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. 2019 Mar 28;9:5302. doi: 10.1038/s41598-019-40457-1

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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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GNDF augments mitosis in caudal nephric duct at the time of ureteric bud outgrowth. (A) Mitotic indices were determined as the percentage of pHH3-positive cells within b-catenin or E-cadherin positive epithelium. Graphs show indices of wild type (WT, black bars) and Gdnf ^hyper/hyper embryos (white bars) in caudal nephric duct (E10.5), ureteric bud epithelium (E11.5) and ureteric bud tips (E15.5). A statistically significant increase in mitotic indices was observed at E10.5 samples (7–9 vibratome sections per embryo, n = 3/genotype, p < 0.05, Student’s t-test). (B) Total cell counts in E11.5 WT and Gdnf ^hyper/hyper ureteric bud tips (p < 0.05). (C) Representative E10.5 WT and (D) Gdnf ^hyper/hyper kidney primordia shown after staining with pHH3 (red) and b-catenin (green). (E) E11.5 WT kidney shows typical T-shaped ureteric bud (E-cadherin, green), but (F) Gdnf ^hyper/hyper epithelium is abnormally enlarged without normal branching pattern. Arrows in (D) and (F) point to cell mass accumulated in the luminal side of the epithelium. (G) Regardless of the genotype, labeling for cleaved Caspase3 does not reveal apoptosis in the kidney primordia of E10.5 embryos (n = 7) while a statistically significant increase is seen at E11.5 in Gdnf ^hyper/hyper epithelium (white bar, n = 5 for WT and 4 for Gdnf ^hyper/hyper, Student’s t-test, p < 0.001). At E15.5 cleaved caspase 3 labeling reveals similar amount of apoptotic cells in the cortex of WT (n = 3) and Gdnf ^hyper/hyper (n = 2) kidneys. (H) An example of WT kidney stained with cleaved Caspase3 (red, n = 5) shows virtually no apoptosis, while (I) numerous apoptotic cells (arrowhead) are detected in the ureteric bud lumen of Gdnf ^hyper/hyper embryos at E11.5 (n = 4). Scale bars 50 µm. Error bars on all graphs represent standard deviation.