Deubiquitylation Machinery Is Required for Embryonic Polarity in Caenorhabditis elegans
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{"title"=>"Deubiquitylation Machinery Is Required for Embryonic Polarity in Caenorhabditis elegans", "type"=>"journal", "authors"=>[{"first_name"=>"Richard J.", "last_name"=>"McCloskey", "scopus_author_id"=>"55513297100"}, {"first_name"=>"Kenneth J.", "last_name"=>"Kemphues", "scopus_author_id"=>"7003789130"}], "year"=>2012, "source"=>"PLoS Genetics", "identifiers"=>{"sgr"=>"84870685075", "issn"=>"15537390", "scopus"=>"2-s2.0-84870685075", "pmid"=>"23209443", "pui"=>"366216288", "isbn"=>"1553-7404 (Electronic)\\r1553-7390 (Linking)", "doi"=>"10.1371/journal.pgen.1003092"}, "id"=>"951da3c4-6eb4-3dc6-947d-ed2881eef732", "abstract"=>"The Caenorhabditis elegans one-cell embryo polarizes in response to a cue from the paternally donated centrosome and asymmetrically segregates cell fate determinants that direct the developmental program of the worm. We have found that genes encoding putative deubiquitylating enzymes (DUBs) are required for polarization of one-cell embryos. Maternal loss of the proteins MATH-33 and USP-47 leads to variable inability to correctly establish and maintain asymmetry as defined by posterior and anterior polarity proteins PAR-2 and PAR-3. The first observable defect is variable positioning of the centrosome with respect to the cell cortex and the male pronucleus. The severity of the polarity defects correlates with distance of the centrosome from the cortex. Furthermore, polarity defects can be bypassed by mutations that bring the centrosome in close proximity to the cortex. In addition we find that polarity and centrosome positioning defects can be suppressed by compromising protein turnover. We propose that the DUB activity of MATH-33 and USP-47 stabilizes one or more proteins required for association of the centrosome with the cortex. Because these DUBs are homologous to two members of a group of DUBs that act in fission yeast polarity, we tested additional members of that family and found that another C. elegans DUB gene, usp-46, also contributes to polarity. Our finding that deubiquitylating enzymes required for polarity in Schizosaccharomyces pombe are also required in C. elegans raises the possibility that these DUBs act through an evolutionarily conserved mechanism to control cell polarity.", "link"=>"http://www.mendeley.com/research/deubiquitylation-machinery-required-embryonic-polarity-caenorhabditis-elegans", "reader_count"=>33, "reader_count_by_academic_status"=>{"Unspecified"=>2, "Professor > Associate Professor"=>1, "Librarian"=>1, "Student > Doctoral Student"=>3, "Researcher"=>9, "Student > Ph. D. Student"=>11, "Student > Master"=>3, "Student > Bachelor"=>1, "Professor"=>2}, "reader_count_by_user_role"=>{"Unspecified"=>2, "Professor > Associate Professor"=>1, "Librarian"=>1, "Student > Doctoral Student"=>3, "Researcher"=>9, "Student > Ph. D. Student"=>11, "Student > Master"=>3, "Student > Bachelor"=>1, "Professor"=>2}, "reader_count_by_subject_area"=>{"Unspecified"=>2, "Engineering"=>1, "Biochemistry, Genetics and Molecular Biology"=>8, "Mathematics"=>1, "Agricultural and Biological Sciences"=>19, "Computer Science"=>1, "Economics, Econometrics and Finance"=>1}, "reader_count_by_subdiscipline"=>{"Engineering"=>{"Engineering"=>1}, "Economics, Econometrics and Finance"=>{"Economics, Econometrics and Finance"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>19}, "Computer Science"=>{"Computer Science"=>1}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>8}, "Mathematics"=>{"Mathematics"=>1}, "Unspecified"=>{"Unspecified"=>2}}, "reader_count_by_country"=>{"Republic of Singapore"=>1, "United States"=>3, "Switzerland"=>3}, "group_count"=>0}

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/287723", "https://ndownloader.figshare.com/files/287798", "https://ndownloader.figshare.com/files/287839", "https://ndownloader.figshare.com/files/287894", "https://ndownloader.figshare.com/files/287939", "https://ndownloader.figshare.com/files/287985", "https://ndownloader.figshare.com/files/288041", "https://ndownloader.figshare.com/files/288111", "https://ndownloader.figshare.com/files/288166", "https://ndownloader.figshare.com/files/288215", "https://ndownloader.figshare.com/files/288261", "https://ndownloader.figshare.com/files/288316", "https://ndownloader.figshare.com/files/288366"], "description"=>"<div><p>The <em>Caenorhabditis elegans</em> one-cell embryo polarizes in response to a cue from the paternally donated centrosome and asymmetrically segregates cell fate determinants that direct the developmental program of the worm. We have found that genes encoding putative deubiquitylating enzymes (DUBs) are required for polarization of one-cell embryos. Maternal loss of the proteins MATH-33 and USP-47 leads to variable inability to correctly establish and maintain asymmetry as defined by posterior and anterior polarity proteins PAR-2 and PAR-3. The first observable defect is variable positioning of the centrosome with respect to the cell cortex and the male pronucleus. The severity of the polarity defects correlates with distance of the centrosome from the cortex. Furthermore, polarity defects can be bypassed by mutations that bring the centrosome in close proximity to the cortex. In addition we find that polarity and centrosome positioning defects can be suppressed by compromising protein turnover. We propose that the DUB activity of MATH-33 and USP-47 stabilizes one or more proteins required for association of the centrosome with the cortex. Because these DUBs are homologous to two members of a group of DUBs that act in fission yeast polarity, we tested additional members of that family and found that another <em>C. elegans</em> DUB gene, <em>usp-46</em>, also contributes to polarity. Our finding that deubiquitylating enzymes required for polarity in <em>Schizosaccharomyces pombe</em> are also required in <em>C. elegans</em> raises the possibility that these DUBs act through an evolutionarily conserved mechanism to control cell polarity.</p> </div>", "links"=>[], "tags"=>["deubiquitylation", "machinery", "embryonic", "polarity"], "article_id"=>116678, "categories"=>["Biochemistry", "Cell Biology", "Genetics", "Developmental Biology"], "users"=>["Richard J. McCloskey", "Kenneth J. Kemphues"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1003092.s001", "https://dx.doi.org/10.1371/journal.pgen.1003092.s002", "https://dx.doi.org/10.1371/journal.pgen.1003092.s003", "https://dx.doi.org/10.1371/journal.pgen.1003092.s004", "https://dx.doi.org/10.1371/journal.pgen.1003092.s005", "https://dx.doi.org/10.1371/journal.pgen.1003092.s006", "https://dx.doi.org/10.1371/journal.pgen.1003092.s007", "https://dx.doi.org/10.1371/journal.pgen.1003092.s008", "https://dx.doi.org/10.1371/journal.pgen.1003092.s009", "https://dx.doi.org/10.1371/journal.pgen.1003092.s010", "https://dx.doi.org/10.1371/journal.pgen.1003092.s011", "https://dx.doi.org/10.1371/journal.pgen.1003092.s012", "https://dx.doi.org/10.1371/journal.pgen.1003092.s013"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Deubiquitylation_Machinery_Is_Required_for_Embryonic_Polarity_in_Caenorhabditis_elegans__/116678", "title"=>"Deubiquitylation Machinery Is Required for Embryonic Polarity in <em>Caenorhabditis elegans</em>", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2012-11-29 01:51:18"}
  • {"files"=>["https://ndownloader.figshare.com/files/534064"], "description"=>"<p>(A) Bars indicate percent embryo lethality. >300 embryos were scored for each genotype in two trials. Error bars indicate standard error of the mean; asterisks indicate significant difference compared to controls, <i>p</i><0.01, determined by Student's t-test. <i>K09A9.4</i> and <i>T24B8.7</i> are negative control DUBs. (B) Un-rooted phylogenetic tree of 25 UCH domains in <i>C. elegans</i> and two human homologs of <i>math-33</i> and <i>usp-47</i>. We did not perform RNAi on the three genes indicated by ‘. Human homologs USP7 and USP47 are included to highlight that there is evolutionary conservation of the DUBs between species. (C) DIC time-lapse of one-cell to four-cell embryos of the indicated genotypes at 22°C. Embryos are oriented with the anterior at the left in this and subsequent figures. The spindle orientation of P1 as it divides is usually longitudinal to the embryo axis, but is transverse in polarity-defective mutants such as <i>par-2(lw32)</i>. The scale bar represents 10 µm.</p>", "links"=>[], "tags"=>["synthetic", "polarity"], "article_id"=>204562, "categories"=>["Biochemistry", "Cell Biology", "Genetics", "Developmental Biology"], "users"=>["Richard J. McCloskey", "Kenneth J. Kemphues"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1003092.g001"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Paralogs_math_33_and_usp_47_have_synthetic_polarity_phenotypes_/204562", "title"=>"Paralogs <i>math-33</i> and <i>usp-47</i> have synthetic polarity phenotypes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-11-29 01:16:02"}
  • {"files"=>["https://ndownloader.figshare.com/files/534188"], "description"=>"<p>(A) Confocal micrographs of embryos immunostained for PAR-3 (green) and PAR-2 (red). Compared to controls, PAR-3 occupies a larger region of the cortex and PAR-2 a smaller region in the <i>math-33(tm3561); usp-47(RNAi)</i> embryo. (B) Averages of PAR-3 and PAR-2 domain sizes in immunostained embryos as a percentage of embryo length. <i>n</i> indicates the number of anterior and posterior domains examined respectively. Asterisks mark statistical significance compared to wild type, <i>p</i><0.01, by Student's t-test. (C) Time-lapse images displaying the dynamic localization of PAR-2::GFP in the indicated genotypes. The embryo genotype of the control row (top) showing wild-type behavior is <i>par-2::gfp; math-33(tm3561)/nT1</i>. <i>math-33(tm3561); usp-47(RNAi)</i> embryos weakly recruit PAR-2 and could be grouped into two classes: those that maintain a PAR-2 domain throughout the cell cycle (class I), and those in which the weak PAR-2 domain is not maintained (class II). Scale bars represent 5 µm.</p>", "links"=>[], "tags"=>["defects"], "article_id"=>204690, "categories"=>["Biochemistry", "Cell Biology", "Genetics", "Developmental Biology"], "users"=>["Richard J. McCloskey", "Kenneth J. Kemphues"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1003092.g002"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Polarity_defects_in_math_33_tm3561_usp_47_RNAi_embryos_/204690", "title"=>"Polarity defects in <i>math-33(tm3561);usp-47(RNAi)</i> embryos.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-11-29 01:18:10"}
  • {"files"=>["https://ndownloader.figshare.com/files/534308"], "description"=>"<p>(A) Embryonic lethality of the indicated genotypes. Error bars indicate the standard error of the mean. The experimental class marked with an asterisk is significantly different from all three controls, <i>p</i><0.05, by Student's t-test. (B) LGL-1::GFP time-lapse images of embryos of the indicated genotypes. Upon knockdown of the DUBs MATH-33 and USP-47, the LGL-1::GFP domain size is smaller than in control embryos and is not maintained, similar to previous results with PAR-2::GFP in <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1003092#pgen-1003092-g002\" target=\"_blank\">Figure 2C</a>, and transverse spindles occur in P1. Scale bar represents 5 µm.</p>", "links"=>[], "tags"=>["mediate", "polarity"], "article_id"=>204808, "categories"=>["Biochemistry", "Cell Biology", "Genetics", "Developmental Biology"], "users"=>["Richard J. McCloskey", "Kenneth J. Kemphues"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1003092.g003"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_PAR_2_function_is_not_required_to_mediate_loss_of_polarity_in_math_33_RNAi_usp_47_RNAi_/204808", "title"=>"PAR-2 function is not required to mediate loss of polarity in <i>math-33(RNAi) usp-47(RNAi)</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-11-29 01:20:08"}
  • {"files"=>["https://ndownloader.figshare.com/files/534418"], "description"=>"<p>(A) Time-lapse confocal images of embryos expressing NMY-2::GFP. The maximum projections of sections through the cortex of the embryo show the localization of NMY-2::GFP foci at three stages. Kymographs illustrate the pattern of myosin clearing and flow over time. About halfway through the kymographs (from top to bottom), large myosin foci transition into smaller myosin puncta. (B) The extent of myosin absence from the posterior (clearing) was measured as a proportion of the embryo length and the maximum level of clearing was recorded; each data point represents one embryo. (C) Confocal stacks of embryos immunostained for NMY-2 (green) and tubulin (red). Arrows indicate the location of the centrosome, red dotted circles mark the position of pronuclei, and the extent of myosin clearing is indicated by arrowheads. Scale bars represent 5 µm.</p>", "links"=>[], "tags"=>["posterior", "anterior", "myosin", "clearing", "defective"], "article_id"=>204913, "categories"=>["Biochemistry", "Cell Biology", "Genetics", "Developmental Biology"], "users"=>["Richard J. McCloskey", "Kenneth J. Kemphues"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1003092.g004"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_posterior_to_anterior_myosin_clearing_is_defective_in_math_33_tm3561_usp_47_RNAi_/204913", "title"=>"The posterior to anterior myosin clearing is defective in <i>math-33(tm3561); usp-47(RNAi)</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-11-29 01:21:53"}
  • {"files"=>["https://ndownloader.figshare.com/files/534587"], "description"=>"<p>(A) Micrographs showing tubulin::GFP in embryos at the time the centrosome was first detected. Centrosomes are marked with a white arrowhead and the position of the paternal pronucleus is indicated by red dotted circles. The DIC micrograph column shows representative embryos of the same genotypes at the time of pseudocleavage (an indirect indicator of the extent of myosin clearing). The white arrows indicate the location of pseudocleavage furrows. (B) Distance in micrometers of the centrosome from the embryo cortex at the time of earliest detection for the indicated genotypes. Each data point represents a single embryo. Black dots are embryos that had normal or weak pseudocleavage, red dots are embryos that displayed no pseudocleavage. Blue bars mark the mean distance. The asterisk indicates a significant difference from WT controls, <i>p</i><0.01, by Student's t-test. (C) The number of embryos lacking pseudocleavage increases as centrosome distance from the cortex increases. Embryos were pooled from five RNAi experiment replicates. Differences were statistically significant according to a Student's t-test, <i>p</i><0.01. (D) The distance of the centrosome from the cortex measured in the indicated genetic backgrounds. <i>p</i><0.01 for column 4 compared to WT distances from multiple experiments, but <i>p</i> = 0.156 for column 4 compared to <i>math-33(tm3561)</i> in multiple experiments. (E) A single experiment examining absence of pseudocleavage resulting from depletion of DUBs. The absence of pseudocleavage phenotype is completely suppressed by <i>dhc-1(RNAi), p</i><0.05. Scale bar represents 5 µm.</p>", "links"=>[], "tags"=>["centrosome", "one-cell"], "article_id"=>205093, "categories"=>["Biochemistry", "Cell Biology", "Genetics", "Developmental Biology"], "users"=>["Richard J. McCloskey", "Kenneth J. Kemphues"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1003092.g005"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Deubiquitylases_regulate_the_position_of_the_centrosome_in_early_one_cell_embryos_/205093", "title"=>"Deubiquitylases regulate the position of the centrosome in early one-cell embryos.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-11-29 01:24:53"}
  • {"files"=>["https://ndownloader.figshare.com/files/534701"], "description"=>"<p>The left panels are representative images of embryos showing the distribution of GFP::MATH-33 and GFP::USP-47 in one-cell embryos. Both proteins are present in the cytoplasm, but MATH-33 is enriched in nuclei. The right panels are images of embryos showing immunostaining of endogenous MATH-33(red) in the cytoplasm and the nucleus; USP-47(green) primarily in the cytoplasm. Scale bar represents 5 µm.</p>", "links"=>[], "tags"=>["usp-47", "enriched", "cortex"], "article_id"=>205198, "categories"=>["Biochemistry", "Cell Biology", "Genetics", "Developmental Biology"], "users"=>["Richard J. McCloskey", "Kenneth J. Kemphues"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1003092.g006"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_MATH_33_and_USP_47_are_not_enriched_at_cortex_or_centrosome_/205198", "title"=>"MATH-33 and USP-47 are not enriched at cortex or centrosome.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-11-29 01:26:38"}
  • {"files"=>["https://ndownloader.figshare.com/files/534764"], "description"=>"<p>(A) Embryonic lethality of <i>math-33(tm3561); usp-47(RNAi)</i> is reduced by either of two <i>rpn-10</i> mutations. Standard error of the mean is indicated by the error bars. <i>n</i>>350 embryos for N2 controls and <i>n</i>>600 for the other genotypes. Asterisks indicate significance compared to <i>math-33(tm3561)</i> single mutants, <i>p</i><0.01, Student's t-test (B) Data showing the suppression of phenotypic defects in early <i>math-33(tm3561);usp-47(RNAi)</i> embryos by <i>rpn-10</i> mutations. (C) Distance in micrometers of the centrosome from the embryo cortex when it is first detectable. <i>rpn-10</i> mutation suppresses the absence-of-pseudocleavage phenotype and the mislocalization of the centrosome compared to <i>math-33(tm3561); usp-47(RNAi)</i> controls. None of the 18 centrosomes observed in column 2 were detached from the paternal pronucleus, indicating that the detachment phenotype was also completely suppressed. Results in column 4 were significantly different in a Student's t-test <i>p</i><0.01 compared to column 1 controls. Data from columns 1, 3, and 4 are also displayed in <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1003092#pgen-1003092-g005\" target=\"_blank\">Figure 5C</a>.</p>", "links"=>[], "tags"=>["suppresses", "lethality", "polarity", "defects"], "article_id"=>205263, "categories"=>["Biochemistry", "Cell Biology", "Genetics", "Developmental Biology"], "users"=>["Richard J. McCloskey", "Kenneth J. Kemphues"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1003092.g007"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Mutation_of_rpn_10_suppresses_lethality_and_polarity_defects_in_math_33_tm3561_usp_47_RNAi_embryos_/205263", "title"=>"Mutation of <i>rpn-10</i> suppresses lethality and polarity defects in <i>math-33(tm3561); usp-47(RNAi)</i> embryos.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-11-29 01:27:43"}
  • {"files"=>["https://ndownloader.figshare.com/files/534840"], "description"=>"<p>(A) Two-cell embryos at interphase (top) illustrate unequal vs. equal first divisions and at P1 mitosis (bottom) show spindle orientations. Maternal genotypes are indicated. Scale bars for each genotype represent 5 µm. (B) Embryonic lethality measured after depleting <i>math-33</i> in <i>usp-46</i> and <i>usp-47</i> mutants. Bars marked with an asterisk are significantly different from the RNAi controls in a t-test, <i>p</i><0.01. Scale bar represents 5 µm.</p>", "links"=>[], "tags"=>["acts", "redundantly"], "article_id"=>205341, "categories"=>["Biochemistry", "Cell Biology", "Genetics", "Developmental Biology"], "users"=>["Richard J. McCloskey", "Kenneth J. Kemphues"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1003092.g008"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_usp_46_acts_redundantly_with_math_33_and_usp_47_/205341", "title"=>"<i>usp-46</i> acts redundantly with <i>math-33</i> and <i>usp-47</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2012-11-29 01:29:01"}
  • {"files"=>["https://ndownloader.figshare.com/files/534914"], "description"=>"<p><i>n</i> = the number of embryos counted for each genotype.</p>*<p><i>math-33(RNAi)</i> lethality different from control lethality by Student's t-test, p<0.05.</p>", "links"=>[], "tags"=>["lethality", "mutants", "depleted", "math-33"], "article_id"=>205411, "categories"=>["Biochemistry", "Cell Biology", "Genetics", "Developmental Biology"], "users"=>["Richard J. McCloskey", "Kenneth J. Kemphues"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1003092.t001"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Embryonic_lethality_of_par_mutants_depleted_of_MATH_33_by_RNAi_/205411", "title"=>"Embryonic lethality of <i>par</i> mutants depleted of MATH-33 by RNAi.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2012-11-29 01:30:11"}
  • {"files"=>["https://ndownloader.figshare.com/files/534947"], "description"=>"<p>T-T indicates that both AB and P1 cells of two-cell stage embryos divided transversely. The RNAi and mutant combinations of different DUB genes shown here are ordered according to the phenotypic penetrance of the transverse mitotic spindle orientations in P1.</p>", "links"=>[], "tags"=>["combinations", "p1", "spindle"], "article_id"=>205449, "categories"=>["Biochemistry", "Cell Biology", "Genetics", "Developmental Biology"], "users"=>["Richard J. McCloskey", "Kenneth J. Kemphues"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1003092.t003"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Effect_of_RNAi_mutant_combinations_of_math_33_usp_46_and_usp_47_on_P1_spindle_orientation_/205449", "title"=>"Effect of RNAi/mutant combinations of <i>math-33</i>, <i>usp-46</i>, and <i>usp-47</i> on P1 spindle orientation.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2012-11-29 01:30:49"}
  • {"files"=>["https://ndownloader.figshare.com/files/534975"], "description"=>"a<p>proportion of embryos in which AB and P1 divided within 10 seconds of one another.</p>b<p>Anaphase spindle orientations in two-cell embryos. T-L is the wild-type pattern with AB oriented transversely (T) and P1 oriented longitudinally (L).</p>c<p>ratio of the area of an optical cross-section of the AB cell at mid-focal plane to the total embryo area in the same section (see <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1003092#s4\" target=\"_blank\">materials and methods</a>).</p>", "links"=>[], "tags"=>["phenotypes", "loss-of-function"], "article_id"=>205476, "categories"=>["Biochemistry", "Cell Biology", "Genetics", "Developmental Biology"], "users"=>["Richard J. McCloskey", "Kenneth J. Kemphues"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1003092.t002"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Polarity_phenotypes_of_math_33_par_loss_of_function_embryos_/205476", "title"=>"Polarity phenotypes of <i>math-33; par</i> loss-of-function embryos.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2012-11-29 01:31:16"}

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Relative Metric

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