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AUXINS

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Auxin: The Growth Hormone

Auxin Research

The auxin concept A historical perspective

curvature

no curvature

From experiments on
coleoptile phototropism,
Darwin concluded that a
growth stimulus is
produced in coleoptile
tip and transmitted to
growth zone.

no curvature

Growth stimulus passes through gelatin but not through


water-impermeable barriers such as mica.

Coleoptile = sheath/protective organ of the youngest leaves in grasses


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Auxin Research
Provided evidence that the
growth promoting stimulus
produced in the tip was
chemical nature.

The growth-promoting substance can diffuse into a gelatin block.

Auxin stimulates the elongation of oat coleoptile sections

primary leaves

coleoptile

Incubation in water for 18h

Incubation in auxin for 18h

Auxin from the greek word auxien, meaning to increase or to grow.


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Structures of auxins
Indole

Structures of three natural auxins

Peas

All plants

Auxin biosynthesis
Synthesis in meristems, young
leaves, developing fruits and
seeds

Mustard and corn


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Synthetic auxins

Often used as herbicides

Auxins in Biological Samples


can be Quantified
Bioassay
Mass

spectrometry
Enzyme-linked immunosorbent assay
(ELISA)

Biosynthesis of IAA

Biosynthesis of IAA from Tryptophan

IPA pathway
The most common of the
tryptophan-dependent pathways

Tryptophan (Trp)
Trp
transaminase

Indole-3-pyruvic acid (IPA)


IPA
decarboxylase

Indole-3-acetaldehyde (IAId)
IAId
dehydrogenase

Indole-3-acetic acid (IAA)

TAM pathway

Tryptophan (Trp)
Trp
decarboxylase

1. Similar to the IPA pathway


2. Species that do not utilize
the IPA pathway possess
the TAM pathway
3. In tomato, there is evidence
for both the IPA and the
TAM pathways

Tryptamine (TAM)
Amine
oxidase

Indole-3-acetaldehyde (IAId)
IAId
dehydrogenase

Indole-3-acetic acid (IAA)

IAN pathway
The IAN pathway may be
important in only three plant
families:

Tryptophan (Trp)
Trp
Monooxygenase

Indole-3-acetaldoxime

1.Brassicaceae (mustard family)


2.Poaceae (grass family)
3.Musaceae (banana family)

Indole-3-acetonitrile (IAN)
Nitrilase

Indole-3-acetic acid (IAA)

IAM pathway
Tryptophan (Trp)
Used by various pathogenic
bacteria, such as Pseudomonas
savastanoi and Agrobacterium

Trp
Monooxygenase

Involves the two enzymes


tryptophan monooxygenase and IAM
hydrolase

Indole-3-acetamide
(IAM)
IAM
hydrolase

Indole-3-acetic acid (IAA)

IAA is also synthesized from Indole or from Indole-3-Glycerol Phosphate


Evidence for the Tryptophan
Independent Biosynthesis of IAA

Figure 19.4.A Tryptophanindependent pathways of IAA


biosynthesis in plants. The tryptophan
biosynthetic pathway is shown on the
left, and mutants discussed in the text
are indicated in parentheses. The
suggested branch points from the
tryptophan (Trp) biosynthetic
pathway are at indole-3-glycerol
phosphate or at indole. IAN and IPA
are two possible intermediates. The
conversion of tryptophan to IPA is
hypothetical. (After Bartel 1997.)

IAA is also synthesized from Indole or from


Indole-3-Glycerol Phosphate
Orange pericarp mutant (orp)

The orange pericarp (orp) mutant of maize is missing both subunits of tryptophan
synthase. As a result, the pericarps surrounding each kernel accumulate
glycosides of anthranilic acid and indole. The orange color is due to excess
indole. (Courtesy of Jerry D. Cohen.)

Multiple Pathways to Degraded IAA

Decarboxylation pathway

peroxidase (oxidation of IAA)


plays a relatively minor role

Nondecarboxylation pathway

The two nondecarboxylation routes


of IAA oxidative degradation, A and
B, are the most common metabolic
pathways.

Auxin Transport

Polar transport

In phloem

chemiosmotic model
auxin

influx

auxin

efflux

Lack of effect of gravity on


basipetal auxin transport
Roots grow from the basal ends
of these bamboo sections, even
when they are inverted.
The roots form at the basal end
because polar auxin transport in
the shoot is independent of
gravity

Chemiosmotic model

Auxin influx

Passive diffusion of the


protonated (IAAH) form
across the phospholipid
bilayer
IAA- permease (through the
membrane enzyme, AUX1),
the initiative passed.

Auxin efflux

In the cytosol (neutral


pH) forming IAA-
auxin efflux carriers (PIN
proteins), retrieving IAAout of the cell

PHYSIOLOGICAL EFFECTS OF AUXIN


Cell

elongation
Phototropism
Gravitropism

Cell elongation

Auxin is synthesized in the shoot apex and the steady


supply of auxin arriving at the subapical region of the stem
or coleoptile is required for the continued elongation of
these cells.
Generally optimal auxin concentration for elongation
growth is typically 10-6 to 10-5 M
Stem elongation is inhibited by high concentrations of
auxin, because the auxin-induced ethylene biosynthesis.
Auxin concentration is low To promote root cell elongation,
but root growth is strongly inhibited by auxin concentrations
that promote elongation in stems and coleoptiles.

Cell extention

From the results of studies on the influence of auxin on cell growth,


showed that there are indications that auxin can:

In conjunction with cell permeability, the presence of auxin


increases the diffusion of water influx into the cell. This is
supported by the Clelands opinion that auxin support the increased
permeability of the entry of water into the cell.

Phototropism
Is the bending of plants
toward light.
It is caused by distribution of
auxin are not equal, the plant
parts that get light have
lower concentration of auxin
than the plant parts in the
dark.

Gravitropism
Is roots to grow
downward into the
soil and shoots to
grow upward
away from soil

Microsurgery experiments demonstrating that the root cap


produces an inhibitor (auxin!) that regulates root gravitropism

Removal of
root cap;
slight
elongation
growth

Horizontal control root


with root cap showing
normal gravitropic
bending

Removal of half of the


cap causes root to bend
toward side with half cap
Auxin is redistributed
laterally in the root cap

Removal of cap from horizontal


root abolishes response to
gravity
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DEVELOPMENTAL EFFECTS OF AUXIN

Auxin Regulates Apical Dominance


Auxin Promotes the Formation of Lateral and
Adventitious Roots
Auxin Delays the Onset of Leaf Abscission
Auxin Transport Regulates Floral Bud Development
Auxin Promotes Fruit Development
Auxin Induces Vascular Differentiation

Auxin regulates apical dominance

Apical dominance = the growing apical bud inhibits the


growth of lateral (axillary) buds
Phaseolus vulgaris (bean)
Removal of apical/
terminal bud
induces growth of
axillary buds

Axillary buds are


suppressed
because of apical
dominance

Applying IAA
to cut surface
prevents
outgrowth of
axillary buds

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Auxin transport regulates floral bud development


Floral meristem depends on auxin
being transported to it from subapical
tissue
In absence of an auxin carrier protein,
meristem is starved for auxin,
disrupting normal leaf and floral
development

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Auxin promotes fruit development

Auxin is produced in pollen, endosperm, and embryo of developing seeds.


Growth of fruit is regulated by auxin produced by seeds, the achenes.
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Auxin Induces Vascular Differentiation

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