Cadmium uptake kinetics and plants factors of shoot Cd concentration

Background and aims

Accumulation of Cd in the shoots of plants grown on Cd contaminated soils shows considerable variation. A previous preliminary experiment established that one major reason for this variation was the rate of Cd influx into the roots (mol Cd cm^?2 root s^?1). However, this experiment did not distinguish between solubilization of soil Cd on the one hand and difference in Cd uptake kinetics on the other. The main objectives of the present study were thus to characterize Cd uptake kinetics of plants continuously exposed to Cd concentrations similar to those encountered in soils. Furthermore we determined the factors responsible for differences in shoot Cd concentration such as net Cd influx, root area-shoot dry weight ratio, shoot growth rate and proportion of Cd translocated to the shoot.

Materials and methods

Maize, sunflower, flax and spinach were grown in nutrient solution with five constant Cd concentrations varying from 0 to 1.0 μmol?L^?1. Root and shoot parameters as well as Cd uptake were determined at two harvest dates and from these data Cd net influx and shoot growth rates were calculated.

Results and conclusions

Cadmium uptake kinetics, i.e. the net Cd influx vs. Cd solution concentration followed a straight line. Its slope is the root absorbing power, α, $ \left( {\alpha ={{{\mathrm{Cd}\;\mathrm{net}\;\mathrm{influx}}} \left/ {{\mathrm{Cd}\;\mathrm{solution}\;\mathrm{concentration}}} \right.}} \right) $ . The α values of spinach and flax were about double that of maize and sunflower (5?×?10^?6?cm?s^?1 vs. 2.5?×?10^?6?cm?s^?1). Spinach and flax had a 3–5 times higher shoot Cd concentration than maize and sunflower. The difference in shoot Cd concentration was partly due to the higher Cd influx but also to a higher translocation of Cd from root to shoot and also to a slower shoot growth rate.     

Maize plant growth and accumulation of photosynthetic pigments at short- and long-term exposure to cadmium

A wide range of cadmium concentrations (from 4 to 200 μM for seedlings and up to 2 mM for germinating kernels) was used to assess Cd toxic effects on maize (Zea mays L.) plants at the different developmental stages: germinating kernels, seedlings (4–9 days), and juvenile plants (34 days). Cd accumulation in plant organs was followed, and its lethal concentration was elucidated. In maize, cadmium was accumulated predominantly in roots; in shoots it was mainly accumulated in the lower leaves, and the higher was leaf position the lower was Cd content in it. At high concentrations (80 and 200 μM), kernels became the substantial cadmium depot. Germinating kernels manifested the lowest sensitivity to cadmium; seedlings were more sensitive; the inhibition of juvenile plant growth attained 90% and more. In the tested range of concentrations, cadmium suppressed shoot mass accumulation harder than that of roots. In 34-day-old plants, water content in shoots was stronger reduced than in roots. Plant death was also manifested earlier in shoots. It was concluded that maize plant sensitivity to cadmium increases with plant growing and that, under conditions of normal mineral nutrition, cadmium inhibits shoot growth more severe than root growth.     

不同镉水平下大麦幼苗生长和镉及养分吸收的品种间差异

利用水培试验研究了不同Cd水平下大麦幼苗的Cd和几种矿质元素吸收、积累、生长和生物学产量的品种间差异 .结果表明 ,1μmol·L-1Cd处理显著降低麦苗株高、绿叶数、叶绿素计读数、地上部和根系干重 ,显著抑制植株对Zn、Mn、Cu的吸收和累积 ;品种之间存在着显著差异 ,无芒六棱受抑制最为严重 ,米麦 114和浙农 1号表现出相对较强的抗性 .麦苗Cd含量和累积量品种之间也有显著差异 ,浙农 1号的Cd含量最高 ,米麦 114最低 .相关分析表明 ,麦苗生物学产量与地上部Cd含量、累积量及根系Cd含量呈显著负相关 ,其中与地上部Cd含量的相关性最强 ,与根系Cd累积量无显著相关 .     

Cadmium uptake by roots: Contribution of apoplast and of high- and low-affinity membrane transport systems

The aim was to measure the respective contributions of apoplast and symplast to the Cd root uptake and to explain the linear component of the symplastic absorption. Two plants were used, maize (Zea mays L.) and two ecotypes of alpine pennycress (Noccaea caerulescens (J. Presl & C. Presl) F.K. Mey.), with contrasted abilities to accumulate Cd. Their roots were exposed to labelled Cd solutions of increasing concentrations. Root Cd was physico-chemically fractioned to obtain the exchanged apoplastic, non-exchanged apoplastic and symplastic pools. For both species, the proportion of Cd retained by the cell walls increased with Cd concentration in the exposure solution (ranging from 0.05 to 50 μmol L^?1), from approximately 30% to 90% of the total root Cd. This was modeled using Freundlich isotherms. The non-exchanged apoplastic Cd was negligible at the highest exposure concentrations, but reached almost 30% of the total root uptake at the lowest ones. The symplastic influx in roots of both species fitted a Michaelis-Menten function associated with a linear one. The linear component of the symplastic influx could reflect absorption through a low-affinity transport system (LATS). The strong adsorption of Cd on root apoplast might act as a driving force to extract the metal from the soil, compete with the symplastic absorption and contribute to the amount of element taken up by the plant, at least in its roots.     

Cadmium distribution in maize inbred lines: Effects of pH and level of Cd supply

In order to investigate the physiological basis of the differential Cd distribution and the degree of variation of this Cd distribution among maize inbred lines, six inbreds designated earlier as ‘shoot Cd excluders’ (B73, H99, and H96) and ‘non-shoot Cd excluders’ (B37, H98, and N28) were grown in nutrient solution culture at different external Cd levels or at different pH. The characterization of the inbreds according to their shoot/root partitioning of Cd was consistent, independent of pH or level of Cd supply. The Cd concentrations in the plants were highest at the highest pH of the solution cultures. Generally, there was a positive correlation between the Cd concentrations in shoots and xylem exudates. It was shown that the Cd concentration in the roots is particularly important in the Cd distribution process. Above a ‘critical’ internal Cd concentration in the roots, specific for each inbred, the ability to retain Cd is strongly diminished. It is concluded that structural and/or physiological characteristics of the roots are involved in Cd partitioning.     

Uptake and long-distance transport of phosphate,potassium and chloride in relation to internal ion concentrations in barley: evidence of non-allosteric regulation

The extent to which uptake and transport of either phosphate, potassium or chloride are controlled by the concentration of these ions within the root, perhaps through an allosteric mechanism, was investigated with young barley plants in nutrient solution culture. Plants were grown with their roots divided between two containers, such that a single seminal root was continuously supplied with all the required nutrient ions, while the remaining four or five seminal roots were either supplied with the same solution (controls) or, temporarily, a solution lacking a particular nutrient ion (nutrient-deficient treatment). Compared with controls, there was a marked stimulation of uptake and transport of labelled ions by the single root following 24 h or more of nutrient dificiency to the remainder of the root system. This stimulation, which comprised an increased transport to the shoot and, for all ions except Cl^-, increased transport to the remainder of the root system, took place without appreciable change in the concentration of particular ions within the single root. However, nutrient deficiency quickly caused a lower concentration of ions in the shoot and the remaining roots. The results are discussed in relation to various mechanisms, proposed in the literature, by which the coordination of ion uptake and transport may be maintained within the plant. We suggest that under our conditions any putative allosteric control of uptake and transport by root cortical cells was masked by an alternative mechanism, in which ion influx appears to be regulated by ion efflux to the xylem, perhaps controlled by the concentration of particular ions recycled in the phloem to the root from the shoot.     

Enhancement of cadmium effects on growth and nutrient composition of birch (Betula pendula) by buthionine sulphoximine (BSO)

Binding of Cd to non-specific metal-binding peptides (phytochelatins)in birch roots has been suggested as an explanation for toleranceto Cd toxicity in birch (Betula pendula). In the present study,the tolerance of birch roots to Cd was further investigatedby using buthionine sulphoximine (BSO) as an inhibitor of phytochelatinsynthesis. Birch seedlings, grown in nutrient solution at pH4.2, were exposed to 0 or 2 µM CdCl₂ combined with 0 or0.1 mM BSO for 6 d. Plant growth (fresh weight increase andshoot to root dry weight ratio) and the nutrient compositionin fine roots, whole roots and shoots were determined. The effectsof Cd on growth confirms the results of earlier studies on birch,suggesting a reduced shoot growth, but preserved or stimulatedroot growth. When Cd and BSO were combined, overall plant growthwas severely reduced. BSO was also shown to aggravate Cd-inducedreductions of root and shoot concentrations of K, Ca and Mgbut to impede the accumulation of Cd. The results suggest that phytochelatins participate in protectingthe root against Cd interferences with growth, possibly by restrictingCd-induced changes in the nutrient composition of the plant. Key words: Betula pendula, buthionine sulphoximine, cadmium, phytochelatins, roots, tolerance     

Potassium uptake efficiency and dynamics in the rhizosphere of maize (Zea mays L.), wheat (Triticum aestivum L.), and sugar beet (Beta vulgaris L.) evaluated with a mechanistic model

Plant species differ in nutrient uptake efficiency. With a pot experiment, we evaluated potassium (K) uptake efficiency of maize (Zea mays L.), wheat (Triticum aestivum L.), and sugar beet (Beta vulgaris L.) grown on a low-K soil. Sugar beet and wheat maintained higher shoot K concentrations, indicating higher K uptake efficiency. Wheat acquired more K because of a greater root length to shoot dry weight ratio. Sugar beet accumulated more shoot K as a result of a 3- to 4-fold higher K influx as compared to wheat and maize, respectively. Nutrient uptake model NST 3.0 closely predicted K influx when 250 mg K kg^?1 were added to the soil, but under-predicted K influx under low K supply. Sensitivity analysis showed that increasing soil solution K concentration (C_Li) by a factor of 1.6–3.5 or buffer power (b) 10- to 50-fold resulted in 100% prediction of K influx. When both maximum influx (I_max) and b were increased by a factor of 2.5 in maize and wheat and 25 in sugar beet, the model could predict measured K influx 100%. In general, the parameter changes affected mostly calculated K influx of root hairs, demonstrating their possible important role in plant K efficiency.     

Long-term effects of exogenous silicon on cadmium translocation and toxicity in rice (Oryza sativa L.)

Most nutrient solution studies on the interactions between silicon (Si) and cadmium (Cd) are short term. Here we reported a long-term experiment in which rice (Oryza sativa L.) was cultured for 105 days and harvested at four different growth stages to measure biomass accumulation and Cd uptake and distribution in shoots and roots. Exogenous Si increased shoot biomass by 61–238% and root biomass by 48–173% when the culture solution was free of Cd. When 2 μmol L^?1 Cd was added, Si supply increased shoot and root biomass by 125–171% and by 100–106% compared to the zero-Si treatment. Increasing the Cd concentration to 4 μmol L^?1 decreased the beneficial effects of Si on root and shoot biomass. Silicon supply decreased shoot Cd concentrations by 30–50% and Cd distribution ratio in shoot by 25.3–46%, compared to the treatment without Si supply. Additionally, lower Si supply or more serious Cd stress would lead to roots with bigger biomass and higher Si concentration. Energy-dispersive X-ray microanalysis showed that both Si and Cd accumulated synchronously in the border and middle of phytoliths of the shoots. We conclude that Si enhances plant growth and decreases Cd accumulation in shoots and thereby helps to lower the potential risks of food contamination.     

Uptake and distribution of cadmium in maize inbred lines

Genotypic variation in uptake and distribution of cadmium (Cd) was studied in 19 inbred lines of maize (Zea mays L.). The inbred lines were grown for 27 days on an in situ Cd-contaminated sandy soil or for 20 days on nutrient solution culture with 10 µg Cd L^-1. The Cd concentrations in the shoots showed large genotypic variation, ranging from 0.9 to 9.9 µg g^-1 dry wt. for the Cd-contaminated soil and from 2.5 to 56.9 µg g^-1 dry wt. for the nutrient solution culture. The inbred lines showed a similar ranking for the Cd concentrations in the shoots for both growth media (r²=0.89). Two main groups of inbreds were distinguished: a group with low shoot, but high root Cd concentrations (shoot: 7.4±5.3 µg g^-1 dry wt.; root: 206.0±71.2 µg g^-1 dry wt.; shoot Cd excluder) and a group with similar shoot and root Cd concentrations (shoot: 54.2±3.4 µg g^-1 dry wt.; root: 75.6±11.2 µg g^-1 dry wt.; non-shoot Cd excluder). The classification of the maize inbred lines and the near equal whole-plant Cd uptake between the two groups demonstrates that internal distribution rather than uptake is causing the genotypic differences in shoot Cd concentration of maize inbred lines. Zinc (Zn), a micronutrient chemically related to Cd, showed an almost similar distribution pattern for all maize inbred lines. The discrepancy in the internal distribution between Cd and Zn emphasizes the specificity of the Cd distribution in maize inbred lines.