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CHAPTER 8

PLANT HORMONES
Biochemistry and
Metabolism

Plants Hormones :
Means intercellular comunication within plants is mediated by
the action of chemical messengers

Signal molecules that individually or cooperatively direct the


development of individual cells,
or /@

Carry information between cells and thus coordinate growth


and development
Naturally occurring organic substances that at low
concentration will influence physiological process
Site of synthesis not clearly localized
More diffuse, cannot always localized to discrete organs

Types of plant hormones


5 recognized groups of plant hormones :
Auxins; Gibberellins; Cytokinins; Abscisic acid;
and Ethylene
More recently a 6 groups :
Brassinosteroids; Polyamines; Jasmonic acid;
and Salicylic acid

(i) AUXIN

First plant hormone to be discovered


Play a major role :
i. in the regulation of plant cell elongation
ii. in the growth responses of plants to
undirectional stimulus, it known as tropism

Natural auxins :
Indole-3-acetic acid (IAA)
4-chloroindole-3-acetic asid
Phenylacetic acid
Indole-3-butyric acid (IBA)

Synthetic auxins:
Naphthaleneacetic acid (NAA)
2-Methoxy-3,6-dichloro-benzoic acid (dicamba)
2,4-Dichlorophenoxyacetic acid (2,4-D)
2,4,5-Trichlorophenoxy-acetic acid (2,4,5-T)
Amount of IAA present depend on a number of
factors :

(i) Type of tissue


(ii) Age of tissues
(iii) Stage of growth
Concentrations exceeding the optimum
characteristically result in reduced growth

Auxin concentration is high enough growth


inhibited compared with control
Exogenous hormone do not show a
significant response
Endogenous hormone content at intact
tissues enough to support elongation
Effect of exogenous supplied auxin only in
tissues that have been removed from the
auxin supply (such as excised segments of
stem and coleoptile)

Auxins : Chemistry, Metabolic and


Transport

1870s ; Darwin and son Francis studies plant


growth phenomena of the growth stimulus in Avena
sativa (oat) coleoptile

Auxin regulated cell enlargement in excised tissues


such as coleoptile
If the tip of a coleoptile removed coleoptile
growth ceased. They found that the tip of the
coleoptile preceived light
If their covered the tips
With metal No growth toward light
With glass Growing towards light
# coleoptile growth towards light is controlled by
coleoptile tip

Wents discovery :
Growth promoting substance from excised coleoptile
tips would diffuse into agar block
Blocks could the used to restore growth in decapitated
coleoptile
If agar block containing auxin was placed on one side
of coleoptile stump coleoptile bent away from the
side containing the block
Curvature occurred because:
i. the increase in auxin on one side stimulated cell
elongation; and
ii. decrease in auxin on the other side caused a
descrease in the growth rate
Refer to fig 16.1 page 400 (Taiz /Zieger)- Plant
Physiology

Transport of auxins
Depend on the developmental stage of
theorgan/tissue of plant
Transport of hormone into or out of tissue/organ
influence level of active hormone within
tissue/organ
Hormone (Auxin) transport of plant is the polarity
of movement
Polar transport expressed as movement in one
direction
This indicate that polarity is not driven by external
forces (gravity etc.) but the property of cells
themselves

When movement is preferentially away from the


morphological apex toward the morphological
base of the transporting tissue, the direction of
movement described as basipetal
Movement in the opposite direction, that it toward
the morphological apex referred as acropetal
When stem or coleoptile is inverted, original
direction is maintained
# Refer to Fig 16.7 page 349 textbook

Fig. 16.7 Polarity in auxin transport in Avena


sativa coleoptile segment
Donor
block

14

C-IAA

14C-IAA

A
B

Receiver
block

B
A

A
B

B
A

14C-IAA

14C-IAA

Normal
orientation

Inverted
sections

Physiologial Effects of Auxin


Auxin has a variety of effect s on plant growth and
morphogenesis such as :
promote the elongation growth of stem and coleoptile
(however, it inhibit root elongation)
promote cell division in stems (but inhibit in lateral buds)
Development of fruit

Effect of auxin depend of factor, including


(i) developmental stage of tissue or organ
(ii) concentration of auxin
(iii) type of auxin (natural @ synthetic)
(iv) involvement of other plant hormones
(v) use of intact versus excised tissue for experiment

1. Cell elongation/enlargement
Stimulate cell elongation in excised tissue
(coleoptile)
Auxin concentration response curve show
increasing response with increasing
concentration of auxin an optimum
concentration is reached
Concentration exceeding the optimum
rduced growth
Concentration high growth inhibited

2. Increases the extensibility of the


cell wall
Increase in cell wall extensibility in coleoptile and young
developing stem
Auxin acts at the plasma membrane @ within the cell
Acid growth hypothesis proposed that auxin activates ATPproton pump located in plasma membrane.
Acid-growth hypothesis to explains auxin stimulated plant
cell elongation and enlargement
According to the hypothesis, auxin causes responsive cells
to extrude proton (H+)actively into the cell wall regions
decrease pH activates wall-loosening enzymes that promote
the breakage of cell wall bond increase wall extensibility

3. Growth responses to directional


stimuli
Auxin mediate effect of light and gravity on plant
growth
Plant oriented to the environment leaves
(houseplant) facing to window; roots growing
toward the earth
Tropic responses i.e. growth in response to
directional light phototropism or gravity
gravitropism
Toward light +ve phototropism
Away from light -ve phototropism

4. Inhibits the growth of lateral buds


@ Apical Dominance
Removal of shoot apex, stimulate growth one
@ more lateral bud
Auxin from apical bud inhibits growth of lateral
(axillary) bud

5. Promote the formation of


lateral roots

Elongation growth of root inhibited by auxin, but


initial of lateral (branch) root and adventitious
root is stimulated by high auxin level
The dividing cell by auxin form a root apex and
lateral root grow through root cortex and
epidermis
Adventitious root develop from a part of the
plant @ other than the normal form of root
branching

6. Delay the onset of leaf


abscission
Process of shedding organs such as leaves, flower and
fruit; known as abscission
Abscission occurs in region called abscission zone;
located near the base petiole.
Organ ages cell wall in abscission layer weaken and
separate
Leaf aging called leaf senescence. The cell wall in
abscission layer digested which cause them to become
soft and weak
The leaf break off at the abscission layer due to stress
Abscission dependent on concentration of auxin on either
side of the abscission layer
As leaf ages release of auxins decline, inducing the
ethylene synthesis abscission stimulated

Environmental stress eg.


Water stress, nutrient
deficiency reduce IAA
Ethylene produced at the
abscission layer
senescence

Cell wall expand and


suberized
Abscission layer

cellulase & pectinase are


produced
Middle lamela degraded
Cells separated, leaves/fruits
shedd

7. Regulates fruit development


Auxin produced in pollen, endosperm and
embryo of developing seeds
Involve in the initial stimulus for fruit growth from
pollination
After fertilization, fruit growth depend on auxin
produced in developing seed
In some species seedless fruits produced
naturally @ by treating the unpollinated flower
with auxin called parthenocarpy
Ethylene known to influence fruit development
The effect of auxin on fruiting mediated through
the promotion of ethylene synthesis

GIBBERELLINS
Gibberellins members of terpenoids
Gibberellins; assigned as GA
most important in higher plants
GA3 extracted from fungal culture
3 principal sites of gibberellin biosynthesis :
(i) Developing seeds (developing endosperm,
cotyledon, scutellum)
(ii) Developing fruits
(iii) Young leaves of developing apical bud and
elongating
shoots
(iv) Apical region of root

PHYSIOLOGICAL ACTION OF
GIBERELLINS
1.Stimulate hyperelongation of
intact stems
Occurs especially in dwarf and rosette plants
Promote elongation in intact plants rather than
excised tissues
Study on dwarf mutants of Oryza sativa, Zea
mays, Phaseolus vulgaris
These mutants exhibit the normal phenotype
when treated with GA3
GA treated enhanced internode elongation

The role of gibberellin in stem elongation come


from the study of rosette plants
Rosette plants compact growth habit, closely
spaced leaves
Failure of internode elongation may result from a
genetic mutation @ environmentally induced
Environmentally limited rosette plant (eg. Brassica
sp. and Spinacea oleraceae) generally do not
flower in the rosette form
Before flowering, plants undergo extensively
internode elongation; known as bolting
Bolting, due to environmental signal eg.
photoperiod or low temperature
Bolting in rosette plants can be induced by
exogenous application of GA

2. Seed germination
Cereal grain seeds like Hordeum vulgare
consists of embryo and non embryonic
regions
Embryo region synthesized GA and induced
release of -amylase to hydrolyze endosperm
(starch)
Non embryonic region if treated with GA will
stimulate to produce -amylase at high
concentration

3. Stimulate mobilization of nutrient


reserve during germination
Occurs of cereal grain
During imbibition water is absorbed by seed to
ensure GA secretion
GA moves from the embryo to the aleurone where
stimulated -amylase secretion ( -amylase
synthesized) and synthesis protease enzyme
Aleurone a layer of cell surrounding the
endosperm in seed
Protease converts an inactive -amylase to the
active form
-amylase and -amylase together digest starch
to glucose which mobilized to meet the metabolic
demands of the growing embryo

-amylase
(inactive)

aleurone
endosperm

-amylase
(active)
starch
-amylase

protease

glucose

scutellum
coleoptile

GA

plumule
radical

embryo

4. Flowering
Flowering is induced by gibberellins
Eg. Pharbitis nil, Chenopodium rubrum will
flower immediately
Many perennial plants capable of flowering
after pass through a juvenile phase
Gibberellins overcome juvenile phase in many
conifers and stimulate precocious flowering
Gibberellins promote maleness in unisexual
flowers while auxin promote femaleness

5. Inhibition of gibberellins
biosynthesis
Growth of stem can be inhibited or reduced by synthetic
chemical that block gibberellin biosynthesis i.e growth
retardant or anti gibberellins (AMO-1618, cycocel,
Phosphan-D, ancymidol)
These compound has commercial application in production
of ornamental plants
Effects:
To reduce stem elongation
Results in shorter and compact plants with darker
green foliage
Spraying the plant (Wheat) with antigibberellin produce a
shorter, stiffer stem and preventing lodging

CYTOKININS
Cytokinin- adenine derivatives
Kinetin- first compound found with cytokinin
activity
Synthetic cytokinin prepared by heating DNA
Zeatin- first natural cytokinin discovered and
most widespread
Isopentenyl adenine (iP)
Dihydrozeatin (diHZ) : leass active than
zeatin
Benzyl adenine (BAP)

1.Cytokinins regulate cell division


Cytokinin is major factor in regulating cell
division in the presence of auxin
Have capacity to initiate division in plant cells
and in quiescent or non dividing cells (tissue
culture)
Initiate cell division by controlling cell cycle
at two points
1. Catalyze transition from G2 phase to
mitosis
2. Control G1 to S phase transition

2. Stimulate cell proliferation


In the case of neoplastic (tumorous growth)
Bacterium Agrobacterium tumefaciens pathogens
that causes tumorous growth on stems known:
crown gall
Crown gall tissue can be excised and maintained
on simple medium without hormone
Have capacity to synthesis auxin and cytokinin
When bacteria invade host tissue, it transfer these
genes
Genes replicated in host cell
Produced elevated level of auxin and cytokinin
Stimulate neoplastic growth

Crown gall

2. Organogenesis
Cytokinin and auxin stimulate organogenesis:
organ formation
Development of shoot and root
Cell culture growth required cytokinin and
auxin
High Cytokinin/auxin ration stimulate root
formation
Low cytokinin/auxin ration stimulate shoot
formation

3. Senescence
Mature leaves and fruits express senescence
Senescence: breakdown of protein, nucleic acids,
other macromolecules, loss of chlorophyll,
accumulation of amino acids
Cytokinins will delay senescence while ethylene
promote senescence
Cytokinins direct nutrient mobilization and retention
by stimulating metabolism in the area of application
Creates a new sink: area that attract metabolites
from region of application

senescence

Experiment : role of cytokinins in


nutrient mobilization
Control

Treatment with cytokinins


kinetin

kinetin

radioactive

C
A
Radioactive spreads
into vascular tissue
for export through
petiole

B
Radioactive
accumulates in the
area treated with
kinetin

Radioactive
retain near poin
of application

-Radioactive labeled nutrient are fed to the


plant
-In control (A) radioactive spreads into vascular
tissue for export through petiole
-In treatment (B) one part/half of leaf is treated
with cytokinin
-Radioactive accumulates in the area treated
with kinetin
-In other treatment (C) cytokinin applied on
part of leaf (right)
-Radioactive retain near point of application
-Cytokinins direct nutrient mobilization and
retention by stimulating metabolism in the area
of application
-creates a new sink area that attract
metabolites from region of application

Other effects of cytokinins

Stimulate cell enlargement


Regulate vascular differentiaiton
Promote axillary bud and release apical
dominance

Abscisic Acid

Abscisic acid is a single compound


Occur in mature, green leaves
Synthesized in cytoplasm of leaf mesophyll cell
and accumulated in chloroplast
Able to move quickly out of leaves sink tissues
Actions: induce storage protein synthesis during
seed development
Regulating stomatal closure during water stress
Also involved in regulating abscission and bud
dormancy

ABA physiological effects


1. Stomatal closure
In drought, leaves will synthesized high level of
ABA
Allowed stomata closure
Water will be stored during drought/water stress

2. Bud/seed dormancy
Woody plants in temperate zone
ABA concentration maximum in early winter and
low end of winter
ABA prevents bud development and seed
germination

ABA actions antagonistic with other hormone:


Inhibits amilase which produced by seed treated
with giberellins
Promote chlorosis which have been inhibited by
cytokinins
Inhibits cell wall elasticity and cell enlargement
by IAA

ETHYLENE
Simple hydrocarbon gaseous : H2C=CH2
Not required for normal vegetative growth
Synthesized primarily in response to stress
Produced in large amounts by tissues
undergoing senescence or ripening
Occurs in all plant organs

Physiological effects of ethylene


Vegetative development
Stimulate elongation of stems, petioles, roots
and floral structure of aquatic and semiaquatic
plants
Ethylene promotes gibberellin synthesis in rice
to promote root and shoot elongation
While in peas, root and shoot elongation
inhibited by ethylene
Stimulate abnormal growth response such as
swelling of stem tissues and downward
curvature of leaves (epinasty)

Fruit Ripening
Stimulate fruit ripening: banana, apple avocado
etc..
Ethylene is autocatalytic
release of ethylene gas by ripening fruits
will in turn stimulate premature climacteric
and ethylene production by other fruits stored
near
Number of qualitative metabolic changes are
initiated in fruit

Changes in fruit ripening:


During ripening promote production of sugars,
which increase sweetness (breakdown of starch
and acid) and odor
Induced rupture of cell membranes and water loss
from tissues: increase cell wall softening by the
action of enzymes
Involved breakdown of chlorophyll and synthesis
of pigments
Synthesis of flavor

Ethylene has important


commercial uses

Storage facilities developed to inhibit ethylene


production and promote preservation of fruits have a
controlled atmosphere of low O2 concentration and
low temperature that inhibits biosynthesis
High concentration of CO2 (3-5%) prevents ethylene
action

Low pressure to remove ethylene and oxygen from


storage chamber
Use of inhibitors of ethylene action
Such as CO2 and Ag+ (silver)
Will delay or prevent ripening
Ethylene has high diffusion rate
Difficult to apply in a gas form
Spray ethylene releasing compound such as
ethepon
When taken up by plant tissue, ethepon is
converted to ethylene

Brassinosteroids
Steroid hormones
Chemical structure similar to animal steroid
hormones
Brassinolides most biologically actve
Functions: brassinosteroids promote stem
elongation in mutant plants, shoot elongation
and ethylene production
Inhibits root growth and development