Mutations in fam20b and xylt1 Reveal That Cartilage Matrix Controls Timing of Endochondral Ossification by Inhibiting Chondrocyte Maturation
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{"title"=>"Mutations in fam20b and xylt1 reveal that cartilage matrix controls timing of endochondral ossification by inhibiting chondrocyte maturation", "type"=>"journal", "authors"=>[{"first_name"=>"B. Frank", "last_name"=>"Eames", "scopus_author_id"=>"6602761639"}, {"first_name"=>"Yi Lin", "last_name"=>"Yan", "scopus_author_id"=>"7404586642"}, {"first_name"=>"Mary E.", "last_name"=>"Swartz", "scopus_author_id"=>"7201969341"}, {"first_name"=>"Daniel S.", "last_name"=>"Levic", "scopus_author_id"=>"49661498500"}, {"first_name"=>"Ela W.", "last_name"=>"Knapik", "scopus_author_id"=>"16947222700"}, {"first_name"=>"John H.", "last_name"=>"Postlethwait", "scopus_author_id"=>"7005868055"}, {"first_name"=>"Charles B.", "last_name"=>"Kimmel", "scopus_author_id"=>"7101813997"}], "year"=>2011, "source"=>"PLoS Genetics", "identifiers"=>{"sgr"=>"80052327147", "doi"=>"10.1371/journal.pgen.1002246", "pui"=>"362471746", "pmid"=>"21901110", "scopus"=>"2-s2.0-80052327147", "issn"=>"15537390", "isbn"=>"1553-7404 (Electronic)\\r1553-7390 (Linking)"}, "id"=>"5e61f696-77de-333d-88ef-b0916beaa743", "abstract"=>"Differentiating cells interact with their extracellular environment over time. Chondrocytes embed themselves in a proteoglycan (PG)-rich matrix, then undergo a developmental transition, termed \"maturation,\" when they express ihh to induce bone in the overlying tissue, the perichondrium. Here, we ask whether PGs regulate interactions between chondrocytes and perichondrium, using zebrafish mutants to reveal that cartilage PGs inhibit chondrocyte maturation, which ultimately dictates the timing of perichondral bone development. In a mutagenesis screen, we isolated a class of mutants with decreased cartilage matrix and increased perichondral bone. Positional cloning identified lesions in two genes, fam20b and xylosyltransferase1 (xylt1), both of which encode PG synthesis enzymes. Mutants failed to produce wild-type levels of chondroitin sulfate PGs, which are normally abundant in cartilage matrix, and initiated perichondral bone formation earlier than their wild-type siblings. Primary chondrocyte defects might induce the bone phenotype secondarily, because mutant chondrocytes precociously initiated maturation, showing increased and early expression of such markers as runx2b, collagen type 10a1, and ihh co-orthologs, and ihha mutation suppressed early perichondral bone in PG mutants. Ultrastructural analyses demonstrated aberrant matrix organization and also early cellular features of chondrocyte hypertrophy in mutants. Refining previous in vitro reports, which demonstrated that fam20b and xylt1 were involved in PG synthesis, our in vivo analyses reveal that these genes function in cartilage matrix production and ultimately regulate the timing of skeletal development.", "link"=>"http://www.mendeley.com/research/mutations-fam20b-xylt1-reveal-cartilage-matrix-controls-timing-endochondral-ossification-inhibiting", "reader_count"=>53, "reader_count_by_academic_status"=>{"Unspecified"=>2, "Professor > Associate Professor"=>4, "Student > Doctoral Student"=>2, "Researcher"=>6, "Student > Ph. D. Student"=>15, "Student > Postgraduate"=>3, "Other"=>1, "Student > Master"=>12, "Student > Bachelor"=>4, "Professor"=>4}, "reader_count_by_user_role"=>{"Unspecified"=>2, "Professor > Associate Professor"=>4, "Student > Doctoral Student"=>2, "Researcher"=>6, "Student > Ph. D. Student"=>15, "Student > Postgraduate"=>3, "Other"=>1, "Student > Master"=>12, "Student > Bachelor"=>4, "Professor"=>4}, "reader_count_by_subject_area"=>{"Unspecified"=>6, "Engineering"=>1, "Biochemistry, Genetics and Molecular Biology"=>3, "Nursing and Health Professions"=>1, "Mathematics"=>1, "Agricultural and Biological Sciences"=>33, "Medicine and Dentistry"=>6, "Pharmacology, Toxicology and Pharmaceutical Science"=>1, "Chemistry"=>1}, "reader_count_by_subdiscipline"=>{"Engineering"=>{"Engineering"=>1}, "Medicine and Dentistry"=>{"Medicine and Dentistry"=>6}, "Chemistry"=>{"Chemistry"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>33}, "Nursing and Health Professions"=>{"Nursing and Health Professions"=>1}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>3}, "Mathematics"=>{"Mathematics"=>1}, "Unspecified"=>{"Unspecified"=>6}, "Pharmacology, Toxicology and Pharmaceutical Science"=>{"Pharmacology, Toxicology and Pharmaceutical Science"=>1}}, "reader_count_by_country"=>{"United States"=>2, "Brazil"=>1, "Mexico"=>1, "France"=>1}, "group_count"=>6}

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/742071"], "description"=>"<p>A,F, RT-PCR on wild-type extracts at ages indicated; B,C,G,H,K, whole-mount and D,E,I,J,L section <i>in situ</i> hybridization on wild-type embryos. RT-PCR demonstrated transcripts for <i>fam20b</i> (A) and <i>xylt1</i> (F) from 3 hpf through 54 hpf. Frontal and lateral views of embryos stained by whole-mount <i>in situ</i> hybridization revealed transcripts for <i>fam20b</i> (B,C) and <i>xylt1</i> (G,H) in developing cartilage elements of the craniofacial and pectoral fin skeletons at 55 hpf. In addition, there was diffuse expression of <i>fam20b</i> in the brain and specific <i>xylt1</i> expression in the forebrain (fb). Horizontal section <i>in situ</i> hybridization localized transcripts for <i>fam20b</i> and <i>xylt1</i> to developing chondrocytes (c) of the ceratohyal at 63 hpf (D,I) and 72 hpf (E,J), but no expression in perichondrium (pc) was detected. Lateral view of whole-mount (K) and horizontal section (L) <i>in situ</i> hybridization revealed expression of <i>xylt1</i> in osteoblasts of the opercle (op) at 72 hpf. Abbreviations: -ctl = negative control; c = chondrocytes; ch = ceratohyal; e = eye; et = ethmoid; fb = forebrain; hpf = hours post-fertilization; hs = hyosymplectic; Mk = Meckel's; op = opercle; pc = perichondrium; pf = pectoral fin; pq = palatoquadrate.</p>", "links"=>[], "tags"=>["genetics and genomics", "developmental biology"], "article_id"=>412442, "categories"=>["Genetics", "Developmental Biology"], "users"=>["B. Frank Eames", "Yi-Lin Yan", "Mary E. Swartz", "Daniel S. Levic", "Ela W. Knapik", "John H. Postlethwait", "Charles B. Kimmel"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1002246.g004", "stats"=>{"downloads"=>1, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_fam20b_and_xylt1_are_expressed_in_chondrocytes_but_not_in_perichondrium_/412442", "title"=>"<i>fam20b</i> and <i>xylt1</i> are expressed in chondrocytes, but not in perichondrium.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-08-25 00:40:42"}
  • {"files"=>["https://ndownloader.figshare.com/files/374388", "https://ndownloader.figshare.com/files/374418", "https://ndownloader.figshare.com/files/374456", "https://ndownloader.figshare.com/files/374494"], "description"=>"<div><p>Differentiating cells interact with their extracellular environment over time. Chondrocytes embed themselves in a proteoglycan (PG)-rich matrix, then undergo a developmental transition, termed “maturation,” when they express <em>ihh</em> to induce bone in the overlying tissue, the perichondrium. Here, we ask whether PGs regulate interactions between chondrocytes and perichondrium, using zebrafish mutants to reveal that cartilage PGs inhibit chondrocyte maturation, which ultimately dictates the timing of perichondral bone development. In a mutagenesis screen, we isolated a class of mutants with decreased cartilage matrix and increased perichondral bone. Positional cloning identified lesions in two genes, <em>fam20b</em> and <em>xylosyltransferase1</em> (<em>xylt1</em>), both of which encode PG synthesis enzymes. Mutants failed to produce wild-type levels of chondroitin sulfate PGs, which are normally abundant in cartilage matrix, and initiated perichondral bone formation earlier than their wild-type siblings. Primary chondrocyte defects might induce the bone phenotype secondarily, because mutant chondrocytes precociously initiated maturation, showing increased and early expression of such markers as <em>runx2b</em>, <em>collagen type 10a1</em>, and <em>ihh</em> co-orthologs, and <em>ihha</em> mutation suppressed early perichondral bone in PG mutants. Ultrastructural analyses demonstrated aberrant matrix organization and also early cellular features of chondrocyte hypertrophy in mutants. Refining previous <em>in vitro</em> reports, which demonstrated that <em>fam20b</em> and <em>xylt1</em> were involved in PG synthesis, our <em>in vivo</em> analyses reveal that these genes function in cartilage matrix production and ultimately regulate the timing of skeletal development.</p> </div>", "links"=>[], "tags"=>["mutations", "cartilage", "matrix", "controls", "endochondral", "ossification", "inhibiting", "chondrocyte", "maturation"], "article_id"=>133927, "categories"=>["Genetics", "Developmental Biology"], "users"=>["B. Frank Eames", "Yi-Lin Yan", "Mary E. Swartz", "Daniel S. Levic", "Ela W. Knapik", "John H. Postlethwait", "Charles B. Kimmel"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1002246.s001", "https://dx.doi.org/10.1371/journal.pgen.1002246.s002", "https://dx.doi.org/10.1371/journal.pgen.1002246.s003", "https://dx.doi.org/10.1371/journal.pgen.1002246.s004"], "stats"=>{"downloads"=>0, "page_views"=>8, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/Mutations_in_fam20b_and_xylt1_Reveal_That_Cartilage_Matrix_Controls_Timing_of_Endochondral_Ossification_by_Inhibiting_Chondrocyte_Maturation/133927", "title"=>"Mutations in <em>fam20b</em> and <em>xylt1</em> Reveal That Cartilage Matrix Controls Timing of Endochondral Ossification by Inhibiting Chondrocyte Maturation", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2011-08-25 01:05:27"}
  • {"files"=>["https://ndownloader.figshare.com/files/741988"], "description"=>"<p>A–E, Flat-mounted, dissected pharyngeal skeletons of Alcian blue/Alizarin red-stained 6 dpf larvae. Both sides of the pharyngeal skeletons are shown because the mosaic nature of the rescue experiment produced asymmetric phenotypes in injected larvae. Compared to uninjected wild-type (A) and mutant (B) controls, <i>fam20b<sup>b1127</sup></i> mutant larvae that were injected with wild-type <i>fam20b</i> cDNA under the control of <i>beta-actin2</i> promoter (C) showed skeletal elements with rescued cartilage matrix staining by Alcian blue and decreased bone matrix staining (asterisk) by Alizarin red. D, Higher magnification of boxed region in C illustrates patches of light Alcian blue staining surrounded by heavy Alizarin red staining (arrow, dashed lines) adjacent to patches of dark Alcian blue staining surrounded by light Alizarin red staining. E, <i>fam20b<sup>b1127</sup></i> mutants embryos similarly injected with <i>fam20b<sup>b1127</sup></i> did not rescue the mutant skeletal phenotype. Abbreviations: UIC = uninjected control.</p>", "links"=>[], "tags"=>["rescues", "mutant"], "article_id"=>412358, "categories"=>["Genetics", "Developmental Biology"], "users"=>["B. Frank Eames", "Yi-Lin Yan", "Mary E. Swartz", "Daniel S. Levic", "Ela W. Knapik", "John H. Postlethwait", "Charles B. Kimmel"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1002246.g003", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Wild_type_fam20b_expression_rescues_the_fam20b_b1127_mutant_phenotype_/412358", "title"=>"Wild-type <i>fam20b</i> expression rescues the <i>fam20b<sup>b1127</sup></i> mutant phenotype.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-08-25 00:39:18"}
  • {"files"=>["https://ndownloader.figshare.com/files/742687"], "description"=>"<p>A–R, horizontal section <i>in situ</i> hybridization of developing ceratohyals. The levels of <i>runx2b</i> transcripts appeared higher in chondrocytes (c) and perichondrium (pc) of the ceratohyal in <i>fam20b<sup>b1127</sup></i> (B) and <i>xylt1<sup>b1128</sup></i> (C) mutants, than in wild types (A) at 72 hpf. Expression of <i>col10a1</i> was abundant in chondrocytes (c) and perichondrium (pc) of the ceratohyal in <i>fam20b<sup>b1127</sup></i> (E) and <i>xylt1<sup>b1128</sup></i> (F) mutants, but was not detectable in wild types (D) at 83 hpf. Levels of <i>ihha</i> and <i>ihhb</i> transcripts were up-regulated in chondrocytes (c) of the ceratohyal in <i>fam20b<sup>b1127</sup></i> (H,K) and <i>xylt1<sup>b1128</sup></i> (I,L) mutants, but were not detectable in wild types (G,J) at 72 hpf. Transcript levels for <i>ptch2</i> were increased in the perichondrium of <i>fam20b<sup>b1127</sup></i> (N) and <i>xylt1<sup>b1128</sup></i> (O) mutants, compared to wild types (M) at 72 hpf, whereas no obvious differences in levels of <i>ptch1</i> expression in perichondria were apparent (P–R). Abbreviations: c = chondrocytes; pc = perichondrium.</p>", "links"=>[], "tags"=>["molecular", "markers", "chondrocyte", "maturation"], "article_id"=>413058, "categories"=>["Genetics", "Developmental Biology"], "users"=>["B. Frank Eames", "Yi-Lin Yan", "Mary E. Swartz", "Daniel S. Levic", "Ela W. Knapik", "John H. Postlethwait", "Charles B. Kimmel"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1002246.g010", "stats"=>{"downloads"=>0, "page_views"=>6, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Early_molecular_markers_of_chondrocyte_maturation_in_fam20b_and_xylt1_mutants_/413058", "title"=>"Early molecular markers of chondrocyte maturation in <i>fam20b</i> and <i>xylt1</i> mutants.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-08-25 00:50:58"}
  • {"files"=>["https://ndownloader.figshare.com/files/742412"], "description"=>"<p>A–H, Alcian blue/Alizarin red-stained 6 dpf ceratohyals. Compared to wild types (A,E), cartilage matrix of <i>fam20b<sup>b1127</sup></i> (B) and <i>xylt1<sup>b1128</sup></i> (F) mutants stained less with Alcian blue, but the perichondria of <i>fam20b</i> and <i>xylt1</i> mutants stained more with Alizarin red (arrows) than in wild types. Similar to <i>uxs1</i> mutants (C,G), <i>uxs1;fam20b<sup>b1127</sup></i> (D) and <i>uxs1;xylt1<sup>b1128</sup></i> (H) double mutants showed a greater decrease in Alcian blue staining of cartilage matrix than the decrease seen in <i>fam20b</i> and <i>xylt1</i> mutants; also, Alizarin red staining (arrows) in <i>uxs1</i> single mutant and <i>uxs1;fam20b</i> and <i>uxs1;xylt1</i> double mutant perichondria was at wild-type levels. Abbreviations: bsr = branchiostegal ray.</p>", "links"=>[], "tags"=>["mutation", "epistatic", "cartilage", "phenotypes"], "article_id"=>412783, "categories"=>["Genetics", "Developmental Biology"], "users"=>["B. Frank Eames", "Yi-Lin Yan", "Mary E. Swartz", "Daniel S. Levic", "Ela W. Knapik", "John H. Postlethwait", "Charles B. Kimmel"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1002246.g007", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_uxs1_mutation_is_epistatic_to_cartilage_and_bone_phenotypes_of_fam20b_and_xylt1_mutants_/412783", "title"=>"The <i>uxs1</i> mutation is epistatic to cartilage and bone phenotypes of <i>fam20b</i> and <i>xylt1</i> mutants.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-08-25 00:46:23"}
  • {"files"=>["https://ndownloader.figshare.com/files/741749"], "description"=>"<p>A–H, Alcian blue/Alizarin red-stained 6 dpf larvae, entire heads viewed laterally (A,B, eyes removed) or flat-mounted, dissected pharyngeal skeletons viewed ventrally (C–H). I–L, lateral and dorsal views of entire 8-month-old heads (I,J), and corresponding lateral view images of Alizarin red fluorescence taken by optical projection tomography (OPT; K,L). Overall gross anatomy of the head and craniofacial skeleton are similar in wild types (A) and mutants (B), although staining of cartilage and bone appeared altered in mutants. Compared to wild types (C), <i>fam20b<sup>b1125</sup></i> (D), <i>fam20b<sup>b1127</sup></i> (E), <i>xylt1<sup>b1128</sup></i> (F), and <i>xylt1<sup>b1189</sup></i> (G) mutants had increased Alizarin red staining of bone (arrows) and decreased Alcian blue staining of cartilage. There was no sided-ness to the phenotype, for defective cartilage and bone appeared symmetrically on left and right sides. Double mutant <i>fam20b<sup>b1127</sup>;xylt1<sup>b1128</sup></i> larvae (H) had Alizarin red staining (arrow) similar to that seen in single mutants, but more severe loss of Alcian blue staining. Relative to wild-type siblings (I), <i>fam20b<sup>b1127</sup></i> mutant adults (J) showed foreshortened upper and lower jaws, hypoplastic midface, and bulging eyes. These overall morphological features were underlain by severe reductions to the size of the mutant craniofacial skeleton (K,L). Abbreviations: A = anterior; bsr = branchiostegal ray; ch = ceratohyal; hs = hyosymplectic; Mk = Meckel's; op = opercle; P = posterior; pq = palatoquadrate.</p>", "links"=>[], "tags"=>["skeletons", "mutant", "zebrafish", "larvae", "matrix", "cartilage"], "article_id"=>412112, "categories"=>["Genetics", "Developmental Biology"], "users"=>["B. Frank Eames", "Yi-Lin Yan", "Mary E. Swartz", "Daniel S. Levic", "Ela W. Knapik", "John H. Postlethwait", "Charles B. Kimmel"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1002246.g001", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Craniofacial_skeletons_of_mutant_zebrafish_larvae_exhibit_increased_bone_matrix_and_decreased_cartilage_matrix_/412112", "title"=>"Craniofacial skeletons of mutant zebrafish larvae exhibit increased bone matrix and decreased cartilage matrix.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-08-25 00:35:12"}
  • {"files"=>["https://ndownloader.figshare.com/files/741865"], "description"=>"<p>A, SSR markers mapped <i>b1127</i> to a 0.2 cM interval on LG20 around z20582, showing 0 cross-overs/862 meioses. B, cDNA sequencing of <i>fam20b<sup>b1127</sup></i> revealed a T991C mutation in the sixth exon; genotype assays confirmed perfect linkage of this mutation to mutant phenotype (0 cross-overs/614 meioses). Similarly, <i>fam20b<sup>b1125</sup></i> had a C1162T mutation in the seventh exon. C, The <i>fam20b<sup>b1127</sup></i> mutation changed a highly-conserved Cys to Arg at aa331 (numbering based upon zebrafish protein), while <i>fam20b<sup>b1125</sup></i> mutation changed aa388 from Gln to STOP, which deleted another highly conserved Cys residue at aa389. D, SSR markers mapped <i>b1128</i> to a 0.7 cM interval on LG3. E, cDNA sequencing of <i>xylt1<sup>b1189</sup></i> revealed a T1600G mutation in the seventh exon; <i>xylt1<sup>b1128</sup></i> contained a G2103A splice donor mutation in the ninth exon, which forced usage of cryptic splice donors, typically causing a tetranucleotide insertion. Genotype assays confirmed perfect linkage of these mutations to mutant phenotypes (<i>xylt1<sup>b1189</sup></i>: 0 cross-overs/632 meioses; <i>xylt1<sup>b1128</sup></i>: 0 cross-overs/738 meioses). F, <i>xylt1<sup>b1189</sup></i> mutation changed a highly-conserved Ser to Ala at aa534 (numbering based upon zebrafish protein), while <i>xylt1<sup>b1128</sup></i> mutation typically frameshifted the coding sequence from aa702. Lines in A, B, D, and E are not to scale. Abbreviations: <i>D.re = Danio rerio; G.ga = Gallus gallus; H.sa = Homo sapiens; X.la = Xenopus laevis</i>.</p>", "links"=>[], "tags"=>["mutants", "reveals", "lesions"], "article_id"=>412235, "categories"=>["Genetics", "Developmental Biology"], "users"=>["B. Frank Eames", "Yi-Lin Yan", "Mary E. Swartz", "Daniel S. Levic", "Ela W. Knapik", "John H. Postlethwait", "Charles B. Kimmel"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1002246.g002", "stats"=>{"downloads"=>1, "page_views"=>13, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Mapping_the_mutants_reveals_lesions_in_fam20b_and_xylt1_/412235", "title"=>"Mapping the mutants reveals lesions in <i>fam20b</i> and <i>xylt1</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-08-25 00:37:15"}
  • {"files"=>["https://ndownloader.figshare.com/files/742814"], "description"=>"<p>A–D, transmission electron micrographs of 84 hpf ceratohyals. At 5600X magnification, chondrocytes in the central region of the wild-type ceratohyal displayed abundant rough ER, Golgi, and mitochondria in the cytoplasm (A). Similar views of <i>xylt1</i> mutant chondrocytes at the same age and comparable positions within the developing ceratohyal showed cytoplasmic clearing and tremendous reduction in biosynthetic organelles (B). Mutant chondrocytes also appeared to be separated by less extracellular matrix. While wild-type (C) and mutant (D) extracellular matrix demonstrated fibrillar collagens at 31000X magnification, mutant matrix showed tighter fibril packing and also contained more amorphous electron-dense structures (*) in pericellular matrix than seen in wild types. Abbreviations: cyt = cytoplasm; ecm = extracellular matrix; nuc = nucleus. Scale bars: A,B = 2 µm; C,D = 500 nm.</p>", "links"=>[], "tags"=>["premature", "chondrocyte", "hypertrophy", "aberrant", "matrix"], "article_id"=>413180, "categories"=>["Genetics", "Developmental Biology"], "users"=>["B. Frank Eames", "Yi-Lin Yan", "Mary E. Swartz", "Daniel S. Levic", "Ela W. Knapik", "John H. Postlethwait", "Charles B. Kimmel"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1002246.g011", "stats"=>{"downloads"=>4, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Ultrastructural_evidence_of_premature_chondrocyte_hypertrophy_and_aberrant_matrix_production_in_xylt1_mutants_/413180", "title"=>"Ultrastructural evidence of premature chondrocyte hypertrophy and aberrant matrix production in <i>xylt1</i> mutants.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-08-25 00:53:00"}
  • {"files"=>["https://ndownloader.figshare.com/files/742900"], "description"=>"<p>A–H, Alcian blue/Alizarin red-stained 6 dpf ceratohyals. Alcian blue staining was comparable between wild-type (A,E) and <i>ihha</i> mutant (B,F) cartilages, but was decreased in <i>fam20b<sup>b1127</sup></i> (C) and <i>xylt1<sup>b1128</sup></i> (G) single mutant, and in <i>fam20b<sup>b1127</sup>;ihha</i> (D) and <i>xylt1<sup>b1128</sup>;ihha</i> (H) double mutant, cartilages. Alizarin red staining of <i>fam20b<sup>b1127</sup></i> (C) and <i>xylt1<sup>b1128</sup></i> (D) chondral bones was abundant, while no such staining was observed in wild types (A,E), <i>ihha</i> mutants (B,F), or <i>fam20b<sup>b1127</sup>;ihha</i> (D) and <i>xylt1<sup>b1128</sup>;ihha</i> double mutants (H). Abbreviations: bsr = branchiostegal ray.</p>", "links"=>[], "tags"=>["mutation", "epistatic", "phenotypes"], "article_id"=>413271, "categories"=>["Genetics", "Developmental Biology"], "users"=>["B. Frank Eames", "Yi-Lin Yan", "Mary E. Swartz", "Daniel S. Levic", "Ela W. Knapik", "John H. Postlethwait", "Charles B. Kimmel"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1002246.g012", "stats"=>{"downloads"=>0, "page_views"=>6, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_ihha_mutation_is_epistatic_to_bone_but_not_cartilage_phenotypes_of_fam20b_and_xylt1_mutants_/413271", "title"=>"The <i>ihha</i> mutation is epistatic to bone, but not cartilage, phenotypes of <i>fam20b</i> and <i>xylt1</i> mutants.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-08-25 00:54:31"}
  • {"files"=>["https://ndownloader.figshare.com/files/742592"], "description"=>"<p>A–F, Alcian blue/Alizarin red-stained ceratohyals. G–I, live Alizarin red fluorescence of <i>fam20b<sup>b1127</sup>;Tg(sp7:EGFP)b1212</i> and <i>xylt1<sup>b1128</sup>;Tg(sp7:EGFP)b1212</i> larvae. J–O, whole-mount <i>in situ</i> hybridization of 4 dpf ceratohyals. No signs of Alizarin red staining were observed at 3 dpf in ceratohyals of wild types (A), or <i>fam20b<sup>b1127</sup></i> (B) or <i>xylt1<sup>b1128</sup></i> (C) mutants. By 4.5 dpf, Alizarin red staining was still absent from ceratohyals of wild-type larvae (D, arrow), although it was detected in ceratohyals of <i>fam20b<sup>b1127</sup></i> (E, arrow) and <i>xylt1<sup>b1128</sup></i> (F, arrow) mutants. Both GFP expression and Alizarin red staining were at background levels in the perichondrium of the ceratohyal in 4.5 dpf wild types (G, arrow), but were obvious in the <i>fam20b<sup>b1127</sup></i> and <i>xylt1<sup>b1128</sup></i> mutant perichondria (H,I, arrow). At 4 dpf, <i>col10a1</i> expression was not detected in the ceratohyal perichondrium of wild types (J, arrow), but was expressed highly in <i>fam20b<sup>b1127</sup></i> (K, arrow) and <i>xylt1<sup>b1128</sup></i> (L, arrow) mutant perichondria. Abbreviations: bsr = branchiostegal ray.</p>", "links"=>[], "tags"=>["osteoblast", "differentiation", "perichondria"], "article_id"=>412965, "categories"=>["Genetics", "Developmental Biology"], "users"=>["B. Frank Eames", "Yi-Lin Yan", "Mary E. Swartz", "Daniel S. Levic", "Ela W. Knapik", "John H. Postlethwait", "Charles B. Kimmel"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1002246.g009", "stats"=>{"downloads"=>0, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Precocious_bone_formation_and_osteoblast_differentiation_in_perichondria_of_fam20b_and_xylt1_mutants_/412965", "title"=>"Precocious bone formation and osteoblast differentiation in perichondria of <i>fam20b</i> and <i>xylt1</i> mutants.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-08-25 00:49:25"}
  • {"files"=>["https://ndownloader.figshare.com/files/742496"], "description"=>"<p>At 6 dpf, Alizarin red staining (see <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1002246#pgen-1002246-g001\" target=\"_blank\">Figure 1C–1G</a>) was visible more often in skeletal elements of <i>fam20b<sup>b1127</sup></i> (A) and <i>xylt1<sup>b1128</sup></i> (B) mutants than in wild types. Statistically significant (*, p<0.05) increases, however, were observed in more chondral bones (pq, hm, ch) than dermal bones (ept, mx). Quantitative analyses of the sum of bone areas in left and right ceratohyals (ch(l) and ch(r), dashed outlines) in whole-mount, ventral images of live Alizarin red fluorescence (C–E) revealed statistically significant (*, p<0.05) increases in <i>fam20b<sup>b1127</sup></i> (F) and <i>xylt1<sup>b1128</sup></i> (G) mutants, compared to wild-type siblings at 6 dpf. Abbreviations: A = anterior; ch = ceratohyal; ept = entopterygoid; hm = hyomandibular; l = left; mx = maxilla; pq = palatoquadrate; P = posterior; r = right.</p>", "links"=>[], "tags"=>["mutants", "perichondral"], "article_id"=>412868, "categories"=>["Genetics", "Developmental Biology"], "users"=>["B. Frank Eames", "Yi-Lin Yan", "Mary E. Swartz", "Daniel S. Levic", "Ela W. Knapik", "John H. Postlethwait", "Charles B. Kimmel"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1002246.g008", "stats"=>{"downloads"=>1, "page_views"=>8, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_fam20b_and_xylt1_mutants_have_increased_perichondral_bone_/412868", "title"=>"<i>fam20b</i> and <i>xylt1</i> mutants have increased perichondral bone.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-08-25 00:47:48"}
  • {"files"=>["https://ndownloader.figshare.com/files/742177"], "description"=>"<p>A–E, Alcian blue-stained 6 dpf ceratohyals. Compared to wild types (A), Alcian blue staining appeared lower in <i>fam20b<sup>b1127</sup></i> (B) and <i>xylt1<sup>b1128</sup></i> (C) mutants. Staining in <i>fam20b<sup>b1127</sup>;xylt1<sup>b1128</sup></i> double mutants (D) was decreased in comparison to single mutants, although <i>uxs1</i> mutants (E) showed the greatest decrease in Alcian blue staining. F, Quantitation of Alcian blue in lysates of 6 dpf larvae revealed statistically significant changes (ANOVA p<0.0001) for each mutant class. Compared to wild-type Alcian blue staining, <i>fam20b<sup>b1127</sup></i> mutants had 50±3.0%, <i>xylt1<sup>b1128</sup></i> mutants had 57±2.0%, <i>fam20b<sup>b1127</sup>;xylt1<sup>b1128</sup></i> double mutants had 39±2.6%, and <i>uxs1</i> mutants had 22±1.5%. G, Simplified pathway of PG synthesis, demonstrating that <i>uxs1</i> is upstream of both <i>xylt1</i>, which adds xylose to a serine (Ser) residue of the core protein, and also <i>fam20b</i>, which phosphorylates (P) xylose in the nascent GAG chain. Abbreviations: CSPGs = chondroitin sulfate proteoglycans; HSPGs = heparan sulfate proteoglycans.</p>", "links"=>[], "tags"=>["alcian", "staining", "mutants"], "article_id"=>412545, "categories"=>["Genetics", "Developmental Biology"], "users"=>["B. Frank Eames", "Yi-Lin Yan", "Mary E. Swartz", "Daniel S. Levic", "Ela W. Knapik", "John H. Postlethwait", "Charles B. Kimmel"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1002246.g005", "stats"=>{"downloads"=>1, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Reduction_of_Alcian_blue_staining_in_fam20b_and_xylt1_mutants_is_less_severe_than_in_uxs1_mutants_/412545", "title"=>"Reduction of Alcian blue staining in <i>fam20b</i> and <i>xylt1</i> mutants is less severe than in <i>uxs1</i> mutants.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-08-25 00:42:25"}
  • {"files"=>["https://ndownloader.figshare.com/files/742266"], "description"=>"<p>A–F, Ventral views of confocal images of embryonic heads immunostained for chondroitin sulfate proteoglycan (CSPG) at 3 dpf (A,B) or heparan sulfate PG (HSPG) at 4 dpf (C–F). Compared to abundant CSPGs detected in wild-type skeletal elements (A), <i>fam20b<sup>b1127</sup></i> mutants showed decreased CSPG immunoreactivity (B). HSPGs were abundant in gill filaments and ventral muscle groups of both wild-type (C) and <i>fam20b<sup>b1127</sup></i> mutant zebrafish (D). HSPG immunoreactivity was reduced greatly around developing chondrocytes of the <i>fam20b<sup>b1127</sup></i> mutant ceratohyal (outline in F), compared to those of wild-type siblings (outline in E). Also, the number of intracellular HSPG aggregates (arrowhead) was increased in <i>fam20b<sup>b1127</sup></i> mutant chondrocytes. Abbreviations: A = anterior; αCS = anti-chondroitin sulfate; αHS = anti-heparan sulfate; c = chondrocyte; ch = ceratohyal; e = eye; gf = gill filament; ih = interhyoideus; imp = intermandibularis posterior; Mk = Meckel's; op = opercle; P = posterior; pq = palatoquadrate.</p>", "links"=>[], "tags"=>["mutants", "reduced", "proteoglycan"], "article_id"=>412639, "categories"=>["Genetics", "Developmental Biology"], "users"=>["B. Frank Eames", "Yi-Lin Yan", "Mary E. Swartz", "Daniel S. Levic", "Ela W. Knapik", "John H. Postlethwait", "Charles B. Kimmel"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1002246.g006", "stats"=>{"downloads"=>1, "page_views"=>8, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_fam20b_mutants_display_reduced_proteoglycan_immunoreactivity_/412639", "title"=>"<i>fam20b</i> mutants display reduced proteoglycan immunoreactivity", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2011-08-25 00:43:59"}

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Relative Metric

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