The plant hormones I

HBC 206 Plant Biochemistry

Dr Fiona C Robertson
Department of Biochemistry
University of Zimbabwe
 Plant hormones
• Organic compounds produced in low
concentrations
• Produced in one part of plant (i.e. source)
• Transported to another part of plant (i.e. target)
• (in some instances the hormone can act as an
autocrine signal, binding to receptors on the
same cell as produced it
• Cause physiological or developmental responses
(stimulatory or inhibitory)
• Also called plant growth regulators
/phytohormones
 Plant hormones

Cytokinin Gibberellic Acid

Auxin
Abscisic Acid

Ethylene

Strigolactone
Brassinosteroid Salicylic Jasmonic Acid
Acid
 Plant hormones regulate all stages of
the plant life cycle
Fruit Germination
ripening
Seed
Embryogenesis dormancy
Growth and
branching

Fertilization and
fruit formation Flower
development
Most hormones affect most stages of
the plant life cycle
We will look at each hormone
and discuss some of its roles

But these are only examples;

most hormones affect most
processes in one way or
another
 Lecture outline
1. How hormones work
2. Hormonal control of vegetative development
Auxin
Cytokinin
Strigolactones
Gibberellins
Brassinosteroids
3. Hormonal control of reproduction
Ethylene
Abscisic Acid

4. Hormonal responses to stress

Salicylic Acid
Jasmonic Acid
 Hormones: Synthesis, transport,
perception, signaling and responses

Production of active
hormone Downstream
effects
Transport

H Signal
transduction Downstream
Binding to effects
receptor
 Synthesis
Conjugation Many biochemical
H
De-conjugation
pathways contribute
H
to active hormone
Synthesis Production Break- accumulation. These
of active down
pathways are tightly
hormone
regulated.
Conjugation can -
• temporarily store a hormone in an inert form
• lead to catabolic breakdown
• Lead to production of the active hormone
 Transport and perception
Hormones can move
• through the xylem or phloem
• across cellular membranes
• through regulated transport
proteins
Transport
Many hormone receptors have
H now been identified, they can be
• membrane bound
Binding to
receptor
• cytoplasmic
• nuclear localized
 Signal transduction
Hormonal signals are transduced by
downstream factors

Signal
transduction
 Responses
Downstream effects
usually involve Transcription
• changes in gene
transcription
• Other effects such as
ion channel
activation
Ion channel
regulation
 Hormones: Synthesis, transport,
perception, signaling and responses
Conjugation
H
De-conjugation Downstream effects
H
Synthesis Production of active Break-down
hormone
Transcription
Transport

Downstream effects
H Signal Ion channel
Binding to transduction regulation
receptor
 Hormones that affect vegetative growth:
elongation, branching and organ formation
Elongation in the
shoot and root of a
germinating soybean
Organogenesis

Germinated
seedling Growth by
branching

Growth by
elongation
Photo courtesy of Shawn Conley
 Auxin
• Growth
• Phototropism and gravitropism
• Branching
• Embryonic patterning
• Stem cell maintenance
• Organ initiation (roots)

Indole-3-acetic acid (IAA),

the most abundant natural
auxin
 Phototropism
Darwin (1890s)

Darwin, C., and Darwin, F. (1881) The power of movement in plants. Appleton and Co., New York.; Photos courtesy of Dr. R.L. Nielsen
 Auxin controls phototropism

Site of signal
Charles Darwin perception
studied the way
seedlings grow
Site of
towards light response

Coleoptile drawing from Darwin, C., and Darwin, F. (1881) The power of movement in plants. Available online.
 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
Untreated Coleoptiles with the base.
coleoptile tips shielded from
bends light or 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.”

Charles Darwin image courtesy of Patche99z

Auxin was purified and shown to
promote growth
 Cell expansion
Auxin acts by loosening the
cellulose microfibrils of cell
walls

Causes cell expansion

 Coleoptiles or shoots transport auxin
to the shaded side
IAA accumulates on
the shaded side of
phototropically
stimulated Brassica
oleracea Increased auxin
hypocotyls. promotes cell
elongation on the
shaded side, causing
bending toward the
Cell length light.

Auxin
concentration

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.
DR5::GUS – an Auxin reporter gene
An inhibitor of auxin transport
prevents phototropism
Auxin’s root-promoting properties
were also known by the 1930s
A more recent experiment in which radish roots were
dipped into auxin and initiated lateral roots at a
frequency proportional to auxin concentration.

Adventitious roots are

initiated from grape
stems treated with
auxin

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 promotes apical dominance
Auxin’s role in apical dominance was
proven with the isolation of auxin
Auxin suppresses
bud outgrowth
Replace apex
Decapitate with agar block:
without or with
auxin.

No auxin
Bud Length

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.
 Auxin controls many aspects of plant
development
Promote and specify Inhibit
lateral organ initiation at branching in
the shoot apical the shoot
meristem

Control patterning
and vascular
development
Promote
Maintain stem cell fate branching in
at the root apical the root
meristem
Wolters, H., and Jürgens, G. (2009). Survival of the flexible: Hormonal growth control and adaptation in plant development. Nat. Rev. Genet. 10: 305–317.
 Auxin signalling pathway
• 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

Catabolism Conjugation

Synthesis IAA Transport Perception TF activation/ Target Biological

(receptor) inactivation genes Functions

Perception Signal transduction Response

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.
Cytokinins
• promote cell divisions in
the shoot trans-zeatin (tZ)

• delay leaf senescence

• regulate nutrient allocation
• promote root nodule
development
Isopentenyl-
• contribute to environmental adenine (iP)

signaling and pathogen

responses
• regulate auxin action and
distribution
 The discovery of cytokinin
Control With kinetin
Cytokinin was
discovered when
researchers were
trying to identify
compounds that
increase the
growth of plant
cells in culture.

In the 1950s, Folke Skoog’s

research group identified a
synthetic cytokinin, kinetin.
Amasino, R. (2005) 1955: Kinetin Arrives. The 50th Anniversary of a newplant hormone. Plant Physiol. 138: 1177 – 1184.
 Cytokinin and auxin control
organogenesis in tissue culture

+ CK

+ auxin + auxin
and CK

Tobacco leaf disc are placed into

sterile culture dishes on medium
containing various hormones. TIME

Images courtesy of Richard Amasino.

 Cytokinins delay leaf senescence
Senescence
(programmed
death of the
leaves) is
dramatically
reduced in
kinetin-treated
leaves or leaves of
plants producing
higher levels of Plants
cytokinins. overproducing
Control plants
cytokinin
Smart, C.M., Scofield, S.R., Bevan, M.W., and Dyer, T.A. (1991). Delayed leaf senescence in tobacco plants
transformed with tmr, a gene for cytokinin production in Agrobacterium. Plant Cell 3: 647-656.
 Cytokinins are a family of related
adenine-like compounds

Isopentenyl trans-zeatin dihydrozeatin cis-zeatin

adenine

Hirose, N., Takei, K., Kuroha, T., Kamada-Nobusada, T., Hayashi, H., and Sakakibara, H. (2008). Regulation of cytokinin
biosynthesis, compartmentalization and translocation. J. Exp. Bot. 59: 75–83.
 Cytokinins act antagonistically to
auxins Promote
CK Auxin
branching
Promote stem Promote in the shoot
cell fate at the lateral organ
shoot apical initiation at the Inhibit
meristem shoot apical branching
meristem in the shoot

Inhibit
Promote Maintain stem Promote branching
differentiation cell fate at the branching in the root
at the root root apical in the root
apical meristem
meristem

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.
Gibberellins
• Growth
• Promotion of elongation
of cells

Gibberellic Acid (GA4)

 Gibberellins were identified as
fungal compounds that promote stem
elongation
Bakanae disease Bakanae disease means
(Gibberella fujikuroi) “foolish seedling”, because
infected plants elongate too
rapidly, and are unable to
Uninfected support themselves; they
plant are also male sterile.

Infected,
hyper-
elongated
plants

A Gibberellin (GA3)
Photo source: Nigel Cattlin, Visuals Unlimited, Inc.
 Catabolism Conjugation

Synthesis IAA Transport Perception TF activation/ Target Biological

GA (receptor) inactivation genes Functions

Synthesis Perception Signal transduction Response

 GA has been described as “an
inhibitor of an inhibitor”
A car doesn’t roll down a
hill when the brake is on
to inhibit it. Releasing the
brake “inhibits the
inhibitor”, allowing
another force (gravity) to
move it.
GA doesn’t promote
growth on its own, but
instead inhibits growth
inhibitors (i.e. releases
the brakes).
Plants that have a mutation in their GA
synthesis genes are severely dwarfed
Wild-type and dwarf
mutant rice

oscps-1 osks-1

WT oscps-1 osks-1
Sakamoto, T., et al. (2004). An overview of gibberellin metabolism enzyme genes and their related mutants in rice. Plant Physiol. 134: 1642-1653.
 Catabolism Conjugation

Synthesis IAA Transport Perception TF activation/ Target Biological

GA (receptor) inactivation genes Functions

Synthesis Perception Signal transduction Response

 Perception and signaling

DELLAs GA

GA inactivates DELLA inhibitors

Without the inhibitory DELLAs, the

plant can respond to growth-
promoting activities.
DELLA-encoding genes have great had
great agricultural impact

The green-revolution
gene reduced height1
from wheat encodes a
mutant DELLA.

Wild-type
wheat
Reduced-height1
wheat

Reprinted by permission of Macmillan Publishers, Ltd. Peng, J., et al. (1999) 'Green revolution' genes encode mutant gibberellin response modulators. Nature 400: 256-261.
The manipulation of GA levels is
tremendously important for
agriculture

One of the most

significant
accomplishments of
20th century science
was the development
of semi-dwarf grain
varieties which are
deficient in GA
synthesis or response.
Distinguished plant breeder and Nobel Laureate
Norman Borlaug 1914-2009
Photos courtesy of S. Harrison, LSU Ag center and The World Food Prize.
 Response

Catabolism Conjugation

Synthesis IAA Transport Perception TF activation/ Target Biological

GA (receptor) inactivation genes Functions

Synthesis Perception Signal transduction Response

 GA promotes growth through cell
expansion and division
GA induces
expression of cell-
cycle regulatory
proteins called
cyclins. Cell
expansion
Cell
division

GA promotes elongation by
Cell
cell wall loosening and
expansion
stabilizing the orientation of
cortical microtubules, which
help direct growth.
GA promotes growth through cortical
microtubule orientation
Circumferential or Uniconazole
hoop-like cortical (GA synthesis Uniconazole
microtubules help Control inhibitor) + GA
promote
unidirectional,
elongation growth.

Inhibition of GA biosynthesis
disrupts cortical microtubule
arrangement while GA restores
arrangement.
Inada, S. and Shimmen, T. (2000). Regulation of elongation growth by gibberellin in root segments of Lemna
minor. Plant Cell Physiol 41: 932-929, by permission of the Japanese Society of Plant Physiologists.
 Summary – hormonal control of
vegetative growth
Plant hormones have diverse effects on
plant growth
Auxin, gibberellic acid and
brassinosteroids contribute to
elongation growth
Auxin, cytokinin and strigolactones
control branching patterns
Growth and branching profoundly affect
crop yields