Hormones

I. Auxin
1. Plant growth phenomena and discovery of auxin
- Phototropism: the bending of plants toward light (by differential growth)
- 1st growth hormone to be discovered
- Mutations are lethal
- Required more(less) continuously
- Darwins and his son did an experiments about phototropism of plants

2. Biosynthesis and Metabolism of auxin

- In mid 1930s, auxin is determined IAA( indole-3-acatic acid)
- IAA is abundant, simple to synthesis
- Compounds which have same biological activities as IAA can be counted as auxin (elongate in coleoptile, stem
sections ; etc)
- Chemical structure is diverse but still have a common (fractional positive charge on aromatic ring and a
negatively charge carboxyl group)
a. Synthesis of IAA in meristems, young leaves and developing fruits and seeds:
+ detected when GUS(beta-glucuronidase) expression occurs by histochemistry
=> auxin is created by group of cells where hydathode will appear
b. Multiple pathways for biosynthesis
+ The IPA path way
+ The TAM pathway
+ The IAN pathway
+ There are some evidence show that plants can synthesize IAA through tryptophan-independent way
 Multiple pathways to synthesize, plants will unlikely lack of auxin.
+ also, IAA can be synthesized from indole/ indole-3-glycerol phosphate
+most in covalently bound form
+ highest concentrations: apical meristems of shoots, young leaves
( there are the primary sites of auxin synthesis) and ubiquitous in plants
c. Multiple ways for biodegradation IAA:
 + peroxidative enzymes ( unclear)
+ decarboxylation pathway : minor roles
+ Nondecarboxylation pathways: most common metabolic pathways
3. Auxin Transport: (polar transport)
- The only plant growth hormone with polar transportation
- Requires energy and gravity:
+ experiment: roots grow even bamboos are upside down.
+ cell-to-cell way: auxin get out of the cell through membrane( efflux), diffuse and get into other cell through
membrane (influx)
- A chemiosmotic model to explain (according to textbook)
+ 1. IAA enters the cell either passively in the undissociated form IAAH or IAA-
+ 2. The cell wall is maintained at acidic pH by the acitiviy of the plasma membrane H+ATPase
+ 3. In the cytosol, which has a neutral pH , the anionic form IAA- predominates
+ 4. The anions exit the cell via auxin anion efflux carriers that are concentrated at the basal ends of each cell
in the longitudinal pathway
- Protein transport

4. Physical effects:
a. Cell elongation
b. Phototropism and gravitropism
5. Developmental effects
- Regulate apical dominance
+ auxin (other parts to apical bud) inhibit the outgrowth of the axillary bud
- Delays the onset of leaf abscission
+ (from the blade) usually prevents the abscission
+ when plants stop producing auxin, abscission is triggered during leaf senescence
- Regulate floral bud development
- Promotes fruit development
- Induces vascular differentiation
- Synthetic auxin has other uses in reality
+ herbicides, seedless fruits etc
6. Auxin signal transduction pathways
a. Auxin receptor: wait for signal to destroy the repressor => growth genes are turned on
- Auxin-binding protein ABP1
- Calcium and intracellular pH (complex)
b. Auxin-induced genes;
- Early genes: when the activation of preexisiting transcription factors simulates the expression of genes
+ cannot be blocked by inhibitor of protein synthesis
+ has 3 main functions (textbook)
- Early genes for growth and development
- Early genes for stress adaptations

II. Cytokinin ( regulator of cell division)

1. Discovery
- In a research of cell division stimulation
- 1961: after lots of experiments, researches, cytokinin was found in plants (zeatin)
2. Chemical structure:
 - Aminopurine ring
- N6 side chain
- Cytokinins Occur in Both Free and Bound Forms
- The Free Base: hormonally active cytokinin
3. Biosynthesis:
- Found the most in root apical meristem then root cap cells, ovules, phloem cells, etc
- Other organisms: bacteria fungi, insects, nematodes.
a. Pathway: (textbook)
- IPT Catalyzes the First Step in biosynthesis (isopentenyl transferase enzyme that transfers the isopentenyl
group from DMAPP to AMP (adenosine monophosphate) to form
isopentenyl adenine ribotide)
+ 1. The tumor is initiated when bacteria enter a lesion and attach themselves to cells.
+ 2. virulent bacterium carries a Ti plasmid in addition to its own chromosomal DNA. The plasmid‘s T-DNA
enters a cell and
integrates into the cell‘s chromosomal DNA.
+ 3. Transformed cells proliferate to form a crown gall tumor.
- Transported from the Root to the Shoot via the Xylem
- There is a signal from the shoot to regulate the amount of hormone transported from roots(not determined)
- Metabolized by Plant Tissues very fast
4. Biological roles:
a. Regulate Cell Division in Shoots and Roots
b. Regulate Specific Components of the
Cell Cycle
III. Abscisid acid
1. Discovery

Plant physiologists presumed that the phenonmena of seed dormancy was caused by a repressive
substances for a long time. After a lot of experiments with various types of plants tissues, particularly
dormant buds, they found that the compound (called dormin) was identical to Absicis acid (ABA). It
represses growth and stomatal opening especially in environmental stresses. Moreover, it regulates
seed maturation and dormancy.

- Mid 1960s, three inhibitors was reported=> chemically identical (ABA)

a. Occurrence
- Ubiquitous hormone in vascular plants
- Is synthesized in almost cells that contains chloroplasts, amyloplasts
b. Chemical structures ( figures in textbook)
- Determines its physiology activity
- Naturually in cis form (by convention known as ABA)
- S enantiomer only response fast to ABA
 - Long term (seed maturation) both enantiomers are active
- Any changes in structure results in loss in activity
2. Biosynthesis of ABA
- In chloroplasts, other plastids

The pathway :

- Begin with IPP ( isopentenyl diphosphate) 5-C

- synthesized by cleavage of the c40 hydroxylated carotenoid called zeaxanthin.
- Zeaxanthin is converted to violaxanthin in by the enzyme zeaxanthin oxidase red EP
violaxanthine is then cleaved to form the compound called xanthoxin C-15, an growth inhibitor
by the enzyme NCED (epoxy carotenoid dioxygen )
- Xanthonin converted to abscisic aldehyde and then abscisic acid mutants defective in ABA
synthesis have proved to be very useful.
a. ABA biosynthesis and concentrations can dramatically fluctuate in different plant species
during:
- Development
- In response to environmental stress
- Caused by :
+ Biosynthesis regulates ABA concentration
+ in cytosol Degradation
+ compartmentatation
+ conjugation
+ transport
b. In identifying many of the steps in ABA synthesis inactivate of ABA occurs through the
formation of hydroxylated products by the action of cytochrome p450 oxidases which are
futher converted to physic acid and dihydrophilic acid. ABA can also be inactivated by the
formation of glucose esters.

These esters are transported to the endoplasmic reticulum or vacuoles for storage and free ABA is
released when required by the action of glucose located in these organelles.
 c. ABA is synthesized in vascular tissue of leaves fron where it is transport by an intercellular
pathway to the ABA resposive tissues like guard cells. Active transport of ABA across cell
membranes takes place by a ATP binding cassette or ABC transporters. These transporter
are involved in active transport of abscisic acid across membranes and is accompanied by
ATP hydrolysis and ABA transporter AtABCG25 has been shown to bring about the export of
ABA from vascular parenchyma cells where it is synthesized another ABC transporter
AtABCG40 is located in the plasma membranes of ABA responsive guard cells and plays a
roll in ABA import leading to stomatal closure import of ABA into cells is necessary because
the soluble ABA receptors are located in the cytoplasm

According to textbook Two ways of ABA transportation: through phloem and xylem.

- In Phloem (more abundant)

+ radioactive ABA in a leaf -> up to stem
 Down to roots (found in 24h)
- In Xylem from roots to shoots ;
+ well- watered plants ABA concentration > than in water-stressed
+ the magnitude depends on plant species
+ Also, ABA can be transferred in a conjugated form, then hydrolysis in leaves (in suggestion),
but the hydrolases haven’t been found yet
- Among plant cell compartments, ABA distribution follows the anion trap concept: (same in
thextbook)
+ Dissociated anion form gather in alkaline compartments( pH>7?)
+ May be redistributed
+ Specific uptake carriers participate in keeping low apoplastic ABA concentration in unstreseed
plants.
3. Seed dormancy
a. Coat-imposed dormancy : Dormancy imposed on the embryo by the seed coat and

other enclosing tissues

b. embryo dormancy; a dormancy that is intrinsic to the embryo and is not due to any influence of
the seed coat or other surrounding tissues.