Sex Determination in Honeybees: Two Separate Mechanisms Induce and Maintain the Female Pathway
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{"title"=>"Sex determination in honeybees: Two separate mechanisms induce and maintain the female pathway", "type"=>"journal", "authors"=>[{"first_name"=>"Tanja", "last_name"=>"Gempe", "scopus_author_id"=>"15047754100"}, {"first_name"=>"Martin", "last_name"=>"Hasselmann", "scopus_author_id"=>"56187651600"}, {"first_name"=>"Morten", "last_name"=>"Schiøtt", "scopus_author_id"=>"24069503500"}, {"first_name"=>"Gerd", "last_name"=>"Hause", "scopus_author_id"=>"6603607913"}, {"first_name"=>"Marianne", "last_name"=>"Otte", "scopus_author_id"=>"8847628700"}, {"first_name"=>"Martin", "last_name"=>"Beye", "scopus_author_id"=>"6603956891"}], "year"=>2009, "source"=>"PLoS Biology", "identifiers"=>{"pmid"=>"19841734", "doi"=>"10.1371/journal.pbio.1000222", "sgr"=>"70350520545", "isbn"=>"1545-7885 (Electronic)\\n1544-9173 (Linking)", "scopus"=>"2-s2.0-70350520545", "issn"=>"15449173", "pui"=>"355524093"}, "id"=>"e644d623-3ef1-30a9-99e3-8d15b020897f", "abstract"=>"Organisms have evolved a bewildering diversity of mechanisms to generate the two sexes. The honeybee (Apis mellifera) employs an interesting system in which sex is determined by heterozygosity at a single locus (the Sex Determination Locus) harbouring the complementary sex determiner (csd) gene. Bees heterozygous at Sex Determination Locus are females, whereas bees homozygous or hemizygous are males. Little is known, however, about the regulation that links sex determination to sexual differentiation. To investigate the control of sexual development in honeybees, we analyzed the functions and the regulatory interactions of genes involved in the sex determination pathway. We show that heterozygous csd is only required to induce the female pathway, while the feminizer (fem) gene maintains this decision throughout development. By RNAi induced knockdown we show that the fem gene is essential for entire female development and that the csd gene exclusively processes the heterozygous state. Fem activity is also required to maintain the female determined pathway throughout development, which we show by mosaic structures in fem-repressed intersexuals. We use expression of Fem protein in males to demonstrate that the female maintenance mechanism is controlled by a positive feedback splicing loop in which Fem proteins mediate their own synthesis by directing female fem mRNA splicing. The csd gene is only necessary to induce this positive feedback loop in early embryogenesis by directing splicing of fem mRNAs. Finally, fem also controls the splicing of Am-doublesex transcripts encoding conserved male- and female-specific transcription factors involved in sexual differentiation. Our findings reveal how the sex determination process is realized in honeybees differing from Drosophila melanogaster.", "link"=>"http://www.mendeley.com/research/sex-determination-honeybees-two-separate-mechanisms-induce-maintain-female-pathway", "reader_count"=>169, "reader_count_by_academic_status"=>{"Unspecified"=>5, "Professor > Associate Professor"=>13, "Researcher"=>43, "Student > Doctoral Student"=>9, "Student > Ph. D. Student"=>34, "Student > Postgraduate"=>7, "Student > Master"=>15, "Other"=>7, "Student > Bachelor"=>20, "Lecturer"=>2, "Lecturer > Senior Lecturer"=>2, "Professor"=>12}, "reader_count_by_user_role"=>{"Unspecified"=>5, "Professor > Associate Professor"=>13, "Researcher"=>43, "Student > Doctoral Student"=>9, "Student > Ph. D. Student"=>34, "Student > Postgraduate"=>7, "Student > Master"=>15, "Other"=>7, "Student > Bachelor"=>20, "Lecturer"=>2, "Lecturer > Senior Lecturer"=>2, "Professor"=>12}, "reader_count_by_subject_area"=>{"Unspecified"=>9, "Environmental Science"=>2, "Biochemistry, Genetics and Molecular Biology"=>25, "Mathematics"=>1, "Agricultural and Biological Sciences"=>122, "Neuroscience"=>2, "Physics and Astronomy"=>3, "Psychology"=>2, "Chemistry"=>1, "Social Sciences"=>1, "Immunology and Microbiology"=>1}, "reader_count_by_subdiscipline"=>{"Neuroscience"=>{"Neuroscience"=>2}, "Chemistry"=>{"Chemistry"=>1}, "Social Sciences"=>{"Social Sciences"=>1}, "Physics and Astronomy"=>{"Physics and Astronomy"=>3}, "Psychology"=>{"Psychology"=>2}, "Immunology and Microbiology"=>{"Immunology and Microbiology"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>122}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>25}, "Mathematics"=>{"Mathematics"=>1}, "Unspecified"=>{"Unspecified"=>9}, "Environmental Science"=>{"Environmental Science"=>2}}, "reader_count_by_country"=>{"Argentina"=>1, "Romania"=>1, "United States"=>3, "Japan"=>2, "United Kingdom"=>2, "India"=>1, "Saudi Arabia"=>1, "Austria"=>1, "Netherlands"=>2, "Sweden"=>2, "Belgium"=>1, "Norway"=>2, "China"=>1, "Brazil"=>2, "Mexico"=>1, "South Africa"=>1, "Italy"=>1, "France"=>1, "Germany"=>3}, "group_count"=>11}

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/879591"], "description"=>"<p>Fragments corresponding to female (A) and male (B) <i>fem</i> mRNAs were independently amplified by RT-PCR and resolved by agarose gel electrophoresis. The weak ∼1,600 bp fragments observed in reactions devoted to amplify the female-specific fragment correspond to the male mRNAs. Differences in the amount of cDNAs in the different samples were adjusted prior to PCR amplifications. For the embryonic stages the hours after egg deposition are indicated. The early blastoderm is formed ∼12 h after egg deposition. L1 and L5 are 1<sup>st</sup> and 5<sup>th</sup> instar larvae, P2 are pupae at medium stage.</p>", "links"=>[], "tags"=>["mrna"], "article_id"=>550048, "categories"=>["Genetics", "Evolutionary Biology"], "users"=>["Tanja Gempe", "Martin Hasselmann", "Morten Schiøtt", "Gerd Hause", "Marianne Otte", "Martin Beye"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000222.g005", "stats"=>{"downloads"=>2, "page_views"=>11, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Developmental_profile_of_fem_mRNA_expression_/550048", "title"=>"Developmental profile of <i>fem</i> mRNA expression.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2009-10-20 00:00:48"}
  • {"files"=>["https://ndownloader.figshare.com/files/880015"], "description"=>"<p>n.p., not performed.</p>", "links"=>[], "tags"=>["instar", "larvae", "treated", "dsrnas"], "article_id"=>550465, "categories"=>["Genetics", "Evolutionary Biology"], "users"=>["Tanja Gempe", "Martin Hasselmann", "Morten Schiøtt", "Gerd Hause", "Marianne Otte", "Martin Beye"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000222.t001", "stats"=>{"downloads"=>1, "page_views"=>7, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Gonad_development_of_5_th_instar_larvae_treated_with_dsRNAs_and_siRNAs_/550465", "title"=>"Gonad development of 5<sup>th</sup> instar larvae treated with dsRNAs and siRNAs.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2009-10-20 00:07:45"}
  • {"files"=>["https://ndownloader.figshare.com/files/435731"], "description"=>"<div><p>Organisms have evolved a bewildering diversity of mechanisms to generate the two sexes. The honeybee (<em>Apis mellifera</em>) employs an interesting system in which sex is determined by heterozygosity at a single locus (the Sex Determination Locus) harbouring the <em>complementary sex determiner</em> (<em>csd</em>) gene. Bees heterozygous at Sex Determination Locus are females, whereas bees homozygous or hemizygous are males. Little is known, however, about the regulation that links sex determination to sexual differentiation. To investigate the control of sexual development in honeybees, we analyzed the functions and the regulatory interactions of genes involved in the sex determination pathway. We show that heterozygous <em>csd</em> is only required to induce the female pathway, while the <em>feminizer</em> (<em>fem</em>) gene maintains this decision throughout development. By RNAi induced knockdown we show that the <em>fem</em> gene is essential for entire female development and that the <em>csd</em> gene exclusively processes the heterozygous state. <em>Fem</em> activity is also required to maintain the female determined pathway throughout development, which we show by mosaic structures in <em>fem</em>-repressed intersexuals. We use expression of Fem protein in males to demonstrate that the female maintenance mechanism is controlled by a positive feedback splicing loop in which Fem proteins mediate their own synthesis by directing female <em>fem</em> mRNA splicing. The <em>csd</em> gene is only necessary to induce this positive feedback loop in early embryogenesis by directing splicing of <em>fem</em> mRNAs. Finally, <em>fem</em> also controls the splicing of <em>Am-doublesex</em> transcripts encoding conserved male- and female-specific transcription factors involved in sexual differentiation. Our findings reveal how the sex determination process is realized in honeybees differing from <em>Drosophila melanogaster</em>.</p></div>", "links"=>[], "tags"=>["mechanisms", "induce", "pathway"], "article_id"=>145925, "categories"=>["Genetics", "Evolutionary Biology"], "users"=>["Tanja Gempe", "Martin Hasselmann", "Morten Schiøtt", "Gerd Hause", "Marianne Otte", "Martin Beye"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000222", "stats"=>{"downloads"=>6, "page_views"=>15, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Sex_Determination_in_Honeybees_Two_Separate_Mechanisms_Induce_and_Maintain_the_Female_Pathway/145925", "title"=>"Sex Determination in Honeybees: Two Separate Mechanisms Induce and Maintain the Female Pathway", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2009-10-20 01:38:45"}
  • {"files"=>["https://ndownloader.figshare.com/files/879824"], "description"=>"<p>(A) Model for the honeybee sex determination pathway that controls both soma and germ cells. The heterozygous or homo-/hemizygous state of the <i>csd</i> gene determines whether Csd protein is active. Active Csd proteins, derived from different <i>csd</i> alleles in females, are splicing factors that direct the processing into female <i>fem</i> mRNAs. Female <i>fem</i> mRNAs (<i>fem<sup>F</sup></i>) are producing active Fem proteins that are required to mediate the splicing of <i>Am-dsx</i> pre-mRNA into the female mRNAs. The Fem protein has an additional positive feedback activity that directs the processing of <i>fem<sup>F</sup></i> mRNAs. Inactive CSD proteins, when derived from homo- or hemizygous <i>csd</i> alleles, result in a splicing of the <i>fem</i> and <i>dsx</i> transcripts, which is the default male state (<i>fem<sup>M</sup></i>, <i>Am-dsx<sup>M</sup></i>). (B) Model for the sex determination pathway in <i>Ceratitis capitata </i><a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1000222#pbio.1000222-Salvemini1\" target=\"_blank\">[37]</a>,<a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1000222#pbio.1000222-Pane1\" target=\"_blank\">[38]</a>. The presence or absence of an unidentified factor <i>M</i> determines sex. In the absence of <i>M</i> the maternal provided <i>Cc-tra</i> gene product establishes an autoregulative loop in which Cc-Tra protein mediates the production of female <i>Cc-tra</i> mRNA. The Cc-Tra protein directs the splicing of <i>Cc-dsx</i> pre-mRNA into the female mode. The presence of <i>M</i> impairs the positive autoregulative loop of the <i>Cc-tra</i> gene products producing a default splicing pattern of <i>Cc-tra</i> transcripts, the male pre-mRNA. The male <i>Cc-dsx</i> mRNA is produced by default. (C) Simplified view of the somatic sex determination hierarchy in <i>D. melanogaster </i><a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1000222#pbio.1000222-Cline1\" target=\"_blank\">[22]</a>. The X∶A ratio determines whether <i>Sxl</i> is activated. Sxl protein in females is a splicing factor that directs the splicing of <i>tra</i> pre-mRNA into the female mode, resulting in the production of active Tra protein in females. Tra protein mediates the processing of female <i>dsx</i> mRNAs. In the absence of Sxl protein all these regulatory decisions do not occur and the male <i>dsx<sup>M</sup></i> is produced by default. The male and female <i>dsx</i> transcripts encode sex-specific transcription factors that have several target genes and are involved in various aspects of sexual differentiation. (D) The evolutionary relationship of the species used in the comparison with their approximate time scale of divergence.</p>", "links"=>[], "tags"=>["regulative", "hierarchy", "honeybee"], "article_id"=>550280, "categories"=>["Genetics", "Evolutionary Biology"], "users"=>["Tanja Gempe", "Martin Hasselmann", "Morten Schiøtt", "Gerd Hause", "Marianne Otte", "Martin Beye"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000222.g007", "stats"=>{"downloads"=>2, "page_views"=>119, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_regulative_hierarchy_of_honeybee_sex_determination_in_relation_to_other_insect_model_species_/550280", "title"=>"The regulative hierarchy of honeybee sex determination in relation to other insect model species.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2009-10-20 00:04:40"}
  • {"files"=>["https://ndownloader.figshare.com/files/879350"], "description"=>"<p>(A–F) Development of the reproductive organ. (A) Normal pair of ovaries, oviducts (od), and unpaired vagina (va) of an untreated female (worker bee). The ovaries are composed of less than five ovarioles (ovl). (B) Normally developed pairs of testes, spermducts (sd), mucus glands (mg), and unpaired endophallus (ep) of a diploid male. The testes consist of hundreds of thickly packed and folded testioles (tl). (C) Male reproductive organ from a female injected with <i>fem</i> siRNAs. The testes are reduced in size and composed of fewer testioles of reduced length. (E) Male reproductive organ from a female treated with <i>csd</i> siRNAs. The testes from these pseudomales are of normal size and structure and appear equivalent to the testes from diploid males (B). (D and F) Normally differentiated reproductive organ from a male treated with <i>fem</i> or <i>csd</i> siRNAs, respectively. Reproductive apparatus was stained with aceto-orcein (reddish colouring), which facilitated the dissection process. Scale bars, 1 mm. (G–L) Differentiation of germ cells in microscopic sections through ovarioles and testicular tubules. (G) Undifferentiated cells in an ovariole of an untreated female. The ovariole is surrounded by an epithelial sheath (white arrowhead). (H) Bundles of spermatids in a testicular tubule (testioles) of a non-injected diploid male. The testicular tubules are composed of spermatocystes containing the spermatids (black arrow) and nurse cells (black arrowhead). (I) Spermatids formed in a fully male differentiated testis of a <i>fem</i> siRNA injected female. (K) Spermatids in a fully male-like developed testis from a <i>csd</i> siRNA treated female. (J and L) Normal testis and germ cell differentiation of <i>fem</i> (J) and <i>csd</i> (L) siRNA injected diploid males. Sections were stained with toluidine blue. Scale bars, 10 µm. (M–S) Development of the inner tibia and tarsus surface of the left hind leg. (M) Normally differentiated pollen brush, pollen comb, and lobe of an untreated female worker. The first tarsal segment displays symmetrically arranged rows of bristles, which are used to brush pollen from the body surface (pollen brush, pb). The upper posterior part of the first tarsal segment forms a lobe (black arrow). Spines at the distal part of the tibia form the pollen comb (grey arrow) in which pollen is detached from the pollen brush. (N) Normally developed tibia and first tarsal segment of non-injected diploid males that lack the symmetrical organization of bristles (pollen brush), the lobe, and the pollen comb. (O) Male differentiated hind leg from a female injected with <i>fem</i> siRNAs. (P) Development of a mosaic intersex upon <i>fem</i> siRNA injections. The posterior part of the first tarsus segment is male, lacks the female-specific lobe (black arrow), and displays bristles in a non-arranged pattern. The anterior part of the tarsal segment is female and shows the symmetrical arrays of bristles. The distal part of the tibia harbours the spines of the pollen comb (grey arrow) indicating a fully developed female structure. (R) Male developed hind leg from a female treated with <i>csd</i> siRNAs. (Q and S) Normally developed hind legs from <i>fem</i> (Q) and <i>csd</i> (S) siRNA injected diploid males. Scale bars, 1 mm.</p>", "links"=>[], "tags"=>["germ", "diploid", "pupae", "knockdown"], "article_id"=>549802, "categories"=>["Genetics", "Evolutionary Biology"], "users"=>["Tanja Gempe", "Martin Hasselmann", "Morten Schiøtt", "Gerd Hause", "Marianne Otte", "Martin Beye"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000222.g003", "stats"=>{"downloads"=>2, "page_views"=>14, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Soma_and_germ_line_development_of_female_and_diploid_male_late_pupae_in_the_knockdown_analysis_of_the_fem_and_the_csd_gene_/549802", "title"=>"Soma and germ line development of female and diploid male late pupae in the knockdown analysis of the <i>fem</i> and the <i>csd</i> gene.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2009-10-20 02:43:22"}
  • {"files"=>["https://ndownloader.figshare.com/files/879228"], "description"=>"<p>(A–C) Reproductive organ development of untreated individuals: (A) A pair of normally developed ovaries (ov) and oviducts (od) from an untreated female. (B) A pair of normally differentiated testes from untreated haploid males consisting of densely packed layers of folded testioles. The paired spermducts are not shown. (C) A pair of normally differentiated testes from untreated diploid males consisting of less densely packed layers of folded testioles. The paired spermducts are not shown. (D–G) Repression analysis of gene <i>GB11211</i> and <i>GB13727</i>. Normally developed gonads of females and haploid males injected with dsRNA devoted to repress the function of gene <i>GB11211</i> (D–E) and <i>GB13727</i> (F–G). (H–I) Repression analysis of the <i>fem</i> gene. (H) Pair of underdeveloped testes from a female treated with <i>fem</i> siRNA. The testes of this female individual are covered with oversized epithelial sheaths. The testioles are reduced in length and number when compared with the haploid (B) or diploid (C) males or the pseudomales after <i>csd</i> siRNA injection (J). The shape and course of spermducts appear normal. (I) Normally developed testes from a haploid male injected with <i>fem</i> siRNAs. (J–K) Repression analysis of the <i>csd</i> gene. (J) Pair of fully developed testes from a female treated with <i>csd</i> siRNAs. The number, length, and arrangement of testioles resemble entirely of those dissected from diploid males (C). (K) Normally developed testes from a haploid male injected with <i>csd</i> siRNA. (L–M) Repression analysis of the <i>GB30480</i> gene. Normally developed gonads of females (L) and haploid males (M) injected with <i>GB30480</i> dsRNA. Gonads were stained with aceto-orcein (reddish colouring of gonads), which facilitated the dissection process. Scale bars, 1 mm.</p>", "links"=>[], "tags"=>["instar", "larvae", "repression", "sdl"], "article_id"=>549682, "categories"=>["Genetics", "Evolutionary Biology"], "users"=>["Tanja Gempe", "Martin Hasselmann", "Morten Schiøtt", "Gerd Hause", "Marianne Otte", "Martin Beye"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000222.g002", "stats"=>{"downloads"=>1, "page_views"=>11, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Reproductive_organ_development_of_5_th_instar_male_and_female_larvae_in_the_repression_analysis_of_SDL_genes_/549682", "title"=>"Reproductive organ development of 5<sup>th</sup> instar male and female larvae in the repression analysis of SDL genes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2009-10-20 02:41:22"}
  • {"files"=>["https://ndownloader.figshare.com/files/879097"], "description"=>"<p>(A) Diagram of genes within the SDL, which is always heterozygous in females as deduced by high resolution genetic mapping <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1000222#pbio.1000222-Hasselmann3\" target=\"_blank\">[19]</a>,<a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1000222#pbio.1000222-Hasselmann4\" target=\"_blank\">[24]</a>. Genes are orientated 5′ to 3′ according to the direction of arrows; the names of functionally characterized genes are underlined. <i>GB30480</i> corresponds to gene Ex4.8–5.8 <a href=\"http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.1000222#pbio.1000222-Beye3\" target=\"_blank\">[16]</a>. (B) Exon and intron structure diagram of genes encoded at SDL. Exons are shown as boxes and introns by connecting lines. The deduced open reading frames are marked in grey and the presumed start and stop codons are indicated.</p>", "links"=>[], "tags"=>["mrna", "producing", "genes"], "article_id"=>549551, "categories"=>["Genetics", "Evolutionary Biology"], "users"=>["Tanja Gempe", "Martin Hasselmann", "Morten Schiøtt", "Gerd Hause", "Marianne Otte", "Martin Beye"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000222.g001", "stats"=>{"downloads"=>1, "page_views"=>6, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Genomic_organization_of_mRNA_producing_genes_of_the_SDL_/549551", "title"=>"Genomic organization of mRNA producing genes of the SDL.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2009-10-20 02:39:11"}
  • {"files"=>["https://ndownloader.figshare.com/files/879453"], "description"=>"<p>(A) The male and female <i>fem</i> and <i>Am-dsx</i> mRNAs of eight 5<sup>th</sup> larval instar pseudomales that have been injected with <i>csd</i> siRNAs. Fragments corresponding to the <i>fem</i> female (∼350 bp) and male (∼1.6 kb) mRNAs and the <i>Am-dsx</i> female (∼1.4 kb) and male (∼500 bp) mRNAs were amplified by RT-PCR and resolved by agarose gel electrophoresis. The fragments obtained from untreated females and males are shown in the 1<sup>st</sup> and 2<sup>nd</sup> lane, respectively. (B) Same analysis as in (A) except that eight pseudomales have been treated with <i>fem</i> instead of <i>csd</i> siRNAs.</p>", "links"=>[], "tags"=>["transcripts", "knockdowns", "induced"], "article_id"=>549906, "categories"=>["Genetics", "Evolutionary Biology"], "users"=>["Tanja Gempe", "Martin Hasselmann", "Morten Schiøtt", "Gerd Hause", "Marianne Otte", "Martin Beye"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000222.g004", "stats"=>{"downloads"=>5, "page_views"=>11, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_processing_of_fem_and_Am_dsx_transcripts_in_the_response_to_the_knockdowns_of_the_csd_and_the_fem_gene_induced_by_RNAi_/549906", "title"=>"The processing of <i>fem</i> and <i>Am-dsx</i> transcripts in the response to the knockdowns of the <i>csd</i> and the <i>fem</i> gene induced by RNAi.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2009-10-20 02:45:06"}
  • {"files"=>["https://ndownloader.figshare.com/files/879720"], "description"=>"<p>(A) Fragments corresponding to the female <i>fem</i> mRNAs of individual 72-h-old embryos were amplified by RT-PCR and resolved by agarose gel electrophoresis. The identity of the female fragments was confirmed by nucleotide sequence analysis. The last lane shows the reactions in which the (pl) <i>fem</i><sup>csd-UTR</sup> mRNA encoding plasmid (pfem<sup>csd-UTR</sup>) was used as a template. The absence of a fragment in this high copy DNA control strongly suggests that our primer oligonucleotides will not amplify fragments corresponding to the injected Fem encoding mRNAs (<i>fem</i><sup>csd-UTR</sup> mRNA). (B) Amplified fragments corresponding to the male <i>fem</i> mRNAs on the same set of samples as described in (A).</p>", "links"=>[], "tags"=>["endogenous", "transcripts", "injection", "fem", "encoding", "mrna", "haploid"], "article_id"=>550172, "categories"=>["Genetics", "Evolutionary Biology"], "users"=>["Tanja Gempe", "Martin Hasselmann", "Morten Schiøtt", "Gerd Hause", "Marianne Otte", "Martin Beye"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000222.g006", "stats"=>{"downloads"=>1, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_processing_of_endogenous_fem_transcripts_in_response_to_the_injection_of_Fem_encoding_mRNA_in_haploid_males_/550172", "title"=>"The processing of endogenous <i>fem</i> transcripts in response to the injection of Fem encoding mRNA in haploid males.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2009-10-20 00:02:52"}
  • {"files"=>["https://ndownloader.figshare.com/files/879974"], "description"=>"<p>Sexual development of late pupae (P3) treated with siRNAs.</p>", "links"=>[], "tags"=>["pupae", "treated"], "article_id"=>550430, "categories"=>["Genetics", "Evolutionary Biology"], "users"=>["Tanja Gempe", "Martin Hasselmann", "Morten Schiøtt", "Gerd Hause", "Marianne Otte", "Martin Beye"], "doi"=>"https://dx.doi.org/10.1371/journal.pbio.1000222.t002", "stats"=>{"downloads"=>4, "page_views"=>7, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Sexual_development_of_late_pupae_P3_treated_with_siRNAs_/550430", "title"=>"Sexual development of late pupae (P3) treated with siRNAs.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2009-10-20 00:07:10"}

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Relative Metric

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