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. 2023 Oct 30;14:6929. doi: 10.1038/s41467-023-42643-2

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© The Author(s) 2023

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 licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence 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 licence, visit http://creativecommons.org/licenses/by/4.0/.

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Fig. 3 — a To probe whether YAP targets are sensitive to dynamic YAP inputs, ES cells expressing LEXY-YAP were exposed to light patterns with different pulse widths during pluripotency exit. b IF staining for Oct4 in LEXY-YAP mESCs upon illumination with chronic or pulsed light inputs demonstrates induction of Oct4 upon illumination as compared to the dark control, with higher Oct4 levels in the oscillatory than chronic YAP export condition. Scale bar: 20 µm. c, d Quantification of nuclear Oct4 (c) and Nanog (d) levels from IF staining as shown in b. Cells were subjected to light pulses inducing constant YAP export durations of 4 h and varying import duration ranging from 15 min to 4 h as indicated. Only cells with moderately high LEXY-YAP/YAP expression levels (>IC50 of dark control) were quantified (see Supplementary Fig. 6b for comparison to cells with lower YAP expression levels). Pulse frequencies with specific YAP export/import cycles (4 h export, 60 min import) significantly upregulate Oct4 and Nanog as compared to the non-light responsive YAP control or compared to chronic YAP export. Quantifications shown only include cells with YAP levels >= IC50 in the dark condition. Note that the chronic light condition is sufficient to induce Oct4 in cells with lower YAP levels (see Supplementary Fig. 6a, b). Note that the induction of Nanog by oscillatory inputs is minor. Shown are mean ± SEM, N = 7 independent experiments, p values comparing LEXY-YAP vs YAP from two-sided unpaired Student’s t test; p values comparing light conditions within the LEXY-YAP group from two-sided paired non-parametric t test. e Possible decoding logic of dynamic YAP inputs through adaptation. For an adaptive system, continuous YAP export would transiently activate YAP effectors (left panel). In contrast, pulsed YAP inputs would induce sequential adaptive YAP responses; this would result in higher total Oct4 induction (center panel) than is seen for chronic YAP export (left panel). The adaptive system gives a maximum output at a specific pulse frequency input (center) compared to faster (right panel) or slower (not shown) frequencies.