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SUMMARY: Plants cope with low phosphorus availability by adjusting growth and metabolism through transcriptomic and proteomic adaptations. We hypothesize that selected genotypes with distinct phosphorous (P) use efficiency covering the breeding history of European Flint heterotic pool provide a tool to reveal general and genotype‐specific molecular responses to P limitation. We reconstructed protein and gene co‐expression networks by weighted correlation network analysis and related these to phosphate deficiency‐induced traits. In roots, low phosphate supply resulted in a decreasing abundance of proteins in the oxidative pentose phosphate pathway and a negative correlation with root and shoot phosphate content. We observed an increase in abundance and positive correlation with root and shoot phosphate content for proteins in sucrose biosynthesis, lipid metabolism, respiration and RNA processing. Purple acid phosphatases, superoxide dismutase and phenylalanine ammonia lyase were identified as being upregulated under low phosphate in all genotypes. Overall, correlations between protein and mRNA abundance changes were limited, with ribosomal proteins and the ubiquitin protein degradation pathway exclusively responding with protein abundance changes. Carbohydrate, phospho‐ and sulfo‐lipid metabolism showed abundance changes at the protein and mRNA levels. These partially non‐overlapping proteomic and transcriptomic adjustments to low phosphate suggest sugar and lipid metabolism as metabolic processes associated with improved P use efficiency specifically in Founder Flint lines. We identified a mitogen‐activated protein kinase‐kinase as a potential genotype‐specific regulator of sucrose metabolism at low phosphate in Founder Flint line EP1. We conclude that, during breedingt of Elite Flint lines, regulation of primary metabolism has changed to result in a distinct low phosphate response in Founder lines. Significance Statement: We reconstructed protein and transcript co‐expression networks in Flint maize lines and related them to phosphate deficiency‐induced traits. Post‐translational regulation was found with protein synthesis and protein degradation, transcriptional and post‐transcriptional regulation was found with metabolic processes. MKK6 homolog was proposed as a genotype‐specific regulator for carbohydrate metabolism and protein synthesis processes. [ABSTRACT FROM AUTHOR]

Summary: Phosphorus (P) is an essential element for plant growth and development. Vacuoles play a fundamental role in the storage and remobilization of P in plants, while our understanding of the evolutionary mechanisms of creating and reusing P stores are limited. Besides, we also know very little about the coordination of intercellular P translocation, neither the inorganic phosphate (Pi) signaling nor the Pi transport patterns. Here we summarize recent advances in understanding the core elements involved in cellular and/or subcellular P homeostasis and signaling in unicellular green algae and multicellular land plants. We also propose further work that might help to uncover the high‐resolution intracellular and intercellular landscape of Pi distribution and signaling in plants. [ABSTRACT FROM AUTHOR]

Phosphorus (P) assimilation and polyphosphate (polyP) synthesis were investigated in Chlamydomonas reinhardtii by supplying phosphate (PO43−; 10 mg P·L−1) to P‐depleted cultures of wildtypes, mutants with defects in genes involved in the vacuolar transporter chaperone (VTC) complex, and VTC‐complemented strains. Wildtype C. reinhardtii assimilated PO43− and stored polyP within minutes of adding PO43− to cultures that were P‐deprived, demonstrating that these cells were metabolically primed to assimilate and store PO43−. In contrast, vtc1 and vtc4 mutant lines assayed under the same conditions never accumulated polyP, and PO43− assimilation was considerably decreased in comparison with the wildtypes. In addition, to confirm the bioinformatics inferences and previous experimental work that the VTC complex of C. reinhardtii has a polyP polymerase function, these results evidence the influence of polyP synthesis on PO43− assimilation in C. reinhardtii. RNA‐sequencing was carried out on C. reinhardtii cells that were either P‐depleted (control) or supplied with PO43− following P depletion (treatment) in order to identify changes in the levels of mRNAs correlated with the P status of the cells. This analysis showed that the levels of VTC1 and VTC4 transcripts were strongly reduced at 5 and 24 h after the addition of PO43− to the cells, although polyP granules were continuously synthesized during this 24 h period. These results suggest that the VTC complex remains active for at least 24 h after supplying the cells with PO43−. Further bioassays and sequence analyses suggest that inositol phosphates may control polyP synthesis via binding to the VTC SPX domain. [ABSTRACT FROM AUTHOR]

The different cortical activity evoked by a mechanical tactile stimulus depends on tactile stimulus patterns, which demonstrates that simple stimuli (i.e., global synchronous stimulation the stimulus area) activate the primary somatosensory cortex alone, whereas complex stimuli (i.e., stimulation while moving in the stimulus area) activate not only the primary somatosensory cortex but also the primary motor area. Here, we investigated whether the effects of a repetitive mechanical tactile stimulation (MS) on somatosensory evoked magnetic fields (SEFs) and cortical oscillations depend on MS patterns. This single‐blinded study included 15 healthy participants. Two types interventions of MS lasting 20 min were used: a repetitive global tactile stimulation (RGS) was used to stimulate the finger by using 24 pins installed on a finger pad, whereas a sequential stepwise displacement tactile stimulation (SSDS) was used to stimulate the finger by moving a row of six pins between the left and right sides on the finger pad. Each parameter was measured pre‐ and post‐intervention. The P50m amplitude of the SEF was increased by RGS and decreased by SSDS. The modulation of P50m was correlated with its amplitude before RGS and with the modulation of beta band oscillation at the resting state after SSDS. This study showed that the effects of a 20‐min MS on SEFs and cortical oscillations depend on mechanical tactile stimulus patterns. Moreover, our results offer potential for the modulation of tactile functions and selection of stimulation patterns according to cortical states. [ABSTRACT FROM AUTHOR]

Spatially resolved absolute electroluminescence (EL) imaging demonstrates the localized EL intensity and the uniformity of solar cells. Combined with two‐dimensional (2‐D) distributed circuit network modeling, detailed and important information that is contained in experimental data can be extracted for in‐depth understanding of solar cell performances. Herein, we measured the absolute EL images of three different solar cells (Si, GaAs, and Cu (In,Ga)Se2 (CIGS)) and observed the different injection‐current‐dependent EL intensities of the defect points (dark or bright) on the solar cells. The origins of these defects were attributed to different defect types according to our established 2‐D distributed equivalent circuit model. The results demonstrated that the combination of absolute EL imaging and distributed circuit modeling yielded accurate quantitative diagnoses of the electrical defects in solar cells. [ABSTRACT FROM AUTHOR]

Summary: Phosphorus (P) is one of the limiting macronutrients for algal growth in marine environments. Microalgae have developed adaptation mechanisms to P limitation that involve remodelling of internal phosphate resources and accumulation of lipids.Here, we used in silico analyses to identify the P‐stress regulator PtPSR (Phaeodactylum tricornutum phosphorus starvation response) in the diatom P. tricornutum. ptpsr mutant lines were generated using gene editing and characterised by various molecular, genetics and biochemical tools.PtPSR belongs to a clade of Myb transcription factors that are conserved in stramenopiles and distantly related to plant P‐stress regulators. PtPSR bound specifically to a conserved cis‐regulatory element found in the regulatory region of P‐stress‐induced genes. ptpsr knockout mutants showed reduction in cell growth under P limitation. P‐stress responses were impaired in ptpsr mutants compared with wild‐type, including reduced induction of P‐stress response genes, near to complete loss of alkaline phosphatase activity and reduced phospholipid degradation.We conclude that PtPSR is a key transcription factor influencing P scavenging, phospholipid remodelling and cell growth in adaptation to P stress in diatoms. [ABSTRACT FROM AUTHOR]

Abstract: With this study the effects of water deficit on radish growth, dry mass production and partitioning, source–sink relations, physiological responses and accumulation of secondary metabolites in storage roots and leaves have been investigated. Three water stress treatments were applied: (Control) 100% of water holding capacity (WHC), (WS50) 50% of WHC, (WS30) 30% of WHC. Water stress was effectively acquired starting from 287 growing degree days (GDD) as demonstrated by the lower values of relative water content (RWC), higher leaf temperature and NIR‐based water indices values in water stressed conditions. Water stress reduced storage root dry weight by 27% at WS50 and 53% at WS30 as well as leaf dry matter accumulation (by 23% and 31% in WS50 and WS30, respectively), expansion (by 28% and 45% in WS50 and WS30, respectively) and specific leaf area (by 7% and 20% in WS50 and WS30, respectively) at 403 GDD. The increasing of leaf‐to‐root mass (L/R) and leaf area‐to‐root mass (LA/R) ratios indicated less dry matter allocation into storage organs under water stress conditions. Besides, water shortage increased leaf greenness as estimated by the higher soil–plant analysis development (SPAD) values (+14% and +20% on average for WS50 and WS30, respectively); other reflectance indices only partially confirmed SPAD readings. Substantially, water limitation did not significantly influence total anthocyanin content, ABTS‐radical scavenging activity and total free phenolic compounds in storage roots, as well as the total free phenolic compounds in leaves. Radish showed a strong plasticity in its adaptation to drought thanks to avoidance mechanisms such as constrained leaf development, increased leaf thickness and adjusted source–sink relationships. [ABSTRACT FROM AUTHOR]

Magnetoencephalography ( MEG) recordings were performed to investigate the inhibitory effects of conditioning stimuli with various types of interstimulus intervals ( ISIs) or intensities on somatosensory evoked magnetic fields ( SEFs) using a 306-ch whole-head MEG system. Twenty-three healthy volunteers participated in this study. Electrical stimuli were applied to the right median nerve at the wrist. Six pulse trains with ISIs of 500 ms were presented in Experiment 1. A paired-pulse paradigm with three kinds of conditioning stimulus ( CON) intensities, 500 ms before the test stimulus ( TS), was applied in Experiment 2. Finally, three CONs 500 or 1000 ms before TS were presented in Experiment 3. Three main SEF deflections (N20m, P35m, and P60m) were observed, and the source activities of P35m and P60m significantly decreased after the 2nd pulse of a six pulse trains. These source activities also significantly decreased with increasing intensity of CON. In addition, these attenuations of source activities were affected by CON- CON or CON- TS intervals. These results indicated that the source activities were modulated by the intensity and ISIs of CONs. Furthermore, P35m after the stimulation were very sensitive to CONs; however, the attenuation of P60m after the stimulation lasted for a longer period than that of P35m. Our findings suggest that the conditioning stimulation had inhibitory effects on subsequent evoked cortical responses for more than 500 ms. Our results also provide important clues about the nature of short-latency somatosensory responses in human studies. [ABSTRACT FROM AUTHOR]

To reveal whether hypocotyl sink activities are regulated by the aboveground parts, and whether physiology and morphology of source leaves are affected by the hypocotyl sink activities, we conducted grafting experiments using two R aphanus sativus varieties with different hypocotyl sink activities. Comet ( C) and Leafy ( L) varieties with high and low hypocotyl sink activities were reciprocally grafted and resultant plants were called by their scion and stock such as CC, LC, CL and LL. Growth, leaf mass per area ( LMA), total non-structural carbohydrates ( TNCs) and photosynthetic characteristics were compared among them. Comet hypocotyls in CC and LC grew well regardless of the scions, whereas Leafy hypocotyls in CL and LL did not. Relative growth rate was highest in LL and lowest in CC. Photosynthetic capacity was correlated with Rubisco (ribulose 1·5-bisphosphate carboxylase/oxygenase) content but unaffected by TNC. High C/ N ratio and accumulation of TNC led to high LMA and structural LMA. These results showed that the hypocotyl sink activity was autonomously regulated by hypocotyl and that the down-regulation of photosynthesis was not induced by TNC. We conclude that the change in the sink activity alters whole-plant growth through the changes in both biomass allocation and leaf morphological characteristics in R . sativus. [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]

Throughout evolution, plants have evolved sophisticated adaptive responses that allow them to grow with a limited supply of phosphate, the preferential form in which the essential macronutrient phosphorus is absorbed by plants. Most of these responses are aimed to increase phosphate availability and acquisition through the roots, to optimize its usage in metabolic processes, and to protect plants from the deleterious effects of phosphate deficiency stress. Regulation of these adaptive responses requires fine perception of the external and internal phosphate levels, and a complex signal transduction pathway that integrates information on the phosphate status at the whole-plant scale. The molecular mechanisms that participate in phosphate homeostasis include transcriptional control of gene expression, RNA silencing mediated by microRNAs, regulatory non-coding RNAs of miRNA activity, phosphate transporter trafficking, and post-translational modification of proteins, such as phosphorylation, sumoylation and ubiquitination. Such a varied regulatory repertoire reflects the complexity intrinsic to phosphate surveying and signaling pathways. Here, we describe these regulatory mechanisms, emphasizing the increasing importance of ubiquitination in the control of phosphate starvation responses. [ABSTRACT FROM AUTHOR]

Xylem sap primarily transports water and mineral nutrients such as nitrogen (N) from roots to shoots in vascular plants. However, it remains largely unknown how nitrogenous compounds, especially proteins in xylem sap, respond to N under- or over-supply. We found that reducing N supply increased amino-N percentage of total N in maize ( Zea mays L.) xylem sap. Proteomic analysis showed that 23 proteins in the xylem sap of maize plants, including 12 newly identified ones, differentially accumulated in response to various N supplies. Fifteen of these 23 proteins were primarily involved in general abiotic or biotic stress responses, whereas the other five proteins appeared to respond largely to N under- or over-supply, suggesting distinct protein responses in maize xylem upon N under- and over-supply. Furthermore, one putative xylanase inhibitor and two putative O-glycosyl hydrolases had preferential gene expression in shoots. [ABSTRACT FROM AUTHOR]

Inorganic phosphate (Pi) is an essential nutrient for plants, and the low bioavailability of Pi in soils is often a limitation to growth and development. Consequently, plants have evolved a range of regulatory mechanisms to adapt to phosphorus‐starvation in order to optimise uptake and assimilation of Pi. Recently, significant progress has been made in elucidating these mechanisms. The coordinated expression of a large number of genes is important for many of these adaptations. Several global expression studies using microarray analysis have been conducted in Arabidopsis thaliana. These studies provide a valuable basis for the identification of new regulatory genes and promoter elements to further the understanding of Pi‐dependent gene regulation. With focus on the Arabidopsis transcriptome, we extract common findings that point to new groups of putative regulators, including the NAC, MYB, ethylene response factor/APETALA2, zinc‐finger, WRKY and CCAAT‐binding families. With a number of new discoveries of regulatory elements, a complex regulatory network is emerging. Some regulatory elements, e.g. the transcription factor PHR1 and the microRNA (miRNA) miR399 and associated factors are well documented, yet not fully understood, whereas other suggested components need further characterisation. Here, we evaluate the contribution of the regulatory elements to the P‐responses and present a model comprising factors directly or indirectly involved in transcriptional regulation and the role of miRNAs as regulators and long‐distance signals. A striking feature is a series of feedback loops and parallel mechanisms that can modify and attenuate responses. We suggest that these mechanisms are instrumental in providing an accurate response and in keeping P‐homeostasis. [ABSTRACT FROM AUTHOR]

• Nodulated lupins ( Lupinus angustifolius cv. Wonga) were hydroponically grown under conditions of low phosphate (LP) or adequate phosphate (HP) to assess the effect of phospho enolpyruvate carboxylase (PEPC)-derived organic acids on nitrogen assimilation in LP nodules. • LP conditions are linked to altered organic acid metabolism, by the engagement of PEP metabolism via PEPC. In LP nodules, the enhanced organic acid synthesis may reduce the available organic carbon for nitrogen assimilation. The diversion of carbon between the organic acid- and amino acid pools was assessed through key nodular enzymes and 14CO2 metabolism. • Under LP conditions, increased rates of organic acid synthesis via PEPC and malate dehydrogenase (MDH), coincided with reduced nitrogen assimilation via aspartate aminotransferase (AAT), aspartate synthetase (AS) and glutamine synthetase (GS)/glutamate synthase (GOGAT) activities. There was a preferential metabolism of nodular 14CO2 into organic acids and particularly into malate. High malate levels were associated with reduced N2 fixation and synthesis of amino acids. • These results indicate that phosphorus deficiency can enhance malate synthesis in nodules, but that excessive malate accumulation may inhibit N2 fixation and nitrogen assimilation. [ABSTRACT FROM AUTHOR]

Previous findings have shown that the human somatosensory cortical systems that are activated by passive nonpainful electrical stimulation include the contralateral primary somatosensory area (SI), bilateral secondary somatosensory area (SII), and bilateral insula. The present study tested the hypothesis that these areas have different sensitivities to stimulation frequency in the condition of passive stimulation. Functional MRI (fMRI) was recorded in 24 normal volunteers during nonpainful electrical median nerve stimulations at 0.5, 1, 2, and 4 Hz repetition rates in separate recording blocks in pseudorandom order. Results of the blood oxygen level-dependent (BOLD) effect showed that the contralateral SI, the bilateral SII, and the bilateral insula were active during these stimulations. As a major finding, only the contralateral SI increased its activation with the increase of the stimulus frequency at the mentioned range. The fact that nonpainful median-nerve electrical stimuli at 4 Hz induces a larger BOLD response is of interest both for basic research and clinical applications in subjects unable to perform cognitive tasks in the fMRI scanner. Hum Brain Mapp 2006. © 2006 Wiley-Liss, Inc. [ABSTRACT FROM AUTHOR]

Expression of selected genes in relation to phosphate (Pi) starvation and sugar sensing was studied in leaves of Arabidopsis. Excised leaf segments with different P status were supplied with combinations of Pi and sugars. Sugar-inducible genes, encoding β-amylase (β-AMY) and chalcone synthase (CHS), were also induced by P deficiency, and were more strongly regulated by sugars when leaf segments originated from P-starved plants. Furthermore, transcript levels of the P-starvation-inducible genes ACP5 (encoding an acid phosphatase), RNS1 (encoding a ribonuclease), and IPS1 (unknown function) increased in response to exogenously applied sugars. Supply of Pi to the leaf segments reversed both P-starvation-induced and sugar-induced gene expression. These interactions reveal a close relationship between P and sugar sensing. To differentiate between hexokinase-dependent and hexokinase-independent sugar sensing the effect of the glucose analogue 2-deoxyglucose and gene expression in the hexokinase-1 deficient mutant, gin2-1, were studied. Both β-AMY and CHS were induced by supplying sucrose to excised leaves but not by 2-deoxyglucose, confirming that these genes are regulated by hexokinase-independent sugar sensing. In the gin2-1 mutant both β-AMY and CHS responded clearly to P starvation excluding that hexokinase-1 mediates the response to P. Similarly, the P-responding genes, IPS1 and RNS1 were repressed by addition of Pi also in the gin2-1 mutant. In conclusion, several phosphate starvation-induced genes are also sugar-induced and hexokinase-independent sugar sensing in Arabidopsis is strongly intensified by phosphate starvation. [ABSTRACT FROM AUTHOR]

Expression of phosphate starvation inducible genes was studied during recovery from phosphate starvation of Arabidopsis thaliana. Genes analysed were ACP5 (encoding an acid phosphatase), RNS1 (encoding an RNase), At4 and IPS1 (both of unknown function), Pht1;7 and Pht2;1 (encoding phosphate transporters). Plants grown at limiting phosphate exhibited reduced growth rate and accumulated anthocyanins, soluble sugars, and starch. Re-supply of phosphate resulted in dramatic uptake of phosphate, increased growth rate and decreased levels of anthocyanins, soluble sugars, and starch in leaf tissue. In both shoots and roots re-supplied with Pi, transcript levels decreased rapidly, and first changes were observed within 30 min. These alterations in gene expression occurred before the content of carbohydrates decreased, indicating that transcriptional regulation was due to phosphate sensing, and not a secondary effect of carbon accumulation. The data reveal different response rates for individual genes and demonstrate that roots and shoots can differ with respect to both timing and genes responding. In general, the changes in transcriptional activity in roots preceded the changes in shoots. Furthermore, transcriptional regulation was observed in isolated roots and shoots. This implies that roots do not strictly require a signal from the shoot, and vice versa, in order to respond to phosphate starvation. [ABSTRACT FROM AUTHOR]

Contents I. Introduction 424 II. The phosphorus conundrum 424 III. Adaptations to low P 424 IV. Uptake of P 424 V. P deficiency alters root development and function 426 VI. P deficiency modifies carbon metabolism 431 VII. Acid phosphatase 436 VIII. Genetic regulation of P responsive genes 437 IX. Improving P acquisition 439 X. Synopsis 440 Summary Phosphorus (P) is limiting for crop yield on > 30% of the world's arable land and, by some estimates, world resources of inexpensive P may be depleted by 2050. Improvement of P acquisition and use by plants is critical for economic, humanitarian and environmental reasons. Plants have evolved a diverse array of strategies to obtain adequate P under limiting conditions, including modifications to root architecture, carbon metabolism and membrane structure, exudation of low molecular weight organic acids, protons and enzymes, and enhanced expression of the numerous genes involved in low-P adaptation. These adaptations may be less pronounced in mycorrhizal-associated plants. The formation of cluster roots under P-stress by the nonmycorrhizal species white lupin (Lupinus albus ), and the accompanying biochemical changes exemplify many of the plant adaptations that enhance P acquisition and use. Physiological, biochemical, and molecular studies of white lupin and other species response to P-deficiency have identified targets that may be useful for plant improvement. Genomic approaches involving identification of expressed sequence tags (ESTs) found under low-P stress may also yield target sites for plant improvement. Interdisciplinary studies uniting plant breeding, biochemistry, soil science, and genetics under the large umbrella of genomics are prerequisite for rapid progress in improving nutrient acquisition and use in plants. [ABSTRACT FROM AUTHOR]

Phosphate (Pi) plays a central role as reactant and effector molecule in plant cell metabolism. However, Pi is the least accessible macronutrient in many ecosystems and its low availability often limits plant growth. Plants have evolved an array of molecular and morphological adaptations to cope with Pi limitation, which include dramatic changes in gene expression and root development to facilitate Pi acquisition and recycling. Although physiological responses to Pi starvation have been increasingly studied and understood, the initial molecular events that monitor and transmit information on external and internal Pi status remain to be elucidated in plants. This review summarizes molecular and developmental Pi starvation responses of higher plants and the evidence for coordinated regulation of gene expression, followed by a discussion of the potential involvement of plant hormones in Pi sensing and of molecular genetic approaches to elucidate plant signalling of low Pi availability. Complementary genetic strategies in Arabidopsis thaliana have been developed that are expected to identify components of plant signal transduction pathways involved in Pi sensing. Innovative screening methods utilize reporter gene constructs, conditional growth on organophosphates and the inhibitory properties of the Pi analogue phosphite, which hold the promise for significant advances in our understanding of the complex mechanisms by which plants regulate Pi-starvation responses. [ABSTRACT FROM AUTHOR]