{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,20]],"date-time":"2026-10-20T01:15:08Z","timestamp":1792458908734,"version":"4.5.0"},"reference-count":47,"publisher":"Oxford University Press (OUP)","issue":"4","license":[{"start":{"date-parts":[[2014,4,1]],"date-time":"2014-04-01T00:00:00Z","timestamp":1396310400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/journals\/pages\/open_access\/funder_policies\/chorus\/standard_publication_model"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2014,4,1]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>B chromosomes are small, heterochromatic chromosomes that are transmitted in a non-Mendelian manner. We have identified a stock of Drosophila melanogaster that recently (within the last decade) acquired an average of 10 B chromosomes per fly. These B chromosomes are transmitted by both males and females and can be maintained for multiple generations in a wild-type genetic background despite the fact that they cause high levels of 4th chromosome meiotic nondisjunction in females. Most curiously, these B chromosomes are mitotically unstable, suggesting either the absence of critical chromosomal sites or the inability of the meiotic or mitotic systems to cope with many additional chromosomes. These B chromosomes also contain centromeres and are primarily composed of the heterochromatic AATAT satellite sequence. Although the AATAT sequence comprises the majority of the 4th chromosome heterochromatin, the B chromosomes lack most, if not all, 4th chromosome euchromatin. Presumably as a consequence of their heterochromatic content, these B chromosomes significantly modify position-effect variegation in two separate reporter systems, acting as enhancers of variegation in one case and suppressors in the other. The identification of B chromosomes in a genetically tractable organism like D. melanogaster will facilitate studies of chromosome evolution and the analysis of the mechanisms by which meiotic and mitotic processes cope with additional chromosomes.<\/jats:p>","DOI":"10.1534\/genetics.113.160556","type":"journal-article","created":{"date-parts":[[2014,1,30]],"date-time":"2014-01-30T05:39:19Z","timestamp":1391060359000},"page":"1007-1016","source":"Crossref","is-referenced-by-count":42,"title":["Discovery of Supernumerary<i>B<\/i>Chromosomes in<i>Drosophila melanogaster<\/i>"],"prefix":"10.1093","volume":"196","author":[{"given":"Elisabeth","family":"Bauerly","sequence":"first","affiliation":[{"name":"Stowers Institute for Medical Research , Kansas City, Missouri 64110"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stacie E","family":"Hughes","sequence":"additional","affiliation":[{"name":"Stowers Institute for Medical Research , Kansas City, Missouri 64110"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dana R","family":"Vietti","sequence":"additional","affiliation":[{"name":"Department of Child Health , University of Missouri School of Medicine, Columbia, Missouri 65201"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Danny E","family":"Miller","sequence":"additional","affiliation":[{"name":"Stowers Institute for Medical Research , Kansas City, Missouri 64110"},{"name":"Department of Molecular and Integrative Physiology , University of Kansas Medical Center, Kansas City, Kansas 66160"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"William","family":"McDowell","sequence":"additional","affiliation":[{"name":"Stowers Institute for Medical Research , Kansas City, Missouri 64110"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"R Scott","family":"Hawley","sequence":"additional","affiliation":[{"name":"Stowers Institute for Medical Research , Kansas City, Missouri 64110"},{"name":"Department of Molecular and Integrative Physiology , University of Kansas Medical Center, Kansas City, Kansas 66160"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2014,4,1]]},"reference":[{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"2536","DOI":"10.1105\/tpc.113.111856","article-title":"Formation and expression of pseudogenes on the B chromosome of rye.","volume":"25","author":"Banaei-Moghaddam","year":"2013","journal-title":"Plant Cell"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"270","DOI":"10.1016\/j.tig.2009.04.002","article-title":"Origin and evolution of Y chromosomes: Drosophila tales.","volume":"25","author":"Bernardo Carvalho","year":"2009","journal-title":"Trends Genet."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"328","DOI":"10.1038\/hdy.1994.140","article-title":"Bewildering Bs: an impression of the 1st B-chromosome conference.","volume":"73","author":"Beukeboom","year":"1994","journal-title":"Heredity"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"505","DOI":"10.3389\/fpls.2013.00505","article-title":"Meiotic behavior of small chromosomes in maize.","volume":"4","author":"Birchler","year":"2013","journal-title":"Front Plant Sci"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1093\/genetics\/129.1.177","article-title":"Fine mapping of satellite DNA sequences along the Y chromosome of Drosophila melanogaster: relationships between satellite sequences and fertility factors.","volume":"129","author":"Bonaccorsi","year":"1991","journal-title":"Genetics"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"E1222","DOI":"10.1073\/pnas.1301690110","article-title":"Binding of Drosophila Polo kinase to its regulator Matrimony is noncanonical and involves two separate functional domains.","volume":"110","author":"Bonner","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1098\/rstb.2000.0556","article-title":"B-chromosome evolution.","volume":"355","author":"Camacho","year":"2000","journal-title":"Philos. Trans. R. Soc. Lond. B Biol. Sci."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1007\/s004120050309","article-title":"Characterization of sequences associated with position-effect variegation at pericentric sites in Drosophila heterochromatin.","volume":"107","author":"Cryderman","year":"1998","journal-title":"Chromosoma"},{"key":"2023030811105774200_"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1016\/S0092-8674(00)80084-7","article-title":"Direct evidence of a role for heterochromatin in meiotic chromosome segregation.","volume":"86","author":"Dernburg","year":"1996","journal-title":"Cell"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"793","DOI":"10.1093\/genetics\/122.4.793","article-title":"Position effect variegation in Drosophila melanogaster: relationship between suppression effect and the amount of Y chromosome.","volume":"122","author":"Dimitri","year":"1989","journal-title":"Genetics"},{"issue":"Pt 2","key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"463","DOI":"10.1242\/jcs.101.2.463","article-title":"Cloning and expression of Drosophila HP1 homologs from a mealybug, Planococcus citri.","volume":"101","author":"Epstein","year":"1992","journal-title":"J. Cell Sci."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"e1000234","DOI":"10.1371\/journal.pbio.1000234","article-title":"Species-specific heterochromatin prevents mitotic chromosome segregation to cause hybrid lethality in Drosophila.","volume":"7","author":"Ferree","year":"2009","journal-title":"PLoS Biol."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1159\/000079283","article-title":"Human supernumeraries: Are they B chromosomes?","volume":"106","author":"Fuster","year":"2004","journal-title":"Cytogenet. Genome Res."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"539","DOI":"10.1007\/BF00355318","article-title":"A species specific satellite DNA family of Drosophila subsilvestris appearing predominantly in B chromosomes.","volume":"103","author":"Gutknecht","year":"1995","journal-title":"Chromosoma"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1002\/bies.950180410","article-title":"Is the Y chromosome of Drosophila an evolved supernumerary chromosome?","volume":"18","author":"Hackstein","year":"1996","journal-title":"Bioessays"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"637","DOI":"10.1093\/genetics\/165.2.637","article-title":"A deficiency screen of the major autosomes identifies a gene (matrimony) that is haplo-insufficient for achiasmate segregation in Drosophila oocytes.","volume":"165","author":"Harris","year":"2003","journal-title":"Genetics"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"440","DOI":"10.1002\/dvg.1020130608","article-title":"There are two mechanisms of achiasmate segregation in Drosophila females, one of which requires heterochromatic homology.","volume":"13","author":"Hawley","year":"1992","journal-title":"Dev. Genet."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"310","DOI":"10.1016\/0168-9525(93)90249-H","article-title":"Requiem for distributive segregation: achiasmate segregation in Drosophila females.","volume":"9","author":"Hawley","year":"1993","journal-title":"Trends Genet."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"1539","DOI":"10.1534\/genetics.106.066407","article-title":"A Distinct type of heterochromatin within Drosophila melanogaster chromosome 4.","volume":"175","author":"Haynes","year":"2007","journal-title":"Genetics"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1007\/BF00327335","article-title":"Spontaneous translocations between B chromosomes and the normal complement in the grasshopper Eyprepocnemis plorans.","volume":"88","author":"Henriques-Gil","year":"1983","journal-title":"Chromosoma"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"e1002209","DOI":"10.1371\/journal.pgen.1002209","article-title":"Gamma-tubulin is required for bipolar spindle assembly and for proper kinetochore microtubule attachments during prometaphase I in Drosophila oocytes.","volume":"7","author":"Hughes","year":"2011","journal-title":"PLoS Genet."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"e1000348","DOI":"10.1371\/journal.pgen.1000348","article-title":"Heterochromatic threads connect oscillating chromosomes during prometaphase I in Drosophila oocytes.","volume":"5","author":"Hughes","year":"2009","journal-title":"PLoS Genet."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1016\/S1360-1385(03)00187-0","article-title":"B chromosomes in plants: Escapees from the A chromosome genome?","volume":"8","author":"Jones","year":"2003","journal-title":"Trends Plant Sci."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1126\/science.273.5271.118","article-title":"Centric heterochromatin and the efficiency of achiasmate disjunction in Drosophila female meiosis.","volume":"273","author":"Karpen","year":"1996","journal-title":"Science"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"5246","DOI":"10.1002\/j.1460-2075.1996.tb00910.x","article-title":"Misdivision analysis of centromere structure in maize.","volume":"15","author":"Kaszas","year":"1996","journal-title":"EMBO J."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1534\/genetics.105.043273","article-title":"Molecular evidence for transcription of genes on a B chromosome in Crepis capillaris.","volume":"171","author":"Leach","year":"2005","journal-title":"Genetics"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"234","DOI":"10.1038\/emboj.2008.283","article-title":"Sgt1, a co-chaperone of Hsp90 stabilizes Polo and is required for centrosome organization.","volume":"28","author":"Martins","year":"2009","journal-title":"EMBO J."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"13343","DOI":"10.1073\/pnas.1204237109","article-title":"Selfish supernumerary chromosome reveals its origin as a mosaic of host genome and organellar sequences.","volume":"109","author":"Martis","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1159\/000339615","article-title":"Accumulation of multiple copies of maize minichromosomes.","volume":"137","author":"Masonbrink","year":"2012","journal-title":"Cytogenet. Genome Res."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1007\/s10577-012-9283-2","article-title":"Multiple maize minichromosomes in meiosis.","volume":"20","author":"Masonbrink","year":"2012","journal-title":"Chromosome Res."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1159\/000166619","article-title":"Homolog pairing and segregation in Drosophila meiosis.","volume":"5","author":"McKee","year":"2009","journal-title":"Genome Dyn."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"680","DOI":"10.1007\/BF02007717","article-title":"Supernumerary chromosomes in Drosophila nasuta albomicana","volume":"41","author":"Ramachandra","year":"1985","journal-title":"Experientia"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1007\/BF00330354","article-title":"Characterization of heterochromatin in the B chromosomes of Drosophila nasuta albomicana","volume":"95","author":"Ramachandra","year":"1987","journal-title":"Chromosoma"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1016\/S1084-9521(02)00138-6","article-title":"Position-effect variegation and the genetic dissection of chromatin regulation in Drosophila.","volume":"14","author":"Schotta","year":"2003","journal-title":"Semin. Cell Dev. Biol."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"8210","DOI":"10.1128\/MCB.24.18.8210-8220.2004","article-title":"cis-Acting determinants of heterochromatin formation on Drosophila melanogaster chromosome four.","volume":"24","author":"Sun","year":"2004","journal-title":"Mol. Cell. Biol."},{"key":"2023030811105774200_","first-page":"211","article-title":"Accumulation of B-chromosomes in Drosophila kikkawai Burla.","volume":"80","author":"Sundaran","year":"1994","journal-title":"Cytobios"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"141","DOI":"10.3897\/compcytogen.v6i2.1769","article-title":"Cytogenetic analysis of B chromosomes in one population of the fish Moenkhausia sanctaefilomenae (Steindachner, 1907) (Teleostei, Characiformes).","volume":"6","author":"Voltolin","year":"2012","journal-title":"Comp Cytogenet"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"1263","DOI":"10.1101\/gad.9.10.1263","article-title":"Position effect variegation in Drosophila is associated with an altered chromatin structure.","volume":"9","author":"Wallrath","year":"1995","journal-title":"Genes Dev."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"93","DOI":"10.4161\/fly.20029","article-title":"H3S10 phosphorylation by the JIL-1 kinase regulates H3K9 dimethylation and gene expression at the white locus in Drosophila.","volume":"6","author":"Wang","year":"2012","journal-title":"Fly (Austin)"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"e1001648","DOI":"10.1371\/journal.pbio.1001648","article-title":"A meiosis-specific form of the APC\/C promotes the oocyte-to-embryo transition by decreasing levels of the Polo kinase inhibitor matrimony.","volume":"11","author":"Whitfield","year":"2013","journal-title":"PLoS Biol."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1007\/BF00293289","article-title":"Supernumerary microchromosomes identified as inverted duplications of chromosome 15: a report of three cases.","volume":"69","author":"Wisniewski","year":"1985","journal-title":"Hum. Genet."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1007\/BF00399391","article-title":"Cytogenetic and clinical studies in five cases of inv dup(15).","volume":"50","author":"Wisniewski","year":"1979","journal-title":"Hum. Genet."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1534\/genetics.106.061424","article-title":"The mechanism of secondary nondisjunction in Drosophila melanogaster females.","volume":"174","author":"Xiang","year":"2006","journal-title":"Genetics"},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"e323","DOI":"10.1371\/journal.pbio.0050323","article-title":"The inhibition of polo kinase by matrimony maintains G2 arrest in the meiotic cell cycle.","volume":"5","author":"Xiang","year":"2007","journal-title":"PLoS Biol."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"e1002203","DOI":"10.1371\/journal.pgen.1002203","article-title":"B chromosomes have a functional effect on female sex determination in Lake Victoria cichlid fishes.","volume":"7","author":"Yoshida","year":"2011","journal-title":"PLoS Genet."},{"key":"2023030811105774200_","doi-asserted-by":"crossref","first-page":"505","DOI":"10.1534\/genetics.110.118778","article-title":"Statistical analysis of nondisjunction assays in Drosophila.","volume":"186","author":"Zeng","year":"2010","journal-title":"Genetics"}],"container-title":["Genetics"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/genetics\/article-pdf\/196\/4\/1007\/49452162\/genetics1007.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/genetics\/article-pdf\/196\/4\/1007\/49452162\/genetics1007.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,7,10]],"date-time":"2023-07-10T04:08:43Z","timestamp":1688962123000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/genetics\/article\/196\/4\/1007\/5935621"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2014,4,1]]},"references-count":47,"journal-issue":{"issue":"4","published-online":{"date-parts":[[2014,4,1]]},"published-print":{"date-parts":[[2014,4,1]]}},"URL":"https:\/\/doi.org\/10.1534\/genetics.113.160556","relation":{},"ISSN":["1943-2631"],"issn-type":[{"value":"1943-2631","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2014,4,1]]},"published":{"date-parts":[[2014,4,1]]}}}