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REVIEWS OPEN ACCESS

ROLE OF IRON IN PLANT GROWTH AND


METABOLISM
Gyana R. Rout and Sunita Sahoo
Department of Agricultural Biotechnology, College of Agriculture, Orissa University of Agriculture & Technology, Bhubaneswar 751 003,
Odisha, India

ABSTRACT
Iron is an essential micronutrient for almost all living organisms because of it plays critical role in metabolic processes such
as DNA synthesis, respiration, and photosynthesis. Further, many metabolic pathways are activated by iron, and it is a prosthetic
group constituent of many enzymes. An imbalance between the solubility of iron in soil and the demand for iron by the plant are
the primary causes of iron chlorosis. Although abundant in most well-aerated soils, the biological activity of iron is low because it
primarily forms highly insoluble ferric compounds at neutral pH levels. Iron plays a significant role in various physiological and
biochemical pathways in plants. It serves as a component of many vital enzymes such as cytochromes of the electron transport
chain, and it is thus required for a wide range of biological functions. In plants, iron is involved in the synthesis of chlorophyll, and
it is essential for the maintenance of chloroplast structure and function. There are seven transgenic approaches and combinations,
which can be used to increase the concentration of iron in rice seeds. The first approach involves enhancing iron accumulation in
rice seeds by expressing the ferritin gene under the control of endosperm-specific promoters. The second approach is to increase
iron concentrations in rice through overexpression of the nicotianamine synthase gene (NAS). Nicotianamine, which is a chelator
of metal cations, such as Fe+2 and zinc (Zn+2), is biosynthesized from methionine via S-adenosyl methionine synthase. The third
approach is to increase iron concentrations in rice and to enhance iron influx to seeds by expressing the Fe+2- nicotianamine
transporter gene OsYSL2. The fourth approach to iron biofortification involves enhancing iron uptake and translocation by
introducing genes responsible for biosynthesis of mugineic acid family phytosiderophores (MAs). The fifth approach to enhance
iron uptake from soil is the over expression of the OsIRT1 or OsYSL15 iron transporter genes. The sixth approach to enhanced iron
uptake and translocation is overexpression of the iron homeostasis-related transcription factor OsIRO2. OsIRO2 is responsible for
the regulation of key genes involved in MAs-related iron uptake. The seventh approach to enhanced iron translocation from flag
leaves to seeds utilizes the knockdown of the vacuolar iron transporter gene OsVIT1 or OsVIT2. The present review discusses iron
toxicity in plants with regard to plant growth and metabolism, metal interaction, iron-acquisition mechanisms, biofortification of
iron, plant-iron homeostasis, gene function in crop improvement, and micronutrient interactions.
Keywords:Iron homeostasis, Iron biofortification, Iron tolerance, Iron toxicity, Micronutrient interaction, Plant metabolism

Introduction and the platinum group elements (Farlex, 2005). Fifty-three of


Heavy metals are environmental pollutants, and their toxicity the 90 naturally occurring elements are considered heavy metals
is a problem of increasing significance for ecological, nutritional, (Weast, 1984), but only few are of biological importance. Based
evolutionary, and environmental reasons. The term heavy metal on their solubility under physiological conditions, 17 heavy metals
refers to any metallic element with has a relatively high specific may be available to living cells, and they could be significant
gravity (typically five times heavier than water) and is often toxic in plant and animal communities within different ecosystems
or poisonous even at low concentrations. This group of heavy (Weast, 1984). Some of the heavy metals (Fe, Cu, and Zn) are
metals includes lead (Pb), cadmium (Cd), nickel (Ni), cobalt (Co), known to be essential for plants and animals (Wintz et al., 2002).
iron (Fe), zinc (Zn), chromium (Cr), arsenic (As), silver (Ag), Other heavy metals such as Cu, Zn, Fe, Mn, Mo, Ni, and Co are

Received: January 7, 2015. Accepted: March 1, 2015. Published on line: May 19, 2015. 1
Correspondence to G.R.R: grrout@rediffmail.com
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essential micronutrients (Reeves and Baker, 2000), excess uptake al., 1997), membrane function (Quariti et al., 1997; Azevedo et al.,
of which by plants results in toxic effects (Blaylock and Huang, 2005), water relations (Kastori et al., 1992), and seed germination
2000; Monni et al., 2000 ) ; these are also referred to as trace (Iqbal and Siddiqui, 1992; Al-Hellal, 1995). The present review
elements due to their presence in trace or in ultra-trace (1 µg kg-1 highlights the physiology of plant growth in the context of iron
or µg L-1) quantities in environmental matrices. Plants experience toxicity, interactions with other micronutrients, interactions with
oxidative stress upon exposure to heavy metals that leads to other heavy metals, and molecular mechanisms of iron toxicity,
cellular damage and disturbance of cellular ionic homeostasis. membrane transport, plant iron homeostasis, iron biofortification,
To minimize the detrimental effects of exposure to heavy metals and genes responsible for iron tolerance.
and their accumulation, plants have evolved detoxification
mechanisms that are mainly based on chelation and subcellular Effects of Iron on Plant Growth
compartmentalization. Phytochelatins (PCs) are a principal class Iron is the third most limiting nutrient for plant growth and
of heavy metal chelators in plants, which are synthesized without metabolism, primarily due to the low solubility of the oxidized
translation from reduced glutathione (GSH) in a transpeptidation ferric form in aerobic environments ( Zuo and Zhang, 2011;
reaction catalyzed by the enzyme phytochelatin synthase (PCS). Samaranayke et al., 2012). Iron deficiency is a common nutritional
Therefore, the availability of glutathione is essential for PCs disorder in many crop plants, resulting in poor yields and reduced
synthesis in plants, at least during their exposure to heavy nutritional quality. In plants, iron is involved in chlorophyll
metals (Yadav, 2010). Iron (Fe) is an essential micronutrient for synthesis, and it is essential for the maintenance of chloroplast
plants whose deficiency presents a major worldwide agricultural structure and function. Being the fourth most abundant element
problem. Moreover, iron is not easily available in neutral in the lithosphere, iron is generally present at high quantities
to alkaline soils, rendering plants iron-deficient despite its in soils; however, its bioavailability in aerobic and neutral pH
abundance. Thirty percent of global cultivated soils are calcareous environments is limited. In aerobic soils, iron is predominantly
with low iron availability, because iron is present in insoluble found in the Fe+3 form, mainly as a constituent of oxyhydroxide
oxidized forms ( Guerinot and Yi, 1994; Mori, 1999 ) . Dicots polymers with extremely low solubility. In most cases, this form
and monocots have developed the Strategy I response (proton does not sufficiently meet plant needs. The visual symptoms of
extrusion to solubilize Fe+3 in the soil, reduction of the solubilized inadequate iron nutrition in higher plants are interveinal chlorosis
Fe+3 by a membrane-bound Fe+3 chelate reductase, and subsequent of young leaves and stunted root growth. In waterlogged soils, the
transport of the resulting Fe+2 into the plant root cell by an Fe+2 concentration of soluble iron may increase by several orders of
transporter) to iron deficiency stress (Römheld and Marschner, magnitude because of low redox potential (Schmidt, 1993). Under
1986; Marschner and Römheld, 1994). This response includes such conditions, iron may be taken up in excessive quantities.
acidification of the rhizosphere by releasing protons, subsequent However, it is potentially toxic and can promote the formation of
induction of Fe+3 chelate reductase activity that reduces Fe+3 to reactive oxygen-based radicals, which are able to damage vital
Fe+2, and acquisition of Fe+2 across the plasma membrane of root cellular constituents (e.g., membranes) by lipid peroxidation.
epidermal cells (Römheld, 1987). Bronzing (coalesced tissue necrosis), acidity, and/or blackening
Two types of causal relationships exist between the high of the roots are symptoms of plants exposed to above-optimal
concentration of heavy metals in soil and the expression of iron levels (Laan et al., 1991). Iron predominantly exists as Fe+3
toxicity symptoms. On one hand, heavy metals compete with chelate forms in the soil, and plants ultimately cannot absorb it
essential mineral nutrients for uptake, thereby disturbing the under various physiological conditions such as high soil pH in
mineral nutrition of plants (Clarkson and Luttge, 1989). On the alkaline soils. Thus, plants growing in high-pH soils are not very
other hand, after uptake by the plant, it accumulates in plant efficient at developing and stabilizing chlorophyll, resulting in the
tissues and cell compartments and ultimately hampers the general yellowing of leaves, poor growth, and reduced yield. However,
metabolism of the plant (Thurman and Collins, 1983; Taylor et plants have developed sophisticated mechanisms to take up
al., 1988; Turner, 1997). Heavy metal accumulation in plants has small amounts of soluble iron. Non-graminaceous plants release
multiple direct and indirect effects on plant growth, and it alters protons, secrete phenolics, reduce Fe+3, and take up iron (Römheld
many physiological functions ( Woolhouse, 1983 ) by forming and Marschner, 1983; Marschner, 1995a; Jeong and Guerinot,
complexes with O, N, and S ligands (Van Assche and Clijsters, 2009; Cesco et al., 2010). Once iron is solubilized, Fe+3 is reduced
1990). They interfere with mineral uptake (Yang et al., 1998; to Fe by a membrane-bound Fe+3 reductase oxidase (Jeong and
Zhang et al., 2002; Kim et al., 2003; Shukla et al., 2003; Drazic Connolly, 2009). Fe is then transported into the root by an iron-
et al., 2004; Adhikari et al., 2006), protein metabolism (Tamas et regulated transporter (IRT1). Ishimaru et al. (2011) reported that

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graminaceous plants rely on an Fe+3 chelation system during the through the action of the ribonucleotide reductase (Reichard, 1993).
secretion of mugineic acid (MA) family phytosiderophores. MAs It is also an active cofactor of many enzymes that are necessary
are secreted to the rhizosphere through TOM1, and they then for plant hormone synthesis, such as ethylene, lipoxygenase,
chelate Fe+3. In rice, the resulting Fe-MA complex is transported 1-aminocyclopropane acid-1-carboxylic oxidase ( Siedow,
by the yellow stripe family transporters (OsYSL15) (Nozoye et 1991), or abscisic acid (compounds that are active in many plant
al., 2011). Rice plants also have the ability to take up the iron development pathways and their adaptive responses to fluctuating
transporter (Ishimaru et al., 2006). Rout et al. (2014) screened environment conditions). Iron deficiency severely affects plant
51 varieties of upland and lowland rice using different levels of development and growth, and excess iron in cells is toxic. Iron
iron (0, 50, 100, and 200 mM) in nutrient solution to study the reactivity with reduced forms of oxygen produces radical elements
toxicity effect. Out of 51 varieties, 16 varieties were tolerant that can cause a loss of integrity and kill the cell. Therefore, there
(>200 mM Fe), 11 exhibited medium tolerance (<200 mM Fe), is an optimal window for the iron concentration to ensure smooth
and 24 varieties were susceptible (<100 mM) to selected iron plant development. Iron is a constituent of all redox systems in
concentrations. Total chlorophyll, total proline, total phenol, total plants, and the best known examples are enzyme systems (heme-
protein, and total carbohydrate content showed variation in both iron structure), including prosthetic groups like cytochromes.
tolerant and susceptible varieties. The oxidative enzymes also Iron plays a role in the porphyrin structure of chlorophyll, and is
showed variation among the tolerant and non-tolerant genotypes. therefore a principal component of chloroplasts. The best-known
Typically, approximately 80% of iron is found in photosynthetic functions of cytochrome are electron transport and the involvement
cells where it is essential for the biosynthesis of cytochromes of cytochrome oxidase in the terminal step in the respiration
and other heme molecules, including chlorophyll, the electron chain. Iron also interacts with non-heme proteins as an iron-sulfur
transport system, and the construction of Fe-S clusters (Briat et al., protein (e.g., ferredoxin, superoxide dismutase). Iron is a major
2007; Hansch and Mendel, 2009). In the photosynthetic apparatus, component of plant redox systems. Because of its physicochemical
two or three iron atoms are found in molecules directly related properties, especially its affinity to active metalloprotein sites, it
to photosystem II (PS-II), 12 atoms in photosystem I (PS-I), five acts as a cofactor in redox reactions that are necessary for oxygen
in the cytochrome complex, and two in the ferredoxin molecule production and use. However, its best-known function is its
(Varotto et al., 2002). Such distributions show that iron is directly structural role in the prosthetic groups of enzyme systems such as
involved in the photosynthetic activity of plants and, consequently, cytochromes, catalases, and peroxidases. These enzymes are also
their productivity (Briat et al., 2007). Iron availability is assumed to the main components of chloroplasts and mitochondria (Mengel
affect the natural distribution of species, and it may limit the growth and Kirby, 1987; Marschner, 1995b). The cytochrome essentially
of fast-growing economically important plants (Chen and Barak, acts as an electron carrier in the respiratory chain. Iron also interacts
1982). Iron can also be potentially toxic at high concentrations. with non-heminic proteins such as those in the iron sulfide group
The ability of iron to donate and accept electrons means that if iron (e.g., ferredoxin, superoxide dismutase). Moreover, iron is also
is free within the cell, it can catalyze the conversion of hydrogen active in protein synthesis (Bennet, 1945; Perur et al., 1961). Bashir
peroxide into free radicals. Free radicals can cause damage to a et al. (2011) identified and characterized the mitochondrial iron
wide variety of cellular structures, and they can ultimately kill transporter (MIT) in rice after screening 3993 mutant lines. The
the cell (Crichton et al., 2002). To prevent that kind of damage, identification of MIT is a significant advance in the field of plant
life forms have evolved a biochemical protection mechanism by iron nutrition, and it should facilitate the cloning of paralogs from
binding iron atoms to proteins. other plant species. Iron deficiency-induced chlorosis is a major
problem in plants, and it affects both yield and crop quality. Plants
Effects of Iron on Plant Metabolism have evolved multifaceted strategies, such as reductase activity,
When the insoluble ferric (Fe3+) form is reduced, it is converted protons, and specialized storage proteins to mobilize iron from the
to a ferrous form in the soil, and is then absorbed by plants. Even environment and to distribute it within the plant. The activation of
though iron is hardly present in living matter (50–100 µg·g-1 dry iron uptake reactions requires an overall adaptation of the primary
matter), it is still an essential element that is critical for plant metabolism because these activities need a constant supply of
life (Guerinot and Yi, 1994), as this element is involved in plant energetic substrates (i.e., NADPH and ATP). Vigani (2012) reported
metabolism. As a critical component of proteins and enzymes, that mitochondria are the energetic centers of the root cell, and they
iron plays a significant role in basic biological processes such as are strongly affected by iron deficiency. They observed that they
photosynthesis, chlorophyll synthesis, respiration, nitrogen fixation, display a high level of functional flexibility, which allows them to
uptake mechanisms (Kim and Rees, 1992), and DNA synthesis maintain the cell viability. Mitochondria contain a large amount

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of metalloproteins that require iron to carry out their function been demonstrated that photoprotection through excessive light
(Bertini and Rosato, 2007). In fact, several enzymes belonging dissipation as well as reversible photosynthesis might lead to
to both the respiratory chain and to the tricarboxylic acid cycle down-regulation and sustained photo inhibitory damage (Morales
are iron-containing proteins. Moreover, crucial steps of the Fe-S et al., 2000; Gogorcena et al., 2001; Donnini et al., 2003). Nenova
cluster assembly for the entire cell take place in the mitochondria, et al. (2006) found that in pea plants, the excess iron resulted in
suggesting an important role for this compartment in iron handling increased pigment concentrations of up to 28%. Some changes
by the cell. Iron deficiency and mitochondria are strongly linked in the pigment ratios were also observed, suggesting that their
because this stress greatly induces Strategy I activities that increased content could not only be attributed to concentration
demand considerable energy, thereby requiring the participation effects due to inhibited growth. Suh et al. ( 2002 ) also found
of mitochondria. Mitochondria represent a crucial target of studies evidence of photoinhibition in pea plants. The tendency of iron to
on plant homeostasis, and it might be of interest to concentrate form chelates and to undergo changes (Fe2+  Fe3++e-) are the two
future research on understanding how mitochondria orchestrate important characteristics underlying its numerous physiological
the reprogramming of root cell metabolism under iron deficiency. effects. The best-known function of iron is in the enzyme systems
Overall, root mitochondria are strongly affected by iron starvation, in which heme or hemin function as prosthetic groups (Mengel
because they require at least forty iron atoms per respiratory unit to and Kirkby, 1987 ) . Here, iron plays a somewhat similar role
function (Vigani et al., 2009). The ABC transporter STA1/AtATM3 to magnesium in the porphyrin structure of chlorophyll. These
has been characterized in Arabidopsis and implicated in the export heme enzyme systems include catalase, peroxidase, cytochrome
of Fe-S clusters (Kushnir et al., 2001). In addition, ferric-chelate oxidase, and various cytochromes, but the role of these enzymes
reductases, encoded by FRO genes in plants, might be involved in in plant metabolism is not completely understood. Research has
mitochondrial iron transport because AtFRO3 and AtFRO8, mainly also provided information about the function of cytochromes in
located in root and shoot veins, respectively, contain mitochondria- electron transport and the involvement of cytochrome oxidase in
targeting sequences (Mukherjee et al., 2006). This suggests that a the terminal step of the respiration chain. Catalase brings about the
reduction-based uptake could also occur in the invaginated inner breakdown of H2O2 to water and ½ O2. This enzyme also plays an
membrane of mitochondria. Moreover, the oxidizing conditions important role in chloroplasts (along with the enzyme superoxide
found in the mitochondrial intermembrane space ( IMS ) are dismutase), photorespiration, and the glycolytic pathway. Cell-
indicative of the need for a reduction-based mechanism (Hu et al., wall-bound peroxidases appear to catalyze reactions during
2008; Abadía et al., 2011). the polymerization of phenol to lignin, and peroxidase activity
Because of iron's redox properties and its ability to form seems to be particularly depressed in iron deficient roots. Cell-
complexes with diverse ligands, this element is a constituent wall formation and lignification were impaired, and phenolics
of many electron carriers and enzymes; therefore, it plays an accumulated in the rhizodermis of iron-deficient sunflower roots
important role in plant metabolism. On the other hand, low (Romheld and Marschner, 1981). Phenolics can be released into
solubility of inorganic iron at physiological pH levels and its the external solution, and, in the case of caffeic acid, may bring
high reactivity in the presence of oxygen, which generates toxic about chelation and reduction of inorganic Fe3+ (Olsen et al.,
hydroxyl radicals, represent a severe difficulty (Hell and Stephan, 1981). This reaction is not only favored by the accumulation of
2003). Soil conditions that result in insufficient or excess iron phenolics, but also by an increase in the reducing capacity of the
uptake are widespread in nature (Snowden and Wheeler, 1993; roots, which accompanies the impairment of lignin synthesis.
Schmidt and Fuhner, 1998; de la Guardia and Alcantara, 2002a). Barton ( 1970 ) observed large quantities of phytoferritin in
Chlorosis of young leaves is often the first visual sign of iron chloroplasts, which confirmed earlier evidence that chloroplasts are
deficiency. It is not only associated with the loss of chlorophyll, rich in iron, containing as much as 80% of the total iron in plants.
as several steps of its biosynthesis depend on iron, but also with Like heme iron, Fe-S proteins play a major role in oxidoreduction,
changes in the expression and assembly of other components of and both binuclear Fe-S clusters (2Fe-2S) and tetranuclear Fe-S
the photosynthetic apparatus (Terry and Abadia, 1986). On the clusters ( 4Fe-4S ) occur. Each cluster is surrounded by four
other hand, evidence exists that suggests that iron excess increases cysteine residues associated with a polypeptide chain (Sandmann
cytochrome b6/f complex in the thylakoid (Suh et al., 2002). In and Boger, 1983). These clusters are known as ferredoxins if they
recent years chlorophyll fluorescence analyses have been applied act exclusively as electron carriers and are characterized by a very
as rapid non-destructive tools to obtain information about the state high negative redox potential. Iron and chlorophyll concentrations
of photosynthetic apparatuses under iron deficiency or toxicity, are often well correlated in green plants. The same metabolic
and this is especially true regarding photosystem II (PS II). It has pathway involved in chlorophyll formation also operates during

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the biosynthesis of heme. Iron appears to control the rate of delta- ferrous form ( Chaney et al., 1972 ) . In anaerobic soils, high
aminolevulinic acid (ALA) formation, which is the precursor of concentrations of ferrous ions may lead to iron toxicity via
porphyrins (Miller et al., 1980). During iron deficiency, a decrease excessive iron uptake. Plants may limit iron uptake under those
occurred in the condensation rate of glycine and succinyl-CoA to conditions by the oxidation of ferrous ions with oxygen, which is
form ALA. In iron-deficient leaves, the photosynthetic apparatus transported from the shoot via aerenchyma (Foy et al., 1978). Iron
remained intact, but the number of photosynthetic units decreased in aerobic soils is mainly present as the ferric ion in precipitates
(Terry, 1980). Iron is also involved in protein metabolism. During (Lindsay et al., 1982) or in soluble chelates (Powell et al., 1980).
iron deficiency, the protein fraction decreases simultaneously with Chaney et al. (1972) reported that dicotyledonous plants might
an increase in the level of soluble organic N compounds. Nitric enhance their capacity for iron uptake, in response to a developing
oxide (NO) is a bioactive molecule that has recently emerged as a deficiency, by increasing their ability to reduce ferric chelates at
cellular messenger in numerous physiological processes in plants the root surface. Plants have evolved two separate mechanisms
(Neil et al., 2003). NO is biologically active at a concentration for the acquisition of iron, which can be referred to as Strategy I
of 1 nmol L -1, and it participates in signaling cascades that and Strategy II. These working hypotheses were first proposed by
drive plant growth and developmental processes (Beligni et al., R mheld and Marschner (1986). Sharma et al. (2013) reported that
2001). It is known that NO displays a high affinity towards, and the increase in the iron content of plants shows an enhancement
reacts with, transition metals. The three redox-related states of upon the treatment of rice plants with plant growth-promoting
biochemically distinguishable NO species, nitroxyl anion (NO-) rhizobacteria. They concluded that application of plant growth-
nitric oxide radical (NO), and nitrosodium cation (NO+), can promoting rhizobacteria strains was an important strategy to
react with the ferric (Fe3+) and/or ferrous (Fe2+) forms of iron combat the problem of iron deficiency in rice crops and humans.
(Stamler et al., 1996). Indeed, the high affinity of NO for iron is Iron is essential for various cellular events in plants, including
a well-characterized example of the study of compounds formed cellular respiration, chlorophyll synthesis, and photosynthetic
between metal ions and other neutral or charged molecules. electron transport. It also serves as a cofactor for a wide range
The complexes formed between iron and NO are called iron- of plant functions (e.g., cytochromes, catalase, and peroxidase
nitrosyl complexes (Stamler et al., 1992; Stamler et al., 1996), isozymes contain iron as a prosthetic group). In Fe-S proteins, iron
and these interactions are central in NO biochemistry. NO can is associated with cysteine, inorganic sulfur, or both (Marschner,
form iron-nitrosyl complexes in vivo with Fe-S and heme protein 1995b). Iron is also required for the synthesis and stabilization of
centers that are important for the biological activity of NO (Wink chlorophyll, which is why the chlorophyll content significantly
et al., 1998). The Fe-S cluster plays an important role in iron decreases in Fe-deficient plants. As a result, iron deficiency has
homeostasis because it can act as an iron sensor that regulates a marked effect on plant growth and product quality. Subcellular
protein expression associated with iron balance such as ferritin organelles, such as chloroplasts and mitochondria, depend on
and the transferring receptor in mammals (Beinert et al., 1999). In iron for their activities. Iron deficiency significantly affects the
addition, Fe-S proteins are viewed as carrier proteins that avoid function of mitochondria, and the disturbance in mitochondrial
the toxicity associated with free iron and allow its delivery at iron homeostasis adversely affects plant growth and development
lower intracellular concentrations (Mansy et al., 2004). The redox- ( Mori et al., 1991; Bashir et al., 2011 a, b, c; Bashir and
related NO species can have simultaneous effects on cellular Nishizawa, 2013; Vigani et al., 2013).
iron metabolism and homeostasis via mechanisms that might Iron is primarily absorbed by plants, and it solubilizes Fe3+ and
involve S-nitrosylation and/or the ligation of NO to Fe-S clusters then reduces it to Fe2+ for absorption or transport into the root. Tong
(Richardson et al., 1995). The uptake of iron, translocation, and et al. (1985) reported that both oxygen and photosynthetic substrates
regulation in higher plants has been reviewed (Kobayashi and are required for iron reduction by roots in apples (Rümheld and
Nishizawa, 2012 ) , and they reported that higher plants have Marschner, 1985). Clarkson and Sanderson (1978) observed that the
developed two distinct strategies to acquire slightly soluble iron root tips 1–4 cm from the apical zone absorbed and translocated
from the rhizosphere. Key molecular components, including more iron than the rest of the root. The roots of grapes under iron
transporters, enzymes, and chelators, have been clarified for both stress displayed enhanced proton release from approximately
strategies, and many of these components are now thought to also the same area (Korcak, 1987). R mheld and Marschner (1985,
function inside the plant to facilitate internal iron transport. 1986) proposed that iron uptake occurs by two species-dependent
strategies. Strategy I refer to iron mobilization by dicots and non-
Distribution of Iron in Plants gramineous monocots in response to iron deficiency stress, and
Iron uptake by plants is fastest when iron is present in the Strategy II refers to that found only with gramineous monocots. The

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chemotaxonomic boundary for the two strategies is not between bacterial and fungal siderophores, and they belong to a class of
dicotyledons and monocotyledons, but between higher plants compounds called mugineic acids (MAs) (Ma and Nomoto, 1996).
and grasses (Römheld, 1987). Plant species exhibiting Strategy I The biosynthetic pathway of MAs, starting from the condensation
show one or more of the following adaptive components: i) iron of three S-adenosyl-methionine molecules that form the precursor
deficiency-induced enhancement of Fe3+ reduction to Fe2+ at the nicotianamine (NA), is well established (Mori and Nishizawa, 1987;
root surface, with preferential uptake of Fe2+ (Chaney et al., 1972); Kawai et al., 1988; Shojima et al., 1990; Ma et al., 1995; Tkahashi
ii) H+ extrusion (Römheld and Marschner, 1984) that promotes et al., 1999; Kobayashi et al., 2001). The Fe (III)-phytosiderophore
the reduction Fe3+ to Fe2+; and iii) in certain cases the release of uptake transporter is called yellow stripe 1 (YS1), after the mutant
reducing and/or chelating substances by the roots. Enhanced Fe3+ phenotype of yellow striped leaves that result from the disruption
reduction under iron-deficiency, the most typical feature of Strategy of the gene encoding this transporter in maize (von Wiren et al.,
I, is characterized by the activation of reductases located on the 1994; Curie et al., 2001). YS1 proteins are distantly related to the
plasma membrane and presumably in the cell wall (apoplast) of oligopeptide transporter (OPT) family of transporters (Yen et al.,
the apical root zones (Bienfait et al., 1983). Bennett et al. (2011) 2001), and have been identified in several grass species (Curie et
reported that reduction of Fe+3 by root ferric chelate reductase al., 2001; Yen et al., 2001; Murata et al., 2006; Inoue et al., 2009;
further enhances iron solubility, since Fe+2 is more soluble than Lee et al., 2009). YS1 is a proton-coupled symporter of Fe (III)-
Fe+3. Reduction also prepares iron for uptake by IRT1 types (i.e., PS complexes (Schaff et al., 2004). The transport activity and
ferrous transporters), which move Fe+2 across the root epidermal specificity of YS1 transporters from maize ( ZmYS1 ) , barley
area to the plasma membrane. All three components of Strategy (HvYS1), and rice (OsYSL15) have been examined using yeast
I (proton pumping, ferric chelate reductase gene expression and growth complementation and radioactive uptake assays in two
enzyme activity, and IRT1 expression) increase markedly when electrode voltage clamp experiments in Xenopus leavis oocytes
plants are grown in iron-deficient conditions. Strategy I is used by (Curie et al., 2001; Roberts et al., 2004; Schaaf et al., 2004; Murata
most plant types, including the model plant Arabidopsis thaliana, et al., 2006; Inoue et al., 2009; Lee et al., 2009).
which facilitated the identification of key Strategy I genes such In the case of graminaceous plants, the plants secrete MAs
as ferric reductase oxidase (FRO2) (encodes root ferric chelate that bind to and solubilize Fe +3 ( Takagi, 1976; Takagi et al.,
reductase (Robinson et al., 1999)) and IRT1 (encodes the ferrous Fe 1984), and the resulting Fe (III)-MA complexes are readily taken
transporter (Eide et al., 1996). Strategy II systems are characterized up by the yellow stripe-like ( YSL ) family of transporters at
by an iron deficiency-induced release of specific Fe3+-chelating the root surface (Curie et al., 2001). MAs are synthesized from
compounds phytosiderophores (Takagi et al., 1984) and a high L-methionine (Shojima et al., 1990). NA synthase (NAS) converts
affinity uptake system (transport protein) for Fe3+ phytosiderophores three molecules of S-adenosyl methionine (SAM) to NA (Higuchi
(Römheld and Marschner, 1986). As a consequence, gramineous et al., 1994, 1995, 1999), which is then converted to a 3'-keto
species exhibiting these two components are highly efficient at intermediate by NA aminotransferase (NAAT) (Kanazawa et al.,
acquiring iron forms sparingly, including soluble inorganic Fe3+ 1993; Takahashi et al., 1999). This 3'-keto intermediate is then
(e.g., iron hydroxide). Chlorotic leaves might send a signal to the reduced to produce deoxymugineic acid (DMA) by the action of
roots inducing them to develop the reactions, but only the roots of DMA synthase (DMAS) (Bashir and Nishizawa, 2006; Bashir et
efficient varieties could pick up and interpret the chlorosis signal . al., 2006). This DMA is then released into the rhizosphere through
Certain plants, namely the grasses, that include most of the the MAs1 transporter (Nozoye et al., 2011, 2013). The genes of
world's staple grains, have evolved a distinct mechanism to acquire MA biosynthetic pathway have been cloned and characterized in
iron from the soil, which is known as Strategy II. This strategy is barley, rice, maize, and wheat (Takahashi et al., 1999; Nakanishi
best described as a chelation, which is similar to that used by et al., 2000; Kobayashi et al., 2001; Inoue et al., 2003, 2008;
many bacteria and fungi (Miethke and Marahiel, 2007), and it may Bashir et al., 2006; Bashir and Nishizawa, 2006). The expression
have arisen as an adaptation to alkaline soils where acidification of these genes is significantly upregulated by iron deficiency
of the rhizosphere is difficult to achieve (Ma and Nomoto, 1996). through transcriptional regulation that is mediated by various
Strong iron chelators called phytosiderophores (PS) are synthesized combinations of cis-acting elements and trans-acting factors
by the plant and secreted into the rhizosphere where they bind Fe+3 (Kobayashi et al., 2003, 2005, 2007, 2009, 2010, 2012; Ogo et al.,
(Tagaki, 1976; Tagaki et al., 1984, 1988). The Fe (III)-PS complex 2008, 2011; Kobayashi and Nishizawa, 2012), and the synthesis
is then taken up into root cells via transporters that are specific and secretion of MAs increases with limited iron availability. A
to the complex (Römheld and Marschner, 1986; von Wiren et diagrammatic representation of iron transport in the rice plant
al., 1994). Phytosiderophores are chemically quite distinct from is presented in Figure 1 (Bashir et al., 2013). Iron deficiency

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responsive elements 1 and 2 (IDE1 and IDE2, respectively) were (1980) reported that the absorption of nutrients by plant leaves
the first identified cis-acting elements controlling gene expression may be similar to that by roots, the main step being transport
in response to micronutrient deficiencies (Kobayashi et al., 2003). through the plasmalemma. As transport through the plasmalemma
Iron is translocated from roots to shoots as a ferric-citrate is an active process, the uptake rate of most plant nutrients is
chelate form, and this is transported to actively growing shoot influenced by the physiological status of the leaf (Mengel and
regions. Morris and Swanson (1981) reported that iron stress Kirkby, 1987). In leaf tissues, in contrast to the root, this active
did not change the redox potential of sap as compared to that uptake process is usually not the limiting step in ion uptake. The
of non-stressed trees. Since iron moves as a citrate complex, its availability of the micronutrient on the leaf surface is directly
stability during transport would be determined, in part, by the related to the water solubility of applied compounds. The
both pH and redox potential. Thus, under iron stress, movement precipitation for the crystallization of exogenous elements leads
as a citrate-chelate should not be affected. Translocation via to immobilization, and the higher retention of inorganic iron in
citrate in the stem exudates of four soybean genotypes was most epicuticular waxes may be connected with its low solubility. When
nearly related to the available iron supply (Brown et al., 1958). the solubility of Fe+2 extends over a wide pH range, this element
The transporters that are responsible for loading iron and citrate is readily oxidized to Fe3+, which tends to precipitate out in the
into the xylem have been recently revealed. Citrate appears to be form of hydroxide even at low pH levels. The formation of such
moved into the xylem, at least in part, via the multidrug and toxic an insoluble product would immobilize the element on the leaf
compound extrusion (MATE) family transporter ferric chelate surface. However, chelating improves iron solubility in water, and
reductase defective 3 (FRD3), which has a mutation that was this property can explain the lesser degree of superficial retention
responsible for the manganese accumulator (man1) phenotype and, consequently, the higher uptake with chelates. In the case
(Rogers and Guerinot, 2002a, b; Durrett et al., 2007). If high levels of pea plants, iron and zinc cross the cuticle more readily as free
of exogenous citrate are provided in the growth medium, iron ions (Ferrandon and Chamel, 1989), although an opposite result
accumulation in the root's vasculature is alleviated, and plants occurred with iron in citrus. The penetration of substances through
become phenotypically normal (Durrett et al., 2007). Kannan the cuticle is a diffusive process that is influenced by temperature

Fig.1. Iron transport in rice reported by Bashir et al. (2013).

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and concentration gradient. Water and solutes penetrate both Various disorders have been attributed in rice grown in iron-rich
stomatous and astomatous cuticles. Penetration through stomatous conditions (pH < 6) in which iron solubility can be very high (e.g.,
cuticles is usually more rapid than through astomatous cuticles, 6000 mg L-1 in flooded acid soils) (Martin, 1968). However, studies
but citrus cuticles show the opposite ( Basiouny and Biggs, have shown iron toxicity in some dry land plants in high pH
1976). Cuticles are 10–20 times more permeable to urea than to soils. Soluble organic-iron complexes, which may be available to
inorganic ions, and the foliar spray of nitrogen compounds can plants, might also maintain iron in solution (Marschner and Barber,
also enhance iron uptake. Regarding corn, the addition of urea 1975) in organic substrata, and this may partly account for the high
and ammonium nitrate increased the effectiveness of foliar- dissolved-iron concentration (approximately 20 mg L-1). Wheeler
applied FeSO4. Moreover, urea and ammonium nitrate had no (1985) observed that the iron accumulation in E. hirsutum shoots
effect on Fe-amino chelate, but they decreased the effectiveness and roots grown in iron-rich conditions point to the feasibility of
of Fe-EDTA. In tomatoes and white beans, the transport was not direct iron toxicity in this plant. Kuraev (1966) reported that the
as efficient, but it involved at least 25% of leaf-absorbed iron. initial toxic effect of high iron inhibits root development, and this
Translocation of foliar-applied iron may be enhanced by chelation was observed at higher iron concentrations (200 mg L-1), which
and by treatment with GA3 or kinetin (6-furfuryl amino purine). may have been due to possible toxicity mechanisms such as the
In corn, ferrous sulfate with chelating agents (EDTA or DTPA) iron-induced production of superoxide (O2-). Tanaka et al. (1966)
considerably increased the translocation of iron out of the treated reported that high iron concentrations may influence the growth
leaf (Ferrandon and Chamel, 1989). These authors suggested that and distribution of various wetland plant taxa. E. hirsutum roots
the metal cation is bound to its ligand within the plant, and that its also have some capacity that is clearly inadequate in high iron
high mobility might correspond to a lesser affinity of chelated iron environments. Many researchers have reported iron toxicity as a
for the negative sites on cell membranes and along vessels. problem in plants, particularly rice (Howeler 1973, Snowden and
Wheeler 1993). Furthermore, Batty and Younger (2003) examined
Biochemistry of Iron Toxicity in Plants the effects of total iron (added as FeSO4 7H2O) on an aquatic reed
Iron is an essential nutrient for plants. It accepts and donates species (Phragmites australis), and they found growth inhibition
electrons, and it plays important roles in the electron transport above a total iron concentration of 1 mg L-1. However, they believed
chains associated with photosynthesis and respiration. However, that the iron might not directly explain the reduction in growth.
iron is toxic when high levels accumulate. It can act catalytically Wang (1986) reported that the effective concentration (EC50) for
via the Fenton reaction to generate hydroxyl radicals, which can duckweed (Lemna minor) was 3.7 mg L-1, and Sinha et al. (1999)
damage lipids, proteins, and DNA. Therefore, plants must respond found that relatively low concentrations of iron (0.5–5.0 mg L-1)
to iron stress in terms of both iron deficiency and iron overload. resulted in damage to the membranes due to lipid oxidation in the
Plant species and growth media-related factors such as plant age, aquatic plant Hydrilla. Although iron is present in both Fe2+ and
hydrogen sulfide accumulation, organic acids, and other reduction Fe3+ forms in soils, it is only taken up by plants in Fe2+ forms. The
products also influence the occurrence of iron toxicity (De Datta et available iron content of tea soils varies from 50 to 100 mg kg-1.
al., 1994; Sharawat, 2004). Some researchers reported that seedlings Moreover, since oxygen diffuses in air 103 to 104 times faster than
may be more sensitive to toxic metals than mature plants (Hodgson, in water or water-saturated soils, oxygen is rapidly depleted by the
1972; Tadano, 1975). Moreover, Borggard (1983) observed that respiration of soil microorganisms and plant roots in waterlogged
iron-rich substrata (especially Fe3+ solids) can fix dissolved soils and continuously wetted soils. With the depletion in oxygen,
phosphorous via the formation of insoluble ferric phosphates or Fe3+ can act as electron acceptor for microbial respiration, and it is
by the adsorption of phosphate onto amorphous ferric solids. This subsequently reduced to Fe2+ (Becker and Asch, 2005). Rout et al.
might be expected to have a particularly adverse effect on Epilobium (2014) used different iron concentrations to screen 51 varieties of
hirsutum (a fen plant species), which has a high relative growth rate upland and low land rice varieties under nutrient culture conditions.
and is largely confined to nutrient-rich, productive habitats (Wheeler They observed that most of the varieties showed low germination
and Giller, 1982). High iron concentrations may also cause iron- rates, shoot and root tolerant indices, and leaf bronzing symptoms at
induced deficiencies of essential nutrients (e.g., Mn, P, K, Ca, 40 mM Fe. The total chlorophyll, proline, polyphenol content, total
and Mg) in plants (Howeler, 1973, Ottow et al., 1982). However, protein, and total carbohydrate content showed variation in both
shoot analyses provided little evidence for such deficiencies in E. tolerant and susceptible varieties.
hirsutum, and direct iron toxicity in a typical wetland plant like
E. hirsutum may seem a surprising possibility, especially in a high Effect of Iron on Plant Morphology
pH environment where iron solubility is likely to be rather low. Numerous visible effects associated with high iron concentrations

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were commonly observed, and some species showed several of the 1991).
following symptoms (Cook, 1990): growth retardation; reduction in Differential Iron Tolerance
leaf size; deepening of green leaf color (particularly in the youngest One approach used to overcome iron toxicity is the precise
leaves); reddening or purpling of stems and older leaves; wilting of study of its effects at the cellular level, and to select putative
shoots; yellowing and dieback of oldest leaves (especially from the tolerant cells that are screened in the presence of different iron
tips or margins); brown or black speckles or larger necrotic patches concentrations that are lethal to unselected cells (Mandal and
on leaves; blackening of leaf tips and stem bases; stiffening of Roy, 2000 ) . Metal tolerance screening for a large number of
stems; root stunting (particularly of adventitious roots); lack of root species demands a relatively rapid and reliable method (Baker
branching; root flaccidity; root blackening (particularly of the apices); and Walker, 1989). The germination response of plants potentially
and formation of precipitates on roots (Snowden and Wheeler, 1993). provides rapid screening for metal tolerance, but it has been found
Furthermore, Hemalatha and Venkatesan (2011) reported that tea to be unsatisfactory. An important mechanism, which excludes
plants that received higher concentrations of iron in soil developed many dry land plants from the wetland environment, is their
toxicity symptoms, and the bottom-most leaves turned coppery red inability to prevent the uptake of reduced toxins such as Fe2+ by
at later stages. They also reported that the upper most leaves of the oxidative precipitation (Martin, 1968). It seems likely that similar
tea plant established in soil containing above 500 mg kg-1 of rice mechanisms may also help explain differential tolerance of
grown in iron-rich conditions iron showed reddish brown spots on wetland plants to available iron (Wheeler et al., 1985; Cook 1990).
the upper epidermis. Albano and Miller (1993) reported that a specific Wheeler et al. (1983) showed that monocotyledonous species were
physiological disorder, bronze speckle, was consistently induced in usually more tolerant of iron than were dicotyledonous species.
First Lady and Voyager marigolds with Fe-DTPA concentrations Monocotyledonous shoots are frequently hollow, which facilitates
greater than 0.018 mM Fe-DTPA (1 ppm) applied to a soilless oxygen diffusion to the roots ( Etherington, 1983 ) . Justin and
medium. The disorder was characterized by specific symptomology Armstrong (1987) indicated that monocotyledonous species (and
that was visually distinguished by speckled patterns of chlorosis and particularly wetland ones) also have more porous root systems
necrosis as well as a downward curling and cupping of the leaves. The than dicotyledonous species. Although dicotyledonous species
percentage of total leaf dry weight affected with symptoms generally (and particularly wetland ones) seem more able to increase root
increased with increasing Fe-DTPA treatments. Symptomatic porosity upon flooding than monocotyledonous ones, they do
leaf tissue had a greater iron concentration than corresponding not attain the high values inherent in monocotyledonous roots.
asymptomatic leaf tissue. This disorder that affected African and Armstrong and Beckett (1987) pointed out that the many wetland
French marigolds was reported in the floriculture industries of the monocotyledonous species also have additional adaptations
United States and Canada (Biernbaum et al., 1988; Carlson, 1988; that help conserve oxygen supplies for apical consumption and
Halbrooks and Alabo, 1990; Albano and Miller, 1993). Furthermore, oxidative detoxification at the root tip, but these adaptations have
symptoms of this disorder include leaf chlorosis, bronze spotting not be found in dicotyledonous species. Hodgson (1972) observed
of the leaf, overall stunting, and delayed flowering (Biernbaum et that high concentrations of extractable manganese and aluminum
al., 1988). Symptoms are initially observed on older, mature leaves, were also frequently found in sites supporting iron-tolerant species,
and then progress to younger growth. Slight chlorotic speckling so it is possible that some plants have co-tolerance to these metals.
progresses to necrotic spotting (pitting), and necrosis of leaf margins The lack of correlation between tolerance and mean extractable
(Vetanovetz and Knauss, 1989). Affected leaf tissue has been reported potassium and magnesium concentrations is notable because there
to have iron and Mn concentrations of 400–2500 ppm, which is is considerable evidence that plants insufficiently supplied with
considered excessive (Biernbaum et al., 1988). Vetanovetz et al. (1989) either of these elements are prone to iron toxicity (Howler, 1973,
have identified this disorder as an iron toxicity of some floriculture Foy et al., 1978; Benckiser et al., 1984). Iron tolerance shows a
crops, including New Guinea impatiens, Sultana impatiens, cutting negative relationship with extractable phosphorus concentrations
and seed geraniums, vinca, and some species of Brassica. Similar and with soil fertility. The least-tolerant species tend to grow in
disorders have been observed in cabbage and tomato transplants as the more fertile sites and areas with higher extractable phosphorus
well as elatior begonias (Vetanovetz and Knauss, 1989). Disorders concentrations. This may be partly because many iron sensitive
attributed to iron toxicity include freckled leaves of sugarcane (Foy species have high relative growth rates (Snowden and Wheeler,
et al., 1978), the grey effect in tobacco (Arnold and Birns, 1987), and 1993 ) . The development of adventitious roots is sometimes
various unnamed disorders in other plants. Visual symptoms of iron regarded as an important adaptation to flood tolerance (Drew,
toxicity have been reported to include necrotic pitting, spotting or 1987 ) . Snowden and Wheeler ( 1993 ) reported iron tolerance
speckling, and bronzing (Foy et al., 1978; Albano and Halbrooks, screening for 43 British native fen plant species. They included

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23 dicotyledons and 20 monocotyledons, and they found evidence iron concentrations. This trend is due to the uniformity of pigment
of a clear relationship between the iron tolerance of a species and synthesis (Wilkinson and Ohki, 1988), and carotenoids also act
the nature of the root precipitate (Cook, 1990). Roy et al. (2003) as important metabolites that aid in protecting the cellular and
examined two rice cultivars IR72 (susceptible) and C14-8 subcellular systems from the cytotoxic effects of these reactive
(tolerant), and in vitro screening indicated the detrimental effects oxygen species (Agarwal et al., 2006) by quenching the singlet
of higher concentrations on plantlet regeneration. However, a few oxygen (Prasad and Bisht, 2011). Amylase is an enzyme that
calli survived on stressed media and regenerated plantlets. Cultivar participates in carbohydrate metabolism. Hofner (1970) found that
C14-8 showed a higher degree of tolerance than IR72 . Prominent amylase activity decreased with increased iron concentrations,
differences, including changes in enzyme activity with respect and this was due to the fact that iron forms complexes in plants
to iron-toxicity were noted. The results indicated that the activity with carbohydrate compounds such as maltose. The assimilation
of esterase, malate dehydrogenase, and lactate dehydrogenase of NH4+ into amino acids occurs via the joint action of glutamine
in C14-8 decreased in iron-stressed media. Decreased activity synthetase and glutamate synthase. Glutamate synthase catalyzes
indicates that the gradual degradation of these enzymes or their the transfer of an amide group from glutamine to alpha-
structural modification under increased iron toxicity levels. The ketoglutarate to yield two glutamate molecules. This reaction
activity of these enzymes increased in IR72 under stressed media. is agreed to be the primary route of nitrogen assimilation in
However, the activity of peroxidase remained almost unaltered in plants (Temple et al., 1998). When the activity decreased, the
C14-8 across the stress gradient. formation of amino acids also decreased, which led to lack of
nitrogen assimilation in plants. Hemalatha et al. (2011) found that
Role of Iron in Enzyme Activity in tea plants, the amylase invertase, aspartate amino transferase,
Iron is an essential mineral for plants that is required for and glutamate synthase were also drastically affected by the
biological redox systems (Asad and Rafique, 2000), and it is excess iron concentrations. They also reported that when iron
also a vital component of many enzymes that play important concentration decreased, the amount of pigments like chlorophyll
roles in the physiological and biochemical processes of plants. and carotenoid were reduced. The polyphenol content, which
It acts as a cofactor of key enzymes involved in plant hormone plays a vital role in tea quality, was also affected by the localized
synthesis and participates in numerous electron transfer reactions formation of a Fe-polyphenol complex. Bashir et al. (2007) studied
( Kerkeb and Connoly, 2006 ) . Plants are subjected to varying the expression patterns and enzyme activities of glutathione
differences in iron availability from the environment because of reductase (GR) in graminaceous plants under iron-sufficient and
their immobility. Therefore, either starvation or excess amounts iron-deficient conditions by isolating barely cDNA clones for
of this element are believed to generate oxidative stress (Abdel- chloroplastic GR (HvGR1) and cytosolic GR (HvGR2). They
Kader, 2007), which subsequently leads to several nutritional found that the GR1 and GR2 sequences were highly conserved
disorders that affect physiology of plant (Becker and Asch, 2005). in graminaceous plants, and based on their nucleotide sequences,
The toxicity of reactive oxygen species depends on the presence HvGR1 and HvGR2 were predicted to encode polypeptides of
of a Fenton's catalyst, such as iron or copper ions, which give rise 550 and 497 amino acids, respectively. Both proteins showed in
to extremely reactive OH- radicals in the presence of H2O2 and vitro GR activity, and the specific activity for HvGR1 was 3-fold
O2- (Chen and Schopfer, 1999). The reactive toxic oxygen species that of HvGR2. HvGR1, HvGR2, and TaGR2 were upregulated
cause damage to DNA proteins, lipids, chlorophyll, and almost in response to iron-deficiency ( Romero-Puertas et al., 2006 ) .
every other organic constituent of living cells (Becana et al., Moreover, HvGR1 and TaGR1 were mainly expressed in shoot
1998). Plants protect cellular and sub-cellular systems from the tissues, whereas HvGR2 and TaGR2 were primarily observed in
cytotoxic effects of these reactive oxygen species with antioxidant root tissues. The GR activity increased in roots of barley, wheat,
enzymes and metabolites such as carotenoids, ascorbic acid, etc. and maize and in the shoot tissues of rice, barley, and maize
(Alscher et al., 1997). Venkatesan et al. (2005, 2006) found that in response to iron-deficiency. Furthermore, the researchers
high concentrations of iron may result in the localized formation observed that GR was not post-transcriptionally regulated in
of iron polyphenol complexes. Monteiro and Winterbourn rice, wheat, and barley. Kaminaka et al., (1998) reported that
(1988) found that excess iron amounts must have catalyzed the OsGR1 expression was mainly observed in roots and calli, rather
generation of active O2 free radical species, which might have than in leaves. Hence, GR may play a role in the iron-deficiency
eventually oxidized the chlorophyll and subsequently led to response in graminaceous plants and in internal iron homeostasis.
decreased chlorophyll content. Arunachalam et al. (2009) reported Romero-Puertas et al., (2006) reported that the expression of
a decrease in chlorophyll and carotenoid content with increased PsGR1 and PsGR2 was differentially regulated in response to

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various stresses such as mechanical wounding and increased imbalances in cation ratios; poor soil aeration; certain organic
temperature. Lipton et al. (2001) reported that GSH may play an matter additions to soil; viruses; and root damage by nematodes
important role in signal transduction for the regulation of gene and other organisms (Wallace and Lunt, 1960; Embleton et al.,
expression in response to different stresses and iron homeostasis. 1973 ) . Lindsay and Schwab ( 1982 ) reported that olive plants
However, an S-nitrosylated form of GSH has been suggested to growing on calcareous soils faced characteristic problems
act as a transport molecule for NO, thereby increasing its half- regarding iron acquisition. The total iron content in these soils
life and allowing for effective biological activity (Lipton 2001). was high, but the fraction available for the plants was insufficient.
GSH is also essential for the activity of glutathione S-transferase, This was caused by the very low solubility of iron oxides at
which specifically interacts with the dinitrosyl-diglutathionyl-Fe alkaline pH conditions that were buffered by the presence of
complex, and it behaves like a storage protein for this complex bicarbonate in these soils. Further, Lopez-Jimenez (1985) reported
both in vivo and in vitro (Maria et al., 2003; Turella et al., 2003). that in young Avocado leaves, the chlorosis increase was related to
decreased iron content in leaves, chlorophyll, and catalase activity.
Iron Deficiency in Plants Iron deficiency is usually manifested as interveinal chlorosis of
Iron is an important component of enzymes that are involved young leaves while the veins remain green, hence termed iron
in the transfer of electrons (e.g., cytochromes and iron-sulfur deficiency chlorosis. The manifestation of the symptoms in
[Fe-S] proteins involved in redox reactions ) . Furthermore, it young leaves is due to the inability to redistribute iron within
is a constituent of non-heme iron proteins that are involved in the plant. However, iron may become more mobile under stress
photosynthesis, N2 fixation, and respiration (Taiz and Zeiger, 1991). conditions (Chaney et al., 1992). One of the best alternatives to
Iron limitation affects one-third of the cultivable land on Earth, prevent iron chlorosis problems is the use of tolerant plant species
and it represents a major concern for agriculture. Iron limitation or genotypes. Differences in tolerance have been found in many
causes the decline of many photosynthetic components, including herbaceous and woody species (Hamze et al., 1986; Kolesh et al.,
the Fe-S protein ferredoxin (Fd), which is involved in essential 1987; Chaney et al., 1992; Cianzio, 1995). Alcantar et al. (2003)
oxidoreductive pathways of chloroplasts (Tognetti et al., 2007). In observed that olive trees are widely grown on the calcareous
this role, it is reversibly oxidized from Fe2+ to Fe3+ during electron soils of the Mediterranean without generally showing problems
transfer. Characteristic symptoms of iron deficiency involve associated with iron chlorosis, and this species is considered
interveinal chlorosis, which is similar to Mg deficiency (Bienfait more tolerant than other fruit species such as peaches or quinces.
and Van der Mark, 1983). Iron-deprived plants usually develop Nonetheless, iron chlorosis has been observed in some olive
interveinal chlorotic symptoms in young leaves as well as poor orchards (Martar et al., 1977; Fernadez-Escobar et al., 1993).
root formation, and when severe, the deficiency leads to growth This could be related to different causes including the following:
retardation, stasis, and death (Kobayashi et al., 2003). Chlorosis location specific poor soil or environmental conditions; use of
has been attributed to the inhibition of chlorophyll synthesis, irrigation, fertilization, or higher plantation density aimed to
which requires the function of iron-containing enzymes (Reinboth increase production; or the introduction of less tolerant varieties.
et al., 2006). However, expression of chlorophyll-binding proteins Soil-based organisms have evolved various adaptive mechanisms
and other photosynthetic components is downregulated with to mitigate the consequences of an iron deficit, and most rely
relative independence of pigment levels (Thimm et al., 2001). on the optimization of metal uptake from scarcely available
Therefore, chloroplasts are primary targets of iron deficiency, sources. Dicotyledon plants and non-graminaceous monocots
resulting in decreased photosynthetic activity and plastidic display the so-called Strategy I response, which is characterized
pigments and proteins (Winder et al., 1995). Both the utilization by the development of root hairs and transfer cells, increased
of ribulose-1, 5-biphosphate by rubisco and its regeneration by proton extrusion to the rhizosphere to improve Fe3+ solubility, and
the Calvin cycle appear compromised in iron-starved plants enhanced accumulation of ferric-chelate reductases and broad
(Arulanathan et al., 1990; Winder et al., 1995). Iron deficiency range metal transporters in the root cell plasma membrane to favor
can occur in agricultural soils at both extremes of the pH range. Fe2+ intake (Eide et al., 1996; Robinson et al., 1999; Thomine
Moreover, several of the following factors contribute either et al., 2000; Bereczky et al., 2003; Mukherjee et al., 2006;). On
individually or in combination to the development of chlorosis: the other hand, grasses, have developed a different response in
low iron supply; calcium carbonate in soil; bicarbonate in soil which phytosiderophores of the mugineic acid family are released
or irrigation water; over irrigation or water logged conditions; into the surrounding soil (Mori, 1999). The screening for iron
high phosphate levels; high levels of heavy metals; low or high deficiency in cultivars using hydroponic systems with bicarbonate
temperatures; high light intensities; high levels of nitrate nitrogen; has been used for herbaceous (Pissaloux et al., 1995; Ahendawi et

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al., 1997; Campbell and Nishio, 2000; Lucena, 2000) and woody as Fe+2 and zinc (Zn+2), is biosynthesized from methionine via
species (Romera et al., 1991a, b; Shi et al., 1993; Sudahono et al., S-adenosyl methionine synthase (SAMS) and NAS (Higuchi et
1994; Cinelli et al., 1995; Alcantara et al., 2000; Nikolic et al., al., 1994). Hell and Stephan (2003) and Takahashi et al. (2003)
2000). Moreover, Pestana et al. (2004) studied the effects of iron reported that all higher plants synthesize and utilize nicotianamine
deficiency in three citrus rootstocks ( Troyer citrange, Taiwanica for the internal transport of iron and other metals. Rice has three
orange and Swingle citrumelo ) using different growth NAS genes: OsNAS1, OsNAS2, and OsNAS3. These genes are
parameters and physiological characteristics. They observed that differentially regulated by iron, and all of them are expressed in
Troyer citrange and Taiwanica orange plants had a similar cells involved in the long-distance transport of iron (Inoue et al.,
degree of tolerance to iron chlorosis, whereas the Swingle 2003). Masuda et al. (2013) suggested that, in addition to their
citrumelo rootstock was more sensitive. roles in phytosiderophore synthesis, NAS and nicotianamine play
important roles in the long-distance transport of iron in rice.
Iron Biofortification in Plants The third approach is to increase iron concentrations in
Iron is an essential micronutrient for plants and animals, rice and to enhance iron influx to seeds by expressing the Fe+2-
and iron deficiency is one of the most prevalent micronutrient nicotianamine transporter gene OsYSL2. Koike et al. ( 2004 )
deficiencies globally. It affects an estimated two billion people identified the rice OsYSL2 gene. This gene is preferentially
(Stoltzfus et al., 1998) and causes 0.8 million deaths annually expressed in leaf phloem cells, the vascular bundles of flowers,
worldwide ( WHO, 2002 ) . Iron deficiency is ranked sixth and developing seeds, which suggests a role in internal iron
among the risk factors for death and disability in developing transport. The iron concentration in OsYSL2 knockdown rice
countries with high mortality rates (WHO, 2002). Micronutrient was 18% lower in brown seeds and 39% lower in polished seeds
supplementation, food fortification, and biofortification are the compared to non-transgenic rice (Ishimaru et al., 2010). Masuda
three basic approaches used to alleviate micronutrient deficiencies. et al. (2013) hypothesized that enhanced expression of OsYSL2
Rice is a particularly suitable target for biofortification because could increase the iron concentration in rice seeds.
iron deficiency anemia is a serious problem in developing The fourth approach to iron biofortification involves enhancing
countries where rice is a major staple crop (WHO 2002; Juliano, iron uptake and translocation by introducing genes responsible
1993 ) . Biofortification can be applied to increase the iron for biosynthesis of MAs. The iron concentrations in the seeds of
concentrations in polished seeds and to achieve the target iron transgenic lines were analyzed after cultivation in the paddy field in
demand for human nutrition. Sperotto et al. (2012) suggested iron-sufficient or low (calcareous) soil (Masuda et al., 2008; Suzuki
conventional breeding or directed genetic modification as two et al., 2008). Three transgenic rice lines, which were transformed
possible ways to develop iron-rich rice plants. Iron concentrations with barley genome fragments involving mugineic acid synthase
in rice have a narrow variation, ranging from 6 to 22 g L-1 as genes (HvNAS1 or HvNAS1 and HvNAAT-A,-B or IDS3) were
compared to 10 to 160 g L-1 in maize and 15 to 360 g L-1 in wheat produced (Higuchi et al., 2001b; Kobayashi et al., 2001; Masuda et
(Gómez-Galera et al., 2010); however, most maize genotypes do al., 2008; Suzuki et al., 2008). Masuda et al. (2008) reported after
not reach the highest values (Ortiz-Monasterio et al., 2007). Based cultivation in iron-sufficient soil, the IDS3 rice lines produced iron
on available the literature, there is scanty information on the concentrations that were 1.4-fold and 1.3-fold higher in the polished
development of rice genotypes with iron-enriched grains through and brown seeds than in non-transgenic rice, respectively.
conventional breeding programs. However, the methodology is The fifth approach to enhance iron uptake from soil is the
well established and genetic variation exists. Masuda et al. (2013) over expression of the OsIRT1 or OsYSL15 iron transporter
reported seven transgenic approaches and combinations, which genes. Lee et al. (2009a) produced transgenic rice that expressed
can be used to increase the concentration of iron in rice seeds. The the rice ferric ion transporter gene OsIRT1 under the control of
first approach involves enhancing iron accumulation in rice seeds the ubiquitin promoter. This rice showed a 13% increase in iron
by expressing the ferritin gene under the control of endosperm- concentration in the brown seeds, while the iron concentration
specific promoters. The iron stored in soybean ferritin is readily in the leaves increased 1.7-fold. Lee et al. (2009a) reported that
absorbed by the human gastrointestinal tract (Lonnerdal, 2009), plants that over expressed OsIRT1 were shorter and had fewer
and iron stored in ferritin is an important source that humans can tillers. As constitutive expression of OsIRT1 may disturb metal
use to avoid iron deficiency (Theil et al., 2012). homeostasis in rice, they suggested that targeted expression
The second approach is to increase iron concentrations in of OsIRT1 using specific promoters might solve this problem.
rice through over expression of the nicotianamine synthase gene Masuda et al. (2013) concluded that OsIRT1 could be used to
(NAS). Nicotianamine, which is a chelator of metal cations, such enhance iron levels in rice grains. Lee et al. (2009b) reported that

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OsYSL15 over expression using the OsActin1 promoter increased varieties, and then identified genes related to iron homeostasis (e.g.,
the concentration of iron in brown seeds by approximately 1.3-fold OsYSLs, OsNASs, OsNRAMP1, OsIRT1, OsZIPs, and APRT)
compared with non-transgenic rice. Gómez Galera et al. (2012) as candidate genes that effect iron concentrations seeds. Sperotto
produced transgenic rice that overexpressed the barley Fe (III)- et al. (2010) analyzed the gene expression profiles of 25 metal-
MA transporter gene HvYS1 under the control of the CaMV35S related genes, including rice homologues of YSL2, NRAMPs,
promoter. The concentration of iron in the transgenic leaves was ZIPs, IRT1, VIT1, NASs, FROs, and NAC5, in eight rice varieties
1.5-fold higher than in the non-transgenic leaves. with different iron and zinc concentrations in the seeds. They also
The sixth approach to enhanced iron uptake and translocation identified putative target genes that contributed to increased iron
is over expression of the iron homeostasis-related transcription and zinc concentrations in rice grains. Xiong et al. (2014) indicated
factor OsIRO2. Ogo et al. (2006) identified an iron deficiency- that IRT1 is a key component of iron uptake from the soil in dicot
inducible basic helix-loop-helix ( bHLH ) transcription factor, plants. They introduced the peanut (Arachis hypogaea L.) AhIRT1
OsIRO2, in rice. OsIRO2 is responsible for the regulation of into tobacco and rice plants using an iron-deficiency-inducible
key genes involved in MAs-related iron uptake (e.g., OsNAS1, artificial promoter. Induced expression of AhIRT1 in tobacco plants
OsNAS2, OsNAAT1, OsDMAS1, TOM1, and OsYSL15) (Ogo et resulted in the accumulation of iron in young leaves under iron-
al. 2007; Ogo et al. 2011). Ogo et al. (2007) introduced OsIRO2 deficient conditions. Even under iron-excess conditions, the iron
under the control of the CaMV35S promoter into rice plants. concentration was also markedly enhanced, which suggests that
Plants with the overexpressed OsIRO2 transcription factor the iron status did not affect the uptake and translocation of iron by
secreted more DMA than non-transgenic rice, and they exhibited AhIRT1 in the transgenic plants. They observed that the transgenic
enhanced iron deficiency tolerance in calcareous soils (Ogo et al., tobacco plants showed improved tolerance to iron limitation in
2007; Ogo et al., 2011). Moreover, the iron concentration in the culture with two types of calcareous soils. The induced expression
transgenic brown seeds increased 3-fold when the transgenic rice of AhIRT1 in rice plants also resulted in high tolerance to low iron
was cultivated in calcareous soil (Ogo et al., 2011). Therefore, this availability in calcareous soils. Induction of AhIRT1 expression into
approach can be used to increase the iron concentrations of seeds tobacco plants also affected zinc and manganese concentrations in
in soils with low iron availability. both roots and young leaves of the transgenic plants as compared to
The seventh approach to enhanced iron translocation from non-transgenic plants during iron starvation. The results suggested
flag leaves to seeds utilizes the knockdown of the vacuolar iron that the accumulation of AhIRT1 in transgenic plants disturbed the
transporter gene OsVIT1 or OsVIT2. Kim et al. (2006) reported zinc, manganese, and copper uptake and/or translocation (Pedas et
that the Arabidopsis vacuolar iron transporter, VIT1, is highly al., 2008).
expressed in developing seeds, and it transports Fe and Mn into
the vacuole. Zhang et al. (2012) reported that disruption of the Interaction of Iron with Other Nutrients
rice VIT orthologs ( OsVIT1 and OsVIT2 ) increased the iron Other nutrients may not only play an important role in reducing
concentrations by 1.4-fold in brown seeds, and it decreased the the effects of iron toxicity but also in the expression of iron tolerance
iron concentrations by 0.8-fold in the source organ flag leaves by various rice cultivars (Sahrawat, 2004). Iron toxicity is a complex
because VIT genes are highly expressed in rice flag leaves. nutrient disorder, and the deficiencies of other nutrients (especially
Bashir et al. (2013) reported that an OsVIT2-knockdown mutant P, K, Ca, Mg, and Zn) are considered when instances of iron toxicity
showed 1.3-fold and 1.8-fold increases in iron concentrations in occur in rice (Ottow et al., 1983). Deficiencies of Ca, Mg, and Mn
the brown and polished rice seeds, respectively. They suggested are not commonly observed in lowland rice, except in acid sulfate
that disruption of OsVIT1 or OsVIT2 enhanced iron translocation soils. Therefore, P, K, and Zn deficiencies deserve special attention
between the source and sink organs, and proposed this as a novel (Yoshida, 1981). In aerobic conditions, iron in the soil is usually
strategy for producing iron-biofortified rice. Zhang et al. (2012) found as oxyhydroxide polymers with very low solubility, which
and Bashir et al. (2013) reported that the cadmium concentration limits the iron supply for plant uptake (especially in calcareous
was also increased following VIT knockdown in rice. Therefore, soils). Therefore, iron deficiency is a yield-limiting factor with major
this approach should be avoided in cadmium-contaminated soils. implications for field crop production in many agricultural regions
The mining of high iron rice varieties or the identification of of the world (Hansen et al., 2006). The expression of chlorosis may
novel target genes is important to the iron biofortification of rice. be due to the deficiency of iron interactions with micronutrients like
Anuradha et al. (2012) found seven quantitative trait loci (QTL) and Zn and Mn (Dixit and Yamdagni, 1983). Ferrihydrite is the inorganic
selection markers related to the iron concentrations in rice seeds. compound usually associated with iron accumulation in ferritin
They mapped the QTL using Madhukar × Swarna indica rice (Chasteen and Harrison, 1999). Ferritins are large proteins with a

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central cavity that can store up to 4500 iron atoms, which can be Genes Responsible for Crop Productivity
released when necessary (Yamashita, 2001). Therefore, these proteins Iron is an essential micronutrient, and the expression of genes
are believed to play a critical role in the cellular regulation of iron involved in iron transport is regulated by iron availability and
storage and homeostasis (Zancani et al., 2004), because expression of demand. Most previous studies have compared the expression of
some plant ferritin isoforms can be induced by iron overload (Fobis- genes in response to iron deficiency (Mori, 1999; Kobayashi and
Loisy et al., 1995; Petit et al., 2001). Chen et al. (1980) identified Nishizawa, 2012; Bashir et al., 2013). However, the expression of
both goethite (α-FeOOH) and lepidocrocite (γ-FeOOH) in rice genes involved in iron transport is also regulated in response to high
root coatings. Furthermore, they found that under iron-deficient iron demand at different developmental stages. Among the abiotic
conditions, phosphorous concentrations decreased in the roots of stresses, iron deficiency constitutes a major factor leading to a
rice cultivars and increased in the shoots of rice plants, which had reduction in crop yield due to the high pH, especially in calcareous
lower Zn concentrations in their roots (Murad and Schwertmann, soils with extremely low iron solubility (Kobayashi et al., 2005).
1984). Selviera et al. (2007) noted increased calcium concentrations They reported that the expression of genes encoding every step in
when the roots were iron-deficient. Several iron transporters are the methionine cycle was thoroughly induced by iron deficiency
able to transport iron along with Zn and other metals (Korshunova in roots, and almost thoroughly induced in the leaves. The rice
et al., 1999; Eckhardt et al., 2001; Gross et al., 2003; Lopez-Millan genes involved in iron acquisition are coordinately regulated by
et al., 2004). The induction of iron transporters under iron-deficient conserved mechanisms in response to iron deficiency, in which iron
conditions resulted in increased zinc uptake. Mn concentrations deficiency inducible (IDE)-mediated regulation plays a significant
decreased under iron deficiency in the roots and shoots of plants, role. A promoter search revealed that iron deficiency-induced
and Mn can be transported by the plant through iron transporters genes associated with iron uptake possessed sequences that were
(Eckhardt et al., 2001; Lopez-Millan et al., 2004). Iron deficiency homologous to the iron deficiency-responsive cis-acting elements
resulted in lower sulfur concentrations in the roots and shoots of plant IDE1 and IDE2 in their promoter regions, which were identified at
varieties, and this is probably a consequence of the iron requirements a higher rate than those showing no induction under iron deficiency.
for sulfur assimilation (Hell and Stephan, 2003). Furthermore, Cu Kobayashi et al. (2005) and Paz et al. (2007) reported that two
concentrations decreased in the roots of both cultivars and in the major mechanisms of iron acquisition studied in plants are based on
shoots of rice plants under iron-deficient conditions. In the Poaceae either the reduction of organic ferric iron chelates via membrane-
family, iron deficiency induces the synthesis and exudation of associated ferrireductase or the release of phytosiderophores that
phytosiderophores (PS), which bind both iron and copper. The Cu-PS bind ferric ions and introduce them into the cell. Plant ferritin is
complex can be transported into root cells by the Fe-PS transporter, synthesized as a precursor containing a unique amino acid sequence
although with lower affinity (Briat et al., 1995). At high iron composed of over 70 residues at the N-terminal, followed by
concentrations, phosphorus concentrations increased considerably a region that is relatively conserved among other ferritins. The
in the roots. Higher amounts of iron oxides that accumulated in the first part of this region is known as transit peptide (TP), which is
roots are capable of absorbing anions such as phosphates (Zhang et responsible for plastid targeting and is processed upon entry into
al., 1999). In shoots, the phosphorous amounts decreased in plants the plastid. The second part is known as the extension peptide,
under iron excess, which suggests that there was limited phosphorous which may be lost in the germination process (Ragland et al., 1990).
translocation to the shoots. Moreover, there was likely limited uptake The first and second ferritins consist of about 40 and 30 amino
of apoplast phosphorous. Howeler (1973) stated that phosphorous acid residues, respectively. The conserved region neighboring
precipitation in the apoplast of the root results in lower phosphorous the extension peptide has been shown to have 40–50% sequence
absorption by the plant. Calcium concentrations decreased under the identity with mammalian ferritins.
excess iron treatment in roots and shoots, and Selveira Vivian et al. Genetic improvement of iron content and stress adaptation in
(2007) found no reports on manganese deficiency levels in the roots plants using the ferritin gene has been reviewed (Goto et al., 2001),
of rice plants at high iron levels. Precipitation of manganese in the and the gene transfer technique has been a revolutionary tool
iron plaque may have resulted in its lower absorption by the sensitive for quickly creating novel varieties. The plants that have had the
cultivar, where the highest iron concentrations were found. Negative ferritin gene introduced possess two particular advantages: 1) the
interactions between iron and manganese have been reported in storage of excess iron and other elements and 2) the cancellation
plants (Fageria, 2003), and the potassium concentration decreased in of oxidative stress. There are two possibilities to create useful
the shoots under iron excess relative to the control (Neue et al., 1998). plants using the exogenous ferritin gene in this review. One is
Sulfur concentrations were lower in the shoots of plants exposed to iron-fortified plants used to overcome iron deficiency in humans,
excess iron. and the other is plants that have resistance to heavy metals,

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pathogens, and other stresses or those that exhibit enhanced iron works as a cofactor for enzymes involved in a wide variety
growth. These genetic improvements could lead to increased iron of oxidation-reduction reactions (i.e., photosynthesis, respiration,
content or stimulated growth. However, the mechanism of iron hormone synthesis, DNA synthesis, etc.). This function makes
storage and the scavenger system for oxidative stress using ferritin iron an essential nutrient, and its deficiency causes iron chlorosis,
gene expression have not been determined. An understanding at a which seriously constrains normal plant development. Iron
physical and molecular level is now necessary to take advantage toxicity in plants is indicated by bronzing characteristics, which
of the ferritin gene at an advanced stage. In non-graminaceous have been observed in plants grown in greater than 100 mM iron
plants, the Fe II-transporter gene and ferric-chelate reductase solutions. Higher iron uptake by plants reduces protein synthesis
gene have been cloned from A. thaliana, but Fe +2-reductase in leaves. Ferritin is considered crucial for iron homeostasis, and
has not been cloned. In graminaceous monocots, the genes for it consists of a multimeric spherical protein called phytoferritin,
mugineic acid (MAs) synthesis, nicotianamine synthase (nas) which is able to store up to 4500 iron atoms inside its cavity
and nicotianamine aminotransferase (naat), have been cloned in non-toxic form. A resistant variety may accumulate a larger
form barley, whereas the Fe+3-MAs transporter gene has yet to be amount of phytoferritin, which forms a complex that reduces iron
cloned. Transferrin absorption in Dunaliella has been reported, toxicity. Large amounts of iron in plants are responsible for protein
suggesting a phagocytic (endocytic) iron-acquisition mechanism. degradation, because iron may lead to the formation of reactive
In rice, the overexpression of OsIRO2 resulted in increased MAs oxygen radicals that are highly phytotoxic and cause protein
secretion, but the repression of OsIRO2 resulted in lower MAs degradation. Iron chlorosis is a widespread problem, especially for
secretion and hypersensitivity to iron deficiency. Northern blots regions where the bioavailability of iron in the soil is low. Usual
revealed that the expression of genes involved in the Fe (III)- remediation strategies consist of amending iron to soil, which
MAs transport system was dependent on OsIRO2 (Mori, 1999; is an expensive practice. Thus, genetically improved chlorosis-
Kobayashi et al., 2005; Ogo et al., 2009). The expression of the resistant rootstocks and new cultivar combinations offer the best
genes for NAS, a key enzyme in MAs synthesis, was notably solution to iron chlorosis, and it is one of the most important
affected by the level of OsIRO2 expression. Furthermore, a lines of investigation needed to prevent this nutritional problem.
microarray analysis demonstrated that OsIRO2 regulates 59 iron- Therefore, there is a need for new methods to diagnose and correct
deficiency-induced genes in roots. Some of these genes, including this nutritional disorder. The use of microarray techniques revealed
two transcription factors that were upregulated by iron deficiency, the changes in gene expression levels due to iron deficiency,
possessed the OsIRO2 binding sequence in their upstream regions. and it has allowed insights into the transcriptional regulation of
OsIRO2 possesses a homologous sequence of the iron deficiency- some functions. These studies have extended our knowledge of
responsive cis-acting element in its upstream region (Ogo et al., stress responses to iron deficiency, and have identified candidate
2009). Three NAS genes have been identified (OsNAS1-3) in genes and processes for further experimentation to increase our
rice, and they are reported to catalyze the formation of NA (Inoue understanding of iron deficiency stress. It is likely that more
et al., 2003). OsNAAT1 converts NA to a 3'-keto intermediate extensive microarray analyses, coupled with suitable annotations
(Inoue et al., 2008), which is further converted to deoxymugineic of completely sequenced genomes, will prove valuable in future
acid (DMA) by OsDMAS1 (Bashir and Nishizawa, 2006; Bashir studies of iron stress responses in plants. Novel gene regulation
et al., 2006). In rice, this is secreted to the rhizosphere by TOM1 networks for iron deficiency responses include iron deficiency-
(Nozoye et al., 2011), where it binds to Fe+3 and is absorbed by inducible basic helix-loop-helix transcription factors (OsIRO2),
OsYSL15 (Inoue et al., 2009). The role of YSL family transporters iron deficiency-responsive cis-acting elements (IDEs), and the
in iron transport is well characterized. The rice genome contains two transcription factors. The identification of the trans-acting
18 putative YSL family genes ( Koike et al., 2004 ) , among factors responsible for the pathways will contribute greatly to an
which OsYSL2, OsYSL15, OsYSL16, and OsYSL18 have been understanding of the regulatory mechanisms, and these factors
characterized (Aoyama et al., 2009; Inoue et al., 2009; Kakei will also prove beneficial to plants under iron-deficient conditions.
et al., 2012; Zheng et al., 2012). OsYSL2 is a Mn-NA and Fe-
NA transporter (Koike et al., 2004). These genes will contribute Acknowledgement
greatly to an understanding of the regulatory mechanisms. The authors wish to acknowledge to the Department of Science
and Technology, Government of Odisha for financial assistance.
Conclusions
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