ࡱ> npmvq`IbjbjqPqP 9::? ***>OOO8OzP4>QVBV"lYlY >>*<>>><>>>   GENETIC ANALYSIS OF PHYSIOLOGICAL EFICIENCY OF NITROGEN IN FOUR WHEAT CULTIVARS AND THEIR HIBRIDS D. KNEZEVIC1, V. KOVA EVI2, A. PAUNOVIC3, M. MADIC3, N. DJUKIC4 Abstract::In diallel crosses (without reciprocals) of four genetically divergent wheat cultivars (Jugoslavija, Osijecanka, Zitnica, and NS Rana 2) the variability and inheritance of physiological nitrogen efficiency (PEN) in wheat plant were investigated. The mode of inheritance, gene effect, combining ability of parent cultivars and F1 and F2 hybrids for physiological nitrogen efficiency (PEN) were studied. The different mode of inheritance: partial dominance, dominance and overdominance for analyzed traits were established. On the base of genetic components of variance was estimated that non-aditive gene action have gene higher impact than additive in the inheritance of PEN. The best general combiner for PEN in both F1 and F2 generation was Zitnica cultivar. The combination Zitnica x NS Rana 2 in both F1 and F2 generation was the best for PEN. Key words: Wheat, nitrogen, efficiency, gene effect, cultivars, hybrids. INTRODUCTION Improvement in wheat yield potential, both before and after the advent of semi-dwarf lines, has been shown to be most strongly associated with improved partitioning of assimilates to grain (Austin et al., 1989). Achieving consistent production at these high levels without causing environmental damage requires improvements in soil quality and precise management of all production factors. The nitrogen (N) plays main role in wheat nutrition because of its importance in protein and nucleic acid synthesis as well plant productivity. The very important for plant productivity is intensities of genotype reaction on nitrogen uptake on environment and its utilization in plants represents nitrogen efficiency utilization (Nielsen and Schjorring, 1983; Kochegarova, 1988; Gamzikova et al., 1991; Knezevic et al., 2007). Although grain protein composition depends primarily on genotype, it is significantly affected by environmental factors and their interactions (Huebner et al., 1997; Tribo et al., 2003). However, the mechanism by which genotype and environmental factors modified the accumulation of the protein fractions is unknown. Degrees of adaptability of plant species and cultivars to suboptimal supplies of mineral element, including N, are different (Agoston and Pepo, 2005; Pepo, 2005; Drezner et al. 2006; Balogh et al. 2006). Modern technology of wheat production mainly based on numerous scientific farming measures as well enormous application of mineral fertilizers. Mainly one third of applied nutrient wheat plants are able use during vegetative period (Clark, 1983; Ivanova and Mateewa, 1983). In the filed practices is very important optimize quantity of fertilizers, decrease expenses of production and improve efficiency of wheat plant of nitrogen absorption, accumulation and reutilization (Saric and Kovacevic, 1981; Klimashevsky and Chernysheva, 1982). Wheat properties are mainly caused by effect of genetic factors in interaction with environment (). The total N content represent indicator of N accumulation in plant (Desai and Bathia, 1978; Djokic et al. 1992) which indicating root system activity and translocation of organic and inorganic matter to top of plant. Physiological nitrogen efficiency (PEN) in plant indicating activity of top of plant and involve of absorbed nitrogen into processes of synthesis. On the base of dynamics nitrogen efficiency absorption and its introduction of organic synthesis are increasing speed of these processes and plant productivity. Genotype specificities of wheat N efficiency are reflect in all phases of plant growth, determined by general and variety specific genes located on three genomes (ABD) in hexaploid wheat (Gamzikova, 1994). By promoting large numbers of progeny in the breeding process there is a chance of identifying phenotypes where favorable interactions among genes permit the expression of higher physiological nitrogen efficiency. The probability of selecting these lines can be enhanced by eliminating inferior agronomic phenotypes and selecting superior physiological phenotypes visually in early generations and by using rapid detection techniques such as IR- thermometry in intermediate generations. Goal of this paper is investigation of variability of physiological nitrogen efficiency (PEN) in wheat genotypes and breeding of wheat with high physiological nitrogen efficiency and its importance for sustainable agriculture. MATERIAL AND METHODS The four divergent winter wheat cultivars (Triticum aestivum ssp. vulgare) and hybrids produced in diallel crosses of Jugoslavija, Osijecanka, Zitnica and NS Rana 2 cultivars were analyzed. By method of half diallel crosses produced hybrid progenies of F1 and F2 generation. Seeds of F1 and F2 hybrids and parents planted in randomized block system in three replications on the experimental field in Center for Small Grains. The seeds planted in 1 m long raws, with distance between raw 0.25 m and with 0.10 m space between each seed in raw. The plants in full maturity stage used for analysis. The nitrogen analyzed by use Kjeldahl methods. The components of genetic variance analyzed by method Jinks (1954) and Mather and Jinks (1971) while the combining abilities analyzed according to Griffing (1956) method 2 mode I. RESULTS AND DISCUSSION The average value of physiological nitrogen efficiency (PEN) in parent cultivars and F1 and F2 hybrids presented in the table 1. Different mode of inheritance for physiological nitrogen efficiency (PEN) was found, namely: overdominance, dominance and partial dominance. Similar results for physiological nitrogen efficiency (PEN) variability reported by Desai and Bahtia, (1978), Djokic et al. (1992). Additive gene effect in both generation of hybrids was lower indicated that dominant gene have higher importance for inheritance of physiological nitrogen efficiency (PEN) table 2. The positive interaction additive x dominant indicated that dominant genes have higher influence than recessive genes in the inheritance of physiological nitrogen efficiency (PEN). The frequency of dominant genes is higher than frequency of recessive gene and distribution of dominant and recessive alleles in F1 and F2 generation are unequal (Tab. 2). The best general combinig ability (GCA) in both generation for physiological nitrogen efficiency (PEN) showed Zitnica cultivar (Tab 3). Progenies obtained in combination in which Zitnica cultivar was one of the parent expressed the highest value for specific combining ability (SCA) (Tab. 3). Table 1. Physiological efficiency of nitrogen (PEN) in wheat genotypes CultivarABCDJugoslavija (A)2525+d22-sd18-sdF1 hybridsOsijecanka (B)25+d 2420-sd21-dLSD 0.05LSD 0.01Zitnica (C)22-sd 23-sd 2621-d4.466.02NS Rana 2 (D)23 i 23 i 20-sd 21F2 hybridsLSD 0.05 = 3.85 LSD 0.01 = 5.09 Table 2. Genetic variance and heritability for Physiological efficiency of nitrogen (PEN) in wheat hybrids VariancePhysiological efficiency of nitrogen (PEN)VariancePhysiological efficiency of nitrogen (PEN)F1F2 F1 F2D 1.892.542 u0.6450.751H143.24328.469 v0.3540.258H237.89818.703EMBED Unknown4.7833.346F4.0879.448 KD/KR1.5813.478E2.381.729*Th2 5.82% H2 / 4H1 0.8760.657*Gh2 7.62%* Th2 = total heritability *Gh2 = genetical heritability Table 3. General and specific combining abilities for Physiological efficiency of nitrogen (PEN) in wheat hybrids Wheat cultivar XLSDABCDSe0.050.01General combining abilityF1PEN0.0540.3260.598-1.0280.7021.3121.730Rank3214F2PEN0.190-0.0541.306-1.0060.8231.5192.031Rank3214Specific combining ability of four wheat cultivarsLSDABCDSe0.050.01APEN-1.2138-2.75440.1867F1 hybridsBPEN-2.24080.0466-4.29510.463.7214..974CPEN-0.98030.4201 1.5872F2 hybridsDPEN-2.52090.13922.80106.123.1444.239 X Jugoslavija (A), Osijecanka (B), Zitnica (C) and NS Rana 2 (D) The analysis of genetic components in F2 showed that partial dominance (EMBED Unknown > 1), negative and positive heterosis was prevalent in the inheritance of PEN. Additive gene effect in F1 generation of hybrids was lower but dominance played a more important role in the inheritance of PEN than additive variance (H1 and H2 are higher than value of parameter D). Also, in F2 generation, nonadditive gene have higher importance than aditive (H1 and H2 are higher than value of parameter D). The positive interaction additive x dominant indicated that dominant genes have higher influence than recessive genes in the inheritance PEN. That is confirmed by the ratio of number of dominant and recessive alleles (Kd/Kr=1.581 in F1, and Kd/Kr=3.478 in F2 generation). These results are in agreement with investigation Pavlovic (1998). The frequency of dominant genes is higher compared to recessive gene and distribution of dominant and recessive alleles in F1 and F2 generation are unequal (Table 2). Analysis of variance for combining ability showed that in both F1 and F2 generation, nonaditive gene effect played more important role in the inheritance of PEN. Average value of domination in F1 and F2 generation indicated that prevail negative over dominance as a mode of inheritance of PEN (Tab. 2). In this investigation estimated variability of PEN, its genetic controls, and perspective new developed genotypes for this traits. PEN is very important for increasing productivity and decreasing of energy expenses and costs of wheat production. Development of genotypes with high capacity of PEN is very important from ecological point view. The best general combining ability (GCA) in both generation for PEN showed Zitnica and Jugoslavija cultivars. The hybrid with the best PEN in both F1 and F2 generation was Zitnica x NS Rana which is promising hybrid for further breeding (Tab. 3). CONCLUSIONS AND FUTURE WORK Improvement of PEN in the plant leading to an increasing grain N content and yield is one of the major tasks of wheat breeding. Since N absorption from soils and translocation from roots and leaves to grains are primarily under genetic control. Breeders need make concept of creation new genotypes with high physiological efficiency of nitrogen as well with high capacity of utilization, translocation and accumulation. Parents can be selected for improved biochemical, physiological and anatomical traits and crossed to high yielding agronomic ally elite materials. Also, development of new genotypes with high physiological efficiency of nitrogen and capacity of nitrogen utilization will contribute to decreasing rate of application of nitrogen fertilizers what is very important for environmental protection and development of sustainable agriculture. In this investigation were developed genotypes, by crossing Zitnica x NS Rana 2 cultivars, which expressed the highest physiological efficiency of nitrogen and represents promising for sustainable agriculture growing, because of its low requirements for fertilizer application. Also we can conclude that major scientific breakthroughs must occur in molecular biology, basic plant physiology, ecophysiology, agroecology, and soil science to achieve the ecological intensification that is needed to meet the expected increase in food demand REFERENCES Agoston T., Pepo P. (2005): Effects of genetic and ecological factors on yield formation in winter wheat production. Cereal Research Communications 33 (1):37-40. Austin R.B., Ford, M.A., Morgan, C.L. (1989): Genetic improvement in the yield of winter wheat: a further evaluation. Journal of Agricultural Science, 112, 295301. Balogh A., Pepo P., Hornok M. (2006): Interactions of cropyear, fertilization and variety in winter wheat management. Cereal Research Communications 34 (1):389-392. Clark, R. B. (1983): Plant genotype differences in uptake, translocation, accumulation and use of mineral elements required for plant growth. Genetic aspect of plant nutrition. The Haque, Boston, Lancaster: Martinus Nijhoff Publ., pp. 49-70. Desai, R. M., Bhatia, C. R. (1978): Nitrogen uptake and nitrogen harvest index in durum wheat. Euphytica: 27:561-566. Djokic, D., Knezevic, D., Kostic, M. (1992): Parameters of accumulation and distribution of dry matter and nitrogen in F1 and F2 generation of wheat. J. of Agric. Sciences 53, 75-85. Drezner G., Dvojkovic K., Horvat D., Novoselovic D., Lalic A., Babic D., Kovacevic J. (2006): Grain yield and quality of winter wheat genotypes in different environments. Cereal Research Communications 34 (1):457-460. Gamzikova, O. I., Kalashnik, N.A. (1988): Genetics of wheat characters against nutrition backgrounds. Novosibirsk, Nauka, Siberian Division, (in Russian) p.128. Gamzikova, O. I., Koval, S.F., Barsukova, V.S. (1991): Effect of genes of short-stemming on utilization of nitrogen by wheat. J. Agrochemistry, 1, 12-17. (in Russian). Gamzikova, O.I. (1994):Genetics of agrochemical wheat traits. Novosibirsk, p.220. (Russian) Griffing, B. (1956): Concept of general and specific combining ability in relation to diallel crossing systems. Aust. J. Biol. Sci., 9, 463-493. Huebner, F.R., Nelsen, T.C., Chung, O.K., Bietz, J.A. (1997): Protein distributions among hard red winter wheat varieties as related to environment and baking quality. Cereal Chem 74, 123-128. Ivanova, T.I., Mateeva, A.V. (1993): Response of winter wheat cultivars to fertilizers. Agrohemiya, 2, 120-137. (in Russian). Jinks, J.L. (1954): The analysis of continuous variation in a diallel crosses of Nicotiana rustica varieties. Genetics, 39, 767-789. Klimashevsky, E. L, Chernysheva, N.F. (1982): The investigation of genetic variability of plant mineral nutrition. Physiology and Biochemestry of Plants, 12, 375-389. (in Russian). Kochegarova, N.F. (1988): Estimation nitrogen efficiency utilization of nitrogen in spring wheat cultivars. Siberian Vestnik of Selskohozyaistvennaya Science, 2, 9-12. (in Russian). Knezevic, D ., Paunovic, A., Madic Milomirka, Djukic Nevena (2007): Genetic analysis of nitrogen accumulation in four wheat cultivars and their hybrids. Cereal Research Communications, Vol. 35 No. 2, 633-336. Mather, K., Jinks, J. L. (1971): Biometrical Genetics. Sec. ed., Chapman & Hall., London, Nielsen, N. E., Schjorring, J. K. (1983): Efficiency and kinetics of phosphorus uptake from soil by various barley genotypes. Plant and soil, 61: 384-390. Pavlovic, M. (1997): Phenotypic variability and inheritance of some parameters of nitrogen nutrition efficiency in wheat hybrids. PhD thesis..Faculty of Agriculture, Zemun,University of Belgrade. Pepo P. (2005): Effect of cropyear, genetic and agrotechnical factors on dry matter production and accumulation in winter wheat production. Cereal Res. Comm. 33,1, 29-32. Saric M., Kovacevic V. (1981): Varietals specifity of wheat mineral nutrition. In: Physiology of wheat. Serbian Academy of Sciences, Special edition, Book DXXXVI, Department of Natural Sci., Book 53 (Jovan Belic, Editor) Belgrade, p. 61-77 (in Serbian). Tribo, E., Martre, P., Tribo-Blondel, A.M. (2003): Environmentally-induced changes of protein composition for developing grains of wheat are related to changes in total protein content. J. Exp. Bot., 54, 1731-1742. ABOUT THE AUTHORS 1 Prof. dr Desimir Knezevic, Faculty of Agriculture, Zubin Potok, University Pristina, Serbia,  HYPERLINK "mailto:deskoa@ptt.yu" deskoa@ptt.yu 2Prof. Dr. Vlado Kovacevic, University J. J. Strossmayer in Osijek, Faculty of Agriculture, Trg Sv. Trojstva 3, HR-31000 Osijek, Croatia; E-mail:  HYPERLINK "mailto:vladok@pfos.hr" vladok@pfos.hr 3 Doc. Dr Aleksandar Paunovic, Associate professor, Agricultural Faculty, Cacak, Cara Dusana 32, 32000 Cacak, Serbia E-mail:  HYPERLINK "mailto:acopa@tfc.kg.ac.yu" acopa@tfc.kg.ac.yu 3 Doc. Dr Milomirka Madic, Associate professor, Agricultural Faculty, Cacak, Cara Dusana 32, 32000 Cacak, Serbia, E-mail:  HYPERLINK "mailto:mmadic@tfc.kg.ac.yu" mmadic@tfc.kg.ac.yu 4Dr Nevena Djukic, 3Faculty of Natural Sciences, Department of Biology Kragujevac, Serbia E-mail:  HYPERLINK "mailto:nevena@kg.ac.yu" nevena@kg.ac.yu     Research People and Actual Tasks on Multidisciplinary Sciences 6 8 June 2007, Lozenec, Bulgaria  Research People and Actual Tasks on Multidisciplinary Sciences 6 8 June 2007, Lozenec, Bulgaria       " . 0 6 8 F H J L N ^ ` b    µ鏃h8BhE56CJhg2hE6h]hCH*^JhC^JmH sH hChC5^Jh]hN;^JmHsHhC^JmHsHh]hC^JmHsHh]hCH*^J hC^Jh]hC^Jh-AfhC5^JhE0J L N   Q R _ (d1$gdNd1$gd&`gdN;`gd+4`gd"E<1$gd&1$#1$`gdCgdC$a$gdC1$HII $ . 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