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.Keywords Potassium availability . Potassium micronutrient interaction . Spatial availability of potassium . K/Mg ratio . Abiotic stress . Biotic stress . Frost resistance . Food quality . K/Cd relations Plant Soil (2010) 335:155-180

Mineral nutrients taken up by the roots are, as a rule, transported in the xylem to the shoot, and photoassimilates transported in the phloem to the roots. According to the Thornley model of photosynthate partitioning, nutrient deficiencies should favour photosynthate partitioning to the roots. Examples are cited to show that this preferential partitioning is dependent on phloem mobility and hence on nutrient cycling from shoot to roots. Thus, root growth is enhanced under nitrogen and phosphorus deficiencies, but not under deficiencies of nutrients of low mobility in the phloem, such as calcium and boron. Enhanced root growth under nutrient deficiency relies on the import of both photosynthates and mineral nutrients. Cycling of mineral nutrients serves a number of other functions. These include the root supply of nutrients assimilated in the shoot (nitrate and sulphate reduction), maintenance of cation-anion balance in the shoot, providing an additional driving force for solute volume flow in the phloem and xylem, and acting as a shoot signal to convey nutrient demand to the root. Cycling of certain mineral nutrients through source leaves has a considerable impact on photosynthate export as demonstrated in impaired export under magnesium, potassium, or zinc deficiencies. Mineral nutrient deficiency can, therefore, affect photosynthate partitioning either directly via phloem loading and transport or indirectly by depressing sink demand.

Magnesium (Mg) deficiency exerts a major influence on the partitioning of dry matter and carbohydrates between shoots and roots. One of the very early reactions of plants to Mg deficiency stress is the marked increase in the shoot-to-root dry weight ratio, which is associated with a massive accumulation of carbohydrates in source leaves, especially of sucrose and starch. These higher concentrations of carbohydrates in Mg-deficient leaves together with the accompanying increase in shoot-to-root dry weight ratio are indicative of a severe impairment in phloem export of photoassimilates from source leaves. Studies with common bean and sugar beet plants have shown that Mg plays a fundamental role in phloem loading of sucrose. At a very early stage of Mg deficiency, phloem export of sucrose is severely impaired, an effect that occurs before any noticeable changes in shoot growth, Chl concentration or photosynthetic activity. These findings suggest that accumulation of carbohydrates in Mg-deficient leaves is caused directly by Mg deficiency stress and not as a consequence of reduced sink activity. The role of Mg in the phloem-loading process seems to be specific; resupplying Mg for 12 or 24 h to Mg-deficient plants resulted in a very rapid recovery of sucrose export. It appears that the massive accumulation of carbohydrates and related impairment in photosynthetic CO2 fixation in Mg-deficient leaves cause an over-reduction in the photosynthetic electron transport chain that potentiates the generation of highly reactive O2 species (ROS). Plants respond to Mg deficiency stress by marked increases in antioxidative capacity of leaves, especially under high light intensity, suggesting that ROS generation is stimulated by Mg deficiency in chloroplasts. Accordingly, it has been found that Mg-deficient plants are very susceptible to high light intensity. Exposure of Mg-deficient plants to high light intensity rapidly induced leaf chlorosis and necrosis, an outcome that was effectively delayed by partial shading of the leaf blade, although the Mg concentrations in different parts of the leaf blade were unaffected by shading. The results indicate that photooxidative damage contributes to development of leaf chlorosis under Mg deficiency, suggesting that plants under high-light conditions have a higher physiological requirement for Mg. Maintenance of a high Mg nutritional status of plants is, thus, essential in the avoidance of ROS generation, which occurs at the expense of inhibited phloem export of sugars and impairment of CO2 fixation, particularly under high-light conditions.

Sumnwarv. An investigation was carried ouit to study the cation-anion balance in different tissues of tomato plants suipplied with nitrate, urea, or ammonium niitrogen in water culture.Irrespective of the form of nutrition, a very close balance was found in the tissues investigated (leaves, petioles, stems, and roots) between total cations (Ca, Mg, K and Na), and total anions (N03-, H2P04-, SO4--, Cl-) total non-volatile organic acids, oxalate, and uronic acids. In comparison with the tissues of the nitrate fed plants, the corresponding ammonium tissues contained lower concentrations of inorganic cations, and organic acids and a correspondingly higher proportion of inorganic anions. Tissues from the urea plants were intermediate between the other 2 treatments. These results were independent of concentration or dilution effects, caused by growth. In all tissuies approximately equivalent amounts of diffusible cations (Ca", \Ig+, K' and Nat), and diffusible anions (No3-, SO4--, H,PO4-, Cl-) and non-volatile organic acids were found. An almost 1:1 ratio occurred between the levels of bound calcium and magnesium, and oxalate and uronic acids. This points to the fact that in the tomato plant the indiffusible anions are mainly oxalate and pectate. Approximately equivalent values were found for the alkalinity of the ash, and organic anions (total organic acids including oxalate, and uronic acids).The influence of nitrate, urea, and ammonitum nitrogen nutrition on the cation-anion balance and the organic acid content of the plant has been considered and the effects of these different nitrogen forms on both the pH of the plant and the nutrient medium and its conseqtuences discussed.Ionic equilibrium in plant tissues is maintained by diffusible and indiffusible ions, in which both organic and inorganic cations and( anions play a part. It is known that this equilibrium is very much dependent on the form of nitrogen nutrition to the plant (6, 9, 14). Previous studies have centered mainly on the comparison of nitrate and ammonium sources on the diffusible ionic balance in leaves, (6,9,14)

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