Mouse TRIP13/PCH2 Is Required for Recombination and Normal Higher-Order Chromosome Structure during Meiosis
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{"title"=>"Mouse TRIP13/PCH2 is required for recombination and normal higher-order chromosome structure during meiosis", "type"=>"journal", "authors"=>[{"first_name"=>"Ignasi", "last_name"=>"Roig", "scopus_author_id"=>"24345435300"}, {"first_name"=>"James A.", "last_name"=>"Dowdle", "scopus_author_id"=>"57198434682"}, {"first_name"=>"Attila", "last_name"=>"Toth", "scopus_author_id"=>"16551533700"}, {"first_name"=>"Dirk G.", "last_name"=>"de Rooij", "scopus_author_id"=>"7007029491"}, {"first_name"=>"Maria", "last_name"=>"Jasin", "scopus_author_id"=>"7005221713"}, {"first_name"=>"Scott", "last_name"=>"Keeney", "scopus_author_id"=>"7005614687"}], "year"=>2010, "source"=>"PLoS Genetics", "identifiers"=>{"pui"=>"359697343", "sgr"=>"77957341191", "pmid"=>"20711356", "scopus"=>"2-s2.0-77957341191", "isbn"=>"1553-7404 (Electronic)\\r1553-7390 (Linking)", "doi"=>"10.1371/journal.pgen.1001062", "issn"=>"15537390"}, "id"=>"0c872dc7-6f01-3c21-9ce1-f2303fc00dbc", "abstract"=>"Accurate chromosome segregation during meiosis requires that homologous chromosomes pair and become physically connected so that they can orient properly on the meiosis I spindle. These connections are formed by homologous recombination closely integrated with the development of meiosis-specific, higher-order chromosome structures. The yeast Pch2 protein has emerged as an important factor with roles in both recombination and chromosome structure formation, but recent analysis suggested that TRIP13, the mouse Pch2 ortholog, is not required for the same processes. Using distinct Trip13 alleles with moderate and severe impairment of TRIP13 function, we report here that TRIP13 is required for proper synaptonemal complex formation, such that autosomal bivalents in Trip13-deficient meiocytes frequently displayed pericentric synaptic forks and other defects. In males, TRIP13 is required for efficient synapsis of the sex chromosomes and for sex body formation. Furthermore, the numbers of crossovers and chiasmata are reduced in the absence of TRIP13, and their distribution along the chromosomes is altered, suggesting a role for TRIP13 in aspects of crossover formation and/or control. Recombination defects are evident very early in meiotic prophase, soon after DSB formation. These findings provide evidence for evolutionarily conserved functions for TRIP13/Pch2 in both recombination and formation of higher order chromosome structures, and they support the hypothesis that TRIP13/Pch2 participates in coordinating these key aspects of meiotic chromosome behavior.", "link"=>"http://www.mendeley.com/research/mouse-trip13pch2-required-recombination-normal-higherorder-chromosome-structure-during-meiosis", "reader_count"=>69, "reader_count_by_academic_status"=>{"Professor > Associate Professor"=>6, "Researcher"=>24, "Student > Doctoral Student"=>2, "Student > Ph. D. Student"=>24, "Student > Postgraduate"=>2, "Student > Master"=>5, "Student > Bachelor"=>1, "Lecturer"=>2, "Lecturer > Senior Lecturer"=>1, "Professor"=>2}, "reader_count_by_user_role"=>{"Professor > Associate Professor"=>6, "Researcher"=>24, "Student > Doctoral Student"=>2, "Student > Ph. D. Student"=>24, "Student > Postgraduate"=>2, "Student > Master"=>5, "Student > Bachelor"=>1, "Lecturer"=>2, "Lecturer > Senior Lecturer"=>1, "Professor"=>2}, "reader_count_by_subject_area"=>{"Biochemistry, Genetics and Molecular Biology"=>11, "Agricultural and Biological Sciences"=>56, "Medicine and Dentistry"=>2}, "reader_count_by_subdiscipline"=>{"Medicine and Dentistry"=>{"Medicine and Dentistry"=>2}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>56}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>11}}, "reader_count_by_country"=>{"Canada"=>1, "United States"=>2}, "group_count"=>1}

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/835686"], "description"=>"<p>(A) Left panel: <i>Trip13</i> expression in different mouse tissues detected by RT-PCR from a commercial set of cDNAs (Origene). β actin expression served as a control. In addition to full-length transcript, a smaller transcript was identified lacking exon 8 (Δ<i>Ex.8 Trip13</i>) that was abundant in adult testis, but was at trace levels or absent in other tissues. The small size difference (87 bp) may explain why this isoform was not previously detected. Middle and right panels: <i>Trip13</i> expression detected by RT-PCR on samples from whole testes of juveniles of the indicated ages (in dpp) and flow-sorted populations from adult testes, enriched for the indicated cell types (spermatogonia (SG), spermatocytes (SC), and spermatids (SD)). RT-PCR analysis during the first, semi-synchronous meiotic wave in juvenile testes detected the longer <i>Trip13</i> transcript in all samples analyzed, including pre-meiotic samples (5 dpp). In contrast, Δ<i>Ex.8 Trip13</i> was detected only in samples containing cells that had already entered meiosis (12 dpp onwards; meiotic prophase starts ∼8 dpp). RT-PCR on flow-sorted, highly enriched cell populations from adult testis detected full-length <i>Trip13</i> transcript in all cell types analyzed, but Δ<i>Ex.8 Trip13</i> was seen only in meiotic (spermatocytes) and post-meiotic cell types (spermatids). <i>Trip13</i> is thus expressed throughout spermatogenesis, but with developmentally regulated exclusion of exon 8 via alternative splicing. (B–D) RNA <i>in situ</i> hybridization to a testis section. The <i>Trip13</i> hybridization signal appears as dark grains in the bright field image of a seminiferous tubule (B) and white grains in the corresponding dark field image (C). A zoomed image of the same tubule (D) shows <i>Trip13</i> hybridization signal overlaying spermatogonia (SG), spermatocytes (SC) and spermatids (SD). Bars  = 20 µm. (E) Schematic showing <i>Trip13</i> splicing variants as well as contribution from exons 5–10 to the AAA+ ATPase domain. Walker A and B motifs (WA and WB) are depicted as green boxes. The Δ<i>Ex.8 Trip13</i> transcript maintains the open reading frame, but lacks sequences for the Walker B ATPase motif. (F) Schematic of the <i>Trip13</i> alleles used in this study. Red boxes indicate gene trap insertion locations. (G) RT-PCR analysis of <i>Trip13</i> expression in whole-testis samples from wild-type and mutant animals. PCR products were detected by ethidium bromide staining (EtBr) or Southern blotting (two exposures of the Southern blot are shown). Only the full-length <i>Trip13</i> transcript is detected in <i>Trip13<sup>sev/sev</sup></i> testes, which may be a consequence of later stage germ cells being absent in this mutant (see <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1001062#pgen-1001062-g001\" target=\"_blank\">Figure 1A</a>).</p>", "links"=>[], "tags"=>["biochemistry/replication and repair", "developmental biology/germ cells", "genetics and genomics/chromosome biology", "genetics and genomics/gene function", "genetics and genomics/nuclear structure and function"], "article_id"=>506048, "categories"=>["Biochemistry", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Ignasi Roig", "James A. Dowdle", "Attila Toth", "Dirk G. de Rooij", "Maria Jasin", "Scott Keeney"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1001062.g001", "stats"=>{"downloads"=>0, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Trip13_expression_in_mouse_/506048", "title"=>"<i>Trip13</i> expression in mouse.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-08-12 01:40:48"}
  • {"files"=>["https://ndownloader.figshare.com/files/837166"], "description"=>"a<p>Best-fit gamma distribution shape parameters, corrected as described <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1001062#pgen.1001062-deBoer1\" target=\"_blank\">[38]</a> for the fact that the gamma distribution assumes theoretical limits of infinitely small and infinitely large inter-focus distances, but the actual range of inter-focus distances that can be detected is limited by the resolution of light microscopy and by the finite length of each SC.</p>", "links"=>[], "tags"=>["parameters", "mlh1", "inter-focus"], "article_id"=>507533, "categories"=>["Biochemistry", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Ignasi Roig", "James A. Dowdle", "Attila Toth", "Dirk G. de Rooij", "Maria Jasin", "Scott Keeney"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1001062.t004", "stats"=>{"downloads"=>0, "page_views"=>8, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Gamma_distribution_parameters_for_MLH1_inter_focus_distances_/507533", "title"=>"Gamma distribution parameters for MLH1 inter-focus distances.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-08-12 02:05:33"}
  • {"files"=>["https://ndownloader.figshare.com/files/837280"], "description"=>"a<p>Significantly different from wild type (p≤0.05, t test).</p>b<p>Significantly different from <i>Trip13<sup>mod/mod</sup></i> (p≤0.05, t test).</p>c<p>Detected with an anti-RAD51 antibody that cross-reacts with DMC1.</p>d<p>Detected with a DMC1-specific antibody.</p><p>n.a., not applicable; n.d., not determined.</p>", "links"=>[], "tags"=>["recombination-associated", "foci"], "article_id"=>507646, "categories"=>["Biochemistry", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Ignasi Roig", "James A. Dowdle", "Attila Toth", "Dirk G. de Rooij", "Maria Jasin", "Scott Keeney"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1001062.t003", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Numbers_of_early_recombination_associated_foci_in_spermatocytes_/507646", "title"=>"Numbers of early recombination-associated foci in spermatocytes.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-08-12 02:07:26"}
  • {"files"=>["https://ndownloader.figshare.com/files/836491"], "description"=>"<p>Spread chromosomes of pachytene (or pachytene-like) spermatocytes from the indicated genotypes were stained with antibodies to SYCP3 and to either BRCA1 (A,D,G), ATR (B,E,H), or γH2AX (C,F,I) to monitor sex body formation. Arrowheads point to the sex chromosomes; in (G, H, I) identities of the X and Y are deduced from axis lengths and synaptic status (i.e., fully unsynapsed). Note that many of the synapsed autosomes show abnormal presence of BRCA1 or ATR foci in the <i>Trip13<sup>sev/sev</sup></i> mutant (G, H) and of γH2AX in both mutants (F, I). Bar = 10 µm.</p>", "links"=>[], "tags"=>["mutant"], "article_id"=>506855, "categories"=>["Biochemistry", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Ignasi Roig", "James A. Dowdle", "Attila Toth", "Dirk G. de Rooij", "Maria Jasin", "Scott Keeney"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1001062.g006", "stats"=>{"downloads"=>3, "page_views"=>11, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Sex_body_formation_in_Trip13_mutant_mice_/506855", "title"=>"Sex body formation in <i>Trip13</i> mutant mice.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-08-12 01:54:15"}
  • {"files"=>["https://ndownloader.figshare.com/files/836812"], "description"=>"<p>(A) Spread chromosomes from a <i>Trip13<sup>mod/mod</sup></i> pachytene spermatocyte stained for DNA (DAPI), SYCP3, and MLH1 to mark the positions of COs. Bar = 10 µm. (B) Quantification of the number of autosomal MLH1 foci per pachytene cell. Means are shown as horizontal lines. Results were as follows (mean ± sd): 24.2±3.1 (wild type, six mice); 23.9±2.3 (<i>Trip13<sup>+/mod</sup></i>, seven mice); 22.6±3.2 (<i>Trip13<sup>mod/mod</sup></i>, four mice). (C) Percentages of bivalents in wild-type and <i>Trip13<sup>mod/mod</sup></i> pachytene spermatocytes with the indicated numbers of MLH1 foci. (D) Spread chromosomes from a <i>Trip13<sup>mod/mod</sup></i> diplotene spermatocyte stained for SYCP3 and SYCP1. Note the presence of achiasmate bivalents, some of which are presented in the zoomed images on the right. Bar = 10 µm. (E) Percentages of achiasmate chromosome pairs at diplonema in wild-type and <i>Trip13<sup>mod/mod</sup></i> spermatocytes. (F) Absence of MLH1 foci in a pachytene-like <i>Trip13<sup>sev/sev</sup></i> spermatocyte. Bar = 10 µm. (G) Quantification of MLH1 focus numbers in pachytene or pachytene-like oocytes. Means are indicated by the horizontal line. Results were as follows (mean ± sd): 23.0±3.2 (wild type); 18.2±5.5 (<i>Trip13<sup>sev/sev</sup></i>). (H) Percentages of bivalents with the indicated numbers of MLH1 foci. This analysis provides a conservative estimate of the defect in <i>Trip13<sup>sev/sev</sup></i> oocytes, because only cells with ≥20 total foci were considered in order to minimize possible secondary effects of differences in the timing of progression through meiosis.</p>", "links"=>[], "tags"=>["cos", "chiasmata", "spermatocytes"], "article_id"=>507176, "categories"=>["Biochemistry", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Ignasi Roig", "James A. Dowdle", "Attila Toth", "Dirk G. de Rooij", "Maria Jasin", "Scott Keeney"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1001062.g008", "stats"=>{"downloads"=>1, "page_views"=>10, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Reduced_COs_and_chiasmata_in_Trip13_deficient_spermatocytes_and_oocytes_/507176", "title"=>"Reduced COs and chiasmata in <i>Trip13</i>-deficient spermatocytes and oocytes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-08-12 01:59:36"}
  • {"files"=>["https://ndownloader.figshare.com/files/836262"], "description"=>"<p>Spread chromosomes from a wild-type pachytene oocyte (A) or a <i>Trip13<sup>sev/sev</sup></i> pachytene-like oocyte (B–E) stained for SYCP3 (green) and DNA (DAPI, blue). Arrows indicate examples of unsynapsed regions, zoomed images of which are shown in (C–E). Bar on (A) applies to (A, B) and represents 10 µm. Bar on (C) applies to (C–E) and represents 1 µm.</p>", "links"=>[], "tags"=>["defects", "trip13-deficient"], "article_id"=>506627, "categories"=>["Biochemistry", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Ignasi Roig", "James A. Dowdle", "Attila Toth", "Dirk G. de Rooij", "Maria Jasin", "Scott Keeney"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1001062.g004", "stats"=>{"downloads"=>1, "page_views"=>11, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Synapsis_defects_in_TRIP13_deficient_oocytes_/506627", "title"=>"Synapsis defects in TRIP13-deficient oocytes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-08-12 01:50:27"}
  • {"files"=>["https://ndownloader.figshare.com/files/836381"], "description"=>"<p>Spread chromosomes from pachytene wild-type spermatocytes (A, C) or pachytene-like <i>Trip13<sup>sev/sev</sup></i> spermatocytes (B, D) were stained for SYCP3 (red), SYCP1 (blue), and in green for either HORMAD1 (A, B) or HORMAD2 (C, D). Previous studies demonstrated that HORMAD1 and HORMAD2 show continuous staining of unsynapsed chromosome axes in leptonema and zygonema and become substantially depleted from axes soon after synapsis occurs, and that this synapsis-associated HORMAD depletion does not occur properly in <i>Trip13<sup>mod/mod</sup></i> spermatocytes <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1001062#pgen.1001062-Wojtasz1\" target=\"_blank\">[20]</a>. <i>Trip13<sup>sev/sev</sup></i> mutants show an indistinguishable defect, with both HORMADs remaining at high levels on synapsed axes. Bar in panel A applies to all main panels and represents 10 µm; bar in the inset to panel A applies to all insets and represents 1 µm.</p>", "links"=>[], "tags"=>["hormad", "proteins", "synapsed", "axes", "requires"], "article_id"=>506740, "categories"=>["Biochemistry", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Ignasi Roig", "James A. Dowdle", "Attila Toth", "Dirk G. de Rooij", "Maria Jasin", "Scott Keeney"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1001062.g005", "stats"=>{"downloads"=>1, "page_views"=>37, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Depletion_of_HORMAD_proteins_from_synapsed_axes_requires_TRIP13_/506740", "title"=>"Depletion of HORMAD proteins from synapsed axes requires TRIP13.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-08-12 01:52:20"}
  • {"files"=>["https://ndownloader.figshare.com/files/837078"], "description"=>"<p>Spread pachytene nuclei were stained for SYCP3, MLH1, and DAPI (to detect pericentromeric heterochromatin), and the positions of MLH1 foci were measured relative to centromeres and expressed as percentage of SC length. Because chromosomes of different size show different distributions of MLH1 patterns, data are presented for groups of similarly sized chromosomes, rank ordered as described in the legend to <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1001062#pgen-1001062-g009\" target=\"_blank\">Figure 9</a>. No significant differences were observed between wild type and <i>Trip13<sup>+/mod</sup></i>, so data for these genotypes were pooled for simplicity; separated data are provided in <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1001062#pgen.1001062.s002\" target=\"_blank\">Figure S2</a>. Frequency distribution histograms are shown separately for MLH1 foci on chromosomes with exactly two foci (graphs i–ii in each panel), chromosomes with only one focus (graphs iii–iv), or for all MLH1 foci (graphs v–vi). Chromosomes with three foci were rare and are omitted from this analysis. For the smallest size ranks (D), bivalents with two foci were rare so these were also omitted (n.a., not applicable). Smoothed curves (Gaussian kernel density estimates) are overlaid to facilitate comparison (dark blue, wild type and <i>Trip13<sup>+/mod</sup></i>; red, <i>Trip13<sup>mod/mod</sup></i>). The number of foci in each category is indicated. The central regions (from 40–75% of bivalent length) are bracketed for chromosomes with two foci, and the percentages of foci within these regions are indicated. Telomere-proximal regions (80–95% of bivalent length) and the percentages of foci within them are indicated in panels Biii, Biv, Ciii, Civ. Asterisks indicate values significantly different from wild type (Fisher's exact test, p<0.05).</p>", "links"=>[], "tags"=>["co", "distributions", "autosomes"], "article_id"=>507441, "categories"=>["Biochemistry", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Ignasi Roig", "James A. Dowdle", "Attila Toth", "Dirk G. de Rooij", "Maria Jasin", "Scott Keeney"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1001062.g010", "stats"=>{"downloads"=>0, "page_views"=>6, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Altered_CO_distributions_on_autosomes_in_Trip13_deficient_spermatocytes_/507441", "title"=>"Altered CO distributions on autosomes in <i>Trip13</i>-deficient spermatocytes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-08-12 02:04:01"}
  • {"files"=>["https://ndownloader.figshare.com/files/835823"], "description"=>"<p>(A–C) Representative epithelial stage IV tubules from periodic acid Schiff (PAS)-stained testis sections from mice of the indicated genotypes. Red arrowheads indicate pachytene spermatocytes with an apoptotic morphology (condensed, darkly staining nuclei). Green arrowheads point to examples of postmeiotic cells (elongating spermatids) seen infrequently in <i>Trip13<sup>mod/mod</sup></i> mutants but not observed in <i>Trip13<sup>sev/sev</sup></i> mutants. (D–F) TUNEL-stained testis sections from mice of the indicated genotypes. In wild type (D), few TUNEL-positive cells (stained brown) were seen, most often spermatogonia. In both <i>Trip13</i> mutants (E, F), numerous TUNEL-positive spermatocytes were found. (G–I) PAS-stained ovary sections from 21-day-old mice. In wild type, several follicles at different stages of development can be observed, including primordial follicles (inset in Panel G). In contrast, <i>Trip13<sup>mod/mod</sup></i> ovaries had no primordial follicles (inset in Panel H) and fewer growing follicles (19.3±4.7 per section vs. 75.8±16.2 in wild type). <i>Trip13<sup>sev/sev</sup></i> ovaries completely lacked follicles (panel I). Each bar applies to all images in the row and represents 20 µm.</p>", "links"=>[], "tags"=>["completion", "meiosis", "males"], "article_id"=>506189, "categories"=>["Biochemistry", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Ignasi Roig", "James A. Dowdle", "Attila Toth", "Dirk G. de Rooij", "Maria Jasin", "Scott Keeney"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1001062.g002", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_TRIP13_is_needed_for_completion_of_meiosis_in_males_and_females_/506189", "title"=>"TRIP13 is needed for completion of meiosis in males and females.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-08-12 01:43:09"}
  • {"files"=>["https://ndownloader.figshare.com/files/836720"], "description"=>"<p>(A–C) Representative pachytene and pachytene-like nuclei of the indicated genotypes stained with anti-SYCP3 (green) and an anti-RAD51 antibody that also cross-reacts with DMC1 (red). Arrows indicate the sex chromosomes. Bar in (A) applies to all micrographs and represents 10 µm. (D–F) Representative leptotene nuclei stained for γH2AX (green) and SYCP3 (red). (G–I) Representative leptotene nuclei stained for SYCP3 (green) and RPA (red). (J, K) Quantification of the numbers of RAD51/DMC1 foci (detected with the cross-reacting anti-RAD51 antibody) or DMC1-specific foci per cell at the indicated stages. Horizontal lines denote the means; n.a., not applicable. See <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1001062#pgen-1001062-t003\" target=\"_blank\">Table 3</a> for summary of means, standard deviations, and results of statistical tests. (L) Western blots to determine γH2AX levels in extracts from whole testes from juvenile males at 11.5 dpp. At this age, the majority of the γH2AX is in response to SPO11-generated DSBs; the residual γH2AX in <i>Spo11<sup>–/–</sup></i> mutants is likely from somatic cells or premeiotic germ cells undergoing DNA replication <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1001062#pgen.1001062-Mahadevaiah2\" target=\"_blank\">[27]</a>. For each mutant genotype, testes were also collected from a wild-type littermate. Blots were stripped and reprobed for β-actin as a loading control. (M) Quantification of numbers of RPA foci per cell in wild type, <i>Trip13<sup>mod/mod</sup></i> and <i>Trip13<sup>sev/sev</sup></i> spermatocytes [color code as in (J)].</p>", "links"=>[], "tags"=>["recombination", "defects", "mutant"], "article_id"=>507086, "categories"=>["Biochemistry", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Ignasi Roig", "James A. Dowdle", "Attila Toth", "Dirk G. de Rooij", "Maria Jasin", "Scott Keeney"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1001062.g007", "stats"=>{"downloads"=>0, "page_views"=>6, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Early_and_late_recombination_defects_in_Trip13_mutant_spermatocytes_/507086", "title"=>"Early and late recombination defects in <i>Trip13</i> mutant spermatocytes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-08-12 01:58:06"}
  • {"files"=>["https://ndownloader.figshare.com/files/416352", "https://ndownloader.figshare.com/files/416375"], "description"=>"<div><p>Accurate chromosome segregation during meiosis requires that homologous chromosomes pair and become physically connected so that they can orient properly on the meiosis I spindle. These connections are formed by homologous recombination closely integrated with the development of meiosis-specific, higher-order chromosome structures. The yeast Pch2 protein has emerged as an important factor with roles in both recombination and chromosome structure formation, but recent analysis suggested that TRIP13, the mouse Pch2 ortholog, is not required for the same processes. Using distinct <em>Trip13</em> alleles with moderate and severe impairment of TRIP13 function, we report here that TRIP13 is required for proper synaptonemal complex formation, such that autosomal bivalents in <em>Trip13</em>-deficient meiocytes frequently displayed pericentric synaptic forks and other defects. In males, TRIP13 is required for efficient synapsis of the sex chromosomes and for sex body formation. Furthermore, the numbers of crossovers and chiasmata are reduced in the absence of TRIP13, and their distribution along the chromosomes is altered, suggesting a role for TRIP13 in aspects of crossover formation and/or control. Recombination defects are evident very early in meiotic prophase, soon after DSB formation. These findings provide evidence for evolutionarily conserved functions for TRIP13/Pch2 in both recombination and formation of higher order chromosome structures, and they support the hypothesis that TRIP13/Pch2 participates in coordinating these key aspects of meiotic chromosome behavior.</p></div>", "links"=>[], "tags"=>["recombination", "higher-order", "chromosome", "meiosis"], "article_id"=>142190, "categories"=>["Biochemistry", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Ignasi Roig", "James A. Dowdle", "Attila Toth", "Dirk G. de Rooij", "Maria Jasin", "Scott Keeney"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1001062.s001", "https://dx.doi.org/10.1371/journal.pgen.1001062.s002"], "stats"=>{"downloads"=>2, "page_views"=>14, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Mouse_TRIP13_PCH2_Is_Required_for_Recombination_and_Normal_Higher_Order_Chromosome_Structure_during_Meiosis/142190", "title"=>"Mouse TRIP13/PCH2 Is Required for Recombination and Normal Higher-Order Chromosome Structure during Meiosis", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2010-08-12 00:36:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/837249"], "description"=>"a<p>Irrespective of whether interstitial asynapsis was also present.</p>", "links"=>[], "tags"=>["anomalies", "spermatocytes"], "article_id"=>507612, "categories"=>["Biochemistry", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Ignasi Roig", "James A. Dowdle", "Attila Toth", "Dirk G. de Rooij", "Maria Jasin", "Scott Keeney"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1001062.t002", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Synaptic_anomalies_in_Trip13_sev_sev_spermatocytes_and_oocytes_/507612", "title"=>"Synaptic anomalies in <i>Trip13<sup>sev/sev</sup></i> spermatocytes and oocytes.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-08-12 02:06:52"}
  • {"files"=>["https://ndownloader.figshare.com/files/836054"], "description"=>"<p>(A–K) Progression of synapsis followed by staining spermatocyte chromosome spreads for SYCP3 (green) and SYCP1 (red). <i>Trip13<sup>mod/mod</sup></i> spermatocytes (E–H) show progression of synapsis (leptonema through pachynema) and subsequent desynapsis (diplonema) that is largely comparable to wild type (A–D). In contrast, <i>Trip13<sup>sev/sev</sup></i> spermatocytes form axial elements (I) and initiate synapsis (J) but never achieve complete synapsis of all homologs (K). The cell in (K) is a representative example of the most advanced stage observed in the <i>Trip13<sup>sev/sev</sup></i> mutant and has characteristics of both zygonema and pachynema (see text). Synapsed X and Y chromosomes are indicated in (C–D) and (G–H); they are unsynapsed in (K). Bar on (A) applies to (A–K) and represents 20 µm. (L) Zoomed image of a selected SC with pericentric fork (i.e., an unsynapsed centromeric end, as detected by CREST staining) and interstitial unsynapsis. Bar = 1.5 µm. Arrowheads in (K, L) point to examples of bivalents whose ends have failed to complete synapsis; arrows point to examples of interstitial asynapsis.</p>", "links"=>[], "tags"=>["synapsis"], "article_id"=>506423, "categories"=>["Biochemistry", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Ignasi Roig", "James A. Dowdle", "Attila Toth", "Dirk G. de Rooij", "Maria Jasin", "Scott Keeney"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1001062.g003", "stats"=>{"downloads"=>1, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_TRIP13_is_required_to_complete_synapsis_in_spermatocytes_/506423", "title"=>"TRIP13 is required to complete synapsis in spermatocytes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-08-12 01:47:03"}
  • {"files"=>["https://ndownloader.figshare.com/files/837205"], "description"=>"<p>Pachytene spermatocyte spreads were immunostained for SYCP3 and lengths of autosomal SCs were measured from wild-type and <i>Trip13<sup>mod/mod</sup></i> spermatocytes. The bivalents in each spread were rank-ordered by length from 1 (largest) to 19 (smallest), then divided into groups of similarly sized chromosomes. Note that each of the chromosomes in each size class makes up essentially the same percentage of total SC length in both genotypes, indicating that the decrease in average total SC length in <i>Trip13<sup>mod/mod</sup></i> spermatocytes is uniform across all autosomes.</p>a<p>Mean ± standard deviation for each chromosome in the indicated size rank.</p>b<p>The mean SC length per chromosome divided by the average total SC length per cell, multiplied by 100.</p>c<p>Total number of chromosomes analyzed.</p>d<p>These values are statistically significantly different from those for wild type (p≤0.05, t test).</p>", "links"=>[], "tags"=>["scs", "shorter"], "article_id"=>507576, "categories"=>["Biochemistry", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Ignasi Roig", "James A. Dowdle", "Attila Toth", "Dirk G. de Rooij", "Maria Jasin", "Scott Keeney"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1001062.t001", "stats"=>{"downloads"=>2, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Autosomal_SCs_are_shorter_on_average_in_Trip13_mod_mod_spermatocytes_/507576", "title"=>"Autosomal SCs are shorter on average in <i>Trip13<sup>mod/mod</sup></i> spermatocytes.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-08-12 02:06:16"}
  • {"files"=>["https://ndownloader.figshare.com/files/836946"], "description"=>"<p>(A) Pachytene spermatocyte spreads were immunostained for SYCP3 (shown in red) and MLH1 (shown in green), then the distances between foci were measured on autosomal bivalents that contain two or more MLH1 foci. Examples of relative and corresponding cumulative frequency plots of gamma distributions are shown. If there is no interference between MLH1 foci, an exponential frequency distribution is expected (gray lines). Deviation from exponential behavior indicates the existence of interference (red and blue lines): short and long distances become more rare and the spacing becomes more even (i.e., distances are tightly clustered). Curves were calculated using an average interfocus distance of 10 and the indicated increasing values for the shape parameter, <b>υ</b>. Adapted from ref. <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1001062#pgen.1001062-Barchi1\" target=\"_blank\">[23]</a>. (B–K) Inter-focus distances in wild type and <i>Trip13<sup>mod/mod</sup></i> spermatocytes. (B–F) show the frequency distributions (step plots) of inter-focus distances as a percentage of the SC length for wild type (blue) and <i>Trip13<sup>mod/mod</sup></i> (red). (G–K) show cumulative plots of the same data, with the number of chromosomes analyzed and the Kolmogorov-Smirnov test p value indicated. The left column of graphs (B, G) pools data for all autosomes. For the remaining columns, the autosomal bivalents in each nucleus were rank-ordered by SC length from largest (1) to smallest (19), then divided into groups of similarly sized chromosomes: the two largest chromosomes are shown in (C, H), the next three in (D, I), and so forth. Autosome size ranks 17–19 are excluded from this analysis because they rarely have more than a single MLH1 focus.</p>", "links"=>[], "tags"=>["mlh1", "inter-focus", "distances"], "article_id"=>507314, "categories"=>["Biochemistry", "Molecular Biology", "Genetics", "Developmental Biology"], "users"=>["Ignasi Roig", "James A. Dowdle", "Attila Toth", "Dirk G. de Rooij", "Maria Jasin", "Scott Keeney"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1001062.g009", "stats"=>{"downloads"=>0, "page_views"=>24, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Reduced_MLH1_inter_focus_distances_in_Trip13_mod_mod_spermatocytes_/507314", "title"=>"Reduced MLH1 inter-focus distances in <i>Trip13<sup>mod/mod</sup></i> spermatocytes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-08-12 02:01:54"}

PMC Usage Stats | Further Information

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Relative Metric

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