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. 2023 Mar 14;16(3):dmm049858. doi: 10.1242/dmm.049858

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© 2023. Published by The Company of Biologists Ltd

This is an Open Access article distributed under the terms of the Creative Commons Attribution License (https://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution and reproduction in any medium provided that the original work is properly attributed.

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Fig. 1. — Spinal neurulation failure and enhanced pSmad-1,5,8 expression in Zic2^Ku/Ku mutants. (A-D) Defective spinal closure, as shown by the enlarged posterior neuropore (PNP) (outlined by white dashed lines; arrows in A,B), in Zic2^Ku/Ku embryos at E9.0 (B) and E9.5 (D) compared with normally developing wild-type (Zic2^+/+) littermates (A,C). (E) Somite level of the rostral limit of the PNP (asterisks in B,D) among 124 Zic2^Ku/Ku embryos at E9.5-10.5. Closure does not usually progress beyond the level of somites 12-17, at which level dorsolateral hinge points (DLHPs) first appear in normal development. (F,G) Transverse sections (Haematoxylin and Eosin-stained) through the rostral PNP of E9.5 Zic2^+/+ (F) and Zic2^Ku/Ku (G) embryos, at the level of the yellow dashed lines in C,D, respectively. Both show equivalent neural fold elevation, but paired DLHPs are present in the Zic2^+/+ embryo (arrows in F) and absent from the Zic2^Ku/Ku embryo (G). The mutant neuroepithelium shows marked apicobasal thickening (double-headed arrow between dashed lines in G). (H-K) Immunohistochemistry for pSmad-1,5,8 on transverse sections through the elevated neural folds of Zic2^+/+ (H) and Zic2^Ku/Ku (I-K) embryos. In non-cultured (NonC) 15-somite-stage embryos (H,I), immunoreactivity is enhanced in the dorsal Zic2^Ku/Ku neuroepithelium (arrows in I), compared with that in Zic2^+/+ embryos, in which expression is mainly in surface ectoderm (arrows in H). Zic2^Ku/Ku embryos exposed for 16 h in culture to vehicle (Con) from E8.5 show pSmad-1,5,8 immunoreactivity comparable to that seen in non-cultured Zic2^Ku/Ku embryos (compare J with I). Dorsomorphin exposure in culture produces markedly reduced pSmad-1,5,8 expression (K), similar to that in non-cultured Zic2^+/+ embryos. NE, neuroepithelium; SE, surface ectoderm. Scale bars: 0.5 mm (A-D); 50 µm (F); 50 µm (I).