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. 2021 Feb 8;7:2. doi: 10.1038/s41526-020-00130-8

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© The Author(s) 2021

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. 2 — a Mechanical negative feedback maintaining YAP activity in a cell. b YAP-mediated 3D organ/tissue formation withstanding gravity. In (a), YAP/TAZ acts as a mechanotransducer and mechanoeffector. As a mechanotransducer, it provides physical inputs, including gravity activation of YAP/TAZ that leads to an expansion of organ size. As a mechanoeffector, it activates YAP, which, in turn, controls F-actin turnover, leading to the suppression of YAP as part of a negative feedback mechanism. F-actin turnover controls the cell/tissue tension that mediates 3D organogenesis. b YAP is essential for the formation of complex 3D organs by coordinating 3D tissue shape (left) and tissue alignment (right). In response to external forces, including gravity, YAP activates (1) ARHGAP18 expression, which mediates (2) contractile actomyosin formation controlling (3) tissue tension. Tissue tension is required for both (4) cell stacking to form a 3D tissue shape and (5) fibronectin assembly required for adjacent tissue alignment, e.g., the alignment of the lens and eye-cup.