19-02-2022

ABP 121 (Fundamentals of Crop Physiology)

Gopal Lal Choudhary

Asstt. Prof.-cum-Jr. Scientist
(Agronomy)

 Nutrient uptake is the process in which minerals enter the cellular material,
typically following the same pathway as water. The most normal entrance portal for
mineral uptake is through plant roots. During transport throughout a plant,
minerals can exit xylem and enter cells that require them. Mineral ions cross plasma
membranes by a chemiosmotic mechanism. Plants absorb minerals in ionic form:
nitrate (NO3-), phosphate (HPO4-2 & H2PO4-) and potassium ions (K+); all have
difficulty crossing a charged plasma membrane.
 Previously it was thought that absorption of mineral salts takes place along with
water absorption. But it is now understood that mineral salt absorption and water
absorption are two different processes.
 Mineral salts are absorbed from the soil solution in the form of ions. They are
chiefly absorbed through the meristematic regions of the roots near the tips.

Contd………….

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 Plasma membrane of the root cells is not permeable to all the ions. It is selectivity
permeable.
 All the ions of the same salt are not absorbed at equal rate but leads unequal
absorption of ions.
 First step in the absorption of mineral salts is the process of Ion exchange which
does not require metabolic energy.
 For the uptake of nutrient by plant roots, first we required to study the process of
nutrient movement in the soil solution.

 Nutrients must reach the surface of a root for plant uptake of essential
elements to occur. There are three major mechanism of movement of ions from soil
to roots. They are
(1) Root interception
 Root interception occurs when a nutrient comes into physical contact with the root
surface. As a general rule, the occurrence of root interception increases as the root
surface area and mass increases, thus enabling the plant to explore a greater amount
of soil.
 Root interception may be enhanced by mycorrhizal fungi, which colonize roots and
increases root exploration into the soil.
 Root interception is responsible for an appreciable amount of calcium uptake, and
some amounts of magnesium, zinc and manganese.
Contd………….

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(1) Root interception

 The CEC of roots for monocots is 10-30 meq/100 g of soil and takes up monovalent
cations more readily and that of dicots is 40-100 meq/100 g of soil and takes up
divalent cations more readily.
 The quantity of nutrients intercepted by roots depends on the soil concentration of
nutrients and volume of soil displaced by root system.
(2) Mass flow
 Mass flow occurs when nutrients are transported to the surface of roots by the
movement of water in the soil (i.e. percolation, transpiration, or evaporation).
 The rate of water flow governs the amount of nutrients that are transported to the
root surface. Therefore, mass flow decreases as soil water decreases.

Contd………….

(2) Mass flow

 Most of the nitrogen, sulfur, copper, boron and molybdenum and part of magnesium
and manganese move to the root by mass flow. Quantity of nutrients transported is
proportional to:
 Rate of flow (volume of water transpired)
 Solution concentration of nutrient
 Nutrients supplied primarily by mass flow are considered mobile nutrients.
Examples: N, S, B

Contd………….

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(3) Diffusion
 Diffusion is the movement of a particular nutrient along a concentration gradient. When
there is a difference in concentration of a particular nutrient within the soil solution, the
nutrient will move from an area of higher concentration to an area of lower
concentration.
 This phenomenon is observed when adding sugar to water. As the sugar dissolves, it
moves through parts of the water with lower sugar concentration until it is evenly
distributed, or uniformly concentrated.
 Diffusion delivers appreciable amounts of phosphorus, potassium, zinc, and iron to the
root surface.
 Diffusion is a relatively slow process compared to the mass flow of nutrients with water
movement toward the root.
 Nutrients supplied primarily by diffusion are considered immobile nutrients. Example:
P, K.

 The processes of the absorption of mineral salts may be of two types viz., passive and active
absorption.
(I) Passive absorption
 When the concentration of mineral salts is higher in the outer solution than in the cell sap of
the root cells, the mineral salts are absorbed according to the concentration gradient by
simple process of diffusion. This is called as passive absorption because it does not require
expenditure of metabolic energy. In passive absorption mineral salt absorption is not
affected by temperature and metabolic inhibitors. Rapid uptake of ions occurs when plant
tissues are transferred from a medium of low concentration to high concentration. The
major hypotheses (theories) that explain the mechanism of passive transport of ions are:
1. Mass flow theory
2. Contact exchange theory
3. Carbonic acid exchange theory
4. Donnan equilibrium theory
Contd………….

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(I) Passive absorption

1. Mass flow theory
 According to this theory ions are absorbed or taken up by the roots along with mass flow of
water under the effect of transpiration.
 An increase in transpiration pull increases the uptake of ions by the roots, (the uptake of
ions by free diffusion). Thus, mass flow of ions through the root tissues occurs due to
transpiration pull in the absence of metabolic energy.
 Russel and Barber (1960) also supported this theory but raised a question whether the effect
of transpiration is direct or indirect.
 Lopushinsky (1964) worked on this problem and studied in tomato the uptake of radioactive
isotopes P32 and Ca45. He found that an increase in the hydrostatic pressure (comparable to
transpiration pull) increases ion uptake. So transpiration effect on salt absorption is direct.
However, both mass flow theory and direct influence of transpiration have been challenged
in view of recent researchers. Both of these fail to explain salt accumulation against osmotic
gradient. Contd………….

(I) Passive absorption

2. Contact exchange theory
 According to this theory, the ions adsorbed on the clay micelles get adsorbed to the root
in exchange for H+ ions, previously, adsorbed on the root.
 As per this theory the ions adsorbed on the surface of root cells and clay particles are
not held tightly and oscillate within small volume of space.
 If the roots and clay particles are in close contact with each other, the oscillation volume
of ions adsorbed on root surface may over by the oscillation volume of ions adsorbed on
clay particles, and the ions adsorbed on clay particle may be exchanged with the ions
adsorbed on root surface directly without first being dissolved in soil solution.
 Ionic exchange phenomenon has been experimentally shown in excised barley roots with
the use of radioactive K+ in which case K+ ions exchange place with non-radioactive K+
ions. Likewise negatively charged Cl- and Br- are also exchanged without disturbing the
electrical neutrality.
Contd………….

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(I) Passive absorption

2. Contact exchange theory
 It is believed that a similar mechanism operates between soil solution and clay
micelle which would allow for a greater absorption of ions from the external
medium than could normally be accepted by free diffusion.

Figure: Contact exchange theory

Contd………….

(I) Passive absorption

3. Carbonic acid exchange theory
 According to this theory, the CO2 released during respiration of root cells combines
with water to form carbonic acid (H2CO3). Carbonic acid dissociates into H+ and an
anion HCO3- in soil solution. These H+ ions may be exchanged for cations adsorbed on
the clay particles. The cations thus released into the soil solution from the clay
particles, may be adsorbed on root cells in exchange for H+ ions or as in ion pairs with
bicarbonate. Thus, the soil solution plays an important role in carbonic acid exchange
theory.

Figure: Carbonic acid exchange theory Contd………….

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(I) Passive absorption

4. Donnan equilibrium theory
 The Donnan equilibrium theory accounts for the effect of fixed or indiffusable ions and
assumes the cooperation of both electrical as well as diffusion phenomenon (Donnan
equilibrium) for control of electrochemical equilibrium. This theory assumes that
certain fixed or indiffusable ions (for which the cell membrane is impermeable) are
present in a cell sap. In order to maintain an internal balance, such ions would require
ions of other charge.
 Suppose one cell is placed in the nutrient solution. Inside the cell membrane there is
concentration of anions to which the membrane is impermeable. Suppose this
membrane is permeable to anions and cations of the outer solution equal number of
anions and cations will move across the membrane till equilibrium is reached. However,
additional cations are needed to neutralize the fixed ions.

Contd………….

(I) Passive absorption

4. Donnan equilibrium theory
 Therefore, concentration of cations will be more inside the cell whereas, concentration
of anions will be more in external solution. Therefore, ions can move inside the cell
without involvement of energy against the concentration gradient in response to
electrochemical potential gradient. When product of anions and cations in the internal
solution is equal to that of anions and cations in the external solution the Donnan
equilibrium is attained.
 When fixed anions are present on the inner side of the membrane, cations in addition to
normal exchange would be absorbed to maintain equilibrium thus increasing the cation
concentration of the internal solution in comparison to external solution. The same type
of mechanism involving additional absorption on anions would operate in case of fixed
cations on the inner side of the membrane.

Contd………….

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(II) Active absorption

 The active transport of ions from the outer space of the cell to the inner space is
generally occurs against the concentration gradient and hence requires metabolic
energy, this energy is obtained from metabolism of the cell either directly or in directly.
The major hypotheses that explain the mechanism of active transport of ions are:
1. Carrier concept theory
2. Cytochrome-pump theory
1. Carrier concept theory
 Van den Honert (1937) proposed the carrier concept theory of mineral salt absorption.
 According to this theory the ion transport process is carried out by means of carriers
which may be organic molecules or vesicles.
 The space in a cell or tissue where mineral ions enter by the usage of metabolic energy
is called inner space.
Contd………….

(II) Active absorption

1. Carrier concept theory
 According to this concept there are
separate carriers for cations and
anions.
 Outside the membrane ion combines
with the carrier forming ion carrier
complex.
 Now complex moves across membrane
and reaches at the inner surface.
 Finally the complex is broken down on
Figure: Carrier ion transport across a membrane
the inner face of the membrane through
involving energy expenditure
the action of phosphatase enzyme, the
ion is released into the cytoplasm.
Contd………….

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(II) Active absorption

1. Carrier concept theory
 The whole process requires the ATPs which are obtained through the respiration. The
ATPs become available to the carrier by action of kinase enzyme, the process is called
phosphorylation. In the process the ADPs are formed and carrier becomes activated
reaches to the outer surface of the membrane and again gets ready to accept the other
ion.
2. Cytochrome-pump theory
 Lundegardh and Burstrom (1933) claimed that there is close relationship between
anion absorption and respiration. They observed that the rate of respiration increases
when plant is transferred from water to salt solution. The increase of respiration rate
due to transfer of plant tissue from water to salt solution is called salt respiration.

Contd………….

(II) Active absorption

2. Cytochrome-pump theory
 Lundegardh (1954) proposed cytochrome pump theory which is based on the following
assumptions.
1. The mechanism of anion and cation absorption is different.
2. Anions are absorbed through cytochrome chain by an active process.
3. Cations are absorbed passively.
 According to Lundegardh’s theory dehydrogenase reactions on the inner surface of the
barrier or membrane produce protons (H+) and electrons (e-). The electrons move
outward through cytochrome chain and anions move inward. At the outer surface of
the barrier the reduced iron of the cytochrome is oxidized losing an electron and
picking up an anion. The released electron unites with a proton and O2 to form water.

Contd………….

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(II) Active absorption

2. Cytochrome-pump theory
 At the inner barrier surface
the oxidised iron of
cytochrome becomes
reduced in dehydrogenase
reactions. The anion is
released on the inside of the
barrier in the last reaction.
Cations are absorbed
passively to balance the
potential difference caused
Figure: Lundegardh’s cytochrome pump theory
by the accumulation of
anions on the inner barrier
surface.

 Absorption of mineral salts is affected by the number of external and internal factors
(A) External factors
(1) Temperature
 An increase in temperature increases the salt absorption. However, beyond 40ºC
temperature there was a decrease in salt absorption which was mainly due to
denaturation of enzymes involved in salt absorption. Temperature changes affect both
passive and active absorption processes. The rate of free diffusion depends on kinetic
energy of diffusing molecules which is dependent on temperature. Low temperature
also reduces rate of biochemical reactions required for active transport.
(2) Soil pH
 The availability of ions in the soil solution is greatly affected by hydrogen ion
concentration or pH of the soil. Change in the hydrogen ion concentration (pH) of the
soil solution affects the availability of ions to the plants.
Contd………….

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(A) External factors

(2) Soil pH
 For example monovalent phosphate H2PO4- which is readily taken up by the plants is
common in acidic soils. However, as soil approaches towards alkaline medium HPO4-2
and PO4-3 forms are available. H2PO4- form is easily taken up by the plants, HPO4-2
form is not easily taken up by the plant and PO4-3 form is not absorbed by the plants.
Hence soils having low pH values are associated with higher absorption of phosphorus.
(3) Light
 The effects of light on opening and closing of stomata and on photosynthesis indirectly
affects salt uptake. Opened stomata increases mass flow of water which also accelerates
salt absorption due to transpiration stream. The energy obtained from photosynthesis
provides energy for active salt absorption and oxygen given off also improves
conditions for active absorption of ions.

Contd………….

(A) External factors

(4) Availability of O2
 The salt absorption is retarded in absence of O2 due to decrease in ion pump
mechanism and oxidation reduction processes.
(5) Interaction of ions
 Absorption of one element is affected by presence of others. Viets (1944) found that the
K+ absorption is affected by presence of Ca++, Mg++ and other polyvalent cations in
external medium. Uptake of phosphorus is lower at higher concentration and vice-
versa.
 Higher concentration of Fe and Mn reduces the uptake of Zn and Cu by crop plants.
 Negative interactions are found between phosphorus and calcium, phosphorus and
zinc, phosphorus and iron, calcium and magnesium, calcium and zinc, magnesium and
zinc, sulphur and molybdenum.
Contd………….

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(A) External factors

(5) Interaction of ions
 Positive interactions are found between nitrogen and phosphorus, nitrogen and
sulphur, nitrogen and zinc, nitrogen and calcium, nitrogen and magnesium, nitrogen
and molybdenum, phosphorus and sulphur, phosphorus and molybdenum, potassium
and phosphorus, potassium and zinc, potassium and sulphur.
(B) Internal factors
(1) Growth
 The growth of plant increases surface area, number of cells, synthesis of new binding
sites or carriers, factors etc. thereby increasing salt absorption.
(2) Aging
 As the root matures it increases the surface area which is favourable for salt
absorption, but due to heavy suberization the mineral salt uptake is greatly reduced.

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