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All organisms require macromolecules (carbohydrates, proteins, fats etc.), water & minerals for growth and development.

METHODS TO STUDY THE MINERAL REQUIREMENTS OF PLANTS

- The technique of growing plants in a defined nutrient
solution (without soil) is known as hydroponics.
- It is demonstrated by Julius von Sachs (Germany, 1860). Hydroponics is used
- The nutrient solution is aerated for optimum growth. for commercial
- Hydroponics is used to identify the essential elements production of
required for plants and their deficiency symptoms. vegetables such as
- In this, plant roots are immersed in nutrient solutions and tomato, seedless
an element is added / removed or given in varied cucumber & lettuce.
concentration.

ESSENTIAL MINERAL ELEMENTS

- More than 60 elements are found in different plants. fixation). Zn2+ is an activator of alcohol dehydrogenase
- Some plant species growing near mining sites accumulate and Mo of nitrogenase during nitrogen metabolism.
selenium, gold etc. Some plants near nuclear test sites take 4. Elements that alter the osmotic potential of a cell: E.g.
up radioactive strontium. Potassium helps in opening & closing of stomata.
- There are techniques to detect the minerals even at a very
Role of Macro- and Micro-nutrients
low concentration (10-8 g/mL).
Essential elements have role in metabolic processes such as
Criteria for Essentiality of an element • Permeability of cell membrane.
• An element must be needed for normal growth and • Maintenance of osmotic concentration of cell sap.
reproduction. In the absence of the element the plants do • Electron transport systems.
not complete their life cycle or set the seeds. • Buffering action.
• The requirement of an element must be specific. i.e., • Enzymatic activity.
deficiency of an element cannot be met by another element. • Constituents of macromolecules and co-enzymes.
• It must be directly involved in the plant metabolism. Nitrogen:
• This is required by plants in the greatest amount.
Only 17 elements are absolutely essential for plant growth
• It is absorbed mainly as NO 3–. Some are also taken up as
and metabolism.
Based on quantitative requirements, essential elements are 2 NO2– or NH 4+.
types: Macronutrients & Micronutrients. • It is essential for all plant parts, particularly the
meristematic tissues and the metabolically active cells.
i. Macronutrients
• It is the major constituents of amino acids, proteins,
- They are present in plant tissues in large amounts (more nucleic acids, chlorophyll, vitamins and hormones.
than 10 mmole Kg–1 of dry matter).
Phosphorus:
- They include carbon, hydrogen, oxygen, nitrogen,
• It is absorbed by plants from soil as phosphate ions (as
phosphorous, sulphur, potassium, calcium & magnesium. H2PO4− or HPO42−).
- C, H & O are mainly obtained from CO2 & H2O. Others
• It is a constituent of cell membranes, certain proteins, all
are absorbed from soil as mineral nutrition.
nucleic acids and nucleotides.
ii. Micronutrients (trace elements) • It is required for all phosphorylation reactions.
- They are needed in very small amounts (less than 10 Potassium:
mmole Kg–1 of dry matter). • It is absorbed as potassium ion (K+).
- They include iron, manganese, copper, molybdenum, zinc, • Essential in meristematic tissues, buds, leaves & root tips.
boron, chlorine and nickel. • It maintains an anion-cation balance in cells.
Higher plants also need sodium, silicon, cobalt, selenium etc. • It is involved in protein synthesis, opening & closing of
Based on functions, essential elements are 4 categories: stomata, activation of enzymes and in the maintenance of
the turgidity of cells.
1. Components of biomolecules & structural elements of
cells: E.g. carbon, hydrogen, oxygen & nitrogen. Calcium:
2. Components of energy-related chemical compounds: • It is absorbed from the soil as calcium ions (Ca2+).
E.g. Mg in chlorophyll and phosphorous in ATP. • It is required by meristematic and differentiating tissues.
3. Elements that activate or inhibit enzymes: E.g. Mg2+ is • During cell division, it is used in the synthesis of cell wall,
an activator for RUBISCO & phosphoenol pyruvate particularly as calcium pectate in middle lamella. It is also
carboxylase (critical enzymes in photosynthetic carbon needed during the formation of mitotic spindle.

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• It accumulates in older leaves. Deficiency Symptoms of Essential Elements

• It is involved in the functioning of the cell membranes. - Deficiency of an essential element causes retarded growth.
• It activates some enzymes and regulates metabolic activities. - Concentration of an essential element below which plant
Magnesium: growth is retarded is called critical concentration. The
• It is absorbed by plants as divalent Mg2+. element is said to be deficient when present below the
• It activates enzymes of respiration & photosynthesis. critical concentration.
• It is involved in the synthesis of DNA and RNA. - The morphological changes due to deficiency or absence of
• It is a constituent of the ring structure of chlorophyll. an element are called deficiency symptoms.
• It helps to maintain the ribosome structure. - Deficiency symptoms vary from element to element.
Sulphur: - The plant parts that show the deficiency symptoms depend
• Plants obtain it as sulphate (SO)42−. on the mobility of the element. For elements that are
• It is present in 2 amino acids (cysteine & methionine). actively mobilized and exported to young developing
• It is the constituent of several coenzymes, vitamins tissues, the deficiency symptoms appear first in the older
(thiamine, biotin, Coenzyme A) and ferredoxin. tissues. E.g. deficiency symptoms of nitrogen, potassium
Iron: and magnesium are visible first in the senescent leaves.
• Plants obtain iron as ferric ions (Fe3+). - In older leaves, biomolecules containing these elements are
• It is required in larger amounts in comparison to other broken down. It makes these elements available for
micronutrients. mobilizing to younger leaves.
• It is a main constituent of proteins involved in the transfer - If the elements are relatively immobile and are not
of electrons like ferredoxin and cytochromes. transported out of the mature organs, the deficiency
• It is reversibly oxidized from Fe2+ to Fe3+ during electron symptoms appear first in the young tissues. E.g. S and Ca
transfer. are part of the structural component of the cell and hence
• It activates catalase enzyme, and is essential for the are not easily released.
formation of chlorophyll. - This aspect of mineral nutrition has great significance in
Manganese: agriculture and horticulture.
• It is absorbed as manganous ions (Mn2+). - The deficiency symptoms include chlorosis, necrosis,
• It activates many enzymes involved in photosynthesis, stunted growth, premature fall of leaves & buds and
respiration and nitrogen metabolism. inhibition of cell division.
• The best defined function of manganese is in the splitting - Chlorosis is the loss of chlorophyll leading to yellowing in
of water to liberate O2 during photosynthesis. leaves. It is due to the deficiency of elements N, K, Mg, S,
Fe, Mn, Zn and Mo.
Zinc:
- Necrosis is the death of tissue, particularly leaf tissue. It is
• Plants obtain zinc as Zn2+ ions.
due to the deficiency of Ca, Mg, Cu, K.
• It activates various enzymes, especially carboxylases.
- Lack or low level of N, K, S & Mo inhibits cell division.
• It is needed in the synthesis of auxin.
Low concentration of N, S, Mo etc. delay flowering.
Copper:
- Deficiency of different elements may cause same
• It is absorbed as cupric ions (Cu2+).
symptoms. Hence, to identify the deficient element, all the
• It is essential for the overall metabolism in plants.
symptoms are studied. Also, different plants respond
• Like iron, it is associated with some enzymes in redox
differently to the deficiency of the same element.
reactions and is reversibly oxidised from Cu+ to Cu2+.
Boron: Toxicity of Micronutrients
• It is absorbed as BO3 3− or B4O72−. - A moderate increase in micronutrients causes toxicity.
• It is required for uptake & utilisation of Ca2+, membrane - Any mineral ion concentration in tissues that reduces the
functioning, pollen germination, cell elongation, cell dry weight of tissues by about 10% is considered toxic.
differentiation & carbohydrate translocation. Such critical concentrations vary widely among different
Molybdenum: micronutrients.
• Plants obtain it as molybdate ions (MoO)22+.
- The toxicity symptoms are difficult to identify. Toxicity
• It is a component of many enzymes such as nitrogenase
levels for an element also vary for different plants.
& nitrate reductase. These enzymes participate in - Excess of an element may inhibit the uptake of another
nitrogen metabolism. element. E.g. Excess of Mn induces deficiencies of Fe, Mg
& Ca because it competes with Fe & Mg for uptake and
Chlorine:
with Mg for binding with enzymes. Mn also inhibits Ca
• It is absorbed as chloride anion (Cl–).
translocation in shoot apex. Thus symptoms of Mn toxicity
• Along with Na+ & K+, it helps in determining the solute
may actually be the deficiency symptoms of Fe, Mg & Ca.
concentration and the anion-cation balance in cells.
Main symptom of manganese toxicity is the appearance of
• It is essential for the water-splitting reaction in
brown spots surrounded by chlorotic veins.
photosynthesis that leads to oxygen evolution.
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MECHANISM OF ABSORPTION OF ELEMENTS

- The inward movement of ions into the cells is called influx in it. Use of radioisotopes of mineral elements also proved
and the outward movement is efflux. that they are transported through the xylem.
- The process of absorption includes 2 main phases:
Soil as Reservoir of Essential Elements
o First phase: Initial rapid and passive uptake of ions into
Weathering and breakdown of rocks enrich the soil with
the apoplast (free space or outer space) of cells. It
dissolved ions and inorganic salts.
usually occurs through ion-channels (trans-membrane
proteins that function as selective pores). Roles of soil:
o Second phase: The ions are taken in slowly into the o It supplies minerals and holds water.
symplast (inner space or cytoplasm) of the cells. It is an o It harbours nitrogen-fixing bacteria and other microbes.
active process (requires energy). o It supplies air to the roots.
o It acts as a matrix that stabilizes the plant.
Translocation of solutes
Deficiency of essential minerals affects the crop-yield. So
- Mineral salts are translocated through xylem along with the
fertilisers should be supplied. Both macro-nutrients and
ascending stream of water.
micro-nutrients form components of fertilisers.
- Analysis of xylem sap shows the presence of mineral salts

METABOLISM OF NITROGEN
Nitrogen Cycle - Nitrate present in the soil is also reduced to nitrogen by the
- Nitrogen is the most prevalent element in living organisms. process of denitrification. It is carried by bacteria
- Plants compete with microbes for the limited nitrogen in Pseudomonas and Thiobacillus.
soil. Thus, nitrogen is a limiting nutrient for natural and Biological Nitrogen Fixation
agricultural eco-systems. - It is the reduction of N2 to NH3 by living organisms in
- The process of conversion of atmospheric nitrogen (N2 or presence of nitrogenase enzyme.
NºN) to ammonia is called nitrogen fixation.
- In nature, lightning and UV radiation provide energy to
- Very few organisms can utilize the nitrogen in the form of
convert nitrogen to nitrogen oxides (NO, NO2, N2O).
N2 in the air.
Industrial combustions, forest fires, automobile exhausts
- Only certain prokaryotic species have Nitrogenase enzyme
and power-generating stations are also sources of
and capability to fix N2. They are called N2- fixers.
atmospheric nitrogen oxides.
- Nitrogen-fixing microbes are 2 types:
- Decomposition of organic nitrogen of dead plants and
o Free-living: E.g. Azotobacter & Beijerinckia (aerobic
animals into ammonia is called ammonification.
microbes), Rhodospirillum & Bacillus (anaerobic),
- Some of this ammonia volatilizes and re-enters atmosphere
cyanobacteria such as Anabaena & Nostoc.
but most of it is oxidised into nitrate by soil nitrifying
o Symbiotic: E.g. Rhizobium (aerobic).
bacteria (Nitrosomonas, Nitrococcus & Nitrobacter-
chemo-autotrophs). These steps are called nitrification. Symbiotic Biological Nitrogen Fixation
2NH3 + 3O2 Nitrosomonas, Nitrococcus 2NO - + 2H
2
+
+ 2H2O - Legume-bacteria relationship: Most prominent. E.g.
Nitrobacter Rhizobium species (rod-shaped) seen in the roots of
2NO2- + O2 2NO3- legumes such as alfalfa, sweet clover, sweet pea, lentils,
- Plants absorb the nitrate and is transported to the leaves. In garden pea, broad bean, clover beans etc.
leaves, it is reduced to form ammonia that finally forms the - The most common association on roots is as nodules.
amine group of amino acids. - The microbe, Frankia also produces N2 fixing nodules on
the roots of non-leguminous plants (e.g. Alnus).
- Rhizobium & Frankia are free-living in soil, but as
symbionts, can fix atmospheric nitrogen.
- Central part of a nodule is red or pink coloured due to the
presence of leguminous haemoglobin (leg-haemoglobin).
Principal stages in the Nodule formation:
o Rhizobia multiply and colonise the surroundings of the
roots and get attached to epidermal and root hair cells.
o Root-hairs curl and the bacteria invade the root-hair.
o An infection thread is produced carrying the bacteria into
root cortex, where they initiate nodule formation.
o The bacteria are released from thread into cells. It leads to
differentiation of specialized nitrogen fixing cells.

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o The nodule establishes a direct vascular connection with - Rhizobia are aerobic under free-living conditions (where
the host for exchange of nutrients. nitrogenase is not operational), but during N2-fixing
events, they become anaerobic (to protect nitrogenase).
Fate of ammonia:
- At physiological pH, the NH3 is protonated to form NH4 +
(ammonium) ion. Most of the plants can assimilate nitrate
and NH4+. But NH4+ is quite toxic to plants and so cannot
accumulate in them.
- In plants, NH 4+ is used to synthesize amino acids by 2 ways:
a. Reductive amination: In this, ammonia reacts with a-
ketoglutaric acid to form glutamic acid.
a- ketoglutaric acid + NH4+ + NADPH
Glutamate dehydrogenase
Glutamate + H2O + NADP
b. Transamination: It is the transfer of amino group (NH2)
from one amino acid to the keto group of a keto acid in
presence of transaminase enzyme. Glutamic acid is the
- Nodule contains nitrogenase enzyme & leg-haemoglobin. main amino acid from which the transfer of NH2 takes
- Nitrogenase (a Mo-Fe protein) catalyzes the conversion of place and other amino acids are formed through
nitrogen to NH3, the first stable product of N2 fixation. transamination. For example,

- Asparagine & glutamine are most important amides

found in plants. They are structural part of proteins. They
are formed from 2 amino acids (aspartic acid & glutamic
acid) by addition of another amino group to each. The
N + 8e- + 8H+ + 16 ATP → 2NH + H + 16ADP + 16P hydroxyl part of the acid is replaced by another NH2- radical.
i
2 3 2 - Since amides contain more nitrogen than the amino acids,
- Ammonia synthesis needs high input energy (8 ATP for they are transported to other parts of the plant via xylem
each NH3). It is obtained from the respiration of host cells. vessels. In addition, along with the transpiration stream the
- Nitrogenase is highly sensitive to the molecular oxygen. So nodules of some plants (e.g. soyabean) export the fixed
it requires anaerobic conditions to protect from oxygen. For nitrogen as ureides. These compounds also have
this, leg-haemoglobin acts as an oxygen scavenger. particularly high nitrogen to carbon ratio.