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Interaction between two rice mitogen activated protein kinases and its possible role in plant defense
Sheikh, Arsheed H
Raghuram, Badmi
Jalmi, Siddhi K
Wankhede, Dhammaprakash P
Singh, Pallavi
Sinha, Alok K
BMC Plant Biology
The canonical mitogen activated protein kinase (MAPK) signaling pathway plays a vital role in carrying out the normal growth and development of the plant. The pathway, connecting the upstreams signal with the downstream target is considered to be linear, mostly starting with a MAPKKK and ending in a MAPK.
Here we report a novel interaction between two rice MAPKs, OsMPK20-4 and OsMPK3 suggesting the complex nature of the pathway rather than a linear one at individual steps. The interaction between OsMPK20-4 and OsMPK3 found by yeast two-hybrid analysis was confirmed in planta by co-immunoprecipitation and fluorescence resonance energy transfer (FRET) assays. The interaction is specific and is phosphorylation independent. The results suggest a role of the interaction between OsMPK20-4 and OsMPK3 in basic plant defense.
The current novel work showing the physical interaction between two plant MAPKs, OsMPK20-4 and OsMPK3 is the diversion from the dogma of a typical MAPK cascade thereby opening a new dimension to the MAPK signal transduction.
MAP kinase; Protein-protein interaction; Pseudomonas syringae pv. tabaci; Rice; Yeast two-hybrid assay
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Next-generation sequencing-based mRNA and microRNA expression profiling analysis revealed pathways involved in the rapid growth of developing culms in Moso bamboo
He, Cai-yun
Cui, Kai
Zhang, Jian-guo
Duan, Ai-guo
Zeng, Yan-fei
BMC Plant Biology
As one of the fastest-growing lignocellulose-abundant plants on Earth, bamboos can reach their final height quickly due to the expansion of individual internodes already present in the buds; however, the molecular processes underlying this phenomenon remain unclear. Moso bamboo (Phyllostachys heterocycla cv. Pubescens) internodes from four different developmental stages and three different internodes within the same stage were used in our study to investigate the molecular processes at the transcriptome and post-transcriptome level.
Our anatomical observations indicated the development of culms was dominated by cell division in the initial stages and by cell elongation in the middle and late stages. The four major endogenous hormones appeared to actively promote culm development. Using next-generation sequencing-based RNA-Seq, mRNA and microRNA expression profiling technology, we produced a transcriptome and post-transcriptome in possession of a large fraction of annotated Moso bamboo genes, and provided a molecular basis underlying the phenomenon of sequentially elongated internodes from the base to the top. Several key pathways such as environmental adaptation, signal transduction, translation, transport and many metabolisms were identified as involved in the rapid elongation of bamboo culms.
This is the first report on the temporal and spatial transcriptome and gene expression and microRNA profiling in a developing bamboo culms. In addition to gaining more insight into the unique growth characteristics of bamboo, we provide a good case study to analyze gene, microRNA expression and profiling of non-model plant species using high-throughput short-read sequencing. Also, we demonstrate that the integrated analysis of our multi-omics data, including transcriptome, post-transcriptome, proteome, yield more complete representations and additional biological insights, especially the complex dynamic processes occurring in Moso bamboo culms.
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Elucidating the evolutionary history and expression patterns of nucleoside phosphorylase paralogs (vegetative storage proteins) in Populus and the plant kingdom
Pettengill, Emily A
Pettengill, James B
Coleman, Gary D
BMC Plant Biology
Nucleoside phosphorylases (NPs) have been extensively investigated in human and bacterial systems for their role in metabolic nucleotide salvaging and links to oncogenesis. In plants, NP-like proteins have not been comprehensively studied, likely because there is no evidence of a metabolic function in nucleoside salvage. However, in the forest trees genus Populus a family of NP-like proteins function as an important ecophysiological adaptation for inter- and intra-seasonal nitrogen storage and cycling.
We conducted phylogenetic analyses to determine the distribution and evolution of NP-like proteins in plants. These analyses revealed two major clusters of NP-like proteins in plants. Group I proteins were encoded by genes across a wide range of plant taxa while proteins encoded by Group II genes were dominated by species belonging to the order Malpighiales and included the Populus Bark Storage Protein (BSP) and WIN4-like proteins. Additionally, we evaluated the NP-like genes in Populus by examining the transcript abundance of the 13 NP-like genes found in the Populus genome in various tissues of plants exposed to long-day (LD) and short-day (SD) photoperiods. We found that all 13 of the Populus NP-like genes belonging to either Group I or II are expressed in various tissues in both LD and SD conditions. Tests of natural selection and expression evolution analysis of the Populus genes suggests that divergence in gene expression may have occurred recently during the evolution of Populus, which supports the adaptive maintenance models. Lastly, in silico analysis of cis-regulatory elements in the promoters of the 13 NP-like genes in Populus revealed common regulatory elements known to be involved in light regulation, stress/pathogenesis and phytohormone responses.
In Populus, the evolution of the NP-like protein and gene family has been shaped by duplication events and natural selection. Expression data suggest that previously uncharacterized NP-like proteins may function in nutrient sensing and/or signaling. These proteins are members of Group I NP-like proteins, which are widely distributed in many plant taxa. We conclude that NP-like proteins may function in plants, although this function is undefined.
Nucleoside phosphorylases; Vegetative storage proteins; Bark storage proteins; Nitrogen cycling; Populus trichocarpa
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AtCTF7 is required for establishment of sister chromatid cohesion and association of cohesin with chromatin during meiosis in Arabidopsis
Singh, Dipesh K
Andreuzza, Sebastien
Panoli, Aneesh P
Siddiqi, Imran
BMC Plant Biology
The establishment of sister chromatid cohesion followed by its controlled release at the metaphase to anaphase transition is necessary for faithful segregation of chromosomes in mitosis and meiosis. Cohesion is established by the action of Ctf7/Eco1 on the cohesin complex during DNA replication following loading of cohesin onto chromatin by the Scc2-Scc4 complex. Ctf7 is also required for sister chromatid cohesion during repair of DNA double strand breaks. Ctf7 contains an acetyltransferase domain and a zinc finger motif and acetylates conserved lysine residues in the Smc3 subunit of cohesin. In Arabidopsis CTF7 is encoded by a single gene and mutations in AtCTF7 cause embryo lethality indicating that the gene is essential.
To study the function of Ctf7 in plants and to determine its role in sister chromatid cohesion, we constructed a conditional allele of AtCTF7 in Arabidopsis using an inducible RNA interference (RNAi) strategy, so as to avoid the embryo lethality caused by mutations in AtCTF7. We found that induction of RNAi against AtCTF7 caused severe inhibition and defects in growth during vegetative and reproductive stages as well as sterility. AtCTF7-RNAi plants displayed chromosome fragmentation and loss of sister chromatid cohesion during meiosis. Immunostaining for the cohesion subunit AtSCC3 showed a marked reduction in association of cohesin with chromatin during meiosis in AtCTF7-RNAi plants.
We find that AtCTF7 is essential for sister chromatid cohesion during meiosis in Arabidopsis and is required for association of cohesin with chromatin in prophase of meiosis.
Plant meiosis; Chromosome organization; Synapsis; Conditional RNAi; Gametogenesis; DNA repair
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Development of an efficient regeneration and transformation method for the new potential oilseed crop Lepidium campestre
Ivarson, Emelie
Ahlman, Annelie
Li, Xueyuan
Zhu, Li-Hua
BMC Plant Biology
Lepidium campestre is an undomesticated oilseed species with a great potential to become a new crop for both food and industrial feedstocks production. Genetic modification is needed for further improving the oil quantity and quality of Lepidium. Studies on in vitro shoot regeneration of Lepidium are very limited and there is no transformation protocol available.
We have investigated the effects of different factors, especially the type, concentration and combination of plant growth regulators (PGRs) on in vitro shoot regeneration of Lepidium. The results showed that the 2,4-D treatment was crucial to shoot regeneration from different explants. The duration of 2,4-D exposure between 2-4 days did not show significant difference in shoot regeneration, while the effect of 2,4-D concentration varied greatly depending on the type of explants and cytokinins used, for example, the low concentration of 2,4-D combined with TDZ significantly increased the regeneration frequency of hypocotyls. Cotyledon and hypocotyl explants responded differently to cytokinin, for example, TDZ was more effective than zeatin in promoting shoot regeneration from hypocotyls, but did not affect the regeneration of cotyledons which was more affected by high concentration of zeatin. The results also showed that NAA was not effective for shoot regeneration. Germination in light increased the regeneration frequency compared to that in dark. After optimization of the different conditions, an efficient regeneration protocol was developed with the regeneration efficiency of 92.7%. Using this protocol, the transformation frequency of 6% in average was achieved. The presence of transgenes in the transgenic lines was confirmed by GUS staining, PCR and Southern blot analyses.
Through systematic investigation of important factors affecting in vitro shoot regeneration, we have developed an efficient regeneration and transformation protocol for the genetic modification of Lepidium campestre. The method may also be applied to the related species.
Lepidium campestre; Regeneration; Transformation; Hypocotyls; 2,4-D; TDZ
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Potential link between biotic defense activation and recalcitrance to induction of somatic embryogenesis in shoot primordia from adult trees of white spruce (Picea glauca)
Rutledge, Robert G
Stewart, Don
Caron, Sébastien
Overton, Cathy
Boyle, Bryan
MacKay, John
Klimaszewska, Krystyna
BMC Plant Biology
Among the many commercial opportunities afforded by somatic embryogenesis (SE), it is the ability to clonally propagate individual plants with rare or elite traits that has some of the most significant implications. This is particularly true for many long-lived species, such as conifers, but whose long generation times pose substantive challenges, including increased recalcitrance for SE as plants age. Identification of a clonal line of somatic embryo-derived trees whose shoot primordia have remained responsive to SE induction for over a decade, provided a unique opportunity to examine the molecular aspects underpinning SE within shoot tissues of adult white spruce trees.
Microarray analysis was used to conduct transcriptome-wide expression profiling of shoot explants taken from this responsive genotype following one week of SE induction, which when compared with that of a nonresponsive genotype, led to the identification of four of the most differentially expressed genes within each genotype. Using absolute qPCR to expand the analysis to three weeks of induction revealed that differential expression of all eight candidate genes was maintained to the end of the induction treatment, albeit to differing degrees. Most striking was that both the magnitude and duration of candidate gene expression within the nonresponsive genotype was indicative of an intense physiological response. Examining their putative identities further revealed that all four encoded for proteins with similarity to angiosperm proteins known to play prominent roles in biotic defense, and that their high-level induction over an extended period is consistent with activation of a biotic defense response. In contrast, the more temperate response within the responsive genotype, including induction of a conifer-specific dehydrin, is more consistent with elicitation of an adaptive stress response.
While additional evidence is required to definitively establish an association between SE responsiveness and a specific physiological response, these results suggest that biotic defense activation may be antagonistic, likely related to the massive transcriptional and metabolic reprogramming that it elicits. A major issue for future work will be to determine how and if suppressing biotic defense activation could be used to promote a physiological state more conducive to SE induction.
Conifer; Gene expression profiling; Microarray analysis; Absolute qPCR; LRE qPCR; Clonal propagation
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Identification and characterization of the zinc-regulated transporters, iron-regulated transporter-like protein (ZIP) gene family in maize
Li, Suzhen
Zhou, Xiaojin
Huang, Yaqun
Zhu, Liying
Zhang, Shaojun
Zhao, Yongfeng
Guo, Jinjie
Chen, Jingtang
Chen, Rumei
BMC Plant Biology
Zinc (Zn) and iron (Fe) are essential micronutrients for plant growth and development, their deficiency or excess severely impaired physiological and biochemical reactions of plants. Therefore, a tightly controlled zinc and iron uptake and homeostasis network has been evolved in plants. The Zinc-regulated transporters, Iron-regulated transporter-like Proteins (ZIP) are capable of uptaking and transporting divalent metal ion and are suggested to play critical roles in balancing metal uptake and homeostasis, though a detailed analysis of ZIP gene family in maize is still lacking.
Nine ZIP-coding genes were identified in maize genome. It was revealed that the ZmZIP proteins share a conserved transmembrane domain and a variable region between TM-3 and TM-4. Transiently expression in onion epidermal cells revealed that all ZmZIP proteins were localized to the endoplasmic reticulum and plasma membrane. The yeast complementation analysis was performed to test the Zn or Fe transporter activity of ZmZIP proteins. Expression analysis showed that the ZmIRT1 transcripts were dramatically induced in response to Zn- and Fe-deficiency, though the expression profiles of other ZmZIP changed variously. The expression patterns of ZmZIP genes were observed in different stages of embryo and endosperm development. The accumulations of ZmIRT1 and ZmZIP6 were increased in the late developmental stages of embryo, while ZmZIP4 was up-regulated during the early development of embryo. In addition, the expression of ZmZIP5 was dramatically induced associated with middle stage development of embryo and endosperm.
These results suggest that ZmZIP genes encode functional Zn or Fe transporters that may be responsible for the uptake, translocation, detoxification and storage of divalent metal ion in plant cells. The various expression patterns of ZmZIP genes in embryo and endosperm indicates that they may be essential for ion translocation and storage during differential stages of embryo and endosperm development. The present study provides new insights into the evolutionary relationship and putative functional divergence of the ZmZIP gene family during the growth and development of maize.
Embryo; Endosperm; Expression profiling; Zinc; Iron; Zinc-regulated transporters; Iron-regulated transporter-like protein (ZIP); Subcellular localization; Yeast complementation; Maize
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Organ-specific remodeling of the Arabidopsis transcriptome in response to spaceflight
Paul, Anna-Lisa
Zupanska, Agata K
Schultz, Eric R
Ferl, Robert J
BMC Plant Biology
Spaceflight presents a novel environment that is outside the evolutionary experience of terrestrial organisms. Full activation of the International Space Station as a science platform complete with sophisticated plant growth chambers, laboratory benches, and procedures for effective sample return, has enabled a new level of research capability and hypothesis testing in this unique environment. The opportunity to examine the strategies of environmental sensing in spaceflight, which includes the absence of unit gravity, provides a unique insight into the balance of influence among abiotic cues directing plant growth and development: including gravity, light, and touch. The data presented here correlate morphological and transcriptome data from replicated spaceflight experiments.
The transcriptome of Arabidopsis thaliana demonstrated organ-specific changes in response to spaceflight, with 480 genes showing significant changes in expression in spaceflight plants compared with ground controls by at least 1.9-fold, and 58 by more than 7-fold. Leaves, hypocotyls, and roots each displayed unique patterns of response, yet many gene functions within the responses are related. Particularly represented across the dataset were genes associated with cell architecture and growth hormone signaling; processes that would not be anticipated to be altered in microgravity yet may correlate with morphological changes observed in spaceflight plants. As examples, differential expression of genes involved with touch, cell wall remodeling, root hairs, and cell expansion may correlate with spaceflight-associated root skewing, while differential expression of auxin-related and other gravity-signaling genes seemingly correlates with the microgravity of spaceflight. Although functionally related genes were differentially represented in leaves, hypocotyls, and roots, the expression of individual genes varied substantially across organ types, indicating that there is no single response to spaceflight. Rather, each organ employed its own response tactics within a shared strategy, largely involving cell wall architecture.
Spaceflight appears to initiate cellular remodeling throughout the plant, yet specific strategies of the response are distinct among specific organs of the plant. Further, these data illustrate that in the absence of gravity plants rely on other environmental cues to initiate the morphological responses essential to successful growth and development, and that the basis for that engagement lies in the differential expression of genes in an organ-specific manner that maximizes the utilization of these signals – such as the up-regulation of genes associated with light-sensing in roots.
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Novel candidate genes influencing natural variation in potato tuber cold sweetening identified by comparative proteomics and association mapping
Fischer, Matthias
Schreiber, Lena
Colby, Thomas
Kuckenberg, Markus
Tacke, Eckhard
Hofferbert, Hans-Reinhard
Schmidt, Jürgen
Gebhardt, Christiane
BMC Plant Biology
Higher plants evolved various strategies to adapt to chilling conditions. Among other transcriptional and metabolic responses to cold temperatures plants accumulate a range of solutes including sugars. The accumulation of the reducing sugars glucose and fructose in mature potato tubers during exposure to cold temperatures is referred to as cold induced sweetening (CIS). The molecular basis of CIS in potato tubers is of interest not only in basic research on plant adaptation to environmental stress but also in applied research, since high amounts of reducing sugars affect negatively the quality of processed food products such as potato chips. CIS-tolerance varies considerably among potato cultivars. Our objective was to identify by an unbiased approach genes and cellular processes influencing natural variation of tuber sugar content before and during cold storage in potato cultivars used in breeding programs. We compared by two-dimensional polyacrylamide gel electrophoresis the tuber proteomes of cultivars highly diverse for CIS. DNA polymorphisms in genomic sequences encoding differentially expressed proteins were tested for association with tuber starch content, starch yield and processing quality.
Pronounced natural variation of CIS was detected in tubers of a population of 40 tetraploid potato cultivars. Significant differences in protein expression were detected between CIS-tolerant and CIS-sensitive cultivars before the onset as well as during cold storage. Identifiable differential proteins corresponded to protease inhibitors, patatins, heat shock proteins, lipoxygenase, phospholipase A1 and leucine aminopeptidase (Lap). Association mapping based on single nucleotide polymorphisms supported a role of Lap in the natural variation of the quantitative traits tuber starch and sugar content.
The combination of comparative proteomics and association genetics led to the discovery of novel candidate genes for influencing the natural variation of quantitative traits in potato tubers. One such gene was a leucine aminopeptidase not considered so far to play a role in starch sugar interconversion. Novel SNP's diagnostic for increased tuber starch content, starch yield and chip quality were identified, which are useful for selecting improved potato processing cultivars.
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Sequencing-based variant detection in the polyploid crop oilseed rape
Wells, Rachel
Trick, Martin
Fraser, Fiona
Soumpourou, Eleni
Clissold, Leah
Morgan, Colin
Pauquet, Jérôme
Bancroft, Ian
BMC Plant Biology
The detection and exploitation of genetic variation underpins crop improvement. However, the polyploid nature of the genomes of many of our most important crops represents a barrier, particularly for the analysis of variation within genes. To overcome this, we aimed to develop methodologies based on amplicon sequencing that involve the incorporation of barcoded amplification tags (BATs) into PCR products.
A protocol was developed to tag PCR products with 5' 6-base oligonucleotide barcode extensions before pooling for sequencing library production using standard Illumina adapters. A computational method was developed for the de-convolution of products and the robust detection and scoring of sequence variants. Using this methodology, amplicons targeted to gene sequences were screened across a B. napus mapping population and the resulting allele scoring strings for 24 markers linkage mapped to the expected regions of the genome. Furthermore, using one-dimensional 8-fold pooling, 4608 lines of a B. napus mutation population were screened for induced mutations in a locus-specific amplicon (an orthologue of GL2.b) and mixed product of three co-amplified loci (orthologues of FAD2), identifying 10 and 41 mutants respectively.
The utilisation of barcode tags to de-convolute pooled PCR products in multiplexed, variation screening via Illumina sequencing provides a cost effective method for SNP genotyping and mutation detection and, potentially, markers for causative changes, even in polyploid species. Combining this approach with existing Illumina multiplexing workflows allows the analysis of thousands of lines cheaply and efficiently in a single sequencing run with minimal library production costs.
SNP; Mutation; Polyploid; Crop
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A receptor-like kinase gene (GbRLK) from Gossypium barbadense enhances salinity and drought-stress tolerance in Arabidopsis
Zhao, Jun
Gao, Yulong
Zhang, Zhiyuan
Chen, Tianzi
Guo, Wangzhen
Zhang, Tianzhen
BMC Plant Biology
Cotton (Gossypium spp.) is widely cultivated due to the important economic value of its fiber. However, extreme environmental degradation impedes cotton growth and production. Receptor-like kinase (RLK) proteins play important roles in signal transduction and participate in a diverse range of processes in response to plant hormones and environmental cues. Here, we introduced an RLK gene (GbRLK) from cotton into Arabidopsis and investigated its role in imparting abiotic stress tolerance.
GbRLK transcription was induced by exogenously supplied abscisic acid (ABA), salicylic acid, methyl jasmonate, mock drought conditions and high salinity. We cloned the promoter sequence of this gene via self-formed adaptor PCR. Sequence analysis revealed that the promoter region contains many cis-acting stress-responsive elements such as ABRE, W-Box, MYB-core, W-Box core, TCA-element and others. We constructed a vector containing a 1,890-bp sequence in the 5′ region upstream of the initiation codon of this promoter and transformed it into Arabidopsis thaliana. GUS histochemical staining analysis showed that GbRLK was expressed mainly in leaf veins, petioles and roots of transgenic Arabidopsis, but not in the cotyledons or root hairs. GbRLK promoter activity was induced by ABA, PEG, NaCl and Verticillium dahliae. Transgenic Arabidopsis with constitutive overexpression of GbRLK exhibited a reduced rate of water loss in leaves in vitro, along with improved salinity and drought tolerance and increased sensitivity to ABA compared with non-transgenic Col-0 Arabidopsis. Expression analysis of stress-responsive genes in GbRLK Arabidopsis revealed that there was increased expression of genes involved in the ABA-dependent signaling pathway (AtRD20, AtRD22 and AtRD26) and antioxidant genes (AtCAT1, AtCCS, AtCSD2 and AtCSD1) but not ion transporter genes (AtNHX1, AtSOS1).
GbRLK is involved in the drought and high salinity stresses pathway by activating or participating in the ABA signaling pathway. Overexpression of GbRLK may improve stress tolerance by regulating stress-responsive genes to reduce water loss. GbRLK may be employed in the genetic engineering of novel cotton cultivars in the future. Further studying of GbRLK will help elucidate abiotic stress signaling pathways.
Gossypium barbadense; Receptor-like protein kinase; Abscisic acid; Arabidopsis thaliana; Abiotic stress tolerance; Transgene
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Copper-containing amine oxidases contribute to terminal polyamine oxidation in peroxisomes and apoplast of Arabidopsis thaliana
Planas-Portell, Joan
Gallart, Marta
Tiburcio, Antonio F
Altabella, Teresa
BMC Plant Biology
Polyamines (PAs) are oxidatively deaminated at their primary or secondary amino-groups by copper-containing amine oxidases (CuAOs) or FAD-dependent amine oxidases (PAOs), respectively. Both enzymes have long been considered to be apoplastic proteins. However, three out of five PAO isoforms in Arabidopsis thaliana are localized in peroxisomes, while the other two PAOs are predicted to be cytosolic. Interestingly, most of these PAOs do not contribute to terminal PA oxidation, but instead are involved in the back-conversion pathway, producing spermidine from spermine and putrescine from spermidine, which in turn is inhibited by putrescine. This opens the question as to whether PAs are catabolized in the apoplast of Arabidopsis and if the terminal oxidation occurs in the peroxisomes. The main objective of this study was to know if these catabolic processes are mediated by CuAOs.
A. thaliana contains ten genes annotated as CuAOs, but only one (ATAO1) has been characterized at the protein level. Reported herein is the characterization of three genes encoding putative Arabidopsis CuAOs (AtCuAO1, AtCuAO2 and AtCuAO3). These genes encode functional CuAOs that use putrescine and spermidine as substrates. AtCuAO1, like ATAO1, is an extracellular protein, while AtCuAO2 and AtCuAO3 are localized in peroxisomes. The three genes present a different expression profile in response to exogenous treatments, such as application of abcisic acid, methyl jasmonate, salycilic acid, flagellin 22 and wounding.
PA catabolism in the Arabidopsis apoplast is mediated predominantly by CuAOs, while in peroxisomes the co-localization of CuAO-dependent terminal catabolism with PAO-back-conversion machineries might contribute to modulating putrescine-mediated inhibition of the back-conversion, suggesting the occurrence of a tight coordination between both catabolic pathways. The expression profile of AtCuAO1-3 in response to different exogenous treatments, together with the different localization of the corresponding proteins, provides evidence for the functional diversification of Arabidopsis CuAO proteins.
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Reduction of photosynthetic sensitivity in response to abiotic stress in tomato is mediated by a new generation plant activator
Wargent, Jason J
Pickup, Douglas A
Paul, Nigel D
Roberts, Michael R
BMC Plant Biology
Yield losses as a result of abiotic stress factors present a significant challenge for the future of global food production. While breeding technologies provide potential to combat negative stress-mediated outcomes over time, interventions which act to prime plant tolerance to stress, via the use of phytohormone-based elicitors for example, could act as a valuable tool for crop protection. However, the translation of fundamental biology into functioning solution is often constrained by knowledge-gaps.
Photosynthetic and transcriptomic responses were characterised in young tomato (Solanum lycopersicum L.) seedlings in response to pre-treatment with a new plant health activator technology, 'Alethea', followed by a subsequent 100 mM salinity stress. Alethea is a novel proprietary technology composed of three key constituent compounds; the hitherto unexplored compound potassium dihydrojasmonate, an analogue of jasmonic acid; sodium benzoate, a carboxylic acid precursor to salicylic acid, and the α-amino acid L-arginine. Salinity treatment led to a maximal 47% reduction in net photosynthetic rate 8 d following NaCl treatment, yet in Alethea pre-treated seedlings, sensitivity to salinity stress was markedly reduced during the experimental period. Microarray analysis of leaf transcriptional responses showed that while salinity stress and Alethea individually impacted on largely non-overlapping, distinct groups of genes, Alethea pre-treatment substantially modified the response to salinity. Alethea affected the expression of genes related to biotic stress, ethylene signalling, cell wall synthesis, redox signalling and photosynthetic processes. Since Alethea had clear effects on photosynthesis/chloroplastic function at the physiological and molecular levels, we also investigated the ability of Alethea to protect various crop species against methyl viologen, a potent generator of oxidative stress in chloroplasts. Alethea pre-treatment produced dramatic reductions in visible foliar necrosis caused by methyl viologen compared with non-primed controls.
'Alethea' technology mediates positive recovery of abiotic stress-induced photosynthetic and foliar loss of performance, which is accompanied by altered transcriptional responses to stress.
Photosynthesis; Abiotic stress; Priming; Tomato; Transcriptomics; Potassium dihydrojasmonate; Sodium benzoate; L-arginine
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Purple Acid Phosphatase5 is required for maintaining basal resistance against Pseudomonas syringae in Arabidopsis
Ravichandran, Sridhar
Stone, Sophia L
Benkel, Bernhard
Prithiviraj, Balakrishnan
BMC Plant Biology
Plants have evolved an array of constitutive and inducible defense strategies to restrict pathogen ingress. However, some pathogens still manage to invade plants and impair growth and productivity. Previous studies have revealed several key regulators of defense responses, and efforts have been made to use this information to develop disease resistant crop plants. These efforts are often hampered by the complexity of defense signaling pathways. To further elucidate the complexity of defense responses, we screened a population of T-DNA mutants in Colombia-0 background that displayed altered defense responses to virulent Pseudomonas syringae pv. tomato DC3000 (Pst DC3000).
In this study, we demonstrated that the Arabidopsis Purple Acid Phosphatse5 (PAP5) gene, induced under prolonged phosphate (Pi) starvation, is required for maintaining basal resistance to certain pathogens. The expression of PAP5 was distinctly induced only under prolonged Pi starvation and during the early stage of Pst DC3000 infection (6 h.p.i). T-DNA tagged mutant pap5 displayed enhanced susceptibility to the virulent bacterial pathogen Pst DC3000. The pap5 mutation greatly reduced the expression of pathogen inducible gene PR1 compared to wild-type plants. Similarly, other defense related genes including ICS1 and PDF1.2 were impaired in pap5 plants. Moreover, application of BTH (an analog of SA) restored PR1 expression in pap5 plants.
Taken together, our results demonstrate the requirement of PAP5 for maintaining basal resistance against Pst DC3000. Furthermore, our results provide evidence that PAP5 acts upstream of SA accumulation to regulate the expression of other defense responsive genes. We also provide the first experimental evidence indicating the role PAP5 in plant defense responses.
Arabidopsis; Plant defense responses; PAP5; Pseudomonas syringae; Phosphate starvation
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Characterization of temperature and light effects on the defense response phenotypes associated with the maize Rp1-D21 autoactive resistance gene
Negeri, Adisu
Wang, Guan-Feng
Benavente, Larissa
Kibiti, Cromwell M
Chaikam, Vijay
Johal, Guri
Balint-Kurti, Peter
BMC Plant Biology
Rp1 is a complex locus of maize, which carries a set of genes controlling race-specific resistance to the common rust fungus, Puccinia sorghi. The resistance response includes the "Hypersensitive response" (HR), a rapid response triggered by a pathogen recognition event that includes localized cell death at the point of pathogen penetration and the induction of pathogenesis associated genes. The Rp1-D21gene is an autoactive allelic variant at the Rp1 locus, causing spontaneous activation of the HR response, in the absence of pathogenesis. Previously we have shown that the severity of the phenotype conferred by Rp1-D21 is highly dependent on genetic background.
In this study we show that the phenotype conferred by Rp1-D21 is highly dependent on temperature, with lower temperatures favoring the expression of the HR lesion phenotype. This temperature effect was observed in all the 14 genetic backgrounds tested. Significant interactions between the temperature effects and genetic background were observed. When plants were grown at temperatures above 30°C, the spontaneous HR phenotype conferred by Rp1-D21 was entirely suppressed. Furthermore, this phenotype could be restored or suppressed by alternately reducing and increasing the temperature appropriately. Light was also required for the expression of this phenotype. By examining the expression of genes associated with the defense response we showed that, at temperatures above 30°C, the Rp1-D21 phenotype was suppressed at both the phenotypic and molecular level.
We have shown that the lesion phenotype conferred by maize autoactive resistance gene Rp1-D21 is temperature sensitive in a reversible manner, that the temperature-sensitivity phenotype interacts with genetic background and that the phenotype is light sensitive. This is the first detailed demonstration of this phenomenon in monocots and also the first demonstration of the interaction of this effect with genetic background. The use of temperature shifts to induce a massive and synchronous HR in plants carrying the Rp1-D21 genes will be valuable in identifying components of the defense response pathway.
Maize; Hypersensitive response; Disease resistance; Temperature sensitive; Light dependent; Rp1; Autoactive R gene
Related Articles
Novel members of the AGAMOUS LIKE 6 subfamily of MIKCC-type MADS-box genes in soybean
Wong, Chui E
Singh, Mohan B
Bhalla, Prem L
BMC Plant Biology
The classical (C) MIKC-type MADS-box transcription factors comprise one gene family that plays diverse roles in the flowering process ranging from floral initiation to the development of floral organs. Despite their importance in regulating developmental processes that impact crop yield, they remain largely unexplored in the major legume oilseed crop, soybean.
We identified 57 MIKCc-type transcription factors from soybean and determined the in silico gene expression profiles of the soybean MIKCc-type genes across different tissues. Our study implicates three MIKCc-type transcription factors as novel members of the AGAMOUS LIKE 6 (AGL6) subfamily of the MIKCC-type MADS-box genes, and we named this sister clade PsMADS3. While similar genes were identified in other legume species, poplar and grape, no such gene is represented in Arabidopsis thaliana or rice. RT-PCR analysis on these three soybean PsMADS3 genes during early floral initiation processes revealed their temporal expression similar to that of APETALA1, a gene known to function as a floral meristem identity gene. However, RNA in situ hybridisation showed that their spatial expression patterns are markedly different from those of APETALA1.
Legume flower development system differs from that in the model plant, Arabidopsis. There is an overlap in the initiation of different floral whorls in soybean, and inflorescent meristems can revert to leaf production depending on the environmental conditions. MIKCC-type MADS-box genes have been shown to play key regulatory roles in different stages of flower development. We identified members of the PsMADS3 sub-clade in legumes that show differential spatial expression during floral initiation, indicating their potential novel roles in the floral initiation process. The results from this study will contribute to a better understanding of legume-specific floral developmental processes.
Development; Floral meristem; MADS-box transcription factor; Soybean
Related Articles
Dynamic compartment specific changes in glutathione and ascorbate levels in Arabidopsis plants exposed to different light intensities
Heyneke, Elmien
Luschin-Ebengreuth, Nora
Krajcer, Iztok
Wolkinger, Volker
Müller, Maria
Zechmann, Bernd
BMC Plant Biology
Excess light conditions induce the generation of reactive oxygen species (ROS) directly in the chloroplasts but also cause an accumulation and production of ROS in peroxisomes, cytosol and vacuoles. Antioxidants such as ascorbate and glutathione occur in all cell compartments where they detoxify ROS. In this study compartment specific changes in antioxidant levels and related enzymes were monitored among Arabidopsis wildtype plants and ascorbate and glutathione deficient mutants (vtc2-1 and pad2-1, respectively) exposed to different light intensities (50, 150 which was considered as control condition, 300, 700 and 1,500 μmol m-2 s-1) for 4 h and 14 d.
The results revealed that wildtype plants reacted to short term exposure to excess light conditions with the accumulation of ascorbate and glutathione in chloroplasts, peroxisomes and the cytosol and an increased activity of catalase in the leaves. Long term exposure led to an accumulation of ascorbate and glutathione mainly in chloroplasts. In wildtype plants an accumulation of ascorbate and hydrogen peroxide (H2O2) could be observed in vacuoles when exposed to high light conditions. The pad2-1 mutant reacted to long term excess light exposure with an accumulation of ascorbate in peroxisomes whereas the vtc2-1 mutant reacted with an accumulation of glutathione in the chloroplasts (relative to the wildtype) and nuclei during long term high light conditions indicating an important role of these antioxidants in these cell compartments for the protection of the mutants against high light stress.
The results obtained in this study demonstrate that the accumulation of ascorbate and glutathione in chloroplasts, peroxisomes and the cytosol is an important reaction of plants to short term high light stress. The accumulation of ascorbate and H2O2 along the tonoplast and in vacuoles during these conditions indicates an important route for H2O2 detoxification under these conditions.
Arabidopsis; Ascorbate; Chloroplast; Glutathione; High light; Reactive oxygen species
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Cadmium uptake and partitioning in durum wheat during grain filling
Harris, Neil S
Taylor, Gregory J
BMC Plant Biology
Concentrations of cadmium (Cd) in the grain of many durum wheats (Triticum turgidum subsp. durum) grown in North American prairie soils often exceed international trade standards. Genotypic differences in root-to-shoot translocation of Cd are a major determinant of intraspecific variation in the accumulation of Cd in grain. We tested the extent to which changes in whole-plant Cd accumulation and the distribution of Cd between tissues influences Cd accumulation in grain by measuring Cd accumulation throughout the grain filling period in two near-isogenic lines (NILs) of durum wheat that differ in grain Cd accumulation.
Roots absorbed Cd and transported it to the shoots throughout the grain filling period, but the low- and high-Cd NILs did not differ in whole-plant Cd uptake. Although the majority of Cd accumulation was retained in the roots, the low- and high-Cd NILs differed substantively in root-to-shoot translocation of Cd. At grain maturity, accumulation of Cd in the shoots was 13% (low-Cd NIL) or 37% (high-Cd NIL) of whole-plant Cd accumulation. Accumulation of Cd in all shoot tissue, including grain, was at least 2-fold greater in the high-Cd NIL at all harvests. There was no net remobilization of shoot Cd pools during grain filling. The timing of Cd accumulation in grain was positively correlated with grain biomass accumulation, and the rate of grain filling peaked between 14 and 28 days post-anthesis, when both NILs accumulated 60% of total grain biomass and 61-66% of total grain Cd content.
These results show that genotypic variation in root-to-shoot translocation of Cd controls accumulation of Cd in durum wheat grain. Continued uptake of Cd by roots and the absence of net remobilization of Cd from leaves during grain filling support a direct pathway of Cd transport from roots to grain via xylem-to-phloem transfer in the stem.
Cadmium (Cd); Durum wheat; Near-isogenic lines (NIL); Grain filling; Uptake; Translocation; Remobilization
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Elongator subunit 3 positively regulates plant immunity through its histone acetyltransferase and radical S-adenosylmethionine domains
DeFraia, Christopher T
Wang, Yongsheng
Yao, Jiqiang
Mou, Zhonglin
BMC Plant Biology
Pathogen infection triggers a large-scale transcriptional reprogramming in plants, and the speed of this reprogramming affects the outcome of the infection. Our understanding of this process has significantly benefited from mutants that display either delayed or accelerated defense gene induction. In our previous work we demonstrated that the Arabidopsis Elongator complex subunit 2 (AtELP2) plays an important role in both basal immunity and effector-triggered immunity (ETI), and more recently showed that AtELP2 is involved in dynamic changes in histone acetylation and DNA methylation at several defense genes. However, the function of other Elongator subunits in plant immunity has not been characterized.
In the same genetic screen used to identify Atelp2, we found another Elongator mutant, Atelp3-10, which mimics Atelp2 in that it exhibits a delay in defense gene induction following salicylic acid treatment or pathogen infection. Similarly to AtELP2, AtELP3 is required for basal immunity and ETI, but not for systemic acquired resistance (SAR). Furthermore, we demonstrate that both the histone acetyltransferase and radical S-adenosylmethionine domains of AtELP3 are essential for its function in plant immunity.
Our results indicate that the entire Elongator complex is involved in basal immunity and ETI, but not in SAR, and support that Elongator may play a role in facilitating the transcriptional induction of defense genes through alterations to their chromatin.
Arabidopsis; Elongator; Plant immunity; AtELP3; Transcription
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Natural variation in floral nectar proteins of two Nicotiana attenuata accessions
Seo, Pil Joon
Wielsch, Natalie
Kessler, Danny
Svatos, Ales
Park, Chung-Mo
Baldwin, Ian T
Kim, Sang-Gyu
BMC Plant Biology
Floral nectar (FN) contains not only energy-rich compounds to attract pollinators, but also defense chemicals and several proteins. However, proteomic analysis of FN has been hampered by the lack of publically available sequence information from nectar-producing plants. Here we used next-generation sequencing and advanced proteomics to profile FN proteins in the opportunistic outcrossing wild tobacco, Nicotiana attenuata.
We constructed a transcriptome database of N. attenuata and characterized its nectar proteome using LC-MS/MS. The FN proteins of N. attenuata included nectarins, sugar-cleaving enzymes (glucosidase, galactosidase, and xylosidase), RNases, pathogen-related proteins, and lipid transfer proteins. Natural variation in FN proteins of eleven N. attenuata accessions revealed a negative relationship between the accumulation of two abundant proteins, nectarin1b and nectarin5. In addition, microarray analysis of nectary tissues revealed that protein accumulation in FN is not simply correlated with the accumulation of transcripts encoding FN proteins and identified a group of genes that were specifically expressed in the nectary.
Natural variation of identified FN proteins in the ecological model plant N. attenuata suggests that nectar chemistry may have a complex function in plant-pollinator-microbe interactions.
LC-MS/MS; Nectar protein; Nectarin; Nicotiana attenuata
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The dual targeting ability of type II NAD(P)H dehydrogenases arose early in land plant evolution
Xu, Lin
Law, Simon R
Murcha, Monika W
Whelan, James
Carrie, Chris
BMC Plant Biology
Type II NAD(PH) dehydrogenases are located on the inner mitochondrial membrane of plants, fungi, protists and some primitive animals. However, recent observations have been made which identify several Arabidopsis type II dehydrogenases as dual targeted proteins. Targeting either mitochondria and peroxisomes or mitochondria and chloroplasts.
Members of the ND protein family were identified in various plant species. Phylogenetic analyses and subcellular targeting predictions were carried out for all proteins. All ND proteins from three model plant species Arabidopsis, rice and Physcomitrella were cloned as N- and C-terminal GFP fusions and subcellular localisations were determined. Dual targeting of plant type II dehydrogenases was observed to have evolved early in plant evolution and to be widespread throughout different plant species. In all three species tested dual targeting to both mitochondria and peroxisomes was found for at least one NDA and NDB type protein. In addition two NDB type proteins from Physcomitrella were also found to target chloroplasts. The dual targeting of NDC type proteins was found to have evolved later in plant evolution.
The functions of type II dehydrogenases within plant cells will have to be re-evaluated in light of this newly identified subcellular targeting information.
Type II NAD(P)H dehydrogenases; Dual targeting; Mitochondria; Peroxisomes; Plastids
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Progressive 35S promoter methylation increases rapidly during vegetative development in transgenic Nicotiana attenuata plants
Weinhold, Arne
Kallenbach, Mario
Baldwin, Ian Thomas
BMC Plant Biology
Genetically modified plants are widely used in agriculture and increasingly in ecological research to enable the selective manipulation of plant traits in the field. Despite their broad usage, many aspects of unwanted transgene silencing throughout plant development are still poorly understood. A transgene can be epigenetically silenced by a process called RNA directed DNA methylation (RdDM), which can be seen as a heritable loss of gene expression. The spontaneous nature of transgene silencing has been widely reported, but patterns of acquirement remain still unclear.
Transgenic wild tobacco plants (Nicotiana attenuata) expressing heterologous genes coding for antimicrobial peptides displayed an erratic and variable occurrence of transgene silencing. We focused on three independently transformed lines (PNA 1.2, PNA 10.1 and ICE 4.4) as they rapidly lost the expression of the resistance marker and down-regulated transgene expression by more than 200 fold after only one plant generation. Bisulfite sequencing indicated hypermethylation within the 35S and NOS promoters of these lines. To shed light on the progress of methylation establishment, we successively sampled leaf tissues from different stages during plant development and found a rapid increase in 35S promoter methylation during vegetative growth (up to 77% absolute increase within 45 days of growth). The levels of de novo methylation were inherited by the offspring without any visible discontinuation. A secondary callus regeneration step could interfere with the establishment of gene silencing and we found successfully restored transgene expression in the offspring of several regenerants.
The unpredictability of the gene silencing process requires a thorough selection and early detection of unstable plant lines. De novo methylation of the transgenes was acquired solely during vegetative development and did not require a generational change for its establishment or enhancement. A secondary callus regeneration step provides a convenient way to rescue transgene expression without causing undesirable morphological effects, which is essential for experiments that use transformed plants in the analysis of ecologically important traits.
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A non-synonymous SNP within the isopentenyl transferase 2 locus is associated with kernel weight in Chinese maize inbreds (Zea mays L.)
Weng, Jianfeng
Li, Bo
Liu, Changlin
Yang, Xiaoyan
Wang, Hongwei
Hao, Zhuanfang
Li, Mingshun
Zhang, Degui
Ci, Xiaoke
Li, Xinhai
Zhang, Shihuang
BMC Plant Biology
Kernel weight, controlled by quantitative trait loci (QTL), is an important component of grain yield in maize. Cytokinins (CKs) participate in determining grain morphology and final grain yield in crops. ZmIPT2, which is expressed mainly in the basal transfer cell layer, endosperm, and embryo during maize kernel development, encodes an isopentenyl transferase (IPT) that is involved in CK biosynthesis.
The coding region of ZmIPT2 was sequenced across a panel of 175 maize inbred lines that are currently used in Chinese maize breeding programs. Only 16 single nucleotide polymorphisms (SNPs) and seven haplotypes were detected among these inbred lines. Nucleotide diversity (π) within the ZmIPT2 window and coding region were 0.347 and 0.0047, respectively, and they were significantly lower than the mean nucleotide diversity value of 0.372 for maize Chromosome 2 (P < 0.01). Association mapping revealed that a single nucleotide change from cytosine (C) to thymine (T) in the ZmIPT2 coding region, which converted a proline residue into a serine residue, was significantly associated with hundred kernel weight (HKW) in three environments (P <0.05), and explained 4.76% of the total phenotypic variation. In vitro characterization suggests that the dimethylallyl diphospate (DMAPP) IPT activity of ZmIPT2-T is higher than that of ZmIPT2-C, as the amounts of adenosine triphosphate (ATP), adenosine diphosphate (ADP), and adenosine monophosphate (AMP) consumed by ZmIPT2-T were 5.48-, 2.70-, and 1.87-fold, respectively, greater than those consumed by ZmIPT2-C. The effects of artificial selection on the ZmIPT2 coding region were evaluated using Tajima's D tests across six subgroups of Chinese maize germplasm, with the most frequent favorable allele identified in subgroup PB (Partner B).
These results showed that ZmIPT2, which is associated with kernel weight, was subjected to artificial selection during the maize breeding process. ZmIPT2-T had higher IPT activity than ZmIPT2-C, and this favorable allele for kernel weight could be used in molecular marker-assisted selection for improvement of grain yield components in Chinese maize breeding programs.
Maize; Isopentenyl transferase 2; Association mapping; Artificial selection
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Different evolutionary histories of two cation/proton exchanger gene families in plants
Pires, Inês S
Negrão, Sónia
Pentony, Melissa M
Abreu, Isabel A
Oliveira, Margarida M
Purugganan, Michael D
BMC Plant Biology
Gene duplication events have been proposed to be involved in the adaptation of plants to stress conditions; precisely how is unclear. To address this question, we studied the evolution of two families of antiporters. Cation/proton exchangers are important for normal cell function and in plants, Na+,K+/H+ antiporters have also been implicated in salt tolerance. Two well-known plant cation/proton antiporters are NHX1 and SOS1, which perform Na+ and K+ compartmentalization into the vacuole and Na+ efflux from the cell, respectively. However, our knowledge about the evolution of NHX and SOS1 stress responsive gene families is still limited.
In this study we performed a comprehensive molecular evolutionary analysis of the NHX and SOS1 families. Using available sequences from a total of 33 plant species, we estimated gene family phylogenies and gene duplication histories, as well as examined heterogeneous selection pressure on amino acid sites. Our results show that, while the NHX family expanded and specialized, the SOS1 family remained a low copy gene family that appears to have undergone neofunctionalization during its evolutionary history. Additionally, we found that both families are under purifying selection although SOS1 is less constrained.
We propose that the different evolution histories are related with the proteins' function and localization, and that the NHX and SOS1 families are examples of two different evolutionary paths through which duplication events may result in adaptive evolution of stress tolerance.
Related Articles
Lr67 and Lr34 rust resistance genes have much in common – they confer broad spectrum resistance to multiple pathogens in wheat
Spielmeyer, Wolfgang
Mago, Rohit
Wellings, Colin
Ayliffe, Michael
BMC Plant Biology
Adult plant rust resistance genes Lr67 and Lr34 confer race non-specific resistance to multiple fungal pathogens of wheat. Induced, susceptible mutants were characterised for both genes.
Three categories of Lr34 mutants were identified that were either partial susceptible, fully susceptible or hyper-susceptible to stripe rust and leaf rust. The likely impact of the mutational change on the predicted Lr34 protein correlated with differences in response to rust infection. Four independent Lr67 mutants were recovered that were susceptible to stripe rust, leaf rust and stem rust pathogens, including one possible hyper-susceptible Lr67 mutant.
Detailed study of Lr34 mutants revealed that subtle changes in resistance response to multiple pathogens were correlated with mutational changes in the predicted protein. Recovery of independent Lr67 mutants indicates that as for Lr34, a single gene at the Lr67 locus is likely to confer resistance to multiple pathogens. The infection phenotypes of Lr67 mutants closely resembled that of Lr34 mutants.
Lr34; Lr67; Rust resistance; Mutants
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Supplement Facts
Serving Size 1 Capsule
Amount Per Serving | % Daily Value | |
Astragalus Root Powder | 470 mg | * |
*Daily Value not established.
Other ingredients: Gelatin, may contain one or more of the following: microcrystalline cellulose (plant fiber), magnesium stearate, silica.
Suggested Use: As a dietary supplement, take one capsule per day with water.
WARNING: Do not take this product if you are pregnant or nursing. Consult with your healthcare provider before use if you are taking prescription medication or if you have an autoimmune condition.

subset: commoncrawl


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Reviving this thread. I am now a Low GM Zerg player and a High M T player. I also cast at local tournaments, if there's any questions you guys have about the game, don't hesitate to ask
There's also a bunch of good players who can help out.
Q: How much saturation should I have at my bases?
A: Full Saturation is exactly 3 drones per mineral patch, this amounts to 24 plus 6 in gas. However, as stated by this link (Will input link later) it has been proven that 16 drones (2 per patch) produces maximum economic efficiency.
TL is literally your go to place for everything Starcraft. As well as boasting user streams (in which many high level well known and not so well known zergs broadcast) they also have a very handy forum and sub forum, as Snex said, browse through the strategy forum, you'll find threads a plenty from build orders to holding off timing attacks and what not.
Zerg Strategy wiki
MrBitter's 12 Weeks with the Pros
********* Super helpful. ********* - MrBitters is a top level zerg player who I'm pretty sure introduced or made popular/refined the whole Infestor/Ling with Mass upgrades and Neural into late game Ultras against T.
Will re-work this.


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Holy smokes. I went to Walgreens today and they had 'get your flu shot today' signs posted EVERYWHERE! At the end of every aisle, in the middle of the aisle and hanging from the ceiling. Even the clerks were wearing tee shirts with the message in huge red letters. I know other stores are doing it too but, geees.
Gave me the creeps.
Walgreens Flu Shots
Started by
, Oct 19 2011 05:16 PM
1 reply to this topic
Posted 19 October 2011 - 05:53 PM
They did not make a new flu vaccine for this year, it is last years for the swine flu, but that has mutated now so its about useless? I would rather invest in astragulus extract and take it for a few months instead! ( Immune builder).
The light shines in the darkness, and the darkness did not overcome it.
John 1:5
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Sunday March 9, 2014
The 14th Annual Marky B Boardfest
- Registration: Starts @ 8:00am in the Evergreen Room
- Time: Race on the King Pine Trail starts @ 9:30am
- Location: King Pine Trail, top to bottom
- Awards: Awards and raffle in the Evergreen Room @ 3:00pm. Awards given to 1st, 2nd and 3rd finishers in all categories.
- Forms Needed: Registration form and entry fee of $45. Entry fee includes lift ticket, lunch voucher, 1 raffle ticket and race fee ($25 with a King Pine season pass).
Description: The 14th annual snowboard race in memory of Mark Bierweiler is the most unique snowboard-only event in the region… a slalom race through a course with hits, rolls and jumps! Funds generated from the race will aid and further supporting the snowboard programs at King Pine. Race format: best two combined times out of three runs. Categories: Male and Female, 10 & under, 11-15, 16-20, 21-27, 28-34, 35-49, & 50+ .


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The chronograph's design-conscious Japan!
Car designer of the Audi Design
Chronograph designed by Satoshi WADA.
'Japan traditional aesthetics, based on
Is the watch was manufactured.
Add 坪屋 if shipping & packing free of charge!
If you would like to receive products from this shop in us you must use a Forwarding Service.
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HFF extended highlights, final scores from 8/17 | News
Title (Max 100 Characters)
HFF extended highlights, final scores from 8/17
(KFVS) - The Heartland Football Friday final scores and extended highlights from August 17 are available.
You can click here to see the HFF Scoreboard with the final scores for Friday.
You can click here to see extended highlights from week 1 that includes the following games:
- Perryville Jamboree
- Heath vs. Ballard Memorial
- Farmington Jamboree
- Malden Jamboree
For everything else Football Friday, you can visit our Heartland Football Friday page.
Copyright 2011 KFVS. All rights reserved.


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40 CFR 421.3 - Monitoring and reporting requirements.
§ 421.3
Monitoring and reporting requirements.
The following special monitoring requirements apply to all facilities controlled by this regulation:
The monthly average regulatory values shall be the basis for the monthly average discharge in direct discharge permits and for pretreatment standards. Compliance with the monthly discharge limit is required regardless of the number of samples analyzed and averaged.
Periodic analysis for cyanide are not required for a facility in the primary beryllium subcategory (subpart O of this part) when both of the following conditions are met:
The first wastewater sample taken in each calandar year has been analyzed and found to contain less than 0.07 mg/1 cyanide.
The owner or operator of the primary beryllium manufacturing facility certifies in writing to the POTW authority or permit issuing authority that cyanide is neither generated nor used in the beryllium manufacturing process employed at that facility.
[49 FR 8790, Mar. 8, 1984, as amended at 55 FR 31697, Aug. 3, 1990]
Title 40 published on 2012-07-01
no entries appear in the Federal Register after this date.
This is a list of United States Code sections, Statutes at Large, Public Laws, and Presidential Documents, which provide rulemaking authority for this CFR Part.
This list is taken from the Parallel Table of Authorities and Rules provided by GPO [Government Printing Office].
It is not guaranteed to be accurate or up-to-date, though we do refresh the database weekly. More limitations on accuracy are described at the GPO site.


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This news from the feisty Historic Inns of Rockland was just too fun not to share. Check out this group of innkeeper's answers to health care reform. When it comes the public vs. private debate over health care reform, the conversation at the four Historic Inns of Rockland, ME revolves around whether to have the massage ensuite or in the spa. Along with their senator, Olympia Snowe, this feisty group of four historically preserved inns is not afraid to cross party lines and has their own idea for a prescription drug plan simplified into two tiers – red or white. Wellness coverage here includes complimentary anti-oxidant chocolates and blueberry juice, pie or pancakes.
Best of all, affordability is assured; a cost-cutting measure is included with 10 percent off all rooms (built into the package) along with spa treatments at RHeal Wellness Center and Day Spa. Stress-relief toys even come free. It's all tongue in cheek, but it measures up to one great relaxing weekend geared toward wellness, stress relief and a form of health care reform that promises to be passed by all those participating. The promise: no one will be denied; all pre-existing conditions will be accepted, and affordability is guaranteed; with four inns, you're sure to find one that fits your budget and all wellness criteria.
The expanded prescription coverage for the Historic Inns of Rockland's Health Care Reform package includes not only cardio-training but stress relief and couples therapy all in one.
Cost-cutting measures have been employed. Participants will receive 10% off additional spa services and room rates for this package (only) offered from November 1, 2009 – May 15, 2010 for $510-750 per couple, excluding tax.
The Historic Inns of Rockland have decided not to allow health care reform to cripple their economy. For more information on how four inns are working together to insure health care reform for Maine and travelers beyond , visit www.HistoricInnsofRockland.com or the LimeRock Inn, Captain Lindsey House, Granite Inn and Berry Manor Inn. For reservations, call 877-ROC-INNS (877-762-4667).


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2 hrs ago | Business Wire
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That's actually not a real stretch for the company as it already controls the general partner of MLP Terra Nitrogen boosted its dividend by 50%.
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How do you build a big market for batteries and other forms of energy storage? Here's a look at the key challenges.
Availability: LG starts Global Rollout of the LG G Flex - Begins with ...
LG today announced that it soon launch the recently announced LG G Flex , the world's first curved flexing smartphone, globally - starting in Singapore and Hong Kong this week.
Sioux City Man Facing Murder Charges Pleads Not Guilty
By: ABC9 News firstname.lastname@example.com 'Tis the season for hanging holiday decorations! But with decorations, come fire hazards.
Agrium says it's not considering potash job cuts
The Agrium head office seen before the start of the company's annual meeting in Calgary, in a May 11, 2012 photo.
Open Burn Ban Being Lifted Across Riverside County, Some Hazardous Fire Areas Reopened
As winter weather descends on Riverside County, Cal Fire/Riverside County Fire Chief John R. Hawkins is lifting the suspension of open burning for all areas in Riverside County effective Wednesday, Dec. 4 at 8 a.m. "The threat of wildfires in Riverside County is a year-round concern," he said in a released statement.
Sabic opens $100m R&D centre in China
The centre significantly enhances Sabic's R&D footprint in Asia, building on the existing infrastructure of its application centers - one in Moka, Japan, and the other in Sungnam, South Korea.
Dow Says Cost of Buffett Exit Doesn't Suit Shareholders
Andrew Liveris , the chairman and chief executive officer of Dow Chemical Co., said he's not going to be hurried into paying too much to redeem $3 billion of convertible preferred shares held by Warren Buffett .
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In the future, soda will not be some grotesque concoction of artificial flavors and sugar.
Dow Chemical Sees Unusually High Options Volume
Shares of Dow Chemical saw unusually large options trading on Monday. Traders purchased 23,485 call options on the stock, American Banking and Market News reports.
Potash Corp. to cut hundreds of jobs
The Saskatoon-based PotashCorp says the decision to cut its workforce is necessary because of soft demand for potash and phosphates, which are used to make fertilizer.
Japan Vs. South Korea: Who Will Win The Lithium-Ion Battery War?
There is at least one reason to believe that Japan has much more potential than South Korea to dramatically increase production of Li-ion batteries in the world in the next five years or so.
Dow to Sell Commodity Chemicals Businesses
Dow Chemical Co identified the commodity chemicals businesses it would sell or spin off as part of a plan to sell assets worth $3 billion to $4 billion.
Cabot Corporation Agrees to Spend Over $84 Million to Control Harmful ...
Boston-based Cabot Corporation, the second largest carbon black manufacturer in the United States, has agreed to pay a $975,000 civil penalty and spend an estimated $84 million on state of the art technology to control harmful air pollution, resolving alleged violations of the New Source Review provisions of the Clean Air Act at its three ... (more)


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Comments about Mavic Crossmax SLR 29Er Wheels 2013:
They are light and really snappy epically on climbs. Were a great addition to my specialized stump jumper comp carbon till I destroyed the rear wheel.
I'm not holding it against them because I weigh 235 and did it at a downhill bike park with my hard tail.
I'm running the stock Rovals now and while I know I wont break them, they feel like riding in sand compared to the light weight Mavics.
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Comments about Mavic Crossmax SLR 29Er Wheels 2013:
Incredible set of wheels. Beat my best race time on a 50 mile course by 40 minutes on these babies. They are so smooth and fast rolling. Don't think I can go back to my King/Stan's as they just aren't as responsive. We'll have to see how they hold up over time, but I am impressed so far. If they were 150 bucks cheaper they would be a great deal, but for 1K they are a bit overpriced. Still-almost carbon weight but should have better durability.
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A Maze'n Math
Description: A Maze'n Math - Navigate the mazes in order to eliminate all the numbers. Simple math makes for great obstacles.
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Rudolf Nureyev: A Life in Dance
This special exhibition is dedicated to the life and work of the legendary dancer and choreographer Rudolf Nureyev (1938–1993). It will showcase more than 80 costumes and 50 photographs from the dancer's personal collection, entrusted to the Centre national du costume de scène, France, by the Rudolf Nureyev Foundation, and will incorporate key loans from active ballet companies.
The Soviet-born Nureyev was a rising star in his native country before he defected to France in 1961. He was soon recognized… Show more
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The Fine Arts Museums of San Francisco, comprised of the de Young in Golden Gate Park and the Legion of Honor in Lincoln Park, is the largest public arts institution in San Francisco. |
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text: China's Ag Boom Creates Tension Between Food Security, Environment Journal of Environmental Quality looks at relationship between China's growing demand for food and environmental quality Published on: Jul 5, 2013 Tweet Post to Your Wall. Email Story RSS Permalink Print China now feeds 22% of the world's population on just 9% of its total arable land, largely due to investments in research, technology and government assistance, says a new collection of papers published in the Journal of Environmental Quality. But those advancements come at an environmental cost, the papers note. Nutrient loss has polluted China's water resources, and China has also dipped deeply into global resource supplies, using in recent years more synthetic nitrogen fertilizer than all of North America and Europe combined, the paper found. "As the country has transitioned over the past four decades from an undeveloped nation into the world's second largest economy, very hefty tolls have been placed on China's natural resources and the environment," says China Agricultural University's Fusuo Zhang, an author on several papers in the collection.Field workers in Guilin. (Photo by Mike Wilson, Farm Futures) He notes that some of the most serious pollution problems are linked directly to "injudicious use" of nutrients for crop and animal production. Soil scientists like the University of Deleware's Tom Sims are taking note of the changes, calling for a triple emphasis on food security, efficient resource use and environmental protection. "It's really important that China be successful in what it's trying to do not only to feed all those people, but also for the sake of the global economy they're part of and the global environment," Sims says. But approaches in combatting the nutrient issues that China faces isn't easy. Some hurdles include a lack of teaching outlets and collaboration in research. Scientists who study nutrient management issues tend to work separately from one another, resulting in "too many conflicting messages," Zhang says, for policy makers, businesses, farmers, and the public about how to solve problems.


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2013 Ram 3500 Regular Cab 4X4
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The 2013 3500 Regular Cab 4X4 is a 2-door, up to 3-passenger PF, available in two trims, the ST LWB and the... read more SLT LWB.
Upon introduction, both trims are equipped with a standard 5.7-liter, V8, 383-horsepower engine. A 6-speed a transmission with overdrive is standard. hide
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Ram Rumble Bee Concept Could Go Into Production
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Julie Reiner of the Flatiron Lounge Launches New Basil Hayden's Cocktails
Filed under: Spirits
Expert mixologist and owner of Manhattan's iconic Flatiron Lounge Julie Reiner wants you to drink bourbon all summer long.
Reiner has created two new cocktails using Basil Hayden's Kentucky Straight Bourbon Whiskey, a spirit she has loved since the age of 18 (scandal!). "I started working in a restaurant in Waikiki when I was 18, and I wanted to know what I was serving. I've always thought Basil Hayden's was very mixable. It's spicy, and that flavor comes through in the cocktail."
Basil Hayden's is indeed spicy, and has twice the rye of regular bourbons. Most notably, the body and flavor are significantly lighter than other bourbons or whiskeys, with a kindly alcohol quantity of just 80 proof. Bourbon and whiskey can sound a bit heavy for summertime imbibing, but after trying it for the first time myself on Wednesday, I'd say Basil Hayden's is remarkably refreshing, and I can see why Reiner chose it for Flatiron Lounge's summer menu.


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Andy Reid and the Kansas City Chiefs have a contract agreement, according to several media reports.
Andy Reid was the head coach of the Philadelphia Eagles from 1999 to 2012, leading the Eagles to the Super Bowl in 2004.
The Chiefs are planning an announcement later today.
View the discussion thread.


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Senate Asks C.I.A. to Share Its Report on Interrogations
By MARK MAZZETTI
The Senate Intelligence Committee has asked the C.I.A. for an internal study that lawmakers believe is broadly critical of the C.I.A.'s detention and interrogation program.
Wednesday, December 18, 2013 Last Update:
The Senate Intelligence Committee has asked the C.I.A. for an internal study that lawmakers believe is broadly critical of the C.I.A.'s detention and interrogation program.
The bipartisan budget deal advanced by the Senate on Tuesday may not be a cure-all for the gridlock in Washington, but it provides a much-needed break in the inaction.
The Iraqi foreign minister, Hoshyar Zebari, urged President Hamid Karzai to get over his differences with the Americans and sign a security deal with the United States.
Investors were watching closely as the Federal Reserve, which is wrapping up its latest policy meeting on Wednesday, prepared to announce its plans for the global economy.
The 40,389-mile Olympic torch relay to the Sochi Games in Russia, which hit a low point this week with the death of a runner, has proven to be an exercise in misfortune.
With just two weeks before he takes office, New York's next mayor, Bill de Blasio, has yet to pick a schools chancellor.
Mr. Biggs gained international celebrity for his role in one of Britain's most famous crimes, the Great Train Robbery of 1963, and his subsequent life on the run.
The oil giant said Mikal C. Watts represented clients who turned out to be "phantoms" in the 2010 Gulf Coast spill.
The prime minister, Mykola Azarov, said the agreement for assistance meant financial crisis had been averted.
Mike Woodson has been able to shield himself after previous defeats by citing injuries and poor shooting, but he mentioned "bad coaching" as a factor in the Knicks' latest loss.
The United States has much to learn from other countries, especially in preparing teachers and paying for schools.
A scientist is trying to find out whether nuts have staying power as a healthy treat.
Ask Chad Robertson of Tartine Bakery about branching beyond white sugar and flour.
A Bergen County community less than two miles from the New York State border has good schools and transportation, and its own utilities.
Many luxury hybrids have been sales duds, but there are higher expectations for the new models.
"Part of being a good C.E.O. is not believing that you know best," says the C.E.O and co-founder.
Find the best job in the New York metro area and beyond.
Opinion »One Man, One Woman. Or More.Room for Debate asks: Should courts recognize a right to polygamy among consenting adults? |
Opinion »Op-Ed: A Blow Against Big BrotherA United Nations resolution could transform governments into protectors of privacy rights. |


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HEAT oven to 450ºF.
ROLL Perfect Parmesan Pizza Dough into 12-inch circle on lightly floured surface. Press onto bottom of 12-inch pizza pan sprayed with cooking spray, stretching dough as necessary to fit. Let stand 10 min.
MIX dressing and garlic; spread onto dough. Top with remaining ingredients.
BAKE on bottom oven rack 17 to 18 min. or until crust is golden brown.


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Much of the infrastructure built for the Fair remains. The city's underground public transportation system, the Metro, opened a year before the Expo, and the dirt that was excavated was used to create the artificial island Île Notre-Dame, which is where the Formula One racetrack and casino (formerly the French Pavilion) are located.
There's also Habitat 67, a gravity defying housing development originally conceptualized using Lego pieces (and it shows), and the Biosphere, a geodesic dome that was originally the American Pavilion.
Another international Pavilion that survived the test of time? Jamaica's -- it's used for private events now.
11. The roads are in bad shape
Blame the weather, the materials used or government ineptitude, but Quebec roads don't seem to hold up as well as those of their American or Ontarian neighbors.
In addition to mammoth potholes and cracks, road surface markings seem to fade quickly. Road conditions are especially bad during spring, when the snow melts and temperatures rise.
And that's to say nothing of Montreal's crumbling interchanges and overpasses, which probably look worse than they really are.
Be prepared to be redirected a fair bit during the summer, because that's when most construction work is done.


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EDMONTON - Quarterback Mike Reilly and the Edmonton Eskimos offence took a big step forward on Sunday against the Toronto Argonauts.
"Now, can we do the same on defence?" asked defensive co-ordinator Greg Marshall.
His defence, which was supposed to be the cornerstone of the team while the offence got up to speed, has been very inconsistent this season as the 1-6 Eskimos have lost five consecutive CFL games.
"We have not held up our end of the bargain," Marshall said. "When we do perform the way we're capable of performing, we play pretty well. But when we haven't, when we've got guys do their own thing, miss assignments, whatever, we have not performed well, obviously.
"We haven't had all three levels (defensive line, linebackers and secondary) play a complete game together," he continued. "The challenge right now is we don't know what we're going to get from play to play, never mind week to week.
"We've very inconsistent and we're missing a lot of tackles and we're just not playing very well. The frustrating part is it's not one area. They've taken their turns."
Marshall said the coaches asked the defensive linemen to be more physical against the run game on Sunday "and, for the most part, they did that and we got some decent pass rush out of them. So they stepped up and played better than they had the weeks before. We didn't play as well in other areas of the defence as we had before."
Obviously, missing star linebacker J.C. Sherritt, who set a CFL record for tackles en route to being chosen the league's top defensive player last year, and rookie cornerback Aaron Grymes didn't help matters against Toronto "because, arguably, those are your two best players on defence," said Marshall.
Replacements Rennie Curran at linebacker and Bryan Williams at cornerback both had their ups and downs on Sunday, which was to be expected for their first CFL starts.
"But those two guys were not isolated," Marshall said. "There were other people who struggled, as well."
On a positive note, defensive end Odell Willis had his best outing of the season with two quarterback sacks and two forced fumbles after being challenged by the coaching staff to pick up his game.
"He was in the backfield and caused two or three fumbles and he was a factor," Marshall said. "That's what we need out of Odell and now, hopefully, we can get more people on board this week doing the same.
In the Toronto game, Marshall pointed out that the Eskimos got the Argos offence off the field quickly on some two-and-outs, at times, but allowed too many big plays.
"We never really established that consistency in the game where we went a whole quarter, a quarter-and-a-half, where the offence just didn't do much against us," he said. "That never happened. A couple of series, we'd be good. A couple of series, we weren't good."
Marshall said the mistakes the Eskimos are making shouldn't be happening at this point in the season.
But the players don't have the answers either.
Cornerback Joe Burnett pointed out that the Eskimos are getting burned on second-and-long plays, but believes the defence is real close to getting its act together.
"There's a few breakdowns that actually bite us in the foot big time and, sometimes, costs us six points," he said. "If we can correct some of those things ... we're right there in the ball game."
Burnett thought he may have tweaked the hamstring he previously injured on Sunday and sat out Tuesday's practice, but expects to play Saturday when the Eskimos take on the Saskatchewan Roughriders at 2 p.m. at Commonwealth Stadium.
Defensive halfback Chris Thompson also took Tuesday off and will be assessed again by the training staff on Wednesday.
Linebacker T.J. Hill doesn't understand why the third quarter continues to be an Achilles heel, with the Argos outscoring the Esks 16-0 during that 15-minute period on Sunday.
"The intensity is there ... the enthusiasm and the motivation is there ... but we're not executing like we should or like how we were doing in the first quarter and the second quarter," he said.
"We have sense of urgency," he added. "It's there. It's just that we've got to do it for 60 minutes.
"We'd be talking about things differently if a couple of things had bounced this way or bounced that way, but the reality is we're 1-6 right now and we've just got to improve out there."


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Banged-up Bulls expect to have Noah, Belinelli back on Saturday
Published On: Mar 28 2013 06:56:45 AM CDT
Updated On: Mar 28 2013 06:56:46 AM CDT
Joakim Noah walked around the practice facility on Wednesday morning with a much less noticeable limp, but the plantar fasciitis in his right foot was troublesome enough to keep him out against the Miami Heat on Wednesday night. Shooting guard Marco Belinelli [abdominal strain], as well as Rip Hamilton [back] also missed the game, but the news was good on Noah and Belinelli, with both expected back Saturday against Dallas.
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Set inside a tiny historic log cabin, Rooster's Nest uses roosters throughout its rustic interior. The patchwork quilts on the beds fit the area's Amish influence, and the homey breakfast, complete with raspberry-stuffed French toast, is a sweet treat. From $110. See reviews and ratings. (877) 386-3302; roostersnest.net
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About The Council
Estate and Business Planning Council of Worcester County
The Estate and Business Planning Council of Worcester County celebrated its 50th anniversary last year. The Council's first meeting was on November 15, 1960.
Since its inception, the Estate and Business Planning Council of Worcester County has provided a forum for members to learn more about estate and business planning, and gain a better understanding of the services offered by various professional disciplines that could be of benefit to their clients.
Our Upcoming Meetings
To see a complete listing of meetings, please visit Meetings page.
Our membership year runs from November through May. Prospective members may submit a new member application on-line or via post at any time. Please note that new member applications submitted during the summer months will be reviewed by the Membership Committee in late summer/early fall. Renewing members may renew their membership on-line or via post at any time for the 2012-2013 year. We welcome new members who are involved in advising clients on estate and business planning issues. To learn more about becoming a member, please visit the Membership page.
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- RFG - Financial Statement Analysis
All financial statement analysis reports for your requested company can be accessed below. The SIC # column is the SIC Code for the industry under which the current company is analyzed. When a report is purchased the SIC Code indicated is the industry that the company displayed will be compared against. If a different industry is desired for comparison, purchase of a separate industry analysis is suggested.
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Financial statement analysis updated: December 8, 2013
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