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. 2019 Nov 1;9:15864. doi: 10.1038/s41598-019-52436-7

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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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Pedigree of the two families analyzed in this study, and morphological and ultrastructural analysis of patient spermatozoa. (a) Filled and open symbols indicate the affected individuals and their unaffected relatives, respectively. A dot in the middle of a symbol indicates a heterozygous carrier. Probands are indicated with black arrows. (b–e) Papanicolaou staining of spermatozoa smears from the seminal fluid of the normal control (NC) (b) and the three patients (P1, P2, and P3) (c–e). Morphological abnormalities of sperm flagella were observed (red arrows), including absent, short, bent, coiled, and irregular flagella. Scale bars = 50 μm. (f–m) Ultrastructure analysis of spermatozoa obtained from the NC and patients by TEM. (f–i) Axonemal cross-sections of sperm flagella in NC, P1, P2, and P3. Absence of central microtubules and peripheral microtubule doublets (red arrow) or disorganization of outer dense fibers (red arrowhead) were observed. (j–m) Longitudinal sections of sperm flagella in NC, P1, and P2 showed that the disorganized fibrous sheath or mitochondrial sheath, and a lack of axonemal CPC in the sperm flagella of patients, and tails with a cytoplasmic mass were shown (green arrowhead). The sperm neck area including proximal centriole, striated column, vault (blue arrowhead) are visible. Scale bars = 0.1 μm.