The Mating Type Locus (MAT) and Sexual Reproduction of Cryptococcus heveanensis: Insights into the Evolution of Sex and Sex-Determining Chromosomal Regions in Fungi
Publication Date
May 20, 2010
Journal
PLOS Genetics
Authors
Banu Metin, Keisha Findley & Joseph Heitman
Volume
6
Issue
5
Pages
e1000961
DOI
https://dx.plos.org/10.1371/journal.pgen.1000961
Publisher URL
http://journals.plos.org/plosgenetics/article?id=10.1371%2Fjournal.pgen.1000961
PubMed
http://www.ncbi.nlm.nih.gov/pubmed/20502678
PubMed Central
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2873909
Europe PMC
http://europepmc.org/abstract/MED/20502678
Web of Science
000278557300031
Scopus
77956631575
Mendeley
http://www.mendeley.com/research/mating-type-locus-mat-sexual-reproduction-cryptococcus-heveanensis-insights-evolution-sex-sexdetermi
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{"title"=>"The mating type locus (MAT) and sexual reproduction of Cryptococcus heveanensis: Insights into the evolution of sex and sex-determining chromosomal regions in fungi", "type"=>"journal", "authors"=>[{"first_name"=>"Banu", "last_name"=>"Metin", "scopus_author_id"=>"26322693600"}, {"first_name"=>"Keisha", "last_name"=>"Findley", "scopus_author_id"=>"26322464900"}, {"first_name"=>"Joseph", "last_name"=>"Heitman", "scopus_author_id"=>"19634810800"}], "year"=>2010, "source"=>"PLoS Genetics", "identifiers"=>{"issn"=>"15537390", "scopus"=>"2-s2.0-77953204250", "sgr"=>"77953204250", "pui"=>"358963166", "isbn"=>"1553-7404 (Electronic)\\r1553-7390 (Linking)", "pmid"=>"20502678", "doi"=>"10.1371/journal.pgen.1000961"}, "id"=>"9e7a0291-1b2e-330c-9fe8-7878d8b39b18", "abstract"=>"Mating in basidiomycetous fungi is often controlled by two unlinked, multiallelic loci encoding homeodomain transcription factors or pheromones/pheromone receptors. In contrast to this tetrapolar organization, Cryptococcus neoformans/Cryptococcus gattii have a bipolar mating system, and a single biallelic locus governs sexual reproduction. The C. neoformans MAT locus is unusually large (>100 kb), contains >20 genes, and enhances virulence. Previous comparative genomic studies provided insights into how this unusual MAT locus might have evolved involving gene acquisitions into two unlinked loci and fusion into one contiguous locus, converting an ancestral tetrapolar system to a bipolar one. Here we tested this model by studying Cryptococcus heveanensis, a sister species to the pathogenic Cryptococcus species complex. An extant sexual cycle was discovered; co-incubating fertile isolates results in the teleomorph (Kwoniella heveanensis) with dikaryotic hyphae, clamp connections, septate basidia, and basidiospores. To characterize the C. heveanensis MAT locus, a fosmid library was screened with C. neoformans/C. gattii MAT genes. Positive fosmids were sequenced and assembled to generate two large probably unlinked MAT gene clusters: one corresponding to the homeodomain locus and the other to the pheromone/receptor locus. Strikingly, two divergent homeodomain genes (SXI1, SXI2) are present, similar to the bE/bW Ustilago maydis paradigm, suggesting one or the other homeodomain gene was recently lost in C. neoformans/C. gattii. Sequencing MAT genes from other C. heveanensis isolates revealed a multiallelic homeodomain locus and at least a biallelic pheromone/receptor locus, similar to known tetrapolar species. Taken together, these studies reveal an extant C. heveanensis sexual cycle, define the structure of its MAT locus consistent with tetrapolar mating, and support the proposed evolutionary model for the bipolar Cryptococcus MAT locus revealing transitions in sexuality concomitant with emergence of a pathogenic clade. These studies provide insight into convergent processes that independently punctuated evolution of sex-determining loci and sex chromosomes in fungi, plants, and animals.", "link"=>"http://www.mendeley.com/research/mating-type-locus-mat-sexual-reproduction-cryptococcus-heveanensis-insights-evolution-sex-sexdetermi", "reader_count"=>69, "reader_count_by_academic_status"=>{"Professor > Associate Professor"=>5, "Student > Doctoral Student"=>5, "Researcher"=>22, "Student > Ph. D. Student"=>15, "Student > Postgraduate"=>2, "Student > Master"=>9, "Other"=>1, "Student > Bachelor"=>4, "Lecturer > Senior Lecturer"=>1, "Professor"=>4}, "reader_count_by_user_role"=>{"Professor > Associate Professor"=>5, "Student > Doctoral Student"=>5, "Researcher"=>22, "Student > Ph. D. Student"=>15, "Student > Postgraduate"=>2, "Student > Master"=>9, "Other"=>1, "Student > Bachelor"=>4, "Lecturer > Senior Lecturer"=>1, "Professor"=>4}, "reader_count_by_subject_area"=>{"Unspecified"=>2, "Environmental Science"=>1, "Biochemistry, Genetics and Molecular Biology"=>6, "Agricultural and Biological Sciences"=>53, "Medicine and Dentistry"=>1, "Chemistry"=>1, "Immunology and Microbiology"=>3, "Earth and Planetary Sciences"=>1}, "reader_count_by_subdiscipline"=>{"Medicine and Dentistry"=>{"Medicine and Dentistry"=>1}, "Chemistry"=>{"Chemistry"=>1}, "Immunology and Microbiology"=>{"Immunology and Microbiology"=>3}, "Earth and Planetary Sciences"=>{"Earth and Planetary Sciences"=>1}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>53}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>6}, "Unspecified"=>{"Unspecified"=>2}, "Environmental Science"=>{"Environmental Science"=>1}}, "reader_count_by_country"=>{"Sweden"=>1, "United States"=>2, "Japan"=>2, "United Kingdom"=>1, "Malaysia"=>1, "Germany"=>1, "Spain"=>1}, "group_count"=>0}
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Figshare 13821 Feb 21:10 UTC

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  • {"files"=>["https://ndownloader.figshare.com/files/849251"], "description"=>"<p><i>C. heveanensis</i> has a multiallelic HD locus (<i>B</i> locus) including the genes <i>SXI1</i> and <i>SXI2</i> with at least 6 alleles among 7 unique strains examined. The PR locus (<i>A</i> locus) of <i>C. heveanensis</i> is at least biallelic and 6 of the strains have the <i>A1</i> allele, while one of the strains, BCC 8398, has the <i>A2</i> allele. The PR locus contains at least the <i>STE3</i>, <i>STE12</i>, <i>MFA1</i>/<i>2</i>, <i>CNB00600</i>, and <i>CNG04540</i> genes.</p>", "links"=>[], "tags"=>["tetrapolar", "mating", "multiallelic", "locus", "biallelic"], "article_id"=>519707, "categories"=>["Microbiology", "Evolutionary Biology", "Genetics", "Molecular Biology"], "users"=>["Banu Metin", "Keisha Findley", "Joseph Heitman"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000961.g009", "stats"=>{"downloads"=>1, "page_views"=>38, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_C_heveanensis_is_likely_to_have_a_tetrapolar_mating_type_system_with_a_multiallelic_mating_type_locus_and_at_least_a_biallelic_pheromone_receptor_locus_/519707", "title"=>"<i>C. heveanensis</i> is likely to have a tetrapolar mating type system with a multiallelic mating type locus and at least a biallelic pheromone/receptor locus.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-05-20 02:41:47"}
  • {"files"=>["https://ndownloader.figshare.com/files/849363"], "description"=>"<p>First, mating-related genes together with other genes were acquired into two unlinked loci. Second, the mating type specific region including only the homeodomain genes in the HD locus and the pheromone and the pheromone receptor gene in the PR locus expanded serially by rearrangements or insertions leading to suppressed recombination. During expansion, some of the genes were relocated to different parts of the genome and others were lost. Third, the two loci were combined to a single locus in one mating type by a translocation event forming a transient tripolar system. Fourth, recombination between the mating types occurred to form the bipolar system. Ongoing inversions and rearrangements fashioned the extant <i>C. neoformans</i>/<i>C. gattii MAT</i> locus. Genes shown with black or white arrows are those found in the <i>C. neoformans</i>/<i>C. gattii MAT</i> locus, those in white exhibit a species-specific phylogeny, and those in yellow are not linked to the extant <i>MAT</i> locus in the pathogenic <i>Cryptococcus</i> species complex.</p>", "links"=>[], "tags"=>["depicting", "bipolar", "locus", "pathogenic", "ancestral", "unlinked", "homeodomain"], "article_id"=>519817, "categories"=>["Microbiology", "Evolutionary Biology", "Genetics", "Molecular Biology"], "users"=>["Banu Metin", "Keisha Findley", "Joseph Heitman"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000961.g010", "stats"=>{"downloads"=>0, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_A_model_depicting_the_evolution_of_the_bipolar_MAT_locus_in_the_pathogenic_Cryptococcus_species_from_ancestral_unlinked_homeodomain_and_pheromone_receptor_loci_/519817", "title"=>"A model depicting the evolution of the bipolar <i>MAT</i> locus in the pathogenic <i>Cryptococcus</i> species from ancestral unlinked homeodomain and pheromone/receptor loci.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-05-20 02:43:37"}
  • {"files"=>["https://ndownloader.figshare.com/files/848599"], "description"=>"<p>C. heveanensis and <i>T. mesenterica</i> genes that are present in the <i>MAT</i> locus of <i>C. neoformans</i> and <i>C. gattii</i> are shown in black, while others are shown in yellow. Syntenic blocks containing more than one gene are shown as pink bars. The <i>T. mesenterica</i> and <i>C. heveanensis</i> homeodomain locus contains two homeodomain genes homologous with <i>SXI1</i> and <i>SXI2</i>; whereas, <i>C. neoformans</i> and <i>C. gattii</i> contain a single homeodomain gene, namely <i>SXI1</i> for α, and <i>SXI2</i> for the <b>a</b> allele. Upper scale bar refers to <i>T. mesenterica</i> and <i>C. heveanensis</i>, while lower scale bar refers to <i>C. neoformans</i> var. <i>neoformans</i>, <i>C. neoformans</i> var. <i>grubii</i> and <i>C. gattii</i>. JGI annotation IDs for <i>T. mesenterica</i> genes are listed in <a href=\"http://www.plosgenetics.org/article/info:doi/10.1371/journal.pgen.1000961#pgen.1000961.s006\" target=\"_blank\">Table S1</a>. The relative order and the orientations of the <i>C. heveanensis</i> and <i>T. mesenterica</i> contigs are unknown.</p>", "links"=>[], "tags"=>["locus", "regions", "alleles"], "article_id"=>519061, "categories"=>["Microbiology", "Evolutionary Biology", "Genetics", "Molecular Biology"], "users"=>["Banu Metin", "Keisha Findley", "Joseph Heitman"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000961.g003", "stats"=>{"downloads"=>0, "page_views"=>9, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Synteny_between_MAT_locus_regions_of_C_heveanensis_T_mesenterica_and_alleles_of_C_neoformans_var_neoformans_C_neoformans_var_grubii_and_C_gattii_/519061", "title"=>"Synteny between <i>MAT</i> locus regions of <i>C. heveanensis</i>, <i>T. mesenterica</i>, and α alleles of <i>C. neoformans</i> var. <i>neoformans</i>, <i>C. neoformans</i> var. <i>grubii</i>, and <i>C. gattii</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-05-20 02:31:01"}
  • {"files"=>["https://ndownloader.figshare.com/files/848784"], "description"=>"<p>(A) Southern blot analysis showing hybridization patterns of <i>SXI1</i> and <i>SXI2</i> gene probes to EcoRV digested genomic DNA from <i>C. heveanensis</i> strains. Lane 1, CBS 569, 2, BCC 8305, 3, BCC 8313, 4, BCC 8384, 5, BCC 8396, 6, BCC 8398, 7, BCC 11754, 8, BCC 11757, 9, BCC 15000. (B) Percent sequence identity plots comparing an ∼12 kb region containing the <i>SXI1</i> and <i>SXI2</i> genes from CBS 569 with the corresponding region from other isolates. The blue box shows the highly dissimilar region corresponding to the N-terminal regions of Sxi1 and Sxi2. The black bars under the genes depict the exons. (C) A representative dot plot comparing the homeodomain region of CBS 569 (x-axis) and BCC 11757 (y-axis).</p>", "links"=>[], "tags"=>["multiallelic"], "article_id"=>519241, "categories"=>["Microbiology", "Evolutionary Biology", "Genetics", "Molecular Biology"], "users"=>["Banu Metin", "Keisha Findley", "Joseph Heitman"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000961.g005", "stats"=>{"downloads"=>0, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Homeodomain_region_of_C_heveanensis_is_multiallelic_and_likely_to_be_limited_to_the_SXI1_and_SXI2_genes_/519241", "title"=>"Homeodomain region of <i>C. heveanensis</i> is multiallelic and likely to be limited to the <i>SXI1</i> and <i>SXI2</i> genes.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-05-20 02:34:01"}
  • {"files"=>["https://ndownloader.figshare.com/files/848410"], "description"=>"<p>(A,B) Hyphae with clamp connections. Scale bars represent 50 µm. Representative clamp connections (CC) are shown in the enlarged images located in the lower left panels (arrows, scale bars  = 10 µm). (A) Differential interference contrast (DIC) image, (B) fluorescence image showing cell walls, septa and nuclei stained with Calcofluor white and Sytox green. (C–E) DIC images of basidial clusters. Scale bars represent 50 µm. (C) A young basidial cluster embedded in the agar, (D,E) mature basidial clusters on the surface of the agar. (F,G) Scanning electron microscopy images of basidial clusters, scale bar in (F) is 50 µm, and in (G) is 10 µm. (H) A developing basidial cluster with two mature and numerous young basidia, scale bar  = 10 µm. (I and J) Globose basidia with cruciate septa. Scale bars represent 10 µm. (K) A basidium and basidiospores, scale bar  = 10 µm. (L) Vegetative yeast cells, scale bar  = 10 µm.</p>", "links"=>[], "tags"=>["reproduction"], "article_id"=>518868, "categories"=>["Microbiology", "Evolutionary Biology", "Genetics", "Molecular Biology"], "users"=>["Banu Metin", "Keisha Findley", "Joseph Heitman"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000961.g001", "stats"=>{"downloads"=>0, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Sexual_reproduction_of_C_heveanensis_/518868", "title"=>"Sexual reproduction of <i>C. heveanensis</i>.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-05-20 02:27:48"}
  • {"files"=>["https://ndownloader.figshare.com/files/422206", "https://ndownloader.figshare.com/files/422272", "https://ndownloader.figshare.com/files/422317", "https://ndownloader.figshare.com/files/422398", "https://ndownloader.figshare.com/files/422438", "https://ndownloader.figshare.com/files/422473", "https://ndownloader.figshare.com/files/422502"], "description"=>"<div><p>Mating in basidiomycetous fungi is often controlled by two unlinked, multiallelic loci encoding homeodomain transcription factors or pheromones/pheromone receptors. In contrast to this tetrapolar organization, <em>Cryptococcus neoformans/Cryptococcus gattii</em> have a bipolar mating system, and a single biallelic locus governs sexual reproduction. The <em>C. neoformans MAT</em> locus is unusually large (>100 kb), contains >20 genes, and enhances virulence. Previous comparative genomic studies provided insights into how this unusual <em>MAT</em> locus might have evolved involving gene acquisitions into two unlinked loci and fusion into one contiguous locus, converting an ancestral tetrapolar system to a bipolar one. Here we tested this model by studying <em>Cryptococcus heveanensis</em>, a sister species to the pathogenic <em>Cryptococcus</em> species complex. An extant sexual cycle was discovered; co-incubating fertile isolates results in the teleomorph (<em>Kwoniella heveanensis</em>) with dikaryotic hyphae, clamp connections, septate basidia, and basidiospores. To characterize the <em>C. heveanensis MAT</em> locus, a fosmid library was screened with <em>C. neoformans/C. gattii MAT</em> genes. Positive fosmids were sequenced and assembled to generate two large probably unlinked <em>MAT</em> gene clusters: one corresponding to the homeodomain locus and the other to the pheromone/receptor locus. Strikingly, two divergent homeodomain genes (<em>SXI1, SXI2</em>) are present, similar to the bE/bW <em>Ustilago maydis</em> paradigm, suggesting one or the other homeodomain gene was recently lost in <em>C. neoformans/C. gattii</em>. Sequencing <em>MAT</em> genes from other <em>C. heveanensis</em> isolates revealed a multiallelic homeodomain locus and at least a biallelic pheromone/receptor locus, similar to known tetrapolar species. Taken together, these studies reveal an extant <em>C. heveanensis</em> sexual cycle, define the structure of its <em>MAT</em> locus consistent with tetrapolar mating, and support the proposed evolutionary model for the bipolar <em>Cryptococcus MAT</em> locus revealing transitions in sexuality concomitant with emergence of a pathogenic clade. These studies provide insight into convergent processes that independently punctuated evolution of sex-determining loci and sex chromosomes in fungi, plants, and animals.</p></div>", "links"=>[], "tags"=>["mating", "locus", "reproduction", "insights", "sex-determining", "chromosomal", "regions", "fungi"], "article_id"=>143394, "categories"=>["Microbiology", "Evolutionary Biology", "Genetics", "Molecular Biology"], "users"=>["Banu Metin", "Keisha Findley", "Joseph Heitman"], "doi"=>["https://dx.doi.org/10.1371/journal.pgen.1000961.s001", "https://dx.doi.org/10.1371/journal.pgen.1000961.s002", "https://dx.doi.org/10.1371/journal.pgen.1000961.s003", "https://dx.doi.org/10.1371/journal.pgen.1000961.s004", "https://dx.doi.org/10.1371/journal.pgen.1000961.s005", "https://dx.doi.org/10.1371/journal.pgen.1000961.s006", "https://dx.doi.org/10.1371/journal.pgen.1000961.s007"], "stats"=>{"downloads"=>0, "page_views"=>17, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/The_Mating_Type_Locus_MAT_and_Sexual_Reproduction_of_Cryptococcus_heveanensis_Insights_into_the_Evolution_of_Sex_and_Sex_Determining_Chromosomal_Regions_in_Fungi/143394", "title"=>"The Mating Type Locus (<em>MAT</em>) and Sexual Reproduction of <em>Cryptococcus heveanensis</em>: Insights into the Evolution of Sex and Sex-Determining Chromosomal Regions in Fungi", "pos_in_sequence"=>0, "defined_type"=>4, "published_date"=>"2010-05-20 00:56:34"}
  • {"files"=>["https://ndownloader.figshare.com/files/849518"], "description"=>"<p><b>T</b>: type strain.</p><p>*: inferred mating type from molecular analysis.</p><p>§: the <i>B</i> locus of this strain was found to be very similar to BCC 8398 (92% for Sxi1 and 94% for Sxi2, with just one amino acid difference in the N-terminal dimerization region of Sxi1 and four amino acid differences in this region of Sxi2), but it could represent a different allele (B7).</p>", "links"=>[], "tags"=>["isolates"], "article_id"=>519983, "categories"=>["Microbiology", "Evolutionary Biology", "Genetics", "Molecular Biology"], "users"=>["Banu Metin", "Keisha Findley", "Joseph Heitman"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000961.t001", "stats"=>{"downloads"=>0, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Description_of_C_heveanensis_isolates_used_in_this_study_/519983", "title"=>"Description of <i>C. heveanensis</i> isolates used in this study.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-05-20 02:46:23"}
  • {"files"=>["https://ndownloader.figshare.com/files/849004"], "description"=>"<p>(A) Comparison plot of the PR locus of CBS 569 and BCC 8398. The red and blue bands represent the forward and reverse matches, respectively. The genes are mostly in the same orientation except <i>MFA1/2</i>. The intensity of the color is proportional to the percent identity of the match, where red bands show regions of high sequence identity, while pink bands indicate lower sequence identity. (B) Dot plot analysis of the PR locus of CBS 569 and BCC 8398. (C) Sequence alignment of pheromone sequences from CBS 569, BCC 8398, <i>C. neoformans</i> var. <i>neoformans</i> MFα, <i>C. neoformans</i> var. <i>neoformans</i> MF<b>a</b>, <i>C. neoformans</i> var. <i>grubii</i> MFα, <i>C. neoformans</i> var. <i>grubii</i> MF<b>a</b>, <i>C. gattii</i> MFα, <i>C. gatii</i> MF<b>a</b> and <i>T. mesenterica</i> pheromone tremerogen a-13. The arrow indicates the N-terminal amino acid of tremerogen a-13 after proteolytic cleavage. (D) Southern blot analysis showing hybridization patterns of <i>STE11</i> and <i>MYO2</i> gene probes to EcoRV digested genomic DNA from <i>C. heveanensis</i> strains. Lane 1, CBS 569, 2, BCC 8305, 3, BCC 8313, 4, BCC 8384, 5, BCC 8396, 6, BCC 8398, 7, BCC 11754, 8, BCC 11757, 9, BCC 15000.</p>", "links"=>[], "tags"=>["Evolutionary biology", "evolutionary biology/microbial evolution and genomics", "genetics and genomics", "genetics and genomics/comparative genomics", "genetics and genomics/genomics", "genetics and genomics/microbial evolution and genomics", "microbiology", "microbiology/cellular microbiology and pathogenesis", "microbiology/medical microbiology", "microbiology/microbial evolution and genomics", "molecular biology"], "article_id"=>519453, "categories"=>["Microbiology", "Evolutionary Biology", "Genetics", "Molecular Biology"], "users"=>["Banu Metin", "Keisha Findley", "Joseph Heitman"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000961.g007", "stats"=>{"downloads"=>0, "page_views"=>35, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Analysis_of_the_pheromone_receptor_region_of_C_heveanensis_isolates_/519453", "title"=>"Analysis of the pheromone/receptor region of <i>C. heveanensis</i> isolates.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-05-20 02:37:33"}
  • {"files"=>["https://ndownloader.figshare.com/files/848531"], "description"=>"<p><i>MAT</i> gene probes (represented by blue bars in the figure) were used to screen the fosmid library of <i>C. heveanensis</i> CBS 569. Sequencing the positive fosmids, represented by black lines, and subsequent assembly of the sequences generated two clusters corresponding to the homeodomain locus (∼80 kb) and the pheromone/receptor locus (∼180 kb). The genes that are present in the <i>MAT</i> locus of the pathogenic <i>Cryptococcus</i> species are shown in black, while others are shown in yellow. The red line indicates an ∼20 kb region sequenced by primer walking.</p>", "links"=>[], "tags"=>["heveanensis", "locus"], "article_id"=>518994, "categories"=>["Microbiology", "Evolutionary Biology", "Genetics", "Molecular Biology"], "users"=>["Banu Metin", "Keisha Findley", "Joseph Heitman"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000961.g002", "stats"=>{"downloads"=>0, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_C_heveanensis_MAT_locus_structure_/518994", "title"=>"<i>C. heveanensis MAT</i> locus structure.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-05-20 02:29:54"}
  • {"files"=>["https://ndownloader.figshare.com/files/849119"], "description"=>"<p>(A) Species-specific profile exhibited by <i>CID1</i> and <i>GEF1</i>, (B) mating type-specific pattern exhibited by <i>STE20</i> and <i>SPO14</i> from <i>C. gattii</i> and <i>C. neoformans</i>, but not from <i>C. heveanensis</i> or <i>T. mesenterica</i>, (C) mating type-specific profile demonstrated by <i>STE3</i> and <i>STE12</i>.</p>", "links"=>[], "tags"=>["genes", "phylogenetic"], "article_id"=>519576, "categories"=>["Microbiology", "Evolutionary Biology", "Genetics", "Molecular Biology"], "users"=>["Banu Metin", "Keisha Findley", "Joseph Heitman"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000961.g008", "stats"=>{"downloads"=>0, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Mating_type_genes_exhibit_three_phylogenetic_patterns_/519576", "title"=>"Mating type genes exhibit three phylogenetic patterns.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-05-20 02:39:36"}
  • {"files"=>["https://ndownloader.figshare.com/files/848693"], "description"=>"<p><i>C. heveanensis MAT</i> locus includes numerous genes that are not present in the <i>MAT</i> locus of <i>C. neoformans</i> or <i>C. gattii</i>. Although many of these genes are on the 4th chromosome of <i>C. neoformans</i> strain JEC21 where <i>MAT</i> is located, they correspond to the opposite telomeric region, suggesting intrachromosomal rearragements during the evolution of the <i>MAT</i> locus. Pink lines/bars depict the synteny between genes located within the <i>MAT</i> locus of <i>C. neoformans</i> and the corresponding genes in <i>C. heveanensis</i>. Blue lines/bars show the syntenic relationship of the genes that are linked to the <i>MAT</i> locus of <i>C. heveanensis</i>, but not part of the <i>MAT</i> locus of <i>C. neoformans</i>. The <i>C. heveanensis</i> genes whose <i>C. neoformans</i> homologs are located on different chromosomes are indicated with purple arrows with chromosome numbers. The genes that do not have apparent <i>C. neoformans</i> homologs are indicated by NCH, which stands for “<u>n</u>o <i><u>C</u>. neoformans </i><u>h</u>omolog”.</p>", "links"=>[], "tags"=>["heveanensis", "locus", "shares", "synteny", "chromosome"], "article_id"=>519154, "categories"=>["Microbiology", "Evolutionary Biology", "Genetics", "Molecular Biology"], "users"=>["Banu Metin", "Keisha Findley", "Joseph Heitman"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000961.g004", "stats"=>{"downloads"=>0, "page_views"=>2, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_C_heveanensis_MAT_locus_region_shares_synteny_with_the_4_th_chromosome_of_C_neoformans_JEC21_/519154", "title"=>"<i>C. heveanensis MAT</i> locus region shares synteny with the 4<sup>th</sup> chromosome of <i>C. neoformans</i> JEC21.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-05-20 02:32:34"}
  • {"files"=>["https://ndownloader.figshare.com/files/848890"], "description"=>"<p>(A) Alignment of the homeodomain regions of <i>S. cerevisiae</i> (Sc), <i>C. cinerea</i> (Cc), <i>C. neoformans</i> var. <i>neoformans</i> (Cnn), <i>C. neoformans</i> var. <i>grubii</i> (Cng), <i>C. gattii</i> (Cg), <i>C. heveanensis</i> (Ch) homeodomain proteins. These proteins are classified as either HD1 or HD2 according to the length of the homeodomain region. There are three extra amino acids between helix 1 and helix 2 of HD1 proteins. <i>C. heveanensis</i> Sxi1 and Sxi2 proteins are of type HD1 and HD2, respectively. (B) Schematic representation of the Sxi1 and Sxi2 domain structures. DD, HD, and NLS denote dimerization domain, homeodomain, and nuclear localization signal, respectively. The types of nuclear localization signal are indicated in parentheses: Bipartite and pat7, pat7 denotes the 7-residue pattern.</p>", "links"=>[], "tags"=>["sxi2", "align", "hd1", "hd2", "homeodomain"], "article_id"=>519337, "categories"=>["Microbiology", "Evolutionary Biology", "Genetics", "Molecular Biology"], "users"=>["Banu Metin", "Keisha Findley", "Joseph Heitman"], "doi"=>"https://dx.doi.org/10.1371/journal.pgen.1000961.g006", "stats"=>{"downloads"=>0, "page_views"=>16, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Sxi1_and_Sxi2_from_C_heveanensis_align_with_HD1_and_HD2_homeodomain_proteins_respectively_/519337", "title"=>"Sxi1 and Sxi2 from <i>C. heveanensis</i> align with HD1 and HD2 homeodomain proteins, respectively.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-05-20 02:35:37"}
PMC Usage Stats 203817 Jul 12:35 UTC

PMC Usage Stats | Further Information

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Relative Metric 93439223 Feb 17:38 UTC

Relative Metric

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F1000Prime 105 Aug 10:57 UTC

F1000Prime | Further Information

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