Review: Nitrogen assimilation in crop plants and its

affecting factors
Bataung Mokhele1, Xianjin Zhan2, Guozheng Yang1,3, and Xianlong Zhang1
1
College of Plant Science and Technology, Huazhong Agricultural University, Wuhan 430070, PR China; and
2
Cash Crops Institute, Hubei Provincial Academy of Agricultural Science, Wuhan 430070, PR China.
Received 12 July 2011, accepted 15 December 2011.
Mokhele, B., Zhan, X., Yang, G. and Zhang, X. 2012. Review: Nitrogen assimilation in crop plants and its affecting factors.
Can. J. Plant Sci. 92: 399405. In this review we discuss mainly nitrogen assimilation in crop plants and factors affecting
the related process. Nitrogen is a major macro-element limiting the growth and development of plants in agriculture. Both
organic and inorganic forms of nitrogen are metabolized in plants; nitrate and ammonia in soil are common forms of
Can. J. Plant Sci. Downloaded from cdnsciencepub.com by 132.191.0.11 on 05/12/23

inorganic nitrogen that can be metabolized in all plants. There are other nitrogen forms, which include amino acids, nitrite
and urea, that are metabolized in plants. Metabolism normally starts with reduction of nitrate to nitrite, and the latter
further reduces to form ammonium with the presence of relevant enzymes. This reaction occurs more rapidly in leaves in
the presence of light. After ammonia is formed, it enters into the biosynthetic pathways of plant cells, such as reductive
amination and transpiration, to produce different amino acids. Amino acids in cells take part in the synthesis of protein
and other nitrogenous compounds that help in body building. Radiation, gaseous factors, the presence of metals, soil pH
and amount of nitrate are some of the environmental factors affecting absorption and reduction of nitrogen in plants. This
review presents a comprehensive understanding of the assimilation process by crop plants of nitrogen and recommends
that favorable surrounding conditions are the prerequisites for plants to absorb and utilize nitrogen efficiently.

Key words: Nitrogen, absorption, assimilation, nitrate reductase, environmental factors

Mokhele, B., Zhan, X., Yang, G. et Zhang, X. 2012. Assimilation de l’azote par les plantes cultivées et facteurs modifiants.
Can. J. Plant Sci. 92: 399405. Dans cette analyse, les auteurs parlent surtout de l’assimilation de l’azote par les plantes
cultivées et des facteurs qui affectent ce processus. L’azote est un élément nutritif majeur qui limite la croissance et le
développement des plantes en agriculture. Les plantes métabolisent cet élément sous sa forme organique ou minérale; les
nitrates et l’ammoniaque sont des formes courantes d’azote inorganique dans le sol que métabolisent toutes les plantes.
L’azote se rencontre toutefois sous d’autres formes, notamment les acides aminés, les nitrites et l’urée, que métabolisent
aussi les végétaux. Habituellement, le métabolisme commence par la réduction des nitrates en nitrites, puis en ammonium
en présence des enzymes pertinents. La réaction survient plus rapidement dans les feuilles quand il y a de la lumière. Après
formation de l’ammoniaque, l’azote pénètre dans les voies de la biosynthèse des cellules végétales notamment l’amination
réductive et la transpiration, ce qui crée différents acides aminés. Dans la cellule, les acides aminés participent à la synthèse
des protéines et d’autres composés azotés qui concourent à l’expansion de l’organisme. Les rayonnements, les gaz, les
métaux, le pH du sol et la concentration de nitrates figurent parmi les facteurs environnementaux qui affectent l’absorption
et la réduction de l’azote par les plantes. Cet article brosse un tableau complet du processus d’assimilation de l’azote par les
plantes et avance que pour absorber et assimiler efficacement l’azote, les plantes cultivées ont absolument besoin de
conditions ambiantes favorables.

Mots clés: Azote, absorption, assimilation, nitrate réductase, facteurs environnementaux

Nitrogen (N) is an essential component of the proteins
that build cell materials and plant tissue. Nitrogen often
comes from fertilizer application, and, although the
atmosphere is mostly made up of N, only legumes such
as soybeans and alfalfa can convert atmospheric N2 to
plant-available forms via a symbiotic biological process
involving Rhizobium bacteria and the plant roots. Plant-
available inorganic forms of N include nitrate (NO 3 );
and nitrite (NO ); as well as ammonium (NH 
4 ). The
2
concept of plant organic N nutrition relies, to a large
degree, on studies of amino acids. Thus, amino acid N is
in many cases used as a synonym for organic N.
However, urea (NH2)2CO is perhaps the most com-
monly used source of organic N applied as a fertilizer.
Nitrogen assimilation into carbon skeletons repre-
sents a physiological process of the utmost importance
for plant growth and development. Inorganic N is
assimilated into amino acids, namely glutamate, gluta-
mine, and asparagine which play a pivotal role as N-
transport compounds in plants (Lea and Miflin 2003).

Abbreviations: AM, arbuscular mycorrhizal; GDH, glutamate

3 dehydrogenase; GOGAT, glutamine-2-oxoglutarate amino
Corresponding author (e-mail: ygzh9999@mail.hzau. transferase; GS, glutamine synthetase; GSA, glutamine synthetase
edu.cn). activity; NR, nitrate reductase; NRA, nitrate reductase activity

Can. J. Plant Sci. (2012) 92: 399405 doi:10.4141/CJPS2011-135 399

 400 CANADIAN JOURNAL OF PLANT SCIENCE
This review highlights recent studies and progress on
N absorption, N assimilation and the enzymes involved,
as well as the factors affecting the processes in crop
plants.
NITROGEN ABSORPTION
The yield of an agricultural crop strongly depends on
the supply of mineral nutrients, particularly N (Sawan
2006). Because N is a constituent of amino acids, which
are required to synthesize proteins and other related
compounds, it plays a role in almost all plant metabolic
processes (Tucker 1999). Nitrogen is regarded the single
most important growth-limiting factor for crops (Shah
2008), and most plants can utilize N as NO 
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3 and NH4 ;
the available inorganic forms of N absorbed by the roots
(Lasa et al. 2002; Wickert et al. 2007; Oh et al. 2008)
from the soil solution, which then undergo complex
systems of assimilation, transformation, and mobiliza-
tion within plants (Oh et al. 2008). Although inorganic
N is predominantly available to plants as nitrate in most
soils, in certain soil and in hydroponic cultures, ammo-
nium can be the major N ion (Lasa et al. 2002). On the
other hand, both natural and agricultural plants can
take up amino acids from the soil (Godlewski and
Adamczyk 2007).
The biological reduction of atmospheric N2 to ammo-
nium (N fixation) provides about 65% of the bio-
sphere’s available N. Most of this ammonium is
contributed by legume-rhizobia symbioses, which are
initiated by the infection of legume hosts by bacteria
(rhizobia), resulting in the formation of root nodules
(Lodwig et al. 2003). Therefore, legumes are unique
among higher plants because they are able to use either
simultaneously or solely the sources of NO 3 from soil
or fertilizer and N2 through symbiotic fixation in
association with rhizobia (da Silveira et al. 2001).
Plant leaves are a sink for N during the vegetative
stage and, afterwards, this N is remobilized for later use.
The method used in determining total N is time
consuming and may be hazardous as it involves the
use of concentrated sulfuric acid. Therefore, leaf tissue
and sap nitrate are used as indicators of the plant’s N
status, and many farmers use this as a measure in
making decisions regarding fertilizer application rates.
These measurements of leaf tissue nitrate determine
nitrate stored in the vacuoles (Fan et al. 2007). This
follows uptake into the roots, where nitrate allocated to
the leaves is temporarily stored in vacuoles and then
remobilized, especially when N supply is insufficient to
meet demand (von der Fecht-Bartenbach et al. 2010), or
during senescence; N is transported mainly via amino
acids. In rice and wheat, up to 80% of grain N contents
are derived from leaves (Kant et al. 2011). Since most
plants perform nitrate vacuole storage and tolerate high
ion concentrations, it is reasonable to assume that
nitrate has an important function as an osmotic agent
(Wickert et al. 2007).
Because N in the soil can become depleted, through
leaching or being used up by the plants, it has to be
added to the soil. Urea is the N fertilizer most
commonly used in agriculture, and about half of all
N used for crop production is applied as urea (Witte
2011).
NITROGEN ASSIMILATION
During the growth and development of plants, N is
moved into and out of proteins in different organs and
transported between organs in a limited number of
transport compounds. Some of the organic N is moved
between compounds via the activity of transaminases
and glutamine-amide transferases, but a significant
portion is released as NH3 and reassimilated via
glutamine synthetase (GS) (Miflin and Habash 2002).
There are two GS isoenzymes (cytosolic GS1 and
plastidic GS2) in most plant species, each having a
different sub-cellular compartmental location (Tobin
and Yamaya 2001; Cao et al. 2008).
Nitrate Nitrogen
Nitrate N is used in various processes, including
absorption, vacuole storage, xylem transport, reduction
and incorporation into organic forms (Wickert et al.
2007). Primary nitrate assimilation takes place predo-
minantly in the roots of the plant, being strongly
dependent on the age and limitation of space for root
growth (Marquez et al. 2007). Nitrate taken up by plants
is reduced to nitrite by nitrate reductase (NR) (Kuoadio
et al. 2007; Cao et al. 2008; Rosales et al. 2011). This
enzyme catalyzes the reduction of nitrate to nitrite with
pyridine nucleotide in N assimilation in higher plants
(Ahmad and Abdin 1999). Since nitrite is highly
reactive, plant cells immediately transport the nitrite
from the cytosol into chloroplasts in leaves and plastids
in roots. In these organelles, nitrite is further reduced to
NH 4 by nitrite reductase (Rosales et al. 2011).
Ammonium Nitrogen
In plants such as Arabidopsis, ammonium originates
from nitrate reduction, direct absorption, photorespira-
tion, gaseous N (N2) fixation or deamination of
nitrogenous compounds, such as asparagine (Wickert
et al. 2007). All inorganic N is first reduced to
ammonium, because it is the only reduced N form
available to plants for assimilation into N-carrying
amino acids (Ruiz et al. 2007).
Ammonia is then assimilated into glutamine and
glutamate, which serve to translocate organic N from
sources to sinks in legumes and non-legumes. The major
enzymes involved are glutamate synthase, or glutamine-
2-oxoglutarate amino transferase (GOGAT), and gluta-
mate dehydrogenase (GDH) (Lam et al. 1996; Frechilla
et al. 2002; Esposito et al. 2005; Wickert et al. 2007).
However, GS has a vital role in NH 4 assimilation and
the activity of the enzyme is considered to be a critical
and possibly rate-limiting step in NH4 assimilation
 MOKHELE ET AL. * N ASSIMILATION IN CROP PLANTS
(Cao et al. 2008). Temple et al. (1998), in subsequent
studies using raidolabeled (13N) and stable (15N) N
isotopes, enzymes inhibitors and mutants of plant N
metabolism, indicated that the primary assimilation of
NH 4 into amino acids occurs through the joint action
of GS and GOGAT.
Although the GS/GOGAT metabolic pathway is the
main route of N assimilation in higher plants, these
plants possess the ability to use alternative routes, such
as the reversible amination of 2-oxoglutarate to produce
glutamate by GDH (Lasa et al. 2002). GDH is one of
the few enzymes capable of releasing amino N from
amino acids to give a keto-acid and NH3 that can be
separately recycled to be used in respiration and amide
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formation, respectively. GDH may be expected to
function in the deaminating direction in tissues that
are converting amino acids into transport compounds
with a low C:N ratio (Miflin and Habash 2002) or the
potential involvement of N remobilization during senes-
cence (Miyashita and Good 2008).
Ammonia can also be incorporated into the amino
acids, glutamine/glutamate using the C-skeletons pro-
duced via other metabolic pathways, such as respiration
and photosynthesis. All these reactions catalyzed by
NR, including the natural NADH/NADPH-driven
nitrate reduction, are best understood by viewing the
enzyme as a redox system with an internal electron
transfer chain (Cambell 1999).
However, ammonium is said to be toxic to plants,
because it can cause proton extrusion, which is asso-
ciated with ammonium uptake, changes in cytosolic
pH and uncoupling of photophosphorylation in plants
(Wang et al. 2007). Due to the fact that the main
metabolic process in leaf senescence consists of nutri-
ent remobilization, toxic free ammonium, which is
also produced during photosynthesis (Simonovic and
Anderson 2008), should be rapidly converted into the
organic form (amino acids) to avoid negative effects and
provide nitrogenous forms suitable for source-sink
transport (Masclaux-Daubresse et al. 2006).
Urea Nitrogen
Urea occurs ubiquitously in nature, and it is a rapidly
available N source for the growth of various organisms,
including bacteria, fungi and plants. Being the most
used N fertilizer in agriculture, urea plays a role as a
primary N source; it is taken up actively by plants from
the soil solution and is also an intermediate of plant
arginine catabolism, involved in N remobilization from
source tissues (Witte 2011). Urease (or urea amidohy-
drolase) and urea amidolase are the two known distinct
types of non-degrading enzymes, which catalyze urea
assimilation after uptake into the plant cells. They both
hydrolyze urea in the cytosol to CO2 and NH3 (Wang
et al. 2008).
401
Nitrogen Assimilation in Arbuscular Mycorrhizal
Fungi
Mycorrhizal associations occur on almost all terrestrial
plants, and are not as plant-specific as other plant-
microbe associations that have formed between some
plants (e.g., legumes) and bacteria (e.g., rhizobia). There
is increasing evidence of a major contribution by ar-
buscular mycorrhizal (AM) fungi to plant N uptake
and assimilation, adding to the fact that the signifi-
cance of these fungi with respect to N nutrition was less
clear, even though the role of mycorrhizal fungi in
phosphorus nutrition of their hosts was well documen-
ted (Faure et al. 1998).
Mycorrhizal fungi, as shown by Singh (2007), have
the ability to assimilate N in a pattern closely similar to
plant roots, and mycorrhizal fungi are known to
produce NR and ammonium-assimilating enzymes.
Nitrate can be reduced inside the AM fungal cells by
the assimilatory reduction pathway. Nitrate mobilized
from soils by an AM fungus could be transferred
directly to the root cells where it will be assimilated
(Ruiz-Lozano and Azcon 1996).
The effect of vesicular-AM fungi in onion on gluta-
mine synthetase activity (GSA) under water stress
( 0.17 MPa) was evident only in roots being compared
to those found in phosphorus fertilized plants; hence
it was concluded that there is a direct effect on
absorption, translocation and assimilation of both N
forms by endomycorrhizal systems (Singh 2007).
Furthermore, it was concluded by Ruiz-Lozano (2003)
that levels of nitrate reductase activity (NRA) were
higher in AM plants under drought stress compared
with non-mycorrhizal plants, and, thus, mycorrhizal
plants were able to tolerate drought in terms of plant
biomass production.
FACTORS AFFECTING NITROGEN ABSORPTION
AND ASSIMILATION
Nitrogen Levels
Plants that exhibit low rates of NO 3 in roots export
most of the absorbed NO 3 to shoots, where it is reduced
and incorporated into amino acids (Aslam et al. 2001).
Some changes in NRA occurred independently of total
N concentrations. Maighany and Ebrahimzadeh (2004),
in a study on intervarietal differences in N content and
nitrate assimilation in wheat under salt stress, found
that NRA was more highly correlated with N levels in
roots than in leaves. In leaves, a direct correlation
between N content and NRA was found only at tillering
and then an inverse relation was observed between
them. Besides, the impact of salt was more evident on
altering the activity of NR and N content in the leaves
than in roots. This suggests that salt stress can alter N
level and NRA, and the effect varies with cultivar,
organ, developmental stage and the degree of salt stress.
Glutamine synthetase activity decreased when ammo-
nium appeared in the medium replacing nitrate as the N
 402 CANADIAN JOURNAL OF PLANT SCIENCE
source. Also, by increasing ammonium concentration,
GSA decreased slightly, mainly at high ammonium
concentration (5 mmol L1), and it was observed that
GDH in the roots of pea plants was stimulated at
low ammonium concentrations (1 mmol L1) (Frechilla
et al. 2002).
Aslam et al. (2001) reported that while the amino
acids directly inhibited enhancement of the activity of
the NO 3 uptake system, independent of NO3 avail-

ability, inhibition of the induction of NRA was due to a
decrease in NO 3 uptake, resulting in decreased NO3

availability. Also, a rapid cycling of amino acids
between roots and shoots occurs whether NO 3 assim-
ilation is localized mainly in the shoots or the roots.
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The amino acids, whether accumulated in the plant
internally or supplied externally, usually down regu-
late the induction of NO 3 uptake and reduction
systems. Because amino acids decreased the concentra-
tion of NO 3 ; this indicates that inhibition was due
to insufficient NO 3 : Aslam et al. (2001) further reported
that enhancement of RNA was not inhibited by amino
acids when the roots were supplied with 10 mM NO 3 :
Therefore, this means NO 3 should always be at ade-
quate levels.
On the other hand, it seems that when plants need to
assimilate large amounts of N, both glutamine and
asparagine are the preferential form in which N is
assimilated and translocated. This is due to the fact
that these molecules show a low C:N ratio, which
represents an advantage for ammonium incorporation
to non-toxic forms (Frechilla et al. 2002).
Gaseous Factors
Low NO2 had no effect on the organic N content of the
plants, or the concentration of organic N in leaves
and roots, with the exception that it slightly increased
the concentration of organic N in the leaves of plants
grown at low nitrate levels. Nitrogen dioxide absorbed
by the plants can be converted into nitrate and nitrite.
As a result it follows that assimilation will take place.
The amount of nitrate taken up per plant and the uptake
rates were not significantly affected by exposure to low
NO2. However, exposure to high NO2 causes increase in
nitrate concentration (Qiao and Murray 1998).
In cucumber leaves, high rates of CO2 assimilation
enhanced nitrate reduction by stimulating the synthesis
and activity of NR, and sugars derived from CO2
assimilation probably act as positive regulatory meta-
bolites. This was a similar in sunflower leaves, where
the expression and GSA were modulated by the rate of
CO2 assimilation after brief exposure to high atmo-
spheric CO2, and photosynthesized sugars are presum-
ably involved as regulatory metabolites (Aguera et al.
2006). On the other hand, N assimilation may regulate
simultaneous CO2 assimilation, because N assimilation
into glutamate is a very important sink for redox
equivalents from the photosynthetic electron flow.
When N is assimilated quantitatively in the root,
ammonium-grown plants save about 7.611.9% of the
overall cost of growth as compared with nitrate-grown
plants; for assimilation in the shoot the difference is
reduced to 3.06.1% because of the direct use of
photons. Leaf NO 3 assimilation into amino acids is
linked to the consumption of carbon skeletons and
photosynthetically generated ATP and reductants
(NADPH) (Guo et al. 2007).
However, Guo et al. (2007) established that elevated
CO2 (or low O2) atmospheric concentrations decrease
rates of photorespiration and initially enhance rates of
photosynthesis and growth by as much as 35% in most
plants (C3 plants). This is in agreement with Bloom et al.
(2010), who reported that atmospheric CO2 enrichment
does not stimulate the growth of wheat plants receiving
NO 3 as a sole N source to the same extent as those
receiving NO 4 ; hence they concluded that elevated CO2
(or low O2) atmospheric concentrations, which are
conditions that decrease photorespiration, inhibit NO 3
assimilation in the shoots of C3 plants.
Ultra-violet B Radiation
Increases in solar UV-B have raised concerns regarding
its damaging impact on crop plants. UV-B radiation can
impair all the major processes of photosynthesis,
including photochemical reactions in thylakoid mem-
branes, enzymatic processes in the Calvin cycle, stoma-
tal limitations to CO2 diffusion, destruction of amino
acid residues, and oxygen-mediated damage to unsatu-
rated fatty acids in plant cell membranes. Current global
terrestrial UV-B radiation ranges between 2 and
12 kJm 2 d1 on a given day, with near-equator and
mid-latitudes receiving higher doses, which includes an
increase of 614% since the 1980s (Surabhi et al. 2009).
Though plants need sunlight for processing their own
food, exposure of high radiation poses a danger to the
health of plants in general.
Cao et al. (2007) stated that the absorption and
assimilation of the important minerals for plant nitrate
were significantly inhibited under UV-B radiation,
especially at high levels; correspondingly, the growth
of seedlings was restrained. Under low levels of UV-B
radiation, GDH activity was enhanced to prevent
ammonia toxity, when the activity of GS and GOGAT
decreased sharply, leading to the accumulation of
ammonia in plants, the high accumulation of which is
toxic. Conversely, under high levels of UV-B radiation,
the structure of GDH or some related genes might be
damaged, decreasing GDH activity.
Both the leaves and roots showed decreased values of
NRA after exposure to UV-B radiation in comparison
with control seedlings (Quaggiotti et al. 2004), in
agreement with data obtained in Vigna unguiculata L.
(Balakumar et al. 1999) and barley (Ghisi et al. 2002).
Metals
Contamination of agricultural soil by heavy metals has
become a critical environmental concern due to their
 MOKHELE ET AL. * N ASSIMILATION IN CROP PLANTS
potential adverse ecological effects. Such toxic elements
are considered to be soil pollutants owing to their
widespread occurrence, and their acute and chronic
toxic effect on plants grown in such soils (Yadav 2010).
Besides being toxic, many studies have revealed their
positive and negative side effects on the metabolic
processes governing growth and development depending
on the quantity of such metals in the soil.
La could promote the uptake and transport of nitrate
in plants, which might be related to the fact that La
has a positive effect on the transpiration of seedlings,
promoting the uptake and movement of water and
mineral nutrients. Low levels of La promoted GDH
activity (Cao et al. 2007), but the amount of nitrate
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taken up from the nutrient solution was dramatically
reduced by Cd. The total nitrate uptake in plants treated
with 50 mM Cd for 10 h was about 20% of the nitrate
uptake in control plants; the decline in net nitrate
uptake was even higher in plants treated with 100 mM
Cd (12% of nitrate uptake in control plants) (Gouia
et al. 2000).
High Al concentrations directly restrict NO 3 uptake
in most plant species studied to date and therefore cause
a deficit in N metabolism. At the same time it has been
observed that Al can act directly on NO
3 assimilation
inside the plant, inhibiting NR activity. The same was
true for excessive application of Mo, leading to reduced
normal plant growth (Ruiz et al. 2007).
The mechanism by which plants take up nutrients
is through the soil, whereby the root hairs release
hydrogen ions (H) into the soil. These hydrogen ions
displace cations attached to negatively charged soil
particles so that the cations are available for uptake by
the roots. Some metals can inhibit uptake of nitrate,
affecting the activity of NR.
Soil Salinity
Soil salinity is a major factor rendering soil unfit for
agriculture. Cordovilla et al. (1999), experimenting on
the effects of NaCl on the growth and N fixation and
assimilation of inoculated and KNO3 fertilized Vicia
faba L. and Pisum sativum L. plants, found that the
salinity response of fababean plants was generally
consistent with results obtained for Cicer arietinum,
Medicago trunculata and other genotypes of V. faba in
that N-fixing plants were more sensitive to salinity than
were N-fertilized plants.
Salt dissolved in irrigation water could interfere with
N use by plants, since salinity has previously been shown
to be a major factor responsible for low N availability
(Debouba et al. 2006), because nitrate reductase activity
in leaves is largely dependent on nitrate flux from roots,
and is severely affected by osmotic shock induced by
NaCl (Silveira et al. 2001). NO 3 levels were highly
decreased in both the leaves and roots in tomato
(Lycopersicon esculentum) seedlings under increasing
salinity. In the leaves, a substantial decrease in nitrate,
of 60 and 80%, occurred at 50 and 100 mM NaCl,
403
respectively. Similar decreases were found for NO 3
contents in the roots (Debouba et al. 2006).
In legumes, salt stress from 50 to 200 mM NaCl
significantly limits productivity by interfering with plant
growth (Cordovilla et al. 1999). It is generally observed
that salt stress promotes the accumulation of ammo-
nium, nitrate, and free amino acids in plants, and
decreases the incorporation of ammonium into amino
acid compounds.
In addition, one of the increasing contaminants is
CIO 3 ; which is a component of industrial waste water
and disinfectants of drinking water. Its release into the
environment causes inhibition of nitrate reductase and
growth due to the production of ClO . However, the
toxic effect was strong if the ambient nitrate concentra-
tion was low (Tischner 2000).
Crop plants have an efficient mechanism to absorb
and utilize N from the soil. However, the key processes
or molecule(s) involved in these processes remain
uncertain. Further research is required to understand
the functions and mechanisms of various environmental
factors acting on the plant’s ability to absorb N.
ACKNOWLEDGEMENTS
The study was supported by the Professional (Agricul-
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the Modern Agro-Industry Technology Research Sys-
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