ISSN: 2320-5407 Int. J. Adv. Res.

11(07), 89-93

Journal Homepage: -www.journalijar.com

Article DOI:10.21474/IJAR01/17210
DOI URL: http://dx.doi.org/10.21474/IJAR01/17210

RESEARCH ARTICLE
DEMYSTIFYING THE ROLE OF NITRATE TRANSPORTERS IN PLANT DEFENSE

Aakanksha Wany, Ashutosh Kumar and Rajesh Jha

School of Sciences, P PSavani University, Surat-394125, Gujarat, India.
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Manuscript Info Abstract
……………………. ………………………………………………………………
Manuscript History Plants are frequently exposed to a variety of soil-borne or airborne
Received: 05 May 2023 diseases during its life, which pose a threat tothe safety and security of
Final Accepted: 09 June 2023 the world's food supply. Due to millions of years of evolution, plants
Published: July 2023 have developed multi-layered defense mechanisms to detect and offend
the invasion of these diverse harmful microbes, such as viruses,
Key words:-
Nitrate Transporters, NUE, Nitrate, bacteria, nematodes, and fungi.Plant defense mechanisms are greatly
Ammonium, Pathogen influenced by nitrogen nutrition. The form of nitrogen nutrition, either
nitrate (NO3-) or ammonium (NH4+), plays a very important role in
plant defense. Both these forms of N, have tight coordination with a
group of transporter proteins called nitrate transporters. The critical role
of NO3- transporters in mediating the absorption and its long-distance
transport have been the subject of several researches; however, little is
known about the transport systems involved in NO 3- re-allocation under
the influence of biotic and abiotic stressors. NO3- transporters are
essential to know how plants react to challenging environmental
conditions. In fact, when less NO3- is sent to the shoot, plants adapt to
environmental stress better. The role of NO3- transporters in helping
plants thrives in several challenging environmental cues. As a matter of
fact, under different stresses, the amount of NO3- that is translocated
from roots to shoots fluctuates, which may have an impact on plant's
NUE either favorably or unfavorably.

Copy Right, IJAR, 2023. All rights reserved.

……………………………………………………………………………………………………....
Introduction:-
Nitric oxide- a critical messenger in plant defense
Upon pathogen attack, a number of biochemical elements are elicited which activates many signaling pathways
(Kaur et al., 2022). When there is no pathogen, plants grow and develop by utilizing the available oxygen. But,
during pathogen infection, reactive oxygen species (ROS) are generated in plant tissues (Singla et al., 2019), which
in turn leads to photo-oxidative damage and internal cellular structures (Mittler, 2017).Among ROS, superoxide
(O2•–) and hydrogen peroxide (H2O2)are rapidly produced. This causes rapid oxidative burst and stimulates genes
involved in cellulardefense and protection during hypersensitive response (HR). HR is another crucial
defensemechanism against the spread of microbial diseases. It is defined as rapid programmed cell death in response
of an infection and is specifically localized. This prevents the transmission of pathogen to other regions of the plant
(Balint‐ Kurti, 2019).

During HR, another important signaling molecule is rapidly produced, i.e., nitric oxide (NO) (Delledonne et al.,
1998). NO functions in both abiotic as well as biotic stress (Khan et al., 2023, Fancy et al., 2017). The relationship

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Corresponding Author:- Aakanksha Wany
Address:- School of Sciences, P PSavani University, Surat-394125, Gujarat, India.
ISSN: 2320-5407 Int. J. Adv. Res. 11(07), 89-93

of NO with nitrogen (N) is well known since many decades. N is an important macronutrient and nitrate (NO 3-) and
ammonium (NH4+) are its two readily available forms taken up by the plants and affects plant’s resistance (Gupta et
al., 2013). Nitrate uptake and metabolism may regulate both NO generation and scavenging (Mur et al., 2013).

However, apart from N, NO has a complicated relationship with other minerals such as phosphorus (P),
potassium(K), zinc (Zn), and magnesium (Mg). Graziano et al., (2002) provided the first evidence suggesting a
protective rolefor NO in plant mineral nutrition when Fe-deficient maize plants were supplied with NO
exogenously. Similarly, the role of NO was also evidenced in plants which grow in nutrient deprivation. During
alterations in nutrient supply, plant tissues induce NO production and therefore, a new mechanism involving the role
of NO in nutrient deficiency and plant growth was explained (Buet et al., 2019). Nitrate is readily absorbed by the
roots and its concentration plays a decisive role in lateral root elongation and development (Alvarez et al., 2012).
NO is directly involved in modulating root growth and have a metabolic connection with nitrate.

Nitrogen Use Efficiency

The demand for the use of chemical fertilizers based on N has steadily increased over time because N is a crucial
ingredient needed for plant growth and total productivity. According to Mohanty et al., (2020), 60–70% of applied N
fertilizers are lost leading to serious environmental problems such as pollution, global warming, biodiversity loss,
and significant plant physiological abnormalities. A good crop yield must be maintained while using very little
fertilizer because the rate at which N is being applied is alarmingly high. Therefore, increasing plants’ nitrogen
usage efficiency (NUE) is one of the natural solutions to solve agricultural difficulties. In other words, a key element
in increasing crop yield is effective N use.

Depending on the specific plant species, NUE is a measurement of the amount of seeds, grains, or fruits produced
per unit of available soil nitrogen. According to Bharati and Mandal (2019), NUE can also be described in terms of
N uptake efficiency (NUpE), N transport efficiency (NTE), N remobilization efficiency (NRE), and N utilization
(assimilation) efficiency (NUtE), all of which are significant determinants of NUE in plants.

NO3- and NH4+ are the two forms of N that are made available to plants. Because NO3- is easily leached, its
availability to plants is mostly limited (Jin et al., 2015).NO 3- acts as a signaling moleculeand triggers the activation
of NO3-related genes involved in its absorption, transport, assimilation, vegetative and reproductive development.
According to Iqbal et al., (2020), plants absorb NO3- from the root, assimilate it, and then move it to the shoot where
it can be remobilized to sink organs. The primary forces behind the absorption of NO 3-to the remobilization stage are
NO3-transporters.

Nitrate transporters and NUE

In fact, various researchers have addressed the mechanisms involved in transport, sensing, and signaling processes
while discussing the relationship between NO3-uptake and transport activities in plants (Fan et al., 2017; Zuluaga
and Sonnante, 2019; Vidal et al., 2020). Plants in order to use N sources, maximize the activities of NO 3-
transporters. These NO3- transporters' functions in the enhancement of plant NUE. The impact of necessary nutrients
and environmental signals especially in biotic stress and the expression and activity of NO3- transporters, however,
is less understood.

Nitrate transporters along withcoordinated expression with other transcription factors, have an indisputable influence
on crop productivity and NUE. Not only N, NO3- transporters have impact on the uptakeand utilization efficiency of
other plant nutrients thereby improving NUE in crop plants (Aluko et al., 2023).

Two main NO3- uptake systems have evolved in plants. The low-affinity transport system (LATS) promotes nitrate
uptake in conditions of high soil nitrate (millimolar concentration; > 0.5 mM), whereas the high-affinity transport
system (HATS) drives nitrate in conditions of low soil nitrate (micromolar range) (Iqbal et al., 2020; Raddatz et al.,
2020). It is well known that four families of NO3- transporters, including chloride channel (CLC), slow anion
channel-associated homologs (SLAC/SLAH), nitrate transporter 1/or peptide transporter NPF (NRT1), nitrate
transporter 2/nitrate-nitrite-porter NRT2/NNP, participate in the uptake and transport of nitrate in plants.

NPF (NRT1), NRT2, NRT3 and their homologs have been identified as the main channels involved in root nitrate
uptake and long-distance transport between and among plant organs (Hsu and Tsay, 2013; Wang et al.,
2021b).Members of the NRT2 family, in contrast to the NRT1 family, are high-affinity NO3- transporters (HATs).

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Seven members of the NRT2 family have been described. Out of the seven identified NRT2 transporters, four
(NRT2.1, NRT2.2, NRT2.4, and NRT2.5) have been shown to be directly involved in the influx of NO 3- into
Arabidopsis root cells (O'Brien et al., 2016).

As a result, enhanced plant growth under N shortages and challenging circumstances depends on NO3- redistribution
in plants (Fan et al., 2017). Stressed plants tend to store more nitrate in their roots than normal plants by absorbing
and transporting less NO3- to the shoot. Zhang et al. (2018) used this term SINAR ("Stress-initiated nitrate allocation
to roots") to describe NO3-allocation to the root during biotic and abiotic stress.

Nitrate Transporters genes are upregulated during pathogen infection

There is a lot of agronomic evidence showing how using excess N fertilizer affects the frequency of plant diseases
(Fagard et al., 2014; Fan et al., 2017). A biotrophic disease that depletes plant nutrients, for instance, powdery
mildew, and excessive N fertilizer use increases its severity. It's interesting to note that Arabidopsis tolerance to
Erwinia amylovora is reduced with decreased N fertilizer use. These results suggest an intricate interplay between N
absorption, metabolism, and disease infection mechanisms. As a result, it is clear that N status influences plant
disease susceptibility or tolerance under particular environmental circumstances (Fagard et al., 2014). The molecular
mechanism underlying how NO3-transporters affect fungal infection or pathogenic assault is unfortunately not
copiously known.

Pike et al., (2014) studied the low-affinity transporter NPF3.2 (in grapevine) and cloned the Arabidopsis ortholog
NPF3.1 in order to examine potential processes behind N uptake by the biotrophic pathogen. In this work, it was
demonstrated that the major and minor veins of leaves' vascular tissues and the powdery mildew pathogen infection
upregulate the expression of NPF3.2 and NPF3.1. Under N-deficient conditions, the deletion of NRT2.1 and NRT2.2
led to an increase in resistance to infection by Pseudomonas syringaepv tomato DC3000 (Camanes et al., 2012).
Additionally, using single and double Arabidopsis mutants, the roles of two potential high-affinity NO3- transporters
in the NRT2 family, NRT2.5 and NRT2.6, were examined in response to the rhizospheric bacterium STM196
(Kechid et al., 2013).

According to the study, STM196-induced root system architecture was eliminated and plant growth was hindered by
mutations in NRT2.5 and NRT2.6. Consequently, Arabidopsis leaves expressing NRT2.5 and NRT2.6 seem to be
essential for the plant to respond to STM196 in a way that is independent of NO3- absorption. Both NRT2.5 and
NRT2.6 must be expressed for STM196 to function properly and stimulate plant development (Kechid et al., 2013).
The importance of NRT2.5 in plant biotic defense was recently demonstrated by T-DNA mutants of NRT2.5, which
displayed greater resistance to Pseudomonas syringaepv. tomato DC3000 inoculation than their wild-type
counterparts (Du Toit et al., 2020; Devanna et al., 2021).

These results have shown the functions of NO3- transporters in the plant response to biotic stress and have also
proposed safe, novel, and long-term approaches to agricultural disease management.The expression of a putative
nitrate transporter, OsNPF4.5, appears to be enhanced by mycorrhizal colonization of rice roots (Wang et al.,
2020c).This outcome enhanced the host plant's growth and yield characteristics. However, OsNPF4.5 inactivation
decreased rice's ability to take up symbiotic nitrogen and reduced the incidence of arbuscules (Wang et al., 2020c).

Clustered Regularly Interspaced Palindromic Repeats (CRISPR)/Cas9 combined with the Cas9 nuclease
(CRISPR/CAS9) system was created to further discover the genes boosting NUE. The use of CRISPR/CAS9 has
made it simple and reliable to modify genes for better plant N uptake. The CRISPR/CAS9 technique has been used
in numerous applications in important crops, such as sorghum, rice, and tomatoes (Ito et al., 2015; Ma et al., 2015).
It is noteworthy that CRISPR/CAS9 primarily alters negative growth regulators rather than overexpressing positive
regulators, opening up possibilities for agricultural breeding (Tiwari et al., 2020).

Conclusion:-
Nitrate transporters have not only been demonstrated to play important roles in plant intake and transport capacity,
but also in enhancing plant N utilization, which has ensured the possibility of fulfilling future global food demands.
Increased NUE and plant development are, in fact, dependent upon enhanced NO 3- absorption and use (NO3-
transport, remobilization, and assimilation) through transporters activity. Majority of studies have successfully
demonstrated how nitrate transporters affect stressful environmental conditions (biotic and abiotic stress both). They
have also discussed how they interact with other plant nutrients, in lab; there aren't many field studies that support

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their roles.The established NO3- transporters were discovered to serve diverse physiological roles in environmental
and dietary stressors, although comparatively few NO3- transporters performing complicated interaction functions
have been identified. Among all nitrate transporters, NRT2.1, NRT2.2, NRT2.5 and NPF2.6 have potential roles in
plant defense. The fundamental processes driving these multipurpose roles in plant defenseare still unknown, and it
is also unclear.

Declaration
Authors declare no conflict of interest.

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