MINERAL NUTRITION

Macronutrients
Nitrogen:
Assimilation of nitrogen:
The major sources of inorganic nitrogen are
-Nitrate
-Ammonium
Most of ammonium are incorporated into organic compounds in the root, while
nitrate is mobile in xylem or stored in vacuoles. Nitrate has also to be reduced into
ammonia to be incorporated into organic compounds.
The importance of reduction and assimilation process of nitrate are as important
as carbon dioxide reduction and assimilation.

Nitrate assimilation and reduction:
the currently accepted nitrate reduction pathway in higher and lower plants is as
follows:
NO
3
-
+ 8H
+
NH
3
+ 2H
2
O + OH
-

Some bacteria uses nitrate as an electron acceptor in anaerobic conditions and
produce nitrogenous gases causing loss of combined nitrogen from soil by denitri-
fication.
Reduction of ammonia is mediated by two separate enzymes:
-nitrate reductase which reduces nitrate into nitrite
-nitrite reductase which reduces nitrite to ammonia.
Nitrate reductase: is a complex high molecular weight enzyme, it have several
prosthetic groups such as FAD and molybdenum.
It is localized at the cytoplasm and requires NADP or NADPH as e-donor.
Its activity can be inhibited by ammonium and certain amino acids or amides. Also
it is very low in molybdenum-deficient plants.
Nitrite reductase: converts nitrite to ammonia-in higher plants- without release of
intermediates.
It has a low molecular weight
In leaves it is associated with chloroplasts where the reduced ferredoxin is the
e-donor and in roots with protoplastids where an unknown e-carrier between
NADPH and the enzyme is present.
It is rarely accumulates in intact plants under normal condition and is present in
much higher concentration in tissue than nitrate reductase. Root nodules are
exception of this role.
Certain herbicides can inhibit nitrate reductase in leaves and correspondingly
increase nitrite level in tissues.
In C
4
plants mesophyll and bundle sheath cells differ in nitrate assimilation as they
differ in CO
2
assimilation. Because nitrate and nitrite reeducates are localized in
the mesophyll and are absent in the bundle sheath cells.

Localization in roots and shoots:
In most plants shoots and roots are capable to reduce nitrate and the
proportion of reduction depends on:
1- level of nitrate supply
2-the plant species
3-the plant age.
and have effect on:
1-mineral nutrition
2-carbon economy of plants
When nitrate supply is low, a high proportion of nitrate is reduced in
roots. When increased supply reduction capacity limited and increasing
proportion of nitrogen is translocated to the shoots as nitrate.
The large carbohydrate requirement for reduction is one of its limiting
factors. The capacity of reduction also differ among different species or
cultivars of a species.
Leaf age: during leaf ontogeny the maximum activity of nitrate reductase is
when the rate of leaf expansion is maximal. In fully expanded leaves the reduc-
tase activity is very low and thus the nitrate level are high.
In roots the nitrate reductase activity is high in expanding cells in apical zones
and declines rapidly towards the basal zones.
Light: there is a close correlation between light intensity and nitrate reduction.
The nitrate reductase mechanism shows that this light effect reflects carbohy-
drate level variation and in supply reducing equivalents (ferredoxin and
NADPH), light also affect the enzyme stability.
Nitrate assimilation and osmoregulation:
In plant species that assimilate most or all nitrate in shoots and synthe-
size organic acid anions in the cytoplasm and stored in the vacuole to
maintain both cation-anion balance and intercellular pH.
This will result in osmotic problems if nitrate reduction were continue af-
ter the termination of leaf expansion.
Several mechanisms exists for excess osmotic solutes removal:
1. Precipitation of excess solutes in an osmotically inactive form. E.g. the
precipitation of calcium oxalate.
2. Retranslocation of reduced nitrogen (amino acids and amides) togeth-
er with phloem-mobile cations to growth areas.
3. Retranslocation of organic acid anions with potassium into roots and
release of an anion after decarboxylation. This can be used for nitrate
uptake in exchange for released anion.
Assimilation of ammonium:
Ammonium and its equilibrium partner ammonia are toxic at quit low
concentrations. NH
3
NH
4
+
+ OH
-

the formation of amino acids, amides and related compounds is the
main pathway for its detoxification.
The main steps in assimilation ammonium ions supplied from roots are
-uptake into root cells with simultaneous release of proton for charge
compensation
-incorporation into amino acids and amides.
Nearly all of the assimilated ammonia is translocated to the shoots as
amino acids, amides and related compounds.
Ammonium assimilation in roots has a large carbohydrate requirement
because of the need for carbon in amino acid and amides.
In order to minimize carbon loss from nitrogen transport -which occurs
through xylem- one or rarely two or more of the nitrogen compounds
(N/C > 0.4) dominate in xylem exudate of the root. These compounds
take the bulk of transport they are:
-the amides glutamine (2N/5C) and asparagine (2N/4C)
-the amino acid arginine (4N/6C)
-the ureide allantion (4N/4C)
The low molecular weight organic compounds used predominantly for
long distance transport or for storage in individual cells differs among
plant families.
Although the different sites of ammonia assimilation, two key enzymes
are involved, glutamine synthetase and glutamate synthase.
In this pathway the amino acid glutamate acts as ammonia acceptor and
glutamine is formed.
Glutamine synthetase: has a very high affinity for ammonia (low K
m
val-
ue) thus it works even on low ammonia concentrations.
-it is activated by high pH and high concentration of both magnesium
and ATP. These three factors are increased in the chloroplast stroma.
-thus in chloroplasts light stimulated nitrate reduction and ammonia as-
similation are coordinated to prevent ammonia toxicity.
Glutamate synthase (GOGAT): catalyze the transfer of amide group (-
NH
2
) from glutamine to 2-oxoglutarate (product of tricarboxylic acid cy-
cle)
-either reduced ferredoxin, NADH or NADPH are required.
-cycle results in two molecules of glutamate one for cycle maintenance
and the other can be utilized for protein biosynthesis.
-when ammonia supply is large both molecules act as ammonia acceptor
and one glutamine molecule released.
Glutamate dehydrogenase: localized principally in mitochondria of root
and leaves and have low affinity to ammonia (high K
m
) inconsistent with
the need to maintain the low intracellular ammonia concentration.
Amino acid and protein biosynthesis:
the organically bound nitrogen of glutamate and glutamine can be utilized for
the synthesis of other amides, ureides, amino acids and proteins.


Role of low molecular weight organic nitrogen compounds:
Intermediates between the assimilation of inorganic nitrogen and high
molecular weight compounds.
Higher plants are neither able to excrete organic nitrogen e.g. as urea.
But they store it in nontoxic forms as urea derivatives e.g. allantion. Nor
they capable for reoxidizing it to nitrate which would be a safe storage
form.
The tripeptide glutathione function in chloroplast redox system and in
long distance transport of reduced sulfur in the phloem.
Several antibiotics such as valinomycin are low molecular weight poly-
peptides.
Some are involved in the long distance transport of certain heavy metals
in xylem.
They are precursors for amine synthesis. Amines are component of the
lipid fraction of the biomembranes.
They are also involved in osmoregulation in higher plants. Compounds
like proline are synthesized and salt or water stress and accumulate to
counter act the osmotic disturbance.
Ammonium versus nitrate nutrition:
The genotypic pattern of plants are:
Calcifugus plants that are adapted to acid soils and low soil redox poten-
tial and have preference for ammonia.
Cacicole plants that are adapted to high pH and calcareous soils and
have preference for nitrate.
The uptake and utilization of ammonia are higher than nitrate at low
temperature for all plants.
Highest growth rates are gained when combined ammonium and nitrate
nutrition or ammonium only.
Effects of the two nitrogen sources on plant growth are expected be-
cause their different effects on cation-anion balance, on root induced
rhizosphere changes and on energy metabolism. Ammonium generally
inhibit cation uptake and can depress growth by inducing magnesium
deficiency. Ammonium also unlike nitrate increases root respiration
which results in enhanced root exudation and hence increased bacterial
growth.
Growth inhibition by ammonium is closely related to the fall in substrate
pH. At low pH the ammonium uptake is not depressed as other cations
which increases the cation anion imbalance.
Nitrogen supply, plant growth and plant composition:
Nitrogen content required for optimal growth 2-5% depending on the plant
species, developmental stage and organ.
Sub-optimal supply:
1. Plant growth retarded
2. Enhanced senescence of older leaves.
Supra-optimal supply:
1. Inhibit root elongation which is unfavorable for nutrient gaining and wa-
ter uptake.
2. Enhance shoot elongation which increases susceptibility to lodging and
yield limiting factor
3. Increase length, width and area of the leaves thus interferes with light
interception
4. Induce changing in phytohormones balance.