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Phosphorus is an essential macronutrient for plant growth and development. Root system architecture (RSA) affects a plant's ability to obtain phosphate, the major form of phosphorus that plants uptake. In this review, I first consider the relationship between RSA and plant phosphorus-acquisition efficiency, describe how external phosphorus conditions both induce and impose changes in the RSA of major crops and of the model plant Arabidopsis, and discuss whether shoot phosphorus status affects RSA and whether there is a universal root developmental response across all plant species. I then summarize the current understanding of the molecular mechanisms governing root developmental responses to phosphorus deficiency. I also explore the possible reasons for the inconsistent results reported by different research groups and comment on the relevance of some studies performed under laboratory conditions to what occurs in natural environments., (© 2021 Institute of Botany, Chinese Academy of Sciences.)

Plant acclimatory responses to phosphate (Pi) starvation stress include the accumulation of carbohydrates, namely sugar and starch. However, whether altered endogenous carbohydrate profile could in turn affect plant Pi starvation responses remains widely unexplored. Here, two genes encoding the large and small subunits of an ADP-glucose pyrophosphorylase (AGP) in rice (Oryza sativa), AGP Large Subunit 1 (AGPL1) and AGP Small Subunit 1 (AGPS1), were functionally characterized with regard to maintenance of phosphorus (P) homeostasis and regulation of Pi starvation signaling. AGPL1 and AGPS1 were both positively responsive to nitrogen (N) or Pi deprivation, and expressed in almost all the tissues except in the meristem and mature zones of root. AGPL1 and AGPS1 physically interacted in chloroplast, and catalyzed the rate-limiting step of starch biosynthesis. Low-N- (LN) and low-Pi (LP)-triggered starch accumulation in leaves was impaired in agpl1, agps1 and apgl1 agps1 mutants compared with the wild-type plants. By contrast, mutation of AGPL1 and/or AGPS1 led to an increase in the content of the major sugar, sucrose, in leaf sheath and root under control and LN conditions. Moreover, the Pi accumulation was enhanced in the mutants under control and LN conditions, but not LP conditions. Notably, the LN-induced suppression of Pi accumulation was compromised attributed to the mutation of AGPL1 and/or AGPS1. Furthermore, the increased Pi accumulation was accompanied by the specific suppression of OsSPX2 and activation of several Pi transporter genes. These results indicate that a balanced level of carbohydrates is vital for maintaining plant P homeostasis., (© 2020 Society for Experimental Biology and John Wiley & Sons Ltd.)

Phosphorus, in its orthophosphate form (P(i)), is one of the most limiting macronutrients in soils for plant growth and development. However, the whole-genome molecular mechanisms contributing to plant acclimation to P(i) deficiency remain largely unknown. White lupin (Lupinus albus) has evolved unique adaptations for growth in P(i)-deficient soils, including the development of cluster roots to increase root surface area. In this study, we utilized RNA-Seq technology to assess global gene expression in white lupin cluster roots, normal roots, and leaves in response to P(i) supply. We de novo assembled 277,224,180 Illumina reads from 12 complementary DNA libraries to build what is to our knowledge the first white lupin gene index (LAGI 1.0). This index contains 125,821 unique sequences with an average length of 1,155 bp. Of these sequences, 50,734 were transcriptionally active (reads per kilobase per million reads ≥ 3), representing approximately 7.8% of the white lupin genome, using the predicted genome size of Lupinus angustifolius as a reference. We identified a total of 2,128 sequences differentially expressed in response to P(i) deficiency with a 2-fold or greater change and P ≤ 0.05. Twelve sequences were consistently differentially expressed due to P(i) deficiency stress in three species, Arabidopsis (Arabidopsis thaliana), potato (Solanum tuberosum), and white lupin, making them ideal candidates to monitor the P(i) status of plants. Additionally, classic physiological experiments were coupled with RNA-Seq data to examine the role of cytokinin and gibberellic acid in P(i) deficiency-induced cluster root development. This global gene expression analysis provides new insights into the biochemical and molecular mechanisms involved in the acclimation to P(i) deficiency.

White lupin (Lupinus albus) is a legume that is very efficient in accessing unavailable phosphorus (Pi). It develops short, densely clustered tertiary lateral roots (cluster/proteoid roots) in response to Pi limitation. In this report, we characterize two glycerophosphodiester phosphodiesterase (GPX-PDE) genes (GPX-PDE1 and GPX-PDE2) from white lupin and propose a role for these two GPX-PDEs in root hair growth and development and in a Pi stress-induced phospholipid degradation pathway in cluster roots. Both GPX-PDE1 and GPX-PDE2 are highly expressed in Pi-deficient cluster roots, particularly in root hairs, epidermal cells, and vascular bundles. Expression of both genes is a function of both Pi availability and photosynthate. GPX-PDE1 Pi deficiency-induced expression is attenuated as photosynthate is deprived, while that of GPX-PDE2 is strikingly enhanced. Yeast complementation assays and in vitro enzyme assays revealed that GPX-PDE1 shows catalytic activity with glycerophosphocholine while GPX-PDE2 shows highest activity with glycerophosphoinositol. Cell-free protein extracts from Pi-deficient cluster roots display GPX-PDE enzyme activity for both glycerophosphocholine and glycerophosphoinositol. Knockdown of expression of GPX-PDE through RNA interference resulted in impaired root hair development and density. We propose that white lupin GPX-PDE1 and GPX-PDE2 are involved in the acclimation to Pi limitation by enhancing glycerophosphodiester degradation and mediating root hair development.

Roots of phosphorus (P)-deficient white lupin exhibit striking changes in morphology and gene expression. In this report we provide further insight into genetic elements affecting transcription of P-deficiency-induced genes. Moreover, we also show that sugars and photosynthates are integrally related to P-deficiency-induced gene expression. White lupin phosphate transporter (LaPT1) and secreted acid phosphatase (LaSAP1) promoter-reporter genes when transformed into alfalfa, a heterologous legume, showed significant induction in roots specifically in response to P-deficiency. In addition, both promoters were active in nitrogen-fixing root nodules but not in ineffective nodules indicating a link between P-deficiency and factors related to nitrogen fixation/metabolism. As sugars play a role in signal transduction during nitrogen assimilation and are required for effective nitrogen fixation, we tested the relationship of sugars to P-deficiency-induced gene expression. Exogenous sucrose, glucose, and fructose stimulated LaPT1 and LaSAP1 transcript accumulation in dark-grown P-sufficient white lupin seedlings. Furthermore, in intact P-deficient white lupin plants, LaPT1 and LaSAP1 expression in cluster roots was strikingly reduced in dark-adapted plants with expression rapidly restored upon reexposure to light. Likewise, interruption of phloem supply to P-deficient roots resulted in a rapid decline in LaPT1 and LaSAP1 transcript accumulation. Similar results were also obtained with a third lupin P-deficiency-induced gene encoding a putative multidrug and toxin efflux protein (LaMATE). Taken together, our data show that the regulation of P-deficiency-induced genes is conserved across plant species and sugars/photosynthates are crucial for P-deficiency signal transduction.

The aim of this study was to characterize the transcriptome of a balanced polymorphism, under the regulation of a single gene, for phosphate fertilizer responsiveness/arsenate tolerance in wild grass Holcus lanatus genotypes screened from the same habitat. De novo transcriptome sequencing, RNAseq (RNA sequencing) and single nucleotide polymorphism (SNP) calling were conducted on RNA extracted from H. lanatus. Roche 454 sequencing data were assembled into c. 22,000 isotigs, and paired-end Illumina reads for phosphorus-starved (P-) and phosphorus-treated (P+) genovars of tolerant (T) and nontolerant (N) phenotypes were mapped to this reference transcriptome. Heatmaps of the gene expression data showed strong clustering of each P+/P- treated genovar, as well as clustering by N/T phenotype. Statistical analysis identified 87 isotigs to be significantly differentially expressed between N and T phenotypes and 258 between P+ and P- treated plants. SNPs and transcript expression that systematically differed between N and T phenotypes had regulatory function, namely proteases, kinases and ribonuclear RNA-binding protein and transposable elements. A single gene for arsenate tolerance led to distinct phenotype transcriptomes and SNP profiles, with large differences in upstream post-translational and post-transcriptional regulatory genes rather than in genes directly involved in P nutrition transport and metabolism per se., (© 2013 The Authors. New Phytologist © 2013 New Phytologist Trust.)

Phosphorus, a crucial macronutrient essential for plant growth and development. Due to widespread phosphorus deficiency in soils, phosphorus deficiency stress has become one of the major abiotic stresses that plants encounter. Despite the evolution of adaptive mechanisms in plants to address phosphorus deficiency, the specific strategies employed by species such as Epimedium pubescens remain elusive. Therefore, this study observed the changes in the growth, physiological reponses, and active components accumulation in E. pubescensunder phosphorus deficiency treatment, and integrated transcriptome and miRNA analysis, so as to offer comprehensive insights into the adaptive mechanisms employed by E. pubescens in response to phosphorus deficiency across various stages of phosphorus treatment. Remarkably, our findings indicate that phosphorus deficiency induces root growth stimulation in E. pubescens, while concurrently inhibiting the growth of leaves, which are of medicinal value. Surprisingly, this stressful condition results in an augmented accumulation of active components in the leaves. During the early stages (30 days), leaves respond by upregulating genes associated with carbon metabolism, flavonoid biosynthesis, and hormone signaling. This adaptive response facilitates energy production, ROS scavenging, and morphological adjustments to cope with short-term phosphorus deficiency and sustain its growth. As time progresses (90 days), the expression of genes related to phosphorus cycling and recycling in leaves is upregulated, and transcriptional and post-transcriptional regulation (miRNA regulation and protein modification) is enhanced. Simultaneously, plant growth is further suppressed, and it gradually begins to discard and decompose leaves to resist the challenges of long-term phosphorus deficiency stress and sustain survival. In conclusion, our study deeply and comprehensively reveals adaptive strategies utilized by E. pubescens in response to phosphorus deficiency, demonstrating its resilience and thriving potential under stressful conditions. Furthermore, it provides valuable information on potential target genes for the cultivation of E. pubescens genotypes tolerant to low phosphorus. [ABSTRACT FROM AUTHOR]

Oryza sativa PHOSPHATE RESPONSE2 (OsPHR2) can promote the uptake and use of phosphorus (P) in rice. We introduced OsPHR2 into the winter wheat (Triticum aestivum L.) variety "Zhengmai0856." OsPHR2 was integrated into the wheat genome with two copy numbers and could be correctly transcribed and expressed. OsPHR2 was mainly expressed in the leaves at the seedling stage. From the jointing to filling stage, OsPHR2 was mainly expressed in the roots, followed by the leaves, with a low expression level in detected the tassels and stems. The transgenic lines exhibited higher P accumulation at each growth stage and increased P uptake intensity in comparison to the wild type under low P and high P conditions. Analysis of the root characteristics showed that the transgenic expression of OsPHR2 increased the maximum root length, total root length, root-to-shoot ratio, and root volume under the conditions of P deficiency or low P. A field experiment showed that the transgenic lines had a higher grain yield than the wild type under low P and high P conditions. The yield of the transgenic lines increased by 6.29% and 3.73% on average compared with the wild type under low P and high P conditions, respectively. Thus, the transgenic expression of OsPHR2 could increase P uptake and yield in wheat, but the effect was more prominent under low P conditions. [ABSTRACT FROM AUTHOR]

Due to the non-uniform distribution of inorganic phosphate (Pi) in the soil, plants modify their root architecture to improve acquisition of this nutrient. In this study, a split-root system was employed to assess the nature of local and systemic signals that modulate root architecture of Brassica napus grown with non-uniform Pi availability. Lateral root (LR) growth was regulated systemically by non-uniform Pi distribution, by increasing the second-order LR (2°LR) density in compartments with high Pi supply but decreasing it in compartments with low Pi availability. Transcriptomic profiling identified groups of genes regulated, both locally and systemically, by Pi starvation. The number of systemically induced genes was greater than the number of genes locally induced, and included genes related to abscisic acid (ABA) and jasmonic acid (JA) signalling pathways, reactive oxygen species (ROS) metabolism, sucrose, and starch metabolism. Physiological studies confirmed the involvement of ABA, JA, sugars, and ROS in the systemic Pi starvation response. Our results reveal the mechanistic basis of local and systemic responses of B. napus to Pi starvation and provide new insights into the molecular and physiological basis of root plasticity. [ABSTRACT FROM AUTHOR]

The high-quality phosphorus (P) rocks on the Earth will be depleted within the next 50 years. P-mobilizing microbes can mobilize soil unavailable P and reduce P fertilizer use in agriculture. Therefore, incubation and field experiments were conducted to study the effects of a new white-rot fungus Ceriporia lacerata HG2011 on soil P mobilization and canola yield and P uptake to understand the potential of this fungus as a biofertilizer. In the soil incubation experiment, available P and phosphatase activity increased, whereas soil pH decreased as the fungal hyphae grew on the soil surfaces and in the soil. Accordingly, Olsen P was positively correlated with the phosphatase activity (r = 0.565, p ≤ 0.01) but negatively with the pH (r = −0.506, p ≤ 0.01 or 0.05). The mechanisms of P mobilization by this fungus could thus be soil acidification to solubilize inorganic P and phosphatase efflux to mineralize organic P. Compared with chemical fertilizers alone, the application of this fungus in combination with chemical fertilizers increased canola yield by 3.94–9.49% in the field. Our findings firstly demonstrated the new use of C. lacerata HG2011 as a biofertilizer in canola cultivation. [ABSTRACT FROM AUTHOR]

Phosphorus (P) limitation is a significant factor restricting crop production in agricultural systems, and enhancing the internal P utilization efficiency (PUE) of crops plays an important role in ensuring sustainable P use in agriculture. To better understand how P is remobilized to affect crop growth, we first screened P-efficient (B73 and GEMS50) and P-inefficient (Liao5114) maize genotypes at the same shoot P content, and then analyzed P pools and performed non-targeted metabolomic analyses to explore changes in cellular P fractions and metabolites in maize genotypes with contrasting PUE. We show that lipid P and nucleic acid P concentrations were significantly lower in lower leaves of P-efficient genotypes, and these P pools were remobilized to a major extent in P-efficient genotypes. Broad metabolic alterations were evident in leaves of P-efficient maize genotypes, particularly affecting products of phospholipid turnover and phosphorylated compounds, and the shikimate biosynthesis pathway. Taken together, our results suggest that P-efficient genotypes have a high capacity to remobilize lipid P and nucleic acid P and promote the shikimate pathway towards efficient P utilization in maize. [ABSTRACT FROM AUTHOR]

Symbiotic nitrogen (N) fixation entails successful interaction between legume hosts and rhizobia that occur in specialized organs called nodules. N-fixing legumes have a higher demand for phosphorus (P) than legumes grown on mineral N. Medicago truncatula is an important model plant for characterization of effects of P deficiency at the molecular level. Hence, a study was carried out to address the alteration in metabolite levels of M. truncatula grown aeroponically and subjected to 4 weeks of P stress. First, GC-MS-based untargeted metabolomics initially revealed changes in the metabolic profile of nodules, with increased levels of amino acids and sugars and a decline in amounts of organic acids. Subsequently, LC-MS/MS was used to quantify these compounds including phosphorylated metabolites in the whole plant. Our results showed a drastic reduction in levels of organic acids and phosphorylated compounds in –P leaves, with a moderate reduction in –P roots and nodules. Additionally, sugars and amino acids were elevated in the whole plant under P deprivation. These findings provide evidence that N fixation in M. truncatula is mediated through a N feedback mechanism that in parallel is related to carbon and P metabolism. [ABSTRACT FROM AUTHOR]

Phosphorus (P) is limiting nutrient in many soils, and P availability may often depend on iron (Fe) speciation. Colloidal iron phosphate (FePO4coll) is potentially present in soils, and we tested the hypothesis that phytate exudation by Pteris vittata might dissolve FePO4coll by growing the plant in nutrient solution to which FePO4coll was added. The omission of P and Fe increased phytate exudation by P. vittata from 434 to 2136 mg kg−1 as the FePO4coll concentration increased from 0 to 300 mM. The total P in P. vittata tissue increased from 2880 to 8280 mg kg−1, and the corresponding increases in the trichloroacetic acid (TCA) extractable P fractions were inorganic P (860–5100 mg kg−1), soluble organic P (250–870 mg kg−1), and insoluble organic P (160–2030 mg kg−1). That is, FePO4-solubilizing activity was positive correlated with TP, TCA P fractions in P. vittata, TP in growth media, and root exudates. This study shows that phytate exudation dissolved FePO4coll due to the chelation effect of phytic acid on Fe; however, the wider question of whether phytic acid excretion was prompted by deprivation of P, Fe, or both remains to be answered. [ABSTRACT FROM AUTHOR]

Background Limitation of plant productivity by phosphorus (P) supply is widespread and will probably increase in the future. Relatively large amounts of P fertilizer are applied to sustain crop growth and development and to achieve high yields. However, with increasing P application, plant P efficiency generally declines, which results in greater losses of P to the environment with detrimental consequences for ecosystems. Scope A strategy for reducing P input and environmental losses while maintaining or increasing plant performance is the development of crops that take up P effectively from the soil (P acquisition efficiency) or promote productivity per unit of P taken up (P utilization efficiency). In this review, we describe current research on P metabolism and transport and its relevance for improving P utilization efficiency. Conclusions Enhanced P utilization efficiency can be achieved by optimal partitioning of cellular P and distributing P effectively between tissues, allowing maximum growth and biomass of harvestable plant parts. Knowledge of the mechanisms involved could help design and breed crops with greater P utilization efficiency. [ABSTRACT FROM AUTHOR]

Background: White lupin (Lupinus albus) is a leguminous crop with elite adaptive ability in phosphorus-deficient soil and used as a model plant for studying phosphorus (P) use. However, the genetic basis of its adaptation to low P (LP) remains unclear. ATPase binding cassette (ABC) transports G subfamily play a crucial role in the transportation of biological molecules across the membrane. To date, identification of this subfamily has been analyzed in some plants, but no systematic analysis of these transporters in phosphorus acquisition is available for white lupin. Results: This study identified 66 ABCG gene family members in the white lupin genome using comprehensive approaches. Phylogenetic analysis of white lupin ABCG transporters revealed six subclades based on their counterparts in Arabidopsis, displaying distinct gene structure and motif distribution in each cluster. Influences of the whole genome duplication on the evolution of L.albABCGs were investigated in detail. Segmental duplications appear to be the major driving force for the expansion of ABCGs in white lupin. Analysis of the Ka/Ks ratios indicated that the paralogs of the L.albABCG subfamily members principally underwent purifying selection. However, it was found that L.albABCG29 was a result of both tandem and segmental duplications. Overexpression of L.albABCG29 in white lupin hairy root enhanced P accumulation in cluster root under LP and improved plant growth. Histochemical GUS staining indicated that L.albABCG29 expression increased under LP in white lupin roots. Further, overexpression of L.albABCG29 in rice significantly improved P use under combined soil drying and LP by improving root growth associated with increased rhizosheath formation. Conclusion: Through systematic and comprehensive genome-wide bioinformatics analysis, including conserved domain, gene structures, chromosomal distribution, phylogenetic relationships, and gene duplication analysis, the L.albABCG subfamily was identified in white lupin, and L.albABCG29 characterized in detail. In summary, our results provide deep insight into the characterization of the L.albABCG subfamily and the role of L.albABCG29 in improving P use. [ABSTRACT FROM AUTHOR]

Homeostasis of phosphorus (P), an essential macronutrient, is vital for plant growth under diverse environmental conditions. Although plants acquire P from the soil as inorganic phosphate (Pi), its availability is generally limited. Therefore, plants employ mechanisms involving various Pi transporters that facilitate efficient Pi uptake against a steep concentration gradient across the plant–soil interface. Among the different types of Pi transporters in plants, some members of the PHOSPHATE TRANSPORTER 1 (PHT1) family, present in the plasma membrane of root epidermal cells and root hairs, are chiefly responsible for Pi uptake from the rhizosphere. Therefore, accurate regulation of PHT1 expression is crucial for the maintenance of P homeostasis. Previous investigations positioned the Pi-dependent posttranslational regulation of PHOSPHATE STARVATION RESPONSE 1 (PHR1) transcription factor activity at the center of the regulatory mechanism controlling PHT1 expression and P homeostasis; however, recent studies indicate that several other factors also regulate the expression of PHT1 to modulate P acquisition and sustain P homeostasis against environmental fluctuations. Together with PHR1, several transcription factors that mediate the availability of other nutrients (such as nitrogen and zinc), light, and stress signals form an intricate transcriptional network to maintain P homeostasis under highly diverse environments. In this review, we summarize this intricate transcriptional network for the maintenance of P homeostasis under different environmental conditions, with a main focus on the mechanisms identified in Arabidopsis. [ABSTRACT FROM AUTHOR]

Nitrogen (N) and phosphorus (P) are two major plant nutrients, and their deficiencies often limit plant growth and crop yield. The uptakes of N or P affect each other, and consequently, understanding N–P interactions is fundamental. Their signaling mechanisms have been studied mostly separately, and integrating N–P interactive regulation is becoming the aim of some recent works. Lupins are singular plants, as, under N and P deficiencies, they are capable to develop new organs, the N2-fixing symbiotic nodules, and some species can also transform their root architecture to form cluster roots, hundreds of short rootlets that alter their metabolism to induce a high-affinity P transport system and enhance synthesis and secretion of organic acids, flavonoids, proteases, acid phosphatases, and proton efflux. These modifications lead to mobilization in the soil of, otherwise unavailable, P. White lupin (Lupinus albus) represents a model plant to study cluster roots and for understanding plant acclimation to nutrient deficiency. It tolerates simultaneous P and N deficiencies and also enhances uptake of additional nutrients. Here, we present the structural and functional modifications that occur in conditions of P and N deficiencies and lead to the organogenesis and altered metabolism of nodules and cluster roots. Some known N and P signaling mechanisms include different factors, including phytohormones and miRNAs. The combination of the individual N and P mechanisms uncovers interactive regulation pathways that concur in nodules and cluster roots. L. albus interlinks N and P recycling processes both in the plant itself and in nature. [ABSTRACT FROM AUTHOR]

Knowing how switchgrass (Panicum virgatum L.) responds and adapts to phosphorus (P)‐limitation will aid efforts to optimize P acquisition and use in this species for sustainable biomass production. This integrative study investigated the impacts of mild, moderate, and severe P‐stress on genome transcription and whole‐plant metabolism, physiology and development in switchgrass. P‐limitation reduced overall plant growth, increased root/shoot ratio, increased root branching at moderate P‐stress, and decreased root diameter with increased density and length of root hairs at severe P‐stress. RNA‐seq analysis revealed thousands of genes that were differentially expressed under moderate and severe P‐stress in roots and/or shoots compared to P‐replete plants, with many stress‐induced genes involved in transcriptional and other forms of regulation, primary and secondary metabolism, transport, and other processes involved in P‐acquisition and homeostasis. Amongst the latter were multiple miRNA399 genes and putative targets of these. Metabolite profiling showed that levels of most sugars and sugar alcohols decreased with increasing P stress, while organic and amino acids increased under mild and moderate P‐stress in shoots and roots, although this trend reversed under severe P‐stress, especially in shoots. Low phosphorus (P) availability limits plant growth in most soils. We performed a multilevel analysis of P‐limitation responses in the perennial bioenergy crop switchgrass. Elucidation of strategies and mechanisms deployed by switchgrass to cope with P‐limitation will help in efforts to optimize P‐efficiency and maximize growth in low P conditions. [ABSTRACT FROM AUTHOR]

The use of arbuscular mycorrhizal fungi (AMF) to enhance soil phosphate uptake is a biological technique considered to cure phosphate deficiency in soils. This study investigated association of Aspilia pruliseta Schweif shrub with AMF in Kenya. The study aims at profiling a tropical shrub with multiple ecological benefits that could reduce addition of chemical phosphatic fertilizer into the soil and reverse negative consequences of eutrophication. Sampling was purposive to have areas with or without Aspilia pruliseta vegetation growing. A small amount (10 g) of the soil from 27 composite samples was used for spore count determination and a similar amount for next generation sequencing. Spore counts varied significantly among soil textural types, sample locations, and soil depth. Sandy loam had the highest spore counts with a mean average of 404 spores. The spore count decreased significantly (P <.05) with the depth of soil from a mean of 514 spores to 185 along the rhizosphere. The intensity of spore morphotypes was significantly higher at P <.05 for soils whose vegetation was covered with Aspilia pruliseta than those without. Aspilia pruliseta vegetation used together with sandy loam soil could culture commercial mycorrhiza fungi production for use in agrisystems. [ABSTRACT FROM AUTHOR]

Phosphate (P) is characterized by its low availability and restricted mobility in soils, and also by a high redistribution capacity inside plants. In order to maintain P homeostasis in nutrient restricted conditions, plants have developed mechanisms which enable P acquisition from the soil solution, and an efficient reutilization of P already present in plant cells. Nitric oxide (NO) is a bioactive molecule with a plethora of functions in plants. Its endogenous synthesis depends on internal and environmental factors, and is closely tied with nitrogen (N) metabolism. Furthermore, there is evidence demonstrating that N supply affects P homeostasis and that P deficiency impacts on N assimilation. This review will provide an overview on how NO levels in planta are affected by P deficiency, the interrelationship with N metabolism, and a summary of the current understanding about the influence of this reactive N species over the processes triggered by P starvation, which could modify P use efficiency. [ABSTRACT FROM AUTHOR]

Background: To improve plant phosphorus (P)-acquisition efficiency to secure sustainable food production, an important step is to increase the concentration of plant-available P in the rhizosphere. Root exudation of organic anions is a key strategy in mobilizing less-available soil P. Scope: This review covers how organic anions (carboxylates) mobilize soil P and research methodologies applied. It then discusses the root-release of organic anions induced by low P availability and their contribution to soil P mobilization and plant P acquisition, and highlights the impact, challenges and perspectives in this research area. Conclusions: The release of organic anions is increased considerably in some plant species, but very little in others under low P availability. Rhizosphere organic anions play important roles in increasing plant-available P, but the contribution is greatly affected by many factors. In future research, improved and ecologically meaningful root exudation sampling methods, the use of mature leaf manganese (Mn) concentration or total 14C exudation as a proxy for rhizosphere carboxylates, case-by-case field experiments, molecular mechanisms underpinning organic anion biosynthesis and efflux under low P availability warrant further attention. Finally, carbon costs and multiple root trait combinations (e.g., root hairs plus root exudation) should be considered in crop breeding programs to generate more P-efficient cultivars. [ABSTRACT FROM AUTHOR]

It was previously shown that plasma membrane (PM) H+‐ATPases of active cluster roots are acclimated to phosphorus (P) deficiency. In the present study, we followed the question whether the qualitative acclimation of PM H+‐ATPase is based on isoform‐specific expression. Therefore, three different PM H+‐ATPase isoforms (LHA1–LHA3) were investigated at the transcriptional level. Our data indicate that LHA1 may play a key role in the qualitative acclimation of PM H+‐ATPase in cluster roots to P deficiency. [ABSTRACT FROM AUTHOR]

Background and aims: Twenty-four species of eucalypts were studied regarding their ability to grow under low P and their responsiveness to P inputs. Methods: Growth and photosynthesis-related parameters were evaluated. Results: Growth of all species was influenced by low P availability. No significant correlation was found between leaf P concentration and biomass, indicating that P concentrations in leaves cannot be solely considered an indication of the responsiveness to P in eucalypts. Species responsive to P-input (high agronomic P efficiency values, APE) were those with low P use efficiency - PUE (here assessed as relative efficiency of P-use, REP) and low P uptake efficiency (PUpE). But, non-responsive species were related to higher P-efficiency under low soil P-availability. Eucalyptus tereticornis, E. cladocalyx, E. globulus and E. camaldulensis were efficient under low-P availability. Whereas, E. crebra and E acmenoides were the most responsive species, with high APE, suggesting that for these species P-inputs are needed to guarantee plant growth. The root:shoot ratio remained constant at different P availabilities, suggesting that biomass allocation towards the root in response to P and greater investment in roots were not correlated with greater PUE. Under limited P, E. robusta and E. botryoides exhibited low foliar P contents and higher root:shoot ratios than those of other species with higher P contents, indicating that greater root investment does not necessarily result in greater PUE. Conclusion: The results suggest that the divergence among species is probably related to different mechanisms, which may improve P-use efficiency. [ABSTRACT FROM AUTHOR]

Background and aims: Phosphorus deficiency often limits crop productivity, while phosphate rock, which is used to produce phosphorus fertilisers, is a non-renewable resource. Formation of cluster roots is an adaptation to nutrient-poor soils in Lupinus species, including L. albus. Lupinus species also produce nodules, which require a large investment of phosphorus. Our aim is to test whether nodulation promotes cluster-root formation in L. albus. Methods: Seedlings of L. albus, either inoculated with rhizobia or non-inoculated, were grown in nutrient solution with a low phosphorus supply. Non-inoculated plants were provided with the same amount of nitrogen in the form of nitrate as the nodulated ones acquired, from both air and nutrient solution, based on preliminary experiments. We measured biomass, phosphorus and nitrogen concentrations as well as photosynthesis just prior to each harvest. Results: Nodulated plants and non-nodulated control plants produced the same amount of biomass. Nodulated plants had, on average, 86% more cluster roots than non-nodulated ones at the four harvests. As hypothesised, nodulation significantly promoted cluster-root formation; it also enhanced rates of photosynthesis. Conclusions: Nodulation promoted cluster-root formation and photosynthesis, presumably because nodules are significant sinks for phosphorus and photosynthates. Our results do not provide evidence for a trade-off between investment of resources in nodules and cluster roots. [ABSTRACT FROM AUTHOR]

White lupin cluster roots are specialized brush‐like root structures that are formed in some species under phosphorus (P)‐deficient conditions. They intensely secrete protons and organic acid anions for solubilization and acquisition of sparingly soluble phosphates. Phytohormones and sucrose modulate cluster root number, but the molecular mechanisms of cluster root formation have been elusive. Here, a novel peptide phytohormone was identified that affects cluster root development. It belongs to the C‐TERMINALLY-ENCODED PEPTIDE (CEP) family. Members of that family arrest root growth and modulate branching in model species. LaCEP1 was highly expressed in the pre‐emergence zone of clusters. Over‐expression of the gene encoding the LaCEP1 propeptide resulted in moderate inhibition of cluster root formation. The primary and lateral root lengths of lupin were little affected by the overexpression, but LaCEP1 reduced cluster rootlet and root hair elongation. Addition of a 15‐mer core peptide derived from LaCEP1 similarly altered root morphology and modified cluster activity, suggesting that a core sequence of the propeptide is functionally sufficient. Stable overexpression in Arabidopsis confirmed the LaCEP1 function in root growth inhibition across species. Taken together, the root inhibitory effects of the LaCEP1 phytohormone suggest a role as of a regulatory module involved in cluster root development in white lupin. [ABSTRACT FROM AUTHOR]

Key message: Hexaploid wheat is more responsive than tetraploid to the interactive effects of elevated [CO2] and low P in terms of carboxylate efflux, enzyme activity and gene expression (TaPT1 and TaPAP).Abstract: Availability of mineral nutrients to plants under changing climate has become a serious challenge to food security and economic development. An understanding of how elevated [CO2] influences phosphorus (P) acquisition processes at the whole-plant level would be critical in selecting cultivars as well as to maintain optimum yield in limited-P conditions. Wheat (Triticum aestivum and T. durum) grown hydroponically with sufficient and low P concentration were exposed to elevated and ambient [CO2]. Improved dry matter partitioning towards root resulted in increased root-to-shoot ratio, root length, volume, surface area, root hair length and density at elevated [CO2] with low P. Interaction of low P and [CO2] induced activity of enzymes (phosphoenolpyruvate carboxylase, malate dehydrogenase and citrate synthase) in root tissue resulting in twofold increase in carboxylates and acid phosphatase exudation. Physiological absorption capacity of roots showed that plants alter their uptake kinetics by increasing affinity (low Km) in response to elevated [CO2] under low P supply. Increased relative expression of genes, purple acid phosphatase (TaPAP) and high-affinity Pi transporter (TaPT1) in roots induced by elevated [CO2] and low P supported our physiological observations. Hexaploid wheat (PBW-396) being more responsive to elevated [CO2] at low P supply as compared to tetraploid (PDW-233) necessitates the ploidy effect to be explored further which might be advantageous under changing climate. [ABSTRACT FROM AUTHOR]

Roots and root-released organic anions play important roles in uptake of phosphorus (P), an essential macronutrient for food production. Oat, ranking sixth in the world's cereal production, contains valuable nutritional compounds and can withstand poor soil conditions. Our aim was to investigate root transcriptional and metabolic responses of oat grown under P-deficient and P-sufficient conditions. We conducted a hydroponic experiment and measured root morphology and organic anion exudation, and analysed changes in the transcriptome and metabolome. Oat roots showed enhanced citrate and malate exudation after 4 weeks of P deficiency. After 10 d of P deficiency, we identified 9371 differentially expressed transcripts with a 2-fold or greater change (P<0.05): 48 sequences predicted to be involved in organic anion biosynthesis and efflux were consistently up-regulated; 24 up-regulated transcripts in oat were also found to be up-regulated upon P starvation in rice and wheat under similar conditions. Phosphorylated metabolites (i.e. glucose-6-phosphate, myo-inositol phosphate) were reduced dramatically, while citrate and malate, some sugars and amino acids increased slightly in P-deficient oat roots. Our data are consistent with a strategy of increased organic anion efflux and a shift in primary metabolism in response to P deficiency in oat. [ABSTRACT FROM AUTHOR]

A potential mechanism to enhance utilization of sparingly soluble forms of phosphorus (P) is the root secretion of malate, which is mainly mediated by the ALMT gene family in plants. In this study, a total of 34 GmALMT genes were identified in the soybean genome. Expression patterns diverged considerably among GmALMTs in response to phosphate (Pi) starvation in leaves, roots and flowers, with expression altered by P availability in 26 of the 34 GmALMTs. One root-specific GmALMT whose expression was significantly enhanced by Pi-starvation, GmALMT5, was studied in more detail to determine its possible role in soybean P nutrition. Analysis of GmALMT5 tissue expression patterns, subcellular localization, and malate exudation from transgenic soybean hairy roots overexpressing GmALMT5, demonstrated that GmALMT5 is a plasma membrane protein that mediates malate efflux from roots. Furthermore, both growth and P content of transgenic Arabidopsis overexpressing GmALMT5 were significantly increased when sparingly soluble Ca-P was used as the external P source. Taken together, these results indicate that members of the soybean GmALMT gene family exhibit diverse responses to Pi starvation. One member of this family, GmALMT5, might contribute to soybean P efficiency by enhancing utilization of sparingly soluble P sources under P limited conditions. [ABSTRACT FROM AUTHOR]

Root structures secreting carboxylates and phosphatases are thought to enhance a plant's phosphorus (P) acquisition. But do closely related species with and without such structures really differ in root exudation, P mobilization, or ecological niche? We investigated this by comparing 23 European Carex species with and without 'dauciform roots' (DRs)., Plants grown in pots with sand were screened for DR formation, phosphatase activities, carboxylate exudation, and utilization of various organic and inorganic P compounds. Ecological niches were compared using ecological indicator values and nutrient concentrations of plant shoots in natural habitats., Species of subgenus Carex formed DRs, while species of subgenus Vignea did not. Species with DRs had higher root diesterase activity than species without DRs, exuded more citrate but less oxalate and less total carboxylates, and allocated less biomass to roots. Species with and without DRs showed similar growth responses to different forms of P and different amounts of P supplied; their natural habitats do not differ in soil fertility or degree of P limitation., Despite some differences in physiological function, DRs did not influence the P acquisition and nutritional niche of European Carex species, suggesting that species with and without DRs do not exhibit distinct P-acquisition strategies. [ABSTRACT FROM AUTHOR]

Phosphorus (P) is one of the essential nutrients for plants, and is indispensable for plant growth and development. P deficiency severely limits crop yield, and regular fertilizer applications are required to obtain high yields and to prevent soil degradation. To access P from the soil, plants have evolved high- and low-affinity Pi transporters and the ability to induce root architectural changes to forage P. Also, adjustments of numerous cellular processes are triggered by the P starvation response, a tightly regulated process in plants. With the increasing demand for food as a result of a growing population, the demand for P fertilizer is steadily increasing. Given the high costs of fertilizers and in light of the fact that phosphate rock, the source of P fertilizer, is a finite natural resource, there is a need to enhance P fertilizer use efficiency in agricultural systems and to develop plants with enhanced Pi uptake and internal P-use efficiency ( PUE). In this review we will provide an overview of continuing relevant research and highlight different approaches towards developing crops with enhanced PUE. In this context, we will summarize our current understanding of root responses to low phosphorus conditions and will emphasize the importance of combining PUE with tolerance of other stresses, such as aluminum toxicity. Of the many genes associated with Pi deficiency, this review will focus on those that hold promise or are already at an advanced stage of testing ( Os PSTOL1, AVP1, PHO1 and Os PHT1;6). Finally, an update is provided on the progress made exploring alternative technologies, such as phosphite fertilizer. [ABSTRACT FROM AUTHOR]

Phosphorus (P) is an important mineral nutrient for plant growth and development. Overexpressing AtWRKY6 (35S:WRKY6-9) was more sensitive and wrky6 (wrky6-1) was more resistant under low Pi conditions. To better understand the function of AtWRKY6 under low phosphate stress conditions, we applied two-dimensional gel electrophoresis (2-DE) to analyse differentially expressed proteins in the shoots and roots between wild type, 35S:WRKY6-9 and wrky6-1 after phosphorus deficiency treatment for three days. The results showed 88 differentially abundant protein spots, which were identified between the shoots and roots of 35S:WRKY6-9 and wrky6-1 plants. In addition, 59 differentially expressed proteins were identified in the leaves and roots of 35S:WRKY6-9 plants. After analysis, 9 genes with W-box elements in their promoter sequences were identified in the leaves, while 6 genes with W-box elements in their promoter sequences were identified in the roots. A total of 8 genes were identified as potential target genes according to the quantitative PCR (QPCR) and two dimension difference gel electrophoresis, (2D-DIGE) results, including ATP synthase, gln synthetase, nitrilase, 14-3-3 protein, carbonic anhydrases 2, and tryptophan synthase. These results provide important information concerning the AtWRKY6 regulation network and reveal potential vital target genes of AtWRKY6 under low phosphorus stress. two dimension difference gel electrophoresis, 2D-DIGE. [ABSTRACT FROM AUTHOR]

Eutrema salsugineum, a halophytic relative of Arabidopsis thaliana, was subjected to varying phosphate (Pi) treatments. Arabidopsis seedlings grown on 0.05 m m Pi displayed shortened primary roots, higher lateral root density and reduced shoot biomass allocation relative to those on 0.5 m m Pi, whereas Eutrema seedlings showed no difference in lateral root density and shoot biomass allocation. While a low Fe concentration mitigated the Pi deficiency response for Arabidopsis, Eutrema root architecture was unaltered, but adding NaCl increased Eutrema lateral root density almost 2-fold. Eutrema and Arabidopsis plants grown on soil without added Pi for 4 weeks had low shoot and root Pi content. Pi-deprived, soil-grown Arabidopsis plants were stunted with senescing older leaves, whereas Eutrema plants were visually indistinguishable from 2.5 m m Pi-supplemented plants. Genes associated with Pi starvation were analysed by RT-qPCR. EsIPS2, EsPHT1; 4 and EsPAP17 showed up-regulated expression in Pi-deprived Eutrema plants, while EsPHR1, EsWRKY75 and EsRNS1 showed no induction. Absolute quantification of transcripts indicated that PHR1, WRKY75 and RNS1 were expressed at higher levels in Eutrema plants relative to those in Arabidopsis regardless of external Pi. The low phenotypic plasticity Eutrema displays to Pi supply is consistent with adaptation to chronic Pi deprivation in its extreme natural habitat. [ABSTRACT FROM AUTHOR]

Phosphorus (P) is a limiting factor for plant growth, especially in highly weathered tropical soils. Plants have several mechanisms to overcome low P availability in soil, such as humic substances, that reduce phosphate (Pi) adsorption on oxide surfaces and enhance soil P availability. However, the direct influence of humic substances on Pi transporters in root cells or the distribution of P species in leaves remains unclear. Tomato seedlings were grown in a sand-vermiculite mixture with low or high P concentrations (10 or 100 mg kg-1 KH2PO4, respectively) and humic acids (0 or 48 mg C L-1) isolated from vermicompost. Plant responses were evaluated in the fifth week by measuring root and shoot weights and P concentration, and differential expression in the roots of the high-affinity Pi transporter genes LePT1 and LePT2. In addition, the distribution of P species in the leaves was assessed using 31P-NMR. Humic acids increased the root biomass and changed the distribution of P species in the leaves. Inorganic phosphate was the major compound in plants supplied with a high P concentration, whereas in plants supplied with a low P concentration, Pi was only identified in plants not treated with humic acids. Glycerophosphodiester and phosphorylcholine accumulated in plants treated with humic acid, indicating a modified metabolic pathway for economical P consumption at low P concentrations. High transcript accumulation of LePT2 was observed in roots treated with humic acids at both P concentrations. Our results show that humic substances are strategically involved in plant adaptation to P availability. [ABSTRACT FROM AUTHOR]

Improving crop nutrient efficiency becomes an essential consideration for environmentally friendly and sustainable agriculture. Plant growth and development is dependent on 17 essential nutrient elements, among them, nitrogen (N) and phosphorus (P) are the two most important mineral nutrients. Hence it is not surprising that low N and/or low P availability in soils severely constrains crop growth and productivity, and thereby have become high priority targets for improving nutrient efficiency in crops. Root exploration largely determines the ability of plants to acquire mineral nutrients from soils. Therefore, root architecture, the 3-dimensional configuration of the plant's root system in the soil, is of great importance for improving crop nutrient efficiency. Furthermore, the symbiotic associations between host plants and arbuscular mycorrhiza fungi/rhizobial bacteria, are additional important strategies to enhance nutrient acquisition. In this review, we summarize the recent advances in the current understanding of crop species control of root architecture alterations in response to nutrient availability and root/microbe symbioses, through gene or QTL regulation, which results in enhanced nutrient acquisition. [ABSTRACT FROM AUTHOR]

Background and aims: The aims were to investigate effects of availability of nitrogen (N), phosphorus (P) and iron (Fe) on root properties of seabuckthorn ( Hippophaë rhamnoides) and to test the hypothesis that seabuckthorn is able to form cluster roots (CRs). Methods: Two sources of seabuckthorn were used: the seabuckthorn cultivar BHi10726 originating from a breeding programme based on H.r. ssp mongolica and carried out in rich agricultural field soil in the black earth (chernozem) region of Russia and the seabuckthorn accession named Pk originating in a natural population of H.r. ssp turkestanica in the mountainous region of northern Pakistan. Three cultivation systems giving different water availabilities were used at two levels each of N, P and Fe. Root morphology of seedlings and clones was characterized and metabolite content in extracts of young and old CRs of Pk was analyzed by proton nuclear magnetic resonance spectroscopy. Results: Availability of N affected growth and distribution of biomass between shoot and root, while P and Fe deficiency modified root system architecture towards more lateral roots. Densely positioned rootlets with a determinate type of growth consistent with the definition of CR were observed under low P and low Fe. Pk formed on average 12 CRs per plant, which was 3 to 4-fold higher compared to BHi10726 also when normalized per root length. Malate and glycine were most abundant of the organic acids and amino acids, respectively, and decreased in old CRs. Conclusions: Seabuckthorn has the ability to form cluster roots especially in Pk and under deficiency of P and Fe. The two sources of seabuckthorn with different histories showed distinctly different root system architectures. The high contents of malate and glycine and their decrease in old CRs may reflect roles in CR metabolism. [ABSTRACT FROM AUTHOR]

Apart from substrate functions, a signaling role of sucrose in root growth regulation is well established. This raised the question whether sucrose signals might also be involved in formation of cluster-roots (CRs) under phosphate (Pi) limitation, mediating exudation of phosphorus (P)-mobilizing root exudates, e.g. in Lupinus albus and members of the Proteaceae. Earlier studies demonstrated that CR formation in L. albus was mimicked to some extent by external application of high sucrose concentrations (25 mM) in the presence of extremely high P supply (1-10 mM), usually suppressing CR formation. In this study, we re-addressed this question using an axenic hydroponic culture system with normal P supply (0.1 mM) and a range of sucrose applications (0.25-25 mM). The 2.5 mM sucrose concentration was comparable with internal sucrose levels in the zone of CR initiation in first-order laterals of P-deficient plants (3.4 mM) and induced the same CR morphology. Similar to earlier studies, high sucrose concentrations (25 mM) resulted in root thickening and inhibition of root elongation, associated with a 10-fold increase of the internal sucrose level. The sucrose analog palatinose and a combination of glucose/fructose failed to stimulate CR formation under P-sufficient conditions, demonstrating a signal function of sucrose and excluding osmotic or carbon source effects. In contrast to earlier findings, sucrose was able to induce CR formation but had no effect on CR functioning with respect to citrate exudation, in vitro activity and expression of genes encoding phosphoenolpyruvate carboxylase, secretory acid phosphatase and MATE transporters, mediating P-mobilizing functions of CRs. [ABSTRACT FROM AUTHOR]

Plants have evolved a plethora of mechanisms to circumvent the potential damaging effects of living under low phosphorus availability in the soil. These mechanisms include different levels of organization, from root-shoot signalling at the whole-plant level to specific biochemical responses at the subcellular level, such as reductions in photosynthesis and the consequent activation of photo- and antioxidant mechanisms in chloroplasts. Some recent studies clearly indicate that severe phosphorus deficiency can lead to alterations in the photosynthetic apparatus, including reductions in CO2 assimilation rates, a down-regulation of photosynthesis-related genes and photoinhibition at the photosystem II level, thus causing potential photo-oxidative stress. Photo-oxidative stress is characterized by an increased production of reactive oxygen species in chloroplasts, which at low concentrations can serve a signalling, protective role, but when present at high concentrations can cause damage to lipids, proteins and nucleic acids, thus leading to irreversible injuries. We discuss here the mechanisms that phosphate-starved plants have evolved to withstand photo-oxidative stress, including changes at the subcellular level (e.g. activation of photo- and antioxidant protection mechanisms in chloroplasts), cellular and tissular levels (e.g. activation of photorespiration and anthocyanin accumulation) and whole-plant level (alterations in source-sink relationships modulated by hormones). Of particular importance is the current evidence demonstrating that phosphate-starved plants activate simultaneous responses at multiple levels, from transcriptional changes to root-shoot signalling, to prevent oxidative damage. In this review, we summarize current knowledge about the occurrence of photo-oxidative stress in phosphate-starved plants and highlight the mechanisms these plants have evolved to prevent oxidative damage under phosphorus limitation at the subcellular, cellular and whole-plant levels. [ABSTRACT FROM AUTHOR]

Phosphorous (P) is one of the most deficient nutrients owing to its fixation and low mobility in acid soils. A total of 324 genotypes were screened in both P sufficient and insufficient conditions to evaluate the P uptake efficiency. Thirty-four responsive genotypes were evaluated in solution culture with two checks, ICGV 86590 and JL 24. The components of variability revealed high magnitude and highly heritable nature of the variation. Highly responsive genotypes to P insufficiency; ICG 14475, ICGV 92188, and M 68 were further tested for phosphorus uptake efficiency at 15, 30, and 45 days after sowing (DAS) along with checks in sand culture in pots. ICG 14475, ICGV 92188, and ICGV 86590 were found superior in both acquisition and utilization of phosphorus due to enhanced root production for P exploration and shoot expansion, respectively. JL 24 was poor in acquisition due to lack of response in root traits under P insufficiency. [ABSTRACT FROM PUBLISHER]

White lupin ( L upinus albus) produces cluster roots, an adaptation to low soil phosphorus ( P). Cluster roots exude large levels of P-solubilizing compounds such as citrate and malate. In contrast, narrow leaf lupin ( L . angustifolius) is closely related to L . albus, but does not produce cluster roots. To examine the different strategies for P acquisition, we compared the growth, biomass allocation, respiratory properties and construction cost between L . albus and L . angustifolius under P-deficient conditions. Both L upinus species were grown in hydroponic culture with 1 or 100 μ M P. Under the P-deficient regime, L . albus produced cluster roots with little change in biomass allocation, while L . angustifolius significantly increased biomass allocation to roots. The rate of cyanide-resistant SHAM (salicylhydroxamic acid)-sensitive respiration was high in cluster roots and very low in roots of L . angustifolius. These results suggest a low alternative oxidase ( AOX) activity in L . angustifolius roots, and thus, ATP would be produced efficiently in L . angustifolius roots. The construction cost was highest in cluster roots and lowest in L . angustifolius roots. This study shows that under P deficiency, L . albus produces high-cost cluster roots to increase the P availability, while L . angustifolius produces large quantities of low-cost roots to enhance P uptake. [ABSTRACT FROM AUTHOR]

Background: Phosphorus (P) is an essential macronutrient for plant growth and development. Several genes involved in phosphorus deficiency stress have been identified in various plant species. However, a whole genome understanding of the molecular mechanisms involved in plant adaptations to low P remains elusive, and there is particularly little information on the genetic basis of these acclimations in coniferous trees. Masson pine (Pinus massoniana) is grown mainly in the tropical and subtropical regions in China, many of which are severely lacking in inorganic phosphate (Pi). In previous work, we described an elite P. massoniana genotype demonstrating a high tolerance to Pi-deficiency. Methodology/Principal Findings: To further investigate the mechanism of tolerance to low P, RNA-seq was performed to give an idea of extent of expression from the two mixed libraries, and microarray whose probes were designed based on the unigenes obtained from RNA-seq was used to elucidate the global gene expression profiles for the long-term phosphorus starvation. A total of 70,896 unigenes with lengths ranging from 201 to 20,490 bp were assembled from 112,108,862 high quality reads derived from RNA-Seq libraries. We identified 1,396 and 943 transcripts that were differentially regulated (P<0.05) under P1 (0.01 mM P) and P2 (0.06 mM P) Pi-deficiency conditions, respectively. Numerous transcripts were consistently differentially regulated under Pi deficiency stress, many of which were also up- or down-regulated in other species under the corresponding conditions, and are therefore ideal candidates for monitoring the P status of plants. The results also demonstrated the impact of different Pi starvation levels on global gene expression in Masson pine. Conclusions/Significance: To our knowledge, this work provides the first insight into the molecular mechanisms involved in acclimation to long-term Pi starvation and different Pi availability levels in coniferous trees. [ABSTRACT FROM AUTHOR]

Phosphorus is an essential nutrient that is required for all major developmental processes and reproduction in plants. It is also a major constituent of the fertilizers required to sustain high-yield agriculture. Levels of phosphate-the only form of phosphorus that can be assimilated by plants-are suboptimal in most natural and agricultural ecosystems, and when phosphate is applied as fertilizer in soils, it is rapidly immobilized owing to fixation and microbial activity. Thus, cultivated plants use only approximately 20-30% of the applied phosphate, and the rest is lost, eventually causing water eutrophication. Recent advances in the understanding of mechanisms by which wild and cultivated species adapt to low-phosphate stress and the implementation of alternative bacterial pathways for phosphorus metabolism have started to allow the design of more effective breeding and genetic engineering strategies to produce highly phosphate-efficient crops, optimize fertilizer use, and reach agricultural sustainability with a lower environmental cost. In this review, we outline the current advances in research on the complex network of plant responses to low-phosphorus stress and discuss some strategies used to manipulate genes involved in phosphate uptake, remobilization, and metabolism to develop low-phosphate-tolerant crops, which could help in designing more efficient crops. [ABSTRACT FROM AUTHOR]

Phosphate (Pi) is an essential nutrient for plants but is normally fixed in soil, which limits plant growth and reproduction. In response to low availability of Pi, shoots and roots react differently but cooperatively to improve Pi acquisition from the rhizosphere and adjust Pi distribution and metabolism within plants. Shoot and root responses are coordinated by the trafficking of various kinds of systemic signals through the vasculature. Mutual communication between different tissues is necessary to integrate the environmental stimuli with the internal cues at the whole-plant level. Different approaches have been used to monitor or manipulate components in the vascular stream to reveal several candidates of systemic signals from roots or shoots, including photosynthates, phytohormones, microRNAs, and Pi. In addition, the downstream signalling pathways mediated by these signals have been discovered. The crosstalk among different signalling pathways has been revealed, showing the complexity of the Pi signalling network. In this review, we summarize the approaches used for studying systemic signalling and discuss recent progress and challenges in investigating the systemic signalling pathway that integrates Pi starvation responses to maintain Pi at physiological concentrations. Knowledge gained from this study may help improve the phosphorus use efficiency of crops. [ABSTRACT FROM AUTHOR]

Plants with tolerance to low-phosphorus (P) can grow better under low-P conditions, and understanding of genetic mechanisms of low-P tolerance can not only facilitate identifying relevant genes but also help to develop low-P tolerant cultivars. QTL meta-analysis was conducted after a comprehensive review of the reports on QTL mapping for low-P tolerance-related traits in maize. Meta-analysis produced 23 consensus QTL (cQTL), 17 of which located in similar chromosome regions to those previously reported to influence root traits. Meanwhile, candidate gene mining yielded 215 genes, 22 of which located in the cQTL regions. These 22 genes are homologous to 14 functionally characterized genes that were found to participate in plant low-P tolerance, including genes encoding miR399s, Pi transporters and purple acid phosphatases. Four cQTL loci (cQTL2-1, cQTL5-3, cQTL6-2, and cQTL10-2) may play important roles for low-P tolerance because each contains more original QTL and has better consistency across previous reports. [ABSTRACT FROM AUTHOR]

As an essential plant macronutrient, the low availability of phosphorus (P) in most soils imposes serious limitation on crop production. Plants have evolved complex responsive and adaptive mechanisms for acquisition, remobilization and recycling of phosphate (Pi) to maintain P homeostasis. Spatio-temporal molecular, physiological, and biochemical Pi deficiency responses developed by plants are the consequence of local and systemic sensing and signaling pathways. Pi deficiency is sensed locally by the root system where hormones serve as important signaling components in terms of developmental reprogramming, leading to changes in root system architecture. Root-to-shoot and shoot-to-root signals, delivered through the xylem and phloem, respectively, involving Pi itself, hormones, miRNAs, mRNAs, and sucrose, serve to coordinate Pi deficiency responses at the whole-plant level. A combination of chromatin remodeling, transcriptional and posttranslational events contribute to globally regulating a wide range of Pi deficiency responses. In this review, recent advances are evaluated in terms of progress toward developing a comprehensive understanding of the molecular events underlying control over P homeostasis. Application of this knowledge, in terms of developing crop plants having enhanced attributes for P use efficiency, is discussed from the perspective of agricultural sustainability in the face of diminishing global P supplies. [ABSTRACT FROM AUTHOR]

The aim of this study was to develop techniques to evaluate soil phosphorus (P) sorption capacity (PSC) and determine critical soil P levels to predict P loss potential for calcareous soils. Seventy-five soils mostly from Northern China were analyzed for soil P using four extraction methods (water, P; carbonate, P; ammonium oxalate, P; and Mehlich 3, P) as well as PSC derived from single-point (PSC) and multipoint sorption ( S) isotherms. Strong correlation was found between PSC and S ( r=0.89, p<0.001). The sum of αCa and βMg as an index of PSC (PSC) was most closely related to the maximum amount of P sorbed ( S) as given by the sum of S and soil initial P setting α=0.039 and β=0.462 ( r=0.80, p<0.001). The degree of P saturation (DPS) was thereafter calculated from PSC (DPS), to which Olsen P (P) was significantly correlated ( r=0.82, p<0.001). In a split-line regression from P against DPS ( r=0.87, p<0.05), a change point was identified at 28.1% DPS, which was equivalent to 49.2 mg kg P and corresponded to a P concentration of 8.8 mg kg. After the change point, a sharp increase in P was observed. Our results reveal a new approach to approximating DPS from Ca and Mg for calcareous soils without the need to generate a S. We conclude that in the absence of an environmental soil test criteria for P, the DPS and P could be used to predict P loss potential from calcareous soils. [ABSTRACT FROM AUTHOR]

Background and Aims Some Lupinus species produce cluster roots in response to low plant phosphorus (P) status. The cause of variation in cluster-root formation among cluster-root-forming Lupinus species is unknown. The aim of this study was to investigate if cluster-root formation is, in part, dependent on different relative growth rates (RGRs) among Lupinus species when they show similar shoot P status. Methods Three cluster-root-forming Lupinus species, L. albus, L. pilosus and L. atlanticus, were grown in washed river sand at 0, 7·5, 15 or 40 mg P kg−1 dry sand. Plants were harvested at 34, 42 or 62 d after sowing, and fresh and dry weight of leaves, stems, cluster roots and non-cluster roots of different ages were measured. The percentage of cluster roots, tissue P concentrations, root exudates and plant RGR were determined. Key Results Phosphorus treatments had major effects on cluster-root allocation, with a significant but incomplete suppression in L. albus and L. pilosus when P supply exceeded 15 mg P kg−1 sand. Complete suppression was found in L. atlanticus at the highest P supply; this species never invested more than 20 % of its root weight in cluster roots. For L. pilosus and L. atlanticus, cluster-root formation was decreased at high internal P concentration, irrespective of RGR. For L. albus, there was a trend in the same direction, but this was not significant. Conclusions Cluster-root formation in all three Lupinus species was suppressed at high leaf P concentration, irrespective of RGR. Variation in cluster-root formation among the three species cannot be explained by species-specific variation in RGR or leaf P concentration. [ABSTRACT FROM PUBLISHER]

Formation of cluster roots is a typical morphological response to phosphorus (P) deficiency in white lupin ( Lupinus albus), but its physiological and molecular mechanisms are still unclear. We investigated the role of auxin in the initiation of cluster roots by distinguishing the sources of auxin, measuring the longitudinal distribution patterns of free indole-3-acetic acid ( IAA) along the root and the related gene expressions responsible for polar auxin transport ( PAT) in different developmental stages of cluster roots. We found that removal of shoot apex or primary root apex and application of auxin-influx or -efflux transport inhibitors, 3-chloro-4-hydroxyphenylacetic acid, N-1-naphthylphthalamic acid and 2,3,5-triiodobenzoic acid, to the stem did not affect the number of cluster roots and the free- IAA concentration in the roots of P-deficient plants, but when these inhibitors were applied directly to the growth media, the cluster-root formation was greatly suppressed, suggesting the fundamental role of root-derived IAA in cluster-root formation. The concentration of free IAA in the roots was higher in P-deficient plants than in P-adequate ones, and the highest in the lateral-root apex and the lowest in the mature cluster roots. Meanwhile the expression patterns of LaAUX1, LaPIN1 and LaPIN3 transcripts related to PAT was consistent with concentrations of free IAA along the lateral root, indicating the contribution of IAA redistribution in the cluster-root development. We proposed that root-derived IAA plays a direct and important role in the P-deficiency-induced formation of cluster roots. [ABSTRACT FROM AUTHOR]