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{"title"=>"A New Component of the Nasonia Sex Determining Cascade Is Maternally Silenced and Regulates Transformer Expression", "type"=>"journal", "authors"=>[{"first_name"=>"Eveline C.", "last_name"=>"Verhulst", "scopus_author_id"=>"35076721400"}, {"first_name"=>"Jeremy A.", "last_name"=>"Lynch", "scopus_author_id"=>"7403674356"}, {"first_name"=>"Daniel", "last_name"=>"Bopp", "scopus_author_id"=>"7003565790"}, {"first_name"=>"Leo W.", "last_name"=>"Beukeboom", "scopus_author_id"=>"7003336203"}, {"first_name"=>"Louis", "last_name"=>"van de Zande", "scopus_author_id"=>"6701442824"}], "year"=>2013, "source"=>"PLoS ONE", "identifiers"=>{"sgr"=>"84878093786", "doi"=>"10.1371/journal.pone.0063618", "pui"=>"368973452", "pmid"=>"23717455", "scopus"=>"2-s2.0-84878093786", "issn"=>"19326203", "isbn"=>"1932-6203 (Electronic)\\r1932-6203 (Linking)"}, "id"=>"f3239a6a-dab1-3ffd-bcf9-181abecb4b11", "abstract"=>"Although sex determination is a universal process in sexually reproducing organisms, sex determination pathways are among the most highly variable genetic systems found in nature. Nevertheless, general principles can be identified among the diversity, like the central role of transformer (tra) in insects. When a functional TRA protein is produced in early embryogenesis, the female sex determining route is activated, while prevention of TRA production leads to male development. In dipterans, male development is achieved by prevention of female-specific splicing of tra mRNA, either mediated by X-chromosome dose or masculinizing factors. In Hymenoptera, which have haplodiploid sex determination, complementary sex determination and maternal imprinting have been identified to regulate timely TRA production. In the parasitoid Nasonia, zygotic transformer (Nvtra) expression and splicing is regulated by a combination of maternal provision of Nvtra mRNA and silencing of Nvtra expression in unfertilized eggs. It is unclear, however, if this silencing is directly on the tra locus or whether it is mediated through maternal silencing of a trans-acting factor. Here we show that in Nasonia, female sex determination is dependent on zygotic activation of Nvtra expression by an as yet unknown factor. This factor, which we propose to term womanizer (wom), is maternally silenced during oogenesis to ensure male development in unfertilized eggs. This finding implicates the upstream recruitment of a novel gene in the Nasonia sex determining cascade and supports the notion that sex determining cascades can rapidly change by adding new components on top of existing regulators.", "link"=>"http://www.mendeley.com/research/new-component-nasonia-sex-determining-cascade-maternally-silenced-regulates-transformer-expression", "reader_count"=>45, "reader_count_by_academic_status"=>{"Unspecified"=>1, "Professor > Associate Professor"=>3, "Researcher"=>7, "Student > Doctoral Student"=>3, "Student > Ph. D. Student"=>13, "Student > Postgraduate"=>1, "Student > Master"=>7, "Student > Bachelor"=>4, "Lecturer"=>1, "Lecturer > Senior Lecturer"=>1, "Professor"=>4}, "reader_count_by_user_role"=>{"Unspecified"=>1, "Professor > Associate Professor"=>3, "Researcher"=>7, "Student > Doctoral Student"=>3, "Student > Ph. D. Student"=>13, "Student > Postgraduate"=>1, "Student > Master"=>7, "Student > Bachelor"=>4, "Lecturer"=>1, "Lecturer > Senior Lecturer"=>1, "Professor"=>4}, "reader_count_by_subject_area"=>{"Unspecified"=>2, "Environmental Science"=>1, "Biochemistry, Genetics and Molecular Biology"=>7, "Agricultural and Biological Sciences"=>34, "Physics and Astronomy"=>1}, "reader_count_by_subdiscipline"=>{"Physics and Astronomy"=>{"Physics and Astronomy"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>34}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>7}, "Unspecified"=>{"Unspecified"=>2}, "Environmental Science"=>{"Environmental Science"=>1}}, "reader_count_by_country"=>{"Argentina"=>1, "Netherlands"=>1, "United Kingdom"=>1}, "group_count"=>2}

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• http://doi.org/10.1371/journal.pone.0114191
• http://doi.org/10.1111/imb.12663
• http://doi.org/10.1111/imb.12333
• http://doi.org/10.1126/science.abb8949
• http://doi.org/10.1073/pnas.1510338112
• http://doi.org/10.1002/ece3.6422
• http://doi.org/10.1126/sciadv.aaz9808
• http://doi.org/10.1111/eea.12808
• http://doi.org/10.7717/peerj.2316
• http://doi.org/10.1111/imb.12352
• http://doi.org/10.1186/s12862-015-0370-9
• http://doi.org/10.1093/jisesa/iex031
• http://doi.org/10.1186/s12863-016-0342-0
• http://doi.org/10.1371/journal.pgen.1008518
• http://doi.org/10.1534/genetics.112.147512
• http://doi.org/10.1186/s12862-015-0340-2
• http://doi.org/10.1016/j.ibmb.2017.09.007
• http://doi.org/10.1111/imb.12522

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Figshare

• {"files"=>["https://ndownloader.figshare.com/files/1548735"], "description"=>"<p>Amplification curves of all samples to determine the exact amount of cycles at which all samples would amplify exponentially. The black square indicates the cycles at which all samples are amplifying exponentially. Each line represents one sample.</p>", "links"=>[], "tags"=>["developmental biology", "embryology", "Evolutionary developmental biology", "Molecular development", "Organism development", "Evolutionary biology", "Evolutionary genetics", "genetics", "epigenetics", "Genomic imprinting", "Molecular genetics", "Gene identification and analysis", "Gene regulation", "gene expression", "Gene networks", "Gene splicing"], "article_id"=>1069085, "categories"=>["Biological Sciences"], "users"=>["Eveline C. Verhulst", "Jeremy A. Lynch", "Daniel Bopp", "Leo W. Beukeboom", "Louis van de Zande"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0063618.g001"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Exponential_phase_of_amplification_/1069085", "title"=>"Exponential phase of amplification.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-05-22 07:57:27"}
• {"files"=>["https://ndownloader.figshare.com/files/1548747"], "description"=>"<p>Panels A – D and E – H indicate the four replicate PCR runs. The maternal origin and age of the embryos is indicated above each set of PCR fragments. White arrow indicates cDNA fragment, black arrow indicates Russia Bait cDNA fragment and dotted arrow indicates gDNA fragment. M is 100 bp molecular standard, ranging from 300 bp (lowest band) to 500 bp (higest band).</p>", "links"=>[], "tags"=>["developmental biology", "embryology", "Evolutionary developmental biology", "Molecular development", "Organism development", "Evolutionary biology", "Evolutionary genetics", "genetics", "epigenetics", "Genomic imprinting", "Molecular genetics", "Gene identification and analysis", "Gene regulation", "gene expression", "Gene networks", "Gene splicing", "pcr", "cdna", "derived", "maternal", "paternal"], "article_id"=>1069097, "categories"=>["Biological Sciences"], "users"=>["Eveline C. Verhulst", "Jeremy A. Lynch", "Daniel Bopp", "Leo W. Beukeboom", "Louis van de Zande"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0063618.g002"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Colony_PCR_showing_the_cDNA_derived_maternal_and_paternal_fragment_of_Nvtra_/1069097", "title"=>"Colony PCR showing the cDNA derived maternal and paternal fragment of <i>Nvtra</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-05-22 07:57:27"}
• {"files"=>["https://ndownloader.figshare.com/files/1548753"], "description"=>"<p>Nuclear spots of <i>Nvtra</i> (green) and <i>Nvcad</i> (red) reveal that there is no apparent imprinting of either the <i>Nvcad</i> or <i>Nvtra</i> locus in early (A) or late (B) blastoderm stage female embryos. In male embryos (C,D) there is moderately less <i>Nvtra</i> expression in early blastoderm stage (C), and significantly less in late blastoderm stage (D).</p>", "links"=>[], "tags"=>["developmental biology", "embryology", "Evolutionary developmental biology", "Molecular development", "Organism development", "Evolutionary biology", "Evolutionary genetics", "genetics", "epigenetics", "Genomic imprinting", "Molecular genetics", "Gene identification and analysis", "Gene regulation", "gene expression", "Gene networks", "Gene splicing", "transcription", "compared"], "article_id"=>1069102, "categories"=>["Biological Sciences"], "users"=>["Eveline C. Verhulst", "Jeremy A. Lynch", "Daniel Bopp", "Leo W. Beukeboom", "Louis van de Zande"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0063618.g003"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Nascent_transcription_of_Nvtra_compared_to_that_of_Nvcad_/1069102", "title"=>"Nascent transcription of <i>Nvtra</i> compared to that of <i>Nvcad</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-05-22 07:57:27"}
• {"files"=>["https://ndownloader.figshare.com/files/1548756"], "description"=>"<p>Nuclei containing only a <i>Nvcad</i> spot are included. Distribution of <i>Nvtra</i> (black bars) or <i>Nvcad</i> (grey bars) spots in 10–11 cycle female embryos are equal indicating that <i>Nvtra</i> and <i>Nvcad</i> expression are similar (A). Some variation in the amount of spots is seen which is due to the nuclear division at that stage. In cycle 12 nuclear division is complete (B) and both <i>Nvtra</i> and <i>Nvcad</i> show primarily diploid expression. In 10–11 cycle male embryos (C) this variation due to nuclear division is also present and after division in cycle 12 <i>Nvtra</i> shows haploid expression similar to <i>Nvcad</i> (D). In male embryos (C, D) the number of nuclei without a <i>Nvtra</i> spot is higher than in female embryos (A, B).</p>", "links"=>[], "tags"=>["developmental biology", "embryology", "Evolutionary developmental biology", "Molecular development", "Organism development", "Evolutionary biology", "Evolutionary genetics", "genetics", "epigenetics", "Genomic imprinting", "Molecular genetics", "Gene identification and analysis", "Gene regulation", "gene expression", "Gene networks", "Gene splicing", "spots", "counted", "nuclei"], "article_id"=>1069106, "categories"=>["Biological Sciences"], "users"=>["Eveline C. Verhulst", "Jeremy A. Lynch", "Daniel Bopp", "Leo W. Beukeboom", "Louis van de Zande"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0063618.g004"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Number_of_spots_of_Nvtra_and_Nvcad_counted_in_a_given_number_of_nuclei_/1069106", "title"=>"Number of spots of <i>Nvtra</i> and <i>Nvcad</i> counted in a given number of nuclei (%).", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-05-22 07:57:27"}
• {"files"=>["https://ndownloader.figshare.com/files/1548758"], "description"=>"<p>The <i>doublesex</i> (<i>dsx</i>) and <i>transformer</i> (<i>tra</i>) gene are conserved, but the upstream signal differs between the species (reviewed in <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063618#pone.0063618-Verhulst2\" target=\"_blank\">[12]</a>). <i>Drosophila</i> has <i>Sexlethal</i> incorporated upstream of <i>tra</i> that splices <i>tra</i> and is regulated by the X chromosome dose <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063618#pone.0063618-Erickson1\" target=\"_blank\">[3]</a>. Other Diptera (e.g. <i>C. capitata</i>, <i>M. domestica</i>) have a M-factor usually on the Y-chromosome that blocks <i>tra</i> auto regulation <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063618#pone.0063618-Pane1\" target=\"_blank\">[4]</a>, <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063618#pone.0063618-Hediger1\" target=\"_blank\">[10]</a>. <i>A. mellifera</i> has a <i>complementary sex determiner</i> (<i>csd</i>) that splices <i>fem </i><a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063618#pone.0063618-Gempe1\" target=\"_blank\">[21]</a> and <i>Nasonia</i> has <i>womanizer</i> (<i>wom</i>) that regulates <i>tra</i> expression. In many insect species, like the dipteran <i>M. domestica</i>, <i>Anastrepha obliqua</i>, <i>C. capitata</i>, <i>Lucilia cuprina</i>, and <i>Sciara ocellaris </i><a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063618#pone.0063618-Concha1\" target=\"_blank\">[13]</a>, <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063618#pone.0063618-Burghardt1\" target=\"_blank\">[35]</a>–<a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063618#pone.0063618-Martn1\" target=\"_blank\">[38]</a>, the hymenopteran insect Apis mellifera <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063618#pone.0063618-Nissen1\" target=\"_blank\">[39]</a> as well as in the lepidopteran insect <i>Bombyx mori </i><a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063618#pone.0063618-Niu1\" target=\"_blank\">[40]</a>, the <i>transformer-2</i> gene is described to be needed for proper <i>tra</i> functioning. * indicates that the ortholog of <i>transformer</i> is called <i>feminizer</i> (<i>fem</i>) in <i>A. mellifera</i>.</p>", "links"=>[], "tags"=>["developmental biology", "embryology", "Evolutionary developmental biology", "Molecular development", "Organism development", "Evolutionary biology", "Evolutionary genetics", "genetics", "epigenetics", "Genomic imprinting", "Molecular genetics", "Gene identification and analysis", "Gene regulation", "gene expression", "Gene networks", "Gene splicing", "determining", "mechanisms", "Diptera"], "article_id"=>1069108, "categories"=>["Biological Sciences"], "users"=>["Eveline C. Verhulst", "Jeremy A. Lynch", "Daniel Bopp", "Leo W. Beukeboom", "Louis van de Zande"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0063618.g005"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Overview_of_the_different_sex_determining_mechanisms_in_Diptera_and_Hymenoptera_/1069108", "title"=>"Overview of the different sex determining mechanisms in Diptera and Hymenoptera.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2013-05-22 07:57:27"}
• {"files"=>["https://ndownloader.figshare.com/files/1548759"], "description"=>"<p>Quantification of nuclear spots of <i>Nvtra</i> and <i>Nvcad</i> in early and late, male and female embryos. In this table, nuclei containing only a <i>Nvcad</i> spot are included.</p>", "links"=>[], "tags"=>["developmental biology", "embryology", "Evolutionary developmental biology", "Molecular development", "Organism development", "Evolutionary biology", "Evolutionary genetics", "genetics", "epigenetics", "Genomic imprinting", "Molecular genetics", "Gene identification and analysis", "Gene regulation", "gene expression", "Gene networks", "Gene splicing"], "article_id"=>1069109, "categories"=>["Biological Sciences"], "users"=>["Eveline C. Verhulst", "Jeremy A. Lynch", "Daniel Bopp", "Leo W. Beukeboom", "Louis van de Zande"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0063618.t002"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Quantification_of_nuclear_spots_/1069109", "title"=>"Quantification of nuclear spots.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2013-05-22 07:57:27"}
• {"files"=>["https://ndownloader.figshare.com/files/1548760"], "description"=>"<p>Number of Russia alleles (R) and alleles (S) in the seven hours old embryos samples. This number is based on the colony PCR (<a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0063618#pone-0063618-g002\" target=\"_blank\">Figure 2</a>). female were mated to Russia Bait males and Russia Bait females were mated to males.</p>", "links"=>[], "tags"=>["developmental biology", "embryology", "Evolutionary developmental biology", "Molecular development", "Organism development", "Evolutionary biology", "Evolutionary genetics", "genetics", "epigenetics", "Genomic imprinting", "Molecular genetics", "Gene identification and analysis", "Gene regulation", "gene expression", "Gene networks", "Gene splicing", "maternal", "paternal", "alleles", "hours"], "article_id"=>1069110, "categories"=>["Biological Sciences"], "users"=>["Eveline C. Verhulst", "Jeremy A. Lynch", "Daniel Bopp", "Leo W. Beukeboom", "Louis van de Zande"], "doi"=>["https://dx.doi.org/10.1371/journal.pone.0063618.t001"], "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Number_of_maternal_and_paternal_alleles_in_seven_hours_old_embryos_/1069110", "title"=>"Number of maternal and paternal alleles in seven hours old embryos.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2013-05-22 07:57:27"}

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Relative Metric

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F1000Prime