TRENDS in Plant Science Vol.7 No.

7 J uly 2002
http://plants.trends.com 1360-1385/02/$ see front matter 2002 Elsevier Science Ltd. All rights reserved. PII: S1360-1385(02)02295-1
309 Review
Stephan Clemens
Leibniz Institute of Plant
Biochemistry, Weinberg 3,
D-06120 Halle, Germany.
e-mail: sclemens@
ipb-halle.de
Michael G.Palmgren
Dept of Plant Biology,
The Royal Veterinary and
Agricultural University,
Thorvaldsensvej 40,
DK-1871 Frederiksberg C,
Denmark.
Ute Krmer
Max Planck Institute of
Molecular Plant
Physiology,
AmMhlenberg 1,
D-14476 Golm, Germany.
The concept of phytoremediation has evoked
considerable interest in plant metal accumulation [1].
Using hyperaccumulation as a means of cleaning up
metal-contaminated soil and water has been
proposed [2] based on well documented observations
that several plant species not only tolerate otherwise
toxic levels of certain metals in the soil but even
hyperaccumulate them in their shoots [3]. Plants
ideal for phytoremediation should posses multiple
traits. They must be fast growing, have high biomass,
deep roots, be easy to harvest and should tolerate and
accumulate a range of heavy metals in their aerial
and harvestable parts. To date, no plant has been
described that fulfils all these criteria. However, a
rapidly growing non-accumulator could be engineered
so that it achieves some of the properties of
hyperaccumulators. Recent progress in determining
the molecular basis for metal accumulation and
tolerance by hyperaccumulators has been significant,
and provides us with a strong scientific basis to
outline some strategies for achieving this goal.
Molecular understanding of plant metal
accumulation determinants has numerous
additional biotechnological implications. For instance,
health-threatening human deficiencies in trace metals
appear to be widespread in developing countries and
possibly worldwide [4]. Engineering Zn accumulation
in edible plant parts might help in enriching diets
for Zn. Conversely, most of the toxic non-essential
elements such as Cd enter the human body via
plant-derived material (i.e. food or tobacco smoke) [5].
Because significant differences in Cd accumulation
are known to occur between plant varieties, molecular
identification of mechanisms governing this process
could result in the development of useful markers,
or of crops with reduced toxic metal content. Finally,
because signal transduction cascades are hypothesized
to have recruited metal homeostasis factors [6], a
better understanding of plant metal handling might
also lead to new insights into other fundamental
aspects of plant physiology.
In this article, we are describing key aspects of our
current understanding of plant metal accumulation
determinants and selected engineering approaches to
enhance metal accumulation capacity. The main focus
is on Zn and Ni as the main hyperaccumulated metals,
on Cu as a particularly toxic micronutrient, and on
Cd, which is used as a model non-essential metal of
significant environmental relevance in many studies.
Metal homeostasis network
The existence of a complex metal homeostasis network
has only recently come into focus [7,8]. It is a common
characteristic of all life forms that some of the elements
present in the environment are accumulated and
others are rejected. The rates of accumulation are
necessarily governed by physiological requirements
rather than toxicity. Trace elements such as copper
and zinc pose a specific dilemma to organisms. Their
ions are essential for a vast number of metabolic
processes yet are potentially dangerous. Both Cu(I)
and Cu(II) ions show an extremely high affinity to
various organic molecules. The redox-activity of Cu,
which is exploited in many electron transfer reactions,
for example, in photosynthesis, can lead to the
generation of oxygen radicals. Among the divalent
cations present in biological systems, Cd
2+
ions,
together with Ni
2+
and Zn
2+
ions, rank behind copper
in their affinity for organic donors such as thiolates
and amines [9]. Zn
2+
, in contrast to Ni
2+
, is widely used
as a cofactor. Zinc ions are key structural or catalytic
components in DNAbinding proteins and hydrolytic
enzymes, respectively. However, uncontrolled binding
can result in the inactivation of proteins.
Thus, tight control and regulation of essential
metal accumulation are vital at the organismal as
well as at the cellular level. Metal requirements of
different organelles vary tremendously because most
Zn- or Cu-requiring proteins are located outside the
cytoplasm. Coordinate function of uptake, buffering,
translocation and storage processes is required to
maintain essential metal concentrations in various
tissues and compartments within the narrow
physiological limits. Furthermore, because control
of accumulation is imperfect, organisms have to
cope with exposure to unwanted elements. Alack of
specificity of uptake and distribution systems also
leads to the accumulation of metals such as Cd, which
is generally considered nonessential.
Some plants can hyperaccumulate metal ions that are toxic to virtually
all other organisms at low dosages.This trait could be used to clean up
metal-contaminated soils.Moreover,the accumulation of heavy metals by
plants determines both the micronutrient content and the toxic metal content
of our food.Complex interactions of transport and chelating activities control
the rates of metal uptake and storage.In recent years,several key steps have
been identified at the molecular level,enabling us to initiate transgenic
approaches to engineer the transition metal content of plants.
Published online: 12 J une 2002
A long way ahead:understanding and
engineering plant metal accumulation
Stephan Clemens,Michael G.Palmgren and Ute Krmer
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310 Review
TRENDS in Plant Science
Xylem transport (c)
Trafficking and sequestration (e)
Mobilization (a)
Uptake and sequestration (b)
Unloading and tissue distribution (d)
M
M
M
Xylem
parenchyma
Unloading
Ion
exchange
Phloem
Symplastic
loading
Transfer
cell
Apoplastic
passage?
M
n+
M
n+
M
M
M
M
Golgi
Nucleus
Vacuole
CW
Cytosolic
metalloenzyme
M
M
M
M
M
M
M M
M
M
M
Plasma
membrane
H
+
H
+
-ATPase M
n+
M
n+
M
n+
M
n+
M
H
+
CW
Plasma
membrane
Other
chelators?
Citric acid cycle
Root
epidermis
Malate
Citrate
Malate
Citrate
Anion channels
Soil
particle
Plasma
membrane
Cytosolic
chelators
Vacuole
Precipitates
CW
M
n+
M
n+
M
n+
M
M
M
M
M
M M M
M
M
M
M
M
M
M
M M
M
M
M
H
+
H
+
-ATPase
H
+
M
MM
M
n+
M
n+
Xylem
Cell wall
binding
Uptake from
the apoplast
Storage in
trichomes
Symplastic
passage
Afundamental aspect of transition metals is that
during their passage through the plant only minute
proportions, if any, are present as free hydrated ions.
Recent work has indicated that in yeast and bacterial
cells, virtually all the cellular Cu and Zn, respectively,
is bound to ligands and that any movement within the
cell involves exchange reactions between ligands [10,11].
This tight control over metal availability within the
cell requires the existence of specific distribution
systems for the trace elements. In the case of Cu,
some of the distributors the metallochaperones
have been identified in yeast, mammals and plants
[7]. For Zn, such molecules have yet to be identified.
Determinants of transition metal accumulation
Generally speaking, accumulation of a given metal is
a function of uptake capacity and intracellular binding
sites. In a multicellular organism, the situation is
complicated by tissue- and cell-specific differences and
also by intercellular transport. Figure 1 illustrates
the processes that are assumed to be influencing metal
accumulation rates in plants: mobilization and uptake
from the soil, compartmentation and sequestration
within the root, efficiency of xylem loading and
transport, distribution between metal sinks in the
aerial parts, sequestration and storage in leaf cells.
At every level, concentration and affinities of chelating
molecules, as well as the presence and selectivity of
transport activities, affect metal accumulation rates.
This tightly knit network of determinants of metal
accumulation makes the development of feasible
engineering approaches targeting the primary limiting
steps a big challenge. Important insights have been
obtained from the study of model metallophytes, often
in comparison to closely related, non-metallophyte
plants. The major model metallophytes being studied
are various Zn-, Ni- or Cd-hyperaccumulating ecotypes
of Thlaspi caerulescens, some Ni-hyperaccumulating
taxa of the genus Alyssum, and Zn-hyperaccumulating
populations of Arabidopsis halleri.
Mobilization
The elements essential for life are also among the
most abundant on Earth [9]. However, the actual
bioavailability of some metals is limited because of
low solubility in oxygenated water and strong binding
to soil particles. Iron, the most extreme example, is
mainly present as Fe(III), which forms insoluble
hydroxides. Availability of Zn, which is required in
similar quantities as Fe, is less restricted, whereas
the bioavailability of some of the target metals in
phytoremediation, particularly Pb, is limited.
With the notable exception of Fe, which is
solubilized by either reduction to Fe(II) or extrusion
of Fe(III)-chelating phytosiderophores [12], little is
known about active mobilization of trace elements
by plant roots. Possibly, mechanisms assisting in the
acquisition of phosphorus contribute to increasing
the bioavailability of certain micronutrients. Both the
acidification of the rhizosphere and the exudation
of carboxylates are considered potential targets for
enhancing metal accumulation. Because organic acid
secretion also protects plant roots fromAl toxicity [13],
transgenic plants were generated that express a
bacterial citrate synthase. These plants were reported
to show elevated citrate production and secretion,
which resulted in enhanced Al tolerance [14]. However,
these results have now been questioned [15]. Alternative
approaches to increase carboxylate secretion include
the engineering of phosphoenolpyruvate carboxylase,
pyruvate dikinase or ATP-citrate lyase expression.
Restricting the overproduction of organic acids to
certainroot zones by using specific promoters, as certain
plants such as Lupinus albus are known to do [16], could
significantly reduce the strain on plant resources.
The same applies for the proposed modulation of
plasma membrane H
+
-ATPase activity [17] to support
rhizosphere acidification. Regulated local increases
in metal availability would also help to circumvent
problems associated with applying chelators such
as EDTAto the soil. The risk of this technique, which
has been applied in the phytoremediation of lead
contaminations [18], is that metal-chelate complexes
are leached into the groundwater.
Clearly, root-colonizing bacteria, as well as
mycorrhiza, have a large impact on the availability
of heavy metals for plant uptake. For instance,
soil microorganisms significantly enhance
Zn accumulation in the shoot of the hyperaccumulator
Thlaspi caerulescens [19]. Consequently, specific
modifications of the rhizosphere could greatly enhance
metal accumulation.
Uptake and sequestration
Following mobilization, a metal has to be captured by
root cells. Metals are first bound by the cell wall, an
ion exchanger of comparatively low affinity and low
selectivity. Transport systems and intracellular
high-affinity binding sites then mediate and drive
uptake across the plasma membrane. Uptake of
metal ions is likely to take place through secondary
transporters such as channel proteins and/or
H
+
-coupled carrier proteins. The membrane
potential, which is negative on the inside of the
plasma membrane and might exceed 200 mV
in root epidermal cells [20], provides a strong
driving force for the uptake of cations through
secondary transporters.
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311 Review
Fig. 1. (left) Molecular mechanisms proposed to be involved in transition metal accumulation by
plants. (a) Metal ions are mobilized by secretion of chelators and by acidification of the rhizosphere.
(b) Uptake of hydrated metal ions or metal-chelate complexes is mediated by various uptake systems
residing in the plasma membrane. Inside the cell, metals are chelated and excess metal is sequestered
by transport into the vacuole. (c) From the roots, transition metals are transported to the shoot via the
xylem. Presumably, the larger portion reaches the xylem via the root symplast. Apoplastic passage
might occur at the root tip. Inside the xylem, metals are present as hydrated ions or as metal-chelate
complexes. (d) After reaching the apoplast of the leaf, metals are differentially captured by different
leaf cell types and move cell-to-cell through plasmodesmata. Storage appears to occur preferentially
in trichomes. (e) Uptake into the leaf cells again is catalyzed by various transporters [not depicted in (e)].
Intracellular distribution of essential transition metals (=trafficking) is mediated by specific
metallochaperones and transporters localized in endomembranes (please note that these processes
function in every cell). Abbreviations and symbols: CW, cell wall; M, metal; filled circles, chelators;
filled ovals, transporters; bean-shaped structures, metallochaperones.
Several cation transporters have been
identified in recent years with the use of molecular
techniques, largely owing to the complementation of
Saccharomyces cerevisiae mutants. Most of the
transporters thought to be involved in the uptake of
micronutrients are in the ZIP(ZRT, IRT-like protein)
and the Nramp (natural resistance-associated
macrophage protein) family [21,22]. For some of them,
expression in roots and up-regulation under deficiency
conditions indicate a role in uptake from the soil.
Direct evidence demonstrating the contribution
of a specific transporter to transition metal
acquisition is scarce.
From yeast studies, it is apparent that most of the
cation transporters show a rather broad substrate
range, enabling even non-essential metals such as Cd
to enter cells. IRT1 expression, for instance, rescues
Fe
2+
, Mn
2+
and Zn
2+
uptake-deficient S. cerevisiae
strains and causes Cd
2+
hypersensitivity [23,24].
The wheat Ca
2+
transporter LCT1 mediates uptake
of Na
+
and Cd
2+
in yeast [25,26]. However, for
phytoremediation, specificity and affinity of transport
systems have to be considered in relation to relative
abundance of different substrates. Non-target elements
such as Ca might out-compete target elements.
Also, little is known about the actual physiological
substrates for uptake. For example, the metals might
be taken up as hydrated ions and in metal-chelate
complexes. The recent molecular identification of
the first Fe-phytosiderophore transporter ys1 from
maize, and the finding that the Arabidopsis genome
contains several ys1-like sequences [27], should help
in addressing this question.
At least three different engineering approaches
to enhance metal uptake can be envisioned. The
number of uptake sites could be increased, the
specificity of uptake systems could be altered so that
competition by unwanted cations is reduced, and the
sequestration capacity could be enhanced by increasing
the number of intracellular high-affinity binding sites
or the rates of transport into organelles. Acorrelation
between uptake capacity and hyperaccumulation was
found for ZIPfamily members in Thlaspi caerulescens.
Under Zn-replete conditions, ZNT1 and ZNT2 are
expressed at significantly higher levels in roots of
different T. caerulescens accessions than in roots of
the non-hyperaccumulating relative Thlaspi arvense
[28,29]. This observation fits well with the
four-to-fivefold higher Zn
2+
uptake measured for
T. caerulescens roots [30]. Thus, overexpression of
uptake systems might result in enhanced accumulation
of transition metals. Substrate specificity can be
engineered. By substituting various conserved
residues in ZIPfamily transporters with alanine
produces mutant versions of IRT1 that apparently no
longer transport Fe
2+
and Mn
2+
yet retain Zn
2+
and
Cd
2+
transport activity [24].
Intracellular binding and sequestration drive the
passage of transition metal ions across the plasma
membrane. Several processes known to contribute to
metal tolerance are associated with metal accumulation
at the same time. S. cerevisiae cells synthesizing
phytochelatins, glutathione-derived metal-binding
peptides, show significantly higher Cd
2+
tolerance
and increased Cd accumulation even at subtoxic
concentrations [31]. Phytochelatin synthesis is the
principal response of plants and many fungi to toxic
metal exposure, and apparently also of certain
animals [3234]. Phytochelatin deficiency is correlated
with Cd, Cu and As hypersensitivity. Therefore,
numerous attempts have been made to boost
phytochelatin formation by overexpressing enzymes
involved in the synthesis of the phytochelatin
precursor glutathione. In some cases, this led to a
slight enhancement of Cd tolerance and accumulation
[35], whereas in others it did not [36]. In spite of the
recent cloning of phytochelatin synthase genes (PCS)
[31,37,38], the effects of PCS overexpression have not
been reported in plants. It remains to be determined
which step limits the rate of phytochelatin formation
and whether engineering phytochelatin synthesis
can indeed be exploited to enhance tolerance and
accumulation. The best target might be the transport
of phytochelatin-metal complexes into the vacuole,
which in fission yeast is mediated by an ABC-type
transporter, Hmt1 [39]. However, in plants the
corresponding proteins have yet to be identified.
Other potential mediators of metal sequestration
and accumulation include members of the
Cation Diffusion Facilitator family (CDF). In
eukaryotic systems, they have been implicated in
moving Zn, Cd and Co from the cytosol into
cellular compartments [40]. Studies on various
hyperaccumulators and their respective close relatives
show a possible correlation between differences in
either expression levels or substrate specificities of
CDFs and the hyperaccumulation phenotype [29,41].
Arabidopsis plants overexpressing the transporter
ZAT become more Zn
2+
tolerant and accumulate
more Zn [42]. Thus, engineering metal-tolerance
mechanisms, such as the formation of metal-chelating
molecules or transporter-mediated vacuolar
sequestration, might result in enhanced accumulation.
Xylem transport
The apoplast continuum of the root epidermis and
cortex is readily permeable for solutes. The cell walls
of the endodermal cell layer act as a barrier for
apoplastic diffusion into the vascular system. In
general, solutes have to be taken up into the root
symplasm before they can enter the xylem [43].
Subsequent to metal uptake into the root symplasm,
three processes govern the movement of metals from
the root into the xylem: sequestration of metals inside
root cells, symplastic transport into the stele and
release into the xylem. The transport of ions into the
xylem is generally a tightly controlled process
mediated by membrane transport proteins [44].
The metal transporters involved in xylem loading
have yet to be identified.
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Inside the root, the presence of elevated amounts of
exogenously supplied histidine, which is able to chelate
Ni
2+
, results in a 50-fold increase in the rate of transport
of Ni into the xylem of Alyssummontanum[45]. In
agreement with this, several Ni hyperaccumulators
respond to Ni
2+
exposure by a large dose-dependent
increase in histidine concentrations in the xylem.
Thus, chelation with certain ligands, for example
histidine, nicotianamine [46] and citrate [47],
appears to route metals primarily to the xylem.
By contrast, chelation with other ligands, such as
phytochelatins or metallothioneins, might route
metals predominantly to root sequestration [48].
Inside the xylem, a pH-dependent equilibrium
exists between low-molecular weight chelators, free
hydrated metal cations and metal chelates in the
mobile transpiration stream, and stationary metal-
binding sites in the cell wall material surrounding the
xylem vessels [49]. Xylem-unloading processes are
the first step in controlled distribution and
detoxification of metals in the shoot, as well as in a
possible re-distribution of metals via the phloem [50].
Apromising engineering approach aimed at
enhancing xylem transport rates of metals is to
increase the steady-state concentrations of a chelator
that promotes root-to-shoot mobility of a metal.
With this aim, transgenic plants have been generated
that contain elevated levels of wild-type or mutant
ATPphosphoribosyltransferases, catalyzing the
first rate-limiting step in the histidine biosynthetic
pathway (U. Krmer et al., unpublished). Because
an elevated histidine content, whether of exogenous
or endogenous origin, is known to increase
Ni tolerance [45], this approach targets at least
two of the determinants of plant metal accumulation.
The identification of novel proteins involved in
root-to-shoot transport of metals or its regulation
should provide researchers with novel targets for
engineering metal accumulation.
Unloading, trafficking and storage
Transition metals reach the apoplast of leaves in the
xylem sap, from where they have to be scavenged by
leaf cells [12]. Transporters mediate uptake into the
symplast, and distribution within the leaf occurs
via the apoplast or the symplast [51]. Trafficking of
metals occurs inside every plant cell, maintaining the
concentrations within the specific physiological ranges
in each organelle and ensuring delivery of metals to
metal-requiring proteins. Metallochaperones, such
as CCH1, [52] and pumps, such as P-type ATPases,
are probably involved in these processes. For instance,
it has been suggested that RAN1 pumps Cu into
post-Golgi vesicles and is required for Cu delivery to
the ethylene receptor and other proteins [53,54].
In the Arabidopsis genome, there are several
additional P-type ATPase genes encoding pumps of
unknown specificity and unknown subcellular
localization. Some of these resemble bacterial
pumps transporting heavy metals such as Ag, Zn,
Co, Cd and Pb [55].
Excess essential metals, as well as non-essential
metals, are sequestered in leaf cell vacuoles [56,57].
Intriguingly, different leaf cell types show pronounced
differential accumulation. The distribution pattern
varies with plant species and element. Zn accumulation
in T. caerulescens leaves is 5.06.5-fold higher in
epidermis cells than in mesophyll cells [58], whereas
in metal-treated A. halleri, the mesophyll cells are
thought to contain more Zn and Cd than the epidermal
cells [59]. Furthermore, trichomes apparently play
a major role in storage and detoxification of metals.
For the phytoremediation candidate species
Brassica juncea, Cd accumulation was reported
to be >40-fold higher in trichomes compared with
the leaf total [60]. Asignificant proportion of
Ni and Zn are found in trichomes of the
hyperaccumulators Alyssum lesbiacum[61] and
Arabidopsis halleri [58], respectively.
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TRENDS in Plant Science
Storage and
detoxification
Symplastic
passage
Apoplastic
passage
Xylem
transport
Phloem
transport
Use
Use
Uptake
Uptake Unloading
Xylem
loading
(c)
Storage and
detoxification Uptake
Remobilization
Mobilization
Symplastic
passage
(a) (b)
(d)
Fig. 2. The path of a transition metal from the soil to the sites of use and storage in the leaf. The
different processes governing metal accumulation rates are depicted in Fig. 1 and described in the
text. Here they are represented by arrows. Red arrows highlight current engineering targets. These
include attempts (a) to enhance mobilization by secretion of organic acids, (b) to increase uptake by
overexpression or deregulation of transporters, (c) to stimulate uptake into the root and translocation
via the xylem by overproduction of intracellular chelators, or (d) to increase the strength of metal sinks
in the leaves by overexpression of storage and detoxification mechanisms. Processes that, according
to our present knowledge, show distinct molecular characteristics in hyperaccumulator plants are
represented by arrows of varying size. A thicker arrow indicates a process displaying higher activity in
metallophytes as compared to non-metallophytes, for example, rates of uptake into the roots via
plasma membrane-localized Zn
2+
transporters. A thinner arrow indicates a process displaying
reduced activity in metallophytes. This applies to sequestration of metals in root tissue.
Molecularly, the factors governing differential
metal accumulation and storage are unknown. It is
likely that vacuolar uptake is important. Therefore,
engineering vacuolar transporters, preferably in
specific cell types, might result in significantly
increased accumulation rates. Alternatively,
creating artificial metal sinks in the shoot by
enhancing the number of metal-binding sites, via
expression of cell wall proteins with high-affinity
metal-binding sites for instance, might suggest
to the plant metal homeostatic network an increased
metal demand in the shoot, resulting in enhanced
root-to-shoot translocation. Also, there might
be a need to concomitantly repress metal sinks
in the roots, for example, by decreasing transport
into root vacuoles.
There are numerous checkpoints along the way
(Fig. 2) as a transition metal makes its way from the
soil to the storage sites in the leaf. We now have some
molecular understanding of the key processes in
metal homeostasis in general, and of the phenotype of
metallophytes in particular. We are beginning to
understand the trafficking of metals at the cellular level
and the roles of increased uptake rates and chelator
synthesis in hyperaccumulation. However, many steps
of the way remain enigmatic. The transport and storage
forms of transition metals are largely unknown. Many
transport systems have yet to be analyzed with respect
to mechanism, localization, function and structure.
Key components, such as proteins involved in xylem
loading and the differential distribution between leaf
cells, have not been identified.
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314 Review
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Acknowledgements
We thank Nina Thue
Rasmussen for figure
preparation. Research in
our laboratories on the
engineering of plant
metal accumulation is in
part funded by an
EU framework V grant
(http://biobase.dk/~palmg
ren/metallophytes.html).
TRENDS in Plant Science Vol.7 No.7 J uly 2002
http://plants.trends.com 1360-1385/02/$ see front matter 2002 Elsevier Science Ltd. All rights reserved. PII: S1360-1385(02)02290-2
315 Review
Elizabeth Ble
Laboratoire des
Phytooxylipines,
IBMP-CNRS-UPR 2357,
28 Rue Goethe,
67083 Strasbourg Cedex,
France.
e-mail: elizabeth.blee@
ibmp-ulp.u-strasbg.fr
Intriguing analogies can be found between the
biosynthesis and the functions of oxylipins in
vertebrates and plants. Oxylipins are produced
during the metabolism of unsaturated fatty acids via
their oxidation by one, two or four atoms of oxygen
catalyzed by cytochrome P450, lipoxygenase (LOX)
and cyclo-oxygenase-like activities, respectively
(Fig. 1). In mammals, oxylipins are formed mainly
via the arachidonic acid cascade (a C20 fatty acid);
they play a major role in inflammatory processes
and, in general, in stress responses to infection,
allergy and exposure to food, drugs and
environmental xenobiotics.
Plants are devoid of the intricate protective
mechanisms provided by the immune system in
animals. Nevertheless, they have evolved an efficient
armory of defense systems against attacks by
pathogens or other life-threatening events.
These means of defense can be constitutive and/or
inducible, and include physical barriers and
production of toxic compounds. Phyto-oxylipins are
believed to play a pivotal role in these defenses
because they act as signal molecules and/or
protective compounds such as antibacterial and
wound-healing agents, or as constituents of cutin
(the framework of the cuticle that protects all the
aerial parts of a plant from its environment). Plants
can respond in different ways to the various stimuli
that result in tissue damage caused by abiotic factors
or pathogens. These stress responses usually include
the production of specific oxylipins, which have many
different biological functions. Therefore, knowledge
of the processes that regulate the biosynthesis and
localization of such bioactive lipid derivatives in
plant cells is crucial to the understanding of the
biology of this still-growing family of metabolites.
Recent studies, including the characterization of new
catalytic activities and the cloning of some important
biosynthetic enzymes, have provided new insights
into these processes. Here, these recent findings
are reviewed and the possible regulation of the
phyto-oxylipins biosynthetic pathways during stress
conditions is discussed.
Phyto-oxylipins are metabolites produced in plants by the oxidative
transformation of unsaturated fatty acids via a series of diverging metabolic
pathways.Biochemical dissection and genetic approaches have provided
compelling evidence that these oxygenated derivatives actively participate
in plant defense mechanisms.During the past decade,interest in this
field was focused on the biosynthesis of jasmonic acid (one branch of
C18 polyunsaturated fatty acid metabolism) and on its relationship to the
other plant defense-signaling pathways.However,recently,antisense
strategies have revealed that oxylipins other than jasmonates are probably
also essential for the resistance of plants to pathogens.
Published online: 12 J une 2002
Impact of phyto-oxylipins in plant
defense
Elizabeth Ble
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