PLANT GROWTH AND MOVEMENT
Growth is a process in which there is increase in the size by cell division and
enlargement accompanied by the formation of new cellular material. It results
in the irreversible increase in size, length area or volume accompanied by
increase in dry weight. In living beings, growth is internal or intrinsic in contrast
to extrinsic growth observed in some non-living objects. Irreversible increase in
size, mass or volume of living beings is an external manifestation of growth.
Further growth is a quantitative phenomenon which can be measured in
relation to time.
Development can be defined as a process in which there is a sequence of
qualitative changes, towards a higher or more complex state. It includes all the
changes an organism undergoes from the time of birth till death. In
development plant shows a regular sequence of seed germination, growth,
differentiation, maturation, seed formation and senescence.
CHARACTERISTICS OF GROWTH
Primary growth is formation of primary permanent tissues and organs. It is
caused by activity of apical and intercalary meristems.
Secondary growth is increase in girth. It occurs by two types of lateral
meristems, vascular cambium and cork cambium.
Unlimited growth is growth that continues throughout life as it occurs in case
of root and stem. Limited growth is that growth which stops after some time,
e.g. leaves, flowers, fruits
In higher plants the growth involves three steps or phases
- Phase of cell division
- Phase of cell enlargement
- Phase of cell maturation
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GROWTH RATES
The expression of increased growth per unit time is called growth rate.
Growth rate shows two types of increase
Arithmeticgrowth : It is a type of growth in which the rate of growth is constant
and increase in growth occurs in arithmetic progression 2,4,6,8 etc. Here after
mitosis, only one daughter cell continuous to divide. Other take part in
differentiation and maturation i.e. root elongating at constant rate. Here a
linear curve is obtained with value
Lt = Lo + rt
Lt : Length at time t
Lo = Length at time ‘zero’
R : growth rate
Geometric growth: It is quite common in unicellular organisms when growth in
nutrient rich medium. Here every cell divides. The daughter grow and divide.
The granddaughter repeat the process and so on. Number of cells is initially
small so that initial growth is slow. Later on, there is rapid growth at
exponential rate. An embryo log or exponential growth. An embryo initially
shows geometrical growth in cells, but later it passes into arithmetic phase
GROWTH CURVE
The exponential growth curve can be represented by equation
Wt = Woert
Wt : Final size
Wo : Initial size at beginning of the time period
r : growth rate
2
t : time of growth
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V.H. Blackman (1919) suggested ‘r’ might be used as a measure of the ability of
a plant to produce new plant material and called it efficiency index
The rate of growth is not uniform for a cell, organ or an organism. If the growth
rates of a plant are plotted against time on a paper, a sigmoid growth curve or
S-shaped growth curve is obtained. It consists of lag phase or slow phase, log
phase or exponential growth phase, rate becomes more rapid and senescence
phase or declining phase. A steady phase or stationary phase(limited growth)
occurs at the end.
MEASUREMENT OF GROWTH
Growth is measured with the help of auxometer, horizontal microscope and
crescograph ( developed by Sir [Link])
Growth can be measured by measuring increase in weight area and volume in
fresh and dry form. In yeast and bacteria, increase in length, number of cells
can be taken into account.
In arc auxometer, when growth occurs, stem increases in length. Due to
pressure of weight wheel rotates gradually. This leads to movement of
indicator on arc scale. This reading on graduated arc provides information
about the rate of growth.
FACTOR INFLUENCING GROWTH AND DEVELOPMENT
1. LIGHT
It controls photosynthesis, transpiration, pigment formation, tissue
differentiation seed germination, etc. Ultraviolet ray are inhibitory. They induce
rosette formation. Red light promotes growth but plant organs are soft. Blue
light produce normal slightly less growth. Low light intensity increases
intermodal length and leaf size. High light intensity decreases their size but
allows more Certain seed germinate only in the presence of light, eg.
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Viscumalba. Seeds of lettuce and tobacco require red light for their
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germination. Certain other seeds e.g. onion cannot grow in the presence of
light. Light induces tissue differentiation in plants. Light induced tissue
differentiation is called photo-morphogenesis. It involves photoreceptor
pigment system called phytochrome. This can be observed by growing
seedlings in dark. The seedlings are pale and weak. The phenomenon is
etiolation and seedlings known as etiolated seedlings.
Etiolated seedlings have:
(i) Pale colour
(ii) Long internodes
(iii) Subterminal hook
(iv) Small pale leaver
(v) No mechanical tissue
(vi) Small and few roots
2. TEMPERATURE
Minimum, optimum and maximum temperature for plant growth are 2 O -5O C.
20O -30O C., 40O -50OC. Lower temperature inactivates enzymes. It increase
density of protoplasm. There is little absorption of water from soil. Internal
water can be frozen. Chilling and freezing injury may occur. Cold storage or
refrigerator is due to inhibition of growth of plant organs, reduced metabolic
activity influences seed germination, growth and seasonal development of
plants. Some plants require low temperature treatment for their growth and
flowering. The phenomenon is called vernalisation.
High temperature of 45OC and above reduces growth due to excessive
transpiration, denaturation of enzymes and coagulation of protoplasm. Its
effect is externally visible as leaf scorch, heat canker and desiccation.
3. WATER
Optimum hydration is essential for cell elongation, cell turgidity and
functioning of metabolic machinery. Growth is reduced even in slight
deficiency of water. It however, promotes differentiation. Water stress
completely stops growth.
4. OXYGEN
It is required for release of energy during respiration. Energy is needed for
anabolic activities of growth
5. SOIL
It provides minerals, some growth stimulants and water
6. CARBON DIOXIDE
Essential for photosynthesis and hence nutrition. However, in higher
concentration carbon dioxide inhibits growth
7. HERIDITARY POTENTIALITY
Growth and differentiation is genetically determined
8. GRAVITY
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It determines the direction of root and shoot growth.
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9. STRESS
Stress factors like excess or deficiency of salts, low or high temperature, excess
and deficiency of water have a detrimental effect on growth.
10. NUTRITION
It provides raw material for growth and differentiation as well as source of
energy.
11. GROWTH REGULATORS
DIFFERENTIATION, DEDIFFERENTIATION, REDIFFERENTIATION
Differentiation: It is the permanent qualitative changes in structure, chemistry
and physiology of cell wall and protoplasm of cells, their tissues and organs. It
is caused by repression of some genes.
Dedifferentiation: It is the process of despecialisation of differentiated living
cells so that they regain the capacity to divide and for new cells. A
dedifferentiated tissue can act as meristem [Link] vascular
cambium, cork cambium, wound cambium. In culture experiments,
parenchyma cells dedifferentiate to produce a mass of dividing cells called
callus.
Redifferentiation : Structural, chemical and physiological specialization of cells
derived from dedifferentiation. It is similar to differentiation of cells and tissues
formed by primary meristems. Secondary phloem, secondary xylem, cork,
secondary cortex are some of the tissues formed through redifferentiation.
PHOTOPERIODISM
Photoperiodism is the effect of duration of light and darkness on the growth,
development and behavior of organisms. It is especially connected with
flowering of plants, formation of underground storage organs, leaf fall etc. the
effect of photoperiods on flowering was discovered by Garner and Allard (
1920)incase of maryland of mammoth variety of Tobacco on the basis of their
response to photoperiods.
Plants are of the following types
1. Long Day plants (LDP)
Plants flowers after receiving light above a critical length e.g. wheat oat,
sugarbeet, spinach, radish, lettuce, poppy.
2. Short Day Plants (SDP)
Plants flower only when they receive light below a critical length e.g.
tobacco, potato, Xanthium, rice, dahlia, chrysanthemum, soya bean
3. Day Neutral Plants (DNP)
Photoperiods have no impact on these plants which come to flower after
completing their vegetative growth e.g. tomato, maize, cotton, sunflower,
cucumber
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4. Short-long Day Plants (SLDP)
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Short photoperiod for initiation of lower and long photoperiods for
blossoming e.g. Campanula medium, Trifoliumrepens.
5. Long-short Day Plants ( LSDP)
Long photoperiods for initiation of flowering and short photo photoperiod
for blossoming e.g. Bryophyllum, cestrum
Day Neutral Plants are more common in tropical areas. They can be made to
flower throughout year. In subtropical and temperate areas short day plants
flower during autumn and spring while long day plants flower during summer.
Critical day length is that continuous duration of light which must always be
exceed in short day light.
Critical dark period is that continuous duration of darkness. Long day plants do
not require any continuous dark period. They are, therefore, called short night
plants (SNP). Short day plants are known as long night plants (LNO)
The phenomenon of perceiving appropriate light periods and obtaining the
stimulus of flowering is known as photoperiodic induction. Photoperiodic
induction requires
(i) A minimum vegetative growth
(ii) A minimum number of appropriate photoperiod
(iii) Photoreceptor pigment called phytochrome
(iv) Fully developed leaves which receive the stimulus of light
VERNALISATION
Vernalisation means ability of low temperature treatment to convert cereal
into spring cereal
Site of vernalisation is apical meristem or all the meistematic cells. E.g. shoot,
tip, embryo tips, root apex
As a result of vernalization, a flowering hormone called vernaline is formed but
vernaline has never been isolated.
Once a plant is vernalized, it can be devernalized by exposing the plant to
temperature of 30OC or above. For establishing vernalization, plant should be
kept at 20OC for 4-5 days
CONDITIONS NECESSARY FOR VERNALISATION
Temperature : 1 – 6OC is the optimum temperature
Duration : 1 to 1.5 months low temperature treatment is necessary
Oxygen : As vernalisation is aerobic process, so it requires O2
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Water : Proper hydration is necessary
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IMORTANCE OF VERNALISATION
Crop can be grown earlier
Plants can be grown in such regions where normally they do not grow
Yield of the plant is increased
Resistance to cold and frost is increased
Resistance to fungal disease
SENESCENCE
It is the process of ageing which is caused by increased entropy, cellular breakdown,
reduced homeostasis, increased metabolic failure and errors of replication as well as
transcription
1. Whole plant senescence
Monocarpic plants begin to undergo senescence along with fruit ripening. Eg.
Bamboos and sago palm
2. Shoot senescence
Certain perennial herbs produce annual aerial shoots for photosynthesis,
bearing of flowers and fruits. With maturity of fruits, the aerial shoot undergo
senescence underground parts perennate. Eg. Ginger, Narcissus, banana
3. Sequential / progressive senescence
Polycarpic evergreen perennial plants show progressive senescence of older
leaves, lateral organs and branches, flower, fruit bearing shoots as new growth
occurs near the tip eg. Mango, eucalyptus
4. Simultaneous/ Synchronous leaf senescence.
In perennial polycarpic deciduous plant all the leaves undergo senescence and
are shed simultaneously in a particular season, commonly autumn E.g. Elm,
maple, mulberry
GROWTH REGULATORS
(a) Characteristics
Plant growth regulators are small, simple molecules secreted in minute
quantities that influence various physiological functions in plants. They are of
diverse chemical composition, as given below
(i) Indole compounds : Indole-3-acetic acid (IAA)
(ii) Adenine derivatives : Kinetin, N6 – furfuryl amino purine
(iii) Derivatives of carotenoids : Abscisic acid ( ABA)
(iv) Gases : Ethylene
(b) Classification
Plant growth regulators are grouped into two categories based on the nature
of their action
(i) Plant growth promoters e.g. Auxins, cytokinins, Gibberellins
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They promotes growth activities like cell division, cell enlargement,
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flowering, fruiting and seed formation, tropic growth movement etc.
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(ii) Plant growth inhibitors e.g. Abscisic acid
They play an important role in plant response to wounds and stress of
biotic and a biotic origin
They are involved in growth inhibiting activities such as dormancy,
abscission etc.
AUXIN
(a) Discovery
Charles Darwin and his son Francis Darwin observe that the coleoptiles of
canary grass responded to unilateral illumination by growing towards the light
source ( phototropic curve)
After a series of experiments, it was concluded that tip of the coleoptiles is the
site of production of a substance, that caused the bending of coleoptiles.
F.W. Went isolated auxin from the tips of coleoptiles of oat seedlings
(b) Isolation
The word auxin is derived from Greek word ‘auxein’ meaning to grow
It was first isolated from human urine
Indole-3-acetic acid (IAA) and Indole butyric acid have been isolated from
plants
Naphthalene acetic acid (NAA) and 2,4 dichlorophenoxy acetic acid (2,4-D) are
synthetic auxins
(c) Physiological effects
Auxins control growth of plant cells.
They also control cell division in vascular cambium and xylem differentiation
They are responsible for apical dominance in plants
They prevent the formation of abscission layer and there by prevent premature
fall of leaves, flowers, fruit etc
(d) Application / Use
Auxin are used to initiate rooting in stem cuttings.
They are used to induce parthenocarpic fruits in tomatoes.
2, 4 D is used as a weedicide/ herbicide to kill dicotyledonous weeds
They promote flowering in pineapples and litchi
Auxins are used to prevent the premature fall of leaves, flowers, fruits etc
GIBBERELLINS
(a) Discovery
The ‘Bakan’ ( foolish seedlings) disease in rice seedling was caused by a fungus
Gibberellafujikuroi
EE. Kurosawa found that the symptoms of the disease could be developed in
uninfected seedling by treating them with sterile filtrate of the fungus
The active principle was later identified to the gibberellin acid.
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(b) Isolation
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There are more than hundred gibberellins reported from various higher plants
and fungi.
They are denoted as GA1, GA2, GA3 and so on
GA3 is the most intensively studied form of GAS
(c) Physiological effects
They cause elongation of the internodes
They promote bolting in rosette plants like cabbage, beet root etc
GA3 initiates synthesis of hydrolases to digest and mobilize the reserve food
materials of the seed to the developing embryo and thus it breaks seed
dormancy
(d) Applications/ Uses
Gibberellins are used to increase the length of grape stalks
They causes the fruits like apple to elongate and improve shape
They delay senescence and hence the fruits can be left on the trees for longer
period; this increases/ extend the market period
GA3 is used to speed up malting process in brewing industry
By increasing the length of internodes in sugarcane, GA3 increases the yield of
sugar cane by about 20 tonnes/acre.
Spraying juvenile conifers with GA3 hastens their maturity and leads to early
seed production.
CYTOKININS
(a) Discovery
Skoog and his co-workers observed that tobacco callus could proliferate only if
the medium contains in addition to auxin, the extracts from yeast, coconut milk
or DNA.
Skoog and miller later identified and crystallized the cytokinesis – promoting
substance and called it kinetin
(b) Isolation
Cytokinins were discovered as kinetic from the autoclaved herring sperm DNA.
Kinetin does not occur naturally in plants and search for natural substances
with cytokinin – like activities led to the isolation of zeatin from corn-kernels
and later a cytokinin from coconut milk
Later several naturally –occuringcytokinins and synthetic compounds with cell
division – promoting activity have been identified.
Natural cytokinins are synthesized in plants in regions where rapid cell division
occurs e.g. root apices, shoot buds, young fruits et
(c) Physiological effects
Cytokinins help in the growth of lateral buds into branches and help to
overcome apical dominance
They also promote adventitious shoot formation as they help to produce new
leaves and chloroplasts in leaves
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They promote nutrient mobilization and help to delay senescence
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(d) Applications
They are used to make the lateral buds grow into branches
They are used to delay leaf senescence
ABSCISIC ACID (ABA)
(a) Discovery
There independent researchers reported three kinds of growth inhibitors:
(i) Inhibitor – B
(ii) Abscisin II
(iii) Dormin
Later all of them were found to be chemically similar and named abscisic acid
(b) Isolation
It was discovered from its role in regulating abscission and dormancy
(c) Physiological effects
It plays an important role in seed development and inducing seed dormancy,
by this it helps the seed to withstand desiccation and other unfavourable
factors.
It stimulates the closure of stomata under conditions of intense solar radiation
and water stress
It increases tolerance of plants to various stress and hence called as stress
hormone
ABA stimulates the formation of abscission layer and abscission of leaves
flowers and fruits
(d) Application
Seeds are treated with ABA to remain dormant during storage
ETHYLENE
(a) Discovery
Cousins confirmed that ripened organs released a volatile substance that
fastened the ripening of the stored banana ; later this volatile substance was
identified as ethylene
(b) Isolation
Ethylene is synthesized in large quantities by the tissues of ripening fruits and
senescing organs
(c) Physiological effect
It promotes horizontal growth of seedlings and swelling on the axis.
It induces apical kook formation (called epinasty) in dicot seedlings
It promotes senescence and abscission of leaves and flowers
It promotes root growth and root hair formation, thereby increasing the
absorptive area.
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It promotes rapid elongation of internodes and petioles of deep water rice
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plants and helps the leaves to be above water level
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(d) Applications
Ethephon is the compound used to supply ethylene, it is an aqueous solution
that is readily absorbed and transported within the plant and it releases
ethylene slowly.
It hastens fruit ripening in tomatoes and apple
It promotes the production of female flowers on a monoecious plant
It accelerates abscission of flower and fruits in cotton, cherry, walnut etc
It is used to initiate flowering and for synchronizing fruit set in pineapples
It breaks seed and bud dormancy and initiates germination in peanut seeds and
sprouting of potato tubers
PLANT MOVEMENT
Plants have the capacity of changing their position in response to external (
environment) or internal stimuli, which are known as plant movement.
The specific region or site where the stimulus is received for changing the position
called perception site or region. The minimum period for which stimulus should be
given for inducing plant movement is called presentation time
Summary of movement flow chart on next page
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