สรีรวิทยาของพืชประยุกต์ ชว310 ethylene... ·...
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Abscisic acid (ABA):
A Seed maturation & antistress signal
Dr.Yuwalee Unpaprom1
สรีรวิทยาของพืชประยกุต ์ชว310
S-Abcisic acid
(S-ABA)
Hormones in PlantSource: Santner et al., 2009 2
Abscisic Acid (ABA)
Physiologists suspected that the
phenomena of seed & bud
dormancy were caused by inhibitory
compounds (dormin).
Winter dormancy
& chilling in
woody plants
Discovery that dormin was
chemically identical to a
substance that promotes the
abscission of cotton fruits, abscissin II
Renamed abscisic acid (ABA)
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Occurrence, Chemical Structure, and Measurement of ABA
ABA is a ubiquitous plant hormone in vascular plants.
It has been detected in
mosses but appears to be absent in liverworts.
ABA has been detected in
living tissues from the root
cap to the apical bud.
It is synthesized in almost all
cells that contain chloroplasts or amyloplasts.
Chloroplasts in plant cells Potato amyloplasts
The chemical structure of ABA
determines its physiological activity
ABA is a 15-C compound that
resembles the terminal portion of
some carotenoid molecules.
Nearly all the naturally
occurring ABA is in the cis form
ABA also has an asymmetric C
atom at position 1´ in the ring,
resulting in the S and R(or+ and -)
enantiomers.
The S and R forms cannot be interconverted in the plant tissue.
ABA is assayed by biological, physical, and chemical methods
A variety of bioassays have
been used for ABA
inhibition of germination
inhibition of GA-induced α-
amylase synthesis
15
+ABA GA
Coleoptile growth inhibition.
Biosynthesis, Metabolism, and Transport of ABA
ABA is synthesized from a
carotenoid intermediate
Biosynthesis takes place in chloroplasts and other plastids.
The pathway begins with
isopentenyl diphosphate (IPP) and leads to the
synthesis of the C40 xanthophyll violaxanthin.
Violaxanthin
Xanthoxal (C15)
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Maize mutants (viviparous; vp) that are blocked at
other steps in the carotenoid pathway; exhibit vivipary
Synthesis of 9-cis-epoxycarotenoid
dioxygenase (NCED) is rapidly
induced by water stress.
Upon rewatering, the ABA
level declines to normal in the same amount of time.
ABA is translocated in vascular tissue ; xylem, phloem
As water stress begins, some of
the ABA carried by the xylem
stream may be synthesized in roots
that are indirect contract with the drying soil.
The major control of ABA
distribution among plant cell
compartments follows the “anion trap” concept.
ABA closes stomata in response
to water stress
Elucidation of the roles of ABA in
freezing, salt, and water stress led to
the characterization of ABA as
a stress hormone.
Biosynthesis of ABA is very
effective in causing stomatal closure.
ABA promotes seed storage reserve
accumulation and desiccation
tolerance
During mid – to late
embryogenesis, when seed ABA
levels are highest, seeds
accumulate storage compounds
that will support seedling growth
at germination.
As maturing seeds
begin to lose water,
specific mRNA
encoding so-called late-
embrogenesis-abundant
(LEA) proteins thought
to be involved in
desiccation tolerance
accumulate in embryos.
ABA closes stomata in response
to water stress
Elucidation of the roles of ABA in
freezing, salt, and water stress led to
the characterization of ABA as a stress hormone.
ABA promotes leaf senescence
independently of ethylene
ABA is involved in
leaf senescence, it
might indirectly
increase ethylene
formation & stimulate abscission.
Leaf segments senesce faster in darkness
than in light, and turn yellow as a result of chlorophyll breakdown.
Ethylene:
The gaseous Hormone
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Coal gas was used for street illumination, trees defoliated
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Coal gas & air pollution affect
plant growth
Ethylene (C2H4) was active component of coal gas
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First indication that ethylene is a natural product of plant tissues:
Published by H. H. Cousins in
1910
Cousins reported; emanation
from oranges caused the premature ripening of bananas
However, oranges synthesized
little ethylene compared to
other fruits
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In 1934, R. Gane & others identified
ethylene chemically as natural product of
plant metabolism
For 25 yrs ethylene was not recognized
After GC was introduced in ethylene in 1959, plant growth regulator was recognized
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STRUCTURE, BIOSYNTHESIS & MEASUREMENT OF ETHYLENE
Ethylene can be produced
by almost all parts of higher
plants
Meristematic & nodal
regions are the most active in ethylene biosynthesis
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The Properties of Ethylene are Deceptively Simple
Molecular weight: 28
Lighter than air
It is flammable and readily
undergoes oxidation; can be oxidized to ethylene oxide
Ethylene oxideEthylene
[o]
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Ethylene oxide can be hydrolyzed to ethylene glycol
Ethylene glycol
odorless,
colorless,
syrupy,
sweet-tasting
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In plants, ethylene can be
oxidized to CO2
Ethylene oxideEthylene
[O] [O]HOOC COOH CO2
Oxalic acid
Ethylene is released easily from
tissues & diffuses through the
intercellular spaces
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Ethylene gas is easily lost from
tissue and may effect other
tissues or organ
Ethylene-trapping system are
used during storage fruits,
vegetable and flowers: KMnO4
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The production of ethylene in
plants is highest in senescing
tissues & ripening fruits
Active at very low conc.- less
than 1 ppm (1µL/L)
Internal ethylene conc. In a ripe
apple: 2500 µL/L
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Regulated Biosynthesis Determines
the Physiological Activity of Ethylene
Plant tissues convert methionine to
ethylene
Yang cycle
Precursor of ethylene:
1-aminocyclopropane-1-carboxylic
acid
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AOA: aminooxyacetic acid
AVG: aminotoethoxy-vinylglycine
SAM
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Fruit ripening: ACC and ethylene
increase
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Stress-induced ethylene production:
drought, flooding, chilling, expose
to ozone, wounding
Auxins-induced ethylene
production: to promote ethylene
synthesis by enhancing ACC
synthase
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Inhibitors of ethylene synthesis:
AVG, AOA, Co2+
Inhibitors of ethylene action: Silver
ions (Ag+), applied as silver nitrate
(AgNO3) or silver thiosulfate
(Ag(S2O3)23-
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Inhibitors of ethylene: CO2 at high
conc. (5-10%)
The volatile compound trans-
cyclooctene (but not cis-) is strong
competitive inhibitor of ethylene
binding (1-methylcyclopropene:
MCP)
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Ethylene Promotes the Ripening of
Some Fruits
Fruit ripening refers to changes in
fruit that make it ready to eat.
Such: softening, starch hydrolysis,
sugar accumulation, disappearance
of organic acids & phenolic
compound
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Climacteric fruits
Nonclimacteric fruits
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Breaks seed & bud dormancy in
some sp.
Induced flowering in the pineapple
family
Enhances the rate of leaf
senescence
Induces the formation of roots &
root hairs
Other Eeffects of Ethylene
Salicylic Acid:
A phenolic phytohormone
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Salicylic acid (SA): from Latin
salix, willow tree (salicin)
A phenolic phytohormone
64
Roles in plant growth and
development
Photosynthesis, transpiration, ion
uptake and transport
Induces specific changes in leaf
anatomy and chloroplast structure
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SA is involved in endogenous
signaling, mediating in plant defense
against pathogens
inducing the production
of pathogenesis-related
proteins
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The signal can also move to nearby
plants by salicylic acid being
converted to the volatile ester, methyl
salicylate.
It is involved in the systemic
acquired resistance (SAR)
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Salicylic acid is biosynthesized from
the amino acid phenylalanine
Sodium salicylatephenylalanine Salicylic acid
Commercially prepared:
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Unripe fruits and vegetables are
natural sources of salicylic acid:
blackberries, blueberries,
cantaloupes, dates, raisins, kiwi
fruits, guavas, apricots, green
pepper, olives, tomatoes, radish
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Salicylic acid is known for its ability to
ease aches and pains and reduce fevers
a key ingredient in many skin-care
products