Regulating Growth and

Development: The Plant

Hormones
CHAPTER OUTLINE
Auxins
Cytokinins
Ethylene
Abscisic Acid
Gibberellins
The Molecular Basis of Hormone Action
The normal development of plant depends on a number of internal and external factors:
1. The internal factors are chemicals.
2. The external factors such as light, temperature, day length and gravity.
The communication among cells, tissues, and organs is important for the growth and
development of an organism.
The regulation and coordination of metabolism, growth, morphogenesis depends on
chemical signals called hormones (to stimulate).
The plant hormones or phytohormones concept had three basic elements:
1. Synthesis of the hormone in one part of the body
2. Transport of the hormone to another part of the body
3. It induces a chemical response to control a specific physiological events
Some of the plant hormone produced in one tissue and acts in another tissue, while
other plant hormone produced and acts in the same tissue.
Plant hormones are synthesized at several locations in the plant body.
Plant hormones are active in very small quantities.
eg: pineapple plant
6 microgram of IAA for 1 kilo of plant material
Plant hormones may stimulates or inhibit the growth and development of plant.
The response of a plant to particular hormones depends on:
1. Its chemical structure.
2. How it is read by the target tissue: i.e
•The function of the same hormone is different in two different tissues.
•The function of the same hormone is different of the same tissue at two different
time of development.
Some plant hormones are able to influence the synthesis and the function of other
plant hormone.
Tissues may require different amount of hormones, such differences are referred to
as differences in sensitivity.
Plant hormones found in the plant conjugated to other substances such as sugar,
these hormones are not active and to be active hydrolysis must occur.
Traditionally, five classes of plant hormones:
1. Auxins
2. Cytokinins
3. Ethylene
4. Abscisic acid
5. Gibberellins
More recently, it has become clear that additional chemical signals are used by plant
such as:
1. Brassinosteroids: required for normal growth of most plant tissues.
2. Salicylic acid (similar to aspirin): defense responses to plant pathogens.
3. Jasmonates: plant growth regulation and defense.
4. Polyamines: essential for growth and development and affect the processes of
mitosis and meiosis.
5. Systemin: long-distance signal to activate chemical defense against herbivores.
6. Nitric oxide: serve as signal in hormonal and defense responses.
 Auxins:
The effect of auxins was discovered by Charles Darwin and his son in 1880.

Darwin’s experiment: in the figure

Darwin’s conclusion:
When seedling are freely exposed to a lateral
light some influence is transmitted from the
upper to the lower part causing the lower part to
bend (elongation).
In 1926, went succeeded in isolating the auxin (to increase).
Indoleacetic acid (IAA) is the principle naturally occurring auxin which is resembles the
amino acid tryptophan.
Plant can produce IAA directly from tryptophan.
Mutants in maize and Arabidopsis that are unable to synthesize tryptophan can still make
IAA.
Auxins produced in:
1. Shoot apical meristem
2. Young leaves
3. Developing fruits and seeds
4. Root tips, most evidence indicate that it is not produced there but is transported there
via the vascular cylinder.
Auxin is the only plant hormone known to be transported polary
Auxin moves only from shoot tip to base not to in the reverse direction, this unidirectional
transport of auxin is called polar transport (from the shoot tip and leaves downward in the
stem and from the root tip toward the base of the root (the root-shoot junction).
The velocity of polar auxin transport is 5-50cm/h
The auxin in plant is transported:
1. Polarly
2. Nonpolarly
Auxin is transported in parenchyma cells of the vascular tissues polar.
In addition to the polar transport of auxin, it has recently been recognized that most of the
IAA synthesized in mature leaves apparently is transported throughout the plant non-polarly
via the phloem at rates considerably higher than those of polar transport.
Most of the auxin reaching the root tip is transported nonpolarly in the sieve tubes of
phloem.
In the root tip, the IAA is redirected polarly in the parenchyma cells.
Polar transport of IAA is mediated by
auxin efflux carries localized at the
basal ends of the conducting cells.
IAA enters the parenchyma cell by
passive diffusion in its protonated form
(IAAH) or/and by active cotransport in
its anionic form (IAA-) via an influx
carriers (Aux1). The anionic form
predominates in the cytosol, which has
a neutral pH. The anions exit the cells
only at the basal end, via efflux carriers
(PIN proteins).
Auxin plays a role in the differentiation of
vascular tissue
Example:
When a stem of plant is wounded in such a
way as to sever and remove portions of the
vascular bundles, new vascular tissues will
from cells in the pith and will connect the
severed bundles.
But if the leaves and buds above the wound
are removed, the formation of new cells
delayed.
When IAA is added just above the wound,
new vascular tissues begins to form.
Auxin plays important role in the joining of
vascular traces from developing leaves to the
bundles in the stem.
Auxin plays a role in the induction and arrangement of leaves
The inhibition of polar auxin transport blocks leaf formation at the vegetative apical
meristem of tomato, adding small amounts of IAA to the shoot tip restores leaf
formation.
Externally applied IAA also was shown to induce flower formation on inflorescence
apices.
Auxin provides chemical signals that communicate information over long distances
The flow of auxin from the growing apical bud inhibits the growth of axillary (lateral)
bus.
Apical dominance: the inhibitory influence of an apical bud upon the lateral buds.
The role of auxin in apical dominance can be demonstrated experimentally. For
instance, when the shoot tip of a bean plant (Phaseolus vulgaris) is removed, the
lateral buds begin to grow. However, when auxin is applied to the cut surface, the
growth of the lateral buds is inhibited.
In wood plants, auxin promotes activity of the vascular cambium.
With expansion of buds and resumption of their growth in the spring, auxin moves
downward in the stems and stimulates the cambial cells to divide, forming secondary
vascular tissues.
Auxin promotes the formation of lateral and adventitious roots
Lateral roots arise from pericycle
Auxin stimulates the pericycle cells to divide
The first practical application of auxin involved its stimulation of the initiation of adventitious
roots in cutting (treating cuttings with auxin).
High concentration of auxin to already-growing roots usually inhibits their growth.
Auxin promotes fruit developing
Without fertilization, the fruit will not developed
Adding auxin to no fertilized or pollinated flower will promote parthenocarpic fruit.
Example: seedless tomato.
Developing seeds are a source of auxin.
Auxin, produced by developing embryos, promotes maturation of the ovary wall and the
development of fleshy fruits.
Synthetic auxins are used to kill weeds
2,4-dichlorophenoxyacetic acid (2,4-D) is use to control weeds on agriculture lands (in
economic terms, this is the major practical use for plant growth regulators).
2,4-D is not broken down in plants as natural auxin, and the resulting of high level of 2,4-
D contribute to lethal effects.
2,4-D is more effective on broad-leaf weeds (these weeds absorb more 2,4-D than
grasses).
2,4-D be inactive by conjugation.
Cytokinins:
1941, van overbeek found that coconut milk contains a potent growth factor that greatly
accelerated the development of plant embryos and promoted the growth of isolated
tissue and cells in the test tube.
1950, skoog experiment: showed that a stem segment of tobacco plant grow initially in a
culture media, but its growth soon slowed or stopped, because the stimulus factor
originally present in the tobacco stem exhausted.
Skoog add IAA: no effect
Skoog add coconut milk: the cell began to divide, and growth of the tobacco stem
resumed.
Skoog try to isolate the growth factor from coconut milk but he failed.
Skoog changing course, he tested a variety of purine-containing substrate (nucleic acid) in
the hope of finding a new source of the growth factor.
The work of skoog in purine substrates led to the discovery by miler, that a breakdown
product of DNA contained material that was highly active in promoting cell division.
Subsequently, miller and skoog, succeeded in isolating the growth factor from DNA
preparation and identifying its chemical structure.
The first growth factor isolated called kinetin and the group of regulators called cytokinins
(because of their involvement in cytokenisis).
Kinetin resembles the purine adenine
Kinetin has a relatively simple structure and biochemists were able to synthesize a number of
related compounds.
Zeatin: is a growth factor isolated from kernels of maize, it occurs naturally in plant.
Zeatin: the most active of the naturally occurring cytokenins.
Cytokenins fund in:
1. Dividing cells
2. Dividing seeds
3. Fruits
4. Leaves
5. Root tips
6. Bleeding sap (the sap that drips out of wounding, injuring).
The importance of cytokenin:
1. Important in plant development research
2. Central to tissue culture methods
3. Extremely important to biotechnology
4. Modify apical dominance (treatment of lateral buds with cytokenin causes buds to grow
even in the presence of auxin).
The cytokenin/Auxin ratio regulates the production of root and shoots in tissue cultures.
In studies of tobacco stem tissues:
1. IAA: tissue culture produces rapid cell expansion.
2. Kinetin: little or no effect.
3. Certain concentration of both IAA and cytokenin (10:1) will increase cell division and
cells remains meristematic (callus).
4. High concentration of auxin and low of cytokenin will give raise to roots.
5. High concentration of cytokenin and low of auxin will give raise to bud and shoots.
Cytokinin delay leaf senescence:
Loss chlorophyll in removed leaves can be delayed by cytokenins.
Amino acids migrate from the yellow parts of the leaf to the cytokenin treated parts.
The detached leaves die after short time, because the cytokenin in these leaves is limited:
the conclusion led to an important unanswered question which is, where cytokenin
produced?
Cytokenin is abundant in dividing tissues but it is synthesized in root tip and transported
through xylem to all other parts of the plant.
Ethylene:
Ethylene discovery: ethylene was known to have effects on plants long before the
discovery of auxin (goes back to 1800).
City streets in Europe were lighted with lamps that burned illuminating gases, this gas
increase the growth of the plants closed to it in the streets.
1901, neljubov demonstrated that ethylene was the active component of illuminating gas
by his experiments in peas.
Ethylene was active at concentrations as low as 0.06 ppm in air.
Ethylene influence many aspects of growth and development in plants including:
1. Growth of most tissue
2. Fruit maturation
3. Fruit and leaf abscission
4. Fruit and leaf senescence
Ethylene is biosynthesis as follows:
The formation of ACC is affected by:
1. High auxin concentration
2. Air pollution damage
3. wounding
These three, stimulate the formation of ethylene.
Ethylene may inhibit or promote cell expansion:
1) inhibition:
Triple response of pea seedling to ethylene:
The effects of increasing ethylene concentration on the growth of dark-grown pea seeding
as shown in triple response include:
1-a decrease in epicotyls elongation
2-thickening of the shoot
3-change in orientation of growth from vertical to horizontal
2) Promotion:
for some semi aquatic species, ethylene increases the stem growth
in deepwater or floating varieties of rice, submergence of young rice plants during
the monsoon season triggers on increase in ethylene biosynthesis this ethylene-
induced intermodal elongation rising flood water
Ethylene plays a role in fruit ripening:
ripening in fruit involves a number of changes:
a) Chlorophyll degraded and other pigments may form, changing the fruit color
b) The fleshy part of the fruit softens as a result of the enzymatic digestion of pectin
(the principle component of the middle lamella of the cell wall)
c) Starches, organic acid, and oil will be converted to sugars
d) Large increase in cellular respiration evidenced by an increased uptake of O2
due to these changes, the ethylene synthesis precedes
the effect of ethylene on fruit ripening has agricultural importance:
1) Promoting the ripening of tomatoes that are picked green and stored in the
absence of ethylene until just before marketing
2) Ethylene is also used to accelerate the ripening of grapes
Whereas ethylene promotes abscission, Auxin prevents abscission
Abscission: the dropping off of leaves, flowers, fruits or other plant parts, usually
following the formation of an abscission zone
Ethylene promotes the formation of the enzymes that cause the cell wall dissolution.
Ethylene is used commercially:
1) To promote fruit loosening in cherries….., which makes the mechanical harvesting
possible.
2) As a fruit-thinning agent in commercial orchards of prunes and peaches
-Abscission is controlled by an interaction of ethylene and auxin
Auxin effect has been used commercially:
e.g: auxin treatment prevent pre harvest drop of citrus fruits
high auxin concentration stimulate abscission (this concentration promote the
ethylene synthesis)
Ethylene apparently plays a role in sex expression in cucurbits
Ethylene plays a major role determining the sex of flowers in some monoecious
treatment with ethylene changes the expression of sex to femaleness
Abscisic Acid (ABA):
This substance has no direct role with abscission
1949, wareing discovered that the dormant buds of
potatoes contain large amount of growth inhibitor
(dormin)
1961, addicott reported the discovery in leaves and
fruits of substance capable of accelerating abscission
(abscisin)
dormin and abscisin compounds now is called abscisic
acid (ABA).
abscisin acid prevents seed germination:
during seed development, ABA level will increase.
high level of ABA stimulates the production of seed
storage proteins and is also responsible for preventing
premature germination.
the breaking of dormancy in many seed is correlated
with declining of ABA level in the seed.
Abscisic acid plays a role as root-to-shoot signal:
When the plants are exposed to abiotic stress that generate water stress or water
deficiency.
The roots respond by increasing the biosynthesis of ABA and releasing it in the xylem
ABA is transported to the leaves
In leaves, the stomata respond to the increased ABA concentration by closing (reducing
the loss of water by transpiration).
Gibberellins:
Gibberellins research was an exclusive product of Japanese scientists.
1926, kurosawa of Japan was studding a disease of rice called foolish seedling disease,
the cause of this disease was a substance produced by a fungus (giberella fujikuroi) which
was a parasitic on the seedlings.
Gibberellins was named and isolated in 1934.
1956, gibberellins was isolated from plants.
Gibberellins occur in all plants.
Gibberellins presents in various amounts in all plant parts of the plant, but the highest
concentration are found in immature seeds.
125 gibberellins now have been isolated and identified chemically.
Most plants contain 10 or more gibberellins.
Gibberellins vary in structure and function.
GA3 is the best studied gibberellin which is extracted from fungi.
The main function of gibberellins is stem and leaf elongation in intact plant by stimulating
both cell division and cell elongation.
Application of gibberellins can cause dwarf mutant
to grow tall.
When GAs hormones is applied to dwarf mutant
plants, these plants become indistinguishable from
normal tall, non mutant plants, indicating that these
mutant are unable to synthesize GA and that tissue
growth requires GA.
The increase in yield of rice and wheat during 1960-
1979 was due to dwarfing genes for both crops.
The advantages of dwarfing genes:
1. Development of shorter plants.
2. More grain production.
3. Little straw production.
4. More resistant to damage by wind and rain.
Gibberellins play multiple roles in breaking seed dormancy and in germination
The seeds of many plants require a period before they will germinate.
Cold or light is needed for the breakdown of seed dormancy.
The treatment of the seeds with GA will promote the growth of the embryo and the
emergence of the seedling.
Gibberellins enhance cell elongation, making it possible for the root to penetrate seed
coat.
Practical application of gibberellins:
Accelerate the seed germination and ensures germination uniformity for the production
of the barley malt used in brewing.
Barley seed has specialized layer of endosperm cells called aleurone (lies just inside
seed coat).
Aleurone: rich with proteins
When seeds begin to germinate, the embryo releases GA, which diffuses to aleurone
and stimulates them to synthesize α-amylase and other enzymes.
The enzymes produced by aleurone digest the stored food in the endosperm.
The digested food will be absorbed by the growing regions of the embryo.
Gibberellins can cause bolting and can effect fruit development.
Bolting: is the growth of an elongated stalk with flowers grows from within the main stem
of plant
Some plant like cabbages and carrots form rosettes before flowering
Rosettes: leaves developed, but the internodes between them do not elongate.
In cabbages and carrots, flowering can be induced by exposure to long day, to cold or
both.
In cabbages and carrots, bolting can be happened by adding gibberellins without
exposure these plants to long day or cold conditions.
Gibberellins can be used for the development of parthenocarpic fruits.
The major commercial application of gibberellins is in the production of table grapes
Example:
Adding gibberellic acids to Thompson seedless
grapes causes larger fruit and much looser clusters,
making them more attractive to the consumers.
The molecular basis of hormone action:
The molecular mechanism by which these chemical (plant hormones) influence the growth,
development and rapid responses at the cellular level
Development: cell division/cell elongation
Differentiation: the result of selective expression of a particular set of genes
Communication: plant hormone coordinates growth and development
The molecular basis of plant hormones is supported in part by numerous observable
influences of plant hormones on the rate of cell division and on the rate and direction of
cell expansion.
the traditional plant hormones, as well as the newly discovered ones, can act either to
stimulate or to repress specific genes within the nucleus. In fact, it appears that many
observable hormone responses are the result of such differential gene expression.
Hormones control the expression of specific genes:
Every cell in the adult plant contains the same DNA (genes) which is present in the zygote.
Every cell selects certain genes and synthesizes mRNA-protein.
The protein could be: enzyme, structural component, ….
According to the expression of different genes in two different cellscortical cell in the root
differ from the mesophyll cell in the leaves in structure and function of the same plant.
The principle of the functioning of genes:

Regulatory transcription factor  switching on or switching off the gene

When the gene is on, transcription and translation will take place and proteins (enzymes) will
be produced.
The role of plant hormone in switching on or switching off the gene:
e.g
The role of GA and ABA in the synthesis of α-amylase which is present in aleurone layer in
barley seeds
1) If aleurone tissue is incubated with radioactive amino acid and various hormonesthe α-
amylase will be formed.
2) Treatment of aleurone tissue with GA abundance amount of α-amylase
3) Treatment of aleurone tissue with ABAlittle amount of α-amylase
Result:
GA-activate α-amylase synthesize
ABAinhibit α-amylase synthesize