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. 2018 Jul 16;8:10703. doi: 10.1038/s41598-018-29109-y

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© The Author(s) 2018

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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Illustration of critically affected genes and their downstream effects that most closely fit the data presented in the present study. ZIP8 deficiency (top), has a major impact on TAL1 and GATA transcription factors (TFs), which appear to function primarily in the hematopoietic stem cells of Slc39a8(neo/neo) GD13.5 yolk sac. This function then causes severe dysregulation of hematopoietic stem cell fate and striking anemia in yolk sac, which is visibly obvious [in Fig. S2 of ref.¹³]. Moreover, hematopoiesis in yolk sac is well known to precede that in liver and then spleen and marrow [cf. Fig. S4 of ref.¹³]. Downstream effects, as development proceeds, include severe anemia and defects in coagulation, innate immunity, and response to inflammation. The striking anemia leads to a hypoxia response which is seen in all tissues examined, but largely in yolk sac. TAL1, T-cell acute lymphocytic leukemia protein-1 TF. GATA, family of six zinc-finger TFs that regulate hematopoietic stem cell fate. Interactions between TAL1 and GATA exist⁶¹ [see text]. ZIP8-deficiency, plus the result of all these downstream changes, alter the expression of nine Cyp genes and 27 Slc genes (excluding Slc39a8); these changes are mostly unique to one tissue, as detailed in Supplementary Table S1. Δ denotes “changes in”. HIF, hypoxia-inducible factor.