{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T20:44:44Z","timestamp":1785789884240,"version":"3.56.0"},"reference-count":36,"publisher":"Oxford University Press (OUP)","issue":"4","license":[{"start":{"date-parts":[[2018,4,1]],"date-time":"2018-04-01T00:00:00Z","timestamp":1522540800000},"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":[[2018,4,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n               <jats:p>Balancer chromosomes are multiply inverted and rearranged chromosomes used in Drosophila melanogaster for many tasks, such as maintaining mutant alleles in stock and complex stock construction. Balancers were created before molecular characterization of their breakpoints was possible, so the precise locations of many of these breakpoints are unknown. Here, we report or confirm the positions of the 14 euchromatic breakpoints on the 2nd chromosome balancers SM1, SM5, CyO, and SM6a. This total includes three breakpoints involved in a complex rearrangement on SM5 that is associated with the duplication of two genomic regions. Unbiased sequencing of several balancers allowed us to identify stocks with incorrectly identified balancers as well as single and double crossover events that had occurred between 2nd chromosome balancers and their homologs. The confirmed crossover events that we recovered were at least 2 Mb from the closest inversion breakpoint, consistent with observations from other balancer chromosomes. Balancer chromosomes differ from one another both by large tracts of sequence diversity generated by recombination and by small differences, such as single nucleotide polymorphisms (SNPs). Therefore, we also report loss-of-function mutations carried by these chromosomes and unique SNP and InDel polymorphisms present on only single balancers. These findings provide valuable information about the structure of commonly used 2nd chromosome balancers and extend recent work examining the structure of X and 3rd chromosome balancers. Finally, these observations provide new insights into how the sequences of individual balancers have diverged over time.<\/jats:p>","DOI":"10.1534\/g3.118.200021","type":"journal-article","created":{"date-parts":[[2018,2,8]],"date-time":"2018-02-08T09:47:16Z","timestamp":1518083236000},"page":"1161-1171","source":"Crossref","is-referenced-by-count":20,"title":["The Molecular and Genetic Characterization of Second Chromosome Balancers in <i>Drosophila melanogaster<\/i>"],"prefix":"10.1093","volume":"8","author":[{"given":"Danny E","family":"Miller","sequence":"first","affiliation":[{"name":"Stowers Institute for Medical Research, Kansas City, Missouri 64110"},{"name":"MD-PhD Physician\u2013Scientist Training Program and"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kevin R","family":"Cook","sequence":"additional","affiliation":[{"name":"Department of Biology, Indiana University, Bloomington, Indiana 47405, and"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Elizabeth A","family":"Hemenway","sequence":"additional","affiliation":[{"name":"Stowers Institute for Medical Research, Kansas City, Missouri 64110"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Vivienne","family":"Fang","sequence":"additional","affiliation":[{"name":"Department of Biology, Davidson College, North Carolina 28035"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Angela L","family":"Miller","sequence":"additional","affiliation":[{"name":"Stowers Institute for Medical Research, Kansas City, Missouri 64110"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Karen G","family":"Hales","sequence":"additional","affiliation":[{"name":"Department of Biology, Davidson College, North Carolina 28035"}],"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":[[2018,4,1]]},"reference":[{"issue":"1","key":"2021052005090501000_bib1","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1534\/genetics.107.083196","article-title":"Molecular population genetics of Drosophila subtelomeric DNA.","volume":"178","author":"Anderson","year":"2008","journal-title":"Genetics"},{"issue":"3","key":"2021052005090501000_bib2","doi-asserted-by":"publisher","first-page":"184","DOI":"10.4161\/fly.24466","article-title":"Genetic decay of balancer chromosomes in Drosophila melanogaster.","volume":"7","author":"Araye","year":"2013","journal-title":"Fly (Austin)"},{"issue":"2","key":"2021052005090501000_bib3","doi-asserted-by":"publisher","first-page":"745","DOI":"10.1508\/cytologia.FujiiJubilaei.745","article-title":"Correspondences between linkage maps and salivary chromosome structure, as illustrated in the tip of chromosome 2R of Drosophila melanogaster.","volume":"Fujii Jubil","author":"Bridges","year":"1937","journal-title":"Cytologia (Tokyo)"},{"key":"2021052005090501000_bib4","volume-title":"The Mutants of Drosophila melanogaster","author":"Bridges","year":"1944"},{"issue":"9","key":"2021052005090501000_bib5","doi-asserted-by":"publisher","first-page":"677","DOI":"10.1038\/nmeth.1363","article-title":"Breakdancer: an algorithm for high-resolution mapping of genomic structural variation.","volume":"6","author":"Chen","year":"2009","journal-title":"Nat. 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