The Story of Auxin

www.plantcell.org/cgi/doi/10.1105/tpc.110.tt0410
Auxin research has been continually
advanced by new technologies

Abel, S., and Theologis, A. (2010). Odyssey of auxin. Cold Spring Harb Perspect Biol. doi: 10.1101/cshperspect.a004572
Darwin (1890s) studied phototropism
– movement towards light

Darwin and others studied

coleoptiles –tissues that
protect monocot leaves
during germination

Darwin, C., and Darwin, F. (1881) The power of movement in plants. Appleton and Co., New York.; Photos courtesy of Dr. R.L. Nielsen
 Cutting off or covering the coleoptile
tip interferes with the response

These experiments showed that

the light signal is perceived at
the tip, although the bending
occurs at the base

Untreated Coleoptiles with tips

coleoptile shielded from light or
bends removed do not bend
Darwin concluded that a signal
moves from tip to base

“We must therefore conclude that when

seedlings are freely exposed to a lateral
light some influence is transmitted from
the upper to the lower part, causing the
latter to bend.”
In the 1930s, auxin was purified and
shown to promote growth
Angle of curvature
is proportional to
amount of auxin in
block

Frits Went collected auxin ...and showed that

from shoot tips into agar the material collected
blocks... in the agar blocks
was the growth-
promoting substance.

This bending assay for the growth-

Indole-3-acetic promoting effect of auxin was used
acid, IAA as a basis for its purification.

Redrawn from Went, F.W. (1935) Auxin, the plant growth-hormone. Bot. Rev. 1: 162-182.
Auxin’s root-promoting properties
were also known by the 1930s

Adventitious roots
are initiated from
grape stems treated
with auxin

A more recent experiment: auxin-

treated radish roots initiate lateral
roots at a frequency proportional to
auxin concentration
µM
IAA
Thimann, K.V. (1938). Hormones and the analysis of growth. Plant Physiol. 13: 437-449. Kerk, N.M., Jiang, K., and
Feldman, L.J. (2000). Auxin metabolism in the root apical meristem. Plant Physiol. 122: 925-932.
Auxin’s role in apical dominance
was also known in the 1930s
Auxin suppresses
bud outgrowth
Replace apex
Decapitate with agar
block: without
or with auxin.

Bud Length

No auxin
Auxin

Thimann, K.V., and Skoog, F. (1934). On the inhibition of bud development and other functions of growth substance in Vicia faba. Proceedings of the Royal Society of
London B. 114: 317-339 with permission; Went, F.W. and Thimann, K.V. (1937) Phytohormones. The Macmillan Company, New York.
Different tissues were recognized to
have different sensitivities to auxin

Auxin concentrations
that promote elongation
in stems can be
inhibitory in roots.

Thimann, K.V. (1938). Hormones and the analysis of growth. Plant Physiol. 13: 437-449.
 Polar transport of auxin was
recognized in the 1930s
When the segment is
inverted, it is unable to
A segment cut from a
transport auxin from
coleoptile can move
base to tip.
auxin from tip to base.

The upper agar block was This experiment, carried out by

loaded with auxin, which H.G. Van der Weij revealed
the segment translocated that auxin in the shoot is
to the lower block. translocated from tip to base.
Adapted from Went, F.W. (1935) Auxin, the plant growth-hormone. Bot. Rev. 1: 162-182.
 What classical studies told us

•The chemical structure of auxin

•That auxin promotes root formation and inhibits bud outgrowth
•That auxin moves through the shoot from tip to base
•That different tissues have different sensitivities
•That auxin affects growth in a concentration and tissue-specific way

What we didn’t know was how

auxin promoted growth, moved in
a polar direction, limited bud
outgrowth and stimulated root
formation.
This lecture describes how the tools
of molecular biology, genetics, and
cell biology have continued the
auxin story, and revealed more
about how auxin regulates growth.
 Auxin: a 21st century perspective

• Auxin homeostasis
• Tools in auxin research
• Polar auxin transport
• Perception and signaling
• Auxin action in whole-plant
processes
• Interactions with other signals
 Auxin signaling pathway

Catabolism Conjugation

Synthesis IAA Transport Perception TF activation/ Target Biological

(receptor) inactivation genes Functions

Auxin’s effects depend upon its synthesis, transport,

perception, signaling, and target gene responses. Most of
these functions are controlled by many genes with differing
cell specificities.

Adapted from Kieffer, M., Neve, J., and Kepinski, S. (2010). Defining auxin response contexts in plant development. Curr. Opin. Plant Biol.13: 12-20.
 Biosynthesis and homeostasis
IAA is produced from The IAOx pathway
tryptophan (Trp) via may be restricted
several pathways and Indole to Arabidopsis and
one Trp-independent its close relatives.
pathway (black arrow).
Tryptophan

Indole-3-
pyruvic acid Tryptamine Indole-3- Indole-3-
(IPA) acetamide acetaldoximine
(IAM) (IAOx)
Indole-3-
acetaldehyde

IAA

Adapted from Quittenden, L.J., Davies, N.W., Smith, J.A., Molesworth, P.P., Tivendale, N.D., and Ross, J.J.
(2009). Auxin biosynthesis in pea: Characterization of the tryptamine pathway. Plant Physiol. 151: 1130-1138
..
 Auxin synthesis is developmentally
and environmentally controlled
Indole

Methyl
Jasmonate Tryptophan
Ethylene
Indole-3-
pyruvic acid Tryptamine Indole-3- Indole-3-
acetamide acetaldoximine
Red / Far-red light
ratio
Indole-3-
acetaldehyde
Temperature

Auxin biosynthesis is
influenced by other hormones IAA
and environmental conditions
Adapted from Quittenden, L.J., Davies, N.W., Smith, J.A., Molesworth, P.P., Tivendale, N.D., and Ross, J.J.
(2009). Auxin biosynthesis in pea: Characterization of the tryptamine pathway. Plant Physiol. 151: 1130-1138
..
 Auxin synthesis promotes shade-
avoidance hypocotyl-elongation
response
Auxin synthesis TAA1 Tryptophan
White light
Shade Indole-3-
pyruvic acid

In wild-type plants
Wild-type taa1 but not loss-of-
function taa1
Wild-type taa1 mutants, auxin
An environment enriched with far-red synthesis is
light (which simulates shading by other
plants) promotes hypocotyl elongation, increased in shade
but not in a mutant blocked in an auxin conditions.
synthesis pathway.

Reprinted from Tao, Y., et al. (2008) Rapid synthesis of auxin via a new tryptophan-dependent pathway
is required for shade avoidance in plants. Cell 133: 164–176, with permission from Elsevier.
Auxin homeostasis is also regulated
by conjugation and degradation
Overexpression of an auxin conjugating enzyme
encoded by a GH3 gene reduces auxin levels in
the plant and causes a dwarfed phenotype.
IAA
Rice

GH3 genes
are auxin- Arabidopsis
(GH3 genes)
induced

GH3.13
Wild-type
overexpression
GH3.13
Wild-type
overexpression
Zhang, S.-W., et al., (2009) Altered architecture and enhanced drought tolerance in rice via the cown-regulation of indole-3-acetic acid by TLD1/OsGH3.13 activation. Plant Physiol. 151:1889-1901.
Staswick, P.E., et al., (2005) Characterization of an Arabidopsis enzyme family that conjugates amino acids to indole-3-acetic acid. Plant Cell 17: 616-627.
 Auxin homeostasis - summary

Catabolism Conjugation

Auxin homeostasis is
maintained by its synthesis,
Synthesis IAA Transport catabolism, conjugation and
transport. All of these processes
are tightly regulated.
 Tools in auxin research

•Auxin quantification from tissue or cell extracts

•Expression of auxin-induced genes

•In situ detection by auxin-specific antibodies

•Orientation of PIN auxin-transport proteins

Chemical analysis of auxin levels
from plant extracts
Chemical analysis by gas
chromatography-mass
spectrometry (GC-MS) is an
important way to quantify auxin.

This experiment
shows an auxin
gradient from tip to
base in young
leaves but not
mature leaves.

Edlund, A., Eklof, S., Sundberg, B., Moritz, T., and Sandberg, G. (1995) A microscale technique for gas chromatography-
mass spectrometry measurements of picogram amounts of indole-3-acetic acid in plant tissues.Plant Physiol. 108: 1043-1047.
 Monitoring auxin response by
reporter gene transcription
TGTCTC GUS (or GFP)

AuxRE
Hours of
The DR5 reporter construct is staining
widely used to monitor auxin-
response level.

DR5 consists of seven repeats of

an auxin-response element
(AuxRE) which is a binding site for
auxin-responsive transcription
factors (ARFs).

Auxin-response level increases

with exogenous auxin (NAA)
Ulmasov, T., Murfett, J., Hagen, G., and Guilfoyle, T.J. (1997). Aux/1AA proteins repress expression of reporter genes
containing natural and highly active synthetic auxin response elements. Plant Cell 9: 1963-1971.
DR5 expression reveals high auxin
activity in the root apex and QC

Reprinted by permission from Macmillan Publishers, Ltd. Blilou, I., et al. (2005) The PIN auxin efflux
facilitator network controls growth and patterning in Arabidopsis roots. Nature 433: 39-44.
 Auxin can be detected by anti-auxin
antibodies Primary root
Anti-IAA antibody
Auxin accumulation during lateral root initiation
Anti-IAA antibody
DR5::GUS

Negative
+ auxin
QC control

Reprinted from Benková, E. et al. (2003) Local, efflux-dependent auxin gradients as a common module for
plant organ formation. Cell 115: 591-602, with permission from Elsevier.
 Auxin flow can be inferred from PIN
protein orientation
A shoot apical
meristem imaged over
time, with the
orientation of PIN
proteins indicated by
arrows.

Auxin transport is facilitated by auxin efflux carriers

including PIN proteins, which can have a polar distribution
on the cell membrane. Analysis of the orientation of PIN
proteins provides an approximate picture of intercellular
auxin transport directions.

Reprinted from Heisler, M.G., et al. (2005). Patterns of auxin transport and gene expression during primordium development revealed by live imaging of the
Arabidopsis inflorescence meristem. Curr. Biol. 15: 1899–1911, with permission from Elsevier.
 Polar Auxin Transport

Auxin moves long distances

through the phloem.

Auxin also moves via auxin

transport proteins.

Auxin normally moves from the tip

of the shoot towards the tip of the
root. At the root tip, auxin changes
direction and moves short
distances up the root again
(basipetally).

Reprinted with permission from Macmillan Publishers, Ltd. Robert, H.S., and Friml, J. (2009) Auxin and other signals on the move in plants. Nat. Chem. Biol. 5: 325-332
. Reprinted from Muday, G.K., and DeLong, A. (2001). Polar auxin transport: Controlling where and how much. Trends Plant Sci. 6: 535–542, with permission from
Elsevier.
The Cholodny-Went theory of tropic
curvature
The Cholodny-Went theory of tropic curvature states that perception of
a stimulus (e.g. asymmetric light or gravity) initiates a lateral relocation
of auxin, resulting in differential growth.
 Coleoptiles or shoots move auxin to
the shaded side
IAA accumulates on Increased auxin promotes
the shaded side of
phototropically
cell elongation on the
stimulated Brassica shaded side, causing
oleracea hypocotyls. bending toward the light.

DR5::GUS

Auxin-induced
Cell length
transcription of the DR5
promoter is elevated on
the dark side of this
phototropically-
Auxin
stimulated Arabidopsis
concentration hypocotyl.

Esmon, C.A. et al. (2006) A gradient of auxin and auxin-dependent transcription precedes tropic growth responses. Proc. Natl. Acad. Sci. USA 103: 236–241. Reprinted
by permission from Macmillan Publishers, Ltd: Friml, J., Wisniewska, J., Benkova, E., Mendgen, K., and Palme, K. (2002) Lateral relocation of auxin efflux regulator
PIN3 mediates tropism in Arabidopsis. Nature 415: 806-809.
 Gravitropism is a response to a
change in orientation relative to
gravity Shoots are
negatively
gravitropic – they
grow away from the
center of gravity
vertical

GRAVITY
VECTOR
horizontal
Roots are positively
gravitropic – they
grow towards the
center of gravity

Luschnig, C., Gaxiola, R.A., Grisafi, P., and Fink, G.R. (1998) EIR1, a root-specific protein involved in
auxin transport, is required for gravitropism in Arabidopsis thaliana. Genes Dev.12: 2175-2187.
 Bending is caused by differential
growth on the upper and lower side
length

time
vertical
time

length
Lower side
horizontal

Upper side

time

Parker, K.E., and Briggs, W.R. (1990) Transport of indole-3-acetic acid during gravitropism in intact maize coleoptiles. Plant Physiol. 94: 1763-1769.
 Greater growth is caused by
increased auxin accumulation
Auxin accumulation
time
vertical

horizontal

horizontal vertical

Parker, K.E., and Briggs, W.R. (1990) Transport of indole-3-acetic acid during gravitropism in intact maize coleoptiles. Plant Physiol. 94: 1763-1769.
Auxin accumulates on the lower side
of roots, but they bend down

DR5::GUS expression in vertically oriented root

Increasing the amount
and root 6 hours after turning horizontal.
of auxin on the lower
side of the root creates
supraoptimal levels,
inhibiting cell elongation
and causing the root to
bend downwards.
Rashotte, A.M., DeLong, A., and Gloria K. Muday, G.K. (2001) Genetic and chemical reductions in protein phosphatase activity alter auxin transport, gravity response,
and lateral root growth. Plant Cell 13: 1683-1697; Thimann, K.V. (1938). Hormones and the analysis of growth. Plant Physiol. 13: 437-449; Luschnig, C., Gaxiola, R.A.,
Grisafi, P., and Fink, G.R. (1998) EIR1, a root-specific protein involved in auxin transport, is required for gravitropism in Arabidopsis thaliana. Genes Dev.12: 2175-
2187.
Many Arabidopsis mutants have
been identified with abnormal
gravitropic responses
eir1 = pin2

WT

Analysis
Analysisofofthese
thesemutants
mutantshas
has
identified several types of auxin
identified several types of auxin
transport
transportproteins,
proteins,asaswell
wellas
as
WT pin2 proteins that regulate their activity
proteins that regulate their activity
and
anddistribution.
distribution.
Reprinted with permission from Macmillan Publishers, Ltd. Muller, A., et al. (1998) AtPIN2 defines a locus of Arabidopsis for root gravitropism control. EMBO J 17 :
6903-6911; Luschnig, C., Gaxiola, R.A., Grisafi, P., and Fink, G.R. (1998) EIR1, a root-specific protein involved in auxin transport, is required for gravitropism in
Arabidopsis thaliana. Genes Dev.12: 2175-2187. Rahman, A., Ahamed, A., Amakawa, T., Goto, N., and Tsurumi, S. (2001). Chromosaponin I specifically interacts
with AUX1 protein in regulating the gravitropic response of Arabidopsis roots. Plant Physiol. 125: 990-1000.
 Auxin transport – chemiosmotic
model
Cell wall
pH 5.5
Indole-3-acetic acid is a charged
anion (IAA-) in the cytoplasm (pH 7). A proton ATPase Cytoplasm
maintains the pH 7
H+
differential pH
In the more acidic cell wall (pH 5.5) gradient IAA-
some is uncharged (IAAH). The
uncharged form crosses the plasma Cell-to-cell
polar
membrane into the cell where it is IAAH IAA- + H+ auxin
transport
deprotonated and unable to exit
other than through specific
IAAH
transporters.

IAA- + H+

Redrawn from Robert, H.S., and Friml, J. (2009) Auxin and other signals on the move in plants. Nat. Chem. Biol. 5: 325-332.
Auxin moves through efflux and
influx carrier proteins
ABCB The PIN family of
proteins contributes to
directional movement
ABCB
of auxin out of the cell.

Cell-to-cell
The AUX1/LAX polar
auxin
influx carriers transport

contribute to
The ABCB
movement of
transporters
IAAH into the ABCB contribute to auxin
cytoplasm.
transport in a diverse
ways. (These were formerly
referred to as MDR or PGP
proteins).
Reprinted with permission from Macmillan Publishers, Ltd. Robert, H.S., and Friml, J. (2009) Auxin and other signals on the move in plants. Nat. Chem. Biol. 5: 325-332.
The PIN proteins are named for the
pin-formed mutant
pin-formed, which has a
mutation in the PIN1 gene,
makes some abnormal leaves
and then a bare inflorescence.

Galweiler, L., Guan, C., Muller, A., Wisman, E., Mendgen, K., Yephremov, A., and Palme, K. (1998). Regulation of polar auxin transport by AtPIN1 in Arabidopsis
vascular tissue. Science 282: 2226 – 2230, reprinted with permission from AAAS.
 The distribution of PIN proteins
contributes to the auxin gradients

Křeček , P., Skůpa , P., Libus, J., Naramoto, S., Tejos, R., Friml J., and Zažímalová, E. (2009) The PIN-
FORMED (PIN) protein family of auxin transporters. Genome Biology 10: 249.
PIN protein distributions contribute
to auxin-mediated patterns

Amongst other things,

they specify the
location of auxin
maxima necessary for
embryonic patterning.

Reproduced with permission from Petrášek, J., and Friml, J. (2009) Auxin transport routes in plant development. Development 136: 2675-2688.
Auxin transport - summary

Regulated movement of auxin is critical

for phototropism, gravitropism,
embryonic pattern formation and
organogenesis.
Auxin transport proteins have been
identified. Factors that control their
position and activity are under
investigation.

Arabidopsis has 6 AUX1/LAX

proteins, 21 ABCB proteins
and 8 PIN proteins - we don’t
yet understand all the
complexity of auxin transport!
 Perception and signaling

Catabolism Conjugation

Synthesis IAA Transport Perception TF activation/ Target Biological

(receptor) inactivation genes Functions
Auxin’s effects are mediated
through at least two types of
receptor

Perception ABP1
(receptor)

IAA

Perception
(receptor) SCFTIR1
AUXIN-BINDING Protein1 is necessary
for cell division and expansion
EtOH- Loss-of-function of ABP1 is
Wild-type
induced embryo-lethal.
control
ABP1-
antisense Plants were produced that
carry an inducible antisense
construct that silences the
ABP1 gene. They were
grown for 12 days in non-
inductive conditions.

At day 12, ABP1 was

silenced and plant growth
was severely restricted.

Braun, N., et al. (2008). Conditional repression of AUXIN BINDING PROTEIN1

reveals that it coordinates cell division and cell expansion during postembryonic shoot
development in Arabidopsis and tobacco. Plant Cell 20: 2746-2762.
 ABP1 mediates rapid auxin
responses at the plasma membrane

AUXIN

Perception of auxin by
ABP1 at the outer
face of the plasma
membrane initiates
signals that lead to
proton-pump
activation, wall
acidification and wall
loosening.

Reprinted from Tromas, A., Paponov, I., and Perrot-Rechenmann, C. (2010). AUXIN BINDING PROTEIN 1: functional and evolutionary aspects
Trends Plant Sci. 15: 436–446 with permission from Elsevier.
 From ABP1 to TIR1

The events downstream of

Perception ABP1 Auxin-Binding Protein1 aren’t
(receptor) well understood yet.

IAA

Now we’ll look at the pathway

Perception
SCFTIR1 downstream of auxin
(receptor) perception by SCFTIR1.
 The auxin-receptor TIR1 is a
component of the SCF ubiquitin
ligase complex SCFTIR1

F-box TIR1
auxin
receptor F-box TIR1
protein
SKP1

SCF complex

CUL1
SCF ubiquitin ligase complex
(Named for SKP1, CUL1 and F-box
proteins)
 Ubiquitin ligase complexes
ubiquitinate target proteins

Ubiquitin The F-box protein

recognizes and binds
Target to the target protein.
Target The complex then
transfers ubiquitin
SKP1 proteins to the target.

CUL1

Ubiquitin by Rogerdodd
Auxin specifically binds to the
SCFTIR1 complex

Radioactive IAA was bound to

Benzoic acid the SCFTIR1 complex in the
Tryptophan presence of various
2,4-D competitors. The specificity of
NAA the interaction is shown by its
IAA competition by various auxins
(IAA, NAA, 2,4-D) but not
related compounds (benzoic
acid, tryptophan).

Reprinted by permission from Macmillan Publishers, Ltd. Dharmasiri, N., Dharmasiri, S., and Estelle, M.
(2005) The F-box protein TIR1 is an auxin receptor. Nature 435: 441-445.
 TIR1 requires auxin to bind its
targets
Auxin has been
said to act like
glue to hold
together TIR1
and its targets.
Auxin Target
 TIR1 requires auxin to bind its
targets

Auxin Target

What are the targets

of SCFTIR1-mediated
proteolysis?

I II III IV

Aux/IAA proteins, short-lived, nuclear proteins

that are repressors of auxin-responses.
Aux/IAA genes are induced by auxin

In the 1980’s, the new

techniques of molecular biology GmAux22
allowed scientists to isolate
genes that were strongly and
rapidly induced by auxin. GmAux28

One family of these auxin-

induced genes is known as the
Aux/IAA genes.
Control 15 30 60 90
Auxin treatment (mins)

Auxin
depleted

Walker, J.C. and Key, J.L. (1982) Isolation of cloned cDNAs to auxin-responsive poly(A)+RNAs
of elongating soybean hypocotyl. PNAS 79: 7185-7189.
 Aux/IAA proteins are very short-
lived nuclear proteins

Many of the
Aux/IAA proteins
have a very
short half-life,
suggesting a
regulatory
function. A fusion
protein with
GUS shows
nuclear
accumulation

PS-IAA6

Abel, S., Oeller, P.W., and Theologis, A. (1994). Early auxin-induced genes encode short-lived nuclear proteins. PNAS 91 : 326-330.
Aux-IAA proteins are destabilized in
the presence of auxin
Auxin increases the Transgenic plants were
Luciferase
proteolytic rate of LUC (LUC) control generated carrying the
IAA1∷LUC. LUC coding region or
LUC fused to LUC fused to an
LUC Aux/IAA1 protein
Aux/IAA1 protein.
I II III IV

In the LUC plants, auxin did

not affect protein stability, but
2 h auxin Measure LUC
activity. when LUC was fused to
Aux/IAA1 50% of the protein
Mock was degraded after 2 hours of
Measure LUC
treatment
activity. auxin treatment.

Zenser, N., Ellsmore, A., Leasure, C., and Callis, J. (2001). Auxin modulates the degradation rate of Aux/IAA proteins. Proc. Nat.l Acad. Sci. USA 98: 11795-
11800.Copyright National Academy of Sciences, USA.
 The conserved “degron” in domain II
confers auxin-instability
A short amino acid
LUC
sequence called a
degron is necessary
and sufficient for
auxin-induced protein LUC
I II III IV
instability.
QVVGWPLVRSYRK
Mutant degron
Changing a single
amino acid (red) in
this conserved LUC
I II III IV
sequence makes the
protein much more QVVGWPPVRSYRK
stable.
Wild-type degron

Ramos, J., Zenser, N., Leyser, O., and Callis, J. (2001) Rapid degradation of auxin/indoleacetic acid proteins requires conserved amino
acids of domain II and is proteasome dependent. Plant Cell 13: 2349 - 2360.
 Aux/IAA proteins - summary

Aux/IAA proteins are

repressors of auxin
signaling that are rapidly
degraded in the presence
Auxin of auxin

INPUT OUTPUT

How do Aux/IAA
proteins repress auxin
signaling?
Aux/IAA proteins repress the activity
of Auxin Response Factors (ARFs)

ARF and Aux/IAA proteins have ARF

DNA- Activation / III IV
sequence similarity in their C- binding repression
terminal regions through which domain domain
they can homodimerize.
Aux/IAA
I II III IV

ARF homodimer

ARF Aux/IAA heterodimer

 Some ARFs are transcriptional
activators and some repressors
The Arabidopsis Confirmed
ARF protein family activators

ARF 2-4, 9 are

confirmed
repressors

We don’t know much about

the ARF repressors yet.

Reprinted from Guilfoyle, T., and Hagen, G. (2007). Auxin response factors. Curr. Opin. Plant Biol. 10: 453 – 460 with permission from Elsevier.
 Model for auxin-regulated
transcriptional control
Low auxin level
Some ARFs are
complexed with
Aux/IAA repressors,
interfering with their
action

High auxin level

Aux/IAAs are
degraded by SCFTIR1-
mediated proteolysis,
relieving repression of
ARFs to activate or
repress gene
expression.
monopteros (arf5/mp) and bodenlos
(iaa12/bdl) mutants look similar

WT mp

Lau, S., Jürgens, G., and De Smet, I. (2008) The evolving complexity of
the auxin pathway. Plant Cell 20:1738-1746.
These mutants look like each other because
the effects of their mutations are similar

No ARF5! Stable IAA12!

Lau, S., Jürgens, G., and De Smet, I. (2008) The evolving complexity of
the auxin pathway. Plant Cell 20:1738-1746.
 Perhaps the diversity of auxin responses
is correlated with the diversity in
signaling proteins
Arabidopsis has 29 Arabidopsis has 23
Aux/IAA proteins ARF proteins

Auxin OUTPUT

INPUT
The number of potential
interactions between ARFs
and Aux/IAA proteins is
quite large, and has the
potential to precisely
TIR1 is related to control auxin-induced gene
five other auxin- expression in a vast
receptor proteins, number of combinations.
AFB1 – AFB5.
 Auxin action in whole-plant
processes
Auxin in Promote lateral organ
Responses to pathogens
initiation at the shoot
action! apical meristem Inhibit
branching in
Light responses the shoot
Cell elongation Integrate growth
signaling pathways
Control patterning
and vascular
development
Maintain stem- Promote
cell fate at the branching
Integrate growth root apical in the root
signaling pathways meristem Response to nutrient
Responses to symbionts – distribution and
nodule formation abundance
Reprinted by permission from Macmillan Publishers, Ltd: NATURE Wolters, H., and Jürgens, G. (2009). Survival of the flexible:
Hormonal growth control and adaptation in plant development. Nat. Rev. Genet. 10: 305–317. Copyright 2009.
 Auxin signalling crosstalk

Each step of auxin signaling is extensively regulated by auxin

activity and also environmental influences such as light, gravity,
nutrient availability and stress, but also other hormones.

Reprinted from Kieffer, M., Neve, J., and Kepinski, S. (2010). Defining auxin response contexts in plant development. Current Opinion in
Plant Biology 13: 12-20. with permission from Elsevier.
 Auxin and cytokinin act
antagonistically
Auxin and cytokinin (CK) act Increasing [Auxin]
antagonistically – auxin tends to
promote roots and CK to promote
shoots. Increasing
[Cytokinin]

Auxin suppresses CK synthesis, Roots

and CK induces expression of

some Aux/IAA repressors and
down-regulates PIN gene
expression.

Shoots
Auxin Cytokinin

Skoog F, Miller CO (1957) Chemical regulation of growth and organ formation in plant tissue cultured in vitro. Symp Soc Exp Biol XI: 118–131; Figure from
Amasino, R. (2005) 1955: Kinetin Arrives. The 50th Anniversary of a newplant hormone. Plant Physiol. 138: 1177 – 1184.
 Ethylene and auxin tend to interact
synergistically
Ethylene promotes Air Ethylene At the same time,
auxin synthesis auxin stimulates
ethylene production

Expression of an auxin-
De novo auxin synthesis was
measured in plants treated with
responsive promoter is
D2O. The rate of auxin synthesis increased with exogenous
increased in the presence of ethylene Auxin Ethylene
ACC, an ethylene precursor.

Swarup, R., et al. (2007). Ethylene upregulates auxin biosynthesis in Arabidopsis seedlings to enhance inhibition of root cell elongation. Plant Cell 19: 2186-
2196. Kang, B.G., Newcomb, W., and Burg , S.P. (1971) Mechanism of auxin-induced ethylene production. Plant Physiol. 47: 504-509.
 Summary

In the past 30 years we have identified many of the molecular characters

in the auxin story, and have a pretty good idea of its major themes, but
the story is far from complete.

Catabolism Conjugation

Synthesis IAA Transport Perception TF activation/ Target Biological

(receptor) inactivation genes Functions

Adapted from Kieffer, M., Neve, J., and Kepinski, S. (2010). Defining auxin response contexts in plant development. Current Opinion in Plant Biology 13: 12-20.
 Ongoing investigations

Where does auxin

elimination fit in? What do all those transport
proteins do? What controls their What are the target
activity and distribution?? genes, and what do
Catabolism Conjugation they do?

Synthesis IAA Transport Perception TF activation/ Target Biological

(receptor) inactivation genes Functions

What regulates What are the other Why so many How do all these
auxin synthesis? TIR1 like proteins ARFs and pieces fit together to
doing, and what Aux/IAAs? make a functioning
does ABP1 do? plant????
Adapted from Kieffer, M., Neve, J., and Kepinski, S. (2010). Defining auxin response contexts in plant development. Current Opinion in Plant Biology 13: 12-20.