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. 2019 Mar 1;9:3226. doi: 10.1038/s41598-019-39530-6

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© The Author(s) 2019

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

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Hbs1 is required for male meiosis. (A) In wild-type cysts, each spermatid contained one nebenkern (dark) to one nucleus (light) (1:1 in ratio) of similar sizes. (B,C) In some of the Hbs1^1−/− mutant cysts, heterogeneous mixture of ratios of nebenkern to nucleus (from 1:1, 1:2, 1:3 to 1:4) and abnormal large nebenkern(arrow heads) with small or micronuclei(arrows) were frequently observed, which was evidence of failure of cytokinesis and chromosome mis-segregation (See Table 1 for the quantitative data). (D) The meiotic figure appeared normal in Hbs1^48−/− mutant cysts. (E) Normal onion stage spermatids were observed in the Hbs1 genomic rescued testis. (F) There was greater percentage of total irregular spermatids in the Hbs1^1−/− testes. (G,H) The wildtype testis (G–G’) exhibited fully formed and well-focused spindle poles, and the condensed chromosomes were either at the metaphase plate (arrowhead) or beginning to separate (arrow). In Hbs1^1−/− testes, the spindle microtubules failed to converge into the poles and were splayed outward, and the chromosome could not be correctly separated (arrow).α-tubulin staining is in red, and DAPI is in blue. Scale bars = 10 μm.