Chapter 7. Plant growth :                                                  Mr.
Pravin Bhosale
Growth is one of the characteristic features of living organisms.
Growth as a phenomenon has two aspects viz. quantitative and qualitative.
Quantitative aspect – It is an increase in the length, breadth, size, volume, body mass or dry
weight and number of cells. Growth as a quantitative change is a final end product of
successive metabolism.
Qualitative aspect – It is the change in the nature of growth where development is an
ordered change or progress while differentiation leads to higher and more complex state.
Growth thus can be defined as permanent, irreversible increase in the bulk of an
organism, accompanied by the change of form.
In multicellular (vascular) plants, growth is indeterminate and occurs throughout the life
indefinitely. It is restricted to some specific region called meristems which are the regions
where new cells are constantly and continuously produced.
Meristems are of three types based on location viz. Apical, Intercalary and Lateral.
7.2 Phases of growth :
The cells in the meristem divide, enlarge and get differentiated.
Corresponding to these three stages, there are three phases of growth :
A.Phase of cell division/ formation :
   •   Cells of meristem are thin walled, non-vacuolated having prominent nucleus and
       granular cytoplasm.
   •   Meristematic cell undergoes mitotic division to form two new cells.
   •   One cell remains meristematic and the other cell undergoes enlargement and
       differentiation.
   •   In this phase, rate of growth occurs at a slower pace (Lag phase).
B. Phase of cell enlargement/ elongation :
   •    The newly formed cell becomes vacuolated, osmotically active and turgid due to
        absorption of water.
   •    The turgidity results in the enlargement of cell - both lengthwise and breadthwise.
   •    In this phase new wall materials and other materials are synthesized to cope up with
        the enlargement.
   •    The growth rate in this phase occurs at an accelerated pace (exponential or Log
        phase).
C. Phase of Cell maturation/ differentiation :
   •    The enlarged cell now becomes specialized to perform specific fuction and attains
        maturity - both morphological and physiological.
   •    In this phase, rate of growth slows down and comes to a steady state (stationary
        phase).
The different environmental and physiological conditions necessary for the growth include
   1.   Water,
   2.   Supply of nutrients,
   3.   Temperature,
   4.   Oxygen,
   5.   Carbon/ Nitrogen ratio,
   6.   Gravitational force,
   7.   Light and
   8.   Growth hormones.
The chief conditions are explained below :
   •   Water is the essential component of protoplasm and maintains turgidity of the cell.
       It acts as aqueous medium for biochemical reactions.
   •   Microelements and Macroelements are nutrients required for the proper growth of
       the plant. Optimum Temperature ranges between 25-350C.
   •   Oxygen is essential for respiration and the release of energy.
   •   Light is very much essential for germination of seed and photosynthesis.
   •   Gravitational force decides the direction of growth of the shoot and root.
7.4 Growth Rate and types of growth :
Growth rate :
   •   It is the increased growth per unit time. It is also called efficiency index.
   •   Rate of growth can be measured by an increase in the size and area of different
       plant organs like leaf, flower and fruits.
   •   The ratio of change in the cell number (dn) over the time interval (dt) is called
       Absolute growth rate (AGR).
   •   Alternatively, it is the measurement and comparison of total growth per unit time.
   •   The AGR, when divided by total number of cells present in the medium, gives
       Relative growth ratio (RGR).
   •   Alternatively, RGR refers to the growth of a particular system per unit time,
       expressed on a common basis or it is the ratio of growth in the given time/ initial
       growth.
   •   AGR and RGR are useful in describing the dynamics of cell growth in culture.
Types of growth :
There are two types of growth viz, arithmetic growth and geometric growth.
a.Arithmetic growth :
   1. Here, rate of the growth is constant and an increase in the growth occurs in
      arithmetic progression. i.e. 2, 4, 6, 8 cms etc.
   2. In this type of growth, the rate of growth is constant.
   3. After mitosis one of the daughter cell continues to divide and the other cell takes
      part in the differentiation and maturation.
   4. e.g. elongation of root at a constant rate, best explains arithmetic growth.
   5. Linear curve is obtained when growth rate is plotted against the time.
   6. Arithmetic growth is expressed mathematically by an equation,
        It is expressed as, Lt = Lo + rt
                    Where Lt = Length at time ‘t’
                            Lo = Length at time ‘Zero’
                             r = Growth rate
                             t = Time of growth
When graph of length (L) is plotted against the time (t), a linear curve is obtained as
indicated in the diagram.
b.Geometric growth :
   1. Cell divides mitotically into two. Here, both the daughter cells continue to divide
      and redivide repeatedly.
   2. Such growth is called geometric growth.
   3. Here, growth rate is slow initially but later on there is a rapid growth at exponential
      rate.
   4. Geometric growth can be expressed mathematically by an equation.
We can also observe quantitative comparison between the growth of living system in two
ways.
7.5 Growth curve :
   •   It is a graphic representation of               •   The total time (period) required
       the total growth against time.                      for all phases to occur, is called
   •   There are three types of curves                     Grand Period of Growth.
       viz, Linear, Exponential and
       Sigmoid.
   •   Arithmetic growth curve is linear
       while Geometric growth curve is
       exponential.
   •   Corresponding to three distinct
       phases of growth, growth rates
       differ.
   •   In Lag phase, growth rate is slow.
   •   In Exponential (Log) phase, growth
       rate is faster and reaches its
       maximum.
   •   In Stationary phase, growth rate
       gradually slows down.
   •   When a graph of rate of growth
       against time is plotted for three
       phases of growth, a sigmoid curve
       is obtained.
7.6 Differentiation, De-Differentiation, Re- Differentiation :
a. Differentiation :
   1. It is maturation of cells derived from apical meristem of root and shoot.
   2. Permanent change in structure and function of cells leading to maturation, is called
      differentiation.
   3. During cell differentiation, cell undergoes few to major anatomical and physiological
      changes
   4. e.g. Parenchyma in hydrophytes develops large schizogenous interspaces for
      mechanical support, buoyancy and aeration.
   5. The maturation is at the cost of capacity to divide and redivide.
b. Dedifferentiation :
    1. The living differentiated cell which has lost the capacity to divide, may regain the
       same as per the need and divide.
    2. Thus, permanent (mature) cell undergoes dedifferentiation and becomes
       meristematic
    3. e.g. interfascicular cambium and cork cambium are formed from parenchyma cells
       between vascular bundles and inner most layer of cortex, respectively.
c. Redifferentiation :
    1. The cells produced by dedifferentiation once again lose the capacity to divide and
       mature to perform specific function.
    2. This is called redifferentiation
    3. e.g. secondary xylem and secondary phloem are formed from dedifferentiated
       cambium present in the vascular bundle.
7.7 Development :
    •   It refers to the ordered or
        progressive changes in shape, form
        and degree of complexity.
    •   It includes all the changes
        occurring in sequence from the
        germination of seed upto the
        senescence or death during life
        cycle of plants.
    •   Thus development includes
        growth, morphogenesis,
        maturation and senescence.
Plasticity
    •   It is the capacity of being moulted , formed or modelled.
    •   It is the ability of plant to form different kinds of structures (i.e to change) in
        response to different environmental (external) or internal stimuli, in various stages
        of life
    •   In many plants, juvenile stage and mature stage show different forms of leaves in
        the same plant e.g. heterophylly in cotton, coriander, larkspur (Delphinium).
    •   The environmental heterophylly is shown by Ranunculus flabellasis (butter cup).
    •   The intrinsic plasticity is found in coriander and cotton.
Heterophylly is exhibited in the same plant in different growth phases or under different
environmental conditions.
7.9 Growth Hormones :
   •   The term ‘hormone’ was coined first by Starling (1906) in animal physiology.
   •   The internal factors that influence growth are called growth hormones or growth
       regulators as they inhibit, promote or modify the growth.
   •   Growth promoters are auxins, gibberellins (GA) and cytokinins (CK).
   •   Growth inhibitors in plants are ethylene and abscissic acid (ABA).
   •   All phytohormones are growth regulators.
According to Thimann and Pincus (1948)
“Plant hormones are organic substances produced naturally in higher plants affecting
growth or other physiological functions at a site remote from its place of production and
active in very minute (optimum) amount”.
Hormones are transported through phloem parenchyma (Phillips 1971).
a. Auxins (Auxien = to grow):
   1. F. W. Went in 1931, used this term first.
   2. Auxin was isolated from urine of a person suffering from Pellagra (Kogl and H. Smit
      1931).
   3. It is synthesized in growing tips or meristematic regions of plants from where it is
      transported to other plant parts.
   4. The most common and important natural auxin is Indole-3-acetic acid (IAA).
   5. Tryptophan is the primary precursor of IAA in plants.
   6. It is the first hormone to be discovered in plants and is primarily responsible for cell
      elongation.
   7. It shows polar transport - Basipetal transport in stem.
   8. Now synthetic auxins like
           • IBA (Indole butyric acid),
           • NAA (Naphthalene acetic acid),
           • 2, 4-D (2,4 dichlorophenoxy acetic acid), etc. are used.
Physiological effects and applications of auxin:
   1. The primary effect is cell enlargment.
   2. In most of the higher plants, growing apical bud inhibits the growth of lateral buds.
       This is called as apical dominance.
   3. Stimulates growth of stem and root.
   4. Induces multiplication of cells, hence used in tissue culture experiments to produce
       callus.
   5. Stimulates formation of lateral and adventitious roots.
   6. These are marketed as synthetic herbicides. e.g. 2, 4-D (2,4 dichlorophenoxy acetic
       acid). It kills dicot weeds without affecting monocot crop plants.
   7. The seedless fruits like orange, lemon, grapes, banana etc. are produced by
       application of auxin (i.e. induced parthenocarpy).
   8. Promote cell division in cambium and also cause early differentiation of xylem and
       phloem.
   9. Promotes root elongation in low concentration and shooting at higher
       concentration.
   10. Also hastens early rooting in propagation by ‘cutting’.
   11. Foliar spray of NAA and 2,4-D induces flowering in litchi and pineapple.
   12. Likewise, it prevents premature fruit drop in apples, pear and oranges, and also
       prevents formation of abscission layer.
   13. Play a role in elongation of cell. It is known to increase rate of respiration.
   14. Break dormancy in seed and promote quick germination.
b. Gibberellins :
   •   It is another growth promoting hormone and is abundant in root tip and developing
       seeds.
   •   It shows non-polar transport through vascular tissue.
   •   Gibberellins were first isolated from the fungus Gibberella fujikuroi by Japanese
       scientist Kurosawa (1926).
   •   He observed that when rice plant was infected by fungus Gibberella fujikuroi, it
       shows extensive stem elongation called ‘bakane disease’.
   •   The crystalline form of Gibberellins were isolated by Yabuta and Sumiki (1938) from
       the fungus culture.
   •   They named it as gibberellin.
   •   It is synthesized in young leaves, seeds, roots and stem tips.
   •   These are synthesized from mevalonic acid.
   •   More than 150 chemical types are known so far. GA3 is most common and
       biologically active form. Chemically it contains a gibbeane ring - a cyclic diterpene
       with four isoprene units.
Physiological effects and application of Gibberellins :
   1. Dormancy of bud can be broken by gibberellin treatment.
   2. It can promote seed germination in cereals like barley and wheat by synthesizing
       hydrolysing enzyme amylase to produce sugar.
   3. The most striking effect of it, is the elongation of stem where internodes increase in
       length.
   4. It also promotes bolting i.e. elongation of internodes just prior to flowering in plants
       those with rosette habit e.g. beet, cabbage.
   5. It causes parthenocarpy in tomato, apple and pear, and flowering in long day plants.
   6. It is used to increase the fruit size and bunch length of grapes.
   7. When gibberellins are applied on genetically dwarf plants like maize, the stem
       rapidly elongates and acquires the height of normal tall varities of maize.
   8. Application of gibberellins overcomes the requirement of vernalization.
   9. Usually, it inhibits growth of root, delays senescence and prevents abscission.
   10. It also breaks dormancy of seed and hastens germination.
   11. Application of gibberellin causes production of male flowers on female plant.
c. Cytokinin :
   •   It is another growth hormone that promotes cell division.
   •   Letham coined the term cytokinin.
   •   The first cytokinin was discovered by Skoog and Miller (1954) during investigation of
       nutritional requirements of callus tissue culture of Nicotiana tabacum (Tobacco).
   •   They observed that the callus proliferated when the nutrient medium was
       supplemented with coconut milk and degraded sample of DNA (obtained from
       herring sperm).
   •   They named it as kinetin. Chemically kinins are 6-furfuryl amino purine.
   •   First natural cytokinin was obtained from unripe maize grains by Letham et al.
   •   It is known as Zeatin.
   •   6-benzyl adenine is a synthetic cytokinin hormone.
   •   Seven different types of cytokinins are recorded from plants.
   •   Natural cytokinins are also reported from plants like Banana flowers, apple and
       tomato fruits, coconut milk, etc.
Physiological effects and applications of cytokinin :
   1. Besides cell division, it also promotes cell enlargment.
   2. High cytokinin promotes shooting.
   3. A low ratio of cytokinin to auxin induces root development but a high ratio causes
       buds and shoot to develop.
   4. Cytokinin and auxin ratio and their interactions controls morphogenic
       differentiation.
   5. Promotes the growth of lateral buds and controls apical dominance by cell division.
   6. Delays the senescence or ageing and abscission processes in plant organs. This was
       reported by Richmond and Lang (1957).
   7. Formation of interfasicular cambium and expansion of cells are other functions.
   8. Also breaks dormancy and promotes the germination of seeds.
   9. Cytokinin reverses apical dominance effect.
   10. Induces RNA synthesis and formation of interfascicular cambium.
d. Ethylene :
   •   It is the only gaseous growth regulator.
   •   Denny (1924) reported ethylene is effective in fruit ripening.
   •   Gane (1934) established that plants naturally synthesize ethylene.
   •   Crocker (1930) proposed that ethylene is the plant hormone responsible for fruit
       ripening.
   •   It is a simple gaseous hydrocarbon with essential role in the fruit ripening.
   •   The most widely used compound as a source of ethylene is ethephon.
   •   It is synthesized in roots, shoot apical meristem, ripening fruits etc.
Physiological effects and application of ethylene :
   1. Promotes ripening of fruits like bananas, apples and mangoes.
   2. Stimulates initiation of lateral roots in plants and breaks the dormancy of bud and
      seed.
   3. Accelerates the abscission activity in leaves, flowers and fruits by forming of
      abscission layer.
   4. Ethylene inhibits the growth of lateral buds and causes apical dominance and
      retards flowering.
   5. Associated with the enhancement of process of senescence of plants organs.
   6. Inhibits flowering in most of the plants except pineapple.
   7. Causes epinasty (drooping) of leaves and flowers.
   8. It increases activity of chlorophyllase enzyme causing degreening effect in banana
      and Citrus fruits.
e. Abscissic Acid :
   •    It is a natural growth inhibiting hormone.
   •    Carns and Addicott (1961-65) observed that the shedding of cotton balls was due to
        a chemical substance abscisin I and II.
   •    Wareing (1963) isolated a substance from buds of Acer that can induce bud
        dormancy and named it dormin.
   •    These two identical chemical substances were given the common name abscissic
        acid.
   •    It is synthesized in leaves, fruits, roots, seeds etc.
   •    Chemically, it is a 15-carbon sesquiterpenoid and is synthsized from mevalonic acid.
Physiological effects and application of ABA:
   1. It promotes abscission of leaves and induces dormancy in many plants.
   2. Controls the dormancy in buds and seeds by inhibiting growth processes.
   3. Accelerates the senescence of leaves, flowers and fruits.
   4. Inhibits and delays cell division and cell elongation and suppresses cambium activity
      by inhibiting mitosis in vascular cambium.
   5. ABA could cause efflux of k+ ions from the guard cells and result in closure of
      stomata. So, it is known as an antitranspirant.
   6. Acts as a stress hormone by inducing the plant to bear the adverse environmental
      conditions.
   7. Inhibits flowering in long day plants but stimulates flowering in short day plants.
7.10 Photoperiodism :
   •    Higher plants reproduce sexually by producing special structures called flowers.
   •    Plants exhibit transition from vegetative growth to reproductive growth during
        which flowers are produced.
   •    Like vegetative growth, reproductive growth is also influenced by several
        environmental and nutritional factors.
   •    Among the environmental factors - light and temperature exert profound influence
        on flowering.
        The influence of light on flowering is known as Photoperiodism
   1. Light as an environmental factor influences germination of seed, vegetative growth,
      photosynthesis, etc.
   2. Light as a factor as three aspects viz, Quality, Intensity and Duration of light.
   3. It is the duration of light that has profound effect on flowering in higher plants.
   4. The term photoperiodism was used by Garner and Allard (1920).
   5. They were studying the flowering behaviour in plants - Soyabean and Meryland
      mamoth variety of tobacco.
   6. They found that soyabean plant flowers during late summer and tobacco variety
      during winter, irrespective of their germination and growing season.
   7. They studied effects of different temperatures, nutrition, soil moisture, etc. in
       respect of flowering. None of these were found to regulate flowering.
   8. However, experimentally they found that the exposure to specific duration of light
       (i.e. photoperiod) had profound influence on flowering.
   9. They examined the effect of day length on flowering by using artificial illumination.
   10. They concluded that the relative length of the day was most crucial in the growth
       and development of flowers to which they coined the term photoperiodism.
Based on the photoperiodic response, plants were classified in three categories viz,
Short Day Plants (SDP),
Long Day Plants (LDP) and
Day Neutral Plants (DNP).
a. Short Day Plants (SDP) :
   1. These plants usually flower during winter and late summer when day length is
      shorter than the critical photoperiod (critical photoperiod is that length of
      photoperiod above or below which flowering occurs).
   2. These are called long night plants because they require long uninterrupted dark
      period/ night for flowering.
   3. If dark period is interrupted even by a flash of light, SDP will not flower.
   4. Some of the short day plants are Dahlia, Aster, Tobacco, Chrysanthemum, Soybean
      (Glycine max), Cocklebur (Xanthium), etc.
   b. Long Day Plants (LDP) :
   1.   Plants that flower during summer are called long day plants.
   2.   They require longer duration of light than the critical photoperiod, for flowering.
   3.   They are called short night plants as they require short dark period.
   4.   When long dark period is interrupted by a brief flash of light, LD plants can flower
   5.   e.g. pea, radish, sugar beat, cabbage, spinach, wheat, poppy, etc.
c. Day Neutral Plants (DNP) :
   1. These plants flower throughout the year round, independent of duration of light
      (photoperiod).
   2. They do not require specific photoperiod to flower.
   3. Therefore, they are called Day neutral plants e.g. Cucumber, tomato, cotton,
      sunflower, maize, balsam, etc.
Phytochrome :
   •    Hendricks and Borthwick (1952) observed that flowering in SD plants is inhibited, if
        dark period is interrupted even by a flash of red light of 660 nm.
   •    If it is immediately followed by far red light (730 nm), then SD plants will flower.
   •    This observation led them to conclude that some pigment system in plant receives
        the photoperiodic stimulus.
   •    These pigment proteins are called phytochromes.
   •    The leaves produce light-receiving proteinaceous pigment called phytochrome that
        induces flowering.
   •    It exists in two interconvertible forms viz, red (Pr) and far red (Pfr).
   •   When Pfr absorbs far red light, it is converted into Pr and vice versa.
   •   These are located in the cell membrane of green cells.
   •   During day time, Pfr accumulates in the plants. It inhibits flowering in SDP but
       initiates flowering in LDP.
   •   During dark period Pfr changes into Pr, it stimulates/ promotes flowering in SDP and
       inhibits flowering in LDP.
Vernalization (Yarovization) :
The requirement of low temperature for development of flowering is called as
vernalization
Temperature as a factor has three cardinal points viz, minimum, optimum and maximum
temperature.
It is a low temperature (chilling) treatment that induces early flowering in plants as was
evidenced by Klippart (1918).
Chouard (1960) defines vernalization as acceleration of the ability to flower by chilling
treatment.
The term vernalization was coined by T.D Lysenko (1928) for the effect of low temperature
on flowering in plants.
It is an influence of temperature on development and flowering.
   1. Many plants such as cereals, crucifers require a period of cold treatment for
      flowering.
   2. It is the method of inducing early flowering in the plants by pretreatment to their
      seeds/ seedlings at low temperature (1-60C for one to one and half months’
      duration).
   3. The site of vernalization is believed to be shoot apical meristem.
   4. Generally, vernalization is effective at seed stage in annual plants.
   5. Vernalization stimulus is also a chemical stimulus named as vernalin.
   6. This can be transferred through grafting (Melcher 1939).
Devernalization – The reversion of vernalization by high temperature treatment.
Advantages of vernalization :
• Crops can be produced earlier.
• Crops can be cultivated in regions where they do not grow naturally.
7.12 Mineral nutrition :
   •   Plant absorbs water, gases, mineral, nutrients, etc. from surroundings.
   •   Green plants for the synthesis of their organic food need inorganic substances
       (elements) which are obtained from soil in the form of minerals.
   •   Minerals constitute most commonly occuring solid, inorganic materials obtained
       from the earth’s crust.
   •   Chemical analysis of plant ash clearly indicates that plant absorbs mineral elements
       from surroundings (soil, air and water) for its use.
   •   About 36 to 40 different elements of periodic table are used as minerals by the
       plants.
   •   These are absorbed in ionic (dissolved) form as PO4 , CO3 , SO4 , etc.,usually through
       roots (regions of elongation and growth).
   •   Sources of minerals :
   •   Plants derive necessary elements from the atmosphere, soil and water.
   •   Carbon enters the plant as atmospheric carbon-dioxide.
   •   Source of hydrogen is water and oxygen comes from air and water.
   •   Carbon, Hydrogen and oxygen are not minerals in origin.
   •   Source of nitrogen is the soil. Plant derives nitrogen from both mineral and non-
       mineral origin.
Classificaion of minerals :
   •   Earlier, on the basis of their requirement minerals were classified as essential and
       non-essential. Essential minerals are those that are indispensible without which
       plants can not complete their life cycle e.g. C, H, O, N, P, etc.
   •   These elements play structural and physiological roles.
   •   Their absence can produce/ cause major deficiency symptoms.
   •   The nonessential elements are not indispensable and they do not produce/ cause
       any deficiency symptoms.
       This classification is absolete (not in use) now.
   •   Based on the quantity requirement, minerals are classified as minor or
       microelements and major or macroelements.
   •   Microelements are required in traces because they function in the catalytic role e.g.
       Zn, Cu, Al, Si, etc. as co-factors.
   •   Macroelements are required in large quantity.
   •   They mainly play the nutritive and structural roles e.g. C, H, O, P, Mg, N, K, S, etc.
   •   C, H, O are non mineral major elements.
   •   This classification is not accepted now.
Now a days minerals are classified on the basis of their biochemical functions.
Symptoms of Mineral deficiency in plants :
    •   Any visible deviation from the normal structure and function of the plant, is called
        symptom or hunger sign.
    •   The concentration of the essential elements below which plant growth is retarded, is
        termed as critical concentration.
    •   The element is said to be deficient when present below the critical concentration.
    •   Certain morphological changes are indicative of the deficiency of particular element.
    •   Deficiency symptoms also depend on the mobility of the elements in the plants.
    •   The deficiency symptoms appear first in young tissues when elements are relatively
        immobile e.g. sulphur, calcium.
    •   When the elements are actively mobilized within the plants and exported to young
        developing tissues, the deficiency symptoms are visible first in the older tissues
        (senescent leaves). e.g. nitrogen, magnesium, potassium.
Some important deficiency symptoms seen in plants are:
• Stunting : The growth is retarded. The stem appears condensed and short.
• Chlorosis : It is the loss or non-development of chlorophyll resulting in the yellowing of
leaves
• Necrosis : It is the localized death of tissue of leaves.
• Mottling : Appearance of green and nongreen patches on the leaves.
• Abscission : Premature fall of flowers, fruits and leaves.
Toxicity of Micronutrients :
   •   Mineral ion concentration which reduces the dry weight of tissues by 10%, is
       considered as toxic.
   •   Toxicity of one element may inhibit the uptake of the other
   •   e.g. Mn competes with Fe, Mg for uptake, but inhibits Ca translocation to shoot
       apex.
   •   Thus, Mn toxicity develops deficiency symptoms of Ca, Mg and Fe.
Minerals salt absorption :
   •   Most minerals in the soil are charged particles hence, they can not pass across cell
       membrane. Hence most of the minerals are absorbed actively with the expenditure
       energy.
   •   Minerals can also be absorbed passively without expenditure of energy.
   •   Mineral ion absorption is independent of water absorption.
Mineral ion absorption can occur in two ways :
a.Passive Absorption :
   1. Movement of mineral ions into the root occurs by diffusion.
   2. Molecules or ions diffuse from a region of their higher concentration to a region of
      their lower concentration.
   3. The movement of mineral ions into root cells as a result of diffusion is without
      expenditure of energy is called passive absorption.
   4. Passive absorption can take place by direct ion-exchange, in direct ion-exchange
      mass flow and Donnan equilibrium.
Donnan equilibrium :
   •   It is based on the assumption that certain negatively charged ions, after their entry
       into the cell, become fixed on the inner side of the cell membrane and cannot
       diffuse outside through the cell membrane.
   •   Therefore, additional mobile cations are required to balance these fixed anions.
   •   Obviously concentration of cations become more due to accumulation.
   •   This kind of passive absorption of anions/ cations from cell exterior against their own
       concentration gradient in order to neutralize the effect of cations/ anions, is called
       Donnan equilibrium.
   b.Active Absorption:
   1. Uptake of mineral ions against concentration gradient, is called active absorption,
      such movement requires an expenditure of energy by the absorbing cell.
   2. This energy is derived from respiration and is supplied through ATP.
   3. When the roots are deprived of oxygen, they show a sudden drop in active
      absorption of minerals.
   4. The mineral ions accumulated in the root hair pass into the cortex and finally reach
      the xylem.
   5. The minerals in the xylem are then carried along with water to other parts of the
      plant along the transpiration stream and are subsequently assimilated into organic
      molecules and then redistributed to other parts of the plant through the phloem
7.13 Nitrogen cycle:
   •   It is series of natural processes by which Nitrogen enters successively from air to
       organisms through soil and back to environment.
   •   Plants use photosynthetic product, the sugars to make proteins.
   •   To do this, they need nitrogen. Unfortunately, it is very innert (nonreactive).
   •   Plants need nitrogen in a reactive form usually as nitrate ions.
   •   Nitrogen is a limiting nutrient in the agricultural ecosystems.
   •   It exists as nitrogen atoms with a strong triple covalent bond (N ≡ N).
   •   A regular supply of nitrogen to the plants is maintained through biological and
       physical process.
Nitrogen fixation :
    •   Atmosphere is the source of nitrogen. It can not be used directly.
    •   It combines with C, H, N and O to form compounds before being used.
    •   Conversion of free nitrogen (N2) of the atmosphere into nitrogenous salts to make it
        available for the plants, is called nitrogen fixation.
It is of two types : Physical and Biological fixation.
Physical Nitrogen fixation:
1.It occurs in several steps and starts with combination of atmospheric nitrogen with oxygen under
the influence of electric discharge and thunder storm produce nitric oxide.
2.The nitric oxide is then oxidized to nitrogen peroxide in the presence of oxygen.
3.During rains, the nitrogen peroxide combines with rain water to form nitrous acid and nitric acid
which come to ground along with rains.
4.On ground, the alkali radicals of soil react with nitric acid to produce nitrites and
nitrates.(absorbable form)
5.Industrial nitrogen fixation. It occurs by HaberBosch nitrate process at high temperature and
pressure.
Ammonia is then converted to urea as it is less toxic.
Biological Nitrogen fixation :
   1.   It is carried out by prokaryotes called as ‘Nitrogen fixers’ or Diazotrophs’.
   2.   It accounts nearly 70% of natural nitrogen fixation.
   3.   Nitrogen fixers are either symbiotic or free living.
   4.   The cyanobacteria fix significant amount of nitrogen in specialized cells called
        heterocysts.
   5. Nitrogen fixation is high energy requiring process and nitrogen fixers use 16
      molecules of ATP to fix each molecule of nitrogen to form ammonia.
          N2 + 8H+ + 8e- + 16ATP         2NH3 + H2 + 16ADP + 16Pi
   6. Ammonia is then converted into amino acids.
Nitrification : Conversion of ammonia into nitrate is called as nitrification
Most of the soil bacteria participate in converting ammonia into nitrate, the form of
nitrogen which can be used by plants and animals.
This involves two steps performed by two different types of bacteria.
1.First a soil bacteria convert ammonia into nitrogen-di-oxide (nitrite)
 eg. Nitrosomonas, Nitrosococcus, etc.
2.Then another type of soil bacterium called Nitrobacter adds a third oxygen atoms to
create nitrate.
These bacteria are chemoautotrophs.
By metabolizing nitrogen along with oxygen, they obtain energy to power their own life
processes.
Symbiotic N2 fixation :
1.The best known nitrogen fixing symbiotic bacterium is Rhizobium.
2.This soil living/ dwelling bacterium forms root nodules in plants belonging to family
Fabaceae e.g. beans, gram, groundnut etc.
Ammonification :
After the death of plants and animals, various fungi, actinomycetes and some ammonifying
bacteria decompose the tissues and convert organic nitrogen into amino acid and then to
ammonia and back into the ecosystem.
Ammonia (NH4+) is now available for uptake by plants and other micro-organisms for
growth.
Nitrogen assimilation :
   •   In soil, nitrogen is present as nitrates, nitrites and ammonia (NH4+).
   •   It is absorbed by the green plants and converted to nitrogenous organic compounds
       like amino acids, DNA, etc. This is known as nitrogen assimilation.
   •   From plants, nitrogen as biomolecules like amino acids, enters food chain and moves
       to animals and then to decomposers through the death of animals.
   •   Nitrates are first converted to ammonia but it is highly toxic and immediately used
       for conversion into amino acids, which are then transported to other parts of the
       plants for synthesis of proteins.
Amino Acid synthesis :
Amino acids are building blocks of proteins.
The amino acids are synthesized through :
Reductive amination :
Ammonia reacts with alpha ketoglutaric acid to form glutamic acid (glutamate).
α Ketoglutaric acid + NH4 + NADPH2                            Glutamate + H2O+ NADP
Transamination :
Amino group of one amino acid (-CHNH2) is transfered to keto position (-CO) of other
carboxylic acid.
Glutamic acid + oxaloacetic acid                       α - Ketoglutaric acid + Aspartic acid
Amides :
1.Ammonia may be absorbed by amino acid to produce amides. The process is called
amidation.
2.The amides are the amino acids having two amino groups.
3.Extra amino group is attached to acidic group (-COOH) in presence of ATP.
Glutamic acid + NH4+ + ATP            alpha glutamine + ADP
Aspartic acid + NH4+ + ATP            Aspargine + ADP
4.Amides like asparagine and glutamine are formed from glutamic acid and aspartic acid
respectively by addition of another amino group to each.
5.Amides are transported to other parts of plants via xylem vessels.
Denitrification :
It is the process in which anaerobic bacteria can convert soil nitrates back into nitrogen gas.
Denitrifying bacteria removes fixed nitrogen
i.e. nitrates from the ecosystem and return it to the atmosphere in inert form.
Denitrifying bacteria includes Bacillus spp., Paracoccus spp. and Pseudomonas denitrificans.
They transform nitrates to nitrous and nitric oxides and ultimately to gaseous nitrogen.
2NO3          2NO2            2NO           N2
Sedimentation : Nitrates of the soil are washed away to the sea or leached deep into the
earth along with percolating water.