Genetic variation for the efficiency of nitrogen uptake and use in

International Journal of Farming and Allied Sciences
Available online at www.ijfas.com
©2013 IJFAS Journal-2013-2-21/900-908
ISSN 2322-4134 ©2013 IJFAS
Genetic variation for the efficiency of nitrogen
uptake and use in Bread
wheat cultivars of Iran and Azerbaijan
Gh. Khalilzadeh1, A.R. Eivazi1*, J. Mozaffari2, Y. Arshad2
1. Agriculture and Natural Resources Research Center of Western Azerbaijan,Urmiya ,Iran
2. Department of Genetics and National Plant Gene-Bank, Seed and Plant Improvement Institute (SPII),
Karaj, Iran
Corresponding author: Ali Reza Eivazi
ABSTRACT: Due to economic and environmental concerns, development of cuttivars that absorb and
use nitrogen more efficiently is becoming an important objective of plant breeding programmes. In order
to assess the feasibility of breeding such cultivars, genetic variation for nitrogen uptake and use
efficiency was studied in 20 bread wheat cultivars of Iran and Azerbaijan, in the centre of origin and
diversity. The selected cultivars were evaluated for yield and quality characteristics and nitrogen use
-1
indices under 0 and 200 kg ha of nitrogen application at Moghan, North Western Iran, during 20072009. High levels of variation were observed among genotypes for recorded traits under both nitrogen
application treatments. Genotype×nitrogen interaction effect was significant for all characters, except for
nitrogen use efficiency (NUE) of grain dry weight (NUEgdw). Cultivars Girmizigul-1, Azamatli-95,
Chamran and Koohdasht produced the hightest grain yield under zero nitrogen application . NUE was
determined by the yield and concentration of nitrogen in grain and stable. Under N0kg treatment, the
contribution of grain yield (79%) to total nitrogen yield was higher than that of grain nitrogen
concentration (21%), while under N200kg treatment the trend was different. The contribution of grain
nitrogen concentration (55%) was relatively higher than grain yield (45%). Cultivars Gobustan,
Girmizigul-1, Azamatli-95, Chamran and Koohdasht showed the highest NUE. Nitrogen uptake efficiency
was a critical factor in both nitrogen application treatments. tIt contributed to 79% and 58% of the
variance at N0 and N200kg, respectively. Based on these observations both high grain yield and high N
uptake efficiency can be considered as two important criteria for selection of high NUE in in wheat.
Cultivars Gobustan, Koohdasht and Chamran showed the highest performance for these characteristics.
Keywords: Bread wheat, grain nitrogen, nitrogen use nitrogen uptake, genetic variation
INTRODUCTION
Nitrogen (N) is one of the major inputs in wheat production systems. During the green revolution, plant
breeding programs have released many Mexican type semi dwarf varities with greater responses to high nitrogen
input.Cultivation of these cultivars drascally increased wheat average yield in the world (Le Gouis and Pluchard,
1996). Thus the consumption of nitrogen fertilizers was increased tremendously in the world, from 13.5 million tons
in 1962 to 84.4 million tons in 2004 (FAO, 2004). Approximatly half of this amount has been applied in developing
countries (FAO, 2004). Today, elevated nitrogen level in water, as result of leaching, is an important component of
agricultural pollution, (Mariotti, 1997)causing major problems in marine ecosystems and eutrophication of
freshwater (London, 2005). However, scientists try to release cultivars with low-input of manure and decrease of
pollution risk to ecosystem (Le Gouis et al., 2000). Unfortunately, nitrogen fertilizersare not used effectively, to the
Intl J Farm & Alli Sci. Vol., 2 (21): 900-908, 2013
extent that the efficiency of nitrogen use on cereal is about 33% in the world (Yerlee and Siddiq, 1994). In order to
enhance the efficiency of crop production system while reducing the agricultural pollutions, plant breeders would
have to introduce varities which minimize pollution risks and maximize yield potential. Therefor, development of
cultivars that could absorb nitrogen more effective and use it more efficiently for grain production will lead to a
significant reduction in nitrogen fertilizers (Le Gouis et al., 2000). Genetic variation for NUE has been dtudied on
wheat (Ortez-Monasterio et al., 1997; Van Sanford and Mackown, 1986; Dhugga and Waines, 1989). Recent
studies have shown that it is possible to develop a framework for the analysis of genotypic variability for crop
nitrogen uptake capacity across a wide range of genotypes (Lemaire et al., 2004; Hirel and Lemair, 2005). There
has been also a number of studies which show improvement in modern wheat varieties for NUE, based on nitrogen
conditions (Sylvester-Bradley et al., 2009). Differences in NUE were primarily determined by greater yield, without
any increasing concentrations of N in plant material (Good et al., 2011). NUE can be defined as the product of
uptake efficiency (total N uptake/applied N through fertilizer) and utilization efficiency (grain yield/total N uptake). At
low N rates, uptake efficiency is dominant as compared to utilization efficiency, whereas utilization efficiency is
relatively more important than uptake efficiency at high nitrogen rates (Ortiz Monasterio et al., 1997). Breeding for
NUE in wheat has produced good results in some Europian countries (Good et al., 2011). There has been a 56%
decrease in total fertilizer use between 1987 and 2007, including a significant decrease in N application per
hectare. The objective of this study was to evaluate commercial cultivars of Iran and Azerbaijan for genetic
variability in N uptake and utilization efficiencies and also to identifiy appropriate screening procedure and
germplasm for these characteristics.
MATERIALS AND METHODS
The experiment was conducted in 2007-2008 and 2008-2009 with 20 bread wheat (T. aestivum L.) cultivars
released in Iran and Republic of Azerbaijan by their wheat breeding programs from 1970 to 2005. Characteristics of
these cultivars are presented in Table 1. Seeds were sown on November 2007 and 2008 in Moghan Agricultural
Research Center, North West of Iran.
Table 1. Names, date of release, heading date and plant height of wheat cultivars
Genotype
Nurlo-99
Gobustan
Ruzi-84
Gunashli
Girmizigul-1
Pirshahin-1
Gimatli2/17
Akinchi-84
Azamatli-95
Tale-38
Hirmand
Arta
Atrak
Darya
Tajan
Shiroodi
Chamran
Zagros
Koohdasht
Moghan-3
Abbreviation
Nur
Gob
Ruz
Gun
Gir
Pir
Gim
Aki
Aza
Tal
Hir
Art
Atr
Dar
Taj
Shi
Cha
Zag
Koh
Mog
Country and year
Azerbaijan-1999
Azerbaijan-2003
Azerbaijan-1984
Azerbaijan-1999
Azerbaijan-1998
Azerbaijan-1995
Azerbaijan-1985
Azerbaijan-1987
Azerbaijan-1995
Azerbaijan-1988
Iran-1991
Iran-2006
Iran-1995
Iran-2006
Iran-1995
Iran-1997
Iran-1997
Iran-1996
Iran-2000
Iran-2006
Heading Date
163
165
160
164
164
164
163
158
166
167
160
160
159
163
159
161
164
155
161
162
Plant height (cm)
75
95
100
90
80
90
80
90
95
90
90
75
75
95
80
80
85
87
95
95
-1
The soil type wasclassified as clay loam (Orthic Luvisol, FAO classification)contained an average of 12 g kg
-1
organic matter, pH 7.5 and EC about 1 ds/m. Soil samples were found to have 67 kg ha (before sowing) and 55
-1
kg ha (after harvest) mineral nitrogen in the upper 60 cm profile (Table 2).
Table 2. Soil characteristics of experimental location
Plot
Check
Treatment
Texture
Clay Loam
Clay Loam
CaCo3
25
28
TN
0.07
0.15
OC
1.04
1.2
SP %
39.5
62.4
K
592
641
P
9.8
12.7
EC
1.06
1.01
pH
7.56
7.45
Depth (cm)
0-60
0-60
901
Intl J Farm & Alli Sci. Vol., 2 (21): 900-908, 2013
The experimental design was a randomized complete block design with three replications with two splits for
-1
nitrogen levels. Control plots did not received nitrogen (N0), while fertilized plots (N+) were treated with 200 kg ha
-1
N as urease, 50 kg ha at each satges of: before sowing, tillering, beginning of stem elongation and grain filling.
2
Each plot, included six rows of 5 m long and 20 cm apart. Wheat seeds were sown on density of 350 grains m .
Date of heading was recorded, as number of days from planting to appearance of stamens on 50% of spikes.
Before harvest 20 main shoots were randomly sampled on all six rows, by cutting at the ground level, and then
oven-dried at 75ْ C for 48 h. These shoots were used to estimate thousand kernel weight (TKW), harvest index (HI),
and grain and straw nitrogen concentrations. The concentration of nitrogen in grain and straw was determined by a
Kjeldahl method (Walinga et al., 1989 ). Grain dry weight was estimated as the sum of plot harvest plus grain
weight of the shoot samples. Nitrogen harvest index (NHI) was calculated as the ratio of grain nitrogen/Total
above-ground nitrogen. Grain nitrogen utilization efficiency is determind by grain dry weight/Total above-ground
nitrogen (May et al., 1991). The contribution of each component to the NUE was calculated according to methods
described by( Moll et al., 1982 ; Dhugga and Waines, 1989). Yn=X1n+ X2n, Yn is logY, and X1n and X2n are two
component logs, ∑(X1nYn)/ ∑Y2n and ∑ (X2nYn)/ ∑Y2n are contribution of each dependent triat (Dhugga and Waines,
1989).
2
Log(grain yield)= log(ears per m )+ log(kernel per ear)+ log(TKW)
Log(NUE)= log(N uptake efficiency)+ log(N utilization efficiency)+ log(HI)
2
When the G×N interaction was significant for a character, we computed the (Wricke, 1962) Equivalence (W g)
2
2
which was computed as: W g=∑ (Xgn – Xg.. – X.n + X..) , where, N is the nitrogen level, Xg.. is the mean of
genotype g in all N levels, X.n is the mean of N level n in all levels, X.. is general mean. Analysis of data analysis
was carried using SPSS and MSTAT-C softwares.
RESULTS AND DISCUSSION
Results
Analysis of variance showed significant differences among genotypes for all traits studied (Table 3). The effect
of Nitrogen application was significant for most of the traits expect for harvest index and stable N yield.
Genotype×Nitrogen (G×N) interaction was also significant for all characters expect for NUEgdw. Mean values of
traits for genotypes under the two N application treatments are presented in Table 4.
Grain yield and yield components
-1
-1
-1
Average grain yield of cultivars decreased from 8852 kg ha at N+ (200 kg ha ) to 7221 kg ha at N0 (Fig.
1a). Some cultivars such as Gunashli, Akinchi-84 and Zagrose showed low grain yield at both nitrogen levels.
Based on the Wricke Equivalence, cultivars Gunashli, Akinchi-84, Hirmand, Zagros and Moghan-3 were
responsible for 42% of the G×N interaction (Table 5) in the contrary, TKW was decreased significantly from N0
(40.1 gr) to N+ (37.7 gr) conditions and cultivars Gobustan, Atra, Tajan and Moghan-3 decreased the most.. These
four cultivars were responsible for about 40% of the G×N interaction variance of TKW (Table 6). Although, harvest
index was not significantly affected (Table 3) by the effects of genotype, the effect of G×N interaction on harvest
indexwas significant (P<0.01). Cultivars Girmizigul-1, Pirshahin-1, Atra, Tajan, Shiroodi and Zagros showed the
highest harvest index and cultivar Akinchi-84 the lowest among genotypes at both N levels (table 4).
Nitrogen use efficiency and its components
Nitrogen uptake and utilization efficiencies, and harvest index are the three components of NUE. In general,
nitrogen uptake efficiency was significantly higher at N+ (1.52) than at N0 (1.0) (table 4 and Fig. 1c). High levels of
variation for N uptake efficiency was obserwed in commercial wheat cultivars of both Iran and Azerbaijan. . Three
cultivars Gobustan, Chamran and Moghan-3 were responsible for 59.5% of the G×N interaction variance, however,
-1
cultivar Chamtan showed the highest N uptake efficiency. Total N utilized ranged from 23.3 to 34.2 kg kg nitrogen
-1
in N+ and 33.3 to 42.0 kg kg nitrogen in N0, and was significantly effected by genotypes and G×N interactions.
The highest N utilization efficiency was observed in cultivars Girmizigul-1, Atra, Shiroodi and Chamran. Cultivar
Girmizigul-1 showed the highest sensitivity to reduced nitrogen level.
902
Intl J Farm & Alli Sci. Vol., 2 (21): 900-908, 2013
Grain nitrogen yield
Grain nitrogen concentration was significantly affected (P<0.01) by Genotype (G) and G×N interaction effects.
It ranged from 1.83 to 2.18% at N0 and 2.21 to 3.55% at N+ treatment. All cultivars had a lower value at N0
(2.06%) than N+ (2.85%). Cultivars Gobustan, Zagros and Moghan-3 contained the highest grain N concentration,
while also showing the highest sensitivity to N reduction (Table 4).
Contribution of component traitsto NUE and grain N yield
Relative contributions of component traits to the variance of the resultant nitrogen efficiency traits were
presented in Table 5. Among three components of NUE, N uptake efficiency contributed the most (79%) followed
by total N utilization (26%) and harvest index (9%). In addition the contribution of grain yield on the nitrogen yield in
significantly higher than nitrogen concentration. Most of grain N yield variance was explained by grain yield at N0
(78%) while contribution of nitrogen concentration (22%) was less important. N uptake efficiency and NHI were
determined as traits contributing to NUEgn, the relative with 74% and 26% of the variance, respectively. Relative
contribution of N uptake and NHI at NUEgn showed impotance of N uptake. N uptake varianc showed at N0 (74%),
N+ (64%) and G×N (53%) while in NHI those values were N0 (26%), N+ ( 36%) and G×N (47%) respectively.
Discussion
Breeding wheat cultivars with high nitrogen uptake capacity and nitrogen use efficiency under low nitrogen
input conditions is a key to reduce nitrogen application in wheat production systems (Austin et al., 1980; Power et
al., 2000). The observation of considerable variation for grain yield and NUE at N0 condition indicates that
significant genetic variation exists in bread wheat cultivars of Iran and Azerbaijan. Thus, this gerplasm can be used
for developing wheat varieties with higher efficiencies of nitrogen uptake and use, and suitable for low input wheat
production system. The significant G×N interaction for grain yield (P<0.5) was also a good evidence for varying
responses of these cultivars at variouse N levels (Earl and Ausubel, 1983; Austin et al., 1980).
A positive relationship between N application levels and the grain yield has alredely been shown in many
studies (Austin et al., 1980; Desai and Bahatia., 1979). In the contrary, ally, a negative correlation between grain
yield and grain nitrogen concentration has also been reported in earlier stidies of bread wheat (Cox et al., 1985;
Stoddard and Marshall, 1990).
Table 3. Mean squares traits of wheat genotypes at two nitrogen fertilizer applications
S.O.V
Mean Squares
df
Year
1
Rep(Y)
4
Nitrogen
1
N×Y
1
Ea
4
Genotype
19
G×Y
19
G×N
19
G×N×Y
19
Eb
Mean
CV%
152
-
Grain
yield
TKW
21.14
*
7.04
ns
159.7
**
2.21
ns
1.31
7.10
**
0.31
ns
2.71
*
0.08
ns
0.053
8036
9.1
10.84
*
3.77
ns
329.00
**
1.50
ns
1.254
163.58
**
0.697
ns
38.07
**
0.311
ns
2.758
38.8
4.3
Harvest
index
Nharvest
index
Grain N
Concentration
Stable N
Concentration
Grain N
yield
0.419 **
0.009 ns
4254.6 ns
0.994 **
0.01 ns
1907.3 ns
37.52 **
0.497 **
643221 **
0.007 ns
0.000 ns
1934.0 ns
0.002
0.004
1120.8
0.283 **
0.114 **
6430.6 **
9.13
95.24
**
43.35
ns
370.32
**
5396.1
**
19.27
ns
21.43
78.63
**
1.81 ns
2.33 ns
0.006 ns
0.001 ns
240.08 ns
35.12
**
1.558
ns
13.47
45.1
8.1
38.40
**
0.319 **
0.144 **
2838.2 **
1.81 ns
0.005 ns
0.001 ns
110.4 ns
13.38
78.2
4.7
0.008
2.46
3.7
0.002
0.533
8.1
376.02
200
9.7
125.4 *
222.9
**
21.9 ns
9.48 ns
Stable
N yield
69.77
ns
2772.5
*
266.7
ns
47.7
ns
379.82
599.84
**
8.06
ns
331.76
**
5.23
ns
70.32
52
16.1
NUEgdw
513.34 *
175.78
ns
4059.04
**
49.5 ns
34.51
175.19
**
7.65 ns
17.1 ns
2.18 ns
13.38
40.2
9.2
NUEgn
(kg kg1)
1066.8
ns
481.82
ns
160890
**
493.07
ns
282.12
1599.6
**
59.17
ns
711.43
**
27.19
ns
94.7
100
9.7
N uptake
Efficiency
(kg kg-1)
N
utilization
efficiency
(kg kg-1)
0.117 ns
150.4 ns
0.024 ns
153.8 ns
15.66 **
2522.1 **
0.04 ns
0.150 ns
0.036
26.71
0.185 **
50.46 **
0.005 ns
1.55 ns
0.08 **
33.85 **
0.004 ns
1.229 ns
0.011
1.26
8.2
3.592
32.6
5.8
ns , Not significant: *, significant at p<0.05: **, significant at p<0.01
903
Intl J Farm & Alli Sci. Vol., 2 (21): 900-908, 2013
Table 4. Mean traits of wheat genotypes at two levels of nitrogen applications
Nitrogen
level
N0
N200
LSD
(5%)
Genotype
Grain
yield
-1
(kg ha )
TKW
(gr)
HI
(%)
NHI
(%)
Grain N
Concentration
(%)
Stable N
Concentration
(%)
Grain N
yield
-1
(kg ha )
Stable
N yield
(kg ha
1
)
NUEgdw
-1
(gr gr )
NUEgn
-1
(gr gr )
N uptake
Efficiency
-1
(gr gr )
N
utilization
efficiency
-1
(gr gr )
Nur
Gob
Ruz
Gun
Gir
Pir
Gim
Aki
Aza
Tal
Hir
Art
Atr
Dar
Taj
Shi
Cha
Zag
Koh
Mog
Nur
Gob
Ruz
Gun
Gir
Pir
Gim
Aki
Aza
Tal
Hir
Art
Atr
Dar
Taj
Shi
Cha
Zag
Koh
Mog
N
G
G×N
5998
8072
6640
7579
8168
6419
6158
7301
8386
8031
6618
6548
7071
7256
6215
7238
8405
7813
8427
6074
7357
10080
8794
8606
9541
7819
8201
8218
9662
9126
8882
8549
8566
9241
8236
9358
9538
8652
10345
8374
292.1
588.2
831.8
34.3
36.5
41.7
47.3
34.3
37.7
46.7
43
38.5
35.7
41.3
42.3
42.8
31.8
40.7
36.7
41.8
39
42.2
46
39.8
29.7
38.7
44
32.8
33.7
44.2
39.2
36.2
36.5
39.8
36.5
34.7
34.8
33.3
36.3
40.8
40
42.8
39.7
0.29
1.34
1.89
42.4
47.1
41.9
46.1
50.1
46.8
46.6
45.5
43.6
40.1
44.9
49.2
46
43
48.8
46.4
45.4
49
40.8
44.1
48.6
46.9
38.4
46.2
46.1
48.3
44.1
38
38.8
39.8
41.4
44.8
48.1
44.3
46.3
50.4
46.7
48
43.1
47
0.77
2.96
4.19
75.3
75.2
73.7
69.2
76
72.8
71
72.2
76.2
70.2
70
72.3
80
67.3
70.8
77.3
76.8
76.5
75.5
71.5
75.5
86
86.3
84
87.5
84.8
80.7
80.7
85.7
82
77.7
81.7
84
82.7
80.7
82.2
84
85.7
84.8
83
1.18
2.95
4.17
2.03
2.06
2.16
2.03
1.83
1.95
2.00
2.14
2.11
2.03
2.01
2.18
2.11
2.15
2.06
2.01
2.14
2.06
2.05
2.12
2.69
3.16
3.00
3.12
2.77
3.08
2.88
2.75
2.79
3.13
2.53
2.51
2.45
2.70
2.75
2.49
2.40
3.21
3.09
3.55
0.012
0.074
0.104
0.35
0.36
0.48
0.46
0.76
0.50
0.43
0.60
0.60
0.58
0.53
0.46
0.49
0.60
0.60
0.37
0.30
0.41
0.50
0.40
0.39
0.73
0.77
0.47
0.52
0.36
0.39
0.32
0.36
0.48
0.82
0.83
0.84
0.51
0.74
0.70
0.45
0.61
0.60
0.69
0.016
0.035
0.049
121
166
142
152
149
126
124
156
177
163
132
142
149
155
127
146
180
161
172
128
197
318
263
268
263
240
235
226
269
285
225
214
209
248
224
233
228
278
319
295
8.5
15.6
22.1
39.2
55.6
48.2
67.7
46.4
46.0
49.4
60.0
54.2
69.2
56.2
54.6
37.2
75.0
52.2
42.0
54.6
48.4
55.4
50.4
63.8
51.4
41.8
50.0
36.8
43.2
56.4
54.6
45.2
62.2
63.8
47.4
40.2
52.4
53.6
50.2
44.0
46.6
56.2
59.9
4.97
6.76
9.56
29.8
40.3
33
37.8
40.7
32.2
30.8
36.5
42
40.2
33.2
32.8
35.3
36.2
31
36.2
41.8
38.8
42.2
30.3
37
50.5
44
43
47.7
39.2
41.2
41.2
48.3
45.5
44.3
42.8
42.8
46.2
40.7
46.8
47.8
43.3
51.5
41.8
1.50
2.95
4.17
62.3
82.8
70.7
76.2
74.2
62.8
62.0
77.8
88.7
81.3
65.8
71.0
74.5
77.7
63.5
72.7
89.8
80.3
85.8
64.2
114.8
153.2
131.3
133.7
131.8
120.0
117.5
113.0
134.2
142.5
112.2
107.0
104.8
124.2
112.0
116.7
114.0
137.5
153.7
142.7
4.24
7.68
10.95
0.8
1.08
0.93
1.08
0.98
0.87
0.88
1.1
1.15
1.15
0.95
0.98
0.93
1.15
0.88
0.93
1.17
1.03
1.13
0.88
1.3
1.85
1.5
1.58
1.5
1.42
1.47
1.42
1.57
1.73
1.45
1.32
1.25
1.52
1.4
1.42
1.37
1.63
1.85
1.77
0.049
0.084
0.118
37.7
36.3
34.2
34.5
42.0
37.5
35.5
34.0
36.2
34.5
35.2
33.3
38.0
31.3
34.5
38.5
35.8
37.3
37.3
34.0
28.3
27.3
29.0
26.8
31.7
27.3
28.2
29.2
30.8
26.2
30.8
32.5
34.2
30.8
29.3
33.0
35.0
26.7
27.5
23.3
1.32
1.53
2.16
904
Intl J Farm & Alli Sci. Vol., 2 (21): 900-908, 2013
Figure 1. Respons of 20 wheat cultivars to nitrogen application as indicated by equivalence for grain yield (a), harvest index
(b), nitrogen uptake efficiency (c), grain nitrogen yield (d), nitrogen harvest index (e) and nitrogen use efficiency. The
contribution of each genotype to the G×N level interaction (equivalence) was indicated when superior to 5%. Black and white
pointes shows Azerbaijani and Iranian cultivars respectively.
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Intl J Farm & Alli Sci. Vol., 2 (21): 900-908, 2013
Table 5. Percentage of Wrick Equivalence for characteristics of grain yield, grain nitrogen yield, harvest index, nitrogen harvest
index, nitogent uptake and use efficiency in 20 genotypes
Genotype
Grain yield
Grain N yield
Harvest index
N harvest index
Grain N
content
N uptake
N utlization
Nurlo-99
1.7
4.2
20.7
36.6
4.5
0.1
0.9
Gobustan
3.0
13.1
0.1
0.6
6.0
12.3
3.7
Ruzi-84
5.9
1.7
3.8
3.7
5.1
0.5
7.3
Gunashli
8.2
0.9
0.2
11.2
5.9
0.1
8.3
Girmizigul-1
1.5
0.6
5.2
1.5
5.4
0.0
1.3
Pirshahin-1
1.2
0.6
2.0
2.4
6.1
0.2
1.3
Gimatli2/17
3.6
0.3
1.6
0.0
5.2
1.0
4.5
Akinchi-84
11.4
6.2
21.3
0.5
4.4
7.9
11.4
Azamatli-95
2.9
0.7
7.9
0.0
4.6
2.0
3.3
Tale-38
6.4
1.9
0.0
2.0
6.0
0.7
7.8
Hirmand
8.7
0.6
3.8
1.5
4.1
0.1
7.8
Arta
2.9
5.5
6.5
0.0
3.6
6.4
3.0
Atrak
0.4
10.5
3.3
13
3.6
7.9
0.5
Darya
2.7
0.6
1.6
13.9
4.2
4.4
3.0
Tajan
3.3
0.2
1.6
0.0
4.6
0.0
2.1
Shiroodi
5.2
1.5
9.5
9.1
4.0
0.2
5.3
Chamran
5.6
17
1.6
2.4
3.4
20.2
4.5
Zagros
14
1.0
0.1
0.1
6.2
1.3
12.7
Koohdasht
1.7
10.5
3.8
0.0
5.8
7.9
1.1
Moghan-3
9.7
22.4
5.5
1.5
7.2
27.0
10.1
Table 6. Contribution of the components traits to the variance of resultant trait in 20 wheat cultivars at each N level and
Genotype×Nitrogen interaction
Resultant trait
Log(NUE)
Log(Grain N yield)
Log(NUEgn)*
Component traits
Log(N uptake efficiency)
Log(total N utilization efficiency)
Log(harvest index)
Log(grain yield)
Log(grain N content)
Log(N uptake efficiency)
Log(N harvest index)
N levels
N0 (0Kg ha-1)
0.79
0.26
-0.09
0.79
0.21
0.74
0.26
N+ (200Kg ha-1)
0.58
0.24
-0.18
0.45
0.55
0.64
0.36
Interaction
G×N
0.58
-0.31
-0.11
0.48
0.52
0.53
0.47
*NUEgn=grain N yield/N supply
Number of recent studies have demonstrated that modern wheat varieties with improved NUE, are primarily
determined by their greater yield, not by their increased concentrations of N in the plant material (Good et al.,
2011). Modern UK wheat varieties have shown a 14% to 18% increase in NUE, depending on the N application
conditions (Sylvester-Bradley et al., 2009). This was vene greater in modern Spanish wheat varieties, ranging
between 24% to 29% increas in NUE (Acreche, 2009). (Ceccarelli, 1996) emphasized on an optimal condition to
select varites for low-input environments. He showed that lines selected for high yield in favorable environments
yield more in medium to high yielding conditions than lines selected in less favorable conditions.
Among yield components, the thousand kernel weight was affected significantly and may be unexpectedly by
application of nitrogen. TKW was reduced in most of the cultivars by N fertilization from N0 to N+ treatment. The
contribution of TKW in grain yield variation was 15% more at N0 comparred to N+ conditions (datas not shown).
Significant differences between N+ and N0 treatments for nitrogen uptake efficiency of a given wheat cultivar
indicates high variation for this trait in wheat varieties of Iran and Azerbaigan. Evaluation of component traits of
NUE showed that, most of variation in this characteristics was due to nitrogen uptake efficiency. This was in
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Intl J Farm & Alli Sci. Vol., 2 (21): 900-908, 2013
agreement with the findings in earlier studies on CIMMYT's semi-dwarf and France's old wheat cultivars (OrtizMonasterio et al., 1997; Le Gouis et al., 2000). Harvest index was less important than total N utilization efficiency in
explaining the variation of NUE (Table 6), especially at N0 condiotions. When N is not the limiting factor, N uptake
and N utilization efficiency are determinant factors. When N is limited in the soil, the ability to absorb N become
important as the capacity of N absorbtion also relates to root system charactersitics (Guingo et al., 1998; Kamara
et al., 2003; Coque and Gallaris, 2006). At low nitrogen application rates, N uptake efficiency is dominant
compared to N utilization efficiency, whereas utilization efficiency is relatively more important than uptake efficiency
at high nitrogen application rates (Ortiz-Monasterio et al., 1997). Among the three main NUE component traits N
uptake efficiency showed the highest contribution to the NUE variance at N0 (79%), N+ (58%) and G×N (58%)
interaction, respectively. Significant differences in traits contributing to N uptake and utilization efficiencies indicats
the existence of a high genetic diversity among wheat cultivars widely grown in Iran and Azerbaigan. Based on the
results of this study we conclude that: a) wheat germplasm of Iran and Azerbaijan can be used as sources of
selection for nitrogen uptake and use efficiencies; b)for selecting varities with high NUE, grain nitrogen yield seem
to be more efficient indicator than nitrogen concentrations in the grain.
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