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Erythropoietin (EPO) regulates the production of red blood cells primarily by preventing apoptosis of erythroid progenitors. More recently, however, EPO has emerged as a major cytoprotective cytokine in several nonhemopoietic tissues in the setting of stress or injury. The underlying mechanisms of the protective responses of EPO have not been fully defined. Here we show that EPO triggers a phosphatidylinositol 3-kinase-(PI3K)-dependent survival pathway that counteracts endothelial cell death. The protection conferred by PI3K relies on the subsequent induction of Bcl-xL, a prosurvival member of the Bcl-2 protein family. In addition, EPO counteracts the upregulation of the pro-apoptotic BH3-only protein BIM, which is induced by serum withdrawal. EPO also activates extracellular signal-regulated kinase 1 and 2 (ERK1/2), which are involved in a Bcl-xL-independent cytoprotective pathway. EPO caused a prolonged activation of nuclear factor (NF)-κB, which was blocked by inhibition of PI3K, but not by inhibition of mitogen-activated protein (MAP)/ERK kinase (MEK), suggesting that EPO-activated NF-κB requires PI3K activity. However, the activation of the NF-κB pathway was not required for the ability of EPO to counteract endothelial apoptosis. Thus EPO promotes survival of endothelial cells through PI3K-dependent Bcl-xL-induction and BIM regulation, as well as through a separate mechanism involving the ERK pathway.
- human umbilical vein endothelial cells
- apoptosis
- extracellular signal-regulated kinase
- pro-apoptotic BH3-only protein
- Bcl-2
erythropoietin (EPO) is a glycoprotein hormone which has a prominent role in erythropoiesis. EPO is produced mainly by the adult kidney in response to hypoxia. Although, it has been documented to effectively promote survival, proliferation, and differentiation of erythroid progenitors, its role is to prevent apoptosis of late erythroid progenitors, which is now considered the primary mechanism of EPO action (46). The anti-apoptotic function of EPO has been the subject of many studies performed in the past few years (20, 24). Recent findings indicate that the anti-apoptotic activity of EPO is not restricted to erythroid progenitor cells but extends to other tissues such as the brain, retina, spinal cord, peripheral nerves, kidney, heart, and vasculature (3, 10, 11, 17, 26, 43, 45). EPO has therefore emerged as a remarkable anti-apoptotic cytokine against potential stress or toxic stimuli.
With respect to the vasculature, EPO acts on both endothelial and smooth muscle cells. EPO receptors are expressed on human vascular endothelial cells from coronary, pulmonary, and cerebral arteries, as well as umbilical vein and dermal vessels (2, 4). The endothelium acts as a barrier and interface between blood and tissue and provides the route for systemically administered EPO to pass into organs. It is therefore important to define the effect of EPO on the endothelium.
Several signal transduction pathways, including the phosphatidylinositol 3-kinase (PI3K) and extracellular signal-regulated kinase 1 and 2 (ERK1/2) pathways, are known to be involved in the anti-apoptotic functions of EPO in erythroblasts (9, 21, 28, 46). In contrast, EPO-mediated neuronal survival involves cross talk between the nuclear factor (NF)-κB and Janus kinase 2 signaling pathways (15). However, the major anti-apoptotic pathway in endothelial cells has not been elucidated. In light of this tissue-protective function of EPO, we set out to determine the mechanisms signaling survival that are evoked by EPO in endothelial cells. Our results show that EPO inhibits apoptosis in endothelial cells by PI3K-mediated upregulation of Bcl-xL and by an independent mechanism involving the activity of the ERK pathway. In addition, EPO represses the induction of the pro-apoptotic Bcl-2 family member BIM in response to serum withdrawal. Our findings implicate a critical role for EPO in balancing Bcl-2 family member expression to prevent cell death.
Human recombinant EPO was purchased from Roche Applied Science (Laval, QC, Canada). Rabbit anti-Akt, anti-phospho-Akt (Ser-473), anti-Bcl-xL anti-phospho-ERK1/2 antibodies, LY-294002, and U0126 were from Cell Signaling Technology (Beverly, MA). Antibodies recognizing ERK1/2 were purchased from Stressgen (Victoria, BC, Canada). Lipofectin, human endothelial serum-free medium (SFM) basal growth medium, Opti-MEM I, l-glutamine, streptomycin, and penicillin were purchased from Invitrogen (Carlsbad, CA). MCDB131 medium and heparin came from Sigma (St. Louis, MO). Recombinant tumor necrosis factor-α (TNF) was purchased from R&D Systems (Minneapolis, MN). The NF-κB inhibitor SN50 and the control peptide SN50M were obtained from Biomol (Plymouth Meeting, PA). The dual-luciferase reporter assay kit and the Renilla luciferase control reporter plasmid pRL-CMV were purchased from Promega (Madison, WI). The NF-κB reporter plasmid pNF-κB-luc, which contains five tandem NF-κB-binding sites upstream of the luciferase gene, was a gift from Dr. Frank Jirik (University of Calgary, Calgary, AB, Canada).
Cell culture.
Human umbilical vein endothelial cells (HUVEC) were obtained from collagenase-digested umbilical veins and cultured in MCDB131 medium supplemented with 20% heat-inactivated bovine calf serum (HyClone Laboratories, Logan, UT), endothelial cell growth supplement (20 μg/ml, Calbiochem, La Jolla, CA), heparin (90 μg/ml), l-glutamine (2 mM) in the presence of penicillin (50 U/ml), and streptomycin (50 μg/ml) in a 5% CO2 environment at 37°C as previously described (27).
Transient transfection of plasmid vectors was performed with Lipofectin in accordance with the manufacturer's instructions. Briefly, 60,000 cells were seeded per well of a 24-well plate 1 day before transfection. For each well, 4 ng of the plasmid pRL-CMV and 210 ng of pNF-κB-luc were diluted to 20 μl with OPTI-MEM I. One microliter of Lipofectin was diluted to 20 μl with OPTI-MEM I. Both solutions were incubated for 45 min at room temperature and mixed. After 20 min, 160 μl OPTI-MEM I was added and the mixture added to cells. After 3 h, the transfection mixture was replaced with human endothelial-SFM basal growth medium containing 0.5% calf serum. Sixteen hours later, cells were used for various experiments. Cell extracts were prepared, and the relative luciferase activity was determined according to the manufacturer's instructions.
Western blot analysis.
Two hundred and fifty thousand cells were seeded per well of a 12-well plate. After attachment, the growth medium was replaced with MCDB131 medium containing 1% calf serum and 0.5% bovine serum albumin. Sixteen hours later cells were subjected to various treatments. Cells were washed twice with cold phosphate-buffered saline and lysed in SDS sample buffer containing 2% SDS and 100 mM dithiothreitol. Samples were sonicated and electrophoresed on 10% polyacrylamide gels. After transfer of proteins to nitrocellulose, membranes were blocked with 5% nonfat milk and incubated with various antibodies according to the manufacturer's instructions. Detection was performed using horseradish peroxidase-coupled secondary antibodies and SuperSignal West Pico chemiluminescent system (Pierce, Rockford, IL).
Viability assay.
For determination of viability, HUVEC were seeded at a density of 10,000 cells per well of a 96-well plate. After attachment, cells were switched to human endothelial-SFM basal growth medium containing 0.5% calf serum for 16 h and subjected to various treatments. Medium was removed and replaced with medium containing 0.0025% neutral red (Sigma) and incubated for 4 h. The medium was removed, and after solubilization with 1% acetic acid in 50% ethanol, the absorbance at 560 nm was determined for each well.
Mitochondrial membrane potential.
The mitochondrial membrane potential was assessed using the fluorometric dye tetramethylrhodamine ethyl ester (TMRE). Cells were grown in 12-well plates at a density of 2.5 × 105 cells/well. The next day, cells were changed to human endothelial-SFM basal growth medium containing 1% calf serum in the presence or absence of EPO for 6 h. In some experiments, inhibitors (SN50, 50 μg/ml; LY-294002, 50 μM; wortmannin, 100 nM; U0126, 10 μM) were added 30 min before EPO exposure. TMRE (20 nM) was added during the last 30 min. Cells were harvested and analyzed by flow cytometry.
Statistical analysis.
Student's t-test was used for statistical analysis between two groups. Data are presented as means ± SE. Statistical significance was taken at a P value of ≤0.05.
EPO promotes endothelial survival.
HUVEC were serum-starved in medium containing 1% calf serum (CS) for 16 h. Cell viability was measured using the neutral red assay. As shown in Fig. 1A, the viability of cells grown in 1% CS decreased by 40% compared with cells grown in complete growth medium containing 20% CS. EPO treatment reversed the loss of HUVEC viability associated with CS withdrawal in a dose-dependent manner (Fig. 1A). To validate the effects of EPO, we analyzed other parameters of apoptosis. As shown in Fig. 1B, serum withdrawal also caused the dissipation of the mitochondrial transmembrane potential (Δψm). EPO prevented the loss of Δψm induced by serum withdrawal (Fig. 1B). We induced apoptosis in HUVEC using ceramide to determine whether the antiapoptotic effects of EPO were dependent on the death trigger. As shown in Fig. 1C, EPO also prevented the loss of viability triggered by ceramide, suggesting more widespread cytoprotective activity against various triggers.
EPO induces activation of PI3K in endothelial cells.
We next determined the pathways involved in the cytoprotective effects of EPO in HUVEC. We first determined the effects of EPO on PI3K. The serine-threonine kinase Akt is activated downstream of PI3K in response to a number of growth factors and is also known to induce potent cell survival pathways (18, 19, 23, 35). Activation of Akt involves dual phosphorylation on threonine at position 308 and serine at position 473 downstream of PI3K (1). Because of the absolute requirement of phosphorylation for Akt activation, we used a phosphospecific antibody that only recognizes the phosphorylated form of the enzyme to determine its activation. Figure 2A shows the time course of EPO-dependent Akt activation. Akt phosphorylation appeared within 5 min of treatment and remained elevated up to 60 min. The phosphorylation of Akt was also determined in the presence of a PI3K inhibitor. Pretreatment of cells with LY-294002 blocked the phosphorylation of Akt induced by EPO, confirming that Akt is a downstream target of PI3K in EPO signaling (Fig. 2B).
EPO induces activation of ERK1/2 in endothelial cells.
We next determined whether EPO activates the mitogen-activated protein kinase (MAPK) pathway in HUVEC. We focused on ERK1/2 MAPK because of their role in promoting cell survival (7, 25, 30). Activation of these kinases is dependent on dual phosphorylation on tyrosine and threonine, which is catalyzed by their upstream activator MEK (44). Accordingly, detection of phosphorylation of ERK1/2 is a convenient assay for ERK activation. Serum-starved HUVEC were treated with 5 U/ml EPO for various times, and Western blot analysis was performed using a phosphospecific ERK antibody. As shown in Fig. 3A, EPO induced phosphorylation of both ERK1 and ERK2 within 5 min of treatment. EPO-induced phosphorylation was sustained up to 90 min. In contrast, we did not detect phosphorylation/activation of other MAPKs such as p38 and c-Jun NH2-terminal kinase by EPO (data not shown). We next evaluated the effect of U0126, a MEK inhibitor on the EPO-induced phosphorylation of ERK1/2. U0126 completely diminished the ability of EPO to induce phosphorylation of ERK1/2 indicating the absolute requirement of the upstream activator MEK for activation of ERK1/2 by EPO (Fig. 3B).
EPO activates NF-κB in endothelial cells.
We also assessed the role of EPO in the induction of NF-κB in HUVEC. NF-κB transcriptionally activates multiple genes involved in cellular immune response, growth, and survival (33). In unstimulated cells, NF-κB resides mainly in the cytoplasm bound to inhibitory κB proteins. After stimulation, the inhibitory κB proteins are phosphorylated and then degraded liberating NF-κB to translocate into the nucleus to initiate gene transcription. Cells were transiently transfected with the reporter plasmid pNF-κB-luc, which contains five tandem NF-κB binding sites upstream of a luciferase gene, to quantitate NF-κB transcriptional activity. Twenty-four hours following transfection, cells were treated with various doses of EPO for 4 h, and luciferase activity was determined. As shown in Fig. 4A, EPO caused a dose-dependent increase in NF-κB activity. The activity was maximal at 5 U/ml of EPO, reaching a threefold induction compared with untreated cells. This activation was comparable to that achieved by TNF, a strong NF-κB inducer. To further assess the kinetics of NF-κB activation by EPO, a time course of activation was performed using 5 U/ml EPO. The activation of NF-κB by EPO was apparent within 2 h and reached maximal levels between 4 and 8 h. The activity remained elevated for 8 h and slowly declined by 10 h (Fig. 4B). To further assess the activation of NF-κB activity by EPO, SN50, a cell-permeable inhibitor of NF-κB, was used. As expected, SN50 significantly blocked the ability of EPO to activate NF-κB, whereas a related control compound SN50M had no effect (Fig. 4C).
In different contexts both MAPK and PI3K have been reported to be required for NF-κB activation (25, 30, 37, 41, 42). To characterize further the pathways that lead to NF-κB activation, cells were pretreated with U0126, LY-294002, or SN50 and then stimulated with 5 U/ml EPO for 4 h. The activation of NF-κB by EPO was suppressed in the presence of SN50 as expected. Likewise, the PI3K inhibitor LY-294002 diminished activation of NF-κB by EPO. In contrast, inhibition of the ERK pathway by use of the MEK inhibitor U0126 was without effect (Fig. 4D). These results indicate that PI3K, but not the ERK pathway, is involved in EPO-mediated effects leading to NF-κB activation in endothelial cells.
PI3K and MEK provide EPO-mediated cytoprotective signals in endothelial cells.
Next, we examined whether the PI3K/Akt, MEK/ERK, and NF-κB pathways are involved in cytoprotection by EPO. We assessed cell viability in the presence of inhibitors of the relevant pathways (LY-294002, U0126, and SN50). Inhibition of PI3K by LY-294002 diminished the ability of EPO to promote HUVEC survival after serum withdrawal, indicating that EPO-provides a prosurvival signal through activation of the PI3K/Akt pathway (Fig. 5A). Furthermore, inhibition of the MEK/ERK pathway using U0126 also blocked the cytoprotective effects of EPO implying that EPO provides yet another survival signal by activation of the MEK/ERK pathway. In contrast, inhibition of NF-κB activation was unable to reverse the cytoprotective effects of EPO (Fig. 5A). The pathway inhibitors were also used to assess the effects of EPO on preserving mitochondrial transmembrane potential. As shown in Fig. 5B, inhibition of PI3K by LY-294002 or the structurally unrelated compound wortmannin abrogated the effects of EPO on preserving the mitochondrial transmembrane potential. Likewise, inhibition of MEK by U0126 also reversed cytoprotection by EPO. In contrast, inhibition of NF-κB by SN50 was without effect (Fig. 5B).
EPO regulates the expression of Bcl-2 family members in endothelial cells.
Given the effect of EPO on Δψm, we next investigated expression of various anti- and pro-apoptotic Bcl-2 family members, including Bcl-2, Bcl-x, BAX, BAD, and BIM. Serum-starved HUVEC were incubated in the absence or presence of 5 U/ml EPO, and the expression of the various Bcl-2 family members was determined by immunoblotting. No changes in the expression of Bcl-2 were detected under these conditions (data not shown). However, EPO led to a time-dependent increase in Bcl-xL protein levels (Fig. 6A). At 4 h EPO induced a 2.5-fold increase in Bcl-xL protein level, and by 16 h this had increased to eightfold (Fig. 6A). No consistent change in the level of BAX, BID Bcl-xS, and BAD was detected, and we did not observe cleavage of BID after serum withdrawal (data not shown). In addition, phosphorylation of BAD was not detected (data not shown). However, there was a large increase in the level of BIM protein after serum withdrawal, and this was counteracted by EPO (Fig. 6B). We next used endothelial cells derived from precursors present in the umbilical cord blood [CBEC, as previously described (31)] to determine whether the induction of Bcl-xL by EPO occurred specifically in HUVEC or whether it also occurred in other types of endothelial cells. As shown in Fig. 6C, EPO also induced Bcl-xL in CBEC in a similar temporal fashion.
PI3K is involved in the upregulation of Bcl-xL by EPO.
We next determined the signaling pathways involved in the induction of Bcl-xL by EPO by examining the effects of inhibitors of the PI3K, MEK, and NF-κB pathways. Inhibition of PI3K in the presence of EPO abolished the effects of EPO on Bcl-xL expression (Fig. 7A). In contrast, neither the blockade of NF-κB nor that of MEK activation attenuated the effects of EPO on Bcl-xL expression (Fig. 7A). Hence, these results suggest that EPO-mediated upregulation of Bcl-xL protein in HUVEC is mediated by PI3K and independent of ERK and NF-κB pathways.
In healthy cells, BAK activity is regulated directly and specifically by Bcl-xL (52). Therefore, to assess the relative importance of Bcl-xL to the antiapoptotic effects of EPO in HUVEC, we treated cells with a peptide bearing the BH3 domain of the pro-apoptotic protein BAK (BAK-BH3). BAK-BH3 binds tightly to Bcl-xL (52). Thus by binding to Bcl-xL, BAK-BH3 should reverse EPO-mediated cytoprotection and inhibit its anti-apoptotic activity. HUVEC were pretreated with BAK-BH3 peptide and then treated with or without EPO for 6 h. Apoptosis was determined by measuring Δψm. As shown in Fig. 7B, preincubation of cells with BAK-BH3 reversed the effects of EPO on maintaining the transmembrane mitochondrial potential, thus implicating a critical role for Bcl-xL in EPO-dependent endothelial protection.
Endothelial cell damage is a critical component in the pathogenesis of a number of vascular pathologies such as endotoxic shock and atherosclerosis (6, 14). Identification of biochemical pathways that can signal anti-apoptotic responses is therefore important because this may provide a basis for implementing a clinical intervention. In this study, we explored apoptosis-inhibiting pathways mediated by EPO in endothelial cells subjected to serum withdrawal. Our data indicate that EPO confers cytoprotection through the activation of the PI3K/Akt pathway with subsequent upregulation of Bcl-xL as well as through the activity of the ERK pathway independent of Bcl-xL. Serum starvation also led to an increase in the level of pro-apoptotic BH3-only protein BIM, and this effect was reverted by EPO.
The Bcl-2 family consists of pro- and anti-apoptotic proteins that play a critical role in regulating cell survival. The pro-apoptotic BH3-only proteins respond to apoptotic stimuli and are thought to activate multi-domain pro-apoptotic members (BAX and BAK), which trigger the intrinsic pathway of apoptosis. In contrast, anti-apoptotic Bcl-2 family members bind and sequester BH3-only molecules and thus prevent activation of BAX and BAK.
As a survival factor, EPO regulates the activities of some of the Bcl-2 family of proteins. EPO maintains levels of anti-apoptotic molecules (Bcl-2 and Bcl-xL) and decreases the amount of pro-apoptotic BAX in various other cell types (43, 46, 49). In this study, we show that EPO induces the expression of Bcl-xL in HUVEC following serum withdrawal. Bcl-2 was detected at low levels or not at all in HUVEC, and EPO treatment did not induce Bcl-2 expression. Bcl-xL, but not Bcl-2, therefore, appears to be important for EPO-mediated cell survival in endothelial cells. EPO is also required for optimal expression of Bcl-xL in erythroid cells (21, 46). Similar results in neurons indicate that upregulation of Bcl-xL may be necessary for EPO to confer its anti-apoptotic effects (50). Therefore, the regulation of Bcl-xL expression is likely to play an important role in the anti-apoptotic effects of EPO.
Although EPO upregulates Bcl-xL expression in a variety of cells, the signaling pathways that promote this response appear to vary in different cell types. In hematopoietic cells, for example, EPO upregulates the expression of Bcl-xL through activation of Janus kinases but independently of PI3K, Ras, and STAT5 (40). However, in another study, EPO was shown to induce expression of the Bcl-xL gene through activation of STAT5 by binding to the Bcl-xL promoter (48). In the present study, we found that EPO upregulates Bcl-xL in endothelial cells through activation of the PI3K pathway but not the MEK/ERK pathway. This finding also differs from EPO-mediated Bcl-xL regulation in erythroid progenitor cells where activation of ERKs is reported to be involved (36).
The BH3 domain of the BH3-only proteins is an interaction domain that enables them to bind to multidomain anti- or pro-apoptotic members antagonizing the survival or activating the pro-apoptotic activities (12, 13, 29). To further clarify the role of Bcl-xL in EPO-mediated endothelial cell survival, we determined whether inhibition of Bcl-xL with a BAK-BH3 peptide induces apoptosis in the presence of EPO (52). We demonstrate that treatment of cells with a BAK-BH3 peptide that specifically binds to Bcl-xL reversed the anti-apoptotic effect of EPO. By inhibiting apoptosis, the anti-death Bcl-2 proteins preserve the integrity of the mitochondria. Consistent with this role, we also show that Bcl-xL inhibition by the BAK-BH3 peptide caused a reduction in the mitochondrial potential. Furthermore, inhibition of PI3K, which is required for the upregulation of Bcl-xL by EPO, also reduced the mitochondrial potential in the presence of EPO.
To determine further the mechanism by which EPO exerts its anti-apoptotic effects, we measured the expression of some of the pro-apoptotic members of the Bcl-2 family. The expression of the pro-apoptotic Bcl-2 family member BIM increased following serum withdrawal in HUVEC. Furthermore, this upregulation of BIM as with the induction of apoptosis was blocked by EPO. Increased expression of BIM has been observed following withdrawal of survival factors in a variety of cells, including fibroblasts, epithelial cells, lymphocytes, and neurons (8, 16, 39, 51), and appears to be a conserved mechanism for inducing apoptosis under these conditions.
In contrast, expression of BAX and BID was not changed following serum withdrawal or exposure to EPO. In contrast, BAX expression was upregulated in an in vivo rat model of renal injury and this was counteracted by EPO (49). BID is activated by proteolytic cleavage following ligation of death receptors, but no loss of full length BID or cleavage products were detected after serum deprivation of endothelial cells. BID may therefore not be involved in apoptosis of HUVEC following serum withdrawal. BAD is regulated through phosphorylation at Ser136 and Ser112, and this promotes its sequestration in the cytoplasm by binding to 14-3-3 proteins. There was no evidence of BAD phosphorylation following exposure of HUVEC to EPO, suggesting that BAD phosphorylation is not one of the mechanisms by which EPO exerts its anti-apoptotic effects in HUVEC. So far there is no evidence to support a role of BAD in growth factor withdrawal-induced apoptosis using targeted gene deletion studies in mice. In contrast to BAD, resistance of BIM-deficient lymphocytes and neurons to cytokine withdrawal indicates that BIM is a critical initiator of this pathway to apoptosis. EPO is able to suppress BIM induction in endothelial cells, thereby providing an important additional survival mechanism.
The NF-κB signaling pathway is critical in regulating the apoptotic response. NF-κB is commonly associated in apoptosis suppression by transactivating the expression of anti-apoptotic genes. EPO activated NF-κB in HUVEC, and this activation was blocked by SN50, a peptide inhibitor derived from p50 subunit of NF-κB, which inhibits translocation of active NF-κB into the nucleus (34). Activation of both the PI3K/Akt and MEK/ERK pathways has been shown to be required for NF-κB activation under various conditions (25, 30, 47, 53). The lack of a role for the MEK/ERK pathway in NF-κB activation has also been reported (5). Here we show that PI3K, but not MEK/ERK, is required for EPO-induced activation of NF-κB in endothelial cells. These observations suggest an absence of cross talk between ERK and NF-κB pathways in the EPO signaling cascade. Furthermore, selective inhibition of NF-κB did not attenuate EPO-mediated endothelial survival. Therefore, in this study, the NF-κB pathway can be dissociated from the survival pathways required to confer cytoprotection by EPO.
EPO activated the MEK/ERK pathway, and suppression of this pathway reversed the anti-apoptotic effects of EPO. One mechanism by which the MEK/ERK pathway protects cells from apoptosis is through ribosomal S6 kinase (RSK), which inactivates the proapoptotic protein BAD through phosphorylation (7). ERK5 in particular has been shown to protect endothelial cells from apoptosis by stimulating phosphorylation of BAD (38). We did not detect BAD phosphorylation after exposure of HUVEC to EPO, therefore, this pathway leading to BAD phosphorylation is likely not required for anti-apoptotic effects of EPO. Another mechanism by which the MEK/ERK pathway confers cytoprotection is through phosphorylation of BIM leading to BIM degradation by the proteasome (32). In addition to promoting BIM turnover, phosphorylation of BIM by ERK1/2 may also prevent BIM from interacting with BAX, presumably preventing activation of BAX and cell death (22). In line with the aforementioned studies, serum withdrawal in HUVEC led to the upregulation of BIM protein. Treatment with EPO reversed BIM upregulation. A schema of the mechanism of the anti-apoptotic action of EPO is illustrated in Fig. 8. Taken together our findings indicate that EPO-mediated survival of endothelial cells is mediated by the PI3K and MEK/ERK pathways as well as through regulation of BIM expression.
This work was supported by grants from the Canadian Institutes of Health Research and the Heart and Stroke Foundation of BC and the Yukon, and funds from Ortho Biotech. A. Karsan is a Senior Scholar of the Michael Smith Foundation for Health Research.
We thank Fred Wong for technical support.
The costs of publication of this article were defrayed in part by the payment of page charges. The article must therefore be hereby marked "advertisement" in accordance with 18 U.S.C. Section 1734 solely to indicate this fact.
- Copyright © 2007 by the American Physiological Society


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Students! We are now on SkidSync! Check out our portal and join to get updates about events and schedules.
Welcome, Visitors! Visiting for Homecoming, Celebration Weekend, or just to check out Skidmore? Welcome! You'll find information about local religious and spiritual communities on our "Local Worship" page (on the menu to the left) and our weekly on-campus schedule below. You are welcome to stop by our offices on the second floor of Case Center. Don't forget to check out Wilson Chapel while you're here, perhaps on your way to explore the beautiful North Woods!
The Office of Religious and Spiritual Life (ORSL) provides opportunities for members of the Skidmore community to develop their religious and spiritual lives, which are an important part of the educational experience.
We encourage enriching dialogue, meaningful ritual, and enduring friendships among people of all religious backgrounds.
We offer:
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Contacting a Chaplain or Spiritual Advisor
The ORSL staff are available to meet with students regardless of religious, spiritual, or secular background. Affiliated advisors and local religious leaders from specific traditions are also available on and off campus. We are happy to help students connect with leaders and communities from their preferred tradition.
Fall 2017 Drop-in Hours
- Tuesdays 1-3 pm and by appointement - Case 203 with Parker Diggory
- Thursdays 3-4:30 and Tuesdays 1:30 - 3 pm - Case 201 with Jennifer Schmid-Fareed
- ORSL advisors are also available most weekdays on campus. Stop by our offices or contact us to make an appointment.
Connect with the ORSL...
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Wilson Chapel
The chapel is a beautiful space for worship and wonder, retreat and reflection. When not scheduled for campus events it is available to the Skidmore community for personal reflection and prayer. Events at the chapel include yoga, meditation, religious prayers and services, and events that explore many forms of spiritual and contemplative practice.
Open Chapel
When not otherwise reserved the chapel is normally open from around 6:30 am to around 10:30 pm during the school year.
To contact the ORSL email Parker Diggory at firstname.lastname@example.com, stop by Case 201-203 (note our new office locations effective September 2017!) or see our staff page for more details.
Fall 2017 Weekly Schedule
All events are open to Skidmore students, staff, and faculty unless otherwise noted. Additional events are often added by our office and related clubs throughout the year. Connect with us on SkidSync to stay up to date!
- Christian Prayer
12:15-12:35 pm, Case 203
- Restorative Yoga w/ Mindful Movement Club
7:30—8:30 pm, Wilson Chapel *students only*
- Mindfulness & Meditation
12:15—12:35 pm, ICC
- Vinyasa Yoga
8—9:15 am, Wilson Chapel
- Open Chapel Drop-in Hours
2:30—4:30 pm, Wilson Chapel
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6:25—8:30 pm, Wilson Chapel
- Mindfulness & Meditation
12:15—12:35 pm, Wilson Chapel
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5—6 pm, Wilson Chapel
- Muslim Prayer
drop-in 12-1:30 pm, Wilson Chapel
- Kabbalat Shabbat Services
5:15-6:15 pm, Wilson Chapel, Student-led
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- Skidmore Christian Fellowship - Ignite!
7pm, ICC, large group gathering
- Havdalah Services 5pm, ICC
- Quaker gatherings
3-4 pm, Wilson Chapel, every other Sunday starting 10/1/17
- Vinyasa Yoga
5—6:15 pm, Wilson Chapel
- Roman Catholic Mass
select Sunday evenings, contact ORSL
Schedule subject to change. See SkidSync or contact kdiggory@skidmore for info or to find out about additional opportunities and local services.

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A sensory experience awaits at the Museum of Ice Cream.
The sold-out Meatpacking District pop-up, open until Sept. 4, features a trippy chocolate room, a sprinkle-filled pool and helium balloons made of sugar. Naturally, the museum offers up ice cream from the likes of Oddfellows and Chinatown Ice Cream Factory.
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The Sheraton Xiamen Hotel (Xiamen Xilaideng Jiudian) is centrally located in Xiamen's newest commercial district. Just a 15-minute drive from Xiamen International Airport and 20 minutes from downtown, the hotel offers guests a magnificent view of the Jiangtou area and neighboring park.
Designed by Hirsch, Bedner and Associates (HBA), the 30-story Sheraton Xiamen Hotel offers luxury furnishings in all of its 360 spacious guestrooms, Executive Club rooms and suites.
Various health and recreational facilities are available including an indoor swimming pool, a full-service health spa, fitness club and hair salon. Guests can also take advantage of the dry cleaning services and wireless broadband Internet access available in most of the hotel's public areas.
Other services include a business center, ticket office and airport pickup service.
Check in: from 15:00 Check out: before 12:00
The hotel does not accept guests below 18 years of age staying alone.
Buffet breakfast CNY 168
No pets allowed.
Payment can be made by credit card at the hotel. You can use the following credit cards:
All rates displayed are based on current exchange rate between CNY and the selected currency.
Children and Extra Beds
The hotel does not accept guests below 18 years of age staying alone.
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Hotel brands | World Hotel | Hotel landmarks | Family Hotel | Boutique Hotel | Business Hotel | Luxury Hotel | Golf Hotel | Spa Hotel
Huangshan MAGNOTEL hotel | Changchun Hilton hotel | Hungary hotels | Kunshan hotels | hotels in Changsha | hotels in Xian | hotel in Nanchang | hotel in Dalian | hotel in Chongqing | Wuhu Mountain View Holiday Hotel | Yingbin Hotel


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Series: Psychic Visions #1
Published by Valley Publishing on June 16th 2011
Genres: Paranormal, Romantic Suspense
Format: eBook
Source: Purchased
Amazon | B&N | Goodreads
What she doesn't want...is exactly what he needs.
Shunned and ridiculed all her life for something she can't control, Samantha Blair hides her psychic abilities and lives on the fringes of society. Against her will, however, she's tapped into a killer—or rather, his victims. Each woman's murder, blow-by-blow, ravages her mind until their death releases her back to her body. Sam knows she must go to the authorities, but will the rugged, no-nonsense detective in charge of tracking down the killer believe her?
Detective Brandt Sutherland only trusts hard evidence, yet Sam's visions offer clues he needs to catch a killer. The more he learns about her incredible abilities, however, the clearer it becomes that Sam's visions have put her in the killer's line of fire. Now Brandt must save her from something he cannot see or understand…and risk losing his heart in the process.
As danger and desire collide, passion raises the stakes in a game Sam and Brandt don't dare lose.
I picked this one because it has my preferred kind of paranormal which is psychic / mind type abilities. Romantic suspense is a favorite genre. It also has fairly high ratings on Goodreads.
Sadly, I did not enjoy it. The characters were not real to me, so I did not have many feelings. Sam's relationships with her co-workers didn't feel believable to me. The plots seemed cliché. I could guess what was going to happen, so also no suspense.
The editing was okay, without any glaring errors to take me out of the story. There was some repeated wording or overused phrases. Everything looked good, but Tuesday's Child just didn't connect for me.
I read all types of books but my favorites are urban fantasy and romantic suspense.I like some romance and some action / adventure / mystery in everything.I like sex in my books but please, only between TWO people.Intimate conversation is equally important to the sex in a book.It lets me see the characters.The other thing I enjoy is humor.I'll be doing a feature monthly called Thrifty Thursday,which reviews books (ebooks)that have been available for free.
Friend me at Goodreads.
Latest posts by Anne (see all)
- Twisted Truths by Rebecca Zanetti - November 17, 2017
- Review tour: Midnight Labyrinth by Elizabeth Hunter - November 16, 2017
- Early review: Perfect Gravity by Vivien Jackson - November 2, 2017


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As your organisation grows, your technology needs also grow and you'll need to share your business data with colleagues and access the latest information through a tablet, smartphone or other mobile devices.
As your organisation grows, your technology needs also grow and you'll need to share your business data with colleagues and access the latest information through a tablet, smartphone or other mobile devices.
So many Software Services providing companies nearby are you still unable to find the perfect one for your business? Don't worry because now your hard work is over, as you are at the right team that is 1313Live.
E-Commerce is the newest craze in the digital market which is valuable for both buyers and sellers. Online stores has replaced the traditional shopping ways and increase the presence on the web
Mobile app development is considered as a challenging operation by many businesses, owing to its platforms and technologies. It's no secret that the demand of mobile app is increasing for each and every business.
Is your business online? Are you struggling to have your website on the first page of search engine results? Is your website getting low traffic? If so, your website needs internet marketing (SEO) to improve the visibility of your business.
Today, the world has changed in perspective of web technology. It is important to have a great web presence to achieve business success. A website represents a business or brand.
1313live services are tailored to each client, whether it's consulting or a full company restructure using technology to enhance the performance of their company. "You say and we deliver" is our motto.
Strategic consultancy
Software Development
Web Development
End-to-End Implementation
Quick starts
Ongoing Support
Force.com development (including APEX, Visualforce, and Sites)
Integration between two sites
We have just completed our project with 1313live, and not only are we very pleased with the site that they have created, we are getting excellent feedback from new clients that are now contacting us. We found working with them very simple, with an informative and straightforward approach, they guided us from A-Z with plenty of ideas and creativity along the way.
Im very happy with my website and my business from search engine enquiries is more than doubled from my old site.The team not only gave me a cool design but also a logo which has really helped with my branding. Having gone to 1313live for a new website originally, I have been in regular contact with them as they are always helping boost my website ranking and advising me how I can better my competition.
Excellent service with helpful technical people on the end of the line to change things as we worked towards our end solution. The ongoing support since completion continues to deliver which is a refreshing change and we await to see how the work put into the website and subsequent organic listing services develop into leads.
1313live designed and built our new responsive e-commerce web site and made a great job of it. Got the design right and with no fuss on the change over from our existing site, we have got the ability to edit the site news and all aspects really easily with the custom features on the back end of the site. Highly recommend the 1313live team for all your website and printing needs.


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by Shirley Times
(Memphis, Tennessee)
My 2nd best advice is to "Never shop when you are hungry" (impulse shopping). Make a list of the items you need before going shopping and stick to your list regardless of price and sale item temptations. Shop only for item needed weekly to avoid stocking items in freezer for later use, ultimately forcing you to eat out or shopping again for quick fix and hectic day food items. Use coupons if you have them. Check weekly sale ads to compare prices and decide where to shop after making your weekly menu list. Avoid items such as hair products, personal hygiene, etc., that can be purchased cheaper at the dollar store for instance.
Join in and write your own page! It's easy to do. How? Simply click here to return to Tell Me How You Save Money On Groceries?.


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Status: Closed
Know more about this business than we do? Cool! Please submit any corrections or missing details you may have.Help us make it right
Signs, banners, decals, letters. Made fast with high quality and great service by next day signs, your internet sign shop.your hometown sign shop busy fax us your order atthe best sign at the best price
Vehicles, Signs, Real Estate, Ca# 762007, Braille, Banner's
Be the first YellowBot user to add photos or videos of Next Day Signs
Next Day Signs can be found at W Pine Ave 205. The following is offered: B2B Signage. The entry is present with us since Sep 9, 2010 and was last updated on Nov 14, 2013. In Lompoc there are 2 other B2B Signage. An overview can be found here.
If you don't see your business listed on YellowBot, please add your business listing.
YellowBot wants to get your input! If you have a comment, find a bug or think of something neat we should do, let us know by emailing us.
© 2007-2017 Solfo, Inc. – All rights reserved


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The resort's popular pool area, including a heated lagoon pool, children's wading pool and two heated spas, has been completely transformed and now also features a new 'water blade' water feature from the pool bridge and two new poolside decks. Mantra French Quarter also offers complimentary WiFi to all guests via a voucher issues from reception upon check in.
Relax at the beach or unwind in a lounge chair by the resort pool. Let the little ones splash to their heart's content in a heated, kid-friendly wading pool, while you cook lunch on one of two poolside barbecues.
Finding a park close to Hastings Street won't be an issue for you. Park in the resort's secure, undercover car park and forget about it. Everything is within walking distance. Please note only 1 car per unit is available.
At Mantra French Quarter you can enjoy everything Noosa is famous for. Relax or get active and hire a paddle board.
TripAdvisor Rating


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Sen. Reid: Aiming to Debate Climate Bill by Summer
WASHINGTON -- Senate Majority Leader Harry Reid (D., Nev.), aims to debate -- and potentially even pass -- a climate change bill that would put a cost on emitting greenhouse gases by the end of summer, his office said Friday.
A climate bill failed to pass in the chamber last year, but with a new President in the White House who wants to cut greenhouse gases 80% from 1990 levels by 2050 and larger majorities in both chambers, Mr. Reid's ambition is in step with President Barack Obama's agenda and campaign rhetoric.
But climate change legislation could face challenges passing the Senate, where concerns about putting a high cost on carbon dioxide emissions crosses both party and regional lines. Democratic Senators from states that have heavy manufacturing, automobile plants and coal-fired power plants could be at odds with Democrats from the West and Northeast.
Mr. Reid said Wednesday the Senate could debate an energy bill within weeks, one that would lay the foundation for cutting emissions by mandating an increasing percentage of renewable energy and establishing a new electricity grid.
Critics of putting an expensive premium on carbon say that such a schedule may be overly optimistic given the global financial crisis and the potential impact putting a cap on greenhouse gases would have across nearly every sector of the economy.
"I don't think they can move a climate bill this year with the economy the way it is," said Bill Kovacs, head of regulatory and environmental affairs at the U.S. Chamber of Commerce. "I don't even think the (Environmental Protection Agency) is going to be able to do it...because they can't risk what's going to happen to the economy if that moves forward."
Separately, the Obama administration is pushing ahead with studies of whether the EPA could regulate carbon dioxide under the Clean Air Act.
EPA chief Lisa Jackson said last week that her office would soon begin drafting rules may start regulating CO2 through power plant permitting. Part of the rule-making process, EPA officials say, is likely to include an "endangerment" finding that could set off a domino effect of Clean Air Act CO2 regulations throughout the economy.
Already, staff have been intensely reviewing and updating an existing endangerment finding made last year by agency officials -- but blocked by the previous administration -- that found carbon dioxide threatened human welfare. "Administrator Jackson is reviewing a number of issues, including those related to climate change," said EPA spokeswoman Roxanne Smith.
Write to Ian Talley at email@example.com


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2017-03-16 12:57
Former IRIB head to run for presidency
TEHRAN, Mar. 16 (MNA) – The former head of Iran's IRIB Seyyed Ezzatollah Zarghami announced on Wednesday that he will run in the upcoming Presidential Election as a rival to President Rouhani.
2015-10-12 12:37
Leader receives IRIB officials
TEHRAN, Oct. 12 (MNA) – Iran's Leader Ayatollah Khamenei received officials of Islamic Republic of Iran Broadcasting earlier this morning.


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Express. Home of the Daily and Sunday Express.
Transfer News: Man United winger deal, Arsenal exit close, Costa's Chelsea bombshell
EXPRESS SPORT brings you the latest transfer news from the Premier League.
Lyon have made Manchester United outcast Memphis Depay their number one target this month - but have had their first bid knocked back.
The Ligue 1 side want to bring Memphis to France to help their title charge.
Lyon sit fourth ten points behind Nice but with a game in hand.
Memphis Depay is on Lyon's radar this month
And their coach Bruno Genesio says he still wants his man.
"I have put a priority on Memphis Depay," said Genesio.
"Things are progressing. We keep on working. It takes time because recruiting a player now is not easy.
"He has an interesting profile for us. But there are several things to consider, including the price."
However Paris Saint-Germain have also entered the race, according to the Sun.
Memphis has struggled since his £25m move from PSV 18 months ago.
He has started just one game this season.
Carl Jenkinson is set to move to Crystal Palace
Carl Jenkinson is closing in on a move to Crystal Palace.
Express Sport exclusively revealed a move was close earlier this week.
And Wenger has confirmed the wheels are in motion for a deal.
Wenger said on Jenkinson's future: "It's a possibility that he will move in the next two or three days."
Diego Costa has been left out of the Chelsea squad for tomorrow's game
Diego Costa's Chelsea future has been thrown into doubt after a bust up with Antonio Conte.
Express Sport understands the pair have clashed over fitness issues and with a big money offer on the table from China.
Costa has now been dropped for tomorrow's clash with Leicester and he could be sold this month.


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February 26, 2009 in Services (E)
[prMac.com] Montreal, Quebec - Mac a Montreal is a highly rated member of the Apple Consultants Network working hand in hand with all official Apple retail stores in the Montreal area (Carrefour Laval and Sainte-Catherine) to provide a wide range of expertise in the design and implementation of Apple solutions.
Mac a Montreal is made up of a group of highly dedicated, highly qualified, and very passionate consultants that can cater to almost every business and SOHO need. We are hybrid specialists that are equally knowledgeable in both Mac and Windows environments and help many businesses either completely switch over to an Apple solution or properly integrate the best of both worlds in a unified network that works reliably. Our consultants can identify opportunities for improving your business through technology solutions that you may never have considered. If you think you may not be maximizing the potential of technology in your business, then you can benefit from our experience.
Mac a Montreal can design customized solutions for updating your office that include plans for easy maintenance and continued upgrades as your business grows and changes. Because your relationship with us is ongoing, feel free to incorporate our expertise into your ongoing plans to keep your office computing needs up-to-date. We provide quality Apple-certified on-site servicing as well as remote support with rapid response times and post-visit assistance.
Some of the services we offer:
* Migration from Windows to Mac
* Integration of Macs in a Windows environment
* Server configuration and network administration
* RAID installations and configuration
* Software installations
* Hardware installations
* iPhone configurations and MobileMe integration
* Parallels, VMWare, and Bootcamp installations and configuration
* Remote support
* Backup solutions
* VPN connectivity
* and much more.
Whatever your question, no matter how big or how small, our fully Apple Certified experts are here to help. We can deal with any scale of request, from a home user setting up their latest Mac through to full large scale server configuration and installation. We pride ourselves on rising to the challenge, as let's face it, life gets boring when its all too easy. Mac a Montreal is a fully certified member of the Apple Consultants Network, making us a safe and reliable option for all your Mac needs, no matter how big or small.
Located in Montreal, Quebec, Mac a Montreal is a highly rated member of the Apple Consultants Network. Approved by Apple in Cupertino, we work hand in hand with all official Apple retail stores in the Montreal area (Carrefour Laval and Sainte-Catherine) to provide a wide range of expertise in the design and implementation of Apple solutions. All Material (C) 2009 Mac a Montreal / All Rights Reserved. Apple, the Apple logo, and all other Apple Products and hardware, are registered trademarks of Apple Computer in the U.S. and/or other countries.


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Gates Cambridge Scholar
Institutional Style Guide
Established in 2000 through a $210 million gift from the Bill and Melinda Gates Foundation, the scholarship offers recipients who reside outside the United Kingdom the opportunity to pursue graduate students at the University of Cambridge. The scholarship program aims to build a global network of future leaders committed to improving the lives of others.
The first class of Gates Cambridge Scholars, announced in 2001, included a UGA student.


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Sunny afternoon in Limburg
Limburg an der Lahn Travel Blog› entry 9 of 15 › view all entries
August 11th, 2007 – by: Zen
Limburg itself was very busy, with the local radio station havong some sort of event there. The modern centre of the town was full of people enjoying a fake beach that had been set up. There was a beachball competition and people were sitting in loungers, watching their kids build sand castles.
For more and better quality pics, have a look at Matt's blog
Join TravBuddy to leave comments, meet new friends and share travel tips!


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Overview: Working with one of Australia's largest fruit and vegetable producers, Comunet customised harvest management solution capturing yield and location data via tablets.
Key technologies: Microsoft, Panasonic, Symbol, Intermec, Preceda
Customer said: "Comunet performed admirably, wanting to treat our business like their own. This sets the springboard to continue the rollout of this solution not only to the berries division but to Costa's other produce streams." - Amapole Phoonie and Matthew Vella, Costa
Read more: Fruit picking goes digital at Costa Group with Microsoft partner Comunet
Overview: Created a mobile app for client's consultants built on the Salesforce Sales Cloud CRM.
Key technology: Salesforce
Customer said: "Time saving is a massive win. For example, previously if we had four jobs to do in Newcastle, we had to send four people to complete them. Now these can be attended by the one consultant. A huge bonus for us." - Stephen Ransley, Managing Director, Tyrrells Property Inspections
Read more: Tquila eliminates paper reports for Tyrrells Property Inspections with Salesforce connected app
Overview: Delivering reliable connectivity across Northern Territory by deploying fibre MPLS Telstra network, Cisco firewall and SIP replacement of PSTN.
Key technologies: Telstra, Cisco
Customer said: "It is hard to think that prior to this network project, we put up with having one person drive over two hours a day and 52,000km per year just to deliver a set of reports." - Mehdi Ghaffari, Marketing & IT Manager, Halkitis
Vendor said: "Being able to trust a partner like Calibre One, not just for design but for delivery as well, is very reassuring." - Michael Robotha, Cisco
Read more: How Telstra partner Calibre One saved Halkitis 52,000km a year
Overview: Developed a Microsoft Dynamics 365 sales, marketing and field services cloud solution to empower South Australian solar firm's diverse workforce.
Key technology: Microsoft
Customer said: "Artis Group were able to demonstrate a comprehensive and relevant solution whilst maintaining the vision of the guiding principles of the project and the mission of the organisation." Karen Jarrett, chief financial officer
Vendor said: "Artis Group have the right experience, capability and people." - Scott Smedley, Microsoft
Overview: Collaboration between SMB Consultants, Vend, Xero and Deputy to transform client from manual system to the cloud for point of sale, accounting and payroll.
Key technologies: Xero, Vend, Deputy
Customer said: "Fresh Flowers Group has been working with SMB Consultants over the past two years. In that short space of time they have helped implement and change the way we do business. We have no hesitation in referring SMB Consultants to similar businesses like minded to our own." - Amanda Felice, Operations Manager, Fresh Flowers Group
Overview: Top 20 Australian law firm undertook a comprehensive workplace transformation enabling a paradigm shift from the keyboard to the cloud.
Key technologies: Comscentre, Microsoft, Zerto, Lenovo, Dell, HPE Aruba, Secureworks, Citrix, ServiceNow
Partner said: "The new network comprises dark fibre services, traditional fibre and ethernet options, giving Maddocks diversity with their last-mile coverage and no single point of failure on any of the data services." - Scott Earle, Comscentre
For more than a century, Brother has won recognition as a brand synonymous with delivering product innovation and customer satisfaction. A Japanese company founded in 1908, Brother has 19 production facilities and 43 sales companies operating in 41 countries in different regions today.
Brother is now a leading business brand that produces quality innovative products for the print and imaging, labelling, mobility and sewing markets. Key products include laser printers, laser and inkjet Multi-Function Centres (MFCs), scanners, mobile printers, labellers, and a wide range of home and industrial sewing machines. A trusted brand worldwide that believes in the "Customer First" approach in all aspects of our business, Brother has continuously met the varied needs of our customers through our comprehensive range of quality business solutions.
Hewlett Packard Enterprise
Hyperconvergence: the path to IT transformation
The hyperconverged space is growing considerably in the Australian and New Zealand market. Our customers in ANZ already recognise hyperconvergence as the next step for their wider IT architecture. They are going beyond traditional use cases for hyperconvergence – VDI and ROBO deployments. Instead, they see hyperconvergence as a viable replacement for their traditional server and SAN environment, and as a simpler way to manage and deploy their IT. Speed, simplicity, flexibility, and cost are all key drivers for a hyperconverged infrastructure. With HPE, customers can realise all of these benefits, regardless of their size or vertical market orientation.
Huawei is a leading information and communications technology (ICT) solutions provider serving 45 of the world's top 50 telecom operators. In Australia, Huawei creates around 600 jobs and works with all major Australian operators. We provide enterprise ICT solutions and mobile devices across the globe and half of the Australian population already use some sort of Huawei product for their telecommunications needs. Huawei's products and solutions have been deployed in over 170 countries and support the communications needs of one third of the world's population.
For more information, please visit: www.huawei.com.au, follow us on Twitter @HuaweiOZ or visit us on YouTube at www.YouTube.com/HuaweiOZ.
As market leader in enterprise application software, SAP (NYSE: SAP) helps companies of all sizes and industries run better. From back office to boardroom, warehouse to storefront, desktop to mobile device – SAP empowers people and organizations to work together more efficiently and use business insight more effectively to stay ahead of the competition. SAP applications and services enable more than 345,000 business and public sector customers to operate profitably, adapt continuously, and grow sustainably.
For more information, visit www.sap.com.
Successful partnerships between organisations enable the development of new capabilities, access to new ideas, faster innovation and quicker times to market. Telstra ICT Partners are seeing returns in the form of increased profitability, new customer acquisition, new product development, enhancement of their capabilities and entry into new markets.
The Telstra Partner Program can help you see the power behind partnerships that enables you and your business to gain advantages in this hyper-competitive environment. Telstra is the partner you need to grow your customer base, accelerate your profit and create demand for your brand throughout Australia.
Veeam® has pioneered a new market of Availability for the Always-On Enterprise™ with solutions that help organisations meet recovery time and point objectives (RTPO™) of less than 15 minutes for all applications and data, through a fundamentally new kind of solution that delivers high-speed recovery, data loss avoidance, verified recoverability, leveraged data and complete visibility.
The Veeam ProPartner Program provides Resellers with the expertise to position, sell and implement Veeam solutions to our mutual customer base. To learn more, visit www.veeam.com/propartner.
Veeam also offers its award-winning software to hosting, cloud and managed service providers through the Veeam Cloud & Service Provider program. For more information, visit www.veeam.com/service-providers.
VMware is a leader in cloud infrastructure and business mobility. Built on VMware's industry-leading virtualization technology, our solutions deliver a brave new model of IT that is fluid, instant and more secure. Customers can innovate faster by rapidly developing, automatically delivering and more safely consuming any application. With 2014 revenues of $6 billion, VMware has more than 500,000 customers and 75,000 partners.
The company is headquartered in Silicon Valley with offices throughout the world and can be found online at www.vmware.com
Arrow ECS ANZ provide specialised, technical-service orientated distribution and support for technologies that drive innovation. We provide partners and end users with solutions that solve modern business challenges through our leading technology solutions; technical pre and post-sales support, our flexible finance options, market development and other channel services. We don't simply distribute technology, we translate it into tangible business benefits. That's how we offer our partners the ultimate channel experience.
Learn more about Arrow ECS ANZ at ecs-anz.arrow.com.
CA Technologies helps customers succeed in a future where every business—from apparel to energy—is being rewritten by software. From planning to development to management to security, at CA we create software that fuels transformation for companies in the application economy. With CA software at the center of their IT strategy, organizations can leverage the technology that changes the way we live—from the data center to the mobile device. Our software and solutions help our customers thrive in the new application economy by delivering the means to deploy monitor and secure their applications and infrastructure.
Explore the CA solutions that can help power your future at www.ca.com
Datto protects essential business data for tens of thousands of the world's fastest growing companies. Our Total Data Protection platform delivers uninterrupted access to data on site, in transit and in the cloud. Through Datto's network of partners, we provide forward-looking technology companies with products and services designed to continually keep business running, combined with unrivalled customer service and unmatched accountability. Datto is headquartered in Norwalk, Connecticut, and has offices in Rochester, Boston, Toronto, London, Singapore, and Sydney.
Learn more at www.datto.com
HP Inc. creates technology that makes life better for everyone, everywhere. Through our portfolio of printers, PCs, mobile devices, solutions, and services, we engineer experiences that amaze. With a strong channel presence across Australia, HP Inc key priority is the success of our partner community. We continue to look at ways we can pair our technologies with our partners capabilities to deliver fantastic solutions to our collective customers
Ingram Micro Australia is a wholly owned subsidiary of Ingram Micro Inc, and Australia's largest distributor of technology products.
As a vital link in the technology value chain, Ingram Micro delivers expertise in technology solutions, mobility, cloud, and supply chain solutions, enabling its business partners to operate efficiently and successfully in the markets they serve. Representing over 130 vendors to a network of more than 8,500 active Australian resellers, Ingram Micro helps business realise the promise of technology. Headquartered in Sydney, Ingram Micro Australia has offices in Brisbane, Sydney, Melbourne and Perth, and Advanced Logistics Centre in Sydney.
For more information see au.ingrammicro.com
Mia Distribution
Born out of the desire to fulfil a rapidly changing telecommunication space, Mia Distribution found its beginning as PABX Warehouse – Australia.
Mia Distribution, quickly gained notoriety as an efficient one stop shop with a complete product offering, it wasn't long before some of the industry's leading vendors sought local knowledge, strategic vision and market understanding to partner with one of the regions fastest growing telecommunications distributors.
A drive and passion to meet the demands of the channel, with the understanding of learning the channels requirements, is the company's core focus.
Nimble Storage is the leader in predictive cloud infrastructure. Its Predictive Cloud Platform gives users the fastest, most reliable access to data. By combining predictive analytics with flash storage, IT teams radically simplify operations in their data centre and in the cloud. Over 10,000 customers in virtually every industry rely on Nimble Storage to power their businesses.
We Eliminate the App-Data Gap
When your applications slow down, you've hit the app-data gap. Faster storage alone isn't enough, because 54% of issues are not directly caused by storage.
To close the app-data gap and radically simplify operations you need the unique capabilities of our Predictive Cloud Platform: Predictive analytics to prevent issues; and Multicloud flash fabric for freedom of choice.
Find out more at www.nimblestorage.com and follow us on Twitter: @NimbleStorage


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How Penis Enlargers Have Changed The Landscape Of Men's Health
A lot has changed in recent years. Judging from what is written in historical books and other writings, major changes have taken place in the different parts of life. Specifically, the aspect of men's health has improved significantly. Apart from the medical technology, a whole lot of transformation is being witnessed in the perception of men's health- even by men themselves.
Traditionally, men are expected to be strong and the "I can handle everything" had gained root. It was almost impossible for a man to admit that he wasn't feeling well- health wise and emotionally. Today, they are now taking an active role in enhancing their well-being. The reason for this is yet to be ascertained. However, it is believed that men have understood that good health is directly related to the ability to enjoy good sex. This shouldn't come as a surprise considering the fact that men value good sex.
Penis enlargement devices became commercially available at the beginning of the 20th century. Originally, the devices promised girth and length increase- and nothing more. With the passage of time, the lucrativeness of the industry became evident. As a result, more and more penis enlargement products were availed in the market. While the focus largely remained on devices capable of increasing penis length and girth, other types of products were discovered. It didn't stop there. A lot of discoveries are still being made today in the male enhancement sector. This is for a pretty good reason - male enhancement products are assisting men to improve their sex life and other aspects of their lives.
Having A Small Penis? Try Out These Sex Positions For Small Penis.
The relationship between overall body health and sexual satisfaction has always been known both to patients and medical practitioners. However, it has been accorded more attention in recent years- through research and testing. A lot of fresh information is being availed regarding the role played by nutrition in matters health. This explains why there is a variety of supplements aimed at supporting male health. They mostly come in the form particular vitamins. Additionally, there are exercises for improving stamina, sex drive as well as libido. Men are urged to feed on the right sexual diet to improve their sex life.
While all these male enhancement products are aimed at improving male sexual health, they achieve much more. When a man is able to perform exceptionally well in the bedroom, other areas of his life experience positive transformation. Indirectly, his confidence levels improve and there is nothing he can't conquer. He begins to witness growth in his business, career and even in relationships. In general, therefore, penis enhancement products have directly and indirectly changed the landscape of men.
The internet has provided a useful platform where men can interact and discuss health issues affecting them. On these forums, ideas are exchanged and opinions passed across. This shows a significant stride towards guaranteeing male health. Men have now taken the front seat to make sure that they are in perfect health conditions. On the same platforms, they discuss issues to do with sexual health. Today, topics associated with sex and ways of improving it are no longer a taboo. Men wishing to purchase penis enhancers receive varying advice from men who have used them as well as experts.
Ride in an Overnight Train
You can always add fun to your sexual life, by adding new styles and ideas. You should always try to break the normal routines of intercourse. Have you been thinking of adventurous love making session, with your partner or wife? If yes, then take her on overnight train trip. You can discover lots of excitement and fun of making love on a moving train. Book a sleeping compartment or coach and have her all night. Try newer and better stunts. You will enjoy making love, while viewing some of the scenic beauty on a moving train, and watching people outside window, when train halts. This is definitely going to be a new experience to both of you.
10 Tips To Improve Men's Health
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ProSolution plus vs vigrx plus review
Five tips for finding the right male supplements
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Remember, pills do not give you more length


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Remember the MonkeyLite SL, the handlebar that changed the mountain bike world? That was a great handlebar, but we set out to make it even better. We considered the riders that live and breathe XC racing and we redesigned our most popular handlebar from the ground up. We refined the TaperWall™ butting, added strength and toughness to the Easton mountain composite and refined the shape. The resulting EC90 SL handlebar sets the new standard in weight, stiffness and strength: no other riser bar compares. At 135-grams, this handlebar is light enough to win the weight contests, yet its strength makes it at home shredding rough singletrack atop a long-travel trail bike.
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Miguel Arias Bardou
Miguel Arias Bardou (1841–1915) was a Spanish born Cuban painter. He is best remembered for colorful landscapes such as Pueblo Campesino (1891) and Bohío y Palma con Riachuelo (1908) which mainly depicted harmonious rural themes.
Bardou was born in Astorga, León, Spain in 1841. He moved to Cuba where he worked as a painter and scenographer in Cuban theatres. In 1885 he made a notable display at the La Caridad Theater in Santa Clara, Cuba. He was later employed as a scenographer in the Alhambra Theater in Havana. In 1966, 1971 and 1980 his paintings were exhibited of the National Museum of Fine Arts in Havana. The Museum of Arts and Sciences (Daytona Beach) is today in possession of one of his earlier paintings, Paisaje Cubano (Cuban Landscape), painted in 1875.
|Wikimedia Commons has media related to Miguel Arias Bardou.
|This article about a Cuban painter is a stub. You can help Wikipedia by expanding it.


text: When purchasing tires for your truck I recommend you buy the 10 ply. I needed a set and had been running Michelins, they came on the truck. Got talked into buying a 6 ply tire and man what a difference a few "ply's" make. Truck feels like it's swaying off the road with the trailer hooked to it. Now I know what happened to me a couple years ago when I got the tail wagging experience one day. Going this morning and getting another set put on, this time 10 ply.


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Natural Sciences
Earth science
Earth science is about change overtime. Earth scientist study what the Earth was like now and before. Earth scientist use their knowledge to find out what was there or why it's there in physical forms of the Earth for historians. It is belonging to a typical study of history for historians and very critical to find out what happend.
"The branch of science that deals with the identification of the substances of which matter is composed; the investigation of their properties and the ways in which they interact, combine, and change; and the use of these processes to form new substances."
Natural science
Natural science is a branch of science that studies the physical world like chemistry, physics, biology and Earth science the sub disiplines.
the study of living organism
2.it helps historians by finding what was living and breathing back then.
This sub discipline helps historians because chemist can help analyze some of the substances that the historians can't.
Physics is
the branch of science concerned with the nature and properties of matter and energy. The subject matter of physics, distinguished from that of chemistry and biology, includes mechanics, heat, light and other radiation, sound, electricity, magnetism, and the structure of atoms.
1. Biology is important because biology studies living things between past and present
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FREE Muscle and Fitness Workouts
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Welcome to VitaWise.com, your Online Nutrition Store!
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Office Artificial Plants & Trees: Inject Style Into Your Workspace with Replica Landscapes
Are you searching for office artificial plants and trees? Would you like to enchance your work environment with office floor displays and planters? Replica Landscapes, one of the UK's leading artificial plants and trees suppliers, can help.


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Young Tibetans Nurture the Notion of Returning Home
Tuesday, March 25, 2008
DHARMSALA, India -- Twenty-three-year-old Samdup Nyima works in a Tibetan bakery in this busy Himalayan settlement in northern India, which is host to thousands of exiled Tibetans, including their spiritual leader, the Dalai Lama.
Nyima was only 7 when he made the perilous journey here with his uncle, crossing the border to Nepal and then to India. He left behind his mother and sister in Lhasa, the Tibetan capital. His uncle returned to Tibet after putting him in the care of a school run by the government-in-exile, headed by the Dalai Lama.
"I don't remember much about the journey, I was too little," Nyima said. "But I do remember walking for a long time on a bridge over a very large river. After my father died, my mother sent me to India. She said, 'I want you to study in a free country and have the blessing of the Dalai Lama.' It has been 16 years, and I have not seen my family since."
Nyima, known as Sam to his friends, grew up like many other Tibetans in a boarding school here called the Tibetan Children's Village. In addition to the regular subjects of math, science and English, the school anchored Nyima with classes in Tibetan history, language and religion.
"I know everything about Tibet -- its mountains, its provinces, its folklore and history. I have thorough knowledge from the books, but I have no memory of Tibet," Nyima said as he skimmed news bulletins about the Tibetan uprisings that had been pasted on the walls of the main square. "I do not remember my house, my friends or what I used to play. My mother calls me every now and then and tells me that one of my childhood friends got married. She tells me the name, but I don't have any image to go with that."
With such tenuous links to his past, Nyima is like thousands of young Tibetans in India who painstakingly nurture the idea of returning to an idyllic land whose pristine images, in posters and calendars, adorn the walls of their homes. He said he will return one day, even as he dreams of opening his own restaurant in southern India.
"I will go back when Tibet is free. And I believe that is around the corner," he said. "Even if I set up a successful restaurant in India, I am still a Tibetan in exile. This is not home, even though I do not remember my home."
Nyima shares his room with three others from his bakery. On one of the walls are posters of Christina Aguilera, Jennifer Lopez and Blink-182, along with photographs of the Dalai Lama. He prays and turns the prayer wheels at the Buddhist temple every morning, takes part in demonstrations for Tibetan independence and occasionally volunteers at the Tibetan Youth Congress.
But since the latest uprising this month, he is a nervous wreck. Telephone calls to his 28-year-old sister, who teaches in Tibet, are not getting through.
"I heard that a lot of young people are being picked up by the police everywhere, and I worry that she may be in trouble," he said.
When he called his mother in February for the Tibetan New Year, she said she was getting old. "She said she wants to see me. If I don't go soon, I may never see her again. I have some duties as a son."
But applying for a visa from the Chinese Embassy in New Delhi might not be easy. After he came to India in 1992, his mother was fearful of disclosing the fact that her son had escaped. She has told the Chinese authorities that her daughter is her only child.


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Learn about this topic in these articles:
...be deduced from the other two laws. As a result, Nernst became one of the earliest wholehearted supporters of Einstein and quantum mechanics. In particular, Nernst was instrumental in organizing the First Solvay Congress in Physics, held in Brussels in November 1911, which was devoted to a thorough evaluation of the new quantum hypothesis by a group of leading European physicists.


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There's no better place to take your kids than to the happiest place on earth. Make their wildest dreams come true as they come face to face with their favorite cartoon characters, have autograph signing and photo sessions with them, and watch them perform amazing song and dance numbers right in front of their eyes. Visit the Animal Kingdom to interact with friendly goats and sheep, visit the Barnstormer at Fantasyland and play games at Buzz Lightyear's Space Ranger Spin. There are way too many exciting things to see and do here so make sure to make the most out of your time.
Explore this unique combination of geologic color and erosional land forms that is 277 river miles long. Known as one of the most visited tourist spots in the world, your kids can also have an amazing adventure-filled day here. They can join their Junior Ranger Program where they can learn about the park's history and discover the park's exotic plants, animals and rocks. Kids also get to learn how to help conserve the environment through the program.
San Diego Zoo is home to hundreds of animals coming from different species. Your kids will go wild as they go from one animal exhibit to another and get up close and personal with the animals that they only used to see on TV. Take your kids to the zoo's KinderNights tour where they can interact closely with the zoo's friendly animals and enroll them in the KinderTots program where the zoo's early childhood education specialists connect animal facts and green practices.
Let your children experience the beauty and grandeur of the Niagara Falls. Aside from having your photos taken at the viewing deck, there are a number of fun activities that you can do with your family. Sign up for the Cave of the Winds Tour for the ultimate water park experience. Go on board the Maid of the Mist boat ride which will take you right up to the falls for an exhilarating, one-of-a-kind experience.
Take a step back into time explore the Revolutionary City where you will feel what it is like to live in this tumultuous period of the country's history. Kids will enjoy the RevQuest: Save the Revolution activity where they get to go undercover and help save America. They can pretend to join the army have a feel of what it is like to live as a soldier. They can also play fun and imaginative colonial games at the Benjamin Powell House.


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DEFINITION of 'Chartered Business Valuator - CBV'
A designation offered by the Canadian Institute of Chartered Business Valuators (CICBV). A Chartered Business Valuator (CBV) is a financial professional who determines the value of all, or part of a business or company. CBVs take into account many factors and methods when determining the value of a business, including economic conditions, tangible and intangible assets and future cash flow.
BREAKING DOWN 'Chartered Business Valuator - CBV'
To become a Chartered Business Valuator in Canada, a candidate must first complete four core courses which include business and securities valuation as well as law and tax, along with two elective courses. A CBV candidate must also accumulate 1,500 hours of business and securities valuation work experience as well as receive a passing grade on the membership entrance exam.


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Video: Kodak Black – Tunnel Vision
In Kodak Black's new video for the Metro Boomin and Southside-produced "Tunnel Vision", a redneck deplorable points a gun at an innocent man and ends up getting righteously choked the fuck out. Make sure you watch until the last few seconds though.
Previously: Rae Sremmurd ft. Kodak Black – Real Chill (Video)


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Q: Can I pay in USD using my credit card ?
A: Yes, we accept USD credit card payments in our website shop. If its your first time using your credit card on the site you will be asked to fill in a 3D Secure box; this is for your own security and is standard practice. Please note that we do not keep records of credit card numbers; all transactions are processed via Realex and 3D Security adds another layer of security which protects the customer and their information; the extra autenthication will be asked for at the checkout. You can read more here:
Please see screengrab copied below as an example of the 3D Secure screen you will meet at the checkout.
Q: Where is Celtic Collections located?
A: We are based in Dublin, the Irish capital city. All customer orders are shipped from there. See the 'Contact Us' for address and specific location.
Q: How long has Celtic Collections been operating?
A: Celtic Collections was established in 1996 and has been releasing great Irish music for twenty years.
Q: How many Albums have been released by Celtic Woman?
A: Celtic Woman has to date released twelve albums; titles below.
'Celtic Woman'.
'Celtic Woman; A New Journey.
'Celtic Woman; A Christmas Celebration'.
'Celtic Woman; Lullaby.
'Celtic Woman; Believe'
'Celtic Woman; Songs from the Heart'
'Celtic Woman; Greatest Journey'
'Celtic Woman; Home for Christmas'
'Celtic Woman; Emerald Musical Gems'
'Celtic Woman; Decade'
'Celtic Woman; Solo'
'Celtic Woman; Destiny'
Q: Who is the red head on the cover of the Celtic Woman albums?
A: The lady on the cover of all the Celtic Woman CD's and DVD's was chosen as a physical representation of a Celtic Woman. She is beautiful yet mysterious, and her physical characteristics, red curly hair and blue/green eyes, are synonymous with our image of a Celtic Woman. She is not, and never has been a performer with the show.
Q: Who designed the costumes worn by the Celtic Woman performers?
A: The costumes worn by the soloist's were designed by Irish Designer Synan O'Mahoney. Well known for his lavish gowns, Synan was the first choice for Celtic Woman costumes.
Q: How many albums have been released by The High Kings?
A: The High Kings have released six albums, titles below.
'The High Kings; The High Kings'
'The High Kings; Memory Lane'
'The High Kings; Live In Ireland'
'The High Kings; Friends for Life'
'The High Kings; Four Friends Live'
'The High Kings; Grace & Glory'
Q: Whose musical legacy are The High Kings carrying forward?
A: The band are carrying a long musical tradition forward but their constituent parts come from specific family relations. Martin Furey is the son of legendary piper and singer Finbar. Brian Dunphy's father Sean performed in the showband era. Finbarr Clancy is from the famous Clancy Family and Liam Clancy was his uncle.


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23 Patten Street Whyalla Stuart 5608 SA
Secure & Tidy Home
3 good sized bedrooms
Neat kitchen with new electric oven
Split system air-conditioning to lounge
Polished floor boards to lounge, hall and bedrooms
Tiled laundry and toilet
Security screen on rear door and security screens on back windows
Large low maintenance yards
Rain water tank and new hot water system
Side Carport with roller door and access to rear yard
Close to schools, shopping complexes and sporting grounds
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Elders Real Estate (RLA62833) Whyalla
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- Please select a date to see the proper prices!
Description of the accommodation:
Rooms, services Soul Vacation Resort and Spa Benaulim is offering 30 nicely furnished rooms to stay in to those seeking accommodation in Benaulim. The café or the bar can be an ideal place for a pleasant evening talk or a meeting. In the safe you can secure your valuables. Wireless Internet connection (WiFi) is available at the hotel, enabling portable devices and smartphones to go online. If you wish to learn more about the exact scope of services provided by Soul Vacation Resort and Spa Benaulim (including those available for an extra fee) or about the equipment of the rooms, you can find the complete list on the information sheet below.
Payment Advance payment by credit card or providing credit card guarantee may be necessary for finalizing the reservation, depending on the chosen date, room and price.
Arrival, opening hours The earliest time to arrive is 14:00, the latest time to leave the rooms is 12:00. You will be able to speak with the personnel in English.
Soul Vacation Resort and Spa Benaulim rooms Rooms
- Please select a date to see the proper prices!
Accommodation services
Free WiFi in shared places
- Pool (Outdoor - seasonally open)
Garden, Bar, Cafe, Safe deposit box
Bathrobe, Hairdryer, Fridge, Desk in the room, Mini-bar, Tea/coffee-maker


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After 43 years of membership, the United Kingdom (UK) voted to leave the European Union (EU) in an unprecedented move.
While the full effects of the so-called Brexit will only be known in the coming months and years as the UK negotiates its exit under the Article 50 of the Treaty on European Union, understanding the next steps and potential implications for insurance buyers and insurance carriers is paramount.
Potential alternatives to EU membership, freedom of movement, the right of insurers and brokers to passport, and changes in the legal and regulatory environments will be defined in the in line with exit negotiations.
This Adviser outlines some key insurance implications that companies should be considering in their contingency plans, including:
The outcome of the referendum means a change in the relationship between the UK and the EU, but the specifics are far from certain at this point. Marsh will be working to further understand what Brexit will mean for clients and will be issuing further guidance as the impact becomes clearer.
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Mohammad Nazmuz Salehin
From PayPal Inc
I am working as a Senior Engineering Manager in PayPal. I have completed Masters in Computer Science and MBA. I have over 15 years of work experience and over 8 years experience to manage engineering teams in multiple locations like Bangladesh, Singapore and USA.
Methodology & Architecture
Build & Manage Engineering Team
I would like to share my experience on managing engineering teams. Forming, storming, norming and performing stages for engineering teams are different from other teams. And we also need to do adjourning more frequently. As I have experience to manage teams in different geographical locations I would also like to share the cultural aspect of managing a team. Will also touch on Agile, Lean management aspects on managing software development teams.


text: "Duto paisa diye jao baba...bhogoban mongol korben."(Please spare some change, my son....and may God bless you for it.) Meghna turned around to see an old lady in a maroon shawl and a saree that in some distant past must have been white in colour. She was old and emaciated, just like so many other old and emaciated women in tattered clothing that turned up round the corners, … [Read more...]


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We create customised weddings and events, our services include styling, coordination and prop hire. We bring your vision to life! Enquire now
Ruth was absolutely amazing! I approached Ruth at short notice and she saved our wedding!
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Tumbleweed Events rescued our wedding day! I hired them to plan and style our wedding ceremony and reception space but when the worst thing happened - the wait staff I had booked didn't turn up to our reception - Adam and Ruth from Tumbleweed Events stepped up and saved the day. We had an empty venue that needed to be set up for 120 guests and no one to set-up or serve guests. Tumbleweed took over and set-up everything. Yet they were only hired to style the event! They went above and beyond and never complained once. Their only concern was making sure that the bride, groom and guests were happy. They did all this and still made the place look like a spectacular fairytale! I can't recommend them enough.
Ruth at Tumbelweed Events was simply a delight to deal with. Ruth understood the look we were going for and was so helpful. We hired an arch, vintage suitcase and table plan chalkboard from her. Her pick up and delivery were on time and on budget. Thank you Ruth for making our wedding look so good!
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