Nutrient sensing and signalling in plants: Potassium and phosphorus

Potassium and phosphorus are important macronutrients for crops but are often deficient in the field. Very little is known about how plants sense fluctuations in K and P and how information about K and P availability is integrated at the whole plant level into physiological and metabolic adaptations. This chapter reviews recent advances in discovering molecular responses of plants to K and P deficiency by microarray experiments. These studies provide us not only with a comprehensive picture of adaptive mechanisms, but also with a large number of transcriptional markers that can be used to identify upstream components of K and P signalling pathways. On the basis of the available information we discuss putative receptors and signals involved in the sensing and integration of K and P status both at the cellular and at the whole plant level. These involve membrane potential, voltage-dependent ion channels, intracellular Ca and pH, and transcription factors, as well as hormones and metabolites for systemic signalling. Genetic screens of reporter lines for transcriptional markers and metabolome analysis of K- and P-deficient plants are likely to further advance our knowledge in this area in the near future.

Diversity in expression of phosphorus (P) responsive genes in Cucumis melo L.

Background: Phosphorus (P) is a major limiting nutrient for plant growth in many soils. Studies in model species have identified genes involved in plant adaptations to low soil P availability. However, little information is available on the genetic bases of these adaptations in vegetable crops. In this respect, sequence data for melon now makes it possible to identify melon orthologues of candidate P responsive genes, and the expression of these genes can be used to explain the diversity in the root system adaptation to low P availability, recently observed in this species. Methodology and Findings: Transcriptional responses to P starvation were studied in nine diverse melon accessions by comparing the expression of eight candidate genes (Cm-PAP10.1, Cm-PAP10.2, Cm-RNS1, Cm-PPCK1, Cm-transferase, Cm-SQD1, Cm-DGD1 and Cm-SPX2) under P replete and P starved conditions. Differences among melon accessions were observed in response to P starvation, including differences in plant morphology, P uptake, P use efficiency (PUE) and gene expression. All studied genes were up regulated under P starvation conditions. Differences in the expression of genes involved in P mobilization and remobilization (Cm-PAP10.1, Cm-PAP10.2 and Cm-RNS1) under P starvation conditions explained part of the differences in P uptake and PUE among melon accessions. The levels of expression of the other studied genes were diverse among melon accessions, but contributed less to the phenotypical response of the accessions. Conclusions: This is the first time that these genes have been described in the context of P starvation responses in melon. There exists significant diversity in gene expression levels and P use efficiency among melon accessions as well as significant correlations between gene expression levels and phenotypical measurements.

Genetic responses to phosphorus deficiency

Background: Phosphorus (P) is an essential macronutrient for plants. Plants take up P as phosphate (Pi) from the soil solution. Since little Pi is available in most soils, P fertilizers are applied to crops. However, the use of P fertilizers is unsustainable and may cause pollution. Consequently, there is a need to develop more P-use-efficient (PUE) crops and precise methods to monitor crop P-status. Scope: Manipulating the expression of genes to improve the PUE of crops could reduce their P fertilizer requirement. This has stimulated research towards the identification of genes and signalling cascades involved in plant responses to P deficiency. Genes that respond to P deficiency can be grouped into 'early' genes that respond rapidly and often non-specifically to P deficiency, or 'late' genes that impact on the morphology, physiology or metabolism of plants upon Prolonged P deficiency. Summary: The use of micro-array technology has allowed researchers to catalogue the genetic responses of plants to P deficiency. Genes whose expression is altered by P deficiency include various transcription factors, which are thought to coordinate plant responses to P deficiency, and other genes involved in P acquisition and tissue P economy. Several common cis-regulatory elements have been identified in the promoters of these genes, suggesting that their expression might be coordinated. It is suggested that knowledge of the genes whose expression changes in response to P deficiency might allow the development of crops with improved PUE, and could be used in diagnostic techniques to monitor P deficiency in crops either directly using 'smart' indicator plants or indirectly through transcript profiling. The development of crops with improved PUE and the adoption of diagnostic technology could reduce production costs, minimize the use of a non-renewable resource, reduce pollution and enhance biodiversity.

The forms of bioavailability of phosphorus in integrated vertical flow constructed wetland with earthworms and different substrates

A sequential extraction method was utilized to analyze seven forms of P in an integrated vertical-flow constructed wetland (IVFCW) containing earthworms and different substrates. The aluminum-bound P (Al-P) content was found to be lower, and the occluded P (Oc-P) content was higher in the IVFCW. The addition of earthworms into the influent chamber of IVFCW increased the exchange P (Ex-P), iron-bound P (Fe-P), calcium bound P (Ca-P), Oc-P, detritus-bound (De-P) and organic P (Org-P) content in the influent chamber, and also enhanced P content uptake by wetland plants. A significantly positive correlation between P content of above-ground wetland plants and the Ex-P, Fe-P, Oc-P and Org-P content in the rhizosphere was found (P < 0.05), which indicated that the Ex-P, Fe-P, Oc-P and Org-P could be bio-available P. The Ex-P, Fe-P, De-P, Oc-P and Ca-P content of the influent chamber was higher where the substrate contained a mixture of Qing sand and river sand rather than only river sand. Also the IVFCW with earthworms and both Qing sand and river sand had a higher removal efficiency of P, which was related to higher P content uptake by wetland plants and P retained in IVFCW. These findings suggest that addition of earthworms in IVFCW increases the bioavailable P content, resulting in enhanced P content uptake by wetland plants.

Moderating mycorrhizas: arbuscular mycorrhizas modify rhizosphere chemistry and maintain plant phosphorus status within narrow boundaries

Pastures often experience a pulse of phosphorus (P) when fertilized. We examined the role of arbuscular mycorrhizal fungi (AMF) in the uptake of P from a pulse. Five legumes (Kennedia prostrata, Cullen australasicum, Bituminaria bituminosa, Medicago sativa and Trifolium subterraneum) were grown in a moderate P, sterilized field soil, either with (+AMF) or without (−AMF) addition of unsterilized field soil. After 9–10 weeks, half the pots received 15 mg P kg−1 of soil. One week later, we measured: shoot and root dry weights; percentage of root length colonized by AMF; plant P, nitrogen and manganese (Mn) concentrations; and rhizosphere carboxylates, pH and plant-available P. The P pulse raised root P concentration by a similar amount in uncolonized and colonized plants, but shoot P concentration increased by 143% in uncolonized plants and 53% in colonized plants. Inoculation with AMF decreased the amount of rhizosphere carboxylates by 52%, raised rhizosphere pH by ∼0.2–0.7 pH units and lowered shoot Mn concentration by 38%. We conclude that AMF are not simply a means for plants to enhance P uptake when P is limiting, but also act to maintain shoot P within narrow boundaries and can affect nutrient uptake through their influence on rhizosphere chemistry.

Soil phosphorus dynamics and phytoavailability from sewage sludge at different stages in a treatment stream

The effect of different stages of sewage sludge treatment on phosphorus (P) dynamics in amended soils was determined using samples of undigested liquid (UL), anaerobically digested liquid (AD) and dewatered anaerobically digested (DC) sludge. Sludges were taken from three points in the same treatment stream and applied to a sandy loam soil in field-based mesocosms at 4, 8 and 16t ha−1 dry solids. Mesocosms were sown with perennial ryegrass (Lolium perenne cv. Melle), and the sward was harvested after 35 and 70 days to determine yield and foliar P concentration. Soils were also sampled during this period to measure P transformations and the activities of acid phosphomonoesterase and phosphodiesterase. Data show that the AD amended soils had the greatest plant-available and foliar P content up to the second harvest, but the UL amended soils had the greatest enzyme activity. Characterisation of control and 16t ha−1 soils and sludge using solution 31P nuclear magnetic resonance (NMR) spectroscopy after NaOH–EDTA extraction revealed that P was predominantly in the inorganic pool in all three sludge samples, with the highest proportion (of the total extracted P) as inorganic P in the anaerobically digested liquid sludge. After sludge incorporation, P was immobilised to organic species. The majority of organic P was in monoester-P forms, while the remainder of organic P (diester P and phosphonate P) was more susceptible to transformations through time and showed variation with sludge type. These results show that application of sewage sludge at rates as low as 4t ha−1 can have a significant nutritional benefit to ryegrass over an initial 35-day growth and subsequent 35-day re-growth periods. Differences in P transformation, and hence nutritional benefit, between sludge types were evident throughout the experiment. Thus, differences in sludge treatment process alter the edaphic mineralisation characteristics of biosolids derived from the same source material.

Considerations on the use of the p-nitrophenyl phosphomonoesterase assay in the study of the phosphorus nutrition of soil borne fungi

The p-nitrophenyl phosphomonoesterase assay (p NPPase) is commonly used to measure cell-wall-associated and extracellular phosphatase activity of soil fungi. p NPPases are usually assayed in the context of fungal nutrition, where inorganic P supply might be enhanced by the mineralisation of monoester organic P sources in the soil. The importance of the assay to the P nutrition of soil fungi is considered based on the evidence currently available including the consistency of methodological approach. The nature of organic P in the soil and the relevance of the assay to some specific soil substrates is discussed, particularly the chemistry and bioavailability of myo-inositol hexakisphosphate and the lower inositol phosphates. The evidence for the long-term stability of p NPPases in the soil is examined in the light of the persistence of p NPPase in soils. The role of persistent extracellular fungal p NPPases in the soil P cycle is discussed. Conclusions from p NPPase based studies must be based upon an appreciation of the constraints of the assay and the complex chemistry of organic P and p NPPase in the soil.

Carbon, nitrogen and phosphorus mineralization in two soils amended with distillery yeast

The possibility of using yeast from alcohol distilleries as a source of nutrients in soil was investigated. The following treatments were used: no fertilization (control), 0.5% (w/w) yeast, 1% (w/w) yeast, and NPK. The decomposition of yeast was monitored for 90 days in two soils. The CO, production and the microbial biomass were increased by art average of 1- to 3-fold by yeast incorporation compared to control. Protease activity also was enhanced 3- to 8-fold in the soils supplemented with yeast compared to control. The phosphatase activities were higher than control only during the first days. While nitrate contents increased in all treatments compared to control, available P only increased in the soils amended with 1%, yeast or NPK by 45-119% and 309-489%, respectively. These results indicate that there exists an excellent potential for the use of yeast in the soil as a source of nitrate and available P for plant nutrition. (C) 2003 Elsevier Ltd. All rights reserved.

Phosphorus bioavailability in diets for growing horses

A study of phosphorus (P) metabolism was carried out using 12 month old Brasileiro de Hipismo breed of horses to determine the P bioavailability available from feeds commonly fed to horses in Brazil. Five different diets were formulated to contain approximately equivalent levels of crude protein and digestible energy, as well as to supply at least 22 g P/horse/day (NRC, 1989). All 5 diets contained 40% Bermuda coastal hay plus 60% concentrate. The 5 different concentrates contained: C1 (corn+ cottonseed meal) C2 (corn grain+soybean meal) C3 (corn+sugarcane yeast), C4 (oat+cottonseed meal), and C5 (oat+soybean meal). The radioisotope (32)p was injected with 30 NIB. Blood, feces and urine were collected for 7 days to evaluate endogenous fecal P and true absorption. Analysis of variance of P intake showed differences due to dietary effects (P < 0.05). Concentrate C3 had the lowest intake (79.68 mg/ kg BW). All of the diets produced positive P retention. Absolute values for P concentrations in plasma, urine, feces and endogenous feces did not vary between diets. Values for endogenous fecal P were independent of the level of P intake, so the correlation between P intake and P endogenous was not significant. P bioavailability values were 50.75; 40.98; 43.50; 51.03 and 57.68% for diets C1 through C5, respectively. However, differences in P bioavailability were found (P < 0.05) between diets. Diets C2 and C3 had lower P bioavailability than the other diets. The P bioavailability of all dietary treatments in this study exceeded NRC (1989) postulations of 35% true P absorption in diets not supplemented with inorganic P. The results of this study indicate that inorganic P supplementation is not needed for growing yearlings fed common Brazilian feeds. Considering the high cost of P supplementation and the risk of environmental P contamination, inorganic phosphorus supplementation for growing yearlings may not be required. (c) 2007 Elsevier B.V. All rights reserved.

Phosphate fertilization and phosphorus forms in an Oxisol under no-till

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Extraction methods and availability of phosphorus for soybean in soils from Parana State, Brazil

In studies on the evaluation of methodologies for the analysis of soil, phosphorus (P) has been the single most studied aspect, due to the complexity of this dynamic element in soil. However, these studies have been limited regarding soil conditions in Paranaa. The present study aimed to evaluate the efficiency of the Mehlich-1, Mehlich-3 and ion exchange resin methods in the evaluation of available P for soybean (Glycine max) in the soils of Paranaa State. Twelve soil samples collected from the upper 0-20 cm were planted with soybean for a period of 42 days in the greenhouse. The ability to extract soil P followed the order of decreasing average amount of extracted P: Mehlich-3 > resin > Mehlich-1. The correlation coefficients between the content of P extracted by Mehlich-1, Mehlich-3 and resin and the amount of P accumulated in the plants were 0.86, 0.90 and 0.93, respectively. Mehlich-1, Mehlich-3 and resin showed similar efficiency in the evaluation of P availability to plants and, under conditions of natural fertility and in soils that had received no application of poorly natural reactive phosphates, can be used to quantify the concentrations of P in the soils of Parana State.

Phosphorus and potassium budget in the soil-plant system in crop rotations under no-till

Soil management and crop rotations can affect P and K budget in soil, decreasing losses, and increasing fertilizer use efficiency. The P and K budget in the soil-plant system at depths up to 60. cm was studied for different soil managements and crop rotations under no-till for three years in Botucatu, São Paulo, Brazil. The investigated crop rotations were: triticale (X Triticosecale) and sunflower (Helianthus annuus) cropped in autumn-winter; pearl millet (Pennisetum glaucum), forage sorghum (Sorghum bicolor), and Sunn hemp (Crotalaria juncea) were grown in the spring, as well as an additional treatment with chiseling followed by a fallow period; and soybean (Glycini max, L., Merril) was cropped in the summer. Each year triticale and sunflower were grown in plots and pearl millet, forage sorghum, Sunn hemp and of chisel/fallow in sub-plots. The triticale/millet rotation led to the largest decrease in available P within the 0-0.60. m layer of the soil profile and the largest K increase within the 0-0.05. m layer. Potassium mobility in the soil profile and the increases in the available K content in the 0.40-0.60. m layer were independent of the management system. Crop rotations with or without chiseling are not effective in preventing soil P losses. There is considerable K leaching below 0.60. m, but chiseling and the use of high K accumulating plants as triticale results in lower K losses. © 2012 Elsevier B.V.

Phosphorus and potassium balance in a corn-soybean rotation under no-till and chiseling

Nutrient use efficiency has become an important issue in agriculture, and crop rotations with deep vigorous rooted cover crops under no till may be an important tool in increasing nutrient conservation in agricultural systems. Ruzigrass (Brachiaria ruziziensis) has a vigorous, deep root system and may be effective in cycling P and K. The balance of P and K in cropping systems with crop rotations using ruzigrass, pearl millet (Pennisetum glaucum) and ruzigrass + castor bean (Ricinus communis), chiseled or not, was calculated down to 0.60 m in the soil profile for 2 years. The cash crops were corn in the first year and soybean in the second year. Crop rotations under no-till increased available P amounts in the soil-plant system from 80 to 100 %, and reduced K losses between 4 and 23 %. The benefits in nutrient balance promoted by crop rotations were higher in the second year and under without chiseling. Plant residues deposited on the soil surface in no-till systems contain considerable nutrient reserve and increase fertilizer use efficiency. However, P release from ruzigrass grown as a sole crop is not synchronized with soybean uptake rate, which may result in decreased yields. © 2013 Springer Science+Business Media Dordrecht.

Congo grass grown in rotation with soybean affects phosphorus bound to soil carbon

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Soil phosphorus availability and soybean response to phosphorus starter fertilizer

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)