Volker Römheld

Different strategies in higher plants in mobilization and uptake of iron

Abstract Higher plants differ considerably in their capability for mobilization of iron in the rhizospere of soils with low iron availability. In the plant kingdom at least two different Strategies exist in the Fe deficiency‐induced root responses which lead to enhancement of both iron mobilization in the rhizosphere and uptake rate of iron. Strategy I is found in all dicots and in monocots, with the exception of the grasses (graminaceous species, e.g. barley, corn, sorghum). Strategy I is characterized in all instances, by an increase in the activity of a plasma membrane‐bound reductase leading to enhanced rates of Fe‐III reduction and corresponding splitting of Fe‐III‐chelates at the plasma membrane. Often, the net rate of H+ extrusion, i.e. acidification of the rhizosphere, is also increased. This acidification facilitates iron uptake by both enhancement of the reductase activity and solubilization of iron in the rhizosphere. An additional mobilization of sparingly soluble iron in the rhizosphere may o...

Physiological adaptations to phosphorus deficiency during proteoid root development in white lupin

Abstract. Release of large amounts of citric acid from specialized root clusters (proteoid roots) of phosphorus (P)-deficient white lupin (Lupinus albus L.) is an efficient strategy for chemical mobilization of sparingly available P sources in the rhizosphere. The present study demonstrates that increased accumulation and exudation of citric acid and a concomitant release of protons were predominantly restricted to mature root clusters in the later stages of P deficiency. Inhibition of citrate exudation by exogenous application of anion-channel blockers such as ethacrynic- and anthracene-9-carboxylic acids may indicate involvement of an anion channel. Phosphorus-deficiency-induced accumulation and subsequent exudation of citric acid seem to be a consequence of both increased biosynthesis and reduced metabolization of citric acid in the proteoid root tissue, indicated by increased in-vitro activity and enzyme protein levels of phosphoenolpyruvate carboxylase (EC 4.1.1.31), and reduced activity of aconitase (EC 4.2.1.3) and root respiration. Similar to citric acid, acid phosphatase, which is secreted by roots and involved in the mobilization of the organic soil P fraction, was released predominantly from proteoid roots of P-deficient plants. Also 33Pi uptake per unit root fresh-weight was increased by approximately 50% in juvenile and mature proteoid root clusters compared to apical segments of non-proteoid roots. Kinetic studies revealed a Km of 30.7 μM for Pi uptake of non-proteoid root apices in P-sufficient plants, versus Km values of 8.5–8.6 μM for non-proteoid and juvenile proteoid roots under P-deficient conditions, suggesting the induction of a high-affinity Pi-uptake system. Obviously, P-deficiency-induced adaptations of white lupin, involved in P acquisition and mobilization of sparingly available P sources, are predominantly confined to proteoid roots, and moreover to distinct stages during proteoid root development.

Integrated soil-crop system management for food security

China and other rapidly developing economies face the dual challenge of substantially increasing yields of cereal grains while at the same time reducing the very substantial environmental impacts of intensive agriculture. We used a model-driven integrated soil–crop system management approach to develop a maize production system that achieved mean maize yields of 13.0 t ha−1 on 66 on-farm experimental plots—nearly twice the yield of current farmers’ practices—with no increase in N fertilizer use. Such integrated soil–crop system management systems represent a priority for agricultural research and implementation, especially in rapidly growing economies.

Boron deficiency-induced impairments of cellular functions in plants

The essentiality of B for growth and development of plants is well-known, but the primary functions of B still remain unknown. Evidence in the literature supports the idea that the major functions of B in growth and development of plants are based on its ability to form complexes with the compounds having cis-diol configurations. In this regard, the formation of B complexes with the constituents of cell walls and plasma membranes as well as with the phenolic compounds seems to be a decisive step affecting the physiological functions of B. Boron seems to be of crucial importance for the maintenance of structural integrity of plasma membranes. This function of B is mainly related to stabilisation of cell membranes by B association with membrane constituents. Possibly, B may also protect plasma membranes against peroxidative damage by toxic O2 species. In B-deficient plants, plasma membranes are highly leaky and lose their functional integrity. Under B-deficient conditions, substantial changes in ion fluxes and proton pumping activity of the plasma membranes were noted. Impairments in phenol metabolism and increases in levels of phenolics and polyphenoloxidase activity are typical indications of B deficiency, particularly in B deficiency-sensitive plant species, such as Helianthus annuus (sunflower). Enhanced oxidation of phenols is responsible for generation of reactive quinones which subsequently produce extremely toxic O2 species, thus resulting in the increased risk of a peroxidative damage to vital cell components such as membrane lipids and proteins. In B-deficient tissues, enhancement in levels of toxic O2 species may also occur as a result of impairments in photosynthesis and antioxidative defence systems. Recent evidence shows that the levels of ascorbic acid, non-protein SH-compounds (mainly glutathione) and glutathione reductase, the major defence systems of cells against toxic O2 species, are reduced in response to B deficiency. There is also increasing evidence that, in the heterocyst cells of cyanobacteria, B is involved in protection of nitrogenase activity against O2 damage.

Research on potassium in agriculture: needs and prospects

This review highlights future needs for research on potassium (K) in agriculture. Current basic knowledge of K in soils and plant physiology and nutrition is discussed which is followed by sections dealing specifically with future needs for basic and applied research on K in soils, plants, crop nutrition and human and animal nutrition. The section on soils is devoted mainly to the concept of K availability. The current almost universal use of exchangeable K measurements obtained by chemical extraction of dried soil for making fertilizer recommendations is questioned in view of other dominant controlling factors which influence K acquisition from soils by plants. The need to take account of the living root which determines spatial K availability is emphasized. Modelling of K acquisition by field crops is discussed. The part played by K in most plant physiological processes is now well understood including the important role of K in retranslocation of photoassimilates needed for good crop quality. However, basic research is still needed to establish the role of K from molecular level to field management in plant stress situations in which K either acts alone or in combination with specific micronutrients. The emerging role of K in a number of biotic and abiotic stress situations is discussed including those of diseases and pests, frost, heat/drought, and salinity. Breeding crops which are highly efficient in uptake and internal use of K can be counterproductive because of the high demand for K needed to mitigate stress situations in farmers’ fields. The same is true for the need of high K contents in human and animal diets where a high K/Na ratio is desirable. The application of these research findings to practical agriculture is of great importance. The very rapid progress which is being made in elucidating the role of K particularly in relation to stress signalling by use of modern molecular biological approaches is indicative of the need for more interaction between molecular biologists and agronomists for the benefit of agricultural practice. The huge existing body of scientific knowledge of practical value of K in soils and plants presents a major challenge to improving the dissemination of this information on a global scale for use of farmers. To meet this challenge closer cooperation between scientists, the agrochemical industry, extension services and farmers is essential.

Mobilization of iron and other micronutrient cations from a calcareous soil by plant-borne, microbial, and synthetic metal chelators

Mobilization of Fe, Zn, Cu, and Mn by various chelators from a calcareous soil was measured using a simple dialysis tube/complexing resin system. Root exudates from Fe-deficient barley increased the concentrations of all four metals in solution by, on average, a factor of 20, and the addition of complexing resin as a sink for heavy metal cations forced steady state solution concentrations to be reached sooner. Root exudates mobilized increasing amounts of the various micronutrients in the following order: Cu<Fe<Zn<Mn. Phytosiderophores isolated from root exudates of Fe-deficient barley mobilized similar amounts of Cu and Zn but somewhat more Fe and considerably more Mn than crude exudate. The synthetic chelators EDDHA and DTPA showed low specificity in micronutrient mobilization, but the microbial siderophore Desferal was relatively more specific, preferentially mobilizing Fe and Mn. The data indicates that phytosiderophores are capable of increasing the amount of complexed cations in solution. Despite their lack of specificity, phytosiderophores were just as effective as Desferal increasing the availability of Fe. Thus, phytosiderophores, as plant-borne chelators, are certainly of significance for the Fe nutrition of cereals grown in calcareous soils.

Root‐induced changes of nutrient availability in the rhizosphere

Abstract Nutrient availability in the rhizosphere differs in many respects from that in the bulk soil. Factors of major importance for the mineral nutrition of plants are root‐induced changes in rhizosphere pH, in reducing capacity of the roots, and in amount and composition of root exudates. These changes are in many instances root‐responses to the nutritional status of plants. Examples are given for zinc, iron, and phosphorus nutritional status in plants. Changes in nutrient availability in the rhizosphere may also be a consequence of alterations in the rhizosphere microflora. Root‐induced changes are important components for the adaptation of plants to extreme chemical soil conditions and for efficient use of soil and fertilizer nutrients.

Roots of Iron-Efficient Maize also Absorb Phytosiderophore-Chelated Zinc.

To investigate the recognition of Zn-phytosiderophores by the putative Fe-phytosiderophore transporter in maize (Zea mays L.) roots, short-term uptake of 65Zn-labeled phytosiderophores was compared in the Fe-efficient maize cultivar Alice and the maize mutant ys1 carrying a defect in Fe-phytosiderophore uptake. In ys1, uptake and translocation rates of Zn from Zn-phytosiderophores were one-half of those in Alice, but no genotypical difference was found in Zn uptake and translocation from other Zn-binding forms. In ys1 and in tendency also in Alice, Zn uptake decreased with increasing stability constant of the chelate in the order: ZnSO4 [greater than or equal to] Zn-desferrioxamine > Zn-phytosiderophores > Zn-EDTA. Adding a 500-fold excess of free phytosiderophores over Zn to the uptake solution depressed Zn uptake in ys1 almost completely. In uptake studies with double-labeled 65Zn-14C-phytosiderophores, ys1 absorbed the phytosiderophore at similar rates when supplied as a Zn-chelate or the free ligand. By contrast, in Alice 14C-phytosiderophore uptake from the Zn-chelate was 2.8-fold higher than from the free ligand, suggesting that Alice absorbed the complete Zn-phytosiderophore complex via the putative plasma membrane transporter for Fe-phytosiderophores. We propose two pathways for the uptake of Zn from Zn-phytosiderophores in grasses, one via the transport of the free Zn cation and the other via the uptake of nondissociated Zn-phytosiderophores.

Iron Inefficiency in Maize Mutant ys1 (Zea mays L. cv Yellow-Stripe) Is Caused by a Defect in Uptake of Iron Phytosiderophores

To determine the Fe inefficiency factors in the maize mutant ys1 (Zea mays L. cv Yellow Stripe), root exudates of Fe-inefficient ys1 and of two Fe-efficient maize cultivars (Alice, WF9) were collected in axenic nutrient solution cultures. Analysis by thin-layer chromatography and high-performance liquid chromatography revealed that under Fe deficiency ys1 released the phytosiderophore 2[prime]-deoxymugineic acid (DMA) in quantities similar to those of Alice and WF9. Under nonaxenic conditions, DMA released by plants of all three cultivars was rapidly decomposed by microorganisms in the nutrient solution. Uptake experiments with 59Fe-labeled DMA, purified from root exudates of either Fe-deficient Alice or ys1 plants, showed up to 20 times lower uptake and translocation of 59Fe in ys1 than in Alice or WF9 plants. The presence of microorganisms during preculture and short-term uptake experiments had no significant effect on uptake and translocation rates of 59Fe in Alice and ys1 plants. We conclude that Fe inefficiency in the maize mutant ys1 is the result of a defect in the uptake system for Fe-phytosiderophores.

The chlorosis paradox: Fe inactivation as a secondary event in chlorotic leaves of grapevine

Abstract Iron chlorosis is a wide‐spread disorder of plants, in particular of those on calcareous soils. Elevated bicarbonate concentrations in the soil solution are considered as a main cause of this chlorosis. In nutrient solution culture experiments a supply of bicarbonate results in inhibited Fe acquisition and a subsequent decreased concentration of Fe in the leaf dry matter. This is indicated by a close positive relationship between chlorophyll and total Fe concentration in the upper leaves. In contrast to nutrient solution experiments, in some pot experiments particularly with calcareous soil, and in field experiments under certain conditions, no such close correlation can be observed and a higher Fe concentration can even be found in young chlorotic leaves than in green leaves. This phenomenon is called “the chlorosis paradox” and it has thus been concluded that, Fe chlorosis might be caused by an Fe inactivation in the plant, in particular in the leaf apoplast, e.g. by an alkalinization process. Reconsideration of published data on this phenomenon of enhanced Fe concentrations in chlorotic leaves, coupled with investigations of grapevine grown on calcareous soil reveal that “the chlorosis paradox” can only be observed in soil culture where severe shoot growth inhibition is already present at an early stage; presumably as a consequence of changes in phytohormone metabolism in connection with inhibited root growth. Obviously, the higher Fe concentration in chlorotic leaves with inhibited expansion growth is a consequence of the diminished dilution of normal high Fe concentrations in young leaves. This high concentration in chlorotic leaves can be observed in spite of a distinct lower Fe content in the individual leaves. The declined concentration of HCl‐extractable Fe in chlorotic leaves is presumably not the cause, but rather the consequence, of Fe chlorosis.