CHAPTER 15: PLANT GROWTH AND DEVELOPMENT
All plant organs are made up of a variety of tissues;
All cells of a plant are descendents of the zygote.
The zygote develops into a mature plant through growth and differentiation resulting in the formation of roots,
leaves, branches, flowers, fruits, and seeds. Then they eventually die.
GROWTH:
Growth is defined as an irreversible permanent increase in size of an organ or its parts or an individual cell.
It is one of the most fundamental and conspicuous characteristics of a living being.
It involves metabolic (both anabolic and catabolic) processes that consume energy. For example, expansion
of a leaf is growth.
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.
Meristematic cells 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.
Open form of growth: The form of growth where new cells are always being added to the plant body by the
activity of the meristem is called the open form of growth.
Primary growth of the plants: It occurs due to Root apical meristem and shoot apical meristem. It causes
the elongation of the plants along their axis.
Secondary growth of the plants (In gymnosperms & dicots): It occurs due to the lateral meristems,
vascular cambium and cork-cambium. These meristems cause the increase in the girth of the organs.
Growth is Measurable:
At cellular level, growth occurs due to increase in the amount of protoplasm.
Increase in protoplasm is difficult to measure directly. So growth is measured by parameters like increase in
fresh weight, dry weight, length, area, volume & cell number. E. g.
a. In cell number: One single maize root apical meristem can give rise to more than 17,500 new cells per
hour.
b. In cell size: Cells in a watermelon may increase in size by up to 3,50,000 times.
c. In length: Growth of a pollen tube.
d. In surface area: Growth in a dorsi-ventral leaf.
Detection of zones of elongation by the parallel line technique.
Zones A, B, C, D immediately behind the apex have elongated most.
Diagrammatic representation of locations of root apical meristem, shoot apical meristem and vascular
cambium.
Arrows exhibit the direction of growth of cells and organ.
Phases of Growth
The period of growth is generally divided into three phases: Meristematic, Elongation and Maturation.
Meristematic phase: It occurs in the meristems at the root apex & the shoot apex. Cells in this region have
rich protoplasm and large conspicuous nuclei. Cell walls are primary, thin & cellulosic with abundant
plasmodesmata.
Elongation phase: It occurs in the cells proximal (just next, away from the tip) to the meristematic zone.
The cells have increased vacuolation, cell enlargement and new cell wall deposition.
Maturation phase: It occurs in the cells further away from the apex, i.e., more proximal to the phase of
elongation. The cells attain their maximal size in terms of wall thickening and protoplasmic modifications.
Growth Rates
The increases growth per unit time is termed as growth rate.
The rate of growth can be expressed mathematically.
The growth rate shows an increase that may be arithmetic or geometrical.
Diagrammatic representation of: (a) Arithmetic (b) Geometric growth and (c) Stages during embryo
development showing geometric and arithmetic phases
Arithmetic growth: In this, following mitotic cell division, only one daughter cell continues to divide while
the other differentiates and matures. It is represented as below:
On plotting the length of the organ against time, a linear curve is
obtained. Mathematically, it is expressed as
Lt = L0 + rt
Lt = length at time ‘t’
L0 = length at time ‘zero’
r = growth rate / elongation per unit time.
Constant linear growth, a plot of length L against time t
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 daughter cells continue and retain the ability of mitotic cell division.
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 (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 W1 = W0 ert
W1 = final size (weight, height, number etc.)
W0 = initial size at the beginning of the period
r = relative growth rate
t = time of growth
e = base of natural logarithms
An idealized sigmoid growth curve typical of cells in culture, and many higher plants and plant organs
Relative growth rate (r) is 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 also be made in two ways:
a. Absolute growth rate: Measurement and the comparison of total growth per unit time
b. Relative growth rate Measurement of growth of the given system per unit time expressed on a
common basis, e.g., per unit initial parameter.
Diagrammatic comparison of absolute and relative growth rates. Both leaves A and B have increased their
area by 5 cm2 in a given time to produce A1, B1 leaves.
Conditions (essential elements) for Growth
1. Water: It is essential for cell enlargement. Turgidity of cells helps in extension growth. Water also provides
the medium for enzymatic activities needed for growth.
2. Oxygen: It helps to release metabolic energy for growth.
3. Nutrients: Macro & micro elements are needed for the synthesis of protoplasm and act as source of energy.
4. Temperature: Plants have an optimum temperature at which growth is maximum. Deviation from this range
could be detrimental to its survival.
5. Light & gravity: Affect certain phases/stages of growth.
DIFFERENTIATION, DEDIFFERENTIATION AND REDIFFERENTIATION
A. Differentiation is the process in which the cells from root apical & shoot-apical meristems and cambium
differentiate and mature to perform specific functions.
In this process, cell walls and protoplasm undergo major structural changes. For example, to form a
tracheary element, the cells would lose their protoplasm. They also develop a very strong, elastic,
lignocellulosic secondary cell wall, to carry water to long distances even under extreme tension.
The capacity of cell division is lost.
B. Dedifferentiation Under certain conditions, living differentiated cells regain the capacity of division. This is
called dedifferentiation. E.g. formation of meristems (interfascicular cambium & cork cambium) from
differentiated parenchyma cells
C. Redifferentiation The dedifferentiated cells can divide and produce cells that again lose the capacity to
divide but mature to perform specific functions. It is called redifferentiation.
Plant growth is open, i.e., it can be indeterminate or determinate. Differentiation in plants is also 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. E.g. cells positioned
away from root apical meristems differentiate as root-cap cells, while those pushed to the periphery mature as
epidermis.
DEVELOPMENT
It is a process that includes all changes in the life cycle of an organism from seed germination to
senescence.
It is the sum of growth and differentiation.
It is also applicable to tissues/organs.
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 due to phases of life: E.g. In cotton, coriander and larkspur, the leaves of the juvenile
plants and mature plants are different in shape.
Heterophylly due to environment: E.g. Difference in shapes of leaves produced in air and those
produced in water (e.g. buttercup).
Factors controlling the development:
Intrinsic factors: Include intracellular (genetic) or intercellular factors (such as plant growth regulators).
Extrinsic factors: Include light, temperature, water, oxygen, nutrition, etc.
Heterophylly in (a) larkspur and (b) buttercup
PLANT GROWTH REGULATORS
The plant growth regulators (PGRs) are small, simple molecules of diverse chemical composition. They
could be:
Indole compounds (indole-3-acetic acid, IAA);
adenine derivatives (N6-furfurylamino purine, kinetin)
derivatives of carotenoids (abscisic acid, ABA)
terpenes (gibberellic acid, GA2)
Gases (ethylene, C2H4)
Based on the functions, PGRs are divided into 2 groups.
a. Plant growth promoters: For growth promoting activities like cell division & enlargement, tropic growth,
pattern formation, flowering, fruiting & seed formation. E.g. auxins, gibberellins and cytokinins.
b. Plant growth inhibiters: Involved in growth inhibiting activities like dormancy & abscission. Take part in
plant responses to wounds & stresses of biotic and abiotic origin. E.g. abscisic acid, Ethylene (Ethylene fits
either of the groups, but it is largely a growth inhibitor).
Physiological Effects of Plant Growth Regulators
Auxins
Discovery: Charles Darwin & his son Francis Darwin observed that the coleoptiles of canary grass responded
to unilateral illumination by growing towards the light source (phototropism). It was concluded that the tip of
coleoptile caused the bending of the entire coleoptile.
F.W. Went isolated Auxin (Greek ‘auxein’: to grow) from tips of coleoptiles of oat seedlings.
The term ‘Auxins’ is used to indicate indole compounds (e.g. indole-3-acetic acid, IAA) and other
natural and synthetic compounds having growth regulating properties.
Auxins are produced by the growing apices of stems & roots, from where they migrate to 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.
Functions of auxins:
They help to initiate rooting in stem cuttings, an application widely used for plant propagation.
Promote flowering. E.g. in pineapples.
To prevent fruit and leaf drop at early stages.
Promote the abscission of older mature leaves and fruits.
Induce parthenocarpy, 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.
Controls xylem differentiation and helps in cell division.
Apical dominance: In higher plants, the growing apical bud inhibits the growth of the lateral (axillary)
buds. It is known as apical dominance. Removal of shoot tips (decapitation) results in the growth of
lateral buds. It is widely applied in tea plantations, hedge-making.
Gibberellins
Discovery: E. Kurosawa reported the appearance of symptoms of the disease, ‘bakane’ (foolish seedling- a fungal
disease of rice seedlings, caused by Gibberalla fujikuroi) in uninfected rice seedlings treated with sterile filtrates of
the fungus. The active substances were identified as gibberellic acid.
There are more than 100 gibberellins in fungi and higher plants. They are denoted as GA1, GA2, GA3 and so
on.
All GAs are acidic.
Gibberellic acid (GA3 or Terpenes) is one of the first discovered and most intensively studied gibberellins.
Functions:
They cause an increase in length of axis. So they are used to increase the length of grapes stalks.
To elongate and improve the shape of fruits such as apple.
They 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 malting process in brewing industry.
Sugarcane stores sugar in stems. Spraying sugarcane crop with gibberellins increases the length of the
stem. It increases the yield by as much as 20 tonnes per acre.
Spraying juvenile conifers with GAs hastens the maturity period. It leads to early seed production.
For bolting (internode elongation just prior to flowering) in beet, cabbages and many plants with rosette
habit.
Cytokinins
Discovery: F. Skoog and co-workers observed that from the intermodal segments of tobacco stems the callus (a
mass of undifferentiated cells) proliferated only if the nutrients medium was supplemented with extracts of
vascular tissues, yeast extract, coconut milk or DNA.
Skoog & Miller later identified and crystallized the active substance and termed as kinetin.
Cytokinins were discovered as kinetin (N6-furfurylamino purine - an Adenine derivative) from the
autoclaved herring sperm DNA.
Kinetin does not occur naturally in plants.
Zeatin (from corn-kernels and coconut milk) is the natural substances with cytokinin-like activities.
There are some synthetic compounds with cell division promoting activity.
Natural cytokinins are synthesized in regions where rapid cell division occurs (root apices, developing
shoot buds, young fruits etc).
Functions:
They play a role in cytokinesis.
They help to produce new leaves, chloroplasts in leaves, lateral shoot growth and adventitious shoot
formation.
They help overcome the apical dominance.
They promote nutrient mobilization which helps in the delay of leaf senescence.
Ethylene (C2H4)
Discovery: Cousins 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.
Ethylene is a simple gaseous PGR.
It is synthesized in large amounts by tissues undergoing senescence and ripening fruits.
Functions:
It has influences on horizontal growth of seedlings, swelling of the axis and apical hook formation in
dicot seedlings.
It promotes senescence and abscission of plant organs especially of leaves and flowers.
It 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.
It breaks seed and bud dormancy, initiates germination in peanut seeds, sprouting of potato tubers.
Promotes rapid Internode/petiole elongation in deep water rice plants. It helps leaves/upper parts of the
shoot to remain above water.
Promotes root growth and root hair formation. This helps the plants to increase their absorption surface.
Used to initiate flowering and for synchronizing fruit-set in pineapples. It also induces flowering in
mango.
Since ethylene regulates so many physiological processes, it is widely used PGR in agriculture. The most
widely used 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 (ABA)
Discovery: During mid-1960s, three independent researchers reported 3 kinds of inhibitors: inhibitor-B,
abscission II & dormin. It was proved that these were chemically identical. Now it was named as abscisic acid.
ABA is the derivatives of carotenoids.
It was discovered for its role in regulating abscission and dormancy. But it also has other wide ranging
effects on plant growth and development.
Functions:
It acts as a general plant growth inhibitor and an inhibitor of plant metabolism.
It inhibits seed germination.
It stimulates the closure of stomata in the epidermis and increases the tolerance of plants to various kinds
of stresses. Therefore, it is also called the stress hormone.
It has an important role in seed development, maturation and dormancy. Seed dormancy by ABA helps
to withstand desiccation and other factors unfavourable for growth.
Interactions of PGRs
For growth, differentiation and development of plants, one or more PGR play individualistic or synergistic
role. Such roles may be complimentary or antagonistic.
PGRs interact to affect events such as dormancy in seeds/ buds, abscission, flowering, senescence,
vernalisation, apical dominance, seed germination, plant movements etc.
In most situations, ABA acts as an antagonist to GAs.
Factors influencing the action of Plant Growth Regulators:
Intrinsic factor: Genomic control.
Extrinsic factors: Light and Temperature.