BLOCK 2: MINERAL NUTRITION

Unit 1: Nutrient Elements and Their Importance

Plants are autotroph i.e. they have ability to synthesize their food themselves. They depend
on inorganic nutrients for synthesis of complex organic compounds. Plants derive these
nutrients from the air, water and soil. Inorganic nutrient acquired in the form of inorganic
ions from the soil are called mineral nutrient. The study of the uptake of mineral nutrients and
their use by plants is called mineral nutrition.
Classification of nutrients:
A. Plant nutrients are classified into three types based on their essentiality in a plant:
essential, non-essential and beneficial elements.
(i) Essential elements: An element is considered essential when it is an intrinsic component
in the structure & metabolism and whose absence causes several abnormalities in plant
growth and development. Criteria of essentiality for plant nutrients were given by Arnon
and Stout (1939). According to them, the nutrients must fulfil all the criteria cited by them to
be considered essential. Those are:
 The plant cannot complete its life cycle in the absence of that element.
 The deficiency created by the element can be corrected by supplying that specific element
only.
 The element must be directly involved in plant metabolism.
Epstein (1972) disregarded the second criterion of irreplaceability of any nutrient given by
Arnon and Stout. He then formulated the new criteria:
 The element is regarded essential if the plant cannot complete its life cycle in the absence
of that element.
 The element must form a part of any molecule or constituent of the plant that is itself
essential for the plant.
Based on the essentiality criteria given by Arnon and stout, 16 elements were considered
essential for plants. But after Epstein’s criteria of essentiality came into light, Nickel was
added to the list in 1999, thus increasing the number of essential elements to 17.
Essential elements are further classified into two types based on their relative concentrations
in plant tissue:
(a) Macronutrients or major elements: These are needed by the plants in comparatively
large quantities i.e. more than 1000 ppm or 1000 mg/kg of dry matter. There are total 9
numbers of macronutrients found in plants which include carbon (C), hydrogen (H), oxygen
(O), nitrogen (N), phosphorous (P), potassium (K), calcium (Ca), magnesium (Mg) and sulfur
(S).
Again the macronutrients are classified into three types:
1. Structural or basic or frame work elements: These include carbon, hydrogen and oxygen,
which constitute about 96% of the total dry matter of a plant.
2. Primary nutrients: Primary nutrients are nitrogen, phosphorous and potassium because they
are most often limiting for crop production.
3. Secondary nutrients: Calcium, magnesium and sulfur are secondary nutrients. They are
rarely limiting and more rarely added to soils as fertilizers.
(b) Micronutrients or trace elements: These are required in trace amount or small quantity
i.e. less than 100 ppm or 100 mg/kg of dry matter. Eight numbers of micronutrients are found
in plants which include iron (Fe), boron (B), copper (Cu), molybdenum (Mo), manganese
(Mn), zinc (Zn), chlorine (Cl) and nickel (Ni).
(Trace elements should not be confused with the tracer elements, which are radioactive
isotopes used to trace metabolic pathways, e.g., 14C, 18O, 32P)
(ii) Non-essential elements: These elements are not required by the plants.
(iii) Beneficial elements: Beneficial elements are not required by all plants but may be
essential for particular plants by optimizing their growth and development. Most of the plants
can live a normal life in the absence of beneficial elements in the soil. Some beneficial
elements like aluminum (Al), chromium (Cr), iodine (I), sodium (Na), cobalt (Co), silicon
(Si), tin (Sn), F (fluorine), vanadium (V), selenium (Se) are required by specific plants. These
minerals play specific roles for example, Silicon is essential for pest resistance, prevent water
lodging and aids cell wall formation in Equisetaceae (Equisetum), Cyperaceae and
Gramineae.
Other examples are: V, Si, I: required by some algae
Na: required by halophytes like Atriplex vasicaria (salt marsh plant)
Al: required by ferns
Co: required by legumes
Si: required by rice and equisetum
B. Some researchers have argued that a classification into macronutrients and micronutrients
is difficult to justify physiologically. Konrad Mengel and Ernest Kirkby have proposed
that the essential elements be classified instead according to their biochemical role and
physiological function. The following shows such a classification, in which plant
nutrients have been divided into four basic groups:
1. Nitrogen and sulfur constitute the first group of essential elements. Plants assimilate these
nutrients via biochemical reactions involving oxidation and reduction to form covalent
bonds with carbon and create organic compounds (e.g., amino acids, nucleic acid and
proteins).
2. The 2nd group is important in energy storage reactions or in maintaining structural
integrity. Elements in this group are often present in plant tissues as phosphate, borate and
silicate esters in which the element group is covalently bound to an organic molecule
(e.g., sugar phosphates).
3. The third group is present in plant tissue as either free ions dissolved in the plant water or
ions electrostatically bound to substances such as the pectic acids present in the plant cell
wall. Elements in this group have important roles as enzyme cofactors, in the regulation
of osmotic potentials and in controlling membrane permeability.
4. The fourth group, comprising metals such as iron, has important roles in reactions
involving electron transfer.

Mineral Functions
nutrient
Group I Nutrients that are part of carbon compounds
N Constituent of amino acids, amides, proteins, nucleic acids, nucleotides,
coenzymes, hexosamines etc.
S Component of cysteine, cystine, methionine. Constituent of lipoic acid,
coenzyme A, thiamine pyrophosphate, glutathione, biotin, 5' -adenylylsulfate
and 3'-phosphoadenosine.
Group II Nutrients that are important in energy storage or structural integrity
P Component of sugar phosphates, nucleic acids, nucleotides, coenzymes,
phospholipids, phytic acid etc. Has a key role in reactions that involve ATP.
Si Deposited as amorphous silica in cell walls. Contributes to cell wall
mechanical properties, including rigidity and elasticity.
B Complexes with mannitol, mannan, polymannuronic acid, and other
constituents of cell walls. Involved in cell elongation and nucleic acid
metabolism.
Group III Nutrients that remain in ionic form
K Required as a cofactor for more than 40 enzymes. Principal cation in
establishing cell turgor and maintaining cell electroneutrality.
Ca Constituent of middle lamella of cell wall. Required as cofactor by some
enzymes involved in hydrolysis of ATP and phospholipids. Act as a
secondary messenger in metabolic regulation.
Mg Required by many enzymes involved in phosphate transfer. Constituent of the
chlorophyll molecule.
Cl Required for photosynthetic reactions involved in O2 evolution.
Zn Constituent of alcohol dehydrogenase, glutamic dehydrogenase, carbonic
anhydrase etc.
Na Involved in regeneration of PEP in C4 and CAM plants. Substitutes for
potassium in some functions.
Group IV Nutrients that are involved in redox reactions
Fe Constituent of cytochromes, non-heme iron proteins involved in
photosynthesis, respiration and nitrogen fixation.
Mn Required for activity of some dehydrogenases, decarboxylases, kinases,
oxidases and peroxidases. Involved with other cation-activated enzymes and
photosynthetic oxygen evolution.
 Cu Component of ascorbic acid oxidase, tyrosinase, monoamine oxidase,
uricase, cytochrome oxidase, phenolase, laccase and plastocyanin.
Ni Constituent of urease and dehydrogenase (in N2-fixing bacteria).
Mo Constituent of nitrogenase, nitrate reductase and xanthine dehydrogenase.
Factors affecting mineral absorption in plants:
(A) External factors
i) Water: As nutrients are supplied from the soil to the plant roots in the water medium,
ii) Soil pH: Soil pH affects the rate of mineral absorption by regulating the ion availability
in the medium. Normal pH favors the rapid absorption of monovalent ions, while
alkaline pH favors bivalent and trivalent ion absorption.
Decrease in soil pH accelerates the anion absorption, whereas increase in soil pH favors
cation absorption.
iii) Soil temperature: The rate of mineral absorption increases with soil temperature but up
to a certain level. Very high temperature inhibits mineral uptake, may be due to
denaturation of proteins and enzymes.
iv) Light: Sufficient light increases the salt absorption. Its effect is not direct, but indirect
through transpiration and photosynthesis.
v) Oxygen: Oxygen deficiency reduces the rate of mineral absorption due to low supply of
metabolic energy.
vi) Interaction with other minerals: Absorption of one type of ions may be affected by
other types because of the competition for the binding sites on the carrier. For example,
absorption of K+ is affected by Ca2+, Mg2+ and other polyvalent ions.
(B) Internal factors
i) Growth: Well-grown plants have higher rate of absorption because of the increased
surface area, number of cells and number of binding sites for minerals.
ii) Ageing: Heavy suberin deposition in ageing root cells reduces the mineral absorption
process.
Critical levels of nutrients: Critical nutrient range is defined as that range of nutrient
concentration below which it is deficient and above which it is present in toxic level in plants.
Critical levels vary with the fertility status of the soil and the crop and the cultivator levels.
Diagnosis and prevention of nutrient deficiency and toxicity requires a further knowledge of
symptomology with critical values of any particular element in plants too.
Essential elements Available form
Micronutrients:
Mo MoO42-
Ni Ni2+
Cu Cu+, Cu2+
Zn Zn2+
Mn Mn2+
B H3BO3
 Fe Fe2+, Fe3+
Cl Cl-
Macronutrients:
S SO42-
P H3PO4- , HPO42-
Mg Mg2+
Ca Ca2+
K K+
N NH4+, NO3-
O H2O, O2, CO2
C CO2
H H2O

Specific roles of essential mineral elements in plants:

Major elements or macro elements
1. Nitrogen:
 Important constituent of protein, nucleic acids, porphyrin, alkaloids, some vitamins,
coenzymes etc.
 Porphyrins are important part of chlorophyll and cytochromes.
 Thus, it plays an important role in growth, metabolism, reproduction and heredity.
Deficiency symptoms:
 Nitrogen deficiency causes yellowing i.e. chlorosis of leaves.
 Older leaves are affected first because nitrogen is very mobile in plants and is readily
transported from older to the rapidly developing younger leaves under conditions of
nitrogen deficiency.
 In many plants like tomato, the stem petiole and the leaf veins become coloured due to
the formation of anthocyanin pigments.
 Plant growth is stunted because protein content is reduced. So cell enlargement and cell
division is decreased.
Toxicity symptoms:
 High levels of nitrate or ammonium accumulation in plants lead to dark green leaves with
more or less distinct chlorosis and necrosis of the leaf tips and margins.
 Excess ammonium leads to whitish brown and excess nitrate leads to brownish necrotic
lesions, often with a chlorotic margin between the dark green leaf tissue and the necrotic
zone in the initial stages. In severe cases, the necrotic lesions can spread over the whole
leaf blade.
 Further symptoms include unusually large leaves with large cells and soft, spongy tissue
with or without marginal chlorotic or necrotic lesions. Such leaves are often attacked by
insects, fungi and bacteria. Unusually long shoots (ex: on fruit trees) are another sign of
excess nitrogen.
2. Phosphorus:
 It is an important component of nucleic acid, phospholipids, coenzymes and ATP etc.
 Phospholipids along with proteins may be important constituent of cell membrane.
 Through nucleic acids and ATP, it plays an important role in protein synthesis.
 Through coenzymes like NAD, NADP and ATP, it plays an important role in oxidation,
reduction and energy transfer reaction of cell metabolism like photosynthesis, respiration
and fat metabolism etc.
Deficiency symptoms: Phosphorus deficiency causes
 Premature leaf fall.
 Dead necrotic areas on leaf or fruit.
 Leaf may turn black to blue green or purple in colour.
 Sickle leaf disease.
Toxicity symptoms:
 Excess phosphorus level in the soil can induce zinc and iron deficiency. It may also cause
Ca, B, Cu & Mn deficiency.
 Wilting of the leaves from leaf tip to base has been observed in wheat.
 Greyish-white leaf tip and marginal necrosis, sometimes only as blotches has been
reported for clover, soybeans, oat & barley.
3. Potassium:
 Although it is not a constituent of important organic compounds in the cell, it is essential
for the process of respiration and photosynthesis.
 It probably acts as an activator of many enzymes involved in carbohydrate metabolism
and protein synthesis.
 It is major contributor to the osmotic potential of plant cells.
 It serves to balance the change of both diffusible and non- diffusible ion.
 It plays an important role in stomatal movement.
 It plays vital role in phloem transportation and cell division.
Deficiency symptoms: Potassium deficiency causes
 Hidden hunger
 Mottled chlorosis of leaves
 Scorch or burning on margins of bottom leaves
 Necrotic areas develop at the tip and margins of leaves which curl downward
 Plant growth remains stunted with marked shortening of internodes.
Toxicity symptoms:
 Excess potassium levels in soil can induce Ca & Mg deficiency.
 Growth retardation, leaf scorch, crust formations on leaves and premature leaf dropping
have been reported as direct effects of excess potassium on citrus species in hydroponic
systems.
4. Calcium:
 It is an important constituent of the middle lamella of cell wall.
 It is essential in the formation of the cell membranes.
 It acts as secondary messenger in metabolic regulation (calcium-calmodulin complex)
 It helps to stabilize the structure of chromosomes.
 It may be an activator for many enzymes.
 It is involved in selective permeability of the cell membrane.
 It promotes carbohydrate translocation, amino acid translocation and root development.
Deficiency symptoms: Calcium deficiency causes

 Rapid disintegration of growing meristematic regions of root, stem and leaves.

 Chlorosis along the margins of the younger leaves.
 Malformation of the younger leaves.
 Bitter pit disease in apple (whole apple surface is pitted with small brown necrotic spots).
 Blossom end rot disease in tomato.
 Tip hooking disease in cauliflower, beet and tobacco (hooking of leaf tip occurs).
 Die-back at margins of leaves.
 Young leaves become hooked.
 Premature flower fall
 Inhibits seed germination
Toxicity symptoms:
 High soil calcium level can induce chlorosis & other symptoms resulting from Boron, Fe,
Mn, Zn, & Cu deficiency can be induced.
5. Magnesium:
 It is very important constituent of chlorophyll.
 It acts as activator for many enzymes in phosphate transfer reaction, particularly in
carbohydrate metabolism and nucleic acid synthesis.
Ex: Hexokinase, phosphorylase, peptidase, carboxylase.
 It plays an important role in binding ribosomal particles during protein synthesis.
Deficiency symptoms: Magnesium deficiency causes
 Causes interveinal chlorosis of leaves.
 Older leaves are affected first.
 Causes sand drown disease in tobacco.
 Dead necrotic patches appear on the leaves.
Toxicity symptoms:
 High soil Mg concentration (Mg2+) has a negative effect on plant growth via disturbed
Ca2+ to Mg2+ ratio.
 The roots of the plants are particularly damaged because they are very susceptible to Ca
deficiency.
 The uptake of Mn by plants can be restricted by heavy doses of Mg fertilizer & this leads
to Mn deficiency.
6. Sulphur:
 It is an important constituent of some amino acids like cysteine, cystine and methionine,
which with other amino acids form proteins.
 Disulphide linkage helps to stabilize the protein structure.
 It is also constituent of vitamins like biotin, thiamine and coenzyme A.
 Sulfa–hydril groups are necessary for activity of many enzymes.
 It is required for ethylene biosynthesis.
 It is a constituent of lipoic acid and ferrodoxin.
 It is an important constituent of nitrate reductase along with copper.
Deficiency symptoms: Sulphur deficiency causes
 Causes yellowing or chlorosis of the leaves.
 Tips and margins of the leaf roll inward.
 Stem becomes flat due to development of sclerenchyma.
 Yellowing of tea leaves i.e. tea yellow disease.
Toxicity symptoms:
 High sulphate level in the soil rarely causes damage.
 In citrus, sorghum and beans at high leaf sulphur levels (0.5-1% sulphur in dry matter),
marginal and intercostal chlorosis followed by necrotic lesions have been observed.
 May induce iron deficiency.
 Local effects caused by SO2 emissions are indicated by brown, dark brown or brownish
red blotches on the leaves. They are often surrounded by a somewhat lighter margin.
Micronutrients
1. Iron:
 It is an important constituent of iron porphyrin-proteins like cytochrome oxidase,
catalases, peroxidase, aconitase etc.
 It is essential for the synthesis of chlorophyll.
 It is a very important constituent of ferrodoxin. Thus plays very important role in
biological nitrogen fixation and primary photochemical reaction in photosynthesis.
 It is an important constituent of nitrite reductase enzyme.
 It is required for ethylene production as the enzyme involved in this is ACC oxidase
which requires Fe2+.
Deficiency symptoms: Iron deficiency causes
 Rapid chlorosis of the leaves (interveinal chlorosis).
 The younger leaves are affected first because iron is sparingly mobile in the plant and is
not easily withdrawn from the older leaves to the rapidly growing younger leaves under
conditions of iron deficiency.
Toxicity symptoms:
 Wilting, death of leaf tips, growth suppression, poor flowering
 Excess iron can induce manganese deficiency. (Ex: rose)
 Dark to bluish green colouration of leaves similar to phosphorous deficiency has been
reported from other plants, likewise inhibition of shoot, leaf and root growth, brown
 discolouration of roots and in severe cases the drying out of the dark green leaves without
previous discolouration
 Excess iron may cause brown mottling of paddy rice known as ‘Akagare type I’ or
bronzing. This is characterized by the development of reddish brown blotches on the
lower leaves which spread continuously.
 Sometimes the whole leaf turns brown or the lower leaves turn dark grey and dies early.
The roots become dark blue to black.
2. Manganese:
 It is an activator of many respiratory enzymes.
 It is necessary for evolution of oxygen during photosynthesis.
 It plays important role in nitrogen metabolism.
Deficiency symptoms: Manganese deficiency causes
 Causes chlorotic and necrotic spots in the interveinal areas of the leaves
 Pahala blight disease in sugarcane.
 Grey spec disease in oat.
 Affect chlorophyll synthesis.
 Excess manganese cause iron deficiency.
Toxicity symptoms:
 The symptoms first appear on the older leaves as brown spots or chlorotic and necrotic
blotches which generally advance from the leaf tips and margins on to the leaf blade. In
dicotyledons this is often accompanied by the curling of the leaf margins (crinkled leaf
disease). These symptoms spread rapidly to the younger parts if the excess is severe.
 Appearance of brown spots on veins, petioles and stems. (Ex: potato)
 Excess manganese inhibit Ca, Mg, Fe, Cu and Zn uptake.
3. Copper:
 It is the constituent of several oxidizing enzymes like ascorbic acid oxidase.
 It is a constituent of plastocyanin.
 It is used as fungicide.
 It is a component of nitrate reductase along with iron.
 It is a component of cytochrome.
 Its higher concentration is toxic to the plant
Deficiency symptoms: Copper deficiency causes
 Causes necrosis of the tip of the young leaves.
 Die-back disease in citrus and other fruit trees.
 Reclamation disease in cereals &leguminous plants.
 Exanthema disease (slit in bark).
 Sickle leaf disease.

Toxicity symptoms:
 The symptoms are similar to those of iron deficiencies; however the typical symptoms
include the unilateral chlorosis on the leaves of dicotyledons.
 Excess copper can manganese deficiency.
4. Zinc:
 It is involved in the biosynthesis of auxin (IAA).
 It acts as activator of many enzymes like carbonic anhydrase, alcohol dehydrogenase,
dehydrogenase, DNA polymerase etc.
Deficiency symptoms: Zinc deficiency causes
 Chlorosis of the older leaves which starts from the tips and margins.
 Mottle leaf disease in apple, citrus, walnut and other fruit trees.
 Khaira disease in rice.
 White bud disease in maize.
 Little leaf disease.
Toxicity symptoms:
 Zinc toxicity symptoms resemble to those of iron and manganese deficiency except the
symptoms are not restricted to the youngest leaves but are also found in older leaves.
 Further symptoms include reddish brown necrotic lesions on the leaf blades and margins
like those induced by other heavy metals and symptoms similar to P deficiencies.
5. Boron:
 It plays important role in protein formation in cell wall.
 It plays important role in photosynthesis and nodulation.
 It maintains solubility of calcium in the cell.
 It influences carbohydrate translocation in cell (specifically sucrose).
 It is required for pollen germination.
 It regulates K/Ca ratio in the plants.
Deficiency symptoms: Boron deficiency causes
 Death of the shoot tip.
 Suppression in flower formation.
 Stunted root growth.
 Top sickness disease in tobacco.
 Hard fruit of citrus.
 Heart rot disease in sugarcane and sugar beet.
 Brown heart disease in turnip.
 Cork formation in apple.
 Rolling of leaves in potato.
 Leaves become coppery in texture.
Toxicity symptoms:
 Toxicity symptoms generally appear at the tips and margins of the older leaves and
gradually advance over the reminder of the leaves.
 The damage usually begins as chlorotic spots, stripes, or blotches in which necrotic
lesions develop. The necrotic lesions are often separated from the green tissue by a
yellowing zone.
6. Molybdenum:
 It is associated with the prosthetic group of the enzyme nitrate reductase &nitrogenase
and thus plays important role in nitrogen metabolism.
 It is essential for nodulation in legumes.
Deficiency symptoms: Molybdenum deficiency causes
 Chlorotic interveinal mottling of the older leaves.
 Inhibition in flower formation.
 Whip tail disease in crucifers like cauliflower.
Toxicity symptoms:
 Golden to yellowish orange chlorotic lesions on leaves. This starts on the youngest
leaves.
 The leaf blades are more or less reduced and the internodes are shortened.
 Further symptoms include the failure of buds to sprout and suppressed growth of new
shoots, thickening of the stem, sprouting of axillary buds and succulence of older leaves.
7. Chlorine:
 It is involved in the photolysis of water during photosynthesis in the form of chloride
ions.
 It is required for cell division in leaf and root.
 It is an important osmotically active solute.
Deficiency symptoms: Chlorine deficiency causes
 Wilting of leaf tips followed by general chlorosis and necrosis.
 Bronzing and reduced growth of leaves.
 Roots become stunted in length but thickened near the tip.
Toxicity symptoms:
 Excess soil chloride ions usually itself causes salt damage i.e. leaf scorch, sometimes with
up-curled leaf margins.
 Whitish brown, yellowish brown to brownish black leaf tip and marginal necrosis have
been observed in graminious plants and deciduous trees, growing near the factories
emitting hydrogen chloride gas.
 In contrast, emission of chlorine leads to bleaching of the leaves followed by the
development of the necrotic blotches.

8. Nickel:
 It is cofactor of the enzyme urease in higher plants.
Deficiency symptoms:
 Due to accumulation of urea in leaves, necrosis of leaf tip occurs.
 Beneficial elements
1. Sodium:
 It is an essential micro nutrient for C4 plants.
 It may be related to transport of pyruvate, an intermediate in C4pathway between bundle
sheath cell and mesophyll cells.
 It stimulates growth through enhanced cell expansion.
 It may partly substitute K+ as osmotically active solute.
Deficiency symptoms:
 Reduced growth.
 Wilting.
 Chlorosis and necrosis of leaves.
 Fail to form flowers.
2. Silicon:
 It occurs in normal soil in abundance as SiO2 and also as contaminant in glass containers,
nutrient salts and atmospheric dust.
 In plants, it accumulates as hydrated silica (SiO2.nH2O) in the cell walls, especially
epidermal cells, endoplasmic reticulum and intercellular spaces.
 Plants of family Equisetaceae requires silicon to complete their life cycle where it may
comprise up to 16% of dry matter.
 It may be beneficial to variety of species of higher plants also, especially grasses, where it
may comprise 1-2% of the dry matter and enhances their growth and fertility.
 It forms complexes with polyphenols & may serve as alternative to lignin in providing
strength to cell walls.
 It may overcome toxicity of many heavy metals.
 It is present in the leaf blade of rice.
Deficiency symptoms:
 Deficiency of silicon makes plants more susceptible to fungal infections and lodging.

Physiology of nutrient uptake in plants: Mineral salts are absorbed in the form of ions
through the meristematic regions of the roots near the tips. There are different theories
regarding the mechanism of nutrient uptake in plants, which come under two broad
categories: passive theory and active theory of mineral absorption.
(A) Passive theory of mineral absorption
This type of absorption occurs due to the physical process and does not require expenditure
of metabolic energy. The major theories that explain the mechanism of passive transport of
ions are:
(a) Simple diffusion theory: Minerals are absorbed from the soil solution by the diffusion
process according to the concentration gradient (from the highly concentrated soil solution to
low concentrated root cell sap). Transpiration, rapid assimilation of minerals in the plants and
their compartmentalization in the vacuoles facilitate the rate of diffusion by generating the
concentration gradient of minerals between the soil solution and root cell sap.
(b) Facilitated diffusion theory: According to this theory, ions diffuse into the cell through
the membrane from their higher concentration to lower concentration mediated by a carrier
protein. It is not energy coupled. Its rate is faster than the simple diffusion. The carrier
protein or the integral membrane protein that is involved in the facilitated diffusion is called
as uniporters.
Uniporters allow one ion or molecule to cross a membrane at one site (Fig. 4.6). Here the
activated carrier protein picks a specific nutrient at one surface and translocate the same and
unload at the other surface. After unloading, the protein again comes back to its position with
conformational change. Ex.: Transportation of monosaccharide, amino acids by the calcium
binding protein.
(c) Ion exchange theory: Ions adsorbed on the cell wall surface or membrane of root cells
get exchanged with the ions of same charge from the external solution by the mechanism of
ionic exchange. For example, the positively charged ions like K⁺, Mg2⁺, Ca2⁺ etc. from the
external solution may be exchanged with the H+ ion adsorbed on the root cell wall surface.
Monovalents enter rapidly than the divalents. Exchange of ions occurs either due to physical
contact or via dissolving of CO2 in soil solution. The former is called as contact exchange
theory and the later as carbonic acid exchange theory.
(i) Contact exchange theory: According to this theory, cations or anions may be absorbed
by plant roots without being dissolved in the soil solution. Ions adsorbed on the surface of
root cells and clay micelles are not held tightly but oscillate within small volume of space.
Ion exchange occurs when the oscillation volume of one ion present on the root surface
overlaps that of another ion present on clay micelle (Fig. 6.1A).
(ii) Carbonic acid exchange theory: According to this theory, CO2 released during
respiration combines with water to form carbonic acid (H2CO3) in the soil solution, which
is then dissociates into H+ and HCO3 -. Cation adsorbed on the clay surface is exchanged
with the H⁺ of the soil solution (Fig. 6.1B).The cation thus released into the soil solution
from the clay particle may be absorbed by the root cells in pairs with HCO 3 - or is
exchanged for another cation on the plasma membrane.

Fig. 6.1: Diagrammatic representation of (A) the contact exchange theory and (B) the carbonic acid
exchange theory
(d) Donnan’s equilibrium theory: This theory was given by F.G. Donnan (1927). He
suggested the presence of an electrical balance existing across the biological membrane (cell
membrane and tonoplast). According to his theory, biological membrane acts as Donnan
phase due to the presence of certain pre-existing ions inside the cell which cannot diffuse
outside through the plasma membrane. Such ions are called as indiffusible or fixed ions
which include macromolecules like proteins, lipids, polymers of DNA, RNA etc. that have
many carboxyl groups (-COOH) and phosphate groups (HPO3-). Positively charged particles
like protons (H+) can dissociate from these groups resulting in the acquisition of negative
charge by the macromolecules.
The indiffusible anions are electrically balanced by the entry of equal number of cations from
outside. Now anions from outside will enter into the cell because of a diffusion gradient.
However, each anion will drag cation of equal charge along with it from outside to balance
the charge. The entry of cations will be against their concentration gradient. The
concentration of cations on the inner side of the membrane will be more as compared to
outside, while the concentration of diffusible anions inside the membrane will be lesser than
that of outside because of the excess of negative charges due to fixed anions. Thus, the
accumulation of ions against a concentration gradient can occur without using metabolic
energy until equilibrium is reached at the membrane which is known as Donnan equilibrium.
At equilibrium, the product of cations and anions in the inner side of the membrane will be
equal to the product of cations and anions on the outer side. The charged particles sometimes
fail to distribute evenly across the two sides of the membrane that means equilibrium may not
be reached. Donnan equilibrium is maintained by coordination of both electrical and
diffusion phenomenon. This results in the accumulation of ions inside the cell in a much
greater concentration than outside.
Example: Before equilibrium After equilibrium
Inside Outside Inside Outside

10 A- 20 K⁺ 10 A- 12 K⁺
10 K⁺ 20 Cl- 18 K⁺ 12 Cl-

8 K⁺ 8 Cl-

8 Cl-
(18 x 8 = 144) (12x 12 = 144)

Suppose there are ten numbers of fixed anions (A-) on the inner side of the membrane which
are electrically balanced by the entry of equal number of cations i.e. K+ from outside.
Likewise, there are 20 K+ and 20 Cl- ions on the outer side of the membrane. Now suppose
eight numbers of Cl- will enter into the cell from outside because of a diffusion gradient, it
will also drag equal number of K+ ions from outside to balance the charge.
At equilibrium, cations inside x anions inside = cations outside x anions outside
18 K⁺ x 8 Cl- = 12 K⁺ x 12 Cl-
(e) Mass flow hypothesis: According to this hypothesis, mineral absorption in plant roots
occurs by mass flow. Rapidly transpiring plants show rapid absorption of ions.
Transpirational pull helps in mass flow of minerals.
This theory does not explain the selective absorption of ions. In rapidly transpiring plants,
divalents like Mg2⁺and Ca2⁺are transported more to the top in respect to the monovalents like
Na+, K+ etc.
(B) Active theory of mineral absorption
According to this theory, absorption of salts takes place in two phases: (i) from soil solution
to outer free space of cell which includes cell wall and intercellular space and (ii) From free
space of the cell to cell interior which includes cytoplasm and cell sap.
The first phase is physical and very rapid, while the second phase is physiological and slow.
Entry of ions into the outer space does not require expenditure of metabolic energy, while
ions enter into the cell interior at the expense of metabolic energy.
The involvement of metabolic energy in the active process is supported by the following
evidences:
(i) Plasma membrane contains ATPase required for the active uptake.
(ii) Inhibitors of respiration inhibit mineral absorption.
(iii) Increase in respiration increases salt or mineral absorption between 10⁰C to 30⁰C
temperature.
(iv) Decrease in oxygen content of surrounding reduces the ion absorption.
(v) Q₁₀ for salt absorption shows that it is not a physical process but a physiological process.
The outer free space is further divided into two parts: water free space and Donnan free
space. Cations and anions of soil solution can freely move into water free space by simple
diffusion. However, Donnan free space has fixed adsorptive sites of carboxylic groups. The
carboxylic groups (-COOH) break to form COO⁻ and H⁺ ions. Cations passing through cell
wall displace H⁺ ions and are held by negatively charged sites by weak electrostatic forces.
Then adsorbed cations may be replaced by other cations.
Ions enter into the cell interior through the biological membranes (cell membrane and
tonoplast). There are several theories explaining active absorption of minerals, some of which
are discussed below.
(a) Ion carrier theory: This theory was proposed by Van den Honert (1937). According to
this theory, active transport of ions across a membrane occurs by mediation of some proteins
located on the membranes called carrier proteins. The carriers possess specific binding sites
for particular ion species, thus enabling selective ion transport through the membrane.
 Fig. 6.2: The ion carrier theory

The binding and releasing of ions to carriers is an energy consuming process, which is
supplied by ATP. At the outer surface of the membrane, the activated carrier binds to the ion
for which it has affinity to form a temporary carrier-ion complex. Then the carrier undergoes
conformational change and releases the ion to the inner side of the membrane. This release is
probably mediated by enzyme phosphatase (fig. 6.2). Phosphatase enzyme splits off the
phosphate group from the carrier complex and makes it inactive. The inactivated carrier is
again activated by ATP in the presence of enzyme kinase. The activated carrier molecule
return back to the outer surface of the membrane for complexing with other ions and the
process continues.

Carrier*(Activated) + Ion (cation or anion) Carrier*-Ion complex

Phosphatase
Carrier*-Ion complex Carrier + Ion

Kinase
Carrier (Inactivated) + ATP Carrier* (Activated) + ADP

(b) Lecithin carrier theory: This theory was proposed by Bennet Clark (1956).According
to him, lecithin (a phospholipid present on membranes) acts like carrier of ions. It carries
cations and anions simultaneously due to possession of opposite charges on its two fractions
i.e. choline group and phosphatidic acid group (Fig. 6.3). Choline group possesses positive
charge and phosphatidic acid group possesses negative charge. Therefore, cations get
attached to the phosphatidic fraction and anions to the choline fraction.
Lecithin binds with the ions on the outer surface of the membrane and releases them on the
inner surface after its dissociation into two fractions mediated by the enzyme lecithinase. The
two fractions reunite to form the functional lecithin in the presence of enzyme choline
acetylase, choline esterase and energy source ATP. The functional lecithin again binds to the
cation and anion on its two fractions and the process continues.
 Fig. 6.3: A schematic representation of lecithin carrier theory

(c) Cytochrome pump theory: This theory was proposed by Lundegardh and Burstrom
(1933). According to this theory, the mechanism of anion and cation absorption is different.
Anions are actively carried by cytochrome chains (heme-containing membrane proteins)
whereas absorption of cations is a passive process. Anion uptake requires metabolic energy.

Fig. 6.4: Diagrammatic representation of cytochrome pump theory

Dehydrogenation reaction occurs in the inner side of the membrane to produce protons (H⁺)
and electron (e⁻) (fig. 6.4). Electrons are carried to the outside through the cytochrome chain
while the protons by simple diffusion. Terminal cytochrome at the outer side of the
membrane gets oxidized by losing the electron. This loss of electron is compensated by the
counter movement of anions to the inner side of the membrane via cytochrome chain.
Terminal cytochrome towards inner side of the membrane now gets reduced by accepting
electron released from the dehydrogenase reaction. Reduction of terminal cytochrome
towards the inner side of the membrane subsequently releases the anion into cytoplasm.
Electron lost towards outer side of the membrane by terminal cytochrome unites with H⁺
released during dehydrogenation reaction to form hydrogen, which is finally accepted by
molecular oxygen to produce water.
Objection:
 This theory explains active absorption of anions only.
 This fails to explain the selective uptake of ions.
 It has been observed that even cations can stimulate respiration.
(d) Secondary active transport: Active uptake of ions requires energy which is mainly
derived from the breakdown of ATP. However, energy may be derived secondarily from
other source. The type of transport where ATP is the energy provider is called as primary
active transport or direct active transport. Likewise, that type of transport where energy is
derived from the electrochemical potential gradient across the membrane created by the
primary active transport of ions out of the cell, is called as secondary active transport.
Electrochemical potential gradient is the combination of concentration gradient and charge
difference present across a membrane that affects the movement of ions. The most common
energy source in secondary active transport is the energy derived from the pumping of
protons across a cell membrane or the proton motive force. There are two forms of secondary
active transport: symport and antiport (Fig. 6.5).

Fig. 6.5: schematic representation of types of transport systems

i) Symport or co-transport: When two ions or molecules cross the membrane at the same
site in the same direction, it is called as symport. Here two different ions bind to the two
active sites of the same channel protein or carrier protein or symporters. Then the carrier
protein undergoes conformational change and rotates to the other side of membrane to
release the ions simultaneously.
Ex.: H+/K+ symporters present in the roots that allow one H+ and one K+ ion into the cell.
Likewise, there is symport of Na+ and glucose in bacterial cells. On the exterior side, the
carrier protein has two binding sites, one for sodium and one for glucose. When both of
these bind to the protein, it undergoes a conformational change allowing the
electrochemical gradient to provide energy for transportation of both of these molecules
into the cell.
ii) Antiport or counter-transport: When two different ions or molecules are being
transported across a membrane at the same time but in opposite direction, it is called as
antiport or counter-transport. Here one specific ion is transported in and the other is
transported out by the same transporter. The transporters carrying out the counter-transport
of ions are known as antiporters or exchangers.
 Ex.: Transportation of Na+ ions to the outer side and K+ to the inner side of the membrane
is mediated by Na+/K+ ATPase pump. Likewise proton-sucrose antiporters operate in the
vacuolar membrane to accumulate sucrose in plant vacuoles.

Difference between active and passive mineral absorption:

Active absorption Passive absorption

i) Movement of ions across the membrane i) Movement of ions across the membrane takes
takes place against the concentration gradient. place along the concentration gradient.
ii) Movement of ions does not proceed ii) Movement of ions proceed towards
towards equilibrium. equilibrium.
iii) Energy consumption occurs during the iii) No energy is consumed.
process.
iv) Enzymes are involved. iv) Enzymes are not involved.
v) Specific carrier proteins are required. v) No carriers are involved.
vi) Rate of mineral absorption is dependent vi) Rate of mineral absorption is independent of
on respiration. respiration.
vii) This cause salt accumulation in the cells. vii) This does not cause salt accumulation in the
cells.

Ion transporters and ion channels:

 Ion transporters are transmembrane proteins that transport ions (or other small molecules)
across a biological membrane to accomplish many different biological functions
including, cellular communication, maintaining homeostasis, energy production, etc.
There are different types of transporters including, pumps, uniporters, antiporters, and
symporters.
 Ion transporters differ significantly from ion channels. Channels are pores that run
through the membrane, whereas transports are proteins that must change shape to switch
which side of the membrane it is open to. Because of this, transporters are much slower at
moving molecules than channels.
 Ion channels and ion transporters move the ions via facilitated diffusion which is a type
of passive transport.
 However, only ion transporters can also perform active transport, which involves moving
ions against their concentration gradient using ATP as energy source. This creates an
electrochemical gradient or concentration gradient which can then be used by secondary
transporters or other proteins as a source of energy to further transport ions.

Genes encoding for ion transporters

Different genes encode different transporters that are involved in ion transportation. Plant
nitrate transporters are encoded at least by four gene families, NRT1 (NPF), NRT2, CLC, and
SLAC1/SLAH. The CHL1 (NRT1) gene of Arabidopsis encodes a nitrate-inducible nitrate
transporter.

Role of mycorrhizae in plant nutrient acquisition

Mycorrhizae are essential in areas where soils are deficient in water and certain nutrients -
conditions that are found in the desert. Even when there is an ample amount of a nutrient, it
may not be readily accessible to the plant. A dramatically larger root system (or
mycorrhizae) permits the plant to obtain additional moisture and nutrients. This is particularly
important in uptake of phosphorus, one of the major nutrients required by plants.

When mycorrhizae are present, plants are less susceptible to water stress. The rapid uptake of
nutrients, for example, ‘P’, by roots leads to the formation of a depletion zone, and fungal
hyphae extend beyond the depletion zones to penetrate and exploit a larger volume of soil to
uptake nutrients. Not only do the fungal threads help to bring water and nutrition into the
plant, but they also can store them for use when rainfall is sparse and temperatures are high.
When organic matter (compost) is added to improve a soil, mycorrhizae are important in
making its nutrients available. The residual organic matter and the hyphae improve the
structure of the soil. Recent research indicates that the fungi even help break down rock,
increasing availability of the essential nutrients within, such as potassium, calcium, zinc and
magnesium.

Role of PGPRs in plant nutrient acquisition

Plant Growth Promoting Rhizobacteria (PGPR) has the ability to increase the availability of
nutrient concentration in the rhizosphere by fixing nutrients, thus preventing them from
leaching out. As an example, nitrogen, which is needed for the synthesis of amino acids and
proteins, is the most limiting nutrient for plants. The mechanisms by which atmospheric
nitrogen is added into organic forms that can be assimilated by plants are exclusive to
prokaryotes. A rare example of a free-living nitrogen-fixing organism is Azospirillum, often
associated with cereals in temperate zones and also reported to be able to improve rice crop
yields.
Some PGPR have the ability to solubilize phosphate, resulting in an increased availability of
phosphate ions in the soil, which can be easily taken up by the plants. Kocuria
turfanensis strain 2M4 isolated from rhizospheric soil was discovered to be a phosphate
solubilizer, an IAA producer, and a siderophore producer.
Lavakush et al. studied the effect of PGPR on nutrient uptake by rice. They used PGPR
strains such as Pseudomonas fluorescens, Pseudomonas putida, and Pseudomonas
fluorescens.
Role of root exudates in plant nutrient acquisition
Root exudates are the general term for organic compounds released from roots to growth
media during plant growth and development, accounting for 5–21% of plant photosynthetic
products (Wang et al., 2021), including low molecular weight primary metabolites (especially
sugars, amino acids and organic acids) and secondary metabolites (phenols, flavonoids and
terpenoids). Root exudates improve soil moisture and nutrient bioavailability and plant
growth by altering rhizosphere physical, chemical or biological properties. Root secretion
activities enable plants to adapt to and gradually change the soil environment in which roots
are exposed.
Ion transportation through xylem
 Elements that traverse the root via the symplasmic pathway are loaded into the xylem by
various transport proteins across the plasma membrane of xylem parenchyma cells.
 As per Lin et al., (2008), nitrate can be loaded into the xylem by members of the NRT1
(nitrate transporter 1) family.
 In addition, anion channels can facilitate the movement of nitrate, sulphate, phosphate
and chloride in the direction of their electrochemical gradients from the cytosol of xylem
parenchyma cells to the xylem.
 Boron is loaded into the xylem by orthologues of the Arabidopsis AtBOR1 transporter,
whose activities are regulated in response to plant B status to ensure appropriate B
concentrations are maintained in the shoot.
 Potassium is loaded into the xylem by voltage-gated, outwardly rectified K-channels
present in the plasma membrane of root pericycle and stelar parenchyma cells, such as
orthologues of the AtSKOR protein of Arabidopsis thaliana.
 Cations that are present in low concentrations in the cytosol of root cells are loaded into
the xylem by active transport mechanisms. Calcium is loaded into the xylem by members
of the P2A-Ca2+-ATPase and P2B-Ca2+-ATPase families and members of the heavy metal
P1B-ATPase family load Zn2+ and Cu2+ into the xylem.
 Similarly, it is thought that Mg2+ and Mn2+ are loaded into the xylem by ATPases,
although the genes encoding these transporters are unknown. Cation carriers have also
been implicated in loading Zn2+ and Fe2+ into the xylem.
 A significant amount of Ca and other potentially cytotoxic elements, such as Zn, Fe and
Na, can also reach the xylem through an apoplasmic route when they are present at high
concentrations in the soil solution.
 Nitrogen is mostly present in the xylem in its inorganic forms, although amino acids and
amides have also been observed. Similarly, phosphate and sulphate are the dominant
forms of P and S in the xylem. Calcium, Mg, Mn and Zn are likely to be transported in
the xylem as cations or cation complexes with organic acids.
 Iron is transported mainly as Fe3+ citrate. In Arabidopsis thaliana, a member of the
multidrug and toxin efflux (MATE) transporter family, AtFRD3, is expressed in the root
pericycle and appears to be involved in loading citrate into the xylem.
 Zinc can also be transported as a histidine complex, and Zn, Cu, Mn and Ni can be
transported as nicotianamine (NA) complexes.
Ion transportation through phloem
 Nitrate can be loaded into the phloem by transporters of the NRT1 family.
 Of the nutrients loaded into the phloem, K is usually present in the highest concentration,
followed by P, Mg and S. Potassium is loaded into the phloem by voltage-gated,
 inwardly-rectified K-channels with electrophysiological properties resembling AtAKT2/3
of Arabidopsis thaliana.
 Sulphur occurs in both the reduced form (e.g., glutathione, S-methylmethionine,
methionine, cysteine) and as sulphate. Sulphate is loaded into the phloem by orthologues
of the Arabidopsis thaliana AtSULTR1;3 transporter, while methionine and cysteine are
likely to be loaded by amino-acid transporters. Sulphate concentrations in the phloem sap
can be as high as those of phosphate.
 Chloride and Na may also be present at high concentrations, but this depends on their
external supply and the plant species. In contrast, the concentration of Ca in the phloem
sap is always very low, regardless of plant species.
 Members of the ZIP family are thought to transport Zn into the phloem. Fe, Mn, Zn and
Cu are probably also loaded into the phloem by YSL proteins. A large proportion of Fe
transported in the phloem is complexed to the Iron Transport Protein.
 Except Ca, concentrations of all solutes are usually several times greater in phloem
exudate than in the xylem exudate.

Strategies to acquire and transport minerals under deficient levels

1) Modulation of the root system architecture according to the nutrient conditions:

Nutrient concentrations affect the length, number, angle and diameters of the primary
roots (PRs) and lateral roots (LRs), and root hair development. These developmental
changes result in nutrient-dependent root system patterns. For example, under P
starvation, PR elongation is inhibited, while LR formation is enhanced concomitantly,
resulting in the formation of a shallow root morphology and increased root surface area.
Phosphate tends to accumulate in the topsoil layer (as it readily binds to soil organic
matter) where it is immobilized. Thus, below-ground morphological responses of plants
to P starvation probably contribute to efficient acquisition of phosphate by exposing a
large root surface area to P. In contrast, for water-soluble nutrients such as nitrate and
sulfate, which are readily leached to deeper soil layers, root development towards lower
layers is preferred.
2) Modulation of transport activity within and among plant organs: Nutrient
transporters regulate nutrient uptake into root cells and subsequent translocation within
the plant body. Nutrient conditions strictly regulate transporter gene expression at both
the transcriptional and post-transcriptional levels. Transporters also undergo post-
translational modification to control transport activities. This regulation at multiple steps
benefits nutrient homeostasis under a wide range of nutrient conditions.
3) Secretion of organic acids by plants
Organic acids in plants regulate the cellular metabolism. During stress conditions, plants
exudates the organic acids into soil as it improves the mineral uptaking capacity and also
provides tolerance to toxic metals. These act as chelating agents and help in carbon
sequestration. OAs are chemically charged so they can balance the excess ions in cells
and regulate cellular pH and osmotic potential. Among the OAs, citric acid solubilizes P
ions in the soil and forms a polymer that quickly diffuses to roots and release P ions after
reduction.
4) Recirculation of mineral nutrients:
The mineral cycle is a biogeochemical and ecological process that regulates the flow,
distribution, and migration of mineral nutrients across the Earth’s surface. The Cycling of
mineral nutrients helps in fulfilling the demand for growth of apical root zones and
reduces fluctuations in external nutrient supply of crops. It is also responsible for phloem
loading and export of photosynthates from source leave. Nutrient deficiency leads to dry
matter partitioning between shoot and roots and help in free solute flow of mineral
nutrients from source leaves to roots.
5) Rhizosphere microbes symbiosis
Rhizosphere is the most important part of a plant growing system with all nutrient
dynamics carried in it. The microbes associated with roots and plant growths are acutely
responsible for plant growth and developmental process. The microbiota colonizing the
rhizosphere contributes to the plant growth, productivity, carbon sequestration, and
phytoremediation. Proteobacteria and Firmicutes make up the biomass of rhizosphere.
The release of organic carbon by the plant roots increases the microbial activity in the
rhizosphere.
6) Adaptation strategy in acidic soils
A balanced nutrient supply protects plants against all forms of stress with proper
application of fertilizer doses. The depletion of nutrients, soil organic matter, and erosion
are the principal forms of nutrient deficiency. Though being an abiotic stress factor, a
limited nutrient stock has to be identified to avoid the restricted nutrient availability. This
can be managed by maintain the soil conditions.
7) Adaptation strategy in toxic soils
The increase in toxic percentages in soil increases the H+. Al and Mn+ concentrations
thereby altering the nutrient absorbing mechanism. The leaching percentages also hike
up, inhibiting the root growth thereby affecting the proper functioning of plants. The
decrease in cation exchange concentration also occurs due to the decrease in calcium,
potassium and magnesium concentrations. Plants evolve to different tolerance strategies
like lowering the internal mineral demands, recirculating the mineral nutrients, and
adapting tissues to tolerate higher toxic conditions. Tolerance is a prime path but
avoidance strategies of the plant help in sustaining under such toxic conditions. The
strategies adopted are like modification of root conditions, increased storage of mineral
nutrients that help in targeting the nutrient deficiencies and overcoming the toxic stress
during no availability of nutrients. For example, silicon helps in decreasing the toxic
effects of heavy metals and provides protection against different fungal diseases by
positively influencing the structure and function of plant cell walls.
Unit 3 Concept of Foliar Nutrition

Foliar nutrition is a technique of feeding plants by applying liquid fertilizer directly to their
leaves. Plants are able to absorb essential elements through their leaves. The absorption takes
place through their stomata and also through their epidermis.
Foliar spray is the application of fertilizers to foliage of the crop as spray solution. This
method is suitable for application of small quantities of fertilizers, especially micronutrients.
Major nutrients can also be applied by this method when there is no adequate moisture in top
layer of soil. Foliar application is not substitute for soil application, but only a supplement to
it.
The purpose of foliar feeding is not to replace soil fertilization. Supplying a plant’s major
nutrient needs (N, P and K) is most effective and economical via soil application. However,
foliar application has proven to be an excellent method of supplying plant requirements for
secondary nutrients (Ca, Mg and S) and micronutrients (Zn, Mn, Fe, Cu, B, and Mo), while
supplementing N-P-K needs for short and/or critical growth stage periods. Primarily, foliar
feeding is intended to delay natural senescence processes shortly after the end of reproductive
growth stages. Foliar feeding targets the growth stages where declining rates of
photosynthesis and levelling off of root growth and nutrient absorption occur, in attempts to
aid translocation of nutrients into seed, fruit, tuber or vegetative production. Secondarily,
foliar feeding can be an effective management tool to favourably influence pre-reproductive
growth stages by compensating for environmentally induced stresses of adverse growing
conditions and/ or poor nutrient availability. Early foliar applications can make an already
good crop better, either by stimulating more vigorous regrowth or maximizing the yield
potential growth stage period. The advantages of foliar feeding in accomplishing the desired
crop responses are two-fold.

Mechanism of Foliar Fertilization

• Nutrients must enter the leaf before entering into the cytoplasm of the leaf cell.
• Nutrients must effectively penetrate the outer cuticle and wall of the epidermal cell.
• Once penetration has occurred, nutrient absorption by the cell is similar to absorption by
the roots.
• Among all the components the cuticle offers the greatest resistance to the nutrients.
Foliar nutrient droplet size for effective entry
• Smaller droplets cover a larger area and increase efficiency of foliar applications.
• However, when droplets are too small (less than 100 microns), a drift might occur.
Role of wetting agents in entry of nutrients
• A wetting agent facilitates the uptake of water during initial wetting and optimizes the
movement of water in the substrate which improves the re-wetting of over-dry plants.
• This is due to the wetting agent’s ability in lowering the surface tension of water which
allows it to spread more easily and evenly in the peat. The wetting agent does not affect
the total water uptake of the substrate so the plants, or more precisely the pots, do not take
up any extra water because of the wetting agent added to the substrate.
Timing of Foliar Applications:
1) Proper Growth Stage: This is one of the most critical aspects of a foliar feeding
program. Foliar applications should be timed to provide needed nutrients during the yield
potential determining time frame of plant development, which will in turn favourably
influence the post-reproductive development stages. Multiple, low rate applications may
show the most favourable responses within these time frames, whether a crop is
nutritionally sound or not. Careful crop growth stage monitoring on a weekly, and
sometimes a daily basis, is essential. A comprehensive plant tissue analysis program
taken just prior to the desired application is also essential to establish levels of plant
nutrients most limiting to crop growth. DRIS (Diagnosis and Recommendation Integrated
System) analysis of tissue tests is the best method of relating tissue nutrient levels to
desired plant needs by ranking plant nutrients in order of most limiting to least limiting.
2) Proper Crop Condition: Generally speaking, crops that are nutritionally sound will be
most likely to respond to foliar feeding. This is due to better tissue quality (allowing for
maximum absorption of nutrients into leaf and stem) and better growth vigour (allowing
for translocatable nutrients to be rapidly moved to the rest of the plant). Crops under heat
or moisture stress show less response to foliar applications due to lower leaf and stem
absorption rates and/or poor vigour. However, foliar feeding does benefit crop
performance and yield if an application was made prior to heat or moisture stress.
Recovery from cold growing conditions and herbicide stress can be hastened with proper
foliar applications. Good recovery of corn suffering from light to moderate hail damage
has been shown where nitrogen-sulphur solutions were foliar applied. Under most
conditions, however, due to the practical and economic limitations on the amount of
nutrients that can be foliar applied to give a favourable growth response, foliar feeding
has a limited rescue capability.
3) Proper Meteorological Conditions: Environmental influences, such as time of day,
temperature, humidity and wind speed influence the physical and biological aspects of
foliar applications. Plant tissue permeability is an important factor in absorption of
nutrients into the plant: warm, moist and calm conditions favour highest tissue
permeability, conditions found most often in the late evening hours, and occasionally in
the early morning hours.
Table: Meteorological condition favouring foliar applications

Time of Day Late evening; after 6:00 p.m.

Early morning; before 9:00 a.m.
Temperature Low temperature 18-19 0C (Ideal 210C)
Humidity Greater than 70 % relative humidity
Wind speed less than 5 mph
Rainfall Within 24 to 48 hours after a foliar application
may reduce the application effectiveness, as not
all nutrient materials are immediately absorbed
into the plant tissue
Significance of foliar nutrition:
1) Foliar application is a corrective nutrition:
Foliar sprays are fast acting and effective for treating nutrient deficiencies. It is much
faster than root uptake. Therefore foliar fertilization is the method of choice when
deficiency symptoms are noted, and prompt correction of deficiencies is required.
Nutrients rapidly absorbed through the foliage, providing the plant with the missing
nutrients, and strengthening it.
2) Foliar nutrition is a complementary nutrition:
Foliar application is a must when soil fertilization is sub-optimal or ineffective. In many
farming practices most of the fertilizers for the growth season are applied as base
dressing. Such practice often result a significant decrease of essential nutrients in the soil
during the season. This may affect plant growth, and reduces yield. A mid-season foliar
sprays of fertilizers will balance and complete plant nutrition, and help to retain healthy
and fruitful growth. A relatively small amount of fertilizers will lead to a dramatic
improvement and will get the crops to optimal growth curves. In some cases uptake of
nutrients by the roots may be impaired due to low temperatures, water logging,
nematodes, or other reasons. Here also, foliar nutrition offers a beneficial complementary
nutrition.
3) This is growth boosting:
A precise timing of foliar nutrition lead to growth and yield improvement. Certain phases
of plant development are highly important in determining the final yield. A foliar
nutrition precisely given during these specific phenology stages ensures optimal growth,
and best possible yield. In relatively low concentrations at these critical stages, foliar
nutrition boosts physiological processes of the plant, ensures optimal development, and
contributes significantly to higher yields and better quality.
4) This can activate particular plant mechanisms:
Foliar application of certain fertilizers is known to induce dormancy breaking in grapes
and deciduous fruit trees. They are also efficient in stimulating flowering, mainly in
mango.

Factors affecting foliar nutrition:

1) Cuticular thickness: Cuticle surface is not smooth, and is covered with waxy layer,
protuberances and structures like trichomes which increase the surface tension. Therefore,
any additive included in the aqueous sprays should reduce the surface tension and
increase the surface area of absorption. These are obtained by wetting agents and
surfactants which are now commercially available for different nutrient elements and
crops.
2) Stage of leaf development: Young leaves have been found to absorb nutrients more
effectively than the old and mature ones.
3) Forms of applied nutrient: The chemical forms in which the nutrients are supplied are
also important. For example, Mg has been found to be absorbed more efficiently by apple
leaves from MgNO3 than from MgCl2.
4) Environmental factors: There are environmental factors like light, temperature and
relative humidity which directly and indirectly affect foliar absorption. The temperature
and relative humidity affect the uptake by providing a thin layer of moisture on the leaf
surface through transpiration.
Fertilizer Materials:

Not all fertilizers are suitable for use as a foliar spray. Qualifications for fertilizer materials
should have high solubility, low salt index and high purity.

Types of fertilizer material

1) Nitrogen Materials: Urea is the most suitable nitrogen source for foliar applications,
due to its low salt index and high solubility in comparison to other nitrogen sources.
IFFCO nano urea is becoming popular as a nitrogen source.
2) Phosphorus Materials: A combination of poly and ortho-phosphates shown to lessen
leaf burn and aid in leaf phosphate absorption. The polyphosphate advantage may also be
due to supplying both ortho and polyphosphate forms.
3) Potassium Materials: Depending on availability, potassium polyphosphates are an
excellent source of low salt index, highly soluble potassium. Potassium sulfate has low
salt index, but a rather low solubility.
4) Secondary and Micronutrient Materials: Foliar application of secondary nutrients can
be highly effective, but because of difficulties associated with leaf tissue absorption and
translocation choosing the correct fertilizer sources for these nutrients becomes very
critical