CHAPTER 7:

Mineral nutrition and ion transport

Section "A"
Introduction
The supply and absorption of specific chemical compounds needed for normal growth and
metabolism of plants is defined as nutrition while the chemical compounds that functions as
raw material for synthesis of different structural and functional substance of plants are
termed nutrients . The inorganic material obtained from soil which are used as raw material
by plants are called mineral nutrients. Absorption utilization and assimilation of inorganic
compound or minerals by plants for synthesis of essential material for their growth,
development, structure and physiology is called mineral nutrition.
Essential elements The mineral nutritional elements are available to the plant as ion present
in soil water absorbed through root. Nutrient elements may occur in the soil (a) in the aqueous
solution, (b) adsorbed on organic or inorganic soil colloids, (c) in the form of an insoluble
inorganic compound, and (d) as a constituent of organic compounds, either as a residue of
plants or animals or in living organisms. The uptake of nutrients by the plant roots is closely
related to the form in which the elements occur. Many factors influence nutrient uptake for
plants. Ions can be readily available to roots or could be "tied up" by other elements or the
soil itself. Soil too high in pH (alkaline) or too low (acid) makes minerals unavailable to
plants.
Liebig (1840) proposed the law of minimum which states that productivity of a soil is
dependent upon the proportionate occurrence of deficient mineral.
Julius von Sachs( 1860), German botanist, demonstrated, for the first time, that plants could
be grown from seedling to maturity in a defined nutrient solution in complete absence of soil.
This technique of growing plants in a nutrient solution is known as hydroponics.
Arnon and Stout (1939) physiologists have given three criteria for plant nutrient
essentiality :1. Some elements are essential for supporting normal growth and reproduction. In the
absence of the element the plants do not complete their life cycle or set the seeds.
2. The requirement of the element must be specific and the element cannot be replaced by
the presence of another element. In other words, deficiency of any one element cannot be
met by supplying some other element.
3. The element must be directly involved in the metabolism of the plant.
Two criteria by Epstein, 2005:-

i.

The element is part of a molecule that is an intrinsic component of the structure or

metabolism of a plant.

ii.

The plant can be so severely deprived of element that it exhibits abnormalities in its
growth, development or reproduction - that is, its performance - in comparison with
plants not so deprived.

For example, Magnesium is a constituent of the chlorophyll molecule and is essential for
photosynthesis, so it cannot be replaced by any other element for the same function . It is also
required as a cofactor by many enzymes involved in cellular respiration and metabolic
path ways.
Sixteen chemical elements are important in which carbon, hydrogen, and oxygen are
supplied by air and water, which comprise about 98% of the fresh weight of the plant, at least
13 other chemical elements, called the essential inorganic nutrients, are needed for plant's
growth and survival. The sixteen chemical elements are divided into two main groups:a. Non-Mineral Nutrients :-The Non-Mineral Nutrients are hydrogen (H), oxygen (O),
and carbon (C). In a process called photosynthesis, plants use energy from the sun to
change carbon dioxide (CO2 - carbon and oxygen) and water (H2O- hydrogen and
oxygen) into starches and sugars.
b. Mineral 13 other chemical elements,which come from the soil, are dissolved in water
and absorbed through a plant's roots called the essential inorganic nutrients, are needed
for plant growth.
On the basis of average concentration in plants , Hoagland (1944) divided essential elements
into two categories:1. Macronutrients: -Macronutrients are generally present in plant tissues in large
concentrations of 1.0-10.0 mg per gram of dry matter. The macronutrients
include Carbon (C) , hydrogen (H), oxygen(O), nitrogen,(N) phosphorous(P),
sulphur(S), potassium (K), Calcium (Ca)and magnesium(Mg).
2. Nitrogen, Potassium, and Phosphorous are obtained from the soil and are the primary
macronutrients. Calcium, Magnesium, and Sulphur are the secondary
macronutrients needed in lesser quantity.
3. Micronutrients or trace elements:-Micronutrients are present in very small amounts in
plant body in concentration of equal or less than 0.1 mg per gram of dry matter. These
include iron (Fe), manganese (Mn) , Copper (Cu), molybdenum (Mo), zinc (Zn),
boron, (B) chlorine (Cl) .Many of them are components or cofactors of
enzymes. Some are essential for electron transfer.
There are some beneficial elements such as sodium, silicon, cobalt and selenium, in addition
to the 16 essential elements named above They are required by higher plants.

Essential elements can also be grouped into following types on the basis of their diverse
functions :i.

Framework elements:- Carbon (C ), Hydrogen (H), Oxygen (O) synthesize the

cellulose and other components of cell walls. Calcium (Ca) is component of middle
lamella.

ii.

Protoplasmic elements :- Carbon (C) , hydrogen (H), oxygen(O), nitrogen,(N)

phosphorous(P), sulphur(S), iron (Fe)and magnesium(Mg) give rise to protoplasmic
constituents like proteins, nucleic acids, hormones, vitamins, phospholipids,,
photosynthetic pigments, etc.

iii.

Osmotic potential :- Some essential elements, e. g., nitrate, potassium, sulphate etc. can
alter the osmotic potential of a cell. Potassium plays an important role in the opening and
closing of stomata and turgor movements in various plants.

iv.

Energy transducers :-Magnesium occurs in chlorophyll and phosphorous in ATP are

essential elements that are components of energy-related chemical compounds in plants,
converting one form of energy into another.

v.

Enzymatic effects:- Some elements that activate or inhibit enzymes, e.g.

K,Ca,Mg,Zn,Mn,Cl,Ni,Cu, etc. Mg2+ is an activator for both ribulose bisphosphate
carboxylaseoxygenase and phosphoenol pyruvate carboxylase, both of which are critical
enzymes in photosynthetic carbon fixation, Zn2+ is an activator of alcohol
dehydrogenase and Mo of nitrogenase during nitrogen metabolism.

vi.

Balancing elements :- Calcium , magnesium potassium counteract the toxic effect of

other minerals by ion balancing.

vii.

Fe, Mn, Cu, Zn and Mo are function as co- enzymes

Section "B"
Role of Macro- and Micro-nutrients and Their Deficiency symptoms
.

Name of
elements

Macronutrient
s

Element
obtained in the
form of

Regions of
plant where
element
required

Functions

Deficiency symptoms

Nitrogen is
required by all
parts of a
plant,
particularly the
meristematic
tissues and the
metabolically
active cells.

Nitrogen is one of
the major
constituents of
proteins, nucleic
acids, vitamins and
hormones,
coenzymes, and
nucleotides.Nitroge
n promotes rapid
growth, increases
leaf size and
quality, hastens
crop maturity, and
promotes fruit and
seed development.

Nitrogen (N)

Nitrogen can
exist in a soil
organic form, or
as an
ammonium ion
(NH4+)nitrite
ion (NO2-),or
nitrate ion
(NO3-).

Phosphorus
(P)

Phosphorus is a
constituent of cell
membranes, certain
Phosphate is
proteins, all nucleic
easily
acids and
redistributed in
nucleotides,
most plants
involved in energy
from one organ
transfer such as
to another and
ATP and ADP. It
Phosphorus is
is lost from
activates
absorbed by the older leaves,
coenzymes for
plants from soil accumulating
amino acid
in the form of
in younger
production used in
phosphate ions leaves,
protein
as H2PO4developing
synthesis;and it is
orthophosphate. flowers and
involved in many
seeds.The
other metabolic
meristem
processes required
region of
for normal growth,
growing plants
such as
is high in
photosynthesis,
phosphorus.
glycolysis,
respiration, and
fatty acid synthesis.

Potassium (K) It is absorbed as Required in

potassium ion
more abundant
(K+).
quantities in
the

Helps to determine
an anion-cation
balance in cells and
activates enzymes

Reduced growth,
yellowing (chlorosis),
reds and purples may
intensify with some
plants, reduced lateral
breaks.

Stunted growth, dark

green leaves with a
leathery texture, and
reddish purple leaf tips
and margins.Maturity is
often delayed.

Small plants, brown

margins on lower leaves,
small weak stems,
lodging of plants, poor

Magnesium
(Mg)

Calcium
(Ca2+)

to metabolize
carbohydrates for
the manufacture of
amino acids and
proteins, opening
and closing of
stomata, activation
meristematic
of enzymes and in
tissues, buds,
the maintenance of
leaves and root
the turgidity of
tips.
cells, facilitates cell
division and growth
by helping to move
starches and sugars
between plant
parts,disease
resistance.

yield and quality.As in N

and P, K+ is easily
redistributed from mature
to younger organs, so
symptoms first appear in
older leaves.

It is used for
fruit and nut
formation and
essential for
germination of
It is absorbed as
seeds. Leaves:
magnesium ion
withdrawn
(Mg2+).
from ageing
leaves and
exported to
developing
seeds

is a critical
structural
component of the
chlorophyll
molecule and is
necessary for
functioning of plant
enzymes to produce
carbohydrates,
sugars and fats.It is
an enzyme activator
in the synthesis of
nucleic acids (DNA
and RNA).

Extensive
interveinalchlorosis whic
h starts with basal leaves
and progresses to younger
leaves (it is mobile).

It is absorbed as
calcium ion
(Ca2+).Adsorbe
d Ca2+ is
important for
soil structure by
promoting the
aggregation of
soil particles.
This improves
water and root
penetration

Calcium is involved
in many plant
processes, including
cell elongation, cell
division,
germination, pollen
growth, activates
enzymes,and is a
structural
component of cell
walls, . One of its
most important

Meristematic
and
differentiating
tissues. It is
immobile
(nontranslocatable)
within plants
and remains in
the older tissue
throughout the
growing

Inhibition of bud growth,

death of root tips,
cupping of mature leaves,
weak growth.,

functions is the
through the soil
season.Calciu maintenance of
and maintains
m is easily
membrane
the stability of
leached.
permeability and
soil particles.
cell integrity.

Sulphur (S)

Stem and root

tips young
Sulfate (SO4=) leaves;
from the soil is remobilized
the primary
during
source of
senescence
S.Sulfur is also (the growth
taken up by
phase in a
leaves in
plant or plant
gaseous form as part -as a leaf
SO2.
from full
maturity to
death).

It is a structural
component of
amino acids,
proteins, vitamins
and enzymes and is
essential to produce
chlorophyll.

General chlorosis of leaf,

including vascular
bundles.Sulfur is not
easily mobilized within
the
plant.Therefore,chlorosis,
occur in young tissues
before older ones.

Most zinc in soil

is found in
different
minerals with
only a small
percentage
Everywhere
being adsorbed
in ionic form on
soil and organic
matter exchange
sites.

It activates various
enzymes especially
carboxylases, part
of carbonic
anhydrase and
various
dehydrogenases,
needed for auxin
synthesis.

Interveinal chlorosis of
the upper (youngest)
leaves. Afterwards, shoot
growth slows down,
giving the affected plant
parts a rosette-like
appearance.

Micronutrient

Zinc -Zn
(immobile)

Manganese
-Mn
(Immobile)

The most
Leaves and
important form seeds
for uptake by
roots is Mn++,
but it is also
commonly
found as oxides
of Mn+++ and
Mn ++++
(Mn2O3,

Manganese in the
plant participates in
several important
processes including
photosynthesis and
nitrogen and
carbohydrate
metabolism.

The absence of
Manganese causes
disorganization of
chloroplast thylakoid
membranes. Chlorosis

MnO2, etc.).

Iron, Fe
(Immobile)

Copper (Cu)

Important
constituent of
proteins involved in
Plants obtain
the transfer of
iron in the form
electrons like
of ferric ions
ferredoxin and
(Fe3+). It is
Everywhere , cytochromes,
required in
collects along reversibly oxidised
larger amounts leaf veins.
from Fe2+ to Fe3+
as compare to
during electron
other
transfer,activates
micronutrients.
catalase enzyme,
required for
synthesis of
chlorophyll.

It is absorbed as
cupric ions
Everywhere
(Cu2+).

Molybdenum Plants obtain it Everywhere,

(Mo)
in the form of
the synthesis
molybdate ions. of proteins is
blocked and
plant growth
ceases. Root
nodule
(nitrogen
fixing) bacteria
also require it.
Seeds may not

Extensive interveinal
chlorosis, starting with
younger leaves (iron is
relatively immobile).

Essential for the

overall metabolism
in plants.More than
half of the copper is
located in the
chloroplasts and
participates in
Dieback of shoots
photo-synthetic
reactions. It is also
found in other
enzymes involved
with protein and
carbohydrate
metabolism.
Essential
component of two
enzymes involved
with nitrogen
metabolism.

Pale green leaves with

rolled or cupped margins.

form
completely,
and nitrogen
deficiency may
occur if plants
are lacking
molybdenum.

Boron (B)

The soluble
form is
primarily boric
acid (B(OH)3).
In neutral to
acid soils, boric
acid has no
charge and can,
therefore, be
easily leached. Leaves and
At higher pH
seeds
values,
conversion to
B(OH)4- occurs.
The resulting
negative charge
on the molecule
causes its
absorption by
soil particles.

Chloride (Cl),
the ionic form of
chlorine used by
Chlorine (Cl) plants, is usually Everywhere
found in soluble
forms and is lost
by leaching..

Boron is required
for uptake and
utilisation of Ca2+,
membrane
functioning, pollen
germination, cell
elongation, cell
differentiation and
carbohydrate
translocation.

Terminal buds are

damaged, leaving a
rosette effect on the plant.
Leaves are thick, curled
and brittle. Fruits, tubers
and roots are discolored,
cracked and flecked with
brown spots.

Involved in osmosis
(movement of water
or solutes in cells),
the ionic balance
necessary for plants
to take up mineral
elements and in
photosynthesis.

Reduced growth; stubby

roots, interveinal
chlorosis, nonsucculent
tissue (in leafy
vegetables)

Hunger Signs:- When plants do not get sufficient amount of one or more essential nutrient
elements ,they show poor growth and develop specific deficiency symptoms Which is known
as hunger signs.
Section "C"

Ion transport The plant absorb minerals nutrient from the soil mostly by the roots but in
some plants by leaves((carnivorous plants, bromeliads, etc. ) in ionized form.Roots absorb
some mineral nutrients selectively , including some which may not be essential for them.
Plants absorb minerals from soil through the root, passes the membrane, in two ways: 1. Passive transport:-Movement of molecules from high to low concentration. Needs no
energy input. Including Passive diffusion and Facilitated diffusion
2. Active transport:- Movement of molecules from low to high concentration (opposite
the flow of diffusion). Needs input of energy (ATP).
Passive transport :- The main theories and concepts of passive transport are :1. Donann equilibrium theory (Donann 1911):- Entry of ions into the cell across the
plasma membrane to maintain electrical equilibrium is known as Donann equilibrium
2. Interception and contact or Ion exchange theory:- Jenny and Overstreet
(1935) Exchange of anions and cations absorbed to the root surface with similarly
charged ions of soil solution is known as ion exchange.
3. Mass flow (Kramer 1956) when solute are transported with the conventional flow of
water from the soil to the plant root under the influence of transpiration .
Mechanism of passive transport
A. Diffusion :-All molecules are in constant motion. As molecules collides into each other,
directions are changed, causing random dispersal of the molecules. The random
movement of molecules results in diffusion.
B. Diffusion is the movement of molecules from an area of high concentration to one of
low concentration.
C. This difference in the concentration of molecules across a space is called
the concentration gradient, is the difference between the concentration of a solute in
one place and its concentration in an adjacent area.
D. Molecules diffuse faster at higher temperatures than at lower temperatures, and smaller
molecules diffuse faster than large molecules.

image703
E. Diffusion occurs due to the kinetic energy (thermal motion or heat) of the molecules.
F. The rate of diffusion is variable and depends on temperature, molecular weight, distance
to travel, solvent density, and size and the type of molecules that are diffusing.
G. In diffusion energy is not required by the cell. Most transport of materials into and out of
cells occurs by diffusion Only small molecules can cross the cell membrane by simple
diffusion.
H. Only after an even distribution of molecules is occured there is no longer any diffusion
because there is no longer a concentration gradient.
I. Diffusion will eventually cause the concentration of molecules to be the same throughout
the space the molecules occupy, the system has reached an equilibrium.
Osmosis :-Movement of water molecules across a semipermeable membrane from an
area of high water concentration (low solute) to an area of low water concentration (high
solute) is known as osmosis.
1. Water crosses the membrane toward the higher solute concentration until the
concentration gradients of both water and solutes even out.
2. The net direction of osmosis depends on the relative concentration of solutes on the two
sides of the cell membrane.
3. Hypotonic:- concentration of solute molecules outside cell is lower than the
concentration of solutes inside the cell cytosol.In a hypotonic condition plant cell only
take up water until the inner and the outer water potential are equal. The solution at the
inside of the plant cell is subject to an additional hydrostatic pressure. Plant cells store
ions, sugars, organic and amino acids and other substances in considerable concentrations

in their vacuoles. The solutes cause an influx of water. In this way can plant cells build up
a large positive internal pressure, the turgor pressure. It has a decisive influence on the
maintenance of the rigidity and stability of plant tissues. Each cell exerts a pressure on its
neighbouring cells. The pressures add up to a large tissue tension.

image704
4. Hypertonic :-concentration of solutes outside cell is higher than concentration of solutes
inside cell cytosol. When a plant cell is immersed in a hypertonic solution ,water is
extracted from the protoplasm, it shrinks. This phenomenon is called plasmolysis. The
process is reversed as soon as the cells are transferred into a hypotonic
solution (deplasmolysis).
5. Isotonic :-. concentrations of solutes outside and inside cell are equal. In isotonic
condition, there is no movement of water into the cell of plant. The cell becomes flaccid
(limp), and the plant may wilt.
Facilitated diffusion

Plasma membranes of plant made up of phospholipid bilayer interspersed with two type
of proteins. It is selective-permeable and allows the passage of specific molecules needed
by cell.

image702

Lipid bilayer is impermeable to some of the vital molecules like glucose, sodium ions,
chloride ions etc. Their transport must therefore be "facilitated" by proteins that span the
membrane and provide an alternative route or bypass. Nonelectrolytes (uncharged
particles) diffuse through membrane at a rate proportional to their solubilities in lipid
and inversely proportional to their molecular size.

Due to which, many important gases ,such as, CO2, NH3,O2 and N2 .cross lipid bilayers
by dissolving in lipid portion of membrane, diffusing to the other lipid- water interface,
and then dissolving in the aqueous phase on the other side of membrane.

Facilitated Diffusion is the movement of specific molecules down a concentration

gradient, passing through the membrane via a specific carrier protein.

It is similar to simple diffusion in the sense that it does not require expenditure of
metabolic energy and transport is again down an electrochemical gradient.

Two major groups of integral membrane proteins are involved in facilitated diffusion:

Ion Channels do not really bind the solute, but are hydrophilic pores formed by channel
proteins, through the membrane that open and allow certain types of solutes to pass
through.

Therefore, ion channels mediate only passive transport (down hill) to allow diffusion of
ions inorganic ions,such as Na+, K+,Ca++ ,CL- down their electrochemical
gradients across a plasma membrane.

In general, channels are quite specific for the type of solute they will transport and
transport through channels is quite a bit faster than by carrier proteins.

Three Essential characteristics: 1. Specific (selective for single nutrient molecule) 2.

Passive (requires no input of energy) 3. Saturates (non-linear dependence on
concentration).

Some ion channels are always open, but others have gates that open to allow ions to pass
or close to stop their passage and control the channel's permeability. The gate can
controlled by voltage or ion concentration, even by light, hormones or other
stimuli. Ion channels :1. Voltage-gated channels
2. Mechanical gated channels
3. Ligand-gated channels

image705

Carrier proteins(also known as permeases or transporters) :- The carrier proteins are

found in all natural membrane and have the two features in common (1) they facilitate
movement of solutes in thermodynamically favoured direction and (2) they display
affinity and specificity for the solute to be transported.

Carrier proteins bind a specific type of solute and are thereby induced to undergo a series
of conformational changes which has the effect of carrying the solute to the other side of
the membrane. The carrier then discharges the solute and, through another
conformational change, reorients in the membrane to its original state. Typically, a given
carrier will transport only a small group of related molecules.
Carrier protein s are classified as follows :- Uniporters ,when a single solute is
transported from one side of the membrane to the other.symporters , when transfer of
one solute depends on the simultaneous or sequential transfer of a second solute in the

same direction, and antiporters , where transfer of one solute depends on the
simultaneous or sequential transfer of a second solute, but in the opposite direction.
Section "D"
Active transport (using ATP energy)When a molecule moves from lower concentration
to higher concentration , requires input of energy (uses ATP), and is known as active
transport. For uncharged substances active transport moves molecules against their
concentration gradients. For charged substances (ions), active transport moves molecules
against concentration + electrical gradients.

image706
Active transport is always mediated by carrier proteins , similar to those of facilitated
diffusion, but these carrier proteins act as pumps, using the energy from splitting ATP to
pump specific molecules against the concentration gradient.Two types of active transport :1. ATP driven active transport systems :- Hydrolysis of ATP takes place by
o Na+ ,K+ transport (sodium pump),Na+ is maintained at low concentrations inside
the cell and K+ is at higher concentrations.
o Ca++ transport
o H+,K+ transport iv ABC transporter superfamily, v osteocast proton pumps.
2. Ion -gradient driven active transport systems orCo-transport ;-The gradients of ions
(cations or anions) established by ATPase or by light driven transport may lead to
secondary active transport of substance like amino acid and sugar. These molecules move
in same direction-Symport or in opposite directions Antiport .

image707
In plants, active transport enables roots to absorb mineral ions from the soil, which are
therefore more concentrated inside plant cells than in the soil. This requires ATP energy
from aerobic respiration, and therefore roots need oxygen to allow mineral uptake and a
waterlogged (thus anaerobic) soil will kill most roots.
Bulk transportLarge molecules, such as polysaccharides and proteins,are too large to
cross the cell membrane and cross the membrane via vesicles, occurs byexocytosis
(substances leaving the cell.) or endocytosis.
.

image708
Mineral nutrients uptakeVascular plants have the vascular system ; trumpet hyphae
in bryophytes. Water and dissolved mineral nutrients enter the plant in two ways:o Apoplastic(or outer space):- It is a non-living continuous system made up of
water-filled cellulosic cell wall and intercellular spaces from epidermis to
xylem.The ions which enter the cell walll of epidermis, cell walls of cortex,
cytoplasm of endodermis, cell walls of pericycle and finally accumulate in xylem

vessels.The passive movement of ions into the apoplast usually occurs through
ion-channels, the trans-membrane proteins that function as selective pores. .
o Symplastic( or inner space ) It is a living continuous system formed by
cytoplasm and plasmodesmata from epidermis to xylem parenchyma. An ion enter
the cell wall of the root hair and pass between the wall and plasma membrane and
moves across the cytoplasm, cortex, endodermis, pericycle through
plasmodesmata endodermis, a layer of cells that they must pass through to enter
the xylem.The entry or exit of ions to and from the symplast requires the
expenditure of metabolic energy, which is an process.
The mineral ions stored in the root hairs pass into xylem from where these are
translocated through xylem along with the ascending stream of water, which is pulled
up through the plant by transpiration stream. Mineral elements brought to the leaves
are subsequently assimilated into organic molecules and are redistributed to other
parts of the plant through the phloem.