Chapter 6 The Plant Hormones

(Phytohormones) p309-366
Plant hormone
Plant growth substance
plant regulator
Plant hormones are naturally occurring
organic substance that can be transported from
the synthetic tissue to a specific target tissue
where, at low concentration,exert a profound
influence on physiological process.
1μgIAA/10000 tips.
1mgIAA/1T leaf.
10mgBR/225kg of pollen.
5 putative plant hormones：IAA、GA、CTK、
ABA and Eth。（6、BR）.
• Plant growth regulators (调节剂) are
normally used to denote synthetic
compounds that exhibit plant hormonal
activity.
Section 1 Auxins
1.1 Discover of auxin
• Charles Darwin and Francis Darwin(1880, Phalaris
canariensis )

• Boysen-
• Jensson(1910)

• F.W. Went(1928)----Avena curvature test. α

 Went called it auxin.
Structure of IAA is Indole-3-acetic acid---
IAA.

-CH2COOH

1.2 Distribution and transportation of IAA in

plant
All parts have auxins, but the highest concentrations locates in
meristematic regions and actively growing organ, such as
coleoptile apices, root tips and the apical buds of
growing stems.

0.3
IAA relative contents

0.2

0.1

0
0 20 40 60
Distance form coleoptile apix to root
tip(cm)

Fig 6-1 IAA concentrations in plant parts

 IAA Polar transport：IAA is only one plant
hormone transported unidirectionally from
apical end to basal end.

IAA transport in root from basal to tip.

Fig 6-2 IAA Polar transport controlled by IAA transport
protein (from Taiz and Zeiger 2006)
Other transport: IAA synthesized in mature
leaf is transported in double-directionally along
the phloem.
• 1.3 Biosynthesis of IAA
• 1.3.1 Biosynthesis location：almost parts of
plant. Especially in coleoptile, young leaf and
developing seed and fertilizing ovary
• 1.3.2 Steps of Biosynthesis
• 1）Tryptophan pathway:
• The deficiency of Zn decreases IAA synthesis
due to decreasing in Trp synthesis.
1） Pathway dependent on tryptophan
COOH
NH2
*Trp decarboxylase
*Trp N
monoxygenase Tryptophan(Trp) CO2
O2 *Trp
transaminase
NOH -NH2 NH2
COOH
N N Trptamine
O (TAM)
Indole-3-acetaldoxime
N
O Indole-3-pyruvic acid(IPA) O2 Amine oxidase
NH2 *IPA -NH2
CO2
decarboxylase
N
N IAld reductase
Indole-3- O OH
acetamide
N N Tol oxidase
(IAM) Indole-3-acetonitrile Indole-3-acetaldehyde(IAld)
N
(IAN) *nitrilase +H2O IAld Indole-3-ethanol
-2H dehydrogenase (tryptophol,TOL)
*IAM
hydrolase COOH

N
(IAA) Indole acetic acid
 Fig 6-3 IAA-over producing
plant grows worse

normal IAA-over
producing
plant
 Fig 6-4 Effects of indole-3-acetonitrile
and IAA on wild-type (wt) and nit1-3
mutant seedlings of Arabidopsis.
A Eight-day-old seedlings grown in the presence
and absence of 30 μM indole-3-actonitrile(B) or
wt Nit1-3 1μM IAA(C). (Control is shown[A].) Note that
wild-type plants show a typical auxin-like response
to both IAA and indole-3-acetonitrile.The nit1-3
motant responeds to IAA but does not exhibit an
auxin-like response to indole-3-acetonitrile. It
B lacks nitrilase and cannot convert indole-3-
acetonitrile to IAA
wt Nit1-3
Nit1(nitrilase mutant) is
insensitive to indole-3-acetonitrile
C
wt Nit1-3
2） Pathway independent on tryptophan

Mutants for tryptophan nutrition deficient

Maize -orp mutant, trp synthase β-subnuit point
mutant, 50 time high IAA-conjugate

An orange pericarp (orp) maize cob showing the expected two-

gene recessive trait, the orange kernels carry both mutant genes
• Arabidopsis-trp1,trp2 and trp3 can’t
synthesize Trp but content 19-30 times as
higher as IAA-conjugates of wild type
• N15-anthranilate(邻氨基苯甲酸）

N15-IAA 39%
N15-Trp 13%
 anthranilate synthase(Trp1,MTR)
anthranliate(邻甲基苯氨酸） Chorismate →phenylalanine ，tyrosine
(分支酸) (苯丙氨酸， 酪氨酸)
Anthranilate
PR-transferase
5-phosphoribosylanthranilate
IBA-conjugates
PR-anthranilate
isomerase
1-(o-carboxyphenylamino)
-1-deoxyribulose-5-P COOH

IGP synthase N
OH
CH2OP Indole-3-butyric acid(IBA)

0H N Nitr IBA sythase

ila
nit1 se
N IGP
Trp synthaseαIndole-3-glyceral N
Trp3 IAN COOH
phosphate
Serine + COOH IAA
indole N
N O
Trp synthaseβ
Trp2,orp a se rase N
sfe
COOH ynt h
s
rp otr a n IPA IAA-conjugates
NH2 T n
ami ?
N Trp rty
• 1.4 Inactive and degradation of IAA
• 1.4.1 IAA-conjugate:

IAA-Sugar

IAA-Amide
• Function of IAA-conjugates
• (1)Storage form.
• (2)Easy transport form.
• (3)Anti oxidation form.
• (4)Level control and detoxification.
1.4.2 IAA oxidation
• Two enzymes: IAA oxidase and peroxidase
• 1.5 Physiological effects and application
of IAA
• (1)、Enhancing elongation of cell and
organ.

Control IAA
 10

- Relative growth rate +

8 Stem
6 Bud
4
2
Root
0
-2

-4
-6

-8
-10

-12
0 101-11 2 103-9 4 105- 7 6 10-75 8 109- 3 10 10
11- 1 12
IAA concentration（mol/L)
Fig 6-5 Effect of IAA concentration on elongation of root,
bud and stem. The growth-optimum concentration is for
stem higher than for bud, then for root. The root growth is
most sensitive to IAA.
 Mechanism for elongation by IAA ：
1）IAA activates genes——mRNA and
protein synthesis——Slow reaction.

IAA
mRNA

protein
Elongation

+IAA +IAA
and mRNA

and Protein
Elongation

Elongation

+IAA+actinomycinD +IAA+cycloheximide

CK CK
 B．Acid growth theory (酸生长理论) .——
rapid reaction：IAA activates ATPase→pH ↓, cell wall
acidification →enzyme activation →cell wall dehydration and
loosening →water absorption。

IAA
Inactive E Microfiber of cellulose
Pentosan
ATP
H+Active E H+
ADP+Pi
other cell wall polysaccharide
Hydrogen bound

Fig6-12 IAA as an effector activates ATPase (H+-pump) in

plasmic membrane
• Plant hormone receptors are a group of
proteins which first bind plant hormones
and make the hormone play a serious
role in physiological functions.
 (2) Enhancing cell division and organogenesis
——Cutting-shoot rooting (插枝生根)
• Cambium cell division and root primordial cell
division in the early spring.

CK IAA

The rooting with 10-100 mgL-1 or 0.5-1% of powder.

(3) Enhancing fruit setting and parthenocarpy
(单性结实)——production of seedless fruit

• Watermelon, popper, tomato, egg plant etc are

sprayed with 2，4-D (dichlorophenoxyacetic acid)
of 10 mgL-1 to produce seedless fruit by
parthenocarpy in early spring time or greenhouse
cultivation.
(4) Maintaining apical dominance

Lateral bud develops

after removing apical
bud.。

Lateral bud poorly

develops after
removing apical bud,
then putting IAA in the Lateral bud
part
(5) Enhancing abscission——flower and fruit
thinning (疏花疏果)
Apple with NAA (naphthalene acetic acid)5～20 mgL-1
or NAD (naphthalene acetamide)25～50 mgL-1.

(6) Inhibiting sprouting of potato .

1% of NAA powder
• (7) Enhancing flowering and controlling
sexual differentiation.
• 14 months old pineapple with 2.4─D 50-100
mgL-1 or NAA 15─20 mgL-1 ( 30ml per plant)
• Cucumber ♀↑.
• (8) As a herbicide to kill dicot with 2·4-D
(1000 mgL-1 ）.
• Conclusion of IAA functions:
• (1)Enhancing elongation of cell and organ
• (2)Enhancing cell division and organogenesis
• (3)Enhancing fruit setting and parthenocarpy
• (4) Maintaining apical dominance
• (5) Enhancing abscission
• (6) Inhibiting sprouting
• (7) flowering and sexual differentiation.
• (8) As a herbicide
 Section 2 Gibberellic acid (GA)
• 2.1 Discover of GA
• Rice “foolish seedling” infected by Gibberellia
fujikuroi. A large family of more than 125
members.
11 12 11 12
20 CH3 H H 1 O OH
1 13 H
9 CH3 2 9 13
2 10 10 CH2
8 14 16 17 16
19CO 8 14 17
3 5 15 5
4 6 3 4 6 15
7 7
18 19 H H CH3 HO H
CH3 H COOH
CH3 18CH3

gibberellane GA3
GA is diterpenes constituting with 4 isopentenal groups
C19 --GA is higher activity than 20C– GA. Those with both
3-OH and 1,2 unsaturation exhibit the highest activity.

2.2 Synthesis and transport of GA

GAs exist universally in plant kingdom and fast

growing parts are higher ontent

GAs are synthesized in shoot,root, flower and fruit.

Fig 6-13 Histochemical localization of GA1
promoter activity indicating CPS expression during
the development of transgenic Arabidopsis
containing the GA1 promoter-GUS gene fusion
pGA1-103.
 Mevalonic acid Isopentenyl pyrophosphate (IPP)
CH2—COOH OH OH
O
3CH2C-OCoA HO—C—CH2 H3C-C=CH-CH2- O- P -O-P-OH
O O
CH2—CH2OH CH2
In microbe 4 molecules
In plant

OPP
-AMO-1618， OPP
Geranylgeranylpyrophosphate
-Phosphor-D GGP

Kaurene
贝壳杉烯 Copalylpyrophosphate CPP
-CCC
various
GA
CHO
Kaurenoic acid COOH COOH GA —7aldehyde
12

Biosynthesis pathway of GA and several retardants (inhibitors for

GA biosynthesis)
Fig 6-14 Inactive form:
2β-hydroxylation and
GA-conjugate: GA-
glucoside.
2.3 Physiological role and application of GA

• (1) Promoting elongation of stem and

division of cell.
• Function in subapices and promote
elongation of whole plant, especially of
mutants and physiological draft.
 Fig 6-15 Effect of
GA3 on stem
elongation of
Progress No.9 dwarf
pea seedlings:(left)
control plants, (right)
plants seven days
physiological draft after treatment with 5
μ g GA3.
 CK 100pg 1ng

Fig 6-16 Promotion of leaf sheath elongation of

Tanginbozu dwarf rice three days after treatment with
GA3.
 Application in stem or leaf-harvested
plants such as celery, lettuce, tea etc with 1-5
mgL-1 .
20-40 mgL-1 of GA3 can accelerates heading
of late season rice or hybrid rice.
Mechanisms:
(1) Cell division ---shorten cell division cycle,
especial interphase in which DNA synthesis.
 (2) Cell elongation---increasing in IAA level
(synthesis, antioxidation and conjugate
dehydration)
---Increasing in cell wall plasticity and
extensibility
16
80
14
12
60
Plasticity %

10

Elongation%
8 +GA
40 +GA -GA
-GA 6
4
20
2
0
0
0 10 20 30
0 10 20 30

Fig 6-17 Cell wall plasticity and elongation induced by GA

Fig 6-18 GA signal transduction
• (2) Breaking dormancy and promoting
germination of seed.
• GA3 0.5-1mg/L for GA-dependant mutants
and GA3100mg.L-1 can substitute red light
for seed germination.

• Mechnism：induction to α-amylase in
aleurone.
 α - Amy
aleurone

GA
endosperm
小 starch
盾
maltose，G
片 S
O2

scutellum Fig 6-19 GA induces α – Amy

and germination
In aerobic condition GA is synthesized in scutellum and
transported into aleurone cell where α – Amy is induced and
transported out to endosperm.The enzyme hydrolyzes starch
into maltose and then glucose and sucrose which is transported
into embryo for seedling growth.
• Application：

• Beer production.
• Breaking bud dormancy of potato with 0.5-
1.0 mgL-1 GA3.
(3) Promoting bolting and flowering
GA substitutes low temperature and
long daytime.

Normal T GA3 Low T

Fig 6-20 GA and flower (after Taiz &zeiger 2006)
• (4) Enhancing fruit setting and parthenocarpy
• Grape, apple and pear are sprayed with GA3 10-20
mgL-1 during flowering stage.

• Grape is sprayed
• with GA3 200-500
• mgL-1 for seedless
• fruits
• (5) Controlling sexual differentiation
• GA3 1000 mgL-1 ，4-5 leaf old to form male
flower in cucumber but female flower in some tree.
• Conclusion of GA function:
• (1) Promoting elongation of stem and division of
cell
• (2) Breaking dormancy and promoting
germination of seed
• (3) Promoting bolting and flowering
• (4) Enhancing fruit setting and parthenocarpy
• (5) Controlling sexual differentiation
 Section3 Cytokinin（CTK）
3.1 Discover of CTK
• 1950s, Skoog and co-workers found
extracts of vascular tissue, coconut milk,
and yeast stimulated cell division---kinetin.
---cytokinin(1965).
• 1963, Letham reported the isolation of
purine with kinetin like properties from
young, developing maize seed--- Zeatin.
 Several CTK structures
6 H CH3
H-N-R1 H-N-CH2-C=C
N6-substituted CH3
derivatives of N N
N N
nitrigenous purine Isopentenyl
N N N N adenine
R2 H
R3 H

H CH3 H CH3
H-N-CH2-C=C H-N-CH2-C- CH
CH2OH H CH2OH
N N
N N

N N N
H N
H Dihydrozeatin
H H
Zeatin
• 3.2 Transportation and Biosynthesis of CTK
• (1) Transportation of CTK
• CTK synthesized in root can be transported
up along with xylem. CTK applied in leaf or
bud surface does not move but that is
injected into phloem can be transported in
double directions.
• (2) Biosynthesis of CTK。 Root tip about
1mm。
Fig 6-18 Crown gall（冠瘿）
tumor on a tomato plant.
The stem of a one-month-old
tomato seedling was wounded
with a needle carrying a culture
of wild-type Agrobacterium
tumefaciens. The crown gall
tumor was photographed one
month later.
 Pathway of CTK biosynthesis
H CH3
Isopentenyl pyrophosphate H-N-CH2-C=C
H CH3 CH3
-3O P -O-CH -C=C N
4 2 2 N
CH3
N N
H-N-H H Isopentenyl
P-O-CH2
N
Isopentenyl O AMP
N AMP synthase
N N OH OH
H
P-O-CH2
O
H CH3
H-N-CH2-C=C H CH2 -OH
OH OH CH3 H-N- CH2-C=C
N CH3
N N
AMP N
N N
H
N N Zeatin
Isopentenyl adenine H H
H
CTK degradation
by conjugates
3.3 Physiological role and application of
CTK
(1) Enhancing cell division and enlargement

callus
CTK
CK

Cotyledon

Why? Gene expression

• (2) Inducing organ differentiation.
• Root and bud differentiation depending on
CTK/IAA ratio
CTK/IAA CTK/IAA CTK/IAA
ratio higher ratio lower ratio middle

CTK/IAA ratio from higher to lower

(3) Delaying senescence H2 O

CTK

Keep flesh：
Strawberry（10 mg L-1 KT）
orange（400 mg L-1 6-BA）
mushroom（100 mg L-1 6-BA）
PSAG12-IPT CK CK The senescence of
IPT transgene plant
and wild parent

CK PSAG12-IPT 7d storage
(4)Inhibiting dominance and enhancing lateral sprout .

Fig 6-20 Transgenic tobaco

expressing the Agrobacterium
tumefaciens ipt gene under the
control of the strong CaMV
35S promoter.
Retard leaf senescence and
early release of lateral buds

(5) Enhancing germination

of light-favored seed
• Conclusion of CTK function:
• (1) Enhancing cell division and enlargement
• (2) Inducing organ differentiation
• (3) Delaying senescence
• (4)Inhibiting dominance and enhancing lateral
sprout
• (5) Enhancing germination of light-favored
seed
 Section 4 Abscisic acid (ABA)
• 4.1 Discover of ABA
• 1963, Addicott --abscisin, Wareing --- dormin.

CH3 CH3 CH3

ABA extensively exists in
plant organs, especially in
OH dormant or abscising parts.
COOH
O CH3

Sesquiterpene-C15
4.2 Biosynthesis of abscisic acid
Two pathway: terpenoid pathway and
carotenoid pathway.
4.2 Biosynthesis of abscisic acid
Isopentenyl pyrophosphate
4.3、Physiological role and application of
ABA
(1) Inducing stomata closure
Drought signal and anti-transpiration substance,10-100
μlL-1 。

control ABA
 K+in channel ABA
ABA
Ca2+permeable
induces channel
stomatal Ca2+
closure ADP+Pi
Ca2+ S-type
H+ anion channel
ATP Ca2+ A-
R-type

e
ri z
Vacuole
anion channel

la
po
De
K+
pH
K+out channel
Fig 6-21 A guard cell model, illustrating the proposed
functions of ion channels in ABA signaling and stomatal
closing. The right of the stomatal shows ion channels and regulators that
mediate ABA-induced stomatal closing. The left cell shows the parallel
effects of ABA-induced [Ca2+]cyt increases that inhibit stomatal opening
mechamisms.
(2) Inducing bud dormancy and inhibiting
seed germination
anti-function to GA.

(3) Enhancing organ abscission and

senescence.
Chlorophyll degradation

(4) Increase in resistance as a stress hormone

Induction osmotin, dehydorin and Lea protein during

maturation or drought stress.
• Section 5 Ethylene（Eth）
• A gaseous hormone
• 5.1 Biosynthesis of Eth
• Synthesis in all part, especially in rolling
apices, ripened fruit, wounding or flooding
plants.
• Auto-catalyze feather.
Fig 6-21 pathway of ethylene synthesis
 Regulation of Eth biosynthesis
stimuli SAM inhibitor

Ripening, senescing, ACC AVG (aminoethoxyvinylglycine)

over-IAA, wounding, synthase AOA (aminooxyacetic acid)
chilling, flooding etc COO-

+
CH2
H2C NH3
C H2C NH -C=O
COO- C
H2C COO-
ACC H2C
MACC
Ripening O2 ACC anaerobic，uncouple Co2+ ,
oxidase high temperature >35oC,
CO2+HCN+H2O scavengers

CH2=CH2
 5.2 Physiological role and application of Eth

• (1) Triple responses (三重反应):

• A typical reaction of plant to ethylene
represents inhibition and swelling of
hypocotyl, inhibition of elongation , and
exaggeration of the curvature (leaf epinasty).
Fig6-22 The triple response to
ethylene of six-day-old
etiolated pea seedlings and
four-day-old etiolated mung
bean seedlings
Fig 6-23 Screen for etr1
mutant of Arabidopsis
Fig 6-24 ethylene receptor
Fig 6-25 ethylene signal
transduction
Epinasty (偏上（性）生长) : the downward
curvature of leaves occurs as the upper side of
petiole grows faster than the lower side.

Normal plant Plant under flood or ethylene

(2) fruit ripening: a regulator, (乙烯利, 2-氯乙基
膦) ClCH2CH2OP (OH)2 (water soluble)
releases ethylene when pH is larger than 4.

500-1000 mgL-1,

CK Ethylene treatment
(3) Inducing abscission
• Eth increases in the activities of Pectinase,
peroxidase and cellulase in abscission zone.
• Application: 600-800 mgL-1 as a defoliant for
cotton. Thinning fruit or flower in tea, grape and
Carya illinoensis(山核桃).
(4) Enhancing female flower formation
• 1-4 leaf old cucumber treated with 100-200 mgL-1.
(5) Enhancing secretion of secondary products
Oak etc treated with 5% of ethylene solution.
 Inhibition of growth, Increased Inhibition of
root water cell elongation,
CO2 assimilation,
uptake cell division
transpiration

Senescence Stomatal closure

Inhibition of axillary bud

growth in apical dominance
Stress condition ABA ?
Promotion of dormancy
(loss of turgor)
?
Developmental
signals
Root,shoot
Ethylene gravitropism

Auxin Stress
Cytokinin
at high at high concentration conditions
concentration activity (auxin herbicides)
Intereaction among IAA,CTK,ethylene and ABA
• Section 6 Other plant growth substance
（in brief）
• 6.1 BR ( Brassinosteroids 油 菜 素 甾 醇 类 or
Brassinolide,油菜素内酯）.
• Promote growth for section or whole
plant ， delaying senescence, increase
resistance and yield. 0.001-0.1mg/L .
• 6.2 Polyamine(多胺)
• Polyamines in plants
• Put(腐胺) NH2(CH2)4NH2
• Cad(尸胺) NH2(CH2)5NH2
• Spd(亚精胺) NH2(CH2)3NH (CH2)4NH2
• Hspd(高亚精胺) NH2(CH2)4NH (CH2)4NH2
• Spm(精胺) NH2(CH2)3NH (CH2)4NH(CH2)3NH2
• Agm(鲱精胺) NH2(CH2)5NH C (NH) NH2

• Promote Growth ，delaying senescence

• 6.3 JA (Jasmonic acid )(茉莉酸，Me-JA)
O O

COOH COOH
（+）JA （-）JA
O
O

COOH
COOH
（-）7-iso-JA （+）7-iso-JA
Inhibit growth，promote senescence and
tuberization
• 6.4 SA（Salicylic acid,水杨酸）
-COOH
-OH

Signal transduction in resistance to diseases of

plant （PRPs-pathogenesis-relative proteins）。
Enhance male flower .
• Section 7 Plant growth regulators
• 7.1 Plant growth-promoters
• Indole derivatives :IPA、IBA,
• naphthalene derivatives :NAA、NOA,
• chlorophenol derivatives: 2.4-D、2.4.5-D,
• 2.4.6- Trichlorophenoxyacetic acid,
• 2.3.6- Trichlorobenzoid acid .
• Cytokinin-like :KT、Diphenylurea 、6—BA。
• 8.2 Plant growth inhibitors
• Compounds resistant to IAAs in function and
inhibit the growth of the apical meristem. The
inhibitory effect can be reversed by IAA, but
not by GA.
• 8.2.1 TIBA (2.3.5-triidobenzoid acid) ：Inhibit IAA
polar transport. 100 mgL-1for soybean to inhibit
overgrowth, to prevent it from lodging and to
promote ripening and yield.
• 8.2.2 Morphactin: change in plant morphology for
potted plants
• 8.3 Plant growth retardants
• Compounds resistant to GAs in function and
inhibit the growth of the subapical meristem.
The inhibitory effect can be reversed by GA,
but not by IAA.

• 8.3.1 B9 (dimathyl aminosuccinamic acid，二甲基氨

基琥珀酰胺酸) 。
• To prevent plant branch from overgrowing and
to increase flower differentiation.
• 8.3.2、 CCC（Chlorochdine chloride ，矮壮素).
• To prevent wheat from logding. 0.15-0.3% in
primary elongation stage.
• 8.3.3 Pix（1. 1-dimethypiperidinium chloride,缩节
安，助壮素).
• Cotton plant with 25-150 mgL-1 during flower stage.
• 8.3.4 PP333 （Paclobutrazol,多效唑) .

• Drafting culture for fruit trees.

• Prevent late-season rice seedling from overgrowing
with 200 mgL-1 150Kg/mu at 1leaf stage.
• 7.4 Warning for application of plant
growth regulators
• (1) Chemical characters and absorption. For
absorption 2，4-D ester is easier than 2，4-D acid,
then 2，4-D salt. But water soluble products is
more convenient to use for farmer.
• (2) Concentration based on propose.
• (3) Safety and no remnants.
• (4) Economic and other culture methods.
• (5) Time of day and neighboring plants