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. 2026 Jan 3;17:753. doi: 10.1038/s41467-025-67446-5

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© The Author(s) 2026

Open Access This article is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License, which permits any non-commercial use, sharing, 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 you modified the licensed material. You do not have permission under this licence to share adapted material derived from this article or parts of it. 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-nc-nd/4.0/.

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Fig. 6 — A Schematic of the four motif mutations on the CD28 intracellular domain. B Dose response curve showing IFNγ and TNFα levels in CD4 T_TCR-MA1 CCR cells incubated with decreasing concentrations of cognate peptide overnight. C NSG mice were engrafted with 1 × 10⁵ A375F and subsequently infused with total 5 × 10⁶ CD4 and CD8 T cells (T_{TCR-MA1-CD8αβ}, T_{TCR-MA1-CD8/CD28}, and T_{TCR-MA1-CD8/CD28mut}) at a ratio of 1:1 on day 12 post engraftment. Graph represents mean ± SEM. (n = 3 mice/group, 2 tumors/mouse, data shown are representative of two independent experiments). D NSG mice were engrafted with 1 × 10⁵ A375F and subsequently infused with total 2.5 × 10⁶ CD4 and CD8 T cells (T_{TCR-Irr.-CD8/28}, T_{TCR-MA1-CD8/CD28}, and T_{TCR-MA1-CD8/CD28mut}) at a ratio of 1:1 on day 12 post engraftment. (n = 3mice/group, 2 tumors/mouse, data shown are representative of two independent experiments.) The graph represents mean ± SEM. E A375F tumor volume assessed at sacrifice 9 days after T cell infusions for each indicated condition same as (D). p value determined by one-way ANOVA with Tukey’s multiple comparison test. The graph represents mean ± SEM. (n = 4 mice for T_{TCR-Irr.-CD8/28} group and n = 5 mice for T_{TCR-MA1-CD8/CD28}, and T_{TCR-MA1-CD8/CD28mut} groups, data shown are representative of two independent experiments). F Growth kinetics of the SK-MEL-37 cell line (left) in the absence (black lines) or presence of CD4 T cells with the indicated CCR constructs (T_{TCR-Irr.-CD8/CD28}, T_{TCR-MA1-CD8αβ}, T_{TCR-MA1-CD8/CD28}, and T_{TCR-MA1-CD8/CD28mut}) at a ratio of E:T 5:1 in a live tumor visualization assay (Incucyte S3). Final lymphocyte counts (right) were obtained after 84 h. The graph represents mean ± SD. p-values determined by one-way ANOVA with Tukey’s multiple comparison test. (n = 3 wells/group). G Non-humanized MISTRG mice were engrafted with 1 × 10⁶ SK-MEL-37 (left) and subsequently infused with a total of 2 × 10⁷ CD4 and CD8 T cells (T_{TCR-Irr.-CD8/28}, T_{TCR-MA1-CD8/CD28}, and T_{TCR-MA1-CD8/CD28mut}) at a ratio of 1:1 on day 21 post engraftment (n = 4 mice, 2 tumors per mouse). SK-MEL-37 tumor volume (right) assessed at 62 days after T cell infusions for each indicated condition. The graph represents mean ± SEM. p-value determined by a two-tailed unpaired t-test. (n = 4 mice/group, 2 tumors/mouse averaged). Panels C,D, and G were created in BioRender. Tang, A. (2026) https://BioRender.com/6wg9k7f.