Field Crops Research 167 (2014) 1930

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Field Crops Research

journal homepage:www.elsevier.com/locate/fcr

Growth, yield, and nitrogen use in the wheat/maize intercropping system in

an arid region of northwestern China
a,b a,b,c, a,b,c b,c
Ying Gao , Pute Wu , Xining Zhao , Zikui Wang
a b
Institute of Soil and Water Conservation, University of Chinese Academy of Sciences and Ministry of Water Resources, Yangling 712100, China Institute of
Water Saving Agriculture in Arid Regions of China, Northwest A&F University, Yangling 712100, China
c
College of Water Resources and Architectural Engineering, Northwest A&F University, Yangling 712100, China
article info abstract

Article history: Relay intercropping of spring wheat and maize is practiced on a large scale in northern China. In order to provide information
Received 5 April 2014 on productivity and nitrogen use for spring wheat/maize intercropping, a two-year experiment under three N levels and three
Received in revised form 30 June 2014 cropping systems (monocultured wheat, monocultured maize, and intercropped wheat and maize) was carried out in the Hetao
Accepted 5 July 2014
Irrigation District of the Inner Mongolia Autonomous Region, to quantify and characterize the growth and yield of the wheat
Available online 30 July 2014
and maize and determine the uptake and use efficiency of nitrogen in two crops.

Keywords:
The land equivalent ratio (LER) of the wheat/maize intercropping system ranged from 1.01 to 1.19, and thus land use
Land equivalent ratio (LER)
efficiency was significantly enhanced by this intercropping system. The dry-matter yield and LAI (leaf area index) of wheat
Expolinear growth equation
Growth delay were higher in border rows than in inner rows and monocultured rows, indicating an effect of border rows in the intercropping
Internal N use efficiency (IE) system. The LAI was considerably lower for the intercropped than the monocultured maize, indicating that intercropping
Relative N yield total (RNT) limited the vegetative growth of maize.

The grain yield and nitrogen uptake of both wheat and maize in two cropping systems were significantly improved with
increasing amounts of N application. The LER of intercropping system tended to increase at higher N levels.

The patterns of maize growth in the monocultures and intercrops were characterized by fitting expo-linear growth equations
to periodic harvest data. Fitted parameters indicated a growth delay in the intercropped maize ranging from 2.39 to 8.24 d
relative to the maize monoculture. The growth delays were shortest at the moderate N rate among three N levels (2.39 d in
2012 and 2.57 d in 2013). This result could be interpreted in terms of the competitive effect of wheat during the seedling phase
of maize, which was weakest at the moderate N rate, causing a comparatively short growth delay. The estimated growth rate
1 1
during the linear growth phase was highest in the intercrop in the high N treatment (417.5 kg ha d in 2012 and 376.7 kg
1 1
ha d in 2013).

The physiological nitrogen use efficiency (IE) was more favorable for monocultured wheat and inter-cropped maize and
decreased as N rates increased. The efficiency of the use of N for the entire system was assessed by comparing the relative
nitrogen yield total to the relative yield total. The relative nitrogen yield total ranged from 1.06 to 1.23, while the relative yield
total ranged from 1.01 to 1.19, indicating that the intercrop used more nitrogen per unit production than did the monocultures.

2014 Published by Elsevier B.V.

1. Introduction
A growing population and rising dietary levels are driving global demand
for increasing agricultural production while arable land
grain production in northwestern China, especially in areas with irrigation but
Corresponding author at: College of Water Resources and Architectural Engi-neering,
Northwest A&F University, No. 3 Taicheng Road, Yangling 712100, Shanxi, China. Tel.: +86 0
029 87012411; fax: +86 0 029 87012210.
E-mail address: gjzwpt@vip.sina.com (P. Wu).
is being lost to urbanization and degradation (Gao et al., 2010). Crop yields
need to be increased by improving the use efficiency of resources such as
water and nutrients (Awal et al., 2006), while meeting the demand for diverse
agricultural commodities. Inter-cropping is often a means to better use of land
and other resources (Rodrigo et al., 2001). Wheat-based intercropping, such as
inter-cropping wheat and maize, is a long-established and major system of
with only one growing season annually due to limi-tations of temperature (Li
et al., 2001).

http://dx.doi.org/10.1016/j.fcr.2014.07.003 0378-
4290/ 2014 Published by Elsevier B.V.
20 Y. Gao et al. / Field Crops Research 167 (2014) 1930

In wheat/maize intercropping
systems, spring wheat is sown in
March interspersed with strips of
bare soil. Maize is then sown in the
open spaces in late April. The two
crops grow together for
approximately 10 weeks before the
wheat is har-vested in July. The
maize can then use the entire field,
both above- and belowground.
Interspecific competition will occur
when two crops are grown together
(van der Meer, 1989), and such
competition usually decreases
survival, growth, or repro-duction
of at least one crop (Crawley,
1997). The two crops frequently
interact at their boundaries, which
can lead to increases or decreases in
growth, development, and even
yields (Zhang and Li, 2003). For
example, the yield from maize
border rows in maize/soybean
intercropping increased at the
expense of the yield of border-row
soybeans (Iragavarapu and Randall,
1996). Interspecific competition is
common, but intercropping
generally offers a yield advantage
due to complementation or
facilitation between the component
crops, including maize/soybean
(West and Griffith, 1992;
Ghaffarzadeh et al., 1994),
sorghum/soybean (Elmore and
Jackobs, 1986), wheat/mungbean
(Mandal et al., 1996),
cotton/groundnut (Waterworth,
1994), wheat/medic (Moynihan et
al., 1996), and wheat/cotton
systems (Zhang et al., 2007), etc.
Wheat/maize intercropping
systems in China are becoming
increasingly popular in the irrigated
area of the Hexi Corridor in Gansu
Province, irrigated areas along the
Huanghe River in Ningxia, and in
the Inner Mongolia Autonomous
Region, China (Zhang and Li,
2003). Farmers have continually
applied an excess of fertilizer in
recent years to achieve high yields
in local agricultural produc-tion.
The application rates of nitrogen
1
(N) average 450 kg N ha in the
irrigated area of northwestern China
(Li et al., 2005), and about 3050%
of the N is leached into the
groundwater (Zhang, 1987).
Improving land use efficiency
benefits for N (protein) yield in
inter-crops could thus not only be
economic importance to farmers,
but also can be beneficial to the
environment. The use of N
fertilizers on this subject in
intercropping thus has received
much concern and needs to be
further investigated in the current
practice.
Efficiency of N use in each
component crop is diagnosed by N
use efficiency (NUE) and is
commonly measured by internal (or
physiological) efficiency (IE),
expressed as kg yield per kg N
uptake (Haefele et al., 2003;
Bronson et al., 2001). Several
studies have shown that intercrops
use soil nutrients more efficiently
than do monocultured crops
because of a higher recovery of N
and increased yields of dry matter
(Hauggaard-Nielsen et al., 2001;
Zhang et al., 2004; Inal et al., 2007;
Li et al., 2004). However,
agronomic N-use efficiency may
also be decreased by intercropping.
Aggarwal et al. (1992), for
example, found that upland rice
intercropped with mungbeans took
up the same amount of N as a
monoculture but produced
significantly less dry matter. Zhang
et al. (2008a) showed that the IE of
cotton in a wheat/cotton
intercropping system was reduced
due to delayed flowering and less
reproductive growth.
A modification of the land
equivalent ratio (LER) (Willey,
1985), expressed as the relative N
yield total (RNT) (Baumann et al.,
2001) or the LER for N yield
(Szumigalski and Van Acker, 2006),
can be used to determine the land-
use advantage of intercropping in
terms of N yield. The RNT is
calculated as the sum of the ratios
of N uptakes by the component
crops in the intercrop to their
respec-tive N uptakes in
monoculture. A value of RNT
exceeding the LER suggests that
intercropping does not use N
efficiently (Zhang et al., 2008a).
Wheat/maize relay strip
intercropping systems are widely
prac-ticed in China, but information
on N uptake and use efficiency is
rather incomplete. More data are
needed on crop productivity and on
N requirements and use to provide a
scientific basis for recom-mending
this practice for large-scale use. The
main objectives of this study were
thus: (1) to quantify the yield and
growth of the component crops, (2)
to describe and characterize the
accumula-tion of dry matter and
determine the differences of growth
over
The field experiment was
arranged in a two factorial
randomized complete block design
with three replications. The
cropping sys-tems and N treatments
were completely randomized within
each block. The cropping systems
were: monocultured wheat (MW),
monocultured maize (MM), and
intercropped wheat and maize
(intercropped wheat (IW) and
intercropped maize (IM)). Each
com-ponent crop was fertilized with
three levels (N0, N1, and N2) of N:
1
0, 120, and 240 kg N ha for wheat
1
and 0, 180, and 360 kg N ha for
maize, respectively.
Fig. 1. Layout of the
wheat/maize intercropping
system. The intercrops were oriented
northeast-southwest in strips 1.7 m
in width, with each strip consisting
time among systems, and (3) to
of six rows of wheat 0.15 m apart
quantify N uptake and compare
adjacent to two rows of maize 0.4 m
responses of N-use efficiency to N
apart (Fig. 1). The distance between
fertilization (at crop and system
adjacent wheat and maize rows was
levels). We hypothesized that
27.5 cm. The wheat and maize
intercrops could exploit N resources
occupied 53% and 47% of the
more efficiently than monocultures.
intercropped area, respectively. The
Our findings could potentially be
monocultures of both crops had the
valuable for designing systems that
same dis-tances between rows. Each
require less N input and for
plot was 5 m wide and 10 m long,
modeling cropping systems.
and all plots were separated by a
2.5-m buffer zone.
2. Materials and methods The cultivar of spring wheat
(Triticum aestivum L.) was Yong-
2.1. Experimental site liang 4, and the cultivar of maize
The field experiments were (Zea mays L.) was Nei-dan
conducted at the Shahaoqu Experi- 314. Both were commonly
mental Station (40 54 N, 107 09 E; used on farms. Wheat was
1036 m a.s.l.) in 2012 and 2013.
The two experiments were
seeded at a density of 6
conducted in different but nearby
fields. The station lies in the west of
the Hetao Irrigation District in the
Inner Mongolia Autonomous
Region, which has a typical
continen-tal climate. The average
annual rainfall is 139222 mm,
average annual temperature is 9.1
C, and mean potential evaporation
is 20002300 mm. This area is also
in the upper reaches of the Yel-low
River. The soils are alluvial silt
sediments, with an upper layer (0
60 cm) of silty loam and a lower
layer (60150 cm) of sandy loam.
The soil begins to freeze by the
middle of November and does not
thaw completely until late April or
early May (Xu et al., 2010). Soil
samples were collected at the start
of the experiments. The physical
and chemical properties of the soil
are listed in Table 1.
2.2. Experimental design
 Y. Gao et al. / Field Crops Research 167 (2014) 1930 21
Table 1

Soil properties at the experimental field.

Soil depth (cm) 3 1 1 1 1

Bulk density (kg dm ) Available N (mg kg ) Available P (mg kg ) Available K (mg kg ) Organic C (g kg )
030 1.492 105.3 55.9 120.0 9.8
3060 1.435 74.1 16.8 78.7 4.5
60100 1.512 51.5 5.6 67.5 2.5

Table 2
Irrigation date and irrigation amount (mm) in 2012 and 2013.

2012 2013
Date IW/M MW MM Date IW/M MW MM
8-May 50 50 2-May 75 75
28-May 75 60 25-May 75 75
17-Jun 62.5 62.5 62.5 20-Jun 75 75 75
25-Jul 50 50 30-Jul 75 75
28-Aug 62.5 62.5 31-Aug 60 60
Total 300 172.5 175 Total 360 225 210

Notes: MW, monocultured wheat; MM, monocultured maize; IW/M, intercropped wheat and
maize.

2
67 plants m , and maize was thinned one week after emergence to a density
2
of 8.33 plants m . The monocultured and intercropped treatments had the
same densities. Wheat was sown on 25 March in 2012 and on 19 March in
2013 and was harvested on 19 July in 2012 and on 15 July in 2013. Maize
was sown on 23 April in 2012 and on 21 April in 2013 and was harvested on
Fig. 2. Rainfall and mean air temperature (T) distribution during the two growing seasons. The
21 September in 2012 and on 20 September in 2013. letter F indicates the flowering of maize.

The N and phosphorus (P) fertilizers for both cropping systems were urea
(46% N) and triple superphosphate (44% P 2 O5 ), respec-tively. Full doses of
P and half doses of N were evenly broadcast before seeding to the
monocultured and intercropped wheat, and the remaining 50% N was top-
dressed at the first irrigation. Full doses of P and half doses of N were
incorporated into the soil of the monocultured and intercropped maize, and
the remaining 50% N was applied equally at the elongation and pre-tasseling
stages with the corresponding irrigations.
2.3. Crop management
Before preparation of the experimental land, 180 mm autumn water in
October was applied in the last year. The amount and date of irrigation during
growing season are presented in Table 2. Weeds were controlled equally in all
treatments by hoeing as needed. No pesticides were applied to either crop.
Climatic data during the two growing seasons (MarchSeptember) are
given in Fig. 2. Total precipitation was 202 and 100 mm in 2012 and 2013,
respectively. Despite the difference in total rainfall between years, higher
levels of irrigation in 2013 compensated for the lower rainfall. The
distributions of temperature and rainfall, however, differed greatly between
years, which could contribute to the differences in the results (see below).
2.4. Measurements
plants were separated into leaves, stems, and ears. All samples were then dried
at 65 C in an oven to constant weight to determine the dry matter (DM).
Grain yields were determined by harvesting at maturity from 1 m of outer
and inner rows of intercropped and monocultured wheat. The grains were sun-
dried and weighed after threshing by hand. Grain yield was determined for
each plot from the grain weight at a moisture content of 12%. The grain yields
of maize were determined by harvesting all plants at maturity within a
sampling area of 3 m row length 0.8 m width (two rows) at the center of
each plot. The kernel number per ear was then counted and grain yield of
maize was based on accounting for 13% water content in the sun-dried grain.
The LAIs of wheat and maize were determined from the same samples as
used for determining DM weight. The number of plants per meter was
2
counted. The total leaf area per plant was deter-mined, and the LAI (m leaf
2
area per m ground area) was calculated as the product of plant density (plants
2 2
m ) and leaf area per plant (m ). The area of an individual leaf for both
wheat and maize was determined as leaf area = leaf length leaf width
coefficient. Leaf length and the widest part of leaves were measured with a
ruler. The coefficient was determined using an area meter (model LI-3100, Li-
Cor, Inc., Lincoln NE) and was estimated as the slope of a regression through
the origin, using measured leaf area as the y variable and the product of length
and width of the leaf as the x variable (Zhang et al., 2008b). The coefficients
were 0.835 for wheat and 0.70 for maize.

Samples of aboveground dry matter were collected from 1-m sections of
outer and inner rows of each plot of intercropped and monocultured wheat at
intervals of 1014 d after emergence. The number of plants was counted to
determine the plant densities of the rows, and then a subsample of 20
randomly selected plants was collected for further analyses. Samples of
intercropped and mono-cultured maize were collected based on plant number,
10 seedlings during early growth and three plants subsequently. Leaf areas
and plant heights were measured for both crops. Then the wheat sam-ples
were separated into leaves, stems, and spikes, and the maize
Plant samples were weighed and ground to pass through a 0.25-mm sieve
prior to nutrient analysis. N contents in the plant dry matter were determined
by a micro-Kjeldahl method after diges-tion in a mixture of concentrated H 2
SO4 and H2 O2 . Blanks were also analyzed to detect any sources of
contamination. Nutrient acquisi-tion was calculated as the product of nutrient
content and biomass in the aboveground tissues of the crops.
Gravimetric soil water samples were collected by auger at the center of
two maize rows in intercropping and monoculture, then oven-dried to measure
the soil water content.
22 Y. Gao et al. / Field Crops Research 167 (2014) 1930

where YMW and YIW are grain yields of monocultured and inter-cropped
1
wheat (kg ha ), respectively, and YMM and YIM are grain yields of
1
monocultured and intercropped maize, respectively (kg ha ). The LER is
similar to relative gain yield total, i.e. the sum of the yields of the two
component crops in the intercrop, each scaled by its yield in the monocultures.

The NHI is frequently used to assess the transport of N from shoots and
leaves to the grains in crops. The NHIs of wheat and maize were calculated as
(Li et al., 2011a):

where Ngrain and Naboveground biomass are the N uptakes by grains and crop
biomasses, respectively, in the intercropped and mono-
cultured treatments.
IE (Internal efficiency of nitrogen use) is defined as (Haefele et al., 2003):

where RNW and RNM are the relative N yield for wheat and maize,
respectively. NIW , NMW , NIM , and NMM represent N uptake of intercropped
and monocultured wheat and of intercropped and monocultured maize,
respectively.
To model the growth pattern of intercropped and monocultured maize,
expolinear equations (Goudriaan and Monteith, 1990) were fitted using non-
linear least squares regressions:

1
where Wt is the dry mass (kg ha ) at t days after seeding (DAS), c m is the
1 1
maximum absolute growth rate (kg ha d ), rm is the initial relative growth
1
rate (d ), and tb is the time (d) at which the growth rate is c m /2. The time
course of the growth rate is logistic with the growth parameters r m and cm .
The parameter rm repre-sents the initial exponential rate of growth of the maize
plants. The parameter tb represents the time at which the exponential growth
goes over into linear growth, and c m represents the rate of linear growth when
the crop has attained maximum soil cover.
The parameters were estimated for each treatment and year fol-lowing a
two-step procedure (Zhang et al., 2007). In step 1, the rm and tb were estimated
using a logarithmic transformation of the observations and of Eq. (5).
Logarithmic transformation empha-sizes the early observations and provides
more accurate estimates of the parameters shaping the early growth, r m and tb .
In step 2,
2.5. Calculations and data analysis Table 3
1
Yields of the treatments in 2012 and 2013 (t ha ).
The effects of the treatment on yield, LER, N harvest index (NHI), N Treatment 2012 2013
content, and total N uptake were determined with analyses of variance Wheat Maize LER Wheat Maize LER
(ANOVAs) in SPSS 17.0 (SPSS Institute Ltd., USA). The data were tested for
N0-MW 6.13 5.17
homogeneity of variance before the statistical anal-ysis. Significant N1-MW 6.93 6.06
differences between treatments were determined using least significant N2-MW 7.00 6.08
difference tests at the 0.05 level. N0-MM 16.50 13.80
We used the LER (land equivalent ratio) to evaluate the yield performance N1-MM 19.00 14.40
N2-MM 19.81 14.45
of strip intercropping and monoculturing as (Willey, 1985): N0-IW/M 6.80 16.43 1.06 6.32 10.66 1.01
N1-IW/M 7.42 18.90 1.03 6.63 14.35 1.05
LER = Y + Y (1) N2-IW/M 7.87 21.56 1.11 8.03 15.32 1.19
IW IM
N application * ** ** **

Y Y Planting * NS ** *
MW MM *
NP NS NS NS
Notes: MW, monocultured wheat; MM, monocultured maize; IW/M, intercropped wheat and
1
maize; N0, N1, and N2 represent 0, 120, and 240 kg N ha for wheat and 0, 180, and 360 kg N
1 *
ha for maize, respectively; NS, not significant at the 0.05 level; , significant at the 0.05 level;
**
, significant at the 0.01 level. The values for the intercrops are on an equivalent basis of
comparable land area of the monocultures.
 N
NHI = grain
N rm and tb were fixed at their estimated values, and cm was esti-mated using
aboveground biomass
Yi untransformed observations and the untransformed Eq. (5). The use of
IEi =
untransformed data in the second step provides
Ni

2
(2) a better estimate of the linear growth rate c m . R and root mean square error
obtained in the second step are reported. An ANOVA with cropping system and
year as main effects was then conducted for each parameter to determine the
significance of the differences between treatments and years in the estimated
parameters.
3. Results
(3) 3.1. Grain yield
1 Due to a significant year effect for the variables evaluated, the effect was
where Yi is the grain yield of crop i (kg harvestable product ha ), and Ni is
1 discussed separately for each year (Table 3).
the N uptake (kg total N uptake ha ) by crop i.
RNT (Relative N yield total) was calculated as (Baumann et al., 2001):
3.1.1. Wheat
RNT = RNW + RNM = N + N (4) Wheat yields were significantly different between the intercrop and the
IW IM
The grain
monoculture in 2012 (P < 0.05) and 2013 (P < 0.01) (Table 3).
N N
MW MM yields were 10.1% and 21.2% higher in the intercrops than in
the monoculture in 2012 and 2013, respectively. No
interaction between N application and cropping system was
found in either year. Grain yields were 11.0% and 15.0%
higher in 2012 and 10.4% and 22.8% higher in 2013 at N1 and
N2, respectively, relative to N0.
cm ln [1 + exp [rm (t tb )]]
3.1.2. Maize
Wt = rm The effect of intercropping on grain yield varied with the rate of N
application. The 2013 grain yields were higher in the inter-
(5) cropped than in the monocultured maize at N2 but lower at N1 and N0. The
2012 yields did not differ significantly between the intercropped and
monocultured maize. N fertilization significantly enhanced the grain yields of
maize (P < 0.01). Grain yields in 2012 were 15.0% and 31.2% higher at 180
1
(N1) and 360 kg N ha (N2), respectively, relative to N0 for intercropped
maize and 15.2% and 20.1% higher, respectively, for monocultured maize.
Grain yields in 2013 were 34.6% and 43.7% higher for intercropped maize
and 4.3% and 4.7% higher for monocultured maize at N1 and N2,
respectively.

3.1.3. LER
The LER of the intercrops varied from 1.03 to 1.11 in 2012 and from 1.01
to 1.19 in 2013 (Table 3).
The LERs for all intercropped treatments
were equal to or higher than unity over both years, indicating
that wheat/maize intercropping improved the land use
efficiency in this experiment. The LERs were significantly
different between N2 and the other rates but not between N1
and N0. The
Y. Gao et al. / Field Crops Research 167 (2014) 1930 23
values of LER in both years were highest at N2 (1.11 in 2012 and 1.19 in
2013).
3.2. Crop growth
3.2.1. Dynamics of the aboveground dry biomass
3.2.1.1. Wheat. The border row had a significantly higher accu-mulation of
dry matter than did inner rows and the rows of monocultured wheat from 57
DAS until maturity in both years (P < 0.05; Fig. 3), indicating an interactive
effect in the intercrop-ping system. From 57 to 117 DAS, the dry-matter
accumulation in 2012 for the N2, N1, and N0 treatments was 27.8%, 41.3%,
and 29.5% higher in the border row than in inner rows, respectively, and
15.9%, 14.4%, and 54.3% higher than in the rows of monocultured wheat,
respectively (Fig. 3AC). The corresponding accumulations of dry matter in
2013 were 28.8%, 23.0%, and 28.7% higher in the border row than in inner
rows, respectively, and 29.0%, 22.3%, and 42.6% higher than in the rows of
monocultured wheat, respectively (Fig. 3DF).
3.2.1.2. Maize. The comparison of the dry-matter weight of indi-vidual
maize plants between the intercrops and monocultures at 63 DAS and 112
DAS in the three N application treatments in 2012 is shown in Fig. 4A and B.
At 63 DAS, before the wheat harvest, biomass accumulations were
significantly different between monocultured and intercropped maize (Fig.
4A; P < 0.05). The dry weights of indi-vidual maize plants were 23.39, 3.35,
and 9.97 g higher in the monoculture than the intercrop at N2, N1, and N0,
respectively. The reduction amplitude in the intercrop was considerably lower
at N1 relative to N2 and N0. The growth of maize during the co-growth stage
in the intercrops was thus severely suppressed, indicating competition
between the two crops. And competitive effects were probably weakest at N1
where the degree of the reduction in dry weight between the intercrop and
monoculture was lowest. At 112 DAS, after the wheat harvest, the effect of
intercropping on the biomass accumulations varied among the N treatments
(Fig. 4B).
The dry weights of intercropped maize plants exceeded those of monocultured
plants at N2 but were lower than those of mono-cultured plants at N1 and N0.
These findings indicate that nutrient supply played an important role in the
compensation of growth in the intercropped maize. The year of 2013 also had
the above similar trends (data not shown).
The growth pattern of maize for the entire 2012 growing sea-son is
presented in Fig. 4C. Dry-matter accumulation in maize had attained a
constant rate at the harvest time of wheat. No further dry matter of maize
accumulated during the last two weeks before har-vest. The growth patterns of
maize among the various N treatments in 2013 were similar to those shown
for 2012.
3.2.2. Expolinear growth curves of maize
Due to a significant year effect (P < 0.01) on aboveground dry matter for
maize, expolinear growth equations (Eq. (5); Fig. 5AF) were fitted to the
data from each treatment in each year to char-acterize the growth of maize in
three parameters: rm , the initial relative rate of growth during the exponential
growth phase of the seedlings; c m , the linear growth rate of a mature canopy
after max-imum light interception is attained; and t b , the time of transition
from the exponential to the linear growth phase (Zhang et al., 2007). All fitted
2
curves for each year are shown in Fig. 5G and H. R of the fitted equations
ranged from 0.923 to 0.997, indicating that the expolinear curve was suitable
for describing the accumulation of dry matter in both intercropped and
monocultured maize (Table 4).
The fitted values of rm for total dry matter were significantly different (P <
0.05) between IM and MM in both years, except for the N1 treatments. The
time of transition from exponential to lin-ear growth was earliest in the N2-
MM treatments (52.90 and 54.78 DAS for 2012 and 2013, respectively) and
latest in the N0-IM treat-ments (65.51 and 67.82 DAS for 2012 and 2013,
respectively). The time of transition from exponential to linear growth in MM
was sig-nificantly earlier than in IM at all N rates over both years (P < 0.05),
suggesting that the time course of dry-matter accumulation in the intercrops
was considerably delayed relative to the monocultures.
Fig. 3. Aboveground biomass in rows of wheat in the intercrops and monocultures at various rates of nitrogen fertilization in 2012 and 2013. (A) N2 in 2012, (B) N1 in 2012,
(C) N0 in 2012, (D) N2 in 2013, (E) N1 in 2013, and (F) N0 in 2013. The values of the aboveground biomass were calculated on an equivalent basis of comparable land area to the monocultures.
24 Y. Gao et al. / Field Crops Research 167 (2014) 1930

Fig. 4. Comparison of the dry-matter weights of individual maize plants between the intercrops and monocultures at the various rates of nitrogen fertilization at (A) 63 d after seeding (DAS) and (B)
112 DAS in 2012. Dynamics of aboveground dry biomass of maize in the various treatments in 2012 (C). IM and MM represent intercropped and monocultured maize, respectively. The values for the
intercrops were calculated on an equivalent basis of comparable land area to the monocultures. The arrow indicates the harvest time of wheat (H W ).

Table 4
Parameters of the expolinear growth equation for the growth of aboveground dry matter in intercropped and monocultured maize at the various rates of nitrogen fertilization in 2012 and 2013.

Year Rate Cropping system 1 1 1 2

rm (d ) tb (DAS) cm (kg ha d ) R
2012 N2 IM 0.450b 57.20a 417.5a 0.981
MM 0.498a 52.90b 387.5b 0.993
N1 IM 0.424a 56.88a 341.7b 0.982
MM 0.435a 54.49b 374.2a 0.970
N0 IM 0.338b 65.51a 373.3a 0.923
MM 0.480a 58.62b 387.5a 0.980
2013 N2 IM 0.208b 61.01a 376.7a 0.931
MM 0.332a 54.78b 301.7b 0.972
N1 IM 0.180a 60.10a 275.0b 0.965
MM 0.199a 57.53b 315.8a 0.997
N0 IM 0.288a 67.82a 233.3b 0.984
MM 0.221b 59.58b 283.3a 0.995

Notes: DAS, days after seeding; IM, intercropped maize; MM, monocultured maize; r m , initial relative growth rate; c m , maximum absolute growth rate; t b , the time of transition from the expolinear
to the linear growth phase; DAS, days after seeding. Different letters within a column and year indicate significant differences in least significant difference tests at P = 0.05.

The delays in the intercropping system were 4.3, 2.39, and 6.89 d in 2012 and 3.2.3. Dynamics of the LAI
6.23, 2.57, and 8.24 d in 2013 at N2, N1, and N0, respectively. The growth 3.2.3.1. Wheat. The LAI of wheat was generally low early in the season and
delay was shortest in the N1 treatment in each year. The estimates of linear peaked around 5767 DAS (Fig. 6). The LAI was signifi-cantly higher in the
growth rates in both years were highest in the N2-IM treatments (417.5 kg border row than in the inner and monocultured rows from 57 DAS until
1 1 1 1
ha d in 2012 and 376.7 kg ha d in 2013) and were lowest in the N1- maturity in 2012. From 47 to 77 DAS, the average LAIs in the border rows at
N2, N1, and N0 were 34.8%, 16.0%, and 33.4% higher, respectively, than in
IM and N0-IM treatments over both years. Also, c m was higher over both
the inner rows and 4.7%, 8.2%, and 26.5% higher, respectively, than in the
years in IM than in MM at N2 instead of at N1 and N0. In addition, the
monocultured rows. These trends were similar in 2013 (data not shown).
parameters were notably different between the two years. In particular, the
time of transition from exponential to linear growth was generally slightly
later in 2013 than in 2012, and the linear growth rate was higher in 2012
1 1 1 1 3.2.3.2. Maize. The LAI was significantly lower in the intercropped than the
(341.7417.5 kg ha d ) than in 2013 (233.3376.7 kg ha d ).
monocultured maize from 65 DAS till maturity (P < 0.05; Fig. 7) in both
years, indicating that intercropping limited the
 Y. Gao et al. / Field Crops Research 167 (2014) 1930 25

Fig. 5. Expolinear growth curves fitted to data of aboveground dry weights for the various treatments of intercropped maize (IM) and monocultured maize (MM) in 2012 and 2013. (A), (B), and (C)
are fitted curves of intercropped maize in two years at N2, N1, and N0, respectively. (D), (E), and (F) are fitted curves of monocultured maize in two years at N2, N1, and N0, respectively. (G) and (H)
are all fitted curves in 2012 and 2013, respectively.

vegetative growth of maize. The LAIs of intercropped maize from 65 to 145
DAS at N2, N1, and N0 were 15.6, 38.9, and 55.5% lower, respectively,
relative to the monocultured maize in 2012 and 23.9, 41.5, and 119.0% lower,
respectively, in 2013. Maximum LAI was lowest in the N0-IM treatment in
each year, indicating that an insufficient supply of N had aggravated the
reduction in vegetative growth.
3.3. Grain N content and nitrogen harvest index (NHI)
The grain N content of both wheat and maize tended to be enhanced as N
rates increased (Table 5). Intercropped wheat had a significantly higher grain
N content in each year than did the mono-culture (P < 0.05). Averaged over
both years, the grain N contents of intercropped wheat at N2, N1, and N0 were
12.4%, 10.8%, and 15.9%

Fig. 6. Growth patterns of leaf area index (LAI) for wheat in different rows in the intercrops and monocultures at the various rates of nitrogen fertilization in 2012. (A) N2,
(B) N1, and (C) N0. The LAIs were calculated on an equivalent basis of comparable land area to the monocultures.
26 Y. Gao et al. / Field Crops Research 167 (2014) 1930

Fig. 7. Growth patterns of leaf area index (LAI) for maize in the various treatments in (A) 2012 and (B) 2013. The values for the intercrops are on an equivalent basis of comparable land area to the
monocultures. DAS, days after seeding.

Table 5
Grain nitrogen (N) content, nitrogen yield, nitrogen harvest index (NHI), and internal use efficiency (IE) of wheat and maize in different treatments in 2012 and 2013.

Year Rate Cropping Wheat Maize

system 1 1 1 1 1 1
Grain N content (g kg ) N yield (kg ha ) NHI IE (kg kg ) Grain N content (g kg ) N yield (kg ha ) NHI IE (kg kg )
2012 N2 IM 24.04a 273.37 0.69a 28.81a 14.23a 459.46 0.67a 46.93b
MM 22.34b 281.31 0.56b 24.88b 12.34b 398.28 0.61b 49.74a
N1 IM 21.27a 243.48 0.65a 30.48b 13.14a 366.15 0.68a 51.62a
MM 20.08b 204.64 0.68a 33.89a 12.42b 385.40 0.61b 49.31b
N0 IM 19.69a 195.86 0.68b 34.74b 12.89a 255.52 0.82a 64.32a
MM 19.11a 157.21 0.74a 38.97a 9.87b 272.05 0.60b 60.64b
2013 N2 IM 22.74a 334.26 0.55a 24.03a 11.03b 262.02 0.64a 58.47a
MM 19.38b 236.89 0.50a 25.65a 11.62a 254.93 0.66a 56.69b
N1 IM 21.01a 230.94 0.60a 28.69b 10.97a 230.88 0.68a 62.14a
MM 18.16b 168.39 0.65a 35.99a 10.81a 263.98 0.59b 54.56b
N0 IM 22.51a 247.89 0.57a 25.50b 10.47a 180.68 0.62a 58.99a
MM 17.46b 159.42 0.57a 32.43a 10.78a 239.74 0.62a 57.58a

Notes: IM, intercropped maize; MM, monocultured maize; Different letters within a column and year indicate significant differences in least significant difference tests at P = 0.05. The values of
nitrogen yields in the intercrops are on equivalent basis of comparable land area of the monocultures.

higher, respectively, relative to monocultured wheat. The response of grain N 1

content for maize varied between years. The grain N content of intercropped
maize was significantly higher (P < 0.05) than that of monocultured maize in
2012, but no consistent sig-nificant differences were observed for 2013. The
higher grain N content of intercropped maize may be related to the NHI,
because intercropped maize also had a significantly higher NHI than did the
monoculture in 2012 (Table 5). No consistent significant differ-ences in NHI
were found between intercropped and monocultured wheat.
The N yield of intercropped wheat in both years was significantly higher
(P < 0.05) than that of monocultured wheat, except for the N2 treatment in
2012 (Table 5). The N yield in maize seedlings was lower in the intercrop
than the monoculture at all N levels (data not shown), but the final N yield
was higher in intercropped than mono-cultured maize at N2 but not at the
other two N levels (Table 5). In addition, the N uptake by maize varied
1 1.06, 1.08, and 1.18 in 2012, respectively and 1.23, 1.14, and 1.18 in 2013,
between years, ranging from 255.52 to 459.46 kg ha in 2012 and from
1
180.68 to 262.02 kg ha in 2013. The former was 13.475.3% higher than
the latter, mainly due to a higher yield of dry matter (18.141.9%).
51.94, and 59.11 kg kg for monocultured maize and 52.70, 56.88, and 61.66
1
kg kg for intercropped maize, respectively (Table 5). The IE was lowest at
N2 for both crops and both cropping systems, and the IE tended to increase as
N levels decreased. The IE of mono-cultured wheat was higher than that of
intercropped wheat over both years, and intercropped maize had a higher IE
than did the monoculture, except in the N2 treatment in 2012.
The relative N yield totals (RNTs) and the relative agronomic yield totals
in the various N treatments in the two years are shown in Fig. 8. The RNW
(relative N yield of wheat) in the intercropping system was much higher than
the relative grain yield, except for the N2 treatment in 2012, indicating that
intercropping negatively affected grain yield much more than N uptake. The
RNM (relative N yield of maize) was slightly lower than relative grain yield in
2013 but not in 2012. RNW and RNM did not differ significantly among the
three N rates. The averaged N yield totals (RNTs) at N2, N1, and N0 were
respectively. The RNTs were generally higher than LERs except for N2
treatment in 2012.

4. Discussion
3.4. N-use efficiency and relative N yields
4.1. Biomass productivity and grain yield
The average IEs over both years at N2, N1, and N0 were 25.26, 34.94, and

35.7 kg kg
1
for monocultured wheat and 26.42, 29.58, 30.12 kg kg
1
for Wheat/maize intercropping had an advantage over
intercropped wheat, respectively and 53.22, monocul-turing; the LER was above unity in all treatments
during the two
 Y. Gao et al. / Field Crops Research 167 (2014) 1930
Fig. 8. Relative nitrogen yield totals (RNTs) and land equivalent ratios (LERs) (or relative agronomic yield totals) of intercropping system at the various rates of nitrogen fertilization in (A) 2012 and
(B) 2013.
years. The LER tended to increase at higher N levels, consistent with the
findings by Li et al. (2011b) who reported that the total produc-tivity of an
intercropping system increased under high soil fertility or high rates of N
application. The LERs were 1.031.19 in our N1 and N2 treatments,
indicating that 319% more land area would be required by a monoculturing
system to equal the yield of an intercropping system. Land use efficiency was
previously reported to be higher in intercrops than monocultures (Yang et al.,
2011; Agegnehu et al., 2006).
The border effect, an indication of interspecific interactions and often
regarded as a cause of the high yield advantage in intercrops (Li et al., 2001),
which has previously been reported by Xiao et al. (2004) and Zhang et al.
(2007), is due to greater light interception (Zhang et al., 2008b) and a better
acquisition of nutrients by border plants (Knrzer et al., 2011). Our results
further indicated that both dry-matter accumulation and the LAI in border
rows were higher than those in both inner and monocultured rows of wheat,
indicat-ing that intercropping promoted the vegetative growth of border
plants. Li et al. (2001) reported that both border-row and inner-row effects
were observed for biomass yields (grain + straw). We did not find such an
increase in biological yield in inner rows. How this increase could occur is
still poorly understood.
The subordinate species in an intercropping system is often veg-etatively
limited, because taller species have a negative effect on neighboring plants
due to their advanced stage of development. Zhang et al. (2008b), for
example, reported that the main stem of cotton plants in a wheat/cotton
intercropping system had fewer leaves than did monocultured plants. Yang et
al. (2011) found that the mean LAI was lower in intercropped than
monocultured maize in a wheat/maize intercropping system with alternate
irrigation. The decrease in the LAI of intercropped maize in our study, in
agreement with the above study, was observed throughout nearly the entire
growing season (Fig. 7). Furthermore, our results con-firmed the
compensatory effect in maize and also indicated that the recovery or
compensation was incomplete when the nutri-ent supply was insufficient
(Zhang and Li, 2003). The plant dry weights of intercropped maize only at N2
exceeded those of mono-cultured maize at later growth stages, in accordance
with the higher linear growth rate in intercropping than in monoculturing at
N2 (Table 4). We should also note that the final yield of maize at N2 was
higher in the intercrop than in the monoculture, as readily accounted for by a
recovery effect at later growth phases for high N rates. However, there may be
alternative interpretation for the yield increase in intercropped maize at N2
because the vegetative inhibi-tion (LAI reduction) of intercropped maize
might result in increased yield. Research on redundancy growth and its
application in field management showed that an appropriate reduction of
redundant
27
growth such as biomass or LAI reduction could lead to more mat-ter and
energy flows into grains, causing yield increase (Han et al., 2005). Plants
respond as an integrated whole to the conditions of intercropping, with
numerous feedbacks between morphogenetic and growth processes operating
within the plant, so the available information can not interpret the
performance in detail. But N man-agement, control of vegetative growth, and
an appropriate balance between vegetative and reproductive growth all clearly
play impor-tant roles in crop production and should be considered jointly in
the future intercropping field management.
Zhang et al. (2007) first applied the expolinear growth equa-tion for
analyzing the growth patterns of cotton in a wheat/cotton intercropping
system. They concluded that the expolinear model was a suitable device for
characterizing crop growth and extract-ing key parameters from data sets,
enabling an incisive analysis of intercropping effects in different growth
phases. By using the expo-linear model, we found that growth was delayed
(2.398.24 d) in intercropped relative to monocultured maize due to the
shading by the taller species (wheat) during the initial growth period. A main
difference in the growth pattern was the low growth delay in the N1 treatment
in each year. This result could be interpreted in terms of the competition
between the two crops; the competitive effect of wheat during the seedling
phase of maize was weakest at this level of fertilization. The large competitive
ability of wheat rela-tive to maize under an insufficient supply of N has been
previously described by Li et al. (2001). And some agronomists and ecolo-
gists have proposed that higher nutrient availability aggravates interspecific
competition, because larger individuals occupy more developmental space,
intercept more light, and assimilate more nutrient resources and water (Grime,
1979; Keddy, 1989; Schippers et al., 1999). We thus inferred that the moderate
N rates could alleviate the competitive intensity to some extent, as in our
study reflected by the lowest degree of reduction in plant dry weights between
intercropped and monocultured maize at N1 rates during the initial stages of
growth (Fig. 4A), hence causing the shortest growth delay.
Irrigation water in the Hetao Irrigation District is diverted from the Yellow
River and is transmitted to the fields by channels. Water from the Yellow
River irrigates the local district but also the middle and lower reaches of the
valley, so irrigation is controlled and managed by the irrigation district
authorities. The availability of irrigation water depends on the times of
discharge from the upper reaches of the river without strictly following the
demands of the crops. Weather conditions thus might play an important role in
crop growth and yield performance. In our study, the crops per-formed
markedly different between the two years (Table 3). There are many
uncertainties on the effects of climate and we cannot
28 Y. Gao et al. / Field Crops Research 167 (2014) 1930

Fig. 9. (AC) Soil water content at different depth (020 cm and 3040 cm) in the middle of two maize rows of intercropping and monoculture at flowering in 2012 and 2013. (D) Comparison of
1
kernel number per ear of maize between 2012 and 2013 in different treatments. IM, intercropped maize; MM, monocultured maize; N0, N1, and N2 represent 0, 180, and 360 kg N ha for maize,
respectively; Different letters indicate significant differences in least significant difference tests at P = 0.05.

provide a complete analysis but we can partly give an interpreta-tion. The 1 1

high temperatures and the late fourth irrigation at the flowering stage of maize
in 2013 (Fig. 2; Table 2) induced a mild water deficit, as reflected by the
significantly lower soil water con-tent (040 cm) for every treatment in 2013
than in 2012 (Fig. 9AC). The development of the maize was thus negatively
affected, causing the lower kernel numbers per ear (Fig. 9D) and yield in
2013.
4.2. N uptake and use
Lemaire et al. (2007) concluded that N uptake by plants was co-regulated
by the soil N supply and by shoot growth based on studies under different
environments (temperate and subtropical) and with various crop species (C3
1
and C4). The total N uptake of wheat ranged from 157 to 334 kg ha
corresponds to the
257 kg ha N uptake of wheat at about 8 t ha grain yield aver-aged over
five regions in China (Liu et al., 2006). The total N uptake of maize ranged
1 1
from 180 to 459 kg ha , consistent with the range (180446 kg ha )
reported by Hao and Lao (2002) but higher than that reported by Li et al.
(2011b, 2003). Intercropped wheat had significantly higher grain N contents
and N uptakes than did mono-cultured wheat, perhaps due to an N border
effect as described by Li et al. (2001, 2003). The plant N uptake by
intercropped maize exceeded that of monocultured maize only at N2. This
finding cor-responds to the growth-recovery effect discussed above, which
also indicates that the compensation was incomplete when nutrient supply was
insufficient.
Liu et al. (2006) reported that the IE ranged from 19.8
1 1
to 66.4 kg kg for wheat and from 15.61 to 75.8 kg kg for
, which maize in China based on estimates by the QUEFTS model. Our
Y. Gao et al. / Field Crops Research 167 (2014) 1930 29

results were within the ranges
1 2001. Competition and crop per-
24.0338.97 kg kg for wheat and
1 The wheat/maize intercropping
46.9364.32 kg kg for maize. The
lower IE as a consequence of N
fertilization is in agreement with
results reported by other authors
(Albrizio et al., 2010; Latiri-Souki
et al., 1998; Raun and Johnson,
1999) and is mainly attributed to
the lower increase in grain yield
relative to N uptake.
Our results showed that
monocultured wheat had an
improved IE relative to
intercropped wheat at the crop
level, while inter-cropped maize
had a higher IE than did
monocultured maize. These
findings may be due to a lower
plant N yield for both monocultured
wheat (18.955.5%) and
intercropped maize (5.232.6%) in
most cases. At the system level, the
RNT ranged from 1.06 to 1.23 (Fig.
8), and the LER ranged on average
from 1.01 to 1.19. The RNT was 3
16% higher than the LER.
Therefore, contrary to our
expectation, the wheat/maize relay
intercropping system used N less
efficiently than the monocultures.
Knrzer et al. (2011) similarly
observed that winter wheat/maize
intercropping system did not
increase the N-use efficiency
relative to the monocultures at the
system level, and the internal N-use
efficiency (IE) was more favorable
for monocultured wheat and
intercropped maize.
We concluded that the
wheat/maize intercropping system
had substantial land productivity.
LER ranged from 1.01 to 1.19, but
a higher potential of intercropping
systems depends on favorable
environmental conditions for the
crops. The study area is severely
cold in winter, unlike most other
wheat/maize intercropping areas, so
wheat intercropping systems must
rely on spring wheat. Com-pared to
the intercropping of winter wheat
and maize in milder environments,
the full potential of intercropping
spring wheat and maize in this area
is likely limited by the temperature
(especially in the autumn). Also, the
growth of the intercropped maize in
our study was delayed by 28 d
relative to the monocultured maize.
Covers of plastic film or planting
on ridges may provide a warmer
environment for the maize for
alleviating the delayed growth and
increasing the potential of this
intercropping system.
system did not use N effi-ciently
relative to monocropping system.
Combining high yield with highly
efficient N management in
intercropping practices is always
challenging. Support to farmers
could be provided by demand-
oriented dosing and proper timing
of N fertilization with crop
simulation models or by
information from geographic infor-
mation systems. These options of
new strategies and mechanisms for
system improvement should be
investigated further.
Acknowledgments
This work was jointly supported
by the Special Foundation of
National Science & Technology
Supporting Plan (2011BAD29B09),
the 111 Project from the Ministry
of Education, the State Admin-
istration of Foreign Experts Affairs
(B12007), and the Supporting Plan
of Young Elites from Northwest A
& F University. Basic oper-ational
costs for this research were supplied
by Northwest A & F University. We
thank the anonymous reviewers for
their construc-tive comments,
which substantially improved the
manuscript.
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