Journal of Experimental Botany

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Journal of Experimental Botany, Vol. 51, No. 343, pp. 221-226, February 2000
© 2000 Oxford University Press

Uptake and retranslocation of leaf-applied cadmium (¹⁰⁹Cd) in diploid, tetraploid and hexaploid wheats

I. Cakmak^1,3, R.M. Welch², J. Hart², W.A. Norvell², L. Oztürk and L.V. Kochian²

¹ Cukurova University, Department of Soil Science and Plant Nutrition, 01330 Adana, Turkey
² US Plant, Soil and Nutrition Laboratory, USDA-ARS Cornell University, Ithaca, NY 14853, USA

Received 25 June 1999; Accepted 7 September 1999

	Abstract

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Uptake and retranslocation of leaf-applied radiolabelled cadmium (¹⁰⁹Cd) was studied in three diploid (Triticum monococcum, AA), four tetraploid (Triticum turgidum, BBAA) and two hexaploid (Triticum aestivum, BBAADD) wheat genotypes grown for 9 d under controlled environmental conditions in nutrient solution. Among the tetraploid wheats, two genotypes were primitive (ssp. dicoccum) and two genotypes modern wheats (ssp. durum). Radiolabelled Cd was applied by immersing the tips (3 cm) of mature leaf into a ¹⁰⁹Cd radiolabelled solution. There was a substantial variation in the uptake and export of ¹⁰⁹Cd among and within wheat species. On average, diploid wheats (AA) absorbed and translocated more ¹⁰⁹Cd than other wheats. The largest variation in ¹⁰⁹Cd uptake was found within tetraploid wheats (BBAA). Primitive tetraploid wheats (ssp. dicoccum) had a greater uptake capacity for ¹⁰⁹Cd than modern tetraploid wheats (ssp. durum). In all wheats studied, the amount of the ¹⁰⁹Cd exported from the treated leaf into the roots and the remainder of the shoots was poorly related to the total absorption. For example, bread wheat cultivars were more or less similar in total absorption, but differed greatly in the amount of ¹⁰⁹Cd retranslocated. The diploid wheat genotype ‘FAL-43’ absorbed the lowest amount of ¹⁰⁹Cd, but retranslocated the greatest amount of ¹⁰⁹Cd in roots and remainder of shoots. The results indicate the existence of substantial genotypic variation in the uptake and retranslocation of leaf-applied ¹⁰⁹Cd. This variation is discussed in terms of potential genotypic differences in binding of Cd to cell walls and the composition of phloem sap ligands possibly affecting Cd transport into sink organs.

Key words: Wheat, genotypic variation, radiolabelled Cd, uptake, translocation.

	Introduction

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There is an increasing concern about the accumulation of cadmium (Cd) in food crops. Increases in the amount of Cd in the agricultural environment is primarily the result of the use of Cd-containing fertilizers, application of Cd-containing sewage sludges and atmospheric deposition of Cd on crop or soil surfaces (Ryan et al., 1982; Nicholson and Jones, 1994).

Plant species and genotypes of a given species show great variation in the uptake of Cd by roots and its translocation from roots into shoots (Petterson, 1977; Florjin and Van Beusichem, 1993; Guo et al., 1995; Hart et al., 1998). In most cases, roots contain the highest Cd concentrations of all organs: the gradient of Cd concentration declines in the order root>leaves>grains or seeds (Wagner, 1993). Retention of Cd in roots reduces its transport into aerial parts of plants. This is desirable, not only because it alleviates Cd injury in plants, but also because it reduces Cd entry into the human food chain. As reported (Grant et al., 1998), restricting Cd transport from roots into shoots reduces Cd concentration in grains much more than in leaves. Possibly as a result of variations in root uptake and root-to-shoot transport of Cd, bread (Oliver et al., 1995) and durum wheat cultivars (Penner et al., 1995; Clarke et al., 1997) differ in grain concentrations of Cd. There are some reports showing that durum wheats accumulate higher amounts of Cd in grain than bread wheats (Meyer et al., 1982), particularly under Zn deficiency (Grant et al., 1998). Higher Cd accumulation in grains of durum wheats may be related to their higher sensitivity to Zn deficiency than bread wheats (Graham et al., 1992; Cakmak et al., 1998).

Accumulation of Cd in aerial parts of plants can also be affected by atmospheric deposition of Cd directly on to the leaf surfaces. As far as is known, there is little experimental evidence concerning uptake of Cd by leaves. In experiments with barley, carrot, kale, wheat, and rye carried out in rural areas of Denmark, it has been reported that 20–60% (or 0.02–0.4 mg Cd kg^-1 dry weight) of the total Cd in plants resulted from direct atmospheric deposition of Cd on to the leaf surfaces (Hovmand et al., 1983). According to Hovmand et al., atmospheric Cd deposited on aerial surfaces of plants can be taken up by leaves and transported within plants, and can represent a significant source of Cd entering the food chain. Atmospheric Cd can be transported up to several hundred kilometres when emitted into the atmosphere (Hovmand et al., 1983). Atmospheric deposition of Cd on to the leaf surfaces of cereals can be important because cereal-based foods are consumed in large amounts, representing 54% of food (i.e. dry matter) consumed world-wide (Graham and Welch, 1996).

In this study, genotypic variation in uptake and export of leaf-applied ¹⁰⁹Cd was investigated using different genotypes of diploid (AA), tetraploid (BBAA) and hexaploid (BBAADD) wheats.

	Materials and methods

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Seeds of three genotypes of diploid (T. monococcum), four genotypes of tetraploid (T. turgidum) and two genotypes of hexaploid (T. aestivum) wheat were surface-sterilized in 0.5% NaOCl for 20 min, rinsed in deionized water and germinated on filter paper. After 2 d, the germinated seedlings were transferred to black polyethylene cups, and the cups were placed in 5.0 l black polyethylene pots containing aerated nutrient solution. The composition of the nutrient solution was as follows (mol m^-3): 0.88 K₂SO₄, 2.0 Ca(NO₃)₂, 0.25 KH₂PO₄, 1.0 MgSO₄, 0.1 KCl, 5x10^-2 FeEDTA, 1x10^-2 H₃BO₃, 1x10^-3 ZnSO₄, 1x10^-3 MnSO₄, 0.2x10^-3 CuSO₄, and 0.02x10^-3 (NH₄)₆Mo₇O₂₄.

Plants were grown for 9 d in a growth chamber under controlled climatic conditions (light/dark regime 16/8 h, temperature 20/15 °C, relative humidity 60/75% and photon flux density of 700–800 µmol m^-2 s^-1).

Seeds of diploid wheats and primitive tetraploid wheats (ssp. dicoccum) were obtained from Dr CI Kling (State Plant Breeding Institute, University Hohenheim, Stuttgart). Seeds of the modern tetraploid (C-1252 and Balcali) and hexaploid (BDME-10 and Bezostaja) wheats were provided by the International Winter Cereals Research Center, Konya, and the Department of Field Crops, Cukurova University, Adana, Turkey. According to our field and greenhouse experiments, all diploid wheats and the hexaploid wheat cultivar Bezostaja used in this study are Zn-efficient (i.e. good growth under Zn-deficient soil conditions), while the other genotypes were Zn-inefficient (poor growth under Zn deficiency), with the exception of the tetraploid wheat Balcali which showed a moderate Zn efficiency (Cakmak et al., 1999a; and unpublished results).

Leaf application of radiolabelled Cd (¹⁰⁹Cd), obtained from the NEN-Life Sciences Products, Boston, MA, was carried out when the plants were 7-d-old. The tip (3 cm) of a mature leaf (the first leaf) was immersed into ¹⁰⁹Cd-containing radiolabelled solution (1.75 ml in 2 ml Eppendorf tubes) for about 10 s. This application procedure was repeated three times at different time intervals during the light phase over a 42 h period. The radiolabelled solution contained 0.01 mol m^-3 Cd as CdSO₄ (42.3 KBq nmol^-1) in 0.01% L77 as a wetting agent (a silicone block co-polymer fluid obtained from the Union Carbide Corporation, New York, USA). Plants were harvested 42 h after the initiation of the Cd application. At harvest, plants were divided into three parts: (i) ¹⁰⁹Cd-treated mature leaf, (ii) remainder of shoot, and (iii) roots. The leaf sections treated with ¹⁰⁹Cd were washed for about 10 min in 5 mol m^-3 CdSO₄ to remove ¹⁰⁹Cd adhering to leaf surface and within the leaf apoplasmic spaces.

Harvested plant parts were dried at 70 °C for the determination of dry matter production, and the ¹⁰⁹Cd activity was measured using a gamma counter (Model Auto-Gamma, Packard Instruments, Meriden, CT) and expressed per gram dry weight.

	Results

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Compared to tetraploid and hexaploid wheats, root and shoot dry weights of diploid wheats were lower (Table 1). Tetraploid and hexaploid wheats did not differ significantly in shoot and root dry weights. Primitive tetraploid wheats (ssp. dicoccum) tended to produce more dry weight than modern tetraploid wheats (spp. durum) (Table 1).

View this table: [in this window] [in a new window]   Table 1. Shoot and root dry weights of different diploid, tetraploid and hexaploid wheats grown for 9 d in nutrient solution Data represent means±SD of five replicates.

 
Wheat species and subspecies differed in uptake and export of ¹⁰⁹Cd. On average, diploid wheats absorbed and translocated more ¹⁰⁹Cd than tetraploid and hexaploid wheats (Table 2). However, there were large variations in the uptake and translocation of ¹⁰⁹Cd among and within wheat species. The accession of T. monococcum, ‘FAL-67’ demonstrated the highest uptake of ¹⁰⁹Cd of all tested accessions and cultivars. The highest variation in uptake of ¹⁰⁹Cd was found within tetraploid wheats. Primitive tetraploid wheats took up much higher amounts of ¹⁰⁹Cd when compared to modern tetraploid wheats. Furthermore, among the modern tetraploid wheats, C-1252 tended to take up more ¹⁰⁹Cd than Balcali (Table 2).

View this table: [in this window] [in a new window]   Table 2 Total absorption and accumulation of 109Cd in the treated leaf, remainder of shoot and roots of different diploid, tetraploid and hexaploid wheats grown for 9 d in nutrient solution Radiolabelled Cd was applied when the plants were 7-ds-old and continued over a 42 h period. Data represent means ±SD of five replicates.

 
Irrespective of the wheat genotypes, greater amounts of retranslocated ¹⁰⁹Cd were found in roots compared to the remainder of the shoot (Table 2). However, there was considerable differences in retranslocation of absorbed ¹⁰⁹Cd among and within wheat species. When compared to other wheats, diploid wheats showed, on average, the highest amount of retranslocated ¹⁰⁹Cd in the root and the remaining parts of the shoot.

The amount of retranslocated ¹⁰⁹Cd from the treated leaf into the root and the remainder of the shoot was poorly related to total absorption (Table 2). For example, among the T. monococcum accessions FAL-67 and FAL-43 had the highest and lowest ¹⁰⁹Cd absorption per unit of dry weight, respectively; but, FAL-43 transported a greater amount of ¹⁰⁹Cd into the rest of shoot and roots. Similarly, hexaploid wheat cultivars BDME-10 and Bezostaja did not greatly differ in their absorption of ¹⁰⁹Cd, but the cultivar BDME-10 contained 3.5-fold more ¹⁰⁹Cd in the remainder of the shoots and 4-fold more ¹⁰⁹Cd in roots (Table 2). The relationship between the amounts of translocated ¹⁰⁹Cd into roots and shoots were significantly (P<0.001) correlated (Fig. 1).

View larger version (14K): [in this window] [in a new window]   Fig. 1. The relationship between the accumulation of 109Cd in roots and shoots in different genotypes of diploid, tetraploid and hexaploid wheats grown for 9 d in nutrient solution. Radiolabelled Cd was applied when the plants were 7-d-old, and continued for a 42 h period. Data represent mean of five replicates.

 
The distribution of total ¹⁰⁹Cd (as a percentage per organ) is presented in Table 3. Irrespective of wheat species, about 5% of total ¹⁰⁹Cd absorption was retranslocated from the treated leaf to other plant parts. The highest percentage of retranslocated ¹⁰⁹Cd was found in roots. Among all the genotypes tested, T. monococcum accession, FAL-43, and hexaploid wheat cultivar, BDME-10, showed the highest proportion of ¹⁰⁹Cd in the roots, namely 7.4% and 6.6%, respectively, while the tetraploid wheat cultivar, Balcali, showed the lowest proportion of retranslocated ¹⁰⁹Cd both in the remainder of the shoot (0.6%) and in the roots (1.7%) (Table 2).

View this table: [in this window] [in a new window]   Table 3. Relative distribution (percentage of total activity of 109Cd per organ) of absorbed 109Cd in the treated leaf, remainder of shoot and roots of different diploid, tetraploid and hexaploid wheats grown for 9 d in nutrient solution 109Cd was applied to mature leaf when plants were 7-d-old, and treatment was continued over a 42 h period. Data represents mean±SD of five independent replicates.

 

	Discussion

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The results presented in this study demonstrate that Cd can be translocated from the source leaf to phloem sink organs (i.e. new leaf and root tissues). This result is in agreement with the results of Welch et al. (Welch et al., 1999). Using a split-root technique, these authors reported that Cd can move from one root to other roots (Welch et al., 1999). In addition, previous reports have shown that Cd deposited on leaf surface from atmospheric sources can be taken up by leaf tissue and transported within the plants (Hovmand et al., 1983), supporting the results presented in this study.

Absorption of leaf-applied ¹⁰⁹Cd and its distribution within the plant greatly differed among and within diploid, tetraploid and hexaploid wheats (Tables 2, 3). Possibly, differences in structure and composition of the cuticle and outer epidermal cell walls could be a major determinant in genotypic variation in ¹⁰⁹Cd uptake. Using the bread and durum wheat cultivars given in Tables 2 and 3, studies from this laboratory have recently found that genotypic differences in the uptake of leaf-applied ⁶⁵Zn were similar to those found with ¹⁰⁹Cd. Uptake of leaf-applied ¹⁰⁹Cd and ⁶⁵Zn was markedly higher in bread, when compared to durum wheat cultivars (B Erenoglu, unpublished results). These results emphasize the importance of leaf surface characteristics on the uptake of leaf-applied divalent cations. Similarly, it has been reported that the uptake of leaf-applied B significantly differed among different fruit trees, and this genotypic variation was ascribed to differences in leaf surface characteristics (Picchioni et al., 1995). Variation in the uptake rate of ¹⁰⁹Cd across the plasma membrane of leaf cells may be a further contributing factor to differential ¹⁰⁹Cd uptake between the genotypes tested in this study. These points need to be clarified in future studies.

Neither the absolute amount (Table 2) nor the relative distribution (Table 3) of the retranslocated ¹⁰⁹Cd were related to the total absorption of ¹⁰⁹Cd by the treated leaf. For example, bread wheats were somewhat similar in total uptake, but differed in the capacity to retranslocate ¹⁰⁹Cd. Also among the T. monococcum accessions, FAL-43 had the lowest uptake capacity for ¹⁰⁹Cd, but was able to translocate the greatest amount of ¹⁰⁹Cd from the treated leaf (Tables 2, 3). These results can be interpreted in terms of genotypic variation in the retention of ¹⁰⁹Cd within tissues of the treated leaf. It is possible that differences in retention and retranslocation of ¹⁰⁹Cd among the genotypes studied can be attributed to differences in the capacity of cell walls to bind ¹⁰⁹Cd in the treated leaf. The importance of cell wall binding of Cd and limiting its translocation into shoot via the xylem is well known for root cells (Wagner, 1993; Grant et al., 1998; and references therein). In different plant species (Guo et al., 1995) and genotypes of a given species (Florjin and Van Beusichem, 1993) there is a large variation in the retention of Cd in root cells, possibly by binding to cell walls. Enhanced production of phytochelatins (PC) in response to Cd application is well documented as a defence against Cd toxicity, particularly in roots (Zenk, 1997). Phytochelatins stimulate vacuolar storage of Cd in roots and prevents xylem transport of Cd from roots into the shoot. It is likely that genotypic variation in binding to leaf cell walls and/or binding to phytochelatins within cells of the leaf receiving ¹⁰⁹Cd may explain the differences found. Alternatively, genotypes may differ in the concentration and composition of phloem sap ligands responsible for transporting Cd into sink organs from source organs. The phloem sap contains various type of ligands facilitating the transport of micronutrient cations from source into sink organs, such as phytosiderophores (or phytometallophores), nicotianamine and citric acid (Welch, 1995). Presently, there is no information available concerning the chemical forms and transport mechanisms of Cd movement in the phloem sap.

Atmospheric deposition directly on leaf surfaces has been considered a major factor in increasing Cd concentration in leaves or grains (Hovmand et al., 1983; Jones et al., 1992; Nicholson and Jones, 1994). In view of the results given in Tables 2 and 3, it is unlikely that the higher Cd accumulation in grains of durum than bread wheats (Meyer et al., 1982) is related to a greater capacity of durum wheats to take up and export Cd deposited on the leaf surface from atmospheric fallout. The absence of the D genome in tetraploid wheats (BBAA) may suggest that the D genome in bread wheats (BBAADD) plays a role in reducing the accumulation of Cd in grains of bread wheats (Meyer et al., 1982). Recently, it has been shown that the D genome carries genes affecting the expression of a high Zn efficiency of synthetic wheats when grown under Zn-deficient conditions (Cakmak et al., 1999b). In the present study there was no evidence about a specific genomic effect on uptake and export of ¹⁰⁹Cd in diploid (AA), tetraploid (BBAA) and hexaploid (BBAADD) wheats. Apparently, uptake and transport of Cd via leaves are controlled by different genes located on different chromosomes of the wheat genomes studied. In addition, there was no relationship between Zn efficiency and uptake and the retranslocation capacity of genotypes. For example, bread wheat cultivar ‘BDME-10’ and durum wheat cultivars ‘C-1252’ and ‘Balcali’ are highly sensitive to Zn deficiency (Cakmak et al., 1998, 1999a), but were distinctly different in uptake and retranslocation of leaf-applied ¹⁰⁹Cd (Tables 2, 3).

In conclusion, these results indicate the existence of substantial genotypic variation in the uptake and retranslocation of Cd when it is applied to or deposited on leaf surfaces from aerial contamination. Depending on genotypes, Cd shows relatively high phloem mobility. Future research will be directed at determining if genotypes with low and high Cd retranslocation capacity from mature leaves, as shown in this study, have correspondingly low and high Cd accumulation capacity in grains. Information about the level of Cd phloem translocation from source to sink organs at the seedling growth stage may be useful in screening for wheat genotypes with reduced ability to accumulate Cd in grain

	Notes

 
³ To whom correspondence should be addressed. Fax: +90 322 3386747. E-mail:cakmak{at}mail.cu.edu.tr

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