Chapter 1 The Biosphere Chapter 2 The Anthroposphere Introduction Air Pollution Water Pollution Soil Plants Chapter 3 Soils and Soil Processes Introduction Weathering Processes Pedogenic Processes Chapter 4 Soil Constituents Introduction Trace Elements Minerals Organic Matter Organisms in Soils Chapter 5 Trace Elements in Plants Introduction Absorption Translocation Availability Essentiality and Deficiency Toxicity and Tolerance Speciation Interaction Chapter 6 Elements of Group 1 (Previously Group Ia) Introduction Lithium Rubidium Cesium Chapter 7 Elements of Group 2 (Previously Group IIa) Beryllium Strontium Barium Radium Chapter 8 Elements of Group 3 (Previously Group IIIb) Scandium Yttrium Lanthanides Actinides Chapter 9 Elements of Group 4 (Previously Group IVb) Titanium Zirconium Hafnium Chapter 10 Elements of Group 5 (Previously Group Vb) Vanadium Niobium Tantalum Chapter 11 Elements of Group 6 (Previously Group VIb) Chromium Molybdenum Tungsten Chapter 12 Elements of Group 7 (Previously Group VIIb) Manganese Technetium Rhenium Chapter 13 Elements of Group 8 (Previously Part of Group VIII) Iron Ruthenium Osmium Chapter 14 Elements of Group 9 (Previously Part of Group VIII) Cobalt Rhodium Iridium Chapter 15 Elements of Group 10 (Previously Part of Group VIII) Nickel Palladium Platinum Chapter 16 Elements of Group 11 (Previously Group Ib) Copper Silver Gold Chapter 17 Trace Elements of Group 12 (Previously of Group IIb) Zinc Cadmium Mercury Chapter 18 Elements of Group 13 (Previously Group IIIa) Boron Aluminum Gallium Indium Thallium Chapter 19 Elements of Group I4 (Previously Group IVa) Silicon Germanium Tin Lead Chapter 20 Elements of Group 15 (Previously Group Va) Arsenic Antimony Bismuth Chapter 21 Elements of Group 16 (Previously Group VIa) Selenium Tellurium Polonium Chapter 22 Elements of Group 17 (Previously Group VIIa) Fluorine Chlorine Bromine Iodine

Trace elements in soils and plants

1. Plant Nutrients. 2. The Soil as a Plant Nutrient Medium. 3. Nutrient Uptake and Assimilation. 4. Plant Water Relationships. 5. Plant Growth and Crop Production. 6. Fertilizer Application. 7. Nitrogen. 8. Sulphur. 9. Phosphorus. 10. Potassium. 11. Calcium. 12. Magnesium. 13. Iron. 14. Manganese. 15. Zinc. 16. Copper. 17. Molybdenum. 18. Boron. 19. Further Elements of Importance. 20. Elements with More Toxic Effects. General Readings. References. Index.

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Principles of plant nutrition

Fifteen or more elements present in rocks and soils normally in very small amounts are essential for plant and/or animal nutrition. By the nature of their low abundance in natural uncontaminated earth materials or plants, they are known as trace elements, minor elements or micro-nutrients. Boron, copper, iron, manganese, molybdenum, silicon, vanadium and zinc are required by plants; copper, cobalt, iodine, iron, manganese, molybdenum, selenium and zinc by animals. In addition essential roles of arsenic, fluorine, nickel, silicon, tin and vanadium have in recent years been established in animal nutrition.

Trace Elements in Soils and Plants

Numerous species of soil bacteria which flourish in the rhizosphere of plants, but which may grow in, on, or around plant tissues, stimulate plant growth by a plethora of mechanisms. These bacteria are collectively known as PGPR (plant growth promoting rhizobacteria). The search for PGPR and investigation of their modes of action are increasing at a rapid pace as efforts are made to exploit them commercially as biofertilizers. After an initial clarification of the term biofertilizers and the nature of associations between PGPR and plants (i.e., endophytic versus rhizospheric), this review focuses on the known, the putative, and the speculative modes-of-action of PGPR. These modes of action include fixing N2, increasing the availability of nutrients in the rhizosphere, positively influencing root growth and morphology, and promoting other beneficial plant–microbe symbioses. The combination of these modes of actions in PGPR is also addressed, as well as the challenges facing the more widespread utilization of PGPR as biofertilizers.

Plant growth promoting rhizobacteria as biofertilizers

In most soils, inorganic phosphorus occurs at fairly low concentrations in the soil solution whilst a large proportion of it is more or less strongly held by diverse soil minerals. Phosphate ions can indeed be adsorbed onto positively charged minerals such as Fe and Al oxides. Phosphate (P) ions can also form a range of minerals in combination with metals such as Ca, Fe and Al. These adsorption/desorption and precipitation/dissolution equilibria control the concentration of P in the soil solution and, thereby, both its chemical mobility and bioavailability. Apart from the concentration of P ions, the major factors that determine those equilibria as well as the speciation of soil P are (i) the pH, (ii) the concentrations of anions that compete with P ions for ligand exchange reactions and (iii) the concentrations of metals (Ca, Fe and Al) that can coprecipitate with P ions. The chemical conditions of the rhizosphere are known to considerably differ from those of the bulk soil, as a consequence of a range of processes that are induced either directly by the activity of plant roots or by the activity of rhizosphere microflora. The aim of this paper is to give an overview of those chemical processes that are directly induced by plant roots and which can affect the concentration of P in the soil solution and, ultimately, the bioavailability of soil inorganic P to plants. Amongst these, the uptake activity of plant roots should be taken into account in the first place. A second group of activities which is of major concern with respect to P bioavailability are those processes that can affect soil pH, such as proton/bicarbonate release (anion/cation balance) and gaseous (O2/CO2) exchanges. Thirdly, the release of root exudates such as organic ligands is another activity of the root that can alter the concentration of P in the soil solution. These various processes and their relative contributions to the changes in the bioavailability of soil inorganic P that can occur in the rhizosphere can considerably vary with (i) plant species, (ii) plant nutritional status and (iii) ambient soil conditions, as will be stressed in this paper. Their possible implications for the understanding and management of P nutrition of plants will be briefly addressed and discussed.

Bioavailability of soil inorganic P in the rhizosphere as affected by root-induced chemical changes: A review

Principles of Plant Nutrition.

The article describes factors and processes which lead to Fe chlorosis (lime chlorosis) in plants grown on calcareous soils. Such soils may contain high HCO3 - concentrations in their soil solution, they are characterized by a high pH, and they rather tend to accumulate nitrate than ammonium because due to the high pH level ammonium nitrogen is rapidly nitrified and/or even may escape in form of volatile NH3. Hence in these soils plant roots may be exposed to high nitrate and high bicarbonate concentrations. Both anion species are involved in the induction of Fe chlorosis. Physiological processes involved in Fe chlorosis occur in the roots and in the leaves. Even on calcareous soils and even in plants with chlorosis the Fe concentration in the roots is several times higher than the Fe concentration in the leaves. This shows that the Fe availability in the soil is not the critical process leading to chlorosis but rather the Fe uptake from the root apoplast into the cytosol of root cells. This situation applies to dicots as well as to monocots. Iron transport across the plasmamembrane is initiated by FeIII reduction brought about by a plasmalemma located FeIII reductase. Its activity is pH dependent and at alkaline pH supposed to be much depressed. Bicarbonate present in the root apoplast will neutralize the protons pumped out of the cytosol and together with nitrate which is taken up by a H+/nitrate cotransport high pH levels are provided which hamper or even block the FeIII reduction. Frequently chlorotic leaves have higher Fe concentrations than green ones which phenomenon shows that chlorosis on calcareous soils is not only related to Fe uptake by roots and Fe translocation from the roots to the upper plant parts but also dependent on the efficiency of Fe in the leaves. It is hypothesized that also in the leaves FeIII reduction and Fe uptake from the apoplast into the cytosol is affected by nitrate and bicarbonate in an analogous way as this is the case in the roots. This assumption was confirmed by the highly significant negative correlation between the leaf apoplast pH and the degree of iron chlorosis measured as leaf chlorophyll concentration. Depressing leaf apoplast pH by simply spraying chlorotic leaves with an acid led to a regreening of the leaves.

Iron availability in plant tissues — iron chlorosis on calcareous soils

Organic matter incorporated into soils may influence soil pH. Until now the detailed processes responsible for a change in soil pH after the incorporation of organic matter were not completely understood. We studied the effects of organic anions, glycine as a representative of amino acids, and glucose as a representative of carbohydrates on soil pH in aerobic incubation experiments. Addition of malate and citrate resulted in an immediate soil pH increase associated with additional CO 2 release. Addition of acetate had no major effect on soil pH but if applied at higher rates decreased soil CO 2 release. Malate added together with ammonium resulted in an immediate soil pH increase, which was followed by a pH decrease associated by an increase in nitrate after 10 days. Also glycine gave an immediate pH increase which was associated with an equivalent decrease of soil organic N extractable with CaCl 2 solution. Addition of glucose gave a slight pH decrease in the first days of incubation followed by an increase to the initial pH value. The results are interpreted in terms of decarboxylation of organic anions, a process which requires one proton per carboxylic group decarboxylated. Under aerobic soil conditions decarboxylation is a major process in organic matter decomposition. The decomposition of carbohydrates in the glycolytic pathway produces carboxylic groups which, after dissociation, may decrease soil pH. As soon as these groups are decarboxylated in the citrate cycle an equivalent amount of protons is required inducing a rise in soil pH. For this reason addition of glucose brought about only a transient soil pH decrease.

Soil pH increase due to biological decarboxylation of organic anions

Maize (Zea mays L.) and sunflower (Helianthus annuus L.) grown on a calcareous soil showed poor growth and/or were chlorotic in spite of abundant Fe in the roots. It has been hypothesized that microbial siderophores chelate Fe (III) in the soil, and that in this form Fe is transported towards the root apoplast. On the calcareous soil, total and apoplastic root Fe concentrations were high, probably because of a high apoplastic pH depressing Fe (III)-reductase activity and thus the Fe2+ supply to the cytoplasm. On the acidic soil, total and apoplastic root Fe concentrations were low, probably because of a low apoplastic pH favouring Fe (III) reduction, hence plants showed no Fe-deficiency symptoms. The main objective of the present work was to investigate the role of microbial soil activity in plant Fe acquisition. For this purpose, plants were grown under sterile and non-sterile conditions on a loess loam soil. Plants cultivated under non-sterile conditions grew well, showed no Fe-deficiency symptoms and had fairly high Fe concentrations in the roots in contrast to plants grown in the sterile medium. Low root and leaf Fe concentrations in the axenic treatments indicated that the production of microbial siderophores was totally suppressed. Accordingly, sunflowers were severely chlorotic and this was associated with very poor growth, whereas in maize only growth was drastically reduced. In maize under sterile conditions, root apoplastic and total Fe concentrations were not as low as in sunflowers, which may have indicated that phytosiderophores produced in maize partly sustained Fe acquisition, but due to poor growth were not as efficient in supplying Fe as microbial activity under natural conditions. It may be therefore assumed that in natural habitats soil microbial activity is of pivotal importance for plant Fe acquisition.

Konrad Mengel

Papers

Principles of plant nutrition

Principles of Plant Nutrition.

Iron availability in plant tissues — iron chlorosis on calcareous soils

Soil pH increase due to biological decarboxylation of organic anions

The central role of microbial activity for iron acquisition in maize and sunflower.