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Objectives: Myostatin, a member of the transforming growth factor-beta (TGF-β) family, plays a key role in skeletal muscle myogenesis by limiting hyperplastic and hypertrophic muscle growth. In cardiac muscle, myostatin has been shown to limit agonist-induced cardiac hypertrophic growth. However, its role in cardiac hyperplastic growth remains undetermined. The aim of this study was to characterise the expression of myostatin in developing myocardium, determine its effect on cardiomyocyte proliferation, and explore the signalling mechanisms affected by myostatin in dividing cardiomyocytes.
Methods: We used quantitative PCR and Western blotting to study the expression of myostatin in cardiomyocytes isolated from rat myocardium at different developmental ages. We determined the effect of recombinant myostatin on proliferation and cell viability in dividing cardiomyocytes in culture. We analysed myostatin's effect on cardiomyocyte cell cycle progression by flow cytometry and used Western blotting to explore the signalling mechanisms involved.
Results: Myostatin is expressed differentially in cardiomyocytes during cardiac development such that increasing expression correlated with a low cardiomyocyte proliferation index. Proliferating foetal cardiomyocytes, from embryos at 18 days of gestation, expressed low levels of myostatin mRNA and protein, whereas isolated cardiomyocytes from postnatal day 10 hearts, wherein the majority of cardiomyocytes have lost their ability to proliferate, displayed a 6-fold increase in myostatin expression. Our in vitro studies demonstrated that myostatin inhibited proliferation of dividing foetal and neonatal cardiomyocytes. Flow cytometric analysis showed that this inhibition occurs mainly via a block in the G1-S phase transition of the cardiomyocyte cell cycle. Western blot analysis showed that part of the mechanism underpinning the inhibition of cardiomyocyte proliferation by myostatin involves phosphorylation of SMAD2 and altered expressions of the cell cycle proteins p21 and CDK2.
Conclusions: We conclude that myostatin is an inhibitor of cardiomyocyte proliferation with the potential to limit cardiomyocyte hyperplastic growth by altering cardiac cell cycle progression.
1. Introduction
The proliferative capacity of mammalian foetal cardiomyocytes diminishes dramatically during early neonatal development [1–3]. From this point onward, most of the adaptive growth needed to accommodate increased cardiac workload is accomplished by myocyte hypertrophy [3]. The inability of mature cardiomyocytes to proliferate, but instead grow by hypertrophy, has serious consequences for patients who suffer from heart failure post-infarct [4] or genetic cardiomyopathies that lead to progressive fibro fatty replacement of cardiac tissue, such as arrythmogenic right ventricular cardiomyopathy [5]. Identifying the key regulators of cardiomyocyte proliferation and understanding the molecular mechanisms involved in limiting the ability of cardiomyocytes to divide, therefore, is crucial if we are to develop new therapies to treat such heart diseases.
The loss in the ability of differentiated adult cardiomyocytes to divide has been the focus of interest for a number of researchers [6,7]. During cardiac development, it has been observed that changes in the proliferative capacity of cardiomyocytes coincide with a biochemical block in cardiac cell cycle progression leading to terminal differentiation [2,8]. In an attempt to stimulate cardiomyocytes to undergo cell division and repair damaged myocardial tissue, considerable efforts have been focussed on identifying putative regulators and molecular mechanisms that alter the expression and activity of cell cycle molecules involved in myocyte proliferation [7].
Myostatin, also known as Growth and Differentiation Factor-8, is a member of the TGF-beta (TGF-β) super family of secreted proteins and a potent inhibitor of skeletal muscle mass [9]. Studies in cattle and mice [9,10] have demonstrated that absence or impaired functioning of myostatin results in hyperplastic and/or hypertrophic growth of skeletal muscle, whilst increased levels of myostatin in the serum and muscles of patients with cachexia and HIV infection is linked to the muscle wasting disease seen in these patients [11]. Recently, a naturally occurring myostatin mutation has been identified in humans such that a male patient showed increased muscle mass compared to other boys of his age [12]. Together, these studies demonstrate that myostatin affects both hyperplastic and hypertrophic skeletal muscle growth. Myostatin mRNA and protein are also expressed in cardiac muscle [13] and it has been shown that myostatin affects adaptive hypertrophic growth in serum-starved neonatal rat cardiomyocytes [14]. However, the effect of myostatin on cardiomyocyte proliferation remains undetermined.
To study the role of myostatin in cardiac muscle proliferation, we firstly determined its expression in foetal cardiomyocytes with high proliferation index, in neonatal cardiomyocytes with lower proliferation index that decreases with age and in adult rat hearts wherein the majority of cardiomyocyte have lost their ability to divide [1–3]. We observed that myostatin is expressed developmentally. Our in vitro studies demonstrated that myostatin limited cardiomyocyte proliferation and caused cell cycle arrest. Myostatin also induced SMAD2 phosphorylation and altered the expression of cell cycle proteins critical for S-phase entry in proliferating cardiomyocytes. These findings highlight the potential of myostatin to limit cardiomyocyte proliferation and suggest that this inhibition involves SMAD2 signalling and cell cycle regulation.
Materials and methods
Wistar rats were obtained from Charles River Laboratories, UK. All animal experiments complied with the United Kingdom Animal Scientific Procedures Act 1986 as well as with the Guide for the Care and Use of Laboratory Animals published by the US National Institutes of Health (NIH Publication No. 85-23, revised 1996) and were approved by the ethical committee of the University of Reading, UK.
Recombinant myostatin
Bacterially-expressed recombinant myostatin was a kind gift from Professor Ravi Kambadur, University of Waikato, New Zealand [13]. Recombinant myostatin, purified from mouse myeloma cell line, NS0, was purchased from R&D Systems. Variations in the relative amounts of correctly folded bioactive protein in purified preparations of the recombinant protein have been described for members of the TGF-β [19] including myostatin [20]. Hence, concentrations of recombinant myostatin from different sources/lots that resulted in greater than 45% inhibition of proliferation in P2 rat cardiomyocytes after 72 h of treatment were empirically determined (see Table 1) and used in this study.
|Myostatin concentration
|Mean cell no.±SD (n=3)
|% Growth
|Recombinant myostatin(E. coli)
|0 μg/ml
|1 μg/ml
|5 μg/ml
|Recombinant myostatin (mouse myeloma, lot EZ11612A, R&D)
|0 ng/ml
|250 ng/ml
|500 ng/ml
|Myostatin concentration
|Mean cell no.±SD (n=3)
|% Growth
|Recombinant myostatin(E. coli)
|0 μg/ml
|1 μg/ml
|5 μg/ml
|Recombinant myostatin (mouse myeloma, lot EZ11612A, R&D)
|0 ng/ml
|250 ng/ml
|500 ng/ml
Proliferating cardiomyocytes were treated with indicated concentrations of myostatin for 72 h. Viable cells were identified by trypan blue exclusion and counted using a haemocytometer. * indicates significant difference to untreated control cultures.
Isolation of primary cardiomyocyte
Cardiomyocytes from, embryonic age 18 (E18), postnatal ages 0, 2, 5 and 10 (P0, P2, P5 and P10, respectively) rat ventricles were obtained by serial enzymatic digestions. Adult (Ad) cardiomyocytes were prepared by Langendorff perfusion [21]. Isolated cardiomyocytes preparations were >95% pure as revealed by staining with cardiac-specific sarcomeric actin (Abcam, UK) antibody.
Cardiomyocyte culture and growth inhibition studies
Isolated E18 and P0 rat cardiomyocytes were seeded at a density of 2×105/well in 6 well dishes and grown in myocyte medium (DMEM/M199 (4:1) containing 5% foetal bovine serum (FBS) for 24 h. Recombinant myostatin was then added to appropriate cultures. At various time points thereafter, myostatin treated and untreated control cardiomyocytes were harvested by trypsinisation for analysis. Viable cells identified by trypan blue exclusion were counted using a haemocytometer.
Measurement of cell death
Propidium iodide (PI) uptake was used as a marker of dying cells that have lost their cell-membrane integrity to determine whether myostatin induced cell death in proliferating cardiomyocytes. Briefly, E18 cardiomyocytes cultured for 72 h with or without myostatin were harvested. Adherent and non-adherent cells from respective cultures were pooled, washed with PBS and incubated in PI buffer (PBS containing 3% FBS and PI; 20 μg/ml) at 4 °C for 20 min. Sorting of PI positive cells was performed using a FACSCalibur flow cytometer (BD Biosciences). Loss of plasma membrane permeability was quantified as the percentage of cells positively stained by PI.
Fluorescence activated cell sorting (FACS) analysis
Two-colour flow cytometric analysis was performed on ethanol-fixed cardiomyocytes. Briefly, foetal cardiomyocytes seeded at a density of 1×106 cells/10 cm2 dish were grown in 5% FBS myocyte media with or without myostatin for 48 h. Cells then were pulse-labelled with 10 μM bromodeoxyuridine (BrdU) for 30 min at 37 °C and prepared for FACS analysis as described [21]. Labelled nuclei were stained with anti-BrdU antibody and PI and analysed using a FACS Calibur flow cytometer and the CellQuest software (BD Biosciences). Flow cytometric measurements were performed on samples containing 10,000–30,000 cells. Cardiomyocytes were gated into the respective cell cycle phases (G0/G1, S and G2/M) as described [21].
Protein synthesis analysis
The effect of myostatin on protein synthesis in cultured cardiomyocytes was evaluated using [14C]-phenylalanine incorporation as an index. For proliferating cardiomyocytes, E18 myocytes were cultured in 5% FBS myocyte media with or without myostatin for 24 h. [14C]-phenylalanine (0.1 μCi/ml) was then added to each culture and allowed to grow for a further 24 h. For non-dividing cardiomyocytes, P2 myocytes cultured in 5% FBS myocyte media for 24 h were serum starved for 48 h. Phenylephrine (PE) (100 μM) with or without 500 ng/ml myostatin (R&D) and 0.1 μCi/ml [14C]-phenylalanine was then added to respective cultures and grown for a further 24 h. Cells were washed with PBS and ice-cold trichloroacetic acid (10%) and the precipitated proteins solubilized in 0.1 N NaOH and 0.01% SDS at 37 °C. Samples were mixed with scintillation fluid (3 ml) and quantified using a liquid scintillation counter.
Western blotting
To assess the expression of myostatin protein in cardiomyocytes isolated from foetal (E18), neonatal (P2, P5 and P10) and adult rat hearts, protein extracts prepared as described previously [21] were analysed by Western blotting by using the anti-myostatin rabbit polyclonal antibody previously described [13]. For analysis of phospho protein and cell cycle proteins, myostatin treated and control E18 cardiomyocytes from triplicate experiments were washed with ice cold PBS and scraped directly in protein lysis buffer. Pooled protein extracts from each treatment group were analysed by Western blotting using antibodies against phospho-SMAD2 (Chemicon), SMAD2/3 (Upstate biotechnology), p21 and CDK2 (Santa Cruz Biotechnology Ltd). GAPDH (Abcam) was used to correct for protein loading. Relative protein abundance was determined by densitometric analysis, in which signal intensities within the linear range on X-ray film were converted to numerical values using Quantity One® densitometric analysis software (Bio-Rad).
Quantitative RT-PCR
Real-time quantitative RT-PCR (qPCR) was used to evaluate myostatin mRNA expression in E18, P2, P5, P10 and adult rat cardiomyocytes, as well as p21 mRNA expression in E18 cardiomyocytes following myostatin treatment. RNA isolation and cDNA synthesis was performed as previously described [22]. Real-time quantitative PCR analysis was performed using ABsolute™ QPCR ROX mix (ABgene) and a GeneAmp® 5700 sequence detector (Applied Biosystems). Myostatin cDNA was amplified using forward primer, 5′-CGCTACCACGGAAACAATCATT-3′, and reverse primer, 5′-GCTTTCCATCCGCTTGCAT-3′ and the amplified product detected using a myostatin-specific Taqman® probe, 5′-(FAM)-CCATGCCTACCGAGTCTGACTTTC-(TAMRA)-3′. p21 cDNA was amplified using forward primer, 5′-AGAGCCACAGGCACCATGTC-3′, and reverse primer, 5′-CGAACAGACGACGGCATACTT-3′ and the amplified product detected using a p21-specific Taqman® probe, 5′-(FAM)-ATCCTGGTGATGTCCGACCTGTTCCAC-(TAMRA)-3′. Real-time PCR analysis was normalized using the rodent GAPDH control reagent kit (Applied Biosystems).
Statistical analysis
At least three independent experiments were performed for all quantitation analysis. Student's t-test was used for paired data analysis and for multiple data analysis one-way analysis of variance followed by Bonferroni t-test was used. Significance was accepted at p<0.05.
Myostatin is expressed developmentally in cardiomyocytes
We and others have reported previously that there is a greater percentage of proliferating cardiomyocytes in E18 rodent hearts compared to P2 hearts and that this decreases with age [1–3]. We determined whether there was a correlation between these observed age-related changes in cardiomyocyte proliferative index and the temporal expression of myostatin, by studying the expression of myostatin in cardiomyocytes isolated from rat hearts at different developmental ages. Fig. 1 shows that myostatin is differentially expressed in rat cardiomyocytes at various developmental stages. Accordingly, E18 foetal cardiomyocytes display low levels of myostatin expression compared to P2 cardiomyocytes which display significantly higher level of myostatin mRNA (Fig. 1a). By postnatal age 10, when the majority of cardiomyocytes have lost their ability to proliferate [1–3], a 6-fold increase in myostatin mRNA expression is observed compared to that in E18 cardiomyocytes (p<0.001). Western blot analysis (Fig. 1b and c) demonstrates that the levels of active myostatin protein correlate strongly with myostatin mRNA levels, up to the age of P10. Thereafter in adult cardiomyocyte, a 2-fold decrease in the myostatin protein levels and a 15-fold decrease in mRNA levels were observed. Since the majority of cardiomyocytes lose their ability to proliferate during early postnatal development [1–3], the increasing expression of myostatin (particularly during the early postnatal period of cardiomyocyte growth) suggests a potential role for myostatin during this developmental growth phase.
Myostatin inhibits cardiomyocyte proliferation
In skeletal muscle, mature myostatin is a potent inhibitor of skeletal muscle myoblast [23] and satellite cell [24] proliferation. Therefore, we determined whether myostatin would inhibit cardiomyocyte proliferation by adding recombinant myostatin to proliferating cultures. Fig. 2 shows that the addition of recombinant myostatin to proliferating E18 (Fig. 2a) and P0 (Fig. 2b) rat cardiomyocyte cell cultures significantly inhibited their proliferation. The inhibitory effect of myostatin on cardiomyocyte proliferation was evident 24 h after treatment and at all time points studied thereafter. Quantitation of our results indicated that proliferation of E18 and P0 cardiomyocytes was inhibited by 29% and 21%, respectively after 24 h of myostatin treatment. The effect of myostatin treatment was sustained since, at 72 h, proliferation in E18 and P0 cultures was inhibited by 46% and 58%, respectively relative to untreated controls. The inhibition was not accompanied by increased cell death as determined by PI cell exclusion staining (Fig. 2c).
It has been observed that myostatin acts in a novel manner in that it blocks proliferation but that cells maintain their ability to carry on dividing once myostatin is withdrawn [25]. Thus, we investigated the ability of E18 cardiomyocytes to resume proliferation once myostatin was removed. E18 proliferating cardiomyocytes were cultured in the myostatin-containing media for 24 h, this media was then replaced with 5% FBS myocyte media and cardiomyocytes cultured for a further 48 h. Inhibition of myocyte proliferation by myostatin was found to be reversible as demonstrated by the reactivation of cardiomyocyte proliferation following replacement of myostatin-containing media with 5% FBS myocyte media (Fig. 2d).
Myostatin blocks G1 to S phase cell cycle progression in cardiomyocytes
Flow cytometric analysis was performed to determine what phase of the myocyte cell cycle myostatin affected. Fig. 3 shows that myostatin inhibited cardiomyocyte proliferation by blocking S phase entry during myocyte cell cycle progression. Results in Fig. 3b indicates that treatment of proliferating cardiomyocytes with myostatin for 48 h caused a 57–86% decrease in the S phase population (p<0.05), and a concomitant increase of 7–14%, and 15–17% (though not statistically significant) increase the G0/G1 and G2/M populations, respectively.
Myostatin treatment alters the expression of cell cycle proteins necessary for G1 to S phase transition
In skeletal muscle myoblasts, myostatin treatment alters the expression of cell cycle proteins, such as p21 and CDK2 [18,23]. We therefore examined whether myostatin treatment affected the expression of cell cycle proteins critical for the G1 to S-phase transition. Fig. 4a and b shows that in proliferating E18 cardiomyocytes, myostatin treatment decreases the expression of CDK2 protein significantly at 48 h. However, by 72 h, as proliferating control cultures become confluent and the rate of cell division ceases, a decreased expression of CDK2 is also observed in untreated cultures and at comparable levels to that observed in non-confluent myostatin-treated cultures (Fig. 4b). Interestingly myostatin mRNA is also upregulated as proliferating cardiomyocyte cultures become confluent (data not shown). Myostatin treatment also increased the mRNA (Fig. 4c) and protein (Fig. 4d and e) expression of the cell cycle inhibitor p21 in the proliferating E18 cultures significantly. A decreased (but insignificant) expression of cyclin E protein was also observed in some myostatin-treated cardiomyocyte cultures (data not shown).
Effects of myostatin on protein synthesis in cardiomyocytes
Recently, we demonstrated that myostatin inhibits PE-induced protein synthesis rate per cell in serum-starved P2 neonatal cardiomyocytes thereby limiting agonist-induced hypertrophy [14]. Thus, we compared the effect of myostatin on protein synthesis in proliferating foetal cardiomyocytes and serum-starved non-proliferating P2 neonatal cardiomyocytes by [14C]-phenylalanine incorporation. Fig. 5a shows that total protein synthesis in E18 cardiomyocytes exposed to myostatin was significantly lower than that in untreated control cultures. However, as shown in Fig. 5b the protein synthesis rate per cell (normally an indication of hypertrophic growth) was not significantly different between myostatin treated and control cultures. This is in contrast to the effect of myostatin on serum-starved P2 cardiomyocytes subjected to PE-induced hypertrophy (Fig. 5c) wherein a significant inhibition on the rate of protein synthesis per cell and subsequently hypertrophic growth [14] is evident within 24 h of myostatin treatment. This suggests that the decrease in total protein synthesis observed in myostatin-treated E18 cultures (Fig. 5a) reflects proliferation inhibition and hence lower cardiomyocyte numbers relative to untreated controls.
Acute myostatin treatment stimulates SMAD2 phosphorylation in E18 cardiomyocytes
Previous studies have demonstrated that members of the TGF-β growth factor family of ligands including myostatin, can elicit a functional response through the classical SMAD2/3 signalling pathway [15–17]. We determined whether myostatin activates a similar signalling pathway in proliferating cardiomyocytes by Western blot analysis. Fig. 6 demonstrates that myostatin induced SMAD2 phosphorylation in cardiomyocytes within 30 min of treatment and that this phosphorylation persisted (although at a reduced level) for up to 1 h. SMAD2 phosphorylation, however, dropped to basal levels by 3 h and no further increase in SMAD2 phosphorylation was evident up to 24 h of myostatin treatment. In contrast, Western blot analysis using anti-SMAD2/3 specific antibody shows that total SMAD2 protein remains unchanged in both myostatin-treated and control cultures.
This study and previous reports [13,14,26] demonstrates that myostatin mRNA and protein are expressed in cardiomyocytes. However, its physiological role in cardiac muscle growth remains unclear since myostatin null mice do not display the profound alterations in cardiac muscle mass evident in skeletal muscle [9]. Our study shows for the first time that myostatin expression is regulated developmentally in rat cardiomyocytes. E18 foetal cardiomyocytes were shown to express low levels of myostatin that gradually increased 6-fold in P10 rat cardiomyocytes. Given that the majority of rat cardiomyocytes cease to proliferate shortly after birth [1–3], it was interesting to note that the increased expression of myostatin coincided closely with the diminishing proliferative index of neonatal rat cardiomyocytes with age. Hence, we studied the effects of myostatin on cardiac proliferation and observed that myostatin is a potent inhibitor of serum-induced embryonic and neonatal cardiomyocyte proliferation. Taken together, these observations support a developmental role for myostatin in limiting cardiomyocyte proliferation.
Using FACS analysis, we have shown that myostatin inhibits cardiac proliferation in vitro through a block in the G1 to S phase cell cycle transition. Progression through the G1-S phase transition is controlled by the activity of the G1-acting cyclin:cyclin-dependent kinase (CDK) complex, cyclin E:CDK2 [27]. CDK2-associated activity plays a key role in regulating entry into and exit from S-phase of the cell cycle. For instance, it has been demonstrated that microinjection of affinity-purified anti-cdk2 antibodies during G1, inhibited entry into S phase [28]. Therefore, limiting CDK2 availability ultimately can lead to a G1 to S cell cycle arrest. Accordingly, we observed that a decrease in CDK2 expression is part of the mechanism through which myostatin acts to inhibit cardiomyocyte proliferation.
The CDK inhibitor p21 has been shown to inhibit cyclin:CDK complexes, including those that promote cell cycle progression through the G1 and S phase [29]. Furthermore, overexpression of p21 has been shown to arrest cells in G1 [30]. Since p21 mRNA and protein is upregulated in myostatin-treated E18 cardiomyocytes, our results suggests that upregulation of p21 is also part of the mechanism through which myostatin elicits a G1/S cell cycle block in cardiomyocytes. Myostatin-mediated inhibition of skeletal muscle myoblast proliferation has also been shown to involve increased expression of p21, decreased CDK2 activity and protein expression that ultimately results in a G1/S cell cycle arrest [24]. These observations highlight similarities in cell cycle regulation by myostatin in both skeletal muscle and cardiac myocytes.
In this study we observe that in proliferating cardiomyocytes, myostatin induced SMAD2 phosphorylation within 30 min of stimulation and subsequently inhibited cardiomyocyte cell division. It is noteworthy that TGF-β, which also signals via the classical SMAD2/3 pathway, has been reported to inhibit proliferation of immature cardiomyocytes [31]. The anti-proliferative effect of myostatin and other TGF-β family members, in conjunction with SMAD signalling, is well recognised in other cell types [15,16,32]. SMAD phosphorylation following binding of these ligands to their respective serine threonine kinase transmembrane receptor leads to the formation of heteromeric complexes with SMAD4. The complex then translocates into the nucleus to bind promoter regions of target genes or interact with various transcription factors to regulate gene expression [33–35]. SMAD2 proteins are able to interact with a diverse range of proteins that do not share a common motif [35,36]. Hence the phenotypic outcome following SMAD activation is therefore likely to depend on the SMAD-binding transcription factors present within the nucleus at the time of their activation. We propose that SMAD2 activation in myostatin treated E18 cardiomyocyte cultures most likely affects proliferation, as very little hypertrophic growth is evident at this age [1–3].
Our study also shows that in adult cardiomyocytes, where adaptive growth occurs mainly by hypertrophy [1], myostatin expression is maintained but at a much lower level. Since the majority of adult cardiomyocytes have lost their ability to divide and express very low levels of key molecules involved in cell division, we propose that high levels of myostatin, perhaps, are not necessary at this age to actively inhibit cell proliferation. We speculate that the expression of myostatin in adult cardiomyocytes might reflect a potential role in modulating hypertrophic growth rather than hyperplastic growth. Indeed, as observed for myostatin [14], adenovirus-mediated expression of growth differentiation factor 15 (GDF15), another member of the TGF-β family, has been shown to antagonise agonist-induced hypertrophy in P2 cardiomyocytes in vitro. Interestingly, this was shown to involve SMAD2/3 activation [37]. Furthermore, overexpression of SMAD2 in serum-starved cultured neonatal cardiomyocytes inhibits agonist-induced cardiomyocyte hypertrophy in vitro[37]. Thus the ability of myostatin to affect both hyperplastic and hypertrophic growth, depending on the physiological status of the cardiomyocyte, might be partly related to the transcriptional pathways influenced following SMAD activation by myostatin.
Myostatin knockout mice studies indicate that the cardiac muscle in these mice is not as grossly enlarged as their skeletal muscle [9,14]. This suggests that other regulators can compensate for the absence of myostatin in cardiac muscle. Nevertheless, relatively smaller cardiomyocytes are observed in myostatin knockout mice compared to their wildtype counterparts [14]. The observation that cardiac-specific overexpression of CDK2 increases cardiomyocyte proliferation and results in significantly smaller cardiomyocytes [38], suggests that the relatively smaller cardiomyocytes observed in myostatin knockout mice might be linked to abnormal proliferation. Thus, it will be interesting to determine whether the proliferative capacity of cardiomyocytes obtained from myostatin knockout mice is enhanced. It also is worth noting that satellite cells isolated from myostatin knockout mice have an enhanced proliferation rate [25] and that their cardiomyocytes display exuberant cell enlargement in response to the hypertrophic agonist, phenylepherine [14].
Our observations emphasise the fact that molecules that regulate cell cycle gene expression play a critical role in the proliferation of cardiomyocytes and that their identification and manipulation might help to extend the proliferative capacity of cardiomyocytes. A role for myostatin in cardiomyocyte growth is further supported by several other observations. Firstly, IGF-1 mediates stretch-induced upregulation of myostatin in neonatal rat cardiomyocytes [26]; secondly, myostatin is induced in cells immediately surrounding a myocardial infarct for up to 30 days post infarct [13]; and finally, myostatin is upregulated dramatically in the hearts of AKT knockout mice, a genetic model of cardiac hypertrophy [14]. These observations, together with those from our study, demonstrate that myostatin plays an important role in modulating cardiac growth. The emerging picture is that myostatin affects both hyperplastic and hypertrophic growth in cardiomyocytes, depending on their physiological and growth status.
We thank Dr. Ian Kavanagh for his help with the protein synthesis studies and Dr. Fleur Moseley, Carmen H. Coxon and Wendy Lockwood for their help with the cardiomyocyte cultures. This project was funded by grants from the British Heart Foundation project grant no PG/04/053/17031 (awarded to G.B. and K.P.) and a BHF Intermediate Fellowship FS/03/024/15491(awarded to K.A.B.).
Jin HJ, Dunn MA, Borthakur D, Kim Y.S. Refolding and purification of unprocessed porcine myostatin expressed in Escherichia coli. Protein Expr Purif.;35:1–10.


text: Revamped Sling.com Launches to Spendthrift PublicMark Wilson11/26/08 11:10amFiled to: WebsitesSling.comSlingBetaHuluTelevisionprogramsStreamingVideoSoftware8EditPromoteShare to KinjaToggle Conversation toolsGo to permalink Sling.com has been in closed beta for some time, but now the Huluesque service is open to the public. The site includes 60 other content partners including networks like FOX and CBS and studios like MGM and Sony. And while you wouldn't expect to see any NBC content on there, you can actually watch shows like 30 Rock that are badged with a Hulu logo. My first reaction is that Sling's quality leaves something to be desired, but I certainly can't complain about the price. [Sling via AlleyInsider]Reply8 repliesLeave a reply

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Most enthusiasts will agree that Packards glory days began in earnest in the late 1920s and ran through the mid-1930s. During this time the... More
1938 Packard Twelve Convertible Victoria
Orange Village, OH
• Sold new by Packard of Manhattan May 22, 1938 • Classic Car Club of America Senior winner • One of only 157 Packard V-12s in... More
1935 Packard 1201 Eight Victoria Model 1201 Convertible Coupe
Bedford Heights, OH
"Ask the Man Who Owns One." For most of Packard's 58-year life, the slogan was an integral part of the company's existence. No mere... More
1941 Packard 160 All Original 28,000 Miles
Bedford Heights, OH
Known as the Nineteenth Series, the 1941 Packard model line continued the styling innovations that were pioneered during the prior year, along with a... More
1937 Packard 115 4-Door Sedan
La Vergne, TN
For sale in our Nashville showroom is simply stunning 1937 Packard 115 sedan. This show quality built 37 Packard for sale exudes luxury and... More
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1950 Packard Eight Deluxe Club Sedan
Canton, OH
1950 Packard Deluxe Eight 23rd series club sedan, 2-dr. fastback. 288 ci./130 hp. Eight cylinder, 3-speed manual transmission with over drive. Finished in beautiful... More
Advertiser since 2004
1930 Packard 733 Packard Eight 733 Coupe Cabriolet
Austin, TX
You are looking at a gorgeous 1930 Packard Eight 733 Coupe Cabriolet. It is a privately owned vehicle being sold on consignment at Stressless... More
1938 Packard Eight
Lenexa, KS
If Style, Elegance, and Grace is what you desire, LOOK NO FURTHER!!! This 1938 Packard Eight 1601 Convertible Sedan in Deep Loyola Maroon and... More
1937 Packard 120 C
Lithia Springs, GA
To survive the Great Depression, independent automaker Packard had to diversify, and that meant selling more cars to more people. Hence, the 120. With... More
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1949 Packard 2301
Celina, OH
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Athol, ID
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1937 Packard Twelve Coupe Roadster
St. Louis, MO
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1955 Packard Caribbean Convertible
Fredericksburg, VA
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Cleveland, OH
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West Chester, PA
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Astoria, NY
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St. Louis, MO
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Norwalk, OH
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Bedford Heights, OH
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Emmerich, Germany
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Pleasanton, CA
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West Chester, PA
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Elsewhere, teenagers are sailing the world single-handed : Dutch girl, Laura Dekker – all of 16 years-of-age, just arrived at home port after a year-long solo crossing of one year. An Aussie teen – also 16 – Jessica Watson did likewise last year.
Then, of course, is the build-up for the America's Cup in 2014 and, anyone down Pattaya way in February, can have an inkling on what these speed machines are capable of achieving.
But, back to Thailand and the forthcoming extravaganza. Yacht-racing in the kingdom is forging ahead, with a junior world champion and many other Thais in top contention. And, now, with events like the Catamaran fest and the many regattas held up and down the country during the year, Thailand's water-ways are the most desirable venues in the world. This is the collective opinion of many sailors who come back year after year.
Events during the week will include the 'Island Record Race' which will always be an identical course each year allowing a record to be set and also speed trials, as some of the fastest beach cats in existence will be taking part in the event.
In successive years, the 'Island Record Race' will be open to windsurfers, kite surfers and also larger multi-hulls creating a true on the water challenge between all sail-driven craft.
The event is organized by the HM the King's own Royal Varuna Yacht Club, situated on a beautiful property south of Pattaya and, as the kingdom's oldest club, has years of experience in managing big international events.
Thai Cat Regatta website


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A Different Type of Tab
I'm sure that we're all very accustomed to the usual left, center, right and decimal tab stops Word has used for years. But, if you're using a newer version of Word, you may have noticed a new addition to the family.
When flipping through your tab choices, did you ever notice this little guy?
Well, I did and had to wonder what it was… Continue reading


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Armored Saint
This Friday, Armored Saint will release their new live album, 'Carpe Noctum'. For a preview, a live video of 'Last Train Home' from Wacken Open Air can be viewed at metalblade.com/armoredsaintRead More
Native Construct
Boston, MA based progressive act Native Construct has just launched the instrumental version of their critically-acclaimed debut release, 'Quiet World'.Read More
Princess Pang
In celebration of Metal Blade Records' 35th anniversary, a recently digitized video from the archives will be launched every Thursday. This week's installment is Princess Pang's 'Trouble in Paradise'.Read More
Necromancing the Stone
To celebrate Valentine's Day (Feb. 14th), Necromancing the Stone has launched a new video for the track 'Bleed for the Night', taken from their latest album, 'Jewel of the Vile'.Read More
New Releases
New Releases
Latest News
Amon Amarth premieres "Raise Your Horns" video via RollingStone.com
- September 19th, 2016 – Launches contest with Grimfrost; set to kick...Read More
Metal Blade Records launches new edition of "6 Bands in 60 Seconds"
- September 15th, 2016 – In conjunction with RED Music, Metal Blade...Read More
Serpentine Dominion's Adam Dutkiewicz launches guitar play-through for new track, "On the Brink of Devastation"
- September 14th, 2016 – Imagine Killswitch Engage at their heaviest,...Read More
Monte Pittman streams new track, "Be Very Afraid", online
- September 13th, 2016 – There are great guitarists, great vocalists...Read More
Good Tiger premieres new video for "Aspirations" via AltPress.com – alongside exclusive interview with Derya "Dez" Nagle
- September 13th, 2016 – Kicks off North American tour with Dance Gavin...Read More
Armored Saint launches PledgeMusic campaign for new live album
- September 9th, 2016 – Photo credit: Stephanie Cabral Having toured...Read More
Allegaeon launches mini studio documentary for new album, 'Proponent for Sentience', online
- September 9th, 2016 – On September 23rd, melodic death metal outfit...Read More
Poison Headache launches live video for "Rot With Me"
- September 9th, 2016 – Set to join Killswitch Engage, Noisem for west...Read More


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Aristokrat Records new boy, Pucado, releases a new joint "Ukwu Nka" as a follow-up to "Bigger, Better, Best" (watch the video) which got – and is still getting – loads of airplay in Lagos. It's an experimental freestyle that will grow on you with each listen.
Crazy instrumentals. LeriQ is climbing up my personal producer charts.
Produced by LeriQ [DOWNLOAD][audio:http://notjustok.com/wp-content/uploads/2013/06/Pucado-Ukwu-Nka.mp3]


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Cool hit sound mixed for Nothing To Hide based on sounds under CC0:
Cymbal Impact.wav by LloydEvans09
Dramatic Bass Hit by snakebarney
Thanks. Great sound effect.
That's just great!!!
brilliant. thank you :)
Great sound! Combined it with another sound in an infomercial-type comedy video as the sound to indicate something "bad" (i.e. "It removes rust! [sound] And grime! [sound]). Thanks for this!
Thank you:D
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Suggest Edits
- Tags
Link to Lauer's In-Town Restaurant
Copy the link below and paste onto your website or blog to display badge.
Recent Visitors
9/23/2013 A diner from Overland Park, KS tried it, liked it, and rated it . They liked the food, liked the service, and liked the ambiance.


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Hasiru Dala is a membership based non-profit organization of waste pickers and other informal waste workers. Hasiru Dala strives to integrate marginalized informal waste workers including waste pickers in the solid waste management framework by utilizing their expertise in the domain. Initiated in 2013, Hasiru Dala now has a membership of over 7500 waste pickers and is innovating services and business models that leverage waste pickers' expertise and entrepreneurship, generating stable livelihoods in the process.
Through its work, Hasiru Dala is creating recognition of waste pickers and informal waste workers as important contributors to urban economy. Their work which includes collection, sorting, grading and transportation of waste for recycling is foundational to both green and circular economies, much needed for mitigation of climate change.
Hasiru Dala has trained waste pickers to upgrade their skills for providing more services to the citizens. Current services include Total Waste Management for bulk generators, Dry Waste Collection and Aggregation, Operation of Bio-gas Plants, and Composting Services.
Geographically Hasiru Dala is based in Bengaluru, and is slowly- slowly expanding to other cities like Nelamangala, Mysuru.
In addition to livelihood creation, Hasiru Dala enables workers' access to social security benefits for improving the quality of life. The admittance to social security benefits include procuring government issued occupational ID cards that give workers a sense of identity as well as access to financial inclusion, health insurance schemes, educational scholarship and loans for their children's education and to run their own enterprises.
Who are waste pickers and informal waste workers? What do they do?
In a neighborhood dumping spot, you must be seeing men and women looking through the garbage. Armed only with old shoes and clothes picked out from waste heaps and a stick to sort through trash, they look for material which has a value and can be sent for recycling. They are waste pickers. They collect discarded materials that have zero value and convert it into a tradable commodity through their labor in extraction, collection, sorting, grading and transporting (Alliance of Indian Waste picker). There are itinerant buyers, whom we refer to as Kabadiwallas. Riding on the bicycle they call the residents to sell the used materials i.e. old newspapers, pet and glass bottles, metal which has no use. There are others, who purchase the collected material by waste pickers and itinerant buyers, they are scrap dealers. In both scrap shops and the hierarchy above there are workers, who sort and grade waste in various categories, they are called sorters. The term 'other informal waste workers' is all encompassing and includes sorters, itinerant buyers, scrap dealers and others. Hasiru Dala is a membership based organization of waste pickers and informal waste workers.
What is the vision and mission of Hasiru Dala?
Vision: An inclusive society for waste pickers and other informal waste workers, free of discrimination based on their caste, religion, occupation, gender and sexuality.
Mission: Creating sustainable opportunities for waste pickers and other informal waste workers in the urban economy and for conservation of our natural habitats.


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Body Lift including implant removal
63 of 1617
Was this photo helpful? {{ voteCount + ' other' + (voteCountOthers == 1 ? '' : 's') }} found this helpful
When this patient was 35 she had a breast lift with implants and a tummy tuck. Her body morphed since her thirties, leaving her with more weight around her abdomen and more breast than she was comfortable with. She no longer wanted to have breast implants. She chose to have implant removal with breast lift and a revision of her tummy tuck.
By uploading this photo, Mary Lee Peters, MD has agreed with RealSelf.com policy that they've secured proper patient consent.


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Ceilidh 27th February
To buy tickets please go to https://www.picatic.com/event14537332686445008
The Blue Factory is at the end of Millers Avenue, on the right hand side. Just follow the signs.
Read our latest annual report
This year's annual report was written by our volunteer Helena Smith and designed by Matthew Smith.
Autumn Gardening Day – 19th September 2015
Music night at Ruby House 21st May 2015
Pictures of the night, by Helena Smith
Spring Gardening Day 21st March 2015
Under the Kauri Tree – fundraiser 12th Dec 2014
Three Course Story
Driving volunteers needed
We need volunteers with cars to pick up fruit and veg from Growing Communities on Leswin Rd (Stoke Newington) at 9 am on Saturdays and drop them off at the church hall (about 30 minutes).
We also need volunteers to pick up from Wholefoods at 9 pm on Sundays drop off at church hall. (about 1 hour).
If you can help, please email email@example.com with subject title Driver.
Patchwork Preserves: fundraising for NLAH
Patchwork Preserves is a collaboration between Growing Communities and North London Action for the Homeless (NLAH). Growing Communities donate their surplus veg to the NLAH kitchen. What isn't used is then turned into lovely organic chutneys and pickles by Patchwork Farmers, staff and service users of NLAH. Recently we've even used beetroots from our own garden! So far we have made Spicy Beetroot and Orange Chutney, Spicy Carrot, Lemon preserve, Crunchy Courgette Pickle, Spicy Tomato Relish, Red cabbage and Ginger Chutney.
For Christmas we sold chutney at Stoke Newington Farmer's Market as well as at Growing Communities Urban Farm Shop. Since the project started we have sold well over 200 jars and all profits go to North London Action for the Homeless.


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This is to Striver7777 who helped me alot when i was a noob to booksie. Telling me what he liked what was funny and so on. Thanks Striver7777!!!!
This is to My lovely (only 6) fans who think im good at what i do. Thanks for becoming my fan and liking me. Thanks you guys!
This is To Maddie Grey. I've only read 2 of your books (War Of Attrition and Baby Blue Eyes) I think their amazing. They have been detailed and well done to your 600 fans! Congrats for making it to the featured writing on the home page!
This is to Megan Mckenzie who again wrote amazing pieces. And congrats Megan for your amazing novel Smack!
And this is to the people who commented, liked or even just read my work. Without you i couldn't have improved my writing and ideas! I appreciate when you comment and give your feedback. I get so excited when i log in and see the orande box that says 'New messages!' or 'New fan rewquests' Thanks for making me who i am.
Love you all and appreciate your help and i enjoy reading your poems, short stories and Novels especially!!
TheMysteriousGirl x
© Copyright 2017 TheMysteriousgirl. All rights reserved.
Status: Finished
Genre: Non-Fiction
Book / Other
Book / Action and Adventure
Book / Action and Adventure
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- 04-01-2014, 09:25 AM Thread Author #1
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Will the Samsung Galaxy Note 3 work in the UK ?!
I am a UK resident and I wish to purchase a Samsung Galaxy Note 3. It's quite expensive in the UK although I was wondering if I bought it from the US, due to the currency conversion rate, it'll come out cheaper.
My main concern is that will the phone still work here in the UK I.e will I still be able to get calls ?! I was asking because of the different carriers and antennas. Although I'm not an LTE person I'll happily use 3G. I'm currently on the 3 carrier in the UK. Thank You
Sent from my HTC One using AC Forums mobile app
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Re: Will the Samsung Galaxy Note 3 work in the UK ?!
I am not sure about US Cellular. Tmobile or AT&T should work, as they are both GSM. Of the two, TMobile has the better chance of working.
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The R&B superstar R Kelly has been cleared of making child pornography after a month-long trial which boiled down to whether a man filmed having sex with an underage girl had a mole on his back.
Mr Kelly was charged six years ago after a graphic videotape allegedly showing him having sex with a 13-year-old girl was handed to Chicago police.
After six hours of deliberations, the jury in Chicago yesterday pronounced the singer not guilty on all 14 charges.
As the verdict was read out, Mr Kelly, 41, dabbed his face with a handkerchief and hugged each of his four lawyers. He could have faced a 15-year jail sentence had he been convicted.
Both Mr Kelly and the girl, now aged 23, had denied they were the people in the 27-minute film. Mr Kelly's defence team argued that it could not have been him on the tape, as he had a large mole on his back and the man in the video did not.
The issue of whether the alleged victim was the girl in the film was also subject to contradictions from witnesses, including relatives and childhood friends.
Mr Kelly, whose real first name is Robert, rose from poverty in Chicago to become a singer, songwriter and producer, winning a Grammy in 1997 for "I Believe I Can Fly". He is also known for raunchy hits such as "Bump N' Grind".
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- Children's Rights
- R&b Singers
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CINCINNATI -- The only thing that stood between Noah Lowry and finishing the best overall performance of his brief career was a blister.
Lowry pitched a season-high 8 1/3 innings, drove in the tying run and scored the go-ahead run as the San Francisco Giants beat the Cincinnati Reds 3-2 on Wednesday night.
The left-hander was two outs from the third complete game of his career and first of the season when a blister on his left index finger forced him out of the game after striking out Ken Griffey, Jr., to start the ninth.
Scott Eyre relieved and, after completing a walk to Austin Kearns that was charged to Lowry, struck out Adam Dunn. LaTroy Hawkins came on and got Sean Casey to fly out to center fielder Randy Winn on the warning track for his sixth save, second with the Giants.
"Lowry was tremendous," Giants manager Felipe Alou said. "He pitched four innings with that blister developing, but he kept going back and going back. We asked him if he wanted to come out, and he said, 'No, I'll just keep icing it and going back.'"
"It's nothing serious," Lowry said. "It'll just take a couple of days to heal, and I'll be ready in five days."
The Giants improved to 5-4 on their season-long 13-game road trip with their fourth win in six games. A victory on Thursday would give them their first sweep of a four-game series in Cincinnati since May 1983.
The Reds, who have won 10 of their last 11 road games, dropped their fourth consecutive home game and have lost seven of their last nine in Cincinnati.
Lowry (9-11) went at least seven innings for the third time in his last three starts, all of which he won after going 1-4 in his previous five. He limited the National League's highest-scoring team to seven hits and two runs, striking out seven. He walked one and matched Jason Schmidt for the team lead in wins while impressing Reds manager Jerry Narron.
"He's a very good pitcher," Narron said. "He's not a thrower. He did a good job changing speeds and hitting both sides of the plate. He kept us off-balance."
Lowry, who was hitting .282 going into the game, also played a key role in San Francisco's three-run fifth that gave the Giants a 3-2 lead.
The Giants broke up Luke Hudson's shutout on Ray Durham's leadoff walk, Todd Linden's double to right-center and Jason Ellison's sacrifice fly.
One out later, Lowry lined a single to left on a 0-2 pitch to drive in Linden from second with the game-tying run. The RBI was Lowry's sixth of the season.
"I was just trying to put the ball in play and make good contact," Lowry said. "He left the ball up a little bit, and I got just enough of it."
"That was the biggest at bat of the game," Narron said. "I know Lowry's a pretty good hitter, but you've got to bear down there."
Lowry moved to third on Winn's double, and after Omar Vizquel was hit by a pitch to load the bases, J.T. Snow walked to drive in Lowry and knock out Hudson. Todd Coffey came in to get Pedro Feliz to ground into an inning-ending forceout.
Hudson (4-6), who won his last three starts, went 4 2/3 innings, giving up four hits and three runs.
The Reds took a 1-0 lead in the first when Rich Aurilia singled with one out, moved to third on Griffey's sharp single to the base of the right-field wall and Kearns' sacrifice fly.
Hudson led off the third with the first of three consecutive singles and scored on Griffey's bases loaded sacrifice fly to give the Reds a 2-0 lead. Kearns lined into an inning-ending double play, and Lowry settled down, allowing two more hits.
"They put some good swings on some good pitches and found some holes," Lowry said. "I decided to stick with my game plan and keep the ball down. I know this place plays short. All four of my pitches were working. I was able to establish my fastball on both sides of the plate, get ahead and stay ahead."
Lowry had thrown 14 1/3 consecutive scoreless innings going into the game. ... Feliz had hits in six consecutive at-bats over two games before flying out to end the fourth. ... Reds SS Felipe Lopez has six hits over the last two games after going 2-for-38. ... With a day game Thursday, Giants IF Deivi Cruz got the night off Wednesday despite setting a career high with five hits on Tuesday. ... Eyre's appearance was his major league-leading 64th.


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Toro Y Moi
The musical revelation of the season is a guy from South Carolina
Chaz Bundick is a young man from South Carolina who everyone is talking about. His musical project goes by the name of Toro Y Moi and last year he released an album entitled Causers of This which seemed to fit in with the pop sensibilities of the time: dreamlike and evocative. These qualities are still present in Underneath The Pine, the sophomore album that comes out on the 22nd of February. There is one major change: whilst Causers of This was electronic, Underneath the Pine is composed with live instruments and sounds like early 80s US radio-friendly pop, of the kind that was born dated considering the electronic pop coming out of Europe at the time.
Bundick has taken music that sounds creepily adult and shifted its shape for the information age, something that Ariel Pink has been praised/accused of doing with rock. So much so that Simon Reynolds devoted an entire page of The Wire to Bundick's new album. The polished sound of Toro Y Moi apparently goes against the lo-fi aesthetics of recent pop music, underground artists now strive for "pop utopia": sound perfection, friendly yet sophisticated melodies.
For the moment this trend is working out, but once Pitchfork will be full of Steely Dan sound-alikes we are sure to be in for another wave of nostalgie de la boue. Fortunately Bundick has talent to match his fashionable production choices and Underneath The Pine is definitely worth a listen.
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Located in San Diego (Downtown San Diego), The Sofia Hotel is minutes from Spreckels Theater and Westfield Horton Plaza.
This romantic hotel is close to San Diego Convention Center and Balboa Park.
Make yourself at home in one of the 211 air-conditioned rooms featuring refrigerators and microwaves. Your room comes with a pillowtop bed. Wired and wireless Internet access is complimentary, and flat-screen televisions with satellite programming provide entertainment. Private bathrooms with showers feature designer toiletries and hair dryers.
Take advantage of recreation opportunities such as a health club, or other amenities including wireless Internet access (surcharge) and concierge services.
Grab a bite to eat at the hotel's restaurant, which features a bar, or stay in and take advantage of room service (during limited hours). Quench your thirst with your favorite drink at a bar/lounge.
Featured amenities include complimentary high-speed (wired) Internet access, a business center, and express check-in. Event facilities at this hotel consist of conference space and meeting rooms. Self parking (subject to charges) is available onsite.
About the The Sofia Hotel
Property Features
There's a restaurant on site. You can enjoy a drink at the bar/lounge. A business center and meeting rooms are available.
Wired Internet is free in public spaces. A health club, tour/ticket assistance, and multilingual staff are also featured at the business-friendly The Sofia Hotel. Parking is available for a fee.
This 3.5-star San Diego hotel is smoke free.
- 1 building
- 211 guestrooms or units
- 7 levels
- 3 conference rooms
- Built in 1926
- Dining venue
- Bar or lounge
- Business facilities
- Conference facilities
- Breakfast available (surcharge)
- Dry cleaning
- Self-service laundry
- Front desk (24 hours)
- Express check-in
- Staff is multilingual
- Storage area for luggage
- Tour and ticket information
- Concierge
- Fireplace in lobby
- Television in lobby
- Elevator
- No smoking on site
Available in all rooms: Free WiFi , Free wired high-speed Internet
Available in some public areas: Wireless internet access ( USD 8.75 per night ), Free wired high-speed Internet
Self parking (surcharge), Valet parking (surcharge)
Room Amenities
The Sofia Hotel offers 211 air-conditioned accommodations with laptop-compatible safes and coffee/tea makers. Pillowtop beds feature premium bedding. Flat-screen televisions come with premium satellite channels. Refrigerators and microwaves are provided. Bathrooms include showers, designer toiletries, complimentary toiletries, and hair dryers.
This San Diego hotel provides complimentary wired and wireless high-speed Internet access. Business-friendly amenities include desks and complimentary weekday newspapers; free local calls are provided (restrictions may apply). Additionally, rooms include irons/ironing boards and blackout drapes/curtains. Housekeeping is offered daily and hypo-allergenic bedding can be requested.
- Pillowtop beds
- In-room dining (limited hours)
- Flat-screen television
- Satellite channels
- Premium channels
- Refrigerator
- Microwave
- Coffee maker
- Luxury bedding
- Hypo-allergenic bedding
- Shower
- Cribs (free)
- Housekeeping (daily)
- Weekday newspapers (free)
- Local calls (free)
- Hair dryer
- Laptop safe
- Designer toiletries
- Iron and ironing board
- Desk
- Blackout drapes
- Climate control (air conditioning)
- Ensuite bathroom
Food & Drink
For a fee, guests can enjoy cooked-to-order breakfast on weekends.
Currant American Brasseri - This restaurant serves brunch, lunch, and dinner. Guests can enjoy drinks at the bar.
Room service (during limited hours) is available.
On weekends, start your day with a cooked-to-order breakfast for a surcharge. You can order up room service during limited hours.
Nearby Things to Do
The hotel provides a 24-hour yoga studio with on-call instructor, and a 24-hour fitness studio with state-of-the-art equipment.
If you need to make special requests for accessibility needs, please note them at check-out when booking your stay.
- Accessible bathroom
- Roll-in shower
- In-room accessibility
Hotel Policies
Check-in time starts at 4 PM
Check-in time ends at midnight
Check-out time is 11 AM
Payment types
Children and extra beds
- Children are welcome.
- Free cribs (infant beds)!
- Pets: USD 25 per pet, per night
- Certain rooms only
- You can request a pet-friendly room by reaching out to the property at the number on the booking confirmation
Charges for extra guests may apply and vary according to property policy.
A credit card for incidental charges and government-issued photo identification are required upon check-in.
Special requests are subject to availability at the time of check-in. Special requests can't be guaranteed and may incur additional charges.
- 2.89 percent destination fee will be charged
- Resort fee: USD 9.85 per accommodation, per night
- Fitness center access
- Fitness/yoga classes
- Sporting facilities or equipment
- Business center/computer access
- Internet access
- Newspaper
- Phone calls
- In-room safe
- In-room coffee
- In-room bottled water
- Additional inclusions
Optional extras
- Breakfast fee: between USD 6 and USD 12 per person (approximately)
- Fee for wireless Internet in public areas: USD 8.75 per night (rates may vary)
- Self parking fee: USD 36.00 per night
- Valet parking fee: USD 36.00 per night (in/out privileges)
- Pet fee: USD 25 per pet, per night
Hotel Name
- Hotel Sofia
- Sofia Hotel
- Sofia Hotel San Diego
- Sofia San Diego
You should know
Pets are welcome in specific rooms only, which can be requested by contacting the property directly at the number on your booking confirmation (please see the Fees section for related charges).
Awards & Affiliations
- The property is a member of Historic Hotels of America.


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Moving content, widgets and apps
Newb Android parent, like 2nd day.
Android 4.04 on Samsung Galaxy Exhilarate.
I was moving the Samsung Kies app off the home screen to the one before it and it just disappeared. I held down the icon til the screen changed to the customize mode, and then tried to drag it to the previous screen. And poof! Gone.
There doesn't seem to be a trash can to locate it and I'm stumped. I still have the Kies Air app, icon, widget or w/e you call it on another page, but I'm wondering where it went. This is not the same app I don't believe; it's the app for the Kies update software that links to PC I think. And where can I find it online or on my phone to restore it? I searched google for it but no luck.
Any help is greatly appreciated. This thing really is like caring for a new baby. lol Can't wait til it grows up or I get as tricky and smart as it already is.
EDIT: I'm reading elsewhere that the app should still be somewhere on the apps list and that it was not uninstalled but the shortcut was just deleted, but I still can't locate it. Is there a Remove or Trashcan folder where it might be found?
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Is this restaurant appropriate for Kids?
Is this restaurant good for lunch?
Is this restaurant good for special occasions?
Is this restaurant romantic?
Does this restaurant offer takeout or food to go?
Does this restaurant accept reservations?
Is this restaurant good for dinner?
Is this a Japanese restaurant?
Are the prices at this restaurant mid-range / moderate?
Is this restaurant good for dining on a budget?
Thanks for helping!
Nice restaurant when you are travelling by. Its not the tastiest sushi but the staff is great, its good price and the food is quite good. And on tuesday night for 25$ its all you can eat:)
Best in town. Delicious food. Fast service and very friendly staff. Well located In the center of town.
If you want good Sushi this is a great place to go. Compared to Sushi restaurants around the U.S. it is only good range. Not a sushi restaurant you will walk away saying.....best sushi ever.
I do recommend going if you are in the mood. Not the cheapest but you really don't want to go to any "cheap" sushi restaurants...
Best sushi restaurant in Rimouski, a bit expensive but totally worth it! I've been there a couple of times and never been disappointed. Fresh meals, fast service and nice ambience. You can also order for takeout
If you crave for sushi and have the luck to be in Rimouski , your only stop for that will be at L'unique sushi bar ... The food is amazing and the so is the service , we didn't have any trouble having some answers from the hostess present at the time and the staff seems to have fun when...
Excellent sushi, a bit pricy but definitely not the most expensive in Rimouski. It is a treat for me. The food has been great everytime I went.
The service was good... but the food... oh god! Never ate sushis like that. Don't go there, it didn't worth the price.
The sushis was excellent and very fresh. Them hosomaki was enormous! And the sake perfect! The "Jardinier" was the best vegetarian sushi I taste since a long time!
best sushi ever... try the "rouge piquant", the "sept-iles" and the "geisha"!
I get it for take out whenever I can
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Important information about purchasing from Amazon in the US.
RØDE Microphones does not authorize Fulfilled By Amazon. We have purchased counterfeit RØDE products using Fulfilled by Amazon and highly recommend that you only purchase RØDE products from authorised dealers.
If you purchase any RØDE microphone from an unauthorized dealer via Fulfilled By Amazon you will not receive any US warranty or technical support.
You can view a full list of authorized US dealers here.
30 Online retailers selling the RØDELink Filmmaker Kit
RØDELink represents the next generation of digital wireless systems. Using a Series II 2.4GHz digital transmission with 128-bit encryption, it is able to constantly monitor and hop between frequencies to maintain the strongest possible signal level at a range of up to 100 metres* (over 100 yards). The RØDELink Filmmaker Kit provides everything you need to start shooting video wirelessly, including a receiver, transmitter and broadcast-grade lavalier microphone. The receiver (RX-CAM) features an OLED display with information on level, battery status (receiver and transmitter), mute and channel selection. The unit can be mounted on a standard camera shoe mount, 3/8" thread or belt-clip, and for added versatility the shoe mount can be located in one of two positions. The lightweight transmitter (TX-BELT) can be fitted to a belt or clothing and features a locking thread to ensure the microphone connection is as secure as possible. RØDE has included an omnidirectional lavalier microphone in the kit to provide the highest possible audio reproduction quality.
* range estimations based on optimal environment
|Transmission type
|2.4GHz Fixed Frequency Agile System
|System dynamic range
|Range (distance)
|Up To 100m
|Frequency range
|35Hz - 22kHz
|Maximum output level
|1v Rms (2.8v P-p)
|Max input signal level
|1v Rms (2.8v P-p)
|Max latency
|Power source
|2 X AA Battery Or MicroUSB
|Output connection
|3.5mm TRS Locking Jack Socket - Dual Mono
|Output impedance
|111mm X 65mm X 52mm
|Power source
|2 X AA Battery Or MicroUSB
|Input connection
|3.5mm TRS Locking Jack Socket
|Input impedance
|111mm X 65mm X 40mm
|Acoustic Principle
|Permanently Polarized
|Active Electronics
|JFET impedance converter
|Address Type
|Frequency Range
|60Hz - 18kHz
|Output Impedance
|Maximum SPL
|110dB SPL (@ 1kHz, 1% THD into 1KΩ load)
|Maximum Output Level
|-33.5dB re 1 Volt/Pascal (21.00mV @ 94 dB SPL) +/- 2 dB @ 1kHz
|Equivalent Noise Level (A-weighted)
|12.00mmH x 4.50mmW x 4.50mmD
[Ease of use] is where the RØDELink scores very highly. I have never seen a wireless system that is so simple to use. All you do is turn it on and it's ready to go. As it's using 2.4GHz band that I talked about earlier, the RØDELink is constantly monitoring and hopping between frequencies in real time without you having to do anything. The battery life is absolutely superb. I used the RØDELink system for more than 30hours and didn't have to change the batteries in either the receiver or transmitter.
The RØDELink system certainly offers a lot of value for money. With great audio quality, ease of use and amazing battery performance, I'm sure a lot of shooters will be very pleased with the RØDELink system.
I found the system to be very easy to use, quick to set up and very straightforward to work with.
Quite simply anyone should be able to set up and work with the Filmmaker Kit, and it's also extremely compact and lightweight to carry around as an added bonus.
5 Star, Highly Recommended
The RØDELink system has an operating range up to 100m (~110 yards), however as with all wireless transmission devices this operating range can be influenced by external factors.
The RØDELink system operates in the 2.4GHz frequency spectrum which is shared with common wireless transmission devices such as WiFi networks, Bluetooth devices, microwave ovens and more. The more 2.4GHz devices that are simultaneously running in the same location as your RØDELink system, the more congested this 2.4GHz frequency spectrum will be, cutting down the signal strength and overall operating range for all 2.4GHz devices. In a very busy 2.4GHz environment you may find your RØDELink system is unable to maintain its maximum operating range.
Signal travelling distance is also another possible factor. This refers to the physical distance between the Receiver (RX) and Transmitter (TX). An unobstructed line of sight between the two devices will provide the shortest possible signal travelling distance, therefore the strongest signal. If the signal needs to travel through, or around walls and other objects, this lengthens your signal travelling distance, reducing overall signal strength.
For more information on the 2.4GHz spectrum with your RØDELink system watch the video below.
Not simultaneously. Each RØDELink System is designed to operate as a pair and you can only receive audio from one transmitter unit at any time.
That said, each RØDELink receiver has up to eight memory locations where it can store paired transmitters so you can have one receiver unit that can connect to eight transmitters without having to re-pair the units.
To switch between paired transmitters (in this example Ch1 and Ch2) on the same receiver, follow these steps:
Yes you can, using a 3.5mm to XLR adaptor such as the RØDE VXLR.
Please note that this will be an unbalanced signal.
Yes you can connect an XLR microphone to the RØDELink Filmmaker kit, however the microphone must be self-powered as the TX-BELT transmitter unit does not provide phantom power.
To connect your XLR microphone you will need an XLR-to-3.5mm cable such as this.
RØDELink kits are shipped pre-paired from the factory, however if you wish to re-pair for any reason just follow these steps:
If you wish to pair multiple RØDELink kits, simply switch off the first set and then pair the second set using the steps above on a different channel.
No. The lav mic included is a different microphone, and does not have a removable adaptor but rather a fixed 3.5mm connector with locking thread. While both are broadcast-grade in performance, the Lavalier supports the MiCon adaptor system for a wide range of compatibility with popular audio devices.
If you have a question or a support enquiry about the RØDELink Filmmaker Kit head over to our contact page to get in touch or locate your local distributor here


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