Gavotte 1
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Bach Gavotte 1 BWV 1012 guitar solo
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The gavotte is a French dance, taking its name from a folk dance of the Gavot, the people of the Pays de Gap region of Dauphiné. Bwv Gavotte 1 And 2 by Johann Sebastian Bach tab. One accurate version. No abusive ads. Recommended by The Wall Street Journal.
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Rate this product's difficulty level: Review Guidelines Explain exactly why you liked or disliked the product. Do you like the artist? Is the transcription accurate? After the first Wolbachia introduction, the infection level in the cell line was about 10 5 times lower than in the DSR adult female. Following five sequential Wolbachia introductions, the resulting Wolbachia infection level was increased 5. To generate the uninfected S2 U line, the S2 I cell line was split and treated with tetracycline Fig.
For transcriptomic comparison, the S2 U cell line was used instead of the naive S2 line due to concerns that the genetic background of the S2 cell line might have been altered during the repeated transfection procedure. To avoid the potential complication caused by transcriptional differences induced by the tetracycline treatment, the microarray analysis was not conducted until six passages after antibiotic treatment.
Diagram illustrating the in vitro infection strategy, tetracycline clearing and microarray analysis of infected S2 I and uninfected S2 U cell lines.
The line illustrates an estimated infection level based upon periodic qPCR assays indicated by hollow circles. The number provides a relative measure, not the absolute number of Wolbachia. The "waves" in the left half of the graph resulted from removal of the infection by host and enrichment of infection by shell vial introduction.
Arrows indicate shell vial introduction of Wolbachia infection. At passage 17 marked with asterisk , the cell line was split and one subline was tetracycline treated. At passage 23 grey shading the cell line was examined via microarray. The Genechip contains 13, probe sets, of which 7, were found to be uninformative signal was absent in all eight replicates. The frequency of uninformative probes is not unexpected considering the use of cells derived from embryos Genechip includes development stage and sex-specific probe sets and results of prior studies use of the same platform results in probe sets with signals below the detectable threshold; [ 25 ].
A complete set of microarray data results is available at Gene Expression Omnibus [ 26 ]. Distribution of the relative expression of 6, genes in the infected S2 I versus uninfected S2 U cell lines. Of the differentially transcribed probe sets, fifty are categorized as genes with unknown functions and are not discussed subsequently.
Orchestral Suite No 3 in D major, BWV 1068 (Gavotte 1 and 2)
The remaining probe sets are identified as belonging to a particular biological process ontology, with 73 down-regulated and up-regulated. After all the differentially expressed genes were enriched into GO terms, the GO terms were ranked based on Z value. A complete set of GO data is available in the support information Additional File 1 and 2. The top five non-redundant Gene Ontology GO terms in biological process which contain the most differentially expressed genes. Up or down, up- or down-regulation after infection, respectively; T, number of genes assigned to this the GO term; M, the subset of T genes that are represented on the microarray; C, the subset of M genes observed to vary in the analysis; Z, significance score.
Therefore, Ance was selected for in vivo expression characterization. In contrast, Ance expression was 1. Wolbachia infection is consistently associated with higher Ance expression in ovaries and lower Ance expression in testes. To examine for an interaction of Ance and Wolbachia -induced CI, crosses were conducted between wild type and Ance -mutant D.
Among these, the majority function in the different process of oogenesis, including ovarian follicle cell development, pole plasm assembly and nurse cell to oocyte cell transport. Up-regulation was not observed in this category Fig. Among these, two important regulatory proteins: In contrast, ird5 and peroxiredoxin were down-regulated. Regulation pattern of heat shock protein Hsp in Wolbachia infected S2 cells. All heat shock proteins in Drosophila are shown.
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The fold change is shown for each gene. Induction of immune response to Wolbachia infection in S2 cells.
Activation of both Toll and Imd pathways result in up-regulation of Dorsal and Relish, which might lead to a specific antimicrobial expression profile against Wolbachia. Negative cross talk between Imd and JNK pathway is also observed [48].
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Classification of differentially expressed genes related to sexual reproduction based on the Gene Ontology GO term. As a simplified model system, the results are likely to under-represent in vivo events Type II error , for reasons including a relatively low Wolbachia infection level and the reduced complexity of the embryonic cell environment in the S2 cell line. However, the use of a simplified system is not an inappropriate approach given the dearth of information currently available for the Wolbachia -host interaction.
An important motivation for subsequent in vivo transcriptional assays was to address the concern that the microarray results may represent an experimental artifact. Specifically, in vivo characterization of Ance expression via qRT-PCR was used to examine the validity of the in vitro system as a predictor of Ance in vivo expression.
Specifically, Wolbachia has been observed to be associated with differentiating sperm [ 27 ], providing a potential role of sperm modification in CI [ 28 , 29 ]. The location of Ance mutation has been mapped to 34E3—5 on the second chromosome [ 24 ]. The recessive lethality associate with the mutation due to the loss of function could be rescued by expression of Ance in the P-transformation assay and genetic complementation test [ 24 ].
Homozygote Ance males are sterile. Their testes lack individualized sperm and have very few actin-based individualization complexes [ 23 , 50 ]. Therefore, premised upon the described involvement of Ance in spermatid differentiation, additional experiments were conducted to examine a model in which Wolbachia modifies the sperm via a pathway involving Ance expression. The latter experiments included characterization of Ance mRNA levels in testes and ovaries and examining egg hatch resulting from CI crosses of Ance flies.
The latter observation is consistent with the higher Ance expression that is observed in the Wolbachia infected S2 cell line, which was derived from embryos. Future experiments should address whether the observed differential Ance expression originates within oocytes, maternally-derived ovary tissues, or both.
Wolbachia infection was also associated with differential Ance expression in testes, but the variation was the reverse of that observed in ovaries. Lower Ance expression was observed in Wolbachia -infected testes relative to testes from uninfected flies. A similar pattern is observed in D. It is useful to note that the results with uninfected D. However, the Wolbachia infection status in the prior reports is not known. However, comparison of the hatch rate resulting from the four incompatible cross types revealed a significantly higher egg hatch rate in crosses of uninfected Ance females and infected wild type males relative to the other incompatible crosses.
It is important to note that only heterozygous Ance flies could be used in this experiment. Therefore, only dominant Ance effects would be observed. Furthermore, the possibility cannot be excluded that additional other sub-lethal mutations are co-segregating with the Ance mutation. A model to describe the observed cross results would require an interaction of Ance with both the rescue and modification mechanisms [ 33 ].
However, observations of Ance expression are not inconsistent with the model proposed above. Specifically, Ance expression differs between Wolbachia -infected and uninfected Drosophila and that the observed variation in expression occurs in opposite directions in female or male flies. We note that a broad range of egg hatch rates have been previously reported in Drosophila crosses examining CI, and the egg hatch reported here is at the lower end of the reported range. Egg hatch can be affected by the experimental conditions under which females are held, and the hatch rates observed here are similar to a prior study in which females were made to oviposit in isolation [ 34 ].
In addition to Ance , the microarray assay identifies multiple genes that vary in expression level between infected and uninfected cells. Wolbachia has been reported to affect host oogenesis [ 35 ]. The latter is not unexpected due to the critical role of oogenesis in the maternal inheritance of Wolbachia and the somatic stem cell tropism of Wolbachia in Drosophila [ 36 ]. Over-expression of l 2 gl and zipper have been previously shown to mimic CI [ 6 ], leading a hypothesis that l 2 gl and zipper play a role in the CI mechanism.
Consistent with this hypothesis, the results presented here show an up-regulation of l 2 gl in the infected cell. Additional variation was observed in sex lethal Sxl , which acts in Drosophila sex determination. Previously, Wolbachia has been shown to suppress sterility in D. The results of the prior studies suggested that " Wolbachia does not bypass or reduce the requirement for Sxl in the germline in a general way, nor increase overall germline Sxl expression" [ 37 ]. Here, microarray data suggests that Wolbachia down-regulates Sxl expression.
Multiple transcripts encoding proteins involved in heat-shock were observed to be down regulated in Wolbachia -infected S2 cells. A common function of heat shock proteins Hsp is as molecular chaperones that act to reduce inappropriate inter-protein interaction [ 39 ]. Prior studies have shown an ability of heat-shock to abate CI, and distinct Hsp isoforms have been reported in comparisons of infected and uninfected Drosophila [ 40 ]. Consistent with the prior study, our results indicate a down-regulation of Hsp in Wolbachia -infected cells. Thus, heat-shock may act to negate the impact of reduced Hsp expression in Wolbachia -infected flies.
Wolbachia infection in S2 cells is associated with the induction of antibacterial peptides. Prior studies observed that Wolbachia did not alter the expression of defensin , diptericin and cecropin in D. Consistent with the prior results, the microarray assays reported here did not detect changes in the expression of these three antimicrobial peptides.
In contrast, Wolbachia -infected cells show higher expression with multiple genes involved in the Toll and Imd immune signaling pathways Fig. Drosophila Rel is critical in Imd signaling pathways and is transcriptionally up-regulated in response to gram negative bacteria challenge [ 43 ]. An activation of the Toll and Imd immune pathways by gram-negative Wolbachia bacteria could be related to structurally characterized peptidoglycans in Wolbachia [ 44 ] that bind to the extracellular peptidoglycan recognition protein PRGP and stimulate the Drosophila innate immune signaling pathway [ 45 ].
An inhibition of additional expression products involved in immunity is also observed in the microarray assay. The expression of ird5 , the Drosophila homolog of IKK , is down-regulated. Prior studies show that the activation of Relish requires ird5 [ 46 ] and that Escherichia coli survive times better in ird5 mutant lines relative to wild-type Drosophila [ 47 ]. Prior studies show negative cross talk between the JNK and Imd signaling pathways [ 48 ].
Peroxiredoxin expression was also lower in infected cells. Inhibition of superoxide production has been demonstrated to be important to survival of Anaplasma phagocytophila , which is another intracellular bacterium and close relative to Wolbachia [ 11 ].
Together, the latter suggest an ability of Wolbachia to evade the host immune responses. The disappearance of Wolbachia from the cell is consistent with microarray data showing activation of the host immune response. In previous studies, when Wolbachia was transferred by microinjection from D. Future experiments should examine the role of immune response in the establishment and maintenance of Wolbachia infection. For example, RNAi technology may be used to interrupt specific immune pathways in S2 cells [ 19 ].
The latter cells may then be examined for an ability to sustain Wolbachia infection. Differences observed in the S2 cell culture system are consistent with prior studies examining the Wolbachia interaction within insect hosts. The utility of the in vitro system is further supported by results of the in vivo characterization of Ance expression and CI phenotype assays. The microarray results also provide rationale for examining additional gene products that have not yet been assigned a function, but that are shown to vary between infected and uninfected cells.
Due to maternal inheritance, progeny from the above cross are Wolbachia infected. Subsequently, the infected Ance mutant line is maintained over the CyO balancer background by selecting for flies with cinnabar eye color and curly wing mutations. For the CI assay, one-day-old males were mated with three-day-old females for 12 hours in an apple juice plate container.
Females were then isolated individually on a yeast coated apple juice plate to collect eggs. CI was determined as the proportion of hatching eggs, as previous reported [ 51 ]. Cells were maintained as described previously [ 52 , 53 ]. Wolbachia infection in the S2 cell line was established using a previously described shell vial technique [ 53 ].
In brief, DSR eggs were collected by standard procedures [ 54 ], homogenized and introduced into S2 cells. To increase the infection level, the shell vial technique was repeated five times sequentially at an interval of every third passage Fig. The resulting Wolbachia -infected S2 cell line is subsequently referred to as S2 I. Eight Affymetrix Drosophila genome chips 1. Image data was quantified using the genechip analysis microarray suite 5. If all eight replicates for a particular probe set were assigned an "absent" value, the probe set was removed from further consideration.
The transcript level of the remaining 6, probe sets were normalized divided by the corresponding chip median and log transformed [ 55 ]. A Bonferroni significance level was used as an initial criterion for rejecting the null hypothesis of a significant treatment effect 0. The change was calculated as the average of four replicates.
The following criteria were used to define the differential expression caused by treatment: To visualize the differential expression pattern, before Z value ranking, all gene ontology terms were further filtered manually with the criteria: If the annotation for interested genes were missed in the gene database of GenMapp, they were examined in Affymetrix NetAffya analysis center [ 61 ].
The protocol was similar to prior qPCR amplification using the single-copy wsp and su fk C genes of bacterial and host origin, respectively [ 65 ]. S2 cells were quantified using a hemocytometer to obtain 10 6 cells. Primers were designed based upon D.