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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
 Went’s 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 14
C-IAA 14C-IAA
block
A B

B A A
Receiver
block B

A B

B
A
14C-IAA 14C-IAA
Normal Inverted
orientation 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 ATP-
proton 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 α-amylase to hydrolyze endosperm
(starch)

 Non embryonic region if treated with GA will


stimulate to produce α-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 α -amylase secretion (α -amylase
synthesized) and synthesis protease enzyme
 Aleurone ⇨ a layer of cell surrounding the
endosperm in seed
 Protease converts an inactive β -amylase to the
active form
 α -amylase and β -amylase together digest starch
to glucose ⇨ which mobilized to meet the metabolic
demands of the growing embryo
β -amylase aleurone
(inactive)
endosperm

β -amylase
protease
(active)
starch
α -amylase

glucose scutellum

coleoptile

GA plumule embryo

radical
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 B
Radioactive
Radioactive spreads Radioactive retain near poin
into vascular tissue accumulates in the of application
for export through area treated with
petiole kinetin
-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

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