MODULE LESSON 4

MINERAL NUTRITION OF PLANTS:

Functions and Deficiency Symptoms of Nutrients, Nutrient
Uptake Mechanisms

Mineral nutrients are elements acquired primarily in the form of

inorganic ions from the soil. Although mineral nutrients continually cycle
through all organisms, they enter the biosphere predominantly through
the root systems of plants, so in a sense plants act as the "miners" of
Earth's crust. The large surface area of roots and their ability to absorb
inorganic ions at low concentrations from the soil solution make mineral
absorption by plants a very effective process. After being absorbed by the
roots, the mineral elements are translocated to the various parts of the
plant, where they are utilized in numerous biological functions. Other
organisms, such as mycorrhizal fungi and nitrogen-fixing bacteria, often
participate with mots in the acquisition of nutrients.
The study of how plants obtain and use mineral nutrients is called
mineral nutrition.

A. Intended Learning Objectives

B. Learning Content

I. Mineral nutrition
• The chemical compounds required by an organism are termed as
nutrients
• Nutrition may be defined as the supply and absorption of
chemical compounds needed for plant growth and metabolism
• For plant growth and metabolism, 17 elements are essential. They
are C, H, O, N, P, K, Ca, S, Mg, Fe, Mn, Zn, B, Cu, Mo, Cl and Ni.

These essential elements are classified as:

1. Non-Mineral elements
2. Macronutrients (Macro elements)
Sub-classification of Macronutrients:
a. Primary
b. Secondary

2. Micronutrients (Trace elements)

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Non-mineral nutrients:
These elements are required by the plants for its photosynthetic
processes. They are: C, H and O.

Macronutrients:
The essential elements which are required by the plants in
comparatively v e r y large amounts are called as primary macro
elements or macro nutrients (N, P, and K). While secondary
macronutrients are required by plants in large amounts (Ca,
S, and Mg). According to another definition minerals found in
>1000 ppm concentration are macronutrients.

Micronutrients:
The essential elements which are required in very small
amounts or traces by the plants are called as micronutrients or
trace elements. According to another definition minerals found in
<100 ppm concentration are micronutrients. They are Fe, Zn, Mn,
B, Cu, Cl and Mo.

Other Beneficial elements :

Silicon (Si) is now transferred from list of beneficial elements
to essential elements. Sodium has beneficial effect, and, in some
case, it is essential. There are some plant species, particularly the
Chenopodiaceae plants and species adapted to saline conditions
that take up this element in relatively high amounts. Na is also
required for turnips, sugar beets and celery. The same is true for
Si, which is an essential nutrient for rice. Cobalt is an essential
element for the growth of the Bluegreen algae, but it has not been
shown to be essential for other algae or for higher plants. It is also
required by certain legumes to fix atmospheric nitrogen. Here,
however the cobalt ion is necessary for the symbiotic bacteria
present in the nodules associated with the roots.

Criteria of essentiality:
The term essential nutrient element was proposed by Arnon and
Stout (1939). According to them an element to be considered essential,
three criteria must be met:
1. A given plant must be unable to complete its life cycle in the
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absence of mineral elements.
2. The function of the element must not be replaceable by another mineral
element.
3. The elements must be directly involved in plant metabolism. For eg.
as a component of an essential plant constituents or it must be required
for a distinct metabolic step such as an enzyme reaction.

Based on the mobility, elements are also classified into three types.
1. Mobile elements : N, P, K, S and Mg
2. Immobile elements : Ca, Fe and B
3. Intermediate in mobility : Zn, Mn, Cu, Mo

II. Classification of plant nutrients based on their b i o c h e m i c a l

r o l e a n d p h y s i o l o g i c a l function:
Essential elements are now classified according to their
biochemical role and physiological function. Based on the
biochemical behavior and physiological functions, plant nutrients
may be divided into four groups.

Nutrient
Uptake Biochemical function
element
s In the form of CO2, HCO- Major constituents of the
1st
group C, 3, H2O, O2, NO3, NH+, organic compounds of the
H, O, N, S plant. Essential elements of
4N2SO42, SO2. The ions atomic groups which are
from the soil solution, the involved in enzymatic
gases from the processes. Assimilation by
atmosphere.
oxidation reduction reactions.

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 2nd In the form of They are important in energy
Group phosphates, boric storage reactions or in maintaining
P, B, Si acid or borate, silicate
from the soil solution. structural integrity. Elements in this
group are often present in plant
tissues as phosphate, Borate and
silicate esters in which the
elemental group is bound to the
hydroxyl group of an organic
molecule (i.e. sugar- phosphates)

(Esterification*). The phosphate

esters are involved in energy transfer
reactions.

3rd In the form of cations Present in plant tissues as either

free ions or ions bound to
Group from the soil solution
substances such as the pectic
K, Na, except chlorine acids present in the plant cell wall.
Mg, Ca, Of particular importance of their
Mn, Cl roles as enzyme cofactors and in
regulation of osmotic potentials.

4th In the form of Present predominantly in a chelated

Group form Incorporated in prosthetic
ions or chelates
Fe, Cu, groups. Enable electron transport by
Zn, Mo from the soil valency change.
solution

(Source: Taiz and Zeiger 2002)

*Esterification: Compounds formed by condensation of an
acid and alcohol with elimination of water ADP + Pi = ATP

Nutrient deficiency visible symptoms :

When minerals are deficient, the growth of the plant is
stunted, or the plant shows other symptoms. The combination of
symptoms observed for deficiency of a particular mineral can be
traced to the roles that mineral plays in metabolism or physiology.
1. Stunted growth is a symptom for many deficiencies, especially
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stunted stems with nitrogen deficiency and stunted roots in
phosphorus deficiency.
2. Chlorosis decreased chlorophyll synthesis or increased
chlorophyll degradation, is observed with magnesium, nitrogen,
and iron deficiencies. Magnesium is the central atom for the
electron cloud of chlorophyll from which electrons flow through the
light reactions.
3. Necrosis, dead spots or zones, is observed when magnesium,
potassium or manganese deficiencies are present.
4. Colour changes such as excessive anthocyanin production is
observed in stems with phosphorus deficiency. They generally
pick up an intense purple colour sometimes extending onto the
leaves.

Nutrient availability shows an interesting dose effect:

The following graph demonstrates how deficiency reduces growth.

As the mineral availability is increased, growth increases. As the mineral
content continues to be increased there is no further increase in growth,
but quality may be continuing to increase. This zone is called the luxury
zone. However, continuing to increase the mineral concentration
ultimately reaches toxic levels and growth is diminished.

The goal of a plant grower is to keep the plant in the sufficient to

luxury zone but never to get as low as deficiency or as high as toxicity
for any one of the macronutrients or micronutrients. The trouble with
that goal knows how wide the luxury zone is in terms of concentrations.
For minerals like boron, the zone is very narrow, and it is easy to achieve
toxic levels or to be in deficiency. For minerals like phosphorus,44 the
luxury zone is quite broad and large amounts can be given and the plants
will respond nicely in spite of that. As a result, it is difficult to overdose
plants on phosphorus.

Specific roles of essential mineral elements :

A. The macronutrients :

1. Nitrogen specific role :

• Nitrogen is important constituent of proteins, nucleic acids,
porphyries (chlorophylls & cytochromes) alkaloids, some
vitamins, coenzymes etc.
• Thus N plays very important role in metabolism, growth,
reproduction and heredity. Deficiency symptoms
• Plant growth is stunted because protein content cell
division and cell enlargement are decreased
• N deficiency causes chlorosis of the leave i.e yellowing older
leaves are affected first
• In many plants eg. tomato, the stem, petiole and the leaf
veins become purple coloured due to the formation of
anthocyanin pigments.
2. Phosphorus :
• It is important constituent of nucleic acids,
phospholipids, coenzymes NADP, NADPH2 and ATP
• Phospholipids along with proteins may be important constituents
of cell membranes
• P plays important role in protein synthesis through nucleic acids
and ATP
• Through coenzymes NAD, NADP and ATP, it plays important
role in energy transfer reactions of cell metabolism eg.
photosynthesis, respiration and fat metabolism etc.

Deficiency symptoms
• P deficiency may cause premature leaf fall
• Dead necrotic areas are developed on leave or fruits
• Leaves may turn to dark green to blue green colour.
Sometimes turn to purplish colour due to the synthesis and
accumulation of anthocyanin pigments.
3. Potassium Specific role :
• Although potassium is not a constituent of important organic
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 compound in the cell, it is essential for the process of
respiration and photosynthesis
• It acts as an activator of many enzymes involved in
carbohydrate metabolism and protein synthesis
• It regulates stomatal movement
• Regulates water balance

Deficiency symptoms
• Mottled chlorosis of leaves occurs
• Neurotic areas develop at the tip and margins of the leaf
• Plants growth remains stunted with shortening of internodes.

4. Calcium :
• It is important constituent of cell wall
• It is essential in the formation of cell membranes
• It helps to stabilize the structure of chromosome
• It may be an activation of may enzymes

Deficiency symptoms
• Calcium deficiency causes disintegration of growing meristematic
regions of root, stem and leaves
• Chlorosis occurs along the margins of the younger leaves
• Malformation of young leaves takes place

5. Magnesium :
• It is very important constituent of chlorophylls
• It acts as activation of many enzymes in nucleic acid
synthesis and carbohydrate metabolism
• It plays important role in binding ribosomal particles during
protein synthesis.

Deficiency symptoms
• Mg deficiency causes mottled chlorosis with veins green and leaf
tissues yellow or white appearing first on older leaves
• Dead neurotic patches appear on the leaves
• In cotton Mg deficiency leads o reddening of leaves and disorder is
called as reddening in cotton. 46
6. Sulphur specific role :
• It is important constituent of some amino acids (cystine,
cysteine and methionine) with which other amino acids form
the protein
• S helps to stabilize the protein structure
• It is also important constituent of vitamin i.e biotin, thiamine
and coenzyme A
• Sulphydryl groups are necessary for the activity of many
enzymes.

Deficiency symptoms
• Deficiency causes chlorosis of the leaves
• Tips and margins of the leaf roll in ward
• Stem becomes hard due to the development of sclerenchyma.

B. Micronutrients:

1. Iron specific role :

• Important constituent of iron porphyrin proteins like
cytochromes, peroxidanes, catalases, etc.
• It is essential for chlorophyll synthesis
• It is very important constituent of ferredoxin which plays
important role in photochemical reaction in photosynthesis and
in biological nitrogen fixation.

Deficiency symptoms
• Iron deficiency causes chlorosis of young leaves which is usually
interveinal.
2. Zinc specific role :
• It is involved in the biosynthesis of growth hormone auxin (indole
3 acetic acid)
• It acts activator of many enzymes like carbonic anhydrase and
alcohol dehydrogenase, etc.

Deficiency symptoms
• Zinc deficiency causes chlorosis of the young leaves which47starts
 from tips and the margins
• The size of the young leaves is very much reduced. This disorder is
called as ‘little leaf disease’
• Stalks will be very short.

3. Manganese :
• It is an activator of many respiratory enzymes
• It is also an activator of the enzyme nitrite reductase
• It is necessary for the evolution of oxygen (photolysis) during
photosynthesis

Deficiency symptoms
• The young leaves are affected by mottled chlorosis
• Veins remain green
• Small necrotic spots developed on the leaves with yellow strips

4. Copper specific role :

• It is an important constituent of plastocyanin (copper containing
protein)
• It is also a constituent of several oxidizing enzymes. Deficiency
symptoms
• Copper deficiency causes necrosis of the tip of the young leaves
• It also causes die-back of citrus and fruit trees
• Also causes reclamation disease or white tip disease of cereals
and leguminous plants.
5. Boron specific role :
• Boron facilitates the translocation of sugars by forming sugar
borate complex.
• It involves in cell differentiation and development since boron is
essential for DNA synthesis
• Also involves in fertilization, hormone metabolism etc.

Deficiency symptoms
• Boron deficiency causes death of shoot tip
• Flower formation is suppressed
• Root growth is stunted
• The other diseases caused by B deficiency is Heart rot of beet,
Stem crack of celery, Brown heart of cabbage, Water core 48
of turnip,
 Internal cork formation in apple, Hen and chicken in grapes.
6. Molybdeneum :
• It is constituent of the enzyme nitrate reductase and thus plays
an important role in nitrogen metabolism
• It is essential for flower formation and fruit set.

Deficiency symptoms
• Molybdenum deficiency causes interveinal chlorosis of older
leaves
• Flower formation is inhibited
• Causes whiptail disease in cauliflower plants.

7. Chlorine : Specific role

• Chlorine has been shown to be involved in the oxygen evolution
in photosystem II in photosynthesis (Cl and Mn are important for
this reaction)
• It raises the cell osmotic pressure
• Chlorine accelerates the activation of amylase which converts
starch into soluble sugars

Deficiency symptoms
• Chlorosis of younger leaves and an overall wilting of the plant
• In some plant species, like tomato, leaves show chlorotic mottling,
bronzing and tissue necrosis

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III. Physiology of nutrient uptake :
Mineral nutrients are found either as soluble fractions of soil solution
or as adsorbed ions on the surface of colloidal particles. Various
theories proposed to explain the mechanism of mineral salt absorption
can be placed in two broad categories:
I) Passive Absorption
II) Active Absorption
Ion uptake is both active and passive :
After several decades of research on this process of ion uptake it is now
believed that the process involves both passive and active uptake
mechanisms.
Whether a molecule or ion is transported actively or passively across
a membrane (casparian band, plasma membrane or tonoplast) depends
on the concentration and charge of the ion or molecule, which in
combination represent the electrochemical driving force.
Passive transport across the plasma membrane, occurs
along with the electrochemical potential. In this process ions and
molecules diffuse from areas of high to low concentrations. It does not
require the plant to expend energy.
Active transport, (in contrast, to passive transport) energy is
required for ions diffusion against the concentration gradient (electro
chemical potential). Thus, active transport requires the cell to expend
energy.

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 Passive transport mechanism:
A) Diffusion: Simple diffusion to membranes occurs with small, non-polar
molecules (i.e. O2, CO2). In this process ions or molecules move from
the place of higher concentration to lower concentration. It needs no
energy.
B) Facilitated diffusion: For small polar species (i.e. H2 O, Ions and amino
acids) specific proteins in the membrane facilitate the diffusion down the
electrochemical gradient. This mechanism is referred to as facilitated
diffusion. Eg.
a) Channel proteins: The specific proteins in the membrane form channels
(channel proteins), which can open and close, and through which ions
or H 2O molecules pass in single file at very rapid rates. A K+ and NH4+
channel also operates by the same process of facilitated diffusion. In
addition, Na+ can also enter the cell by this process.
b) Transporters or Co-transporters or carriers: Another mechanism involves
transporters or co-transporters responsible for the transport of ions and
molecules across membranes. Transporter proteins, in contrast to
channel proteins, bind only one or a few substrate molecules at a time.
After binding a molecule or ion, the transporter undergoes a structural
change specific to a specific ion or molecule. As a result, the transport
rate across a membrane is slower than that associated with channel
proteins.
Three types of transporters have been identified:
1. Uniporters: transport one molecule (i.e. glucose, amino acids) at a time
down a concentration gradient.
2. Antiproters: catalyze movement of one type of ion or molecule against its
concentration gradient. This is coupled with the movement of a different
ion or molecule in the opposite direction. Examples of antiporter
transport are H + /Na+ and H+ /Ca+2 transport into the vacuole.
3. Symporters: catalyze movement of one type of ion or molecule against its
concentration gradient coupled to movement of a different ion or molecule
down its concentration gradient in the same direction. The high H +
concentration in the apoplast provides the energy for symporter transport
of NO3- and the other anions.
Therefore, the energy for antiporter and symporter transport
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originates from the electric potential and/or chemical gradient of a
secondary ion or molecule, which is often H+.

Active transport mechanism: Larger or more-charged molecules have

great difficulty in moving across a membrane, requiring active transport
mechanisms (i.e., sugars, amino acids, DNA, ATP, ions, phosphate,
proteins, etc.). Active transport across a selectively

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permeable membrane occurs through ATP-powered pumps that transport
ions against their concentration gradients. This mechanism utilizes
energy released by hydrolysis of ATP. The Na+-K+ ATP pump transports
K+ into the cell and Na+ out of the cell, another example is the Ca+2 -ATP
pump.
Thus, it can be understood from the above discussion that the
ion transport mechanisms operate both actively and passively. For
some of the ions the uptake mechanism is active and for some others
it is passive.

Intended Learning Assignment/activity

The assignment and activity to improve the learnings on plant

nutrition will be posted in your Google Classroom.

Reference: Lincoln Taiz and Eduardo Zeiger 2006, Plant Physiology,

Sinauer Associates, Inc. Publishers, Sunderland, Massachusetts.

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