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that the elements nitrogen phosphorus, potassium, magnesium, calcium and iron were
indispensable for the plants. In the absence of any one of these elements the growth of shoots or
roots are stunted.
The essential elements are classified into two broad categories called (i) Macronutrients
and (ii) Micronutrients.
The macronutrients are carbon, hydrogen, oxygen nitrogen, phosphorus, sulphur,
potassium, calcium, magnesium are generally present in plant tissues in concentrations of 1 to
10mg per gram of dry matter.
The micronutrients or trace elements are manganese, copper, molybednum, zinc, boron
and chlorine, recently some other such elements have also been discovered, e.g. cobalt,
vanadium and nickel.
The microelements are required in very low quantity. i.e., about 0.1 mg per gram of dry matter.

Role of Essential Elements

The most important role of the elements is to participate in various metabolic activities such as
regulation of permeability of cell membranes; some elements are required for maintenance of
osmotic pressure of cell sap. While others take part in an electron transport system.

3.4 MACRONUTRIENTS

3.4.1- Carbon, Hydrogen and Oxygen

These are not minerals in origin but are discussed here because they enter into the composition of
practically all organic compounds present in the plant and accounts for a major part of the dry
weight. The significance of water (H2O) can be felt when it is said that water is the liquid of life.
The source for carbon and oxygen is atmosphere and for hydrogen it is water.

3.4.2-Nitrogen
Functions of Nitrogen: The sources of nitrogen are soil and atmosphere. About 78% of
nitrogen is found is atmospheric air but this is of no use to plants in its free state. This enters in
the plants through stomata along with other gases and comes out in the same state unused. The
plants can take nitrogen from the soil in the form of nitrates, nitrites and ammonium salts. The
chief sources of nitrate are sodium nitrate, potassium nitrate, ammonium nitrate and calcium
nitrate. Besides, there are certain highly specialized organism called nitrogen fixers, such as
bacteria and cyanobacteria. They fix atmospheric nitrogen into the soil in the form of nitrites
(No2) and Nitrates (No3).
Nitrogen deficiency symptoms: Nitrogen deficiency causes yellowing of older leaves
(Chlorosis). The plant growth is stunted as protein content, cell division and cell enlargement are
decreased. It also causes dormancy of lateral buds, late flowering, purple coloration and
wrinkling of cereal grains.

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3.4.3-Sulphur
Functions of Sulphur: Sulphur is the constituent of amino acids, vitamin B, coenzyme A and
volatile oils. It is absorbed from the soil as sulphate iron. Through the different amino acids it
participates in protein synthesis. Sulphur affects an increase in nodule formation in root of
leguminous plants.
Sulphur deficiency symptoms: Sulphur deficiency causes yellowing (i.e. chlorosis) of leaves
younger leaves are affected first trips and margins of leaves roll inward a hard woody stem due
to development of sclerenchyma. Sulphur starvation results in shortage of protein.

3.4.4.Phosphorus
Functions of Phosphorus: Phosphorus is absorbed by the plant from the soil in the form of
phosphate ions. It is one of the most important element for the plants.
Phosphorus is vital structural component of the nucleic acids nucleoprotein, phytin,
phospholipids, sugar phosphates, ATP, NADP and numerous phosphorylated compounds. It is an
essential element participating in the skeleton of Plasma membrane.
Phosphorus deficiency symptoms: Phosphorus deficiency causes decrease in the rate of
protein synthesis. It causes premature leaf fall and purple anthocyanin pigmentation. The leaves
become dark blue green in colour and brown necrotic areas are developed on leaves and petioles.
The growth of root and shoot is extremely restricted. Flowering is delayed.

3.4.5. Calcium
Functions of Calcium: This element is always found in green plants. The middle lamella of
the cell wall consists of calcium pectate. Only because of these elements the permeability of the
protoplasm is maintained calcium affects the hydration of colloids. Calcium is believed to be
important in regulating metabolic activities as it activates certain enzymes.
Calcium deficiency symptoms: Calcium deficiency causes disintegration of growing
meristematic regions of root, stem and leaves. Chlorosis generally occurs along the margins of
younger leaves. It also causes malformation of the younger leaves.

3.4.6. Potassium
Functions of Potassium: Potassium is the only monovalent cation essential for plant growth.
This element is usually found in the growing regions of the plant. It is one of the constituents of
protoplasm. Potassium is essential for the formation of sugar and starch and also for their
translocation throughout the plant. It is also needed in cell division, reduction of nitrate,
development of chlorophyll, stomata movements etc.
Potassium deficiency symptoms: Potassium deficiency inhibits synthesis of proteins, which
results in the accumulation of organic nitrogenous compounds in the plant cells Carbohydrate
metabolism is checked. The rate of respiration increases. Mottled chlorosis of leaves occurs
Necrotic areas are developed at the tips and margins of leaves.

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3.4.7. Magnesium
Functions of Magnesium: It is a constituent of chlorophyll and therefore essential for the
formation of this pigment. Magnesium activates enzymes is respiration and photosynthesis. It
plays an important role in synthesis of ATP from ADP and inorganic phosphates.
Magnesium deficiency symptoms: Magnesium deficiency causes interveinal chlorosis of the
leaves. stem becomes yellowish green, often hard and woody. Deficiency symptoms develop on
the older leaves and proceed systematically towards the younger leaves.

3.4.8. Iron
Functions of Iron: Iron is normally absorbed in the ferrous, form though it can be absorbed in
the ferric form as well. It plays an important role in the formation of chlorophyll, constitution of
the chlorophyll. It plays the role of catalyst. Iron in found in ferrodoxin, FRS, flavoprotein and
the iron porphyrin protein, which include cytochromes peroxidases and catalases. It therefore
plays an important role in respiratory mechanism.
Iron deficiency symptoms: Iron deficiency causes rapid chlorosis of the leaves which is
generally interveinal chlorosis may produce a mottled pattern of the leaf may show complete
bleaching, or often become necrotic.

3.5 MICRONUTRIENTS

3.5.1 Manganese
Functions of Manganese: Manganese activates many enzymes which are involved in
photosynthesis, respiration and nitrogen metabolism. It also plays some role in the synthesis of
chlorophyll and in the transfer of electron from H2O to photo-oxidized chlorophyll in
photosynthesis (Homann, 1967)
Manganese deficiency symptoms: Manganese deficiency causes chlorosis which is distinct
from that of iron deficiency. The leaf takes mottled appearance. The chloroplast loss chlorophyll
and starch grains and become yellow green in colour. Dead tissue spots are found scattered over
the leaf.

3.5.2 Copper
Functions of copper deficiency: The element is required is very small quantity. It is very
toxic when present is larger quantity. It acts as catalyst in oxidation reduction reactions, since it
is a constituent of certain oxidising and reducing agents. Copper helps in formation of starch. It
is required for the overall metabolism in plants.
Copper Deficiency symptoms: Copper deficiency causes necrosis of the tips of young leaves.
Both vegetative and reproductive growths are reduced. In crops, the younger leaves wither and

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show marginal chlorosis of the tips. Grain formation is more severely restricted than vegetative
growth.

3.5.3. Zinc
Functions of Zinc: Zinc helps in the formation of chloroplasts. It functions as activator of
certain enzymes. e.g. carbonic anahydrase, alcohol dehydrogenase, hexose kinase etc. Zinc in
required in synthesis of auxins.
Zinc deficiency symptoms: Zinc deficiency causes reduced stem growth due to decreased
synthesis of auxin. It causes chlorosis of older leaves which starts from tips and margin. The
absence of zinc also suppresses seed formation and causes malformation in fruiting trees.

3.5.4. Boron
Functions of Boron: Boron differs from the other micronutrients in that there is no evidence to
suggest its connection with the enzyme systems and it also differs in that it is absorbed as an
onion, i.e. borate and tetraborate, rather than a cation, like the other metallic nutrients.
It is necessary for translocation of sugars and involved in the reproduction and germination of
pollens. It is concerned with water reactions in cells and regulates the intake of water into the
cell. It also affects flowering and fruiting, cell division, metabolism, active salt absorption,
photosynthesis etc.
Boron deficiency symptoms: Boron deficiency causes death of the shoot tip. Flowers
formation is suppressed, root growth is stunted and shoot apices die. Fruit become of small size
and root nodules in leguminous plants are not formed and leaves become coppery in texture.

3.5.5. Molybdenum
Functions of Molybdenum: The main role of molybdenum in plants has been found in the
nitrogen metabolism. It acts as an activator for the enzyme nitrate reductase. It also helps in
formation of proteins. This is absorbed by plant from soil in the form of molybdenum ion (Mo2).
Molybdenum deficiency symptoms
Molybdenum deficiency causes chlorotic interveinal mottling of the older leaves. This may cause
nitrogen deficiency, as it is component of enzymes involved in nitrogen metabolism. It also
inhibits the flower formation.

3.5.6 Chlorine
Functions of chlorine
Chlorine helps in determining solute concentration and anion cation balance in cells. It is
required for cell division in roots and leaves. Chloride ions are essential in the transfer of
electrons from H20 to photo oxidised chlorophyll in photosynthesis.
Chlorine deficiency symptoms: The deficiency of chlorine in plants causes wilting of leaves.

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3.6 ABSORPTION OF MINERAL SALT

Besides water, the plant absorbs from the environment consideration quantities of mineral salts,
gases and various other salts. All these are absorbed in the form of aqueous solutions. The
mineral salts are absorbed from the external solution by the roots.
In plants mineral absorption, also called mineral uptake. In plants, the entrance portal
for mineral uptake is usually through the roots. (Roots, 2005) Some mineral ions diffuse in-
between the cells. In contrast to water, some minerals are actively taken up by plant.

Most of the elements required by the plants are absorbed by them from the soil. The clay
particles of the soil are present in the form of colloids. The micelles of colloidal clay are usually
negatively charged. These charges are balanced by the binding of positively charged ions
(cations) which are taken up from the soil solution. In acidic soil H+ and in alkalins soil ca2++ are
the principal cations associated with the clay particles. In the acidic soil the particles may also
take up and binds potassium, ammonium and other cations. This reversible binding of cations, a
property possessed by clay particles is known as cation exchange. The soil also contains the
anions like Cl-, So4-, HCO3-, H2PO4-, No3-, and OH-. Most anions except the phosphate ions leach
out of soil rapidly.

Passive Absorption: In most cases, the movement of mineral ions into the root occurs by
diffusion. Molecules or ions diffused from a region of their higher concentration to a region of
their lower concentration. As these substances diffuse they exert a pressure. The movement of
mineral ions into root cells as a result of diffusion is called passive absorbtion.

1. Mass flow theory (Bulk Flow): According to this theory ions are taken up by the roots
along with mass flow of water under the influence of transpiration. Russel and Barber (1960)
also supported this theory but raised a question whether the effect of transpiration is direct or
indirect. Lopushimsky (1964) worked in this problem and studied the uptake of radioactive p32
and Ca45, they found that an increase in the hydrostatic pressure (comparable to transpiration
pull) increases ion uptake. So transpiration effect on salt absorption is direct. However, both
mass flow theory and direct influence of transpiration have been challenged in view of recent
research. Both of these fail to explain salt accumulation against osmotic gradient.

2. Ion exchange: In ionic exchange mechanism anions or cations from within the cells are
exchanged for anions or cations of equivalent charge of the external solution in which the tissue
is immersed.
The phenomenon has been experimentally confirmed in excised barley roots in which radioactive
K+ ions exchange place with the non-radioactive K+ ions. A similar exchange mechanism
operates between soil solution and clay micelles. The ions get accumulated against a
concentration gradient without the participation of metabolic energy because cations and anions
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of the external medium get exchanged with H+ and OH- ions, which always remain absorbed on
the surface of the membrane. H+ and OH- are readily available from water.
The process of ionic exchange has been explained by two theories 1 the contact exchange theory
and 2-carbonic acid exchange theory. According to the contact exchange theory an ion may be
absorbed by the plant root without being first dissolved in the soil solution. An ion absorbed
electrostatically to a solid particle such as a plant root or clay micelle, is not held too tightly, but
oscillates within a certain small volume of space. An exchange of ions takes place when the
oscillation volume of one ion overlaps to oscillation volume of another ion. The soil solution,
however, plays an important part in the carbonic acid theory in the it provides the medium for the
exchange of ions between the roots and the clay micelles. Carbon dioxide released in respiration
combines with water to form carbonic acid in the soil solution. Carbonic acid dissociates into
(H+) and (HCO-) ions. A cation absorbed to the clay surface may be exchanged with H+ of the
soil solution. This cation them may diffuse to the root surface in exchange for H +. The cation
them may diffuse to the root surface in exchange for H+. The cation may also be absorbed as ion
pairs with bicarbonate. Thus ion exchange mechanism would allow for greater absorption of ions
from the external medium than could normally be accepted for by the free diffusion.

Fig.3.1 Figure explaining cation exchange theory (Contact exchange theory)

3. Donnan equilibrium: This theory explains the passive accumulation of ions that are non-
diffusible, which may be present on one side of the membrane (Donnan, 1927). Unlike diffusible
ions, the membrane is not permeable to non-diffusible ions. Such ions are called fixed ions. They
may be anions or cations. In which there are no fixed ions, there are equal number of anions and
cations on both sides of the membrane at equilibrium. But in Donnan equilibrium, in order to
balance the charge of the fixed ions (anions) more ions of the order charge (cations) would be
required.
For example there is a membrane that separates a cell from the external medium and
allows exchange of some ions and not others. To the inner side of this membrane there are
anions, which are fixed and non diffusible and therefore the membrane becomes impermeable to
these anions. In such a situation for equilibrium to be reached additional cations are needed to
balance the negative charges of the anions that are structurally formed to the inner side of the
above membrane.
According to the theory, Donnan equilibrium is attained if the product of anions and
cations in the internal solution becomes equal to the product of anions and cations in the external
solution, depicted by the equation as follows:

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Cations inside, Ci+ Anions outside, Ao-

Cations outside Co+ Anions inside, Ai-

For example, a membrane which is permeable to Mg+ and CI- ions and to X- ions present
inside the cell. Here, membrane has 6X- fixed ions on the inner side. CI- ions make across the
membrane by diffusion along the concentration gradient. The concentration of anions on the
inner side is now more than that of cations. In order to balance electrochemical equilibrium
within the cell sap, Mg++ ions move across the membrane against the concentration gradient.
Similarly, if these are fixed cations the anions shall move against the concentration gradient to
bring about equilibrium.

Fig.3.2 Contact exchange theory

Objection to Passive Absorption Concept

Certain strong objections have been raised against passive absorption concept of salt uptake. A
few of them are:
(i) In actual process, the rate of absorption of minerals is too rapid to be explained by passive
absorption.
(ii) No theory of passive absorption adequately explains absorption and accumulation of salts or
ions against the osmotic gradients (or against the laws of diffusion). However, cases of extra
accumulation of K+ ions within the cells (like 1000 times as against the surrounding medium) are
now frequently known in Nitella translucens, Chara australis and Hydrodictyon africanum
(Hober, 1945 and Raven, 1967).
(iii) It has been experimentally demonstrated that there is a close relationship between salt uptake
and metabolic activities. This may be supported by the following examples:
a) A quantitative relationship has been found between anion absorption and respiration.
b) A close relationship between salt accumulation and respiration is found in all cases.
Hopkins (1956) observed that salt accumulation is slowed, and even prevented completely,
with the decrease in the oxygen content of the nutrient medium.

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c) The active phase of salt absorption is inhibited by the absence of oxygen, i.e. oxygen is
required during salt uptake.
d) There is a close relationship between metabolic activites and ability to absorb and
accumulate solutes.
e) The metabolic inhibitors influence the salt absorption Lundegadh (1955) reported that salt
uptake is inhibited by oxidiase inhibitors azides, carbon monoxides and cyanides (all
metabolic inhibitors)
f) Salt uptake has been found to stimulate and increase the rate of respiration. This increased
respiration has been termed as salt induced respiration.
g) Factors like pH, light, oxygen tension and growth affect the salt absorption suggesting that
there is some essential role of metabolic activities in salt uptake.

Active Absorption: According to active absorption concept of salt uptake, it is believed that
this process is supported by metabolic energy, thus the absorption of ions, involving use of
metabolic energy is called active absorption. There have been modification from time to time to
discuss the nature of participation of metabolic energy and that is why several theories have been
proposed.

Active absorption of solutes or theory of salt accumulation

According to this theory Hoagland (1923) suggested that absorption of solutes takes place
against higher concentration of salts. The cells near the tips of roots also have the capacity of
accumulating ions (Hoagland and Broyer 1936). If the initial salt content of the root cells is low
and if other conditions are favorable, the concentration of ions in the absorbing cells may greatly
be increased than that present in soil solution. This involves an expenditure of energy. This
energy in supplied by the respiratory activity of the absorbing cells. The rate of accumulation is
often influenced therefore by the previous metabolic status of the absorbing cells.
The phenomenon of salt accumulation seems confined largely to cells which have the
capacity for cells division and growth, Meristematic cells and cells in the early stages of
enlargement are particularly active in absorbing ions. As cells lose their capacity for growth they
also lose their capacity of mineral salt accumulation. As already referred earlier that
accumulation of salt requires expenditure of energy which is supplied by respiratory activity of
cells.
Hoagland and Broyer (1936) found that if excised roots (young roots) are immersed in
dilute solutions of mineral salts through which N2 is bubbled little or no accumulation of salts
occurs in the root cells. If on the other hand O2 is bubbled through the solution, a rapid
accumulation of salts within root cells takes place. Lack of O2 checks aerobic respiration and
prevents absorption of ions. The accumulation of ions of root cells and their retention within
these cells in a free condition requires an expenditure of energy which is supplied by the process
of respiration.

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Salt accumulation is also affected by the rate of photosynthesis because on this process
depends the supply of carbohydrates which are the respiratory substrates for efficient respiration.
So any factor which reduces photosynthesis, also reduces salt accumulation.
The fact that salt accumulation by root cells is dependent on respiration, suggests that
temperature may have also marked effect on salt accumulation process.

1. Carrier concept: Ions, which are accumulated in cells, may move into the inner space
against concentration and for this movement additional energy is required. This additional
energy is derived directly or indirectly through metabolism. This theory of active absorption has
been supported by various evidences which show that active ion uptake is carried our by carrier
mechanism for both influx and efflux of ions.
Unlike ion channels, the carrier proteins do not have pores. The membrane does not allow
the ions to pass through as it is. The activated ions combine with carrier proteins and form ion-
carrier complex, which is capable of moving across the membrane. The complex moves across
the membrane and reaches the inner surface. Here, the complex breaks and release ions into the
cytoplasm of the cell. Carriers are specific, and combine with particular types of ion.

2. Ion Movement into the Root: Mineral nutrients absorbed by the root are carried to the
xylem. This absorption takes place by two pathways. They are (i) Apoplast pathway and (ii)
Symplast pathway.
(i) Apoplastic pathway- This pathway essentially involves diffusion and mass flow of water
from cell to cell through spaces between cell wall polysaccharides.
The ions that enter the cell wall of epidermis move across cell wall of cortex, cytoplasm
of endodermis cell wall of pericycle and finally accumulate in xylem vessels.
(ii) Symplast pathway- In this pathway, ions that enter the cytoplasm of epidermis move across
the cytoplasm, cortex, endodermis and pericycle through plasmodesmata, and finally reach to
xylem vessels.

Fig.3.3 Apoplastic and symplastic pathways of ion absorption

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Table-1 Differences between Passive Absorption and Active Absorption

S.No. Passive Absorption Active Absorption

1 This processes physical driving force, Here the driving force is energy derived
which is non-metabolic form metrabollic processes.
2 This type of absorption of ions and This type of absorption is not
molecules is sponataneous and proceeds spontaneous and does not proceed
towards equilibrium towards equipbrium
3 Such absorption of a substance occurs The active absorption of a substance
across a protoplasmic membrane from its occurs across a protoplasmic membrane
higher to lower chemical potential. from its lower to higher chemical
potential i.e. against concentration
gradient.
4 In passive absorption the energy yielding When energy yielding metabolic
metabolic processes are not involved. processes are weakened, active transport
system is also checked.
5 The passive transport takes place through The active transport takes place across the
the protoplasmic layer in between the cell protoplasmic membrane (i.e. plasma
wall and the vacuole. membrane, tonoplast, etc.)

3.7 SUMMARY

In this unit we discussed about the mineral nutrition. The plants require some inorganic salts like
Potassium, Calcium, Iron, Sulphur etc for their growth. These inorganic substances occur in soil
in form of solution. This inorganic nutrition of plants is commonly known as Mineral nutrition.
An essential elements are without which the plant cannot complete its life cycle. In fact,
all elements found in a plant are not essential for its growth and life cycle. If the plant grows
normally, the elements are non-essential and if it does not grow normally, it means that the
element is truly essential.
The essential elements are classified into two broad categories called (i) Macronutrients and (ii)
Micronutrients.
The macronutrients are carbon, hydrogen, oxygen nitrogen, phosphorus, sulphur,
potassium, calcium, magnesium. The micronutrients or trace elements are iron, manganese
copper, molybdenum, zinc, boron and chlorine, Recently some other such elements have also
been discovered, e.g. cobalt, vanadium and nickel.
Besides water, the plant absorbs from the environment consideration quantities of mineral
salts, gases and various other salts. All these are absorbed in the form of aqueous solutions. The
mineral salts are absorbed from the external solution by the roots. In plants mineral absorption,
also called mineral uptake. In plants, the entrance portal for mineral uptake is usually through

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