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. 2023 Nov 29;14:7863. doi: 10.1038/s41467-023-43670-9

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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 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. 3 — a, b qPCR analysis of Spink1/SPINK1 expression a in fetal mouse livers and b from human hepatocyte differentiation model at different developmental stages. ES Embryonic stem cell, EN Endoderm, HP Hepatic progenitor cell, LP Liver progenitor cell, PH Premature hepatocytes, HEP Mature hepatocytes. c, d Analysis of a human liver development single-cell atlas²⁶ showed c SPINK1, hepatic progenitor and mature hepatocyte markers expression in HB/FH and AH clusters. Dot size = gene expression frequency; color intensity = expression level, and d UMAP of (top) CytoTRACE predicted order (more to less differentiated) and (bottom) SPINK1 expression (low to high), indicated by color from blue to red. e SPINK1 expression in TCGA-LIHC cohort, stratified by histological grading (Edmondson-Steiner 4-tier). NT non-tumor, G1 well differentiated, G2 moderately differentiated, G3 poorly differentiated, G4 undifferentiated. f–g Patients in TCGA-LIHC cohort were ranked based on SPINK1 expression. High SPINK1 = top 10% SPINK1; low SPINK1 = bottom 10% SPINK1. f Heatmap showing the clustering of SPINK1 expression with hepatic progenitor and mature hepatocyte markers. g SPINK1 high tumors showed enrichment of stem cell-related gene sets^28,29 and undifferentiated cancer, via Gene Set Enrichment Analysis (GSEA). NES = Normalized enrichment scores; FDR = False discovery rate. h (Left) TCGA-LIHC dataset analysis of SPINK1 expression with high and low transcriptomic stemness score (TSS). (Right) Kaplan-Meier curve showing the overall survival in different TSS/SPINK1 groups. i (Left) SPINK1 protein expression in paired T and NT HCC clinical biopsies were examined by immunohistochemistry and expression intensities were scored. Kaplan-Meier analysis of the overall survival in patients with high and low SPINK1 scoring in their tumors. (Right) Representative SPINK1 staining images in paired T and NT samples of two HCC patients. Scale bar in low magnification = 100 μm; high magnification = 50 μm. j SPINK1 transcriptomic level in paired T and NT of NASH-related HCCs. a 2 replicates; b n = 1 experiment; j n = 15 paired T and NT samples. Data were expressed as mean ± s.e.m. Statistical analysis: g Kolmogorov–Smirnov test, h two-sided Unpaired Student’s t-test, h, i log-rank test, or j two-sided Paired Student’s t-test. e Data was expressed as a range (min to max), median as the centre, and distribution shape was shown. Statistical analysis: one-way ANOVA. Source data are provided as a Source Data file.