Chapter 7

Plant Growth and Mineral Nutrition

Plant growth:
• Growth is a permanent, irreversible increase in the size and
the change of form of an organism.
• It is one of the characteristic features of living organisms.
• Quantitative aspect -an increase in the length, breadth, size,
volume, body mass or dry weight and number of cells.
• Qualitative aspect- Development and differentiation.
• In multicellular plants, growth is indeterminate.
• It is localized at meristems.
• Meristems are of three types based on location viz. Apical,
Intercalary and Lateral.

Phases of growth: Fig.7.1

A. Phase of cell division/ formation:
The thin walled, non-vacuolated cells of meristem with
prominent nucleus and granular cytoplasm divide by mitosis
to produce new cells. The growth rate is slow (Lag phase).
B. Phase of cell enlargement / elongation:
The newly formed cells absorb water and become turgid and
enlarged. The growth rate is accelerated (exponential or Log
phase).
C. Phase of Cell maturation/ differentiation:
The enlarged cell becomes mature and is specialized to
perform specific function. The growth rate is steady
(stationary phase).
 Conditions for Growth:
1. Water is the essential as a major constituent of
protoplasm and to maintain turgidity of the cell.
2. It also acts as aqueous medium for biochemical
reactions.
3. Several microelements and macro elements are
required for the proper growth of the plant.
4. Optimum Temperature ranges between 25-350C.
5. Oxygen is essential for respiration and the release of
energy.
6. Light is very much essential for germination of seed
and photosynthesis.
7. The Gravitational force decides the direction of growth
of the shoot and root.

Growth Rate (efficiency index)

It is the increased growth per unit time. It can be measured
by an increase in the size and area of different plant organs
like leaf, flower, and fruits.

Absolute growth rate (AGR) is the ratio of change in the cell

number (dn) over the time interval (dt) the measurement and
comparison of total growth per unit time. AGR = dn/dt

Relative growth ratio (RGR) is the AGR, divided by total

number of cells present in the medium. RGR = AGR/ n AGR
and RGR describe the dynamics of cell growth in culture.

Types of growth :
Arithmetic growth and Geometric growth.
• Arithmetic growth has a constant growth rate and an
increase in the growth occurs arithmetically. It shows a linear
curve. For example, elongation of roots. It is expressed
mathematically as,
Lt = Lo + rt (Where Lt = Length at time‘t’
Lo = Length at time ‘Zero’,
r = Growth rate and
t = Time of growth) Fig.7.2

• Geometric growth initially has a slow growth rate which later

becomes exponential. It can be expressed mathematically as,
W1 = Wo ert (Where W1= Final size, Wo = initial size, r = growth
rate, t =time and e = base of natural logarithm) Fig.7.3

Fig.7.4 In the above example, two leaves ‘A’ and ‘B’ are of
different sizes but show same absolute increase in area in a
given time. Both leaves grow and increase their area by 5cm2
to produce ‘A’ and ‘B’ leaves. ‘A’ leaf of 5cm2 in size grows
5cm2/ day then its RGR would be 100%. If the leaf is 50cm2 in
size and the growth rate/day is 5cm2 then its RGR would
become 10%.

Growth curve: Fig.7.5 Fig.7.6

• Growth curve is a graphic representation of the total
growth against time.
• It can be Linear, Exponential and Sigmoid.
• The Arithmetic growth curve is linear while the Geometric
growth curve is exponential. A sigmoid curve is obtained when a
graph of rate of growth against time is plotted for three phases of
growth.
• In the Lag phase, the growth rate is slow. In the Exponential
(Log) phase, growth rate is faster and reaches its maximum. In
the Stationary phase, growth rate gradually slows down.
• The total time required for all phases to occur, is called the
Grand Period of Growth.
Differentiation, De-Differentiation, Re- Differentiation:
• Differentiation is the maturation of cells derived from apical
meristem of root and shoot. It involves permanent change in
structure and function of cells. For example, the parenchyma in
hydrophytes develops large schizogenous interspaces for
mechanical support, buoyancy and aeration.

• Dedifferentiation is to regain the lost ability to divide by a

living differentiated cell. Thus, permanent cell dedifferentes to
become meristematic. For example,the interfascicular
cambium and cork cambium.

• Redifferentiation is the loss of ability to divide by a

dedifferentiated cell and mature to perform specific function. For
example, secondary xylem and secondary phloem.

Development: Fig.7.7
• It is the ordered, progressive changes in shape, form and
degree of complexity.
• It includes all the changes occurring during the life cycle of
plants. Such as growth, morphogenesis, maturation and
senescence.

Plasticity:
• It is the ability of plants to form different kinds of structures in
response to different environmental or internal stimuli, in
various phases of life.
• For example, in the juvenile and mature stages, some plants
show different forms of leaves. The phenomenon is called
heterophylly. Fig.7.8
• Intrinsic plasticity is seen in cotton, coriander, larkspur
(Delphinium).
• Ranunculus flabellasis (butter cup) shows environmental
heterophylly.
 Growth Hormones/ Growth Regulators/
Phytohormones:
• They are the internal factors that inhibit, promote or modify
growth.
• Growth promoters are auxins, gibberellins (GA) and
cytokinins (CK). Growth inhibitors in plants are ethylene and
abscissic acid (ABA).
• They 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 every minute
(optimum) amount.
• They are transported through phloem parenchyma.

a) Auxins :
• Synthesis-In growing tips of plants.
• The natural auxin- Indole-3-acetic acid (IAA)
• The precursor of IAA in plants - Tryptophan.
• Primary function- cell elongation.
• Synthetic auxins- IBA (Indole butric acid), NAA (Naphthalene
acetic acid), 2, 4-D pichloro (Phenoxy acetic acid)

Physiological effects and applications of auxin:

• Auxins cause cell enlargement.
• Apical dominance: Inhibition of the growth of lateral buds.
• Stimulation of stem and root growth.
• Induction of multiplication of cells in tissue culture to produce
callus.
• It stimulates formation of lateral and adventitious roots.
• 2, 4-D (2,4 dichlorophenoxy acetic acid) kills dicot weeds.
• Production of seedless fruits like orange, lemon, grapes,
banana etc.by induced parthenocarpy.
• Stimulation of cell division in cambium and also cause early
differentiation of xylem and phloem.
• Auxins promote rooting at low concentration and shoot
formation at higher concentration.
• It causes early rooting ‘cuttings’.
• NAA and 2,4-D as foliar spray induces flowering in litchi and
pineapple.
• It prevents premature fruit fall in apples, pear and oranges by
preventing formation of an abscission layer.
• Auxins break dormancy in seed and promote quick
germination.

b) Gibberellins :
• Synthesis- In young leaves, seeds, roots and stem tips and
abundant in root tip and developing seeds.
• It shows non-polar transport through vascular tissue.
• First isolated from the fungus Gibberella fujikuroi
• Rice plants infected with Gibberella fujikuroi show extensive
stem elongation called ‘bakane disease’.
• Precursor- Mevalonic acid.
• GA3 is the most common and biologically active form.

Physiological effects and application of Gibberellins :

• Breaking of bud and seed dormancy.
• Stimulation of seed germination in cereals (barley and wheat)
by synthesizing hydrolysing enzymes.
• Stem elongation due to increase in length of internodes.
• Promotes bolting and flowering in plants with rosette habit e.g.
beet, cabbage.
 It causes Parthenocarpy in tomato, apple and pear
• Stimulation of flowering in long day plants.
• It is used to increase the fruit size and bunch length of
grapes.
• Increase in the height of genetically dwarf plants like maize.
• It can overcome the requirement of vernalization.
• It inhibits growth of root, delays senescence and prevents
abscission.
• It causes production of male flowers on female plants.

c) Cytokinin :
 Cytokinin is made from coconut milk -Kinetin.
 Cytokinin from maize – Zeatin Chemical nature - 6-furfuryl
amino purine.

Physiological effects and applications of cytokinin :

• It promotes cell division and cell enlargment.
• A low cytokinin to auxin ratio induces rooting while a high ratio
causes shooting.
• It delays the senescence.
• It breaks dormancy and promotes seed germination.
• Cytokinin reverses apical dominance and promotes the growth
of lateral buds. It induces RNA synthesis and formation of
interfascicular cambium.

d) Ethylene :
 It is the only gaseous plant hormone.
 Precursor of ethylene - Ethephon.
 Synthesis - in roots, shoot apical meristem, ripening fruits
etc.

Physiological effects and application of ethylene :

 It promotes ripening of fruits like bananas, apples and
mangoes.
 It stimulates initiation of lateral roots in plants.
 It breaks the dormancy of bud and seed.
 It accelerates the abscission in leaves, flowers and fruits
by forming an abscission layer.
  It causes apical dominance and inhibits the growth of
lateral buds.
 It enhances the process of senescence of plants.
 It inhibits flowering in most of the plants except pineapple.
 It causes epinasty (drooping) of leaves and flowers.
 It increases activity of chlorophyllase enzymes causing
degreening effect in banana and Citrus fruits.

e) Abscissic Acid :
• It is a natural growth inhibiting hormone.
• Synthesis - in leaves, fruits, roots, seeds etc.
• Chemical nature - A 15-carbon sesquiterpenoid
 Precursor - Mevalonic acid.

Physiological effects and application of ABA:

• It induces dormancy in buds and seeds by inhibiting growth
processes.
• It accelerates the senescence of leaves, flowers and fruits and
their abscission. It inhibits and delays cell division and cell
elongation.
• It suppresses cambium activity.
• It causes efflux of k+ ions from the guard cells and results in
stomatal closure. Thus, it functions as an antitranspirant and
acts as a stress hormone.
• It inhibits flowering in long day plants but stimulates flowering
in short day plants.

Photoperiodism :
• The transition from vegetative growth to reproductive growth in
higher plants is influenced by light, temperature and several
other environmental and nutritional factors.
• The influence of light on flowering – Photoperiodism.
• The influence of temperature on flowering – Vernalization.
• Three aspects of light viz, Quality, Intensity and Duration of
light influence germination of seed, vegetative growth,
photosynthesis, etc.
• Garner and Allard (1920) whille studying the flowering
behaviour in Soyabean and Meryland mamoth variety of tobacco
concluded that the relative length of the day was most crucial in
the growth and development of flowers. They found that every
plant has a critical photoperiod above or below which flowering
occurs.
• They used the term photoperiodism and classified plants in
three categories asShort Day Plants (SDP), Long Day Plants
(LDP) and Day Neutral Plants (DNP).

a. Short Day Plants (SDP) Fig.7.9

• These plants usually flower during winter and late summer
when day length is shorter than the critical photoperiod.
• They require long uninterrupted dark periods/ night for
flowering.
• If the dark period is interrupted even by a flash of light, SDP
will not flower. Hence, they are also called long night plants.
• Examples of SDP - Dahlia, Aster, Tobacco, Chrysanthemum,
Soybean (Glycine max), Cocklebur (Xanthium), etc.

b. Long Day Plants (LDP):

• Long day plants flower during summer and they require longer
duration of light than the critical photoperiod, for flowering.
• They are called short night plants as they require a short dark
period.
• Examples of LDP are pea, radish, sugar beet, cabbage,
spinach, wheat, poppy, etc.

c. Day Neutral Plants (DNP):

• These plants flower throughout the year and do not require
specific photoperiod to flower. Hence, they are called Day
neutral plants e.g. Cucumber, tomato, cotton, sunflower, maize,
balsam, etc. Phytochrome :
• Hendricks and Borthwick (1952) suggested the involvement of
some pigment proteins in plants that receives the photoperiodic
stimulus and named them as phytochromes.
• Phytochromes are produced by leaves and are located in the
cell membrane of green cells.
• They exist 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. 660nm (red) Pr Pfr 730nm (far red)

During day time, Pfr inhibits flowering in SDP but initiates

flowering in LDP. During the dark period Pfr changes into Pr and
stimulates/ promotes flowering in SDP and inhibits flowering in
LDP.

Vernalization:
• It is a low temperature (chilling) treatment that induces early
flowering in plants. For example, plants such as cereals, crucifers
require a period of cold treatment for flowering.
• Early flowering in the plants can be induced by pretreatment to
their seeds/ seedlings at low temperature (1-60C for one to one
and half months’ duration). The site of vernalization – The shoot
apical meristem.
• Vernalization stimulus A chemical named vernalin.
• The reversal of vernalization by high temperature treatment is
called devernalization.
• Vernalization can be applied to produce crops earlier and
also to cultivate crops in regions where they do not grow
naturally.
Mineral nutrition :
• Plants obtain the inorganic substances from soil in the form of
minerals.
• Minerals are absorbed in ionic form as PO4 , CO3 , SO4 , etc.
• On the basis of their requirement minerals are classified as
essential and non-essential.
• Based on the quantity requirement, minerals are classified as
minor or microelements and major or macroelements.
• Microelements are required in traces e.g. Zn, Cu, Al, Si, etc.
• Macroelements are required in large quantities. e.g. C, H, O,
P, Mg, N, K, S, etc. C, H, O are non mineral major elements.

Symptoms of Mineral deficiency in plants :

• 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 called symptoms, indicate the
deficiency of a particular element.
• Some important deficiency symptoms seen in plants are:
1. Stunting (retarded growth),
2. Chlorosis(yellowing of leaves),
3. Necrosis (localized death of tissues),
4. Mottling ( green and non-green patches on the leaves) and
5. Abscission (Premature fall of flowers, fruits and leaves).

*Please refer Table 7.10

Toxicity of Micronutrients :
• A mineral is considered as toxic when its ionic concentration
reduces the dry weight of tissues by 10%,
• Toxicity of one element may inhibit the uptake of the other e.g.
Mn competes with Fe, Mg for uptake while inhibits Ca
translocation to shoot apex.
• Hence, Mn toxicity develops deficiency symptoms of Ca, Mg
and Fe.

Minerals salt absorption :

• It occurs in two ways :
a. Passive Absorption occurs when the mineral ions diffuse
from a region of their higher concentration to a region of
their lower concentration, without expenditure of energy.
• It can take place by direct ion-exchange, indirect ion-
exchange, mass flow and Donnan equilibrium .
• Donnan equilibrium - The 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 involves the uptake of mineral ions

against concentration gradients using ATP derived from
respiration. When the roots are deprived of oxygen, they
show a sudden drop in active absorption of minerals.
• The mineral ions accumulated in the root hair pass into the
cortex and finally reach the xylem.

Nitrogen cycle:
• It is a series of natural processes by which aerial Nitrogen
enters the bodies of living organisms through soil and is returned
to environment. Plants need nitrogen in a reactive form usually
as nitrate ions. Fig7.11
• Aerial nitrogen is inert and is supplied to the plants through
biological and physical fixation.

Nitrogen fixation :
• It is the conversion of free nitrogen (N2) of the atmosphere into
nitrogenous salts to make it available for the plants is called
nitrogen fixation.
• N2 fixation may occur by physical or biological processes.

Biological Nitrogen fixation is carried out by prokaryotes

called ‘Diazotrophs’, which may be symbiotic or free living.
• The cyanobacteria fix a significant amount of nitrogen in
specialized cells called heterocysts.
• Nitrogen fixation is a high energy requiring process and nitrogen
fixers use 16 molecules of ATP to fix each molecule of nitrogen
to ammonia which is then converted into amino acids.
N2 + 8H+ +8e- + 16ATP 2NH3 +H2 + 16ADP + 16Pi

• Nitrification is conversion of ammonia into nitrates by

chemoautotrophs. Ammonia Nitrosomonas, Nitrosococcus
Nitrites Nitrobacter Nitrates.

• Symbiotic N2 fixation is exhibited by the Rhizobium

bacteria which are soil bacteria that form root nodules in plants
belonging to family Fabaceae e.g. beans, gram, groundnut etc.

• Ammonification: Actinomycetes and some ammonifying

bacteria decompose the dead bodies of plants and animals and
convert organic nitrogen into amino acids and then to ammonia.
Ammonia (NH4+) is now available for uptake by plants and other
micro-organisms for growth.

• Nitrogen assimilation is the conversion of nitrates, nitrites

and ammonia obsorbed from the soil by the green plants into
nitrogenous organic compounds like amino acids, DNA. From
plants, nitrogen enters food chain and moves through animals to
decomposers.

• Amino Acid synthesis occurs through Reductive

amination and Transamination.

• Amides are the amino acids having two amino groups formed
by amidation, e.g. asparagine and glutamine are formed from
glutamic acid and aspartic acid respectively by addition of
another amino group to each.
• Denitrification is conversion of soil nitrates back into nitrogen
gas by the anaerobic bacteria. Denitrifying bacteria include
Bacillus spp., Paracoccus spp. and Pseudomonas denitrificans.

• Sedimentation is the leaching of soil nitrates to the sea or

deep into the earth along with percolating water.