Chapter-13 Plant Growth and Development

Development= is the sum of two processes Growth + Differentiation

13.1 Growth: - “Irreversible and permanent increase in size of an organ or
its parts or even of an individual cell” is termed as growth.
 “Growth is regarded as one of the most fundamental and conspicuous
Characteristic of living being.”
 Growth is accompanied by metabolic processes (both anabolic and
catabolic) that occur at the expense of energy.

Que: How would you describe the swelling of piece of wood when placed in
water?
Ans: it is not real growth because wood increase in size temporary it is
reversible.

13.1.1 Plant Growth generally is Indeterminate

 Reason: Due to the presence of meristems at certain locations in the
plant body.
 This ability where new cells are always being added to the plant body by
the activity of the meristem is called open form of growth.

 Que: Meristem ceases to divide?

Ans:- Under specific conditions meristematic cells
cease to divide e.g. floral conversation, environmental
dormancy, in leaves.

 Primary vs. Secondary Growth

 Primary Growth: Done by the root apical meristem

and shoot apical meristem. It is responsible for the
elongation of the plants along their axis.
 Secondary Growth: Done by the lateral meristems
(vascular cambium and cork-cambium) which appear
later in life. These increase the girth of the organs, in
which they are active. E.g. in gymnosperms and dicot
stems and roots.
13.1.2 Growth is Measurable

 At the cellular level, growth is essentially an increase in the amount of

protoplasm. Since this is difficult to measure directly, we use several
parameters:
 Increase in fresh weight or dry weight.
 Length (e.g., pollen tube).
 Area (e.g., leaves).
 Volume and cell number.

Notable Examples:
 Maize root apical meristem: Can give rise to more than 17,500 new cells
per hour.
 Watermelon cells: May increase in size by up to 350,000 times.
 Pollen tube growth is measured in length.
 Dorsiventral leaf: increase in surface area.

13.1.3 Phases of Growth

The period of growth is divided into three phases:

1. Meristematic Phase
 Occurs at the root apex and shoot apex.
 Characteristics: Cells are rich in protoplasm, possess large conspicuous
nuclei, and have thin, cellulosic primary cell walls with abundant
plasmodesmatal connections.

2. Phase of Elongation
 Located proximal (just next, away from the tip) to the meristematic
zone.
 Characteristics: Increased vacuolation, cell enlargement, and new cell
wall deposition.

3. Phase of Maturation
 Located proximal to the phase of elongation.
 Characteristics: Cells attain their maximal size in terms of wall thickening
and protoplasmic modifications. Most of the tissues and cell types
represent this phase.
13.1.4 Growth Rates

 Definition:- “The increased growth per unit time is termed as growth

rate.”
 The growth rate:-two types :- may be (i) artihmetic or (ii) geometric.

(i) Arithmetic growth:- “following mitotic cell division only one daughter
cell continues to divide while the other differentiates and matures.”
E.g. a root elongating at a constant rate (Fig.1)

 On plotting the length of the organ against time a linear curve is

obtained.
 Arithmetic growth is expressed as
Lt=Lo+rt
 here, Lt= Length at time ‘t’
Lo= Length at the time ‘zero’
And
r= growth rate/elongation per unit
time.
[Fig.1 Constant Linear Growth ]
 (ii) Geometric Growth

 In most systems, the initial growth is slow (lag phase), followed by a

rapid increase (log or exponential phase), and finally slows down due to
limited nutrient supply (stationary phase). (fig.2)

[Fig.2 an idealised sigmoid growth curve

typical of cells in culture and many higher
plants and plant organs]

 Sigmoid Curve: A plot of the growth parameter against time yields a

typical S-curve. This is characteristic of living organisms growing in a
natural environment.
 Equation: W1 = W0 ert
* W1 = Final size (weight, height, number, etc.)
* W0 = Initial size
* r = Growth rate. Here r is relative growth rate and is also the measure of
the ability of plant to produce new plant material (efficiency index)
* t = Time of growth
* e = Base of natural logarithms

Que: Can you think of more similar examples?

Ans:- Bacterial Growth curve

Que:- what kind of curve can you expect in a tree showing seasonal
activities?
Ans:- Sigmoid Curve

Quantitative Comparison of Growth

* Absolute Growth Rate: Measurement and comparison of total growth per

unit time.
* Relative Growth Rate: The growth of the given system per unit time
expressed on a common basis (e.g., per unit initial parameter).

Que. Leaves A and B both are different sizes but shows similar absolute
growth to give Leaves A1 and B1. However one of them shows much higher
relative growth rate. Which one and why?
 RGR (Relative Growth Rate) = Absolute Growth/ Initial Area.
 For leaf A, the initial area is 5 cm² and for leaf B, it is 50 cm².
 Therefore, we can calculate the relative growth rates as follows
 For leaf A: RGRA=5 cm2/5 cm2=1 cm2/cm2 (or 100%)
 For leaf B: RGRB=5 cm2/50 cm2=0.1 cm2/cm2 (or 10%)
 Thus, leaf A shows a much higher relative growth rate compared to leaf
B.

13.1.5 Conditions for Growth

 Plant growth is influenced by a variety of conditions, including internal

(intrinsic) and external (extrinsic) factors. These factors collectively drive
the development of the plant.
Factors Categorization

● Intrinsic Factors: (i) Genes: Genetic makeup of the plant and (ii)PGRs:
Plant Growth Regulators (Hormones).
● Extrinsic Factors: Light, Oxygen, Temperature, Water, Mineral Nutrients,
and Gravity.

Key Environmental Factors

(i) Light

● Required for tissue differentiation and the synthesis of photosynthetic

pigments.
● Necessary for the process of photosynthesis.
● It determines the direction of root and shoot growth (phototropism).

(ii) Temperature

● Optimum Temperature: 20–35°C.

● Temperatures above 45°C damage the protoplasm.
● Denaturation of enzymes takes place at high temperatures, which
retards growth.

(iii) Water

● Maintains the turgidity of plant cells.

● Essential for enzymatic activity in the protoplasm.
● Required for cell elongation.

(iv) Oxygen

● Essential for cellular respiration to release energy required for growth

activities.

(v) Mineral Nutrients (Macro and Micro essential elements)

 Typically required for the synthesis of protoplasm and act as a source of

energy.

(vi) Gravity

● Determines the direction, orientation, and movements of the root, stem,

and branches borne over them (geotropism
13.2 Differentation, Dedifferentiation and Redifferentiation

 Differentiation:- The cells derived from root apical and shoot-apical

meristems and cambium differentiate and mature to perform specific
functions. This act leading to maturation is termed as differentiation.

 During differentiation, cells undergo few to major structural changes

both in their cell walls and protoplasm.
 For example, to form a tracheary element, the cells would lose their
protoplasm. They also develop very strong, elastic, lignocellulosic
secondary cell walls, to carry water to long distances even under
extreme tension.

 Dedifferentiation:-
“The living differentiated cells that by now have lost the capacity to divide
can regain the capacity of division under certain conditions. This
phenomenon is termed as dedifferentiation.”

 For example, formation of meristems – interfascicular cambium and

cork cambium from fully differentiated parenchyma cells.

 Redifferentiation:-“such meristems/tissues are able to divide and

produce cells that once again lose the capacity to divide but mature to
perform specific functions is called rediiferentiation”
 Example: - a woody dicotyledenous plant that are the products of
redifferentiation.

Que:- How would you describe a tumour? What would you call the
parenchyma cells that are made to divide under controlled laboratory
conditions during plant tissue culture?
Ans: - A tumour is an abnormal, uncontrolled mass of tissue resulting
from excessive, autonomous cell division.
 In plant tissue culture, parenchyma cells induced to divide and form
an unorganized mass under controlled conditions are
called dedifferentiated cells, which collectively form a callus.
 NOTE:-
 The growth in plants is open, i.e., it can be indeterminate(meristematic
tissue) or determinate(for flowers, leaves, and in environmental stress)
 Now, we may say that even differentiation in plants is open, because
cells/tissues arising out of the same meristem have different structures at
maturity.
 The final structure at maturity of a cell/tissue is also determined by the
location of the cell within. E.g. Cells positioned away from root apical
meristems differentiate as root-cap cells, while those pushed to the
periphery mature as epidermis.

Que: Can you add a few more examples of open differentiation

correlating the position of a cell to its position in an organ?
Ans: shoot meristem cells becoming leaf primordia or vascular tissue and
cambium cells forming xylem (inward) or phloem (outward)

15.3 DEVELOPMENT
 Definition: - “All the changes that an organism goes through during
its life cycle from germination of seed to senescence.”
 Diagrammatic representation of the sequence of processes which
constitute the development of cell of a higher plant is given in fig. 3.
 It is also applicable to tissues/organs.

[Fig.3 Sequence of the developmental process in a plant cell]

 Plasticity

 Definition:- “ Plants follow different pathways in response to

environment or phases of life to form different kinds of structures. This
ability is called Plasticity.”
 E.g. heterophilly phenomenon
 In some plants the leaves of Juvenile plants are different in shape from
those in mature plants. E.g. In cotton, coriander and larkspur(fig.4).
Represent the heterophyllous development due to phases of life
 In some plants difference in shapes of leaves produced in air and those
produced in water. E.g. In Buttercup represent the heterophyllous
development due to environment (Fig.4)

[Fig.4 Heterophylly in (a) Larkspur and (b) buttercup]

Note: - Development (Both growth and differentiation) is under the control of

intrinsic and extrinsic factors.
 Intrinsic factor includes both intracellular(genetic) or intercellular
factors( Plant growth regulators)
 Extrinsic factors include light, temperature, water, oxygen and nutrient
etc.
15.4 Plant Growth Regulators

15.4.1 Characteristics
 PGRs are small, simple molecules of diverse chemical composition.
 Plant growth regulators are variously described as plant growth
substances, plant hormones or phytohormones.

Hormone Chemical nature

Auxins indole compounds (indole-3acetic acid, IAA)
Gibberellins Terpens (gibberellic acid, GA3)
Cytokinine Adenine derivatives(N6-furfurylamino purine, kinetin)
Abscisic acid, ABA Derivatives of carotenoids
Ethylene(C2H4) Gases

 PGRs can be divided into two groups based on their functions in plant
body:
 (i) Plant Growth Promoters: - involved in growth promoting activity: e.g.
cell division, cell enlargement, pattern formation, tropic growth,
flowering, fruiting and seed formation. E.g. auxins, gibberellins,
cytokinins.
 (ii) Plant Growth Inhibitors: - involved in growth inhibiting activity: e.g.
seed dormancy and abscission e.g. ABA, ethylene.
 Note: - The gaseous PGR ethylene could fit either of the groups but it is
largely an inhibitor of growth activities.

15.4.2 The Discovery of Plant Growth Regulators

 Each of the five major groups of PGRs has been discovered accidently.
(i) Auxins:-
 Discovered by:- Charles Darwin and his son Francis Darwin
 Observation: - they observed that coleoptiles of canary grass responded
to unilateral illumination by growing towards the light source
(phototropism).
 After a series of experiments, it was concluded that the tip of coleoptiles
was the site of transmittable influence that caused the bending of entire
coleoptile(fig.5)
 Isolated by: - F.W. Went from tips of coleoptiles of oat seedlings.
 (ii) Gibberellins:-

 Discovered by:- [Link](1926)

 Observation: - The ‘Bakane’ (foolish seedling) disease of rice seedlings,
was caused by fungal pathogen Gibberella fujikuroi
 [Link] reported the appearance of symptoms of the disease in rice
seedlings when they treated with sterile filtrates of the fungus.
 The active substance was later identified as gibberellic acid.

(iii) Cytokinins:-

 Discovered by:- [Link] and Miller

 Observation:- [Link] and his co-workers observed that from the
intermodal segment of tobacco stems the callus (a mass of
undifferentiated cells) proliferated only if, in addition to auxins the
nutrient medium was supplemented with one of the following: extracts
of vascular tissues, yeast extract, coconut milk or DNA.
 Miller et al. (1955) later identified and crystallised the cytokinesis
promoting active substance that they termed kinetin.

(iv) Abscisic acid:-

 Discovered by:- During 1960s three independent researches reported

the purification and chemical characterisation of three different kinds of
inhibitors: inhibitor-B, abscission II, and dormin. Later all the three were
proved to be chemically identical.
 It was named abscisic acid.

(Iv) Ethylene:-

 Discovered by:- H.H. Cousins (1910) confirmed the release of a volatile

substance from ripened oranges that hastened the ripening of stored
 unripened bananas . Later this volatile substance was identified as
ethylene, a gaseous PGR.

15.4.3 Physiological Effects of Plant Growth Regulators

[Link] Auxins

 Auxins(from Greek ‘auxein’: to grow) was first isolated from human

urine.
 Produced by the growing apices of the stems and roots from where they
migrate to the regions of their actions.
 Natural auxins like IAA (indole-3-acetic acid) and IBA (indole butyric acid)
have been isolated from plants.
 NAA(naphthalene acetic acid) and 2,4-D (2,4-dichlorophenoxyacetic
acid) are synthetic auxins
 All these auxins have been used extensively in agricultural and
horticultural practices.
 Effects:-
 They help to initiate rooting in stem cuttings- an application widely used
for plant propagation.
 Promote flowering e.g. in pineapples.
 They help to prevent fruit and leaf drop at early stages but promote the
abscission of older mature leaves and fruits.
 In higher plants, the growing apical bud inhibits the growth of the lateral
(axillary) buds, a phenomenon called apical dominance.
 Removal of shoot tips(decapitation) usually results in the growth of
lateral buds (Fig.6) therefore It is widely applied in tea plantations,
hedge-making.
 Auxins also induce parthenocarpy, e.g. in tomatoes.
 They are widely used as herbicides.
 2, 4-D widely used to kill dicotyledonous weeds, and does not affect
mature monocotyledonous plants.
 It is used to prepare weed-free lawns by gardeners.
 Auxin also controls xylem differentiation and helps in cell division.
[Link] Gibberellins

 There are more than 100 gibberellins reported from widely different
organisms such as fungi and higher plants.
 They are denoted as GA1, GA2, and GA3 and so on.
 However, Gibberellic acid (GA3) was one of the first gibberellins to be
discovered and remains the most intensively studied form.
 All GAs are acidic.

 Effects:-

 They cause an increase in length of axis and this ability is used to

increase the length of grape vines.
 Gibberellins cause fruits like apple to elongate and improve its shape.
 They also delay senescence. Thus the fruits can be left on the tree longer
so as to extend the market period.
 GA3 is used to speed up the malting process in brewing industry.
 Sugarcane stores carbohydrate as sugar in their stems. Spraying
sugarcane crops with gibberellins increases the length of the stem, thus
increasing the yield by as much as 20 tonnes per acre.
 Spraying juvenile conifers with GAs hastens the maturity, thus leading to
early seed production.
 Gibberellins also promote bolting (internode elongation just prior to
flowering) in beet, cabbage and many plants with rosette habit.

[Link] Cytokinins

 Cytokinins have specific effects on cytokinesis, and were discovered as

kinetin(a modified form of adenine, a purine) from the autoclaved
herring sperm DNA.
 Kinetin does not occur naturally in plants. Search for natural substances
with cytokinin-like activities led to the isolation of zeatin from corn-
kernels and coconut milk.
  Effects:-
 Since the discovery of zeatin, several naturally occurring cytokinins, and
some synthetic compounds with cell division promoting activity, have
been identified.
 Natural cytokinins are synthesised in regions where rapid cell division
occurs, for example root apices, developing shoot buds, young fruits etc.
 It helps to produce new leaves, chloroplasts in leaves, lateral shoot
growth and adventitious shoot formation.
 Cytokinins help overcome the apical dominance.
 They promote nutrient mobilisation which helps in delay of leaf
senescence.

[Link] Ethylene

 Ethylene is a simple gaseous PGR.

 It is synthesised in large amounts by tissues undergoing senescence and
ripening fruits.

 Effects:-
 Horizontal growth of seedlings, swelling of the axis and apical hooks
formation in dicot seedlings.
 Promotes senescence and abscission of plant organs especially of leaves
and flowers.
 Ethylene is highly effective in fruit ripening. It enhances the respiration
rate during ripening of the fruits.
 Respiratory climactic:- rise in the rate of respiration
 Ethylene breaks seed and bud dormancy, initiates germination in peanut
seeds, sprouting of potato tubers.
 Ethylene promotes rapid internode/petiole elongation in deep water
rice plants.
 It helps leaves/upper parts of the shoot to remain above water.
 Ethylene also promotes root growth and root hair formation, thus
helping the plants to increase their absorption surface.
 Ethylene is used to initiate flowering and for synchronising fruit-set in
pineapples.
 It also induces flowering in mango.
  Ethylene is one of the most widely used PGR in agriculture.

Ethephon
 The most widely used compound as source of ethylene is ethephon.
 Ethephon is aqueous solution is readily absorbed and transported
within the plant and releases ethylene slowly.
 Ethephon hastens fruit ripening in tomatoes and apples and
accelerates abscission in flowers and fruits(thinning of cotton, cherry
and walnut)
 It promotes female flowers in cucumbers thereby increasing the
yield.

[Link] Abscisic acid

 Abscisic acid (ABA) was discovered for its role in regulating abscission
and dormancy.
 It acts as a general plant growth inhibitor and an inhibitor of plant
metabolism.
 ABA inhibits seed germination.
 ABA stimulates the closing of stomata and increases the tolerance of
plants to various kinds of stresses. Therefore it is called the stress
hormone.
 ABA plays an important role in seed development, maturation and
dormancy.
 By inducing dormancy, ABA helps seeds to withstand desiccation and
other factors unfavourable for growth.
 In most situations, ABA acts as an antagonist to GAs.

 Plant hormones role in growth and development of plant could be

complimentary or antagonistic. These could be individualistic or
synergistic.
 Similarly, there are number of events in the life of a plant where more
than one PGR interact to affect that event, e.g. dormancy in seeds/buds,
abscission, senescence, apical dominance, Etc.
 Remeber, The role of PGR is of only one kind of intrinsic control. Along
with genomic control and extrinsic factors, they play an important role in
plant growth and development.
 Many of the extrinsic factors such as temperature and light control plant
growth and development via PGR.
 Some of such events could be vernalisation, flowering , dormancy, seed
germination, plant movement etc.