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

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

Zinc deficiency is a well-documented problem in food crops, causing decreased crop yields and nutritional quality. Generally, the regions in the world with Zn-deficient soils are also characterized by widespread Zn deficiency in humans. Recent estimates indicate that nearly half of world population suffers from Zn deficiency. Cereal crops play an important role in satisfying daily calorie intake in developing world, but they are inherently very low in Zn concentrations in grain, particularly when grown on Zn-deficient soils. The reliance on cereal-based diets may induce Zn deficiency-related health problems in humans, such as impairments in physical development, immune system and brain function. Among the strategies being discussed as major solution to Zn deficiency, plant breeding strategy (e.g., genetic biofortification) appears to be a most sustainable and cost-effective approach useful in improving Zn concentrations in grain. The breeding approach is, however, a long-term process requiring a substantial effort and resources. A successful breeding program for biofortifying food crops with Zn is very much dependent on the size of plant-available Zn pools in soil. In most parts of the cereal-growing areas, soils have, however, a variety of chemical and physical problems that significantly reduce availability of Zn to plant roots. Hence, the genetic capacity of the newly developed (biofortified) cultivars to absorb sufficient amount of Zn from soil and accumulate it in the grain may not be expressed to the full extent. It is, therefore, essential to have a short-term approach to improve Zn concentration in cereal grains. Application of Zn fertilizers or Zn-enriched NPK fertilizers (e.g., agronomic biofortification) offers a rapid solution to the problem, and represents useful complementary approach to on-going breeding programs. There is increasing evidence showing that foliar or combined soil+foliar application of Zn fertilizers under field conditions are highly effective and very practical way to maximize uptake and accumulation of Zn in whole wheat grain, raising concentration up to 60 mg Zn kg−1. Zinc-enriched grains are also of great importance for crop productivity resulting in better seedling vigor, denser stands and higher stress tolerance on potentially Zn-deficient soils. Agronomic biofortification strategy appears to be essential in keeping sufficient amount of available Zn in soil solution and maintaining adequate Zn transport to the seeds during reproductive growth stage. Finally, agronomic biofortification is required for optimizing and ensuring the success of genetic biofortification of cereal grains with Zn. In case of greater bioavailability of the grain Zn derived from foliar applications than from soil, agronomic biofortification would be a very attractive and useful strategy in solving Zn deficiency-related health problems globally and effectively.

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Enrichment of cereal grains with zinc: Agronomic or genetic biofortification?

Zinc (Zn) is an essential component of thousands of proteins in plants, although it is toxic in excess. In this review, the dominant fluxes of Zn in the soil-root-shoot continuum are described, including Zn inputs to soils, the plant availability of soluble Zn(2+) at the root surface, and plant uptake and accumulation of Zn. Knowledge of these fluxes can inform agronomic and genetic strategies to address the widespread problem of Zn-limited crop growth. Substantial within-species genetic variation in Zn composition is being used to alleviate human dietary Zn deficiencies through biofortification. Intriguingly, a meta-analysis of data from an extensive literature survey indicates that a small proportion of the genetic variation in shoot Zn concentration can be attributed to evolutionary processes whose effects manifest above the family level. Remarkable insights into the evolutionary potential of plants to respond to elevated soil Zn have recently been made through detailed anatomical, physiological, chemical, genetic and molecular characterizations of the brassicaceous Zn hyperaccumulators Thlaspi caerulescens and Arabidopsis halleri.

Zinc in plants

Cellular Mechanisms of Aluminum Toxicity and Resistance in Plants

summary
Aluminium is the most important growth-limiting factor in many acid soils throughout the world. Physiological effects of Al toxicity and mechanisms of tolerance are not well understood. An initial uptake of Al is confined to the apoplasm. Aluminium complexes (whose exact identification is beyond current experimental techniques) enter the cytosol slowly and only after prolonged exposure. Electrochemical properties of the cell wall Donnan free space as well as the plasma membrane of root cells are altered by the presence of Al ions that are polyvalent cations in acidic environments. The primary Al effects are very fast (taking only seconds to several minutes to develop) and may therefore occur while Al is still in the Donnan free space and on the apoplasmic side of the plasma membrane. Resumption of root growth upon removal of Al ions supports such a claim. Aluminium affects membrane permeability for both electrolytes and non-electrolytes; it reduces accumulation of divalent cations (especially Ca and Mg) by interfering with the membrane transport. Aluminium alters the pattern of Ca2+ fluxes across the plasma membrane, thus supposedly disturbing symplasmic Ca2+ homeostasis. Supplemental Ca2+ can greatly alleviate deleterious Al effects. Frequently observed changes in the secretory activity of root cells exposed to Al are mediated through altered cell Ca2+ homeostasis; they result in cessation of cell wall growth and stoppage of root elongation. Involvement of calmodulin in the Al-related phenomena is suggested to be indirect, at least in the initial stages of the Al treatment when Al is likely to be confined to the apoplasm. The role of growth substances, at least auxins and cytokinins, in the Al toxicity syndrome appears to be related to the Ca-Al interactions that may alter the pattern of auxin transport as well as cytokinin biosynthesis and transport. Disturbance of the cell Ca2+ homeostasis appears to be an important feature of ion-related environmental stresses in general (salt, heavy metals, aluminium).

Role of calcium in aluminium toxicity

Seed nutrient reserves may be important for an early establishment of crops on low-fertility soils. This glasshouse pot study evaluated effects of seed Zn content on vegetative growth of two wheat (Triticum aestivum L.) genotypes differing in Zn efficiency. Low-Zn (around 250 ng Zn per seed) and high-Zn seed (around 700 ng Zn per seed on average) of Excalibur (Zn efficient) and Gatcher (Zn inefficient) wheats were sown in a Zn-deficient siliceous sand fertilised with 0, 0.05, 0.2, 0.8 or 3.2 mg Zn kg -1 soil. After 3 weeks, plants derived from the high-Zn seed had better root and shoot growth; the cv. Excalibur accumulated more shoot dry matter than the cv. Gatcher. After 6 weeks, greater root and shoot growth of plants grown from the high-Zn seed compared to those from the low-Zn seed was obvious only at nil Zn fertilisation. A fertilisation rate of 0.2 mg Zn kg -1 soil was required for achieving 90% of the maximum yield for plants grown from the high-Zn seed compared to 0.8 mg Zn kg -1 soil for plants derived from the low-Zn seed. The critical Zn level in youngest expanded leaves for 90% maximum yield was 16 mg Zn kg -1 dry matter for both genotypes. Zn-efficient Excalibur had greater net Zn uptake rates compared to Zn-inefficient Gatcher after 3 weeks but they were not different at the 6-week harvest. Zinc-deficient plants had greater net uptake rates of Cu, Mn, B, P, and K but a reduced uptake rate of Fe. It is concluded that higher seed Zn content acted similar to a starter-fertiliser effect by improving vegetative growth and dissipating differences in Zn efficiency of wheat genotypes.

Importance of seed Zn content for wheat growth on Zn-deficient soil. I: Vegetative growth

Genotypes of plants vary widely in their tolerance of Zn-deficient soils, both in Zn uptake and utilization. Tolerance, here termed Zn efficiency because it appears to involve in most cases more efficient extraction of soil Zn, is heritable and may therefore be exploited in the breeding of crop plants. This review considers the extent of variability in Zn eficiency in species of crop and natural plants, the physiological and biochemical nature of the mechanisms of efficiency, what is known of the genetics of inheritance, screening techniques for identifying Zn-efficient types in breeding programs, and the agronomic arguments for a breeding solution to the problem of Zn-deficient soils.

Genotypic Variation in Zinc Uptake and Utilization by Plants

Ten Triticum aestivum and two Triticum turgidum conv. durum genotypes were grown in chelate-buffered nutrient solution at Zn supplies ranging from deficient to sufficient (free Zn activities from 2 to 200 pM, pZn from 11.7 to 9.7). The critical level of Zn ion activity in solution for healthy growth of wheat plants was around 40 pM. Genotypes differed in the growth response: those classified as Zn-efficient suffered less reduction of shoot growth and did not change the rate of root growth at a Zn supply quite deficient for Zn-inefficient genotypes. Root growth of Zn-inefficient genotypes increased at deficient Zn supply. The shoot/root ratio was the most sensitive parameter of Zn efficiency; Zn-efficient genotypes showed less reduction in the ratio when grown at deficient compared to sufficient Zn supply. Classification of wheat genotypes into Zn-efficient and Zn-inefficient groups after screening in chelate-buffered nutrient solution corresponded well with classification obtained in field experiments on Zn-deficient soil.

Wheat genotypes differ in Zn efficiency when grown in chelate-buffered nutrient solution

Two wheat (Triticum aestivum L.) genotypes (Zn-efficient Excalibur and Zn-inefficient Gatcher) were grown from low-Zn (around 250 ng Zn/seed) and high-Zn seed (around 700 ng Zn/seed) in a Zn-deficient siliceous sand fertilised with 0, 0.05, 0.2, 0.8 or 3.2 mg Zn kg-1 soil. At maturity, plants derived from the high-Zn seed had bigger grains and produced more grains than plants grown from the low-Zn seed when fertilised with nil Zn. Plants grown from high-Zn seed produced more grain dry matter per unit of Zn absorbed by the above-ground parts, transported a larger proportion of absorbed Zn to the grain, and approached the maximum harvest index with the fertilisation rate of 0.05 mg Zn kg-1 compared to 0.2 mg Zn kg-1 soil required for plants derived from the low-Zn seed. The Zn-efficient cv. Excalibuir had greater fertiliser efficiency when fertilised with 0.05 mg Zn kg-1 soil and a greater harvest index at nil Zn treatment compared to the Zn-inefficient cv. Gatcher. Zn-deficient plants produced seed with higher concentrations of all inorganic nutrients determined except Zn. There was a positive relationship between grain dry weight and Zn concentrations in youngest expanded blades sampled at 6 weeks of growth. It is concluded that crops grown from seed containing higher Zn content have a distinct advantage which culminates in greater yield when grown in soils of low Zn status.

Zdenko Rengel

Papers

Role of calcium in aluminium toxicity

Importance of seed Zn content for wheat growth on Zn-deficient soil. I: Vegetative growth

Genotypic Variation in Zinc Uptake and Utilization by Plants

Wheat genotypes differ in Zn efficiency when grown in chelate-buffered nutrient solution

Importance of seed Zn content for wheat growth on Zn-deficient soil