text:
HSulf-2, an extracellular endoglucosamine-6-sulfatase, selectively mobilizes heparin-bound growth factors and chemokines: effects on VEGF, FGF-1, and SDF-1
© Uchimura et al; licensee BioMed Central Ltd. 2006
Received: 23 July 2005
Accepted: 17 January 2006
Published: 17 January 2006
Heparin/heparan sulfate (HS) proteoglycans are found in the extracellular matrix (ECM) and on the cell surface. A considerable body of evidence has established that heparin and heparan sulfate proteoglycans (HSPGs) interact with numerous protein ligands including fibroblast growth factors, vascular endothelial growth factor (VEGF), cytokines, and chemokines. These interactions are highly dependent upon the pattern of sulfation modifications within the glycosaminoglycan chains. We previously cloned a cDNA encoding a novel human endosulfatase, HSulf-2, which removes 6-O-sulfate groups on glucosamine from subregions of intact heparin. Here, we have employed both recombinant HSulf-2 and the native enzyme from conditioned medium of the MCF-7-breast carcinoma cell line. To determine whether HSulf-2 modulates the interactions between heparin-binding factors and heparin, we developed an ELISA, in which soluble factors were allowed to bind to immobilized heparin.
Our results show that the binding of VEGF, FGF-1, and certain chemokines (SDF-1 and SLC) to immobilized heparin was abolished or greatly diminished by pre-treating the heparin with HSulf-2. Furthermore, HSulf-2 released these soluble proteins from their association with heparin. Native Sulf-2 from MCF-7 cells reproduced all of these activities.
Our results validate Sulf-2 as a new tool for deciphering the sulfation requirements in the interaction of protein ligands with heparin/HSPGs and expand the range of potential biological activities of this enzyme.
Heparan sulfate proteoglycans (HSPGs) abundantly exist in the ECM and on the cell surface of most cells [1, 2]. They are composed of a restricted set of core proteins to which one or more glycosaminoglycan chains (GAG) are covalently attached. It is established that HSPGs are involved in a wide range of biological functions through their ability to bind to and modify the activities of a diverse repertoire of ligands including growth factors, morphogens, cytokines, chemokines, proteases, lipases, matrix proteins, and cell adhesion molecules [3–6]. Heparan sulfate (HS) chains and heparins (chemical analogues of these chains) consist of repeating disaccharide units of glucuronic/iduronic acid and glucosamine that are subject to a complex set of modifications involving deacetylation, epimerization, and sulfation. Four different sites of sulfation are found in heparin/HS: the N-, 3-O, and 6-O positions of glucosamine and the 2-O position of the iduronic acid residue. Heparin is highly-sulfated throughout the polymer chain whereas in HS these modifications are concentrated mainly in the S-domains, which consist of contiguous clusters of N-sulfated disaccharide units variably sulfated at the other positions [7–9]. Interspersed with the S domains are regions with low (transition zones) or zero sulfation; however, the transition zones contain a considerable fraction of the 6-O-sulfates in HS [10]. The ligand binding activities of HSPG/heparin depend on patterns of sulfation along the chains [2].
HS chains are dynamically regulated in development and during tumor progression [11, 12]. Since these changes are central to the ligand binding properties of HSPGs, there is considerable interest in mechanisms that generate diversity of the chains. One such mechanism is through regulated expression of enzymes involved in the biosynthesis of heparan sulfate, for example, the sulfotransferases that modify HS chains [13]. Another potential mechanism is through the action of extracellular endosulfatases that remove specific sulfation modifications from intact GAG chains. The first enzyme identified in this category was QSulf-1, which was discovered in quail embryo [14]. Stimulated by this work, we cloned cDNAs encoding Sulf-1 and a new member of the family, Sulf-2, in mouse and human [15]. We showed that both Sulf-1 and Sulf-2 are secreted into conditioned medium when they are expressed in Chinese hamster ovary (CHO) cells. Both possess endoglucosamine-6-sulfatase activity against intact heparin with an optimum at neutral pH. The enzymes liberate sulfate groups from the C-6 position of glucosamine residues on trisulfated -IdoA(2-OSO3)-GlcNSO3(6-OSO3)- disaccharide units of heparin. A similar activity for QSulf-1 has now been confirmed on HS chains [16, 17]. This trisulfated disaccharide structure occurs within the S-domains and is known to be a key element in many of the protein ligand interactions of heparin and HS (see below).
The transcripts corresponding to QSulf-1 and its rat orthologue (RFP-Sulf-1) demonstrate complex spatiotemporal regulation during embryonic development [14, 18] and QSulf-1 plays an essential role in Wnt-dependent differentiation of somites into muscle in quail [14]. In addition, in vitro assays have demonstrated that QSulf-1 promotes both Wnt [14] and bone morphogenetic protein [17] signaling via its sulfatase activity. Less is known about the distribution and function of the Sulfs in adult tissues. However, some interesting correlations have been revealed in tumors. Lai and colleagues have reported downregulation of Sulf-1 transcripts in human ovarian cancer and a subset of hepatocellular carcinomas [19–21]. These workers and others have stressed the potential role of the enzyme in down-modulating certain signaling pathways involved in cell proliferation, since over-expression of Sulf-1 reduces signaling by FGF-2, HB-EGF, or HGF [19–22]. In striking contrast to the results in ovarian cancer, increased levels of SULF1 or SULF2 transcripts are observed in other human cancers including breast and pancreatic carcinomas [23–25]. Upregulation of Sulf-2 at both the transcript and protein levels has been established in two mouse models of mammary carcinoma [25]. Furthermore, cultured human breast carcinoma cells release enzymatically active Sulf-2 into conditioned medium. The upregulation of Sulf-2 raises the possibility that it may be involved in promoting tumor progression [24, 25]. Indeed, we have shown that Sulf-2 has pro-angiogenic activity, a relevant function for contributing to tumorigenesis [25].
Even before the discovery of the Sulfs, the glucosamine-6-sulfate modification of HS was the focus of considerable interest. Thus, it has been shown that suppressing the expression of heparan sulfate 6-O-sulfotransferases via gene silencing strategies has pronounced developmental consequences in both Drosophila and Zebrafish development [26, 27]. Secondly, binding studies with heparin/HS fragments and chemically modified heparins point to an essential contribution of the glucosamine-6-sulfate modification to their interaction with various protein ligands, including FGF-1 [28], FGF-10 [29], PDGF [30], VEFG [31], hepatocyte growth factor [32], lipoprotein lipase [33], herpes simplex glycoprotein C [34], noggin [17], and L- and P-selectin [35]. Here, we have taken advantage of recombinant Sulf-2 generated in 293 cells and of native Sulf-2 obtained from conditioned medium of a breast carcinoma cell line to explore the range of activities for this enzyme. We have established an ELISA to examine the effects of Sulf-2 from these two sources on the interaction of several growth factors and chemokines with immobilized heparin/HS. We found that pre-treatment of heparin with Sulf-2 significantly reduced its reactivity with certain ligands but not others ("pre-binding effect") and the enzyme also dissociated heparin-ligand complexes ("post-binding effect") raising the possibility that it may be involved in mobilizing HS-bound ligands in the ECM and basement membranes.
Effect of recombinant HSulf-2 on the binding of VEGF165, FGF-1 and FGF-2 to heparin
Estimated dissociation constants for the interaction of ligands with immobilized heparin-BSA and heparan sulfate (HS)-BSA
Kd value (nM)
Comparison of rHSulf-2 requirements for mobilization of heparin-bound ligands
Concentration required for 50% release of heparin-bound ligand
rHSulf-2 (μg/mL)
CXCL10 (IP-10)a
Effects of HSulf-2 on chemokine binding to heparin-BSA
Analysis of the activity of native HSulf-2 in MCF-7 conditioned medium
Effects of HSulf-2 on heparan sulfate interactions
The Sulfs are 6-O endosulfatases that act on the trisulfated disaccharide unit (-IdoA(2-OSO3)-GlcNSO3(6-OSO3)-) which is the most common unit in heparin but is largely confined to the S-domains of HS [15–17]. As the heterogeneous pattern of sulfation within S domains is known to dictate the binding specificity of many proteins for heparin/HS [7], the Sulfs could potentially regulate those interactions with a dependence on the presence of trisulfated disaccharides within the binding motif. Here, we have employed Sulf-2 as a tool to explore the binding requirements of several proteins, some of which had been previously characterized and others whose binding requirements were largely unknown. Previous work has shown that QSulf-1 treatment of soluble recombinant form of an HSPG (Glypican-1) reduced its ability to bind a Wnt ligand [16]. Also, the interaction of Noggin with cell surface HSPGs was diminished by the overexpression of QSulf-1 in the cells [17]. In the present study we have taken advantage of two sources of soluble HSulf-2: recombinant enzyme purified from stably transfected 293 cells and a natural form of the enzyme in CM of a human breast carcinoma cell line (MCF-7). It should be noted that we have not been able to obtain expression of high level of Sulf-1 in CM of transfected 293 cells, nor have we yet identified a soluble form of this enzyme from a native source.
We have previously shown that native Sulf-2 in MCF-7 CM has arylsulfatase activity [25]. In the present study, we have used HPLC to demonstrate that MCF-7 CM possesses endosulfatase activity against intact heparin. This result has also been established by an alternative approach using mass spectroscopy [38]. Employing an ELISA based on binding of protein ligands to heparin-BSA conjugate, we found that rHSulf-2 strongly modulated the binding of VEGF165, FGF-1, and SDF-1 to porcine intestinal heparin. In the case of FGF-1, the inhibitory effect of Sulf-2 was also demonstrated for heparan sulfate chains as well, using a HS-BSA conjugate. All of the effects obtained with the recombinant enzyme were observed for MCF-7 CM, and we confirmed that the activity in the CM was due to Sulf-2. The porcine intestinal heparin used in this study contained 70% trisulfated disaccharides, 18% disulfated disaccharides, 7% monosulfated disaccharides and 5% unsulfated disaccharides (data not shown). Based on our previous work [15] and the present study, we estimate that about 80% of the total trisulfated disaccharides in heparin-BSA was converted into disulfated disaccharides by Sulf-2. Thus, only about 22% of the overall sulfate moieties were removed by the enzyme treatment, arguing against the possibility that the reduced binding of selective protein ligands to treated heparin was due to a reduction in overall charge. As heparan sulfates have limited S-domains [7], Sulf-2 would be predicted to have an even smaller effect on the global sulfation of HSPGs. A further indication of the selectivity of Sulf-2 is that there is no relationship between the estimated Kd for binding of each ligand to heparin-BSA (Table 1) and the susceptibility of the interaction to the enzyme (Table 2).
The Sulf-2 effects on FGF-1 binding were anticipated based on a number of correlative studies [28, 39], which have implicated the trisulfated disaccharide motif as a binding element in HS chains. Our VEGF165 results are compatible with a previous study in which chemical 6-O-desulfation of heparin strongly weakened its ability to interact with VEGF165 [31]. However, our findings provide the first direct evidence that 6-O-sulfation of heparin within the context of the trisulfated disaccharide motif is essential for the interaction. The majority of VEGF isoforms are able to bind to heparin/HS [40], and it is anticipated that our results will generalize to these other forms as well. With respect to SDF-1, it has long been known that this highly basic chemokine binds relatively strongly to heparin [41], but the fine specificity of the interaction has not previously been explored. SLC and IL-8 showed a partial sensitivity to the desulfation effects of Sulf-2 on heparin. This partial susceptibility implies that the 6-O-sulfation of the trisulfated disaccharide can contribute but is not absolutely required for a measurable binding interaction with heparin. One important avenue for future investigation is based on our observation that Sulf-2 strongly modulated the interactions of two chemokines (SDF-1 and SLC) with heparin. SDF-1 has been implicated in a diverse range of processes such as lymphocyte chemotaxis, stem cell homing and retention, tissue repair, angiogenesis, and organ-specific metastasis [42], whereas SLC is important in lymphoid organ homeostasis and inflammation [43]. Thus, Sulf-2 though effects on the ECM-association of chemokines could have important roles in a number of normal and pathophysiologic processes. HSulf-1 with an apparently indistinguishable enzymatic activity [15] may exert similar effects on HSPG-bound growth factors and chemokines.
It is well established that one function of heparan sulfate proteoglycans in the ECM and basement membranes is to sequester protein ligands away from signal transduction receptors [44, 45]. A known mechanism to regulate the mobilization of such factors is through the action of heparanase, an endo-ß-D-glucuronidase that degrades heparan sulfate chains into relatively large fragments [46]. This enzyme is present in a number of normal cell types (e.g., leukocytes, platelets, cytotrophoblasts) and is upregulated in several cancers. Expression of heparanase in a number of settings elicits angiogenesis, apparently through the release of HSPG-bound angiogenic factors [44, 47]. Our finding that Sulf-2 can reverse the association between heparin and angiogenic factors (e.g., VEGF) suggests the possibility of functional parallels between these two enzymes. Sulf-2 may mobilize VEGF or other angiogenic factors sequestered in the ECM and increase their bioavailability to endothelial cells that express the appropriate signaling receptors. The ability of rHSulf-2 to promote angiogenesis in the chick chorioallantoic membrane (CAM) assay is consistent with such a scenario [25]. Thus, the observed upregulation of Sulf-2 in mammary carcinomas and its secretion from these cells could directly contribute to tumor angiogenesis and thus tumor growth. Sulf-2 derived from a cancer cell could also conceivably mobilize HSPG-bound growth factors that then act back on the cancer cell and thus trigger its own proliferation. The ability of QSulf-1 to promote Wnt signaling within the cells that express the enzyme provides a plausible paradigm for this type of autocrine pathway [14, 16]. It should be borne in mind that in the context of some other signaling pathways (e.g., FGF-2 and HGF), Sulf-2 is likely to exert a negative effect.
As demonstrated in the present study, Sulf-2 can modulate the interaction of a number of important bioactive proteins with heparin/HS. A major finding of our study is that Sulf-2 completely disrupts the interaction of certain factors with heparin. As expected, the factors for which Sulf-2 strongly blocks binding to heparin in the pre-binding format are the most susceptible to reversal effects in the post-binding assays (VEGF, FGF-1, and SDF-1). These observations indicate that, irrespective of the engagement of a protein ligand, Sulf-2 has access to the 6-O-sulfate group of the trisulfated disaccharide units of intact heparin. Recombinant Sulf-2 provides a valuable tool for investigating the specific sulfation requirements for heparin/HS interactions of interest. Furthermore, future application of the assays described herein will certainly expand the range of interactions that can be modulated by Sulf-2 and should enhance our understanding of the potential functional roles served by this enzyme in situ.
The following materials were obtained commercially from the source indicated. Heparin conjugated with bovine serum albumin (Heparin-BSA), BSA, heparinases (I, II and III) were from Sigma (St Louis, Mo., USA); heparan sulphate (porcine intestinal mucosal) from Organon (Oss, The Netherlands); DEAE-Sephacel from Sigma (Poole, UK); Phenyl-Sepharose CL-4B from Amersham Biosciences (Little Chalfont, UK); Protein A Sepharose beads from RepliGen (Needham, MA); recombinant human VEGF165, human CXCL12 (SDF-1), anti-human VEGF antibody (Ab), anti-human FGF-1 Ab, anti-human FGF-2 Ab, anti-human CXCL12 Ab, anti-CXCL8 Ab and anti-human CCL21 Ab from R&D Systems. All are polyclonal antibodies produced in goats. Recombinant human FGF-1 (acidic FGF), human FGF-2 (basic FGF), human CCL21 (SLC) and CXCL8 (IL-8) were from Leinco Inc. A biotinylated swine anti-goat IgG (H+L) antibody and a streptavidin conjugated with alkaline phosphatase were from Caltag. FLAG-tagged versions of HSulf-2 and the inactive mutant designated ΔCCHSulf-2 were produced in 293 cells and purified as previously described [25]. Polyclonal rabbit anti-bodies against HSulf-2 (H2.1 and H2.3) were generated and prepared as previously described [25].
Heparan sulfate-BSA conjugate
Porcine intestinal mucosal heparan sulfate was coupled, via its reducing end, to BSA based on the procedure of Najjam et al. [48]. Briefly, 2 mg of sodium cyanoborohydride was added to a mixture of 50 mg of HS and 2 mg of BSA in 2 ml of 0.2 M potassium phosphate buffer pH 8. The mixture was then incubated for 2 days at 37°C. After dilution to 10 ml with 0.15 M NaCl, 20 mM phosphate, pH7.4, it was applied to a 5 ml column of DEAE-Sephacel equilibrated in the same solution. After washing with 0.15 M NaCl solution, followed by buffered 0.4 M NaCl, to remove reagents and any non-conjugated BSA, the HS-containing material was then step-eluted with 0.5 ml aliquots of buffered 1.5 M NaCl. HS-containing fractions were identified by spotting aliquots onto Whatman filter paper and staining with Azure A. The pooled fractions were adjusted to 3 M in (NH4)2SO4, by addition of solid (NH4)2SO4, and then applied to a 2 ml column of Phenyl-Sepharose CL-4B equilibrated in 3 M (NH4)2SO4. After washing with 3 M (NH4)2SO4 to elute free HS, the bound HS-BSA conjugate was step-eluted with 0.5 ml aliquots of 0.15 M NaCl, 20 mM phosphate pH7.4. Fractions containing conjugated BSA were identified by their absorbance at 280 nm. The final product ran as a diffuse band, stainable with both Coomassie Blue and Azure A, close to the top of a 10% polyacrylamide SDS-PAGE gel and with a higher apparent MW than free BSA. The BSA content of the conjugate was assessed by absorbance at 280 nm. The HS content was determined by digestion with a mixture of heparinases I, II and III, and measurement of the resulting increase in absorbance at 232 nm. It was estimated that the average level of conjugation was 1–2 HS chains/BSA molecule.
Preparation of recombinant Sulf-2 and CM from MCF-7 cells
FLAG-tagged versions of active Sulf-2 (rHSulf-2) and inactive enzyme (rΔCCHSulf-2) were produced in stably transfected kidney 293 cells as described previously [25]. The purities of rHSulf-2 and rΔCCHSulf-2 were estimated to be 90% and 80%, respectively based on Coomassie blue staining of SDS-PAGE gels. The amount of Sulf-2 protein was quantified by the BCA assay (Pierce). The amounts of rHSulf-2 and rΔCCHSulf-2 were equalized by an ELISA employing an anti-FLAG M2 antibody (Sigma) as the capture reagent and a mix of H2.1 and H2.3 antibodies for detection. The human breast carcinoma cell line, MCF-7, was grown in RPMI-1640 containing 10% fetal calf serum in a 150 mm-flask. Cells (80% confluent) were rinsed with PBS and OptiMEM (Invitrogen) and then incubated in 25 ml of OptiMEM at 37°C in 5% CO2 for 72 h. The CM was collected and then 100-fold concentrated on a Centricon-30. Subsequently, the concentrated CM was dialyzed into 50 mM HEPES, pH 8.0.
ELISAs for "pre-binding" and "post-binding" effects of HSulf-2
The experimental design for these ELISA is schematically shown in Fig 2A. To immobilize heparin, 100 ng/ml of heparin-BSA in phosphate-buffered saline (PBS) was added to the wells (100 μl/well) of a 96-well plate (Immulon 2HB, Dynex Labs). The plate was placed at 4°C overnight. The wells were washed 3 times with PBS containing 0.1% Tween-20 (PBS-T) and then blocked with 3% BSA (Sigma) in PBS-T at 25°C for 2 h. To determine the effects of Sulf-2 on the ligand-binding activities of heparin ("pre-binding" effects), the wells were washed as above and then incubated with 100 μl of a reaction mixture containing 5 μmol of HEPES, pH 7.6, 1 μmol of MgCl2 and enzyme (rHSulf-2, rΔCCHSulf-2 or MCF-7 CM) at 37°C for 4 h. In the cases where we carried out dose response studies with protein ligands, incubation with the enzyme was overnight. The wells were washed 3 times with PBS-T and then incubated with 25 μl of heparin-binding factors (25 nM VEGF, 12.5 nM FGF-2, 12.5 nM FGF-1, and 100 nM CXCL12) in PBS at 25°C for 30 min. The wells were washed 3 times and incubated with 1 μg/ml of each primary antibody in 0.1% BSA/PBS (100 μl/well) at 25°C for 1 h. The wells were washed as above and incubated with 1.2 μg/ml of a biotinylated secondary antibody in 0.1% BSA/PBS (100 μl/well) at 25°C for 30 min. Then, the wells were washed and incubated with 2 μg/ml of an alkali phosphatase-conjugated streptavidin in 0.1% BSA/PBS (100 μl/well) at 25°C for 30 min. Finally, the wells were incubated with PNPP (Pierce) in cacodylate buffer, pH 9.2 (100 μl/well) at 25°C for 5 minutes, and then OD 405 nm was read on a microplate reader. For the analysis of "post-binding" effects (i.e. dissociation of heparin-ligand complexes), 25 μl of each heparin-binding factor was added to plates coated with heparin-BSA after blocking with 3% BSA. The plates were incubated at 25°C for 30 min to form the heparin-factor complex. The wells were washed as above and then incubated with 100 μl of the enzyme reaction mixture as described above. After washing the wells, the amount of bound factor was determined with the antibody detection procedures described above. For the "pre-binding" study of FGF-1 binding to heparan sulfate-BSA, 1 mg/ml of heparan sulfate-BSA was coated on a 96-well plate (100 μl/well). Other procedures are the same as described above for the heparin-BSA assay. Representative data are shown from one of 2–3 independent experiments in all cases.
Western blotting
A purified FLAG-tagged HSulf-2 (10 ng) or concentrated MCF-7 CM (10 μl) was separated by electrophoresis on a reducing SDS-7.5% polyacrylamide gel (Bio-Rad), and blotted onto ProBlott™ membrane (Applied Biosystems, Foster City, CA). The membrane was blocked with 5% skim milk/PBS-T for 1 h and then incubated overnight with an anti-HSulf-2 peptide antibody (see below) at a concentration of 1 μg/ml in 5% skim milk/PBS-T at 4°C. The membrane was washed and incubated with horseradish peroxidase-conjugated goat anti-rabbit IgG (Jackson ImmunoResearch Laboratories Inc, West Grove, PA) (0.2 μg/ml) for 1 h. Bound antibodies were visualized with SuperSignal West Pico Chemiluminescent reagent (Pierce).
Immunoprecipitation of HSulf-2 in MCF-7 CM
A mixture of anti-HSulf-2 peptide antibodies, H2.1 (5 μg) and H2.3 (5 μg) [25], or a rabbit IgG (10 μg), 10 μl of a 50% (v/v) suspension of protein A-Sepharose (IPA-400HC, RepliGen) and 100 μl of 100-fold concentrated CM of MCF-7 cells were mixed in a total volume of 1 ml with 150 mM NaCl and 50 mM Tris-HCl, pH 7.4. The mixture was rocked overnight at 4°C. The immunocomplexes bound to the protein A beads were removed by centrifugation. The "pre-binding" and "post-binding" ELISA activities remaining in the supernatant were assayed as described above.
Endoglucosamine-6-sulfatase assay
Endoglucosamine-6-sulfatase activity of rHSulf-2 and MCF-7 CM was determined by previously established procedures [15]. The standard reaction mixture contained 5 μmol of HEPES, pH7.8, 1 μmol of MgCl2, 5 μg of porcine intestinal heparin (Sigma), and a purified FLAG-tagged HSulf-2 or a 100-fold concentrated MCF-7 CM in a total volume of 100 μl. The mixture was incubated at 37°C. The reaction was stopped by heating at 100°C for 2 min. The heparin was digested into disaccharides by a mix of heparinase I, heparinase II and heparinase III [15]. The disaccharides were then analyzed by HPLC on a Partisil-10 SAX column (Whatman, Fairfield, NJ) [15]. The enzymatic activity was defined by calculating the increase in the moles of unsaturated disulfated disaccharide, IdoA(2S)-Glc(NS), in the digested heparin.
List of abbreviations used
Chinese hamster ovary, CM, conditioned medium
extracellular matrix
fibroblast growth factor
heparan sulfate
heparan sulfate proteoglycan
- HSulf-2:
human Sulf-2
- rHSulf-2:
recombinant flag-tagged human Sulf-2
- rΔCCHSulf-2:
recombinant inactive mutant of human Sulf-2
serial analysis of gene expression
stromal cell-derived factor
vascular endothelial growth factor.
This work was supported by National Institutes of Health (NIH) Grants R01 GM57411 and R37 GM23547 (SDR), P01 AI53196 (ZW), a Mizutani Foundation Grant for Glycoscience (SDR), a UCSF Comprehensive Cancer Center Intramural Award from the Alexander and Margaret Stewart Trust (SDR). KU and MM-T are Research Fellows of the Japan Society for the Promotion of Science.
Authors' Affiliations
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- Kamimura K, Fujise M, Villa F, Izumi S, Habuchi H, Kimata K, Nakato H: Drosophila heparan sulfate 6-O-sulfotransferase (dHS6ST) gene. Structure, expression, and function in the formation of the tracheal system. J Biol Chem. 2001, 276: 17014-17021. 10.1074/jbc.M011354200.PubMedView ArticleGoogle Scholar
- Bink RJ, Habuchi H, Lele Z, Dolk E, Joore J, Rauch GJ, Geisler R, Wilson SW, den Hertog J, Kimata K, Zivkovic D: Heparan sulfate 6-o-sulfotransferase is essential for muscle development in zebrafish. J Biol Chem. 2003, 278: 31118-31127. 10.1074/jbc.M213124200.PubMedView ArticleGoogle Scholar
- Kreuger J, Salmivirta M, Sturiale L, Gimenez-Gallego G, Lindahl U: Sequence analysis of heparan sulfate epitopes with graded affinities for fibroblast growth factors 1 and 2. J Biol Chem. 2001, 276: 30744-30752. 10.1074/jbc.M102628200.PubMedView ArticleGoogle Scholar
- Ashikari-Hada S, Habuchi H, Kariya Y, Itoh N, Reddi AH, Kimata K: Characterization of growth factor-binding structures in heparin/heparan sulfate using an octasaccharide library. J Biol Chem. 2004, 279: 12346-12354. 10.1074/jbc.M313523200.PubMedView ArticleGoogle Scholar
- Feyzi E, Lustig F, Fager G, Spillmann D, Lindahl U, Salmivirta M: Characterization of heparin and heparan sulfate domains binding to the long splice variant of platelet-derived growth factor A chain. J Biol Chem. 1997, 272: 5518-5524. 10.1074/jbc.272.40.24850.PubMedView ArticleGoogle Scholar
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- Kucia M, Jankowski K, Reca R, Wysoczynski M, Bandura L, Allendorf DJ, Zhang J, Ratajczak J, Ratajczak MZ: CXCR4-SDF-1 signalling, locomotion, chemotaxis and adhesion. J Mol Histol. 2004, 35: 233-245. 10.1023/B:HIJO.0000032355.66152.b8.PubMedView ArticleGoogle Scholar
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This article is published under license to BioMed Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.


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To honor open access week, I'll be highlighting a different open access eBook platform each day. The purpose is to create awareness of various open access ebook collections.  If you have collections to suggest, please let me know.
The OAPEN
OAPEN (Open Access Publishing in European Networks) is a collaborative initiative to develop and implement a sustainable Open Access publication model for academic books in the Humanities and Social Sciences. The OAPEN Library aims to improve the visibility and usability of high quality academic research by aggregating peer reviewed Open Access publications from across Europe.
Titles are available in English, German, French, Italian, Latin, and other languages. The peer reviewed titles are submitted by a variety of publishers, including Taylor & Francis, Palgrave, Brill, Berg and a long list of university presses, including Hong Kong, Manchester, Firenze, Amsterdam, and Universitätsverlag Göttingen.  According to the OAPEN September newsletter, they had over 400,000 downloads in the past 12 months. Continue reading Open Access eBooks: Part One, the OAPEN

subset: commoncrawl


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The aim of treatment of pneumonia is to cure the infection and prevent complications. Treatment approach depends on several factors such as your age and general health, the possible causative organism or organisms, and the setting — community or health care — where the infection developed, and severity of infection. Most people with pneumonia can be treated at home.
Treatment approach for pneumonia may include:
1. Medications: Antibiotics, antiviral medicines, fever and cough medicines may be used to treat pneumonia.
2. Other Treatment
3. Hospital Admission
Most people with community-acquired pneumonia do not require hospital care. You may be admitted if you have any two of these indicators of severe pneumonia.
Prevention will always triumph over cure, and when it comes to keeping pneumonia away from your kids, it is not all that difficult.read more
A recent Chinese study has revealed how living in the cities could increase the risk of pneumonia, a number of factors have given rise to this.read more


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Women of the Wise Oak (book 2)
Publishing "Women of the Wise Oak" a historical fantasy sequel to "Children of the Wise Oak" (2016) Professional editing, cover, & printing
From North Devon to London
To take 250 GCSE students from North Devon to London for a two day educational experience.
Samphire Festival 2016
A brand new music festival in a beautiful coastal location within Exmoor National Park. Two days, two nights, two stages.
Behind the Lines
Andrew 'Cotty' Cotton aims to surf the biggest waves on the planet this season and set a new world record for biggest wave ever ridden. Support him on ... more
Triplicity Music & Arts Festival
Let's make Triplicity Music & Arts Festival a sustainable annual gathering. A spiritual home for the Psychedelic underground in the UK.
The Boat on The Hill
AHOY! We are looking for funds to complete the conversion of a 30ft wooden sailing boat into a fantastic nautical Glamping escape.
Limited Edition Book: Mountain Marathon Man
Mountain Marathon Man: Hardback book, which gives an account of the first athlete to run a solo circumnavigation of the world.
New Open-air Theatre in The Valley of the Rocks
Pleasure Dome Theatre launches this summer bringing exceptional open-air entertainment to The Valley of the Rocks. Macbeth 2nd-20th August.
Save North Devon Hedgehog Rescue
We're fundraising for North Devon Hedgehog Rescue to build a sanctuary for 50 hedgehogs. Without our help, the rescue centre may close.
Mend Exmoor!
An ancient trading route through the heart of Exmoor has become a boggy quagmire, but due to its location your help is needed to repair it.
Eva Mama Medical Aid For Refugees
Hi! I am Linda Mizun, a junior doctor with surgical training background. I am currently volunteering in refugee camps in dire need of...
North Devon and Torridge/West Devon Green Party fundraiser
Help fund our election campaign by spreading our policies across our 2 constituencies with the freepost leaflet, printing 100,000+ is costly
Revive Minehead's Community Christmas Extravaganza
A community organised late night Christmas shopping event in Minehead in early Dec . Organised in Association with Revive Minehead
With your help we can launch The Elizabeth Seafood and Tapas Bistro aboard a 1922 boat to be moored at Watchet Marina Somerset.
Music Therapy Training
To study for three years at the University of West England to gain a Masters Degree in Music Therapy and to then practise in West Somerset.
Wayne Green's Recording Project
With your support I could finally record my original songs and release my first EP. Listen to my song "Breathe" on Soundcloud.
My debut album by Charley Charlesworth!
Hi I'm Charley, I am a singer/songwriter, I live near the sea and play folk/punk/pop music. I would like to make a record!
Wigfest Charity Fund Raising Event
We are a fund raising event where all profits from the event go to cancer and locally related charities, in memory of a few friends who...
Save Winkleigh Tower
RAF Winkleigh WWII Control Tower is a scheduled monument, derelict since 1990. You can help save and restore it!
Victorian railway carriage
We have spent forty years carefully restoring this unique 1st Class carriage, and now need to finish the interior with its luxurious...
Broadlands Retirement Home
Registered Charity Broadlands Retirement home needs to replace old and failing kitchen equipment.
Get ND's Beach Soccer boys to the Nationals!
The local lads from North Devon are going to the Beach Soccer National Championships!
North Devon - Retrocade Cafe
Help Me Open Retrocade Cafe - A family run retro 80's style arcade with a cafe.
Minehead's Mighty Christmas Extravaganza
A community organised late night Christmas shopping event in Minehead on Friday 11th Dec . Organised in Association with Revive Minehead


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Home > About RIC > RIC in the News > RIC Launches Prince Family Esprit! Program
Published on August 20, 2012
CHICAGO, IL—Aug. 16, 2012—The Rehabilitation Institute of Chicago (RIC), ranked the #1 rehabilitation hospital in America, launched the Prince Family Esprit! Program designed to offer ongoing education and wellness opportunities for employees. RIC launched this unique program with a large kick-off celebration at its flagship hospital in Chicago and at each of its 50 sites of care throughout Illinois, Indiana and Michigan.
The Prince Family Esprit! Program is made possible through a generous gift from the Frederick Henry Prince Memorial Fund to recognize, honor and develop the strengths and talents of RIC's dedicated staff.
"The Prince family, longstanding and generous friends to RIC, has provided support for a specific purpose: to enhance RIC employee morale," Joanne C. Smith, M.D., President and CEO at RIC. "Funding for a program like this is not only incredibly generous, it's very special and uncommon! We can't thank the Prince family enough for their generous investment in the spirit of RIC's talented people, which will make RIC a better place in everything we do!"
The Prince Family Espirit! Program will focus on three main areas—the mind, body and spirit. Employees will have access to live or online courses that provide continuing learning and growth opportunities that inspire the "mind." Additionally, RIC will launch a number of new wellness initiatives that help employees live longer, stronger, healthier lives. RIC is also implementing new recognition programs and awards that recognize employees' talent and hard work, boosting their "spirit." To celebrate the program's launch, employees were given pedometers to measure their steps and encouraged to take a minimum of 10,000 steps per day to maintain good heart health.
As the day-to-day champions of RIC's vision of Advancing Human Ability™, RIC's committed professionals remain the driving force behind the organization's industry-leading patient treatments and research. Through its strategic focus on fostering employee talent and leadership, RIC has long cultivated an environment of excellence that translates directly into patients' outcomes of regained ability after a traumatic injury such as a brain or spinal cord injury or stroke. The Prince gift builds on this tradition, enabling RIC to make the fullest possible investment in its staff and, thus, in its patients.
"We know that having happy, healthy employees translates to providing better care for patients, so an investment into the well-being and morale of our employees is an investment in our greater vision," said Laura Ferrio, RN, Vice President, Patient Care Services and Chief Nurse Executive, and "RIC is the best place to work!"
The Rehabilitation Institute of Chicago (RIC) is the nation's leading provider of comprehensive physical medicine and rehabilitation care to patients from around the world. Founded in 1953, RIC has been designated the "No. 1 Rehabilitation Hospital in America" by U.S. News & World Report every year since 1991. RIC sets the standard of care in the post-acute market through its innovative applied research and discovery programs, particularly in the areas of neuroscience, bionic medicine, musculoskeletal medicine and technology transfer. The AbilityLab — RIC's new, state-of-the-art research hospital, will open in March 2017.
Journalists contact: email@example.com, 312-238-6016.
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345 E. Superior Street - Chicago, IL 60611 | 1-800-354-REHAB (7342) | 312-238-1000
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Alice + Olivia black sweater from Pretty Little Liars seen with Mona Vanderwaal (Janel Parrish)
As Seen on Mona Vanderwaal (Janel Parrish) in Pretty Little Liars - Season 7 Episode 10
Mona Vanderwaal (Janel Parrish) is seen with the black Alice + Olivia Bow Embroidered Sweater in Pretty Little Liars, Season 7, Episode 10 - The DArkest Knight. Find Sweaters, Clothing and Women's Fashion from Pretty Little Liars below.
Behind that glamorous prom dress is a pretty girl with an ugly secret. In Pretty Little Liars Season 6, Spencer Hastings (Trojan Bellisario) returns with even more mysteries, terror, and dark secrets. Now it's up to her and the girls to stand up to their tormenter "A" and get their lives back. They might be graduating high school but secrets don't end with the school year.
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Merritt Capital Partners, Real Estate - Developers, listed under "Real Estate - Developers" category, is located at 8235 Douglas Ave 1030 Dallas TX, 75225 and can be reached by 2149879808 phone number. Merritt Capital Partners has currently 0 reviews.This business profile is not yet claimed, and if you are the owner, claim your business profile for free. If you are not the owner you can invite this business to join by email or by phone.Browse all Real Estate - Developers in Dallas TX. Discover census data for Dallas, TX.
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This article takes a close look at what disruptive manufacturing processes are involved in getting Sir Richard Branson's Virgin Spacehip Fleet operational.
Tag Archive: flexible manufacturing
Permanent link to this article: http://www.apriso.com/blog/2016/04/how-the-vss-unity-is-bringing-disruptive-technology-to-space/
Mar 22 2016
Adaptability is Transforming Robotics
Permanent link to this article: http://www.apriso.com/blog/2016/03/adaptability-is-transforming-robotics/
Mar 15 2016
Automotive NPIs Keep Accelerating – Can You Keep Up?
Permanent link to this article: http://www.apriso.com/blog/2016/03/automotive-npis-keep-accelerating-can-you-keep-up/
Mar 08 2016
Is the Automotive "Factory of the Future" Closer Than We Think?
The future might be closer than you think – here is a glimpse at what one automotive manufacturer has in planned for their next plant opening.
Permanent link to this article: http://www.apriso.com/blog/2016/03/is-the-automotive-factory-of-the-future-closer-than-we-think/
Permanent link to this article: http://www.apriso.com/blog/2016/02/the-advantages-of-procrastinating-in-a-fast-paced-manufacturing-world/


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Hotfrog Ads
Is Aussie Seafood House incorrect or missing key business information? Update your Hotfrog listing today.
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Hotfrog Australia provides information regarding Aussie Seafood House in Capalaba QLD. Aussie Seafood House is located at 3230 Old Cleveland Rd, Capalaba QLD 4157 and provides Food & Beverage Products services. Contact them on (07) 32453003.
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California - In Stock
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Orders over $59 ship free in the
continental United States.
A few products do not qualify for free shipping and are designated with "Does Not Ship Free" in the product detail. Those products are charged actual shipping charges. View Details
If you are unsatisfied with any item for any reason you can return the product within 60 days and we will refund your purchase price on that item, excluding shipping charges. View Details
For the fast delivery we ship from CA and PA warehouses! You will see exact arrival dates at checkout. View Details
Ideal for the weekend get away!
This kit is ideal for bottle conditioning your keg with 2oz of corn sugar at room temperature for about a week before you take your homebrew on a road trip.
Or you can use our jumper line and transfer clear beer from your 5 gallon keg to your 2.5 gallon keg leaving all the yeast behind. This way when you go to the party your beer is serving sparkling clear.
Use the C02 injector to dispense beer when ever you need more co2 pressure to serve.
A little trick: Get yourself a 5 gallon rubbermaid cooler. This keg fits like a glove inside. Pack the cooler with ice all around and you'll have cold beer for a couple days.
Another trick is to drill a hole in the cooler lid and feed the beverage line assembly through it (you will have to unscrew a hose clamp...not too hard.) You can arrange somthing similar for the C02 injector as well.
Be gadgety! Be a homebrewer!
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Comfortable and stylish, this outdoor swivel glider chair provides you with a place to enjoy fresh air and sunshine. The swivel base makes side to side movement easy while the gliding feature allows you to gently move back and forth for a relaxing movement. Reversible cushions always keep this piece looking great and fresh. Create a space in your yard where you can entertain and relax with this swivel glider chair.
The Cassley Swivel Glider Chair with Reversible Cushion from Rotmans. We proudly serve the Worcester, Boston, MA, Providence, RI, and New England area. Product availability may vary. Contact us for the most current availability on this product.
Create an outdoor oasis with furniture for your yard, patio or deck from this collection. Add a sofa and loveseat combination to create an outdoor living room perfect for entertaining or relaxing alone. A sun lounger is fun way to add comfortable seating to your outdoor space. Pieces from this collection feature hand-woven all-weather wicker for a look that will last.
The Cassley collection is a great option if you are looking for Casual
Coastal Outdoor Furniture in the Worcester, Boston, MA, Providence, RI, and New England area.
These images are representative of this group as a whole and may differ from this item.
Browse other items in the Cassley collection from Rotmans in the Worcester, Boston, MA, Providence, RI, and New England area.
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Tropical Fish Keeping - Aquarium fish care and resources (http://www.tropicalfishkeeping.com/forum.php)
- Introduce Yourself (http://www.tropicalfishkeeping.com/introduce-yourself/)
- - New here (http://www.tropicalfishkeeping.com/introduce-yourself/new-here-114834/)
New here
Hi everyone! I just joined this forum. I love aquariums and hope to learn and share with others. :)
Hello and welcome to the forum! :-D
Bonjour et bienvenue
Welcome to TFK :-D
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In four novels and a collection of short stories, Leïla Marouane has become a voice for the Algerian women's rights movement, exploring themes of marriage, sex, and identity in the context of the religious and cultural divide of the Maghreb/Western Europe region. She fearlessly takes on the taboo, as her skill with comedy renders even the most troubling political or religious issues accessible. Born in Algeria in 1960, Marouane escaped persecution towards her writing by moving to Paris in 1990. This is her second novel to be translated into English, following The Abductor, which was published by Quartet Books in the UK almost a decade ago.
With The Sexual Life of an Islamist in Paris—vividly translated by Alison Anderson—Marouane skillfully constructs a light, comedic plot behind which hides a dizzying maze of questions that, like an Escher staircase, form an endless loop. The story starts out as a comedy of errors starring Mohamed Ben Mokhtar, an Algerian immigrant living in a Paris suburb who is also a forty-year-old virgin. Once a devout Muslim, Mohamed has left the faith and decided to free himself from his oppressively devoted mother by moving to Paris and Westernizing himself. He changes his name to Basile Tocquard, lightens his skin and straightens his hair, and sets out to become a famous poet and seduce as many blonde women as he possibly can. The results are hilarious, as in this great passage in which Mohamed naively plans out his new apartment:
I determined where the bookshelf would go, along with the desk, facing the window, above the foliage that would inspire poetry to make Antonin Artaud and Octavio Paz week in unison in their graves, and then of course the bed, maybe a palatial king-size, where I would roll about with creatures to tempt angels and demons alike; a bathroom in tones of green and yellow, two sinks side by side, an oval tub that could easily seat two adults and into which I would plunge each of my future conquests and myself along with them; a separate toilet with a bookshelf that went right to the ceiling, where I would place my collections of Diplo and Politics, my graphic novels, and the girlie magazines I intended to acquire . . .
However, Basile (also called Mohamed, and sometimes Momo) is something of a paradox: although he claims that he only has eyes for Western women, the women he becomes involved with are all Muslim, Algerian, and just the sort of woman his mother would approve of. Each of these women inevitably thwarts Mohamed's plans to bed them, leaving him continually frustrated. As time wears on and Mohamed's "conquests" multiply—he is always certain that a wild night of sexual abandon is right around the corner—we begin to question Mohamed's reliability as a narrator. He starts to lose chunks of time, staying up all night entertaining his girlfriends—or does he?—and sleeping all afternoon. Then, suddenly, he jumps ahead three months as though it were the next day. A mysterious writer, Loubna Minbar (or Louisa Machindel, or is it Lisa Martinez?) appears as a common link between everyone Mohamed meets. A manuscript Mohamed believes to be written by her appears in his apartment, throwing the reality of the events of the entire story into question.
For the unobservant reader, it would be easy enough to miss the clues that Marouane—who coincidentally shares the same initials as the mysterious writer—has sprinkled throughout the story, details that seem too murky and puzzling for such a lighthearted, frivolous story. Even so, it would be impossible to escape a growing suspicion that Mohamed is not really the narrator but the protagonist in someone else's story. Each chapter begins in the other writer's voice, saying "he said," before returning to Mohamed's narrative; every so often Mohamed addresses an unnamed figure in the second person; and he is reading a novel called The Sultan of Saint-Germain that seems to be based on his cousin's life, or even his own. The central mystery revolves around the chameleon-like Loubna Minbar, who may or may not be Mohamed's concierge, and who exacts her revenge on men by stealing their life stories for her novels and driving them mad in the process. Because, while Marouane's novel masquerades as a man's story, it's really a story about women, about the countless Algerian women who have had to make the humiliating journey to freedom in the Western world on the currency of their wits and their bodies. Ignored by Mohamed in his quest for sexual liberation, they become the heroines of the story, enjoying the same freedom that eludes him. As their stories cleverly illustrate, it may be a man's world, but women get the last laugh.
Like its title, The Sexual Life of an Islamist in Paris intends to provoke. And if you're not careful it may also send you in circles, trying to get to the bottom of that staircase. As one of Marouane's characters warns Mohamed, "Just reading her is enough to send you round the bend."
Prose translators will likely disagree, but I believe translating poetry requires a significant level of talent, a commitment to the text, and near mania, all of which suggests that the undertaking is the greatest possible challenge. The task is to. . .
The biggest issues with books like The Subsidiary often have to do with their underpinnings—when we learn that Georges Perec wrote La Disparition without once using the letter E, we are impressed. Imagine such a task! It takes a high. . .
Following The Infatuations, Javier Marías's latest novel seems, like those that have preceded it, an experiment to test fiction's capacity to mesmerize with sombre-sexy atmospheres and ruminative elongated sentences stretched across windowless walls of paragraphs. Thus Bad Begins offers his. . .
Death by Water, Kenzaburo Oe's latest novel to be translated into English, practically begs you to read it as autobiography. Like The Changeling, as well as many other works not yet released in English, Death by Water is narrated in. . .
Jocelyne Saucier's Twenty-One Cardinals is about the type of unique, indestructible, and often tragic loyalty only found in families. For a brief but stunningly mesmerizing 169 pages, Twenty-One Cardinals invited me in to the haunting and intimate world of the. . .
We know so very little; so little that what we think to be knowledge is hardly worth reckoning with at all; instead we ought to settle for being pleasantly surprised if, on the edge of things, against all expectations, our. . .
Many of Virginie Despentes's books revolve around the same central idea: "To be born a woman [is] the worst fate in practically every society." But this message is nearly always packaged in easy-to-read books that fill you with the pleasure. . .


text: hi how are we all.. ok ive got a few questions if anybody is able to help me out.. i noticed yesterday my Geophagus brasiliensis has had babies. i noticed them moving them from the rock into the pot i have set up, what i noticed was all white stuff under the rock which im assuming is the spawn ??? the mother is very good, father is good aswell how ever he keeped taking them out of the pot mother putting them back in, then she guarded them all night from the father lol, today she has just now moved them out in the open, i cant tell how many there are coz they are sooo sooo tiny.. the question is: do i leave both parents in the tank with them , the tank is a 6 foot spilt into 2 section so there pretty much 3 foot room with plants pots rocks etc in there.... will the parents harm the young babies at all ??? what do i feed the young ones or when do i start feeding them, i dont even know how old they are as stated only first noticed them yesterday when parents started moving them. any help would be great as im new to this whole breeding thing.... kind regards hope to hear from someone soon


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Barack Obama, our next President
There are 1496585 comments on the Hampton Roads Daily Press story from Nov 5, 2008, titled Barack Obama, our next President. In it, Hampton Roads Daily Press reports that:"The road ahead will be long. Our climb will be steep," Obama cautioned. Young and charismatic but with little experience on the national level, Obama smashed through racial barriers and easily defeated ...
Join the discussion below, or Read more at Hampton Roads Daily Press.
#900687 May 10, 2013
Newly released emails are raising additional questions about the Obama administration's story-line on the Benghazi attack, showing top-level efforts to downplay terrorism ahead of public statements by UN Ambassador Susan Rice.
Since: Apr 13
#900688 May 10, 2013
Guess the UN was a bunch of dopes too. Huh?
Saint Petersburg, FL
#900689 May 10, 2013
You have to admit Realtime - Obama, Hillary and Susan Rice do what they do best LIE... AND YOU TOO!!
#900690 May 10, 2013
Benghazi: Fox News' distraction
If you havent yet seen Wag the Dog, go watch it right now. Ill wait. And dont worry, this article isnt actually about Benghazi, but something that really matters.
The movie is about how the art of manipulation is used by the media to distract people from events of actual importance. It was cited regularly when the GOP had their Clinton witch hunt. The GOPs latest witch hunt over Benghazi is simply more wagging of the dog. Fox News is trying really, really hard to make it into Obamas Watergate to keep us all distracted from some incredibly populist moves in the Senate.
#900691 May 10, 2013
Will Fox Correct The Record On The GOP's False Benghazi E-mail Claims?
The New York Times, however, obtained a copy of the e-mail in question and reported that the phrase conservatives are putting in the mouth of the State Department official -- "Islamic terrorists" - is in fact not used to describe the perpetrators of the attack. Rather, the official describes the perpetrators as having ties to "Islamic extremists" -- a distinction with a difference, according to the Times report:
[A] copy of the e-mail reviewed by The New York Times indicates that A. Elizabeth Jones, the senior State Department official who wrote it, referred to "Islamic extremists," not terrorists.
The distinction is important, administration officials said, because while the White House did not initially characterize the attack as terrorism, senior officials, including Ambassador Susan E. Rice, acknowledged the possibility that extremists had been involved in the assault.
Fox News is no stranger to carrying water for the Republican Party, and the network has led the charge to push Benghazi cover-up conspiracies. But now that the latest GOP line on Benghazi has been exposed, will Fox inform its viewers?
Saint Petersburg, FL
#900692 May 10, 2013
If NOT for obama, Benghazi would NOT be an issue you idiot. It was obama's war in libya you fool...
Since: May 11
#900693 May 10, 2013
So, you think social security & Medicare work the same way??
#900694 May 10, 2013
No, the truth.
#900695 May 10, 2013
Obama lies two weeks before Benghazi:
Al-Qaeda is on the run and Bin Laden is dead!
Mission accomplished!
How wonderful Obama followed the French into Libya!
Reality and happening today:
The U.S. military has alerted two elite military units in Europe to be on standby if needed to respond to a deteriorating security situation in Tripoli, Fox News has learned.
Read more: http://www.foxnews.com/politics/2013/05/10/us...
#900696 May 10, 2013
Eman said that Obama was on the golf course "during" the raid and that was a lie. I don't get the need to lie__particularly when most of the posters her know it's a lie.
I guess if that's all one has__one would be better off over on the game threads.
Since: Apr 13
#900697 May 10, 2013
This is what really got me about Carney's tap dance this afternoon.
He tried on several occasions to get the heat off himself and this administration by eluding to Republicans "politicizing" Benghazi.
I was literally willing someone in the media to ask, "Wasn't it this administration who politicized Benghazi?"
Unfortunately, they didn't Sure wish I had been there as part of the press.
Saint Petersburg, FL
#900698 May 10, 2013
You have to admit sonicfiller - Obama and Hillary do what they do best LIE.
#900699 May 10, 2013
You mean the thesis approved by Harvard that points out uncomfortable facts about illegal immigration?
#900700 May 10, 2013
the UN?
The Bush administration yesterday said it would not wait for the UN security council to approve an attack on Iraq....
i think the next illogical FOXbot move would be to try to lay the Iraq war solely at the feet of the democrats who voted for it. never mind all of the republicans who voted for it, just stick to the FOX propaganda meme and you'll do just fine.
Since: May 11
#900701 May 10, 2013
You are really pathetic. You lied. You know it.
You knew who Obama met with yet you implied he met with just some officials. You thought that meeting was at 11PM DC time.
You know it.
#900702 May 10, 2013
The entire Obama Libyan operation was a "Wag the Dog" plan to divert attention from the rotten economy.
#900703 May 10, 2013
Hey thread propaganda Queen!
How nice of you to vindicate Richard Nixon.
Watergate must have also been a distraction due to your libtarded logic.
#900704 May 10, 2013
Immigration activists are praising the Heritage Foundation for accepting the resignation of researcher Jason Richwine, who argued in recent years that groups of non-white immigrants have inherently lower IQs that could make it impossible for them to adequately assimilate in America. Heritage distanced itself from his past statements, but stood by his work co-authoring a recent study claiming immigration reform would cost $6.3 trillion in large part because undocumented immigrants would fail to improve their education or income levels.
We welcome Mr. Richwine's resignation, but are still waiting for the Heritage Foundation to renounce the findings of his research, which like his claims about Latino IQ's are false and unbecoming of a research institution like the Heritage Foundation, Arturo Carmona, executive director of Latino advocacy group Presente.org said in a statement.Hiring racists like Richwine is unacceptable, but whats worse is putting their organization's name on a report laden with faulty research.
Journalist-turned-activist Jose Antonio Vargas also released a statement through his immigration advocacy group Define American.
Racism and xenophobia have no place in the debate on immigration reform, period," Vargas said. "I hope this is a lesson for all sides that what the public is looking for is a fair and honest debate on immigration reform, not long discredited racial theories designed divide us rather than unite us.
"Often imitated"
Since: Jul 07
never duplicated
#900705 May 10, 2013
He had 2 days to reschedule his golf game, stupid.
#900706 May 10, 2013
It was a video!
Lies, Liars/Democrats.
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Ortho, ortho trisubstituted benzene and pyridine-based synthons and building blocks. Pharmaceuticals that contain this motif include Entacapone, Dasatinib, Levobupivacaine, Treprostinil, and Fenoldopam.
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Josie Loren
The Cuban actress tends to steer towards minidresses and skirts that show off her great legs (um, can you blame her?). Whether going with a pop of color in fuschia or keeping it understated in a LBD (little black dress), this star rarely misses a (style) step.
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At Jl. Bypass Ngurah Rai No. 120F, Simpang Siur – Kuta, Bali beside of Harley Davidson in front of Mall Bali Galeria.
There are various Packages Monthly Quota for SMART OFFICE tailored to your business needs.
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Get other interesting bid from us for Monthly Quota Package with Bonus Using of Meeting Room, LCD Projector and creation of Website Marketing ter-SEO (Sarch Engine Optimization).
C STAR VOUCHER is available FREE of CHARGE to the Tenant Office, Affiliate Partner, Business Partner, Associate Partner, Member Club and Member Group Company of C STAR in Indonesia and overases.
C STAR VOUCHER is a Voucher Discount given by merchant, the object tourism, vendor, partner etc and we provide free of charge for you that should be the full rights of tour guide or tour & travel.
We also present you with Meeting Room which has capacity of 30 people with a very affordable fee that is only IDR 50.000,-/head/hour or only IDR 11.000/head/hour for The Member Club Meeting at high speed of Free Wi-Fi and also complimentary drink such as mineral water, tea or coffee.
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C STAR will issue 27 franchises untl the end of the year 2017 outside Member Group Company of C STAR.
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The Worcester Chorus Sings Rossini & Brahms
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The glorious sounds of the Worcester Chorus, in concert with the newly-restored Hook organ, bring a Rossini mass to stirring life in the ensemble's latest performance, which also boasts a four-handed piano piece and a quartet of fine guest vocalists. Rossini may've been best known for his 39 operas, but he also produced sacred music, like this melodic Petite messe solennelle, towards the end of his life. The acclaimed chorus, led by artistic director Christopher Shepard, is also set to sing Brahms' Nänie, based on a mournful poem by German author Friedrich Schiller. Soprano Kathleen Theisen, alto Elizabeth Russo, tenor Thomas Mooney and bass Gregg Lauterbach join the ensemble as soloists, as do pianists Olga Rogach and Sima Kustanovich and organist Malcolm Halliday. There's a pre-concert lecture one hour prior to the performance.
* Additional fees apply. No coupon or promo codes necessary to enjoy the displayed discount price.
All offers for Worcester Chorus: Rossini & Brahms have expired.
The last date listed for Worcester Chorus: Rossini & Brahms was Friday March 7, 2014 / 8:00pm.
Currently at Mechanics Hall
Vivaldi's The Four Seasons by Venice Baroque Orchestra
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One of the world's premier period-instrument ensembles, the Venice Baroque Orchestra is famed for… More
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Running, a Love Story
I would try and click-bait you, but I'm bad with titles so this is what it's going to be. (Side note: the photo above is the trail I run on just about every morning.)
This month marks 11 years I have been running seriously. I would go out and run around my neighborhood in junior high and run in gym class and run during other sports, but August 1st, 2003 I went to my first high school cross-country practice as a baby freshman. I wish iPhones existed then so I could pull some old Instagram photos from that time. Anyway, running three miles seemed DAUNTING to me. Flash forward 11 years, three miles are junk miles to me*.
*this isn't to say three miles are junk miles – if that's all you can/want to run, be proud! You should be proud of any amount you run – you're still beating everyone on the couch.
I have ran through it all: most notably, ligaments ripping away from my knee cap, years of physical therapy, wearing a knee brace, running on insanely taped-up sprained ankles – things I don't talk about a lot because I've forgotten about them. It may be why I have such a laissez-faire attitude towards injuries – I know they can (usually) always be overcome. I obviously really love something if I'm still out there, pounding out 30 miles a week.
Running has gotten me through every break-up of my life, fights with my friends and parents, deaths, career insecurity – it's been my longest relationship and has definitely been the best therapist I've had. I can't tell you how many tears I have shed before, during and after runs when it has had nothing to do with running.
Running a marathon is something I have toyed around with for over a year now. I decided right before my birthday that this year was the year. "You have to want to run a marathon," is what Emily told me. I finally realized that half-marathons were no longer a challenge for me – I could run a half-marathon drunk. That day basically told me I was pretty fit and I needed to challenge myself more.
I love running for the same reasons I love surfing or yoga: because I love pushing my body to the limit. I want to see what I'm capable of. That's why this November, I'll be running my first full-marathon, the Savannah Rock n' Roll Marathon.
I've ran Savannah's half-marathon before. It was probably one of the worst half-marathons I've ran because I was so out of distance running shape. The farthest I had run prior to that day in 2011 was 7 or 8 miles in over a year and a half. I had such an awful race I decided half-marathons weren't for me and I backed off running for a little over a year, just doing maybe 15 miles a week, max.
In the past two years, I've seen how far I have come as a runner. If I don't love running full marathons, at least I'll have done one and I can stop feeling like a fake distance runner for never running a full. I know that I'll always have 13.1.
As a note, I'm also excited for three other races I have coming up: Montour Trail Half-Marathon the first weekend in September (it fit in perfectly with my training schedule – I'm doing Hal Hidgens), Ragnar DC the second weekend in September and the Key West Half-Marathon (as an official blogger!) in January. The Key West race will be good because after I run the full, I can take a few weeks off then have another race coming up so I can't get lazy.
Chat with me:
What are some goals you have for yourself? Have you ever run a marathon? What thing (not a person) do you a love affair with?
45 comments on "Running, a Love Story"
I'm excited you are challenging yourself even more and going for the full! I have no doubt you will be fantastic and have a great race! I love everything you said about what running is to you; that's exactly what it is for me, too! best therapy out there.
Heather @fitncookies recently posted…I mustache you a question…
Key West race should be so fun. Congrats on everything
Ang @ Nutty for Life recently posted…Thinking Out Loud: Weird Edition
Ahhh I can't wait for you to run your marathon! My running story started very similarly to yours. I didn't like it all that much and now I absolutely love it. SO COOL that you're going to Key West as a blogger!
Sarah recently posted…Baltimore Tourists
TELL ME ABOUT KEY WEST! Take me with you as your official cheering squad!!! But actually this post was beautiful. I used to feel the same way about running but not anymore. Started to do more harm than good for ME but different strokes for different folks so instead you can run it for both of us and I'll be cheering you on the whole way!!
Davida @ The Healthy Maven recently posted…Summer Lately
i'll be a sherpa! can i come? and i agree.. such a beautiful story and relationship with running
Lindsay recently posted…5 Food and Fitness Tips and Tricks
Wow this is awesome! I love that you are creating challenges for yourself and have found running as an outlet. While you are onto 11 years of running, I am onto Month 2 (lol) and definitely enjoying how good it feels to pound out a mile or two. I hope to run a 5K in October and a 10K in the distance after that.. you inspire me to just let my feet pound it out and let my brain have alone time. Can't wait to follow you through your training plan, races, and the big day !!
Marisa @ Uproot from Oregon recently posted…Green Julius Smoothie
good luck! i'm totally in awe of anyone who can run more than 3 miles – that's my limit :)
Lindsay @ The Lean Green Bean recently posted…Bean Bytes 102
Well, Lindsay, I would say that you have a pretty decent excuse!!! lol
Susie @ SuzLyfe recently posted…Run, Sleep, Eat, Repeat #MIMM
I definitely support and encourage runner to attempt running a marathon–it gives you a whole new respect for your body and the sport that you never can comprehend until you try it. It's like going through pregnancy–it just changes the way that you perceive what you are capable of!
Susie @ SuzLyfe recently posted…Run, Sleep, Eat, Repeat #MIMM
That's a beautiful running trail. We just moved from a creepy neighborhood (as in, took my dog for safety every time) to a nice, safe area. The peace of mind when going out for a run is completely liberating!
I have a love affair with cats. I grew up afraid of them, then I decided I wanted one. Now we have two. I would get 20 more if my husband would allow me.
Jessica recently posted…9 Ways to Write Attention-Grabbing Post Titles
Congrats on the decision to run a marathon!! Goals for myself are to try CrossFit and increase Bar Method from 4x weekly to 5.
Charlotte @ Commitness to Fitness recently posted…If Socrates Had Been A Blogger (Thinking Out Loud/ Top 8 List)
I'm running the Nike Women's Half Marathon in SF in 2 months!! Good luck with your marathon. If you have any training tips, I would love to see them on your blog (because I have no idea what I'm doing).
Lisa recently posted…Vegan Banana Nut Muffins
Also, props to you girl for being able to power through a half-marathon drunk! I definitely can't do that!
Lisa recently posted…Vegan Banana Nut Muffins
Marathon #2 in 2015. Already paid and registered, so no looking back! I did my first one when I was 30 and I actually hated it. I took me over 6 months to realize I wanted to try again. I have a love affair with food. I truly do. xoxo, ganneban
ganeeban recently posted…Rose & Thorn [13]
Ahhh how cool! You're gunna rock it. You and running are obvi ready to take things to next level ;)
I <3 running :)
Matt @ Runner Savvy recently posted…Back At It
You are freaking amazing girl! I'm seriously inspired :) Can I just say that I NEVER run and I decided to hop on the tread and run 3 miles yesterday….my whole body is in pain. I'm sore in places I didn't know existed! Gotta take it slow next time :)
Alexis @ Hummusapien recently posted…In My Grocery Bag: Trader Joe's
Big props buddy- I despise running and even a km is a success in my book!
Arman @ thebigmansworld recently posted…Marinated Pesto Potato Salad
Ahhh how exciting! I trained for a half marathon last fall, got up to 12 miles and backed off because I was too scared to be limping around on my wedding day. Now after having been injured for over 6 months I want to get back out there and I desperately want to sign up for a half marathon….not sure if I'm being overly cautious holding back
Kelli @Hungry Hobby recently posted…4 Mile Run and Weekend Things
Awwwr love it, lady. I really wish I could develop that kind of love of running, buuuuuuuut 3 miles and I hit a wall. That being said, I do have other fitness loves that totally do that for me, so I know how it is :) I"m excited for youuuuuuuu!
Amanda @ .running with spoons. recently posted…. vegan chocolate chip cookie dough {bites, bars, spoonfuls} .
[…] I love running. Why would I ever be jealous of non-runners? People are "jealous" of my will-power, my […]
[…] Well, this would have been a useful resource for me if I were still running a marathon next month but for the next time, I'm going to use Lesley's Marathon Packing […]
[…] my first marathon this year, running has defined me for over the past decade. I have days where I love running so much it's gross and I days where I wish I had never laced up a pair of shoes in my life. […]


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The Life and Legacy of Hank Williams
An audio timeline through the short life of Hank Williams who was born exactly 90 years ago.
Country and Americana artists talk about his legacy and why he's still relevant today. Featuring contributions from Emmylou Harris, Rodney Crowell, Willie Nelson, Duane Eddy, The Mavericks, Gretchen Peters, Darrell Scott, The Cowboy Junkies, Aoife O' Donovan and Glasgow band Woodenbox.
Ricky Ross looks at why "I'm So Lonesome I Could Cry" is one of the most powerful and covered songs of all time. Featuring an exclusive re-working of the track, in session, by New York singer-songwriter Tift Merritt.
Hank Williams died at the age of 29. During his lifetime he recorded 129 songs but left a legacy of over 700.
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Hank Williams - Cold Cold Heart
This video clip of Cold Cold Heart juxtaposes the sad story behind the song with a light hearted TV performance.The clip is taken from the BBC Arena documentary Hank Williams: Honkey Tonk Blues.


text:
Chilling and thrilling
Soaking, spilling over me
A spring time shower
Blue skies and white clouds
Win or lose the oarsmen grunt
Henley Regatta
Wet maple leafs fall
Reds, rusts, browns and yellows
Photographers click
Short and chilly days
Scarves, mittens and woolly hats
Children wish for snow
By Dibs
© Copyright 2017 dibbledabble. All rights reserved.
Status: Finished
Genre: Poetry
I've got nothing terribly witty to say (what do you mean, "You're used to it?", excellent!
These are delightful, Dibs - and you have a new format for your thoughts, your writing! Way to go! Way nice.
Warm regards, Connie
Some lovely thoughts Dibs. Good write.
i applaude you these are marvelous ... i have tried this style and slunk away again you show such skill very very good these are
4 seasons....68 syllables....1 like....very cool
nice job Dibs ...
Very nice. I love Haiku. I hope you'll read mine as well. It's called Marvin Haiku. Thanks. Peace.
Book / Romance
Book / Fantasy
Book / Children Stories
Paste the link to picture in the entry below:
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Love for Sale
The Graduate
Pimpmaster General
It's A Family Affair
The Graduate
Duration: 9m50s / Size: 28.5 MB
Pimpmaster General
Duration: 8m10s / Size: 20.2 MB
It's a Family Affair
Duration: 10m38s / Size: 31.4 MB
Love For Sale
Ten documentary films made for the BBC. Based in two brothels just outside the city limits of Carson City, Nevada; The Moonlight Bunny Ranch and Madam Kitty's Cathouse. Nevada is the only state where prostitution is legal. These documentaries challenge the prejudices associated with prostitution. There are no drugs and dark streets. These working girls are hip and glamorous.They live in the desert, under bright blue skies and work in air-conditioned parlours. Jane stayed at the brothels for two years filming and getting inside the lives of the girls, madams and their clients.
© 2017 Jane Hilton. All rights reserved. |
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hey everyone, I'm another one of those Radiohead fanboys that was turned on to Max/MSP from the 'legendary' vid of Jonny going crazy during his Go To Sleep solo (http://www.youtube.com/watch?v=xyKCwDJ7ZUc). I've been trying to find a kind of glitch code for that and stumbled upon this…however it doesn't work. I have found that this patch is not written in the same format as other ones ive found. i was wondering, could anyone tell me what is wrong with it? it's attached in a document. I have Max in Realtime btw and would just like to utilize this kind of patch for live guitar solos, etc. etc. I was wondering if this was obsolete code or there are some fundamental flaws im missing? And how the fixed code looks exactly..
im sure you guys are annoyed by us programming n00bs. but if u could all really help me, id really really appreciate it. sorry if my english is bad its not my first language (or even second haha)
these are 2 max text patches in one text file, that wont open in maxmsp unless you
make 2 textfiles out of them.
the line
#P pop;
always closes a max text file, that is where oyu have to split it.
Ah I see. However it still doesn't work. It says I'm missing paranthesis and stuff. damn! hahaha
Hi, Yes, still works and very interesting and cool patch ! :
just move away "— fridgebuzz_off" in the last line , and open from Clipboard
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