Characterization of the PHO1 gene family and the responses to phosphate deficiency of Physcomitrella patens.

PHO1 was previously identified in Arabidopsis (Arabidopsis thaliana) as a protein involved in loading inorganic phosphate (Pi) into the xylem of roots and its expression was associated with the vascular cylinder. Seven genes homologous to AtPHO1 (PpPHO1;1-PpPHO1;7) have been identified in the moss Physcomitrella patens. The corresponding proteins harbor an SPX tripartite domain in the N-terminal hydrophilic portion and an EXS domain in the conserved C-terminal hydrophobic portion, both common features of the plant PHO1 family. Northern-blot analysis showed distinct expression patterns for the PpPHO1 genes, both at the tissue level and in response to phosphate deficiency. Transgenic P. patens expressing the beta-glucuronidase reporter gene under three different PpPHO1 promoters revealed distinct expression profiles in various tissues. Expression of PpPHO1;1 and PpPHO1;7 was specifically induced by Pi starvation. P. patens homologs to the Arabidopsis PHT1, DGD2, SQD1, and APS1 genes also responded to Pi deficiency by increased mRNA levels. Morphological changes associated with Pi deficiency included elongation of caulonemata with inhibition of the formation of side branches, resulting in colonies with greater diameter, but reduced mass compared to Pi-sufficient plants. Under Pi-deficient conditions, P. patens also increased the synthesis of ribonucleases and of an acid phosphatase, and increased the ratio of sulfolipids over phospholipids. These results indicate that P. patens and higher plants share some common strategies to adapt to Pi deficiency, although morphological changes are distinct, and that the PHO1 proteins are well conserved in bryophyte despite the lack of a developed vascular system.

Uncoupling phosphate deficiency from its major effects on growth and transcriptome via PHO1 expression in Arabidopsis.

Inorganic phosphate (Pi) is one of the most limiting nutrients for plant growth in both natural and agricultural contexts. Pi-deficiency leads to a strong decrease in shoot growth, and triggers extensive changes at the developmental, biochemical and gene expression levels that are presumably aimed at improving the acquisition of this nutrient and sustaining growth. The Arabidopsis thaliana PHO1 gene has previously been shown to participate in the transport of Pi from roots to shoots, and the null pho1 mutant has all the hallmarks associated with shoot Pi deficiency. We show here that A. thaliana plants with a reduced expression of PHO1 in roots have shoot growth similar to Pi-sufficient plants, despite leaves being strongly Pi deficient. Furthermore, the gene expression profile normally triggered by Pi deficiency is suppressed in plants with low PHO1 expression. At comparable levels of shoot Pi supply, the wild type reduces shoot growth but maintains adequate shoot vacuolar Pi content, whereas the PHO1 underexpressor maintains maximal growth with strongly depleted Pi reserves. Expression of the Oryza sativa (rice) PHO1 ortholog in the pho1 null mutant also leads to plants that maintain normal growth and suppression of the Pi-deficiency response, despite the low shoot Pi. These data show that it is possible to unlink low shoot Pi content with the responses normally associated with Pi deficiency through the modulation of PHO1 expression or activity. These data also show that reduced shoot growth is not a direct consequence of Pi deficiency, but is more likely to be a result of extensive gene expression reprogramming triggered by Pi deficiency.

Expression analyses of three members of the AtPHO1 family reveal differential interactions between signaling pathways involved in phosphate deficiency and the responses to auxin, cytokinin, and abscisic acid.

The PHO1 protein is involved in loading inorganic phosphate (Pi) to the root xylem. Ten genes homologous to AtPHO1 are present in the Arabidopsis thaliana (L.) Heyn genome. From this gene family, transcript levels of only AtPHO1, AtPHO1;H1 and AtPHO1;H10 were increased by Pi-deficiency. While the up-regulation of AtPHO1;H1 and AtPHO1;H10 by Pi deficiency followed the same rapid kinetics and was dependent on the PHR1 transcription factor, phosphite only strongly suppressed the expression of AtPHO1;H1 and had a minor effect on AtPHO1;H10. Addition of sucrose was found to increase transcript levels of both AtPHO1 and AtPHO1;H1 in Pi-sufficient or Pi-deficient plants, but to suppress AtPHO1:H10 under the same conditions. Treatments of plants with auxin or cytokinin had contrasting effect depending on the gene and on the Pi status of the plants. Thus, while both hormones down-regulated expression of AtPHO1 independently of the plant Pi status, auxin and cytokinin up-regulated AtPHO1;H1 and AtPHO1;H10 expression in Pi-sufficient plants and down-regulated expression in Pi-deficient plants. Treatments with abscisic acid inhibited AtPHO1 and AtPHO1;H1 expression in both Pi-sufficient and Pi-deficient plants, but increased AtPHO1;H10 expression under the same conditions. The inhibition of expression by abscisic acid of AtPHO1 and AtPHO1;H1, and of the Pi-starvation responsive genes AtPHT1;1 and AtIPS1, was dependant on the ABI1 type 2C protein phosphatase. These results reveal that various levels of cross talk between the signal transduction pathways to Pi, sucrose and phytohormones are involved in the regulation of expression of the three AtPHO1 homologues.

Effectiveness of fused magnesium potassium phosphate for marandu grass

The current high price of KCl and great dependence on importation to satisfy the Brazilian demand indicate the need for studies that evaluate the efficiency of other K sources, particularly those based on domestic raw material. For this purpose, a greenhouse experiment was conducted with samples of a sandy clay loam Typic Haplustox, in a completely randomized 4 x 3 x 2 factorial design: four K rates (0, 60, 120, and 180 mg kg-1), three sources (potassium chloride (KCl), fused magnesium potassium phosphate (FMPP) and a mixture of 70 % FMPP + 30 % KCl) and two particle sizes (100 and 60 mesh), with three replications. Potassium fertilization resulted in significant increases in shoot dry matter production and in K concentrations, both in soil and plants. The K source and particle size had no significant effect on the evaluated characteristics. Potassium critical levels in the soil and the shoots were 1.53 mmol c dm-3 and 19.1 g kg-1, respectively.

Les biomarqueurs dans la maladie de Fabry: relation entre Sphingosine-1 phosphate et Lyso-Gb3

Problématique : La maladie de Fabry est une maladie métabolique à stockage lysosomal. C'est une maladie héreditaire à transmission récessive qui concerne l'enzyme alpha-Galactosidase A. Le gène de l'alpha-Galactosidase A (GLA) se trouve au niveau du bras long du chromosome X «carté en Xq21.33-Xq22 ». L'enzyme muté ne recouvre plus son rôle catabolisateur et il ne métabolise pas le substrat globotriaosylceramide (Gb3). Par conséquence le Gb3 s'accumule dans tous les tissus. Dans les parois des vaisseaux sanguins le Gb3 s'accumule dans l'endothelium, la tunique interne des vaisseaux sanguins. Ce déficit métabolique se traduit par l'épaississement de la paroi vasculaire, des processus d'infarctus et ischémies du tissu cardiaque, rénal et cérébral. L'implication cardiaque de la maladie de Fabry est décrite chez plus de 78% des patients affectés par la maladie et se manifeste par une hypertrophie cardiaque du ventricule gauche. Toutefois, il n'existe pas de relation étroite entre hypertrophie cardiaque et le Gb3. La sphingosine 1-phosphate à été identifiée dans notre laboratoire et proposée comme facteur de croissance à l'origine du remodelage cardiovasculaire. De plus, la Globotriaosylsphingosine (Lyso-Gb3) à été aussi proposée comme facteur vasoactif chez les patients Fabry. Objectif : L'identification d'un biomarqueur pour le diagnostic et le suivi thérapeutique de la maladie de Fabry représente une domaine d'investigation active en recherche scientifique. Le Gb3 plasmatique ou dans les urines, la biopsie rénale ou cardiaque qui est mis en évidence grâce à la microscopie électronique sous forme de corps concentrique lamellaires, constituent les biomarqueurs classiques de la maladie de Fabry. Dernièrement, le Lyso-Gb3 et le Sphingosine-1 phosphate (S1P) ont été proposés comme marqueurs du remodelage cardiovasculaire. Le but de ce travail est de rassembler et de discuter la littérature concernant ces nouveaux marqueurs et, d'étudier une possible interaction entre Lyso-Gb3 et le S1P. Méthodologie : Rassembler la littérature scientifique et analyser l'implication de ces marqueurs dans la maladie de Fabry et leur effets cardiovasculaires. De plus, un travail expérimental est effectué. Ce travail consiste en l'identification d'une relation possible entre le Lyso-Gb3 et le S1P. Résultats : Avec ce travail on a cherché à actualiser et mettre à jour les notions concernant les biomarqueurs qui prennent place dans cette pathologie et les connaissances qu'on possède à ce jour sur les manifestations cardiovasculaires et neurologiques.La recherche d'un biomarqueur prime par le fait qu'un nombre considerable de patients est sous-diagnostiqués pour la maladie de Fabry et que entre les taux de substrat enzymatique accumulé dans les tissus et l'hypertrophie cardiaque, on peut constater une discordance. Grâce à ce travail expérimental, on a exclue la possibilité d'un effet précurseur du lyso-Gb3 pour le S1P. Nous avons montré que le Lyso-Gb3 est reconnu par les récepteurs du S1P avec des effets commun pour les S1P1-3 et différents pour le S1P2. Les taux du Lyso-Gb3 et du S1P doivent être mesuré chez les patients Fabry et une stratégie thérapeutique doit prendre en compte le rapport S1P/Lyso-Gb3.

Mobility of inorganic and organic phosphorus forms under different levels of phosphate and poultry litter fertilization in soils

The eutrophication of aquifers is strongly linked to the mobility of P in soils. Although P mobility was considered irrelevant in a more distant past, more recent studies have shown that P, both in organic (Po) and inorganic forms (Pi), can be lost by leaching and eluviation through the soil profile, particularly in less weathered and/or sandier soils with low P adsorption capacity. The purpose of this study was to determine losses of P forms by leaching and eluviation from soil columns. Each column consisted of five PVC rings (diameter 5 cm, height 10 cm), filled with two soil types: a clayey Red-Yellow Latosol and a sandy loam Red-Yellow Latosol, which were exposed to water percolation. The soils were previously treated with four P rates (as KH2PO4 ) to reach 0, 12.5, 25.0 and 50 % of the maximum P adsorption capacity (MPAC). The P source was homogenized with the whole soil volume and incubated for 60 days. After this period the soils were placed in the columns; the soil of the top ring was mixed with five poultry litter rates of 0, 20, 40, 80, and 160 t ha-1 (dry weight basis). Treatments consisted of a 4 x 5 x 2 factorial scheme corresponding to four MPAC levels, five poultry litter rates, two soils, with three replications, arranged in a completely randomized block design. Deionized water was percolated through the columns 10 times in 35 days to simulate about 1,200 mm rainfall. In the leachate of each column the inorganic P (reactive P, Pi) and organic P forms (unreactive P, Po) were determined. At the end of the experiment, the columns were disassembled and P was extracted with the extractants Mehlich-1 (HCl 0.05 mol L-1 and H2SO4 0.0125 mol L-1) and Olsen (NaHCO3 0.5 mol L-1; pH 8.5) from the soil of each ring. The Pi and Po fractions were measured by the Olsen extractant. It was found that under higher poultry litter rates the losses of unreactive P (Po) were 6.4 times higher than of reactive P (Pi). Both the previous P fertilization and increasing poultry litter rates caused a vertical movement of P down the soil columns, as verified by P concentrations extracted by Mehlich-1 and NaHCO3 (Olsen). The environmental critical level (ECL), i.e., the P soil concentration above which P leaching increases exponentially, was 100 and 150 mg dm-3 by Mehlich-1 and 40 and 60 mg dm-3 by Olsen, for the sandy loam and clay soils, respectively. In highly weathered soils, where residual P is accumulated by successive crops, P leaching through the profile can be significant, particularly when poultry litter is applied as fertilizer.

PA21, a New Iron-Based Noncalcium Phosphate Binder, Prevents Vascular Calcification in Chronic Renal Failure Rats.

Chronic renal failure (CRF) is associated with the development of secondary hyperparathyroidism and vascular calcifications. We evaluated the efficacy of PA21, a new iron-based noncalcium phosphate binder, in controlling phosphocalcic disorders and preventing vascular calcifications in uremic rats. Rats with adenine-diet-induced CRF were randomized to receive either PA21 0.5, 1.5, or 5% or CaCO3 3% in the diet for 4 weeks, and were compared with uremic and nonuremic control groups. After 4 weeks of phosphate binder treatment, serum calcium, creatinine, and body weight were similar between all CRF groups. Serum phosphorus was reduced with CaCO3 3% (2.06 mM; P ≤ 0.001), PA21 1.5% (2.29 mM; P < 0.05), and PA21 5% (2.21 mM; P ≤ 0.001) versus CRF controls (2.91 mM). Intact parathyroid hormone was strongly reduced in the PA21 5% and CaCO3 3% CRF groups to a similar extent (1138 and 1299 pg/ml, respectively) versus CRF controls (3261 pg/ml; both P ≤ 0.001). A lower serum fibroblast growth factor 23 concentration was observed in the PA21 5%, compared with CaCO3 3% and CRF, control groups. PA21 5% CRF rats had a lower vascular calcification score compared with CaCO3 3% CRF rats and CRF controls. In conclusion, PA21 was as effective as CaCO3 at controlling phosphocalcic disorders but superior in preventing the development of vascular calcifications in uremic rats. Thus, PA21 represents a possible alternative to calcium-based phosphate binders in CRF patients.

The emerging importance of the SPX domain-containing proteins in phosphate homeostasis

Plant growth and development are strongly influenced by the availability of nutrients in the soil solution. Among them, phosphorus (P) is one of the most essential and most limiting macro-elements for plants. In the environment, plants are often confronted with P starvation as a result of extremely low concentrations of soluble inorganic phosphate (Pi) in the soil. To cope with these conditions, plants have developed a wide spectrum of mechanisms aimed at increasing P use efficiency. At the molecular level, recent studies have shown that several proteins carrying the SPX domain are essential for maintaining Pi homeostasis in plants. The SPX domain is found in numerous eukaryotic proteins, including several proteins from the yeast PHO regulon, involved in maintaining Pi homeostasis. In plants, proteins harboring the SPX domain are classified into four families based on the presence of additional domains in their structure, namely the SPX, SPX-EXS, SPX-MFS and SPX-RING families. In this review, we highlight the recent findings regarding the key roles of the proteins containing the SPX domain in phosphate signaling, as well as providing further research directions in order to improve our knowledge on P nutrition in plants, thus enabling the generation of plants with better P use efficiency.

Mycorrhizal effectiveness on physic nut as influenced by phosphate fertilization levels

In recent years, physic nut (Jatropha curcas L.) has attracted attention because of its potential for biofuel production. Although it is adapted to low-fertility soils, physic nut requires soil acidity corrections and addition of a considerable amount of fertilizer for high productivity. The objective of this research was to evaluate the effectiveness of arbuscular mycorrhizal fungi (AMF) (control without AMF inoculation, Gigaspora margarita inoculation or Glomus clarum inoculation) on increasing growth and yield of physic nut seedlings under different rates of P fertilization (0, 25, 50, 100, 200, and 400 mg kg-1 P soil) in greenhouse. The experiment was arranged in a completely randomized, block in a factorial scheme design with four replications. The physic nut plants were harvested 180 days after the beginning of the experiment. Mycorrhizal inoculation increased physic nut growth, plant P concentration and root P uptake efficiency at low soil P concentrations. The P use quotient of the plants decreased as the amount of P applied increased, and the P use efficiency index increased at low P levels and decreased at high P levels. Mycorrhizal root colonization and AMF sporulation were negatively affected by P addition. The highest mycorrhizal efficiency was observed when the soil contained between 7.8 and 25 mgkg-1 of P. The physic nut plants responded strongly to P application, independent of mycorrhizal inoculation.

Exudative mineral losses after serious burns: a clue to the alterations of magnesium and phosphate metabolism.

Hypomagnesemia and hypophosphatemia are frequent after severe burns; however, increased urinary excretion does not sufficiently explain the magnitude of the mineral depletion. We measured the mineral content of cutaneous exudates during the first week after injury. Sixteen patients aged 34 +/- 9 y (mean +/- SD) with thermal burns were studied prospectively and divided in 3 groups according to the extent of their burn injury and the presence or absence of mineral supplements: group 1 (n = 5), burns covering 26 +/- 5% of body surface; group 2 (n = 6), burns covering 41 +/- 10%; and group 3 (n = 5), burns covering 42 +/- 6% with prescription of magnesium and phosphate supplements. Cutaneous exudates were extracted from the textiles (surgical drapes, dressings, sheets, etc) surrounding the patients from day 1 to day 7 after injury. Mean magnesium serum concentrations decreased below reference ranges in 12 patients between days 1 and 4 and normalized thereafter. Phosphate, normal on day 0, was low during the first week. Albumin concentrations, normal on day 0, decreased and remained low. Urinary magnesium and phosphate excretion were within reference ranges and not larger in group 3. Mean daily cutaneous losses were 16 mmol Mg/d and 11 mmol P/d (largest in group 2). Exudative magnesium losses were correlated with burn severity (r = 0.709, P = 0.003). Cutaneous magnesium losses were nearly four times larger than urinary losses whereas cutaneous phosphate losses were smaller than urinary phosphate losses. Mean daily losses of both magnesium and phosphate were more than the recommended dietary allowances. Exudative losses combined with urinary losses largely explained the increased mineral requirements after burn injury.

Competitive sorption and desorption of phosphate and citrate in clayey and sandy loam soils

The increase of organic acids in soils can reduce phosphorus sorption. The objective of the study was to evaluate the competitive sorption of P and citrate in clayey and sandy loam soils, using a stirred-flow system. Three experiments were performed with soil samples (0-20 cm layer) of clayey (RYL-cl) and sandy loam (RYL-sl) Red Yellow Latosols (Oxisols). In the first study, the treatments were arranged in a 2 × 5 factorial design, with two soil types and five combinations of phosphorus and citrate application (only P; P + citrate; and citrate applied 7, 22, 52 min before P); in the second, the treatments were arranged in a 2 × 2 factorial design, corresponding to two soils and two forms of P and citrate application (only citrate and citrate + P); and in the third study, the treatments in a 2 × 2 × 6 factorial design consisted of two soils, two extractors (citrate and water) and six incubation times. In the RYL-cl and RYL-sl, P sorption was highest (44 and 25 % of P application, respectively), in the absence of citrate application. Under citrate application, P sorption was reduced in all treatments. The combined application of citrate and P reduced P sorption to 25.8 % of the initially applied P in RYL-cl and to 16.7 % in RYL-sl, in comparison to P without citrate. Citrate sorption in RYL-cl and RYL-sl was highest in the absence of P application, corresponding to 32.0 and 30.2 % of the citrate applied, respectively. With P application, citrate sorption was reduced to 26.4 and 19.7 % of the initially applied citrate in RYL-cl and RYL-sl, respectively. Phosphorus desorption was greater when citrate was used. Phosphorus desorption with citrate and water was higher in the beginning (until 24 h of incubation of P) in RYL-cl and RYL-sl, indicating a rapid initial phase, followed by a slow release phase. This suggests that according to the contact time of P with the soil colloids, the previously adsorbed P can be released to the soil solution in the presence of competing ligands such as citrate. In conclusion, a soil management with continuous input of organic acids is desirable, in view of their potential to compete for P sorption sites, especially in rather weathered soils.

Looming Scarcity of Phosphate Rock and Intensification of Soil Phosphorus Research

In recent years, many researchers have claimed that world reserves of rock phosphate were getting depleted at an alarming rate, putting us on the path to scarcity of that essential resource within the next few decades. Others have claimed that such alarmist forecasts were frequent in the past and have always been proven unfounded, making it likely that the same will be true in the future. Both viewpoints are directly relevant to the level of funding devoted to research on the use of phosphate fertilizers. In this short essay, it is argued that information about future reserves of P or any other resource are impossible to predict, and therefore that the threat of a possible depletion of P reserves should not be used as a key motivation for an intensification of research on soil P. However, there are other, more compelling reasons, both geopolitical and environmental, to urgently step up our collective efforts to devise agricultural practices that make better use of P than is the case at the moment.

LOSS OF EXTRACTION CAPACITY OF MEHLICH-1 AND MONOCALCIUM PHOSPHATE AS A VARIABLE OF REMAINING P AND ITS RELATIONSHIP TO CRITICAL LEVELS OF SOIL PHOSPHORUS AND SULFUR

The Mehlich-1 (M-1) extractant and Monocalcium Phosphate in acetic acid (MCPa) have mechanisms for extraction of available P and S in acidity and in ligand exchange, whether of the sulfate of the extractant by the phosphate of the soil, or of the phosphate of the extractant by the sulfate of the soil. In clayey soils, with greater P adsorption capacity, or lower remaining P (Rem-P) value, which corresponds to soils with greater Phosphate Buffer Capacity (PBC), more buffered for acidity, the initially low pH of the extractants increases over their time of contact with the soil in the direction of the pH of the soil; and the sulfate of the M-1 or the phosphate of the MCPa is adsorbed by adsorption sites occupied by these anions or not. This situation makes the extractant lose its extraction capacity, a phenomenon known as loss of extraction capacity or consumption of the extractant, the object of this study. Twenty soil samples were chosen so as to cover the range of Rem-P (0 to 60 mg L-1). Rem-P was used as a measure of the PBC. The P and S contents available from the soil samples through M-1 and MCPa, and the contents of other nutrients and of organic matter were determined. For determination of loss of extraction capacity, after the rest period, the pH and the P and S contents were measured in both the extracts-soils. Although significant, the loss of extraction capacity of the acidity of the M-1 and MCPa extractants with reduction in the Rem-P value did not have a very expressive effect. A “linear plateau” model was observed for the M-1 for discontinuous loss of extraction capacity of the P content in accordance with reduction in the concentration of the Rem-P or increase in the PBC, suggesting that a discontinuous model should also be adopted for interpretation of available P of soils with different Rem-P values. In contrast, a continuous linear response was observed between the P variables in the extract-soil and Rem-P for the MCPa extractor, which shows increasing loss of extraction capacity of this extractor with an increase in the PBC of the soil, indicating the validity of the linear relationship between the available S of the soil and the PBC, estimated by Rem-P, as currently adopted.

Novel acid phosphatase in Candida glabrata suggests selective pressure and niche specialization in the phosphate signal transduction pathway.

Evolution through natural selection suggests unnecessary genes are lost. We observed that the yeast Candida glabrata lost the gene encoding a phosphate-repressible acid phosphatase (PHO5) present in many yeasts including Saccharomyces cerevisiae. However, C. glabrata still had phosphate starvation-inducible phosphatase activity. Screening a C. glabrata genomic library, we identified CgPMU2, a member of a three-gene family that contains a phosphomutase-like domain. This small-scale gene duplication event could allow for sub- or neofunctionalization. On the basis of phylogenetic and biochemical characterizations, CgPMU2 has neofunctionalized to become a broad range, phosphate starvation-regulated acid phosphatase, which functionally replaces PHO5 in this pathogenic yeast. We determined that CgPmu2, unlike ScPho5, is not able to hydrolyze phytic acid (inositol hexakisphosphate). Phytic acid is present in fruits and seeds where S. cerevisiae grows, but is not abundant in mammalian tissues where C. glabrata grows. We demonstrated that C. glabrata is limited from an environment where phytic acid is the only source of phosphate. Our work suggests that during evolutionary time, the selection for the ancestral PHO5 was lost and that C. glabrata neofunctionalized a weak phosphatase to replace PHO5. Convergent evolution of a phosphate starvation-inducible acid phosphatase in C. glabrata relative to most yeast species provides an example of how small changes in signal transduction pathways can mediate genetic isolation and uncovers a potential speciation gene.

Spatio-temporal transcript profiling of rice roots and shoots in response to phosphate starvation and recovery.

Using rice (Oryza sativa) as a model crop species, we performed an in-depth temporal transcriptome analysis, covering the early and late stages of Pi deprivation as well as Pi recovery in roots and shoots, using next-generation sequencing. Analyses of 126 paired-end RNA sequencing libraries, spanning nine time points, provided a comprehensive overview of the dynamic responses of rice to Pi stress. Differentially expressed genes were grouped into eight sets based on their responses to Pi starvation and recovery, enabling the complex signaling pathways involved in Pi homeostasis to be untangled. A reference annotation-based transcript assembly was also generated, identifying 438 unannotated loci that were differentially expressed under Pi starvation. Several genes also showed induction of unannotated splice isoforms under Pi starvation. Among these, PHOSPHATE2 (PHO2), a key regulator of Pi homeostasis, displayed a Pi starvation-induced isoform, which was associated with increased translation activity. In addition, microRNA (miRNA) expression profiles after long-term Pi starvation in roots and shoots were assessed, identifying 20 miRNA families that were not previously associated with Pi starvation, such as miR6250. In this article, we present a comprehensive spatio-temporal transcriptome analysis of plant responses to Pi stress, revealing a large number of potential key regulators of Pi homeostasis in plants.