Spermatid Cyst Polarization in Drosophila Depends upon apkc and the CPEB Family Translational Regulator orb2
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{"title"=>"Spermatid Cyst Polarization in Drosophila Depends upon apkc and the CPEB Family Translational Regulator orb2", "type"=>"journal", "authors"=>[{"first_name"=>"Shuwa", "last_name"=>"Xu", "scopus_author_id"=>"37070112900"}, {"first_name"=>"Sanjay", "last_name"=>"Tyagi", "scopus_author_id"=>"56748561600"}, {"first_name"=>"Paul", "last_name"=>"Schedl", "scopus_author_id"=>"7005983204"}], "year"=>2014, "source"=>"PLoS Genetics", "identifiers"=>{"pmid"=>"24830287", "sgr"=>"84901642366", "scopus"=>"2-s2.0-84901642366", "doi"=>"10.1371/journal.pgen.1004380", "pui"=>"373197057", "issn"=>"15537404"}, "id"=>"2695d884-e6c5-39a0-8a90-32c3fe269706", "abstract"=>"Mature Drosophila sperm are highly polarized cells--on one side is a nearly 2 mm long flagellar tail that comprises most of the cell, while on the other is the sperm head, which carries the gamete's genetic information. The polarization of the sperm cells commences after meiosis is complete and the 64-cell spermatid cyst begins the process of differentiation. The spermatid nuclei cluster to one side of the cyst, while the flagellar axonemes grows from the other. The elongating spermatid bundles are also polarized with respect to the main axis of the testis; the sperm heads are always oriented basally, while the growing tails extend apically. This orientation within the testes is important for transferring the mature sperm into the seminal vesicles. We show here that orienting cyst polarization with respect to the main axis of the testis depends upon atypical Protein Kinase C (aPKC), a factor implicated in polarity decisions in many different biological contexts. When apkc activity is compromised in the male germline, the direction of cyst polarization within this organ is randomized. Significantly, the mechanisms used to spatially restrict apkc activity to the apical side of the spermatid cyst are different from the canonical cross-regulatory interactions between this kinase and other cell polarity proteins that normally orchestrate polarization. We show that the asymmetric accumulation of aPKC protein in the cyst depends on an mRNA localization pathway that is regulated by the Drosophila CPEB protein Orb2. orb2 is required to properly localize and activate the translation of apkc mRNAs in polarizing spermatid cysts. We also show that orb2 functions not only in orienting cyst polarization with respect to the apical-basal axis of the testis, but also in the process of polarization itself. One of the orb2 targets in this process is its own mRNA. Moreover, the proper execution of this orb2 autoregulatory pathway depends upon apkc.", "link"=>"http://www.mendeley.com/research/spermatid-cyst-polarization-drosophila-depends-upon-apkc-cpeb-family-translational-regulator-orb2", "reader_count"=>21, "reader_count_by_academic_status"=>{"Professor > Associate Professor"=>1, "Researcher"=>6, "Student > Doctoral Student"=>1, "Student > Ph. D. Student"=>6, "Student > Postgraduate"=>1, "Student > Master"=>5, "Professor"=>1}, "reader_count_by_user_role"=>{"Professor > Associate Professor"=>1, "Researcher"=>6, "Student > Doctoral Student"=>1, "Student > Ph. D. Student"=>6, "Student > Postgraduate"=>1, "Student > Master"=>5, "Professor"=>1}, "reader_count_by_subject_area"=>{"Environmental Science"=>1, "Biochemistry, Genetics and Molecular Biology"=>1, "Agricultural and Biological Sciences"=>18, "Neuroscience"=>1}, "reader_count_by_subdiscipline"=>{"Neuroscience"=>{"Neuroscience"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>18}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>1}, "Environmental Science"=>{"Environmental Science"=>1}}, "reader_count_by_country"=>{"India"=>1}, "group_count"=>0}

Scopus | Further Information

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/1498735"], "description"=>"<p>A) Frequency of testes that have at least one spermatid cyst polarized in the wrong orientation so that clustered spermatid cyst nuclei are found in the apical region instead of towards the base of the testis. <i>apkc<sup>48</sup></i> (<i>apkc<sup>ex48</sup></i>), <i>06403</i> (<i>apkc<sup>06403</sup></i>) and <i>apkc<sup>55</sup></i> (<i>apkc<sup>ex55</sup></i>) are <i>apkc</i> mutant alleles. <i>34332</i> and <i>35140</i> are UAS-<i>apkc</i> RNAi lines (dsRNA targeting sites marked in <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1004380#pgen-1004380-g003\" target=\"_blank\">Fig. 3A</a>), while the Gal4 drivers are <i>nos</i> or MTD. MTD has three GAL4 drivers, <i>pCOG-Gal4</i> which has an <i>otu</i> promoter, plus two different <i>nos</i> drivers, <i>NGT-40</i> and <i>nanos</i>-<i>Gal4</i>. Error bars are standard deviation based on three experiments. Total number of testes scored for each genotype are (from left to right): Fig. A1: <i>apkc<sup>ex48</sup></i>: 36; <i>apkc<sup>06403</sup>/+</i>:33; <i>apkc<sup>ex55</sup></i>:27; MTD/UAS-<i>apkc:34332</i>:43; <i>nos-GAL4/</i>UAS-<i>apkc:35140</i>: 18; MTD/UAS-<i>apkc:35140</i>:16; <i>orb<sup>36</sup>/+</i>:55; <i>orb<sup>36</sup></i>: 50; <i>orb2<sup>ΔQ</sup></i>: 86; WT: 65. Fig. A2: <i>apkc<sup>06403</sup>/+</i>: 38; <i>apkc<sup>06403</sup>/</i>+; <i>orb2<sup>ΔQ</sup></i>/+:40; <i>apkc<sup>06403</sup>/</i>+; <i>orb2<sup>36</sup></i>/+: 49; Fig. A3: <i>apkc<sup>ex48</sup>/+</i>: 44; <i>apkc<sup>ex48</sup>/</i>+; <i>orb2<sup>ΔQ</sup></i>/+: 39; <i>apkc<sup>ex48</sup>/</i>+; <i>orb2<sup>36</sup></i>/+: 39; <i>apkc<sup>ex55</sup>/</i>+: 27; <i>apkc<sup>ex55</sup>/</i>+; <i>orb2<sup>ΔQ</sup></i>/+: 28; <i>apkc<sup>ex55</sup>/</i>+; <i>orb2<sup>36</sup></i>/+: 37. B-D) Whole mount antibody staining of wild type or <i>apkc</i> mutant testes. Blue, DNA; red, Orb2. B) Spermatid nuclei clusters (arrow) are found in the spermatogonial region of the testis in <i>apkc</i> hypomorphic alleles. The example shown here is <i>apkc<sup>ex55</sup></i>. D) In the hypomorphic alleles, most spermatid nuclei clusters are located as in wild type on the basal side of testes. D) In MTD/UAS-<i>apkc:35140</i> testes, multiple spermatid nuclei clusters are found in the apical region (arrows). (Not all spermatid cysts express Orb2 and/or are in focus). E) In wild type testes spermatid cyst nuclei are never seen at the apical end of the testis. F) aPKC (green) and Orb2 (red) in a misoriented elongating <i>apkc<sup>ex55</sup></i> spermatid cyst. Note that the Orb2 comet head is present. G) aPKC (green) and Orb2 (red) in a misoriented elongating MTD/UAS-<i>apkc:35140</i> testis. Note that the Orb2 comet is absent. This is also the case for correctly oriented MTD/UAS-<i>apkc:35140</i> spermatid cysts (not shown). All images are orientated with apical side of the testes to the left and basal to the right. Scale bars: 50 µm.</p>", "links"=>[], "tags"=>["developmental biology", "morphogenesis", "Pattern formation", "Cell differentiation", "Molecular development", "genetics", "gene expression", "Gene regulation", "Animal genetics", "Gene function", "Molecular genetics", "cysts", "elongate"], "article_id"=>1027648, "categories"=>["Biological Sciences"], "users"=>["Shuwa Xu", "Sanjay Tyagi", "Paul Schedl"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1004380.g002", "stats"=>{"downloads"=>0, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Spermatid_cysts_elongate_in_the_wrong_direction_in_apkc_or_orb2_mutants_/1027648", "title"=>"Spermatid cysts elongate in the wrong direction in <i>apkc</i> or <i>orb2</i> mutants.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-05-15 02:46:54"}
  • {"files"=>["https://ndownloader.figshare.com/files/1498753"], "description"=>"<p>A) Wild type spermatid cyst showing that Orb2 protein (red) and <i>orb2</i> mRNA (green) are distributed in the characteristic comet pattern and display extensive co-localization, particularly in the comet head. B) In <i>orb2<sup>ΔQ</sup></i> spermatids that elongate in the incorrect direction, Orb2<sup>ΔQ</sup> protein (red) and <i>orb2<sup>ΔQ</sup></i> mRNA (green) are distributed uniformly in the cyst and there is no evidence of a comet head (expected position is indicated by arrow). All images are orientated with apical to the left and basal to the right. Scale bar: 20 µm.</p>", "links"=>[], "tags"=>["developmental biology", "morphogenesis", "Pattern formation", "Cell differentiation", "Molecular development", "genetics", "gene expression", "Gene regulation", "Animal genetics", "Gene function", "Molecular genetics", "autoregulates", "localization"], "article_id"=>1027661, "categories"=>["Biological Sciences"], "users"=>["Shuwa Xu", "Sanjay Tyagi", "Paul Schedl"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1004380.g007", "stats"=>{"downloads"=>0, "page_views"=>8, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_orb2_autoregulates_the_localization_and_translation_of_orb2_mRNA_/1027661", "title"=>"<i>orb2</i> autoregulates the localization and translation of <i>orb2</i> mRNA.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-05-15 02:46:54"}
  • {"files"=>["https://ndownloader.figshare.com/files/1498742"], "description"=>"<p>A) The <i>apkc</i> transcription unit as annotated in <a href=\"http://flybase.org/reports/FBgn0261854.html\" target=\"_blank\">http://flybase.org/reports/FBgn0261854.html</a>. Transcripts expressed from four <i>apkc</i> promoters (P1–P4) plus a collection of alternatively spliced exons and UTRs are predicted to generate more than ten mRNAs. Green bar, protein coding exon; grey bar, UTR; solid black line, intron. <i>apkc-RA</i> CPE sites are labeled in red. Positions of primers used for RT-PCR experiments to identify different <i>apkc</i> mRNA species are marked in dark blue. The two dsRNA sequences used in the <i>apkc</i> RNAi knockdown experiments are marked in pink. Probes for the <i>apkc</i> Orb2 EMSAs, com-GS and RA-GS, are indicated above the gene in brown. B) <i>apkc</i> transcripts expressed in testes were detected by RT-PCR using the primer pairs as indicated. Not all of the primer pairs are specific to one of the four promoters, but some are specific to particular splice forms. C) <i>apkc-RA</i> mRNA can be immunoprecipitated with Orb2 antibodies from WT and <i>orb2<sup>ΔQ</sup></i> testis extracts. β-Gal antibody (a negative control) didn't immunoprecipitate <i>apkc-RA</i> mRNA. D) Orb2 antibody immunoprecipitated <i>orb2</i> mRNA from wild type testes while the control β-Gal antibody did not.</p>", "links"=>[], "tags"=>["developmental biology", "morphogenesis", "Pattern formation", "Cell differentiation", "Molecular development", "genetics", "gene expression", "Gene regulation", "Animal genetics", "Gene function", "Molecular genetics", "mrnas", "orb2"], "article_id"=>1027650, "categories"=>["Biological Sciences"], "users"=>["Shuwa Xu", "Sanjay Tyagi", "Paul Schedl"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1004380.g003", "stats"=>{"downloads"=>0, "page_views"=>14, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_apkc_RA_and_orb2_mRNAs_associate_with_Orb2_in_vivo_/1027650", "title"=>"<i>apkc-RA</i> and <i>orb2</i> mRNAs associate with Orb2 <i>in vivo</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-05-15 02:46:54"}
  • {"files"=>["https://ndownloader.figshare.com/files/1498751"], "description"=>"<p>A) In wild type testes, Orb2 and <i>apkc-RA</i> accumulate in a comet pattern in elongating spermatid cysts located towards the apical side of the testis. B and C) In <i>orb2<sup>ΔQ</sup></i> spermatid cysts that elongate in the incorrect direction, Orb2<sup>ΔQ</sup> protein and both <i>apkc-RA</i> mRNA (detected with the <i>apkc-RA</i> probe, B) and “bulk” <i>apkc</i> mRNA (detected with the <i>apkc-com</i> probe, C) are distributed randomly instead of localizing in the comet pattern. Arrows in A indicate comet heads. In B and C the arrows indicate the expected position of the comet head. In the merged images on the far right, Red: Orb2; Green: <i>apkc</i> mRNA; Blue: DNA. D & E) aPKC protein (arrow) in <i>orb2<sup>ΔQ</sup></i> spermatids elongating in the correct (D) or incorrect (E) orientation. <i>orb2<sup>ΔQ</sup></i> spermatids elongating in the incorrect direction lack aPKC protein stripes at the tip of the elongating flagellar axonemes (compare arrows in D and E). In the merged images on the far right, Red: Tublin; Green: aPKC; Blue: DNA. F & G) <i>apkc-RA</i> (detected with the <i>apkc-RA</i> probe) and “bulk” <i>apkc</i> mRNAs (detected with the <i>apkc-com</i> probe) are expressed in <i>orb2<sup>36</sup></i> spermatids but are not localized. In the merged image on right, Red: Orb2, Green: <i>apkc-RA</i> or bulk (<i>apkc-com</i>) mRNA, and Blue: DNA. All images are orientated with apical side of the testes to the left and basal to the right. Scale bar in A–D: 20 µm; in E and F: 50 µm.</p>", "links"=>[], "tags"=>["developmental biology", "morphogenesis", "Pattern formation", "Cell differentiation", "Molecular development", "genetics", "gene expression", "Gene regulation", "Animal genetics", "Gene function", "Molecular genetics", "polarized", "accumulation", "mrna", "apkc", "spermatid"], "article_id"=>1027659, "categories"=>["Biological Sciences"], "users"=>["Shuwa Xu", "Sanjay Tyagi", "Paul Schedl"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1004380.g006", "stats"=>{"downloads"=>0, "page_views"=>13, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_orb2_is_required_for_polarized_accumulation_of_apkc_RA_mRNA_and_aPKC_protein_in_spermatid_cysts_/1027659", "title"=>"<i>orb2</i> is required for polarized accumulation of <i>apkc-RA</i> mRNA and aPKC protein in spermatid cysts.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-05-15 02:46:54"}
  • {"files"=>["https://ndownloader.figshare.com/files/1498731"], "description"=>"<p>A) Stages of spermatogenesis. Germline cysts are in yellow, while the somatic cyst cells are labeled in dark pink (posterior or head cyst cell) and light pink (anterior or tail cyst cell) respectively. SC, stem cell; SMG, spermatogonia; SMC, spermatocyte; SPT, spermatid. B–E) aPKC expression during spermatogenesis. B) Localized aPKC is not observed in pre-meiotic (arrowhead) or meiotic (arrow) cysts. C) When the spermatid cyst polarizes, aPKC puncta (green) are clustered on the distal side of the polarizing cyst while the spermatid nuclei localize on the proximal side of the cyst. D, E) aPKC stripes at the tip of the growing flagellar axoneme (arrow). Insert in D is a zoom-in view of the marked window. All images are orientated with the apical end of the testes to the left and the basal to the right. Scale bar: 50 µm.</p>", "links"=>[], "tags"=>["developmental biology", "morphogenesis", "Pattern formation", "Cell differentiation", "Molecular development", "genetics", "gene expression", "Gene regulation", "Animal genetics", "Gene function", "Molecular genetics", "spermatid"], "article_id"=>1027644, "categories"=>["Biological Sciences"], "users"=>["Shuwa Xu", "Sanjay Tyagi", "Paul Schedl"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1004380.g001", "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_aPKC_expression_in_spermatid_cysts_/1027644", "title"=>"aPKC expression in spermatid cysts.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-05-15 02:46:54"}
  • {"files"=>["https://ndownloader.figshare.com/files/1498750"], "description"=>"<p>A and B) Green, phalloidin labeled Actin; Blue, DNA; Red, Orb2. A) Clustered nuclei of spermatid cysts (arrowheads) that have completed elongation and just assembled individualization complexes (ICs) (arrows) are found in the spermatogonial/early spermatocyte region of <i>orb2<sup>ΔQ</sup></i> testis. B) White arrows point to two ICs in an <i>orb2<sup>ΔQ</sup></i> testis that are moving in opposite directions in the middle of the testis (yellow arrows indicate directions of motion). C–E) Green, Bol; blue, DNA. C) Like Orb2, the translation factor Bol is localized a comet pattern during flagellar axoneme elongation in wild type testes <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1004380#pgen.1004380-Xu1\" target=\"_blank\">[29]</a>. D) A bi-polar <i>orb2<sup>36</sup></i> spermatid cyst that has Bol concentrated at both elongating ends, while the nuclei are in the middle of the cyst (arrows). E) A partially elongated <i>orb2<sup>36</sup></i> spermatid cyst in which spermatid nuclei and Bol protein are scattered randomly. Dotted line outlines one among four spermatid cysts in the figure. Yellow arrows point out a few of the scattered nuclei. All images are orientated with apical side of the testes to the left and basal to the right. Scale bar: 50 µm.</p>", "links"=>[], "tags"=>["developmental biology", "morphogenesis", "Pattern formation", "Cell differentiation", "Molecular development", "genetics", "gene expression", "Gene regulation", "Animal genetics", "Gene function", "Molecular genetics", "cyst"], "article_id"=>1027658, "categories"=>["Biological Sciences"], "users"=>["Shuwa Xu", "Sanjay Tyagi", "Paul Schedl"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1004380.g005", "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_orb2_is_required_for_cyst_polarization_/1027658", "title"=>"<i>orb2</i> is required for cyst polarization.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-05-15 02:46:54"}
  • {"files"=>["https://ndownloader.figshare.com/files/1498768", "https://ndownloader.figshare.com/files/1498770", "https://ndownloader.figshare.com/files/1498771", "https://ndownloader.figshare.com/files/1498772", "https://ndownloader.figshare.com/files/1498773"], "description"=>"<div><p>Mature <i>Drosophila</i> sperm are highly polarized cells—on one side is a nearly 2 mm long flagellar tail that comprises most of the cell, while on the other is the sperm head, which carries the gamete's genetic information. The polarization of the sperm cells commences after meiosis is complete and the 64-cell spermatid cyst begins the process of differentiation. The spermatid nuclei cluster to one side of the cyst, while the flagellar axonemes grows from the other. The elongating spermatid bundles are also polarized with respect to the main axis of the testis; the sperm heads are always oriented basally, while the growing tails extend apically. This orientation within the testes is important for transferring the mature sperm into the seminal vesicles. We show here that orienting cyst polarization with respect to the main axis of the testis depends upon atypical Protein Kinase C (aPKC), a factor implicated in polarity decisions in many different biological contexts. When <i>apkc</i> activity is compromised in the male germline, the direction of cyst polarization within this organ is randomized. Significantly, the mechanisms used to spatially restrict <i>apkc</i> activity to the apical side of the spermatid cyst are different from the canonical cross-regulatory interactions between this kinase and other cell polarity proteins that normally orchestrate polarization. We show that the asymmetric accumulation of aPKC protein in the cyst depends on an mRNA localization pathway that is regulated by the <i>Drosophila</i> CPEB protein Orb2. <i>orb2</i> is required to properly localize and activate the translation of <i>apkc</i> mRNAs in polarizing spermatid cysts. We also show that <i>orb2</i> functions not only in orienting cyst polarization with respect to the apical-basal axis of the testis, but also in the process of polarization itself. One of the <i>orb2</i> targets in this process is its own mRNA. Moreover, the proper execution of this <i>orb2</i> autoregulatory pathway depends upon <i>apkc</i>.</p></div>", "links"=>[], "tags"=>["developmental biology", "morphogenesis", "Pattern formation", "Cell differentiation", "Molecular development", "genetics", "gene expression", "Gene regulation", "Animal genetics", "Gene function", "Molecular genetics", "spermatid", "cyst", "polarization", "depends", "cpeb", "translational"], "article_id"=>1027671, "categories"=>["Biological Sciences"], "users"=>["Shuwa Xu", "Sanjay Tyagi", "Paul Schedl"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1004380.s001", "https://dx.doi.org/10.1371/journal.pgen.1004380.s002", "https://dx.doi.org/10.1371/journal.pgen.1004380.s003", "https://dx.doi.org/10.1371/journal.pgen.1004380.s004", "https://dx.doi.org/10.1371/journal.pgen.1004380.s005"], "stats"=>{"downloads"=>22, "page_views"=>15, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Spermatid_Cyst_Polarization_in_Drosophila_Depends_upon_apkc_and_the_CPEB_Family_Translational_Regulator_orb2_/1027671", "title"=>"Spermatid Cyst Polarization in <i>Drosophila</i> Depends upon <i>apkc</i> and the CPEB Family Translational Regulator <i>orb2</i>", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2014-05-15 02:46:54"}
  • {"files"=>["https://ndownloader.figshare.com/files/1498745"], "description"=>"<p>A, B) Testes were probed with <i>apkc-com</i> or <i>apkc-RA</i> specific fluorescent oligos. Panels highlight <i>apkc</i> expression at different stages of spermatogenesis. A) Left panel: mRNAs complementary to the <i>apkc-com</i> mRNA probe (specific for an exon that is present in all of the predicted <i>apkc</i> mRNA species – “bulk” <i>apkc</i> mRNA) were detected in the stem cell/spermatogonial/early spermatocytes region of the testis. Middle and right panel: “Bulk” <i>apkc</i> mRNA complementary to the <i>apkc-com</i> probe is also present at moderate levels in spermatocytes (SMC) and spermatids (SPT) (white arrows). In elongating spermatid cysts bulk <i>apkc</i> mRNA is localized in a comet pattern (yellow arrow). B) Left panel: Unlike “bulk” <i>apkc</i> mRNA, <i>apkc-RA</i> is not expressed in stem cells or spermatogonial cysts. Middle panel: <i>apkc-RA</i> is also absent in spermatocytes (SMC, arrowhead). Low levels of <i>apkc-RA</i> are first detected in spermatid cysts (SPT, white arrow) after meiosis is complete but prior to cyst polarization. Right panel: High levels of <i>apkc-RA</i> are present during the elongation phase (yellow arrow). At this stage <i>apkc-RA</i> accumulates in the characteristic comet pattern. C) aPKC protein stripes are found at the proximal (away from the elongating frontier) edge of the <i>apkc-RA</i> comet head. D) “Bulk” <i>apkc</i> mRNAs (detected with the <i>apkc</i>-<i>com</i> probe) and E) <i>apkc-RA</i> mRNAs co-localize with Orb2 in a comet pattern. The most extensive co-localization of mRNA and protein appears to be in the comet head. F) aPKC protein stripes accumulate at the proximal edge of the Orb2 protein comet head. All images are orientated with the apical end of the testis is to the left and basal end to the right. Scale bar: 20 µm.</p>", "links"=>[], "tags"=>["developmental biology", "morphogenesis", "Pattern formation", "Cell differentiation", "Molecular development", "genetics", "gene expression", "Gene regulation", "Animal genetics", "Gene function", "Molecular genetics", "accumulation", "apkc", "orb2"], "article_id"=>1027653, "categories"=>["Biological Sciences"], "users"=>["Shuwa Xu", "Sanjay Tyagi", "Paul Schedl"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1004380.g004", "stats"=>{"downloads"=>0, "page_views"=>9, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Localized_accumulation_of_apkc_mRNA_aPKC_and_Orb2_protein_/1027653", "title"=>"Localized accumulation of <i>apkc</i> mRNA, aPKC and Orb2 protein.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2014-05-15 02:46:54"}

PMC Usage Stats | Further Information

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  • {"unique-ip"=>"19", "full-text"=>"14", "pdf"=>"6", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"1", "supp-data"=>"3", "cited-by"=>"0", "year"=>"2015", "month"=>"1"}
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  • {"unique-ip"=>"8", "full-text"=>"6", "pdf"=>"2", "abstract"=>"0", "scanned-summary"=>"0", "scanned-page-browse"=>"0", "figure"=>"3", "supp-data"=>"0", "cited-by"=>"1", "year"=>"2017", "month"=>"1"}
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Relative Metric

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