267P. Ahmad and M.N.V. Prasad (eds.), Abiotic Stress Responses in Plants: Metabolism, Productivity and Sustainability, DOI 10.1007/978-1-4614-0634-1_15, © Springer Science+Business Media, LLC 2012

15

Abstract

Several environmental factors infl uence the mineral uptake of plants, including pH, redox potential, and the presence of xenobiotics as well as the temperature and salinity. Changes in environmental conditions affect the biological and physiological response of plants. Most important targets of plants are maintained by the ion homeostasis and mineral uptake in stress conditions. Plants may use the different procedures to regulate homeostasis. In general, Na salt stress decreases the levels of cationic nutrients such as K, Ca, and Mg due to competition for ion transport sites. Drought and salinity stress are physiologically related and the tolerance mechanisms overlap. Metals can interfere with mineral nutrition and change the concentration and composition of plant nutrients. Besides, met-als may also alter the conformation of proteins, including transporters, or regulator proteins. Herbicides may disrupt the function and integrity of the cell membrane, and signifi cant ion losses can occur. However, molecular mechanisms and genetic basis of interactions between abiotic stress and mineral uptake is lacking. Thus, future studies will focus on these aspects. In this chapter, the effects of some common stressors, such as salinity, drought, metals, herbicides, and on nutrient uptake are elucidated.

Keywords

Abiotic stress • Salinity • Drought • Heavy metal • Herbicide • Mineral uptake • Metal

Uptake of Mineral Elements During Abiotic Stress

Fatih Duman

F. Duman (�) Department of Biology, Faculty of Science , Erciyes University , Kayseri 38039 , Turkey e-mail: fduman@erciyes.edu.tr

268 F. Duman

1 Introduction

Sixteen chemical elements are known to be important for plant growth and survival. Mineral nutrients are inorganic elements in food, which the body cannot synthesize. These chemicals may be divided into two groups, namely, minerals and non-minerals. Non-mineral nutrients include hydrogen (H), oxygen (O), and carbon (C). These nutrients are found in the air and water. The other 13 mineral nutrients are found in the soil and dis-solved in water, so they are absorbed through the roots. Mineral nutrients are classifi ed as macro-nutrients and micronutrients. There are two groups of macronutrients, namely, primary and secondary nutrients. The primary nutrients are nitrogen (N), phosphorus (P), and potassium (K), which are needed in large amounts for plant growth and survival. The secondary nutrients are calcium (Ca), magnesium (Mg), and sulphur (S). Since the amount of these minerals in the soil is usually suffi cient for growth, fertilization is not always needed. Micronutrients are elements, which are essential for plant growth but are required in much smaller amounts than primary nutrients. The micronutrients are boron (B), cop-per (Cu), iron (Fe), manganese (Mn), molybde-num (Mo), zinc (Zn), and chloride (Cl). Although micronutrients only comprise 5% of the biomass of a plant, they are essential for growth. For example, Zn plays a key role in enzymatic cataly-sis of reactions, which require an electrophile. In addition, Fe, Mn, Cu, and Mo are responsible for redox transformations (Merchant 2006 ) . Although the exact functions of B are not fully understood, it is known to be necessary for numerous impor-tant processes, such as protein synthesis, sugar transport, and respiration (Hänsch and Mendel 2009 ) . The uptake, translocation, and exclusion of these mineral nutrients are related to many environmental factors, such as salinity, acidity, water shortage, and the presence of xenobiotics. In this chapter, we discuss the effects of some common stressors, such as salinity, drought, met-als, herbicides, and on nutrient uptake.

2 Abiotic Stressors Effecting Mineral Uptake

2.1 Salt Stress

Salinity is the major environmental factor, which limits plant growth and productivity. Salinity changes the biological properties of plants, such as facilitating their retention and acquisition of water and maintenance of ion homeostasis (Parida and Das 2005 ) . Grattan and Grieve ( 1999 ) showed that soils have extremely high ratios of Na + /Ca 2+ , Na + /K + , Ca 2+ /Mg 2+ , and Cl − /NO

3 − under saline

conditions, which leads to ion toxicities (e.g., Na + and Cl − ) and ionic imbalance. In Parida and Das’ review ( 2005 ) of salt tolerance and the effects of salinity in plants, they discussed three main issues, which are related to ion homeostasis during salt stress (1) selective accumulation or exclusion of ions, (2) control of ion uptake by roots and transport into leaves, and (3) compart-mentalization of ions at both the cellular and whole-plant levels. However, plants do not have the ability to tolerate high concentrations of salt in the cytoplasm. As a result, they sequester excess salts in vacuoles or compartmentalize the ions in different tissues (Zhu 2003 ) .

Khan ( 2001 ) showed that plants, which are exposed to salt stress exhibit increased levels of Na + and Cl − and decreased levels of Ca 2+ , K + , and Mg 2+ . Similarly, Lee and Liu ( 1999 ) demonstrated that Na + and Cl − concentrations in Ulva fasciata increased when exposed to increasing salinity but decreased the total and water-soluble concentra-tions of Ca 2+ . However, Ferreira et al. ( 2001 ) reported that salinity did not affect the Mg 2+ con-tent in the stems and roots of Psidium guajava L., but decreased it in the leaves. Although K is not incorporated into chemical compounds in plants, it is an important regulator of development. K regu-lates not only the electrical and water balance but also enzymatic processes, such as protein and car-bohydrate synthesis. High levels of Na are cyto-toxic. During salt stress, K + uptake is blocked due to accumulation of Na + . Under normal conditions,

26915 Uptake of Mineral Elements During Abiotic Stress

the cytosolic concentration of K + is higher than that of Na + . Maintaining this balance is essential for plant life (Zhu 2003 ) . Na + competitively inhib-its K + uptake (Grattan and Grieve 1998 ) . During salt stress, K + uptake is disrupted in root cells. Intracellular K + and Na + homeostasis is important for many biological activities, such as enzyme activity, maintenance of the membrane potential, and regulation of cell volume. Unlike animal cells, plant cells do not have Na + -ATPases or Na + /K + -ATPases (Hasegawa et al. 2000 ) . Therefore, excess Na + must be extruded or compartmentalized in vacuoles. During salt stress, plants also need a mechanism to regulate turgidity. Many studies have shown the importance of K in regulating tur-gidity (Guardia and Benlloch, 1980 ; Mengel and Arneke, 1982 ; Hsiao and Läuchli 1986 ) .

Low levels of Ca in growth media cause defects, such as deterioration of the cell mem-brane, loss of cellular components, and eventu-ally cell and tissue death. Kaya et al. ( 2002 ) showed that Ca defi ciency induces high concen-trations of NaCl in strawberry. Calcium ions ameliorate the effect of salt stress by competing with sodium ions for membrane-binding sites. Tuna et al. ( 2007 ) studied the effects of calcium sulphate (CaSO

4 ) on nutrient uptake in tomato

plants, which were grown in pots under salt stress and concluded that CaSO

4 increased concentra-

tion of Ca 2+ , N, and K + and reduced the concen-tration of Na + in the leaves. Patel et al. ( 2011 ) also tested the effects of supplemental Ca on the nutrient levels in Caesalpinia crista L. (Fabaceae) in salinized soil in a greenhouse. They demon-strated that salt stress reduces N, P, K, and Ca content in tissues; however, the addition of Ca restored the levels of these nutrients. In general, as external Ca concentrations increase, Na uptake and concentrations decrease while Ca uptake and concentrations increase because Ca 2+ interferes with non-selective cation channels and restricts Na + uptake. In addition, as the salt concentration in the root zone increases, the requirement for Ca 2+ increases. However, the uptake of Ca 2+ from the soil may be reduced as a result of ion interac-tions, precipitation, and increased ionic strength. These factors reduce the activity of Ca 2+ in solu-tion, which reduces the availability of Ca 2+ (Grattan and Grieve 1998 ) .

In plants, nitrogen comprises 80% of the total absorbed nutrients (Marschner 1995 ) . In general, salinity reduces N accumulation and saline envi-ronments are associated with nitrogen defi ciency (Siddiqui et al. 2010 ) . For example, tomato (Cerda and Martinez 1988 ) and barley (Shen et al. 1994 ) plants have reduced nitrogen content when they are grown under salt stress. This is not surprising since an increase in Cl − uptake and accumulation is often accompanied by a decrease in NO

3 − concentration in shoots. Tuna et al.

( 2007 ) suggested that this decrease may be due to inhibition of nitrogen uptake by NO

3 − /Cl −

interaction at ion transport sites. Many nitrogen- containing compounds, such as amino acids (e.g., proline and glycine betaine), protect against salt stress. For example, Siddiqui et al. ( 2010 ) showed that supplemental N limited the effects of salt stress on Brassica genotypes in salinized envi-ronments. In addition, the chemical form of N, such as NO

3 or NH

4 , affects the interaction

between salt and N content. Martinez and Cerdá ( 1989 ) demonstrated that Cl − uptake decreased in cucumber ( Cucumis sativus ) when only NO

3 −

was added but Cl − accumulation increased when half the NO

3 − was replaced with NH

4 + .

We do not have enough data about the interac-tion between salinity and P to understand the underlying mechanism of changes in P uptake (Patel et al. 2009 ) . The availability of P depends on the salinity and ionic strength of soil. In addi-tion, P concentrations in the soil solution are tightly controlled by sorption processes and by the low solubility of Ca–P minerals (Violante et al. 2002 ) . Champagnol ( 1979 ) showed that the addition of P to saline soils increased crop growth and yield. However, Grattan and Grieve ( 1999 ) reviewed studies about the relationship between salinity and P concentration in tissues and con-cluded that salinity increased the P concentration in tissues in sand or solution cultures but not soil.

In general, Na salt stress decreases the levels of cationic nutrients (K, Ca, and Mg). Ruiz et al. ( 1997 ) studied the effects of salt stress on mineral uptake in citrus plants and showed that NaCl salinity reduced Mg 2+ concentrations in the leaves. There is a strong competition between Mg 2+ and Ca 2+ for plasma membrane binding sites in roots; however, Ca has a higher affi nity

270 F. Duman

for these sites than Mg 2+ (Marschner 1995 ) . As a result, plants grown in the presence of high levels of Ca have lower Mg content. Loupassaki et al. ( 2002 ) reported similar effects of salt stress on olive cultivars. However, Barhoumi et al. ( 2007 ) demonstrated that salinity did not affect Mg lev-els in Aeluropus littoralis , a perennial halophyte that retains Mg.

Several factors infl uence the metal uptake capacity of plants, including pH, redox potential, and concentration of surrounding metals as well as the temperature and salinity of the surrounding water (Leblebici et al. 2011 ) . In addition, Na + ions release Cd from the sediment into water, thereby increasing the concentration of soluble Cd (Greger et al. 1995 ) . Greger et al. ( 1995 ) also showed that when Potamogeton pectinatus grows in highly saline water, low levels of free Cd ions in the water correlate with a low Cd uptake capac-ity. At high salinity, increased concentrations of NaCl reduce the uptake of metals in Spirodela polyrrhiza (Leblebici et al. 2011 ) . Many other studies also have shown that high salinity decreases metal uptake (Munda and Hudnik 1988 ; Wang and Dei 1999 ) , which may be due to the formation of complexes of metal and chloride ions (Mamboya et al. 2009 ) . Similarly, Greger et al. ( 1995 ) designed an experimental study to determine the infl uence of salinity on Cd uptake in submerged macrophytes by using Potamogeton pectinatus , and showed that increasing salinity decreases Cd uptake in P. pectinatus from water, most likely due to the formation of Cd complexes with chloride and sulphate. In contrast, in the presence of sediment, increasing salinity increases Cd uptake because Na + and Mg 2+ displace Cd 2+ in sediment colloids. However, Fritioff et al. ( 2005 ) did not fi nd any effect of salinity on Pb accumu-lation. This may be because Pb does not form a complex with chloride and has a high binding affi nity for organic matter. Manousaki et al. ( 2008 ) also reported a similar effect of increased soil salinity on increased cadmium uptake in Tamarix smyrnensis , which is a halophytic plant, and speculated that this effect may be related to a higher mobility of metals in the sediment or higher water uptake, which would increase the metal fl ux into the plant. In another study on

T. smyrnensis , Kadukova and Kalogerakis ( 2007 ) observed that high salt concentrations in soil decreased the accumulation of Pb in the roots but increased it in the leaves. Collectively, these stud-ies show that salt–metal interactions are very complex. As a result, further research is needed to elucidate these interactions in more detail.

2.2 Drought Stress

Drought (continuous water defi cit) is one of the most important factors that affect the growth, development, and survival of plants. Drought stress usually occurs because of insuffi cient water availability in the soil; however, it can also occur due to excessive loss of water by transpiration or evaporation (Jaleel et al. 2009 ) . Drought stress causes signifi cant physiological changes, such as stomatal closure, decreased photosynthetic activ-ity, and changes in cell wall elasticity. Drought and salinity stress are physiologically related and the tolerance mechanisms overlap. Drought dis-turbs the nutritional status of plants by altering ion concentrations in tissues. Since soil moisture is reduced during drought stress, the rate of diffu-sion of nutrients from the soil matrix to the absorbing root surface is also reduced (Hu et al. 2007 ) . In addition, nutrient transport from the roots to the shoots decreases due to reduced tran-spiration rates.

Ca 2+ is involved in plant drought resistance. When Ca is applied to leaves, it enhances their ability to conserve water (Shao et al. 2008 ) . In addition, Ma et al. ( 2009 ) showed that Ca 2+ alters the degree of hydration of the plasma membrane and improves the cohesion of cell walls, which increases the viscosity of the protoplasm and the resistance of cells to dehydration. Berkowitz et al. ( 2000 ) demonstrated that Ca 2+ inhibits the infl ux of K + to guard cells during water stress by affecting inward rectifying K + channels.

In addition to Ca, K is responsible for osmo-regulation in plant cells during drought stress (Roberts 1998 ) . Mohsenzadeh et al. ( 2006 ) showed that K levels in Aeluropus lagopoides increased signifi cantly under mild and moderate drought stress. Zhang et al. ( 2006 ) demonstrated that K

27115 Uptake of Mineral Elements During Abiotic Stress

transport increases during the early stages of adaption to stress in plants. K channels are impor-tant to regulate the water status of plants (Roberts 1998 ) . However, there are some contradictory reports that drought stress decreases K levels. For example, Thiec and Manninen ( 2003 ) suggested that K levels decrease during drought stress due to changes in nutrient compartmentalization.

Martínez et al. ( 2004 ) proposed that Na + may play a positive role in response to water stress, because Na + absorption increases in plants that are subjected to drought stress on a non-saline substrate. Similar observations have been reported (leaf Na + concentration was signifi cantly increased by simultaneous exposure to drought) in halophyte species, such as Sesuvium portulac-astrum , Atriplex halimus , and Ipomoea pes-caprae (Slama et al. 2008 ; Martínez et al. 2005 ; Sucre and Suárez 2011 ) . These studies suggest that Na + may have a direct or indirect positive infl uence on the accumulation of other elements, which are involved in osmotic regulation. In addition, Na plays a specifi c role in parasitic plants, such as Cuscuta attenuate . Kelly and Horning ( 1999 ) showed that parasitic plants accumulate higher levels of Na than their hosts do, to draw water and nutrients from their hosts.

After drought, there is usually a nitrogen defi -ciency in the ground. There is a strong relationship between water availability and N absorption. In general, the availability, uptake, and utilization of N by plants increases as the soil moisture content increases (Albrizio et al. 2010 ; Kibe et al. 2006 ) . However, Payne et al. ( 1995 ) reported that total nitrogen and NO

3 levels increase during drought

stress in pearl millet ( Pennisetum glaucum ). Song et al. ( 2010 ) demonstrated that N and P have strong interactive effects on plant growth and that the water supply was the primary determinant of this interaction. Water defi cit reduces the net CO

2

assimilation rate, which may be due to inhibition of ribulose bisphosphate synthesis and ATP syn-thase activity (Tezara et al. 1999 ) . Decreased ATP synthesis in chloroplasts may also be caused by a low availability of free inorganic phosphate in the cytoplasm (Guida Dos Santos et al. 2006 ) . Thus, water defi cit decreases the levels of free phosphate in the cytoplasm (Pieters et al. 2001 ) .

Increased levels of Mg in leaves can help maintain water content during drought stress. Mahouachi ( 2009 ) investigated the changes in the concentration of mineral nutrients in Musa acuminate plants, which were grown in increas-ingly dry soil. These drought-stressed plants showed a signifi cant accumulation of Mg (higher 28% compared to control). Since Mg 2+ is an essential element in chloroplasts, a defi ciency can decrease photosynthetic activity. Mahouachi ( 2009 ) proposed two possible explanations for this accumulation (1) the increased concentration of mineral nutrients may be associated with the translocation of ions from old to young leaves or (2) the increased ion concentration in the root zone may be due to a concentration effect, which was produced by the reduction in soil moisture. However, in this case, xylem fl ux should still be active enough to move ions to functioning leaves.

The effects of metals on water content in soil solutions are controversial. Tipping et al. ( 2003 ) argued that the interaction between metals, such as Cu and Zn, and drought is the most important factor in soil acidifi cation. Since the strength of metal binding to natural organic matter is inversely related to pH, acidifi cation may release metals into the soil. On the other hand, Zn avail-ability may promote water conservation by decreasing the transpiration rate. Disante et al. ( 2011 ) showed that, at least in the short term, Zn may promote water conservation. Similarly, Gadallah ( 2000 ) demonstrated that Zn stimulates the accumulation of osmotically active solutes, such as soluble sugars, to facilitate water uptake. In addition, Disante et al. ( 2011 ) suggested that Zn affects stomatal dynamics by inhibiting water channels and reducing K + uptake.

2.3 Heavy Metal Stress

2.3.1 Cadmium Cadmium (Cd) is a phytotoxic element because it can interfere with mineral nutrition and change the concentration and composition of plant nutri-ents. Cd 2+ may interfere with nutrient uptake by altering plasma membrane permeability and may

272 F. Duman

also alter the conformation of proteins, including enzymes, transporters, or regulator proteins, due to its strong affi nity for sulfhydryl and carboxylic groups (Assche and Clijsters 1990 ; Gonçalves et al. 2009 ) . Many studies have focused on the effect of Cd on nutrient uptake and translocation in plants (Zhang et al. 2002 ; Dong et al. 2006 ; Gonçalves et al. 2009 ) . However, there is no con-sensus about the effects of Cd on mineral nutrient uptake because there are contradictory reports about species or cultivar differences, and interac-tions between metals and plant tissues (Liu et al. 2003 ) . For example, Cui et al. ( 2008 ) showed that copper (Cu), iron (Fe), and zinc (Zn) inter-fere with Cd in rice. The addition of Cu signifi -cantly decreased Cd uptake by the shoots and roots of rice. However, Zn uptake decreased sig-nifi cantly as the amount of Cd and Cu increased. In another study, Liu et al. ( 2003 ) studied the interaction of Cd with fi ve mineral nutrients (Fe, Zn, Cu, Mn, and Mg) and found signifi cant dif-ferences between rice cultivars. In addition, Gonçalves et al. ( 2009 ) studied the interaction between Cd 2+ and mineral nutrients in potato ( Solanum tuberosum ) both in vitro and in hydro-ponic culture. Although Cd 2+ did not affect the content of mineral nutrients in hydroponically grown plantlets, it decreased the content in in vitro plantlets. Nada et al. ( 2007 ) conducted an experimental study with hydroponically grown almond ( Prunus dulcis ) with 25–150 m M Cd 2+ for 14 days and showed that all concentrations of Cd 2+ reduced the concentration of Ca 2+ , Mg 2+ , and K + in the leaves, but only the highest concentra-tion of Cd 2+ decreased the concentration of K + and Mg 2+ in the roots.

The concentration of Cd 2+ also affects its inter-action with minerals. For example, when Brassica chinensis was exposed to 0.1 m g mL −1 Cd 2+ , the concentration of Zn increased; however, at higher concentrations of Cd 2+ , the concentration of Zn decreased (Wong et al. 1984 ) . At low levels, Cd may hyperpolarize the plasma membranes on the surface of roots, thereby increasing the transmembrane potential (Gonçalves et al. 2009 ) . This hypothesis is consistent with some studies with metal transporters, which belong to the natural resistance-associated macrophage

protein (NRAMP) and zinc-regulated transporter/ iron-regulated transporter-related protein (ZIP) families (Guerinot 2000 ) . These transporters may increase the concentration of mineral nutrients. Dong et al. ( 2006 ) showed that the main toxic effects of Cd 2+ result from its interaction with essential elements, especially those with the same valence, such as Cu 2+ , Fe 2+ , Mn 2+ , and Zn 2+ . In contrast, Liu et al. ( 2003 ) demonstrated close relationships between Cd 2+ and Fe 2+ , Cd 2+ and Zn 2+ , and Cd 2+ and Cu 2+ in both roots and leaves of rice.

Yu and Zhou ( 2009 ) revealed an antagonistic relationship between Cd and P in a study on Mirabilis palaja . As a result, they advised that the addition of P to Cd-contaminated soil may be an effective way to immobilize Cd. Similarly, Jiang et al. ( 2007 ) suggested that external P can decrease the bioavailability of Cd and Zn. Yang et al. ( 1999 ) determined the effect of P on the accumulation of Cd and Zn in suspension-cultured wheat cells. They observed that the addition of P into the cul-ture medium reduces Cd and Zn bioaccumulation. Earlier studies showed that Cd exposure decreases nitrate reductase activity in Nicotiana tabacum (Dguimi et al. 2009 ) and Oryza sativa (Huang and Xiong 2009 ) , which increases the concentration of free amino acids and decreases the concentra-tion of soluble proteins and nitrates in plant tis-sues. Although few studies have focused on the interactions between sulphur availability and Cd exposure, it is known that Cd plays an important role in the synthesis of thiol-based complexing substances of phytochelatins by upregulating glu-tathione biosynthesis (Astolfi et al. 2004 ) . Fan et al. ( 2010 ) showed that excessive S signifi cantly decreases the accumulation of Cd in brown rice by decreasing Cd availability and increasing glu-tathione in rice leaves. In contrast, Nocito et al. ( 2002 ) demonstrated that S can increase Cd avail-ability and concentration in plants. Eventually, relationship between S and Cd is controversial. Previous studies showed that the presence of Cd in the culture medium decreases the concentration of K and Ca in different plant organs (Yang and Lee 2002 ; Ghnaya et al. 2005 ) . This decrease may be due to the competition of Cd 2+ with Ca 2+ and other cations for entry into plant cells.

27315 Uptake of Mineral Elements During Abiotic Stress

2.3.2 Arsenic Inorganic arsenic (As) compounds are used in industrial and agricultural applications as well as aquatic weed control. Previous studies have shown that As may compete directly with nutri-ents or alter metabolic processes in plants. Since inorganic As(V) and phosphate are chemically similar, As(V) can act as a phosphate analogue and be transported into the cell (Meharg and Macnair 1990 ) . Intracellular As(V) can interfere with essential cellular processes, such as oxida-tive phosphorylation and ATP synthesis (Tripathi et al. 2007 ) . Many studies have demonstrated competition between As and P (Mkandawire et al. 2004 ; Mkandawire and Dudel 2005 ; Karadjova et al. 2008 ) . Meharg and Hartley-Whitaker ( 2002 ) explained that this competition arises because As and P use the same transport system.

Tu and Ma ( 2005 ) tested the effects of As on essential macronutrients (P, K, Ca, and Mg) and micronutrients (Fe, Mn, Cu, Zn, B, and Mo) in Pteris vittata , which is an As hyperaccumulator. At low levels of As, the levels of P and K increased along with As. These authors reported a negative correlation between As accumulation and the Ca level in P. vittata , which may refl ect the ability of Ca to prevent As toxicity. In addition, they showed that Mg has similar patterns of accumulation and distribution as Ca in the fronds. Although As reduced the concentration of Fe and Zn in the fronds, As did not signifi cantly affect the concen-tration of Cu and Mn. Vance et al. ( 2003 ) reported that P can combine with other cations, especially aluminium (Al) and Fe, in complexes under acidic conditions. As a result, the As-induced reduction in micronutrients (excluding Mn) in the fronds was probably due to As phytotoxicity. Another study in tomato ( Lycopersicum esculen-tum ) showed that inorganic As(V) signifi cantly decreased the concentration of both macronutri-ents (K, Ca, and Mg) and micronutrients (B, Cu, Mn, and Zn) (Carbonell-Barrachina et al. 1998a ) . Carbonell-Barrachina et al. ( 1998b ) determined the effects of different chemical forms and con-centrations of As on its uptake and nutrition in an aquatic plant ( Spartina alternifl ora ). The applica-tion of organic As species signifi cantly increased the concentration of Ca in leaves. These increased

Ca levels conferred protection from metal and metalloid toxicity. However, the application of inorganic As species signifi cantly increased the concentration of Cu in the roots and shoots of S. alternifl ora . Due to the high phytotoxicity of monomethylarsonic acid (MMAA), the concen-tration of several essential macronutrients (e.g., P, K, Ca, and Mg) and micronutrients (e.g., B, Cu, and Fe) decreased signifi cantly. The increase in Ca levels could have been due to the protective action of Ca against As toxicity.

2.3.3 Mercury Previous studies have mostly focused on mercury (Hg) accumulation and the effect of Hg on the antioxidant activities of plant enzymes. As a result, few studies have investigated the effects of Hg on uptake of nutrient elements (Rodríguez et al. 2009 ) . Hg has a high affi nity for sulfhydryl groups, which can disrupt the function of essential proteins and, consequently, alter plant development. Hg also can restrict water channels in higher plants by altering membrane permeability (Patra et al. 2004 ) . In plants, Hg can replace some nutritional ele-ments, such as Mg, Zn, and Mn. This may be the main effect of Hg on nutrient uptake (Patra et al. 2004 ) . In previous studies, Hg alters the uptake and translocation of mineral nutrients (Gupta and Chandra 1998 ) ; however, Hg uptake decreases as nutrient levels increase (Göthberg et al. 2004 ) . Gupta and Chandra ( 1998 ) showed that increasing concentrations of Hg in the culture medium of Vallisneria spiralis signifi cantly decreases the concentration of N, P, and K. In addition, the trans-location of P, S, and K to the leaves increased as the concentration of Hg increased.

Rodríguez et al. ( 2009 ) investigated the uptake of Hg and levels of some micronutrients and macronutrients in hydroponically grown Chilopsis linearis . Hg only affected Fe, Mn, and Zn micronutrients and K, P, and S macronutri-ents. The concentration of Zn in the roots was reduced by 62% and 49% by 50 and 100 m M Hg, respectively. In addition, 50 m M Hg increased the concentration of Fe in the roots whereas 100 m M Hg decreased the concentration of Fe in the roots. Moreno-Jiménez et al. ( 2006 ) reported that Hg increased the concentration of Fe in the

274 F. Duman

roots of Marrubium vulgare by more than 40% but reduced the translocation of Fe.

2.3.4 Chromium Chromium (Cr) is a non-essential and toxic ele-ment for plants. It does not have any specifi c uptake mechanisms. However, Cr uptake is dependent on its chemical form. For example, Cr(VI) is taken up actively, whereas Cr(III) is taken up passively through the carriers for essen-tial anions, such as sulphate, and stored in the cell wall (Zayed and Terry 2003 ) . Shanker et al. ( 2005 ) reviewed the known relationships between Cr and plants and concluded that a high concen-tration of Cr is associated with a low concentra-tion of mineral nutrients, such as Ca, K, Mg, P, B, and Cu. Cr can interfere with the uptake of other ionically similar elements, such as Fe and S. Moral et al. ( 1996 ) studied the effect of Cr on the concentration of mineral nutrients in tomato and showed that Cr negatively affects Fe absorption. In the case of Fe, the reduction in its uptake may be due to competition with chemically similar ions. In addition, Shanker et al. ( 2003 ) suggested that nutrient uptake may decrease due to inhibition of the activity of plasma membrane H + ATPase. Despite the general consensus in the literature, there is still some disagreement about whether Cr increases Fe content (Barceló et al. 1993 ) .

Cr has a signifi cant effect on the N content of plants. Kumar and Joshi ( 2008 ) concluded that Cr (VI) adversely affects N content by interfering with key enzymes in nitrogen metabolism. Similar fi ndings have been reported for Miscanthus sinensis (Arduini et al. 2006 ) and Nelumbo nucifera (Vajpayee et al. 1999 ) . Dube et al. ( 2003 ) studied the interaction between Cr and P in citrul-lus. They showed that increasing concentrations of Cr were associated with increasing concentra-tions of P in citrullus leaves. This accumulation of P might be due to the direct interference of Cr with the metabolism of P in plants.

2.3.5 Nickel Although nickel (Ni) is essential for plants at low concentrations (Gajewska and Sklodowska 2007 ; Baccouh et al. 2001 ) , it is phytotoxic at high con-centrations (Duman and Ozturk 2010 ) . Excess Ni

also affects nutrient absorption by roots (Rahman et al. 2005 ) . Ahmad et al. ( 2011 ) assessed the effect of Ni on the accumulation of macronutri-ents (K, Ca, and Mg) and micronutrients (Zn, Mn, Fe, and Cu) in different parts of sunfl ower ( Helianthus annuus ). They showed that Ni stress substantially decreases all macronutrients and micronutrients in sunfl ower leaves and achenes. Specifi cally, high concentrations of Ni decreased the concentrations of Ca, Mn, and Fe in achenes. In addition, increasing concentrations of Ni decreased the concentration of N, K, Zn, Mn, and Cu in achenes. However, Ni did not affect the concentration of P or Mg. Similarly, Ali et al. ( 2009 ) reported that Ni reduced the N, P, and K content in Brassica napus . Kähkönen and Kairesalo ( 1998 ) also demonstrated that Ni inhib-its nutrient metabolism in Elodea canadensis . Moreover, Gajewska and Sklodowska ( 2007 ) suggested that Ni competitively displaces Ca ions from the Ca binding site in the oxygen-evolving complex. Ni has similar chemical characteristics as other mineral nutrients, such as Ca, Mg, Mn, Fe, Cu, and Zn. In addition, Ni is absorbed and transported by the same transport system as that for some other micronutrients, such as Cu and Zn (Ahmad et al. 2011 ) . As a result, high levels of Ni may inhibit the absorption of these nutrients.

2.3.6 Lead Lead (Pb) competes with divalent cations for trans-port into roots. This might be due to direct compe-tition between Pb and other essential nutrients for the same binding site. Therefore, the concentration of micronutrients, such as Mn and Cu, may decrease in the presence of Pb. Sinha et al. ( 2006 ) studied the effects of Pb on the uptake and translo-cation of essential nutrients in cabbage ( Brassica oleracea ). They demonstrated that as the concen-tration of Pb increased, the concentration of Zn increased whereas those of P, S, Fe, Mn, and Cu decreased in various parts of the cabbage plant. Geebelen et al. ( 2002 ) and Diaz-Aguilar et al. ( 2001 ) observed a similar relationship between Pb and P and suggested that Pb forms insoluble com-plexes with P. Many other studies also have shown that excess Pb decreases the concentration of Fe in plants (Kannan and Keppel 1976 ; Paivoke 2002 ) .

27515 Uptake of Mineral Elements During Abiotic Stress

2.3.7 Copper Previous studies have shown that the addition of Cu to plant growth media may affect the uptake of other mineral nutrients (Ke et al. 2007 ; Puig et al. 2007 ) . For example, Bouazizi et al. ( 2010 ) inves-tigated the accumulation and toxicity of Cu and determined the relationship between Cu accumu-lation and plant nutrients, such as Fe, K, Ca, and Zn in Phaseolus vulgaris . They concluded that the Fe, Zn, and K content decreased as a result of Cu accumulation, which refl ected a change in nutrient homeostasis. Ke et al. ( 2007 ) compared the effects of Cu and other mineral nutrients on mineral uptake in a population of Rumex japoni-cas that grew near a copper mine with another population that grew in an uncontaminated area. The population that grew in copper-contaminated soil evolved a tolerance of not only high levels of Cu but also a lack of nutrients. As Yang and Romheld ( 2002 ) suggested in a study on Elsholtzia splendens that Cu tolerance may be related to the ability to maintain high levels of other mineral nutrients while under Cu stress. Indeed, the nutri-ent composition of plants that grew in the con-taminated area exhibited less variation than those growing in the uncontaminated area.

Several studies have reported that mineral nutrients can affect the uptake and accumulation of Cu in plants (Nenova and Stoyanov 1999 ; Xiong et al. 2002 ) . For example, Fe defi ciency in the growth medium of pea ( Pisum sativum ) increased the concentration of Cu (Cohen et al. 1998 ) . Similarly, Xiong et al. ( 2002 ) showed that Fe defi ciency in the culture medium of plants stimulates Cu accumulation, while excess P reduces Cu accumulation. These results suggest that Cu has an antagonistic relationship with var-ious mineral nutrients. Likewise, P antagonizes the absorption and metabolism of several trace elements. However, some studies do not support this hypothesis. For example, Cambrollé et al. ( 2011 ) reported a positive correlation between the accumulation of Cu and P in Glaucium fl a-vum , which suggested that P plays an important role in controlling Cu accumulation and trans-port. In addition, Cu signifi cantly inhibits nitro-gen metabolism. Xiong et al. ( 2006 ) studied Cu-induced disruption of nitrogen metabolism in

Chinese cabbage ( Brassica pekinensis Rupr.) and demonstrated that Cu exposure increases Cu con-centration and decreases nitrate reductase (NR) activity in the roots and shoots. In addition, Mazen ( 2004 ) showed that Cu increases the con-centration of free amino acids. Alaoui-Sossé et al. ( 2004 ) investigated the effect of Cu on ion con-centrations and growth in cucumber ( Cucumis sativus ) and showed that Cu inhibits leaf expan-sion and reduces the net assimilation rate, which may be due to decreased levels of K and Mg, respectively.

2.3.8 Zinc Zinc (Zn) is essential for plant growth; however, it is phytotoxic at elevated levels. Extracellular Zn stress can disrupt the nutrient balance in plant cells. Jiang et al. ( 2007 ) showed that external application of P effectively protects plants from Zn toxicity by forming P–Zn complexes. Thus, the Zn-induced decrease in P content might enhance Zn toxicity in plants. In general, earlier studies showed that Zn exposure decreases nutri-ent content (Wang et al. 2009 ; Bonnet et al. 2000 ) and suggested that excess Zn might competitively inhibit the uptake of these elements. Since excess Zn kills root cells (Chang et al. 2005 ) , injured roots might have a reduced capability to assimi-late nutrients.

2.4 Herbicide Stress

In modern agriculture, herbicides are widely used to control weeds. Although herbicides are gener-ally used in small amounts, they are potent, and, consequently, they have potentially signifi cant risks in aquatic ecosystems. In addition, there are many different kinds of herbicide that are used in different areas. Furthermore, intensive herbicide use has caused signifi cant soil and water pollu-tion. In addition to their desired effects on target organisms, herbicides also have undesirable effects on non-target organisms (Duman et al. 2010 ) . For example, Pandey et al. ( 2005 ) showed that hydroquinone, which is a phytotoxin, dis-rupts the cellular membrane integrity of Chara zeylanica , which is a non-target organism.

276 F. Duman

The severity of the toxic effects of pesticides can be mitigated by nutrients in the local environ-ment. For example, Battah et al. ( 2001 ) showed that high concentrations of phosphate alleviated the toxic effects of thiobencarb ( S -4-chlorobenzyl diethyl (thiocarbamate)) on the growth and pho-tosynthetic activity of Anabaena variabilis . Conversely, Das and Debnath ( 2006 ) showed that herbicides stimulate the growth and activity of aerobic non-symbiotic N

2 -fi xing bacteria and

increase the amount of N and P in the rhizosphere. Qian et al. ( 2009 ) also reported that exogenous nitric oxide protects Chlorella vulgaris from the toxicity of herbicides by reducing the damaging effects of oxidants and increasing the transcrip-tion of related genes. In addition, the duration of herbicide treatment is another important factor affecting their toxicity. For instance, Pandey et al. ( 2005 ) demonstrated that the N, P, and K content of plants decreases as the duration of herbicide treatment increases.

Previous studies showed that the application of high concentrations of herbicides to plants may harm their cell membrane integrity (Duman et al. 2010 ; Wendt-Rasch et al. 2003 ) . Specifi cally, reactive oxygen species may disrupt the function and integrity of the cell membrane and cause irreparable damages to cellular functions (Nemat-Alla and Hassan 2006 ) . As a result, signifi cant ion losses can occur. Sinha ( 2002 ) studied the effect of hexachlorocyclohexane (HCH) alone and in combination with Fe on the cellular integ-rity of Hydrilla verticillata . Increasing concen-trations of HCH increased K + leakage; however, this leakage was lessened in the presence of Fe. In addition, the accumulation of HCH decreased in the presence of high concentrations of Fe, while the accumulation of Fe increased in the presence of high concentrations of HCH.

The interactions between herbicides and met-als in soil are very complex. For example, the amount of dissolved organic carbon compounds in growth media affects these interactions. The presence or absence of herbicides in soil can affect the uptake of mineral nutrients. For example, Chen et al. ( 2004 ) showed that 2,4-dichlorophenol (2,4-DCP) increases the concentration of water-soluble Cu and Zn. Teisseire et al. ( 1999 ) investigated a synergistic effect between Diuron

(3-(3,4-dichlorophenyl)-1,1-dimethylurea) and Cu and revealed that the combination may prevent the toxic effects of Diuron. Some herbicides, such as glyphosate can create complex with diva-lent ions, such as Cu 2+ (Sheals et al. 2003 ) , Zn 2+ , Mn 2+ , Ca 2+ , and Mg 2+ . Therefore, herbicides may decrease the availability of nutrients. In addition, Rengel and Wheal ( 1997 ) demonstrated that chlorsulfuron decreases the uptake of micronutri-ents (Zn, Cu, and Mn) in wheat genotypes. Similarly, Azmat et al. ( 2006 ) showed that atrazine decreases the levels of Na and K in Vigna radita .

3 Conclusion and Future Perspective

Plant cultivation and consumption are important for human survival. However, water and soil resources around the world are increasingly con-taminated by xenobiotic compounds. For exam-ple, in many parts of the world, soil is becoming increasingly arid as a result of many factors, such as wild irrigation and misuse of fertilizers. Consequently, hunger is a signifi cant global prob-lem. Every year, 15 million children die of hun-ger. It may be possible to solve, or at least alleviate, this problem by using soil and water resources more effectively. Unless plants are able to obtain suffi cient nutrients and water from their growth media, they cannot survive. In general, environmental stresses negatively affect plant growth. Consequently, understanding the interac-tion between plant nutrients and stressors is criti-cal. Currently, there is ample knowledge about the interactions between stressors and mineral nutrients; however, information about the under-lying molecular mechanisms and genetic basis of these interactions is lacking. Thus, future studies should focus on these aspects.

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