Journal of Soil Science and Plant Nutrition (2024) 24:4952–4964

https://doi.org/10.1007/s42729-024-01884-w

ORIGINAL PAPER

Ammonium Nitrate Fertilization Increases the Crude Protein Content

and Wheat Grain Yield in Subtropical Conditions
Ezequiel Helbig Pasa1 · Cristiano Weinert1 · Júlia Peralta Ferreira2 · Hector Tavares Ferreira2 ·
Frantiesco Pereira Martinez2 · Tiago Pedó2 · Mateus da Silveira Pasa2 · Filipe Selau Carlos3

Received: 21 November 2023 / Accepted: 14 June 2024 / Published online: 26 June 2024
© The Author(s) under exclusive licence to Sociedad Chilena de la Ciencia del Suelo 2024

Abstract
The objective was to evaluate the relationship between yield components, quality and estimated grain yield of wheat under
ammoniacal and nitric nitrogen fertilization in mild winter conditions. The treatments consisted factorial 2 × 3 + 1 at local
1 (L1), where two nitrogen (N) sources, urea and ammonium nitrate (AN), and three rates of N, in two crop seasons
2021 and 2022, and the a factorial 2 × 4 + 1 at local 2 (L2) with a 2022 crop season. The parameters assessed were tillers
per plant (TP), plant height (PH), spike insertion height (SIH), spike size (SS), spikelet per spike (SkS), grain per spike
(GS), estimated yield, dry matter accumulation (DMY), hectoliter weight (HLW) and crude protein content (CPC). It was
observed that the yield components increased with increasing N rates, with some increasing significantly when AN was
used compared to urea. In relation to grain yield, it increases with the use of N, showing superiority when using AN in
relation 17.6% urea in L1-Y2022 at a rate of 160 kg ha− 1 and 21 and 16.6%, respectively at rate of 40 and 80 kg ha− 1 in
L2-Y2022. The CPC present an average superiority of 12.1% when using AN in the early cultivar, and increases with the
increase in N rates. Grain yield is increased by N supplementation and is rate dependent. Yield response to ammonium
nitrate is greater than to urea. Crude protein content is increased as N rate increases. The rate with the highest dry matter
accumulation is 160 kg N ha− 1.

Keywords Triticum aestivum · Urea · Ammonium nitrate · Efficiency

1 Introduction and have a high influence on plant growth and crop pro-
duction, resulting in a strong relationship between fertilizer
The estimative from FAO is that world food production utilization and crop production (Tilman et al. 2002; Wang et
must increase over 60% to feed world population in 2050, al. 2016). Among the nutrients, N is required in large quanti-
which is continuously growing (Alexandratos and Bruins- ties and its deficiency limits plant growth and crop produc-
mae 2012). This growth is followed by an increasing fertil- tion (Sadras and Lawson 2013). The N corresponds to about
izer utilization in agriculture (Bakken et al. 2017; Ren et al. 70% of fertilizers used worldwide, where most crop systems
2023). Such as water, nutrients are of paramount importance need this element, therefore contributing to the importance
of N fertilizer use to improve yield and impact on environ-
ment sustainability (Li et al. 2009; Wang et al. 2016).
The impacts of the use of N fertilizers on the environ-
Ezequiel Helbig Pasa
ezequielpasa@gmail.com ment and crop yield are elucidated by nitrogen use effi-
ciency (NUE), an index that is affected by N losses in the
Filipe Selau Carlos
filipeselaucarlos@hotmail.com soil. Currently, around 90% of ammonia (NH3) and 70% of
nitrous oxide (N2O) emissions originate from agricultural
1
Postgraduate Program in Soil and Water Management and activities (Boyer et al. 2002; Ren et al. 2023). NH3 and N2O
Conservation, Federal University of Pelotas, Capão do Leão, are derived from reactions of N losses, through NH3 vola-
Brazil
tilization processes and denitrification, which are increased
2
Federal University of Pelotas, Capão do Leão, Brazil by N fertilizers and unfavorable N application conditions,
3
Soil Science Department, Federal University of Pelotas, resulting in NUE of around 30 to 50% of the N applied
Capão do Leão, Brazil

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Journal of Soil Science and Plant Nutrition (2024) 24:4952–4964 4953

(Drury et al. 2017; Li et al. 2009). This low efficiency has microorganisms (Bouwmeester et al. 1985). After the min-
been associated with climate changes which may negatively eralization of OM, the nitrification process is responsible for
impact the environment in the long term (Pan et al. 2022), the conversion of NH4+ into NO3−, showing greater activity
such as the emission of greenhouse gases that contribute at temperatures around 26ºC (Beck 1983). The availability
to global warming (Mezzari et al. 2023). Furthermore, low of NO3− in soils is limited in conditions of high rainfall, due
NUE results in lower N availability to plant nutrition and to the low binding of the compound to soil adsorption sites,
development (Harper 1994), since N is the major compo- resulting in an intense leaching process (Brady et al. 2010).
nent of several organic compounds, such as proteins, nucleic Although mineralization is not intense in mild winter condi-
acids, alkaloids enzymes, and is also associated with energy tions (Maslov et al. 2022), under these conditions there may
transfer molecules like adenosine diphosphate (ADP) and be a predominance of NH4+ concentration in the soils.
adenosine triphosphate (ATP) (Javed et al. 2022). The relationship of NH4+ and NO3− in the soil affects
Adequate N nutrition contributes to the satisfactory the development of several crops, therefore, the response
development of crops. It is reported that the development of wheat to N fertilization from different sources may
and growth of yield components interfere in agricultural vary according to temperature, as observed by Wang et al.
production (Benincasa et al. 2022; Slafer et al. 2009), often (2016). These authors observed a better wheat performance
being genetically determined (Sadras and Slafer 2012). in response to the use fertilizers based on ammonium-N and
Since yield components are mainly genetically and physio- nitrate-N, in relation to the ammonium-N source, which was
logically driven, its variability is low, but yield components attributed to the lower rate of nitrification (NH4+ -> NO3−)
of modern cultivars have shown a higher response to N fertil- of ammonium-N source. This fact may occur because wheat
ization (Ahrends et al. 2018; Hernández-Ochoa et al. 2023). prefers absorption NO3− (Wang et al. 2016), or due to
According to Benincasa et al. (2022), the reason it occurs is the extent that NH4+ accumulates in the soil, it may have
because yield components are interdependent and complex, toxic effects in the plants, due to the large energy spent to
since the improvement of one component may reduce other, ammonium efflux, in order to maintain NH4+ cytoplasmic
making it difficult to improve yield. This fact highlights the concentration (Howitt et al. 2000). The regulation of NH4+
importance of understanding yield components plasticity, concentration is fundamental, as the accumulation of this
aiming the most correct timing for agronomical intervention ion can uncouple electron transport from photophosphory-
(Hernández-Ochoa et al. 2023). However, yield components lation (Peltier and Thibault 1983), reducing the photosyn-
like grain weight show some stability even with genotype thetic rate (Cramer and Lewis 1993). According to Cramer
and environment variations (Bradshaw 1965; Slafer et al. and Lewis (1993), the assimilation of NH4+ competes for
2014), being influenced only when source: sink relationship more carbohydrates, resulting in a reduction in the propor-
is unbalanced (Benincasa et al. 2022). According to Piepho tion of shoots and roots due to the partitioning of carbohy-
et al. (2014) most of grain yield are due to genetic contribu- drates within the plant. Under ideal temperature conditions
tion and less from management, but Ahrends et al. (2021) for nitrification, Wang et al. (2016) observed greater conver-
observed high variability and low stability of yield in the sion of NH4+ to NO3−, thus reducing the toxic effect and
absence of N, and when N was added the yield gain was minimizing the differences observed between AN and urea.
more stable. The stability obtained with N use is because as The reported differences between AN and urea result
the plant grows the nutrient demand increases, which is pro- from the solubilization of urea making only NH4+ avail-
vided by additional N supply (Harper 1977) and allowing a able to the plant, which can result in toxic effects (Cramer
proper plant development to achieve higher yields. and Lewis 1993; Howitt et al. 2000; Wang et al. 2016). The
Considering the higher nutrient demand as plant grows use of AN as a N supplement results in balanced solubiliza-
and develop and N as one of the most required nutrients, tion of NH4+ and NO3− in the soil (Zhu et al. 2021), even
especially in cereals as wheat, the improvement of N fer- in conditions of absence or low nitrification rate. The bal-
tilization efficiency is primordial to achieve an appropriate anced availability of NH4+ and NO3− impacts an increase of
source: sink balance and achieve high yields. As the plants 40 to 70% in the N accumulated in wheat, barley and rice
uptak N from different physiological mechanisms through plants, due to the synergism in the absorption between an
the roots, as ammonium (NH4+) or nitrate (NO3−), they anion and a cation, contributing positively to the develop-
show different responses in their uptake, because its avail- ment and productivity of grains (Cramer and Lewis 1993;
ability varies according to environment factors like nitrifica- Furthermore, crops that prefer NO3−, such as wheat, have
tion, temperature, mineralization, and rainfall (Jackson and greater tolerance to higher concentrations of this ion in the
Bloom 1990). In the soil, N comes from the mineralization soil, as evidenced by Crawford & Glass (1998) in tomato
of Organic matter (OM), a process that is more intense under plants, that the best plant development occurs in soils that
higher temperature conditions, due to the greater activity of have up to three times more NO3− concentration compared

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4954 Journal of Soil Science and Plant Nutrition (2024) 24:4952–4964

to NH4+ concentration. As temperature significantly influ- (Alvarez et al. 2013). The weather conditions during the
ences the ammonium: nitrate ratio in the soil, our study experiments are shown in Fig. 1.
was performed in milder winter conditions, with an average
temperature of 13.7ºC (Fritzsons et al. 2015), evaluating the 2.2 Treatments and Experimental Design
effect of N sources in the production components of modern
wheat cultivars is a key component to achieve higher yields The experiments were performed in areas previously pre-
and improve N use efficiency. pared with a heavy disc harrow, which were both fallow with
Therefore, nitrogen fertilizers based on NH4+ and NO3−, summer grasses, and after harvesting the experiments, they
such as ammonium nitrate, provide an adequate balance were cultivated with soybeans to represent the cultivation
of ions for the N uptake by wheat plants, improving their system in the region. In L1-Y2021 and L1-Y2022 the chosen
development and increasing attributes such as the number cultivar was Tbio Audaz®, a medium sized and early cycle
of tillers per plant and grains per ear, contributing to greater cultivar and in L2-Y2022, it was used Tbio Astro®, a super-
wheat yield in mild winter conditions. Based on this hypoth- early and short size cultivar (Biotrigo 2023). Plant density
esis, the objective of this study was to evaluate the effect was 330 plants m− 2 in L1-Y2021 and L1-Y2022, and 250
of N sources and rates on plant development and dry mat- plants m− 2 in L2-Y2022. A randomized block design was
ter accumulation, as well as on the production components, used, in a 2 × 3 + 1 factorial (2 N sources x 3 N rates + 1 con-
quality and grain yield of modern wheat cultivars grown in trol) in location 1 and a 2 × 4 + 1 factorial (2 N sources + 4 N
mild winter conditions. rates + 1 control) in location 2, both with 4 replications per
treatment. Nitrogen sources (factor 1) were urea [46% N in
the form of an amide (NH2)] and ammonium nitrate [27%
2 Materials and Methods N in the form of ammonium (NH4+) and nitrate (NO3−)]. N
rates were 0, 40, 80, and 120 kg N ha− 1 in L1-Y2021, and 0,
2.1 Site Description 40, 80, and 160 kg N ha− 1 in L1-Y2022 and 0, 40, 80, 120
and 160 kg N ha− 1 in L2-Y2022. The plots were made up of
The experiment was performed in two growing seasons in nine 4 m long rows spaced 0.17 m from each other, where
South Brazil. In the 2021 growing season it was carried out the 7 central rows were used for evaluations, leaving the 2
only in the municipality of Capão do Leão (location 1) – RS extreme rows and 0.5 m at each end as borders. Therefore,
(31º48’07,9”S and 52º30’17,9”W, altitude 45 m), and in the the used plot area was 3.57 m².
2022 growing season it as also carried out in the municipality Experiments were sown on June 2nd (L1-Y2021), June
of Pelotas (location 2) – RS (31º31’50,7”S and 52º14’10,8”W, 13th (L1-Y2022), and June 15th (L2-Y2022), using a 200 kg
altitude 26,7 m). From this point on, location (L) and year ha− 1 of a formulated fertilizer 5-20-20 (N–P2O5–K2O) as
(Y) will be referred to as L (1; 2)-Y202 (1;2), so for example base fertilization, applied in the sowing furrow. Topdressing
L1-2021, refers to location 1 and growing season 2021. The nitrogen fertilization was split in two timings, where 50%
soil of the experimental field is classified as Luvisol (Wrb was applied at GS 21 (beginning of tillering) and 50% at GS
2015). The soil chemical properties (0–20 cm) of the experi- 31 (first node visible), according to growth scale proposed
mental field at the inception of the experiment are shown in by Zadoks et al. (1974). Cultural practices were performed
Table 1. according to recommendations of the Brazilian Commission
The climate of the region is Cfa (humid subtropical), of Wheat and Triticale Research (2018).
according to Köppen’s classification, consisting of rainfall
concentration during hot periods, without a defined dry
season, with low chance of frost events, and hot summer

Table 1 Chemical soil analysis of the experimental field, in the years of 2021 and 2022
O.M. (%) C (%) N (NH4+ + pH (H2O) pH (SMP) P (Mehlich H + Al K (mg Ca (cmolc Mg (cmolc Al (cmolc V
−1
NO3−) (mg ) (mg dm (cmolc dm− 3) dm− 3) dm− 3) dm− 3) (%)
−3
kg− 1) ) dm− 3)
Local 1–2021
2 1.1 8.1 5.6 6.5 10.8 2.7 93.5 3 1.3 4.5 63
Local 1–2022
2.1 1.2 8.7 6.5 6.9 15 1.6 73 4.3 2.2 8.3 80
Local 2–2022
1.2 0.7 5.8 5.9 6.7 22.3 2 105 1.7 1.7 5.7 65
O.M.: organic matter

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Journal of Soil Science and Plant Nutrition (2024) 24:4952–4964 4955

Fig. 1 Climatic conditions during

the experiments in 2021 (a) and
2022 (b)

2.3 Analysis replicates of 100 grain of each plot and expressed propor-
tionally to 1000 grain.
2.3.1 Yield Components and Estimated Yield Grain yield was estimated using yield components pre-
viously described using the Eq. 2: EY = [(NSSM X GS X
The determination of plant height (PH) and spike insertion TGW)/100], where EY = estimated yield; NSSM = number
height (SIH) was performed using a graduated ruler in three of spikes per square meter; GS = number of grains per spike,
plants of the useful portion of the plot. For tillers per plant and TGW = thousand grain weight, in grams.
(TP), spike size (SS), spikelet per spike (SkS), grain per
spike (GS), and grain per spikelet (GSk) five plants random 2.3.2 Shoot Dry Matter Yield (DMY)
sampled at harvest were used. The number of spikes per
square meter (NSSM) was obtained by counting the number The quantification of shoot dry matter was performed by
of spikes in two linear meters of the central portion of each removing whole plants with a cut close to the ground in
plot, then converting it to square meters, through the fol- an area of 0.5 × 0.5 m (0.25 m2) per plot, prior to harvest
lowing equation: NSSM = NS /RS, where NSSM = Number (GS 91). The material was properly identified in the field
of spikes per square meter (m²); NS = Average number of and dried in an oven at 55º C for 72 horas, until reaching a
spikes per linear meter; RS = row spacing (m). The thou- constant mass, and then weighed to determine the total dry
sand grain weigh (TGW) was determined by weighing 8 matter mass. Then, from this value was subtracted the grain

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4956 Journal of Soil Science and Plant Nutrition (2024) 24:4952–4964

yield and obtained shoot dry matter, which was expressed 3 Results
in Mg ha-1.
3.1 Yield Componentes
2.3.3 Hectoliter Weight
The increase in N rates was significant for all yield compo-
The hectoliter weight (HLW) was determined using a hecto- nents evaluated, however, the N sources did not show the
liter measuring system (Comag – Equipamentos Industriais) same behavior. Considering the TP no significant interac-
with a capacity of a quarter of a liter, through the average of tion was observed between the factors in any location and
two measurements of each plot. Each sample was weighed year studied. Considering the simple effects N source, AN
and proportionally converted to one hundred liters. The showed higher TP in L1-Y2022 and L2-Y2022 (Fig. 2) in
results were expressed in Kg hL-1. relation to urea, with values of 1.3 and 1.4, respectively for
AN, and 1.2 for urea in both locations. The TP increased
2.3.4 Crude Protein Content (CPC) linearly in response to N rate in L1-Y2021 and quadratically
in L1-Y2022 and L2-Y2022 (Fig. 2). A similar behavior was
The crude protein content was determined by grinding a observed for PH and SIH for N rate effect while differences
grain sample of each experimental unit (plot), then subject- between N source were found only in L2-Y2022, with higher
ing it to digestion with acid (H2SO4) and heat, prior to dis- PH and SIH values found with AN (Tables 1 and 2). Interac-
tillation using the Kjeldahl method. A nitrogen-to-protein tion between the factors was observed for SS in L1-Y2022,
conversion factor of 6.25 was used, according to Hasten- where a quadratic rate effect was observed with AN. SS was
pflug et al. (2011). higher with urea in the rate of 40 kg ha− 1 and with AN in
the 160 Kg ha− 1 N rate (Table 2). The simple effect of N rate
2.4 Statistical Analysis on SS was observed in L1-Y2021 and L2-Y2022, where SS
increased linearly as the N rate increased (Tables 2 and 3).
The statistical analysis was performed using the software Only the simple effect of N rate was observed on SkS
R (R Core Team, 2020). Regarding the significance, the in the L1-Y2021 and L2-Y2022, with a linear increase on
data were analyzed by means of the F test. When ANOVA SkS in both situations (Tables 2 and 3). Regarding the GS
revealed significant effect of treatments, mean comparison and GSk, no effect of N source was observed in L1-Y2021
and polynomial regression were performed, for qualitative and L1-Y2022, but an N rate effect was found in both cases,
and quantitative data, respectively. with a quadratic response (Table 2). However, in L2-Y2022,
an interaction between the factors was found, where in the
N rate of 40 kg ha− 1, the AN showed a higher GS than urea.
The N rate effect for both variables was quadratic (Tables 2
and 3). In relation to TGW a significant interaction between
the factors was observed only in L2-Y2022, with increasing
linear in relation to the increase in N rate, with urea supe-
riors to AN at N rate 120 and 160 Kg ha− 1. For L1-Y2021
and L2-Y2022, no variation was observed with N source
and rate.

3.2 Ears per Square Meter and Grain Yield

The use of AN or urea did not show a significant differ-

ence for NSSM in any of the evaluated locations. In both
locations, a significant difference was observed with the
increase in N rates. For L1-Y2021 and L2-Y2022, the
NSSM increased linearly in response to the N rate, with the
highest values being observed, respectively, in rates of 120
and 160 kg N ha− 1. While in L1-Y2021 and L2-Y2022 the
highest value of NSSM was observed at the highest rates,
in L1-Y2021 the highest value observed was at the rate of
Fig. 2 Number of tillers per wheat plant subjected to different rates of 122.1 kg N ha− 1, being 478.7 ears m− 2, due the quadratic
N with urea and ammonium nitrate (AN), in the year 2021 at site 1 (L1- behavior observed at this location and year (Fig. 3).
Y2021) and in the year 2022 at sites 1 (L1-Y2022) and 2 (L2-Y2022).

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Journal of Soil Science and Plant Nutrition (2024) 24:4952–4964 4957

Table 2 Plant height (PH), spike insertion height (SIH), spike size (SS), spikelet per spike (SkS), grain per spike (GS), grain per spikelet (GSk)
and thousand grain weight (TGW) of wheat fertilized with different rates of urea and ammonium nitrate (AN), in the location 1, in 2021 and 2022
PH SIH Ss SkS GS GSk TGW
L1-Y2021
N source (NS)
Urea 81.5 74.7 6.8 14.9 22.9 1.6 37.7
AN 81.7 75.1 6.6 15.6 23.0 1.5 37.3
Dose (D)
0 73.9 68.3 5.6 12.4 15.6 1.4 37.3
40 80.6 74.0 6.5 15.0 22.2 1.5 37.6
80 85.0 77.7 7.3 16.7 26.5 1.6 37.7
120 87.0 79.7 7.4 17.0 27.6 1.6 37.3
p value
NS 0.910 0.771 0.314 0.186 0.979 0.603 0.278
D < 0.001 < 0.001 < 0.001 < 0.001 < 0.001 0.462 0.715
NS*D 0.988 0.984 0.903 0.691 0.906 0.951 0.081
p regression
Linear < 0.001 < 0.001 < 0.001 < 0.001 < 0.001 - -
Quadratic 0.191 0.226 0.140 0.044 0.028 - -
L1-Y2022
N source (NS) Urea AN
Urea 78.5 70.5 - - 19.1 34.6 1.9 35.9
AN 80.0 71.4 - - 19.2 35.3 1.8 36.4
Dose (D)
0 71.9 63.8 7.5 7.5 18.7 28.8 1.5 36.9
40 78.2 70.1 8.00 a 6.9 b 19.1 34.7 1.8 36.0
80 82.5 74.4 8.1 8.0 19.5 38.9 2.0 36.1
160 84.5 75.6 8.1 b 9.9 a 19.1 37.3 1.9 35.6
p value
NS 0.091 0.270 0.514 0.674 0.616 0.543 0.266
D < 0.001 < 0.001 < 0.001 0.264 < 0.001 0.002 0.233
NS*D 0.068 0.238 < 0.001 0.408 0.663 0.898 0.826
p regression
Linear < 0.001 < 0.001 ns < 0.001 ns < 0.001 < 0.001 -
Quadratic < 0.001 < 0.001 ns 0.006 ns 0.002 0.012 -
a
Means followed by different letters, lowercase in the columns and uppercase in the rows, are significantly different according to Duncan’s test
at p < 0.05

Considering grain yield, significant interaction between 338.75 kg de N ha− 1 for urea corresponding to yields of 5.2
factors was found in L1-Y2022 and L2-Y2022 (Fig. 4b and and 6.4 Mg ha− 1, respectively.
c), while only the simple effect of N rate was observed in
L1-Y2021 (Fig. 4a). In L1-Y2021, the grain yield increased 3.3 Dry Matter Yield (DMY)
linearly as a response to increased N rate, with no signifi-
cant difference observed between the sources used (Fig. 4a). Regarding the use of AN and urea, no significant differences
In the L1-Y2022, a quadratic behavior was observed of were observed between sources in any of the locations or
both sources, i.e., urea and AN, where superiority of AN years evaluated. The increase in N rates significantly inter-
was observed at 800 kg ha− 1 in relation to urea, at a rate of fered with DMY, showing quadratic behavior in all locations.
160 kg N ha− 1 (Fig. 4b). Through quadratic behavior, the In the L1-Y2021, the highest DMY value was observed at
maximum rate of technical efficiency (RMTE) was deter- a rate of 120 kg N ha− 1, with RMTE at a rate of 140.67 kg
mined at 134 kg N ha− 1 for AN and 100 kg N ha− 1 for urea, N ha− 1, with dry matter accumulation of 10.8 Mg ha− 1. For
with grain yield of 7.0 Mg ha− 1 and 6.2 Mg ha− 1, respec- L1-Y2022 and L2-Y2022, the RMTE for dry matter was
tively. In L2-Y2022 grain yield of AN was 1100 kg ha− 1 145 kg N ha− 1 and 131,5 kg N ha− 1, corresponding to 9.9
and 700 kg ha− 1 higher than urea in the N rates of 40 and and 8.5 Mg ha− 1, respectively (Fig. 5). The highest DMY
80 kg N ha− 1. Both N sources showed a quadratic response values in L1-Y2022 and L2-Y2022 were 10.2 and 8.9 Mg
to N rate, where the RMTE was 173.5 kg N ha− 1 for AN and ha− 1, respectively, both at the highest rate, 160 kg N ha− 1.

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4958 Journal of Soil Science and Plant Nutrition (2024) 24:4952–4964

Table 3 Plant height (PH), spike insertion height (SIH), spike size (SS), spikelet per spike (SkS), grain per spike (GS), grain per spikelet (GSk) and
thousand grain weight (TGW) of wheat fertilized with different rates of urea and ammonium nitrate (AN), in the location 2, in 2022a
PH SIH SS SkS GS GSk TGW
L2-Y2022
N source (NS) Urea AN Urea AN Urea AN
Urea 76.2 B 67.5 B 8.0 17.5 - - - - - -
AN 78.0 A 69.6 A 8.1 17.9 - - - - - -
Dose (D)
0 68.1 61.1 6.8 16.9 21.0 21.0 1.2 1.2 34.5 34.5
40 77.6 68.0 8.1 17.6 30 b 35.2 a 1.7 b 2.0 a 36.6 36.9
80 78.7 69.9 8.2 17.8 33.8 34.3 2.0 2.0 37.4 37.2
120 79.3 71.0 8.3 18.1 39.5 38.7 2.2 2.2 40.0 a 37.3 b
160 81.8 72.7 8.7 18.2 36.9 36.0 2.1 2.0 40.9 a 37.5 b
p value
NS 0.047 0.018 0.669 0.095 0.720 0.453 0.007
D < 0.001 < 0.001 < 0.001 0.010 < 0.001 < 0.001 < 0.001
NS*D 0.810 0.362 0.991 0.610 0.014 0.033 0.022
p regression
Linear < 0.001 0.085 < 0.001 < 0.001 < 0.001 < 0.001 < 0.001 < 0.001 0.000 0.004
Quadratic < 0.001 0.003 0.057 0.428 < 0.001 < 0.001 < 0.001 < 0.001 0.770 0.060
a
Means followed by different letters, lowercase in the columns and uppercase in the rows, are significantly different according to Duncan’s test
at p < 0.05

in both years evaluated, showing no significant effect of the

treatments used in the experiment (Fig. 6b).

3.5 Crude Protein Content (CPC)

In the years evaluated at site 1, a significant interaction was

observed between the use of AN and urea and N rates, show-
ing variation in CPC. At site 2, no significant difference was
observed between the sources used, with only the rate of
N used being significant. In L1-Y2021, a higher CPC was
observed with the use of AN compared to urea, at N rates
of 80 kg N ha− 1 and 120 kg N ha− 1, representing 1.4 and
1.8% more CPC relative to urea, respectively. Regarding
Fig. 3 Number of spikes per square meter (NSSM) of wheat plants the N rate effect within N source, for both sources a linear
subjected to increasing N rates, in the year 2021 at site 1 (L1-Y2021) increase on CPC was found (Fig. 7a). Similar behavior was
and in the year 2022 at sites 1 (L1-Y2022) and 2 (L2-Y2022). observed in L1-Y2022, but in this case the CPC increased
linearly only when AN was used as a source of N, while a
3.4 Hectoliter Weight (HLW) quadratic effect was observed in response to urea. The CPC
in L1-Y2022 showed 11% superiority with the use of AN at
The use of N rates with urea and AN showed a significant N rate of 160 kg N ha− 1 in relation urea, with CPC values of
interaction for HLW in L1-Y2021, where with use of urea 18.1% and 16.1%, respectively, for AN and urea (Fig. 7b).
there was a quadratic response to the increase N rate and a Only the simple effect of N rate was observed in L2-Y2022,
linear response with the use of AN. In both sources used, a which showed a quadratic trend, with a CPC of 17.7%, at a
decrease in HLW was observed as the N rates increased. Con- N rate of 160 kg de N ha− 1.
sidering the effect of N source within N rate, the only signif-
icant effect was observed in the rate of 40 kg N ha− 1, where 3.6 Pearson Correlation
HLW with urea was higher than AN (Fig. 6a). The behavior
observed in L1-Y2022 was different from L1-Y2021, as In the Pearson correlation, it was observed a high degree
only the single effect of the N rate was observed, showing a of association between PH and SIH, between GS and GSk
linear decrease in HLW as the N rate increased. The results and between TP and NSSM (Fig. 8). GY showed a positive
found in L2-Y2022 differed from those observed in site 1 relationship with all parameters, but TGW, with a higher

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Journal of Soil Science and Plant Nutrition (2024) 24:4952–4964 4959

Fig. 5 Dry matter yield of wheat plants subjected to different rates of

N, in the year 2021 at site 1 (L1-Y2021) and in the year 2022 at sites 1
(L1-Y2022) and 2 (L2-Y2022)

Fig. 4 Estimated yield of wheat grains under rates of nitrogen in the

form of urea and ammonium nitrate (AN), in years 2021 (a) and
2022 (b) at local 1, and 2022 (c) at local 2. Asterisks (*) at the top of
the graph represent significant differences by Tukey’s test (p < 0.05)
between N source within each rate

degree of association with GS. However, a negative rela- Fig. 6 Hectoliter weight of wheat grains under rates of nitrogen in the
tionship was observed between GY and HLW. Note that form of urea and ammonium nitrate (AN), in the year 2021 (a) and in
2022 (b) at site 1 (L1-Y2022) and site 2 (L2-Y2022). Asterisks (*) at
HLW showed a negative relationship with almost all param- the top of the graph represent significant differences by Tukey’s test
eters, except with TGW. CPC showed a higher relationship (p < 0.05) between N source within each rate
with GS, with a negative relationship with HLW and DMY
(Fig. 8).

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4960 Journal of Soil Science and Plant Nutrition (2024) 24:4952–4964

Fig. 8 Pearson’s correlation between variables analyzed in wheat

under different rates and sources of nitrogen fertilizer in Southern Bra-
zil. TP: plant height, PH: plant height, SIH: spike insertion height, SS:
spike size, SkS: spikelet per spike, GS: grain per spike, GSk: grain
per spikelet, TGW: Thousand grain weight, GY: grain yield, HLW:
hectoliter weight, DMY: dry matter yield, CPC: crude protein content,
NSSM: number of spikes per square meter. Cells with “X” are not
statistically correlated (p < 0.05)

stem elongation (Luo et al. 2020; Wu et al. 2020), which

favors the crop, since a higher stem elongation improves
assimilate reserve, which may be translocated to grains dur-
ing grain filling (Espindula et al. 2010; Kumar et al. 2017)
and improve yield. The quadratic behavior observed in
L1-Y2022 and L2-Y2022 is probably due to the increase
in the maximum rate used, which may increase lodging, as
observed by other authors (Teixeira Filho et al. 2010; Zhang
et al. 2017b). This effect is associated to the secondary cell
wall formation limitation in response to excessive N, result-
ing in lower lodging resistance (Luo et al. 2020; Wu et al.
2017; Zhang et al. 2017b). However, in a study carried out
Fig. 7 Crude protein content in wheat grains under rates of nitrogen in by Benincasa et al. (2022) plant height was not significantly
the form of urea and ammonium nitrate (AN), in the year 2021 (a) and
2022 (b) at local 1, and 2022 (c) at local 2. Asterisks (*) at the top of affected, which may indicate a different genotype or envi-
the graph represent significant differences by Tukey’s test (p < 0.05) ronment response.
between N source within each rate The increase of TP, NSSM and GS in L1-Y2022 and
L2-Y2022 with increasing N rates is a consequence of better
4 Discussion N supply. According to Zhang et al. (2022), N is fundamen-
tal during tillering, being responsible for TP determina-
The positive linear behavior of PH and SIH in L1-Y2021 tion, spikes, and GS in other cereals like rice. However, in
occurs when the rate used is beneficial to the crop, since wheat, Benincasa et al. (2022) observed a variation of TP
high N rates may result in negative effects, as observed by in response to N, which was not statistically significant. N
Teixeira Filho et al. (2010) and Zhang et al. (2017a). This fertilization effect might vary among cultivars, since geno-
behavior is related to the adequate N availability to stimulate type characteristics may change plant ability to uptake,

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Journal of Soil Science and Plant Nutrition (2024) 24:4952–4964 4961

assimilate, and convert N to vital functions, thus affecting by causing grain abortion in the spike extremity, possibly
grain yield (Luo et al. 2020). This could explain the varia- due to reduction of non-structural carbohydrate supply
tions observed between cultivars in 2022, with a quadratic (NSCs) at the beginning of grain filling. These components
behavior for NSSM in L1-Y2022 and linear in L2-Y2022. significantly interfere in yield. Among them, in our study
The higher TP when used AN in L1-Y2022 and L2-Y2022 the GSk showed positive relationship with GY and contrib-
this is due to wheat plants having higher concentrations of uted the most to yield increase when used AN as source N
cytokinin, as supplementation with NO3- and NH4+ stimu- in L1-Y2022 and L2-Y2022, relative to urea. This observa-
lates the production of this hormone, since cytokinin is tion corroborates with the results found by Benincasa et al.
responsible for the elongation and emission of tillers in (2022), which observed that the higher yield was strongly
cereals (Wang and Below 1996), resulting in greater tiller correlated to number of grains per area (strongly corre-
emission. lated to grains per spike) and not to grain weight. The same
Significant differences between AN and urea were found authors also observed a strong relationship among grains
by Wang et al. (2016), which attributes the lower conversion per spike, spikelet per spike, and grains per spikelet, simi-
of NH4+ to NO3-, the lower nitrification due to milder tem- larly as we observed in our study.
peratures (such as the case in the present study), since when The higher yield observed with AN relative to urea was
used fertilizers as urea, the NH4+ resulting from its solubili- also observed by Heuermann et al. (2021) in rapeseed,
zation ends up accumulating in the soil, then increasing its which also observed a positive yield response to 120 kg
soil levels to values that might be toxic to plant growth and de N ha− 1, as we observed in L1-Y2021, regarding N rate
development. According to Britto and Kronzucker (2002), increase. Wang et al. (2016) observed that N accumulation
this toxicity originates from the large amount of energy in the soil as ammonium caused a negative effect on wheat
required in the ammonium efflux to lower its cytoplasmatic growth and development, reducing its yield compared to
concentration (Howitt et al. 2000). Opposite, with AN, the nitrate accumulation.
soil concentration of NO3- and NH4+ are quite similar, result- The quadratic behavior observed of DMY with increas-
ing in a balanced plant uptake, reducing the toxic effect, and ing N rates is a due to a direct effect of this nutrient on
contributing to improved yield (Zhu et al. 2021). However, vegetative growth, such as higher stem elongation and leaf
the balanced availability of NO3- and NH4+ can increase sizes, as observed in other studies (Luo et al. 2020; Wu et
the accumulated N in wheat, barley and rice plants by up to al. 2020), showing a positive relationship with PH and SIH.
70%, due to the synergism in the uptake between a cation A higher dry matter accumulation is observed in response
and an anion (Cramer and Lewis 1993; Duan et al. 2005). to more leaf expansion (Hernández-Ochoa et al. 2023;
Under conditions of low availability of mineral N in the soil Wang et al. 2016). We have observed that the maximum
associated with low levels of OM in the soil, the toxic effects DMY were obtained in rates lower than 160 kg N ha− 1, i.e.,
of NH4+ accumulation may occur more frequently, as wheat (L1-Y2021), 145 kg N ha− 1 (L1-Y2022), and 131.5 kg N
has a preference for uptake NO3- (Wang et al. 2016), when ha− 1 (L2-Y2022), suggesting that the reduction in dry mat-
ion availability is low in the soil, fertilization only based ter accumulation is a response of negative effects of excess
on NH4+ can result in less plant development compared to N. Besides, the decrease of DMY might be a response to
the use of fertilizers based on NH4+ and NO3-. Furthermore, plant lodging and death favored by higher N rates (Luo et
when the fertilization is performed at higher temperature al. 2020; Wu et al. 2017; Zhang et al. 2017b).
conditions, the nitrification rate is increased, which leads to The maximum response to N depends on the plant abil-
less significant differences between the use of urea and AN ity to translate the N uptake into growth and yield, which
(Wang et al. 2016). is a factor strongly genetically driven (Islam et al. 2021).
The increase of SkS, GS and GSk with increasing N rates According to Zhang et al. (2017b), a higher N availability
is probably associated with the higher N availability, as soils induces more vegetative growth and weaker stems, which
with low levels of OM present limitations to the develop- are more sensitive to lodging and end up reducing yield com-
ment of crops (Carlos et al. 2022), especially grasses, which ponents and yield. Although we expected a higher DMY in
require a greater amount of N for their development (Sriv- response to AN, the data did not confirm this hypothesis and
astava et al. 2018; Ray et al. 2020). The higher N avail- it might be explained by a higher influence of weather con-
ability induce lower grain abortion in the spike, as observed ditions than N source on N accumulation in vegetative tis-
by Ma et al. (2022) in corn, where the lower N availability sues (Heuermann et al. 2021). Another possible explanation
being responsible to reduce flower primordia and negatively for the lack of differences between N sources is the higher
impacting number of grains per spike. According to Oury nitrification rate, because according to Wang et al. (2016),
et al. (2016), the reason it occurs is because N deficiency the differences between urea and AN at higher temperatures
reduces pollination, increases pollination asynchrony, and are irrelevant.

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4962 Journal of Soil Science and Plant Nutrition (2024) 24:4952–4964

The reduction of HLW in L1-Y2021 and in L1-Y2022 is become available to plant growth when N is deficient (Li
reported in other studies, such as in the study of Prando et and Wang 2003).
al. (2012, 2013), which attribute this effect to the increase According to Hachiya et al. (2017), the oxidation of
in number of spikes per area and grains per spike, resulting NO3- to NH4+ catalyzed by nitrate reductase demands more
in higher competition for photoassimilates and grain filling, energy compared to direct uptake of NH4+. The increment
and ultimately hectoliter weight. We have also observed a of CPC when AN was used, observed in our study, might
negative relationship between HLW and GS, and with SS, have occurred due to a higher rate of N uptake, thus com-
SKs, GSk, GY, and NSSM, confirming that as you increase pensating the higher energetic demand, and contributing to
grain yield there is a direct negative effect on HLW. Oppo- protein accumulation. Both NO3- and NH4+ in excess are
site, Hernández-Ochoa et al. (2023) found an increase of harmful to plants but when they balanced plant performance
HLW in the N rate of 120 kg N ha-1, relative to the absence is boosted. Another fact that may contribute to the higher
of N fertilization, but it may be a genotype effect since they CPC with AN is the greater competition for carbohydrates
have studied several old and modern German cultivars. due to the assimilation of NH4+, since according to Cramer
In L2-Y2022 we have not found significant treatment and Lewis (1993), in high concentrations of NH4+ there is a
effects on HLW, similarly as found by Ali et al. (2019) reduction in the dry matter ratio of root and part aerial due
and Ferreira et al. (2021). The different response among to the lower amount of carbohydrates destined for the devel-
the two locations to N rate is probably a cultivar effect, as opment of the plant structure. However, when temperatures
previously reported in the study of Hernández-Ochoa et al. are higher, the nitrification rate is accelerated resulting in
(2023). According to Ferreira et al. (2021), the absence of higher conversion of NH4+ from urea to NO3- in the soil
HLW variation in response to N rates is due to higher phe- (Wang et al. 2016), and the differences of protein content
notypic stability of the cultivar. This characteristic was also between N sources would be more difficult to occur.
reported by Abera et al. (2020), highlighting that yield and
grain quality varies according to genotype response to culti-
vation conditions. Therefore, it is possible to suppose that in 5 Conclusion
L2-Y2022 the cultivar used showed a phenotypic stability,
as well as an appropriate nutrient uptake and translocation, The development and accumulation of dry matter in wheat
to properly supply the assimilates necessary to the higher plants increases with N rates, being greater at a rate of
number of grains as yield increased. 160 kg N ha− 1, showing no significant difference between
The increase in crude protein content associated with the use of ammonium nitrate in relation to urea. The compo-
increasing N rates, is similar to the results found by Hernán- nents Tillers per plant, Plant height, Spike insertion height
dez-Ochoa et al. (2023). The low OM content reduces the and Grains per spike are more influenced by the increases in
availability of N in the soil, and consequently the develop- N rates, with the highest values at the rate of 160 kg N ha− 1.
ment of crops (Carlos et al. 2022), associated with this, the The attributes Grains per spikelet, Spikelet per spike and
AN presents less loss due to volatilization of NH3, improv- Thousand grain weight are less influence by N variation.
ing the efficiency of nitrogen use (Souza et al. 2020), which In both cultivars, the crude protein content increased with
may result in greater accumulation of CPC. This effect was the N rates and was higher at a rate of 160 kg N ha− 1, being
observed in our study in location 1 in both years, but not in higher in the early cultivar with the use of ammonium nitrate
location 2, probably as a genotype effect since cultivars of in relation to urea. Grain yield is increased by N supplemen-
both locations were different, and genotype response to N is tation and is rate dependent, with better results when ammo-
variable (Abera et al. 2020). nium nitrate is used as N source relative to urea. Among the
The increase of CPC is a response to AN is probably due rates analyzed, the rate of 160 kg N ha− 1 using ammonium
to the solubility of this fertilizer be in inorganic compounds nitrate showed better grain yield for both cultivars.
readily absorbed by the plant (Ochieng et al. 2021). Accord- Thus, ammonium nitrate fertilizer can be an efficient
ing to Wang et al. (2016) the application of NO3- promotes nitrogen fertilization strategy in intensive, high-yield wheat
an increase in the levels of nitrate, ammonium, and amide systems in mild winter conditions.
concentration, in comparison to the single NH4+ supply. In
conditions of NO3- accumulation in the soil or its addition Declarations
through NO3- based fertilizers result in higher uptake by
wheat plants and being accumulated in the aerial portion Conflict of Interest On behalf of all authors, the corresponding author
states that there is no conflict of interest.
(Wang et al. 1998; Wang et al. 2016). This accumulation is
possible because the vacuole has the capacity to store nitrate
in large quantities, to later be released in the cytoplasm and

13
Journal of Soil Science and Plant Nutrition (2024) 24:4952–4964 4963

References Duan YH, Zhang YL, Shen QR, Chen HY, Zhang Y (2005) Effect of
partial replacement of NH4+ by NO3– on nitrogen uptake and uti-
lization by different genotypes of rice at the seedling stage. J Plant
Abera D, Tana T, Dessalegn T (2020) Effects of blended NPSB fertil-
Nutr Fert 11:160–165. https://doi.org/10.11674/zwyf.2005.0204
izer rates on yield and grain quality of durum wheat (Triticum
Espindula MC, Rocha VS, Souza MAD, Grossi JAS, Souza LTD
turgidum L.) varieties in Minijar Shenkora District, Central Ethi-
(2010) Nitrogen application methods and doses in the develop-
opia. Ethiop J Agric Sci 30:57–76
ment and yield of wheat. Ciênc Agrotec 34:1404–1411. https://
Ahrends HE, Eugster W, Gaiser T, Rueda-Ayala V, Hüging H, Ewert
doi.org/10.1590/S1413-70542010000600007
F, Siebert S (2018) Genetic yield gains of winter wheat in Ger-
Ferreira LAR, Silva SR, Lollato RP, Ferreira EB, Kölln OT (2021).
many over more than 100 years (1895–2007) under contrast-
Wheat nitrogen utilization efficiency and yield as affected by
ing fertilizer applications. Environ Res Lett 13:10. https://doi.
nitrogen management and environmental conditions. EJFA.
org/10.1088/1748-9326/aade12
https://doi.org/10.9755/ejfa.2021.v33.i11.2788
Ahrends HE, Siebert S, Rezaei EE, Seidel SJ, Hüging H, Ewert F,
Fritzsons E, Wreve MS, Mantovani LE (2015) Altitude and tempera-
Gaiser T (2021) Nutrient supply affects the yield stability of major
ture: study of the thermal gradient in the state of Rio Grande do
European crops—A 50 year study. Environ Res Lett. https://doi.
sul. Rev Bras Climat
org/10.1088/1748-9326/abc849
Hachiya T, Sakakibara H (2017) Interactions between nitrate and
Alexandratos N, Bruinsma J, FAO (2012) World Agriculture
ammonium in their uptake, allocation, assimilation, and signal-
towards 2030/2050: the 2012 revision. https://doi.org/10.22004/
ing in plants. J Exp Bot 68:2501–2512. https://doi.org/10.1093/
ag.econ.288998
jxb/erw449
Ali SA, Tedone L, Verdini L, Cazzato E, De Mastro G (2019) Wheat
Harper JL (1977) The Population Biology of plants. Population biol-
response to no-tillage and nitrogen fertilization in a long-term faba
ogy of plants
bean-based rotation. https://doi.org/10.3390/agronomy9020050.
Harper JE (1994) Nitrogen metabolism. Phys Determ Crop Yield
Agronomy
285–302. https://doi.org/10.2134/1994.physiologyanddetermina-
Alvarez CA, Stape JL, Sentelhas PC, Gonçalves JLM, Sparovek G
tion.c19
(2013) Koppen’s climate classification map for Brazil. Meteor Z
Hastenpflug M, Braida JA, Martin TN, Ziech MF, Simionatto CC,
22:711–728
Castagnino DS (2011) Dual-purpose wheat cultivars submitted to
Bakken LR, Frostegård A (2017) Sources and sinks for N2O, can
nitrogen fertilization and cutting regimes. ABMVZ 63:196–202.
microbiologist help to mitigate N2O emissions? Environ Micro-
https://doi.org/10.1590/S0102-09352011000100029
biol 19:4801–4805. https://doi.org/10.1111/1462-2920.13978
Hernández-Ochoa IM, Gaiser T, Hüging H, Ewert F (2023) Yield
Beck T (1983) Mineralization of soil nitrogen in laboratory incubation
components and yield quality of old and modern wheat cultivars
experiments. Z Pflanzenernähr 120:71–78
as affected by cultivar release date, N fertilization and environ-
Benincasa P, Reale L, Cerri M, Tedeschini E, Tosti G, Falcinelli B,
ment in Germany. Field Crops Res. https://doi.org/10.1016/j.
Rosati A (2022) Nitrogen fertilization levels and timing affect the
fcr.2023.109094
plasticity of yield components in bread wheat (Triticum aestivum
Heuermann D, Hahn H, Von Wirén N (2021) Seed yield and nitrogen
L). Field Crops Res. https://doi.org/10.1016/j.fcr.2022.108734
efficiency in oilseed rape after ammonium nitrate or urea fertil-
Biotrigo. Biotrigo genética - Tbio Audaz (2023) Disponível em:
ization. Front Plant Sci. https://doi.org/10.3389/fpls.2020.608785
<https://biotrigo.com.br/cultivares. Acesso em: 24 de junho de
Howitt SM, Udvardi MK (2000) Structure, function and regulation of
2023.
ammonium transporters in plants. BBA– Biomembr 1465:152–
Bouwmeester RJB, Vlek PLG, Stumpe JM (1985) Effect of environ-
170. https://doi.org/10.1016/S0005-2736(00)00136-X
mental factors on ammonia volatilization from a urea-fertilized
Islam S, Zhang J, Zhao Y, She M, Ma W (2021) Genetic regulation of
soil. SSSA 49:376–381. https://doi.org/10.2136/sssaj1985.03615
the traits contributing to wheat nitrogen use efficiency. Plant Sci.
995004900020021x
https://doi.org/10.1016/j.plantsci.2020.110759
Boyer EW, Goodale CL, Jaworsk NA, Howarth RW (2002) Anthro-
Jackson L, Bloom AJ (1990) Root distribution in relation to soil nitro-
pogenic nitrogen sources and relationships to riverine nitrogen
gen availability in field-grown tomatoes. Plant Soil 128:115–126.
export in the northeastern USA. Biogeochemistry 57:137–169.
https://doi.org/10.1007/BF00011100
https://doi.org/10.1023/A:1015709302073
Javed T, Singhal RK, Shabbir R, Shah AN, Kumar P, Jinger D, Siuta
Bradshaw AD (1965) Evolutionary significance of phenotypic plas-
D (2022) Recent advances in agronomic and physio-molecular
ticity in plants. Adv Genet 13:115–155. https://doi.org/10.1016/
approaches for improving nitrogen use efficiency in crop plants.
S0065-2660(08)60048-6
Front Plant Sci. https://doi.org/10.3389/fpls.2022.877544
Brady NC, Weil RR, Brady NC (2010) Elements of the nature and
Kumar M, Raina SK, Govindasamy V, Singh AK, Choudhary RL, Rane
properties of soils
J, Minhas PS (2017) Assimilates mobilization, stable canopy
Britto DT, Kronzucker HJ (2002) NH4+ toxicity in higher plants:
temperature and expression of expansin stabilizes grain weight
a critical review. J Plant Physiol 159:567–584. https://doi.
in wheat cultivar LOK– 1 under different soil moisture conditions.
org/10.1078/0176-1617-0774
Bot Stud 8:1–13. https://doi.org/10.1186/s40529-017-0169-7
Carlos FS, Kunde RJ, de Sousa RO et al (2022) Urease inhibitor
Li SX, Wang ZH, Hu TT, Gao YJ, Stewart BA (2009) Nitrogen in
reduces ammonia volatilization and increases rice grain yield
dryland soils of China and its management. Adv Agron 101:123–
under irrigation delay. Nutr Cycl Agroecosyst 122:313–324.
181. https://doi.org/10.1016/S0065-2113(08)00803-1
https://doi.org/10.1007/s10705-022-10203-7
Luo L, Zhang Y, Xu G (2020) How does nitrogen shape plant archi-
Cramer MD, Lewis OAM (1993) The influence of nitrate and ammo-
tecture? J Exp Bot 71:4415–4427. https://doi.org/10.1093/jxb/
nium nutrition on the growth of wheat (Triticum aestivum L.) and
eraa187
maize (Zea mays L.) plants. Ann Botany 72:359–365. https://doi.
Ma B, Wang J, Han Y, Zhou C, Xu T, Qu Z, Qian C (2022) The response
org/10.1006/anbo.1993.1119
of grain yield and ear differentiation related traits to nitrogen lev-
Drury CF, Yang X, Reynolds WD (2017) Combining urease and
els in maize varieties with different nitrogen efficiency. Sci Rep.
nitrification inhibitors with incorporation reduces ammonia and
https://doi.org/10.1038/s41598-022-18835-z
nitrous oxide emissions and increases corn yields. J Environ Qual
Maslov MN, Maslova OA (2022) Soil nitrogen mineralization and its
46:939–949. https://doi.org/10.2134/jeq2017.03.0106
sensitivity to temperature and moisture in temperate peatlands

13
4964 Journal of Soil Science and Plant Nutrition (2024) 24:4952–4964

under different land-use management practices. https://doi. Souza EF, Soratto RP, Sandaña P, Venterea RT, Rosen CJ (2020) Split
org/10.1016/j.catena.2021.105922. Catena application of stabilized ammonium nitrate improved potato
Mezzari MF, Veloso M, Santos RND, Valente GB, Carlos FS, Bayer yield and nitrogen-use efficiency with reduced application rate
C (2023) Mitigation of greenhouse gases emission affected by in tropical sandy soils. Field Crops Res 254:107847. https://doi.
no-tillage and winter cover crops in a subtropical paddy rice eco- org/10.1016/j.fcr.2020.107847
system. Rev Bras Ciênc Solo. https://doi.org/10.36783/1806965 Srivastava RK, Panda RK, Chakraborty A, Halder D (2018) Enhanc-
7rbcs20220137 ing grain yield, biomass and nitrogen use efficiency of corn by
Ochieng’ IO, Gitari HI, Mochoge B, Rezaei-Chiyaneh E, Gweyi- varying sowing dates and nitrogen rate under rainfed and irri-
Onyango JP (2021) Optimizing maize yield, nitrogen efficacy and gated conditions. Field Crops Res 221:339–349. https://doi.
grain protein content under different N forms and rates. JSSPN org/10.1016/j.fcr.2017.06.019
21:1867–1880. https://doi.org/10.1007/s42729-021-00486-0 Teixeira Filho MCM, Buzetti S, Andreotti M, Arf O, Benett CGS
Oury V, Caldeira CF, Prodhomme D, Pichon JP, Gibon Y, Tardieu F, (2010) Doses, sources and time of nitrogen application on irri-
Turc O (2016) Is change in ovary carbon status a cause or a conse- gated wheat under no-tillage. Pesq Agropec Bras 45:797–804.
quence of maize ovary abortion in water deficit during flowering? https://doi.org/10.1590/S0100-204X2010000800004
Plant Physiol 171:997–1008. https://doi.org/10.1104/pp.15.01130 Tilman D, Cassman K, Matson P, Naylor R, Polasky S (2002) Agri-
Pan SY, He KH, Lin KT, Fan C, Chang CT (2022) Addressing cultural sustainability and intensive production practices. Nature
nitrogenous gases from croplands toward low-emission agri- 418:671–677. https://doi.org/10.1038/nature01014
culture. Npj Clim Atmos Sci 5:43. https://doi.org/10.1038/ Wang X, Below FE (1996) Cytokinins in enhanced growth and til-
s41612-022-00265-3 lering of wheat induced by mixed nitrogen source. Crop Sci
Peltier G, Thibault P (1983) Ammonia exchange and photorespira- 36:121–126. https://doi.org/10.2135/cropsci1996.0011183X003
tion in Chlamydomonas. Plant Physiol 71:888–892. https://doi. 600010022x
org/10.1104/pp.71.4.888 Wang ZH, Miao YF, Li SX (2016) Wheat responses to ammonium and
Piepho HP, Laidig F, Drobek T, Meyer U (2014) Dissecting genetic nitrate N applied at different sown and input times. Field Crops
and non-genetic sources of long-term yield trend in German offi- Res 199:10–20. https://doi.org/10.1016/j.fcr.2016.09.002
cial variety trials. Theor Appl Genet 127:1009–1018. https://doi. Wrb F (2015) IUSS Working Group WRB World Reference base for
org/10.1007/s00122-014-2275-1 Soil resources 2014. Update
Prando AM, Zucareli C, Fronza V, Oliveira EADP, Panoff B (2012) Wu L, Zhang W, Ding Y et al (2017) Shading contributes to the reduc-
Forms of urea and nitrogen levels in topdressing on the physi- tion of stem mechanical strength by decreasing cell wall synthesis
ological quality of wheat seeds. Rev Bras Sementes 34:272–279. in japonica rice (Oryza sativa L). Front Plant Sci 8:881. https://
https://doi.org/10.1590/S0101-31222012000200012 doi.org/10.3389/fpls.2017.00881
Prando AM, Zucareli C, Fronza V, Oliveira FÁD, Oliveira Júnior A Zadoks JC, Chang TT, Konzak CF (1974) A decimal code for the
(2013) Productive characteristics of wheat according to nitrogen growth stages of cereals. Weed Res 14:415–421
sources and levels. Pesq Agropec Trop 43:34–41. https://doi. Zhang W, Wu L, Ding Y et al (2017a) Nitrogen fertilizer application
org/10.1590/S1983-40632013000100009 affects lodging resistance by altering secondary cell wall synthe-
R Core Team (2020) R: a language and environment for statistical sis in japonica rice (Oryza sativa L). J Plant Res 130:859–871.
computing. R Foundation for Statistical Computing, Vienna https://doi.org/10.1007/s10265-017-0943-3
Ray K, Banerjee H, Dutta S, Sarkar S, Murrell TS, Singh VK, Majum- Zhang M, Wang H, Yi Y, Ding J, Zhu M, Li C, Zhu X (2017b) Effect of
dar K (2020) Macronutrient management effects on nutrient nitrogen levels and nitrogen ratios on lodging resistance and yield
accumulation, partitioning, remobilization, and yield of hybrid potential of winter wheat (Triticum aestivum L). PLoS ONE.
corn cultivars. Front Plant Sci 11:1307. https://doi.org/10.3389/ https://doi.org/10.1371/journal.pone.0187543
fpls.2020.01307 Zhang Z, Wang Y, Wang Z, Ashraf U, Mo Z, Tian H, Pan S (2022)
Ren B, Huang Z, Liu P, Zhao B, Zhang J (2023) Urea ammonium Precise delivery of nitrogen at tillering stage enhances grain yield
nitrate solution combined with urease and nitrification inhibitors and nitrogen use efficiency in double rice cropping systems of
jointly mitigate NH3 and N2O emissions and improves nitrogen South China. https://doi.org/10.1016/j.fcr.2022.108736. Field
efficiency of summer maize under fertigation. Field Crops Res Crops Res
296:108909. https://doi.org/10.1016/j.fcr.2023.108909 Zhu Y, Qi B, Hao Y, Liu H, Sun G, Chen R, Song S (2021) Appro-
Sadras VO, Lawson C (2013) Nitrogen and water-use efficiency of priate NH4+/NO3–ratio triggers plant growth and nutrient uptake
Australian wheat varieties released between 1958 and 2007. Eur J of flowering Chinese cabbage by optimizing the pH value
Agron 46:34–41. https://doi.org/10.1016/j.eja.2012.11.008 of nutrient solution. Front Plant Sci. https://doi.org/10.3389/
Sadras VO, Slafer GA (2012) Environmental modulation of yield fpls.2021.656144
components in cereals: heritabilities reveal a hierarchy of phe-
notypic plasticities. Field Crops Res 127:215–224. https://doi. Publisher’s Note Springer Nature remains neutral with regard to juris-
org/10.21273/JASHS.116.5.861 dictional claims in published maps and institutional affiliations.
Slafer GA, Kantolic A, Appendino M, Miralles D, Savin R (2009)
Crop development: genetic control, environmental modulation Springer Nature or its licensor (e.g. a society or other partner) holds
and relevance for genetic improvement of crop yield. In: Sadras exclusive rights to this article under a publishing agreement with the
VO, Calderini DF (eds) Crop physiology: applications for genetic author(s) or other rightsholder(s); author self-archiving of the accepted
improvement and agronomy. Academic, San Diego, pp 277–308 manuscript version of this article is solely governed by the terms of
Slafer GA, Savin R, Sadras VO (2014) Coarse and fine regulation such publishing agreement and applicable law.
of wheat yield components in response to genotype and envi-
ronment. Field Crops Res 157:71–83. https://doi.org/10.1016/j.
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