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Proper regulation of transcription is essential for cells to acquire and maintain cell identity. Transcriptional activation plays a central role in gene regulation and can be modulated by introducing transcriptional activators such as transcription factors. Activators act on their specific target genes to induce transcription. Reprogramming experiments have revealed that as cells become differentiated, some genes are highly silenced and even introduction of activators that target these silenced genes does not induce transcription. This can be explained by chromatin-based repression that restricts access of transcriptional activators to silenced genes. Transcriptional activation from these genes can be accomplished by opening chromatin, in addition to providing activators. Once a de novo transcription network is established, cells are differentiated or reprogrammed to a new cell type. Emerging evidence suggests that actin in the nucleus (nuclear actin) and nuclear actin-binding proteins are implicated in these transcriptional regulatory processes. This review summarizes roles of nuclear actin and actin-binding proteins in transcriptional regulation. We also discuss possible functions of nuclear actin during reprogramming in the context of transcription and chromatin remodeling.
Regulation of gene expression is critically important for all living organisms. Gene expression is spatially and temporally regulated during development. For example, genes necessary for brain formation need to be expressed in the brain progenitor cells in development. If those genes are not properly expressed in progenitor cells or expressed in unrelated cells, normal brain formation is impaired. Proper gene expression is controlled by transcriptional activators. Activators, such as transcription factors and chromatin remodelers, directly act on genes to allow transcription by RNA polymerases. Each activator has distinct target DNA sequences or sites, thereby enabling gene-specific transcriptional activation. In general, different cell types contain different activators to comprise their own gene expression networks. Due to these transcriptional regulations, specialized cells remain in the same lineage and do not switch to unrelated cell lineages.
Nuclear reprogramming enables cells to reset transcriptional patterns of specialized cells and establish those of embryonic cells [1]. Reprogramming was first demonstrated by nuclear transfer experiments, where differentiated somatic cell nuclei were transferred to enucleated eggs and the reconstructed cloned embryos then developed to term [2, 3]. Successful reprogramming of transcription programs in cloned embryos has been shown using many different cell types, suggesting that the transcriptional programs of any cell type can be modified towards another cell type. This concept has been further reinforced by the success of factor-mediated reprogramming, creating induced pluripotent stem (iPS) cells [4–6] and transdifferentiation by overexpression of transcription factors [7–9]. During iPS cell production, key transcription factors for pluripotency are overexpressed in somatic cells and establish a new transcription network resembling or almost identical to embryonic stem (ES) cells in the reprogrammed cells. Similarly, overexpression of tissue-specific transcription factors triggers transdifferentiation [9]. Reprogramming experiments thus tell us (1) the importance of transcriptional activators for cell differentiation/dedifferentiation and (2) that gene expression can be altered even in highly specialized cells.
Increasing evidence suggests that actin in the nucleus (nuclear actin) and nuclear actin-binding proteins (ABPs) play an important role in transcriptional activation and transcription [10–12]. Actin has been known as a major component of the cytoskeleton and as a key player in many cellular processes including cell migration, division, and shaping. Actin continuously changes its polymerized states, at least in the cytoplasm; monomeric actin (G-actin) polymerizes at the barbed end to form filamentous actin (F-actin). Significant amounts of actin are also found in the nucleus [13]. Nuclear actin has been identified as an important component of transcriptional machineries and chromatin remodeling complexes. Similarly, numerous actin-binding proteins are present in the nucleus [14] and implicated in transcription and chromatin remodeling [15, 16]. In addition, recent studies indicate that nuclear actin and actin-binding proteins play vital roles in transcriptional activation during cell differentiation and reprogramming [17–19].
This review describes our current knowledge about mechanisms of transcriptional activation and transcriptional reprogramming, followed by discussion of roles of nuclear actin and actin-binding proteins in these cellular events.
Transcription from previously silenced genes can be induced by introduction of transcriptional activators (Fig. 1). In a special type of experimental design, nuclear gene expression of one kind of cell is switched to that of an embryo or other cell type, referred to as transcriptional reprogramming. Transcriptional reprogramming is achieved by different approaches, such as induced pluripotency, cell fusion, and nuclear transfer to eggs/oocytes [20]. However, it is generally accepted that the efficiency of transcriptional reprogramming is low [21]. This is due to the fact that gene expression is often repressed by layers of silencing mechanisms to maintain transcriptionally quiescent states [22]. Gene silencing seems to be progressively more difficult to reverse as cells become increasingly differentiated. This is exemplified by sequential epigenetic modifications for Oct4 silencing during ES cell differentiation [23, 24] and differential gene reactivation from the inactive X chromosome between epiblast stem cells and differentiated mouse embryonic fibroblasts [25]. Moreover, mechanisms and extents of gene silencing are different depending on each gene. Lahn and his colleagues [26, 27] have proposed that genes that are not expressed can be classified into two categories based on resistance to transcriptional activation during cell-fusion-induced reprogramming (Fig. 1); (1) genes that are silent in one cell type are expressed (activatable) when that cell is fused with another cell type in which that gene is already active, and (2) genes that are silent in one cell type remain inactive (occluded) when that cell is fused to another cell type in which that gene is active [26, 27]. The former activatable genes are not expressed because of the absence of transcriptional activators and/or the presence of repressors and introduction of activators is enough to induce transcription. The latter occluded genes are proposed to be inactive due to chromatin-based repression mechanisms that maintain silent states regardless of whether transcriptional activators are present. Full activation of occluded genes normally requires multiple cell divisions and a longer time exposure to a cellular milieu that supports transcription from occluded genes than activatable genes, implying that de-repression of chromatin-based inhibition proceeds gradually. Although the classification has been carried out in the context of cell-fusion-mediated reprogramming, this concept seems to be generally applicable to other reprogramming systems and cellular events. For example, addition of retinoic acid (RA) to cells can cause rapid induction of transcription from RA-responding genes such as Hox genes. These transcribed genes are most likely activatable ones since they can respond to activators (RA receptor in this case). On the other hand, in iPS experiments, expression of many embryonic genes from somatic cells requires several days and cell divisions, although known transcriptional activators of these embryonic genes are highly expressed in the somatic cells transduced [28, 29]. This result argues that silenced embryonic genes in somatic cells are likely to be occluded genes. It is important to know the states of gene silencing when we study gene activation and transcriptional reprogramming.
Reprogramming experiments have revealed that resistance to transcriptional activation differs in different genes. They also bring up an intriguing question of what kinds of factors can overcome this resistance. In activatable genes, transcriptional activators are enough to rapidly induce transcription. This also suggests that those genes need to be accessible to activators. In other words, genes may have permissive chromatin states. Permissive chromatin is characterized by the presence of active histone marks such as histone acetylation, and the absence of repressive marks such as histone H3 lysine 9 (H3K9) methylation and DNA methylation [30]. Alternatively, it can be free from nucleosome occupancy [31, 32]. It is also possible that activatable genes are bound by paused RNA polymerase II (Pol II) [33] and introduction of activators enhances the transcriptional elongation step from paused Pol II. In fact, Pol II pausing is seen in many developmentally important genes for quickly achieving gene activation since these genes tend to be occluded by nucleosome formation without paused Pol II [34]. Thus, Pol II itself can work as a factor that precludes gene occlusion. Once occlusion happens, additional factors are necessary to induce transcriptional activation from such genes. In occluded genes, activators cannot access DNA-binding sites, such that RNA polymerase cannot be recruited to these genes. Heterochromatin formation may prevent access of activators. It is often associated with repressive marks such as DNA methylation, histone H3K9 di- or trimethylation, and repressors such as histone deacetylases and heterochromatin proteins. Reprogramming experiments have shown that these repressive modifications are removed before transcriptional reprogramming [35–37] and that removal of these modifications prior to reprogramming enhances its efficiency [24, 38, 39]. These results suggest that repressive modifications leading to compacted chromatin impede gene activation. It is hence plausible that addition of chromatin modifiers that can remove these repressive modifications is effective in reprogramming silenced genes [40–44]. Moreover, activation of silenced genes during reprogramming is enhanced by a chromatin remodeling factor, BAF complex [45], and addition of trans-activating domains to transcription factors [46, 47]. These factors allow efficient access for transcriptional activators to target genes. In conclusion, expression from occluded genes can be achieved by opening chromatin, in addition to providing transcriptional activators.
Mechanisms of transcriptional activation and reprogramming have been described so far. A transcriptional activation process is affected by many nuclear events. Nuclear actin plays key roles in such events including basal transcription by all three RNA polymerases (Pol II case in Fig. 2) [48–51], chromatin remodeling [11, 52, 53], pre-mRNA processing [12, 54], and gene movement [55]. Moreover, recent studies indicate that nuclear actin is directly or indirectly involved in transcriptional activation (Fig. 3) [56–58]. Here, possible roles of nuclear actin in those cellular events are discussed.
Involvement of nuclear actin in transcription was described for the first time in the early 1980s [59]. Injection of antibodies against actin into nuclei of amphibian oocytes induced retraction of lampbrush chromosomes, reminiscent of transcriptional cessation [59]. Recent studies using more sophisticated in vitro and in vivo transcription assays confirmed the concept of this early study. Actin is found in the preinitiation complexes (Fig. 2) and is important for transcriptional initiation by Pol II [49]. The phosphorylated C-terminal domain of Pol II is associated with actin [51]. Actin is also known to interact with the nascent transcripts [54] and heterogeneous ribonucleoproteins (hnRNPs) mediate binding of actin to the nascent RNAs [51]. hnRNPs and actin binding is necessary for transcription elongation by Pol II since a competing peptide that disrupts hnRNPs–actin interaction inhibited transcription elongation when examined by run-on assay [60]. The hnRNPs-actin interaction is important for the association of histone acetyltransferase PCAF and Ser2- and Ser2/5-phosphorylated Pol II [61]. This study [61], together with the work using Chironomus tentans [62], also brings a key concept that actin takes part in establishing permissive chromatin through recruitment of the histone acetyltransferase during transcription. Ser2 phosphorylation of Pol II is mediated by positive transcription elongation factor b (P-TEFb). Actin, especially monomeric actin (G-actin), facilitates recruitment of P-TEFb to Pol II [63], further affirming the importance of nuclear actin in transcription elongation. Collectively, nuclear actin is directly involved in RNA polymerase-mediated transcription through enhancing preinitiation complex formation and transcription elongation. One model of nuclear actin- and actin-binding proteins-mediated transcription is shown in Fig. 2 and our model is explained in the figure legend. Details about nuclear actin in transcription are discussed in recent insightful reviews [10, 12, 64].
Actin and actin-related proteins (Arps) have been found in many chromatin remodeling complexes [11, 16, 65, 66]. These remodelers play key roles in various nuclear events such as transcription, DNA repair, DNA replication, and nuclear organization. Since the functions of ARPs have been extensively reviewed [16, 65, 66], this paper discusses possible roles of actin in chromatin remodeling. An early discovery that nuclear actin participates in chromatin remodeling was made in the late 1990s. β-actin and ARP4 (also known as BAF53) are found in the mammalian SWI/SNF-like BAF chromatin remodeling complexes and tightly associate with the ATPase Brg1 in the complex [52]. The BAF complex requires actin for its full ATPase activity and its association with chromatin [52]. A recent study suggests that interaction between β-actin and ARP4 is important for the integrity of the BAF complex since interruption of heterocomplex formation by β-actin and ARP4 accelerates degradation of Brg1 [67]. The heterocomplex formation by G-actin and ARP4 is supported by biochemical studies [68]. Interestingly, Fenn et al. [68] has revealed that ARP4 is able to enhance depolymerization of actin filaments. This result brings up the possibility that polymerized states of nuclear actin may influence assembly or activity of the BAF complex. In fact, the BAF complex can bind to actin filaments [69]. It would be interesting to know the causal relationship between nuclear actin polymerization and activities of BAF complex. Another chromatin remodeling complex that contains actin and ARP4 is the INO80 complex. The catalytic ATPase subunits of both INO80 and BAF complexes contain the important domain for binding to ARPs and actin, referred to as the helicase-SANT-associated (HSA) domain [70]. In the INO80 complex, ARP4, ARP8, and actin are bound to the HSA domain in the core ATPase INO80 and ARPs directly interact with histones so that the INO80 complex can gain access to nucleosomes [71]. Thus, important progress has been made to decipher the roles of nuclear actin in chromatin remodeling. Interdisciplinary approaches using structural biology and molecular cell biology will accelerate our understanding of nuclear actin in chromatin remodeling.
The implication of nuclear actin in basal transcription has been demonstrated. Interestingly, transcriptional regulation by actin is not limited to polymerase-mediated transcription itself. There seem to be several different mechanisms to regulate transcriptional activation. A first clear example of participation of nuclear actin in gene activation has been demonstrated by the Treisman group. Activation of serum response factor (SRF), a transcription factor that regulates many serum-inducible and muscle-specific genes, coincides with F-actin accumulation in cells [72]. Studies using actin mutants that do not polymerize or enhance polymerization revealed that G-actin is the regulator of SRF activity and activation of MAL, a coactivator of the SRF transcription factor [73, 74]. G-actin binds to MAL in nuclei. This actin binding inhibits MAL's function as a transcription activator and enhances export of MAL to the cytoplasm in a serum-starved condition [56]. After serum stimulation, F-actin is accumulated in the cytoplasm and this should lead to a decreased G-actin pool, thereby increasing active MAL free from G-actin binding. Thus, nuclear actin, in concert with cytoplasmic actin, regulates transcriptional activation by sequestering an activator (Fig. 3a). This transcriptional regulation by actin polymerization-mediated control of the nucleo-cytoplasmic distribution of transcription regulators does not seem limited to MAL's case [75, 76]. Since many signaling molecules affect cytoplasmic actin polymerization, this sort of an indirect effect of actin on gene transcription through sequestration might be seen in other signaling pathways.
Secondly, nuclear actin is involved in induction of HoxB transcription by retinoic acid (RA) treatment [57]. It is known that RA receptors are important for expression of Hox gene and HoxB expression depends on a Prep1–Pbx1 complex that works as a transcription activator [77]. Proteomic analysis revealed that Prep1 binds to Pol II and nuclear β-actin [78]. Prep1 also interacts with nuclear N-WASP (neuronal Wiskott–Aldrich Syndrome Protein) that enhances actin polymerization [57]. Depolymerization of actin, N-WASP knockdown, and overexpression of an actin mutant that does not polymerize inhibit HoxB transcription after RA treatment. Interestingly, chromatin immunoprecipitation (ChIP) analysis has revealed that elongating Pol II, Prep1, actin, and N-WASP are recruited to the HoxB enhancer in an actin polymerization-dependent manner. These results suggest that actin polymerization is required for HoxB gene activation possibly by mediating recruitment of the transcription complex to the regulatory region. It would be interesting to examine whether recruitment of the transcription complex to the HoxB coding region is similarly regulated by actin polymerization and whether N-WASP-mediated actin polymerization or other nuclear actin polymerization is critical for this recruitment. Therefore, nuclear actin seems important for recruiting active transcription machineries during gene activation (Fig. 3b).
Another interesting phenomenon during gene activation is translocation and accumulation of cytoplasmic actin to nuclei [19]. Actin dynamically shuttles between the nucleus and cytoplasm. Actin export from nuclei is accomplished by Exportin 6 [79]. However, it has been elusive whether cytoplasmic actin is actively imported into nuclei, although we know that cofilin is important for actin import in special circumstances, such as stress. Recently, importin 9 has been identified as a key molecule that actively imports cytoplasmic actin to nuclei possibly with cytoplasmic cofilin [80]. Moreover, this active maintenance of nuclear actin by importin 9 is necessary for maximal transcriptional activity for cells [80]. In accordance with this report, translocation of cytoplasmic β-actin to nuclei is observed during differentiation of human promyelocytic leukemia (HL-60) cells towards macrophages; this entails activation of many genes for successful differentiation [19]. During differentiation, association of nuclear actin with Pol II is observed. ChIP-on-chip assays revealed a striking increase of nuclear actin binding to gene promoters (25 to 827 genes). Knockdown of β-actin inhibits Pol II binding to promoters, suggesting that nuclear translocation of actin during differentiation allows efficient recruitment of Pol II to target genes (Fig. 3c). Interestingly, when cells become quiescent, nuclear β-actin is depleted and Pol II binding to transcription sites is destabilized [81]. This report further supports the idea that nuclear actin levels are an important determinant of transcriptional activity.
As mentioned above, nuclear actin has been identified in many chromatin remodeling complexes and chromatin modifiers. One study shows involvement of oligomeric actin in chromatin remodeling and gene activation [58]. Transcription from Toll-like receptor (TLR)-responsive genes is inhibited by nuclear receptor co-repressor (NCoR) complexes and clearance of this NCoR complex from promoters is necessary for transcriptional activation from TLR target genes. The NCoR complex includes Coronin 2A, an actin filament-binding protein. Nuclear actin binds to Coronin 2A and this binding triggers clearance of repressive NCoR complexes from the promoters of target genes, thereby inducing transcriptional activation. These results suggest that transcriptional activation by nuclear actin is achieved by removing gene silencing complexes from chromatin (Fig. 3d).
Nuclear actin-binding proteins as well as nuclear actin play a crucial role in transcription and transcriptional activation. One of the most studied nuclear actin-binding proteins is an isoform of myosin I (nuclear myosin I; NMI) [82, 83]. NMI is involved in transcription by RNA polymerase I [48, 84, 85] and II [86] and interacts with the chromatin remodeling complex WSTF-SNF2h [87]. Many other nuclear myosins have been identified and their diverse nuclear roles have been found [10]. Since functions of nuclear myosins are extensively summarized in recent reviews [10, 12, 88], this review will focus on other actin-binding proteins that can be involved in transcription and transcriptional regulation.
Members of the Wiskott–Aldrich syndrome protein (WASP) family are key factors of actin polymerization by regulating the actin-related protein 2/3 (Arp2/3) complex, an actin nucleator [89–91]. The WASP family consists of two classes of proteins; WASPs (WASP and N-WASP) and WAVEs (WAVE1, WAVE2, and WAVE3). Mutant WASP was identified as the causative gene of the Wiskott–Aldrich syndrome and WASP is expressed in hematopoietic cells while N-WASP is ubiquitously expressed. Although the WASP family proteins were originally identified as cytoplasmic proteins that regulate cortical actin filaments, nuclear WASPs and WAVEs have also been detected [17, 92–94]. N-WASP forms a nuclear protein complex containing non-Pou-domain octamer-binding protein (NonO), polypyrimidine-tract-binding-protein-associated splicing factor (PSF) and Pol II in human 293T cells [93]. Moreover, nuclear N-WASP regulates Pol II-mediated transcription through its interaction with NonO, as well as through the induction of actin polymerization. Interestingly, NonO has been shown to bind to hyperphosphorylated Pol II [95] and N-WASP can also be associated with elongating Pol II [57]. Therefore, it is tempting to speculate that N-WASP plays a role in the elongating step of transcription (Fig. 2). Since N-WASP is also found on gene promoters and binds to unphosphorylated Pol II [93], it is possible that N-WASP is also involved in transcriptional initiation or recruitment. In hematopoietic cells, WASP is expressed instead of N-WASP. WASP translocates into nuclei during T cell differentiation [17]. Nuclear WASP binds to histone modifying enzymes such as RBBP5, a histone H3K4 tri-methyltransferase, and JMJD2A, a H3K9/H3K36 tridemethylase, and RNA Pol II. WASP also binds to the transcription factor SP1. This SP1 binding seems important for recruiting WASP to specific target genes for transcriptional activation. Collectively, WASP plays an important role in transcriptional activation of genes required for T cell differentiation by regulating active histone marks and possibly transcription. Moreover, the actin nucleating Arp2/3 complex is found in nuclei and is required for a full Pol II activity [96]. Arp2/3 and F-actin bind to the gene regions to which WASP is bound [17]. The WASP family proteins have an ability to mediate various protein–protein interactions and this characteristic may help them to associate with many nuclear proteins and nuclear actin.
Cofilin/ADF (actin depolymerizing factor) enhances depolymerization of actin by severing F-actin [97]. Cofilin binds to G-actin and forms a heterodimer. This actin-cofilin complex is imported into nuclei [80, 98]. Cofilin-1 in nuclei seems to bind to the elongating type of RNA polymerase II and actin [99]. ChIP analysis revealed that cofilin-1 is associated with gene coding regions, but not with the promoter. Since G-actin is important for transcriptional elongation [63], cofilin-1 may provide a G-actin pool for Pol II elongation. Considering the cofilin's F-actin severing function, actin polymers may exist adjacent to transcribing genes to provide enough cofilin-1 enrichment. Such polymeric actin might be produced by N-WASP and Arp2/3 functions and further accelerated by profilin (Fig. 2). The regulation of nuclear actin polymerization with the help of cofilin and profilin during transcription is summarized in [64].
A list of other nuclear actin-binding proteins is summarized in [14, 15, 100, 101]. The number of nuclear actin-binding proteins is likely to expand in the future [102, 103]. It is therefore important to define the relationship among these proteins. Most of the actin-binding proteins listed here are connected in the context of transcription and actin polymerization. Such classification will help to get a better view of the nuclear actin network and possibly nucleoskeleton [88]. It is also important to bear in mind that actin-binding proteins can have actin-independent roles in nuclei. This is exemplified by the profilin case, where profilin and actin seem to be recruited differentially to chromosomes [104]. We have recently found another such case in which a nuclear actin-binding protein can bind to the transcription apparatus without an actin-binding domain (unpublished data).
Nuclear actin serves as an important factor for gene activation by several different mechanisms. Most of the cases discussed so far deal with transcriptional activation from activatable genes, but not from occluded genes. This is because introduction of transcriptional activators is enough to induce transcription in these cases and nuclear actin functions in this context. Being different from these examples, transcriptional reprogramming entails activation of occluded genes. Transcriptional reprogramming can be induced in a direct and rapid manner by transplanting somatic nuclei into the nucleus of a Xenopus oocyte called the germinal vesicle (GV) [105]. Reprogramming in eggs/oocytes is proposed to be the most efficient way to induce pluripotency in somatic nuclei [105, 106] and activation of Oct4, an embryonic gene that was previously identified as an occluded gene in somatic cells in cell fusion experiments [26], is induced as early as 20 h after nuclear transfer as judged by the reporter expression [107]. In agreement with this, transcription from many embryonic genes including Oct4, Sox2, and Nanog is induced within a few days in somatic nuclei transplanted into Xenopus oocytes [108]. Thus, oocytes seem to have an ability to reprogram occluded genes that were defined in somatic–somatic or even somatic–ES cell fusion experiments [26]. Nuclear transfer to oocytes therefore provides an opportunity to evaluate unique oocyte factors that can overcome gene occlusion. Xenopus oocytes contain enormous amounts of actin in nuclei [109–111] and this nuclear actin has been linked to transcription from the oocyte genome [59]. Moreover, nuclear actin is found in transplanted nuclei [18]. Interestingly, polymerized actin is formed in nuclei transplanted to Xenopus oocytes, resembling nuclear actin seen in the GV of oocytes [18]. Disturbance of this nuclear actin polymerization impairs activation of Oct4 transcription in transplanted nuclei, suggesting that nuclear actin polymerization is important for transcriptional activation of Oct4. Thus, nuclear actin seems involved in reversal of an occluded gene (Fig. 4), but we still do not know which step during reversal of silencing is caused by nuclear actin. One possibility is that nuclear actin accelerates chromatin remodeling by changing its polymerized states (Fig. 4). In fact, enhanced binding of actin to Oct4 by overexpression of actin-binding protein Toca-1 coincides with enhanced BAF complex binding to Oct4 during reprogramming [18]. It is also possible that actin polymers play a role in the clearance of transcriptional repressors as is the case of the NCoR complex (Fig. 4) [58]. We also ought to take into account the possibility that nuclear actin enhances recruitment or elongation of Pol II.
Possible participation of nuclear actin and actin-binding proteins in nuclear reprogramming has been suggested not only in the Xenopus nuclear transfer system, but also in other reprogramming systems. Firstly, actin-containing BAF remodeling complex is crucial for embryonic gene activation during reprogramming by the iPS route and egg extract treatment [45, 112] and is proposed to convert inaccessible chromatin of embryonic genes in somatic cells to accessible states for transcriptional activators [45]. It would be interesting to examine the function of nuclear actin in concert with the BAF complex during reprogramming. Secondly, Polycomb repressive complex (PRC) is well known as a chromatin remodeler that silences developmentally important genes such as Hox genes and is essential for normal development. It is also necessary for reprogramming somatic nuclei towards pluripotency in cell-fusion and iPS experiments [113, 114]. Ezh2 is a catalytic subunit of PRC2. Interestingly, Ezh2 is also present in the cytoplasm and regulates cytoplasmic actin polymerization [115]. Although it is not clear whether Ezh2 regulates nuclear actin polymerization in a similar manner, PRC is an interesting candidate for examining its relation to nuclear actin. Thirdly, the translationally controlled tumor protein (TCTP) is involved in regulating many cellular processes such as cell proliferation and apoptosis and is necessary for development. TCTP has been shown to enhance activation of pluripotency genes including Oct4 in transplanted nuclei into Xenopus oocytes [116]. TCTP is also implicated in nuclear reprogramming in bovine oocytes although the mechanisms are still unknown [117]. Intriguingly, TCTP contains cofilin-like actin-binding site and binds to G-actin [118] and Xenopus TCTP is also associated with F-actin [119]. The observed positive effects of TCTP on transcriptional reprogramming therefore might be through altering nuclear actin polymerization. Lastly, actin in mammalian oocytes seems to have a great impact on reprogramming. Incorporation of oocyte actin into somatic nuclei is observed during incubation in porcine oocyte extracts [120, 121], which are known to induce a part of early reprogramming events [122]. This implies that actin is likely to be involved in an early step of reprogramming in oocytes. In accordance with this idea, different groups have recently reported that treatment of nuclear transferred embryos with actin depolymerizing reagents greatly affects development of these embryos; Latrunculin A, instead of Cytochalasin B, significantly improves cloning efficiency [123–125]. It is unclear whether this positive effect is caused just by reduced cytotoxicity of the actin depolymerizing reagent used or by improved some reprogramming aspects related to actin polymerization states. Answering this question may advance our understanding on nuclear actin in reprogramming.
Several studies have thus implied that nuclear actin is implicated in nuclear reprogramming. However, functions of nuclear actin in reprogramming are obscure. In this section, we propose some speculative roles of nuclear actin during reprogramming. When we define nuclear reprogramming as a phenomenon in which differentiated cells are reversed to an embryonic state, reprogramming is composed of several distinct processes, such as extinction of differentiation gene expression, initiation and continuous expression of embryonic genes, and establishment of embryonic cell properties including high DNA repair activities, high telomerase activities, and embryonic cell-specific nuclear architectures, etc. One plausible role of nuclear actin in the reprogramming process is that actin is required for transcription of embryonic genes. Many embryonic genes need to be activated and continuously expressed during reprogramming. Nuclear actin might support efficient transcription from those genes by accelerating Pol II initiation and elongation. Also, nuclear actin-mediated chromatin remodeling might be important for reprogramming. Nuclear reprogramming entails activation of occluded genes, which are silenced due to the chromatin-based repression mechanisms (Fig. 4). The BAF, INO80, and Tip60 chromatin remodeling complexes that contain actin can help to establish open or unstable chromatin states. Nuclear actin may affect the activities of such remodeling complexes to induce transcription from occluded genes (Fig. 4). Therefore, it is interesting to investigate the binding of nuclear actin and such remodeling complexes to embryonic genes during reprogramming. In fact, the BAF and Tip60 complexes have been shown to maintain pluripotency by regulating gene expression in ES cells [126–128]. It is also noteworthy that reprogramming towards an embryonic state is accompanied not only by activation of embryonic genes but also by repression of differentiation genes. A recent study, in which β-actin knockout cells are used, showed that β-actin seems to have both gene-activating and gene-repressing activities [129]. Although this idea needs to be further tested, the new actin's function in repressing genes can be explained in concert with chromatin remodeling complexes. The actin-containing BAF complex also plays a role in silencing genes in ES cells [126]. Therefore, nuclear actin might be involved in silencing differentiation genes through chromatin remodeling during reprogramming.
Apart from roles of nuclear actin in transcription, nuclear actin-mediated DNA repair might be involved in the reprogramming process. Embryonic cells and iPS cells possess high DNA repair capacities and DNA repair pathways have to be active for successful reprogramming in the iPS route [130]. Moreover, DNA repair events are likely to be induced in nuclear transfer embryos [131]. A recent study has shown that polymeric actin is required for proper DNA double-strand break repair [132]. Together, it may be worth pursuing the relationship between nuclear actin polymerization and DNA repair during reprogramming and in reprogrammed embryonic cells.
Importantly, we also need to take account of the fact that actin may affect mechanical properties of the nucleoskeleton. Some nucleoskeleton structures are cell type-specific, such that metastatic cancer cells exhibit abnormal chromatin organization [88]. Embryonic cells, as well as reprogrammed cells, seem to possess a specific nucleoskeleton structure since they have a different lamin composition from differentiated cells [133, 134] and exhibit a different chromocenter compartment [135]. Changes of nucleoskeleton organization can influence expression of a large number of genes (>1,000 genes) [136] and hence establishment of a proper nucleoskeleton is important. Actin may coordinate nucleoskeleton organization processes through interactions with its many binding partners in nuclei. Interesting insights might be gained by examining the effects of altering actin polymerization on chromatin reorganization during reprogramming.
As mentioned above, multiple steps are required for reprogramming somatic cells. Nuclear actin seems to play a role in many steps during reprogramming; this argues that actin might be an important player in reprogramming. This is attributed to multifunctional properties of nuclear actin. Such properties of actin probably result from the fact that it has a myriad of binding partners and can affect activities of its binding molecules. To reveal how nuclear actin participates in each step of reprogramming will expand our understanding of reprogramming and nuclear actin biology.
Transcriptional activation is a fundamental cellular process essential for living organisms. Transcription can be induced by adding activators such as transcription factors. However, in some cases, genes are highly silenced (occluded) with layers of repressing mechanisms, which cannot be easily overcome by activators due to the lack of access to these silenced genes, requiring transcriptional reprogramming that includes de-repression of silenced chromatin states. Nuclear actin seems to play important roles in transcriptional activation from both activatable and occluded genes. This intriguing feature of nuclear actin may be attributed to its significant function in basal transcription and chromatin remodeling. Furthermore, nuclear actin and nuclear actin-binding proteins participate in other nuclear processes that potentially relate to transcriptional activation and reprogramming such as nucleoskeletal activities [88] and gene movement upon activation [55, 137, 138]. Recent studies have shown that more and more proteins that regulate actin dynamics are found in nuclei [102, 103]; these imply that our understanding of the functions ascribed to nuclear actin may be extended. For obtaining a global view of the relationship between nuclear actin and actin-binding proteins, it would be interesting to investigate the association of actin and actin-binding proteins with genes in a genome-wide level by ChIP-seq analysis. A genome-wide study using Drosophila cells has shown that actin is associated with active euchromatin regions [139], in agreement with the general view that actin works as a positive regulator of transcription. Examining the genome-wide binding of nuclear actin and actin-binding proteins in different cell types and different conditions such as during differentiation will help us to achieve an overview of actin-mediated regulation of gene transcription. This may also answer the important question as to how gene expression is regulated by nuclear actin in a gene-specific manner. Nuclear actin is not just a byproduct of abundant cytoplasmic actin. A better understanding of nuclear actin in transcriptional activation and reprogramming will lead on to revealing its unknown functions in cellular and developmental contexts.
We thank Drs. R.P. Halley-Stott, M. Teperek, and J. Jullien for their careful reading of the manuscript. We apologize to authors whose interesting works could not be cited due to the focused scope of this review. K.M. is a Research Fellow at Wolfson College in the University of Cambridge and is supported by the Herchel Smith Postdoctoral Fellowship. The Gurdon laboratory is supported by grants from the Wellcome Trust and MRC.
PubMed Central Canada is a service of the Canadian Institutes of Health Research (CIHR) working in partnership with the National Research Council's national science library in cooperation with the National Center for Biotechnology Information at the U.S. National Library of Medicine(NCBI/NLM). It includes content provided to the PubMed Central International archive by participating publishers.


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35 Wallpapers with Exclusive Visual Arts for making this Valentine's Day more Romantic
We all love to see creative and different visual arts whatever the topics. And if the arts or design do match with our specific theme or occasion then it gives more joy to us.
For the lovers there is no other best occasion then Valentine's Day. Most of the lovers fixed the day for expressing their feelings of love. So here I collect 36 gorgeous collections of wallpapers to decorate your desktop on Valentine's Day to make it more Romantic for the lovers.
Amazing Realistic Street Arts Part – II
Few months ago we made a post on amazing street arts for GeekSucks, which got lots of response from our visitors. There were varieties of responses; some appreciated the post and some showed their doubt on those realistic street arts. It is true that, some of arts were really unbelievable to trust.
Now in this time, we came again with some new unbelievable realistic street arts which are also called pavement art. This time you will get some source links with the image. Hope you will enjoy this second part of amazing street arts like previous.
Amazing Realistic Street Arts
Art on the public place or in the streets is not a new trend. In between 1999 and 2000, this art was firstly getting huge popularity among the world. Previously this art work was done by the artists on their own wish, to impress the viewers. But now some of companies or corporate houses are investing on this work for their product or brand promotion. Around the world, here is a few number of street artist and in between them Julian Beever is a popular name.
Here you'll find some of creative, amazing and realistic street arts. I think it will impress you.

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GORGEOUS HOME!!!...with spectacular upgraded 4 bedrooms and 3 bathrooms, 2187 living sqft located in a center of Westminster. Custom double entry doors lead to spacious living room with beautiful Cathedral Vaulted Ceiling and, modern recessed lights. Laminated- wood flooring throughout most rooms. Fully upgraded kitchen with gorgeous craftsmanship cabinets and granite counter tops, plus large dining area opens up to family room. Attached 2 car garage. Beautiful front and back yards landscaped with palms trees and a lot of fruit trees. Conveniently close to Little Saigon, main shopping malls, good school and Freeway 405 and 22. READY TO MOVE IN….. IT'S A MUST SEE!, 1 near corner of Goldenwest and Westminster
Interior Features Include:
Cathedral-Vaulted Ceilings, Granite Counters, Recessed Lighting
Fireplace Family Room, Gas, Decorative
Window Cov: Drapes/Curtains
Air Cond: Central
Heating: Central Furnace
Floor Type: Laminate, Tile
Exterior Features Include:
Fencing Blockwall
Acreage: 0.1000 Acres
Lot: Back Yard, Front Yard, Garden, Landscaped, Lot Over 40000 Sqft
Construction Stucco
Foundation Slab
Parking Driveway, Direct Garage Access, Garage - Front Entry
Parking Spaces 2.00
Garage Capacity: 2
Other Features Include:
Subdivision: Other (OTHR)
Appliances Dishwasher, Gas Stove
Amenities: Playground
Water: Public
Pool: Association
Security Features Automatic Gate, Gated Community
Community Features Sidewalks, Street Lighting, Suburban
View None
The information being provided by CARETS (CLAW, CRMLS, DAMLS, CRMLS, i-Tech MLS, and/or VCRDS) is for the visitor's personal, non-commercial use and may not be used for any purpose other than to identify prospective properties visitor may be interested in purchasing.
Any information relating to a property referenced on this web site comes from the Internet Data Exchange (IDX) program of CARETS. This web site may reference real estate listing(s) held by a brokerage firm other than the broker and/or agent who owns this web site.
The accuracy of all information, regardless of source, including but not limited to square footages and lot sizes, is deemed reliable but not guaranteed and should be personally verified through personal inspection by and/or with the appropriate professionals. The data contained herein is copyrighted by CARETS, CLAW, CRISNet MLS, DAMLS, CRMLS, i-Tech MLS and/or VCRDS and is protected by all applicable copyright laws. Any dissemination of this information is in violation of copyright laws and is strictly prohibited.
CARETS, California Real Estate Technology Services, is a consolidated MLS property listing data feed comprised of CLAW (Combined LA/Westside MLS), CRISNet MLS (Southland Regional AOR), DAMLS (Desert Area MLS), CRMLS (California Regional MLS), i-Tech MLS (Glendale AOR/Pasadena Foothills AOR) and VCRDS (Ventura County Regional Data Share).
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The summer is well and truly over; the nights are getting darker and there is a nip in the air during the evening that means that those nights that we can enjoy dining outside are numbered.
However, this doesn't mean there is nothing to see or do in Nice in autumn. In fact, for some, it is the most charming time of year and can certainly be in the running for the most beautiful season. The red leaves on the trees, the golden glow of the brush and those sunsets that turn the world the pink and purple - for me, this is the city of Nice at its best.
From classical to disco, live music begins the month of October. Saturday 1st October sees the Discollection Tour 2016 hit Nice so that you can relive your disco years with this legendary concert. There will performances from Village People, Boney-M, Precious Wilson and many, many more. On the same evening the elegant Opera de Nice will be the venue for a classical concert of Mendelssohn and Mahler, performed by the philharmonic orchestra.
Romance is in the air during the second weekend of the month as the Nice Wedding Fair takes place in the Palais des Congres. The first wedding show of the year along the French Riviera, you will find a collection of dresses, flowers, make-up and venues all under one roof. Try the latest collections of wedding dresses, discover new menus,check out the caterers and choose the designer who will decorate your dream venue.
The city of Nice is well-known for hosting some of the biggest music concerts in France and October is no exception. Sunday 9th October is a date for your diary as Ben Harper and The Innocent Criminals are in town to perform at the Palais Nikaia at one of only eight dates in the country. Ben Harper, three-time grammy award winner, is known for his guitar skills, vocals and stunning live performances, and now he rejoins the band 'The Innocent Criminals' for this tour.
Culture hits Nice in the middle of October with the International Contemporary Art Fair. art3f will be presented in the Palais des Expositions and aims to enhance the relationship between contemporary artists and art lovers.
During the month, the city of Nice will also be hosting several free classical concerts where the philharmonic orchestra of Nice will be performing outside in Place Saint Roch. Saint Jean Cap Ferrat will be holding their annual Braderie event from the 21st until the 24th October, where you can grab some of the best bargains on the Cote d'Azur...as well as some very special tasting menus in the local restaurants.
You may prefer just to sit in a cafe or bar on the Promenade des Anglais and watch the world go by, or even to visit one of the many art galleries and museums. Whatever you feel like doing, don't forget to check out our hotel and restaurant listings for the best deals, and fall for autumn in Nice.
Location: Nice


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I went to adjust the APC cache size on one of my Ubuntu 10.04 LTS (Lucid) servers today. This particular server runs a single app, and it only uses 12M of cache, so I wanted to change the cache size from the default 32M to 16M. I put
apc.shm_size=16M in
/etc/php5/conf.d/apc.ini and restarted Apache… and to my surprise, Apache didn't come back up. I found this error in the error log:
Which (via Google) led me to this very thorough blog post.
TL;DR, for older versions of APC like the version that ships with Ubuntu 10.04 LTS (APC version 3.1.3p1), you cannot use the "M" suffix. The correct syntax is
According to this bug report, it looks like APC >= 3.1.4 expects the M/G suffix, so this shouldn't be an issue with Ubuntu 12.04 LTS (Precise).


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This news was a few days back, but, if you didn't know, I'll post a snippet of it.
Emma Watson's "HeforShe" Speech as UN Women Goodwill Ambassador
UN Women
Actor and UN Women Goodwill Ambassador Emma Watson gave a game-changing speech at the UN about feminism and the need for gender equality. Her speech kicked off the HeforShe campaign, which encourages men to join the fight for gender equality. Full transcript: http://owl.li/BSbad
Some of our favorite excerpts:
"I started questioning gender-based assumptions when at eight I was confused at being called "bossy," because I wanted to direct the plays we would put on for our parents—but the boys were not. When at 14 I started being sexualized by certain elements of the press. When at 15 my girlfriends started dropping out of their sports teams because they didn't want to appear "muscly." When at 18 my male friends were unable to express their feelings."
"Men—I would like to take this opportunity to extend your formal invitation. Gender equality is your issue too."
"If men don't have to be aggressive in order to be accepted women won't feel compelled to be submissive. If men don't have to control, women won't have to be controlled. Both men and women should feel free to be sensitive. Both men and women should feel free to be strong… It is time that we all perceive gender on a spectrum not as two opposing sets of ideals.
If we stop defining each other by what we are not and start defining ourselves by what we are—we can all be freer and this is what HeforShe is about. It's about freedom."
"You might be thinking who is this Harry Potter girl? And what is she doing up on stage at the UN. It's a good question and trust me I have been asking myself the same thing. I don't know if I am qualified to be here. All I know is that I care about this problem. And I want to make it better."
"In my nervousness for this speech and in my moments of doubt I've told myself firmly—if not me, who, if not now, when. If you have similar doubts when opportunities are presented to you I hope those words might be helpful."
– UN Women Goodwill Ambassador Emma Watson.
Before I had time to read more about it, I thought… "here comes another feminist debate that will just be cut short eventually".
But, found out that it was more of gender equality rather than being one sided.
There has always been sort of expectations out of your gender.
In my timeline, that was back when I was still in school.
When I delved into the working world, people were more open and weren't exactly (sure, there's still some to a certain degree) following the stigma of men and women having to act or behave in certain ways.
Sure, people try to blend in depending in different situations, but, outside of that "box" such as at work and whatnot… it's pretty different.
Times are changing.
I have seen more men crying in the open than women.
Those "definitions" are still floating around.
It's a matter of whether you'd care what others would say about it.
Maybe I'm one of those who'd do what I want, when I know what I want.
In a sense, that uncertainty scares people.
I've been there.
In a feminist point of view, I've been told that I couldn't get the job I wanted to take up because it was "meant" to be for men.
Gave excuses on why they wanted men instead of women for the job.
I was just starting out, a fresh graduate and someone who was still so naive and emotionally weak from the reality of the harsh world ahead of me.
The good thing is that, I turned out to be a fierce tiger, who can take charge and the people around me knows that all too well.
The "bad" thing is, in the view of other people, I'm not being "womanly".
I don't care though.
What is their view to mine?
If I had stuck to their view, I wouldn't have been happy with my own choices and the dreams I wanted to reach back then.
It would've turned out to be filling in the voids of other people's "choices" based on what the different genders should do.
In my humble point of view (based on my own experience) – apologies in advance, if that doesn't work well in your community…
There has always been freedom.
People just tend to keep themselves within the bubble of society's norm.
Then, blame others for their choices.
You did have a choice to do sports as a girl.
Why should you care whether you'd be deemed or seen as more of an athletic build rather than that of a model build who most people see as "perfect"?
Media fed that to you.
So did the brainwash from school, your own community, the circle of friends, etc.
Even with all those negativity around you, you still had the choice to break free from that and be different.
Be yourself.
Love yourself.
Do what you want.
Who's.. or even what's stopping you?
The ideals?
You want to know what's the reality of gender inequality?
The one that piss me off the most (based on my own experience), that I get paid less than those who work lesser than me in my industry.
But, well… I worked it out eventually.
It is NOT a myth that there is a glass ceiling for women.
It's fine though because it used to be like so.. but, as I've said before, I changed.
Fought for what I wanted.
The stigma of women's rights and standing is based on what they were brought up, and influenced as they grow older, to think.
It's as real as the air that you breathe.
One example: women tend to think that they don't deserve the credit and spread the credits to their colleagues even when they do most of the work.
Whilst men tend to take it all for themselves.
It came naturally to me as I grew up as well.
But, when I got angry at the known unjust issue in my own life situation, I did what the men did instead.
It was until much later than I realised that this was one of the examples that successful women state that most women don't even realise they're doing to undermine their abilities, skills, etc.
Yet, not everyone wants to do so even after explaining of the other side.
A small percentage might try to change.
Whilst others tend to be "an old dog" who prefers to stick to their regime, yet, keep whining about the same problem continuously.
Oh well~
LOL.. I didn't mean to talk crap… but.. just thought I'd say something about it.
❤ Icesabel


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Trauma can damage anyones creativity.
Recently, I've gone through a lot. Very traumatic events have taken place that resulted in mourning the lose of relationships of 10 plus years. Unfortunately, there was no way to avoid the lose if I wanted to regain my spiritually emotional freedom. The lost effected my husband and I so much that we had to move. The simplest way to explain it is that we discovered we were being spiritually abused. I won't go into great detail regarding that but I will say church abuse is real but we still love God's church and believe in the local church.
We aren't those people who have incredibly bad experiences with the church and God's people and turn our backs on everything related to Jesus. However; this experience has shown me that it's real and I can understand to a degree why people go through those type of seasons where they need the grace of God to meet them heavily.
Throughout this experience I have been noticing some elements of my life that have been oppressed by other people's opinions and thoughts of myself personally and creatively that distorted my actions. When I first began to blog in 2006 many of my blogs were about how I was processes what God was doing in my life and how I wanted to offer encouragement.
With all that is going on I am realizing that the value I add to this community is encouragement. I don't know what traumatic experiences you have gone through personally but I do know that trauma will completely alter the way you view your own creativity and ability to be produce.
One of of the major issues with the creative soul is that it is stained by the opinions and thoughts of others. Don't get me wrong! Creative critique is good and needed. It takes a humble person to allow others to give input on what they have created from there souls.
However; many of our creative entrepreneurial souls need to be restored from the opinions of others. We have stopped creating freely. There is an overload of blogs, books, and YouTube videos about the five steps to monetizing this and that. The concepts are very alluring and suck you in. But is that what you are called to. I believe in monetizing. I believe in marketing. I believing in developing yourself. What I have been a victim of is adding the value others want me to add to others and not the value I've been created to add to others.
Our creative soul are in need of healing. We have become victims of the Instagram, follow, like, scaleable systems of this world and that is not how our creative souls were designed to work. Johnathan Golden has a new book releasing in Be You. Do Good.: Having the Guts to Pursue What Makes You Come Alive in the books he speaks about purpose. When we think of the work we do we must shift our focus away from the money, exposure, and thoughts of others. He states, "All of our work is a means of making God's world on earth a bit more like heaven".
Our creative souls will experience healing the more we do work that matters. You can pray, hustle and repeat but is that the hustle your supposed to be grinding at.
How have trauma tainted or flourished your creativity?
-Reflections for a person suffering with post traumatic growth.


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Fashion is the science of appearance, something one should feel good about. One Has to be remembered rather than just noticed.Bridal Fashion is the best reliable online source of Latest Pakistani fashion bridal wears for all functions of weddings & others dresses & outfits for different occasions. We can mange your dreams within your budget.
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This year, small- and medium-sized businesses are at a strong clip because of Facebook. In fact, the social network now has a total of 15 million Pages run by small businesses globally, up from 13 million at the end of 2012.
MediaPost reports that in between the end of 2012 and March 2013, 2 million more companies established Brand Pages. Dan Levy, Facebook's director of small business, revealed the updated figure. He also said that local businesses are playing a key part in page growth, with active local pages up 40% in 2012.
You can read the full article here: http://www.mediapost.com/publications/article/195051/facebook-reaches-15-million-smb-pages.html


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Thanks to its Polygiene® system, the fabric remains clean for longer without releasing odors caused by bacteria, without harming the environment.
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- Polartec® Thermal Pro: Distinctive surface textures.
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View more related products to:Buff ® - Men´s clothing - Face warmers
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Ontario Fishing Forum > Ontario Fishing Forum > Ontario Sport Fishing Forum > Regional Moving with Packers and Movers Bangalore
|Moderated by: administrator
|Regional Moving with Packers and Movers Bangalore
| Rate Topic
|Posted: Fri Aug 26th, 2016 04:53 am
|1st Post
|Movers and packers gurgaon @ http://www.shiftingexpert.in/packers-and-movers-gurgaon.html
movers and packers Bangalore @ http://www.shiftingexpert.in/packers-and-movers-bangalore.html
Bangalore is the city of opportunity. Quoting that word for the town won't be a wrong abbreviation. Several lenders from about the country arrive at Bangalore to investigate their career better. There are lots of MNC's and IT organizations in the city that makes it the plastic area of India. There's number scarcity of career possibility in the town ergo folks from about the country concerns the capital of Karnataka. Bangalore being a major town presents resident for many individuals therefore regional shifting in Bangalore can be a very popular phenomenon. Persons have a tendency to change locally in the town according to their ease level.
Regional shifting is especially seen in the city where you will find large numbers of migrants. And this notion is well applicable to the town of Bangalore as well. There are numerous students and small experts who come here to examine their career better and because of this they change within the town as well. Coming to regional moving then there's a misconception related to this shifting. Lots of people think that regional shifting is a simple process and there's no difficulty level in it. But this is simply not the actuality. In actual terms regional shifting is as much complicated as any moving and shifting because all of the processes which are involved in these type of moving are same as others.
Moving is a frantic issue to do. There are so many complex measures that can be followed in order to allow it to be a successful task. To begin with arranging of all things that should be moved is a complex thing to do. And it is really because this will allow an individual as what to maneuver alongwith and what not. After it comes the loading process. It is reported to be the hardest amongst all. Perhaps the shift will undoubtedly be effective or not it is likely to be determined by the packing. Packing and unloading would be the techniques under transportation and by the end there comes arranging of goods. If one wants to get rid from these complicated steps and desire to end moving regional from Bangalore at a positive note then employ packers and movers in Bangalore for the exact same and appreciate the local relocate Bangalore.
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|Current time is 09:05 pm
|Ontario Fishing Forum > Ontario Fishing Forum > Ontario Sport Fishing Forum > Regional Moving with Packers and Movers Bangalore


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"I've been on this planet a while and never been this contented," noted Conan O'Brien while playing Hohokum during a 2013 visit to E3. "I spent six weeks with the Dalai Lama and I didn't feel this good."
The game is simple: you go around and collect seeds which turn into kites which you and your friends fly around on, and during the Conan segment, O'Brien's colorful and perhaps too-blissful gaming experience is soundtracked by the fluid, sunshine-through-clouds instrumental thump of "L" by Tycho. There are no doubt those that saw the bit and were intrigued by what they heard, and although Tycho takes up a few slots on Hohokum's official soundtrack, his songs so easily outshine and outrank the compilations other artists that it almost doesn't seem fair.
Not that Scott Hansen is an asshole, mind you. He's not a showboater: he's a San Francisco graphic artist who just so happens to also have a successful music career. What separates and differentiates him from his contemporaries is his ability to structure his Boards of Canada-indebted ambient-dance numbers in a way that is relatable and immediate, keeping his actual chord progressions astoundingly simple but embellishing each production with enough atmosphere and tiny details that each track feels like a personal epic, a custom-designed soundtrack scoring you and your life. Fans have responded in kind, turning out in droves to his live shows and going so far as to put his heralded 2014 effort Awake in the Top 25 of the Billboard 200—an astounding feat for a post-rock instrumental set.
What a surprise it must be for some fans than to hear a new song called "Epoch" without warning and then find out that Tycho's latest full-length (of the same name) was surprise-dropped less than two weeks later. Some call surprise album-releases gimmicky, while others view it as "appointment listening". It's a stunt no matter how you look at it, but in truth, the sudden release of Epoch is the least interesting thing about it. For a man who so commandingly has cornered his own space in the post-spectrum, Epoch pushes Hansen's sound forward in a way that few could have anticipated: it's more aggressive and yet more pop-friendly at the same time, showing off new angles of his sonic while clearly retaining that relatable open-air accessibility that has enraptured tens of thousands of fans.
As the slowly-warming tones of Epoch's opener "Glider" wrap the listener in a sense of immediate familiarity, a skittery, fast-paced drum beat soon enters, and it's clear that for all the goodwill Hansen has developed since his debut album dropped a decade prior, he's starting to set his sights beyond the restraints of his genre, recognizing that even in the indie-instrumental electronica circuit, there are still valleys that haven't been explored, and he's more than willing to spend some of his capital to arrive at his new destination with his fans in tow.
After all, once "Horizon" kicks on, with its '80s synthpop opening lick, it's clear that what Hansen is going for with Epoch is something that he wouldn't have been able to accomplish with his previous albums: there's a little bit more propulsion to the proceedings, his basslines firing more rapidly and his usually-warm synths now dipped in a cool silicone bath, forcing Hansen to trade in the pulsing heart that's driven Tycho's best work for a sleekness that Awake only started to hint at, and even with that in mind, he still fights hard to achieve that deep emotional resonance that his songs are capable of conjuring, even in the midst of all the new challenges Hansen has given himself.
While tracks like "Slack" and the sparse guitar strummer "Source" could easily have been placed on Tycho's previous two full-lengths, "Receiver" is the kind of moody, Auntumn-hued number that fans have been pining for since 2011's Dive: a slower, more deliberate kind of song that shows restraint, using its minimalist tones to lull the listener into a state of ease before eventually soundtracking an emotional scene for a yet-to-be-made Jason Reitman melodrama.
To casual fans, the changes between Awake and Epoch may hardly be detectable, but the confidence to which Hansen and his band attack the new compositions recalls the change in tone that his contemporary Mark McGuire ran into between his 2014 steel-grey sky guitar instrumental set Along the Way and the far more rocking 2015 set Beyond Belief, wherein McGuire realized that even within the confines of his genre, there was room for great textural expansion, turning up the fuzz guitar but still remaining relaxing and relatable in new and exciting ways.
Similarly for Tycho, a sparse xx-styled guitar closer like "Field" isn't necessarily out of left field for Hansen and co., but as a capper for his shortest, sweetest, and most concise album to date, it feels lived-in and considered, inspiring his fans while making everyone else wonder where this newfound confidence is going to propel Tycho next.
Bruce Sudano "When Cinderella Dies" (video) (premiere) // Music
Gunner & Smith 'Byzantium' (album stream) (premiere) // Music
Andrew Bryant "Robert Downey Jr.'s Scars" (video) (premiere) // Music
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Bette Smith "I Found Love" (audio) (premiere) // Music
Treasuring Memories of Paul McCartney on 'One on One' Tour
// Notes from the Road
"McCartney welcomed Bruce Springsteen and Steven Van Zandt out for a song at Madison Square Garden.READ the article


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Dagestan State Technical University
Dallaeva D.S.,Brno University of Technology |
Bilalov B.A.,Dagestan State Technical University |
Gitikchiev M.A.,Dagestan State Technical University |
Kardashova G.D.,Dagestan State Technical University |
And 4 more authors.
Thin Solid Films | Year: 2012
A process for ion-plasma formation of aluminum nitride (AlN) nanolayers on nitrided sapphire (Al2O3) substrates is presented. The method is based on the direct current magnetron sputtering of a high-purity aluminum target in the presence of an argon-nitrogen gas mix and high frequency-activated nitrogen plasma. The method, combined with ion etching, produced matched layers by nitration of Al2O3 in the (0001) plane, and formation of high quality AlN epilayers on this surface was observed. The processing characteristics and morphology dependence on synthesis parameters were studied using atomic force microscopy. © 2012 Elsevier B.V.
Altuhov V.I.,North-Caucasian Federal University |
Kasyanenko I.S.,North-Caucasian Federal University |
Sankin A.V.,North-Caucasian Federal University |
Bilalov B.A.,Dagestan State Technical University |
Sigov A.S.,Moscow State Technical University
Semiconductors | Year: 2016
A simple but nonlinear model of the defect density at a metal–semiconductor interface, when a Schottky barrier is formed by surface defects states localized at the interface, is developed. It is shown that taking the nonlinear dependence of the Fermi level on the defect density into account leads to a Schottky barrier increase by 15–25%. The calculated barrier heights are used to analyze the current–voltage characteristics of n-M/p-(SiC)1–x(AlN)x structures. The results of calculations are compared to experimental data. © 2016, Pleiades Publishing, Ltd.
Guseinov R.V.,Dagestan State Technical University |
Rustamova M.R.,Dagestan State Technical University
Russian Engineering Research | Year: 2013
Helical bits are generally used for machining holes. Means of improving their design are developed. Recommendations are made for more rapid machining of small-diameter holes. © 2013 Allerton Press, Inc.
Mitarov R.G.,Dagestan State Technical University |
Kallaev S.N.,Russian Academy of Sciences |
Omarov Z.M.,Russian Academy of Sciences |
Abdulvakhidov K.G.,Southern Federal University
Physics of the Solid State | Year: 2015
The temperature dependence of the heat capacity of the PbFe0.5Nb0.5O3 multiferroic has been studied in the temperature range of 130–780 K. An excess heat capacity has been observed in the temperature range of 250–780 K. It has been shown that the temperature dependence of the excess heat capacity is associated with the manifestation of the Schottky effect in three-level systems. © 2015, Pleiades Publishing, Ltd.
Verdiyev M.G.,Dagestan State Technical University |
Nabiev Sh.Sh.,Dagestan State Technical University |
Kamnev N.A.,Dagestan State Technical University
Journal of Physics: Conference Series | Year: 2012
The study is devoted to the results of experimental searches of the disperse dye flows of inkjet drawn on paper in an electric field that can be used to make improvements to technology of improving the quality and the print admissible capacity of various printed materials in inkjet technology.
Ramazanov S.M.,Dagestan State Technical University |
Kurbanov M.K.,Dagestan State Technical University |
Safaraliev G.K.,Dagestan State Technical University |
Bilalov B.A.,Dagestan State Technical University |
And 2 more authors.
Technical Physics Letters | Year: 2014
Epitaxial layers of the (SiC)1 - x(AlN)x solid solution with x ≈ 0.12 and x = 0.64 without macroscopic structural distortions were grown using a new technique. It was established that the compositional dependences of crystal lattice parameters of the epitaxial films obey the Vegard's law with an error of ~0.03. This confirms that there is formation of isomorphic substitutional solid solutions in the SiC-AlN system. © 2014 Pleiades Publishing, Ltd.
Kallaev S.N.,Russian Academy of Sciences |
Mitarov R.G.,Dagestan State Technical University |
Omarov Z.M.,Russian Academy of Sciences |
Gadzhiev G.G.,Russian Academy of Sciences |
Reznichenko L.A.,Southern Federal University
Journal of Experimental and Theoretical Physics | Year: 2014
The heat capacity of Bi1 - x Re x FeO3 (Re = La, Eu, Ho; x = 0, x = 0.05) multiferroics has been studied in the temperature range of 120-800 K. The substitution of a small amount of rare-earth elements for bismuth leads to a significant increase in the heat capacity in the broad temperature range studied. It is established that the temperature dependence of the excess heat capacity is related to the Schottky effect for three-level states certain that appear as a result of structure distortions in the rare-earth-doped compositions. © 2014 Pleiades Publishing, Inc.
Sanaev N.K.,Dagestan State Technical University
Russian Engineering Research | Year: 2014
The performance of bushes in the 4Ch8.5/11 diesel engine is optimized on the basis of mathematical experiment design. Their wear resistance may be improved by finishing on the basis of special roller burnishing, rather than honing. © 2014, Allerton Press, Inc.
Gasanov V.M.,Dagestan State Technical University
Journal of Engineering Physics and Thermophysics | Year: 2011
The surface diameters of gas bubbles at the liquid-gas interface whose burst leads to the formation of aerosol from a thin surface microlayer of thickness 1 μm and less have been determined experimentally. Precisely the anomalous concentrations of such a microlayer are responsible for the fractionation of substances in the process of ocean-atmosphere exchange. ©2011 Springer Science+Business Media, Inc.
Kallaev S.N.,Russian Academy of Sciences |
Omarov Z.M.,Russian Academy of Sciences |
Mitarov R.G.,Dagestan State Technical University |
Bilalov A.R.,Russian Academy of Sciences |
And 2 more authors.
Journal of Experimental and Theoretical Physics | Year: 2010
The heat capacity, thermal expansion coefficient, and deformation of the PLZT-9/65/35 compound are studied in the temperature range 150-800 K. Diffuse anomalies are detected in the temperature dependences of the heat capacity and thermal expansion coefficient over wide temperature ranges of 250-650 and 330-550 K, respectively. The anomalous behavior of the heat capacity in the temperature range 250-650 K is shown to be caused by the appearance of two-level states (Schottky anomaly). The results obtained are discussed along with the data of structural and dielectric studies. © 2010 Pleiades Publishing, Ltd.


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The house is lovely, on beautiful, flower filled grounds which were all set up for our croquet game. Hamish was the perfect host - and made 12 poached eggs (freshly laid by their friendly chickens) at once. Will definitely come again!
Getting you more information on this room
Getting you more information on this room
"We had the double room at the top of the house, which was very quiet."


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The most complex academic work associated with research and writing and of course one of the most time consuming is a dissertation. It is submitted when a candidate is obtaining a degree or professional qualification, so it is a final step of graduation. It usually consists of several chapters and contains a huge volume of information and research results. Writing a dissertation is a stressful process and requires deep topic understanding, attention, and well-developed writing skills. That is why the development of different habits and staging your work on it are important. In this article, we decided to share main dissertation writing approaches which will help you to succeed.
10 main dissertation writing approaches
1. Academic and language skills building
It is important to have academic and language skills developed before you have started writing. Writing a dissertation usually requires not only knowledge of English grammar and spelling, but also other languages, like modern French, German, Italian, or dead one (Latin, Greek) or even programming languages, if your study is related to IT. Moreover, the volume of such paper is usually more than 40,000 of words and you should be able to cope with it. The dissertation is based on the research and findings, that is why you should be ready to analyze and present everything you are planning to include.
So you need to develop your research and writing skills, before you plan to write a dissertation.
2. Develop time management skills
Writing a dissertation is very time-consuming and stressful. Moreover, you will probably work or study during writing, that is why you will be feeling a lack of time for sleeping and/or having a rest. So it is very important to streamline your activities and set up schedules for writing, working learning, sleeping and other things you need to do during the day. If you have not done this before, try to develop your time management skills using different methods and approaches.
3. Understand the aim of dissertation
Before you began the process you need fully understand why it is needed and what will happen after you have written your dissertation. The main goal of dissertation writing is to demonstrate that you have achieved a goal you have set for your research. In the dissertation, you are conducting an original research to showcase that you are able to contribute in your discipline in future. Moreover, you show that you are able to design a research, do all needed tests, collect data and document your findings.
4. Decision making and topic choice
The dissertation is a serious academic work which will have a huge impact on your future. Not everyone can cope with it that is why you need to think twice before the start. In case you think that you need a dissertation and able to write it, you need to choose a right topic. The topic must be not only original and contribute to scholarship, but also the one you feel you are able to research and write about.
5. Audience understanding
Before you will start writing your dissertation, you should learn who will be reading and reviewing it. The first person you need to pay attention to is your advisor who will help you while writing. The others are committee, your colleagues, other students, and people who are interested in your area of study. Your writing should be understandable for all these groups of people and possess not only excellent language and writing skills but also demonstrate your understanding of the topic and your impact in the area.
6. Structure planning
The structure of the dissertation should correspond with the standards in your area. You will definitely need to write an outline of your paper before beginning, so you will know what information you need to collect and how it is better to organize it. Of course, it will have a basic structure, like in research paper, but giving more information about the background of the research, methods used for the research and evidence. Before you start writing, show your outline and plans to peers and advisers, to make sure that you have not missed something.
7. Research and data collection
When conducting a research you need to use only trusted sources and latest scientific findings. They are available in libraries and different online resources. Some of them you will need to purchase. However you may buy a digital version, it is usually cheaper than paper. The papers, books or other sources you are planning to cite always add to your cited works list not to lose this information or forget about them. This also will save you a lot of time at the end of writing.
8. Writing style and unnecessary information removal
Follow writing style which is applied in your area, use specific terms and constructions. Avoid slang, colloquial and informal language usage; follow an academic style of your area. Watch correct grammar and spelling, if needed think of hiring a professional editor after you have completed your paper. While writing the paper try to think of what information, researches, and results are not necessary for your paper, what you should remove to make your text stronger. Of course, don't forget about citing to present others findings and avoid plagiarism.
9. Editing and last steps
After you have completed writing you need to do proofreading and correct grammatical, spelling and style mistakes. Try to do read-through several times, before reading through, try to think of the list of words you can technically misspell or misuse. Make sure that you have used terms correctly. Try to ask your colleague or friend to proofread your paper and give you a review. If you are not confident about your writing, order editing service from the professional.
10. Presentation practice
Usually, the presentation of your paper before the scientific committee is required. Most people don't have much experience in speaking and presenting their works in front of public that is why practicing presentation is important.
To help yourself create a plan of your presentation, work on graphical materials, and your speech and of course think about questions committee may ask you. Ask your friends or family to listen to you and tell what to change or add to make the presentation better. This will help you to get prepared for the presentation.
These simple things will help you to make your dissertation writing more effective and go through this complex time-consuming process with no issues.


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At Silent Gliss�we try to accommodate your exact requirements whenever they are technically possible.
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Senior music majors Angelica Osorio and Michael Choy presented their senior projects May 18 in Morgan Auditorium in front of family, friends and music enthusiasts alike.
Archive | Arts, Etc.
Peel sheds layers through 'Catharsis'
Some students spend their free time on the Internet, others spend it out with their friends. But in this case, sophomore psychology major Matthew Peel spends his free time writing and publishing poetry and prose.
Students sit in the director's chair
Students of the Theatre Arts 351 class premiered their directing capabilities in "Theatre On-Stage: New Directions in Directing" May 17-25 in the Jane Dibbell Cabaret and Dailey Theatre.
Campus and community arts events for the week of May 24, 2013.
New spin on after school activities
During a warm-up session, Soetan Osifeso, 19, tries a few tricks while Jose Carerra, 18, and Soyemi Osifeso, 15, watch and wait for their turn to show off.
Barbershop performances bring harmony
Exceptional harmonies, matching attire and talented vocalists entertained an estimated audience of 70 as Morgan Auditorium hosted the Barbershop Harmony Concert on May 9.
Huzzah for fun at the Renaissance Faire
Terry Clark engages in some lively conversation with two fair-goers in one of the various vendors' tents at the Southern California Renaissance Pleasure Faire on Saturday.
Final choral concert delights
The ULV Chamber Singers and Gospel Ensemble performed their last concert of the semester Tuesday in Morgan Auditorium.
Book Review: 'S-Word' gives the whole story
Imagine waking up, getting ready for class, leaving your home and seeing the letters S-L-U-T written all over your car. In Chelsea Pitcher's story in "The S-Word," this is a reality for a 17-year-old girl.
Tale of love destined for the Fringe
Students in the theater department are taking a new twist on the traditional senior project by breaking free from the educational realm and entering the world of professional theater as they enter the 2013 Hollywood Fringe Festival.


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Gear Bag, 9 in Long x 8 in Wide x 6 in Deep, Zipper Closure, BlackProduct Description: Gear Bag, 9 in Long x 8 in Wide x 6 in Deep, Zipper Closure, BlackPit Stop USA offers Battery Tender Zipper Pouch - Large 10in x 12in 500-0140 at low everyday prices. Be sure to check out all our products from Deltran Battery Tender including Battery Tender Battery Charger Components
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Golden Triangle Tour is providing you with a thrilling experience that would make you travel the three awesome cities of India- the sophisticated city of Delhi and the culturally rich city of Agra and Jaipur.
Experience the awesome traditions, cultures and experience the perfect combination of ancient and modernized India. Visit the great monuments and structures representing the architectural beauty of India which has been the main tourist attraction for people from different parts of the world. To enhance your experience we are providing you our best staff whose first priority is your comfort and safety.
Arriving at the international airport you will be greeted and accompanied to a hotel by our representatives in an air conditioned and private vehicle. They will help you in checking in and then a private guide will be provided to you whose priority will be to escort you all through your sightseeing tour in the capital city- Delhi. You will be taken to the famous Red Fort which looks like a red jewel and Jama Masjid which is one of the largest mosques in the world and certainly the largest in India. You will also be driven past some of the eminent culturally important landmarks like India Gate and Raj Ghat. You will also be taken to explore the World Heritage Site declared by UNESCO- the Qutub Minar which is an exquisite combination of red sandstone and marble and then will be taken to the Humayun's Tomb. In case you feel like shopping you would be taken to a great mall to cram yourself with stuff of your choice. You will then be taken to the hotel for your overnight stay.
After having a splendid breakfast you will be escorted to Agra in a private vehicle. It would be a 3 hour drive and you will reach there by lunch. After checking in at a hotel, your private guide will take you to an enthralling experience through the maze of Agra Fort. Agra Fort is the melding of Hindu and Islamic architecture and was built by Emperor Akbar on the banks of mighty Yamuna. You will then be escorted to one of the Seven Wonders of the World, the World Heritage Site declared by UNESCO- the Taj Mahal.
You will reach there by evening and as the sun set you will be stunned and perplexed to see its unreal beauty. The rays of the setting sun provide it with an ethereal beauty and its reflection is the most beautiful thing you have ever set eyes on. You will be enthralled with the historic aspect related to the construction of Taj Mahal and can further admire its beauty from the opposite bank of Yamuna River. The tour of city Agra will be concluded with your visit to Itmad-Ud-Daula. Then you will be taken to the hotel for the overnight stay.
The journey to the pink city or Jaipur will be started early morning with a brief stop at the former Mughal capital Fatehpur Sikri. Akbar was a great Mughal emperor who created the amazing architectural structure of sandstone, the city of Fatehpur Sikri which is a must watch. You will also encounter Buland Darwaza in Fatehpur which bags the tittle of the largest gateway and is also referred to as the "Gate of Magnificence". You will be taken to encounter the beautiful Jodha Bai Palace, Birbal Bhawan, Panch Mahal etc. After reaching Jaipur you will be taken to explore the pink city after checking in at a hotel. The City Palace which houses amazing courtyards, luscious gardens and exquisite artefacts will enhance your experience of sightseeing. You will also be taken to see the world Heritage Site- Jantar Mantar and the grandiose of Nana Sahab ki Haweli. Then we will go to the hotel for overnight stay.
After having a fulfilling breakfast in the morning we will start with a 5 hour road trip to Jaipur. We will check in at a hotel on arrival. After having lunch and brief amount of rest there we will walk you through the royal buildings of Chandra Mahal and Mubarak Mahal. Chandra Mahal has a museum showcasing dazzling royal costumes and weapons of Rajputs. You can stroll through the markets and spend your evening relaxed and freely. The driver will escort you back to the hotel at your call.
After having a delectable breakfast you will be taken to visit Lakshmi Narayan Temple or Birla Temple. It has a beautiful array of religious figures on the outer façade of the temple. After lunch you will be taken for a 6 hour drive back to Delhi.
You are also provided an amazing opportunity to visit the rich traditional pilgrimage site of Pushkar home to numerable temples. Among these temples the most significant one is Brahma temple the temple dedicted to Lord Brahma. Pushkar fair is a world renowned ceremony and the festivals of Pushkar are unique to this place and are a worldwide attraction.
The holy lake here is considered to be divine and religious tourists or Sadhus believe that taking a dip in this lake two days around full moon will free them of all their sins. The colourful and established carnival of Pushkar is the platform for dancers, magicians, musicians, snake charmers, acrobats etc. You will gorge yourself up with the mouth-watering Rajasthani cuisine.