UTICAJ Ni NA KONCENTRACIJU I RASPODELU NEOPHODNIH ELEMENATA U BILJKAMA GRAŠKA

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1 Letopis nau nih radova Godina 29 (2005), broj 1, strana UDK: : Originalni nau ni rad Original scientific paper UTICAJ Ni NA KONCENTRACIJU I RASPODELU NEOPHODNIH ELEMENATA U BILJKAMA GRAŠKA Petrovi, N 1., Maksimovi Ivana 1, Kevrešan, Ž. 2 REZIME Putem mikro ogleda ispitan je uticaj 5 doza Ni: 0 (kontrola), 20, 40, 80 i 120 g Ni/ha na hemijski sastav biljaka graška, sorte Moravac. Nikal je pre setve inkorporiran u zemljište. Biljke su gajene do faze tehnološke zrelosti. Utvr en je posebno hemijski sastav korena, stabla, listova, mahuna i zrna. Uo eno je da su primenjene doze Ni zna ajno uticale na nakupljanje i distribuciju ispitivanih neophodnih elemenata. Smanjenje i/ili pove anje udela ili transporta mineralnih elemenata iz korena u nadzemne delove je bilo specifi no, i zna ajno je zavisilo od koncentracije pristupa nog Ni u zemljištu i konkretnog neophodnog elementa. Klju ne re i: grašak, organ biljke, doze nikla, koncentracija, distribucija, neophodni elementi UVOD Neophodnost Ni za bakterije, biljke, životinje i oveka uo eno je od strane brojnih autora (Eskew et al., 1983; Klucas et al., 1983; Solomons, 1984; Brown et al., 1987). Zna aj Ni kod viših biljaka naro ito se isti e pri njihovom gajenju u prisustvu uree (Gerendas et al., 1999), i/ili kod leguminoznih vrsta (Walker et al., 1985). Biljke kod kojih je azot primenjen u obliku uree, na primer, kod soje, duvana i paradajza zahtevaju prisustvo Ni u hranljivom supstratu, pošto je Ni neophodna komponenta enzima ureaze (Polacco et al., 1977a; Polacco, 1977b; Shimada i Ando, 1980). U slu aju nedostatka Ni u hranljivom supstratu pove ava se koncentracija uree do nivoa koji esto remeti biljni metabolizam, a ne retko dolazi do nakupljanja uree u tkivima, u toksi nim koncentracijama (Eskew et al., 1983; Eskew et al., 1984). Na neophodnost Ni, posebno kod leguminoza ukazuje injenica da se azot kod ovih biljaka u najve oj meri transportuje u obliku ureida (alantion i alantionska kiselina 1 Dr Novica Petrovi, red. prof., dr Ivana Maksimovi, van. prof., Poljoprivredni fakultet i Nau ni institut za ratarstvo i povrtarstvo, Novi Sad 2 Mr Žarko Kevrešan, asist., Tehnološki fakultet, Novi Sad 31

2 (Walker et al., 1985). Istovremeno treba imati u vidu i uticaj Ni na obrazovanje kvržica i aktivnost enzima nitrogenaze (Cammack, 1995). Brojna ispitivanja, tako e, ukazuju da Ni može da ima vidnu ulogu u sintezi fitoaleksina, ime uti e na otpornost biljaka prema bolestima (Graham i Webb, 1991). U uslovima nedostatka Ni kod nekih vrsta smanjuje se aktivnost malat-dehidrogenaze, seme brže sazreva i biljke prinudno stare (Brown et al., 1987; 1990). Niže koncentracije Ni kod nekih biljaka, na primer, kod žita, podsti u proces klijanja semena (Dalton et al., 1985; Brown et al., 1990; Krogmeier et al., 1991). Raspon kocentracija Ni nedostatka i suviška je kod ve ine vrsta relativno širok, za razliku od drugih mikroelemenata, na primer Cu (Thiel i Finck, 1973; Hodenberg i Finck, 1975). Kod životinja i oveka nedostatak Ni uti e na strukturu i funkciju membrana (Welch, 1981), menja metabolizam Fe (Nielson, 1984), te aktivnost enzima ciklusa Krebsa (Stangl i Kirchegssnesr, 1996). Suvišak Ni je sli no drugim teškim metalima, toksi an za biljke (Bingham et al., 1986), životinje i oveka (Hammondi Foulkes, 1986; Nieboer i Nriagu, 1992). Kod biljaka Ni u suvišku remeti procese fotosinteze i disanja (Yang et al., 1996), mineralnu ishranu (Barceló i Poscenreider, 1990), zatim funkciju membrana (Pandolfini et al., 1992), usporava usvajanje vode (Pandolfini et al., 1992), sintezu hlorofila (Braune i Dietz, 1995), a time i rastenje i razvi e biljaka (Bingham et al., 1986). Imaju i navedeno u vidu smatralo se da je od interesa da se ispita uticaj nižih koncentracija Ni na nakupljanje i raspodelu neophodnih elemenata kod graška, sorte Moravac. MATERIJAL I METOD RADA Ispitivanja su obavljena na oglednom polju Nau nog instituta za ratarstvo i povrtarstvo u Novom Sadu, na Rimskim Šan evima, u toku i godine. Ogledi su postavljeni u 4 ponavljanja, sa veli inom osnovne parcele 25 m 2. Ispitano je 5 doza nikla 0, 20, 40, 80 i 120 g Ni/ha. Nikal je dodat u obliku NiSO 4 x 6H 2 O putem inkorporacije u zemljište pre setve graška, sorte Moravac. Tokom rastenja i razvi a biljaka primenjivane su uobi ajene agrotehni ke mere koje se koriste za ovu vrstu biljaka. U fazi tehnološke zrelosti sa svake parcele uzeto je po 20 biljaka za analizu. Biljke su nakon toga deljene na: koren, stablo, listove, mahune i zrno. Koren je pran prvo vodovodnom vodom, a zatim ispran destilovanom vodom. Uzorci su osušeni u sušnici, na temperaturi od 70 0 C, do konstantne mase, a potom je odre enja masa suve materije. Ukupan azot odre en je metodom Kleldahl-a, sadržaj nitrata spektrofotometrijski koriš enjem fenol-disulfonske kiseline. Koncentracija fosfora odre ena je amoniummolibdat-vanadat metodom, a udeo kalijuma plamenfotometrijski. Koncentracija kalcijuma, magnezijuma, gvož a, mangana, cinka, bakra i nikla utvr ena je standardnom atomskom apsorcionom spektrofotometrijom. Dobijeni rezultati statisti ki su obra eni analizom varijanse, i izra unavanjem najmanje zna ajnih razlika izme u aritmeti kih sredina (LSD) za verovatno u od 5%. 32

3 REZULTATI I DISKUSIJA Nikal prisutan u hranljivom supstratu u pristupa nom obliku za biljke, može da inhibira ili podsti e usvajanje i nakupljanje neophodnih mineralnih elementa (Sela et al., 1988). Uticaj se naj eš e zasniva na antagonizmu ili sinergizmu izme u jona Ni i jona drugih elemenata. Smanjenje i/ili pove anje intenziteta usvajanja i transporta neophodnih mineralnih elemenata, iz korena u nadzemne delove, zavisi od brojnih spoljašnjih i unutrašnjih inilaca. Od egzogenih faktora posebno se isti e obezbe enost hranljivog supstrata neophodnim mineralnim elementima, temperatura i vlažnost zemljišta, a od endogenih biljnih vrsta, genotip i dr. (Yang et al., 1996). Ispitivane doze Ni kod biljaka graška, sorte Moravac su zna ajno uticale na koncentraciju i distribuciju NO 3 iako nisu zna ajno uticale na kocentraciju i raspodelu ukupnog N (Sl. 1). Prou avane doze Ni razli ito su uticale na koncentraciju NO 3 u ispitivanim organima. Tako na primer, pri dozi Ni od 40, ili 80 g/ha uo eno je intenzivnije nakupljanje NO 3 u korenu, pri nivou od 20, ili 40 g/ha u listovima, dok je pri najve oj ispitivanoj dozi Ni (120 g Ni/ha) utvr eno zna ajno smanjenje sadržaja NO 3 u reproduktivnim organima. Istovremeno je uo eno da su doze Ni specifi no uticale na distribuciju NO 3 u ispitivanim organima, a time i na njegov metabolizam. Ve i broj autora je ukazao na ulogu Ni u metabolizmu N kod biljaka (Brown et al., 1990; Krogmeier et al., 1991). Kod biljaka paradajza je utvr eno da akumulacija i distribucija N zna ajno zavisi od obezbe enosti biljaka Ni. Sa pove anjem koncentracije Ni u hranljivom supstratu pove ao se udeo N u korenu i stablu, a smanjio u listovima (Palacios et al., 1998). Sli ni rezultati su dobijeni kod crvene deteline (Abdel-Sabour, 1991), dok je kod je ma uo eno zna ajno smanjenje sadržaja NO 3 u prisustvu Ni (Brown et al., 1990). Primenjene niže doze Ni, 20 ili 40 g Ni/ha, podstakle su nakupljanje P u korenu, a najve a doza (120 g Ni/ha ) je smanjila koncentraciju P u listu i mahuni (Sl. 1). Efekat Ni na nakupljanje i raspodelu P ne zavisi samo od koncentracija Ni u hranljivom supstratu, ve i od gajene biljne vrste. Tako na primer, Ni kod paradajza umanjuje, a kod engleskog ljulja podsti e usvajanje i transport P iz korena u nadzemne delove, dok kod kupusa, kukuruza i bele deteline primenjene koncentracije Ni nisu zna ajno uticale na sadržaj P (Yang et al., 1996; Palacios et al., 1998). Ispitivane doze Ni nisu zna ajno uticale na koncentraciju K u korenu, stablu i listovima, dok se udeo K u reproduktivnim organima biljaka graška zna ajno smanjio posebno pod uticajem najve e ispitivane koncentracije Ni (Sl. 1). Doze Ni zna ajno su uticale na distribuciju K. Naime, pri koncentraciji Ni od 40 ili 80 g/ha uo eno je intenzivnije nakupljanje K u listovima, a smanjen sadržaj u reproduktivnim organima. Yang et al. (1996) isti u povoljan uticaj nižih koncentracija Ni na absorpciju K u korenu kupusa, engleskog ljulja, bele deteline, kukuruza, pšenice (Pandolfini et al., 1996). Palacios et al. (1998) isti u da je Ni kod paradajza pove ao absorpciju i udeo K u korenu, a smanjio u stablu i granama. Sli ne vrednosti su dobijene kod genotipa pšenice tolerantnog prema suši (Pandolfini et al., 1996). Ve i broj autora ukazuje da Ni na koncentraciju K u korenu biljaka prvenstveno uti e preko promene permeabilnosti plazma membrane (De Vos et al., 1989). Ve i broj autora, kod razli itih biljnih vrsta, je uo io da Ni, može zna ajno da uti e na dinamiku sadržaja i metabolizam neophodnih elemenata, posebno Ca (Gyôri et al., 33

4 Sl. 1. Uticaj Ni na koncentraciju i distribuciju N, NO 3, P, K, Ca i Mg u razli itim tkivima graška ( koren, stablo, list, mahuna, zrno), izraženo u % u suvoj materiji. Vertikalne linije predstavljaju LSD za α=0.05. Fig. 1. Effect of Ni on concentration and distribution of N, NO 3, P, K, Ca and Mg in different pea tissues ( root, stem, leaf, pod, seed), expressed in % in dry matter. Bars represent LSD for α=

5 1996). Tako Palacios et al. (1998) su utvrdili da Ni kod paradajza zna ajno smanjuje nakupljanje Ca u nadzemnom delu, a posebno u korenu. Dobijeni rezultati u ovim ispitivanjima ukazuju da niže koncentracije Ni u hranljivom supstratu nisu zna ajno uticale na koncentraciju i distribuciju Ca kod graška (Sl. 1). S tim u vezi ve i broj autora ukazuju da dobijeni rezultati esto mogu da budu protivre ni i da dobijene vrednosti osim od doze Ni i biljne vrste, zavise od brojnih drugih inilaca, na primer, genotipa, koncentracije Ni u hranljivom supstratu, vrste ispitivanog elementa (Yang et al., 1996). Ispitivane doze Ni zna ajno su uticale na nakupljanje i distribuciju Mg kod graška (Sl. 1). Dobijeni rezultati ukazuju da izme u ispitivanih doza Ni u hranljivom supstratu i koncentracije Mg u korenu postoji pozitivna, a u listu negativna korelacija. Koncentracija Mg u stablu, mahunama, a posebno u zrnu nije zna ajno zavisila od primenjenih koncentracija Ni. Me utim, Palacios et al. (1998) su utvrdili zna ajno smanjenje koncentracije Mg kako u korenu tako i u nadzemnim delovima paradajza tretiranog niklom, a Chamel i Heuman (1987) i Tang i Miller (1991) isti u da Ni može kompetativno da inhibira usvajanje Mg. U prisustvu viška Ni u hranljivom supstratu kod ovsa, je ma i paradajza uo eno je smanjenje koncentracije ukupnog Fe. Sli ni rezultati su utvr eni i kod kupusa (Yang et al., 1996). S tim u vezi ve i broj autora ukazuje da Ni u suvišku kod brojnih vrsta može da izazove nedostatak Fe (Bingham et al., 1986). Prepostavlja se da Ni u suvišku indukuje hlorozu prvenstveno inhibicijom translokacije Fe iz korena u nadzemne delove (Aller et al., 1990). Palacios et al. (1998) su utvrdili da kod paradajza sa pove anjem koncentracije Ni u hranljivom supstratu, udeo Fe zna ajno opada. Sli ne vrednosti su dobijene i kod kupusa (Yang et al., 1996). Niže doze Ni povoljno uti u na nakupljanje Fe u grašku (Sl. 2). Doze od 20 do 80 g Ni/ha izazvale su pove anje koncentracije ukupnog Fe u korenu, stablu, listovima i mahunama. Osim u zrnu, doza od 120 g Ni/ha nije zna ajnije uticala na koncentraciju Fe. Ispitivane doze Ni su me utim, uticale na distribuciju Fe. Na primer, pri primeni 20 do 80 g Ni/ha uo eno je zna ajno smanjenje koncentracije Fe u listovima. Na osnovu dobijenih rezultata može se zaklju iti da je nakupljanje Fe u reproduktivnim organima bilo najmanje pri najve oj ispitivanoj dozi Ni. Nikal, posebno u suvišku, uti e na koncentraciju i distribuciju Mn iz korena u nadzemne delove brojnih vrsta biljaka (Palacios et al., 1998; Brune i Dietz, 1995; Yang et al., 1996). Stoga ve i broj autora smatra, da za odre ene uslove gajenja biljaka, koncentracija Mn u biljkama može da se koristi kao pouzdan pokazatelj suviška Ni u hranljivom supstratu (Reuter i Robinson, 1986; Jones et al., 1991). Gajenjem graška pri nižim dozama Ni u hranljivom supstratu uo en je njegov suprotan efekat na koncentraciju i distribuciju Mn (Sl. 2). Naime, u ovim ispitivanjima je potvr en povoljan efekat doze Ni od 20 do 80 g Ni/ha na udeo Mn u korenu, stablu i listu. Istovremeno ispitivanja ukazuje da su prou avane doze Ni u najve oj meri podstakle nakupljanje Mn u korenu, a da nisu zna ajno uticale na njegovu distribuciju iz korena u nadzemne delove biljaka, posebno u reproduktivne organe. U brojnih vrsta absorpcija, nakupljanje i distribucija Cu, a posebno Zn zna ajno zavisi od pristupa nih koncentracija Ni za biljke. Tako e, potrebno je da se usvajanje Ni u najve oj meri odvija istim putevima kao i usvajanje Zn i Cu (Mortvedt et al., 1991; Dang et al., 1990). Kompeticija pri usvajanju izme u pomenutih jona je uo ena kod 35

6 razli itih vrsta (Bagatto et al., 1993; Yang et al., 1996), a antagonizam pri usvajanju Zn i Ni, i/ili Cu i Ni je uo en u brojnim ispitivanjima, naro ito pri prou avanju ve ih koncentracija Ni. Sl. 2. Uticaj Ni na koncentraciju i distribuciju Fe, Mn, Zn i Cu u razli itim tkivima graška ( koren, stablo, list, mahuna, zrno), izraženo u mg kg -1 suve materije. Vertikalne linije predstavljaju LSD za α=0.05. Fig. 2. Effect of Ni on concentration and distribution of Fe, Mn, Zn and Cu in different pea tissues ( root, stem, leaf, pod, seed), expressed in mg kg -1 of dry matter. Bars represent LSD for α=

7 Koncentracija i distribucija Zn kod graška tako e je zavisila od primenjenih doza Ni u hranljivom supstratu (Sl. 2). Prou avane doze Ni pove ale su koncentraciju Zn u korenu i stablu, dok nisu zna ajnije uticale na koncentraciju Zn u listu, mahunama ili zrnu. Ispitivane doze Ni, posebno 40 i 80 g/ha zna ajno su uticale na raspodelu Zn u ispitivanim organima, pove avši njegov sadržaj u stablu, smanjuju i ga u mahunama i zrnu. Ispitivane doze Ni, sli no njihovom uticaju na status Zn, uticale su i na nakupljanje, a posebno na distribuciju Cu (Sl. 2). Naime, primenjene doze Ni u korenu i stablu su pove ale, a u mahunama i zrnu smanjile koncentraciju Cu. Sli no koncentraciji, ispitivane doze Ni uticale su i na raspodelu Cu, smanjuju i transport Cu iz korena u nadzemne delove, posebno u reproduktivne organe. Kod pasulja Ni pri koncentraciji od 1,0 do 2,0 µg/l pove ao je udeo Fe, dok nije zna ajno uticao na akumulaciju Ca, Mg, Mn i Zn (Piccini i Malavolta, 1992). Me utim kod je ma ispitivane koncentracije Ni nisu zna ajno uticale na translokaciju Fe iz korena u nadzemne delove, dok su znatno umanjile premeštanje Cu, Zn, Mn i Co (Agarwala et al., 1977). Kona no treba imati u vidu da suvišak Fe, Cu, Co i/ili Zn u hranljivom supstratu u brojnih vrsta može da inhibira absorpciju i nakupljanje Ni, na primer kod soje (Cataldo et al., 1978). ZAKLJU AK Ispitivane doze Ni nisu zna ajno uticale na koncentraciju i distribuciju ukupnog N. Udeo i raspodela NO 3 zavisila je od koli ine Ni u hranljivom supstratu. Nakupljanje P u ispitivanim organima graška zna ajno je zavisilo od koli ine Ni u podlozi. Pri nižim dozama Ni udeo P pove ao se u korenu, a pri najve oj zna ajno smanjio u listu i mahuni. Ispitivane doze Ni nisu zna ajno uticale na koncentraciju K i Ca u korenu, stablu i listovima, dok su udeo K u reproduktivnim organima zna ajno smanjile, posebno najve a primenjena doza Ni. Dobijeni rezultati ukazuju da izme u primenjenih doza Ni i koncentracije Mg u korenu postoji pozitivna, a u listu negativna korelacija. Koncentracija Mg u stablu, mahunama i zrnu nije zna ajno zavisila od doze Ni. Niže doze Ni povoljno su uticale na nakupljanje i raspodelu Fe. Najmanje nakupljanje Fe u reproduktivnim organima utvr eno je pri najve em sadržaju Ni u supstratu. Nikal je u najve oj meri podstakao nakupljanje Mn u korenu, a nije zna ajno uticao na njegovu translokaciju iz korena u nadzemne delove biljaka. Primenjene doze Ni pove ale su koncentraciju Zn u korenu i stablu, dok nisu zna ajno uticale na njegov udeo u listu, mahunama i zrnu. U korenu i stablu Ni je pove ao, a u mahunama i zrnu smanjio koncentraciju Cu. Prisustvo Ni u hranljivom supstratu umanjilo je transport Cu iz korena u nadzemne delove, posebno u reproduktivne organe. 37

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9 23. Jones, Jr., J.B., Wolf, B., Mills, H. (1991): Plant Analysis Handbook. MicroMacro Publishing, Inc., Athens, GA. 24. Klucas, R.V., Hanus, F.J., Russell, S.A. and Evans, H.J. (1983): Nickel A micronutrient elements for hydrogen-dependent growth of Rhizobium japonicum and for expression of urease activity in soybean leaves. Proc. Natl. Acad. Sci. USA 80, Krogmeier, M.J., McCarry, G.W., Shogren, D.R. and Bremner, J.M. (1991): Effect of nickel deficiency in soybean on the phytotoxicity of foliar-applied urea. Plant and Soil. 135, Mortvedt, J.J., Cox, F.R., Shuman, L.M., Welch, R.M. (1991): Micronutrients in Agriculture. 2nd ed. Soil Science Society of America Book Series, Madison, WI. 27. Nieboer, E., Nriagu, J.O. (1992): Nickel and Human Healt: Current Perspectives. John Wiley and Sons, New York, NY. 28. Nielson, F.H., (1984): Ultratrace elements in nutrition. Annu. Rev. Nutr., 4, Palacios, G., Gómez, I. Carbonell-Barrachina, J. Navarro Pedreno, Mataix, J. (1998): Effect of Nickel Concentration on Tomato Plant Nutrition and Dry Matter Yield. Joutnal of Plant Nutrition, 21, (10), Pandolfini, T., Gabbrielli, R., Compavini, C. (1992): Nickel toxicity and peroxidase activity in seedlings in Triticum aestivum L. Plant Cell Environ. 15, Pandolfini, T., Gabbrielli, R.,Ciscato, M. (1996): Nickel toxicity in two durum wheat cultivars differing in drought sensitivity. J. Plant Nutr. 19, Piccini, D.F., Malavolta, E. (1992): Effect of nickel on two common bean cultivars. J. Plant Nutr., 15, Polacco, J.C. (1977a): Nitrogen metabols in soybean tissue culture: II Urea utilization and urease synthesis require Ni 2+. Plant Physiol. 59, Polacco, J.C. (1997b): Is nickel a universal component of plant urease? Plant Sci. Lett. 10, Reuter, D.J., Robison, J.B. (1986): Plant Analysis. An Interpretation Manual. Inkata Press, Melbourne. Australia. 36. Sela, M., Tel-Or, E., Fritz, E., Huterman, A. (1988): Localization and toxic effects of cadmium, copper, nickel and uranium in Azzolla. Plant Physiol. 88, Shimada, N., Ando, T. (1980): Role of Nickel in Plant Nutrition (2). Effect of nickel on the assimilation of urea by plants. Jpn. J. Soil Sci. Plant Nutr. 51, Solomons, N.W. (1984): The other trace minerals: manganese, molybdenum, vanadium, nickel, silicon, and arsenic. Curr. Top. Nutr. Diss., 12, Stangl, G., Kirchgessner, M. (1996): Effect of nickel deficiency on various metabolic parametars of rats. J. Animal Physiol. Animal Nutr. 75, Tang, T., Miler, M.D. (1991): Growth and tissue composition of rice growth in soil treated with inorganic copper, nickel, and arsenic. Commun. Soil Sci. Plant Anal., 22, Thiel, H., Finck, A. (1973): Ermittlung von Grenzwerten optimaler Kupferversorgung für Hafer Sommergerste. Z. Pflanzenernähr. Bodenkd. 134, Walker, C.D., Graham, R.D., Madison, J.T., Cary, E.E., Welch, R.M. (1985): Effects of nickel deficiency on some nitrogen metabolites in cowpeas (Vigna unguiculata L. Walp). Plant Physiol. 79, Welch, R. M. (1981): The biological significance of Ni. J. Plant Nutr. 3, Yang, X., Baligar, V.C., Martens, D.C., Clark, R.B. (1996): Plant tolerance to nickel toxicity: II Nickel effects on influx and transport of mineral nutrients in four plant species. J. Plant Nutr., 19, (2),

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