RESEARCH PEOPLE AND ACTUAL TASKS ON MULTIDISCIPLINARY SCIENCES 6 – 8 JUNE 2007, LOZENEC, BULGARIA GENETIC ANALYSIS OF PHYSIOLOGICAL EFICIENCY OF NITROGEN IN FOUR WHEAT CULTIVARS AND THEIR HIBRIDS D. KNEZEVIC1, V. KOVAČEVIĆ2, 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 F 1 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; Triboï 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 – – RESEARCH PEOPLE AND ACTUAL TASKS ON MULTIDISCIPLINARY SCIENCES 6 – 8 JUNE 2007, LOZENEC, BULGARIA 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 F 1 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 Cultivar A B C D +d -sd -sd Jugoslavija (A) 25 25 22 18 F1 hybrids Osijecanka (B) 25+d 24 20-sd 21-d LSD 0.05 LSD 0.01 -sd -sd -d Zitnica (C) 22 23 26 21 4.46 6.02 NS Rana 2 (D) 23 i 23 i 20-sd 21 F2 hybrids LSD 0.05 = 3.85 LSD 0.01 = 5.09 – – RESEARCH PEOPLE AND ACTUAL TASKS ON MULTIDISCIPLINARY SCIENCES 6 – 8 JUNE 2007, LOZENEC, BULGARIA Table 2. Genetic variance and heritability for Physiological efficiency of nitrogen (PEN) in wheat hybrids Variance Physiological efficiency of Variance Physiological efficiency of nitrogen (PEN) nitrogen (PEN) F1 F2 F1 F2 u D 1.89 2.542 0.645 0.751 v H1 43.243 28.469 0.354 0.258 D H2 37.898 18.703 4.783 3.346 H1 F 4.087 9.448 KD/KR 1.581 3.478 *Th2 E 2.38 1.729 5.82% *Gh2 H2 / 4H1 0.876 0.657 7.62% 2 2 * Th = total heritability¸ *Gh = genetical heritability Table 3. General and specific combining abilities for Physiological efficiency of nitrogen (PEN) in wheat hybrids Wheat cultivar X LSD A B C D Se 0.05 0.01 General combining ability F1 PEN 0.054 0.326 0.598 -1.028 0.702 1.312 1.730 Rank 3 2 1 4 F2 PEN 0.190 -0.054 1.306 -1.006 0.823 1.519 2.031 Rank 3 2 1 4 Specific combining ability of four wheat cultivars LSD A B C D Se 0.05 0.01 A PEN -1.2138 -2.7544 0.1867 F1 hybrids B PEN -2.2408 0.0466 -4.295 10.46 3.721 4..974 C PEN -0.9803 0.4201 1.5872 F2 hybrids D PEN -2.5209 0.1392 2.8010 6.12 3.144 4.239 X Jugoslavija (A), Osijecanka (B), Zitnica (C) and NS Rana 2 (D) The analysis of genetic components in F2 showed that partial dominance ( H1 D > 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 F 1 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). – – RESEARCH PEOPLE AND ACTUAL TASKS ON MULTIDISCIPLINARY SCIENCES 6 – 8 JUNE 2007, LOZENEC, BULGARIA 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. 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ABOUT THE AUTHORS 1 Prof. dr Desimir Knezevic, Faculty of Agriculture, Zubin Potok, University Pristina, Serbia, [email protected] 2Prof. Dr. Vlado Kovacevic, University J. J. Strossmayer in Osijek, Faculty of Agriculture, Trg Sv. Trojstva 3, HR-31000 Osijek, Croatia; E-mail: [email protected] 3 Doc. Dr Aleksandar Paunovic, Associate professor, Agricultural Faculty, Cacak, Cara Dusana 32, 32000 Cacak, Serbia E-mail: [email protected] 3 Doc. Dr Milomirka Madic, Associate professor, Agricultural Faculty, Cacak, Cara Dusana 32, 32000 Cacak, Serbia, E-mail: [email protected] 4Dr Nevena Djukic, 3Faculty of Natural Sciences, Department of Biology Kragujevac, Serbia E-mail: [email protected] – –
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