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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
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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
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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).
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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
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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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