Published September 24, 2012

Change in Nitrogen Requirement with Increasing

Grain Yield for Winter Wheat
Shanchao Yue, Qingfeng Meng, Rongfang Zhao, Youliang Ye, Fusuo Zhang,
Zhenling Cui,* and Xinping Chen*

Soil Fertility & Crop Nutrition

ABSTRACT
Overestimates of N requirements have led to excessive N application and serious environmental pollution in intensively managed
agricultural systems. A database comprising 1395 measurements was developed from 2000 to 2011 using 88 on-farm and station
experiments conducted in five key winter wheat (Triticum aestivum L.) domains in northern China. The database was created to
evaluate the relationship between aboveground N uptake and grain yield with different N treatments and to quantify N requirements
per Mg grain at different grain yield levels. Across all sites, wheat grain yield ranged from 1.6 to 11.8 Mg ha–1. The nitrogen
requirement per megagram grain yield (Nreq.) increased with N supply from 20.8 kg under the treatment without N fertilizer to
25.7 kg under the excess N treatment. For the optimal N fertilizer treatment, the average Nreq. was 24.3 kg and it declined with
increasing grain yield. For the yield ranges between <4.5 and 6.0 to 7.5 Mg ha–1, Nreq. decreased from 27.1 to 24.5 kg due to increasing
harvest index (HI; from 0.39–0.46) and decreasing grain N concentration (from 2.41 to 2.21%). For the yield ranges between 6.0
to 7.5 and 9.0 to 10.5 Mg ha–1, Nreq. decreased from 24.5 to 22.7 kg due to decreasing grain N concentration (from 2.21 to 2.00%).
For the yield ranges between 9.0 to 10.5 and >10.5 Mg ha–1, Nreq. changed little due to stability in grain N concentrations and HI. In
conclusion, the N requirement of a crop was affected by both the amount of N supplied and the grain yield.

N itrogen requires judicious management in inten-

sively managed cropping systems due to its requirement
in attaining optimal yield offset by the cost of the fertilizer as
well as deleterious environmental effects when applied in excess.
Recent literature on improving the recovery efficiency of N in
crop production has emphasized the need for greater synchrony
throughout the growing season between crop N demand and N
supply from all sources (Cassman et al., 2002; Cui et al., 2008,
2010). However, overestimates of N requirements in intensive
agricultural regions, especially under high-yielding conditions,
have led to excessive applications of N fertilizer and are associated
with widespread environmental problems (Guo et al., 2010; Le et
al., 2010; Liu et al., 2011). For example, in 43 high-yielding maize
studies that achieved >15 Mg ha–1 grain yield in China, average
N fertilizer input was 747 kg ha–1, whereas N removal by the
harvested grain was only 292 kg ha–1 (Chen et al., 2011). Nitrogen
inputs that are higher than the crop uptake limit our understand-
ing of N requirements in high-yielding systems, make demonstra-
tions of technologies more difficult, and mislead producers into
believing that greater resource inputs produce higher grain yield.
S. Yue, Q. Meng, F. Zhang, Z. Cui, and X. Chen, Center for Resources,
Environment and Food Security, China Agricultural Univ., Beijing 100193,
China; R. Zhao, College of Environmental Science and Engineering,
Zhongkai Univ.of Agriculture and Engineering, Guangzhou 510225, China;
Y. Ye, College of Resources and Environmental Sciences, Henan Agricultural
Univ., Zhengzhou 450000, China. Received 20 June 2012. *Corresponding
author (cuizl@cau.edu.cn; chenxp@cau.edu.cn).
Published in Agron. J. 104:1687–1693 (2012)
Posted online 14 Sept. 2012
doi:10.2134/agronj2012.0232
Copyright © 2012 by the American Society of Agronomy, 5585 Guilford
Road, Madison, WI 53711. All rights reserved. No part of this periodical may
be reproduced or transmitted in any form or by any means, electronic or
mechanical, including photocopying, recording, or any information storage
and retrieval system, without permission in writing from the publisher.
Based on extensive on-farm investigations from 1997 to 2007
(Cui et al., 2010), farmers typically applied >300 kg ha–1 N for
winter wheat in northern China, while N uptake was only about
160 kg ha–1. Understanding the N uptake–yield relationship and
quantifying N requirements would be of great benefit for optimiz-
ing N fertilization for winter wheat production, particularly for
high-yielding system.
Most previous studies that have been conducted to quantify the
N uptake–yield relationship for winter wheat and have reported
variations in grain yield-N uptake requirements among regions
(environments), genotypes, and N management strategies (Ortiz-
Monasterio et al., 1997; Pathak et al., 2003; Liu et al., 2006;
López-Bellido et al., 2008; Barraclough et al., 2010; Gaju et al.,
2011; Giuliani et al., 2011; Pask et al., 2012). In an investigation
of 10 winter wheat varieties in Mexico, the N requirement per Mg
grain (on the basis of 15% moisture) ranged from 23.0 to 37.0 kg
(Ortiz-Monasterio et al., 1997). In an examination of 16 winter
wheat varieties in the United Kingdom (UK) and France, the
N requirement per Mg grain ranged from 14.4 to 31.1 kg under
low and high N treatments, respectively (Gaju et al., 2011). Based
on experiments conducted from 1985 to 1995 across widely
differing environments in China, the average N requirement
of winter wheat was 24.6 kg for yields that ranged from 0.35 to
8.73 Mg ha–1 (Liu et al., 2006). To accurately estimate crop N
requirements, we need a better understanding of the factors that
influence variation in N requirements and we need to determine
the degree of influence that occurs in a certain region.
Nitrogen management influences crop N uptake; the response
in N uptake to N fertilizer can be described by a curve that includes
three phases designated as the minimum, deficiency adjusted, and
excess consumption zones (Fowler, 2003). Despite the fact that N
Abbreviations: HI, harvest index; Nreq, the nitrogen requirement per megagram
grain yield; NHI, nitrogen harvest index.

A g r o n o my J o u r n a l • Vo l u m e 10 4 , I s s u e 6 • 2 012 1687
requirements with and without N fertilization have been studied
(Ortiz-Monasterio et al., 1997; Ferrise et al., 2010), few studies have
examined different N levels, particularly under optimal N supply.
Therefore, the estimated N requirements may represent overestimates
or underestimates under unsuitable N fertilizer applications.
Crop N requirements can also be affected by grain yield, the
HI, and the N concentration at harvest, which are needed to
calculate aboveground N uptake. Various studies have found
negative relationships between grain N concentration and grain
yield in winter wheat (Triboi et al., 2006; Barneix, 2007; Bogard
et al., 2010). This indicates that N requirements decline as grain
yield increases. However, whether the trend in this decrease is an
equidistant, accelerated, or phasic process is still not clear.
Previous studies that estimated wheat N requirements often
involved site-specific field experiments, mostly at research stations.
For example, studies on the N requirements of winter wheat
in China have been mainly conducted for grain yields up to
8.7 Mg ha–1 (Liu et al., 2006). The concern is whether this value
represents the current range of yield and the N requirements
of modern, high-yielding varieties in China. In this study, we
developed a large database for winter wheat that covers a wide
yield range (1.6–11.8 Mg ha–1), which is higher than the wheat
yield studied by Liu et al. (2006). Four N-application treatments
were defined for use in this study: a control without N fertilizer
(N = CK), optimal N fertilizer input based on in-season root-zone
N management (N = Opt.), below optimal N input (N < Opt.),
and above optimal N input (N > Opt.). Previous research has
shown that N = Opt. treatments based on in-season root-zone
N management can maximize winter wheat grain yield while
minimizing the environmental impacts of N fertilization (Cui
et al., 2008; Ju et al., 2009). The objectives of this study were to
evaluate the relationship between aboveground N uptake and
grain yield under different N treatments and to quantify the N
requirement per Mg grain for different wheat grain yield levels
under optimal N application conditions.
MATERIALS AND METHODS
Data Sources
Eighty-eight field experiments were conducted in five key
winter wheat domains in northern China between 2000 and
2011. These studies were conducted in Dongbeiwang (DBW)
in Beijing; Quzhou (QZ) in Hebei Province; Huimin (HM),
and Tai’an (TA) in Shandong Province; Lankao (LK), Wenxian
(WX), Xiangyun (XY), and Zhaobao (ZB) in Henan Province;
and Hongtong (HT) and Yongji (YJ) in Shanxi Province
(Table 1). All the field experiments had already been published
in research journals or master’s theses (Zhao et al., 2006; Cui
et al., 2009; Guo, 2011; Ye et al., 2011, 2012; Yue et al., 2012).
The specific details of these field experiments are not reported
in this paper because the aim of this study was to quantify the N
requirement per Mg grain for winter wheat under a wide range
of environmental conditions. In total, 1395 measurements of
winter wheat grain yield, total dry matter, and N concentration
at maturity were collected. Table 1 details the number of
experimental sites, varieties, N treatments, data points, and yield
ranges in the study areas.
All of the experimental sites, except the experiment in QZ2,
included at least three N treatments: no N as a control (CK), an
optimal N rate (Opt.) based on either in-season root zone or yield
goal N management (Cui et al., 2008; Ju et al., 2009), and the
farmer’s nitrogen practice (FNP). Nitrogen treatments at sub- or
supra-optimal N rates were applied in experiments with five or six
N treatments, including Opt.– 30 kg N ha–1; Opt. + 30 kg N ha–1;
Opt. + 60 kg N ha–1; 40, 50, 70, 75, 125, 130, and 150% of Opt.
according to the experimental design at each site. The experiment in
QZ2 received two N treatments: Opt. and FNP. For the in-season
root-zone N-management strategy, the wheat-growing season is
divided into two periods: from planting to the stem elongation stage,
and from the stem elongation stage to the mature stage. The amount
of N fertilizer applied at the beginning of each growing period was
determined by deducting the amount of soil Nmin (NH4+–N +
NO3−–N) in the root zone from the target N value, which was
estimated based on the yield target and crop N uptake. Detailed
descriptions of the target N value and the soil Nmin for all field
experiments have been previously reported (Zhao et al., 2006; Cui et
al., 2009; Guo, 2011; Ye et al., 2011, 2012; Yue et al., 2012).
All of the experiments consisted of a randomized complete
block design with three or four replications. Plot area varied
from 20 to 300 m2 . A typical winter wheat–summer maize
rotation system was adopted at all of the experimental sites.
Winter wheat was planted in mid-October and harvested
in mid-June of the following year. Based on soil P and K test
results, all of the experiments received appropriate amounts of
triple superphosphate (0–150 kg P2O5 ha–1) and potassium

Table 1. Number of experimental sites, varieties, N treatments, data points, and yield range in different regions of northern China.
Location Year No. of experiments Variety no. N treatment No. of data points Range of yield Reference
Mg ha–1
Beijing DBW 2000–2005 1 2 3 288 1.6–7.3 Zhao et al., 2006

Hebei QZ1 2008–2011 1 2 5 80 2.1–8.0 Yue et al., 2012

QZ2 2009–2010 1 4 2 84 4.9–10.1 Guo, 2011

Shandong HM1 2004–2007 12 3 5, 6 304 3.4–8.6 Yue et al., 2012

HM2 2004–2005 41 7 3 123 3.2–8.3 Cui et al., 2009
TA 2005 9 4 3 45 7.4–10.5 Cui et al., 2009

Henan LK 2008 1 2 5 30 9.7–11.8 Ye et al., 2012

WX 2008, 2010 2 3 5, 6 81 7.3–11.3 Ye et al., 2012
XY 2008 1 15 3 135 6.1–11.8 Ye et al., 2011
ZB 2008 1 15 3 135 5.5–11.5 Ye et al., 2011

Shanxi HT 2005 9 1 3 45 4.0–8.5 Cui et al., 2009

YJ 2005 9 1 3 45 2.6–7.2 Cui et al., 2009

1688 Agronomy Journal • Volume 104, Issue 6 • 2012

chloride (0–120 kg K 2O ha–1) before planting. Urea was applied
before winter wheat was planted and as a top dress at the stem
elongation stage according to the specific N treatment. No
obvious water, weed, pest, or disease stress was observed at any of
the sites during the growing season.
dry matter. The nitrogen harvest index (NHI) is defined as the N
accumulation in the grain proportion of the total aboveground
N uptake. The N requirement per Mg grain (Nreq.) is defined as
the amount of aboveground N needed to produce 1 Mg grain (on
the basis of 15% moisture).

Sampling and Laboratory Procedures Data Analysis

Plant measurements followed similar protocols at all sites.
Wheat plants from an area at least 3 m2 in the middle of each plot
were harvested manually at maturity. Dry weights of stems and
grain were determined after separation and oven-drying at 60°C
in a forced-draft oven. Subsamples that passed through a 1-mm
screen in a sample mill were mineralized using H2SO4–H2O2
and plant N concentrations were determined using the standard
Kjeldahl method (Horowitz, 1970).
Grain yields were reported at a standard moisture content
of 0.15 kg H2O kg–1 fresh weight. Grain and straw N
concentrations were reported on the oven dry basis. The HI is
defined as the grain dry weight proportion of the aboveground
All data from the N treatments in the various experiments
were divided into four groups: no N as a control (N = CK);
below optimal N application (N < Opt.), which included
Opt.– 30 kg N ha–1 and 40, 50, 70, and 75% of Opt.;
an optimal treatment based on in-season root-zone N
management or a yield goal (N = Opt.); and above optimal N
application (N > Opt.), which included Opt. + 30 kg N ha–1;
Opt. + 60 kg N ha–1; 125, 130, and 150% of Opt.; and FNP.
The relationship between aboveground N uptake and grain
yield was fitted as linear, quadratic, and power models using
SigmaPlot 10 for Windows (www.sigmaplot.com). The power
model produced the best fit.

Table 2. Descriptive statistics of grain yield, harvest index (HI), concentrations of N ([N]) in grain and straw, total N uptake, and
nitrogen harvest index (NHI) at maturity of winter wheat.
Data set Parameter n† Mean SD Min. 25% Q Med. 75% Q Max.
All Yield, Mg ha–1 1395 6.7 2.2 1.6 5.3 6.5 8.3 11.8
HI 1395 0.45 0.05 0.28 0.42 0.46 0.49 0.58
[N] in grain, % 1395 2.11 0.27 1.15 1.95 2.12 2.28 3.04
[N] in straw, % 1395 0.53 0.15 0.11 0.44 0.53 0.62 1.05
N uptake, kg ha–1 1395 159 50 32 129 158 192 305
NHI 1395 0.77 0.06 0.54 0.74 0.77 0.81 0.94
Nreq., kg‡ 1395 23.7 3.3 14.4 21.7 23.6 25.7 37.5

N = CK Yield, Mg ha–1 356 5.8 2.5 1.6 3.6 5.7 7.8 11.4
HI 356 0.45 0.05 0.28 0.41 0.45 0.49 0.56
[N] in grain, % 356 1.91 0.22 1.22 1.78 1.92 2.06 2.63
[N] in straw, % 356 0.42 0.13 0.17 0.31 0.41 0.50 0.86
N uptake, kg ha–1 356 121 54 32 74 122 159 246
NHI 356 0.79 0.05 0.60 0.77 0.80 0.83 0.91
Nreq., kg 356 20.8 2.6 14.4 19.1 20.6 22.4 31.1

N < Opt. Yield, Mg ha–1 146 6.7 1.8 3.8 5.5 6.3 7.2 10.8
HI 146 0.47 0.04 0.33 0.44 0.48 0.50 0.58
[N] in grain, % 146 2.01 0.24 1.26 1.87 2.06 2.17 2.49
[N] in straw, % 146 0.56 0.13 0.20 0.46 0.57 0.65 0.93
N uptake, kg ha–1 146 152 39 68 129 145 169 269
NHI 146 0.76 0.06 0.57 0.73 0.76 0.80 0.91
Nreq., kg 146 22.8 2.5 16.7 21.1 22.9 24.1 29.3

N = Opt. Yield, Mg ha–1 429 7.2 1.9 3.2 5.8 6.7 8.6 11.8
HI 429 0.46 0.04 0.29 0.42 0.46 0.49 0.58
[N] in grain, % 429 2.18 0.20 1.43 2.06 2.18 2.30 2.80
[N] in straw, % 429 0.53 0.10 0.29 0.46 0.53 0.59 1.05
N uptake, kg ha–1 429 172 41 86 143 164 199 281
NHI 429 0.77 0.05 0.59 0.75 0.78 0.80 0.88
Nreq., kg 429 24.3 2.1 17.9 22.7 24.3 25.7 33.7

N > Opt. Yield, Mg ha–1 464 7.1 1.9 3.2 5.6 6.8 8.5 11.8
HI 464 0.45 0.05 0.32 0.42 0.46 0.49 0.58
[N] in grain, % 464 2.23 0.26 1.15 2.09 2.23 2.38 3.04
[N] in straw, % 464 0.61 0.13 0.11 0.52 0.60 0.71 1.03
N uptake, kg ha–1 464 179 41 86 149 175 206 305
NHI 464 0.75 0.06 0.54 0.72 0.75 0.79 0.94
Nreq., kg 464 25.7 3.3 17.8 23.4 25.2 27.5 37.5
† n = number of observations; SD = standard deviation; Q = quartile; Med. = median; Min. = minimum; Max. = maximum.
‡ Nreq.: The nitrogen requirement per megagram of grain.

Agronomy Journal • Volume 104, Issue 6 • 2012 1689

 RESULTS AND DISCUSSION
General Overview of the Data Set
Considering all 1395 data points that were collected among
all of the experiments, winter wheat grain yield (at 15% moisture)
ranged from 1.6 to 11.8 Mg ha–1, with an average of 6.7 Mg ha–1
(Table 2). The mean grain yield was 42% higher than China’s
average wheat yield of 4.7 Mg ha–1 and 123% higher than the
world’s average wheat yield of 3.0 Mg ha–1 in 2010 (FAO, 2011).
The HI ranged from 0.28 to 0.58, with an average of 0.45 (Table
2). Fifty percent of all observations had HI within a narrow range
of 0.42 to 0.49, which is in agreement with the 0.4 to 0.6 range
reported for modern varieties of most grain crops (Hay, 1995).
The average grain N concentration was 2.11%, while the straw N
concentration was 0.53%. However, the N concentration varied
tremendously in both grain (1.15–3.04%) and straw (0.11–1.05%)
due to the wide range of environmental and N supply conditions
(Table 2). This wide range in grain N concentrations was similar
to the 1.23 to 4% that was reported by Asseng et al. (2002). On
average, N uptake by aboveground plant dry matter was 159 kg
ha–1, and the grain NHI was 0.77 (Table 2).
Overall, the relationship between aboveground N uptake and
winter wheat grain yield could be described by a positive power
function, and 85.6% of the variation in aboveground N uptake
was explained by grain yield (Fig. 1a).The average N requirement to
produce 1 Mg winter wheat grain was 23.7 kg (range from 14.4 to
37.5 kg). This requirement was lower than the 24.6 kg estimated by
Liu et al. (2006). The difference may be attributable to the higher
winter wheat grain yield in this study compared to the study by
Liu et al. (2006), in which the winter wheat yield ranged from 0.35
to 8.73 Mg ha–1.
Grain Yield, Nitrogen Concentration, and Nitrogen
Requirement at Different Nitrogen Treatments
Average winter wheat grain yield was 5.8 Mg ha–1 (n = 356)
without N fertilizer, whereas average grain yield was 7.2 Mg ha–1
(n = 429) in the N = Opt. treatment (130 kg N ha–1 on average,
55 kg N ha–1 before planting and 75 kg N ha–1 at the stem
elongation stage). Treatments that were below Opt. (74 kg N ha–1
on average, 32 kg N ha–1 before planting and 42 kg N ha–1 at the
stem elongation stage) had a 7% yield reduction compared to the
N = Opt. treatment. On the other hand, adding more N fertilizer in
Fig. 1. Relationship of aboveground N uptake and winter wheat grain yield for (a) all N
treatments and (b) N = Opt. treatment. The solid line represents the relationship. The <4.5 Mg ha–1 and was stable at about 0.53%
dashed lines represent the prediction band (P = 0.95). *** Significant at the 0.001.
1690
the N > Opt. treatments (288 kg N ha–1 on average, 122 kg N ha–1
before planting and 166 kg N ha–1 at the stem elongation stage) did
not produce increased yield compared to the N = Opt. treatment.
Across all of the experiments, aboveground N uptake averaged
121, 152, 172, and 179 kg N ha–1 in the N = CK, N < Opt.,
N = Opt., and N > Opt. treatments, respectively (Table 2). For the
N = Opt. treatment, the overall relationship between aboveground
N uptake and winter wheat grain yield could be described by a
positive power function (Fig. 1b), and 89.3% of the variation in
aboveground N uptake was explained by grain yield. The curve
produced by the optimal N supply was higher than that for all of the
N treatments, which indicates that the aboveground N uptake–yield
relationship was affected by the application of N fertilizer.
For the N = Opt. treatment, the average grain yield was
7.2 Mg ha–1 with an average HI (0.46). Average Nreq. was 24.3 kg
and the N concentrations were 2.18% in grain and 0.53% in straw
(Table 2). When less N fertilizer was applied in the N < Opt.
treatment, Nreq. decreased from 24.3 to 22.8 kg and the N
concentrations declined by 8% in grain (from 2.18 to 2.01%) and
increased by 6% in straw (from 0.53 to 0.56%). For the N = CK
treatment, average Nreq. was 20.8 kg, which was the lowest value
observed across all of the N treatments. This was because it had the
lowest N concentrations in both grain and straw (1.91% in grain
and 0.42% in straw). With the addition of more N fertilizer in the
N > Opt. treatment compared to the N = Opt. treatment, Nreq.
increased from 24.3 to 25.7 kg because the N concentration had an
increase of 2% in grain (from 2.18 to 2.23%) and 15% (from 0.53
to 0.61%) in straw. This indicates that when excess N consumption
occurred, the excess N did not lead to greater grain yield or a greater
N concentration in the grain, but instead accumulated in the straw.
Similar results showed that under excess N application, the grain N
concentration reached a maximum and then remained stable, while
the straw N concentration continued to increase (Barneix, 2007).
Nitrogen Requirements for Different Winter
Wheat Grain Yield Levels Under Optimal
Nitrogen Application Conditions
To further clarify the aboveground N uptake–grain yield
relationship, all of the data from the N = Opt. treatment were
grouped into six yield ranges: <4.5 Mg ha–1 (n = 24, 4.0 Mg ha–1
on average), 4.5 to 6.0 Mg ha–1 (n = 103, 5.3 Mg ha–1 on
average), 6.0 to 7.5 Mg ha–1 (n = 147,
6.7 Mg ha–1 on average), 7.5 to 9.0 Mg ha–1
(n = 64, 8.1 Mg ha–1 on average), 9.0 to
10.5 Mg ha–1 (n = 66, 9.8 Mg ha–1 on
average), and >10.5 Mg ha–1 (n = 25,
11.0 Mg ha–1 on average) (Table 3). Using
all of the data for the N = Opt. treatment,
the average HI values were 0.39, 0.44, 0.46,
0.46, 0.47, and 0.48 for the six wheat grain
yield ranges, respectively (Fig. 2a). Average
NHI remained stable at about 0.77 for
all of the yield ranges (Fig. 2b). The grain
N concentrations were 2.41, 2.25, 2.21,
2.13, 2.00, and 2.06% for the six yield
ranges, respectively (Fig. 2c); the straw N
concentration was 0.47% for the yield range
for the other five yield ranges (Fig. 2d).
Agronomy Journal • Volume 104, Issue 6 • 2012
Table 3. Descriptive statistics of grain yield (Mg ha –1) of the N = Opt. treatment at different yield ranges.
Yield ranges n† Mean SD Min. 25% Q Med. 75% Q Max.
<4.5 24 4.0 0.4 3.2 3.9 4.1 4.3 4.5
4.5–6.0 103 5.3 0.4 4.5 5.0 5.3 5.7 6.0
6.0–7.5 147 6.7 0.4 6.0 6.3 6.6 7.0 7.5
7.5–9.0 64 8.1 0.5 7.5 7.7 8.1 8.6 8.9
9.0–10.5 66 9.8 0.4 9.0 9.6 9.9 10.2 10.5
>10.5 25 11.0 0.3 10.5 10.8 11.1 11.3 11.8

Total 429 7.2 1.9 3.2 5.8 6.7 8.6 11.8

† n = number of observations; SD = standard deviation; Q = quartile; Med. = median; Min. = minimum; Max. = maximum.

Average Nreq. was 27.1, 25.0, 24.5, 23.8,

22.7, and 22.5 kg for the six wheat grain
yield ranges, respectively (Fig. 3).
In this study, decreases in the N
requirement of winter wheat in response to
increases in grain yield could be classified
into three response stages. For yield ranges
from <4.5 to 6.0 to 7.5 Mg ha–1, increases
in grain yield were mainly caused by both
improved HI values and higher aboveground
biomass. Yield increased by 68% (from
4.0 to 6.7 Mg ha–1) when aboveground
biomass increased by 39% (from 8.9 to
12.5 Mg ha–1) and HI increased by 18%
(from 0.39 to 0.46). The Nreq. decreased
from 27.1 to 24.5 kg due to the increase
in HI from 0.39 to 0.46 and the decrease
in grain N concentration from 2.41% to
2.21%. For the yield range from 6.0 to 7.5
to 9.0 to 10.5 Mg ha–1, the increase in grain
yield mainly resulted from the increase in
aboveground biomass and the maintenance Fig. 2. (a) Harvest index, (b) N harvest index, (c) grain N concentration, and (d)
straw N concentration of the N = Opt. treatment at maturity of winter wheat for
of a stable HI of 0.47. Nreq. decreased from different yield ranges in the northern China. Solid and dashed lines in this figure
24.5 to 22.7 kg due to a decrease in grain indicate median and mean, respectively. The box boundaries indicate the 75 and 25%
quartiles, the whisker caps indicate 90th and 10th percentiles, and the circles indicate
N concentration from 2.21 to 2.00%. the 95th and 5th percentiles.
For the yield range from 9.0 to 10.5 to
>10.5 Mg ha–1, Nreq. changed little because
the grain N concentration and HI were stable.
In this study, for the N = Opt. treatment, the average grain
yield was 7.2 Mg ha–1 and the N requirement per Mg grain ranged
from 17.9 to 33.7 kg, with an average of 24.3 kg. However, this
N requirement was greater than the value (19.8 kg) for winter
wheat in the UK (Barraclough et al., 2010) and was smaller than
the value (25.9 kg) reported in Mexico (Ortiz-Monasterio et
al., 1997). This is because the grain yield was greater in the UK
(9.27 Mg ha–1) and lower in Mexico (6.4 Mg ha–1). Our results
show that the N requirement decreased from 27.1 kg with an
average grain yield of 4.0 Mg ha–1 to 22.5 kg, with an average grain
yield of 11.0 Mg ha–1. Hou et al. (2012) found similar trends,
whereby N requirements decreased with increasing grain yield in
spring maize.
In our study, the N requirement declined with increasing
grain yield because of an increase in HI and a decrease in the N
concentration, especially in the grain N concentration. An inverse Fig. 3. Nitrogen uptake per Mg wheat grain of the N = Opt.
relationship has been reported between grain N concentration treatment at different yield ranges in the northern China.
Solid and dashed lines in this figure indicate median and mean,
and grain yield in winter wheat (Triboi et al., 2006; Barneix, respectively. The box boundaries indicate the 75% and 25%
2007; Bogard et al., 2010). Grain N concentration or grain protein quartiles, the whisker caps indicate 90th and 10th percentiles,
concentration is a critical trait that determines the nutritional and the circles indicate the 95th and 5th percentiles.

Agronomy Journal • Volume 104, Issue 6 • 2012 1691

value, processing properties, and market value of wheat grain.
Therefore, one critical question is how wheat grain yield can be
increased while maintaining acceptable grain N concentrations.
Our results from the N = Opt. treatment show that average
HI increased from 0.39 in the <4.5 Mg ha–1 yield range to 0.48
in the >10.5 Mg ha–1 yield range. These values were lower than
the HI value (0.55) for winter wheat in the UK (Barraclough et
al., 2010). The difference in HI between northern China and
the UK may be caused by differences in climatic conditions. The
climate in northern China is continental, which includes cold
winters and hot summers. In comparison, the climate in the UK
is oceanic, which includes moderately cool summers and relatively
warm winters, with a narrow annual temperature range. High
temperatures in late May limit grain filling; temperatures over
30°C are common and the maximum temperature can be as high
as 40°C in northern China (Porter and Gawith, 1999; He et al.,
2001). Despite the fact that the highest HI value (0.48) was lower
than the theoretical maximum value of 0.64 for winter wheat
(Foulkes et al., 2011), the scope for further improvements in HI
may be limited in wheat. For example, the highest HI value was
0.53 for winter wheat cultivars released in France from 1946 to
1992 (Brancourt-Hulmel et al., 2003), while the highest HI value
was 0.50 for winter wheat cultivars released in northern China
from 1960 to 2000 (Zhou et al., 2007). To further increase the
yield potential of winter wheat, particular attention should be
given to dry matter production, especially postanthesis, while
maintaining HI at values above 0.5 (Foulkes et al., 2011; Reynolds
et al., 2011; Ye et al., 2011).
In this study, the grain N concentration decreased from
2.41% in the < 4.5 Mg ha–1 yield range to 2.00% under the 9.0
to 10.5 Mg ha–1 yield range. The negative relationship between
grain N concentration and grain yield in winter wheat had been
well established (Triboi et al., 2006; Barneix, 2007; Bogard et
al., 2010). The physiological basis of this negative correlation can
be explained by two hypotheses: (i) competition between C and
N for energy; and (ii) an N dilution effect caused by C-based
compounds (Bogard et al., 2010). The degree of this inverse
relationship varies depending on genetic and environment factors
that include water and N availability (Triboi et al., 2006). One
way to possibly counteract this negative correlation is to transfer
genes for high grain protein concentrations from related species.
For example, a gene designated Gpc-B1, which is not functional
in modern wheat cultivars, was recently identified in the wild
emmer wheat (Triticum turgidum ssp. dicoccoides) (Distelfeld et
al., 2006; Uauy et al., 2006a). Functional Gpc-B1 introgression has
the potential to improve grain N concentration through a more
efficient remobilization of N in straw. However, functional Gpc-B1
introgression could be associated with reduced grain weight in
some varieties (Uauy et al., 2006b; Brevis et al., 2010; Brevis
and Dubcovsky, 2010; Kumar et al., 2011; Carter et al., 2012).
Therefore, a dedicated breeding effort is required to counter this
potential negative effect of the Gpc-B1 gene.
An alternative approach to counteract this negative correlation
is to change agronomic management practices. Experimental and
modeling results have shown that wheat grain N concentrations
are mostly source-determined (Martre et al., 2003). To increase
grain N concentrations, we need to increase the size of the grain N
source. For example, the N stored in vegetative organs at anthesis
and N uptake from the soil during the postanthesis period (Triboi
1692
et al., 2006; Foulkes et al., 2009). Several studies have reported
that the addition of postanthesis N through dry fertilizer material
or foliar N application can increase the grain N concentration
without reducing yield (Bly and Woodard, 2003; Dupont et al.,
2006). Therefore, we can increase both the level and stability of
wheat and grain N concentrations through wheat breeding and
better management practices.
CONCLUSIONS
Uncertainties in estimates of wheat N demand, such as among
regions, N supplies, and wheat grain yield levels, have affected the
development of an optimal N management index. A database
comprising 1395 measurements collected from sites in which
grain yield ranged from 1.6 to 11.8 Mg ha–1 was used to quantify
N requirements. Average N requirement increased with N supply.
Excess N application did not lead to greater grain yield or greater
grain N concentration; instead, N accumulated in the straw. A
decrease in the N requirement per Mg grain yield was observed
with increasing grain yield, which was attributable to an increasing
HI and dilution effects from declining grain N concentrations.
The average N requirement in the N = Opt. treatment was
24.3 kg, which was greater than the value (19.8 kg) for winter
wheat in the UK and smaller than the value (25.9 kg) in Mexico.
In practice, to accurately estimate the N requirements of crops,
we should take into account the effects of different N supplies and
grain yield levels.
ACKNOWLEDGMENTS
We thank the National Basic Research Program of China (973,
Program: 2009CB118606), Special Fund for Agro-scientific
Research in the Public Interest (201103003), the Innovative
Group Grant of National Natural Science Foundation of China
(30821003) and the Key Project of Twelfth Five-year National
Plan (2011BAD11B05).
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