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Low Nitrogent availability

Individual Study 2019.
Low Nitrogen Availability in Plant: Transport
Mechanism and its Effects in Root Shape and Leaf
Adillatul Lathiifatun Jannah.
Abstractitrogen is a essensial macronutrient which is required for plant growth and
development. Mostly plant absorb nitrogen from soil in form of nitrat and ammonium. Since the
concentration of nitrogen in the soil is heterogen, plants do some mechanism to confront this
problem. Some mechanism that plants do are through the regulation of nitrogen acquisition
efficiency and changing in morphology and physiology process. This review provides information
about some plant responses in low nitrogen availability: (i) nitrogen transporter and transport
mechanism in low nitrogen availability, (ii) low nitrogen effect in root shape, (iii) low nitrogen effect
in leaf component.
Keywords leaf, nitrogen, root, transporter.
Nitrogen is being a major food for
plants and it plays the most important role
in plant. This is because all processes in
plants are associated with protein, where
nitrogen is the important constituent of
protein. Mitrogen also the major
constituent of nucleotides, chlorophyll and
many cellular component (Lawlor, 2002).
Some function of nitrogen in plant are
imparting the green colour (Sorin et al.
2016), promoting the growth and root
morphology (Razaq et al. 2017), promote
young leave and stem (Oskarsson et al.,
2006), increasing biomass and crop
productivity (Massignam et al., 2009), and
also being found as component of
secondary metabolites in plant (Cartelat et
al., 2005). The distribution of nitrogen in
the soil is heterogen, in response to this
heterogenous distribution plants do
nitrogen use efficiency that is defined into
two ways, that are omponents, Nitrogen
utilization efficiency and N acquisition
(uptake) efficiency, this mechanism is
facilitated by influx transporters located on
the plasma membrane, and the latter by
alterations in growth and development in
response to local and systemic nitrogen signals
(Forde, 2014 and Krapp et al. 2014). Another
mechanism that plants do are trough changing in
morphology such as root architecture and in
physiology process such as producing
secondary metabolite. This review aims to
summarize some plant responses in low
nitrogen availability. This review mostly focus
on plant transport mechanism, effect in root
shape and leaf component.
Nitrogen Transporter and Transport
Plants acquire nitrogen from the soil
through their roots in inorganic form such as
nitrate (NO3-) and ammonium (NH4+).
Ammonium and nitrate are taken up into root
cells activelly by different transporters which
is localized in different sets of plasma
membrane. Ammonium is transported by
(Ludewig et al. 2007), while nitrate is
transported by two families of transporters,
Individual Study 2019.
(NRT 1 family) and NRT 2 family (Krapp
et al. 2014)
Nitrate consentration in the soil can
be vary from micromolar to millimolar, to
confront with low condition of nitrogen in
soil plant have evolved nitrate transport
system called High Affinity Transport
System (HATS), in Arabidopsis thaliana
the gene that contribute in this sytem are
NRT 2 family (Orsel et al., 2002) and AMT
1.1 (Yuan et al. 2007), that system is
opponent of Low Affinity Transport
System (LATS) which active in high
consentration of Nitrogen using NRT 1.2
gene (Huang et al. 1999). And also NRT
1.1 which have role in both of types of
transport system (Ho et al.2009).
According to the study by Riveras
et al. (2015), one of mechanism that
activates NRT 2.1 is through [Ca2+]cyt
signal (Figure 1), increasing of calcium
activates gene expression of AtNRT 2.1 via
TGA1/TGA4 which is triggered by
Phospholipase C (PLC). They also found
that AtNRT 1.1 is the first regulator of that
mechanism, which activates PLC.
Contribution of AtNRT 1.1 in that pathway
is because AtNRT has a role as nitrate
sensor. Ho et al. (2009) found that AtNRT
1.1 is involved in primary nitrate response
of AtNRT 2.1 gene and serves as a nitrate
sensor by evaluating CHL1 deletion
mutant (chl1-5) and single amino acid
substitution mutant (chl1-9).
Study by Ma et al. (2015) also
found another way which activate another
HATS genes (NRT 2.4 and NRT 2.5)
through the calcium sensor CLB7. This is
realated to that Ca2+ is one of second
messenger in nitrate signaling pathway,
their study found that in the mutation of
cbl7 significantly decreased the expression
of NRT2.4 and NRT2.5 under low N
availability (Figure 1). Another study that
indicate the activates of HATS gene is found by
Tabata et al. (2014), which found that under low
ENCODED PEPTIDEs (CEPs) is translocated
to the shoot, then CEPs are perceived by CEP
RECEPTOR1/2 (CEPR1/2) that increase
AtNRT2.1 gene (Figure 1), but the way of
CEPR1/2 activates AtNRT2.1 is not
described in the study.
Figure 1. Three kind of pathway that activate High
Affinity Transport System (HATS) gene in low
nitrogen availability.
Low Nitrogen Effect in Root Shape
Low concentration of nitrogen in soil
can make change in root. This is because root is
the first organs that senses mineral starvation in
the soil. One of consequence is the change of
root shape. Ma et al. (2014) indicate that low
nitrogen availability reduce auxin content in the
aerial organs and stimulate accumulation of
auxin in lateral root primordia, this stimulation
is promoted by expression of TAR2 which
expressed in the pericycle and vascular of the
Individual Study 2019.
root maturation zone, this case stimulate
the development of lateral root emergence,
make the root has more branch (Figure 2).
Figure 2. Illustration of low nitrogen
concentration effect in root. Source: Zhang et
al, 2007
Low Nitrogen Effect in Leaf
Beside changing root shape, low
nitrogen condition in plant also have effect
in leaf composition. One of physiological
effect is decreasing the chlorophyll content
that cause chlorosis that accelerated leaf
senescence. Wei, Meng (2015) indicate that
nitrogen deficiency decrease chlorophyll,
essential amino acids, such as Lys, Phe,
isoleucine (Ile), tryptophane (Trp), leucine
(Leu), and valine (Val) content, and
nonessential amino acids, consisting of
glutamic acid (Glu), aspartic acid (Asp),
Gly, arginine (Arg), and proline (Pro)
content. Differently with chlorophyll and
amino acids, in low nitrogen availability
another leaf component such as some
secondary metabolites content will
increase. Benard et al. (2011), found that
phenolic compounds including chlorogenic
acid and rutin is increasing in Tomato leaf.
But in different plant the type of compunds
that accumulated will be different. The
of chlorophyll and
polyphenolics content can be used as
indicator of plant nitrogen status (Cartelat
et al., 2005).
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