First lesson: Introduction of Plant Nutrition

I. Scope and Evolution of plant nutrition

* The beneficial effect of adding elements to soils to improve plant growth

and plant yield had been known in agriculture for more than 2000 years ago,
even 150 years ago it was still a matter of scientific controversy as inorganic
elements for plant growth.
* First time, Justus Von Liebig (1803 – 1873) collected the scattered
information concerning to importance of inorganic elements for plants
growth and summarized them and thus the mineral nutrition of plants was
established as scientific subject
* These achievements led to a rapid increase in using mineral fertilizers by
the end of 19th century. In Europe, the amounts of potassium salts, and
super-phosphate and inorganic nitrogen ware used in agriculture to improve
plant growth and crops yield.
* Especially, the beginning of 20th century the growing rate of the world
population required more foods thus the farmers had to try to exploit fully
the bio-energetic potential of soils as well as the yield potential for the
cultivated species and cultivars by more application of mineral nutrients.
* The progress made at the middle of 20th century was the method of study
of mineral nutrition by the solution culture, this method led to new way in
cultivation called hydroponic (agriculture without soil that will be very
useful for next future
* Mineral nutrients are essential for plant growth and development, and it is
an area importance of both basic and applied sciences. Impressive progresses
has been made during last decades of 20th century in our understanding og
mechanisms of nutrient uptake and their functions in plant metabolism, at

1
the same time there have also been advances in increasing crop yields by the
supply of mineral fertilizer application.
* Up to now, Agronomists have taken into account the between mineral
nutrients and the yield formation of crops has close relationship many
experiments have been carried out to study this area. The results of studies
had shown that crop yield is depended on applied mineral nutrients.
* However, now in agriculture we have to face with many disadvantages
such as soil and underground water pollution by application inorganic
fertilizers, losses of mineral elements applied to the soils because low rate of
mineral nutrients applied to soil is taken up by plants, in generally: 40 – 50%
for phosphorus and nitrogen. What way can we improve this rate? This is
very difficulty question for us that indicating this area has many secretes
needed to discover.
II. Position of the subject

* The subject of mineral nutrition of plants is both basic and applied

sciences for training to specialists in agriculture and it is related to many
other areas such as plant physiology (structure of cells, photosynthesis,
respiration, water absorption..), biochemistry of plants as assimilation,
formation of mineral nutrients in the plants, organic matters contain in
the plant organs
* Other fields also related to mineral nutrients as organic and inorganic
chemical subjects.
III. Scope of study.

Mineral nutrition of plants solves to following topics:

a. Classification of mineral nutrients of plants

2
 b. Mechanisms of mineral uptake in individual cell and
translocation of mineral elements in the plants
c. Functions and assimilations of mineral nutrients by plants,
deficiencies and toxicities
d. Plant and soil relationships.
e. Mineral nutrition and diseases and pests
PART A: NUTRITIONAL PHYSIOLOGY
I. The definition and classification of mineral nutrients
1.1. The essential mineral elements:

They are defined as elements that are necessary for plant to complete its life
cycle (normal growth and development) and for which no other elements can
be substituted and have defined role in plant metabolisms. An essential
element is not only a constituent of an essential metabolic process but also is
needed for an enzymatic function such as nitrogen, phosphorus, potassium,
calcium, magnesium, sulfur, boron, chlorine, iron, manganese, zinc, copper,
molybdenum and nickel.
1.2. Beneficial elements: (B. E.)

B. E. are elements that promote plant growth in many plant species but are
not absolutely necessary for completion of the plant life cycle, they are
silicon (Si), sodium (Na), cobalt (Co) and selenium (Se).
1.3. Classification of essential elements

* In the essential elements still consisting of three other elements, they are
carbon, oxygen and hydrogen coming from carbon dioxyte and water under

3
reaction of green plant photosynthesis. These elements are called organic
elements they are present in 95% in biomass of plants.
* Other elements among essential elements, that plants uptake from the
soils, are called the mineral nutrients or nutrient elements
1.4. Classification of mineral nutrients
a. On the basic of the amounts found in the plants: The mineral nutrient
elements can be classified three following groups:
* Macronutrients: N, P, K, Ca, Mg, S, Na, and Si with content of
10-1 to 10-4 % in DW and consist of 99,95% of the total
* Micronutrients: Cl, Fe, B, Mn, Zn, Cu, Mo, and Ni with content
of 10-5 – 10-7 % in DW
* Trace elements: As, I, Se, Cd, Pb with content of lower 10 -8 %
in DW
b. On the basic physiological and biochemical functions in the plants:

* Structural functions, they are a part of the organic compounds that build
plant organs such as N, S and P
* Containing energy and to sustain of cell structure as P, Bo, Si
* Fortifying the activation of enzymes and regulation of omosic-
permeability of plant cells
* Chains of electrical transport as Fe, Cu, Zn and Mo
c. On basic the translocation in the plants:

* Re-transport elements (mobile elements) in the plants: N, K, Mg, P, Cl,

Na, Zn, and Mo, with those elements, if they are deficiency, they can
remove from old organs to young organs and deficient symptoms are in old
leaves first

4
 * Non-retransported elements in the plants are Ca, S, Fe, Cu, and Bo, with
those elements if they are deficiency, they can not remove from old leaves to
young leaves and deficient symptoms are young leaves first.
Another classification of mineral elements following:
Table 1.1
Classification elements higher plants lower plants
Macronutrients N, P, S, K, Mg, Ca + + (exception Ca for fungi)
Micronutrients Fe, Mn, Zn, Cu + + (exception B for fungi)

Micronutrients and Na, Si Co + +

Beneficial elements

II. Ion uptake mechanic of individual cells and root shoot distance
transport
2.1. General:

The results obtained from both lower and higher plants demonstrate that ion
uptake is characterized by following:
a. Selectivity: Certain mineral elements are taken up easily while others
are impossible to transport into membrane cells
b. Accumulation: The concentration of mineral elements in the plant
cell can be much higher than in the external solution
c. Genotype: There are distinct differences among plant species in ion
uptake characteristics. Those results pose many questions, for example, how
do individual cells and higher plants regulate ion uptake? Its ion uptake is a
reflection of demand or is ions that don’t play role in plant metabolism or
that are even toxic also taken up?

5
2.2. Pathway of solutes from the external solution into the cells
2.2.1. Influx into the apoplasm

* Movement of low molecular solutes (ions, organic acids, amino acids,

sugars..) from the external solution into the cell walls of the individual cells
or roots are a non-metabolic passive process driven by diffusion or mass
flow. The cell walls can interact with solutes and thus may facilitate or
restrict further movement to the uptake sites of the plasma membrane of
cells or roots
* Primary cell walls consist of a network of the celluloses, and hemi-
celluloses, this network contains pores that differ in size. For root hairs
of a maximum diameter of 3.5 – 3.8nM (35- 38 A0) has been calculated.
The maximum values for the pores in plant wall cells are in range of
5nM (50A0) comparison hydrated ions such as K+, Ca++ are small shown
in table 2.2
Diameter (nm)
Rhizodermal cell wall (Maize) 500 – 3000
Cortical cell wall 100 – 200
Pores in cell wall < 50
K+ o.66
Ca++ 0.82
Thus hydrated ions are ease to move into the free space of cells
* In contrast to mineral nutrients, high molecule weight solutes (as metal
chelates, fulvic acids or viruses) are mostly restricted or prevented by the
diameter of pores from entering free space of root cells.
* In this network, a variable proportion of the pectins consist of
polygalactorunic acid, originating mainly from the middle lamella.

6
According to both in roots and in the cell wall of other plant tissues called
apoplasm the carboxylic groups (R. COO-) act as cation exchange in roots,
cations from the external solution can accumulate in a non-metabolic step in
the free space, whereas anions are repelled (impossible). Because of the
negative charge, the appoplasm does not provide free space for the
movement of charged solutes.
* Hope and Steven (1952) introduced the terms apparent free space (AFS)
this comprises the water free space (WFS), which is freely accessible to
ions charged and uncharged molecules, and the Donan free space (DWS),
where cation exchange and anion repulsion take place. Ion distribution
within the DFS is characterized by the typical Donan distribution which
occurs in soils at the surface of negative charge particles. Divalent cations
such as Ca2+ are bound to these cation exchange sites. Plant species differ
considerably in cation exchange capacity (CEC) that is in the number of
cation sites located in cell walls.
* As a rule, the CEC of dicotyledonous species is much higher than that
of monocotyledonous species
Plant CEC (mg/100 g in DW)
Wheat 23
Maize 29
Bean 54
Tomato 62
DW: dry weight
* A positive correlation can be observed between the CEC and the ratio of
Ca++ to K+ contents in different plant species. Effective competition between
H+ or mono and polyvalent aluminum species or both with magnesium for
binding sites in the apoplasm of roots is main factor responsible for the

7
depression in Mg++ uptake and appearance of Mg++ deficiency in annual
plants (Rengel, 1990) and forest tree species grown on acid mineral soils
* The importance of cations binding in the AFS for uptake and shoot
transport is shown by experiments with same plant species but with different
binding form of a divalent cation as Zn ++ when it is supplied in form of
inorganic salt, the Zn content not only of the roots but also of the shoots is
several times higher than when Zn is supplied in a chelate (ZnETDA).
Because of the permeation of chelated zinc is restricted by pores of the AFS.
* With heavy- metal cations, binding in apoplasm can be quite specific. Cu
can be bound in form of inorganic nitrogen – containing groups of either
glucoproteins or proteins of enzymes as phosphate peroxydases in the cell
wall. This cationic binding in the apoplasm can contribute significantly to
the total of the cationic content of roots
* The root apoplasm may also serve as storage pool for heavy metals as iron
zinc, which can be mobilized. The size of this storage pool for Fe probably
plays as role for genotypical differences in sensitivity to iron deficiency in
Soybean
2.2.2. Passive transport into cytoplasm and vacuole

* Despite some selectivity for cationic binding in the cell wall, the main sites
of selectivity in the uptake of cations and anion as well as solutes in general
are located in the plasma membrane of individual cells (outer layer of
membrane)
* Plasma membrane is an effective barrier against the diffusion of solutes
either from apoplasm into the cytoplasm (influx) or from cytoplasm into
apoplasm and the external solution (efflux). It is also the main site of active
transport in both directions.

8
* The other main barrier to diffusion is the tonoplast (membrane of vacuole).
In the most mature plant cells, the vacuole comprises more than 80-90% of
the cell volume acting as central storage space for ions and other solutes
(sugars, secondary metabolites). It can be demonstrated that the plasma
membrane and tonoplast function as effective barrier to diffusion and
exchange ions Fig 2.3. Most of the Ca++ taken up within 30minutes (influx)
is still exchange (efflux) and is almost located in the AFS. In contrast only a
small fraction of the K+ is exchangeable within 30 minutes period, most of
K+ having been transported across the membranes into cytoplasm and
vacuoles
* Solutes transport into micro-organs as mitochondria and chloroplast must
be also regulated by their membranes that differ with cytoplasmic
membranes. The capability of bio-membranes for solutes transport and their
regulation is closely related to their chemical composition and molecular
structure that mentioned in plant physiology subjects.
2.2.3. Solutes transport across membrane
a. Transport and energy demand

* Membranes are sites of selectivity and transport against the concentration

gradient of solution
* The concentration of K+ in maize roots (vacuole) is higher than that 80
times in the external solution. In contrast to Na+ concentration in roots is
lower than that in the external solution. It is generally agreed that such
selectivity and accumulation require both energy supply as a driving force
and specific binding sites. Carriers or permeases in the membranes mostly
likely proteins as the sulfate-binding protein isolated from microorganisms.

9
 * A direct coupling between selective carriers with transported ion is
demanded consumption of energy phosphates in the form of ATP in this
processes, ATP was supplied from respiration in mitochondria (oxidative
phosphorylation):
E + A + ATP --------------- EA + ADP + Pi
ADP + Pi -------------------- ATP
Table 2.2.3 Changes in the ion concentration of the external solution and in the
vacuoles of maize and soybean

External concentration (mM)

Concentration in the vacuole root (mM)

After 4 days

Ion Innitial Maize Bean Maize Bean

Potassium 2.00 0.14 0.67 160 84
Calcium 1.00 0.94 0.59 3 10
Sodium 0.32 0.59 0.58 0.6 6
Phosphate 0.25 0.06 0.09 6 12
Nitrate 2.00 0.13 0.07 38 35
Sulfate 0.67 0.61 0.82 14 6
Energy is also required for activation of carriers, in the binding to the carrier,
the membrane transport of the carrier-ion-complex or release of the ion from
the carrier at the internal surface of membrane. This method was presented
by Fisher et. al (1970). Studying the uptake of K+ by roots of various plant
species was demonstrated a close relationship between K + uptake and
ATPase activity.

10
 * The energy demand for ion uptake by roots is considerable, especially
during rapid vegetation growth. Of the total respiration energy consumption
for ion uptake is required up to 36% with increase in plant age, this
proportion declines ATP demand for growth and maintenance of biomass.
* The energy demand for ion uptake by roots is considerable, especially
during rapid vegetation growth. Of the total respiration energy consumption
for ion uptake is required up to 36% with increase in plant age, this
proportion declines ATP demand for growth and maintenance of biomass.
Table .2.2.4. Respiratory energy costs for ion uptake in roots of carex
Proportion of total ATP demand required for Plant age (days)
(%)
40 60 80
Ion uptake 36 17 10

Growth 39 43 38
Maintenance of biomass 25 43 52
b. Active and passive transport: Electronic pumps, carriers and ion
channels

* Solutes transport across membranes is not necessarily an active process

solutes may be more concentrated on one side of the membrane and thus
diffuse from a higher to a lower concentration (or chemical potential). This
downhill transport across membrane is passive transport. In cells, downhill
transport of ions across the plasma membrane may be maintained by a
lowering of the ion acting in the cytoplasm. For example, Due to adsorption
at charged groups (R.COO- or R.NH3+ of protein) or to incorporation into
organic structure (P into nucleic acids and N in protein..)

11
 In contrast, membrane transport against the gradient of potential, energy
(uphill) must be linked directly or indirectly to an energy–consuming
mechanism, a Pump in membrane Fig 2.7
* To determine whether an ion is activity transported across, however,
the activity or concentration of the ion either side of the membrane (the
chemical potential gradient and the electrical potential gradient across the
membrane must be known, a micro-electrodes inserted into the vacuoles, a
strongly negative electrical potential can be measured between the vacuole
and external solution.
* The concentration at which cations and anions on two sides of a
membrane are in electrochemical equilibrium, it can be calculated according
to the Nerst equation:
Concentration inside (vacuole)
E (mv) = -58 log -------------------------------------------------
Concentration outside (external solution)
According to this equation at a negative electro-potential of-59mv,
monovalent cations as K+ or Cl- would be in electrochemical equilibrium if
their concentration in the vacuole were 10 times higher or 10 times lower
(Cl-) than in the external solution. For divalent cations or anions the
difference between chemical and electrochemical equilibrium differs by
even more than the factor of 100.
* In cells of higher plants, the electrical potential differences between
vacuole and the external solution are generally higher than -50mv. Thus as a
rule, in terms of electrophysiology only anion uptake into the vacuoles
would always require an active transport process. The only cations that
would require active transport for its uptake is K+ at low external solution

12
* In recent years, impressive progress has been made in understanding both
the mechanisms leading to the formation of electro- potential across
membranes and the importance of these potential for cell growth and
function. Progress was possible by new techniques allowing measurements
of membrane potential. Some principles of ion transport in membranes are
shown in Fig 2.8
Table.2.2.5. Experimental determination and calculation ion
concentration (mM) according to the electrical potential differences in
roots of Pea and Oat
Pea roots (-110mV) Oat roots (-84mV)

Ion Experimental Calculated Experimental Calculated

Potassium 75 74 66 27
Sodium 8 74 3 27
Calcium 2 10800 3 1400
Chloride 7 0.014 3 0.038
Nitrate 28 0.027 56 0.076
+
* An ATP-driven H pump (proton motive force; Poole, 1978) transports
H+ through the membrane from internal to external membrane, in the plasma
membrane from the cytoplasm to the apoplasm, thus creating a gradient in
pH and electro-potential. Transport of cations and anions along the gradient
is mediated either by ion selective carriers or channels Fig. as in plant
physiology. This model also takes into account another important membrane
function that is a receptor of external and internal signals and the
transformation of these signals into membrane transport processes.
* There is general agreement that in plants proton pumps are located in
both plasma membrane and tonoplast and their main function is the

13
regulation of the pH in cytoplasm. Those pumps transport H + from the
cytoplast either through the plasma into apoplasm or through the tonoplast
into vacuole leading to the typical pH differences between these
compartments. As a rule the pH of cytoplasm is 7.3 – 7.6, vacuole 4.5 – 5.9
and apoplasm about 5.5. This transport of proton in plant cells is the primary
energized process and provides the driving force for secondary energized
transport process of cations and anions along the electrochemical gradient
* The membrane transport of amino acids and sugars follows the same
principles it is driven by membrane bound proton pumps
* Membrane bound ATPase is considered as the master enzyme (Serrano,
1990). The activity of plasma membrane bound ATPase is particularly high
in root hairs where it consumes as much as 25-50% of the cell ATP, so that,
low carbohydrate levels in roots` not only decrease the movement of proton
into external medium but also decrease the pH of cytoplasm.
* Cations are transported downhill along the electrical potential gradient
(about -120 - -180mV) across the plasma membrane in to cytoplasm
mediated by carriers in membrane
For anions, transport across the plasma membrane a proton – anion co-
transport complex (or sumport) operates using electrical potential difference
and chemical (pH difference) gradient for proton as the driving force. The
evidence for co-transport at the plasma membrane of root cells has been
presented for Cl- (Ullrich and Novaky, 1990) phosphorus (Dunlop, 1999)
At tonoplast, the existence of two proton pumps is established an H +
ATPase and an inorganic pyrophosphatase, PPiase (Fig. 2.9). Both proton
pumps are phosphohydrolases using either ATP or inorganic pyrophosphate
as energy source. Mg is essential for both pumps

14
 The proton pumps at the tonoplast are necessary to maintain of a high pH
and provide the driving force for cation transport into the vacuole as antiport
More recently, ion channels in plant cell membranes has been
established, thus channels are included in carrier models of membrane
transport of ion, ion channels are unique among transport protein in their
ability to regulate ion flux, those channels permit rapid passive permeation
(uniport) of ions through membranes. Open channels catalyze the
permeation of 106 to 108 ions per second this rate is at least three of five
faster than carrier-mediated transport of ions. However, ion channels are
closed most of the time, and their number per cell seem to be rather small
There are many discussion about functions of there ion channels in plant
cells as they are important for osmosis-regulation such as guard cell of
leaves
Beside specific functions of ion channels are ion uptake by root, their
role in taking up with divalent cations is not clear
Although ion channels in membrane allow rapid passive transport of
solutes, but the dimension of there channels are not suited for permeation of
macro-molecules as proteins
2.3. Radial transport of ions and water across the roots

* There are two parallel pathway for movement of ions and water across
cortex toward xylem, one is passing through the apoplam (cell walls and
intercellular and another is passing from cell to cell in the symplasm
(Fig.2.31) page 66. As a rule the apoplam pathway of ions is constrained by
casparian band in the wall of the endodermal cells. This band as effective
barrier for radial movement of water has been questioned. At least in young

15
roots, although the hydrophobic properties of casparain band, water seems to
permeate this band readily.
* Depending on plant species and root zone, the apoplam pathway may
be constrained or blocked by suberinazation of zhizodermics. However,
there some different views on the functions of the exodermic as effective
barrier for transport of water and solutes in apoplasm of the root cortex.
* The endodermic is also not perfect barrier against the apoplamic
pathway of ion fro cortex into the xylem. However, for certain solute
apoplasm as polyvalent cations the apoplamic pathway migh be restricted in
the apical root zone
* For mineral nutrients, the symplasmic pathway plays the key role
either beginning of the zhizodermic and root hair at the exodermic or at the
endodermic. Radial transport in the symplasm from cell to cell required
bridge across the cell wall, that connect the cytoplasm of next cells
* The mechanism of symplasmic transport of solutes seems to be
mainly diffusion facilitated by radial water flux. Inhibition of cytoplasm
streaming may have no effect on the radial transport of K+. On the other
hand, in radial transport of phosphate, various metabolic steps such as
incorporation into organic compounds (ATP, sugar phosphates) are involved
before it is released as inorganic phosphate into the xylem. During radial
transport of ions, competition occurs between accumulation in the vacuole
and transport in the symplasm. This competition depends on the mineral
nutrient and its concentration in the vacuole of root cells along the pathway.
* At the low concentration (low salt root). This competition is reflected
by a typical high accumulation rate in the root and a delay in the
translocation of the ion from the roots to the shoots of plants, thus when the

16
supply of a mineral nutrient is suboptimal, the roots usually have higher
content than that in the shoots (restricted translocation).
* Radial transport of water and hydraulic conductivity and radial
transport of ions is strongly affected by maturation of xylem vessels along
the root axis
2.4. Release of ions into xylem

* After radial transport in the symplasm into xylem, most of the ions and
organic solutes are released into xylem (amino acids, organic acids), the
release of ions and organic solutes (xylem loading) is not well understood,
the key role of a proton pump needed energy (ATP) from respiration at the
plasma membrane. Protons are pumped into xylem and xylem has pH value
of 5.2 to 6.0 depending on plant species, source of nitrogen supply
* Anions may be secreted either by co-transport with the proton or along
the electrical potential gradient created by the proton pump. As the oxygen
tension is usually lower in the stele than in the cortex, xylem loading pump
is more quickly inhibited at decreasing oxygen tension in the root
environment
* Selective inhibitors of protein synthesis strongly impair xylem loading
of mineral nutrients such as K+ without affecting their accumulation in the
roots
III. Long distance in the xylem and phloem its regulation
3.1 General

*The long distance transport of water, solutes and low molecular weigh
organic compounds take place in the vascular system of xylem and floem.

17
This transport from the roots to the shoot occurs mainly in the nonliving
xylem vessels
* Xylem transport is driven by the gradient in root pressure and by the
gradient in the water potential. As a reference, the water potential of pure
free water is defined as having a water potential of zero. According for the
water potential is usually negative. The gradient in water potential between
roots and shoots is quite clear, during the day when the stomata are open.
Values become less negative in the order following atmosphere >> leaf
cells>xylem>root cells>external solution. Solutes flow in the xylem from
roots to the shoots up to the leaf cells. Under certain conditions in the shoots
counter-flow of water in the xylem may also occur
* In contrast to the xylem, long distance transport in the phloem takes place
in the living tube cells. The direction of transport is determined mainly by
the nutritional requirements of the various plant organs or tissue and occurs
from source to sink. In addition, phloem transport is an important
component in cycling of mineral nutrition between shoots and roots. Mineral
elements are transported by bidirectional. During long distance transport,
mineral nutrients and organic solutes are transferred between the xylem and
phloem by extensive exchange processes
3. 2 Xylem transport
a. Composition of the xylem

* The composition and concentration of mineral elements and organic

solutes in the xylem depend on factors as plant species, mineral element
supply to the roots, Assimilation of mineral nutrients in the roots and
nutrient recycling. Compositions of solutes are also strongly influenced by
the degree of dilution by water and therefore are dependent on the

18
transpiration rate and the time of the day. In soybean during reproductive
stage, xylem volume flow declines and concentration of some mineral
nutrients in the sap decreases and other increases
* The form and proportion of the various nitrogen fraction in the xylem
sap depend on the form of nitrogen supply (NO 3- or NH4+ N2 fixation), main
site of NO3- reduction (root or shoot) and the proportion of recycled
nitrogen. The concentration of organic acids in the xylem depends mainly on
the cation – anion uptake ratio by roots and the form of nitrogen supply. In
the xylem of annual species high concentration of sugars may also occur for
example, in soybean, sugars may account for 15% of the total organic
carbon in the sap.
* Phytohormones are an normal compounds of xylem sap such as
cytokinines are mainly synthesized in the roots. The increase in ABA
concentration in the xylem sap of nitrogen deficient plants
b. Interaction along the pathway unloading

* Along the pathway of solute transport in the nonliving xylem vessels (in
the apoplasm), from the roots to the leaves, important interactions take place
between solutes and both cell walls of the vessels. The major interactions are
exchange adsorption of polyvalent cations in the cell walls, uptake and
c. Unloading in leave
* Most the solutes and water are transported in the xylem vessels into the
leaves, here water is transported in the major veins to sites of rapid
evaporation such as leaf margins or from vein endings mainly symplasmic
movements toward stomata.

19
 * Depending on the concentration, composition of solutes in the xylem
entering leaf and here rate of water loss by transpiration, the solute
concentration may be enriched many fold as leaf edges
* Of course, the xylem import of solutes in to leaves and the evaporation
of water do not necessarily lead to the accumulation of the solutes in the leaf
apoplasm. In fast – growing plants with only low nutrient supply, the solute
concentration in the xylem declines sharply from roots to the leave and
within leaf blade from the base to the tip
d. Effect of transpiration rate on uptake and translocation

+ The rate of water flux across the root (short-distance transport) and in the
xylem vessels (long-distance transport) is determined by the root, the
pressure and the rate of transpiration. An increase in the transpiration rate
may or may not enhance the uptake and translocation of minerals elements
in the xylem. The uptake and translocation of mineral elements depend on
following factors.
i) Plant age, in seedlings and young plants with a low leaf surface area,
effects of transpiration are not absent. Water uptake and solute
transport in the xylem to the shoots are determined mainly by root
pressure. The age and size of the plants increase. The relative
importance of mineral elements increases.
ii) Time of day: In leave, up to 90% of the total transpiration is stomata
during light period, Transpiration rates and thus potential
enhancement of uptake and translocation of mineral elements are much
higher than during the dark period.
iii) External concentration

20
Table 3.5.Effect of transpiration rate on uptake and translocation of
potassium and sodium from nutrient solution in Sugar Beet
Potassium Sodium
External
concentration (mM) Low High Low High
transpiration transpiration
Uptake rate (μmol/plant after 4h)
1K+ + 1Na+ 4.6 4.9 8.4 11.2
10K+ + 10 Na+ 10.3 3.9 12.0 19.1
Translocation rate (μmol/plant after 4h)
1K+ + 1Na+ 2.9 3.0 2.0 3.9
10K+ + 10 Na+ 6.5 7.0 3.4 8.1

iv) Type of mineral nutrients

* It is well known that an increase in the concentration of mineral elements

in the nutrient medium may fortify the effect of transpiration rate on the
uptake and translocation of mineral elements; usually translocation rate are
more responsive than uptake rate
* As a rule, transpiration enhances the uptake and transport of uncharged
molecules to greater extend than that of ions, the uptake and translocation of
M-E in the form of molecular is of great importance in the cases of Boron
(acid boric)

3.3 Phloem transport (Long-transport)

21
a. Principle of transport apoplasm

* Long-distance transport in the phloem takes place in living cells. The sieve
tubes (Fig 3.9 page 94). Rule of the transport mechanism in the phloem is
based on the principle of the osmometer.
* MÜnch suggested that solutes such as sucrose are concentrated in the
phloem of leaves and the water is sucked into the phloem, creating a positive
internal pressure, this pressure induce a mass flow in the phloem to the sites
of lower positive pressure caused by removal of solutes from the phloem.
Flow rate and direction of the flow are closely related to release at the sink
* For mineral nutrients the main sites (sources) for phloem. Loading are
located in the stems and leaves as components of either mineral nutrient
supply to growth sinks (shoot, fruits, and roots) or of recycling of nutrition.
b. Composition of the phloem sap

+ Phloem sap has a high pH (7-8) and contains high concentrations of

solutes. On average 15-20% dry matter.
Main components is usually sucrose up 90%, among other organic solutes
amino acids (up 90% of solids) are usually present in high concentration.
The glutamine CONH2-CH2-CH2-CH-COOH and asparazine CONH2-CH2-
CH-COOH NH2
NH2 may consist of 90% fraction whereas NO 3- and NH4+ concentrations are
usually very low. Organic acids, anions such as citrate, malate COOH-CH-
CH2-COOH OH
are also high content in the phloem sap. Other organic compounds are also
found on phloem sap such as vitamins, hormones, proteins and ATP.

22
+ Of mineral elements, K+ is usually present in highest concentration,
followed by phosphorus, Mg2+ sulfur (Glutathione > Methionine > Cystein
and as sulfate Cl- and Na+ may also be present at high concentration.
c. Transfer between the xylem and phloem

* In the regulation of long-distance transport, exchange of solutes between

the two systems is very important. From concentration differences have
shown Table 3.8. A transfer from phloem to xylem can occur down hill
(along concentration gradient). Through the plasma membrane of the sieve
tubes, if an adequate concentration gradients exists.
* In contrast, for most organic and inorganic solutes a transfer from xylem to
phloem is usually an uphill transport. The xylem to phloem transfer is very
importance for the mineral nutrition of plants.
* Because xylem transport is directed (Fig 3.11) mainly to the sites (organs)
of highest transpiration, which are usually not the organs for highest demand
for mineral nutrition. This transfer of organic and inorganic solutes can take
place all along pathway from root to shoot. In stems sites of intensive xylem
to phloem transfer are nodes. Such as for mineral nutrients (K+) and for Type
of mineral nutrients
* As a rule, transpiration enhances the uptake and transport of uncharged
molecules to greater extend than that of ions, the uptake and translocation of
M-E in the form of molecular is of great importance in the cases of Boron
(acid boric)

23
3.3 Phloem transport (Long-transport)
* Principle of transport apoplasm

+ Long-distance transport in the phloem takes place in living cells. The sieve
tubes (Fig 3.9 page 94). Rule of the transport mechanism in the phloem is
based on the principle of the osmo-meter.
Munch suggested that solutes such as sucrose are concentrated in the phloem
of leaves and the water is sucked into the phloem, creating a positive internal
pressure. This pressure induces a mass flow in the phloem to the sites of
lower positive pressure caused by removal of solutes from the phloem. Flow
rate and direction of the flow are closely related to release at the sink
+ For mineral nutrients the main sites (sources) for phloem. Loading are
located in the stem and leaves as components of either mineral nutrient
supply to growth sinks (shoot, fruits, and roots) or of recycling of nutrition.
+ Phloem sap has a high pH (7-8) and contains high concentrations of the
solutes. On average 15-20% dry matter Table 3.8 (96)
3.4. Remobilization of mineral nutrients

+ Import and export of mineral nutrients occur during the life of plant
organs. As a rule, ageing is associated with higher rates of export of mineral
nutrients than rates of import and thus, decrease in net content or more
precisely in amount per organ such as a leaf, called redistribution and re-
translocation.
+ Remobilization is based on a range of different physiological and
biochemical process utilization of mineral nutrient stored in vacuoles (K, P,
Mg, amino-N).The breakdown of storage proteins (in vacuole of the
mesophyll cells of legumes).

24
+ Remobilization of mineral nutrients is important during the growth and
development of plants. Seed germination; Periods of insufficient supply to
the roots during vegetative Growth; Reproductive growth, and the period
before leaf drop.
a. Seed germination

* During the germination of seeds (or storage organs such as tubers) Mineral
nutrients are remobilized within seed tissue and trans-located in the phloem
or xylem or both, to the developing roots or shoots. The seedlings will grow
for at least several days without an external supply of mineral nutrients. In
seeds many mineral nutrients (K, P, Mg, and Ca) are usually bound to phytic
acids, thus remobilization of these mineral nutrients take place.
* The remobilization of highly phloem-mobile mineral nutrients can lead to
such a rapid decline in their content in the vegetative shoot that rapid
senescence is induce
* Remobilization is highly selective for mineral nutrients
* The degree of remobilization of micronutrients and calcium in
reproductive stage is higher compared with that in vegetative stage.
- The extent of remobilization of micronutrients strongly depends on
their contents in the fall expanded leave. For example: Wheat, during
grain development Cu content in leaves drops 70%.
- The extent of remobilization of mineral nutrient is attracting attention
in connection with selection and breeding of phenotypes of high
“Nutrient efficiency”. Genotypes that grow well on soils of low
nutrient Cu availability not only may ??? Higher rate of uptake and
translocation of a particular mineral nutrient but may also show higher

25
 nutrient efficiency at cellular level. Including high rates of
remobilization from older to younger leave, seeds and storage organs.
b. Vegetative stage

+ During vegetative growth, nutrient supply to the roots is often insufficient

(In condition of low soil content) and interrupted (when there is lack or
excess of soil moisture). Remobilization of mineral nutrients from mature
leaves to areas of new growth is the key importance for the completion of
the life cycle of plants.
+ The intensive remobilization takes place; however, the remobilization
differs between mineral nutrients. This is well reflected distribution of
visible deficiency symptoms in plants. Deficient symptom occur mainly
young leaves. That reflects insufficient remobilization.
+ The extent of remobilization is also important for diagnosis of the nutrition
status of plants. Leaves and other plant organs respond to an insufficient
supply of mineral nutrient to the roots by a rapid increase in remobilization
of that nutrients
c. Reproductive stage

- Remobilization of mineral nutrient is particularly important during

reproductive growth when seeds, fruits and storage organs are formed. At
this growth stage, root activity and decreasing carbon hydrate supply to the
roots. The mineral nutrient contents of vegetative parts quite often decline
sharply during the reproductive stage. Fig 3.13 (109).
* Remobilization is highly selective for mineral nutrients
* The degree of remobilization of micronutrients and calcium in
reproductive stage is higher than that compared with in vegetative stage

26
* The extent of remobilization of micronutrients strongly depends on their
contents in the full expanded leaves. For example wheat, during grain
development, Cu content in leaves drops 70%
* Genotype that grow well on soils of low nutrient availability not only may
have higher rate of uptake and translocation of a particular mineral nutrients
but also high rate of remobilization from older to younger leaves, seeds and
storage organs.
d. Period before leaf drop

* Remobilization of mineral nutrient (except Ca, Mn) from leave to woody

part is a typical feature of leave drop species and is closely related to the
discoloration of leaves in the autumn.
* As a rule, which annual species, extent of mobilization is high for
nitrogen, potassium, phosphorous, Zn, whereas leaf contents of Ca, B, Fe,
Mn increase until leaf drop.
IV. Soil and plant tissue analysis as indicator of plant nutritional status

* Up to now, in many countries to evaluate ability of supplying an mineral

nutrients from he soil for the plants is based on both its total content and
available content in the soil , but this evaluation is not true, because soil
analysis involves chemical determination of the content of a nutrient in a soil
sample, this technique depends strongly on method of sampling soil, a major
limitation of this technique is that it reflects the levels of nutritional potential
to plant root but fails to evaluate the uptake conditions and the amounts of
nutrients actually taken up by plant whereas the rate of mineral uptake of
an element from the soil depends on many factors such as soil pH,
temperature, its concentration and content of oxygen of soil if these

27
conditions are not favor plants would not take up a mineral element even its
content in soil is high, whereas the content of an element in plant leaf
reflects exactly nutritional status of the plant. So that, soil and tissue analysis
are often important to evaluate the nutritional status of plants, it is also
useful for development of the schedule for fertilization in agriculture. It may
be used to increase or decrease certain nutrients during the various stage of
growth to enhance the yield of economic part of the crops (tube, grain). To
obtain this information, plant tissue analysis must to be done
* Adequate use of plant tissue analysis requires an understanding of the
relationship between plant growth or yield and mineral content of plant
tissue samples (Bouma, 1983). The relationship is showed in Fig

28
 - When nutrient content in a tissue sample is low, Growth or yield is
reduced.
- In this deficiency zone of the curve, an increase in the tissue
concentration of the mineral is correlated with an increase in growth
or yield. As the nutrient level in the tissue sample increases further a
point is reached at rich increases in mineral content are no longer
correlated with increases growth or yield, this region is often called
the adequate zone.
The narrow transition between deficiency zone and adequate zone reveals
the critical concentration of nutrients, which may be defined as the minimal
tissue content of nutrient that is correlated with maximal growth or yield.
The tissue nutritional content increases beyond the adequate zone, growth or
yield begins to decline due to toxicity.
- Plant tissue analysis has been applied to many different plant tissues,
organs, including leaves of different ages and positions, stems, roots,
seeds, fruits and grain. It has been generally observed that change in
nutrient content brought about by changes in the nutrient supply is
closely correlated with the nutrient status of leaves (Bouma. 1983).
Leaf analysis may therefore provide a better indicators or nutrients
deficiency.
- Plant analysis should therefore make it possible to correct for a
deficiency in a crop before it results in a nutrient disorder that could
cause a reduction in growth or yield.
- Plant analysis has been useful in the development of fertilizer
programs in sugarcane production and in the production of forage
crops in pastures, where plant nutrient content is important for animal
nutrition.

29
V. Crop yields can be improved by addition of chemical or organic
fertilizers

* Chemical fertilizers applied to the soil generally contain salts of the

macronutrients such as: N, P, and K.
* Chemical fertilizers contain only one of these three nutrients are termed
“Straight” fertilizers for example super phosphate, ammonium nitrate.
Fertilizers that contain two or more of those key mineral nutrients are called
compound or mixed fertilizer. With increase of the agricultural production,
the consumption of both macro and micro nutrients also is increase thus; the
macro and micro nutrients must be added to the soil as fertilizers.
* Of course, micro nutrients must be added to the soil to correct a preexisting

deficiency. For example, tone soil in the US are deficient in boron and
copper (Mengel and Kirkby, 1979) and can benefit from micro nutrient
supplementation. Chemical fertilizers may also be applied to the soil to
modify soil pH. The pH soil can affect the availability of various mineral
nutrients. Lime addition to the soil can raise pH of the acid soil; an addition
of sulfur element can decrease the soil pH of alkaline soil.
* Disadvantage of chemical fertilizer

In some last decades Green revolution and a revolution in organic fertilizer

were shown Pioneer poles in Agriculture production especially, crop yields
has been increase several times, and Human being overcome starvation
(Shortage of Food). But chemical fertilizers have also caused some
disadvantage in Agriculture.
- Contamination of soil and underground water because of high
concentration of NO3-.

30
 - Toxic for Agricultural products
- Decreasing biodiversity
- Deterioration of soil.
* In contrast to chemical fertilizers that contain inorganic salts, organic
fertilizers contain mineral nutrients in form of complex organic molecules.
Organic fertilizers originate from the waste and residues of plant and animal
life. They are often added to the soil mainly to improve physical structure of
soil rather than for their contribution of mineral nutrients. Organic fertilizers
such as manures can improve soil water retention during drought and
drainage in wet weather. Most of the nutrients in organic fertilizers are
bound in organic molecules. Before they can be taken up by the plant, the
mineral nutrients must be released from the organic compounds usually by
the action of soil microorganisms. Therefore, the nutrients may become
slowly available to plant over a period of weeks or months. This slow
availability is useful for the plants because low release can reduce nutrient
loss due to leaching by water moving through the soil.
* Organic farming: is form of Agriculture that relies only on organic
fertilizer and avoids the use of chemical fertilizers and pesticides. Crops
grown in this way are higher quality than those grown with chemical
fertilizers and healthy for human consumption
* Organically grown crops are often very vigorous (strongly), because of the
improvement of the soil quality (soil water tension) and supply of nutrient,
however, there is no scientific evidence shown that mineral nutrients in
chemical fertilizers and released by organic fertilizers are different quality
and more healthy for human consumption
Organic farming has many advantages.

31
 Growth of
yield C%
maximum

* Advantage of organic fertilizers

- Without contamination of soil and underground water
- For cleanly Agricultural products and sustainable Agriculture
- Maintaining biodiversity
- Enhancement of soil fertility.
Organic farming has many advantages as mentioned about it was carried out
in the last when beginning of Agricultural production, but term of organic
farming only started in 30’s of last century. By 1940, the Market for organic
products has grown at a rapid pace to reach 46 billion USD in 2007, about
32.2 million ha worldwide now are farmed with organic fertilizers. One
international organization was found called (IFOAM) International
federation of organic Agricultural movements
* Organically grown crops are often very vigorous, because of the
improvement of the soil quality (soil water tension) and supply of nutrient,
however, there is no scientific evidence shown that mineral nutrients in
chemical fertilizers and released by organic fertilizers are different quality.
VI. Mineral Nutrition and Yield Response
a. General

* Many factors are required for plant growth such as light, CO2, water and
mineral nutrients, increasing the supply of any those factors from deficiency
range, enhances rate of plant growth and crop yield. The yield response
curves of a particular mineral nutrient shown fig.6.1
* The curve of relationship between yield and supply of one M.E. also
depends on other factors and other M. N. fig.6.2
* Under field condition the interactions between water availability and
nitrogen supply are very important. In maize with increasing nitrogen supply

32
and different soil moisture levels the curve of yield obtained is similar to
curve between potassium and nitrogen. The decrease of yield can occur
when increasing nitrogen supply in combination with low soil moisture
levels because of some following factors
i. delay in stomata response to water deficiency
ii. high water consumption of vegetative biomass and the corresponding
the higher risk of drought stress in period of grain formation.
* Increase in shoot-root dry weight ratio with increasing of nitrogen supply
an effect mostly in C3 species.
* Yield curve can differ not only between vegetative and reproductive
organs but also between the yield components of harvested products. In most
crops, both quantity (dry meter yield = tons per hectare) and quality (Protein,
Sugar contents and mineral nutrients) are important yield components. Fig
6.3
- Maximum quality can be obtained both before (curve 1) or after
(curve 2), the maximum dry weight yield has been reached.
b. LAI and Net photosynthesis

* When the Nutrient supply is suboptimal the Leaf Growth Rate, (LAI. Leaf
area index) can be limited, thus rate of net photosynthesis is low, and
insufficient for leaf expansion or both. For example, with suboptimal supply
of nitrogen and phosphorous, plant suffering from nitrogen deficiency,
elongation rates of leaves may decline.
* The effect of nitrogen deficiency on leaf expansion differs between plant
species. In cereals (monocotyledons) cell expansion is inhibited to day and
night. In dicotyledonous, the inhibition is much more severe during the
daytime.

33
* The similar results for the nitrogen effect in dicotyledonous have been
obtained for phosphorus. It was found that, phosphorus deficiency severely
inhibited leaf growth only during daytime and without the night.
* Mineral nutrient deficiency can also delay plant development for example,
in barley; number of days to reach the booting stage is about twice as much
for Mn deficiency.
* Mineral nutrient can also influence the net photosynthesis in many ways,
the direct involvement of some mineral elements in the electron transport
chain (Fe) in the thylacoid membranes. Mineral nutrients are required for
protein synthesis, pigments of chloroplast as chlorophyll. For example, up to
75% of the total organic nitrogen is located in the chloroplasts mainly
protein enzymes. In spinach, about 24% the total nitrogen is contained in the
thylacoid membrane of photosynthesis organs. Thus nitrogen supple also
effects on the amount of thylacoids membrane per unit leaf area; a mineral
nutrient deficiency can also depress net photosynthesis by influencing the
CO2 fixation reaction and entry of CO 2 through the stomata, sugar synthesis,
and transport in chloroplasts is decreased causing low yield. The deficiency
of an M. Element of immobile will first become evident in younger leaves.
Although precise mechanism of the mobilization process is not understood,
Growth hormones such as cytokinines appear.
VII. Functions of M. nutrients
7.1. General functions

* Functions in structure of plant organs such as N. P.S and Mg, N are an

important part of many organic compounds and P Mg also are a part of
organic compounds
* Roles in metabolic processes of plants

34
 + M.E. effects on omosic potential of plant cells and hydratation on
cytoplasm such as K+ and Mg+2 affects on direction and volume of the
metabolic processes
+ Enhancing action of metabolic enzymes Zn Mo increase nitrate and
nitrite reductases
+ A part of phytohormones as N in NAA and Cytokinine that regulate
plant growth
* M. E. fortify resistant of diseases and environmental stress. B, Mo, Zn
increase the water absorption in dry soil.
* Inadequate supply of an essential M. E. results in a nutritional disorder and
deficient symptoms. In hydroponic culture, an appeared symptom can be
observed by the removal of one essential element. In soil, plants grown,
diagnosis can be more complex because some plant diseases (by virus or
bacteria have the similar symptoms to the deficient symptoms of M. E
* With moveable M. E such as N, P, P, Mg, they can move from leaf to leaf,
when deficiency, these elements move from old to young leaves thus
deficient symptoms occur in older leaves fist. In contrast, the immoveable
M. E as Ca, Fe B when deficiency, symptoms appear in young leaves fist
7.2. Nitrogen
7.2.1. Assimilation of nitrogen

* NO3- and NH4+ are the major of inorganic nitrogen taken up by the roots of
higher plants. Most of NH+4 have to be incorporated into organic compounds
in the roots, whereas nitrate is mobile in the xylem and can also be stored in
the vacuoles of root and storage organs. This accumulation in vacuoles can
be considerable importance for anion and cationic balance, for osmotic
regulation and for quality of vegetable and forage plants. However, in order

35
to be incorporated into organic compounds and to fulfill its essential
functions as a plant nutrient, NO3- has to be reduced to NH4+. Nitrate
reduction is important for nitrogen nutrition of plants
* The high content of nitrate in the plant cells is nontoxic for plants but toxic
for human being and animals
a. Nitrate reduction and assimilation in the plant

* Nitrate is main source of available nitrogen to higher plants

* In most temperate soils ammonia is rapid converted to nitrate by
nitrification by bacteria
* In order to incorporate in to organic compounds and to fulfill its function
in plant metabolism, nitrate must be taken up by cell and then it is reduced
into ammonia
4 NADH2 or NADPH2
* NO3- + 8H+ + 8e- ------------------ NH3 + H2O ------ NH4+ + OH-
This reaction takes place by two steps and is mediated by two enzymes
* The fist, nitrate is reduced to nitrite, this reaction occurs in cytosol and is
mediated by nitrate reductase involving the 2 electron reduction of nitrate to
nitrite, the electron donor of this enzyme in higher plants is coemzyme of
NADH or NADPH, Mo is bound to this enzyme, in condition of Mo
deficiency, the activity of nitrate reductase on cell is much reduced.
NADH2 or NADPH2
NO3- + 2H+ + 2e- ---------------------- NO2- + H2O
* The second step is reduction nitrite to ammonia, in the cytosol, nitrate is
reduced to nitrite in high concentration can be toxic for cytosol, thus nitrite
is rapid transported into plastids-chloroplasts in green leaves. In chloroplasts

36
contain a ferredoxin – nitrite reductase, this enzyme catalyzes the six
electron reduction of nitrite to ammonia:
NO2- + 6H+ + 6e- ----------------- NH4+ + H2O
Ferredoxin (Fd) is the electron donor in photosynthetic tissues as in
following reaction:
NO2- + 6Fd red + 6H+ + 6e- ------------NH4+ + 6 Fd ox + 2H2O
This enzyme is located in chloroplasts in leaves or cytoplasts of roots
* In the C4 plants both nitrate and nitrite reductases are located in the
mesophyll cells, those cells utilize light energy for nitrate reduction and
assimilation. Bundle cells for CO2 reduction, it is probably lead to use of
efficiency nitrogen in C4 more than in C3 plants (Fig 2.1 Dissertation 13)
b. Ammonium assimilation

* Whereas nitrate is stored in vacuoles without toxic for plant cells,

ammonia and ammonium are toxic at quite low concentration. They do not
accumulate in plant cells and are rapidly incorporated into organic
compounds, the formation of amino acids, amides with both ammonium
taken up from the soil by roots and ammonia derived from the nitrate
reduction or N2 fixation
* Nearly all of the ammonium taken up has to be assimilated in the roots and
assimilated nitrogen is transported in the xylem as amino acids and amides
to the shoots. In some plant species, small proportion of the ammonium may
be transported to and assimilated in the shoots
* Ammonium assimilation in roots has large requirement for carbon
skeletons, these carbon skeletons are provided by tri-carboxylic acid cycle
(Crebs cycle)

37
* The assimilation of ammonia-nitrogen into carbon compounds can take
place through 2 possible pathways
* The first pathway is the reductive amination of α ketoglutarate into
glutamate :
α ketoglutarate + NADH 2 + NH3 ----------glutamate + NAD+ + H 2O,
this reaction is catalyzed by glutamate dehydrogenase in mitochondria.
Aspartate and alanin can be produced by this pathway
* The second pathway, in order to reduce carbon skeletons costs for
transport from roots to shoots, the bulk nitrogen is assimilated on roots and
is transported to shoots in form of nitrogen-rich compounds with C/N ratio >
o.4 by following reaction:
Glutamate (Aspartate) + NH3 + ATP→ Glutamine (asparazine) + ADP + P i
(H3PO4). Then glutamine or asparazine can be converted to glutamate or
aspartate by reaction:
Glutamine + α ketoglutarate → 2 Glutamates
* Ammonia can be further incorporated into other amino acids by
transmination reactions, these reaction are carried out by enzymes known as
aminotrasferases as following reaction:
Glutamate + pyruvate (oxoloacetate) → Alanin (aspartate) + α ketoglutarate
* Although plants can contain up to 200 different amino acids but only about
20 of them are required for protein synthesis, these pathway produces amino
acids to satisfy the demand for protein synthesis in the plants
* M. Elements, especially nitrogen have an important role in protein
synthesis. Ammonium and nitrate as source of nitrogen supply is better for
plant growth

38
* Ammonium and nitrate comprise about 80% of the total cations and anions
taken up by plants form of nitrogen supply has inverse impact on the uptake
of other cations and anions because of interaction between ions
* Ammonium or nitrate is suitable for the plants it is dependent on many
factors such as plant species, soil pH, growing stage of plants.
* Urea can be taken up directly by roots or aerial parts, after taking up by
roots it is rapidly hydrolyzed by urease in roots or after translocation to the
shoots

7.2.2: Important of N for plants

a. Nitrogen supply, plant growth and plant composition.
* Nitrogen is the most limited factor for crops yield in the tropics as well as
in the temperate regions, because most cultivated soils are deficient in this
element
* In the plants, nitrogen is a key element in many of important compounds
that build plant cells and organs such as proteins, nucleic acids, amino acids,
nucleotides, chlorophyll and phytohormones
* In green leaves, nitrogen is present in chlorophyll, this pigment converted
sunlight energy into chemical energy in form of organic compounds to
create crop yields. In this process, if nitrogen absorption of crop has trouble
that threatens all metabolic operations in the plants, restrains growth and
crop yields, especially, low quality of agricultural products
* Depending on plant species, development stages, and organs, the nitrogen
content required for optimal growth is between 2-5% of the plant dry weight.
Nitrogen supply is low, plant growth is limited or restrained, and deficient
symptom appears such as chlorosis in older leaves. In severe cases these
leaves become completely yellow and then die by stopping synthesis of

39
chlorophyll. Young leaves remain green longer because they receive
nitrogen form of soluble from older leaves
* Under low nitrogen supply, the total of leaf surface area is low lead to
reduce net photosynthesis. An increase of nitrogen supply not only delays
chlorosis but also stimulates plant growth:
* Plants grown with excess nitrogen usually have dark green leaves and big
canopy, potato grown in this condition, shoot growth is more rapid but the
underground tubes are small, excess nitrogen also causes to split fruits in
tomato, flowering and formation of seeds of several crops are retarded
* In cereals, in condition of excess nitrogen, enhancement of the stem
elongation leads to falling down causing low crop yields
Table.7.1. Nitrogen supply as NH4NO3 on development of rice leaves
Nitrogen supply Leaf blade (cm)
(mg/liter)
Length Width Area Thickness
5 49 0.89 30.6 4.9
20 65.1 1.13 47.8 4.1
200 63.3 1.25 56.1 3.8

* Nitrogen changes plant composition much more than any other mineral
elements. The dry matter production of plants is increased by nitrogen
application, the total nitrogen content increases and content of other
compounds is changed. Table following:
Table.7.2. Effect of nitrogen supply as NH 4NO3 on D.M. production and
compositions of Ryegrass
Organic matter Nitrogen supply (g per pot)
0.5 1.5 2.0
Dry matter (g/pot) 14.9 26.5 26

40
Total nitrogen (% DW) 2.0 3.6 4.2
Sucrose (% in DW) 7.7 7.1 6.3
Polyfructosans (% DW) 10.0 1.8 1.1
Starch (% in DW) 6.1 2.1 1.0
Cellulose (% in DW) 14.4 13.9 17.6
DW: dry weight DM: dry matter
* Increase of nitrogen application enhances protein synthesis and formation
of chloroplast results in increase of lipid, chlorophyll, carotene content in
leaves. An increase in the protein, B vitamins as riboflavin, thiamine and
nicotinic acid content of vegetative or grain of cereal is also correlated to
nitrogen supply.
* Thus more and low nitrogen supply are not useful for plant growth and
yields the question is what is suitable rate for nitrogen applied in one crop, a
suitable dose of applied nitrogen for crop species is important not only
reduction of cost in Agriculture but also increase of crop yield and quality
b. Nitrogen in A as a limiting factor of crop yield.

* Soils are more commonly deficient in nitrogen than any other elements,
one calculation shown that with soils on the world, the supplement of
nitrogen from the soil is range from 30 to 100kg in form of inorganic
nitrogen per year, this rate is very low compared to demand of nitrogen for
intensive crop production for example in Viet Nam…….
* The low nitrogen content in the soil is caused by some following factors:
Nitrogen removal by crops, the amount of nitrogen plants taken up depends
on strongly crop species and yields for example with rice, in local variety in
grain yield of 2.8 tones per hectare, nitrogen removal is 82kg and with new
high yielding rice in grain yield of 8.0 tones per hectare, nitrogen removal is

41
152kg per hectare. As a rule, nitrogen requirement is increased in new plant
varieties with high yield
* Amount of the nitrogen removal also depends on organs of crops, Table
following

Table.7.3 Amount of nitrogen removal by plants

Crop Yield Nitrogen removal
(tone/ha) (kg/ha)
Barley (grain) 2.2 40
Barley (straw) 2.5 17
Wheat (grain) 2.7 56
Wheat (straw) 3.8 22
Oat (grain) 2.9 55
Maize (grain) 9.5 150
Maize (straw) 11.0 110
Sugarcane 75.0 110
Potato (tubers) 27.0 90
Tomato (fruits) 50.0 130
Soybean (seeds) 3.4 210
Tea (dry leaves) 1.3 60
Bananas (fruits) 45.0 78
c. Loss of nitrogen

42
* Loss of nitrogen in form of inorganic fertilizers applied or form of
mineralization processes from the soil by
i) Denitrification, this process takes place when inorganic fertilizers applied
in high temperate and dry soil conditions
ii) Leaching down of nitrate into the depth soil layer amount of nitrogen
leached varies considerable between soils and it depends on the climate, type
of soil and quantity of nitrogen present in the soil, leaching of nitrogen and
other nutrients is very strongly in humid temperate, tropical areas and severe
on light textured soils table following
iii) Removal nitrogen by plants
Table.7.4. Rate of leaching nitrogen from the soils
Soil type Clay content (%) Nitrogen leaching
(kg/ha/year)
Sandy soil <3 12-52
Sandy loam 16 0-27
Loam 28 9-44
Clay 39 4-44
Leaching nitrogen is mainly in form of NO3-
* Rate of nitrogen leaching also depends on crop species, low density of
crops such as maize leaching losses are higher than for high density of crops
as grassland
* A considerable loss of NH3 may occur when NH4 salts are applied to
calcareous soils. High pH soil should not be treated with NH4 salts and urea
* Recent investigations have shown that plant itself may also release
gaseous nitrogen into atmosphere mainly in form of NH3, with soybean
planted under field conditions (high temperature and transpiration) this crop
releases nitrogen in form of NH3 up to about 45kg/ha/year (fig2.4 25)

43
* In total of nitrogen applied to the soil, in which of 25% changed to soil
organic matters, 40-50% taken up by plants, 25% lost from the soil
(denitrification 20% and 5% leached).
7.2.3. Nitrogen application for plants

* Nitrogen application for crop is compulsory to get a desired yield use of

effective fertilizers in increasing of agricultural production is fully
recognized. Clear relationship has been established between increasing of
yield cereals and nitrogen supply. An input of 1kg of chemical nitrogen
produced more than 10kg of additional cereal grain
* Agricultural production in the world over some last decades also showed
that in order to increase crop yield to meet demand food for human beings
during the world population increased, nitrogen supply plays the most
important role, according FAO (Food Agriculture Organization), nitrogen
consumption was increased from 72.5 million tones in 1993/94 to 82.9
million tones in 1997 and production of cereal grain reached the record of
1909 million tons. From last years of 20th century up to now, nitrogen
consumption on the world has not been increased considerable about nearly
90 million tones
* Now, nitrogen and other chemical fertilizers consumption on the world
have two trends:
i) Strong reduction of chemical nitrogen consumption has been taken place
in developed countries as in Europe and North America
ii) In contrast, consumption of chemical nitrogen is still increased in
developing countries as in Asia, South America, and Africa, in this regions
whereas the rates of the population are still high and in order to satisfy the
great demand food they have to carry out intensive cultivation with input of

44
nitrogen fertilizers but farmers face the environmental pollution. In Viet
Nam, from 1980/ 1981 to 2009/2010 use of nitrogen fertilizers increased 10
times (35000 tons to 3.5 million tons and total of foodstuffs increased from
11.5 to about 38 million tons
* The crops are inefficient in their uptake and use of chemical nitrogen
fertilizers with only 50% of the total nitrogen applied taken up by non-
legume crops and 30-40% by rice. This practical becomes big problem for
rice production in Viet Nam, the low efficiency of nitrogen take up causes
many difficulty in my country as mentioned about
* Nitrogen doses of application are very different among crops the demand
of nitrogen to produce one unit of agriculture product is change, so that
depending on kind of crops, desired yields C4 is more than that C3 and
particular condition
* Doses of nitrogen are also depended on content of soil available nitrogen
and external factors, thus an efficient dose of nitrogen application depends
on many factors and it must be determined by experiment before
recommending for every crop on one certain soil
* Soil pH has much more effects on efficiency of applied nitrogen creating
optimum pH for soils plays important for nitrogen uptake by plants. Soil pH
of lower 4.5- 5.0 can be very damaging for most plants, it causes deficiency
with nitrogen and other nutrients such as P, Mg and toxic for AL, Fe. Mn.
pH range of 5.5 – 6.5 is satisfactory for most plants. The optimum pH for
high yield depends on soil type and rotation but general pH of the neutral to
slightly alkaline are useful for nitrogen uptake and for uptake other nutrients.
In this range of pH are also good for activity of effective micro-organisms
* All of liming materials may be used to increase pH in acid soils, lime or
dolomite containing Mg have double benefit

45
* Time of nitrogen application for one crop is dependent on growth and
development stages. In cereals, most nitrogen demand is the young stage of
growth for example rice and maize. In fruit trees, the more demand of
nitrogen is needed after harvesting as citrus trees, lichen plants, late nitrogen
application can cause expansion stem elongation lead to fall dawn and
reduce cereal yield
* Determination of time nitrogen application can reduce loss nitrogen by
leaching and de-nitrification
* Method of applying nitrogen for crop depends on form of nitrogen, kind of
crops and growth stages but general, most nitrogen is applied to soil at
young stages a small part of nitrogen application about 10% of the total (Phu
Nguyen Van., 2002) can be supplied by spraying on the leaves called foliar
application.
7.3. Sulfur
7.3.1. General

* Although atmospheric SO2 is taken up and utilized by aerial part of higher

plants, but the most important source of sulfur is sulfate taken up by the root
in form of SO42- at relative low rates and long transport of sulfate occurs
mainly in the xylem.
* Similar to nitrate reduction of sulfate is necessary for the incorporation of
sulfur into amino acids, proteins and coenzymes. In green leaves, ferredoxin
is a reductant for sulfate.
* Unlike nitrate, sulfate can also be assimilated without reduction and
incorporated into essential organic structures such as sulfolipids in
membrane or polysaccharides.
7.3.2. Sulfate assimilation and reduction

46
* First step of sulfur assimilation is the activation of sulfate ion by ATP. In
this reaction the enzyme ATP sulfurylase catalyzes the replacement of two
phosphate groups of the ATP by the sulfuryl group, which leads to the
formation of Adenosine phophosulfate APS and pyrophosphate.

* For the sulfate reduction, the activated sulfate of APS (Adenosine

phosphor sulfate) is transferase of APS or sulphotransferase to thiol (R-SH)
group of a suitable carrier. This transfer of sulfate to thiol group mediated by
sulphotransferases, is associated with a reduction of sulfate (SO4=) to sulfite
(SO3=) then reduction of carrier-bound-sulfite to sulfide (S-2) may involve
sulfite reductases in chloroplasts, reducted feredoxin is electron donor.
* In higher plant, the enzymes of sulfate reduction are located in the
chloroplasts. In C4 plants the bundle sheath, chloroplasts are main sites of
sulfate assimilation whereas the mesophyll chloroplasts are the sites of
nitrate assimilation.
* Sulfate reduction is several times higher in green leaves than in roots and
in leaves the reaction is strongly stimulated by light. Sulfate reduction in the

47
leaves leads to export in the phloem of reduced sulfur compounds such as
Glutathione (GSH)

7.3.3. Metabolic functions of sulfur

* Sulfur is a part of amino acids such as Cystein, Cystin, Methionine and

hence of proteins

Cysteine

Cystin

* Sulfur is a structural part of compounds such as coenzymes and secondary

plant products or acts as a functional group (R-SH) directly involved in
metabolic reactions. Under normal condition, the tripeptide glutathione
accounts for more than 90% of this fraction.
* Synthesis of glutathione occurs in two steps

48
 - First step: Glutamylcystein is produced from glutamate and
cysteine.
- Second step: Glycine is coupled to glutamylcysteine, mediated by
glutathione synthetase, an enzyme required magnesium for
activity.
Fig.7.3

+
Glutamate Cysteine

Glutamylcysteine

Glutathione -

Synthetase + Mg2+

GSH (Glutathione)

* In the plants, GSH content in leaves is usually higher than that in roots,
and in leaves, more than 50% of it is located in the chloroplast.
* GSH has role of anti-oxydation
* GSH may also function as storage pool of reduced sulfur.

49
* Reduced sulfur is a structural constituent of some coenzymes and
prosthetic groups such as Feredoxin, biotin (vitamin H), thiamine
pyrophosphate (B1). In many enzymes and coenzymes such as Urease,
Sulphotransferase and coenzyme A, the SH groups act as functional group
for synthesis of fatty acids.

Krebs
cycle

* The most important sulfur containing compounds of the secondary

metabolism are glycosinolates. They contain sulfur both on a sulphydryl and
sulfo group, and storages in vacuoles.
* The role of secondary sulfur compounds is not fully understood. They act
as defense substants.
* Sulfur in form of non-reduction as sulfate-ester is a component of
sulfolipids and is thus structural part of all biological membrane.
7.3.4. Sulfur supply. Plant growth and plant composition

* Sulfur requirement for optimal growth varies between 0.1- 0.2% of the dry
weight of the plants. The requirement increases in order Gramineae <
Leguminosae < Cruciferae. On average, proteins from legumes contain less
sulfur than protein from cereals, the N/S ratio being 40/1 and 30/1.
* With nitrogen deficiency also condition under of sulfur deficiency, shoot
growth is more depressed than root growth, interruption of the sulfur supply

50
within a few days, decreases net photosynthesis. Leaf area is reduced in
sulfur deficient plants.
* Chlorophyll content of leaves drastic decreases and inhibits protein
synthesis in sulfur deficient plants.
* In sulfur-deficient-plants, inhibition of protein synthesis is related to an
accumulation of solute organic nitrogen and nitrate, amides.
* Inhibition of protein synthesis during sulfur deficiency leads to chlorosis.
Similar to it does during nitrogen deficiency. The distribution of sulfur in
sulfur deficient plants is also affected by the nitrogen supply. Symptoms of
sulfur deficiency may occur either in young or in old leaves. In legumes,
during the early stages of sulfur deficiency, nitrogenase activity in root
nodules is much more depressed than photosynthesis.
* In sulfur-deficient-plants not only the protein content decreases but also
the sulfur content in the proteins.
* The lower sulfur content of proteins influences nutritional quality
considerable. Methionine (non-substitution amino acids) is an essential
amino acid in human nutrition and often of limiting factor on diets in which
seeds are a major source of protein. Further more, a decrease in the cysteine
content of cereal grains reduces the Baking quality of flour.
* In highly industrialized areas, the sulfur requirement of plants is often met
fully by atmospheric SO2 pollution. When industrial SO2 emissions have
been decreased, deficiency is becoming more widespread in agricultural
areas. For example, in Northern Europe in decade of 90 th, sulfur deficiency
in crop production is common in rural area, especially in high rainfall areas
such as in humid tropics, in temperate climates and in highly leached soils,
under these condition the application of nitrogen fertilizers in form of urea is
ineffective unless sulfur is applied simultaneously.

51
In Viet Nam

7.4. Phosphorus
7.4.1. General

* Unlike Nitrate and sulfate, phosphate is not reduced in plants but it

remains in highest oxidized form. After uptake mainly as H2PO4- it remains

as inorganic phosphate (Pi) or a hydroxyl group to a carbon chain

as a simple phosphate ester (sugar phosphate)
For example: Glucose phosphate, fructose phosphate

Fructose
6-phosphate
And attached to another phosphate by the energy rich pyrophosphate bound
ADP, ATP ( ~ )

ATP

52
* Rates of exchange between and are very high, for example, taken
up by roots is incorporated within a few minutes into organic but there
after is released again as into xylem.
7.4.2. Functions of phosphorus in the plants
a. Phosphorus as a structural element

* The function of phosphorus as a part of macro-molecules as Nucleic

acids, units of DNA, RNA molecules. DNA are carries of genetic
information, and units of RNA, it is translation of genetic information. In
both DNA and RNA, phosphate form a bridge between nucleotide units to
form acid nucleic

53
Structure of Nucleic acid (DNA, RNA)
* The bridging form in phospholipids of bio-membranes and bio-membranes
relate to selective diffusion of plant cells

b. Role in energy transfer

* Although present in cells only in low concentrations, Phosphate esters (C-

) and energy rich phosphates ( ~ ) represent the metabolic
machinery of the cells up to 50 individual esters formed phosphate and sugar
alcohols have been identified as Glucozo-6-phosphate.
+ The energy required for biosynthesis of starch or for ion uptake is supplied
by an energy rich compounds. Mainly ATP and ADP; the release of energy
is following role:

This structure of ATP is not exactly

54
 AMP

ADP

ATP

ATP   ADP  H 3 PO4  6500KCal / mol

ATPase

ADP   AMP  H 3 PO4  750Kcal / mol

ADPase

* Energy liberated during respiration or photosynthesis is used for synthesis

of the energy rich pyrophosphate bound, the phosphorylation of reaction is
following:
ADP + H3PO4 ATP (as in photosynthesis), as well as in oxidation of
organic matters in respiration, this reaction is mediated by protein kinases
* PEP carboxylase is one of the key enzymes in CO 2 fixation of C4 plants
regulated by phosphorylation in light reactions, and phosphorylation
increases activity of PEP carboxylase.
c. The regulatory role of inorganic phosphate and compairmentation

* Phosphate Pi (inorganic) controls some key enzyme reactions. Thus, it is

essential for the regulation of metabolism in the cytoplasm and chloroplast.

55
The activity of the phospho-fructose kinase is increased by Pi. Fruit ripening
of tomato plants can be delayed in phosphorus deficient condition.
* The vacuole of higher plants acts as storage pool or “non metabolic pool”
of , in condition of adequate supply of phosphorus 85-95% of the total Pi
is located in the vacuoles. In contrast, in leaves of phosphorus-deficient
plants all of is found in the cytoplasm and chloroplast in “metabolic
pool” under phosphorus deficiency in leaves, Pi concentration in the
cytoplasm may drop from 5mM to less than 0.2mM and the levels of energy-
rich-phosphate drop 20-30%.
* In leaves, photosynthesis is strongly affected by the Pi concentration in the
stroma of the chloroplasts. In the available light, a Pi concentration for
optimal photosynthesis in chloroplasts is range of 2.0 to 2.5 M. Due to the
large demand of Pi for phosphorylated intermediates of photosynthesis in
leaves of phosphorus-deficient-plants, the Pi concentration may drop to 50%
dark-light transition.
* An increase in external Pi concentrations up to 1mM stimulated net
photosynthesis but decreased incorporation of the fixed carbon into starch,
and starch synthesis inhibited in about 5mM of Pi concentration (two
mechanism page 270)
* Under phosphorus deficiency shoot growth is much more depressed than
photosynthesis.
d. Phosphorus fractions and the role of phytate

Major phosphorus fractions in the vegetative plant organ also increase.

Mainly in leaves, with more increase in supply only as mainly storage
form of phosphorus in highly vacuolated tissue increase. However, both

56
higher and lower plants may also store phosphorus in two forms such as
polyphosphate and phytate those compounds are important for stage of seed
germination.
* The storage of phosphate in inorganic polyphosphates is common among
lower plants (bacteria, fungi…) it has been also found in higher plants.
Polyphosphates may function as energy storage compounds and controlling
level in the metabolic pool of the cells. They also function as cation
exchangers.
* Phytate is mainly storage form of phosphorus in grains and seeds. Phytate
is salt of phytic acid, Myoinositol
Myoinositol + 6 H3PO4 → Phitic acid (formulation)
* The soluble Calcium-Magnesium salt of phytic acid is termed phytic acid
also has Zn, Fe. In legume seeds and cereal grains the main phytate are
potassium-magnesium salt. The proportion of K+, Mg2+ and also Ca2+
associated with phytic acid may vary considerably between plant species.
* Phytate in form of K-Mg-Ca salt is major form of phosphorus in pollen
grains.
* In legume plants and cereals, during the early stages of seed and grain
development the content of phytate is low but rises quickly during the period
of rapid starch synthesis. In contrast, the content of Pi during early stages of
seed and grain development is general low and further declines during rapid
phytate formation. When phosphorus supply to the roots increases after
anthesis stage, phytate is only phosphorus fraction increases in grains.
* Some phosphorus is associated with the starch fraction and is incorporated
into the starch grains
* The function of phytate in seed germination is quite important in the early
stage of seedling growth; the embryo has a long requirement for mineral

57
nutrients such as “Magnesium for phosphorylation and protein synthesis, K +
for cell expansion and phosphorus for incorporation in membrane lipids and
nucleic acid”
* Phytates have attracted considerably attention by nutritionists. These
compounds interfere absorption of mineral elements, especially Zn, Fe and
calcium, thus they cause nutritional deficiencies in both monogastric animals
and human, especially children. This problem can be improved (solved) by
an increase the zinc/phytate ratio in seeds and grains through the foliar
application
7.4.3. Phosphorus requirement, Growth of plant and plant composition

* The phosphorus requirement for optimal growth is in range 0.3-0.5% of

the plant dry matter during vegetative growth stage and probability
phosphorus toxicity is in content of higher than 1% in dry matter. However,
many tropical legumes are more sensitive and toxicity may occur at
phosphorus content in shoot dry matter of 0.3-0.4% (in Cajanus cajan) and
0.6-0.7% (in Vigna mungo).
* Deficiency of phosphorus, leaf expansion, leaf surface area and number of
leaves (Lynch et al, 1991) is reduced. Inhibition of leaf expansion, the
contents of protein and chlorophyll per unit leaf area are not much affected,
even the chlorophyll content is increased under phosphorus deficiency, and
leaves have a darker green color as well as leaf expansion are more retarded
than chloroplast and chlorophyll formation but photosynthesis efficiency per
unit of chlorophyll is much lower in phosphorus deficiency leaves (Lauer et
al, 1989).
* Under condition of phosphorus limitation, not only shoot growth rate is
retarded but also the formation of reproductive organs. Initiation of flower is

58
delayed; the number of flowers decreased and seed formation restricted in
specially; senescence of leaves is another factor limiting seed yield in
phosphorus deficiency plants.
With information discussed about P is especially needed to legume plant
species, high carbohydrates plant species sugarcane, Citrus cultivation
Table 8.1. Effect of phosphorus deficiency on some Growth parameters and
contents of phosphorus and hydrate carbon in Soybean.
Treatments
Parameters
High P Low P
1. Leaf area (dm2) 12.1 1.8
2. Number of primary trifoliate 7 4
3. Shoot/Root dry weight ratio 4.2 3.02
4. Chlorophyll (mg/dm2) 3.02 2.08
5. P content leaves (Pi) (mg/g dry weight) (P
4.43 0.28
organic)
6. P content (total P mg/g dry weight stem and
5.84 1.14
petioles)
7. P content (total P mg/g dry weight roots) 10.54 1.29
8. Ratio total root P/total shoot P 0.54 1.57
Starch 0.4 12.8
9. Carbohydrates of leaves (g/dm2 leaf)
Sucrose 0.7 0.2
Starch 23 160
10. Carbohydrates of roots (mg/g fresh weight)
Sucrose 16 177

7.5 Magnesium
7.5.1. General

59
* Mg is small divalent cation with a hydrated ionic radius of 0.42nm a very
high hydration energy of 1908 J/mol, it’s rate of uptake can be strongly
depressed by other cations such as K+, NH4+, Ca2+ and Mn2+ as well as H+ in
low pH. Thus magnesium deficiency enhanced by competing cations is
fairly widespread phenomenon.
* Functions of magnesium in plants are mainly related to its capacity to
interact with strongly nucleophilic ligands (phosphoryl group) through ionic
bonding and to act as a bridging element, the most bounds involving
magnesium are ionic and some covalent such as in the chlorophyll
molecules. A sustainable proportion of the total magnesium is necessary for
regulation of cellular pH and the cation-anion balance.
7.5.2. Binding form and compartment

* In Green leaves a major function of magnesium, is its role as the central

atom of chlorophyll molecules as mentioned
in plant physiology. Fig 7.1
* The proportion of the total magnesium
bound to chlorophyll depends very strongly
on the Mg supply, from 6% in leaves of
plants with high to 35% in leaves of plants
with magnesium deficiency. Under low light
condition, this proportion might even be higher than 50%. Depending on
mineral nutritional status, this proportion is common from 6% to 25%,
another 5-10% of the total Mg in leave is bound to pectate in the cell wall.
The remaining 60-90% is extractable with water. In most instances, plant
growth is depressed and visual symptoms of Mg deficiency occur when
proportion of Mg in chlorophyll exceeds 20-25%.

60
* Similar to inorganic phosphorus (Pi), the concentration of Mg is not bound
in organic structures but located in the metabolic pool, in cytoplasm and
chloroplast is range of 2-10mM.
* In the “metabolic pool” the Mg distribution between the cytosol and
chloroplast has to be well regulated. In isolated chloroplast photosynthesis is
strongly inhibited even by 5mM in the external solution. Inhibition of
photosynthesis by high Mg concentration in the “metabolic pool” may occur
in plants under drought stress.
7.5.3. Functions of Mg in the plants

a. Role of Mg for Chlorophyll and protein synthesis

* Insertion of Mg into the porphyry structure as the first step of chlorophyll
biosynthesis is catalyzed by Mg-chelatese, chlorophyll break down also
requires two enzymes as Mg-dechelatase and Chlorophyllase.
* Mg has also an essential function as a bridging element between 2
ribosome subunits (50s and 30s ribosome subunits) to create ribosome units
(70s); this unit is an important part of protein-biosynthesis organ. When the
level of free Mg (Mg2+) is deficient or high level of K+, those subunits are
dissociated, protein bio-synthesis is retarded, Mg also is required for RNAr
polymerase thus hence for the formation of RNA molecules.
* Net synthesis of RNA immediately stops in condition of Mg deficiency
and synthesis continues rapidly after the addition of Mg. In contrast, Protein
synthesis remains unaffected for more than 5 hours but after that it rapidly
declines, a concentration of at least 0.25-0.4mM Mg 2+ is necessary at
cytoplast to maintain protein synthesis.
b. Roles of Mg in enzyme activation of phosphorylation and photosynthesis.

61
* There are many enzymes and enzyme reactions required and are strongly
promoted by Mg. Glutathione synthetase, PEP carboxylase, the enzymes
need presence of Mg (as phosphatases, ATPases) for transfer phosphate
group or carboxyl group, for example with ATP.
* The complex of Mg-ATP is formed to stability above pH of 6 and can be
utilized for transfer of the energy-rich-phosphoryl group; ATPase and Mg-
ATPase are bound to plasma membrane in root.
* The synthesis of ATP (phosphorylation ADP + H 3PO4 ATP) has
requirement for Mg as a bridging component between ADP and the enzyme,
more Mg supply can stimulate for the photophosphorylation, the addition of
Ca2+ severely inhibits this reaction.
* Another key reaction of Mg is the modulation of RuBP carboxylase in
stroma of chloroplasts. The activity of this enzyme depends on both Mg and
pH; Mg also shifts the pH optimum of the reactions towards physiological
range.
* One of the key enzymes with a high Mg requirement and pH optimum is
Fructose 1-6 biphosphatase in chloroplast. It regulates between starch
synthesis and export of trios phosphate.
* Activity of glutamine synthetase also is enhanced by Mg. This enzyme
regulates ammonia assimilation within chloroplast
* The accumulation of non-structural carbohydrate also occurs in leaves of
Mg-deficient plants, and causes increase of dry matter content of these
leaves, indicating more depression of the starch degradation than
photosynthesis. The accumulation of carbohydrates in leaves of Mg-
deficient-plants is correlated with a decrease in carbohydrate content at sink
sites (storage organs) such as pod, root, grain…
7.5.4. Mg supply plant growth and composition

62
* The requirement for optimal plant growth
is in the range 0.15-0.35% of dry weight of
vegetative parts, the chlorosis of the full
expanded leaves is most visible symptom of
Mg-deficiency, the leaf area unit,
chlorophyll unit as well as the proportion of
protein nitrogen and the rate of
photosynthesis are low, proportion of non
protein nitrogen increase in condition of Mg
deficiency. Appearance of slight of Mg-
deficient-symptom is not affect to reduce the final yield.
* In the some last decades, the Mg-deficient-symptoms of plants are
widespread in forest ecosystems in central Europe and associated with other
stress factors are pollution soil, acidification.
* When Mg is deficient the export of carbohydrates from source (leaves)
sink sites depressed (tuber, pod, grain), effect on economic yield of crops
* In some cases, the high Mg contents of plants improve the nutritional
quality of plants for example Grass Tenany low Mg-content makes serious
disorder of ruminants. An increase of Mg-content is ease to achieve by foliar
application of Mg fertilizers.
With information discussed about, Mg is especially needed to annual crops
as rice, maize, soybean vegetable crops, their time total of growth and
development is short
7. 6. Calcium
7.6.1. General

63
*Ca is a divalent cation with hydrated ionic radius of 0.4nm, in the
appoplasm, a part of Ca is bound in structures another part is exchangeable
at the cell walls and external surface of the plasma membrane. A high
proportion of Ca might be required in vacuoles whereas its concentration in
the cytosol is extremely low.
* The most function of Ca is related to structure of macro-molecules as
calcium peptates that links from cell to cell
* Ca is most presence in cell walls and plasma membranes, with high level
supply, Ca content can reach more than 10% DW of plant especially in
soybean.
* In recent years a new function of Ca has been discovered this is its
function as a second messenger in the signal conduction between
environmental factors and plant responses in growth and development.
7.6.2. Binding form and compartment

* Unlike to other macronutrients, a high proportion of the total Ca is in the

plant tissue is located in cell walls, this distribution shown that an abundance
of binding sites for Ca in the cell walls as well as of the restricted transport
of Ca in to cytoplasm, between lamella it is bound to R-COO - group of
pectins . In dicotyledons such as sugar beet, soybean when levels of Ca
supply is low up to 50% of the total Ca can be bound as pectates in storage
tissue of apple fruits the cell wall bound fraction of Ca can make up as much
as 90% the total
* Increase of Ca supply, in many species, the proportion of calcium oxalate
increases
* High Ca concentration is found in the middle lamella of the cell walls, at
the external plasma membrane and vacuoles. In contrast, Ca concentration in

64
cytoplasm is very low only in the range of 0.1- 0.2Mm of free Mg 2+ because
of prevention of precipitation of Pi, competition with Mg2+ for binding sites
* The low Ca free concentration in cytoplasm is caused by low permeability
of membranes for calcium, the different in Ca concentration between
vacuole and cytoplasm is up to 105 times
7.6.3. Functions of Ca in the plants
a. Stabilization of the cell wall:

* The peptate in the middle lamella is essential for strengthening of the cell
walls and plant tissues this role of Ca is clearly reflected in the close positive
correlation between CEC of the cell wall and Ca content in plant tissues
required for optimal growth
* The leaves of plant receive high levels of Ca during growth or grown
under conditions of high light intensity, the high proportion of Calcium
pectate in the cell walls is very importance for the resistance of the plant
tissues to fungal and bacterial infections, for the ripening of fruits.
b. Function in membrane stabilization

 The main function of Ca in Membrane and cell wall stability is

reflected by many ways such as increase of leakage of low-molecular-
weight solutes from the cells of Ca deficient tissues.
 Ca stabilizes cell membranes by bridging phosphate and carboxyl
group of phospholipids, there can be an exchange between Ca and
other cations (K+, Na+, H+) in these binding sites, the exchange of
plasma membrane-bound calcium, for example by high external
concentration of Na is main factor involved in salinity stress

65
  To fulfill its function at the plasma membrane, calcium must always
be present in the external solution, whereas it regulates the selectivity
of ion uptake and prevent soluble leakage from the cytoplasm
 In calcium deficient tissues, the impairment of membrane integrity
lead to increased respiration rates which are related to enhanced
leakage of respiratory substrates from vacuoles in to cytoplasts
 Ca stimulates a number of membrane-bound enzymes, particularly
ATPases at plasma membrane of roots of certain plant species
c. Function in cation – anion balance and osmo-regulation

* In vacuolated of leaves in particular, a large proportion of calcium is

located in the vacuoles, where it might contribute to the catio-anion balance
* The formation of soluble calcium oxalate is also important for the osmo-
regulation of cells and provides a means of salt accumulation in vacuoles of
nitrate feed plants without increasing the osmotic pressure in the vacuoles
7.7. Potassium
7.7.1. General

* Potassium is taken up in form of K+ and is monovalent cation

* Uptake of K+ is highly selective process
* In the soil, most potassium is also in form of K +, so that it is very ease to
leach down or runoff by water. Thus under cultivated soils, potassium
deficiency is most common for most plants. In Viet Nam fertility soils of
potassium is outside band of the red river delta or first cultivated soil
* Potassium is high mobility in plant at all levels within individual cell and
long distant transport in the xylem and phloem
* Potassium is most abundant cation in the cytoplast of leaf cells

66
* Due to its high concentration in the cytoplast and chloroplast, it neutralizes
the soluble and insoluble macromolecular anion and stabilizes pH between
7-8 this is optimal pH for most enzyme reactions
* In most cases, cytoplast and chloroplast potassium, concentrations are
maintained in the range of 100-200mM, in these compartments called
metabolic pool in contrast, K concentration in vacuole may vary 10-
200mM even reach 500mM in the guard cell of the leaves
* For the fast transport of K + between different compartments and cells
within tissues. K+ channels are required in membrane, this channels is open
or close in response to environmental signals
7.7.2. Functions of potassium in the plants
a. Enhancement of enzyme activation

* Many enzymes are both completely dependent on potassium and

stimulated by K+
* All enzyme proteins are highly hydrated and stabilized by bounding with
water molecules (hydratation protein) and only this form, enzyme proteins
are fulfill their functions in metabolic processes
* Potassium activates starch synthetase in the range of 50-100mM K +, the
higher concentration may have inhibitory effect
* Another function of K+ is activation of membrane-bound proton pumping
ATPase, this function not only facilitates K+ uptake, transport but also is
important for mineral nutrients and osmo-regulation
* The dominant role of K+ is maintenance of high cytoplamic pH
b. Function in protein biosynthesis

67
  K+ is required for Protein synthesis in higher concentration than for
enzyme activation, its concentration is about 50mM, K+ is related to
several steps of Protein synthesis, including the binding of RNA t to
ribosomes
 In green leaves, chloroplasts account for about haft of leaf RNA and
leaf protein. C3 plants main of chloroplast protein is
RuBPcacboxylase, thus synthesis of this enzyme is impaired under K +
deficiency and responds rapidly to re-supply of K+, in K+ deficiency
plants, the soluble nitrogen compounds (amino acid, amide and
nitrate) even number of polypeptedes are increased in contrast, protein
content is reduced
c. Function in Photosynthesis

 In higher plants, potassium effects on photosynthesis at different

levels, K+ is necessary for the establishment of the transmembrane pH
gradient needed for the synthesis of ATP in photosynthesis (Photo-
phosphorylation)
 The role of K+ in CO2 fixation, an increase of K+ concentration to
100mM in the cytoplasts, CO2 fixation increases three fold
 For maintenance of high pH in stroma, low supply of potassium, K +
concentration reduces causing photosynthesis decrease, this decrease
can be overcome by high concentration in chloroplast, thus under low
supply K+, corresponding lower leaf content, the rate of
photosynthesis is lower, an increase of potassium concentration in
leaves increases both rate of photosynthesis, RuBPcarboxylase

68
 activity as well as photorespiration, probably due to a stronger
depletion at the catalytic sites of the enzyme
 An increase of K+ content causes decrease of dark respiration and
higher respiration rates are main feature of K + deficiency. K+
nutritional status may also affect photosynthesis in leaves by its
function in stomata regulation
d. Function in cell expansion

Cell expansion involves the formation of a large central vacuole

occupying 80-90% of the cell volume there are two major requirements
for cell expansion: an increase in cell wall extensibility and solute
accumulation to create an internal osmotic potential in young cell for
growing
 The enhancement of stem elongation by GA is also dependent on K +
supply, the highest elongation rate is obtained by both GA and K+
applied
 K+ in association with inorganic anions or organic acid anion is the
main solute required in the vacuoles for cell extension, the extent to
sugars and other low molecular weight organic solutes contribute to
the osmotic potential, this effect is dependent on potassium nutritional
status
e. Regulation of stomata movement

 In most plants, in the guard cells, during stomata movement an

increase in the K+ concentration in the guard cells increases their
osmotic potential and results in the uptake of water from the next cells

69
 leading to increase turgor in the guard cells and thus stomata opening
during the day time, the accumulation of K+ in the guard cells during
the day time is driven by proton pump ATPase bound on plasma
membrane, activation of this pump deceases pH guard cell apoplast,
K+ accumulation in the vacuole has to be balanced by counter-anion
mainly CL- or malate acid depending on plant species
 Closing of the stomata is caused by darkness or ABA and associated
with rapid efflux of K+ and anion from the guard cells, close of
stomata by ABA derive from roots is response of the plants to stress
conditions of environments as drought soils
g. Function in cation-anion balance
h. Enhancement of resistance of the disease and falling down

In cereal crops, K+ enhances photosynthesis of hydrate carbon especially,

the cellulose molecules fortifying solidarity of cell wall and lead to increase
of resistance for fugal disease such as pyricularia and heminthosporium
ozyrea, most common diseases in rice and for falling down
7.7.3. Potassium supply for plants

* After nitrogen, potassium is the mineral nutrient required in largest amount

by plants. The potassium requirement for optimal plant growth is range 2-
5% in DW of vegetative parts, when K+ is deficiency, plant growth is
retarded and rate of re-translocation of K+ is enhanced from mature leaves
and stems and under severe deficiency, these organs become chlorosis and
necrosis depending on light intensity to which leaves are exposed
* When soil water is limited, loss of turgor and wilting are typical symptoms
of K+ deficiency. K+ sufficiency condition, plants are lower response to

70
drought stress by: (a) role in stomata regulation, which controls water
regime of higher plants, (b) maintaining the high tissue water content even
under drought conditions and enhancement of resistance with low
temperature.
* By function in synthesis, transport of starch and sugar, K+ is necessary to
increase in cereal crop yields such as maize, potato, rice as well as
sugarcane, sugar beet
* Plants receiving an inadequate supply of potassium are often more
sensitive to frost or low temperature damage by reducing movement of
metabolic matters in the plant cells
Table 7.7 Relationship between potassium supply and tube yield and
damaged by frost in potato by Grewel and Singh (1998)
Potassium supply Tube yield Potassium content of leaves Percentage of leaf
(Kg/ha) (tons/ha) (mg/g in DW) damaged by frost
0 2.39 24.4 30
42 2.72 27.6 16
84 2.87 30.0 7

In Viet Nam, all cultivated soils are needed to supply potassium by intensive
cultivation lead to rapidly decrease K +, especially, lowland rice, maize, tube
crops and potassium is useful in low temperature condition and slope soils.
Two types of the potassium fertilizers are used KCL or K 2SO4 or complex
NPK and most of them are applied at the beginning stage of development of
crops such as formation of seeds, fruits and tubers
VIII. Functions of the micronutrients in the plants
8.1. Iron
a. Iron in the soil and forms of iron uptake by plants

71
* Iron makes up 5% by weight of the Earths crust and sustainable present in
all the soils. Fe is main present in ferromagnesian silicates such as olivine,
augite and biotite,
* In the physiological pH range of the soils, the concentrations of ionic Fe 3+
and Fe2+ are below 10-15M. The content of the soluble Fe in soils is extremely
low compared to the total content
* The soluble inorganic forms include Fe3+, Fe(OH)2 and Fe2+ . In well
arable soils, Fe2+ contributes very little to the total soluble inorganic Fe
except under soil high pH soil conditions
* Chelate of Fe (III) are the dominant form of soluble iron in the soils and
nutrient solutions, as an role Fe(II) is taken up preferentially compared with
Fe(III)
b. Importance of iron for plants

* The most important of iron is present in hence proteins and the well known
are the cytochromes, they are an important part of the electronic transport
chains of oxidative phosphoryllation and photo phosphoryllation reactions
* Other hence enzymes are catalases and peroxidases. Under conditions
of iron deficiency, the activity of both enzymes declines, in non-hence
proteins, iron is coordinated to the thiol group of cysteine of inorganic sulfur
as claster. The most well known one is feredoxin, which acts an electron
transmitter in many basic metabolic processes in the plants.
* Low chlorophyll content (chlorosis) of young leaves is the most visible
symptom of iron deficiency.

72
c. Iron deficiency, toxicity and application for plants

* As a rule, iron deficiency has much less effect on leaf growth, cell number
per unit area or number of chloroplasts per cell than the size of the
chloroplasts and protein content per chloroplasts. Only with severe iron
deficiency, the cell division is inhibited, thus leaf growth reduced. However,
under condition of iron deficiency, protein synthesis in chloroplasts is much
more impaired than in the cytoplasm
* In general, C4 species require a high iron supply than C3 plants, but their
critical deficiency contents are similar, about 72 mg/kg Fe in C 3 plants and
66 mg/kg Fe in C4 plants. In legumes, iron demand for nodule development
is particularly high
* Iron deficiency is a worldwide problem in crop production on calcareous
soils. Iron deficiency might also limit CO2 fixation
* On the other hand, iron toxicity is a serious problem in crop production on
waterlogged soils. It is the second most severe yield-limiting factors in
wetland rice as in Asia, especially in waterlogged rice of Viet Nam the
critical toxic content is about 500mg/kg Fe in leaf dry weight but very much
dependent on other factors as the content of other mineral elements
* Iron toxicity may also take place under dry soil lead to enhancement of
drought damage in photosynthetic tissue caused by iron catalyzed formation
of oxygen free in the chloroplasts
* So that ironic supply for crops to improve growth of crops is related to
both of iron supply on iron deficient soils and reduction of iron toxicity on
waterlogged soils. Iron toxicity in waterlogged soils is dominant problem in
wetland rice in most countries of Asia as in Viet Nam

73
8.2. Manganese
a. Mn in the soil and forms of Mn uptake by plants

* Mn is present in various rocks, especially in ferromanganesian materials,

the total Mn levels may differ considerably between soils. Mn content range
of 200 and 3000ppm is the most common
* The most important Mn in the soil fractions are Mn +2 and Mn oxides, in
which Mn is present in divalent or tetravalent form and the most important
fraction in plant nutrition is Mn+2 . In addition, reduced Mn contributes to
plant supply.
b. Importance of manganese for plants

* Mn acts as cofactor, activating about 35 different enzymes (Burnell, 1988).

Most of them catalyze oxidative and reductive reactions, decarboxylation,
hydrolytic reactions in respiration (Crebs cycle)
* Mn deficiency similar to Mg deficiency, in both cases, interveinal
chlorosis occurs but contrast to Mg deficiency, Mn symptoms are first
visible in the young leaves, whereas Mg deficient symptoms are older leaves
first
* In condition of Mn deficiency, the Mn content of the leaves and both the
seed yield and oil content are reduced. But the seed protein content is
increased in this condition
* Lower lignin contents in Mn deficient plants are a reflection of the
requirement for Mn in various steps of lignin biosynthesis
c. Mn deficiency, toxicity and application for plants

74
* Mn deficiency is abundant in plants growing in soils derived from parent
materials of low Mn content and in highly leached tropical soils as well as in
soils of high pH containing free carbonate
* Mn deficiency can be corrected by soil or foliar application of MnSO4
* The high Mn content in seeds by both spray MnSO 4 to parent plants and
by soaking seeds in MnSO4, this solution can considerably improve plant
growth and seed yield on Mn deficient soils
* Among plant species and cultivars differ considerably in sensitively to Mn
deficiency when grown on low Mn soils. Oat, wheat, soybean are sensitive
whereas maize and rye are not sensitive. But the critical deficient content of
Mn in plants is similar, between 10 and 20mg/kgMn in DW of full expanded
leaves. Below the critical deficiency, the content of dry matter, net
photosynthesis and chlorophyll content decline rapidly, whereas rates of
respiration and transpiration remain unaffected. The grain number and yield
are declined in Mn-deficient plants causing by a combination of both low
pollen fertility and shortage in carbohydrate supply for grain filling
* The environmental factors affect critical toxic content (CTC) of Mn, at
high temperatures the CTC of Mn in leaves is often higher than that at low
temperature and Mn toxic can be fortified by high supply of Mg (Kirkby and
Mengel, 1987)
* In many experiments have shown that, Mn toxic symptoms occur as
interveinal chlorosis similar to iron in young leaves
* Rate of 30kg per ha as MnSO 4 is applied in deficient soil of Mn by foliar
application or soaking seeds

75
8.3. Copper
a. Cu in the soil and forms of Cu uptake by plants

* Cu is present in the soil almost in divalent form and this form is available
for plant nutrition. The largest fraction of Cu is present in the crystal lattices
of primary and secondary minerals. In addition, Cu occurs in organic
compounds
* The Cu levels of total in the soils are range of 5-50ppm (McLaren. 1989).
The Cu concentration of the soil solution is very low in the range of 10-8 to
60.10-8 M
* Increasing of pH dues to stronger Cu adsorption causing a decrease of
level of Cu in the soil solution
b. Importance of Cu for plants

* The most important function of Cu is in redox reactions, in this reaction,

oxydase of Cu enzymes reacts directly with molecular oxygen
* Cu has a high affinity for peptide and sulphydryl group, and thus, to
cystein-rich proteins as well as for carboxylic and phenolic groups
* In the soil solution, in roots and in the xylem, more than 98-99% of the Cu
is present in complex form
* According to Sandman and Boger (1993) three different forms of protein
exist, in which Cu is the metal component (Cu-protein). Under Cu
deficiency the activity of these Cu enzymes decreases rapidly, but not all,
these decreases are correlated to metabolic change and inhibition of plant
growth
* In general, more than 50% of the Cu total located in chloroplasts is bound
to plastocyanin. This Cu protein contains on Cu atom per molecule.

76
Plastocyanin (PC) is a component of the electron transport chain of
photosystem I
c. Cu deficiency, toxicity and application for plants

* Cu deficiency is well known in many crops. In cereal crops, the deficiency

appears first leaf tips at the tillering stage. The tips are white and the leaves
become narrow. Under Cu deficiency there is close relationship between Cu
content of leaves and plastocyanin content and thus activity of PSI
(photosynthesis system I) is reduced whereas Chlorophyll content is slightly
affected. Compared to PSI, activity of PSII is less depressed by Cu
deficiency
* In the chloroplast of Cu deficiency, the inhibition of electron transport is
further concentrated by lack of two polypeptides in the chloroplast
membrane. Which are necessary to maintain suitable membrane fluidity to
ensure the mobility of plastoquinone (PQ) molecules to transport electrons
between two photosystems
* Cu deficiency can be corrected by foliar application with CuSO 4 solution
or soaking seeds in this solution
* Contrast to iron, toxic iron takes place most on the waterlogged soils Cu
toxicity is scare in most crops
8.4. Zn
a. Zn in the soil and forms of Zn uptake by plants

* In the soil Zn is present in the range of 10-300ppm in many different

minerals the level of Zn in the soils is very much related to the parent
materials

77
* The 60% of the soluble Zn in the soil occurs in soluble Zn organic
complex
* Zinc is taken up mainly in divalent cation (Zn2+) at high pH and is long
distance transported in the xylem
b. Importance of Zn for plants

* Zn plays both catalytic function and structural role in enzyme reactions

* In last decade the role of Zn in biosynthesis of protein molecules as in
DNA replication and in the regulation of gene expression has been attracted
interest by researchers. The changes in metabolism caused by Zn deficiency
are quite complex
* There is a large of number of enzymes in which Zn is a part component of
the enzyme structure, in these enzymes Zn has two functions: catalytic and
structural.
* In enzymes with Zn catalytic function Zn atom is coordinated to four
ligands, three of them are amino acids with histidine being the most
frequent, followed by Glu-NH2 Asp-NH2 and a water molecule is the fourth
ligand
* In enzymes with Zn structural function such as alchohol dehydrogenase,
and the proteins involved in DNA replication and gene expression, Zn atoms
are coordinated to the S-group of four cysteine residues forming a structure
of high stability
* In higher plants, under aerobic conditions ethanol formation takes place
mainly in root apices. In Zn deficient plants, alcohol dehydrogenase activity
decreases but the consequences for plant metabolism are not affected
* In lowland rice flooding stimulates activity of root alchohol
dehydrogenase twice more in Zn Sufficient plants compared to Zn-deficient

78
plants, this enzyme contains a single Zn atom, which catalyzes the hydration
of CO2 and converses CO2 to HCO3-, Zn deficiency may have negative effect
on the rate of Photosynthesis in C4 compared with C3 plants
c. Zn deficiency, toxicity and application for plants

* Zn deficiency is widespread among plants grown in highly weathered acid

soils, and in calcareous soils, Zn deficiency is often associated with iron
deficiency, translocations of Zn to the shoot are inhibited by high
concentration of bicarbonate HCO3-
* The most visible symptoms of Zn deficiency are reduced growth due to
shortening of internodes and a drastic decrease of leaf size. Under severe Zn
deficiency the die shoots are widely occurred, Symptom of chlorosis and in
order leaves of Zn deficient plants. Shoot growth is usually more inhibited
than root growth.
* In the leaves, the critical deficient levels are below of 15- 20mg/g Zn in
DW, grain and seed yields are depressed and the total dry matter production
by Zn deficiency. Maize, cotton and apple are much more sensitive than
wheat, oat, or pea
* When the Zn supply is large, Zn toxicity can be induced in non-tolerant
plant, Zn toxicity leads to chlorosis in young leaves. The critical toxic levels
in leaves of crop are as low as 100mg/g DW to more than 300mg/g DW.
Increasing soil pH by liming is the most effective process decreasing both
Zn content and Zn toxicity. Zn deficiency in plants grown in calcareous soil
can be corrected by application to the soil of inorganic Zn salt such as
ZnSO4 (Horst, 1993)

79
8.5. Molybdenum
a. Mo in the soil and form of uptake by plants

* The total Mo content of most agriculture soils lies between 0.6-3.5ppm

and average of 2.0ppm, the available content is about 0.2ppm, the fraction of
Mo in the soil solution as well as the soil Mo may be very different between
soils.
* The maximum Mo absorption is obtained at pH4 in form of molybdate by
plants, Mo uptake can be reduced by competitive effects of SO 4 2- , Mo may
also associated with Fe oxide minerals in adsorption
b. The importance of Mo for plants

 The requirement of plants for Mo is lower than that for any other
nutrients. Functions of Mo as plant nutrient are related to the valence
changes and as a metal component of enzymes
 In higher plants only a few enzymes have been found to contain Mo
as a cofactor. In these enzymes Mo has both structural and
catalytically functions and is directly involved in redox reactions.
They are nitrate reductase, nitrogenase, xanthine oxydasse-
degydrogenase and Mo requirement strongly depends on the form of
nitrogen supply. When Mo is limiting, accumulation in root nodules
may lead to low Mo content in the shoots and seeds of legume plants
c. Mo deficiency, toxicity and application for plants

* In soil with low Mo availability, the effect of Mo application to legumes

depends on the form of nitrogen supply, some experiments had shown that
Mo application increased the Nitrogen content and seed yield only in the

80
plants without or insufficient supply of nitrogen, they indicate that on soils
with low Mo availability it is possible to replace the nitrogen application to
legume by application of Mo fertilizer combined with bacteria rhizobial
infection.
* nitrate reductase activity (NRA) is low in leaves of Mo deficient plants, in
nitrate fed plants there is close relation between Mo supply, NRA of leaves
and yield of spinach
* Mo requirement for plant growth is strongly depends on nitrogen supplied
as nitrate or ammonium as mentioned about function of Mo in nitrate
reduction, the ammonium fed plants are low responses to Mo. Mo-deficient
plants seem to be more sensitive to low temperature stress and waterlogged
soils
* Mo deficiency also has striking effects on pollen formation in maize,
grains is small and poor germination
* Depending on plant species and source of nitrogen supply, the critical
deficient levels of Mo vary between 0.1 and 0.1μg/g leaf DW. In Mo
deficient plants, chlorosis in young leaves are common as well local
chlorosis and necrosis along the main veins of mature leaves
* Mo deficiency is widespread in legumes and other plant species in acid
mineral soils with large content of reactive iron oxide hydrate
* Large-seeded cultivars application of high Mo availability during the seed-
filling stage is very effective in seed production in low available Mo soils
* Mo is highly phloem- mobile, foliar application is to correct Mo
deficiency. In legumes Mo applied as spray in early growth stages, it is
translocated in to the nodules and very effective in increasing final yield
* The toxicity levels of Mo is 0.1 1000 1μg/g Mo in DW

81
* Under conditions of Mo toxicity, leaves are yellow and discoloration of
shoot tissues occur. But non-toxic levels in plants are advantageous for seed
production, whereas such as levels of toxic Mo are dangerous for animals
and for ruminants in particular. Mo content range of 5-10 mg/kg in DW of
leaves is high enough to cause toxicity called molybdenosis. This occurs in
Western part of U.S, in Australia and in New Zealand. Molybdenosis is
caused by imbalance of Mo and Cu in the ruminant diet. The strong effect of
sulfate on molybdate uptake can be used effectively in depressing the Mo
content in the plants to levels that are nontoxic for both plants and animals.
8.6. Boron
a. B in the soil and form of uptake by plants

* The B content of the soils is range of 20-200 ppm, most B soil is

unavailable for plants. The B available fraction is from 0.4-5ppm. B is
present in some minerals. Soluble B in the soil consist mainly of boric acid
B(OH)3 and it is also for plants
* B uptake is close related to the pH and the external B concentration over a
wide concentration range
* B distribution in the plants is controlled by transpiration stream
b. The importance of B for plants

* The role of B in plants nutrition is still less understood compared with all
other Mineral Elements
* What is known of B requirement mainly come form studies of what
happens when without or re-supplied after deficiency

82
* Leaf metabolism and composition can be affected by B deficiency, its
effect on cytokine synthesis in the root tip. When no supply of B both
production and export of cytokinie in to the shoots reduce
c. B deficiency, toxicity and application for plants

* B deficiency id widespread leading to nutritional disorder in the plants.

Under high rainfall condition and high temperature areas as in Viet Nam B is
strongly leached from the soils as B(OH)4-- . B available also decreases in
under drought conditions
* Plant species differ in their capacity to take up B when grown in the same
soil which reflects differences in the requirement of B for growth. The
critical deficient levels increases from 5-10mg to 80-100mg/kg in DW. For
evaluation of B critical levels, the elongation rate of young leaves is a much
more suitable parameter than shoot dry weight
* Symptoms of B deficiency in the shoots are noticeable at the buds or
young leaves, which become discolored and may die, interveinal chlorosis
on manure leaves may appear. Drop of buds, flowers and young fruits is also
main symptom of B deficiency, especially in citrus crops
* B deficiency reduces even failures of seed and fruit set are well known.
However, even when seed yield is not depressed in plants grown in a low B
soil, the production of seed or fruits might have a low quality.
* Application of B both to soil and leaves can be done to improve flower and
fruit set in fruit trees or in soybean. Amount of B applied varies from 0.3 to
3kg/ha depending on requirement and sensitive of the crops to B toxicity.
For example in Viet Nam
* B toxicity is most common in arid and semiarid regions that plants
growing on soil coming from parent materials of marine origin or related to

83
the use of irrigation water high in B. The critical toxic contents in leaves are
in range of 100 with corn, 400 with cucumber, 1000 with squash and 100-
270mg/kg in DW wheat species
* Typical toxic symptom on mature leaves is marginal or tip chlorosis or
both and necrosis. Visual symptom of B toxicity on leaves might appear at
much lower B content than required for depression of grain yield as in wheat
8.7. Titanium
a. Titanium in the soils

* Ti is trace element it is present in the soils and in the plants. Ti is common

mineral rocks and is range in concentration of 0.03 to 1.4%, in minerals, Ti
occurs mainly in the tetravalent oxidation state such as a major component
of oxides, titanates or silicates
* The dioxide of titanium is present in nature in different modification rutile,
brookite, antase, and leucoxene
* The Ti content depends on soil type, the sandy soils, organic light soil are
usually low in Ti contents and clay, clay loamy soils or forest soils have a
high level of Ti content. The Ti content of surface soils generally ranges
from 0.1 to 0.9% except for organic soils in Soviet Union. In soils with low
available titanium, Ti supply with small amount by spraying can satisfy the
demand of Ti for plants (Aline et al., 1986 and Phu Nguyen Van., 2002)
* Ti minerals are known to be most stable minerals in the soil environment,
and the Ti level reported in soil solutions is very low in 0.03mg/l, the
solubility of Ti in the soils is very limited

84
b. Titanium in the plants

* Ti is trace element that is found in many plant parts. The present in the
plants showed that it has some roles in metabolic processes of the plants, but
some last decades no clear evidence of the biochemical function of Ti had
appeared. In the plants, Ti content is range of 15-1500ppm in DW depending
on plant species, organs.
* Recent years, studies of Chapman (1982), Pais et al. (1996) and Phu
Nguyen Van (2002) had showed some roles of Ti in plant metabolisms
* Ti has catalytic function in N 2 fixation by symbiotic microorganisms in
legume species and in photo oxidation of nitrogen compounds by higher
plants as well as in some processes of photosynthesis, an increase of
chlorophyll content in tomato plants grown in culture solution after spraying
with Ti chelate solution
* The results of studies carried out by Phu Nguyen Van (2002) had showed
that Ti spray in form of chelate combining with Mg increased leaf
chlorophyll content, protein total content in biomass of wheat barley and
spinach
* Ti also enhances iron and zinc uptake and reduces phosphorus uptake also
obtained in some experiments of Phu Nguyen Van (2001)
* Research of Yaghoubi et al (2000) showed that effects of Ti4+ added at
level of 55μg daily to animal diets, after 2 weeks body weight of animals
increased 15% than that of the control, this finding have showed that Ti 4+
enhances the animal growth. The function of Ti antibiotic has been
discovering by some studies these findings have been paying attention by
many scientists

85
* The titanium content of the plant biomass is sharply increased by spraying
Ti citrate in concentration of 20-50ppm (Phu Nguyen Van 2002)
PART B. MINERAL NUTRITION, ROOT SYSTEMS AND SOIL
RELATIONS
IX. Role of the plant root system in taking up mineral nutrition and its
interaction with the soil

* The ability of plants to absorb both water and mineral nutrients from the
soil is related to their capacity to develop an extensive root system. There
was detailed study of the winter rye by H. J. Ditmer in 1937 showed: when
root system of a rye plant that had grown for 16 weeks was examined, it was
found to consist of 13.106 root axis and lateral roots. The surface area of
roots and root hairs contribute about 67% of the total surface area
* The volume of the soil in contact with the roots is also important for
the uptake of mineral nutrients. In other studies of wheat by J.E. Weaver in
1926, it was found that in the same environments the roots would grow to
depths of 100 – 200 cm and extend to distances of 30 to 60 cm. Maximum
depth of about 50m have been reported for prosopis trees . The total lenght
of the root systems of trees planted 0.5m apart has been estimated to reach
12 – 18 km, of winter rye had been estimated 15.000km. These features are
useful for taking up water and mineral nutrients from the soils, but to
express beneficial features of root systems depending strongly on many
technical in agriculture applied as water showering water, fertilization,
oxygen supply productive soils and so on.

86
X. Some mineral nutrients enter the roots at the apical region, other
can be taken up the along entire root surface

* The point of entry of minerals into the root system has been a topic of
considerable interest. Some researchers have claimed that nutrient whereas
others believe that absorption takes place over the entire root surface.
Experimental evidence has supported both possibilities, depending on the
nutrient being is investigated, in barley the absorption of potassium,
phosphorus and ammonium can take place freely at all location of the root
surface (Clarkson and Hanson, 1980). In contrast, the uptake of the calcium
in this crop is restricted to the apical region. Iron uptake may carry out either
at the apical region as in barley or over the entire root surface as in maize.
* In the soil, nutrient movement to the root surface can occur by both bulk
flow and diffusion. Bulk flow occurs when nutrients are carried in the
convective flow of water moving through the soil toward the root. The
amount of nutrient provided to the root by bulk flow is dependent on the rate
of water flow through the plant and the amount of nutrient in the soil
solution.
* Under conditions of high rates of water flow and high concentration of
mineral nutrients in soil solution, bulk flow can play an important role in
nutrient supply.
* Measurements of the mineral content of the root tissue and of the soil
solution in the chamber provide information about mineral movement in the
soil and uptake by root.
* Diffusion takes place when mineral nutrients move from a region of
higher concentration to a region of lower concentration. Nutrient uptake by
roots lowers the concentration of nutrients at the root surface, creating

87
concentration gradients in the soil solution surrounding the root. Diffusion
of nutrients depressed their concentration gradient can increase nutrient
availability at the root surface.
* When nutrient uptake by root is high and the nutrient concentration in
the soil is low, bulk flow can supply only a small fraction of the total
nutrient requirement (Mengel and Kirkby, 1979). Under these conditions,
movement of most nutrients to the root surface is limited by diffusion. When
diffusion is too slow to maintain high nutrient concentration near the root, a
nutrient depletion zone adjacent to the root surface is formed. Optimal
nutrient depends on both the capacity for nutrient uptake and on the growth
characteristics of the root system
XI. Soil and mineral Nutrients

The solid phase, liquid phase and gases phase, all of the phases are related to
the supply of the soil surrounding plant roots can be considered as a
heterogenous (complex) material containing the solid nutrients to the root
surface. The inorganic particles of the solid act as a reservoir of nutrients
such as K, Ca, Mg, and Fe, also associated with solid phase are organic
particles containing nitrogen, P, and S. The liquid phase of the soil
constitutes soil solution, which contains dissolved mineral ions and acts as
the medium for ion movement to the root surface. Gases such as oxygen and
carbon dioxide may be dissolved in the soil solution, but their exchange in
root cells occurs in the gaseous phase present in the air space between soil
particles. Supply oxygen to the roots is essential for cellular respiration, the
source of metabolic energy that drives mineral uptake processes

88
XII. Negative charges on the surface of soil particles affect the
adsorption of mineral cations and anions
a. Soil particles, both organic and inorganic have negative charges on
their surface.
b. The organic particles found in humus originate from the products of
the decomposition of dead tissue from plants and animals by the
microbes of the soil
c. The negative surface charge of organic particles result from
dissociation of hydrogen ions from carbolic acid (H2CO3)
d. Negative surface charges of soil particles are important in the
absorption of mineral cations to the surface of the particles. This
absorption is significant because of the high ratio of surface area to
volume of both organic and inorganic soil particles, and it is an
important factor in soil fertility
e. Mineral cations adsorbed on the surface of soil particles are not easily
lost when the soil is leached by water and hey provide a nutrient
reserve available to plant root.
g. Mineral elements absorbed in this way can be replaced by other
cations in a process known as cationic exchange

Soil Mg2+ + H+ Ca2+ →Soil H+ + NH4+ + Mg2+

Particles NH4+ soil solution particles Ca2+ Soil solution
h. The degree to which a soil can adsorb and exchange ions is termed its
cation exchange capacity (CEC), a soil with higher CEC will
generally supply more minerals to the roots
i. In contrast to mineral cations, which are absorbed on the surface of
the soil particles, the soil particles mineral anions are usually repelled

89
 by the negative charge of the soil particles and may remain dissolved
in the soil solution, thus the anion exchange capacity (AEC) of most
agricultural soils is small compared to their (CEC). Among the most
commonly required anions, nitrate, chloride are not adsorbed by the
soil particles and remain in the soil solution, that is related to
technique of using, plants are easy to taking but they also are easy to
leach by water (how to use them for efficient?). On the other hand,
chloride deficient is rarely observed under nutrient conditions.
Phosphate may bind to soil particles containing aluminum or iron, in
this case, the positively charged iron and aluminum will have OH - that
can be exchanged with sulfate, phosphate.
XIII. Relationship between the Nutrient availability, soil microbes, root
growth and soil pH

The reaction of the soil (pH) has direct effect upon the availability of both
essential and nonessential elements in the soil
a. A low soil pH favors the weathering of rocks and the release of
ions such as K+, Mg+2, Ca+2 and Mn+2 as well as trace elements
as Mo, at low pH values , the salts present in the soil as
carbonates sulfates and phosphates are more soluble.
Increasing solubility facilitates absorption by root. Rainfall
levels and decomposition of organic compounds in the soil are
major factors in lowering the soil pH. CO 2 produced as a result
of the decomposition of organic material equilibrates with soil
water in the reaction:
CO2 + H2O → H+ + HCO3-
Forming hydrogen ion

90
Microbial decomposition of organic materials also produces ammonia and
SO3, which can be oxidized in the soil to form strong acids. On the other
hand, in acid regions, the weathering of rock can cause the release of
minerals that act as bases in the soil and result in an increase in pH soil.
However, most major elements becomes less available in acid soils this is
true of N, P, K, and S. The simply considerations of availability are made
more complex by interactions between the supply of a particular element,
microorganisms and the requirement of the plants. Thus phosphorus may be
deficient in alkaline, calcareous soils because of its precipitation as
insolutable phosphates AlPO4 or Ca3(PO4)2, on the other hand, deficiency
may be increased by the enhanced uptake of mycorrhiz.
b. Fungi generally predominate in the soil adjacent to the roots in
the acid pH range. Whereas at higher pH values, the bacteria
may become more prevalent.
c. Fungi growing in side the plant roots and in the surrounding
soil often facilitates mineral uptake by the plants. The capacity
of the root system to absorb nutrients is improved by the
present of external fungi
d. Root growth of plants is generally favored at slightly acid soils

XIIII. The effects of environmental factors on mineral nutrients uptake

by plants
14.1. General

* The mineral nutrient uptake is depended on many environmental factors

such as concentrations of mineral nutrients in the soils (soil solution),
temperature, soil pH, the content of soil oxygen, physical features of the soil

91
as well as the relationships between ions in the soils and so on. If these
factors are suitable, the plants would take up mineral nutrients easily, in
contrast, the environmental conditions are not favor, thus mineral nutrient
uptake will be very difficulty. Even through, plants have deficient symptoms
and result to reduce crop yield even fertility soils.
* The optimal environmental conditions are necessary for high ratio of
mineral nutrient uptake and for maximum crop yields, but it is impossible,
because of environmental factors and their effects on taking up mineral
elements are not to separate and very complement.
14.2. Temperature

* The temperature affects both active and negative mineral uptake, the
maximum T. for taking up mineral elements with most plants is 30 – 35 0c.
The higher T. of 350c the growth of root hairs is depressed even died and
activation of metabolic enzymes in the plant cell are also reduced or retarded
because of decomposition of enzyme structures
* When the temperature is lower than 150c mineral uptake of plants will be
also restricted and stopped depending on plant species, soil types. This effect
is caused by strong reduction of respiration and is not enough bio-energy for
taking up mineral nutrition. On the other hand, at low T. leads to most
mineral matters in the soil to become imsoluable
Thus mineral availabilities are low in the soil.
* From limiting of 150c – 350c, an increase of T. mineral nutrient uptake is
enhanced and optimal temperature of soil for mineral uptake is 25-28 0c with
most of plants. In agriculture, in the winter, in order to improve mineral up
take plants would be cover by nylon or planted in the greenhouse

92
* An increase of temperature in the root zone has much greater effect and
increase the ion concentration in the xylem. Temperature determines the rate
of ion transport in the symplasm and the release into xylem and water moves
accordingly along the water potential gradient. There are distinct differences
between a root and a simple osmo-meter. An increase in the root temperature
results in an increase in the K+ concentration but a decease in the calcium
concentration of the xylem. This shift in K+/Ca2+ ratio might reflect effect of
T. either on membrane selecting or on the relative importance of apoplasmic
pathway of radial transport of Ca 2+ and water. Similar change in the K +/Ca2+
translocation ratio is usually also observed at different soil temperate. This
term effect may have important for the calcium nutrition of plants and might
explain the enhancement of calcium deficiency symptoms at elevated root
temperature, despite a slight increase Ca2+ concentration of the leaf tissue.
14.3. pH soil

The soil reaction affects mineral nutrient uptake by following reasons

* Soil pH affects on nutrient availabilities that directly effects on the
mineral uptake of root system. A low pH soil favors the weathering of rocks
and release of ions as K+, Mg2+, Ca2+ and Mn2+, N, P, S as mentioned about.
* It is not favors for Root growth of plants is generally favored at slightly
acidic pH values (5.5 to 6.5)
* The rate of release of ion into the xylem is closely related to root
respiration, a lack of oxygen strongly depresses the volume flow of ion in to
xylem, the oxygen deficiency seems to effect on release of ions into xylem
and root hydraulic conductivity.
* The activations of micro-organisms in the soil are affected by pH values.
Fungi generally predominate in the acid pH range, whereas at higher pH

93
values, bacteria may become more prevalent. On the other hand, soil pH
affects directly on the type of mineral elements taken up by plants by
following reaction:

Neutral pH
R- CH –COOH ----------------------------- R—CH ----COO-
│ │
NH2 NH3+
* In condition of neutral pH value, protein in bio-membrane of root cell is
balance in electronic, that total + electronic = total – electronic
But in the acidic pH value this balance is changed:

Acidic pH soil
R—CH ----COO- ------------------------------------ R----CH---COOH
│ │
NH3+ NH3+
Mean the total of positive electronic of the protein in root cells is higher than
the total of negative electronic. Thus root cells take up mineral elements in
form of anions more than cations
* In contrast, in condition of the alkaline soil, reaction:

Alkaline pH soil
R--- CH ----- COO- --------------------------- R-----CH------COO- + H2O
│ │
NH3+ NH2
In this condition, root cells is uptake more mineral nutrients in form of
cations than that anions. This is very useful in practical agriculture in

94
 selecting form of nitrogen for fertilization, form of nitrogen applied
depends on plant crops and type of soils and pH of soils should be
controlled by calcium sulfate in order to increase pH in acid soils.
14.4.Oxygen in the soil

* Oxygen in the soil is necessary for respiration of root systems that provide
bio-energy such as ATP for taking up mineral nutrients and other bio-
processes of the plants.
* Oxygen is effect on mineralization process of organic compounds, that
releases mineral elements for uptake by plants
* The content of soil oxygen from 10 to 12 % is suitable and benefic for
taking up mineral nutrients and other benefic bio-processes of both plant
roots and soil. If the content of oxygen in soil lowers than 10%, leading the
root systems are not supplied enough bio-energy for mineral uptake and for
water absorption, mineral nutrient uptake would be strongly depressed, on
he other hand, in this condition respiration of roots can release toxic matters
as lactate or ethylic. The content of oxygen in the soil is lower 0f 5%,
bioprocesses of root system are disordered and restricted with respiration,
mineral uptake and water absorption.
14.5. Soil solution concentration

* As a rule an increase in the external ion concentration leads to an increase

in the concentration of ion in the xylem, the concentration gradient between
the external solution and xylem decreases, diffusion of ion is restricted. The
low water potential in the external solution and the small concentration
gradient between the external solution and xylem also reduces mineral
uptake by plants.

95
* The increase in the exudates concentration of the mineral nutrients with
the rise in external concentration from 1.0 to 10.0mM does not compensate
for the decrease in the exudation volume flow, thus in contrast to the
accumulation in roots, rate of root pressure- driven shoot transport of
mineral nutrients can decline at high external concentration
Vanhoff quotation
14.6. Interaction between ions

In the preceding sections, for study of simplicity the uptake and transport
of a particular ion was treated as a singular process, regulated only by the
physiochemical properties of the ion and the metabolic activities of the
ion and the metabolic activities of individual cell and root. However in
the external solution (soil or nutrient solution) both cations and anions
are present in different concentrations and form and there are many
interactions between them during their uptake.
a. Competition

+ For ion transfer from the external solution to the cytoplasm binding at
transport sites in the plasma membrane is an important step. Competition
between ions of the same electrical charge can be take place. Assuming that
the number of binding sites is small in relation to concentration of
competing ions or capacity of the electronic proton pumps is limited or both.
Such as competition occurs particularly between ions with similar
physiochemical characteristics (electronic and ion diameter) for example,
between the alkali cations as K+ and rubidium (Rb+) although Rb+ can not
replace K+ in its functions in plant metabolism

96
+ Of monovalent cations, competition between K+ and NH4+ is difficult to
explain simply by competition for binding sites at the plasma membrane
(Table 2.16 page 39). Whereas NH4+ is quite effective in competing with K +,
inhibition of NH4+ uptake by K+ is not observed. This seems surprising but
Melgel et. al (1976) obtained similar results with rice, these Authors suppose
that has a sustainable proportion of the annonium nitrogen in the form of
NH4+, but NO3- also permeates the plasma membrane, exporting H + in the
external solution. Inhibition of NH4+ for taking up with negative charges
(anions) is not observed, within individual cell or in the whole plant that is
cations – anion relationship.
+ Cations as K+ and Ca2+ compete quite effectively with Mg2+ and strongly
depress the uptake rate of Mg2+ even in the soil with high concentration of
Mg2+. The competition between ions mostly takes place among cations.
Table 2.17(39)
+ Competition and limited selectivity of binding sites at the plasma
membrane are also observed for anions such as competition between sulfate
and Mo, sulfate and As. Increasing sulfate concentration in the root medium
strong depress Mo, As uptake that is of benefic effect for plant growth and
animals nutrition on soil with toxic level of Mo and As but may become a
critical factor on low Mo soil.
b. Antagonistic relationship between cations as Se and Sulfate is quite
practical importance. Se requirement of human and animals and increasing
concern on Se level in certain soils, increasing level of sulfate very
effectively decreases Se uptake and Se content in plants
+ As and phosphates are taken up by the same transport system in both
lower and higher plants, leading to high uptake and as toxicity in plants
growing on soil with high As levels

97
+ Antagonistic interaction between K+ and Mg2+ when the concentrations of
two cations are high in the soils and a deficient symptom of Mg takes place.
+ The competition between NO3- and Cl- during uptake is of great
importance for crop production. The competing effect of Cl- for NO3- uptake
can be used to decrease NO3- content of some plant species as vegetative
crops, the crops tend to accumulate large amounts of NO3-. On the other
hand, in saline soils the competing effect of Cl- on NO3- uptake may depress
nitrogen nutrition of the plant under these conditions, increasing nitrate
supply can be an effective mean to improve nitrogen nutritional status of the
plants and prevent Cl- toxicity sensitive plant species
* In almost cases, external NH4+ depresses strongly rate of NO3- uptake,
in contrast, externally supplied nitrate has little or no effect on net uptake
of NH4+. According by supplying NH4NO3, NH4+ is usually taken up
much more than NO3- and at higher external NH4+ concentration, uptake
of NO3- is suppressed until the NH4+ concentration is considerable
lowered.
XV. Foliar application (leaf spray fertilization)

The foliar application is method to supply mineral nutrients by spraying

nutritional solution on canopy of plants. It has been popular and
advantage in agriculture in most European Union countries since last
decades of 20th century.
a. Application of mineral nutrients to the leaves can enhance
absorption

* The mineral nutrients can be applied to the leaves as sprays a process

known as foliar application. Leaves can absorb the applied nutrients, and

98
this method may often have agronomic advantages over the application
of nutrients to the soil.
* absorption of mineral nutrients by plant leaves is negative uptake
process, thus mineral elements penetrate to leaf cell according to
regulation from high to enter low concentration. The solution
concentration on the leaves must be higher than that in leaf cells. In this
pathway molecules of mineral nutrients go through hole systems of plant
leaves, there holes have diameter about 1nm and density of 1010hole/
dm2, so that all soluble mineral nutrients (chemical fertilizer) are easy to
enter the leaf cells. But this pathway is not used for chelating and organic
fertilizers.
* There are many advantages of this way compared with soil application
&. When the mineral nutrients are applied on leaves, they are taken up
very fast only 1 or 2 minutes compared with soil application after several
hours. So that the foliar application is useful for supplying nutrients in
stage of nutritional imbalance of the plants, for example, the crops
change from the stage of vegetative growth to the stage of reproduction
as with cereal and fruit species, these stages, crops need to supply more
mineral nutrients but root systems are impossible to take up because
growth of root systems are retarded, if fertilizers applied to the soil
would not be efficiency, the foliar application in these stages is very
useful method. That opens an advantage in agriculture called precise
fertilization. On the other hand, foliar application is also used in stage of
rapid growth of plants
& To reduce pollution of soil and underground water in areas of
agriculture because the ratio of mineral nutrients taken up is high
reaching to 95% compared to 40 to 50 only with soil application

99
 &This method is very effective in condition of restricted uptake of a
mineral nutrients from soil, for example, the foliar application of mineral
nutrients such as Fe, Mn and Cu may be more efficient than application
through the soil, rather in condition, surface layer of the soil is drought
and impossible to supply fertilizers in soil, foliar application is prevalent
method
& Improvement of agricultural products is by increasing
micronutrients, the short of micronutrients in agricultural products causes
to reduce ability of protection of human beings and animals with
diseases. We inflect (catch) so many diseases. Foliar application is great
method to increase mineral elements in plant organs, and improve quality
of A. Products for heath of human and animals. Many experiments had
been carried out by Phu Nguyen Van et al 2002 shown that a mineral
nutrient that is sprayed on leaves its content will increase sharply in the
organs of plant. Even protein content also increases by late nitrogen foliar
application in soybean, oat and wheat (Phu Nguyen Van, 2002)
b. Disadvantages of foliar application

Although foliar application has been advantages in agriculture but it still

has some disadvantages following:
i. The small amount of mineral nutrients taken up through leaves is not
enough for demand of mineral nutrients of plants that is true with
most macro nutrients as N, P, K, Ca. these elements, foliar application
only supports 10% of the demand total. Thus with these elements, F.
A. is only method to subside for soil application. But with
microelements and trace elements, F. A. is enough to supply for
demand of plants

100
ii. Efficiency of foliar application depends on bio-stage and leaf structure
of the plants. Foliar application only has efficiency if it is applied at
late of vegetative stage and beginning of reproductive stage, this stage
of nutritional imbalance takes place. this method is not useful with
plants that their leaves are covered by thick lipid layer
iii. Efficiency of F. A. is still depended strongly on the weather condition
as sunlight, to and rain. Nutrient uptake by plant leaves is more
effective when the nutrient solution remains on the leaves as a thin
film, to promote the production of a thin film, the nutrient solution are
often supplemented with surfactant chemicals such as detergent to
reduce surface tension.
iv. Nutrient movement into the plant seems to involve diffusion through
the cuticle and uptake by leaf cell. For foliar application of nutrients
may be not successful, damage to the leaves can take place if mineral
nutrients are applied on a hot day, when the evaporation is high, salt
may accumulate on the leaf surface and cause burning or scorching.
This problem can be prevented by spraying mineral nutrient solution
in low concentration and on cool days or in the evening on the other
hand, F. A. of Mineral nutrients also is not done by rainy days. Foliar
application has proved practical with cereals as wheat, oat and rye.
Seed protein content in these crops which is important for baking and
animal feeding can be enhanced by the F. A. of nitrogen during late
stages of growth…………….introduce some results in dissertation
thesis

101
c. Leaching of M. N. from the leaves

* leaching can be defined as the removal of inorganic and organic solutes

from the aerial parts of plants by actions such as rain, irrigation, dew, fog
* There four categories distinguished
& Solutes actively excreted to the external surfaces of stem leaves
pods as organic acids (malic acid)
& Excretion of inorganic solution on tips and margins of leaves
& Leaching in damaged leaf areas
& leaching rate increases with leaf age, with leaf age the permeability
of membranes increases, thus the concentration of inorganic and organic
solutes in the apoplasm of the leaf tissue rise resulting concentration
gradient across the cuticle favors leaching by runoff water at the leaf surface.
Under field condition in wetland rice, maximum amount of nitrogen or
potassium in the shoot is leached upto30% during rainy season this loss has
to be considered in studies of nutrient uptake and translocation through out
the growth plants
& Mechanical damage of leaves increases loss of inorganic and organic
solutes, CL- 10 to 70% loss
& Stress conditions such as prolonged darkness, water shortage, high
temperature, air pollution, acid rain
XVI. Relationship between mineral nutrients and plant diseases, pests
1. General

* The effects of mineral nutrients on plant growth and yield are usually
explained in terms of the functions of these elements in plant metabolism.
However, mineral nutrients may create the secondary influences on the

102
growth and yield of crops by effecting changes plant morphology, anatomy
and particularly chemical compositions, thus mineral elements may increase
or decrease the resistance or the tolerance of plants to pathogens and pests.
* The resistance is mainly determined by the ability of the host plants to
limit penetration, development and reproduction of the invading
microorganisms or limiting the feeding of pests
* The tolerance is characterized by the ability of the host plant to maintain
its growth despite the infection or pest attack depending on mineral nutrient,
the nutritional status of plants, host species, type of pathogen and pest
* The resistance and tolerant of plants are considerably influenced by
environmental factors such as T, moisture, M. N. of external solution and so
on
* The close correlation between nitrogen supply and leaf diseases, nitrogen
supply is increased, the incidence of the leaf fungi rises in all cultivars of
Barley
* Usually, a balanced nutrient supply which ensures optimal plant growth is
also considered optimal for plant resistance this is ideal situation in
agriculture, the practically agriculture in Viet Nam is true with this
condition, imbalance in supplying M.N. causes many diseases and pests in
many crops.
2. Fungal diseases
a. Principles of infection

* As a role, germination of spores on leaf, shoot and root surfaces is

stimulated by the presence of plant exudates, the flow of exudates
contributes to success or failure of infection in most fungal diseases. The
rate flow and composition of exudates depends on cell concentration

103
* The concentrations of sugar and amino acids are high in leaves, for
example, when K+ is deficiency, nitrogen supply is high, the most
compositions of the exudates are sugar and amino acids which facilitates to
infect fungi in to host plants
* The moisture on the leaf surface and molecule structure of plant cell walls
play importantly for fungal inflection
* Most fungi and bacteria invade the apoplasm by releasing pectolytic
enzymes, which dissolve the middle lamella. The activity of these enzymes
is strongly inhibited by Ca2+ that explains the high content of calcium in
tissues, their resistance to fungal and bacterium diseases are increased.
* Phenolic compounds play a key in early stages of infection as well as
lignin and suberin biosynthesis. The synthesis of these compounds is a
response to protect plants from inflection.
b. Role of Silicon

* Grasses and wetland rice are mainly silicon accumulation plants, The
silicon supply increases, the silicon content of the leaves rises corresponding
decline in sensitive to fungal diseases. The effect of silicon on resistance of
young leaves is substantial and its application should almost eliminate the
effects of enhanced sensitive to rice with fungi diseases when high levels of
nitrogen are applied this finding is very useful for rice fertilization in Viet
Nam with two fungal diseases
* Formation of a physical barrier in epidermal cells in leaves against the
penetration of the fungal spores or feeding of pests in our country now Si
fertilizer has been applied more in rice production

104
c. Role of Nitrogen and Potassium

* The role of nitrogen in increasing susceptibility of host plants to fungal

diseases, this effect is related to both the nutritional requirements for fungi
and changes in the anatomy and physiology of the host plant in response to
nitrogen. Nitrogen enhances growth rate and during the vegetative stage of
growth the proportion of total nitrogen changes from young to mature tissues
and in high proportion in young tissues, which is more sensitive. In addition
an increase in amino acid concentration in the apoplast and leaf surfaces
seems to have a greater influence than increase in sugar concentration on the
germination
* When plants are supplied with high amounts of nitrogen, the activity of
some key enzymes of phenol metabolism causing content of some phenolics
and lignin may also be lower
* The anatomical and biochemical changes together with the increase in the
content of the low molecules weight organic nitrogen compounds as
substrates for fungi are main factors increasing severely of fungal diseases.
In general, relationship between nitrogen supply and infection of fungal
diseases is more complex
* With potassium, situation is less complex, K + decreases the susceptibility
of host plants both fungal and bacterium diseases and in contrast, The DW
yield is increased in high K+ supply (Fig. 11.8. page 445), this effect of K+ is
confined from range of K+ deficiency to sufficiency plants. According this
effect, plants with potassium deficiency are more severe disease than plants
with K+ sufficiency plants
* The high susceptibility of K+ -deficiency plants to diseases is related to the
metabolic functions of potassium. In deficient plants, the synthesis of high

105
molecule weight compounds (protein, starch and cellulose) is impaired and
low-molecular-weight organic compounds accumulate, these compounds are
as feed for fungi and bacteria
* In plants receiving suboptimal calcium supply the risk of both calcium
deficiency-related physiological disorders and disease susceptibility may
increase.
* The relationship between potassium and resistance is more complex in
seeds and fruits that are supplied with potassium mainly by re-translocation
from vegetative organs thus in suboptimal K+ supply the late fruits or seeds
may be more severe diseases than that in early fruits or seeds.
d. Role of Calcium and Other Mineral Nutrients

* The calcium content of plant tissues affects the resistance of gungal

diseases in two ways
i. First, calcium is essential for the stability of bio-membranes, when
the calcium content is low, the efflux of low-molecular weight compounds
(sugar, amino acids) fro cytoplasm into apoplasm is enhanced that are feed
for infection
ii. Second, calcium polygalacturonates are required in the middle
lamella for cell wall stability. Many fungi and bacteria invade plant tissues
by producing pectolytic enzymes such as polygalacturonase this enzyme
dissolved the middle lamella. The activity of this enzyme is sharply inhibited
by calcium this finding is beneficial for crop production in Viet Nam.
* Plant tissues with low calcium are much more susceptible than tissues with
normal calcium levels to diseases during storage. During storage the fruits
are more susceptive not only physiological disorders but also to fungal
diseases that cause fruit rotting

106
* The effects of micronutrients on diseases has been known, the defense
mechanisms of plants by producing phenolics and lignin are the most well
understood
* The micronutrients such as boron, manganese and copper play a key role
in phenol metabolism and lignin synthesis. These organic compounds
prohibited infection of fungi. In condition of Zn deficiency, a leakage of
sugars onto the leaf surface of Hevea brasilensis species increases severity
of infection. Copper has been extensively used as a fungicide for most fruit
species but the amounts required are at least 10-100 times higher than those
required by plants and it is used by spraying
* The crop production in many developed countries has showed that
micronutrient fertilizer application by spraying increases crop yield by
achievements both improvement of the crop nutritional status and depression
of diseases
* In our country, the farmers try to exploit potential of soils and high yield
of new cultivars by supplying high rates of only macronutrients such as N, P
and K even only N, P and lees rate of K and without application of
micronutrients lead to imbalance of nutrition causing severe diseases for
most crops. In order to increase and maintain crop yields by reducing
diseases, we have to change this situation by balanced fertilization including
macro and micronutrient even thought mixture with beneficial elements
3. Bacterial and Viral Diseases
a. Bacterial diseases

* Bacterial diseases can be divided into three main types: i) leaf spot
diseases, ii) soft rots, iii) vascular diseases.

107
* Leaf spot diseases for example bacterial leaf blight Xanthomonas oryzea
the most common disease in wetland rice. Bacteria enter the host plant
through the stomata, thus the epidermal layer is ineffective barrier to
infection. The bacteria spread and multiply in the intercellular space. The
effects of the mineral nutritional status of the host plant on inflection are
similar to its effect on fungal inflection. Thus when P and K are deficient,
severely diseases are enhanced but not clear with nitrogen deficiency.
* Soft rots, spreading of bacteria within the host tissue as many fungal
diseases, is facilitated by polygalacturinases and related pectolytic enzyme.
The resistance of plants is closely related to their calcium content. In
infected tissues the activity of pectolytic enzymes is very high. Calcium is
inhibited activity of this enzyme and depression of diseases
* Bacterial vascular diseases spread within plants through the xylem they
lead to plug of the vessels. Calcium is effective in both susceptible and the
resistant plants, plants applied adequate of calcium is more resistance with
this disease. The resistant cultivars have high calcium and magnesium
content but lower potassium content
b. Viral diseases

* Multiplication of viruses is occurred in living cells and their nutritional

requirements are amino acids and nucleotides. We have not so much
information in those diseases.
* As a role, nutritional factors which are suitable for plant host growth also
favour for viral multiplication, especially with nitrogen and phosphorus.
Relationship between M.N. and viral diseases is not clear by some reasons
* In nutritional deficiency plants, stimulation of plant growth is by mineral
nutrient supply lead to elimination of symptoms of viral diseases because

108
plants overcome diseases or symptoms are hidden, for example sugar beet
yellow of potato leaf roll virus symptoms might disappear by applying
nitrogen in large even severely disease.
* When grown in Mn deficiency soils, the percentage of infected plants
decreased from75% to 40 %
4. Pests

* Pests are animals as insects, mites which are harmful to cultivated plants.
In contrast to viral and bacterial pathogens, they has digestive and excretory
systems, visual factors as color of leaves are important for recognition
* The main types of the host plant resistance to pests are:
i) physical surface properties, ( color, surface properties, hairs)
ii) mechanical (fibers, silicon)
iii) chemical/biochemical content of stimulants, toxins, repellents. Mineral
nutrients can affect all three factors
 In general, young or rapidly growing plants are more to surfer attack
by pests than are old or slow growing plants, there is often positive
correlation between nitrogen application and pest attack, low
percentage of plants attacked by pests in condition of nitrogen
deficiency.
 In contrast potassium deficiency plants suffered much more attack
than potassium sufficiency plants
 A high content of amino acids in the plants is an important component
for the severity of attack by pests. The high content of amino acids is
reflected by high nitrogen supply and impaired protein synthesis due
to deficiency of potassium or Zn

109
Table.16.1.Effects of Fertilizes Applied on a Soil low in Available
Potassium on Infection of Oat by Eulecunium refulum
Fertilizer
K + Mg N + P + K +Mg Mg P + N +Mg
Number of lice per 0.72 0.82 4.32 8.87
10 cm2 stem section
* The close positive between nitrogen supply, amino acid content and N/C
ratio of plants and attack by pests. This interaction is very complex
* The negative between Boron content and attack of leaves of oil palm
seedlings by red spider mites
Table 16.2. Relationship between Boron supply and intensity of red spider
mites attack on oil palm
Correlation between feeding holes and
Cyanidin content
B supply Mites Feeding holes Feeding holes cyanidin content
(mg/l) (No/m2) ( No/cm2) ( No/cm2 ) ( μg/l)
0 1.8 67 60-65 2-5
0.5 1.7 60 30-50 10-18
5.0 1.2 30
50 1.0 20 20-30 20-32
500 0.9 17
1000 0.9 12
Hock (2003)
* Epidermal cell walls containing silicon act as a mechanical barrier to
sucking and biting insects. The physical properties of leaf surface are also
great of importance in regulating the severity of attack by insects

110
6. Direct and indirect effects of fertilizes application on diseases and
pets

* Under field conditions, fertilizer application affects plant diseases and

pests directly because of mineral nutritional status of the plants and
indirectly by producing dense stands and alteration in light interception and
humidity within a crop stand. In addition the timing of application is an
important factor especially for nitrogen, for example nitrogen applied at the
stage of grain formation can cause severely with fungi of P. Oryea in
lowland rice. This is true with ammonium nitrogen
* Silicon reduces severe for crops by attacking pests. Solubility of silicon in
soils is dependent on many factors as pH, silicon fertilization and form of
nitrogen fertilizer applied
Table. 14.4. Effect of form of nitrogen fertilizer on silicon content and
incidence of Erysiphe graminis in Wheat
Silicon content Disease incidence
(% SiO2 in leaf DW) (% leaf area affected)
Nitrogen form CaCO3 CaCO3
Ca(NO3)2 1.2 27.5
(NH4)2SO4 2.3 18.0
* There are many reports of chloride fertilizer application in macronutrient
as KCL lead to reduce various diseases. Chloride might act directly in the
plants by improving the water balance and thereby tolerance to diseases or
indirectly in the soil inhibition of nitrification or enhancement of Mn
mobilization

111
* The effect of both level and timing of nitrogen supply on the infection are
clear, the most severe infection occurring during large nitrogen applied at
early plant growth. Split application of nitrogen is decreased infection

Practical part
Contents of five lessons
Lesson one: Determination of the total nitrogen content in plant samples
Lesson two: Determination of phosphorus and potassium contents in the
plant samples
Lesson three: Principles of removal of M. E. from nutritional solutions
Lesson four: Classification of deficient symptoms among mineral nutrients
Lesson five: Cultivation of nutritional solution

Lesson 1: Determination of the total nitrogen content in plant samples

1. Principles of the method
* Nitrogen in the plant organs is in both forms organic and incorporated
compounds. In organic compounds as protein, amino acid, nucleic acid, in
incorporated forms as nitrogen bases urea, ammonium and so on
* Nitrogen in form of organic compounds is mineralized by density sulfuric
acid (H2SO4) and converted into (NH4)2SO4, after that NH4+ is distilled by
NaOH, gas NH3 is collected by boric acid (H3Bo). Total nitrogen is
determined by Kjeldahl method

112
2. Mineralization of nitrogen in plant samples
A exactly weight of 0.2 – 0.25 g grounded plant sample is to put into
mineralization vase + 1g catalizer (selen) + 5ml density H2SO4, taking
sample overnight, then sample is boiled at 250oC in 20 minutes and
increasing to 360 or 420oC when the sample is turned to white color,
finishing mineralization
After 20 minutes later, mineralized sample is putted more 50ml distilled
water and turn into 100ml vase for distilling by complex Kjeldahl
The similar process with control vase
3. Distilled nitrogen by complex Kjeldahl
Checking in all complex for working well
Putting 20ml of boric acid solution (4%) into containing vase
25ml of mineralized solution is putted in to distilling vase, clearing by
distilled water
Start working distilled system
Putting about 25ml NaOH 10M into distilling vase
Boiling distilling vase to distil NH4+
Finishing distil NH4+ when containing vase is level of 150ml
Use of stander acid HCL to neutralize solution in containing vase when the
solution is turned to pink color
Similar process with control
4. Calculation for results
(a-b). Ntc.14 .100.k
N% in DW = -----------------------
1000.m
In which:

113
 a. volume of stander acid solution consumed for neutralizing
sample solution
b. volume of stander acid solution consumed for neutralizing
control vase
Ntc concentration of stander acid
14. molecule weight of nitrogen
m weight of sample in solution volume use of distilling
k. correlation of dry well
5. Equipments and chemicals
a.Mineralized vase and oven
Complex of ammonium Kjeldhal volume of 250ml
Vase, buret
Balancer of 0.0002g exactly
b. Chemicals H2SO4 (d = 1.84 no NH4+)
NaOH 10M
Complex catalizer of K2SO4 and Se is grounded well
Indicator Tasiro
H3Bo 4 %

Lesson two: Determination of Potassium and phosphorus contents in the

plant samples

Lesson three: Removal mineral element from nutritional solution

1. Purpose
Every mineral nutrient has different functions in the plants its requirement is
also different at stage of plant growth. In order to determinate the roles of
one mineral element at the certain stage of plant growth, this mineral

114
element is necessary to removal from nutritional solution to study response
to growth or response to field even to deficient symptoms
2. Principles
Removal of one mineral element to study its functions in the plants, we have
to calculate substitution of the chemicals that contain this element by other
chemicals but this substitution does no change content of the other element
For example: Cnop nutritional solution consists of :
Ca(NO3)2: 0.1g
KH2P04: 0.25
MgSO4 : 0.25
KCl: 0.125
FeCl3 0.0123g
When K is removed from this solution, substitution of the chemicals
containing K is KH2P04 and KCl by NaH2PO4 and NaCl
* Substitution of 0.25g KH2P04 by NaH2PO4 is calculated:
In 136g NaH2PO4containing 31g P
In 0.25 KH2P04 containing x (g) P

0.25 x 31
X P = --------------------- = 0.06g
136
In 120g NaH2PO4 containing 31g P
Y(g) NaH2PO4 containing 0.06g P
120 x 0.06
Y(g) = ----------------- = 0.23g
31

115
So that 0.25g NaH2PO4 is substituted by 0.23g NaH2PO4
* Substitution of 0.125g KCL by NaCl is calculated:
In 74,6g KCL containing 35.5g Cl
0.125g KCl containing X(g) Cl
35.5 x 0.125
→ X Cl = ------------------ = 0.06 g
74.6
In 58.5g NaCl containing 35.5g Cl
Y(g) NaCl containing 0.06g
58.5 x 0.06
→ Y(g) = ------------- = 0.1
35.5
So that 0.125g KCL is substituted by 0.1g NaCl
Homework:
1. In 1000ml of Henrighen solution containing mineral elements
Ca(NO3)2.4H2O: 0.708g
KH2P04 : 0.136g
MgSO4 .7H2O: 0.123g
KCL : 0.075g
FeCl3 : 0.025g
In order to remove K from this nutritional solution:
KH2P04 and KCl are substituted by NaH2PO4 and NaCl
How do you calculate for this substitution?
2. In 1000ml of Prianisnikop nutritional solution containing mineral
nutrients:
NH4NO3: 0.2408g
MgSO4 .7H2O: 0.123

116
 CaHPO4: 0.127g
FeCl3 .6H2O: 0.025g
CaSO4.2H2O: 0.344g
KCl: 0.160g
In order to remove K from this nutritional solution:
KCl is substituted by NaCl
How do you calculate for this substitution?
3. In 1000ml of Richter nutritional solution containing mineral nutrients:
Ca(NO3)2.4H2O : 0.72
KH2P04 : 0.20g
KNO3: 0.20g
MgSO4 .7H2O: 0.25g
FeCl3 .7H2O: 0.04g
In order to remove K from this nutritional solution:
KH2PO4 and KNO3 are substituted by NaH2PO4 and NaNO3
How do you calculate for this substitution?
4. In 1000ml of the Pfefer containing mineral nutrients:
Ca(NO3)2.4H2O : 1.92
KH2P04 : 0.33g
KNO3: 0.33g
KCL: 0.16
MgSO4 .7H2O: 0.33g
One drop of FeCl3 5% solution
In order to remove K from this nutritional solution:
KH2PO4, KCl and KNO3 are substituted by NaH2PO4, NaCl and
NaNO3
How do you calculate for this substitution?

117
Lesson four: Classification of deficient symptoms among mineral
nutrients

Lesson five: Plant cultivation without soil

I. Experiment 1: Plant cultivation in nutritional solution
1. Principle of the method
The growth and development of the plants are only depended on factors such
as water, mineral nutrients, sunlight, oxygen, CO 2 and so on and do not
depend on soil. Soil is only as medium for plants to stand well, therefore the
plants can be completely growth and development without soil if they are
supplied enough those environmental factors.
The growth and development of the plants even crop yields depend strongly
on pH of the nutritional solution, value of the optimal pH for most plants is 6
to 7.5. The EC of the nutritional solution reflects amount of mineral
nutrients taken up by plants, modifying the EC to supply mineral nutrients
exactly is necessary. The value of the optimal EC for most crops is
0.5mS/cm – 2.5mS/cm. In technique of cultivation in nutritional solution,
pH and EC is usually changed because mineral ions in solution are taken up.
When the value of the pH and EC is higher than critical level, they are

118
needed to modify to critical level for improving plant growth and
development or high yield.
2. Materials, equipments, nutritional solutions
* Seeds, young plants such as tomato, salat, cucumber …
* pH meter, EC meter
* Nutritional solutions
a. In 1000 ml of Knop nutritional solution containing
Ca(NO3)2.4H2O: 1.0g
KH2PO4: 0.25g
MgSO4: 0.25g
KNO3: 0.25g
KCl: 0.125g
Fe-EDTA: 0.01g pH of 6-7.3
b. (FA0) nutritional solution
KNO3:`` 0.28g/l MnSO4.4H2O: 0.0025g/l
MgSO4: 0.498g/l H3B3: 0.0025g/l
Ca(NO3)2.4H2O: 1.074g/l ZnSO4: 0.0025g/l
KH2PO4: 0.135g/l CuSO4 5H2O: 0.0008g/l
KOH 0.023g/l Na2Mo2H2O: 0.0012g/l
K2SO4: 0.254g/l pH: 6-7.5
Fe-EDTA: 0.01g/l
3. Steps of doing
The bottom of the box is covered by black nylon
Cover of the box is made hole with size similar to size of the holder
Putting the medium and nutritional solution in to box
2- 3 Seeds are sowed in holder in depth of 1-1.5cm
Young plants are planted 1-2 steams in depth of 5-6cm

119
Taking care, not put under the rain and put for enough sunlight, checking in
and supply water
Measurement of pH and EC by machine every week
Determination of the data such as the rate of germination, the number of
emergence roots of the young plants
High, the number of the leaves
Discussion in results
Experiment 2 Plant cultivation in medium containing nutritional
solution
The similar to experiment 1, only different the medium is contained straw
ash or clean sand

120
121