Effects of Eu and Ca on yield and mineral nutrition of wheat (Triticum aestivum) seedlings
Introduction
Rare earth elements (REE) are widely distributed in the environment. Despite of the name they are not quite rare in the nature. The abundance of cerium in soil is almost equal to that of copper and zinc. The abundance of the most rare REE, lutecium and thulium in the majority of soils is comparable to that of cadmium and selenium (Tyler, 2004). While REE are abundant in soil, their concentration in plants is usually low (Fu et al., 2001, Zhang et al., 2002, Tyler, 2004). The biogeochemical behaviour of REE in the soil–plant system is not fully understood. Until recently, REE have not been characterized as essential plant nutrients and environmentally hazardous metals. It is quite possible that both toxic and beneficial effects of REE are similar to those of other trace metals.
REE have been used in agriculture since the seventies of last century. Particularly in China REE-based fertilisers are widely used to increase the yield and quality of crops (Xiong, 1995, Zhang and Shan, 2001, Xu et al., 2002, Shan et al., 2005). Pot and field experiments have been performed to demonstrate beneficial influence of REE on plant growth and soil properties. Interestingly, beneficial effects of REE on plants and soils have been reported mainly by the Chinese researchers. It was found that REE stimulate the synthesis of chlorophyll (Guo, 1988), promote seedling development (Wu et al., 1983, Chang, 1991), and improve the bioavailability of Ca and Mn in soil (Chang, 1991).
Notice however that reported data on field and laboratory tests is often contradictory and inconsistent. For example, Maheswaran et al. (2001) studied the effects of lanthanum (La) and cerium (Ce) on seed germination using various rates ranging from 0 to 0.16%. They reported that both shoot and root lengths were not improving with an application of REE. Application of La or Ce to the shoots or roots generally reduced corn and mungbean growth rate (Diatloff et al., 1999). The authors concluded that in view of toxicity of REE to these crop plants, it seemed unlikely that previously reported beneficial effects could result from direct influence of these elements on the growth processes of a particular plant. Such beneficial action may result from various indirect effects that influence agronomic performance of crop plants. Moreover, it seems that positive response is possible when the soils contain low quantities (less than 10 mg kg−1) of available REE (Maheswaran et al., 2001). These contradictory observations may also be explained by different levels of applied REE. The inhibition may result from disturbances in enzymatic mechanisms (peroxidase and catalase enzymes) involved in the elimination of hydrogen peroxide in a plant cell (Nardi et al., 2004).
A considerable number of publications are dedicated to REE distribution in different cereal crops (Wu et al., 1983, Dong et al., 1992, Liu et al., 1997, Wang et al., 2001, Wang et al., 2004, Zhang et al., 2002, Chu et al., 2003). Concentrations of La and Ce in plants are 10–100 times higher than concentrations of other REE. Probably, this is the reason why so many publications are devoted to these elements in comparison to other REE.
Europium (Eu) is one of the most interesting REE. In different environmental conditions Eu can change its valence state and thus, can serve as an indicator of biogeochemical processes. There have been few reports on the behaviour of Eu in different plant species as well as on beneficial or toxic effects of this element on plants.
Calcium is an essential plant nutrient. As a divalent cation (Ca2+) it is required for numerous processes in higher plant cells (White and Broadley, 2003). REE ions having effective ion radius close to that of Ca2+, may replace Ca or interact with it in the variety of physiological functions (Tyler, 2004). Data on the interference of REE with calcium metabolism were reported (Pickard, 1970, Rengel, 1994). In particular, after an initial sorption of REE to the cell wall they may be transported into the cell through Ca channels and/or by Ca transporters, thus, blocking Ca2+ uptake (Cheng et al., 1999, Weltje, 2002). Tammeba et al. (2004) showed that Ca uptake can be completely inhibited by a voltage-dependent Ca channel antagonist like La3+. Diniz and Volesky (2005) found that the uptake of REE was accompanied by simultaneous release of Ca ions. On the other hand, it was shown that REE can stimulate Ca2+ influx in various animal cells (Cheng et al., 1999). However, the mechanism of the stimulation is still unclear. Nevertheless, this points to an active Ca–REE exchange.
Since Eu may be competitive with Ca for organic ligands, it would be of interest to assess interactions between Ca and Eu in plants. Because seedling stage is thought to be crucial throughout the whole life of a plant, our efforts were concentrated on the initial stages of plant development. The question addressed in this study is whether an increase of Eu, Ca and a combination of the two elements in the growth medium can influence the plant yield and uptake of Eu, Ca and other elements by wheat seedlings during first days of plant growth.
Section snippets
Materials and methods
Seeds of Triticum aestivum wheat were germinated for 4 days on a filter paper wetted with distilled water. The seeds were divided into ten groups. Ca(NO3)2 (1 g L−1) was added to the germination medium on first (group 1), second (group 2) and third (group 3) day of seed germination. Eu(NO3)3 (0.01 mg L−1) was added to the germination medium on first (group 4), second (group 5) and third (group 6) day of germination. A mixture of calcium nitrate and europium nitrate (in the same concentrations that
Soil
In our experiment, pH of the soil was rather stable (6.1). Mean concentrations of elements in soil are shown in Table 1. No statistically significant difference in the soil element concentrations between different pots was observed.
Four-day-old seedlings
All 4-day-old wheat seedlings germinated in Eu- and Ca-rich media looked normal and healthy. Moreover, an addition of Eu to the germination medium appeared to be favourable for the development of seedling roots. While leaf mass of the 4-day-old seedlings was
Conclusions
Wheat seedlings germinated in Eu-rich medium are able to accumulate large amounts of Eu. The rates of Eu uptake and translocation from roots to the upper parts of a plant varied with time and depended on the stage of the plant development. Eu concentration was significantly reduced during the plant growth in normal soil. We suggest that Eu release out of wheat seedlings is connected with water evaporation. An increase of Eu content in a plant affected plant yield and concentrations of
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