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To regulate their internal environments, organisms must adapt to varying ion levels in their diet. Adult Drosophila were exposed to dietary salt stress, and their physiological, survival, and gene expression responses monitored. Insects continued to feed on NaCl-elevated diet, although levels >4% wt/vol ultimately proved fatal. Affymetrix microarray analysis of flies fed on diet containing elevated NaCl showed a phased response: the earliest response was widespread upregulation of immune genes, followed by upregulation of carbohydrate metabolism as the immune response was downregulated, then finally a switch to amino acid catabolism and inhibition of genes associated with the reproductive axis. Significantly, the online transcriptomic resource FlyAtlas reports that most of the modulated genes are predominantly expressed in hindgut or Malpighian (renal) tubule, implicating these excretory tissues as the major responders to salt stress. Three genes were selected for further study: the SLC5 symporter CG2196, the GLUT transporter CG6484, and the transcription factor sugarbabe (previously implicated in starvation and stress responses). Expression profiles predicted by microarray were validated by quantitative PCR (qPCR); expression was mapped to the alimentary canal by in situ hybridization. CG2196::eYFP overexpression constructs were localized to the basolateral membrane of the Malpighian (renal) tubules, and RNAi against CG2196 improved survival on high-salt diet, even when driven specifically to just principal cells of the Malpighian tubule, confirming both this tissue and this transporter as major determinants of survival upon salt stress. Accordingly, CG2196 was renamed salty dog (salt).
- Malpighian tubule
- microarray
- stress response
salt stress can disrupt internal homeostasis, so most organisms can robustly defend their internal milieu against fluctuations in external salt over a limited range. Work, particularly in yeast (1, 5, 6, 29, 34, 35) and in plants (3, 20, 23, 35, 36, 41), suggests a variety of mechanisms, particularly the synthesis or regulation of osmolytes, such as glycerol; the increase of K+ uptake and Na+ efflux at the plasma membrane; and Na+ accumulation at the vacuole (involving proton pumping ATPases, Na+-efflux pumps, plasma membrane proteins that operate as sensors of turgor loss and signaling cascades that regulate these responses).
The worm, Caenorhabditis elegans, is a simple multicellular organism compatible with FACS-like high-throughput screening. In C. elegans, massive salt loading caused upregulation of glycerol 3-phosphate dehydrogenase (25), so allowing the accumulation of glycerol as a protective osmolyte, exactly as in yeast (24), where the high osmolarity glycerol stress pathway (HOG) is under control of the Hog1 kinase (24). Trehalose is another key osmolyte in bacteria, yeast and plants (17); and in C. elegans, osmotic stress increases expression of the synthetic enzyme trehalose 6-phosphate synthase (tps1 and tps2), with a corresponding increase in trehalose levels (26).
In insects, mechanisms of salt tolerance are less well understood. In the aquatic larvae of the mosquito Aedes aegypti, there is an interplay between salinity, feeding, and hemolymph levels of the diuretic amine 5-hydroxytryptamine (12). Most larval mosquitoes live in fresh or brackish water, and so they are specialized for uptake of ions as a scarce resource. The size of the ion-scavenging anal papillae varies inversely with the ionic content of the water, suggesting that these are key osmoregulatory organs (13, 31, 42). By contrast, some species of insect survive in and around saline lakes and show particular specializations for handling of ions (like magnesium) that are present in massive excess (7). The insect body plan is thus adaptable to a wide range of environmental salt levels.
Although Drosophila does not normally encounter extremes of salinity, its advantages as a genetic model in which gene expression can both be studied and manipulated (15, 16) make it an attractive species to study. Previously, mutation of the inebriated putative neurotransmitter/osmolyte transporter (ine) was shown to confer reduced tolerance to dietary salt (21). Overexpression of either of the two isoforms of ine (ine-P1 or ine-P2) rescued this osmotic stress response defect. The osmotic stress-sensitive phenotype of the ine mutants was suggested to result from an inability to accumulate osmolytes within the Malpighian tubule and hindgut. Osmotic stress can also be induced by dehydration and rehydration. In Drosophila populations selected for enhanced desiccation resistance (2, 19), flies excrete some sodium during desiccation but retain ∼85% of the whole body sodium content, 83% of potassium, and 60% of chloride (19), implying that defense against desiccation is not purely by loss of ionic osmolytes.
The aim of this study was to investigate the response of adult Drosophila to acute salt stress, using Affymetrix microarrays and subsequent molecular, transgenic, and physiological validation of genes implicated in the response.
Flies were kept on standard medium in tubes at 25°C, 12:12 h photoperiod, and 55% relative humidity. Wild-type (Oregon R) flies were used for obtaining genomic DNA, cDNA, and for performing fluid secretion assays. Microinjections to produce transgenic lines were performed in a w1118 background. Transgenes under control of yeast upstream activating sequences (UAS) were driven ubiquitously by crossing to the Act5C-GAL4/CyO line (line 4414 from Bloomington Drosophila stock center) and in Malpighian tubule principal cells with the c42-GAL4 line (33).
Generation of transgenic RNA interference flies.
Inserts of ∼500 bp, directed against nonconserved regions of genes of interest, were cloned into the pWIZ vector (27) and germ-line transformed into w1118 embryos, according to standard procedures. This produced transgenic flies, in which hairpin ds-RNA could be expressed in cells of choice under control of appropriate GAL4 driver lines (8).
Dietary salt substitution.
For the preparation of the "salt" food, the required salt (NaCl or KCl) was dissolved in 100 ml of normal food just after its preparation, mixed, and the diet left to set. Where appropriate, indigo carmine dye (at a concentration of 1.66 g/l) was also added immediately after the diet preparation, but before the food set, to provide a visible marker of ingestion. Diet was freshly prepared to avoid any changes in the concentration of the salts due to evaporation. For dissection, we anaesthetized flies by chilling them briefly on ice before dissecting out tubules in Schneider's medium (Invitrogen).
Seven-day-old flies were transferred either into food with 4% NaCl added or normal food. The flies were left for 4, 8, 16, or 32 h to feed, and after that 100 flies from each treatment were anaesthetized on ice and homogenized in 300 μl of TRIzol. This whole procedure was repeated two more times to produce three biological replicates for each sample. All the samples were stored directly at −80°C and processed according to standard protocols. Samples of 20 μg total RNA were reverse-transcribed, then in vitro transcribed, according to Affymetrix standard protocols. The quality of the complementary RNA (cRNA) was also checked on an Agilent RNA Bioanalyzer, with a sample in which the broad cRNA peak exceeded the height of the low-molecular-weight degradation peak taken to be satisfactory. Samples were then run on the Affymetrix Drosophila genome I array under standard conditions: the experiment thus comprised 24 arrays. Quality control was performed as described in a previous paper from our lab (39).
As well as routine analysis by Affymetrix proprietary software (MAS 5.0), array data were analyzed using FunAlyse, a pipeline based on the Bioconductor package, and using robust multichip average for low-level normalization with subsequent calculation of rank products under random permutations. This method provides reliable estimates of fold change, significance, and false discovery rate (FDR), and a sharply improved performance in experiments with limited numbers of arrays (10). Significant changes were assessed for pairs of samples (salt vs. normal diet) at each time point, and gene ontology terms overrepresented in gene lists identified by iterative group analysis (iGA) (9).
Quantitative PCR.
Real-time quantitative reverse transcription PCR (qPCR) was used for further validation of the microarray results. Seven-day-old adult flies were collected and placed 1) in normal food, 2) in 4% NaCl food, and 3) in vials empty of food. After 8 h, flies were collected and anaesthetized in ice, and the tissue of interest was dissected. Total RNA was extracted (Qiagen RNeasy Kit) and cDNA prepared according to standard protocols (Invitrogen, Superscript II). Experiments were controlled against genomic DNA contamination with a no-reverse-transcriptase sample and were normalized against rp49. The samples were replicated independently six to eight times. Data were then expressed as fold difference of stress-treated samples compared with controls ± SE.
Cell type-specific RNA interference knockdown of gene expression.
For downregulation of CG2196 expression, CG2196-RNA interference (RNAi) flies were crossed to the c42-GAL4 (tubule principal cell specific) or actin-GAL4/CyO (ubiquitous expression) driver lines, and the offspring from the crosses were used to validate the extent of knockdown by qPCR. Six repeats were used for each of the different fly lines, and the experiments ware repeated twice more (three biological repeats) in total. Normalization of the results was performed against rp49 controls.
In situ hybridizations.
An expressed sequence tag (EST) clone of the CG2196 gene was obtained (GH19680 EDM1 133-6918917 Drosophila Gene Collection Clone 1) and used to prepare digoxygenin-labeled DNA probes as described previously (18). In situ hybridization was performed as described previously (18).
Survival of RNAi flies.
RNAi male flies were crossed with c42 virgin female flies in cages and for 10 days, eggs were collected each morning, counted, and transferred to vials with food containing 4% NaCl. The percentage of flies surviving long enough to reach the adult stage was compared with the parental lines. At least 12 vials from each line were collected. Two repeats of the full experiment were performed.
Generation of CG2196::eYFP-overexpressing flies.
Flies overexpressing a construct of the CG2196 gene tagged with enhanced yellow fluorescent protein (eYFP) were generated using the Gateway system. The entry and destination vectors used were obtained from the Drosophila Gateway Vector collection (37). The open reading frame of the gene was obtained from an EST clone and inserted in-frame into the Gateway destination vector pTWV, so encoding fluorescently COOH-terminal tagged CG2196, and flies germ-line transformed as described previously. The construct is under control of the UAS promoter, so allowing cell-specific expression under control of appropriate GAL4 driver lines.
Expression in S2 cells.
Drosophila S2 cells were grown in Schneider's medium according to standard protocols. Genes of interest were cloned the destination vector pAWV (Drosophila Gateway Vector Collection). This vector contains a constitutive ActinC5 promoter. S2 cells were transfected according to the Invitrogen protocol as described previously (32). Approximately 3 × 106 cells were used in each transfection. The cells were harvested in PBS and were allowed to attach to poly-l-lysine-treated glass slides. Cells were counterstained with the nuclear stain DAPI and washed repeatedly before being viewed in a Zeiss 510 Meta confocal microscope.
For fluid secretion experiments, significance of results was determined with an unpaired Student's t-test, taking the critical value of P as 0.05 (two-tailed). For microarray experiments, the FDR for significance was taken as 5%.
Do flies eat salty food?
Before studying the effects of dietary salt loading, it is important to establish that the diet remains palatable; otherwise, the effects of salt loading will be obscured by those of starvation or desiccation. Accordingly, flies were exposed to diet labeled with blue dye and containing various loadings of NaCl, for various times and scored for blue color in their guts. The results of this experiment can be seen in Fig. 1B. After 1 h with the food containing no extra NaCl, approximately half of the flies had blue dye in their gut (Fig. 1A). Flies in the food containing 4% or 5% NaCl also appeared with blue dye in their gut, which means that they had also eaten the food. After 3 h, almost all of flies had blue dye in their guts. After 6 h, all the flies in the food without NaCl supplementation, and at least half of those on 4 and 5% NaCl food show a full blue gut (Fig. 1A), confirming that, although food intake is reduced on high-salt diet, unpalatability is not a major issue.
Survival on high-NaCl diet.
Having established that flies continue to ingest diet containing high levels of NaCl, it is appropriate to examine survival. These results are shown in Fig. 1C. From the graph it is evident that flies fed with 3 or 4% NaCl food survive much better than those that are starved. These results are comparable to those reported previously (21), although our work reports a greater range of salt concentrations and monitors survival over a time course, rather than just at 4 days. Additionally, our wild-type stock survived marginally better on high salt; at 5% NaCl (wt/vol) equivalent to 0.85 M, 50% of flies survived to 4 days compared with 0% reported previously for 0.8 M (21). Although the difference is significant (probably reflecting strain differences), both experiments show a similar pattern of reduction of survival with increasing NaCl concentration. Flies fed with NaCl food up to 5% survive significantly better than the starved flies, but those exposed to higher salt levels die almost as fast as flies deprived of food altogether. This suggests that short-term salt-handling mechanisms can offer partial protection against salt levels elevated by up to 5% for at least a few days. Together with the previous results, a behavioral adaptation to high salt is apparent: insects provided with high-salt diet continue to feed (and thus survive better than insects with no food) but feed less avidly than flies on normal diet. In the subsequent microarray time-course experiment, shorter times (0–8 h) can thus be taken as representative of acute salt exposure, whereas longer time points (16–32 h) might be expected to show additional effects of reduced food intake.
Microarray of acute NaCl stress.
From these results, a microarray experiment was devised comparing matched sets of flies transferred from normal diet to either normal or 4% salt diet and sampled after 4, 8, 16, and 32 h. The numbers of genes both up- and downregulated increased steadily with time, as summarized in Fig. 2. The changed genes are listed in Supplementary Table S1,1 and iGA of all changed gene families is shown in Table 1. The iGA gene list shows some compelling features. When viewed as a time series, the response can be resolved into several phases. Initially, there is a prominent immune response, with multiple genes (metchnikowin; drosocin; diptericin; attacin; immune-induced molecules 1, 10, and 23; PGRP-SB1) upregulated at 4 h and downregulated at 8 and 16 h. At 8 and 16 h, the major response is an upregulation of genes associated with carbohydrate metabolism. Conspicuously, about half of the Drosophila α-glucosidase (maltase) gene family is upregulated, together with an α-amylase and the zinc finger transcription factor sugarbabe (sug), a transcription factor that has previously been implicated in starvation and sugar stress responses (45). Sugarbabe upregulation after feeding is thought to downregulate sugar metabolism (45), so its downregulation here is consistent with upregulation of carbohydrate metabolism.
Sugar metabolism as a general response to salt loading.
The marked changes in genes involved in carbohydrate metabolism are evocative of studies of high-salt diet in other species. In both yeast and C. elegans, salt loading causes the accumulation of glycerol as a protective osmolyte (25) (24). In yeast (24) the HOG stress pathway is under control of the Hog1 kinase (24). Although the Drosophila Hog1 homolog mpk2 does not change significantly in our experiment (Supplementary Table S1), this does not preclude a posttranscriptional modulation of glycerol accumulation, so we cannot absolutely exclude it as a mechanism in insects.
Trehalose is another key osmolyte in bacteria, yeast, and plants (17); and in C. elegans, osmotic stress increases expression of the synthetic enzyme trehalose 6-phosphate synthase (tps1 and tps2), with a corresponding increase in trehalose levels. Interestingly, although no early changes in trehalose metabolism are found in the array, there are two significant changes at 16 h: the synthetic enzyme trehalose 6-phosphate synthase 1 (tps1) is downregulated, whereas the catabolic α-trehalase (CG16965) is upregulated. Accordingly, trehalose does not appear to be a key osmolyte in response to acute salt stress in this insect. Rapid conversion of glucose to trehalose on passage through the insect midgut has classically been seen as a way of maintaining a favorable concentration gradient for glucose absorption from the gut (38), so it is possible that, in insects, trehalose has lost its primitive role as an osmolyte.
Another key change in osmotically stressed worms is the upregulation of chaperones of the heat shock protein (hsp) family, presumably to maintain protein folding under changing ionic strength. However, hsps are underrepresented in the change list in Drosophila (Supplementary Table S1): only hsp68 is found to change significantly and then only at a single time point. The impression is thus gained that Drosophila, with its relatively sophisticated osmoregulatory system, can defend its internal environment effectively against dietary salt loading for an extended period (of several days), before death ensues.
Ion transport and salt loading.
Ion transport genes, perhaps the most obvious candidates for change, are relatively modestly represented; one putative sodium/halide cotransporter is upregulated, while another does not change hugely. However, the major insect epithelial transport genes [Na+, K+ ATPase, V-ATPase, Na+/H+ exchangers, Na+/K+/2Cl− cotransporters (40)] do not feature in the list.
Which tissues are critical in salt response?.
These microarray experiments were performed on whole flies; although this design reduces the sensitivity of the experiment to tissue-specific changes of expression (11), it also avoids preconceptions as to the major tissues involved in salt response. However, since the work was performed, the FlyAtlas online resource has become available, allowing basal expression levels to be ascertained for any Drosophila gene across 14 tissues and life stages (11). So, although our original data were obtained from whole flies, it is possible to see from expression in FlyAtlas whether the 19 most significantly changed genes (as called by MAS 5.0) are housekeeping (generally expressed), or whether they are tissue specific, and whether any particular tissues emerge as common candidates for salt response. The results (Table 2) are remarkable; not only are both up- and downregulated genes all highly tissue-specifically expressed, but they almost all show highest expression in either the hindgut or Malpighian (renal) tubules, the two key osmoregulatory tissues of the excretory cycle (4), with some others specific to the midgut. Our model for response to high levels of salt, therefore, is a massive upregulation of carbohydrate metabolism (perhaps reflecting sharply increased energy demands) in the hindgut and Malpighian tubules, perhaps under control of sugarbabe (a transcription factor previously implicated in dietary stress response, and which is predominantly expressed in the midgut, tubules, hindgut, and fat body), while other tissues are relatively unaffected.
Validation of selected genes.
To encompass a range of the responses induced by salt loading, three genes were selected for further study: the SLC5 cotransporter CG2196 (annotated in FlyBase as a sodium/halide symporter), the GLUT4/8-like sugar transporter CG6484, and the transcription factor sugarbabe. According to FlyAtlas (Table 3A), basal expression of CG2196 is virtually tubule-specific, whereas CG6484 is massively upregulated in midgut, and sugarbabe is rather more widely expressed. The expression patterns of CG2196 and CG6486 were validated further by qPCR, confirming the massive expression of these genes in the tubules and midgut respectively (Table 3B). Whole fly quantitative PCRs for the three genes were performed for the 8 h time point and compared with the time-course microarray data (Fig. 3). In all cases, there was reasonable agreement. CG2196 and CG6484 were consistently and increasingly upregulated over time, whereas sugarbabe was consistently downregulated. Thus, based on this limited sample the array, FlyAtlas and qPCR data are concordant.
Role of CG2196/salty dog.
CG2196 was selected for further study, as it was predicted to represent a sodium/halide symporter by the computer annotation of the Drosophila genome, and because it was highly specifically expressed in the Malpighian tubules, one of the key tissues for response to salt (Table 2). According to Drosophila tradition for whimsical but informative names, the gene was named salty dog (salt). Firstly, the expression of salt within the tubule was established by in situ hybridization (Fig. 4). Expression was seen in the principal cells (not the stellate cells) along the length of the tubule but particularly in the main segment. The initial segment was not stained. This localization is reasonable, because the principal cells are the major active ion-transporting cell type in this tissue (15). No staining was seen in other tissues or with the sense control.
Salt localizes to the basolateral plasma membrane.
Drosophila S2 cells were transiently transfected with a salt::eYFP construct using the calcium phosphate transfection method. The S2 cells present a clear staining around the plasma membrane (Fig. 5A). Also, there seems to be staining in vesicles or conglomerations of the protein inside the cell, as frequently observed when proteins are overexpressed. So at least in the S2 cells, the CG2196-eYFP protein is a membrane protein that appears mainly in the plasma membrane.
Homozygous CG2196-eYFP flies (insertion in the first chromosome) were crossed with c42-GAL4 or Act5C-GAL4 flies, and expression of the chimeric protein was assessed (Fig. 5). These GAL4 lines drive expression of UAS constructs in either tubule principal cells [where salt is normally expressed (Fig. 4)] or ubiquitously, respectively. When CG2196-eYFP is driven in tubule principal cells of the main and lower segment by c42-GAL4, it is directed to basolateral membrane infoldings (Fig. 5, B and C). Since this promoter does not drive expression in stellate cells or in the initial segment of the tubule, no staining is seen in these.
With the Act5C-Gal4 promoter, the staining in the principal cells is exactly the same as for the c42 promoter. The progeny of the cross (Act5C-GAL4 flies) × (CG2196-eYFP flies) expresses the CG2196-eYFP construct in the principal cells of the main and lower segment again in the foldings of the basolateral membrane and perhaps in a part of the endoplasmic reticulum, while in the stellate cells of the middle segment no staining is observed (Fig. 5D). Thus salt is expressed in the ion-transporting cells of the Malpighian tubule, on the basolateral membrane, consistent with its presumed function as an uptake symporter.
Modulation of salt expression in tubules impacts on organismal survival after salt challenge.
Flies transgenic for a UAS-salt-RNAi construct were generated, and extent of knockdown was assessed by qPCR. UAS-salt-RNAi flies were crossed with the Act5C-GAL4 and c42-GAL4 driver lines, and expression levels of salt in Malpighian tubules compared with parental controls. The results of this experiment are shown in Fig. 6. The construct produces a significant (Student's t-test, taking P < 0.05 as significant) knockdown in salt expression, in both Act5C>salt-RNAi (18.8 times lower than parental RNAi flies), and c42>salt-RNAi (17.7 times less than the salt expression in RNAi flies and 10.7 times less than the salt expression in c42 flies).
Salt is upregulated in response to salt challenge, so does RNAi-mediated salt knockdown in just the tubule reduce survival of the whole organism when challenged with dietary salt? The progeny of both overexpression and knockdown crosses to the principal cell-specific driver c42 was raised on food containing 4% NaCl, and survival compared with parental controls. The results are shown in Fig. 7. As can be seen from these results, in both repeats overexpression of salt had no consistent effect on survival. Surprisingly, c42>salt-RNAi flies survive better in food containing 4% NaCl than either parental line (RNAi or c42). The percentage of RNAi-c42 flies surviving long enough is more than double of that of any of the parents, and this result is statistically significant (Student's t-test, two-tailed). So, surprisingly, the upregulation of salt on salt challenge observed by microarray and qPCR appears to be counteradaptive.
These results provide the most comprehensive view on salt stress in an insect to date. Previous studies associated salt stress (NaCl and KCl) with the inebriated neurotransmitter/osmolyte transporter (ine) (21), and the studies of dehydration and rehydration (2, 19) demonstrated that Drosophila melanogaster is a very strong osmoregulator. Here, dietary salt stress induced a clear and reproducible sequence of changes in gene expression involving immune/stress response, carbohydrate metabolism, and ion transport pathways. In the longer term (>16 h), effects analogous to starvation were seen, as the reproductive axis was shut down in both males and females. It is interesting to see sugarbabe, a transcription factor associated with dietary sugar loading, also changing in this study; it suggests that sug may play a general role in response to dietary stress.
Salt is a highly tubule-specific gene, expressed on the basolateral plasma membrane of the tubule, and which is strongly up regulated in response to dietary salt. However, this response is inappropriate, because c42>salt-RNAi flies actually survive better on high salt. The most probable reason for this behavior is that the overexpression of salt under salt diet is a miscalculated response of the organism. Such inappropriate responses are commonly encountered in human health and disease: for example, the cytokine storm elicited in healthy humans exposed to influenza can be more dangerous than the original infection. We speculate that flies might sense increasing Na+ levels associated with dietary salt stress and increase Na+ excretion by the tubule by upregulating basolateral Na+ uptake. However, it is possible that the osmolyte cotransported with Na+ by salt is not well handled by the tubule, and so RNAi against salt actually improves the situation.
What does salt/salty dog actually encode? Although it is evidently a member of the well-characterized SLC5 (sodium/solute symporter) gene family (43, 44), the answer is surprisingly vague. The Drosophila genome project (flybase.bio.indiana.edu) annotates it as a sodium/iodide symporter. An NCBI BLASTP search against human sequences produces a closest match (E value 3e-87) to SLC5A12, and a second extremely close match (E value 2e-82) to SLC5A8. The former cotransports short chain fatty acids, lactate, and nicotinate with sodium and is found in kidney, small intestine, and skeletal muscle; whereas the latter cotransports short chain fatty acids, lactate, nicotinate, and iodide with sodium and is found in thyroid, kidney, and intestine (43, 44). However, CLUSTALX alignment of the protein sequences with the human SLC5 members (Fig. 8) places salt between SLC5A6, a sodium/vitamin cotransporter, and SLC5A7, a sodium/choline cotransporter. Even this list is not exhaustive, as SLC5 members are also thought to facilitate the transfer of water and organic solutes such as urea. So, based on structure, salt can be considered to be a Na+/solute cotransport but cannot be ascribed a particular transport substrate. Clearly, then, the exact function of salty dog will require extensive further work, as functional workup of SLC5 family members is known to be problematic (43, 44), However, as SLC5 members are uptake transporters, and salty dog is localized to the basolateral membrane of the Malpighian tubule, its upregulation in response to dietary salt would clearly increase sodium clearance from the blood. Although this overexpression does not appear sufficient to confer enhanced survival on the organism on salt challenge (Fig. 7), the observation that knockdown of the gene is actually beneficial (Fig. 7) is intriguing.
Perhaps the most compelling aspect of the work is the clear phasing of different classes of response over the experimental period (Table 2), particularly the early induction, then repression, of an immune response. The tubule, although classically considered to be fluid-transporting and osmoregulating tissue (14), actually plays multiple roles in the organism (16). Recently, it has been shown to mount a robust, autonomous immune response to pathogens (28), mediated by the canonical Toll/imd immune pathways (22). So the machinery of the immune response is already in place in the tubules; it is intriguing that it is invoked so early in the salt stress response. We speculate that this finding implies a generality in stress signaling, at least in this tissue; irrespective of the precise nature of the stress, a set of key adaptive genes is quickly upregulated, then selectively turned off as the nature of the stress becomes apparent.
Of course, in nature Drosophila does not face quite the problems of excess salt as, for example, do mosquito larvae that live in hypersaline lakes. But, importantly, the results provide a baseline against which to assess the specialized responses of insects whose evolutionary history has equipped them with the ability to osmoregulate and thrive under continuous osmotic stress.
This work was supported by general funds of the University of Glasgow. The microarray experiment was funded by the UK Biotechnology and Biological Sciences Research Council's Investigating Gene Function initiative.
↵1 The online version of this article contains supplemental material.
Address for reprint requests and other correspondence: J. A. T. Dow, Integrative & Systems Biology, Faculty of Biomedical and Life Sciences, Univ. of Glasgow, Glasgow G12 8QQ, UK (e-mail: j.a.t.dow{at}bio.gla.ac.uk).
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 © 2009 the American Physiological Society


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Mycobacterium bovis Survivability in Salt/Minerals Fed to Cattle. John B. Kaneene , DVM, MPH, PhD Joseph Hattey , MS Carol Bolin, DVM, PhD James Averill, DVM, PhD. Introduction.
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Survivability in Salt/Minerals
Fed to Cattle
John B. Kaneene, DVM, MPH, PhD
Joseph Hattey, MS
Carol Bolin, DVM, PhD
James Averill, DVM, PhD
To determine the survivability of M. bovis on salt and mineral blocks over time under normal weather conditions over a 12 month period
Longitudinal Experimental Design
Samples are taken from the top face of the salt block using sterilized scouring pads on a specified schedule.
Sample Collection Schedule
Four Plain Salt and Four Mineral Blocks were used
Pure cultures of M. bovis and M. fortuitum were applied to the surfaces and kept outdoors in BL-3 confinement cages for a year
Samples from the block surfaces were collected at specified intervals and tested for the presence of the organisms
Michigan Department of Agriculture and Rural Development
Center for Comparative Epidemiology, Michigan State University

subset: commoncrawl


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|Indian Railways News =>
|Topic started by RailEnquiry Admin on Jan 10, 2017 - 14:10:31 PM
Title - Booking / Cancellation made easier on IRCTC AppPosted by : RailEnquiry Admin on Jan 10, 2017 - 14:10:31 PM
Now you can easily book, cancel and file TDR on the new IRCTC rail connect app. As of now this app was available as IRCTC connect which from now will be IRCTC Rail Connect. Till date only ticket booking were allowed through mobile app, however if you need to cancel or file TDR it was a cumbersome process.
Now IRCTC has connected App and its Website it will be easier to book or cancel tickets. You can either book your ticket on App or website and can cancel from any if required. Since now data is synced therefore same information will available on website and on App in your history. Earlier these facilities were not available due to which users often use website more for bookings.
Also once logged-in to the app you do not need to enter the ID repeatedly; it will just ask you the pin number to log-in multiple times.


text: You may have missed this story out of California last week. A student brought a shotgun to school with the intent of killing some other students who had bullied him. He shot one student, and two other students and a teacher were injured. What the headlines missed is that the teacher and a colleague managedRead More


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> Career Resources
> Interviews > Industry Updates > Banking / Finance > Money Moves
Leading bancassurance expert establishes friendships before finances
When it comes to enjoying a rewarding career, matching personal attributes with aspirations is often the golden rule. In order to truly leverage intellectual assets, it is wise to follow a logical professional path rather than battle to succeed in an industry which doesn't quite fit.
Simon Chung, insurance specialist head, Hong Kong, HSBC Insurance (Asia-Pacific) Holdings Limited lives by this maxim and 20 years ago began his career as an insurance agent in Canada, where many new immigrants from Hong Kong lived. At that time, the immigrant population rarely considered the importance of protection schemes or sought adequate coverage.
Mr Chung says the beauty of insurance is that: "It pays the most when you need it the most. Unfortunately, this concept is too intangible so many people ignore this important element when planning financially."
With his product knowledge and local experience, Mr Chung helped friends in the Chinese community in Canada set up basic retirement saving plans. He made home visits to potential clients, usually friends of friends, three to four times every week to extend his own people network.
"I enjoyed the moments when I got to know people and they developed from strangers into friends. Many people need a lot of assistance when they move to a new country, for example, how to complete tax returns. In Canada, I was delighted to offer support even though I was not rewarded financially for my efforts," he says.
Once trust was established, Mr Chung delved into the specific assurance needs of his friends, offering tailor-made suggestions to suit individual needs. Since many people were reluctant to discuss the importance of life insurance, or simply oblivious of its significance, Mr Chung would always raise the issue with the people around him.
"There are three important situations we should all plan for: retirement, premature death and disability," Mr Chung advises. "Generally, people are sufficiently prepared for their retirement days. However, the possibility of the other two situations, which have significant impact on their families, is commonly underestimated."
Subject to change
Despite the lack of full public understanding of the array of insurance services on offer, people are now more open to insurance products and services because bancassurance initiatives are growing at a tremendous rate. In response to client demand for comprehensive insurance products, banks have successfully promoted the notion of all-round protection to large client bases.
According to Mr Chung, bancassurance offers clients convenient access to a wide range of integrated financial services and product transparency. This in turn has resulted in higher overall levels of customer satisfaction.
"Most importantly, customers have confidence in banks," Mr Chung remarks. "This is particularly true for HSBC, an organisation that has established its reputation based on a strong foundation," he adds.
Unlike insurance companies, which traditionally target the narrow markets of their agents, bancassurers have the added advantage of a mass distribution market, which is effectively their existing clientele.
"Since our customers already understand and trust HSBC, they are often willing to discuss their protection needs when our staff approach them," Mr Chung observes. "It is the difference the brand name makes."
Likewise, it is the quality assurance system that gives HSBC insurance its elevated status. Indeed, as an internal measure to guarantee employee integrity, the company regularly assesses the wealth management solutions recommended by every sales person in the branch network.
"We have a responsibility to ensure our team members are offering the most appropriate solutions for each client, instead of concentrating on high sales figures,"
Mr Chung explains. "The QA process requires extensive manpower and resources, but it is worthwhile for our long-term benefit and for clients who have trusted HSBC," he says.
Expert team
For the foreseeable future, Mr Chung is concentrating on product diversification and training at the bank, in sessions which ensure bancassurance sagacity and expertise are shared with everyone in the HSBC team.
At the start of 2008, HSBC adopted a new business model for bancassurance. Today, when insurance services form part of a larger financial plan, branch wealth management experts take care of the whole deal. "Before this new business model was launched, financial planning managers were the dedicated insurance experts in the branches and their main function was to provide professional insurance services to the bank customers. Now, to enhance convenience and promote one-stop solutions to customers, our insurance specialist team will provide their branch counterparts with the necessary specialist knowledge in designing and delivering the best possible insurance and retirement planning solutions," he explains.
Currently, the insurance specialist team has some 10 members and this number is expected to grow three-fold within months. Candidates with at least five years of bancassurance experience are invited to join the expanding team.
"Integrity is a prerequisite for a truly rewarding career. A proven track record is often a great indicator of this," Mr Chung concludes. "Additionally, good interpersonal skills are essential, especially when training, mentoring and coaching other team members."
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Do you really think McNeil is going to find wrongdoing on the part of one of his homeboys? Nah, me neither.
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Posted on March 9, 2017
TALLAHASSEE — State Attorney Jack Campbell has launched an investigation into the alleged actions of the Tallahassee Mayor after receiving a complaint letter from a retired Jefferson County law enforcement officer and investigator.
Read letter here.
In his letter, dated March 2, Paul Henry, who said he is also a member of the Jefferson County Grand Jury, requested a Leon County Grand Jury investigate Mayor Andrew Gillum for allegedly purchasing "several thousands of dollars of political software via public funds" and possibly committing grand theft under Florida Statute (FS) 812.014 and official misconduct under FS 838.022
He said, "Andrew Gillum, in his capacity as Mayor of the City of Tallahassee, a public servant, did violate FS 812.014 (2)(c) "Grand Theft" and FS 838.022 "Official Misconduct" by using at least $3,915 of public funds to obtain personal benefit, to-wit political campaign software and falsified the justification for said software."
"In doing so," he continued, "Gillum deprived taxpayers of their right to this property (money) for no lawful purpose, and appropriated it for his own use."
Henry also wrote "Gillum had evidenced guilty knowledge in the matter by acknowledging the misuse of the money and offering to repay it."
He also conducted an online public records search of purchases to the NGP Van software and included his findings in his letter.
This letter was forwarded by Campbell to Leon County Sheriff Walt McNeil, who's office will conduct the investigation.
On February 23, Tallahassee Reports broke the story that Tallahassee Mayor Andrew Gillum had allegedly spent approximately $5,000 in city funds to purchase software from democrat campaign vendor, NGP VAN. That software was allegedly used in campaign email for Gillum in which he mixed "Office of the Mayor" letterhead with his campaign correspondence.
Do you really think McNeil is going to find wrongdoing on the part of one of his homeboys? Nah, me neither.
Exactly, McNeil used to work for the City. Was he still there when Gillum was first elected? If so, major conflict of interest…typical in this town, though.
Finally! Make him own it!
And thank you Mr. Henry!
I get the grand theft, but we can't dismiss the misconduct of office by using the office of the mayor for personal purposes. How do you pay that back?
The expenditure of public funds for a personal benefit is a crime AND an ethical violation. An admission and apology don't change that fact or establish a defense. We know the ethical standards of our mayor, now we're going to see what our new sheriff is made of.
Excellent investigative reporting – no one else has provided a copy of Mr Henry or Mr Campbell's letters. Good going TR!
Mis-appropriation and grand theft of government funds for the software purchase and use of his and one or more COT employees time in sending out over 60 emails for his personal benefit!
Good letter that accurately spells it out!
I believe FDLE would be the better entity to investigate as I see conflicts of interest with McNeil/LCSO/Gillum.
#1 Any local entity is too close for comfort.
#2 Sheriff McNeil was the prior chief of TPD which is part of the City of Tallahassee.
#3 McNeil and Gillum have a common political campaign advisor/friend (Sean Pittman). What if Pittman advised Gillum on this incident? The political associations between NcNeil and Gillum are a conflict.
#4 I believe FDLE would be better equipped to forensically do a data/IT/computer investigation. I believe all the commissioners emails and the entire city's system should be investigated for contamination, breaches, etc.
Who was responsible for deciding which entity did the investigation?
Your last question would make a great subject for a new story on this controversy. I too believe that it should be investigated by FDLE to avoid the obvious conflicts of interest.
Daniel's denial right before TR's presentation of irrefutable evidence tells me they knew what they were doing all along.
I'm very disappointed to hear Sheriff McNeil will be conducting the investigation of Gillum. This examination needs to be from an outside, state-run law entity rather than any city-affiliated agency. Somehow I doubt the Sheriff's investigation will result in any more than a wink and a cursory-but-meaningless slap on the wrist.
However, one can hope – and mine is that Mr. Gillum will find that he has torpedoed not just his fledgling gubernatorial campaign, but his entire political career also. Hopefully, Gillum's hard-leftist obama-wanna-be, Van-Jones-clone dreams will never come true, and his local career will be as well.
The Fox guarding the hen house.
This only came to light because he has a good chance of beating RICK SCOTT and because he's black and a FAMU Rattler! This bullcrap ass story is to stop a bigger and better plan!


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Friday, March 4, 2011
Freebie Quickpage :: Beary Lovely
I used the kit, Beary Lovely by Farrah's Creations for the quickpage.
She has it available in both full size and tagger size.
You can find them HERE.
Download link is below the preview.
Thanks for visiting!
3 comments:
- CBH said...
Thank you so much for this post. I want to let you know that I posted a link to your blog in CBH Digital Scrapbooking Freebies, under the Page 4 post on Mar. 05, 2011. Thanks again.
- March 5, 2011 at 9:48 AM
- CraftCrave said...
Just a quick note to let you know that a link to this post will be placed on CraftCrave in the DigiFree category today [06 Mar 12:00am GMT]. Thanks, Maria
- March 5, 2011 at 5:30 PM
This is a great QP. Thanks for sharing. Lola
- March 5, 2011 at 8:39 PM
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Daily briefs of the top news stories impacting Christians around the world.
In today's edition:
- Southern Baptists, Jewish Christians: God Gave Israel to the Jews
- Pope Delivers Lessons on Religion, Reason, Church Beliefs
- Doctor 'Steals' Dalit's Kidney In India
- Christian Faces Trial Over Bible-Based Pamphlets
Southern Baptists, Jewish Christians: God Gave Israel to the Jews
Both the spokesperson for the Southern Baptist Convention and Jews for Jesus ministry point to the Bible as the reason why a recent national poll found that the majority of the South believes God gave Israel to the Jews, The Christian Post reports. "The reason that 7 out of 10 evangelicals believe that God gave the land of Israel to the Jewish people is simply because it is stated as such in the Bible, and evangelicals believe the Bible to be God's Word," wrote SBC vice president for convention relations, Dr. Kenyn Cureton. Cureton expects the number to be higher among Southern Baptists. Late last month, the Pew Forum on Religion & Public Life released a survey that found 56 percent of those in the South believe that God gave Israel to the Jews. An even higher number (69 percent) of white evangelical Protestants subscribed to this belief. Jews for Jesus spokesperson Susan Perlman referenced Romans 11:29 which says "the gifts and his call are irrevocable" and concluded, "therefore so must our support for the survival of Israel be irrevocable."
Pope Delivers Lessons on Religion, Reason, Church Beliefs
Returning to the city where he once taught theology, Pope Benedict XVI offered a fundamental lesson in what the church believes and why it should proclaim the faith clearly in today's anxious and violent world. In a sermon before an estimated 300,000 people in Regensburg Sept. 12, Catholic News Service reports the pope said it was necessary to recognize the modern "pathologies" associated with reason and religion and the ways that "God's image can be destroyed by hatred and fanaticism." In light of these distortions, Christians need to "state clearly the God in whom we believe and proclaim confidently that this God has a human face. Only this can free us from being afraid of God, which is ultimately at the root of modern atheism. Only this God saves us from being afraid of the world and from anxiety before the emptiness of life," he said.
Doctor 'Steals' Dalit's Kidney In India
While undergoing surgery for pain in her uterus, a Dalit woman was robbed of her kidney by her doctor, says a Gospel for Asia release on ASSIST News Service's site. So far, according to news reports, her efforts at justice have been met with resistance and scorn because of her "untouchable" caste. Dr. K.P. Yohannan, president and founder of Gospel for Asia, says the case is another sad example of how India's Dalits are denied even the most basic rights. "Not only are they denied their rights, they are openly abused with no consequences for those who oppress them," he added. Maya was diagnosed in December 2003 with stones in her uterus. Her husband, Phool Singh, knew the impoverished couple did not have the money for her surgery. However, a friend referred Maya to a Dr. Richharia, who was able to get her in for surgery on December 20, 2003. But after she returned home, the pain continued. Her husband took her back for a checkup, but the doctor assured her that the pain would disappear after a few months. For almost two years, Maya continued to suffer. Finally, in January of this year, the couple went to another doctor for a second opinion. He suggested that she have an ultrasound. Those results showed that Maya's left kidney was missing! The next day, the couple went back to Dr. Richharia's clinic to inform him of their problem. He refused to see them, and the staff physically assaulted them for criticizing his reputation.
Christian Faces Trial Over Bible-Based Pamphlets
Evangelical Christians in Britain are concerned about the prosecution of a protester who was arrested at a homosexuality rally while disseminating leaflets quoting the biblical view about homosexuality, CNSNews.com reports. Stephen Green, who heads a small but active lobby group called Christian Voice, is scheduled to appear before a Cardiff court on September 28, after pleading not guilty to a charge of "threatening, abusive or insulting behavior." Although some Christian Voice campaigns and tactics have caused unease among some Christians, Green on this occasion won support. The Evangelical Alliance, a 160-year-old group representing more than a million British evangelicals, said it was worried that the case may "erode" Christians' freedom to proclaim their faith, as well as freedom of speech and religious liberties.


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Golf Team Continue to Win
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- Published: Tuesday, 30 June 2015 08:29
The Birkdale Golf Team returned triumphant from Surrey yesterday.
They had a number of notable results playing in the Independent Schools Golf Association (ISGA) HG Trophy competition at Burhill Golf Club.
The Team finished 2nd overall in the gross and won the Nett contest. Individual successes included Tom Bradley (L6), who finished 2nd in the U18s and star golfer Barclay Brown (S3/Y9) who won the U15s.
Many congratulations to all the team for an excellent performance. They are clearly a challenge to be reckoned with by the other teams!


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Guinness World Record Face-Off: Me vs. Mr. Giggles the Clown
April 16, 2012
In the 55 years that people have been setting and breaking
Guinness World Records
, we've seen some pretty epic displays of one-upmanship. Some of the most captivating talent categories have even become battlegrounds for intense rivalries. And just last week, a new rivalry was born.
Me vs. Mr. Giggles the Clown.
Not to be confused with countless other clowns who call themselves Mr. Giggles, this Mr. Giggles also happened to be the World Record holder for the 'Most High-fives in a Minute'. As someone with an incurable fear of huggers, the high-five has always been my go-to greeting. My technique a point of pride. And I've been wanting to break a world record, any record, for a very long time. In other words: it was on.
There were only three rules. You can't high five the same person twice. High fives had to be over the head and slap the hand. And if you missed a hand, you could slap it again. No problems there.
On April 19th, also known as
National High Five Day
, I got to work. My team at Spike set everything up in advance. A line of people waited with hands raised, and the Guinness World Record adjudicator was on-hand. 23 seconds later, I blasted Mr. Giggles into anonymity with a grand total of 107 high fives. His previous record? 101 high fives.
Katie Linendoll High-Fives Into Guinness World Records
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Bootylicious amateur nympho rides her boyfriend's cock in the gym
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Another great mating from Joyee Flynn
This story is the sequel to Midnight Matings: Skunk and a Trunk. White Knight and Black Fur is the story of Aldis and Gawain. Aldis is a skunk shifter with low self-esteem and an awesome job. He travels to Florida to check-up on and bring his cousin Dor his stuff. The first person he meets is Gawain, a rhino shifter with a lot of secrets and a long-time fantasy of sex with a skunk shifter.
This story has hot, steamy love scenes, conflict, and some knights from Camelot.
This is the second story of the McMurray Clan grouping. To keep following the McMurray Clan check out Unmated at Midnight: Rumble and Churr. Rumble and Churr [Unmated at Midnight] (Siren Publishing Classic ManLove)
White Knight and Black Fur [Unmated at Midnight] (Siren Publishing Classic ManLove)
[Siren Classic ManLove: Erotic Alternative Paranormal Romance, M/M, shape-shifters, public exhibition, HEA] When his cousin is forced to mate by UPAC, Aldis uses the excuse of bringing Dor's stuff to his new home to check and make sure everything is really okay. What he didn
White Knight and Black Fur [Unmated at Midnight] (Siren Publishing Classic ManLove) is one of best selling in Erotica category.


text:
VEGH Karoly wrote:
On Thu, Oct 02, 2003 at 07:08:23AM -0600, skippi wrote:Greetings all.I have acquired a new motherboard. It's an Intel D865GBF. The onboard video controller is Intel Extreme Graphics 2, 82865G Graphics and Memory Controller Hub, 865G chipset. Is this supported under the Xfree that comes with Woody? Or am I going to be having to do lots of building from source? I have found ahave a look at: http://support.intel.com/support/graphics/linux/graphics.htmIt says you'll need X 4.3, but it also said this to my i845G VGA, and yet it works fine on my SID:
Just in case you end up needing X 4.3, here is the source that I use: deb http://penguinppc.org/~daniels/sid/i386/ ./That is probably your best bet. Otherwise, I would recommend downloading the sources from:
http://necrotic.deadbeast.net/xfree86/I would get, xfree86_4.3.0.orig.tar.gz, and the latest sid branch (updated daily). Unpack them and, assuming you have the debian package building tools installed, go the parent directory of the source tree and issue a dpkg-buildpackage. Go to sleep or watch a movie, and in a few hours you have yourself a set of Debian packages for XFree86 4.3.
The good thing about this approach is that you don;t have to worry about confilcts once the official packages make it into Sid. If you have installed a tarball from xfree86.org, it is possible things will get srewed up when you try to upgrade later.
Just my $.02 -Roberto
Description: PGP signature


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Sense of humor, his smile, his wit, intelligenceAnd his dick. Okay had to say it.
I'll tell you what I find sexy:The curve of a woman's hip.Her eyes.Her lips.The back of her neck, which I love to kiss.Her lower back.A smooth, shapely behind and legs.I also find a confident woman very sexy. And certain accents turn me on like crazy (my wife is a Latina).
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prepkid:See the TER link below....
Sense of humor, his smile, his wit, intelligenceAnd his dick. Okay had to say it.
Call me LOLOLOL
I kinda like a freaky girl, one that wears something erotic. I prefer some form of sexy lingerie, specifically a fishnet outfit that has accessible holes to her vagina and butthole. A good example is, when I step inside, she's already into it, playing with herself with toys, making good moans, and setting the evening to a memorable session.A girl that's really into it knows how to set the atmosphere and put on a climactic performance that will win five stars from the theater. That is sexy to me. She convinced me that she is in to me, and puts on a good effort or act to please me.
Genuine laughterIntelligent conversation with an opinionEye contactConfidenceA come hither look
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text: New York 9780670012435 0670012432 Viking Denenberg Barry Barry Denenberg Shipwrecks--North Atlantic Ocean 2017-11-08 Titanic (Steamship) Shipwrecks--North Atlantic Ocean Titanic (Steamship) North Atlantic Ocean. Shipwrecks Juvenile works en Experience the Titanic's doomed voyage in this unique presentation of fact and fiction Notable Social Studies Trade Books for Young people, 2012 Titanic sinks! Presents the true story of the building and launching of the Titanic and offers a "you are there" feeling when the steamship struck an iceberg and sank on her maiden voyage. 706018465 A letter from the publisher / [Mary Hudson] -- White Star names new luxury liner Titanic : original article, Monday, April 27, 1908 -- Fifteen-year-old dies in fall from Titanic's hull : orginal article, Monday, April 25, 1910 -- Titanic launch dazzling success! : original article, Monday, June 12, 1911 -- Captain Smith takes command of the Titanic : original article, Monday, April 1, 1912 -- "Queen of the seas" : original article, Monday, April 8, 1912 -- Sailing day! : original article, Monday, April 15, 1912 -- Transcribed from the journal of S.F. Vanni, chief correspondent, Modern Times -- The Titanic hour by hour : original article, Monday, April 22, 1912 -- Firsthand accounts from the actual survivors in the lifeboats : original article, Monday, April 22, 1912 -- "Captain Courageous" : original article, Monday, April 29, 1912 -- A closing note from the publisher -- A word from the author. Historical fiction Unknown 'gen' value: ejh Middle School. 2011 2011 706018465 nyu


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Flag Floor Stand Flagpole Flagstaff Cast Iron Military 1 1/4" Pole EUC
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Excellent used condition Military Issue Flagstaff Stand.Features: Size: 12" diameter National Stock Number (NSN): 8345-00 Weight: Approximately 19lbs. Fits 1 1/4" flagpole Snap & button fly Cast Iron Polished gilt finish
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text: Fantasy Flight Games premiered the Game of Thrones board game at Gen Con, with much success! According to their reports, they've sold out all of the copies of the board game they had with them. It should also be appearing in game shops by now. A review of the game can be found here.


text:
The policewoman accused in a Newcastle upon Tyne court of being a £100-an-hour prostitute is alleged to have had up to 20 clients a week, not 20 a day (Page 20, December 9).
The Daily Mirror tries to correct errors and clear up confusion as soon as possible. Readers should phone 020 7293 3953, fax 020 7293 3975, email firstname.lastname@example.com or write to Readers' Editor, Daily Mirror, 1 Canada Square, Canary Wharf, London E14 5AP. The Press Complaints Commission (PCC) is the independent body which governs the newspaper industry. It has a Code of Practice we adhere to. This, along with details on how to complain, is available from Halton House, 20/23 High Holborn, London EC1N 2JD. Website www.pcc.org.uk or email email@example.com Telephone: 020 7831 0022 Helpline: 0845 600 2757.


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Henrik Ibsen Early life. Early Early Life. March 20, 1828 - 23 May 1906 T he oldest of five children Born to Marichen Altenburg and Knud Ibsen Wealthy family Bankrupt. Work and Travel. Family life influences work At fifteen, Ibsen forced to leave school Child at 18
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By Josh Walsh
By Josh Walsh
By: Joel Johnston
Lucy Stone
Susan B. Anthony
Elizabeth Cady Stanton
Feminism is defined as "the doctrine advocating social, political, and all other rights of women equal to those of men."
In the 1800s, and even into the1900s, women were not treated equally to men.
Although one might think that only women support feminism, men also supported this, though to a lesser extent.
Before the 1840s, even women graduating was unheard of. The first woman recorded graduating from college was in 1840.
Defined as "a body of philosophies and ethical perspectives that emphasize the value of human beings, individually andcollectively, and generally place more importance on rational thought than on strict faith or adherence to principle."
The Humanism movement is attributed with moving scientists as well, as many began to call themselves Humanists in the 19th century. It also likely had a part in developing more atheist movements, as it had a part in the questioning of faiths.
"I am not even quite sure what women's rights really are. To me it has been a question of human rights"
Ibsen likely used Nora as an "everyman," not just as a female example.
Since women were oppressed at the time and not given equal rights, they could be seen reasonably as a good example of the humanist view.
However, feminism was not only fueled by the Enlightenment. Many Christians believed in equality as well. These feminists insisted that women's rights were God-given and should not be curtailed by human custom or law.


text:
- Search CSN:
- Members: Login to search all areas
- Not a member? Click here to search public areas
Jun 04, 2005 - 6:24 am
Has anyone had, or heard about repair of the eustachian tube(s) after radiation treatment? Still with fluid filled ears 2 years out of treatment. Also, a neck that is becoming "wooden" to the touch. Thanks for any advice.
Jun 13, 2005 - 2:02 pm
I too have fluid filled ear 1 year, 3 months after last rad treatment. Since then have had tube inserted (did no good) and had mastoid surgery. Was told after last surgery there was nothing else they could do for my ear/hearing. My neck is also tightening and getting a wooden feeling. Apparently these are the side effects that no one tells your about. Sorry I can't help, but know you aren't alone out there.
Jul 07, 2005 - 11:19 pm
Hi, an update to the first message. ENT told us tubes are too far into the face to repair. There was a sinus infection along with the ear fluid. Augmentin has stopped the sinus problem. Waiting on the ear thing. Thanks to All. Hang in there, we will try to answer anything we have experience with.


text:
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21 January, 2015
Oracle Bets Big On Cloud Computing Services
Dubai: Oracle, like other technology peers, is betting big on cloud computing as it transits from a hardware supplier and developer to a complete cloud solutions provider.
Even though cloud offerings posted a robust growth of 140 per cent in the second quarter, Oracle's dominance in the database market is not exactly leading to its dominance in the cloud space as it still accounts for around five per cent of overall revenues.
"Our strategy in cloud is working and we are going to invest more. We had a tremendous growth in cloud offerings in the Middle East. We have added 300 new and expansion software as a service [SaaS] cloud customers in the first quarter and second quarter," said Oracle CEO Mark Hurd, who was in Dubai on Tuesday.
"We are in a very early innings and in a transition phase. The second quarter SaaS growth was 80 per cent in Europe, Middle East and Africa (EMEA)," he said.


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Leidos to Present at the J.P. Morgan Aviation, Transportation and Industrials Conference
Live Audio Webcast Available on March 8th from 8:00am to 8:40am ET
Mar 03, 2016, 16:15 ET from Leidos Holdings, Inc.
RESTON, Va., March 3, 2016 /PRNewswire/ -- Leidos Holdings, Inc. (Leidos) (NYSE: LDOS), a national security, health, and engineering solutions company, will present at the J.P. Morgan Aviation, Transportation and Industrials Conference in New York City.
James Reagan, Chief Financial Officer, will present at 8:00am Eastern time on Tuesday, March 8th, 2016.
A live audio webcast of the event will be available on the Leidos Investor Relations website at http://investors.leidos.com, where related materials will be posted prior to the presentation. A replay of the webcast will be available following the presentation at the same link listed above for 90 days afterward.
About Leidos Leidos is a science and technology solutions leader working to address some of the world's toughest challenges in national security, health and engineering. The Company's 18,000 employees support vital missions for government and the commercial sector, develop innovative solutions to drive better outcomes and defend our digital and physical infrastructure from 'new world' threats. Headquartered in Reston, Virginia, Leidos reported annual revenues of approximately $5.09 billion for the twelve months ended January 1, 2016. For more information, visit www.Leidos.com.
Investor Relations:
Kelly P. Hernandez 571.526.6404 firstname.lastname@example.com
SOURCE Leidos Holdings, Inc.


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- HubPages»
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Minnesota Timberwolves Career Point Scoring Leaders
Kevin Garnett
Sam Mitchell
The Minnesota Timberwolves have only been an NBA franchise since 1989, and have made the playoffs eight times with zero championships to date. The team has mostly struggled, but has a few times put some quality teams on the hardwood. Here is a short list of the T-Wolves' all-time leading scorers in their (to date) brief history:
Kevin Garnett (19,041)—Forward Kevin Garnett began his career in Minnesota and played for the Timberwolves from 1995 to 2007. He was basically the face of the franchise for most of their existence to date. Garnett leads the T-Wolves all-time with 19,041 total points, and he scored over 1500 points eight times. Garnett's best season was 2003-04, when he had a career high 1987 points (24 per game), which also led the NBA that season. That year, Garnett also led the league in field goals (804), field goal attempts (1611), and also defensive rebounds (894) and total rebounds (1139).
Sam Mitchell (7161)—Forward Sam Mitchell had two stints with Minnesota, from 1989 to 1992, and again from 1995 to 2002. Mitchell ranks second in franchise history with 7161 points, and his best season was 1990-91, when he scored a career best 1197 points (15 points per game).
Wally Szczerbiak (6777)—Forward Wally Szczerbiak began his career for the Timberwolves, and played there from 1999 to 2006. Szczerbiak ranks third in the team's history with 6777 total points, and his best season was 2001-02. That year, Szczerbiak scored a career high 1531 points (19 per game), and also had a career best three-point shooting percentage (.455).
Doug West (6216)—Guard/forward David West played for Minnesota from 1989 to 1998, and he ranks fourth in Timberwolves' history with 6216 total points. West had his best season in 1992-93, when he scored a career best 1543 points (19 points per game).Tony Campbell (4888)—Forward/guard Tony Campbell played just a few seasons in Minnesota, from 1989 to 1992, but in that short time racked up 4888 points, which ranks Campbell fifth in franchise history for scoring. Campbell had the three best seasons of his career for the T-Wolves, and his best one was 1989-90. That year, Campbell scored a career high 1903 points (23 per game).
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33 outdoor bench plans - free diy plans, An outdoor bench is an excellent addition to any landscape or garden project. they can be used to delineate space, and provide a tranquil area for reading and relaxing..
Ana white | outdoor storage bench - diy projects, Build two side panels as shown above. the easiest way to do this would be to start with one of the legs. mark the leg 1 1/2″ in (you could just use a 1×2 since it.
Plans To Build Outdoor Storage Bench
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How to build a shed - free shed plans - build-it-yourself!, Last updated dec. 26, 2016 this page contains information on how to build a shed and storage shed plans. here are a couple of things to consider.


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This bridge is located 7 miles northeast of Rogue River on East Evans Creek Road (turn right on Covered Bridge Road)
[this section is currently under maintenance]
The mission of this organization is to ensure that the historic covered bridge, which had stood since 1892, is replaced in all its original glory. We will work to fairly represent the community of Wimer and to be a gathering point for comments, concerns, and fund-raising opportunities. We plan to work closely with the county and other organizations to ensure that the funds necessary are acquired and the re-building proceeds in a timely manner.
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No, this is not how to grow udders and look fashionable like Bessie the Cow. (although udders Are pretty swanky)
Have you ever felt like your Diet/Nutrition and Exercise Program was less effective than it could be? -That your effort in was not equal (!=) to the results you got out? -Like you were wasting time?
You Were wasting time.
One of the many small "Dieting" Gems overlooked by many:
Any minute or second of the day spent with your blood sugar at less than optimal was Wasted Time on your program.
The main culprit: 3 big meals in the day; and possibly unevenly spaced.
A better strategy to keep your metabolism rolling from the moment you get up until just before bed is to chop your Total Calories (you know exactly how much you should be eating for your weight, lean body mass, etc. each day, –right?) into:
5 Smaller Meals spaced at ~Equal Intervals throughout the day.
More Consistent Nutrition/Blood Sugar == More Consistent Metabolism == More Consistent Fat Loss (all things being equal).
If you're drinking the right amount of water and you've gotten hungry, or godforbid, Ravenously Hungry -> Yup, you guessed it: your metabolism has already partially gone into Starvation Mode. (don't let that happen)
Use your time wisely and try 5 smaller meals instead of 3!
Note: Please make sure you're not using 5 meals as an excuse to eat more than you should. "Dieting" in quotes because diets really don't work.
PS: Do yourself a favor and Rinse Your Teeth off by swishing water around your mouth after each meal. Your dentist will thank you.
Photo Credits:
"Highland cow", by Shelley Cunningham
"cow", by Diego Medrano,


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a recording anemometer.
Historical Examples
The hut is towards the left-hand side and the anemograph is on the hill.
The Home of the Blizzard Douglas Mawson
The anemograph screen was blown over and smashed beyond all repair.
The Home of the Blizzard Douglas Mawson
In a seventy-five-mile wind, Hodgeman had several fingers frost-bitten this morning while attending to the anemograph.
The Home of the Blizzard Douglas Mawson
a self-recording anemometer
An anemometer that uses a recording device to document wind speed and direction continuously.
Tagged: a
Read Also:
- Anemology
the study of the movements of the winds. noun (archaic) the study of winds
- Anemography
noun (meteorol) the technique of recording wind measurements
- Anemometer
any instrument for measuring the speed of wind. Historical Examples The anemometer is an instrument for measuring the force and velocity of the wind. Orthography Elmer W. Cavins This is done by the familiar weather-vane and the anemometer. St. Nicholas Magazine for Boys and Girls, Vol. 5, July 1878, No. 9 Various A little way […]
- Anemometry
the science of measuring the speed of wind. noun (meteorol) the technique of measuring wind speed and direction


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In this ticket we propose an addition to the GRIDSPEC (https://cf-pcmdi.llnl.gov/trac/ticket/63) convention to allow it to express multiple stagger locations at once. The multiple staggers are expressed by allowing GRIDSPEC tile files to contain multiple copies of each tile each with a slightly different size from the original. There are two issues that this proposal covers. The first is how to store the stagger coordinates. The second is how to put a variable on a given stagger.
To store the stagger coordinates we propose allowing more than one copy of the coordinates in the tile file. The coordinate variables and dimensions for the center stagger are present as usual. The staggered versions of the coordinates are also in the file. The attribute "gridspec_stagger" attached to the staggered coordinate variables is used to indicate their stagger. A set of standard stagger names used in this attribute could be added to CF.
The following is an example of a tile containing center and corner coordinates:
netcdf tile1 {
nx = 45;
ny = 45;
nxCorner = 46;
nyCorner = 46;
double lon(nx, ny);
lon:standard_name = "longitude;
lon:units = "degrees_east";
double lat(nx, ny);
lat:standard_name= "latitude";
lat:units = "degrees_north";
double lonCorner(nxCorner, nyCorner);
lon:standard_name = "longitude";
lon:units = "degrees_east":
double latCorner(nxCorner, nyCorner);
lat:standard_name = "latitude";
lat:units = "degrees_north";
When referencing a staggered version of the coordinate data the user does the typical thing and uses the "coordinates" attribute to specify which of the coordinates in the tile file the data is on. This in addition to the global variable gridspec_tile_name already in the GRIDSPEC convention gives the information needed to access the correct coordinates in the correct tile file. If the user needs to know which stagger their data is on, they can find that by looking at the "gridspec_stagger" attribute attached to the coordinates.
The following is an example of putting a data field (Q) on the corner stagger.
float Q(nxCorner,nyCorner);
Q:coordinates="lonCorner latCorner";


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Pondering nearing 50? THAT IS A situation You Can't Miss TO CHECK OUT
About author
Comments are closed.
It is important to check out a healthy diet plan after 50 to deal up with the changes the body faces. Read this article to learn about the sorts of ingredients and the total amount to be studied as you age.
If you are given above 50 or nearing that, then it's time to keep count on the sorts of food item you eat. There will be a whole lot of changes that you may see for yourself in your daily life, and it's essential that you take about some minor advancements in your eating habits.
The increase that you may need after 20 is very much indeed numerous than what you for truly have already been acquiring your lifestyle. You needn't get which entails any drastic improvements requirements food habits.
All you have to accomplish is to improve the amount of well-balanced meals that you take in gradually. Growing older is an all natural process, nicer looking a part of it; there will end up being a great deal of physical and physical alterations that may happen to your own.
These changes might impact your organs invariably, in fact, it is better if you realize begin taking some swift and the while you are nearing 50.
You might find a rise in muscle lack and a stark difference impurities redistribution of surplus fat. It'll grow to be difficult to keep the physical body weight and therefore, the health.
Read further to learn about different varieties and are food to come to become when you near to the half dollar bill you will ever have in addition to the right total be consumed.
Comments are closed.


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Discussion in 'Engine Compartment' started by sungoa2010, Jul 20, 2011.
Total capacity of cooling system of linea is approx 7.5 ltr
I have observed that coolant dries up faster if it is over diluted, that is, if we keep adding only distilled water. It doesn't dry up faster if we add the 60-40% mixture. I don't know whether this has any logic, but seems true. May be because of this, the engine heats too much thereby triggering the "noisy fan" too frequently.
Though TASS people keep saying "we have added concentrated coolant, you keep adding water when the level goes down", I decided not to dilute the coolant anymore and during my recent long trip, I bought a litre of coolant at KVR dream vehicles, kannur.
Praflu is available in sealed one litre bottle. Cost Rs. 244.00
fiatlover +1 to that, i totally agree to your point. Coolant level drops quickly if it is over dilluted.
Mods Note - Threads Merged.
Most TASS fill the full coolant say as in my case 4 ltrs of paraflu concentrated and then add water after running for some Kms and so on till the level stabalise.So i must have added around 2.5 litres of water.Full 7.5 liters mixtures doesn't go inside as some coolant/water remains in the radiator even after flushing.
Very true amit.I recently got my 3rd service done.Cost was Rs312/- since my car has only run 8000kms so no major work only changed Air filter and Ac filter.My OEM filter had a black spongy type material covered on it but its not on the new one that i changed.Cost cutting?? Or i got fooled??
the black sponge is the foam, the filter has foam + paper element.
i will have to check whether on the new filters this foam is missing. strange though. it traps a lot of dust.
OEM filter that came with car had foam in the bottom while replacement OEM filter's do not have foam at the bottom they have a 1mm cotton instead.
It have been the same from 2009, don't know the reason behind this as new car still come with foam at bottom but replacement filter come without foam.
OEM replacement filter's are very poorly made with irregular sealing gasket height, in the last service I had to reject 4 filters for same reason.
Usually TASS people take one or two day for servicing. So we cannot be there while they do the work. How do we then trace about the kind of work that people are doing. For example how do we know that they have actually changed the filter.
Amit im planning to change to sum after market Air filter.Hate the current ones.Requires frequent cleaning.Any suggestions??
Separate names with a comma.


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Beowulf. Terms to Define :. Thanes – retinue of fighting men gathered by the king to protect him, warriors Mead – fermented drink made from water, honey, malt and yeast
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Youthful, prime Old age
Proud, arrogant Humble
Courageous Unable to take


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Once the Krippendorf-Dittman Shoe Company, this loft conversion features 17 unique floor plans showcasing open loft spaces with features including gas fireplaces and ceilings as high as 20 feet. Building has floor-to-ceiling windows, incredible Fitness Center and a Tavern/Pub Room, laundry facilities, wireless internet, covered parking, secured entry and Guest Suite. Service based amenities including maid service and valet dry cleaning. Pet friendly.
634 Sycamore St.
Copyright © 2017 Downtown Cincinnati Inc. All rights reserved.
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Programming the Science of Crystallography. PLATON, a Multipurpose Crystallographic Tool A.L.Spek, Utrecht University. How to find the data belonging to a given reflection hkl in a linear list of reflection data. Reflection data assumed in array REFL(I,J) Option #1: linear search
Download Policy: Content on the Website is provided to you AS IS for your information and personal use and may not be sold / licensed / shared on other websites without getting consent from its author.While downloading, if for some reason you are not able to download a presentation, the publisher may have deleted the file from their server.
PLATON, a Multipurpose Crystallographic Tool
A.L.Spek, Utrecht University
Reflection data assumed in array REFL(I,J)
Memory efficient but time inefficient
Addresses array requires worst case dimensioning
Array element ADDR(H,K,L) contains the address I
for the data in REFL(I,J)
No worst case dimensioning needed


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Hi, my name is Dilara. I received my Bachelor's degree from the University of Wisconsin-Parkside in which I studied history and some political science. I received my teacher certification from Trinity International University which I studied secondary social studies and became licensed to teach social studies for grades 6-12. I later added my secondary special education license from Roosevelt University.
I am a licensed teacher in the areas of social studies and special education. I ha... ... More
Hello. My name is Shane and I am currently hoping to tutor college students in beginning and upper level math classes. This includes pre-technical math, technical math, prealgebra, algebra, geometry, trigonometry, and calculus. In the future, I am hoping to move forward to tutoring differential equations, statistics, linear algebra, and some abstract mathematics.
My experience in mathematics is as follows:
In high school, I ended with AP calculus and scored a 5 on the AP exam. I had a v... ... More
As a law enforcement officer for more than 20 years, I can help you understand the laws of Wisconsin, and law enforcement in Wisconsin. I am currently an investigator with a local Law Enforcement agency in SE Wisconsin. I have a B.S. degree in Criminal Justice from UW-Platteville. ... More
I am Wisconsin state certified through the Department of Public Instruction in these grade levels up to 8th grade. If you require personal confirmation of that fact you may find the public information there on my licensure on the Wisconsin DPI website. ... More
I am a graduate of Illinois State University, certified with a Type 03 certificate, and specialized in mathematics, language arts, and reading. I currently work at a middle school. Last year, I worked as a long-term sub for a 7th grade classroom, teaching regular math, Pre-Algebra, and Algebra 1 (a Freshman-level course). This year I help out with Algebra 2 as well. Although my degree is focused on middle school, I am more than able to help out with K-9! I have worked with a pr... ... More
ACT English is an important tool for gauging a student's knowledge and ability to use this language. It also forms one of the core scores that college and university acceptance committees look at when considering applications. Good ACT totals are therefore necessary, especially when higher education is planned. They also can make a difference in upper levels of business. It all comes down to communication--the better the student's English acuity, the better her or his future is likely to be. ... More


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Painting & Decorating in Rochester (Kent)
Are you looking for experienced painters and decorators in Rochester?
SJ Group offers a professional painting and decorating service to customers in Rochester that can completely transform your property. Our decorators wealth of experience in painting and decorating ensures our customers always receive an unrivalled service. Our commitment to quality guarantees that flawless results are always achieved.
Before any work is undertaken, we will have an in-depth discussion about the requirements for the room or rooms being decorating. When required, we can offer our professional advice for the work if you need any ideas for styles and colours. Our commitment to providing a personal service ensures that our customers are always delighted with our service.
From painting your bedroom to decorating your entire home, you can rely on our professional painters to provide a smooth, stylish and professional service. It can be a disruptive process when having your home decorated, therefore we endeavour to leave your home in a reasonable condition after each days work.
Our painting and decorating service includes:
All types of paper hanging
Wood Graining
If you're looking for a professorial paint job, we are one call away, we have all the equipment for interior and exterior decorating. Are you looking for experienced painters and decorators in Chatham or Gillingham?
Get in contact with us for a FREE, no obligation quote and advice.


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A scientist receives a letter from a colleague but their plans are adverted when the plane's engines fail, stranding our protagonist in the wilds of the jungle. Will you survive?
- Decent level design
- Good story line (at least to begin with)
- Some unique things
- Huge lack of story
- The only scares in the story are chase scenes pretty much
- The puzzles were most of the times very easy and boring
"Best custom story award".
How?! Okay, it's from 2011 and stories are generally much better nowdays as both the authors and the level editor improves, but still, this one as the best story?
First of all, it feels like you guys made a story line which were decently good, but after just 1-2 maps you completely gave up on it and focused only on the maps and added a chase scene in half of them.
The maps are nothing really special. Most of them are very short, undetaild and just looks pretty bland in generall and the puzzles and most of the scripts are pretty simple and boring.
On a side note, when I saw pictures of a jaguar monster and heard a jaguar sound in the forest map, I kinda expected a custom monster to appear. I was wrong. A bit disapointed but on the other hand you didn't say you added a custom monster.
I do understand why this story is so unique and popular, but to me it's another decent story. It's not bad, but it's not special either.
6/10 - Decent
Really god, but nothing Extraordinary.
Very impressive mod. The atmosphere is excellent, with several very creepy sequences and scary moments. The story line and puzzles were also very unique. Ending wasn't the best though,
This is an amazing custom story I rate it 1000/10 I highly recommend you play this because it is hard for me to find a good custom story and this is not good, not great, not amazing, but spectacular
A very good mod for sure. It certainly has some pretty scary moments, and I liked the story and puzzles as well. It was a little short, and I wouldn't say it did anything extraordinary, but it's still a highly enjoyable mod that deserves your attention.
There are better mods out there, but this one's still pretty awesome.
Wauw, for 2011 custom story, its worth playing it! Its a okay-good story and cool athmospere! It is worh your time, check it out!
This is a great story! It deserves the best custom story award.
This custom story its one of the earliest examples of good map designing and good story. The atmosphere is good, and there are little to no jumpscares and thats very good. The story its extremely well crafted, but could be a little deeper and a lot creepier.
The notes and mementos are fine, the voice acting its interesting.
Were this mods fail is to deliver the eeriness of the so called aztec temple, if the developers could use a little more aztec imagery to reflect the culture, they would have reach a far better mood of dready isolation in a foreing and strange land. The assets are the same as the A:TTD, and this is good, couse it shows that a good story doesn't need for custom content.
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No body can tell you with certainty what will happen.
According to the National Housing Federation forecasts, house prices
in England will fall this year and next (2010) before recovering and stabilising in 2011. The National Building Society on their part has said there is a reasonable chance, house prices could end the year higher than they started in 2009. Still another group the Royal Institution of Chartered Surveyors, said its members in Wales said they expected house prices to rise and the number of house sales to increase. who to believe?
Heh, think about the barbecue summer we were promised this year before you believe any predictions about house prices and bear in mind long term forecasts are unreliable.
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