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Chapter 6 The Plant Hormones
(Phytohormones) p309-366
Plant hormone
Plant growth substance
plant regulator
Plant hormones are naturally occurringorganic 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.
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1gIAA/10000 tips.1mgIAA/1T leaf.
10mgBR/225kg of pollen.
5 putative plant hormonesIAAGACTKABA and Eth6BR.
Plant growth regulators () arenormally used to denote synthetic
compounds that exhibit plant hormonal
activity.
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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.
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Went called it auxin.
Structure of IAA isIndole-3-acetic acid---IAA.
N
-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
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0
0.1
0.2
0.3
0 20 40 60Distance form coleoptile apix to roottip(cm)
IAArelativecontents
coleoptile apices, root tips and the apical buds of
growing stems.
Fig 6-1 IAA concentrations in plant parts
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IAA Polar transportIAA is only one plant
hormone transported unidirectionally from
apical end to basal end.
IAA transport in root from basal to tip.
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Fig 6-2 IAA Polar transport controlled by IAA transport
protein (from Taiz and Zeiger 2006)
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Other transport: IAA synthesized in mature
leaf is transported in double-directionally alongthe phloem.
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1.3 Biosynthesis of IAA
1.3.1 Biosynthesis locationalmost parts of
plant. Especially in coleoptile, young leaf and
developing seed and fertilizing ovary
1.3.2 Steps of Biosynthesis
1Tryptophan pathway: The deficiency of Zn decreases IAA synthesis
due to decreasing in Trp synthesis.
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1 Pathway dependent on tryptophan
Tryptophan(Trp) CO2
-NH2
CO2
O2
-NH2
O2
+H2O
-2H
(IAA) Indole acetic acid
N
COOH
NH2
N
NH2
*Trp decarboxylase
*Trp
transaminase
N
COOH
O
Indole-3-pyruvic acid(IPA)
IAld reductase
Tol oxidaseN
O
N
OH
Trptamine
(TAM)
Amine oxidase
*IPAdecarboxylase
Indole-3-acetaldehyde(IAld)
N
COOH
IAld
dehydrogenase
Indole-3-ethanol
(tryptophol,TOL)
NNOH
Indole-3-acetaldoxime
N
N
Indole-3-acetonitrile
(IAN) *nitrilase
N
O
NH2
Indole-3-
acetamide
(IAM)
*Trp
monoxygenase
*IAM
hydrolase
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normal IAA-over
producing
plant
Fig 6-3 IAA-over producingplant grows worse
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Fig 6-4 Effects of indole-3-acetonitrile
and IAA on wild-type (wt) and nit1-3
mutant seedlings of Arabidopsis.
Eight-day-old seedlings grown in the presence
and absence of 30 M indole-3-actonitrile(B) or
1M 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
lacks nitrilase and cannot convert indole-3-acetonitrile to IAA
A
B
C
wt
wt
wt
Nit1-3
Nit1-3
Nit1-3
Nit1(nitrilase mutant) is
insensitive to indole-3-acetonitrile
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An orange pericarp (orp) maize cob showing the expected two-
gene recessive trait, the orange kernels carry both mutant genes
2 Pathway independent on tryptophanMutants for tryptophan nutrition deficient
Maize -orp mutant, trp synthase -subnuit point
mutant, 50 time high IAA-conjugate
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Arabidopsis-trp1,trp2 and trp3 cantsynthesize Trp but content 19-30 times as
higher as IAA-conjugates of wild type
N15-anthranilate(
N15-IAA 39%N15-Trp 13%
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N
N
anthranliate( Chorismate phenylalanine tyrosine() ( )
5-phosphoribosylanthranilate
1-(o-carboxyphenylamino)
-1-deoxyribulose-5-P
N
CH2
OP
0H
OH
N
N
COOH
NH2
N
COOH
O
N
COOH
IAASerine +indole
Indole-3-glyceral
phosphate
IGP synthase
Trp synthase
Trp synthase
Trp3
Trp2,orp
IGP
Trp
PR-anthranilate
isomerase
Anthranilate
PR-transferase
anthranilate synthase(Trp1,MTR)
IPA
IAN
Trp s
ynthas
e
aminotransferase
rty?
Nitrilasenit1
N
COOH
Indole-3-butyric acid(IBA)
IBA sythase
IBA-conjugates
IAA-conjugates
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1.4 Inactive and degradation ofIAA
1.4.1 IAA-conjugate:
IAA-Amide
IAA-Sugar
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Function of IAA-conjugates
(1)Storage form.
(2)Easy transport form.
(3)Anti oxidation form.
(4)Level control and detoxification.
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1.4.2 IAA oxidation
Two enzymes: IAA oxidase and peroxidase
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1.5 Physiological effects and application
of IAA (1)Enhancing elongation of cell and
organ.
Control
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-12-10-8-6-4-202468
10
0 1 2 3 4 5 6 7 8 9 10 11 12
-Relativegrowthrate
+
10-11 10-9 10- 7 10- 5 10- 3 10- 1
IAA concentrationmol/L)
Root
Bud Stem
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.
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Mechanism for elongation by IAA 1
IAA activates genesmRNA andprotein synthesisSlow reaction.
IAAmRNA
protein
Elongation
+IAA
+IAA+actinomycinD
CK
+IAA
+IAA+cycloheximide
CKElongation
andmR
NA
Elongation
andPro
tein
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BAcid growth theory () .
rapid reactionIAA activates ATPasepH , cell wallacidification enzyme activation cell wall dehydration andloosening water absorption
Fig6-12 IAA as an effector activates ATPase (H+-pump) in
plasmic membrane
H+ H+
Microfiber of cellulose
Pentosan
other cell wall polysaccharide
Hydrogen bound
Inactive EIAA
Active EATP
ADP+Pi
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Plant hormone receptors are a group ofproteins which first bind plant hormones
and make the hormone play a serious
role in physiological functions.
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(2) Enhancing cell division and organogenesis
Cutting-shoot rooting () Cambium cell division and root primordial cell
division in the early spring.
IAACK
The rooting with 10-100 mgL-1 or 0.5-1% of powder.
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(3) Enhancing fruit setting and parthenocarpy
()production of seedless fruit
Watermelon, popper, tomato, egg plant etc aresprayed with 24-D (dichlorophenoxyacetic acid)
of 10 mgL-1 to produce seedless fruit by
parthenocarpy in early spring time or greenhouse
cultivation.
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Lateral bud develops
after removing apical
bud.
Lateral bud poorly
develops afterremoving apical bud,
then putting IAA in the
part
Lateral bud
(4) Maintaining apical dominance
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(5) Enhancing abscissionflower and fruitthinning ()Apple with NAA (naphthalene acetic acid)520 mgL-1
or NAD (naphthalene acetamide)2550 mgL-1.
(6) Inhibiting sprouting ofpotato .
1% of NAA powder
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(7) Enhancing flowering and controlling
sexual differentiation.
14 months old pineapple with 2.4D 50-100mgL-1 or NAA 1520 mgL-1 ( 30ml per plant)
Cucumber .
(8) As a herbicide to kill dicot with 24-D(1000 mgL-1 .
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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
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Section 2 Gibberellic acid (GA) 2.1 Discover of GA
Rice foolish seedling infected by Gibberellia
fujikuroi. A large family of more than 125members.
12
3 54
18
20
CH3
CH3
CH3
6
8
7
10 9
19 CH3HH
11 12
13
14
15
16 17
CH3
HH
gibberellane
1
2
3 54
18CH3
O
6
8
7
10 9
COOHH
H
11 12
13
14
15
16 17
CH2
OHH
19CO
GA3
GA is diterpenes constituting with 4 isopentenal groups
HO
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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.
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Fig 6-13 Histochemical localization of GA1promoter activity indicating CPS expression during
the development of transgenic Arabidopsis
containing the GA1 promoter-GUS gene fusion
pGA1-103.
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In plant
Copalylpyrophosphate CPP
CHO
GA127aldehyde
3CH2C-OCoA
OCH2COOH
HOCCH2
CH2CH2OH
OH OH
H3C-C=CH-CH2- O- P -O-P-OH
CH2
O O
4 molecules
Geranylgeranylpyrophosphate
GGP
OPP
Kaurene
Kaurenoic acid
various
GA
-CCC
-AMO-1618
-Phosphor-D
OPP
In microbe
Biosynthesis pathway of GA and several retardants (inhibitors for
GA biosynthesis)
COOH COOH
Mevalonic acid Isopentenyl pyrophospha(IPP)
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Fig 6-14 Inactive form:
2-hydroxylation and
GA-conjugate: GA-glucoside.
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2.3 Physiological role and application of GA
(1) Promoting elongation of stem anddivision of cell.
Function in subapices and promote
elongation of whole plant, especially ofmutants and physiological draft.
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Fig 6-15 Effect of
GA3 on stemelongation of
Progress No.9 dwarf
pea seedlings:(left)
control plants, (right)plants seven days
after treatment with 5
g GA3
.
physiological draft
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CK 100pg 1ng
Fig 6-16 Promotion of leaf sheath elongation of
Tanginbozu dwarf rice three days after treatment with
GA3.
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Application in stem or leaf-harvested
plants such as celery, lettuce, tea etc with 1-5mgL-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
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(2) Cell elongation---increasing in IAA level(synthesis, antioxidation and conjugate
dehydration)---Increasing in cell wall plasticity and
extensibility
0
20
40
60
80
0 10 20 30
+GA-GA
02
468
10121416
0 10 20 30
+GA-GA
Plasticity
%
E
longation%
Fig 6-17 Cell wall plasticity and elongation induced by GA
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Fig 6-18 GA signal transduction
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(2) Breaking dormancy and promoting
germination of seed.
GA3 0.5-1mg/L for GA-dependant mutants
and GA3100mg.L-1
can substitute red lightfor seed germination.
Mechnisminduction to -amylase inaleurone.
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Application Beer production. Breaking bud dormancy of potato with 0.5-
1.0 mgL-1 GA3.
(3) P ti b lti d fl i
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Normal T GA3 Low T
(3) Promoting bolting and flowering
GA substitutes low temperature andlong daytime.
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Fig 6-20 GA and flower (after Taiz &zeiger 2006)
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(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
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(5) Controlling sexual differentiation
GA3 1000 mgL-1 4-5 leaf old to form male
flower in cucumber but female flower in some tree.
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Conclusion of GA function: (1) Promoting elongation of stem and division of
cell
(2) Breaking dormancy and promotinggermination of seed
(3) Promoting bolting and flowering
(4) Enhancing fruit setting and parthenocarpy
(5) Controlling sexual differentiation
S ti 3 C t ki iCTK
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Section3 CytokininCTK3.1 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.
S l CTK t t
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Several CTK structures
H-N-R1
N
N
N
N
R2
R3
N6-substituted
derivatives of
nitrigenous purineN
N
N
N
HH
H-N-CH2-C=C
CH3
CH3
H
N
N
N
N
HH
H-N-CH2-C=C
CH3
CH2OH
H
N
N
N
N
HH
H-N-CH2-C- CH
CH3
CH2OH
H
H
Isopentenyl
adenine
ZeatinDihydrozeatin
6
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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 orbud surface does not move but that is
injected into phloem can be transported in
double directions.
(2) Biosynthesis of CTK Root tip about1mm
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Fig 6-18 Crown gall
tumor on a tomato plant.
The stem of a one-month-old
tomato seedling was wounded
with a needle carrying a cultureof wild-type Agrobacterium
tumefaciens. The crown gall
tumor was photographed one
month later.
Pathway of CTK biosynthesis
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Pathway of CTK biosynthesis
N
N
N
N
H
H-N-H
OP-O-CH2
OH OH
Isopentenyl pyrophosphate
AMP
Isopentenyl
AMP synthase
-3O4P2-O-CH2-C=C
CH3
CH3
HCH2-C=C
CH3
CH3
H
N
N
N
N
H
H-N-
OP-O-CH2
OH OH
IsopentenylAMP
CH2-C=C
CH3
CH3
H
H-N-
N
N
N
NH
HIsopentenyl adenine
CH2-C=C
CH2
CH3
H
H-N-
N
N
N
NH
H
-OH
Zeatin
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CTK degradation
by conjugates
3 3 Physiological role and application of
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3.3 Physiological role and application of
CTK(1) Enhancing cell division and enlargement
CTKcallus
Cotyledon
CK
Why? Gene expression
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(2) Inducing organ differentiation.
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Root and bud differentiation depending onCTK/IAA ratio
CTK/IAA
ratio higherCTK/IAA
ratio lower
CTK/IAA
ratio middle
CTK/IAA ratio from higher to lower
(3) Delaying senescence
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(3) Delaying senescenceH2O
CTK
Keep flesh
Strawberry10 mg L-1 KT
orange400 mg L-1 6-BA
mushroom100 mg L-1 6-BA
P IPT CK CK Th f
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PSAG12-IPT
PSAG12-IPT
CK CK
CK
The senescence of
IPT transgene plantand wild parent
7d storage
(4)Inhibiting dominance and enhancing lateral sprout .
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Fig 6-20 Transgenic tobacoexpressing the Agrobacterium
tumefaciens iptgene 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
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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)
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Section 4 Abscisic acid (ABA)
4.1 Discover of ABA
1963, Addicott --abscisin, Wareing --- dormin.
CH3
COOH
CH3CH3
CH3
OH
O
Sesquiterpene-C15
ABA extensively exists in
plant organs, especially in
dormant or abscising parts.
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4.3Physiological role and application of
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ABA(1) Inducing stomata closureDrought signal and anti-transpiration substance,10-100
lL-1
control ABA
ABACa2+permeable
K+in channelABA
i d
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Ca2+
Ca2+
p
channel
Vacuole
Depolarize
K+
K+out channel
A-
S-type
anion channelR-type
anion channel
pH
Ca2+H
+
ATP
ADP+Pi
Fig 6-21 A guard cell model, illustrating the proposed
functions of ion channels in ABA signaling and stomatalclosing. The right of the stomatal shows ion channels and regulators thatmediate ABA-induced stomatal closing. The left cell shows the parallel
effects of ABA-induced [Ca2+]cyt increases that inhibit stomatal opening
mechamisms.
induces
stomatal
closure
(2) Inducing bud dormancy and inhibiting
seed germination
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seed germination
anti-function to GA.
(3) Enhancing organ abscission andsenescence.
Chlorophyll degradation
(4) Increase in resistance as a stress hormone
Induction osmotin, dehydorin and Lea protein during
maturation or drought stress.
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Section 5 EthyleneEth
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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.
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Fig 6-21 pathway of ethylene synthesis
Regulation of Eth biosynthesis
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MACCACC
SAM
AVG (aminoethoxyvinylglycine)
AOA (aminooxyacetic acid)
inhibitor
Ripening, senescing,
over-IAA, wounding,
chilling, flooding etc
stimuli
CH2=CH2
O2
CO2+HCN+H2O
Ripening anaerobicuncouple Co2+ ,
high temperature >35oC,
scavengers
ACC
oxidase
ACC
synthase
H2C
H2C
CNH3
+
COO-H2C
H2C
CNH
COO-
COO-
CH2
-C=O
5.2 Physiological role and application of Eth
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(1) Triple responses (): A typical reaction of plant to ethylene
represents inhibition and swelling ofhypocotyl, inhibition of elongation , and
exaggeration of the curvature (leaf epinasty).
Fig6-22 The triple response to
ethylene of six-day-old
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y y
etiolated pea seedlings andfour-day-old etiolated mung
bean seedlings
Fig 6-23 Screen foretr1
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mutant of Arabidopsis
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Fig 6-24 ethylene receptor
Fig 6-25 ethylene signal
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transduction
Epinasty () : the downward
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curvature of leaves occurs as the upper side ofpetiole grows faster than the lower side.
Normal plantPlant under flood or ethylene
(2) fruit ripening: a regulator, (, 2-
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) ClCH2CH2OP (OH)2 (water soluble)releases ethylene when pH is larger than 4.
Ethylene treatment
500-1000 mgL-1,
CK
(3) Inducing abscission
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Eth increases in the activities ofPectinase,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().(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,
CO2 assimilation,
Increased
root water
uptake
Inhibition of
cell elongation,
cell division
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Cytokininat high
concentration
Auxinat high concentration
activity (auxin herbicides)
Stressconditions
Ethylene
ABAStress condition(loss of turgor)
Developmental
signals
Senescence Stomatal closure
Root,shootgravitropism
Inhibition of axillary bud
growth in apical dominance
Promotion of dormancy
transpirationp
?
?
Intereaction among IAA,CTK,ethylene and ABA
Section 6 Other plant growth substance
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in brief 6.1 BR ( Brassinosteroids or
Brassinolide,.
Promote growth for section or whole
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plant delaying senescence, increaseresistance and yield. 0.001-0.1mg/L .
6.2 Polyamine()
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Polyamines in plants Put() NH2(CH2)4NH2 Cad() NH2(CH2)5NH2 Spd() NH2(CH2)3NH (CH2)4NH2 Hspd() NH2(CH2)4NH (CH2)4NH2
Spm() NH2(CH2)3NH (CH2)4NH(CH2)3NH2 Agm() NH2(CH2)5NH C (NH) NH2
Promote Growth delaying senescence
6.3 JA (Jasmonic acid )(Me-JA)
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O
COOH
O
COOH
O
COOH
O
COOH
+JA -JA
-7-iso-JA +7-iso-JA
Inhibit growthpromote senescence and
tuberization
6.4 SASalicylic acid,
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-COOH
-OH
Signal transduction in resistance to diseases of
plant PRPs-pathogenesis-relative proteins
Enhance male flower .
Section 7 Plant growth regulators
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7.1 Plant growth-promoters
Indole derivatives :IPAIBA,
naphthalene derivatives :NAANOA, chlorophenol derivatives: 2.4-D2.4.5-D,
2.4.6- Trichlorophenoxyacetic acid,
2.3.6- Trichlorobenzoid acid .
Cytokinin-like :KTDiphenylurea 6BA
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8.3 Plant growth retardants
C d i t t t GA i f ti d
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Compounds resistant to GAs in function andinhibit 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)
To prevent plant branch from overgrowing andto increase flower differentiation.
8.3.2 CCCChlorochdine chloride ). To prevent wheat from logding. 0.15-0.3% in
primary elongation stage.
8.3.3 Pix1. 1-dimethypiperidinium chloride,
)
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). Cotton plant with 25-150 mgL-1 during flower stage.
8.3.4 PP333 Paclobutrazol,) .
Drafting culture for fruit trees. Prevent late-season rice seedling from overgrowingwith 200 mgL-1 150Kg/mu at 1leaf stage.
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