Plant Hormones
Sean R Cutler, Department of Botany and Plant Sciences, Center for Plant Cell
Biology, & Institute for Integrative Genome Biology, University of California, Riverside,
California, USA
David C Nelson, Department of Botany and Plant Sciences, Center for Plant Cell
Biology, & Institute for Integrative Genome Biology, University of California, Riverside,
California, USA
Plant hormones coordinate physiology and devel-
opment between organ systems, which can often
be separated by large distances. In this article,
we focus on the set of nine small molecule plant
hormone families that are present throughout
land plants: auxins, gibberellins, jasmonates, sal-
icylates, brassinolides, strigolactones, cytokinins,
abscisic acid, and ethylene. We cover the basic
aspects of their physiological roles, biosynthesis,
signal transduction and agricultural relevance.
We note commonalities in the molecular mech-
anisms of plant hormone perception and signal
transduction; for example, hormone-stabilised
protein–protein interactions, that are often linked
directly to ubiquitylation of downstream effector
proteins are prevalent. Plant hormone signalling
pathways preferentially exploit soluble receptors
over transmembrane receptors; brassinosteroids
are the only hormone family known to act through
a classical plasma membrane-anchored receptor
kinase, a modality common in animal signal trans-
duction pathways. Plant hormones provide a
complementary set of molecular solutions to the
biological problem of communicating information
over long distances in multicellular organisms.
Introduction
Plant form is established by highly controlled patterns of cell
division, cell elongation and differentiation, which continuously
respond to abiotic and biotic environmental cues. Because
plants have sessile lifestyles, this dynamic adaptation to the
environment is a necessity. For example, plant shoots typically
grow towards light (phototropism) to maximise photosynthesis,
eLS subject area: Plant Science
How to cite:
Cutler, Sean R and Nelson, David C (March 2017) Plant
Hormones. In: eLS. John Wiley & Sons, Ltd: Chichester.
DOI: 10.1002/9780470015902.a0002091.pub2
eLS © 2017, John Wiley & Sons, Ltd. www.els.net
Advanced article
Article Contents
• Introduction
• Discovery and Functions of Plant Hormones
• Hormone Perception
• Hormone Crosstalk
• Uses of Synthetic Plant Hormones
• Conclusion
Online posting date: 20th March 2017
and stop growing when resources such as water or nitrogen
become limiting. How do plants coordinate their growth with
environmental signals? The answers to this simple question
could occupy an entire textbook, but conceptually the processes
involved can be broken down into two major components: (1) the
perception of external or internal stimuli, followed by (2) a physi-
ological response, such as altered cell expansion or transcription.
In the case of phototropism, light is perceived by special proteins
called photoreceptors that instruct plant cells which direction has
the greatest intensity of photons. After this initial sensory event,
the rates of cell growth are controlled by the local action of a
plant hormone called auxin (Briggs, 2014). This article focuses
on the action of plant hormones, but it is important to remember
that hormones function as part of a much larger set of integrated
systems for controlling cellular and organismal physiology.
What makes a molecule a hormone? The term hormone was
coined in 1905 by medical researchers who defined them as ‘the
chemical messengers which speeding from cell to cell along the
blood stream, may coordinate the activities and growth of differ-
ent parts of the body’ (Tata, 2005). This initial, animal-centric
definition has been expanded to include chemical messengers
of many types, but many researchers refer to plant hormones
as phytohormones. Moreover, the concept of diffusible plant
growth substances pre-dated the hormone concept and histori-
cally some have preferred the terms ‘plant growth substance’
or ‘plant growth regulator’ to plant hormone, which is now in
common usage. Plant hormones are molecules made by cells in
response to an environmental or developmental signal that can act
at a distance from their site of synthesis. In addition, hormones are
potent and have biological effects at low (nanomolar to micromo-
lar) concentrations. Plant hormones are frequently small organic
molecules (<500 Da) that can diffuse easily between cells.
Plant hormones play integral roles in coordinating growth
and development between organ systems that can sometimes
be separated by large distances. Such action at a distance is
a defining property of hormones that can be elegantly demon-
strated through grafting experiments. For example, mutant plants
that are deficient in the biosynthesis of strigolactones (SLs)
display an excess shoot branching phenotype due to abnor-
mally active axillary buds; grafts of SL-deficient shoots onto
wild-type, SL-producing roots restores normal shoot branching
patterns. This simple experiment demonstrated the existence
of a root-to-shoot transmissible signal that regulates branch-
ing long before it was determined to be SL (Domagalska and
1
 Plant Hormones

O OH O
H
OC OH
H
HO N
H O
O OH N
H O
HO O
Gibberellin A4 Indoleacetic acid (+)-7-iso-Jasmonyl-L-Isoleucine

O O

H H
OH H H O
O OH OH
O
O
Abscisic acid Ethylene (+)-Strigol O

HO
OH
H OH
O OH

HO
HO NH
H H
H N N
HO O
H N N
O
epi-Brassinolide Salicylic acid trans-Zeatin

Figure 1 Structures of plant hormones.

Leyser, 2011). Hormone movement is typically mediated by
active transport by both influx and efflux carrier proteins located
in cellular membranes. Brassinosteroids (BRs), described below,
provide a counterexample of long-distance action. Because BRs
are hydrophobic molecules with low aqueous solubility, they act
close to their sites of synthesis, but are still classified as hormones.
Many animal hormones are large molecules such as peptides,
and a number of systemically acting plant peptide signals have
also been described. A few examples include systemin, which is
produced locally at the sites of wounding and acts throughout the
plant body to modulate defence responses in the Solanaceae fam-
ily (Ryan, 2000); a recently described family of CEP (C-terminal
encoded peptides) that participate in signalling nitrogen status
perceived by roots to plant shoots (Tabata et al., 2014); and
phytosulfokines, five-amino acid peptides that promote cellular
differentiation (Shinohara et al., 2007). Although plant peptide
hormones are important signals, some have species-specific activ-
ities due to rapid divergence of primary amino acid sequence.
Interestingly, all of the currently known receptors of plant pep-
tide signals are receptor-like kinases. Hundreds of receptor-like
kinases and predicted small proteins are encoded by each plant
genome, suggesting that there may be many more peptide plant
growth regulators (PGRs) to be found. Indeed, a recent explosion
of research is making it clear that plant peptide signals are impor-
tant growth regulators in plants. Here, however, we focus on a
core set of nine small molecule hormones that can be found in
essentially all land plants (Figure 1).
How are plant hormones discovered? In general, plant
hormones have been identified through a research strategy
sometimes humorously referred to as ‘grind-and-find’; that
is, large amounts of plant tissues are homogenised and their
chemical constituents separated (fractionated) using chemi-
cal methods such as high performance liquid chromatography
(HPLC). These fractions are then tested using bioassays, for
example cell growth, which indicate the presence of a candidate
hormone. The chemical complexity of bioactive fractions is
iteratively reduced through a series of purification steps that
are coupled with bioassays, until chemical methods such as
nuclear magnetic resonance (NMR) spectroscopy can be used to
deduce the structures of the few remaining candidate molecules.
Finally, organic synthesis of the putative hormone is conducted
to validate the deduced structure. This process is often called
bioassay- or activity-guided fractionation and is the process that
has been used to isolate most plant hormones. This method is
labour-intensive and requires large amounts of starting materials
to obtain sufficient quantities for hormone identification. For
example, brassinolide (the first BR) was identified in extracts
prepared from 40 000 g of pollen that was collected from several
acres of Brassica napus using honeybees (Grove et al., 1979).
Given how technically difficult it is to isolate plant hormones,

2 eLS © 2017, John Wiley & Sons, Ltd. www.els.net


it is probable that others exist. The discovery of new hormones
remains an exciting area of plant research.
How are the biological functions of hormones established?
While chemical methods have been powerful for isolating and
defining plant hormones, genetic analysis has been instrumental
for determining how plant hormones are made, stored, deacti-
vated, perceived, and function at the molecular level. Geneticists
generally use mutants defective in either hormone biosynthesis
or hormone response to understand hormone biology and to infer
hormone functions. For example, plants harbouring mutations
in the locus GA2 in Arabidopsis thaliana,, which encodes an
enzyme called Ent-Kaurene Synthase, are unable to make gib-
berellins (GAs) (Yamaguchi et al., 1998). These biosynthesis
mutants have dwarf stature and often flower later than wild-type
plants, which indicates that GAs are important for both growth
and for the induction of flowering (Hedden and Thomas, 2016).
A distinctive feature of a biosynthetic mutant is that its pheno-
types can be rescued by adding back the hormone that is missing.
For example, the dwarf phenotype of a ga2 mutant plant can be
rescued by treatment with exogenous GA.
Response- or signalling-defective mutants can still synthe-
sise a particular hormone but do not respond appropriately to
it. In fact, in many cases, signalling-defective mutants have
increased amounts of the unperceived hormone, due to reduced
negative feedback inhibition of biosynthesis. Signalling mutants
are useful for defining the genes and proteins required for sig-
nal transduction. For example, to understand how ethylene is
perceived, geneticists identified a series of ethylene resistant
(Etr) mutants (Bleecker et al., 1988). Ethylene induces a triple
response in dark-grown seedlings consisting of a shortened
hypocotyl, shortened root, and apical hook. A population of
mutant seedlings was easily screened in the dark in the presence
of ethylene for ethylene-insensitive mutants that literally stood
out above the rest. Characterisation of the Etr class of mutants
led to the identification of ethylene receptors and a detailed
molecular understanding of how ethylene elicits changes in
cellular function (Chang et al., 1993). In principle, geneticists
can identify all of the genes necessary for the biosynthesis or
response to a hormone by isolating a large number of mutants.
However, genetic methods, while powerful, can be limited
because plant genomes often contain multiple, closely related
copies of genes that function interchangeably; as a consequence
of this redundancy, mutations in some genes will not cause
detectable phenotypes. In addition, lethal or infertile phenotypes
may prevent the identification of some genes. In some cases,
synthetic chemicals that selectively target subsets of otherwise
redundant factors or temporarily inhibit gene function can
overcome these problems (Park et al., 2009).
Discovery and Functions of Plant
Hormones
Ethylene (ET)
The old adage ‘one rotten apple spoils the whole barrel’ origi-
nates from the action of the gaseous hormone ethylene. The role
eLS © 2017, John Wiley & Sons, Ltd. www.els.net
Plant Hormones
of ethylene as a plant hormone was discovered accidentally in
the mid-nineteenth century when it was observed that plants cul-
tivated in the proximity of gas lamps displayed growth defects
including small stature. The natural gas burnt in the lamps con-
tains trace amounts of ethylene, which was identified as the
bioactive component in the early twentieth century. Ethylene is
often described as the ‘ripening hormone’ because it is criti-
cal for the induction of senescence during fruit ripening and
other plant biological processes. The loss of leaves and flower
petals due to organ abscission, are also forms of senescence that
are controlled by ethylene in most plant species. Mutant tomato
plants with defective ethylene perception display a ‘never ripe’
phenotype in which fruit ripening and organ abscission do not
occur (Lanahan et al., 1994). Other prominent effects of ethy-
lene include mediation of responses to biotic or abiotic stress,
inhibition or stimulation (depending on plant species) of organ
elongation and flowering, and sex determination of flowers (Wang
et al., 2002). Ethylene, being a gas, is well suited to indicating
enclosed environments. For example, during submergence, deep-
water rice internodes elongate at nearly one foot per day to keep
their shoots above water; submergence is signalled by the accu-
mulation of high ethylene, which would otherwise diffuse rapidly
into air (Bailey-Serres et al., 2012).
The biosynthetic pathway for ethylene formation in plants
was elucidated in the late 1970s. Ethylene is synthesised in
plants by a simple pathway from the amino acid methion-
ine. A key intermediate in the biosynthesis is the substance
1-aminocyclopropane-1-carboxylic acid (ACC), which is
synthesized by the enzyme ACC synthase (ACS) using
S-adenosyl-L-methionine as a substrate (Wang et al., 2002).
ACS is a highly regulated enzyme that plays a central role in
determining ethylene levels. ACS, like many proteins, has a
short half-life and is unstable under typical growth conditions.
However, conditions such as stress or the transition to senescence
stabilise ACS by preventing its degradation, triggering increased
ethylene synthesis (Chae and Kieber, 2005).
Auxin (IAA or IBA)
While studying phototropisms of coleoptiles in the nineteenth
century, Charles Darwin, concluded that a signal from the tip was
transported to the tissue below to induce bending. Years later this
principle was identified as indole-3-acetic acid (IAA). IAA is the
predominant naturally occurring representative of the auxin class
of plant hormones, which also includes the auxin indole-3-butyric
acid (IBA). A common joke heard at plant hormone conferences
is that auxins ‘do everything.’ Indeed, auxins have been impli-
cated in a wide variety of processes such as cell elongation,
apical dominance, tropic growth responses stimulated by light
and gravity, and the initiation of lateral and adventitious root for-
mation (Woodward and Bartel, 2005). IAA is derived from the
amino acid tryptophan. There are multiple pathways to form IAA,
however a simple three-step pathway involving the Yucca family
of cytochrome P450s most likely plays the predominant role in
most land plants (Zhao, 2010). Auxins are also stored as sugar
or amino acid conjugates, providing an alternative source of free
auxin.
3
 Plant Hormones
Gibberellins (GAs)
The first GAs were isolated as bioactive molecules produced by
the plant pathogenic fungus Gibberella fujikuroi, which causes a
disease in rice called ‘bakane’. This translates to ‘foolish seedling
disease,’ an apt description because infected seedlings lodge (fall
over) due to extensive growth triggered by fungal-produced GAs.
Endogenous plant-produced GAs were isolated in the late 1950s,
and currently more than 100 GAs are known. The main bioactive
forms in plants are GA1 , GA3 , GA4 , and GA7 (GA4 is shown
in Figure 1). GAs are constructed from a repeating 5-carbon
unit called isopentyl diphosphate that is synthesised largely in
the plastids. The pathway to active GAs is complex and involves
many enzymes, but a key step is controlled by ent-kaurene syn-
thase; mutant plants lacking this enzyme are dwarfs. GAs are
involved in stem elongation, stimulation of germination, induc-
tion of enzymes upon seed germination, flowering, transition
of developmental stages (e.g. juvenile–adult), sex determina-
tion of flowers, and other plant physiological processes (Yam-
aguchi, 2008). During the 1960s, the adoption of high-yielding
semi-dwarf wheat varieties bred by Norman Bourlag stimulated
the ‘Green Revolution’, which saw a near doubling of yields in
Pakistan, India and eventually other countries. Bourlag’s varieties
are less sensitive to GA due to alterations of a key protein involved
in GA response related to GID1, a key GA receptor (Peng et al.,
1999). The higher yield of semi-dwarfs arises in part because
the plants lodge less than conventional varieties. GA-deficient
strains of rice and other crops have been bred and are now com-
monplace in modern agriculture (Hedden, 2003). As described
below, chemical inhibitors of GA biosynthesis can be used to
induce semi-dwarf phenotypes and realise similar yield improve-
ments.
Cytokinins (CKs)
Kinetin, the first known cytokinin (CK), was isolated in the 1950s
as a degradation product of herring sperm DNA (deoxyribonu-
cleic acid) by Skoog and Miller, who were at the time systemat-
ically searching for compounds that could stimulate cell division
in cultured plant cells. Trans-zeatin, the main endogenous CK,
was isolated later in the 1960s from maize (Amasino, 2005).
Kinetin and trans-zeatin are N6 -substituted adenosine deriva-
tives, of which many have been isolated from plants. A key
step in the biosynthesis of CKs is the modification of adenine
by the enzyme adenosine phosphate-isopentyltransferase (IPT)
(Kamada-Nobusada and Sakakibara, 2009). Together with aux-
ins, CKs are the most essential plant growth factors necessary for
cell proliferation and organogenesis.
Abscisic acid (ABA)
Abscisic acid (ABA) was isolated from plant tissues in the early
1960s by multiple groups; one group set out to identify molecules
that control organ abscission and isolated ABA, which they called
‘abscisin II’, while another identified it as a growth inhibitor
in dormant tree buds. The latter group called their molecule
‘dormin’, which is a better name because ABA plays a rela-
tively minor role in organ abscission; nonetheless, the name ABA
4 eLS © 2017, John Wiley & Sons, Ltd. www.els.net
stuck. ABA is derived from ß-carotene, the pigment that makes
carrots orange. There are several enzymatic steps in between
ß-carotene and ABA. A key step in ABA biosynthesis involves
9-cis epoxycarotenoid dioxygenases (NCEDs), which cleave a
C40 carotenoid into a C15 product that is processed further into
ABA. NCEDs are critical regulators of ABA levels and are tran-
scribed at high levels during many forms of abiotic stress (Nam-
bara and Marion-Poll, 2005).
ABA is a ubiquitous hormone in land plants that plays a central
role in regulating transpiration because it stimulates stomatal
closure. Mutant plants that are deficient in ABA biosynthesis
wilt easily and have reduced resilience to water stress in com-
parison to wild-type plants. Another critical role for ABA is
in the establishment and maintenance of physiological seed
dormancy, which blocks germination under suboptimal condi-
tions or times. ABA-deficient plants can display a viviparous
phenotype in which seeds germinate while still attached to the
mother plant; in agriculture, such pre-harvest sprouting can
cause substantial decreases in yield and grain quality, and is par-
ticularly problematic for wheat. ABA also affects bud dormancy,
inhibits elongation growth, accelerates senescence, and mediates
responses to stress, in particular, drought, salt and cold stress
(Cutler et al., 2010). Treatment of plants with high levels of ABA
triggers ethylene synthesis, which in turn causes organ abscission.
Jasmonate (JA)
Methyl jasmonate (MeJA) was first identified as a fragrance
component of jasmine flower extracts used in perfumes and
was later shown to have important roles in wounding and other
defense responses. JA is a member of a large family of molecules
derived from C18 fatty acids called the octadecanoates. These
include JA and 12-oxophytodienoate (OPDA), which regulate
developmental processes (e.g. tuber formation, establishment of
pollen fertility, mechanosensing, and responses to biotic and
abiotic stress). Although this class of hormones is generally
referred to as jasmonates (JAs), JA is not the bioactive hor-
mone; rather JA is conjugated to different amino acids, most
frequently isoleucine, which yields jasmonyl-Ile (JA-Ile), the
form of the hormone that interacts with the SCFCOI1 receptor
(see below). The JA signalling system can be hijacked by some
plant pathogens. For example, the plant bacterial pathogen Pseu-
domonas syringae synthesises a mimic of JA-Ile called corona-
tine, which forces guard cells to open. This allows P. syringae
to gain access to plants through the stomata (Melotto et al.,
2006).
The key regulatory step in the biosynthesis of JA and other
octadecanoates is the oxidation of C18 fatty acids by a family
of enzymes called lipoxygenases (LOXs), which act on lipids
released from plastid membranes by lipases after mechanical
damage (Wasternack and Hause, 2013). Related enzymes in
mammals control the biosynthesis of C20 fatty acid derivatives,
such as arachidonic acid, which get processed into prostaglandins
and other inflammatory hormones. The C18 and C20 fatty acids
precursors catabolised by LOXs are released by conditions that
disrupt membranes; these metabolites often signal damage to
membranes and trigger appropriate protective responses.

Strigolactones (SLs)
SLs were first discovered in the 1960s as compounds in root exu-
dates that trigger germination of noxious weeds in the Oroban-
chaceae family of root parasites. The impact of these weeds on
crop yields is tremendous; Striga spp. alone, from which SLs
derive their name, collectively cause billions of US dollars in
annual crop losses and are a growing challenge to food security
in sub-Saharan Africa. This raised the obvious question of why
plants would release a chemical into the soil that makes them
vulnerable to parasitism. In 2005, it was found that SLs pro-
mote symbiotic interactions with arbuscular mycorrhizal fungi,
which trade mineral nutrients for carbon. Fitting this role, SL
exudation increases under nutrient-poor conditions in which myc-
orrhizal associations are most beneficial. A few years later, SLs
were revealed to be a root-derived, upwardly mobile signal that
suppresses shoot branching, which had been suspected since the
1990s based upon grafting studies with more axillary growth
(max) mutants. Since then, the list of hormonal roles for SL has
continued to grow and currently includes regulation of root archi-
tecture, senescence and secondary growth.
SLs are derived from carotenoids through the consecutive
action of a carotenoid isomerase (D27) and two carotenoid
cleavage dioxygenases (CCD7/MAX3 and CCD8/MAX4),
producing carlactone. Further action by a cytochrome P450
(CYP711A1/MAX1) leads to SL or carlactonoic acid, which has
SL activities but a noncanonical SL structure.
Brassinosteroids (BRs)
Brassinolide was the first BR to be identified. It was isolated
from oilseed rape (B. napus) pollen and its structure was elu-
cidated in 1979. As its name implies, BRs are a plant steroid
(or phytosteroid) derived from a cholesterol-like molecule called
campesterol. BRs have been implicated in a variety of processes,
including stimulation of growth, enhancement of tolerance to
biotic and abiotic stress and cell differentiation. Mutants deficient
in BR synthesis have dwarfism and display defects in photomor-
phogenesis.
Salicylic acid (SA)
Salicylic acid (SA) is the natural precursor of aspirin (acetylsali-
cylic acid) and has been used medicinally for millennia. In the
1970s, plant biologists observed that the application of SA to
plants could protect them from infection by pathogens and mimic
a natural defence system known as systemic acquired resistance
(SAR). Plants unable to accumulate SA are highly susceptible
to infections by many pathogens. SA is produced at high levels
around sites of infection and can act at a distance to trigger protec-
tion against future infection. For example, SA triggers the produc-
tion of pathogenesis-related (PR) proteins, some of which have
antimicrobial activities. There are multiple biosynthetic routes to
SA; however, most SA is derived from chorismate, the precursor
of the aromatic amino acids phenylalanine and tyrosine.
eLS © 2017, John Wiley & Sons, Ltd. www.els.net
Plant Hormones
Hormone Perception
Hormone signalling requires detection of a hormone by receptor
proteins, as well as a way to transduce that event to initiate
a signalling cascade, for example by altering the transcription
of a set of genes. Genetic analyses over the last three decades,
primarily conducted in A. thaliana and rice, has led to a detailed
understanding of the signal transduction cascades for each of the
nine major hormones. A few major themes that distinguish plant
hormone signalling have emerged from these analyses.
In mammals, many signal transduction events are initiated
by plasma membrane-localised G-protein coupled receptors
(GPCRs), of which there are hundreds encoded by a typical
genome. GPCR activation of a heterotrimeric G-protein complex
often initiates signal amplification by secondary messengers
such as cyclic AMP (adenosine monophosphate) or calcium.
Plants, however, have very few GPCRs. Instead, most hormone
signalling in plants is mediated by soluble proteins. Of the
nine hormones, only three (BR, ET and CK) are perceived by
membrane-localised receptors and of these, the BR receptor is
the only one localised to the plasma membrane. In mammals,
many small molecule hormones, such as retinoic acid, are per-
ceived by proteins in the Nuclear Hormone Receptor family,
which lack homologues in plant genomes; instead, plants have
co-opted other families of proteins to accomplish similar tasks
(Lumba et al., 2010).
Plant hormone perception is often linked to the stabilisation
of protein–protein interactions. This can take two forms. First,
a hormone receptor may undergo conformational changes in
response to binding its ligand that enhance its affinity for a
partner protein(s). This is known as allosteric regulation and
is the case with GA, ABA, and SL signalling. Alternatively,
the hormone may act as a ‘molecular glue’ that stabilises the
interaction between two proteins that function as co-receptors.
This is seen in auxin, JA, SA, and BR signalling.
For several plant hormones, the hormone-induced formation of
a protein complex regulates the stability of important signalling
factors. Hormone-activated proteolysis is a common feature of
the auxin, JA, GA, SL, and SA signalling pathways (Figure 2).
In each mechanism, an F-box protein serves as an adapter com-
ponent that confers specificity for a particular set of protein sub-
strates to an SCF (Skp1-Cullin-F-box) ubiquitin ligase complex.
Upon association with the SCF complex, the substrate is polyu-
biquitylated and then degraded by the proteasome. Degradation
may happen very rapidly; JAZ (jasmonate ZIM-domain) proteins,
for example, have a half-life of 1.4 min following JA treatment
(Pauwels et al., 2010).
Relief of transcriptional repression is another important theme
in plant hormone signalling. Many of the SCF substrates of
hormone signalling pathways are components of a transcriptional
co-repressor complex, although their interaction with DNA may
be indirectly mediated by association with transcription factors.
Thus, proteolysis of the SCF substrate can result in transcriptional
activation.
5
 Plant Hormones

TIR1
Auxin Aux/IAA
AFB
JA-Ile COI1 JAZ
GA SLY1 GID1 DELLA
D53
SL MAX2 D14
SMXL6/7/8

‘Molecular glue’ signaling mechanism Ub

Ub

Ub
Target Ub

H Target
Ub Ub
H
F-box F-box
E2 E2
Skp1 Rbx1 Skp1 Rbx1

Cullin1 Cullin1

Ub
Allosteric signaling mechanism Ub

Ub
Target Ub

Receptor
Target
H Receptor
Ub H Ub

F-box F-box
E2 E2
Skp1 Rbx1 Skp1 Rbx1

Cullin1 Cullin1

Before hormone perception After hormone perception

Co-repressor
complex

Target
OFF ON

TF Hormone-responsive gene TF Hormone-responsive gene

Figure 2 Hormone signalling pathways. The signalling mechanisms of Auxin, JA, GA, and SL involve hormone-activated targeting of protein substrates
through SCF (Skp1-Cullin-F-box) complexes. SCF complexes are a type of E3 ubiquitin ligase that consist of Skp1, Cullin1, and an F-box protein that confers
substrate specificity. E3 ubiquitin ligases work in concert with E1 (not shown) and E2 ubiquitin ligases to attach ubiquitin or polyubiquitin chains to proteins.
Polyubiquitylated proteins are rapidly degraded by the 26S proteasome. Two types of SCF-mediated plant hormone signalling mechanisms have been
observed. The hormone may function as molecular glue that facilitates binding between an F-box protein and its target substrate. Effectively these proteins
are co-receptors. Alternatively, a receptor protein may undergo conformational changes following ligand binding or processing (allosteric regulation) that
promote formation of a complex with an F-box protein and a target. In either case, the target is polyubiquitylated and degraded. The targets of the auxin,
JA, and GA pathways function as transcriptional regulators, and it is thought that this is also a function of SL targets. None of the known targets directly
bind DNA; instead, they interact with transcription factors and modulate their activity. The targets of auxin, JA, and likely SL signalling are associated with
transcriptional repressor complexes. Therefore, degradation of the target protein can relieve transcriptional repression.

Auxins through interaction with the TPL/TPR class of transcriptional

Auxin functions through an elegant mechanism that controls the
levels of downstream transcriptional effectors by regulating their
proteolysis. Aux/IAA proteins block the transcriptional activation
activity of ARF (auxin response factor) transcription factors,
which directly bind to the promoters of auxin-responsive genes,
co-repressors. Auxin binds at the interface between the F-box
protein TIR1 (or the related AFB F-box proteins) and Aux/IAAs,
thus acting as a glue to bring the SCFTIR1 complex together with
its Aux/IAA substrates (Tan et al., 2007). Effectively, TIR1/AFB
and Aux/IAAs function as co-receptors. Aux/IAAs are then
polyubiquitylated and degraded by the proteasome, relieving

6 eLS © 2017, John Wiley & Sons, Ltd. www.els.net


Aux/IAA-mediated repression of ARF activity and producing
transcriptional responses to auxin (Mockaitis and Estelle, 2008).
The auxin response system was surprising in its simplicity. After
it was understood in molecular detail, it became apparent that
other hormone pathways utilise molecular glues that control inter-
actions between SCFs and their target proteins.
Jasmonates
JA perception is conceptually very similar to auxin perception as
it relies on a molecular glue-type mechanism in which JA con-
trols transcriptional repressor levels through ubiquitin-mediated
proteolysis. However, most of the proteins involved are unique
to JA responses. JAZ proteins repress the activity of MYC
family transcription factors, which bind to promoter elements
found in JA-regulated genes, by interaction with MYC and
a TPL/TPR-interacting protein known as NINJA. JA-Ile sta-
bilises a complex between SCFCOI1 and JAZ proteins, leading to
their rapid polyubiquitylation and degradation, and the relief of
JA-responsive transcriptional repression. Thus, the functions of
JA and auxin signalling components are highly similar although
they do not share sequence homology; COI1 is analogous to
TIR1/AFB, JAZ proteins are analogous to Aux/IAAs, and MYC
proteins are analogous to ARFs (Wasternack and Hause, 2013).
Gibberellins
GA perception triggers hormone-activated proteolysis, similar
to auxin and JA signalling, but features an allosteric signalling
mechanism. The GA receptor GID1 is an 𝛼/𝛽-hydrolase super-
family protein that undergoes conformational changes following
GA binding that promotes association with DELLA proteins and
the F-box protein SLEEPY1. DELLAs are polyubiquitylated and
degraded, giving rise to GA growth responses. In Arabidopsis,
different members of the DELLA family have more specialised
roles in different aspects of growth. DELLAs can act as either
positive or negative transcriptional regulators but do not bind
DNA directly. They do so either through working with a transcrip-
tion factor to activate target gene expression, or through seques-
tering transcriptional activators or repressors. DELLAs have been
implicated in interactions with transcriptional regulators found
in several hormone pathways, providing a means for crosstalk
(Davière and Achard, 2016).
Strigolactones
Similar to GAs, the SL receptor D14 is an 𝛼/𝛽-hydrolase protein.
Unlike GID1, however, D14 retains and requires hydrolytic activ-
ity for its signalling function. D14 has very slow hydrolytic activ-
ity upon SL in vitro, which is due to a two-phase action. It rapidly
catalyses cleavage of the enol-ether linked butenolide compo-
nent of the SL molecule, which is then covalently attached to a
catalytic His residue for an extended duration. In this activated
state, D14 undergoes a conformational change that is stabilised
through association with the F-box protein MAX2. D14 also
associates with downstream target proteins in the SMXL/D53
family, which are polyubiquitylated by SCFMAX2 and degraded.
The function of SMXL/D53 proteins is currently uncertain, but a
eLS © 2017, John Wiley & Sons, Ltd. www.els.net
Plant Hormones
popular hypothesis is that they function as transcriptional repres-
sors through interactions with TPL/TPR proteins.
Salicylic acid
SA perception was the last of the major hormones to be elucidated
and still has unresolved questions. SA controls the transcription
of many defence genes, for example the PR genes mentioned
above through NON-INDUCER of PR1 (NPR1), a key factor iden-
tified genetically as being necessary for SA-mediated induction
of PR genes. NPR1 interacts with the transcription factor TGA2,
which represses SA-mediated transcription. When NPR1 binds
to TGA2, it converts it into an activator, thereby de-repressing
transcription. NPR1 protein levels increase upon SA treatment
and it translocates to the nucleus to activate transcription. NPR1
can directly bind SA and synthetic SA mimics in a test-tube with
high affinity; along with related proteins it is believed to be an SA
receptor; however, this is still under investigation. Interestingly,
NPR1, like the ethylene receptor, requires Cu++ to bind its ligand.
It has been proposed that SA induces conformational changes in
NPR1 that may be sufficient for nuclear translocation and subse-
quent transcriptional activation. Although the details remain to be
worked out, it is clear that NPR1 and related proteins participate
in SA perception.
Abscisic acid
ABA is perceived by a large family of related proteins called
PYRABACTIN RESISTANCE 1 (PYR1)/PYR1-like (PYL) or reg-
ulatory component of ABA receptor (RCAR). Most angiosperm
genomes contain between 12–24 ABA receptors, making them
the single largest family of receptors for a single hormone. ABA
levels are usually low under normal growth conditions but rise
during environmental stress or during appropriate developmen-
tal states such as seed development. ABA action is mediated
by a three-part signalling module that links cellular ABA lev-
els to protein kinase activity. PYL receptors directly bind ABA,
which induces a conformational change in the PYLs that enables
them to bind a group of protein phosphatases (clade A protein
phosphatase type 2Cs, or PP2Cs). ABA-bound PYLs bind in the
PP2C active sites and block the access of substrates to the active
catalytic activity; thus ABA inhibits PP2C activity by trigger-
ing a conformational change in the PYL receptors. The PP2Cs
are intrinsically active and, unless inhibited by PYL receptors,
continuously dephosphorylate a small family of stress-activated
kinases called the SnRK2s (SUCROSE NON-FERMENTING
RELATED SUBFAMILY 2 KINASEs). Upon PP2C inhibition,
SnRK2s auto-activate and then control the activity of downstream
effectors by phosphorylation. For example, ABI5 and its related
DNA-binding proteins (the ABFs) activate transcription after
phosphorylation by the SnRK2s. One unresolved question is why
land plants require so many receptors for a single hormone. One
hypothesis is that there may be other endogenous ligands that acti-
vate the different subfamilies (Peterson et al., 2010). Recent data
suggests that the ABA catabolite (breakdown product) functions
as a weak selective receptor agonist (Weng et al., 2016). Addi-
tionally, differences in ABA affinity between receptors have been
documented; this may facilitate dose-dependent responses over a
wider range than possible with a smaller number of receptors.
7
 Plant Hormones
Ethylene
Ethylene binds directly to a family of homologous ER (endoplas-
mic reticulum)-membrane localised proteins that were initially
defined by the founding member ETR1. Ethylene receptors con-
tain a Cu++ ion, which coordinates ethylene binding (Rodríguez
et al., 1999). Dominant mutations that lock ETR1 in its active
state inhibit ET signalling; thus, ETR1 is a negative regulator
of ET responses. In contrast to the majority of signalling mech-
anisms, ET binding inactivates ET receptors. In the absence of
ethylene, the receptors activate the protein kinase CTR1 (CON-
STITUTIVE TRIPLE RESPONSE 1), which negatively regulates
ethylene action (Kieber et al., 1993). The function of CTR1 is to
phosphorylate the C-terminus of ethylene insensitive 2 (EIN2),
which keeps EIN2 in an inactive state. Upon ethylene binding,
CTR1 activity is reduced and EIN2 becomes dephosphorylated;
this then allows an unknown protease to cleave the C-terminus of
EIN2, which contains a nuclear localisation signal that facilitates
its translocation into the nucleus (Qiao et al., 2012). Once in the
nucleus, the C-terminus of EIN2 stimulates the transcription of
ethylene regulated genes through interactions with ETHYLENE
INSENSITIVE 3 (EIN3) and related transcription factors. Thus,
ET controls a simple circuit: it binds to its receptors, which lowers
CTR1 kinase activity, which in turn leads to cleavage and nuclear
translocation of the C-terminus of EIN2. There are still unre-
solved questions in ethylene signal transduction regarding how
cleavage of EIN2 occurs after ET perception, and how CTR1 is
inactivated by ET receptors.
Cytokinins
CKs are perceived by a family of ER membrane-localised pro-
teins related to the Arabidopsis gene AHK4 (ARABIDOPSIS
HISTIDINE KINASE 4). CK receptors are derived from bacterial
two-component regulatory systems, which are involved in many
ligand sensing systems such as bacterial chemotaxis towards
nutrients. CK receptors were presumably co-opted from pro-
teins present in the endosymbionts that ultimately evolved into
organelles. CK receptors contain an extracellular ligand-binding
domain called the CHASE (CYCLASE HISTIDINE KINASE
ASSOCIATED SENSORY EXTRACELLULAR) domain that is
found in a large family of bacterial ligand sensors. Although
two-component regulators have an ancient bacterial lineage, the
CK-binding CHASE domain appears to be restricted to land
plant lineages. CK binding to the CHASE domain leads to acti-
vation of kinase activity and receptor autophosphorylation. Once
activated, CK receptors phosphorylate a group of proteins called
AHPs (ARABIDOPSIS HISTIDINE-CONTAINING PHOSPHO-
TRANSFER PROTEINs), which subsequently transfer phosphate
to a subfamily of transcriptional regulators called ARRs (ARA-
BIDOPSIS RESPONSE REGULATORs). ARRs directly bind
to the promoter regions of CK-regulated genes (Kieber and
Schaller, 2010).
Brassinosteroids
Of all of the plant hormone receptors, the BR receptor displays the
most ‘animal’-like signalling mechanism. BR binds directly to
8 eLS © 2017, John Wiley & Sons, Ltd. www.els.net
BRI1 (BRASSINOSTEROID INSENSITIVE 1), which is localised
in the plasma membrane. The ligand-binding domain of BRI1
is located on the extracellular side of the membrane, but an
internal kinase domain becomes activated upon BR binding. BR
perception involves features reminiscent of the ‘molecular glue’
mechanism described above, in that BR binding is shared equally
between the co-receptors BRI1 and BAK1 (BRI1-ASSOCIATED
KINASE 1). BR-stimulated BRI1-BAK1 association leads to acti-
vation of the BRI1 intracellular kinase domain by transphos-
phorylation. Once activated, this kinase domain phosphorylates
and activates BSK protein kinases, which in turn phosphorylate
and activate BSU protein phosphatases, which dephosphorylate
and inactivate BIN2, a negative regulator of BR action. BIN2
normally phosphorylates the BZR transcription factors, which
inactivates their function. Thus, BR perception leads to inactiva-
tion of BIN2 kinase activity, accumulation of dephosphorylated
BZR1, and subsequent activation of BR-regulated genes (Kim
and Wang, 2010). BRI1 and BAK1 both contain extracellular
leucine rich repeats (LRRs), which are present in several hundred
Arabidopsis proteins. Extracellular LRRs appear to be involved
in many aspects of molecular recognition, involved in pathogen
perception. In most cases, LRR proteins recognise peptide lig-
ands rather than small molecules.
Hormone Crosstalk
Plant hormones often influence the behaviour of other hormone
pathways, in what is broadly known as crosstalk. Crosstalk can
take many forms, ranging from regulating another hormone’s
abundance, transport, or signalling efficacy, to modulating the
activity of a shared signalling node. To briefly describe one
example from the control of shoot branching, SLs inhibit the polar
transport of auxin by reducing the amount of the auxin efflux
carrier PIN1 at the plasma membrane. However, the crosstalk
relationship is not one-way, as auxin positively regulates expres-
sion of SL biosynthesis genes. Another classic example of plant
hormone crosstalk is the antagonistic control of seed germina-
tion by ABA and GA. ABA promotes a state of seed dormancy,
whereas GA promotes germination. The balance between the two
hormone pathways determines the germination outcome. ABA
inhibits the expression of GA biosynthesis genes. In addition,
GA and ABA pathways converge on MFT (MOTHER OF FT
AND TFL1); GA inhibits expression of MFT through influencing
DELLA abundance, and the ABA-activated transcription factors
ABI3 and ABI5 have opposing effects on MFT expression (Xi
et al., 2010).
Hormones also crosstalk with nonhormonal pathways and their
activity is influenced by light, temperature and biotic interac-
tions. For example, either light or cold (stratification) stimulate
expression of GA biosynthesis genes in imbibed Arabidopsis
seeds, and cold can also promote sensitivity to exogenous GAs.
Gating of hormone responses by the circadian clock has also
been observed; at some times of day plants may be more recep-
tive to a hormone than others. In one study, a synthetic pro-
moter that reports auxin signalling activity (eDR5) was fused to
the luciferase reporter gene (LUC). Transgenic plants carrying
this transcriptional reporter showed different responses to auxin
 Plant Hormones

Cl N S O
N
O S
O
Cl OH
2,4-D Acibenzolar-S-methyl
OH
N S O
N
N N N
H H N Cl
N
Thidiazuron Paclobutrazol

NH2
OH
OH
O P
O
OH Cl O
NH2
Ethephon AVG

O N
O
S
N
H O

Quinabactin MCP

Figure 3 Structures of hormone mimics and inhibitors used in agriculture.

application at night versus in the day, with corresponding changes
in hypocotyl elongation responses to auxin treatment (Covington
and Harmer, 2007).
Uses of Synthetic Plant Hormones
Given the power of plant hormones to control plant physiology
and development, synthetic hormones and hormone inhibitors
have become part of the modern chemical arsenal used in agri-
culture. Plant hormones and synthetic mimics are often broadly
classified by industrial scientists as PGRs (Figure 3). Because
of increasing pressures on agricultural productivity imposed by
climate change, population growth, and post-industrial lifestyles,
chemical strategies that improve plant performance during
drought and other stressful environments are actively being
pursued. To date, agrochemicals are used primarily to control
insects, weeds, and biotic pathogens like fungi. Synthetic modu-
lators of plant hormone pathways may create new opportunities
for modulating crop physiology.
Auxins
Modern agriculture relies heavily on the use of synthetic herbi-
cides for weed control. The first synthetic herbicide, 2,4-D, was
developed in the 1940s and remains in widespread use (Peter-
son, 1967). 2,4-D binds to auxin receptors and activates auxin
responses. It is applied at high levels that cause misregulation
of development and cellular physiology, which ultimately causes
plant death. As 2,4-D is crop selective, it does not kill all plants. In
general, plants in the grass family such as maize, wheat and rice,
tolerate 2,4-D because they are able to detoxify it. For this reason,
2,4-D is the active ingredient in many products sold to control
dicot weeds (such as dandelions) in household lawns. Many syn-
thetic auxins have been developed since the discovery of 2,4-D
(Grossmann, 2010).
Ethylene
As Ethylene is a gas, its physical application is more challenging
than other PGRs. To address this limitation, the chemical ethep-
hon was developed; this molecule is hydrolysed into an unstable
product that releases ethylene. Ethephon stimulates defoliation
in cotton and other crops. Other important applications of the
ethylene pathway involve compounds that antagonise its action,
preventing fruit ripening or abscission. For example, when sil-
ver (Ag++ ) ions bind to ethylene receptors in place of the nat-
urally bound Cu++ ions, the receptors are inactivated. Based
on this principle, silver thiosulfate is used in horticulture to

eLS © 2017, John Wiley & Sons, Ltd. www.els.net 9

 Plant Hormones
prolong the shelf-life of cut flowers by preventing abscission.
The ethylene-like gas MCP, binds to ethylene receptors and cova-
lently inactivates them (Pirrung et al., 2008). MCP can be applied
to apples and other fruits after harvest to delay ripening. Ethy-
lene biosynthesis can also be blocked using compounds such as
aminoethoxyvinylglycine (AVG), a substrate-mimic inhibitor of
ACS that is used in agriculture.
Strigolactones
One of the most commonly sought applications for SLs is to
trigger suicidal germination of parasitic weeds, which cannot
survive for more than a few days in the absence of a host. In their
natural form SLs are too expensive to synthesise and unstable in
the field to be of practical use. Many synthetic analogs of SLs
have been developed to overcome these problems while retaining
a high degree of bioactivity. Although this approach is showing
promise in field trials, widespread adoption has not yet happened.
Gibberellins
Natural GAs have been used by horticulturists, gardeners, and
malters to stimulate germination for many years. The most impor-
tant agricultural connection to GA however is to reduce growth
using GA biosynthesis inhibitors. Numerous inhibitors of GA
biosynthesis, such as paclobutrazol, have been developed that
reduce stem or internode elongation, thus providing a chemical
means to reduce lodging. Other important applications for GA
inhibitors include managing the size of fruit trees and cotton.
Salicylic acid
SA plays a critical role in mounting protective responses against
pathogens. The synthetic SA-mimic acibenzolar-S-methyl (also
known as BTH, for its benzothiadiazole substructure) was devel-
oped as an agent that can be applied to crops to specifically
protect against fungal pathogens. Because it activates SAR, how-
ever, it provides broad spectrum defense activity. Acibenzolar
was developed by screening through large compound collections
for molecules that activate PR-gene expression, the outputs of SA
and SAR (Görlach et al., 1996). Other classes of synthetic SA
mimics have been described, but primarily acibenzolar-S-methyl
is used in agriculture.
Cytokinins
Synthetic CKs have found use in agriculture as defoliants, a broad
class of compounds that cause shedding of leaves. Thidiazuron
is one CK developed for this purpose that binds to CK receptors
(Hothorn et al., 2011). CKs induce abscission via ethylene by sta-
bilising ACS, a key regulatory enzyme in ethylene biosynthesis
(Chae and Kieber, 2005).
Abscisic acid
Quinabactin, a synthetic mimic of ABA, has been developed
as a modulator of transpiration. It shows promise as the parent
molecule of a new class of agrochemical agents that may improve
crop yields during drought and mitigate the effects of abiotic
10 eLS © 2017, John Wiley & Sons, Ltd. www.els.net
stress (Helander et al., 2016). ABA itself is currently used in
grape vineyards to increase consistency in the onset of coloura-
tion in table grapes.
Conclusion
We have limited our discussion of plant hormones to several
well-studied small molecules, but there are many other plant
growth regulating substances currently known and others that
remain to be discovered. The pace of discovery of signalling
mechanisms has increased rapidly with the advent of genomic
tools, as exemplified by the rapid delineation of the SL pathway in
the 8 years since its recognition as a plant hormone. The capacity
for discovering new hormones has also increased with techno-
logical gains in the sensitivity of tools for chemical detection
and identification. Unravelling crosstalk between plant hormone
pathways continues to be a major challenge for the field due to its
inherent complexity.
References
Amasino R (2005) 1955: kinetin arrives: the 50th anniversary of a
new plant hormone. Plant Physiology 138: 1177–1184.
Bailey-Serres J, Lee SC and Brinton E (2012) Waterproofing
crops: effective flooding survival strategies. Plant Physiology 160:
1698–1709.
Bleecker AB, Estelle MA, Somerville C and Kende H (1988) Insensi-
tivity to ethylene conferred by a dominant mutation in Arabidopsis
thaliana. Science 241: 1086–1089.
Briggs WR (2014) Phototropism: some history, some puzzles, and a
look ahead. Plant Physiology 164: 13–23.
Chae HS and Kieber JJ (2005) Eto Brute? Role of ACS turnover
in regulating ethylene biosynthesis. Trends in Plant Science 10:
291–296.
Chang C, Kwok SF, Bleecker AB and Meyerowitz EM (1993) Ara-
bidopsis ethylene-response gene ETR1: similarity of product to
two-component regulators. Science 262: 539–544.
Covington MF and Harmer SL (2007) The circadian clock regulates
auxin signaling and responses in Arabidopsis. PLoS Biology 5:
e222.
Cutler SR, Rodriguez PL, Finkelstein RR and Abrams SR (2010)
Abscisic acid: emergence of a core signaling network. Annual
Review of Plant Biology 61: 651–679.
Davière J-M and Achard P (2016) A pivotal role of DELLAs in
regulating multiple hormone signals. Molecular Plant 9: 10–20.
Domagalska MA and Leyser O (2011) Signal integration in the con-
trol of shoot branching. Nature Reviews. Molecular Cell Biology
12: 211–221.
Görlach J, Volrath S, Knauf-Beiter G, et al. (1996) Benzothiadiazole,
a novel class of inducers of systemic acquired resistance, activates
gene expression and disease resistance in wheat. Plant Cell 8:
629–643.
Grossmann K (2010) Auxin herbicides: current status of mechanism
and mode of action. Pest Management Science 66: 113–120.
Grove MD, Spencer GF, Rohwedder WK, et al. (1979) Brassinolide,
a plant growth-promoting steroid isolated from Brassica napus
pollen. Nature 281: 216–217.

Hedden P (2003) The genes of the Green Revolution. Trends in
Genetics 19: 5–9.
Hedden P and Thomas SG (2016) Annual Plant Reviews, The Gib-
berellins. Chichester, UK: John Wiley & Sons, Ltd.
Helander JDM, Vaidya AS and Cutler SR (2016) Chemical manipu-
lation of plant water use. Bioorganic and Medicinal Chemistry 24:
493–500.
Hothorn M, Dabi T and Chory J (2011) Structural basis for cytokinin
recognition by Arabidopsis thaliana histidine kinase 4. Nature
Chemical Biology 7: 766–768.
Kamada-Nobusada T and Sakakibara H (2009) Molecular basis for
cytokinin biosynthesis. Phytochemistry 70: 444–449.
Kieber JJ, Rothenberg M, Roman G, Feldmann KA and Ecker JR
(1993) CTR1, a negative regulator of the ethylene response path-
way in Arabidopsis, encodes a member of the raf family of protein
kinases. Cell 72: 427–441.
Kieber JJ and Schaller GE (2010) The perception of cytokinin: a story
50 years in the making. Plant Physiology 154: 487–492.
Kim T-W and Wang Z-Y (2010) Brassinosteroid signal transduction
from receptor kinases to transcription factors. Annual Review of
Plant Biology 61: 681–704.
Lanahan MB, Yen HC, Giovannoni JJ and Klee HJ (1994) The never
ripe mutation blocks ethylene perception in tomato. Plant Cell 6:
521–530.
Lumba S, Cutler S and McCourt P (2010) Plant nuclear hormone
receptors: a role for small molecules in protein-protein interac-
tions. Annual Review of Cell and Developmental Biology 26:
445–469.
Melotto M, Underwood W, Koczan J, Nomura K and He SY (2006)
Plant stomata function in innate immunity against bacterial inva-
sion. Cell 126: 969–980.
Mockaitis K and Estelle M (2008) Auxin receptors and plant devel-
opment: a new signaling paradigm. Annual Review of Cell and
Developmental Biology 24: 55–80.
Nambara E and Marion-Poll A (2005) Abscisic acid biosynthesis and
catabolism. Annual Review of Plant Biology 56: 165–185.
Park S-Y, Fung P, Nishimura N, et al. (2009) Abscisic acid inhibits
type 2C protein phosphatases via the PYR/PYL family of START
proteins. Science 324: 1068–1071.
Pauwels L, Barbero GF, Geerinck J, et al. (2010) NINJA connects
the co-repressor TOPLESS to jasmonate signalling. Nature 464:
788–791.
Peng J, Richards DE, Hartley NM, et al. (1999) “Green revolution”
genes encode mutant gibberellin response modulators. Nature 400:
256–261.
Peterson GE (1967) The discovery and development of 2, 4-D.
Agricultural History 41: 243–254.
Peterson FC, Burgie ES, Park SY, et al. (2010) Structural basis
for selective activation of ABA receptors. Nature Structural and
Molecular Biology 17: 1109–1113.
Pirrung MC, Bleecker AB, Inoue Y, et al. (2008) Ethylene recep-
tor antagonists: strained alkenes are necessary but not sufficient.
Chemistry and Biology 15: 313–321.
Qiao H, Shen Z, Huang S-SC, et al. (2012) Processing and subcellular
trafficking of ER-tethered EIN2 control response to ethylene gas.
Science 338: 390–393.
eLS © 2017, John Wiley & Sons, Ltd. www.els.net
Plant Hormones
Rodríguez FI, Esch JJ, Hall AE, et al. (1999) A copper cofactor for the
ethylene receptor ETR1 from Arabidopsis. Science 283: 996–998.
Ryan CA (2000) The systemin signaling pathway: differential activa-
tion of plant defensive genes. Biochimica et Biophysica Acta 1477:
112–121.
Shinohara H, Ogawa M, Sakagami Y and Matsubayashi Y (2007)
Identification of ligand binding site of phytosulfokine receptor by
on-column photoaffinity labeling. Journal of Biological Chemistry
282 (1): 124–131.
Tabata R, Sumida K, Yoshii T, et al. (2014) Perception of root-derived
peptides by shoot LRR-RKs mediates systemic N-demand signal-
ing. Science 346: 343–346.
Tan X, Calderon-Villalobos LIA, Sharon M, et al. (2007) Mechanism
of auxin perception by the TIR1 ubiquitin ligase. Nature 446:
640–645.
Tata JR (2005) One hundred years of hormones. EMBO Reports 6:
490–496.
Wang KL-C, Li H and Ecker JR (2002) Ethylene biosynthesis and
signaling networks. Plant Cell 14 (Suppl): S131–S151.
Wasternack C and Hause B (2013) Jasmonates: biosynthesis, per-
ception, signal transduction and action in plant stress response,
growth and development. An update to the 2007 review in Annals
of Botany. Annals of Botany 111: 1021–1058.
Weng J-K, Ye M, Li B and Noel JP (2016) Co-evolution of hormone
metabolism and signaling networks expands plant adaptive plastic-
ity. Cell 166: 881–893.
Woodward AW and Bartel B (2005) Auxin: regulation, action, and
interaction. Annals of Botany 95: 707–735.
Xi W, Liu C, Hou X and Yu H (2010) MOTHER OF FT AND
TFL1 regulates seed germination through a negative feedback
loop modulating ABA signaling in Arabidopsis. Plant Cell 22:
1733–1748.
Yamaguchi S, Sun T-P, Kawaide H and Kamiya Y (1998) The GA2
locus of Arabidopsis thaliana encodes ent-kaurene synthase of
gibberellin biosynthesis. Plant Physiology 116: 1271–1278.
Yamaguchi S (2008) Gibberellin metabolism and its regulation.
Annual Review of Plant Biology 59: 225–251.
Zhao Y (2010) Auxin biosynthesis and its role in plant development.
Annual Review of Plant Biology 61: 49.
Further Reading
Buchanan BB, Gruissem W and Jones RL (2015) Biochemistry and
Molecular Biology of Plants. Chichester, UK: John Wiley & Sons,
Ltd.,.
Cutler S and Bonetta D (2011) Plant Hormones: Methods and Pro-
tocols. New York, NY, USA: Humana Press.
McCourt P (1999) Genetic analysis of hormone signaling. Annual
Review of Plant Physiology and Plant Molecular Biology 50:
219–243.
Taiz L, Zeiger E, Møller IM and Murphy A (2015) Plant Physiology
and Development. Sunderland, MA, USA: Sinauer Associates,
Incorporated.
11