Agricultural and Forest Meteorology 308-309 (2021) 108616

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Agricultural and Forest Meteorology

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

The efficient use of radiation, water, and nitrogen uptake by

low-nitrogen-tolerant broomcorn millet (Panicum miliaceum L.) increased
grain yield in the Loess Plateau, China
Chunjuan Liu a, c, Xiangwei Gong b, c, Ke Dang b, Panpan Zhang d, Qinghua Yang b,
Xiping Deng a, *, Baili Feng b, *
a
College of Life Sciences, Northwest A & F University, Yangling, Shaanxi 712100, PR China
b
College of Agronomy, State Key Laboratory of Crop Stress Biology in Arid Areas/Northwest A & F University, Yangling, Shaanxi 712100, PR China
c
College of Agronomy/Shenyang Agricultural University, Shenyang, Liaoning 110866, PR China
d
College of Life Sciences/Yulin University, Yulin, Shaanxi 719000, PR China

A R T I C L E I N F O A B S T R A C T

Keywords: The development of highly productive, low-N-tolerant cultivars is important to facilitate green and sustainable
Broomcorn millet agriculture. However, it is unclear how low-N-tolerant cultivars achieve high grain yields under field conditions.
Dry matter accumulation In this study, the physiology facilitating the strong growth and efficient resource use of four low-N-tolerant
Low-N-tolerance
Panicum miliaceum L. cultivars was compared with two low-N-sensitive cultivars using low and typical N
Photosynthetic characteristics
Resource use efficiency
fertilization applications (0, 75, and 150 kg N ha− 1) on the semiarid Loess Plateau, China in 2019 and 2020.
Compared to normal N fertilization, low N fertilization treatments significantly decreased plant photosynthetic
capacity by regulating leaf stomatal closing, resulting in reduced dry matter accumulation and transportation,
growth inhibition, weakened resource utilization, and consequently, a 28.9% decrease in grain yield. Compared
to the low-N-sensitive cultivars, the low-N-tolerant cultivars had improved growth, leaf photosynthetic capacity,
and dry matter transportation. These improved physiological characteristics resulted in more effective resource
capture in the low-N-tolerant cultivars. This included higher and more efficient radiation use (24.4% and 26.9%),
water use (63.5% and 48.2%), and N uptake (37.0% and 33.7%) in 2019 and 2020, respectively. Furthermore,
the low-N-tolerant cultivars had increased productivity, with higher grain yields (38.6-41.6%) and harvest index
values (16.7-28.2%) under low N conditions in the two years of the study. Overall, the results of this study
determined the effect of low N input on broomcorn millet productivity and illustrated the strategies by which
low-N-tolerant cultivars increased grain yield in semiarid field conditions.

LDM largest dry mass of the stem

NPE N productive efficiency
Abbreviations NUE crop N use efficiency
Ci intercellular CO2 concentration NUPE N uptake efficiency
DMM dry mass of the same vegetative organ at maturity PAR photosynthetically active radiation
DTA dry matter transported Pn photosynthetic rate
DTR dry matter transportation rate RUE radiation use efficiency
GCR contribution rates of vegetative organs (stem, leaf, sheath) to SWC soil water content
the grains Tr transpiration rate
GDM dry mass of the grain WUE water use efficiency
gs stomatal conductance
HI harvest index
IPAR PAR intercepted by the plant canopy

* Corresponding author.
E-mail addresses: dengxp@ms.iswc.ac.cn (X. Deng), fengbaili@nwsuaf.edu.cn (B. Feng).

https://doi.org/10.1016/j.agrformet.2021.108616
Received 17 February 2021; Received in revised form 6 July 2021; Accepted 15 August 2021
Available online 25 August 2021
0168-1923/© 2021 Elsevier B.V. All rights reserved.
C. Liu et al. Agricultural and Forest Meteorology 308-309 (2021) 108616

1. Introduction produce higher grain yields than N-inefficient cultivars under the same
N fertilizer treatment (Sinclair and Rufty, 2012). Grain yield may be
By the 2050s, the global population is estimated to reach 9.7 billion further improved through the combination of genetically modified cul­
(Gerland et al., 2014). To meet the increasing demands of the growing tivars and increased N availability. Lian et al. (2020) reported that ge­
population for meat and dairy products, crop production must be notypes of wheat from the 2000s and the 2010s in China showed an
increased. The usage of nitrogen (N) fertilizer has risen approximately increased NUE under abiotic stresses (water deficiency and N limita­
10-fold over the last 50 years to ensure food security and crop yields tion). The practical potential of genetically improved crop cultivars
worldwide (Chen et al., 2015; Zhang et al., 2016). China is the largest cannot be achieved if their NUE has not been explored. Therefore,
consumer of N fertilizer, with farmers becoming accustomed to overuse cultivating low-N-tolerant and N-efficient crop cultivars is the most
for obtaining high yields; however, such practices simultaneously lead efficient method of improving yields, especially in arid and semiarid
to severe biodiversity loss and environmental problems such as global regions.
warming, greenhouse gas emissions, and lake eutrophication (Guo et al., Crop productivity depends on not only sufficient nutrition but also
2010; Zhang et al., 2013). Hence, it is critical to explore strategies to crucial environmental factors, such as soil water absorption and solar
decrease crop N application rates whilst maintaining high crop yields radiation. Nutrients and water are usually co-limited, and insufficient
and product quality. fertilizer use becomes even more serious in drylands due to water supply
Nitrogen plays an important role in plant growth since it is a con­ limitations. When a rational application of nitrogen fertilizer is used, the
stituent of cell walls, membranes, amino acids, proteins, and nucleic ratio of water consumed by plant transpiration to that lost by soil
acids (Kusano et al., 2011; Marschner, 2011). Efficient N uptake and evaporation increases from 0.32 to 0.65, reducing water loss by one-
utilization in grain production requires effective N transport, assimila­ third and a significantly increasing crop water use efficiency (WUE)
tion, and redistribution processes. Although crop yields are significantly (Li et al., 2009). Photosynthesis is strongly affected by photosyntheti­
improved by increasing N application, the higher rates of N application cally active radiation (PAR) (Zhu et al., 2010). Plant dry matter accu­
have sharply decreased crop N use efficiency (NUE) (Fan et al. 2012). mulation reflects the efficiency of photosynthesis through the
NUE is a complex term, and mainly involves N uptake efficiency (NUPE) conversion of intercepted radiation into biomass (Ashraf and Harris,
(Moll et al., 1982), and N productive efficiency (NPE) (Gallais and Hirel, 2013). Although the Loess Plateau region of China receives abundant
2004). It is well known that N-efficient cultivars of various crops sunlight, poor radiation use efficiency (RUE) is common because of
inappropriate agricultural cultivation and management practices (Qin
Table 1 et al., 2018; Zhang et al., 2019b). Hence, rational N fertilization is
The soil fertility data in 0–20 cm soil layer before sowing in 2019 and 2020. crucial to guarantee food supply and sustainable social development in
dryland agricultural areas by improving the conversion efficiency of
Year pH Organic Total Total Total
matter nitrogen (g phosphorus (g potassium (g environmental resources into crop yield.
content (g kg− 1) kg− 1) kg− 1) Broomcorn millet (Panicum miliaceum L.) is one of the oldest Chinese
kg− 1) domesticated cereals. It has viable agronomic traits including a short
2019 8.6 4.80 0.31 0.47 19.43 growth cycle, low water requirements, high nutrient use efficiency, and
2020 8.6 5.12 0.35 0.56 22.36 good resistance to various stresses (Liu et al., 2016; Yuan et al., 2020).

Fig. 1. The chlorophyll content (SPAD value) of broomcorn millet under different N application rates in 2019 and 2020. N0, N75, and N150 represent N fertilizer
application rates of 0, 75, and 150 kg N ha− 1 for broomcorn millet, respectively. Different letters indicate significant differences (P < 0.05) among different varieties.
The error bars indicate the standard errors of the means (n = 3).

2
C. Liu et al. Agricultural and Forest Meteorology 308-309 (2021) 108616

Fig. 2. The net photosynthetic rate and transpiration rate of broomcorn millet under different N application rates in 2019 and 2020. N0, N75, and N150 represent N
fertilizer application rates of 0, 75, and 150 kg N ha− 1 for broomcorn millet, respectively. Different letters indicate significant differences (P < 0.05) among different
varieties. The error bars indicate the standard errors of the means (n = 3).

Moreover, this minor grain crop is a vital food source and forage species,
popular for cultivation in the Great Wall of China region, Russia, India,
the Middle East, and Europe (Zhang et al., 2019a). The release of its
genome sequence by the China Agricultural University (Shi et al., 2019),
Purdue University, and Chinese Academy of Sciences (Zou et al., 2019)
has attracted more interest from researchers. For example, scientists are
breeding broomcorn millet varieties that use soil N more efficiently (Xu
et al., 2012). Although the physiological and molecular mechanisms of
low-N-tolerant cultivars have been extensively studied, most experi­
ments were carried out under laboratory conditions (Shah et al., 2017;
Luo et al., 2019).
In this study, we hypothesized that low-N-tolerant broomcorn millet
cultivars would maintain high resource (radiation, water, and N) use
efficiency and productivity stability under field conditions. We con­
ducted a field study with the following objectives: i) explore the changes
in the leaf photosynthetic characteristics and dry matter accumulation
and transportation in different low-N-tolerant broomcorn millet culti­
vars provided with low N input over time; ii) compare the resource
capture and use efficiency and grain yield in order to provide a theo­
retical basis for breeding broomcorn millet cultivars with low-N-
tolerance and developing green agriculture.
2. Materials and methods
2.1. Location description
This field study was conducted in 2019 and 2020 at the Modern
Agricultural Science and Technology Demonstration Park in Yulin,
Shaanxi, China, located 1,120 m a.s.l. (109◦ 21′ 46.0′′ E longitude,
37◦ 56′ 26.0′′ N latitude). This area is characterized by a semiarid conti­
nental monsoonal climate, and the average annual air temperature and
precipitation in the test region were 8.3◦ C and 365 mm, respectively
(Fig. S1). The soil had a loess-like loam texture. The soil fertility data
recorded in the 0–20 cm soil layer before sowing are presented in
Table 1.
2.2. Field management and research design
The field study was conducted in a split-plot design: 20 m2 (5 m × 4
m) with 0.40 m row spacing with three replications. The main plots were
N fertilization (urea, N > 46%) rates, and the subplots were the culti­
vars. The following N levels were used: (1) 0 kg N ha− 1 (N0), (2) 75 kg N
ha− 1 (N75), and (3) 150 kg N ha− 1 (N150). N150 was the control based
on local farmers’ traditional fertilization practices. Four low-N-tolerant
cultivars broomcorn millet cultivars (86, 184, 235, and 298) and two
low-N-sensitive cultivars (230 and 111) were chosen based on previous
laboratory experiments (Liu et al., 2020). Seeds were sown on June 11,
2019, and June 12, 2020, and mature grains were harvested from
September 20 to October 7, 2019, and 2020, depending on the fullness
of the grains. The plant density was 400,000 plants ha− 1. Before planting
each year, uniform fertilization at 100 kg P2O5 ha− 1 and 75 kg K2O ha− 1
was applied to all plots as basal fertilizer, and no fertilizer or artificial
irrigation was applied during the growth period.

Fig. 3. The stomatal conductance and intercellular CO2 concentration of broomcorn millet under different N application rates in 2019 and 2020. N0, N75, and N150
represent N fertilizer application rates of 0, 75, and 150 kg N ha− 1 for broomcorn millet, respectively. Different letters indicate significant differences (P < 0.05)
among different varieties. The error bars indicate the standard errors of the means (n = 3).

3
C. Liu et al.
Fig. 4. The dry matter accumulation of broomcorn millet under different N application rates in 2019 and 2020. N0, N75, and N150 represent N fertilizer application
rates of 0, 75, and 150 kg N ha− 1 for broomcorn millet, respectively. The error bars indicate the standard errors of the means (n = 3)
2.3. Sampling and measurements
2.3.1. Chlorophyll content and photosynthetic characteristics
At the jointing, heading, flowering, and grain-filling stages, three
individual plants were selected from each plot, and a Minolta SPAD-502
chlorophyll meter (Tokyo, Japan) was used to measure the chlorophyll
content (SPAD value) between 09:00 and 11:00 on cloudless days using
the topmost fully opened leaves. At the same time, the net leaf photo­
synthetic rate (Pn), transpiration rate (Tr), stomatal conductance (gs),
and intercellular CO2 concentration (Ci) were measured under a PAR of
1,200 μmol m− 2 s− 1 using a CIRAS-3 portable photosynthesis system
(Massachusetts, USA). All measurements were conducted at 400 μmol
mol− 1 atmospheric CO2 concentration and 30◦ C leaf temperature.
2.3.2. Dry matter accumulation and transportation
Similar plants were randomly selected at the different growth stages
from each plot, cut to ground level, and dried at 75◦ C for 48 h to
determine dry matter accumulation. The amount of dry matter trans­
ported (DTA, g plant− 1), the dry matter transportation rate (DTR, %),
and the contribution rates of vegetative organs (stem, leaf, sheath) to the
grains (GCR, %) were calculated using the following equations (Gong
et al., 2020):
DTA = LDM − DMM,
DTA
DTR = × 100%,
LDM
and
DTA
GCR = × 100%,
GDM
where LDM (g plant− 1) is the largest dry mass of the stem, leaf, or
sheath; DMM (g plant− 1) is the dry mass of the same vegetative organ at
maturity; and GDM (g plant− 1) is the dry mass of the grain.
4
Agricultural and Forest Meteorology 308-309 (2021) 108616
2.3.3. IPAR and RUE
It was assumed that 50% of the total incident solar radiation is PAR,
and the amount of PAR intercepted by the plant canopy (IPAR, MJ m− 2)
was computed using the following exponential function (Gong et al.,
2021):
∑ ( )
IPAR = 0.5R 1 − e− kLAI ,
where k is the light extinction coefficient (set at 0.65) (Dunan et al.,
1999); R is the daily solar radiation (MJ m− 2 d− 1); and LAI is the leaf
area index.
We calculated RUE (g MJ− 1) according to Liu et al. (2010) using the
following formula:
RUE = Biomass yield/IPAR,
where biomass yield is the dry matter accumulation during the entire
growth period.
2.3.4. WUE
The soil water content (SWC, %) was measured after harvest at soil
depths of 10 cm, 20 cm, 30 cm, 40 cm, 50 cm, 60 cm, 70 cm, 80 cm, 100
cm with an auger (5.5 cm diameter). The soil samples were oven-dried at
105◦ C to a constant weight.
The WUE (kg ha− 1 mm− 1) of broomcorn millet was calculated as
follows:
WUE = Grain yield/evapotranspiration
The evapotranspiration (mm) during the entire growing season was
determined as described by Gong et al. (2021).
2.3.5. NUE
The N contents of the stems, leaves, sheaths, and spikes at maturity
were measured using an automatic Kjeldahl N determination apparatus
(K9860, Hanon, China). The plant N uptake and NUE were determined
with the following equations (Zhang et al., 2019b; Gong et al., 2021):
 C. Liu et al.
Table 2
Dry matter distribution of broomcorn millet under different N application rates in 2019 and 2020.
Year N Treatment Cultivar Jointing stage Heading stage Flowering stage Filling stage Maturity stage

Stem Leaf Sheath Stem Leaf Sheath Spike Stem Leaf Sheath Spike Stem Leaf Sheath Spike Stem Leaf Sheath Spike

2019 N0 86 16.45bc 56.65ab 26.90ab 41.30a 31.62bc 15.53b 13.64a 45.18ab 26.18a 10.32a 18.32c 39.58ab 15.47b 7.47a 37.48bc 22.83ab 6.84b 3.55ab 66.78bc
184 17.32ab 54.85ab 27.82ab 40.17ab 29.40c 20.63a 13.33abc 47.52a 23.78bc 10.09a 18.61bc 42.34a 15.20bc 7.23a 35.23c 22.99a 7.71a 3.64ab 65.65c
235 15.55c 52.47b 31.98a 38.66abc 31.37bc 21.76a 12.14bc 45.99ab 23.41c 10.68a 19.92abc 40.98a 14.36c 7.24a 37.42bc 23.75a 6.81b 3.80a 65.64c
298 15.04c 53.38b 31.57a 37.38c 33.55ab 18.52ab 13.45ab 43.57bc 25.44ab 10.98a 20.02abc 37.78b 15.01bc 7.11a 40.09ab 22.60ab 6.54bc 3.21b 67.65abc
230 18.19a 56.54ab 25.26b 35.93c 34.89a 19.14a 13.12abc 41.19c 27.20a 10.32a 21.29a 36.49b 17.11a 5.45b 40.95a 21.18b 6.01c 3.21b 69.60a
111 17.70ab 58.45a 24.18b 37.84bc 34.40a 18.74ab 11.98c 43.27bc 26.40a 10.13a 20.20ab 37.43b 16.92a 5.96b 39.68ab 21.66b 6.98b 3.33ab 68.03ab
N75 86 19.29c 50.83abc 29.88ab 39.50ab 30.59b 18.01ab 14.65a 46.85a 23.85b 9.50a 19.80abc 40.40abc 14.77abc 6.63b 38.19a 24.07bc 7.22a 3.62a 65.10ab
184 23.69a 48.13bc 28.17ab 39.68ab 29.54b 19.70a 14.32ab 44.86ab 26.78a 9.65a 18.71cd 41.92ab 16.03a 7.25ab 34.81b 23.26c 7.12a 3.78a 65.85a
235 21.41b 51.36ab 27.23b 38.63b 30.93b 18.20ab 15.05a 46.37ab 25.35ab 9.65a 18.64d 43.14a 15.26ab 6.64b 34.96b 23.58c 7.06a 3.56a 65.80a
298 22.26ab 47.10c 30.64a 39.81ab 30.70b 16.44bc 15.37a 46.30ab 24.12ab 10.17a 19.40bcd 38.50c 14.13bc 7.29ab 40.08a 24.91abc 6.51b 3.68a 64.91ab
230 17.64d 52.36a 29.99ab 38.97b 33.87a 15.16c 14.00a 44.32b 25.65ab 9.34a 20.69a 39.45bc 13.55c 6.97ab 40.03a 25.53ab 6.35b 3.80a 64.32ab
111 17.54d 54.27a 28.18b 41.57a 33.27a 14.42c 12.57b 44.05ab 26.15ab 9.51a 20.30ab 39.93bc 14.00bc 7.32a 38.74a 26.47a 6.74ab 3.70a 63.09b
N150 86 22.01a 49.43a 28.56c 38.59a 31.57ab 16.65bc 15.57a 50.64a 24.19a 9.10b 16.07b 41.83a 15.09a 5.98c 37.09b 24.50a 6.31b 3.40c 65.78a
184 22.97a 47.62a 29.41bc 37.23a 32.22ab 18.69a 14.81a 48.41abc 24.42a 9.49b 17.68a 39.40bc 14.06b 6.62ab 39.92a 24.07a 7.25a 4.07a 64.61a
235 21.33a 46.77a 31.90a 37.93a 32.52ab 16.82b 15.16a 49.37ab 23.19a 8.98b 18.47a 40.82ab 12.76c 6.51ab 39.90a 23.95a 6.99a 3.59c 65.47a
298 22.49a 46.53a 30.98ab 39.21a 31.64ab 15.23c 15.94a 48.66abc 23.36a 10.26a 17.72a 38.47bc 15.19a 6.21bc 40.13a 25.66a 6.07b 3.55c 64.72a
230 20.59a 47.86a 31.55ab 36.66a 33.37a 16.23bc 16.01a 46.59c 24.28a 10.60a 18.54a 37.71c 15.19a 6.77a 40.33a 25.13a 5.59c 3.86b 65.41a
111 22.50a 45.52a 32.32a 38.43a 31.29b 17.36ab 15.49a 47.77bc 22.79a 10.58a 18.86a 39.25bc 14.24ab 6.73a 39.77a 25.48a 6.13b 3.50c 64.89a
2020 N0 86 19.79c 53.77a 26.44a 25.18bc 38.46cd 24.21b 12.15a 47.34cd 25.86a 12.23bc 14.57bc 47.38bc 12.50bc 7.75a 32.37ab 27.12a 6.68b 4.11bc 62.09b
184 20.33bc 53.53a 26.14a 26.97ab 37.11e 24.35b 11.57a 44.72e 25.73a 13.04a 16.52a 45.85c 12.64abc 7.64ab 33.87a 25.02ab 7.62ab 4.72ab 62.63b
235 20.71abc 52.43a 26.85a 26.33ab 39.30c 23.12bc 11.25ab 46.281d 26.01a 12.79ab 14.92abc 46.96bc 12.05c 7.12ab 33.87a 21.72c 7.73ab 5.05a 65.50a
5

298 20.34bc 53.78a 25.88a 23.76c 38.23d 26.30a 11.71a 47.87c 23.32b 12.92ab 15.88ab 50.94ab 12.24bc 7.65ab 29.17bc 23.82bc 7.30ab 4.59ab 64.29ab
230 22.54a 52.66a 24.79a 27.07a 41.47a 21.38c 10.08b 52.69a 23.31b 11.41d 12.59d 50.54ab 13.33a 6.87b 29.26bc 25.36ab 8.45a 3.61c 62.58b
111 21.86ab 52.02a 26.12a 25.54abc 40.41b 22.90bc 11.16ab 49.84b 24.82a 11.58cd 13.76cd 51.79a 12.98ab 7.34ab 27.89c 24.77b 7.92a 4.27bc 63.05b
N75 86 21.53b 51.96a 26.51b 26.40b 34.19c 25.04ab 14.37a 43.89bc 22.74c 15.35ab 18.02ab 44.54b 14.31a 7.72a 33.43ab 28.07a 7.08b 5.36ab 59.50b
184 21.32b 49.44b 29.23a 26.90b 35.21bc 23.86bc 14.03ab 39.66e 26.57a 16.16a 17.61ab 41.36c 14.81a 8.21a 35.63a 24.98b 7.64b 5.80a 61.57ab
235 20.96b 49.42b 29.61a 27.16b 35.80b 23.95b 13.08bc 41.46d 25.12ab 15.06b 18.35ab 42.45bc 15.42a 8.04a 34.08ab 24.85b 7.59b 5.66a 61.90a
298 23.29a 51.59ab 25.11b 26.17b 35.31bc 25.74a 12.78c 42.83c 22.62c 15.94a 18.61a 42.16bc 14.74a 8.42a 34.68ab 24.17b 7.83ab 5.27ab 62.73a
230 22.38ab 52.49a 25.13b 29.38a 37.43a 22.73c 10.46d 47.68a 24.29bc 12.83c 15.20c 47.70a 11.57b 6.33b 34.40ab 25.40b 8.85a 4.13c 61.62ab
111 23.93a 51.63ab 24.44b 26.90b 36.98a 24.10b 12.01c 44.74b 24.81ab 13.29c 17.15b 40.89b 11.89b 7.53a 32.12b 25.28b 8.08ab 4.69bc 61.95a
N150 86 20.30bc 51.17a 28.52c 26.90b 34.45bc 23.05b 15.60a 39.30b 26.16ab 16.38a 18.16b 40.11b 17.80ab 8.61ab 32.69ab 26.47ab 9.07a 6.34a 58.12c
184 19.96c 50.46a 29.58abc 25.48d 33.52c 25.89a 15.11ab 38.59b 28.43a 15.22bc 17.76b 39.34b 17.06a 8.78ab 34.05a 26.67ab 9.22a 6.09a 58.02c

Agricultural and Forest Meteorology 308-309 (2021) 108616

235 22.27a 48.13bc 29.60abc 27.22b 34.53bc 23.12b 15.13ab 39.14b 27.04ab 15.69ab 18.12b 44.31a 16.56ab 9.42ab 34.68a 27.32a 9.06a 5.70ab 57.93c
298 20.44bc 49.14b 30.42ab 26.68bc 34.78b 23.68b 14.86abc 39.57b 25.76b 15.01bc 19.66a 44.25a 15.92b 8.44b 31.33bc 24.71bc 8.90a 5.90a 60.49b
230 21.75ab 48.97bc 29.28bc 29.28a 35.05ab 21.82c 13.84c 42.12a 25.95b 14.71cd 17.22b 44.03a 16.23ab 9.65a 29.88c 23.27c 8.17b 4.77c 63.79a
111 21.23abc 47.85c 30.92a 26.12cd 35.89a 23.76b 14.23bc 41.66a 26.21ab 13.96d 18.17b 40.89b 15.86b 9.14ab 30.96bc 24.33c 8.92a 5.05bc 61.71b
Variation source
Year (Y) *** ns *** *** *** *** *** *** * *** *** *** *** *** *** *** *** *** ***
N Treatment (N) *** *** *** *** *** *** *** *** ns *** *** *** *** *** ** *** *** *** ***
Cultivar (C) * ** ** ns *** *** *** *** *** *** *** * ** *** ns *** *** *** ***
Y×N *** *** ns ** *** *** * *** *** *** *** *** *** *** *** *** *** *** **
Y×C *** ** * *** ns *** *** *** *** *** *** *** *** ns *** *** *** *** **
N×C *** ** *** *** *** *** * ns *** *** * * *** *** * ** *** ns **
Y×N×C *** ns *** ns ** *** ns ** ns * *** ns * *** ns ns ** * ***
1
Values followed by a different letter within the same column are significantly different at P < 0.05. N0, N75, and N150 represent N fertilizer application rates of 0, 75, and 150 kg N ha− for broomcorn millet, respectively.
*, **, and *** significant at the 0.05, 0.01, and 0.001 probability levels, respectively. ns, no significant difference.
C. Liu et al. Agricultural and Forest Meteorology 308-309 (2021) 108616

Table 3
Dry matter translocation and contribution rate to grain yield of every organ in broomcorn millet under different N application rates in 2019 and 2020.
Year N Treatment Cultivar Stem Leaf Sheath

DTA (g plant )
–1
DTR (%) GCR (%) DTA (g plant )
–1
DTR (%) GCR (%) DTA (g plant–1) DTR (%) GCR (%)

2019 N0 86 1.24c 12.71d 10.91b 1.21b 33.31d 6.51c 0.44c 27.64bc 2.31cd
184 2.82a 24.87a 14.67a 1.21b 29.94e 6.30c 0.59a 30.49ab 3.39a
235 1.95b 18.79b 10.69b 1.22b 32.81de 7.73b 0.48bc 26.21c 2.66bc
298 0.98d 11.32de 6.50d 1.21b 35.44c 6.80bc 0.54ab 33.20a 3.02ab
230 0.68e 9.82e 6.14d 1.44a 45.53a 9.62a 0.09e 8.51e 0.89e
111 1.24c 16.90c 8.79c 1.34ab 40.75b 9.96a 0.23d 19.76d 1.81d
N75 86 2.00c 21.89a 10.63b 1.39b 31.06b 6.09a 0.47ab 23.08c 2.04c
184 2.43b 18.74b 9.89c 1.73a 35.03a 7.03a 0.53a 23.23c 3.13ab
235 2.98a 22.37a 12.78a 1.61a 34.48ab 6.85a 0.49ab 23.96bc 1.97c
298 0.67e 6.36d 3.22f 1.28bc 33.17ab 6.91a 0.54a 26.99ab 2.60bc
230 1.05d 10.51c 6.34d 1.18c 35.45a 7.04a 0.43b 27.99a 2.73abc
111 0.75e 9.12cd 5.30e 1.16c 32.65ab 7.03a 0.55a 29.43a 3.51a
N150 86 2.86b 17.99b 12.51a 2.28c 42.24b 8.32b 0.46c 21.37c 1.84c
184 2.95b 19.45ab 11.00b 1.72e 32.02c 6.61c 0.48c 18.95c 1.91c
235 3.42a 22.14a 12.31a 1.31f 27.36d 5.42c 0.66a 26.98b 2.39b
298 2.22c 14.95c 9.83c 2.76a 49.43a 11.00a 0.63ab 27.65b 2.52b
230 1.52e 12.29c 8.59d 2.52b 51.68a 11.17a 0.54bc 25.01b 2.62ab
111 1.73d 12.87c 7.22e 2.02d 42.35b 8.42b 0.69a 30.46a 2.83a
2020 N0 86 5.43c 31.25c 24.47c 1.65a 36.06a 7.43a 1.04a 36.51a 4.66a
184 7.66b 39.99b 33.06b 1.79a 34.09a 7.64a 1.04a 30.82bc 4.41a
235 9.75a 52.19a 44.68a 1.62a 33.71a 7.43a 0.76b 25.98d 3.51b
298 7.94b 44.58ab 36.74ab 1.25b 29.23ab 5.80a 0.77b 28.74cd 3.55b
230 5.73c 40.82b 37.36ab 0.94c 25.26b 6.04a 0.72b 37.98a 4.69a
111 7.14b 45.82ab 33.69a 1.21b 31.10ab 7.42a 0.76b 34.47ab 4.66a
N75 86 8.29a 35.68a 31.11abc 3.70a 49.49a 13.85a 1.18b 29.16ab 4.43b
184 8.19a 36.08a 36.69c 3.68a 45.40a 12.01b 1.13bc 25.12b 3.69b
235 8.38a 38.44a 29.68bc 3.84a 48.28a 13.49ab 1.08bc 26.06b 3.80b
298 8.23a 41.64a 33.11abc 3.17b 45.93a 12.73ab 1.42a 35.94a 5.40a
230 7.33a 41.79a 37.32a 0.70d 19.73c 4.59c 0.67d 28.38ab 4.43b
111 8.06a 42.09a 36.37ab 1.15c 24.55b 5.32c 0.91c 30.70ab 4.14b
N150 86 6.83c 28.97b 21.62c 4.52a 43.97a 14.28a 0.95c 19.14d 2.67d
184 7.18c 28.46b 20.96c 4.49a 41.91a 13.13ab 1.40b 25.18c 4.09c
235 5.38d 23.10b 16.33d 3.87b 39.24a 11.74bc 1.84a 32.78b 5.58b
298 10.54a 41.53a 33.78a 3.77bc 41.35a 12.05bc 1.30b 26.89c 4.14c
230 8.18b 40.39a 29.13b 3.20cd 42.80a 10.43c 1.93a 43.99a 6.87a
111 8.70b 39.62a 30.50b 3.05d 38.40a 11.42c 1.81a 39.72a 6.04b
Variation source
Year (Y) *** *** *** *** ns *** *** *** ***
N Treatment (N) *** *** *** *** *** *** *** ns ***
Cultivar (C) *** *** ** *** *** *** *** *** ***
Y×N *** *** *** *** *** *** *** *** ***
Y×C *** *** *** *** *** *** ns ** ***
N×C *** *** *** *** *** *** *** *** ***
Y×N×C *** *** *** *** *** *** * *** ***

Values followed by a different letter within the same column are significantly different at P < 0.05. N0, N75, and N150 represent N fertilizer application rates of 0, 75,
and 150 kg N ha− 1 for broomcorn millet, respectively. *, **, and *** significant at the 0.05, 0.01, and 0.001 probability levels, respectively. ns, no significant
difference.

N uptake = Dry matter accumulation × N content, 2.4. Statistical analyses

NUE = Grain yield/N uptake,
NHI = Grain N uptake/plant N uptake,
NUPE = N uptake/N application rate,
and NPE = Grain yield/N application rate,
where N uptake (kg ha− 1) is the whole-plant N uptake (sum of the stem,
leaf, sheath, and spike N contents), NUE (kg kg− 1) is the N use efficiency,
NHI (%) is the N harvest index, NUPE (kg kg− 1) is the N uptake effi­
ciency, and NPE (kg kg− 1) is the N productive efficiency.
The data from each year were subjected to analysis of variance
(ANOVA) using the SPSS 19.0 program (SPSS Inc., Chicago, USA) to
evaluate the effects of cultivar, N treatment, and their interactions with
all measured parameters. Origin Pro 2018 was used to prepare all fig­
ures. Mean values were compared using Duncan’s multiple range test (P
< 0.05).
3. Results

2.3.6. Grain yield and harvest index 3.1. Photosynthetic characteristics

Broomcorn millet was manually harvested at 12–14% moisture
content, and the grain yield in each plot was recorded according to the
actual harvested area. Five plants were randomly selected from each
plot and their 1,000-grain weights were determined. The harvest index
(HI) is the ratio of grain yield to biomass yield.
Low N treatments (N0 and N75) decreased the chlorophyll content
(SPAD value) (Fig. 1), Pn, Tr (Fig. 2), gs, and Ci (Fig. 3) of broomcorn
millet by 7.0%, 14.9%, 31.5%, 19.2%, and 29.1%, respectively,
compared with the N150 treatment over the average of two years and six
cultivars. The mean SPAD, Pn, Tr, gs, and Ci values of 86, 184, 235, and
298 (low-N-tolerant cultivars) were 6.2%, 17.1%, 20.9%, 14.9%, and
19.6% higher, respectively, than those of 230 and 111 (low-N-sensitive

6
C. Liu et al.
Fig. 5. The intercepted photosynthetically active radiation (IPAR) and radiation use efficiency (RUE) of broomcorn millet under different N application rates in 2019
and 2020. N0, N75, and N150 represent N fertilizer application rates of 0, 75, and 150 kg N ha− 1 for broomcorn millet, respectively. Different letters indicate
significant differences (P < 0.05) among different varieties.
cultivars) under low-N conditions.
3.2. Dry matter accumulation and transportation
Significant differences were observed in response to different N
application rates at all growth stages (Fig. 4). At maturity, the average
dry matter accumulation of 86, 184, 235, 298, 230, and 111 in the N75
treatment were 21.2%, 25.4%, 27.4%, 15.7%, 20.1%, and 27.9% higher,
respectively, than those in N0. Further, the average dry matter accu­
mulation of 86, 184, 235, 298, 230, and 111 in the N150 treatment were
42.1%, 43.7%, 50.8%, 41.3%, 53.1%, and 60.2% higher, respectively,
than those in N0.
The effects of low N stress on the dry matter distribution to the stem,
leaf, sheath, and spike in 230 and 111 were greater than those in 86,
184, 235, and 298 (Table 2). The dry matter accumulation in the spike
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Agricultural and Forest Meteorology 308-309 (2021) 108616
increased to promote grain filling with the reproductive growth of the
plant.
At the early growth stage, low N treatments restricted the develop­
ment of a large photosynthetic source and significantly weakened the
transfer of photosynthetic products from the stems, leaves, and sheaths
to the grains (Table 3). On average, DTA values under low N stress
corresponding to the stem, leaf, and sheath decreased by 14.0%, 86.6%,
and 48.0%, respectively, compared to those under N150. Further, the
DTR and GCR were 13.9% and 1.2% lower from the leaves and 28.0%
and 7.9% lower from the sheaths, respectively. Across the two study
years, the low-N-tolerant cultivars transferred significantly more
photosynthetic products from stems, leaves, and sheaths to grains than
the low-N-sensitive cultivars did under the N0 treatment. This was
observed by the 23.4% higher DTA, 5.3% higher DTR, and 1.8% higher
GCR stem dry matter accumulation. Similarly, these values were 72.9%,
C. Liu et al.
Fig. 6. The soil water content (SWC) of broomcorn millet under different N application rates in 2019 and 2020. N0, N75, and N150 represent N fertilizer application
rates of 0, 75, and 150 kg N ha− 1 for broomcorn millet, respectively. The error bars indicate the standard errors of the means (n = 3).
15.2%, and 15.3% higher from the leaf sheath and 43.3%, 4.3%, and
1.5% higher from the sheath, respectively in the low-N-tolerant cultivars
than in the low-N-sensitive cultivars under the N0 treatment (Table 3).
3.3. IPAR and RUE
The IPAR and RUE of the broomcorn millet canopy were significantly
decreased due to low N fertilization (Fig. 5). The IPAR in the N0 and N75
treatments decreased by 9.9% and 12.7% in 2019 and 2.0% and 4.5% in
2020, respectively, compared to that in the N150 treatment. In addition,
low-N stress decreased the RUE by 38.9% in 2019 and 49.7% in 2020
under N0, and 13.0% in 2019 and 29.6% in 2020 under N75 compared
with N150. Among the different cultivars, the RUE values of low-N-
tolerant cultivars were significantly higher than those of the low-N-
sensitive cultivars (P < 0.05).
3.4. SWC and WUE
SWC was significantly affected by low N application rates and
exhibited spatial variation (Fig. 6). During the plant maturity stage in
both years, the SWC in the N0 and N75 treatments decreased dramati­
cally and was lower than that in N150 in most of the 0–100 cm soil
layers. For the 0–100 cm soil profiles, the mean SWC values of the low-
N-tolerant cultivars were 45.8% higher than those of the low-N-sensitive
cultivars under low N conditions. SWC strongly influenced the varia­
tions in other water characteristics. Compared to the N150 treatment,
WUE was 20.7% and 40.0% lower in the N0 and N75 treatments
(Table 5). The mean WUE of the low-N-tolerant cultivars (12.5 kg ha− 1
mm− 1) was significantly higher than that of the low-N-sensitive cultivars
(8.0 kg ha− 1 mm− 1).
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Agricultural and Forest Meteorology 308-309 (2021) 108616
3.5. N uptake and NUE
During the entire 2019 growth period, there was a 97.4% and 42.4%
reduction in the total N uptake in the N0 and N75 treatments, respec­
tively, compared to the N150 treatment, while in 2020 the decreases
were 108.2% and 48.9% (Fig. 7). There were significant differences in
the N uptake between the six cultivars. On average, the N uptake of the
low-N-tolerant cultivars were 49.1% higher in 2019 and 51.9% higher in
2020 than those of the low-N-sensitive cultivars.
The changes in N uptake affected NUE to some extent. NUE and NHI
were 20.3% and 5.5% higher, respectively, under N0 and 8.6% and 4.8%
higher, respectively, under N75 than under N150 across the two years of
the study (Table 4). Increased NUE and NHI, indicated an enhanced
translocation of N from vegetative organs to grains. Additionally, there
were no significant differences in average NUE and NHI values between
the low-N-tolerant cultivars and the low-N-sensitive cultivars under the
N0 and N75 treatments. However, the low-N-tolerant cultivars had
significantly higher NUPE (34.8%) and NPE (38.8%) values than the
low-N-sensitive cultivars did across the study years and N treatments.
3.6. Grain yield and HI
Low N fertilization consistently and significantly decreased produc­
tivity compared to N150 in both study years. The 1,000-grain weight
and grain yield decreased by 7.2% and 36.5% in 2019, respectively, and
by 5.8% and 30.2% in 2020 under low N stress (Table 5). Furthermore,
the six cultivars showed significant differences in grain yield. On
average, the yield of the low-N-tolerant cultivars was 50.1% higher in
2019 and 40.1% higher in 2020 than that of the low-N-sensitive culti­
vars under low N conditions. More specifically, cultivars 184 and 111
C. Liu et al.
Fig. 7. The N uptake of broomcorn millet under different N application rates in 2019 and 2020. N0, N75, and N150 represent N fertilizer application rates of 0, 75,
and 150 kg N ha− 1 for broomcorn millet, respectively. Different letters indicate significant differences (P < 0.05) among different varieties. The error bars indicate
the standard errors of the means (n = 3).
produced the maximum and minimum grain yields, respectively. The
same trends were observed for the HI. Hence, the productivity response
of low-N-tolerant cultivars to N fertilization was superior to that of low-
N-sensitive cultivars.
4. Discussion
4.1. Leaf photosynthetic characteristics and dry matter accumulation
The low N treatments noticeably altered the leaf gas exchange pa­
rameters, including decreasing SPAD (7.0%), Pn (14.9%), and Tr
(31.5%) (Figs. 1 and 2) compared with the N150 treatment, indicating
that low N stress weakens the plant assimilate supply capacity by
reducing photosynthetic physiological metabolic activity. During the
latter stages of growth, the low N treatments markedly induced leaf
stomatal closing, as shown by the decreased gs and Ci (Fig. 3), which
contributed to a decline in Pn and Tr values. The gs strongly affects the
exchange of CO2 and H2O between plant leaves and the atmospheric
environment (Ning et al., 2018), and increasing gs is a physiological
mechanism by which plants stimulate the transformation of CO2 into
carbohydrates. The close relationship between Pn and gs showed that
low N fertilization weakened the functioning of the photosynthetic
apparatus and resulted in decreased CO2 assimilation (Fang et al., 2018).
Notably, under low N conditions (N0 and N75), the photosynthetic pa­
rameters of the low-N-tolerant cultivars were significantly higher than
those of the low-N-sensitive cultivars (P < 0.05). Specifically, SPAD, Pn,
Tr, gs, and Ci were 6.2%, 17.1%, 20.9%, 14.9%, and 19.6% higher,
respectively. This increase may reflect the intensity of the photosyn­
thetic utilization ability of different broomcorn millet cultivars under N
deficiency. The photosynthetic capacity of leaves is mainly dependent
on the nutrient level of the substrate because the photochemical reaction
is characterized by the oxidation of water molecules in the light-driven
reactions, and the released electrons are transported to the thylakoid
cavity, where ATP is eventually formed (Moriwaki et al., 2019). These
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Agricultural and Forest Meteorology 308-309 (2021) 108616
results implied that the low-N-tolerant genotypes maintained a stronger
ability to absorb and assimilate light than the low-N-sensitive genotypes.
The main reasons for these differences may be related to the biological
characteristics of the leaves, such as their greater ability to use light
energy and increased CO2 assimilation (Britto and Kronzucker, 2002;
Luo et al., 2013).
Dry matter accumulation and distribution are integrated reflections
of individual plant accumulation quantity and plant population quality
and may influence the response of grain yield to the availability of nu­
trients (Vega et al., 2000). Low N input significantly restrained the
optimization of dry matter accumulation and translocation in the
aboveground tissues of broomcorn millet (Table 2, Fig. 4). The DTA,
DTR, and GCR from the stem, leaf, and sheath were significantly lower
than those in the N150 treatment in the two low N treatments (Table 3).
More importantly, the low-N-tolerant cultivars transferred significantly
more photosynthetic products from the stems, leaves, and sheaths to
grains than the low-N-sensitive cultivars, consistent with the results of a
previous study (Zhao et al., 2005). Some cultivars such as alfalfa that
were tolerant of N deficiencies maintained adequate photosynthesis to
increase the redistribution of nutrients to the leaves (Fiasconaro et al.,
2013). Our results indicated that increased dry matter characteristics
were consistently associated with leaf photosynthetic capacity and the
concentration of soil N (Table S2). The vigorous physiological meta­
bolism of the low-N-tolerant cultivars helped to enhance the transfer of
photosynthetic products.
4.2. Resource use efficiency and grain yield
Low N treatments result in less productive crops than nutrient-
abundant conditions often due to less efficient capture or use of light
(Moriwaki et al., 2019). The productivity disadvantage of broomcorn
millet under low N stress was attributed primarily to a combination of
these two factors as significant decreases in IPAR and RUE were
observed (Fig. 5). This is due to the lower leaf area index restraining the
C. Liu et al. Agricultural and Forest Meteorology 308-309 (2021) 108616

Table 4 Table 5
N use efficiency (NUE), N harvest index (NHI), N uptake efficiency (NUPE) and Grain yield, HI, and WUE of broomcorn millet under different N application
N productive efficiency (NPE) of broomcorn millet under different N application rates in 2019 and 2020.
rates in 2019 and 2020. Year N Cultivar 1000− grain Grain HI WUE
Year N Cultivar NUE(kg NHI(%) NUPE NPE(kg Treatment weight (g) yield (kg (kg
Treatment kg− 1) (kg kg− 1) ha− 1) ha− 1
kg− 1) mm− 1)

2019 N0 86 18.18a 51.62bc − − 2019 N0 86 8.0714a 3647.8b 0.26b 12.27b

184 16.48ab 43.53c − − 184 8.2161a 4239.7a 0.30a 14.61a
235 16.70ab 58.34ab − − 235 7.966ab 3553.6bc 0.26b 11.14c
298 15.80b 51.60bc − − 298 7.466b 3350.0c 0.25b 10.95c
230 16.86ab 55.62ab − − 230 7.454b 2291.3e 0.18c 7.13d
111 17.25ab 63.50a − − 111 7.694ab 2621.0d 0.20c 7.77d
N75 86 16.61ab 45.31bc 3.67bc 60.83b N75 86 8.262ab 4562.4b 0.29b 15.59b
184 14.89b 42.77c 4.52a 67.16a 184 8.7801a 5037.0a 0.31ab 17.22a
235 15.77b 48.73bc 3.82b 60.24b 235 8.301ab 4517.7b 0.31ab 15.16b
298 17.87a 50.17b 3.50c 62.24ab 298 7.823b 4667.8ab 0.32a 14.56b
230 17.80a 60.34a 2.33e 41.49c 230 8.071b 3112.0c 0.24c 9.70c
111 15.06b 59.35a 2.80d 42.12c 111 7.825b 3158.8c 0.23c 9.50c
N150 86 13.76bc 49.95bc 2.50a 34.40ab N150 86 8.656a 5160.3ab 0.26ab 16.71a
184 17.37a 60.37a 2.12c 36.66a 184 8.825a 5499.2a 0.28a 16.96a
235 13.53bc 51.96abc 2.49a 33.72b 235 8.846a 5058.6b 0.25bc 15.21b
298 14.49b 54.34ab 2.31b 33.44b 298 8.270a 5016.5b 0.27ab 15.20b
230 13.91bc 48.67bc 1.92d 26.68c 230 8.386a 4001.4c 0.24cd 11.35c
111 12.97c 44.30c 2.06c 26.61c 111 8.428a 3990.9c 0.22d 11.81c
2020 N0 86 11.87abc 30.86a − − 2020 N0 86 8.196a 3930.9b 0.27a 9.62c
184 11.88abc 34.48a − − 184 8.319a 4706.3a 0.27a 11.78a
235 12.82a 35.32a − − 235 8.071a 4570.8a 0.27a 11.21b
298 11.31c 31.42a − − 298 7.783a 3938.7b 0.25a 9.36c
230 12.67ab 34.93a − − 230 7.221b 2847.1c 0.20b 6.60e
111 11.51bc 32.13a − − 111 7.962a 3048.5c 0.21b 7.16d
N75 86 9.61bc 30.19a 6.30b 60.47b N75 86 8.449ab 4535.3b 0.23ab 10.71b
184 9.73bc 32.29a 7.37a 71.68a 184 8.715a 5375.8a 0.24a 12.80a
235 10.09bc 30.64a 7.01a 70.48a 235 8.198bc 5286.1a 0.24a 12.12a
298 10.56ab 32.81a 6.09b 64.26b 298 8.073bc 4819.3b 0.25a 10.87b
230 11.29a 32.04a 4.51d 50.94c 230 7.915c 3820.5c 0.23ab 8.25c
111 9.24c 30.10a 5.15c 47.35c 111 8.228bc 3551.1c 0.21b 7.84c
N150 86 8.73b 32.37a 4.34b 37.84b N150 86 8.696b 5675.9b 0.23bc 14.55b
184 8.74b 31.04a 4.80a 41.89a 184 9.099a 6283.8a 0.24ab 16.57a
235 8.55b 29.18a 4.65a 39.73b 235 8.494bc 5959.3b 0.23bc 14.07b
298 9.56a 30.66a 4.15b 39.60b 298 8.105c 5939.4b 0.25a 14.35b
230 8.25b 34.06a 3.58c 29.48c 230 8.334bc 4422.7c 0.22cd 10.54c
111 8.29b 31.44a 3.48c 28.80c 111 8.321bc 4320.7c 0.20d 10.09c
Variation source Variation source
Year (Y) *** *** *** *** Year (Y) ns *** *** ***
N Treatment (N) *** * *** *** N Treatment (N) *** *** *** ***
Cultivar (C) *** * *** *** Cultivar (C) *** *** *** ***
Y×N *** ns *** ns Y×N ns * ** ***
Y×C ns * *** * Y×C ns ** *** ***
N×C *** *** *** *** N×C ns * ** ***
Y×N×C *** *** * * Y×N×V ns * ns ***

Values followed by a different letter within the same column are significantly
different at P < 0.05. N0, N75, and N150 represent N fertilizer application rates
of 0, 75, and 150 kg N ha− 1 for broomcorn millet, respectively. *, **, and ***
significant at the 0.05, 0.01, and 0.001 probability levels, respectively. ns, no
significant difference.
Values followed by a different letter within the same column are significantly
different at P < 0.05. N0, N75, and N150 represent N fertilizer application rates
of 0, 75, and 150 kg N ha− 1 for broomcorn millet, respectively. *, **, and ***
significant at the 0.05, 0.01, and 0.001 probability levels, respectively. ns, no
significant difference.

canopy to intercept PAR (Table S1). In semiarid areas with abundant
sunlight, such as the Loess Plateau, suitable N application may signifi­
cantly enhance light reflection from the surface to the canopy and
reduce light leakage loss, thereby promoting leaf photosynthetic ca­
pacity of leaves and driving crop growth (Gong et al., 2021). Further­
more, under low N conditions, the mean RUE values were 24.4% higher
in 2019 and 26.9% higher in 2020 in the low-N-tolerant cultivars than in
the low-N-sensitive cultivars, while no significant differences in IPAR
were observed among the six cultivars. This suggested that the much
higher RUE of the low-N-tolerant cultivars is mainly attributed to their
increased dry matter accumulation under low N stress conditions. Pro­
ductive plant transpiration is the physiological basis for the efficient use
of light in food crop photosynthetic systems (Hou et al., 2015). There­
fore, the higher stomatal photosynthetic performance of the
low-N-tolerant cultivars, and the potential light compensation resulted
in a much longer reproductive growth stage, directly increased
assimilation and photosynthate translocation, and ultimately promoted
crop canopy radiation capture (Lindquist et al. 2005).
Rational nutrient inputs increase dryland agricultural production
through increasing transpiration and decreasing evaporation (Li et al.,
2009; Jia et al., 2020). In our study, the low N treatments resulted in
significantly lower SWCs than the N150 treatment in the two years
(Fig. 6). This was mainly due to increased water loss by transpiration
and increased evaporation (data not published) (Li et al., 2009). In a
previous study, the broomcorn millet grew slowly and took up less water
from the soil under low N fertilization, thus enhancing water loss by
evaporation; however, the smaller leaf area exposed more soil surface
(Table S1), increased the soil temperature and decreasing the SWC
(Zhang et al., 2017). In this study, low N treatments resulted in
decreased WUE due to the reduced ability of plants to utilize soil water
via exchange of water between the different soil layers (Table 5).
Generally, efficient transpiration in crop photosynthetic systems is the

10
C. Liu et al.
physiological basis for effective water use, especially in semiarid re­
gions. The SWC and WUE values for the low-N-tolerant cultivars (SWC,
9.6%; WUE, 12.5 kg ha− 1 mm− 1) were significantly higher than those for
the low-N-sensitive cultivars (SWC, 6.6%; WUE, 8.0 kg ha− 1 mm− 1)
under low N conditions. It appears that the morphological and physio­
logical parameters (leaf growth strategies) of the low-N-tolerant culti­
vars are more effective than those of the low-N-sensitive cultivars for
managing low N stress (Table S1). Low-N-tolerant cultivars may also
create an impermeable barrier to limit water loss via soil evaporation
while increasing physiologically significant canopy transpiration (Lian
et al., 2020).
Nutrient capture and use efficiency are of interest to researchers
attempting to determine the capacity of an area for sustainable agri­
cultural production. Low N stress strongly reduced N uptake in N0 and
N75 than in N150 and decreased broomcorn millet biomass by 45.7% in
2019 and by 102.8% in 2020 (Table 4). This was likely due to the
reduced plant nutrient availability and uptake along with changes in the
soil environment, which significantly limited plant growth and photo­
synthesis (Lone and Khan, 2007). Moreover, NUE and NHI values
decreased as the N application rate increased across the two years of the
study. This is indicative of a negative relationship between fertilizer rate
and mineralized-N use efficiency, which contributes to rapid N losses
when N input surpasses crop assimilation capacity (Zhu et al., 2016).
However, adequate N application may enhance grain yield.
The N uptake and NUE were significantly higher in the low-N-
tolerant cultivars than in the low-N-sensitive cultivars (Fig. 7 and
Table 4). This result agrees with that of the study by Santos et al. (2015),
who reported that species and/or genotypes with higher uptake effi­
ciency for a certain nutrient have a greater ability to obtain that nutrient
under conditions of low nutrient availability. This may be explained by
the greater absorption area and capacity of the roots of low-N-tolerant
cultivars with less viscous sap in their xylem leading to increased
nutrient uptake and transfer (Xu et al., 2012).
The low-N-tolerant cultivars had significantly higher NUPE (34.8%)
and NPE (38.8%) values than the low-N-sensitive cultivars across the
years and N treatments. Both NUPE and NPE are defined by the nutrient
uptake efficiency from the soil and depend on nitrate and ammonium
transport at the root level (Masclaux–Daubresse et al., 2008). At the
whole-plant level (aboveground and belowground parts), NUPE and
NPE depend on the plant’s storage capacity in its cells and on the
transport from the root to the shoot, where assimilation occurs
(Masclaux-Daubresse et al., 2008; Chardon et al., 2010). Taken together,
these results indicate that broomcorn millet tolerates low N conditions
by adapting to the ecological environment through optimizing its
developmental characteristics to make better use of the available
nutrient resources. Knowledge-based N management practices generally
improve NUE by providing better synchronization of crop N demand
with N supply and therefore have been adopted for increasing yield
while decreasing nitrous oxide emission and other N losses (Xia et al.,
2017). The greenhouse gas emissions of low-N-tolerant broomcorn
millet needs further study.
The low N treatments significantly decreased broomcorn millet
productivity due to the growth and resource use efficiency disadvan­
tages over the two years of the study. For example, low N stress
decreased the 1,000-grain weight by 7.2% in 2019 and 5.8% in 2020,
and the grain yield by 36.5% in 2019 and 30.2% in 2020 compared with
the N150 treatment (Table 5). This indicated that low N input decreased
the leaf photosynthetic metabolism source and reduced the dry matter
accumulation and transportation sinks, thereby weakening the ecolog­
ical resource utilization efficiency (Li et al., 2017). However, the
low-N-tolerant cultivars produced higher grain yields and HIs than the
low-N-sensitive cultivars mainly because of their better use of light,
water, and nutrients. In other words, the sufficient environmental N
supply under field conditions may have stimulated the physiological
processes in low-N-tolerant cultivars that transport dry matter from the
vegetative organs (stem, leaf, and sheath) to the reproductive organs
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Agricultural and Forest Meteorology 308-309 (2021) 108616
(grains), eventually resulting in a higher HI (Moriwaki et al., 2019). This
study shows that we need to make full use of the value of low-N-tolerant
cultivars to achieve green and sustainable agriculture in the future.
5. Conclusion
Low N stress significantly decreased photosynthetic activity and dry
matter accumulation in broomcorn millet and reduced its IPAR, soil
water uptake, and the capacity to take up nutrients, thereby resulting in
decreased crop productivity. The low-N-tolerant cultivars had signifi­
cantly higher leaf photosynthesis and resource capture (dry matter
transportation) compared to the low-N-sensitive cultivars. The positive
effects of efficient radiation, water, and nutrient use strategies were the
key mechanisms by which low-N-tolerant cultivars enhanced crop pro­
duction (45.1% higher grain yield and 22.5% higher HI). In summary,
the results of this study provide an important theoretical basis for
breeding low-N-tolerant broomcorn millet and promoting its populari­
zation and application in semiarid areas similar to the Loess Plateau,
China.
Declaration of Competing Interest
The authors declare no conflict of interest.
Acknowledgments
The authors gratefully acknowledge the support received from the
Yulin Academy of Agricultural Science. This work received financial
support from the National Natural Science Foundation of China
(31860340 and 31371529), China Agriculture Research System of MOF
and MARA (CARS-06-A26), Minor Grain Crops Research and Develop­
ment System of Shaanxi Province (NYKJ-2021-YL(XN)40), Shaanxi
Province Key Research and Development Project (2018TSCXLNY0301),
and National Science and Technology Supporting Plan
(2014BAD07B03).
Supplementary materials
Supplementary material associated with this article can be found, in
the online version, at doi:10.1016/j.agrformet.2021.108616.
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