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. 2017 Sep 25;8:587. doi: 10.1038/s41467-017-00597-2

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© The Author(s) 2017

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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Fig. 5 — The absence of a brain generates an abnormal neural network in the entire animal body. a, b Acetylated-tubulin (Tub) immunoexpression for Ctrl (a) and BR⁻ (b) animals. There three types of nerve fibers: (i) commissural fibers (dorsoventral axis, long arrows); (ii) longitudinal fibers (anteroposterior axis, short arrow); and (iii) internal neuropil (no defined axis, unfilled triangles). Animals developed without a brain show normal commissural and longitudinal nerve fibers (turquoise long arrows in b), with some alterations (magenta long arrow), but a dense internal neuropil (yellow unfilled triangles). c, d Tub-immunoexpression for BR⁻ treated with cholinergic drugs: scopolamine (c) and carbachol (d). Scopolamine treatment was not able to rescue the aberrant internal network (yellow unfilled triangles in c), and carbachol-treated animals exhibited a chaotic nerve patterning (magenta and yellow arrows in d). e Ectopic HCN2-WT expression (injected in both cells at two-cell stage) fixed the BR⁻–induced internal nerve branching. f Quantification of the mean OD of internal neuropil and statistical comparisons among untreated/uninjected Ctrl and untreated/uninjecetd BR⁻ (BR⁻, without drug treatment nor ion channel misexpression), scopolamine-treated BR⁻ (BR⁻ + scopolamine), carbachol-treated BR⁻ (BR⁻ + carbachol), HCN2-WT both-sides injected BR⁻ (BR⁻ + HCN2 WT), and HCN2-WT LR side-injected BR⁻ (BR⁻ + 1/2 HCN2 WT) embryos (one-way ANOVA, P < 0.01). No significant differences after a posteriori analysis were detected among the different Ctrl groups. Data represent the mean OD units and s.d. of two independent replicates. Number in bars indicates n or number of animals analyzed for each group. P values after post-hoc Bonferroni’s test are indicated as **P < 0.01, *P < 0.05, ns P > 0.05. g, h. Ectopic HCN2 expression in only one LR side fixes the BR⁻-induced internal nerve branching. Tub-immunoexpression on β-gal-reacted sections (dark deposits) in a 1/2 HCN2-WT BR⁻, showing both the contralateral uninjected side (g) and the injected (h) of the same embryo. Aberrant neural network was completely rescued (turquoise arrows), exhibiting a similar nerve pattern to the Ctrl group in both sides. a-e, h: Rostral is upper right and dorsal is up. g: Rostral is upper left and dorsal is up. Scale bar, 100 μm