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. 2021 Apr 23;9:662903. doi: 10.3389/fcell.2021.662903

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Copyright © 2021 Hwang, Nawaz, Choi, Wang, Hussain, Nawaz, Lopez-Giraldez, Jeong, Dong, Oh, Bilguvar, Mane, Lee, Bystroff, Lifton, Ahmad and Chung.

This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

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Flagellar ultrastructure is disorganized, and axonemal proteins express abnormally in Dnah2-null sperm. (A,B) Scanning electron microscopy images of Dnah2^+/Δ (A) and Dnah2^Δ/Δ (B) sperm. Dnah2^Δ/Δ sperm display absence of (left) or an irregular (right) mitochondrial helical sheath in the midpiece of the sperm tail, along with abnormal head shapes. Arrowheads indicate annulus. H, head; MP, midpiece; and PP, principal piece. (C,D) Transmission electron microscopy images of Dnah2^+/Δ (C) and Dnah2^Δ/Δ (D) sperm. Representative cross-section images of TEM reveal conformational defects in the midpiece of Dnah2^Δ/Δ sperm. Incomplete 9+2 axonemal structure (left), delocalized outer dense fibers (ODF, arrow) and microtubule doublet (middle, arrowhead), and lack of the axonemal components (right) were observed in the Dnah2^Δ/Δ sperm. Inner (yellow asterisk) and outer (white asterisk) dynein arm structure were observed from microtubule doublet occasionally (left, inset; scale bar = 50 nm). Microtubule doublets are numbered with the absence of the corresponding microtubule doublet in red. M, mitochondria. (E) Confocal fluorescence images of axonemal proteins in WT and Dnah2^Δ/Δ sperm. Dnah2^Δ/Δ sperm do not express another IDA component (DNAH1, magenta) and a radial spoke protein (RSPH3, green) but an ODA protein (DNAH9, yellow). Flagella microtubules (AcTub, white) and sperm heads (Hoechst, blue) serve as controls. The corresponding DIC images are shown together for Dnah2^Δ/Δ sperm.