Plant Hormones IV:

ABA, Ethylene,
Chapter 21
 Learning objectives
• Know the pathways for the synthesis of
abscisic acid and ethylene.

• Understand the physiological effects of

these three plant hormones.

• Understand the signalling pathways by

which these hormones exert their effects.
 Abscisic acid
• Unlike auxin, cytokinins, and gibberellins,
only one molecule represents the hormone
abscisic acid (ABA).

• This hormone was named based upon the

erroneous assumption that it was
responsible for the abscission of leaves
and other tissues.
 Abscisic acid
• Figure 21.1
 Abscisic acid
• The primary functions of ABA include:
– Prevention of precocious germination
(precocious germination - refers to seed germination before
the fruit has released seed)
– Initiation and maintenance of seed dormancy
– Stomatal control
– Protection of cells from desiccation (extreme dryness)

• Other functions may include:

– Induction of storage proteins in seeds
– Heterophylly (presence of differently shaped leaves on the same
shoot or on different shoots of the same plant)
– Initiation of secondary roots
– Flowering and senescence
 Definition: Seed Dormancy
• Seed dormancy is a condition of plant seeds that prevents germinating under
optimal environmental conditions
• Living, non dormant seeds germinate when soil temperatures and moisture
conditions are suited for cellular processes and division; dormant seeds do not.
• One important function of most seeds is delayed germination, which allows
time for dispersal and prevents germination of all the seeds at same time. The
staggering of germination safeguards some seeds and seedlings from suffering
damage or death from short periods of bad weather or from transient
herbivores; it also allows some seeds to germinate when competition from
other plants for light and water might be less intense. Another form of delayed
seed germination is seed quiescence, which is different than true seed
dormancy and occurs when a seed fails to germinate because the external
environmental conditions are too dry or warm or cold for germination. [ Many
species of plants have seeds that delay germination for many months or years,
and some seeds can remain in the soil seed bank for more than 50 years
before germination
 ABA synthesis
• ABA is synthesized from the -
carotene violaxanthin, typically in
mature leaves.

• Synthesis begins in the

chloroplast, beginning with G3P
and pyruvate, and ending with
xanthoxin.

• Synthesis in completed in the

cytosol (fluid portion of a cell's
cytoplasm, which lies outside the
organelles and other insoluble
components of the cytoplasm)
where xanthoxin is converted to
ABA.
 ABA synthesis
• Figure 21.2
 ABA synthesis
• Based upon the expression of genes
encoding enzymes involves in ABA
synthesis, there is also evidence that ABA
is synthesized in:
– Guard cells
– Phloem parenchyma
– Xylem parenchyma
 Transport and storage of ABA
• ABA is highly mobile within plants.

• ABA tends to accumulate in sink tissues,

including seeds.

• During periods of water stress, ABA can

be stored in roots.
 Transport and storage of ABA
• ABA may also be stored in chloroplasts.

– In the cytosol, the pH favors the

protonated form of ABA (i.e., ABAH)

– In a chloroplast not undergoing

photosynthesis, the stromal pH of 7.5
favors the formation of deprotonated
ABA (i.e. ABA-), trapping the ABA in
the chloroplast.

– During photosynthesis, the stromal

pH decreases to a value of 6.5,
facilitating the formation of ABAH and
the release of ABA to the cytosol.
 Transport and storage of ABA
• Figure 21.4
 Control of ABA activity
• ABA is metabolized quickly in plants.

• One pathway to control ABA activity

involves the formation of a glucose ester.

• The primary pathway involves the

oxidation of ABA to phaseic acid (PA)
followed by reduction to dihydrophaseic
acid (DPA).
 Functions of ABA
• ABA regulates the maturation of the embryo and
the germination of seeds.
– During embryogenesis, ABA levels peak during the
maturation stage.
– ABA imposes dormancy on the embryo, preventing
vivipary (precocious germination).
– ABA also induces desiccation of the seed, which also
contributes to seed dormancy.
A period in an organism's life cycle when growth, development is
temporarily stopped. This minimizes metabolic activity and therefore
helps an organism to conserve energy.
 Functions of ABA
• ABA also has important roles in the
response to water stress.
– ABA accumulates in water-
stressed leaves where it inhibits
stomatal opening.
– Some of the ABA comes from
storage pools in the cell
apoplast.
– The acidification of the
chloroplast caused by water
stress allows stored ABA to be
released and transported to the
guard cells.
– Increased rates of ABA
synthesis help to maintain
stomatal closure during water
stress.
 Functions of ABA
• Figure 21.5
 Functions of ABA
• ABA also has important roles in
the response to water stress.
– The transport of ABA from
roots to shoots is triggered
by decreases in soil
moisture.
– This feed-forward signal
initiates stomatal closure
before there is a change in
leaf water potential.
– The presence of ABA
reduces stomatal
conductance, helping to
preserve water.
 Functions of ABA
• Figure 21.6
 Functions of ABA
• Additional roles for ABA include:
– Lateral or secondary root development.
– A possible role in the suppression of flower
formation.
 ABA signalling
• One possible ABA receptor that has been
identified is the ABAP1 protein, located in
the plasma membrane of aleurone cells.
(outermost cell layer of the endosperm)
Three other ABA receptors may include:
– The chloroplast protein magnesium
protoporphyrin-IX chelatase H subunit.
– The flowering-time control protein FCA.
– A G-protein identified as GCR2.
 ABA signalling
• The signalling pathway for ABA includes
the following components:
– Ca2+ is an important second messenger,
especially in guard cells, where it activates
membrane anion channels.
– Transcription factors known as ABA response
element binding factors activate ABA-induced
genes that promote the synthesis of
osmolytes, or compatible solutes.
 ABA signalling
• The signalling pathway for ABA includes
the following components:
– ABA may regulate specific aspects of gene
expression in seeds.
– Protein phosphorylation, via kinases and
phosphatases, is regulated by ABA.
 Ethylene synthesis
• Figure 21.8
 Ethylene
• Ethylene is a small molecule that has key
roles in plant growth and development.

• Ethylene can influence root and shoot

growth, but is primarily associated with
stress, senescence, and ripening.

• Ethylene can be found in all plant tissues,

although the concentration can vary.
 Ethylene synthesis
• Ethylene is synthesized by a three step pathway.
– The precursor methionine is converted to S-adenosylmethionine
(SAM) by the enzyme SAM synthetase.
– SAM is converted to 1-aminocyclopropane-1-carboxylic acid
(ACC) by ACC synthase.
– ACC is converted to ethylene by ACC oxidase.
 Ethylene synthesis
• Sustained synthesis of ethylene requires the
sulfur-containing by-product of ACC synthesis to
be recycled to methionine.

• The recycling of methionine occurs via the Yang

cycle.
 Ethylene synthesis
• Figure 21.9
 Ethylene synthesis
• Ethylene synthesis is influenced by:
– Auxin
– Wounding
– Water stress
– Temperature
– Inhibitors of RNA and protein synthesis

• The effects of these factors on synthesis occurs

at the transcriptional level via ACC synthase.
 Control of ethylene activity
• Ethylene activity can be controlled by:
– Oxidation to carbon dioxide
– Conversion to ethylene oxide
– Conversion to ethylene glycol

• Ethylene activity can simulated with 2-

chloroethylphosphonic acid (ethephon),
which decomposes to form ethylene.
 Functions of ethylene
• Ethylene is involved in the ripening and
senescence of fruits.

• The synthesis of ethylene in fruits differs between

climacteric and non-climacteric fruit.
(Climacteric fruit continue to ripen after harvest, non-climacteric fruit do not)

• Ethylene stimulates the elongation of above ground

vegetative tissues but inhibits root elongation.

• Gibberellins can antagonize the effects of ethylene

on root and shoot growth.
 Functions of ethylene
• Ethylene is involved in epinasty, a downward
curvature of leaves caused by excessive cell
elongation on the adaxial (upper) side of the
leaf.

• Epinasty is a typical response for flood-sensitive

plants.

• Ethylene also:
– Promotes seed germination
– Inhibits bud break
– Reduces apical dominance
 Ethylene receptors and signalling
• A well known effect of ethylene is the triple
response observed in etiolated seedlings.
– Inhibited hypocotyl and elongation
– Radial swelling of hypocotyl
– Exaggerated curvature of plumular hook
• The various triple response mutants that exist
were useful in determining the ethylene signalling
pathway.

• seedlings grown in the dark develop long hypocotyls and their cotyledons remain closed around the
epicotyl in an apical hook. This is referred to as skotomorphogenesis or etiolation. Etiolated
seedlings are yellowish in color aschlorophyll synthesis and chloroplast development depend on
light. They will open their cotyledons and turn green when treated with light.