Abscisic Acid

• This hormone antagonizes the actions of the other hormones that we discussed earlier. It
plays a role towards the end of the life of plants and is associated with different kinds of stress
and seed dormancy. So this is a hormone that inhibits growth in general (produces dormancy,
closure of the stomates, increases during stress conditions and so on).

• It is made of 15 carbons, and it can be present in cis or trans form. The cis form is the active
form and it is naturally found. The cis form can be present as S (more active) or R. Cis and
trans forms are interconvertible.
It is a cyclic compound, and it is derived from isopentenyl pyrophosphate. Recall the IPP is
essential as a precursor for GA, cytokinins, steroids, carotenoids, brassinosteroids… so it is
used to synthesize many important biologically active compounds. But at the same time, IPP
is involved in the synthesis of hormones that delay growth like ABA. This is interesting
because the intermediate (IPP) is the same. This is controlled depending on the
developmental stage of the plant. The synthesis takes place mainly in the chloroplast.

• Biosynthesis of Abscisic Acid:

2 molecule of IPP (IPP gives rise to cytokinins) à Geranyl Pyrophospahte à Farnesyl
Pyrophosphate (15C) (gives rise to Brassinosteroids) à GernaylGeranyl Pyrophosphate (20C)
(gives rise to GA) →→ violaxanthin (40C). This compound is an example of xanthophylls
(oxygen containing carotenoids) → Neoxanthin (also 40C) (it is a potent inhibitor / can inhibit
growth) → xanthoxal (15 C) and this compound can act as a growth inhibitor (similar
properties of ABA). Then oxidation of xanthoxal à ABA aldehyde, the immediate precursor
of ABA. This synthesis occurs mainly in the chloroplasts as well as in other plastids in the
plants.

• Transport:
Once ABA is synthesized, it will be transported. The transport occurs in the xylem as well as
in the phloem. Though it is more abundant in the phloem compared to the xylem.

• Inactivation:
- By Oxidation: ABA oxidase will oxidize ABA to Phaseic Acid (PA). This intermediate can still
perform some ABA functions (→ incomplete degradation of ABA). Among these activities:
stomatal closure and inhibition of production of the amylases / hydrolytic enzymes that are
stimulated by GA (so it counteracts GA).
Further oxidation of Phaseic Acid will lead to 4’-duhydrophaseic acid (DPA). This oxidized form
of ABA is completely inhibited.
 - By conjugation: it is a kind of storage form. An example of a conjugated molecule is glucose.
Glucose has OH groups that can form ester linkages with the carboxyl groups of ABA → ABA-
β-D-glucosylester. This is an inactivated form. Esterase can break this bond, resulting in the
active form of ABA (so this is reversible form of inactivation). Usually, the stored form is in
the vacuole.

• Physiological effects of ABA:

1) Stomatal Closure:
ABA concentrations can fluctuate dramatically depending on the developmental stage of
the plant.
Under water stress, the concentration of ABA increases to 50x in stomates within few
hours → Permeability of membrane is affected → K+ will leave the guard cells → water
potential decreases → water will leave → closure of stomata
If stomata do not close under water stress, wilting will occur.
Wilt mutants are plants that cannot produce ABA → they wilt whenever subjected to
water stress. If ABA was applied to these plants, the effect will be reversed.
Equisitum is a naturally occurring plant that does not produce ABA → their stomates
remain open all of the time.

2) Bud Dormancy:
Mentioned before.

3) Seed Dormancy:
When a seed is undergoing dormancy, level of ABA Is around 100 times more than a seed
that is not in dormancy. This shows that ABA could be released from sources where it is
stored rather than synthesized directly.
For seeds to germinate, there are 2 conditions that must be available: water and oxygen.
Sometimes, even if these are available, and suitable temperature is available, the seed
will not germinate and it will undergo dormancy. Why? Dormancy might be due:

i. Coat-induced dormancy: the seed is surrounded by a seed coat that inhibits its
germination. The embryo cannot germinate unless the seed coat is removed. The
radical is the first part of the seed that will leave the coat. The coat might:
- Acts as a mechanical constraint (very hard coat, ex: peaches and apricot)
- Prevents water uptake
- Prevents gas exchange (seed coat is permeable to gases)
- Prevents the escape of inhibitors like ABA (retention of growth inhibitors)
- Source of inhibitors itself.
 ii. Embryo Dormancy: the embryo itself contains certain inhibitors, and these inhibitors
are mostly ABA. You can find these inhibitors surrounding embryo or in cotyledons…
Sometimes if you remove the cotyledons, the seeds germinate. In fact, it is the ratio
of ABA to GA that determines whether germination will occur or not (GA promotes
germination while ABA inhibits it).
There are also two types of dormancy in the seed:

i. Primary dormancy: the seed will be cultivated or produced by the plant as a dormant
seed. So the seed is dormant when it is released from the plant.
ii. Secondary dormancy: some seeds become dormant after they are released from the
plant. The plant seed could germinate if it is taken directly from the plant. But if you
take it and store it under unfavorable conditions, it will become dormant with time.
Sometime, you can overcome the dormancy by chilling period (exposing it to 0-10 C). Far
red light can also stop dormancy. So there are certain environmental factors that result
in the buildup of a level of hormones that promote growth, and this will overcome
dormancy. Sometimes, the seeds released are dormant, but when they are stormed in
conditions of water, they will moisten and germinate.

Recall that in germinating barley seeds, GA will induce formation of α-amylase, which will
cause mobilization of starch. ABA inhibits the synthesis of these GA induced hydrolytic
enzymes. How does ABA inhibit germination?

This is done by 2 ways: One is that ABA can induce the production of a protein called VP1,
and this protein will repress the synthesis GA regulated enzymes. The other way, if you
recall the MYB transcription factor that stimulates the production of enzymes like
amylase, ABA represses the production of GA-MYB → hydrolytic enzymes are not
produced. So the role of ABA is either the inhibition of the transcription factor MYB or by
the production of the VP1 protein.

• One might expect that ABA would be involved in the abscission of fruits and leaves à not
true. The abscission of leaves is due to the hormone ethylene rather than ABA.
• At the beginning of plant development, there are more GA and cytokinins produced.
• At the end of plant’s life and development, more ABA is produced.