PLANT GROWTH AND DEVELOPMENT

• Development is the sum of two processes: growth and differentiation.

• Growth, differentiation and development are very closely related events in the life of a plant.
• Development of a mature plant from a zygote (fertilised egg) follow a precise and highly ordered
succession of events.
• During this process a complex body organisation is formed that produces roots, leaves, branches,
flowers, fruits, and seeds, and eventually they die .
• The first step in the process of plant growth is seed germination.
• The seed germinates when favourable conditions for growth exist in the environment.
• In absence of such favourable conditions the seeds do not germinate and goes into a period of
suspended growth or rest.
• Once favourable conditions return, the seeds resume metabolic activities and growth takes place.
• Development in plants (i.e., both growth and differentiation) is under the control of intrinsic or
internal and extrinsic or external factors.
• Intrinsic factors includes both intracellular (genetic) or intercellular factors (chemicals such as plant
growth regulators)
• Extrinsic factors includes light, temperature, water, oxygen, nutrition, etc.
Germination and seedling development in bean
 GROWTH

• Growth is regarded as one of the most fundamental and conspicuous characteristics of a

living being.
• Growth can be defined as an irreversible permanent increase in size of an organ or its parts
or even of an individual cell.
• Generally, growth is accompanied by metabolic processes (both anabolic and catabolic), that
occur at the expense of energy.
• Example - expansion of a leaf is growth.
 How would you describe the swelling of piece of wood when placed in water?
• The process involving absorption of water by a substance (solid particles) is referred to as imbibition.
Swelling of a piece of wood when placed in water is due to imbibition. Imbibition is a reversible process.

Reason for not considering a swelled piece of wood as growth

• Growth is characterised by an irreversible, permanent increase in the size of an individual cell, organ or
its parts, an organism over a period of time.
• Swelling up of the piece of wood is not considered as growth.
• There is no increase in the dry weight of the wood.
• Also, the temporary increase in size due to imbibition can be reversed.
 Plant Growth Generally is Indeterminate

• Plant growth is unique because plants retain the capacity for unlimited growth throughout their
life.
• This ability of the plants is due to the presence of meristems at certain locations in their body.
• The cells of such meristems have the capacity to divide and self-perpetuate.
• The product, however, soon loses the capacity to divide and such cells make up the plant body.
• This form of growth wherein new cells are always being added to the plant body by the activity of
the meristem is called the open form of growth.
What would happen if the meristem ceases to divide? Does this ever happen?
Meristematic tissues
Meristematic tissues are the tissues which have the ability to divide.
• Root and shoot apical meristems are responsible for the primary growth of the plants
and principally contribute to the elongation of the plants along their axis.
• In dicotyledonous plants and gymnosperms, the lateral meristems, vascular cambium
and cork-cambium appear later in life. These are the meristems that cause the
increase in the girth of the organs in which they are active. This is known as secondary
growth of the plant.

Cessation of meristematic division

If root and shoot apical meristems stop dividing, the primary growth of roots and shoots
will get hampered. The seed after germination wouldn't grow into seedling and further.
Non-functional lateral meristem would lead to absence of secondary growth in the plant.

Stoppage of meristematic activity

Meristematic tissues, after primary growth, lose their ability of division when they get
differentiated and specialised to perform a particular function. Such specialised tissues
are termed as permanent tissues.
 Growth is Measurable

• Growth, at a cellular level, is principally a consequence of increase in the amount of protoplasm.

• Since increase in protoplasm is difficult to measure directly, one generally measures some quantity which
is more or less proportional to it.
• The various parameters to measure growth are : increase in fresh weight, dry weight, length, area,
volume and cell number.
• Examples
• Growth expressed as increase in cell number - One single maize root apical mersitem can give rise to
more than 17,500 new cells per hour
• Growth expressed as increase in size of the cell - Watermelon may increase in size by upto 3,50,000
times.
• Growth expressed as Increase in length - Growth of a pollen tube
• Growth expressed as increase in surface area - Growth in a dorsiventral leaf
 Phases of Growth

The period of growth is generally divided into three phases :

1) Meristematic phase
• The constantly dividing cells, both at the root apex and the shoot apex, represent the
meristematic phase of growth.
• The cells in this region are rich in protoplasm, possess large conspicuous nuclei. Their cell walls
are primary in nature, thin and cellulosic with abundant plasmodesmatal connections.
1) Phase of elongation
• The cells proximal (just next, away from the tip) to the meristematic zone represent the phase
of elongation.
• Increased vacuolation, cell enlargement and new cell wall deposition are the characteristics of
the cells in this phase.
1) Phase of maturation
• Further away from the apex, i.e., more proximal to the phase of elongation, lies the portion of
axis which is undergoing the phase of maturation.
• The cells of this zone, attain their maximal size in terms of wall thickening and protoplasmic
modifications
 Growth Rates
• The increased growth per unit time is termed as growth rate.
• Rate of growth can be expressed mathematically.
• The growth rate shows an increase that may be
(1) arithmetic or
(2) geometrical
1) Arithmetic growth
• In arithmetic growth, following mitotic cell division, only one daughter cell
continues to divide while the other differentiates and matures.
• The simplest expression of arithmetic growth is exemplified by a root
elongating at a constant rate.
• On plotting the length of the organ against time, a linear curve is obtained.
• Mathematically, it is expressed as
2) Geometrical growth
• In most systems, the initial growth is slow (lag phase), and it increases
rapidly thereafter – at an exponential rate (log or exponential phase).
• Here, both the progeny cells following mitotic cell division retain the ability
to divide and continue to do so.
• However, with limited nutrient supply, the growth slows down leading to a
stationary phase.
• If we plot the parameter of growth against time, we get a typical sigmoid or
S-curve .
• A sigmoid curve is a characteristic of living organism growing in a natural
environment.
• It is typical for all cells, tissues and organs of a plant.
• The exponential growth can be expressed as

• Here, r is the relative growth rate and is also the

measure of the ability of the plant to produce new
plant material, referred to as efficiency index.
• Hence, the final size of W1 depends on the initial
size, W0
Quantitative comparisons between the growth of living system can be made in two ways :
(i) Absolute growth rate - measurement and the comparison of total growth per unit time is called
the absolute growth rate.
(ii) Relative growth rate - The growth of the given system per unit time expressed on a common
basis, e.g., per unit initial parameter is called the relative growth rate.

• In the given figure , both leaves A and B have increased their area by 5cm2 in a given time to
produce A1 , B1 leaves.

• The absolute growth rate of both the leaves is the

same. Both A and B increase in their area by 5 cm2.
• Relative growth rate is more for leaf A ,since it
almost doubles.
 Conditions for Growth

• Water - The plant cells grow in size by cell enlargement which in turn requires water. Turgidity of cells
helps in extension growth. Thus, plant growth and further development is intimately linked to the water
status of the plant. Water also provides the medium for enzymatic activities needed for growth.
• Oxygen - Oxygen helps in releasing metabolic energy essential for growth activities.
• Nutrients - Nutrients as very essential elements for growth. Macro and micro essential elements are
required by plants for the synthesis of protoplasm and act as source of energy.
• Temperature - Every plant organism has an optimum temperature range best suited for its growth. Any
deviation from this range could be detrimental to its survival.
• Light and gravity - Environmental signals such as light and gravity also affect certain phases/stages of
growth.
 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 a very strong, elastic, lignocellulosic secondary cell walls, to carry water to long
distances even under extreme tension.
TRACHEARY ELEMENT:
LOSE THE PROTOPLASM
STRONG
ELASTIC
TO CARRY WATERR TO LONG DISTANCE
 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 secondary meristems – interfascicular cambium and cork cambium from
fully differentiated parenchyma cells

REDIFFERENTIATION

• The cells of secondary meristems/tissues are able to divide and produce cells that once again lose the
capacity to divide but mature to perform specific functions. Such cells are called redifferentiated and the
phenomenon is called redifferentiation.
• Example in dicot plants interfascicular cambium give rise to secondary xylem and secondary phloem,
Cork cambium produces cork or phellem and secondary cortex or phelloderm .
• List some of the tissues in a woody dicotyledenous plant that are the products of redifferentiation.
- Secondary xylem , secondary phloem, cork or phellem and secondary cortex or phelloderm

• How would you describe a tumour?

- Tumour is a product of dedifferentiation

• What would you call the parenchyma cells that are made to divide under controlled laboratory
conditions during plant tissue culture?

- It is dedifferentiation and the cells are dedifferentiated cells

 Differentiation in plants is open
• 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.
• For example, cells positioned away from root apical meristems differentiate as root-cap cells,
while those pushed to the periphery mature as epidermis.
 DEVELOPMENT

• Development is a term that includes all changes that an organism goes through during its
life cycle from germination of the seed to senescence.

Sequence of developmental processes in a plant cell

 Plasticity

• 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 heterophylly in cotton, coriander and larkspur.
• Heterophylly in different phases of life - The leaves of the juvenile plant are different in shape from
those in mature plants. Eg . Larkspur
• Heterophylly in response to environment - Difference in shapes of leaves produced in air and those
produced in water in buttercup also represent the heterophyllous development due to environment.
This phenomenon of heterophylly is an example of plasticity

Heterophylly in larkspur
Heterophylly in buttercup
PLANT GROWTH REGULATORS ( PLANT HORMONES,PHYTOHORMONES)

 The plant growth regulators (PGRs) are small, simple molecules of diverse
chemical composition.
 According to the chemical composition they are grouped into:
(1) Indole compounds (indole-3-acetic acid, IAA)
(2) Adenine derivatives (N6-furfurylamino purine, kinetin)
(3) Carotenoid derivatives (abscisic acid, ABA)
(4) Terpenes (gibberellic acid, GA3)
(5) Gases (ethylene, C2H4).
 The PGRs can be broadly divided into two groups based on their functions in a plant body.
(1) Plant growth promoters
 Involved in growth promoting activities, such as cell division, cell enlargement, pattern formation,
tropic growth, flowering, fruiting and seed formation.
 E.g., auxins, gibberellins and cytokinins.

(2) Plant growth inhibitors

 Involved in various growth inhibiting activities such as dormancy ( state of reduced activity . Eg seed
dormancy ) and abscission ( shedding of leaves,fruits,flowers etc).
 Play an important role in plant responses to stresses of biotic and abiotic origin.
 Eg . Abscisic acid
 The gaseous PGR, ethylene, could fit either of the groups, but it is largely an inhibitor of growth
activities.
 The Discovery of Plant Growth Regulators
1) Auxin
 Charles Darwin and his son Francis Darwin observed
that the 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 coleoptile was the site of transmittable influence
that caused the bending of the entire coleoptile.
 Auxin was isolated by F.W. Went from tips of coleoptiles
of oat seedlings.
2) Gibberellic acid
 The ‘bakanae’ (foolish seedling) disease of rice seedlings,
was caused by a fungal pathogen Gibberella fujikuroi.
 E. Kurosawa (1926) reported the appearance of symptoms
of the disease in rice seedlings when they were treated
with sterile filtrates of the fungus.
 The active substances were later identified as gibberellic
acid.
 3) Kinetin
 F. Skoog and his co-workers observed that from the internodal segments of tobacco
stems the callus (a mass of undifferentiated cells) proliferated only if, in addition to
auxins the nutrients medium was supplemented with one of the following: extracts of
vascular tissues, yeast extract, coconut milk or DNA.
 Miller (1955), later identified and crystallised the cytokinesis promoting active
substance that they termed kinetin.
4) Abscisic acid
 During mid-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
(ABA).
 5) Ethylene

 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.
Physiological Effects of Plant Growth Regulators
Auxins

 Auxins (from Greek ‘auxein’ : to grow) was first isolated from human urine.
 The term ‘auxin’ is applied to the indole-3-acetic acid (IAA), and to other natural
and synthetic compounds having certain growth regulating properties.
 They are generally produced by the growing apices of the stems and roots, from
where they migrate to the regions of their action.
 Auxins like IAA and indole butyric acid (IBA) have been isolated from plants.
 NAA (naphthalene acetic acid) and 2, 4-D (2, 4-dichlorophenoxyacetic) are
synthetic auxins.
 All these auxins have been used extensively in agricultural and horticultural
practices.
 They help to initiate rooting in stem cuttings, an application widely used for plant
propagation.
 Auxins 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 most 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.
 It is widely applied in tea plantations, hedge-making.
 Auxins also induce parthenocarpy ( formation of fruits without fertilisation)
e.g., in tomatoes.
 They are widely used as herbicides. 2, 4-D, widely used to kill dicotyledonous weeds, 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.
 APICAL DOMINANCE IN PLANTS

a) Plant with apical bud intact

b) Plant with apical bud removed

- Growth of lateral buds into
branches after decapitation
 Gibberellins
 Gibberellins are another kind of growth promoting PGR.
 There are more than 100 gibberellins reported from widely different organisms
such as fungi and higher plants.
 They are denoted as GA1, GA2, 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.
 They produce a wide range of physiological responses in the plants.
 Their ability to cause an increase in length of axis is used to increase the length of
grapes stalks.
 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 crop with gibberellins increases the
length of the stem, thus increasing the yield .
 Spraying juvenile conifers with GAs hastens the maturity
period, thus leading to early seed production.
 Gibberellins also promotes bolting (internode elongation just
prior to flowering) in beet, cabbages and many plants with
rosette habit.
 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. (DNA is isolated which is
sterilized using steam at high pressure which is called autoclaved herring sperm DNA. From this, kinetin
was isolated which has ability to induce cell division. )
 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.
 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 the delay of leaf senescence.
 Ethylene
 Ethylene is a simple gaseous PGR.
 It is synthesised in large amounts by tissues undergoing
senescence and ripening fruits.
 Influences of ethylene on plants include horizontal growth of
seedlings, swelling of the axis and apical hook formation in
dicot seedlings.
 Ethylene 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.
 This rise in rate of respiration is called respiratory climactic.
 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.
 Since ethylene regulates so many physiological processes, it is one of the most
widely used PGR in agriculture.
 The most widely used compound as source of ethylene is ethephon.
 Ethephon in an 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, walnut).
 It promotes female flowers in cucumbers thereby increasing the yield.
 Abscisic acid
 Abscisic acid (ABA) was discovered for its role in regulating abscission and dormancy.
 It also has other wide ranging effects on plant growth and development.
 It acts as a general plant growth inhibitor and an inhibitor of plant metabolism.
 ABA inhibits seed germination.
 ABA stimulates the closure of stomata and increases the tolerance of plants to various
kinds of stresses.
 Therefore, it is also 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.
 The role of PGR could be complimentary or antagonistic.
 These could be individualistic or synergistic.( When two or more hormones act together and
contribute to the same function)
 Similarly, there are a 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.
 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 movements, etc.