Cd uptake by intact wheat plants

Abstract. An investigation of the uptake of Cd by intact wheat plants has been carried out. The accumulation of Cd in the roots as a function of time and external Cd concentration was determined. Since the transport of Cd to the grains and shoots was found to be small, only uptake into the root was examined further. An appreciable fraction of the Cd absorbed by the roots can be desorbed and consequently constitutes the freely diffusible and exchangeable fraction.
The effect of temperature and of metabolic inhibitors on the absorption provides evidence that another Cd fraction is taken up by metabolically mediated processes. The absorption of this Cd fraction was found to be inhibited by Cu and Zn. The inhibitory effect of Cu and Zn on Cd absorption was comparable.     

Nature and magnitude of calcium uptake by excised roots of Vetch and Barley

Summary Generally, Ca uptakes were higher and K uptakes lower for excised vetch roots as opposed to those of barley under similar conditions. This behavior was considered to be related to root ion exchange properties and Dorman theory, since the ion exchange capacity of vetch roots is considerably higher than that of barley. Metabolic inhibitors were very effective against K uptakes in contrast to Ca uptakes by both vetch and barley indicating that Ca uptake was primarily non-metabolic under these experimental conditions. The rapidity with which most of the Ca previously taken up was removed by a Ba solution suggests that, for short time periods, a quasi-reversible ion exchange relationship existed between Ba and Ca. Fundamental differences between Ca and K washout curves are attributed to this primarily non-metabolic process. It was concluded that uptake of Ca by excised vetch and barley roots is basically of a surface nature.Senior author currently Assistant Plant Physiologist at the Connecticut Agricultural Experiment Station, New Haven, CT, 06504.     

The Relation between the Rate of Nutrient Uptake by Excised Barley Roots and their Content of Sucrose and Reducing Sugars

A re-examination of the results of previous experiments hasshown that the positive regression of the rate of nutrient uptakeby excised barley roots on sugar content is attributable tothe reducing sugar fraction. Partial regression coefficientsof the rates of uptake of potassium, nitrogen, and phosphoruson reducing sugar content were all positive; those on sucrosecontent were either not significant or negative. These results were confirmed for potassium by experiments inwhich the sugar content of the roots was varied by varying thenitrogen supply to the plants from which the roots were taken,by shading them or by supplying sucrose to the solution in whichthe excised roots were held.     

Characteristics of Zinc Uptake by Barley Roots

The linear uptake of zinc by excised barley roots (Hordeum distichon L.) in the time range from 20–120 min is not continued over periods of 20–28 h. In concentration dependent uptake experiments with intact barley roots three phases in the range up to 1.38 mM zinc could be detected independent of the tested uptake period. The kinetic constants increased with higher phases and the transition points were lowered with increasing time. The presence of copper did not inhibit the uptake of zinc competitively. On the contrary a slight stimulation in the uptake rates was observed indicating an interaction with the transition site. It is concluded that zinc and copper are taken up by separate mechanisms.     

THE EFFECT OF BENZIMIDAZOLE ON CATION UPTAKE BY PLANT ROOTS

Klingensmith , M. J. (Colgate U., Hamilton, N.Y.) The effect of benzimidazole on cation uptake by plant roots. Amer. Jour. Bot. 48(8): 711–716. Illus. 1961.—A number of benzazole compounds were examined for their effect on cation uptake. Benzimidazole was found to almost double the uptake of potassium by excised barley roots in a 6-hr period. Chlorobenzimidazole also enhanced the absorption of potassium but not to the same extent. This stimulation of potassium accumulation was found to be insensitive to cyanide but was dependent on the temperature of the ambient solution. There was also an increased accumulation of potassium by roots of intact barley plants with benzimidazole treatment without interference with subsequent transport of the potassium. Benzimidazole also stimulated uptake of sodium and calcium by excised barley roots but not at identical levels. Results are discussed in the light of various theories of ion absorption.     

Ion-exchange aspects of toxic metal uptake by Indian mustard

Uptake of lead (Pb), copper (Cu), zinc (Zn), and cadmium (Cd) as +2 ions by excised roots of Indian mustard was demonstrated to be an ion-exchange process with existing Ca or protons released to the solution. This initial reaction at the root-aqueous interface is important in the uptake of these toxic metals from contaminated soil. Ethylene diamine tetraacetic acid (EDTA)-amended soil for phytoremediation has Pb in anionic form as [Pb-EDTA]2-, which was not taken up by excised roots. In nonliving B. juncea, Pb2+ was translocated from a solution through a cut stem to petiole and leaves much more quickly than anionic [Pb-EDTA]2-. However, in living plants [Pb-EDTA]2- was more quickly translocated from a solution through roots and petiole to leaves than Pb2+. The final amount of uptake on roots of the living plants was the same for both forms of Pb. The present results are important toward understanding the mechanism of phytoremediation of toxic metal-contaminated soil for two reasons: 1) the initial process, uptake of metal ions by roots, was shown to occur by cation exchange and 2) since [Pb-EDTA]2- was not sorbed by excised roots, other factors such as transpiration and active transport are important in applications using EDTA-amended soils contaminated by Pb.     

Effect of Removal of a Part of the Root System on the Subsequent Growth of the Root and Shoot

Removal of up to 50 per cent. of the roots of barley and ryehas no effect on the growth-rate of the root which is the sameas in the intact plant. In contrast the growth-rate of the shootdecreases as more roots are removed. When more than 50 per cent.of the roots are removed, root growth declines but not so rapidlyas that of the shoot. Similar results are obtained by the removalof lateral roots of tomato but root growth begins to declinewhen 40 per cent. of the lateral roots are removed. The uptake of potassium by barley plants with proportions ofthe root system excised is closely proportional to the dry-matterincrease when the nutrient supply is not limiting. In conditionsof low nutrition the potassium uptake is less than the dry-matterincrease.     

Efficiency of the first steps of sulfate utilization by maize hybrids in relation to their productivity

The genetic variability of the efficiency of the first steps of sulfate utilization and its correlation with productivity were evaluated in nine maize hybrids. ³⁵SO₄²⁻ uptake by excised roots, uptake by intact plant roots, translocation to leaves, and ATP sulfurylase in leaves were taken into account. Uptake rate by roots of intact plants did not show any pulse within 7 to 12 days from emergence, in contrast with the previously observed behaviour of excised roots during root elongation. The uptake rate of intact plants was positively correlated with that of excised roots, but the variability within the nine genotypes tested was less. Productivity was positively correlated with sulfate uptake by both intact plant and excised roots, the level of significance being higher in the first case. Translocation to leaves and ATP sulfurylase activity were not correlated to productivity. Therefore, in the case of sulfate, the grain yield of commonly cultivated maize hybrids appeared to be controlled more by the root uptake step than by the activation and translocation steps.     

Boron uptake by excised barley roots: I. Uptake into the free space

At 2 C, all boron accumulated by excised barley roots (Hordeum vulgare L. cv. Herta) remains in the free space; i.e. active uptake is nil at this temperature. Three component fractions of free space B were apparent: (a) a surface contaminant film of B on blotted roots, (b) water free space B, and (c) B reversibly bound in the cell walls. A stoichiometric release of H⁺ from the roots in the presence of B indicated that B was bound by borate complexes with polysaccharides in the cell walls. Polysaccharide-borate complexes are much less stable than those of monosaccharides, and the bound B fraction could be readily removed by rinsing the roots in the presence of a monomeric polyol possessing the necessary cis-diol configuration. Cell wall material separated from excised barley roots had a B binding capacity 66% greater than that of intact roots.     

Further studies on cation uptake from bentonite suspensions by excised barley roots

Summary The net uptake of Na, K, Li, and Ca or Mg by excised barley roots was studied from bi-ionic and tri-ionic bentonite suspensions. The net uptake of Li from Li-Ca system progressed linearly with progressive Li levels and was related to the concentration of soluble lithium. Calcium in this system was taken up only at the 100 per cent Ca level. At lower Ca levels calcium was lost from the roots to the suspensions. In K-Mg and Na-Mg systems the net uptake of Na or K by the excised roots was related to the concentration of the cation in the solution phase. Magnesium uptake took place at 80 and 100 per cent Mg levels. It was much less than that of K or Na at similar levels. At lower levels of Mg the roots lost some of their initial Mg contents to the suspensions. In the Na-K-Mg system magnesium was not taken up by the excised roots. Sodium uptake was not practically affected by the Mg level, but K uptake was greatly enhanced by magnesium.     

A Comparison of the Uptake and Translocation of Some Organic Herbicides and a Systemic Fungicide by Barley: II. RELATIONSHIP BETWEEN UPTAKE BY ROOTS AND TRANSLOCATION TO SHOOTS

The relationship between uptake by barley roots and translocationto shoots has been examined under different conditions for sixherbicides and a systemic fungicide (four triazines, diuron,2,4-dichlorophenoxyacetic acid and ethirimol). For all thesecompounds a large proportion of the material taken up by rootsdoes not appear to move readily to the shoots. Elution withwater of the roots of intact plants which had previously beenplaced in labelled solutions of these compounds enabled thematerial released to be separated into three fractions differingin their rates of diffusion out of the roots. There was reasonablygood correlation between the concentration of the most readilydiffusible fraction in the roots and the concentration in thetranspiration stream. The results obtained were consistent withthe postulate that lipophilic compounds can diffuse into thevacuoles of the cortical cells where they are available fortransport to the shoots, whereas for lipophobic compounds materialreaching the shoots originates largely from the free space inthe roots.     

Effect of associative bacteria on element composition of barley seedlings grown in solution culture at toxic cadmium concentrations

The response of barley seedlings to inoculation with associative rhizobacteria Azospirillum lipoferum 137, Arthrobacter mysorens 7, Agrobacterium radiobacter 10 and Flavobacterium sp. L30 was studied in hydroponic and quartz sand cultures in the presence of 50 microM CdCl2. Cadmium caused severe inhibition in the growth and uptake of nutrient elements by the plants. Inoculation with the bacteria slightly stimulated root length and biomass of hydroponically grown Cd-treated seedlings. The bacteria increased the content of nutrients such as P, Mg, Ca, Fe, Mn and Na in roots and or shoots of the plants grown in the absence of Cd. Positive changes in the element composition caused by the bacteria were less pronounced in Cd-treated plants, whereas the total amount of nutrients taken by the inoculated plants was generally increased significantly. The content of Cd in the inoculated plants was unchanged, except increased in roots upon addition of A. lipoferum 137. Inoculation did not affect the activity of peroxidase, alpha-mannosidase, phosphodiesterae, alpha-galactosidase, and concentration of sulfhydryl compounds used as biochemical markers of stress in plant roots. The results showed that associative bacteria were capable of decreasing partially the toxicity of Cd for the barley plants through the improvement in uptake of nutrient elements.     

Effect of calcium on the absorption and translocation of heavy metals in excised barley roots: Multi-compartment transport box experiment

Summary The effect of Ca on the absorption and translocation of Mn, Zn and Cd in excised barley roots was studied using a multi-compartment transport box technique. A radioisotope (⁵⁴Mn,⁶⁵Zn or^115mCd)-labelled test solution was supplied to the apexes of excised roots and the distribution pattern in the roots was examined in the absence or presence of Ca. Results obtained were as follows. Addition of Ca to the test solution reduced the absorption of Mn and inhibited drastically its translocation in excised roots. With increasing concentrations of Ca in test solutions, its inhibitory effects on the absorption and translocation of Mn became severe. Similar results were observed for the absorption and translocation of Zn. Ca in the test solution decreased the absorption and inhibited drastically the translocation of Zn; as in the case of Mn, higher concentrations of Ca had severe effects on these functions. It was also evident that the addition of Ca to the test solution reduced the absorption of Cd at all levels of Cd concentration (1, 10, and 100 μM). Cd absorption decreased with increasing concentrations of Ca in the test solution. However, Ca accelerated the translocation of Cd in excised roots supplied with test solutions containing up to 10μM Cd. At 100μM Cd, addition of Ca caused a negligibly small acceleration of Cd translocation. The accelerating effect of Ca on Cd translocation, especially “xylem exudation”, decreased markedly with the addition of 2,4-dinitrophenol, but not with the addition of chloramphenicol or p-chloromercuribenzene sulphonic acid. When barley plants were supplied with only CaSO₄ during the entire growing period, that is, plants were not supplied with nutrient solution on the last day of this period, Ca had no accelerating effect on Cd translocation in excised roots.     

Effect of cadmium ion excess over cell structure and functioning of Zea mays and Hordeum vulgare

Zea mays (maize) and Hordeum vulgare (barley) plants were analyzed in order to study the variation in response to Cadmium (Cd) toxicity based on development of leaf symptoms, effect in dry matter production, Cd uptake, lipid peroxidation and effect on cell ultrastructure in leaves and roots. Cd accumulation in roots of Z. mays and H. vulgare was 18–50 times higher than in the aerial parts. Malondialdehyde (MDA) content was more affected in the roots of both Z. mays and H. vulgare than in shoots (60 and 56–51 and 40%, respectively). At ultrastructural level, in Cd treated seedlings, a decline in the vacuolar content of barley roots cells and maize leaf cells was observed. Results corroborate that these gramineous crops can uptake and accumulate substantial amounts of Cd especially in roots. Therefore, H. vulgare and Z. mays could have a phytostabilization potential and thereafter could be tested in phytoremediation technologies.     

ON THE EFFECTS OF DMSO IN CATION TRANSPORT BY EXCISED BARLEY ROOTS

Dimethyl sulfoxide (DMSO) in concentrations of up to 10% by volume stimulates the uptake of zinc by excised barley roots. In the same concentration it severely depresses uptake of sodium and of rubidium. It does not seem to affect the permeability of the membrane since roots treated with desorption solutions which were 10% in DMSO did not lose more of the preferred ion than did roots desorbed in solutions not containing DMSO. Oxygen utilization (measured in the Warburg respirometer) was reduced when DMSO was present. It is suggested that DMSO is a poisoning agent which interferes with cation transport by attacking some aspect of metabolism and not by influencing the permeability of the membrane.     

Influence of Potassium and Nitrogen Fertilization on Parasitism by the Root-knot Nematode Meloidogyne javanica

The influence of various c oncentrations of K⁺, nitrogen sources, and inoculation with root-knot nematode Meloidogyne javanica were evaluated in tomato plants. Increased potassium concentration increased top and root fresh weights of intact plants and fresh weights of excised roots. Nitrate-fertilized plants weighed more than plants receiving ammonium independent of the K level in the medium. Nematode counts on roots were not affected by nutritional differences in intact or excised roots. In intact roots a high percentage of males was recorded at low K⁺ levels, whereas in excised roots the proportion of males in the population rose as the K⁺ levels increased. Inoculated intact roots accumulated K⁺ when the level of potassium supply was low; infected excised roots contained less K⁺ than did nematode-free roots.     

Impacts of plant roots on soil CO cycling and soil–atmosphere CO exchange

Carbon monoxide (CO) plays a major role in tropospheric chemical dynamics. Accordingly, global CO budgets have been reasonably well documented. Atmospheric CO consumption by soils contributes significantly to these budgets, with the magnitude of the sink generally considered to reflect a balance between microbial uptake and abiological production. However, assays of live fine roots showed that diverse intact plants produced carbon monoxide at net rates ranging from 2 to 3000 mµg gdw⁻⁻¹ d⁻⁻¹. CO production was greater for legumes than nonlegumes, and primarily associated with nodules. Excised roots from woody and herbaceous plants produced CO at comparable rates. CO production rates were similar for roots of intact plants and roots excised from those plants. The magnitude of net CO fluxes from roots was determined in part by the balance between simultaneous production and consumption. Surface sterilization of roots indicated that CO consumption was due, in part, rhizoplane CO-oxidizing bacteria, but maximum CO consumption rates were typically only a small fraction of net production rates. Assays in a Brazilian agroecosystem indicated that root CO production affects soil–atmosphere CO exchange. Estimates of global CO production rates indicated that roots contribute about 170–260 Tg CO to the soil atmosphere annually, an amount comparable to current estimates of atmospheric CO uptake by soils, and much larger than estimates of net abiological soil CO production.     

Bacteria-mediated uptake of choline sulfate by plants. Bacterial effectiveness

The ability of bacteria to cause rapid uptake of choline sulfate in plants, i.e., effectiveness, was studied using Pseudomonas tolaasii and excised roots of barley (Hordeum vulgare L.). Once effective, bacteria remained so after being killed by treatments which cause little damage to their outer structure. However, effectiveness was destroyed by disruption of the cell wall, protein reagents, a mild heat treatment or removal of Mg²⁺. Effective bacteria adsorbed choline sulfate. This adsorption had characteristics similar to those of bacterial effectiveness (magnesium requirement, high substrate specificity). These results indicate that a proteinaceous structure on the bacterial surface binds and, somehow, transfers choline sulfate to the plant.     

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.     

Absorption and translocation of Cd in bush beans (Phaseolus vulgaris)

A series of experiments was conducted to examine some factors affecting the absorption and translocation of Cd in young bean plants ( Phaseolus vulgaris L. cv. Bulgarian). Absorption of Cd by roots was reduced in the presence of other cations of increasing valency or ionic radii. Reduced absorption was also found in the presence of EDTA. Concentration of Cd in exudates from excised stems increased with increased passage of Cd solutions and approached the concentration in the external medium (4.5 μ M Cd). This was apparently associated with saturation of adsorption sites in the stems. The stem behaved as a cation exchange column resulting in a chromatographic distribution of Cd towards the top of the plant. These experiments indicate that Cd existed in the xylem fluid as a free or weakly complexed cation. Additional experiments showed that the total amount of Cd absorbed by bean plants was elevated by inducing higher transpiration rates. The effect of water flux on Cd transport indicated apoplastic flow to the stele.