn or rtiliztion 7.93 7.45 2 n ut 7.63 6.50 7.43 6.40 7.13 6.35 7.05 6.65 7.06 6.91 6.74 6.85 7.31 6.48 3 r ut 5.45 5.03 5.67 5.62 6.45 5.98 6.70 6.35 5.23 5.65 6.21 6.53 6.07 5.74 Tl 5 Mn 20.45 18.75 21.25 19.00 21.07 20.00 22.48 20.93 19.60 20.13 20.54 21.70 21.31 19.67 Mn or uts 7.69 6.89 5.91 20.49 Explntions, s Tl 1
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