Journal of Integrative Agriculture 2021, 20(2): 482–493

Available online at www.sciencedirect.com

ScienceDirect

RESEARCH ARTICLE

The effect of solar radiation change on the maize yield gap from the
perspectives of dry matter accumulation and distribution
YANG Yun-shan1, 2*, GUO Xiao-xia1*, LIU Hui-fang3, LIU Guang-zhou2, LIU Wan-mao1, MING Bo2, XIE
Rui-zhi2, WANG Ke-ru2, HOU Peng2, LI Shao-kun1, 2

1
The Key Laboratory of Oasis Eco-agriculture, Xinjiang Production and Construction Corps/College of Agronomy, Shihezi
University, Shihezi 832003, P.R.China
2
Key Laboratory of Crop Physiology and Ecology, Ministry of Agriculture and Rural Affairs/Institute of Crop Sciences, Chinese
Academy of Agricultural Sciences, Beijing 100081, P.R.China
3
College of Agronomy, Ningxia University, Yinchuan 750021, P.R.China

Abstract
The uneven distribution of solar radiation is one of the main reasons for the variations in the yield gap between different
regions in China and other countries of the world. In this study, different solar radiation levels were created by shading and
the yield gaps induced by those levels were analyzed by measuring the aboveground and underground growth of maize.
The experiments were conducted in Qitai, Xinjiang, China, in 2018 and 2019. The maize cultivars Xianyu 335 (XY335)
and Zhengdan 958 (ZD958) were used with planting density of 12×104 plants ha–1 under either high solar radiation (HSR)
or low solar radiation (LSR, 70% of HSR). The results showed that variation in the solar radiation resulted in a yield gap
and different cultivars behaved differently. The yield gaps of XY335 and ZD958 were 8.9 and 5.8 t ha–1 induced by the
decreased total intercepted photosynthetically active radiation (TIPAR) of 323.1 and 403.9 MJ m–2 from emergence to the
maturity stage, respectively. The average yield of XY335 was higher than that of ZD958 under HSR, while the average
yield of ZD958 was higher than that of XY335 under LSR. The light intercepted by the canopy and the photosynthetic
rates both decreased with decreasing solar radiation. The aboveground dry matter decreased by 11.1% at silking and 21%
at maturity, and the dry matter of vegetative organs and reproductive organs decreased by 9.8 and 20.9% at silking and
by 12.1 and 25.5% at physiological maturity, respectively. Compared to the HSR, the root weights of XY335 and ZD958
decreased by 54.6 and 45.5%, respectively, in the 0–60 cm soil layer under LSR at silking stage. The aboveground and
underground growth responses to different solar radiation levels explained the difference in yield gap. Selecting suitable

Received 13 April, 2020 Accepted 30 June, 2020

YANG Yun-shan, E-mail: 18093656011@189.cn; Correspondence
LI Shao-kun, Tel/Fax: +86-10-82108891, E-mail: lishaokun@caas.
cn; HOU Peng, Tel/Fax: +86-10-82108595, E-mail: houpeng@
caas.cn
*
These authors contributed equally to this study.
© 2021 CAAS. Published by Elsevier Ltd. This is an open
access article under the CC BY-NC-ND license (http://
creativecommons.org/licenses/by-nc-nd/4.0/).
doi: 10.1016/S2095-3119(20)63581-X
 YANG Yun-shan et al. Journal of Integrative Agriculture 2021, 20(2): 482–493 483

cultivars can increase maize yield and reduce the yield gaps induced by variation of the solar radiation levels in different
regions or under climate change.

Keywords: maize, solar radiation, yield gap, cultivar

conditions during the whole growth period. Ensuring suitable

light time and light intensity are beneficial for promoting
1. Introduction photosynthesis and yield formation (Li et al. 2002). In the
absence of other limiting factors in the environment, maize
Through the expansion of planting area, cultivar replacement yield is closely related to the intercepted photosynthetically
and the introduction and intensification of field cultivation active solar radiation, and a simple linear relationship exists
management measures, the global grain yield has increased between them (Muchow et al. 1990; Braconnier 1998;
significantly in the past 50 years (Cassman 1999; Foley et al. Hammad et al. 2016). With the decrease of intercepted solar
2005; Neumann et al. 2010). Nevertheless, food security radiation, the yield decreases (Muchow et al. 1990; Hammad
is under threat as natural resources are reduced and food et al. 2016). At the stage of maize reproductive growth, a
production stagnates due to many factors, especially climate decrease in solar radiation will restrict the establishment of
change (Senapati and Semenov 2020). The most effective reproductive organs, and lead to declines in leaf area index,
way to ensure global food security is to improve the yield photosynthetic rate and aboveground and underground
potential, increase the adaptation of agricultural production biomass (Mbewe and Hunter 1986; Zhang et al. 2007; Cui
to climate change and close the yield gap between current et al. 2013; Ren et al. 2016; Gao et al. 2017). At the same
and potential yields (Krandikar and Risbey 2000; Li et al. time, the rate of grain filling is decreased, the durations of
2014; Senapati and Semenov 2020). the grain filling peak and active filling days are shortened,
Potential yield refers to the yield obtained by a suitable the effective filling period of maize is limited, and the grain
crop cultivar under good growth conditions, without being size and weight are decreased, eventually leading to the
restricted by water, fertilizers or the stress of pests and yield decrease (Andrade and Ferreiro 1996; Chen et al.
diseases (Yang and Liu 2014; Liu et al. 2015). Under optimal 2014; Yang et al. 2016; Gao et al. 2018).
management conditions, the yield potential of a particular Agriculture is a human activity that is intimately linked
crop cultivar depends on meteorological factors, such as to climate, and farmers are always faced with an array of
solar radiation and temperature at a specific location (Van changing contexts. A major concern in understanding the
Ittersum and Rabbinge 1997; Liu et al. 2016). A previous impacts of climate change is the extent to what agriculture
systematic analysis of global maize, rice, wheat, and will be affected and its adaptation to climate change (such
soybean yields concluded that if 95% of the world’s planting as solar radiation) (Krandikar and Risbey 2000). In recent
areas reached climate potential, their yields would increase years, solar radiation intensity has been declining globally
by 50, 40, 60, and 20%, respectively (Licker et al. 2010). as a result of climate change (Stanhill and Cohen 2001; Che
Due to the limitation of soil, pests, management measures, et al. 2005; Abakumova et al. 2008; Liu et al. 2012). Solar
and farmers’ enthusiasm, the actual yields of crops are far radiation in China has also shown a decreasing trend due
lower than the potential yields of local crops (Liu Z J et al. to environmental pollution (Shi et al. 2008; Qi et al. 2019;
2017). The existence of a yield gap between different Zhang Q et al. 2019; Chen and Pang 2020). At the same
regions is mainly driven by meteorological factors, especially time, due to differences in economic development between
solar radiation. There are great differences in solar radiation regions, the environmental situation varies as well as the
in different regions of China, especially between the eastern degree of declination in solar radiation among regions (Liu
and western regions (Yang et al. 2011; Hou et al. 2014). et al. 2015; Liu S et al. 2019; Zhao et al. 2020). Previous
Solar radiation in the western regions is more abundant than studies on yield gap have focused on the effects of field
that in the eastern regions in China. As a result, there is a management systems on crop yield, such as the effects of
great yield gap between the eastern and western regions weed and pest stress (Hochman and Horan 2018), water
(Xu et al. 2017; Hou et al. 2018). management (Mohammadi-Ahmadmahmoudi et al. 2020)
As a driving factor for crop growth and development, and fertilization (Rhebergen et al. 2020). The effect of solar
solar radiation is closely related to plant morphogenesis and radiation on the yield gap is mostly simulated by models,
yield formation (Demetriades-Shah et al. 1994; Bansouleh which have indicated that the yield gap decreases due to the
et al. 2009). Maize is a typical C4 crop with short sunlight decrease in simulated yield potential after a solar radiation
and high light efficiency, and it needs good solar radiation decline and the actual yield increase under optimized
484 YANG Yun-shan et al. Journal of Integrative Agriculture 2021, 20(2): 482–493

cultivation management measures (Liu et al. 2012; Tao et al. 2019). The amount of the photosynthetically active
et al. 2015; Zhao et al. 2015). However, few studies have radiation reaching the top and bottom of the canopy were
focused on the effect of solar radiation on the yield gap measured with a Sunscan (SUNSCAN, Delta-T, UK). The
through field experiments. In addition, the previous studies total intercepted photosynthetically active radiation (TIPAR)
mainly focused on the effect of shading on maize growth, from the period of emergence to the maturity stage was
but did not incorporate the quantification of radiation. In this calculated according to the following formula:
study in the region with the best solar radiation resources
TIPAR (MJ m–2)=(1– B )×C
in China, we used shading to mimic the low solar radiation A
conditions in other regions of China, and explored the yield where A is photosynthetically active radiation (PAR)
gap caused by the experimental decrease of solar radiation above the canopy, B is transmitted photosynthetically
and its internal physiological mechanisms. The results active radiation at the bottom of the canopy and C is total
will provide a theoretical basis for reducing the yield gap accumulated photosynthetically active radiation from the
between regions and under the conditions of solar radiation period of emergence to the maturity stage. PAR was
declines caused by climate change. measured with a diagonal orientation every 30 cm above
the ground (Liu G et al. 2019). The fraction of intercepted
2. Materials and methods PAR was measured on clear days during the silking stage,
using a line quantum sensor (SunScan, Delta-T) in 2018
2.1. Experimental design and 2019. The three ear leaves were chosen at the silking
stage for photosynthesis measurements in 2018 and 2019.
The experiment was conducted in Qitai (Xinjiang Uygur The measurements of gas exchange were carried out on
Autonomous Region, China, 43°49´27´´N, 89°48´22´´E) clear days at 12:00–13:00 using a programmable, open-
in 2018 and 2019. Meteorological data for the 2018 flow gas exchange portable system (LI-6400, Li-Cor Inc.,
and 2019 maize growing seasons were obtained from a Lincoln, NE, USA) operated at a 500-µmol s–1 flow rate, and
“Watch Dog” Data Logger (Spectrum Technologies, Inc., the CO2 concentration of the reference chamber (CO2-R)
USA) located in the experimental field. The two-year was set at 400 μmol mol−1. Light inducements were done
meteorological data are summarized in Table 1. The effect of by keeping the leaves in the chamber under PAR=2 000
solar radiation on yield gap was studied by using single factor μmol m−2 s−1 until parameter readings were stable (Sofo
experiments involving two widely planted high-yield maize et al. 2009; Zhang Y et al. 2019).
cultivars, namely Xianyu 335 (XY335) and Zhengdan 958 Sufficient water was applied to prevent water stress.
(ZD958). The planting density was 12×104 plants ha–1. On the first day after sowing, all experimental plots were
Solar radiation intensity was 70% of natural solar radiation irrigated (15 mm), and starting from 60 d after sowing,
after shading (low solar radiation, LSR), and that level was single water applications of 58 mm were delivered at 9–10 d
compared with natural solar radiation (high solar radiation, intervals throughout the growing season for a total of nine
HSR). Shading was imposed from the three-leaf stage until applications; the total irrigation amount was about 540 mm
maturity. The experimental plots were 11 m×10 m in size, (Zhang et al. 2017). Base fertilizers were applied as 150 kg
and adjacent plots were separated by a walkway with a N ha– 1 from urea, 225 kg P2O5 ha–1 (super phosphate) and
width of 1 m. Shading nets of different shading levels were 75 kg K2O ha–1 (potassium sulfate) prior to sowing. An
designed and fabricated. A detachable shade shed was built additional 300 kg N ha–1 was applied during the growing
using a shade net, the top of which was positioned about season to ensure a non-limiting supply of nutrients. Weeds,
1.5 m away from the maize canopy, which ensured that the diseases and pests were controlled manually in the plots to
microclimatic conditions (other than the solar radiation) in the eliminate confounding effects.
shade shed were consistent with the unshaded portions of
the field (Yang et al. 2019). A photo of the field experiment 2.2. Sampling and measurement
is shown in Fig. 1.
Previous studies found that shading did not affect Dry matter At the silking and physiological maturing stages,
the quality of incident light in the maize canopy (Yang three consecutive plants were measured in the center row of

Table 1 Mean daily maximum temperature (Tmax), minimum temperature (Tmin), diurnal temperature variation (Td), solar radiation,
relative humidity (RH), and precipitation (Pre) during the maize growing season at Qitai Farm, Xinjiang, China in 2018 and 2019
Year Tmax (°C) Tmin (°C) Td (°C) Solar radiation (MJ m–2 d–1) RH (%) Pre (mm)
2018 24.0 10.3 17.2 9.41 53.4 221.0
2019 25.9 11.5 18.7 9.75 52.6 138.5
 YANG Yun-shan et al. Journal of Integrative Agriculture 2021, 20(2): 482–493 485

differences in yield between the two different years, and

the average yield in 2019 was higher than that in 2018.
The yield gap between HSR and LSR in 2018 was greater
than the gap in 2019, and the TIPAR showed the same
trend. TIPAR of XY335 and ZD958 decreased by 323.1
and 403.9 MJ m–2, respectively, and the average yield gaps
were 8.9 and 5.8 t ha–1 (by averaging 2018 and 2019 data).
In other words, the reductions of average TIPAR were 26.3
and 33.5%, and the average yield reductions were 41.8 and
30.6%, respectively. The average yield of XY335 (21.4 t
ha–1) was higher than that of ZD958 (18.8 t ha–1) under
HSR, so the yield gap between the two cultivars was 2.6 t
ha–1 under HSR. The average yields of XY335 and ZD958
Fig. 1 Maize fields under natural and 70% natural solar
radiation levels. under LSR were 12.5 and 13.0 t ha–1, respectively, so the
yield gap between the two cultivars was 0.5 t ha–1, which
was far less than the yield gap under HSR. These results
each plot. All organs were cut off in their natural state within indicated that XY335 was more sensitive to changes in
the canopy. The organs were separated into the stem, leaf, solar radiation, while ZD958 had better adaptability to the
sheath, tassel, ear, and bract, and dried to constant weight decreased solar radiation (Fig. 2).
at 85°C (Liu G Z et al. 2017).
Roots In each plot, the roots of three adjacent plants from 3.2. Effect of solar radiation change on aboveground
the same inside row were removed from the soil by digging dry matter accumulation and distribution in maize
at the silking stage. Each root system was excavated with
a certain volume of soil, depending on the average area that At silking and maturity stages, the amounts of dry matter
the plant occupied, at a depth of 60 cm; and this sample was in each organ decreased as solar radiation decreased,
then divided into three layers of 20 cm each. The soil from and the decreases in XY335 were greater than those in
each of these three layers was kept in a separate numbered ZD958. The aboveground dry matter of XY335 and ZD958
nylon bag for subsequent manual root washing and picking, decreased by 13.1 and 9.1% at silking and by 22.4 and
and after washing and picking, the roots were scanned 19.6% at maturity, respectively (by averaging 2018 and
with a scanner (Epson V800, Indonesia) and analyzed by 2019 data). The proportions of stem, leaf and sheath dry
an analysis program (WinRhizo Pro Vision 5.0, Canada) to matter increased with decreasing solar radiation, and the
obtain the root length. Finally, the roots were dried at 85°C proportions of ear and tassel dry matter decreased with
to a constant weight to determine the root dry weight. decreasing solar radiation at silking and maturity stages.
Yield At physiological maturity, an area of three central rows The dry matter proportions of stem, leaf and sheath were
and 5 m long was harvested manually from the center of each greater than those of ear and tassel.
plot and grain weight was measured. A portable moisture At the silking stages in 2018 and 2019, the stem, leaf,
meter (PM8188, Kett Electric Laboratory, Tokyo, Japan) was sheath, ear, and tassel dry matter weight proportions of
used to determine the moisture content of the grains. Grain XY335 were 43.3, 31.1, 17.3, 5.5, and 2.9% under HSR,
yield was then determined at 14% grain moisture content. respectively, while they were 43.4, 32.2, 16.9, 5.0, and
Statistical analysis Statistical calculations were performed 2.6% under LSR during the two years. The stem, leaf,
using the Microsoft Excel 2010 Software, plots were sheath, ear, and tassel dry matter weight proportions of
generated with SigmaPlot, and statistical analyses were ZD958 were 37.5, 32.7, 17.5, 6.2, and 6.4% under HSR,
performed using the SPSS 21.0 Software (IBM, Armonk, respectively, while they were 38.8, 34, 16.5, 5.1, and 5.7%
NY, USA). under LSR (Fig. 3-A and B). The overall reductions of the
dry matter of vegetative organs and reproductive organs
3. Results were 9.8 and 20.9%.
At the maturity stages in 2018 and 2019, the dry matter
3.1. Yield gap of maize induced by solar radiation weight proportions of stem, leaf, sheath, ear, bract, and
change tassel of XY335 were 18.7, 11, 5.4, 59.7, 4.75, and 0.5%
under HSR, respectively, while they were 20, 11.7, 5.9, 57.9,
The difference in solar radiation level caused a variation in 4.2, and 0.5% under LSR. The dry matter weight proportions
maize yield, which was HSR>LSR. There were significant of stem, leaf, sheath, ear, bract, and tassel of ZD958 were
486 YANG Yun-shan et al. Journal of Integrative Agriculture 2021, 20(2): 482–493

HSR LSR
A 25 B 25
2018 2019
a a
a
20 20 b
5.7
c 3.5
d
12.2 8.1
Yeild (t ha–1)

Yeild (t ha–1)
15 b 15
b
10 10

5 5

0 0
XY335 ZD958 XY335 ZD958
C 1 600 D 1 600
2018 2019
a
a a a
1 200 1 200
TIPAR (MJ m–2)

TIPAR (MJ m–2)

353.0
b 474.2 b 293.2 b 333.6
c
800 800

400 400

0 0
XY335 ZD958 XY335 ZD958

Fig. 2 Maize yield and the total intercepted photosynthetically active radiation (TIPAR) from the period of emergence to the maturity
stage of Xianyu (XY335) and Zhengdan 958 (ZD958) under high solar radiation (HSR) and low solar radiation (LSR) in 2018 and
2019. Error bars represent the standard deviations of three replicates. Means with different letters are significantly different at
the level of 0.05.

15.6, 12, 5.6, 61.6, 4.4, and 1% under HSR, respectively, under both treatments in 2018 and 2019 (Fig. 4). The
while they were 17.3, 14.9, 6.1, 56.3, 4.2, and 1.3% under distributions of light for XY335 under HSR and LSR matched
LSR. In 2018 and 2019, the proportions of vegetative logarithmic functions, which were y=53.785ln(x)+75.767 and
organs (stem+leaf+sheath) of XY335 were 35.1 and 37.5% y=55.598ln(x)+23.094, in 2018, and y=67.299ln(x)+71.286
under HSR and LSR, respectively, and they were 33.2 and and y=82.481ln(x)–21.238, respectively, in 2019. The
38.3% for ZD958. The proportions of reproductive organs distributions of light for ZD958 under HSR and LSR
(ear+bract+tassel) of XY335 were 64.9 and 62.5% under matched logarithmic functions of y=67.720ln(x)–9.187 and
HSR and LSR, respectively, and they were 66.9 and 61.8% y=42.583ln(x)+26.297, in 2018, and y=54.737ln(x)+17.798
for ZD958. Compared to HSR, the proportions of vegetative and y=53.781ln(x)+17.987, respectively, in 2019. The
organs of XY335 and ZD958 were decreased by 17 and 7.1% averages of photosynthetically active radiation transmission
under LSR, respectively, in 2018 and 2019. The proportions of HSR and LSR for XY335 were 21.3 and 36.3%, and 23.9
of reproductive organs of XY335 and ZD958 were decreased and 31.5% for ZD958, respectively, in 2018. The averages
by 25.5 and 25.5% under LSR, respectively, in 2018 and 2019. photosynthetically active radiation transmission of HSR
The reductions of the dry matter of vegetative organs and and LSR for XY335 were 20.6 and 31.1%, and those of
reproductive organs were 12.1 and 25.5%. Comparing the ZD958 were 30.3 and 32.3%, respectively, in 2019. These
data of the two cultivars under HSR, the decrease in XY335 results indicated that the average light interception by the
was greater than that in ZD958. The reductions of the dry canopy of the population decreased when the solar radiation
matter of vegetative organs and reproductive organs in 2018 decreased. The average photosynthetically active radiation
were greater than that in 2019 (Fig. 3-C and D). transmission gaps of HSR and LSR for XY335 were 15.0
and 10.5% in 2018 and 2019, respectively, while they were
3.3. Distribution of light in the canopy 7.6 and 2.0% for ZD958. These results indicated that the
photosynthetically active radiation transmission gaps of the
Light transmission increased with increasing plant height two treatments in 2018 were greater than those in 2019, and
 YANG Yun-shan et al. Journal of Integrative Agriculture 2021, 20(2): 482–493 487

Ear Stem Leaf Sheath Bract Tassel

A B
50 50
2018 Silking stage 2019 Silking stage
Dry matter accumulation (t ha–1)

Dry matter accumulation (t ha–1)

40 40

30 30
a
20 a ab 20 b bc c
ab b
10 10

0 0
HSR LSR HSR LSR HSR LSR HSR LSR
XY335 ZD958 XY335 ZD958
C D
50 50
2018 Maturity stage 2019 Maturity stage
Dry matter accumulation (t ha–1)

Dry matter accumulation (t ha–1)

a a a
40 40 bc ab
b c
30 b 30

20 20

10 10

0 0
HSR LSR HSR LSR HSR LSR HSR LSR
XY335 ZD958 XY335 ZD958

Fig. 3 Dry matter accumulation and distribution among various organs of Xianyu 335 (XY335) and Zhengdan 958 (ZD958) under
high solar radiation (HSR) and low solar radiation (LSR) in 2018 and 2019 at the silking and physiological maturity stages. Error
bars represent the standard deviations of three replicates. Means with different letters are significantly different at the level of 0.05.

they were greater for XY335 than for ZD958. layer decreased with decreasing solar radiation. In 2018
and 2019, the average dry weights of XY335 and ZD958
3.4. Photosynthetic rate of maize induced by solar decreased by 54.6 and 45.5% under decreased solar
radiation change radiation. The significant difference in the effect of solar
radiation on the root dry weight was mainly found in the
The photosynthetic rates of XY335 and ZD958 decreased 0–20 cm soil layer. In 2018 and 2019, the average root dry
with decreasing solar radiation at the silking stage. In 2018, weights for XY335 and ZD958 in the 0–20 cm soil layer under
the photosynthetic rates of the XY335 and ZD958 cultivars LSR were 55.4 and 45.9% lower than those under HSR,
were 34.5 and 36.9 μmol m s under HSR, and 30.1 and −2 −1
respectively (Fig. 6-A–D). The root length of the 0–20 cm soil
30.7 μmol m−2 s−1 under LSR, respectively. Compared to layer decreased more than that of the 40–60 cm soil layer.
the HSR, the photosynthetic rates of XY335 and ZD958 In 2018 and 2019, the root lengths of the two cultivars in the
decreased by 12.6 and 16.9%, respectively, under the LSR. 0–20 cm soil layer decreased significantly under reduced
In 2019, the photosynthetic rates of XY335 and ZD958 solar radiation, and the average root lengths for XY335 and
were 38.6 and 35.9 μmol m s under HSR, and 28.4 and −2 −1
ZD958 in the 0–20 cm soil layer under LSR were 50.9 and
34.6 μmol m−2 s−1 under LSR, respectively. Compared to 46.3% lower than HSR, respectively (Fig. 6-E–H). Under
the HSR, the photosynthetic rates of XY335 and ZD958 the different solar radiation conditions, the root dry weight in
decreased by 26.9 and 3.7% under the LSR (Fig. 5). 2018 was greater than that in 2019, but the difference was not
significant. However, the root length in 2019 was significantly
3.5. Distribution of roots greater than that in 2018. Root dry weights and root lengths
of the two cultivars were significantly different, and both were
The root dry weight and root length of the 0–60 cm soil greater in XY335 than in ZD958, especially under HSR.
488 YANG Yun-shan et al. Journal of Integrative Agriculture 2021, 20(2): 482–493

4. Discussion radiation on increased yield is greater than the effect of

temperature (Yang et al. 2018; Wang et al. 2019). Variations
4.1. Effect of solar radiation change on maize yield in resources and environment result in yield gaps among
gap maize growing regions (Lobell et al. 2009). Due to the
special geographical environment of China, there are
Recent studies have shown that solar radiation and varying degrees of regional differences in solar radiation and
temperature resources are significantly related to maize temperature resources, resulting in the existence of yield
grain yield, and the negative impact of decreased solar gaps between different regions (Yang et al. 2011; Hou et al.

HSR LSR

A 400 B 400
2018-XY335 2018-ZD958
350 350
y=53.785ln(x)+75.767 y=67.720ln(x)–9.187
Plant height (cm)

Plant height (cm)

300 R2=0.985** 300 R2=0.916**
250 250
200 200
150 150
y=55.598ln(x)+23.094 y=42.583ln(x)+26.297
100 100
R2=0.922** R2=0.888**
50 50
0 0
0 10 20 30 40 50 60 70 80 90 100 0 10 20 30 40 50 60 70 80 90 100
Transmission (%) Transmission (%)
C 400 2019-XY335 D 400 2019-ZD958
350 y=67.299ln(x)+71.286 350 y=54.737ln(x)+17.798
R2=0.980** R2=0.946**
Plant height (cm)

Plant height (cm)

300 300
250 250
200 200
150 y=82.481ln(x)–21.238 150 y=53.781ln(x)+17.987
100 R2=0.968** 100 R2=0.969**
50 50
0 0
0 10 20 30 40 50 60 70 80 90 100 0 10 20 30 40 50 60 70 80 90 100
Transmission (%) Transmission (%)

Fig. 4 Distribution of photosynthetically active radiation transmission every 30 cm in the canopy of Xianyu 335 (XY335) and Zhengdan 958
(ZD958) under high solar radiation (HSR) and low solar radiation (LSR) in 2018 and 2019. **, significant at the level of P<0.01.

HSR LSR
A B
50 50
Photosynthetic rate (μmol m–2 s–1)

Photosynthetic rate (μmol m–2 s–1)

2018 2019
a
40 a 40 a a
a
b b
b
30 30

20 20

10 10

0 0
XY335 ZD958 XY335 ZD958

Fig. 5 Effects of photosynthetic rate at the silking stage in leaves under high solar radiation (HSR) and low solar radiation (LSR)
in 2018 and 2019. XY335, Xianyu 335; ZD958, Zhengdan 958. Error bars represent the standard deviations of three replicates.
Means with different letters are significantly different at the level of 0.05.
 YANG Yun-shan et al. Journal of Integrative Agriculture 2021, 20(2): 482–493 489

HSR LSR
Root dry weight (g/plant)
0 2 4 6 8 10 12 0 2 4 6 8 10 12 0 20 40 60 80 0 20 40 60 80
A B C D
a a a a
0–20 b b b b
Soil depth (cm)

c c c c
20–40 c c d c

c c d c
40–60 c 2018-XY335 c 2018-ZD958 2019-XY335 c 2019-ZD958
d
Root length (m/plant)
E F G H
a a a a
0–20 b b b b
Soil depth (cm)

c c c c
20–40 c cd d d

c d e d
40–60 c 2018-XY335 d 2018-ZD958 e 2019-XY335 e 2019-ZD958

Fig. 6 Distributions of root dry weight and root length of Xianyu 335 (XY335) and Zhengdan 958 (ZD958) under high solar radiation
(HSR) and low solar radiation (LSR) at the silking stage in 2018 and 2019. Error bars represent the standard deviations of three
replicates. Means with different letters are significantly different at the level of 0.05.

2012; Xu et al. 2017). The results of this study showed determinant of the plant growth potential and closely related
that when the level of solar radiation was decreased, the to the yield formation. Root growth is closely related to solar
effective solar radiation interception during the growth radiation, maize root growth under high solar radiation is
period of the maize decreased, and the photosynthetic rates better than that under low solar radiation, and the responses
of XY335 and ZD958 at the silking stage also decreased of roots of different genotypes to solar radiation are different,
with decreasing solar radiation, which led to the decreases resulting in varying yield gaps (Son et al. 1988; Amos and
in maize yield and caused the yield gaps. The difference Walters 2006; Niu et al. 2019). The results of this study
in yield gap between different years was mainly due to the confirmed this view. With the reduction of solar radiation, the
difference in TIPAR in the two years. root dry weight and root length in each soil layer decreased.
Canopy light distribution is important for crop The 0–20 cm soil layer had a larger rate of decrease, and
photosynthesis, which is a key factor for crop production the decrease rate for the two cultivars was XY335>ZD958
(Gu et al. 2014; Kromdijk and Long 2016). Dry matter (Fig. 6). Root growth of XY335 was greatly affected by solar
production is the physical basis of maize grain yield (Hou radiation, which might result in a larger yield gap. In this
et al. 2020), and photosynthesis is the basis of biomass study, the total root lengths of the two cultivars in the LSR
formation. In this study, light transmission in the canopy treatments in 2019 were found to be significantly higher than
increased with increasing plant height and the average those in 2018. Therefore, the absorption and utilization of
amount of light intercepted by the canopy decreased when water and nutrients might increase, which could increase
the solar radiation was reduced, which indicated that under the yield and reduce the yield gap between HSR and LSR
the low solar radiation, the light captured by the canopy in 2019 (Fig. 6). In addition, the photosynthetically active
was weak. The photosynthetic rates at the silking stage radiation transmission gaps between HSR and LSR in
decreased under reduced solar radiation, as did the amounts 2018 were greater than that in 2019, and the reduction of
of dry matter of each organ and dry matter accumulation, aboveground dry matter accumulation in 2018 was greater
thus resulting in yield gaps (Zhu et al. 2016; Gu et al. 2017; than that in 2019; all of which might have been the reason
Khan et al. 2017; Yao et al. 2017; Liu G et al. 2020). for the greater yield gap in 2018.
The aboveground and underground components of
4.2. The aboveground and underground growth maize complement each other to form the overall functional
responses to different solar radiation levels system of the crop, which is self-adaptive and -adjusting
to the external environment. When the solar radiation
The root draws water and nutrients from the soil to supply decreases, the plant supplies photosynthates for nearby
the growth of the ground of the plant, which is the key organ growth, so the aboveground growth is better while
490 YANG Yun-shan et al. Journal of Integrative Agriculture 2021, 20(2): 482–493

underground growth is severely limited (Casper and water management (Mohammadi-Ahmadmahmoudi et al.
Jackson 1997; Chen et al. 2002; Miralles et al. 2011). This 2020), fertilization (Rhebergen et al. 2020), and social
mechanism is consistent with the results of the present economy (Yang and Liu 2014), also affect the yield gaps
study. At silking, the root weights of XY335 and ZD958 under between regions (Liu W et al. 2020). In addition, abiotic
LSR were decreased by 54.6 and 45.5%, respectively, and factors can constrain yield as well, such as climate change
the aboveground dry matter decreased by 13.1 and 9.1%, (Liu et al. 2012, 2016), temperature and rainfall (Xu et al.
which meant that the reduced level of solar radiation had a 2017; Abdulai et al. 2020). Considering these factors,
greater impact on the underground root growth. Meanwhile, selecting the most appropriate cultivars is more important for
compared to the HSR, with the decrease of the radiation reducing the regional yield gaps (Andrea et al. 2018). In this
level, the dry matter weight of each organ on the ground study, different cultivars responded differently to changes
decreased, and the reductions of the dry matter of vegetative in solar radiation. The difference in yield of XY335 caused
organs and reproductive organs were 9.8 and 20.9% at silking by the reduced solar radiation was greater than for ZD958,
stage, and 12.1 and 25.5% at the physiological maturity indicating that XY335 was more sensitive to changes in solar
stage, respectively, so the dry matter of reproductive organs radiation. However, the average yield of XY335 was higher
was greater than that of vegetative organs. This pattern than that of ZD958 under HSR (Fig. 1). In this respect,
was consistent with previous studies which indicated that XY335 can be used to improve production potential in
under LSR, the development of maize ears (a reproductive areas with abundant light resources. Previous studies on
organ) was limited, and the growth of tassel and ears was light response curves of different cultivars at the silking
unbalanced, which would lead to an increase in male and and grain filling stages have indicated that XY335 has a
female separation during the flowering period and thus greater ability to utilize strong light and ZD958 has a greater
decrease grain yield (Cui et al. 2015; Yang et al. 2016). ability to utilize weak light under the same planting density
in the same area (Liu 2016). In this study, the average
4.3. Yield gap between different regions induced by yield of ZD958 was higher than that of XY335 under LSR,
solar radiation indicating that ZD958 had better adaptability to LSR.
Therefore, cultivars similar to XY335 should be selected
Previous studies have indicated that adequate light in areas with high solar radiation resources, and on the
conditions can increase grain yield (Li et al. 2010; Chen contrary, cultivars similar to ZD958 should be selected in
et al. 2014). The east-west distance spanning the maize areas with low solar radiation resources to increase grain
ecological regions in China is large and this results in a yields and reduce yield gaps.
significant difference in solar radiation, especially between
the west and east regions, and leads to the differences in 5. Conclusion
planting densities and final grain yields (Li and Wang 2008;
Xu et al. 2017; Yang et al. 2019; Hou et al. 2020). In this In this study, we found that yield gaps could be induced
study, shading was adopted to mimic regions of low solar by solar radiation change and different gaps were found
radiation conditions, especially the eastern region of China. between cultivars XY335 and ZD958. Under HSR, grain
One of the most effective measures for maximizing maize yield of XY335 was higher than that of ZD958, while the
yield has been adjusting the plant density to different climatic opposite trend was found under LSR. This indicated that
conditions (Xu et al. 2017). In this study, the variation of XY335 could be used in areas with abundant light resources
solar radiation that was tested resulted in yield gaps ranging and ZD958 should be used under low solar radiation
from 5.8 to 8.9 t ha–1, which were smaller than the actual conditions to increase grain yields and reduce the yield gap.
conditions in the field. This difference might be because the The light intercepted by the canopy, photosynthetic rates,
higher yield gap between the western and eastern regions the amounts of dry matter of different organs, and root dry
is not only related to solar radiation change, but also closely weight and length all decreased at reduced solar radiation
related to the actual planting density (Xu et al. 2017; Hou levels, and the decreases in XY335 were greater than those
et al. 2020). The planting density in the western region is in ZD958, resulting in different yield gaps. The reduced level
higher than that in the eastern region (Xu et al. 2017; Zhang of solar radiation had a greater impact on the underground
D et al. 2019). root growth than aboveground growth, and it had a greater
impact on reproductive organ growth than vegetative organ
4.4. Reducing yield gap by using suitable maize growth, which explained the difference in the yield gaps.
cultivars Selecting suitable cultivars can increase yields and reduce
yield gaps that are driven by variations of solar radiation in
Other factors, such as cultivars, weeds, plant diseases, different regions or under climate change.
 YANG Yun-shan et al. Journal of Integrative Agriculture 2021, 20(2): 482–493 491

Acknowledgements between the plant root and shoot. Journal of Shaoguan

University (Natural Science Edition), 12, 64–71. (in Chinese)
Cui H Y, Camberato J J, Jin L, Zhang J. 2015. Effects of shading
This work was financially supported by the National
on spike differentiation and grain yield formation of summer
Key Research and Development Program of China
maize in the field. International Journal of Biometeorology,
(2016YFD0300110, 2016YFD0300101), the National
59, 1189–1200.
Natural Science Foundation of China (31871558) and the Cui H Y, Jin L B, Li B, Dong S T, Zhang J W. 2013. Effects of
National Basic Research Program of China (973 Program, shading on dry matter accumulation and nutrient absorption
2015CB150401). of summer maize. Chinese Journal of Applied Ecology, 24,
3099–3105. (in Chinese)
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Managing editor WANG Ning