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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]

Phosphorus (P), an essential macronutrient required for plant growth and development, is often limiting in natural and agro-climatic environments. To cope with heterogeneous or low phosphate (Pi) availability, plants have evolved an array of adaptive responses facilitating optimal acquisition and distribution of Pi. The root system plays a pivotal role in Pi-deficiency-mediated adaptive responses that are regulated by a complex interplay of systemic and local Pi sensing. Cross-talk with sugar, phytohormones, and other nutrient signaling pathways further highlight the intricacies involved in maintaining Pi homeostasis. Transcriptional regulation of Pi-starvation responses is particularly intriguing and involves a host of transcription factors (TFs). Although PHR1 of Arabidopsis is an extensively studied MYB TF regulating subset of Pi-starvation responses, it is not induced during Pi deprivation. Genome-wide analyses of Arabidopsis have shown that low Pi stress triggers spatiotemporal expression of several genes encoding different TFs. Functional characterization of some of these TFs reveals their diverse roles in regulating root system architecture, and acquisition and utilization of Pi. Some of the TFs are also involved in phytohormone-mediated root responses to Pi starvation. The biological roles of these TFs in transcriptional regulation of Pi homeostasis in model plants Arabidopsis thaliana and Oryza sativa are presented in this review. [ABSTRACT FROM AUTHOR]

We surveyed the iron nutrition-responsive transcriptome of Chlamydomonas reinhardtii using RNA-Seq methodology. Presumed primary targets were identified in comparisons between visually asymptomatic iron-deficient versus iron-replete cells. This includes the known components of high-affinity iron uptake as well as candidates for distributive iron transport in C. reinhardtii. Comparison of growth-inhibited iron-limited versus iron-replete cells revealed changes in the expression of genes in chloroplastic oxidative stress response pathways, among hundreds of other genes. The output from the transcriptome was validated at multiple levels: by quantitative RT-PCR for assessing the data analysis pipeline, by quantitative proteomics for assessing the impact of changes in RNA abundance on the proteome, and by cross-species comparison for identifying conserved or universal response pathways. In addition, we assessed the functional importance of three target genes, VITAMIN C 2 (VTC2), MONODEHYDROASCORBATE REDUCTASE 1 (MDAR1), and CONSERVED IN THE GREEN LINEAGE AND DIATOMS 27 (CGLD27), by biochemistry or reverse genetics. VTC2 and MDAR1, which are key enzymes in de novo ascorbate synthesis and ascorbate recycling, respectively, are likely responsible for the 10-fold increase in ascorbate content of iron-limited cells. CGLD27/At5g67370 is a highly conserved, presumed chloroplast-localized pioneer protein and is important for growth of Arabidopsis thaliana in low iron. [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]

An R2R3 MYB transcription factor, OsMYB2P-1, was identified from microarray data by monitoring file expression profile of rice (Oryza sativa ssp. japonica) seedlings exposed to phosphate (Pi)-deficient medium. Expression of OsMYB2P-1 was induced by Pi starvation. OsMYB2P-1 was localized in the nuclei and exhibited transcriptional activation activity. Over expression of OsMYB2P-1 in Arabidopsis (Arabidopsis thaliana) and rice enhanced tolerance to Pi starvation, while suppression of OsMYB2P-1 by RNA interference in rice rendered the transgenic rice more sensitive to Pi deficiency. Furthermore, primary roots of OsMYB2P-1-overexpressing plants were shorter than those in wild-type plants under Pi-sufficient conditions, while primary roots and adventitious roots of OsMYB2P-1-over expressing plants were longer than those of wild-type plants under Pi-deficient conditions. These results suggest that OsMYB2P-1 may also be associated with the regulation of root system architecture. Over expression of OsMYB2P-1 led to greater expression of Pi-responsive genes such as Oryza sativa UDP-sulfoquinovose synthase, OsIPS1, OsPAP1O, OsmiR399a, and OsmiR399j. In contrast, over expression of OsMYB2P-1 suppressed the expression of OsPHO2 under both Pi-sufficient and Pi-deficient conditions. Moreover, expression of OsPT2, which encodes a low-affinity Pi transporter, was up-regulated in OsMYB2P-1- over expressing plants under Pi-sufficient conditions, whereas expression of the high-affinity Pi transporters OsPT6, OsPT8, and OsPT1O was up-regulated by over expression of OsMYB2P-1 under Pi-deficient conditions, suggesting that OsMYB2P-1 may act as a Pi-dependent regulator in controlling the expression of Pi transporters. These findings demonstrate that OsMYB2P-1 is a novel R2R3 MYB transcriptional factor associated with Pi starvation signaling in rice. [ABSTRACT FROM AUTHOR]

Motor dominance is well established, but sensory dominance is much less clear. We therefore studied the cortical evoked magnetic fields using magnetoencephalography (MEG) in a group of 20 healthy right handed subjects in order to examine whether standard electrical stimulation of the median and ulnar nerve demonstrated sensory lateralization. The global field power (GFP) curves, as an indication of cortical activation, did not depict sensory lateralization to the dominant left hemisphere. Comparison of the M20, M30, and M70 peak latencies and GFP values exhibited no statistical differences between the hemispheres, indicating no sensory hemispherical dominance at these latencies for each nerve. Field maps at these latencies presented a first and second polarity reversal for both median and ulnar stimulation. Spatial dipole position parameters did not reveal statistical left-right differences at the M20, M30 and M70 peaks for both nerves. Neither did the dipolar strengths at M20, M30 and M70 show a statistical left-right difference for both nerves. Finally, the Laterality Indices of the M20, M30 and M70 strengths did not indicate complete lateralization to one of the hemispheres. After electrical median and ulnar nerve stimulation no evidence was found for sensory hand dominance in brain responses of either hand, as measured by MEG. The results can provide a new assessment of patients with sensory dysfunctions or perceptual distortion when sensory dominance occurs way beyond the estimated norm. [ABSTRACT FROM AUTHOR]

Background: Phosphorus (P) is an essential macronutrient for all living organisms. Maize (Zea mays) is an important human food, animal feed and energy crop throughout the world, and enormous quantities of phosphate fertilizer are required for maize cultivation. Thus, it is important to improve the efficiency of the use of phosphate fertilizer for maize. Results: In this study, we analyzed the maize root response to phosphate starvation and performed a transcriptomic analysis of the 1.0-1.5 cm lateral root primordium zone. In the growth of plants, the root-to-shoot ratio (R/L) was reduced in both low-phosphate (LP) and sufficient-phosphate (SP) solutions, but the ratio (R/L) exhibited by the plants in the LP solution was higher than that of the SP plants. The growth of primary roots was slightly promoted after 6 days of phosphate starvation, whereas the numbers of lateral roots and lateral root primordia were significantly reduced, and these differences were increased when associated with the stress caused by phosphate starvation. Among the results of a transcriptomic analysis of the maize lateral root primordium zone, there were two highlights: 1) auxin signaling participated in the response and the modification of root morphology under low-phosphate conditions, which may occur via local concentration changes due to the biosynthesis and transport of auxin, and LOB domain proteins may be an intermediary between auxin signaling and root morphology; and 2) the observed retardation of lateral root development was the result of co-regulation of DNA replication, transcription, protein synthesis and degradation and cell growth. Conclusions: These results indicated that maize roots show a different growth pattern than Arabidopsis under low-phosphate conditions, as the latter species has been observed to halt primary root growth when the root tip comes into contact with low-phosphate media. Moreover, our findings enrich our understanding of plant responses to phosphate deficits and of root morphogenesis in maize. [ABSTRACT FROM AUTHOR]

Adequate phosphorus (P) nutrition during early stages is critical for maize growth. Our objective was to evaluate the relative contribution of seed P reserves and exogenous P to maize nutrition during early growth stages. Seedlings were grown with labeled nutrient solution (P). Seedlings were harvested periodically over the course of the three-week study. Initially, 87% and 77% of the total C and N in seeds were located in the endosperm, whereas 86% of seed P was located in the scutellum as phytate. Up to the 7th day after sowing, 96% of phytate was hydrolyzed. Hydrolyzed forms of P were temporarily stored in the seed before being translocated to growing organs, suggesting that the hydrolysis of phytate was not a limiting step for P supply to seedlings. Significant P uptake by roots was observed from the 5th day after sowing on. Both sources of P supplied roots and leaves, with a slightly higher proportion of P from seed reserves going to leaves rather than to roots. Of total seed P, 60% and 92% was exported towards newly growing seedlings till 7th and 17th days after sowing and ceased to be a significant source of P for growth thereafter. We conclude that although both P supply processes overlap in time, seed P was the main P source during early growth stages. [ABSTRACT FROM AUTHOR]

Plants acquire phosphorus in the form of phosphate (Pi), the concentration of which is often limited for plant uptake. Plants have developed diverse responses to conserve and remobilize internal Pi and to enhance Pi acquisition to secure them against Pi deficiency. These responses are achieved by the coordination of an elaborate signaling network comprising local and systemic machineries. Recent advances have revealed several important components involved in this network. Pi functions as a signal to report its own availability. miR399 and sugars act as systemic signals to regulate responses occurring in roots. Hormones also play crucial roles in modulating gene expression and in altering root system architecture. Transcription factors function as a hub to perceive the signals and to elicit steady outputs. In this review, we outline the current knowledge on this subject and present hypotheses pertaining to other potential signals and to the organization and coordination of signaling. [ABSTRACT FROM AUTHOR]

Background and Aims The formation of root cortical aerenchyma (RCA) reduces root respiration and nutrient content by converting living tissue to air volume. It was hypothesized that RCA increases soil resource acquisition by reducing the metabolic and phosphorus cost of soil exploration. Methods To test the quantitative logic of the hypothesis, SimRoot, a functional–structural plant model with emphasis on root architecture and nutrient acquisition, was employed. Sensitivity analyses for the effects of RCA on the initial 40 d of growth of maize (Zea mays) and common bean (Phaseolus vulgaris) were conducted in soils with varying degrees of phosphorus availability. With reference to future climates, the benefit of having RCA in high CO2 environments was simulated. Key Results The model shows that RCA may increase the growth of plants faced with suboptimal phosphorus availability up to 70 % for maize and 14 % for bean after 40 d of growth. Maximum increases were obtained at low phosphorus availability (3 µm). Remobilization of phosphorus from dying cells had a larger effect on plant growth than reduced root respiration. The benefit of both these functions was additive and increased over time. Larger benefits may be expected for mature plants. Sensitivity analysis for light-use efficiency showed that the benefit of having RCA is relatively stable, suggesting that elevated CO2 in future climates will not significantly effect the benefits of having RCA. Conclusions The results support the hypothesis that RCA is an adaptive trait for phosphorus acquisition by remobilizing phosphorus from the root cortex and reducing the metabolic costs of soil exploration. The benefit of having RCA in low-phosphorus soils is larger for maize than for bean, as maize is more sensitive to low phosphorus availability while it has a more ‘expensive’ root system. Genetic variation in RCA may be useful for breeding phosphorus-efficient crop cultivars, which is important for improving global food security. [ABSTRACT FROM PUBLISHER]

A proper concentration and turnover of inorganic phosphate (P[sub i) is essential to maintain cellular processes. Consequently, plants have mechanisms to control P[sub i] homeostasis and to alleviate P[sub i] limitation. The MYB-related transcription factor, PHR1, is important for gene induction during P[sub i] starvation. PHR1 belongs to a family, characterised by the presence of a GARP- and a coiled coil domain. We propose that this family, with 15 members in Arabidopsis thaliana (L.) Heynh., be termed the GCC-family. In this study, transgenic plants overexpressing one member, GCC7, and a T-DNA knockout mutant, gcc7, are characterised. We find overexpressor plants to accumulate more P[sub i] in shoots, irrespective of the P[sub i] supply. Therefore, GCC7 was characterised in relation to P[sub i starvation. We conclude that GCC7 is not strictly required fora P-starvation response since the gcc7mutant respondstoP[sub i] limitation. However, overexpression of GCC7 strongly interferes with the P-starvation response, abolishing induction of several P-responsive genes including AT4, IPS1andseveral P-transportergenes, where as GCC7 does not directly interfere with the PHR1(GCC1) dependent regulation of miR399d. Thus GCC7 influences P-accumulation and P-dependent gene regulation, but GCC7 has a function entirely different from PHR1. [ABSTRACT FROM AUTHOR]

The article cites a research study that describes a chromatin immunoprecipitation (ChIP) protocol for the unicellular green alga Chlamydomonas reinhardtii. It is said that the protocol can analyze nucleosome occupancy, histone modifications and transcription factor binding sites. The optimization of conditions for crosslinking, chromatin fragmentation and antibody titer determination are being described in the study.

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]

Background: Phosphorus (P) is often a limiting mineral nutrient for plant growth. Many soils worldwide are deficient in soluble inorganic phosphate (Pi), the form of P most readily absorbed and utilized by plants. A network of elaborate developmental and biochemical adaptations has evolved in plants to enhance Pi acquisition and avoid starvation. [ABSTRACT FROM PUBLISHER]

An appropriate cellular phosphate (Pi) concentration is indispensable for essential physiological and biochemical processes. To maintain cellular Pi homeostasis, plants have developed a series of adaptive responses to facilitate external Pi acquisition and to limit Pi consumption and to adjust Pi recycling internally when the Pi supply is inadequate. Over the past decade, significant progress has been made toward understanding such regulation at the molecular level. In this review, the focus is on the molecular regulators that mediate cellular Pi concentrations. The regulators are introduced and organized according to their original identification procedures, by the forward genetic approach of mutant screening or by reverse genetic analysis. These genes are involved in Pi uptake, allocation or remobilization or are upstream regulators, such as transcriptional factors or signalling molecules. In the future, integration of current knowledge and exploration of new technology is expected to offer new insights into molecular mechanisms that maintain Pi homeostasis. [ABSTRACT FROM PUBLISHER]

Abstract  Plants count on a wide variety of metabolic, physiological, and developmental responses to adapt their growth to variations in mineral nutrient availability. To react to such variations plants have evolved complex sensing and signaling mechanisms that allow them to monitor the external and internal concentration of each of these nutrients, both in absolute terms and also relatively to the status of other nutrients. Recent evidence has shown that hormones participate in the control of these regulatory networks. Conversely, mineral nutrient conditions influence hormone biosynthesis, further supporting close interrelation between hormonal stimuli and nutritional homeostasis. In this review, we summarize these evidences and analyze possible transcriptional correlations between hormonal and nutritional responses, as a means to further characterize the role of hormones in the response of plants to limiting nutrients in soil. [ABSTRACT FROM AUTHOR]

Maize (Zea mays) is the most widely cultivated crop around the world; however, it is commonly affected by phosphate (Pi) deficiency in many regions, particularly in acid and alkaline soils of developing countries. To cope with Pi deficiency, plants have evolved a large number of developmental and biochemical adaptations; however, for maize, the underlying molecular basis of these responses is still unknown. In this work, the transcriptional response of maize roots to Pi starvation at 1, 3, 6, and 10 d after the onset of Pi deprivation was assessed. The investigation revealed a total of 1179 Pi-responsive genes, of which 820 and 363 genes were found to be either up- or down-regulated, respectively, by 2-fold or more. Pi-responsive genes were found to be involved in various metabolic, signal transduction, and developmental gene networks. A large set of transcription factors, which may be potential targets for crop breeding, was identified. In addition, gene expression profiles and changes in specific metabolites were also correlated. The results show that several dicotyledonous plant responses to Pi starvation are conserved in maize, but that some genetic responses appear to be more specific and that Pi deficiency leads to a shift in the recycling of internal Pi in maize roots. Ultimately, this work provides a more comprehensive view of Pi-responses in a model for economically important cereals and also sets a framework to produce Pi-specific maize microarrays to study the changes in global gene expression between Pi-efficient and Pi-inefficient maize genotypes. [ABSTRACT FROM PUBLISHER]

Phosphorus (P) and sulfur (S) are two macronutrients that photosynthetic organisms require in relatively large amounts despite their levels in the environment often being limited. Accordingly, to adapt to random changes in macronutrient concentrations, plants and algae must sense and respond in a coordinated fashion. The unicellular green alga Chlamydomonas reinhardti is a widely used model organism for the study of P and S stress responses. Herein, we review the current knowledge of P and S nutrient stress responses, highlighting the roles of P and S key global-regulator proteins in mediating signals that link P and S detection to different chloroplast nutrient stress responses. [ABSTRACT FROM AUTHOR]

Abstract  Using a radiogasometric method the rates of photorespiratory and respiratory decarboxylations of primary and stored photosynthates in the leaves of two groups of C3 species, differing in the ability of starch accumulation, were determined. One group included starch-accumulating (SA) species with rates of starch synthesis on the average 38 % the rate of photosynthesis [Solanum tuberosum L., Arabidopsis thaliana (L.) Heynh, Helianthus annuus L., and Plantago lanceolata L.]. The second group represented starch-deficient (SD) species with rates of starch synthesis less than 8 % the rate of photosynthesis (Secale cereale L., Triticum aestivum L., Hordeum vulgare L., and Poa trivialis L.). In SA species the rate of respiration in the dark was significantly higher than in SD species. No differences were found in the rates of photosynthesis, photorespiration, and respiration under irradiation. Thus, the degree of inhibition of respiration by irradiation was in SA species higher than in SD species. It is concluded that starch does not provide substrates for respiratory and photorespiratory decarboxylations in irradiated photosynthesizing leaves. [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]

Sugars in plants, derived from photosynthesis, act as substrates for energy metabolism and the biosynthesis of complex carbohydrates, providing sink tissues with the necessary resources to grow and to develop. In addition, sugars can act as secondary messengers, with the ability to regulate plant growth and development in response to biotic and abiotic stresses. Sugar-signalling networks have the ability to regulate directly the expression of genes and to interact with other signalling pathways. Photosynthate is primarily transported to sink tissues as sucrose via the phloem. Under phosphorus (P) starvation, plants accumulate sugars and starch in their leaves. Increased loading of sucrose to the phloem under P starvation not only functions to relocate carbon resources to the roots, which increases their size relative to the shoot, but also has the potential to initiate sugar-signalling cascades that alter the expression of genes involved in optimizing root biochemistry to acquire soil phosphorus through increased expression and activity of inorganic phosphate transporters, the secretion of acid phosphatases and organic acids to release P from the soil, and the optimization of internal P use. This review looks at the evidence for the involvement of phloem sucrose in co-ordinating plant responses to P starvation at both the transcriptional and physiological levels. [ABSTRACT FROM PUBLISHER]

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]

Ribonuclease LX (RNaseLX) from tomato (Solanum lycopersicum L.) belongs to the RNase T2/S-RNase superfamily of plant endoribonucleases and this is a report on the characterization of the RNaseLX gene and its encoded protein as a member of the phosphate starvation response in tomato. RNaseLX gene sequences were cloned by a PCR-assisted approach. RNaseLX promoter sequences contained the conserved binding motif of the transcription factor PHR1 known to mediate phosphate starvation-dependent gene expression. The increase of RNaseLX transcript levels in roots during phosphate starvation correlated with high promoter activity in transgenic plants carrying a PromLX::uidA gene construct and pointed to transcriptional control of RNaseLX expression. Histochemical staining for β-glucuronidase activity and immunodetection of RNaseLX protein revealed striking RNaseLX expression in main and lateral root tips of phosphate-starved transgenic plants, specifically in epidermal cells, as well as in lateral and adventitious root primordia. Induced RNaseLX expression in roots correlated with stimulated growth and elongation of primary and lateral roots during phosphate deprivation. Phosphate-starvation-induced RNaseLX transcript levels in roots were not modulated by auxin or ethylene. These data indicate that the role of intracellular RNaseLX in the phosphate starvation response is connected with specific RNA turnover processes at the root tip. [ABSTRACT FROM PUBLISHER]

The availability of genome sequences makes it possible to develop microarrays that can be used for profiling gene expression over developmental time, as organisms respond to environmental challenges, and for comparison between wild-type and mutant strains under various conditions. The desired characteristics of microarrays (intense signals, hybridization specificity and extensive coverage of the transcriptome) were not fully met by the previous Chlamydomonas reinhardtii microarray: probes derived from cDNA sequences (~300 bp) were prone to some nonspecific cross-hybridization and coverage of the transcriptome was only ~20%. The near completion of the C. reinhardtii nuclear genome sequence and the availability of extensive cDNA information have made it feasible to improve upon these aspects. After developing a protocol for selecting a high-quality unigene set representing all known expressed sequences, oligonucleotides were designed and a microarray with ~10,000 unique array elements (~70 bp) covering 87% of the known transcriptome was developed. This microarray will enable researchers to generate a global view of gene expression in C. reinhardtii. Furthermore, the detailed description of the protocol for selecting a unigene set and the design of oligonucleotides may be of interest for laboratories interested in developing microarrays for organisms whose genome sequences are not yet completed (but are nearing completion). [ABSTRACT FROM AUTHOR]

Reduced net photosynthesis (Pn) and decreasing shoot and root biomass are typical effects of phosphorus deficiency in plants. Lower biomass accumulation could be the result of reduced Pn (source limitation), but may also be due to direct negative effects of low P availability on growth (sink limitation). Because of the principal importance of root growth for P uptake, this study specifically examined the question whether source or sink limitations were responsible for reduced root growth rates under P deficiency. Rice plants were grown in nutrient solutions with four levels of P supply and at two light treatments and the effect of P×light treatments on growth and carbohydrate distribution was observed. Plants had up to 70% higher Pn when grown with natural (high) light compared with low light. Higher Pn, however, did not lead to additional growth under P deficiency, suggesting that assimilate supply from source leaves to roots was not a limiting factor under P deficiency. This was supported by observations that root starch concentrations increased in P-deficient roots. The comparison of two genotypes with different tolerance to P deficiency showed that the more tolerant one preferentially distributed P to roots where the additional P stimulated root growth and, ultimately, P uptake. The results therefore suggest that source limitation is of little importance under P deficiency. Even at highly sub-optimal tissue P concentrations of below 0.7 mg P g−1 dry weight, plants were able to produce enough assimilates to sustain growth rates that were directly limited by low P availability. [ABSTRACT FROM PUBLISHER]

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]

The influence of phosphorus (P) and nitrogen (N) supply on biomass, leaf area, photon saturated photosynthetic rate ( Pmax), quantum yield efficiency (α), intercellular CO2 concentration ( Ci), and carboxylation efficiency (CE) was investigated in Vicia faba. The influence of P on N accumulation, biomass, and leaf area production was also investigated. An increase in P supply was consistently associated with an increase in N accumulation and N productivity in terms of biomass and leaf area production. Furthermore, P increased the photosynthetic N use efficiency (NUE) in terms of Pmax and α. An increase in P supply was also associated with an increase in CE and a decrease in Ci. Under variable daily meteorological conditions specific leaf nitrogen content (NL), specific leaf phosphorus content (PL), specific leaf area (δL), root mass fraction (Rf), Pmax, and α remained constant for a given N and P supply. A monotonic decline in the steady-state value of Rf occurred with increasing N supply. δL increased with increasing N supply or with increasing NL. We tested also the hypothesis that P supply positively affects both N demand and photosynthetic NUE by influencing the upper limit of the asymptotic values for Pmax and CE, and the lower limit for Ci in response to increasing N. [ABSTRACT FROM AUTHOR]

This review focuses on the biosynthesis of pigments in the unicellular alga Chlamydomonas reinhardtii and their physiological and regulatory functions in the context of information gathered from studies of other photosynthetic organisms. C. reinhardtii is serving as an important model organism for studies of photosynthesis and the pigments associated with the photosynthetic apparatus. Despite extensive information pertaining to the biosynthetic pathways critical for making chlorophylls and carotenoids, we are just beginning to understand the control of these pathways, the coordination between pigment and apoprotein synthesis, and the interactions between the activities of these pathways and those for other important cellular metabolites branching from these pathways. Other exciting areas relating to pigment function are also emerging: the role of intermediates of pigment biosynthesis as messengers that coordinate metabolism in the chloroplast with nuclear gene activity, and the identification of photoreceptors and their participation in critical cellular processes including phototaxis, gametogenesis, and the biogenesis of the photosynthetic machinery. These areas of research have become especially attractive for intensive development with the application of potent molecular and genomic tools currently being applied to studies of C. reinhardtii. [ABSTRACT FROM AUTHOR]

In C4 plants, photorespiration is decreased relative to C3 plants. However, it remains unclear how much photorespiratory capacity C4 leaf tissues actually have. We thoroughly investigated the quantitative distribution of photorespiratory organelles and the immunogold localization of the P protein of glycine decarboxylase (GDC) in mesophyll (M) and bundle sheath (BS) cells of various C4 grass species. Specific differences occurred in the proportions of mitochondria and peroxisomes in the BS cells (relative to the M cells) in photosynthetic tissues surrounding a vein: lower in the NADP-malic enzyme (NADP-ME) species having poorly formed grana in the BS chloroplasts, and higher in the NAD-malic enzyme (NAD-ME) and phosphoenolpyruvate carboxykinase (PCK) species having well developed grana. In all C4 species, GDC was localized mainly in the BS mitochondria. When the total amounts of GDC in the BS mitochondria per unit leaf width were estimated from the immunogold labeling density and the quantity of mitochondria, the BSs of NADP-ME species contained less GDC than those of NAD-ME or PCK species. This trend was also verified by immunoblot analysis of leaf soluble protein. There was a high positive correlation between the degree of granal development (granal index) in the BS chloroplasts and the total amount of GDC in the BS mitochondria. The variations in the structural and biochemical features involved in photorespiration found among C4 species might reflect differences in the O2/CO2 partial pressure and in the potential photorespiratory capacity of the BS cells. [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]

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

Photosynthetic traits of two-year-old Japanese larch seedlings ( Larix kaempferi Carr.) grown at elevated CO2 concentrations were studied in relation to structural changes in the needles. Seedlings were grown at two CO2 concentrations, 360 (AC) and 720 (EC) μmol mol−1 at high and low nutrient supply rates, high N (HN) and low N (LN). The photosynthetic capacity fell significantly in EC+LN, but increased significantly in EC+HN. Since the mesophyll surface area exposed to intercellular space per unit leaf area (Ames/A) is correlated with the photosynthetic rate, we measured Ames/A for larch needles growing in EC. Changes of Ames/A in both EC+HN and EC+LN were very similar to the changes in photosynthetic capacity. This suggests that the changes of Ames/A in EC probably caused the changes in the photosynthetic capacity. The changes of Ames/A in EC were attributed to changes in the mesophyll cell size and mesophyll cell number. The photosynthetic capacity in EC can be explained by taking morphological and structural adaptations into account as well as biochemical factors. [ABSTRACT FROM AUTHOR]

Plants have evolved an array of responses that adapt their growth to conditions of limited phosphate (Pi) supply. These involve biochemical and developmental changes that improve Pi acquisition and recycling, and protect against the stress of Pi starvation. The induction of these responses requires a sophisticated regulatory system that integrates information on external and internal plant Pi status and the details of this regulatory system are only just beginning to be elucidated. In this review, the current knowledge of this regulatory system is summarized, the hallmark of which is the central role of transcription factor PHR1 in the co‐ordinated regulation of many phosphate‐starvation‐responsive genes. The role of hormonal signalling is also described, including auxins, ethylene and, particularly, cytokinins in the regulation of Pi‐starvation responses. [ABSTRACT FROM PUBLISHER]

We previously isolated 31 cDNAs corresponding to pdi [inorganic phosphate (Pi) deficiency-inducible] genes through the differential screening of a cDNA library constructed from the mycelium of Pholiota nameko strain N2 cultured in Pi-depleted medium. Among the cDNAs, pdi251, pdi263 and pdi315 were analyzed here. The deduced amino acid sequences of pdi251, pdi263 and pdi315 showed high similarity to fungal hydrophobins and genes corresponding to these cDNAs encoding P. nameko hydrophobins were designated pnh1, pnh2 and pnh3, respectively. PNH1, PNH2 and PNH3 had a conserved spacing of eight cysteine residues in hydrophobins and a hydropathy pattern characteristic of class I hydrophobins. Phylogenetic analysis showed that PNH1, PNH2 and PNH3 were phylogenetically similar and significantly related to the hydrophobin POH1 specifically expressed in fruiting bodies of Pleurotus ostreatus. Northern blot analysis indicated that, under conditions of Pi deficiency, pnh2 and pnh3 were also induced in strains N4 and N301. [ABSTRACT FROM AUTHOR]

Thirty-one cDNAs corresponding to pdi genes [inorganic phosphate (Pi) deficiency-inducible genes] were previously isolated through the differential screening of a cDNA library constructed from the mycelium of Pholiota nameko. Among the cDNAs, pdi370 was analyzed here. The deduced amino acid sequence showed high similarity to fungal ribonucleases (RNases) and contained two signature sequences conserved in T2 family RNases: CAS1 and CAS2. Genomic DNA harboring the pdi370 gene was isolated from a genomic library of P. nameko. Sequence analysis showed that the pdi370 gene is interrupted by 16 introns and that the promoter region contains two cis-acting sequences found in Pi deficiency-induced genes from Saccharomyces cerevisiae, together with several known functional elements, such as a TATA box. RNase activity in the mycelium and culture filtrate increased 5.6-fold and 5.2-fold, respectively, under Pi-deficient conditions. Staining for RNase activity showed that at least four RNases are induced and secreted under the conditions. The N-terminal sequence of one of them agreed with that of the pdi370 gene product. [ABSTRACT FROM AUTHOR]

The effect of Pi on the properties of phosphoenolpyruvate carboxylase (PEPC) from Amaranthus hypochondriacus, a NAD‐ME type C4 plant, was studied in leaf extracts as well as with purified protein. Efforts were also made to modulate the Pi status of the leaf by feeding leaves with either Pi or mannose. Inclusion of 30 mM Pi during the assay enhanced the enzyme activity in leaf extracts or of purified protein by >2‐fold. The effect of Pi on the enzyme purified from dark‐adapted leaves was more pronounced than that from light‐adapted ones. The Ki for malate increased >2.3‐fold and >1.9‐fold by Pi in the enzyme purified from dark‐adapted leaves and light‐adapted leaves, respectively. Pi also induced an almost 50–60% increase in Km for PEP or Ka for glucose‐6‐phosphate. Feeding the leaves with Pi also increased the activity of PEPC in leaf extracts, while decreasing the malate sensitivity of the enzyme. On the other hand, Pi sequestering by mannose marginally decreased the activity, while markedly suppressing the light activation, of PEPC. There was no change in phosphorylation of PEPC in leaves of A. hypochondriacus due to the feeding of 30 mM Pi. However, feeding with mannose decreased the light‐enhanced phosphorylation of PEPC. The marked decrease in malate sensitivity of PEPC with no change in phosphorylation state indicates that the changes induced by Pi are independent of the phosphorylation of PEPC. It is suggested here that Pi is an important factor in regulating PEPC in vivo and could also be used as a tool to analyse the properties of PEPC. [ABSTRACT FROM PUBLISHER]

Phosphoenolpyruvate carboxylase (PEPC), which catalyses the carboxylation of phosphoenolpyruvate using HCO3– to generate oxaloacetic acid, is an important enzyme in the primary metabolism of plants. Although the PEPC genes (ppc) comprise only a small gene family, the function of each gene is not clear, except for roles in C4 photosynthesis and CAM. Three PEPC genes (Nsppc1–3) from the C3 plant Nicotiana sylvestris were used to investigate their roles and regulation in a C3 plant, and their regulation by phosphorus depletion in particular. First, the induction of PEPC by phosphorus depletion was confirmed. Next, Nsppc1 was determined to be mainly responsive to phosphorus deficiency at the transcriptional level. Further studies using transgenic tobacco harbouring a chimeric gene consisting of the 2.0 kb promoter region of Nsppc1 and the β‐glucuronidase (GUS) reporter showed that PEPC is transcriptionally induced. It was also found that sucrose had a synergistic effect on the induction of PEPC by phosphorus deficiency. A series of transgenic tobacco containing 5′‐deletion mutants of Nsppc1 promoter::GUS fusion revealed that the –539 to –442 bp Nsppc1 promoter region, relative to the translation start site, was necessary for the response to phosphorus deficiency. Gain‐of‐function analysis using a construct containing three tandem repeats of the –539 to –442 bp region confirmed that this region was sufficient to induce the phosphorus‐deficiency response in tobacco. [ABSTRACT FROM PUBLISHER]

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]

Membrane-spanning transport proteins are responsible for the selective passage of most mineral nutrients and metabolites across cellular and intracellular membranes. This review's focus is on summarising the current state of research covering the molecular regulation and biochemical mechanisms involved in the transport of phosphorus, an often growth-limiting nutrient, in vascular plants. Physiological data illustrating the tight control of Pi homeostasis on the cellular as well as on the organism's level are discussed together with the recent results on molecular transport mechanisms. [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]

Implicates the Arabidopsis PHABULOSA (phb) and PHAVOLUTA (pvh) genes in the perception of radial positional information in the leaf primordium. Effect of phb and pvh mutations on leaves and leaf-like organs; Discussion on phb transcript expression; Description on model proposed for phb in the development of leaf polarity.

Examines the role of the mutant alleles of KANADI (KAN) in the regulation of organ polarity in Arabidopsis. Floral defects of KAN in the Landsberg erecta and Wassilewskije genetic backgrounds; Composition of KAN; Evidence supporting the hypothesis that KAN functions as a transcription factor.