WJGO

World Journal of Gastrointestinal Oncology

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DOI: 10.4251/wjgo.v16.i2.475

World J Gastrointest Oncol 2024 February 15; 16(2): 475-492 ISSN 1948-5204 (online)

ORIGINAL ARTICLE

Basic Study Comprehensive analysis of the protein phosphatase 2A regulatory subunit B56E in pan-cancer and its role and mechanism in hepatocellular carcinoma

Hong-Mei Wu, Yuan-Yuan Huang, Yu-Qiu Xu, Wei-Lai Xiang, Chang Yang, Ru-Yuan Liu, Di Li, Xue-Feng Guo, Zheng-Bao Zhang, Chun-Hua Bei, Sheng-Kui Tan, Xiao-Nian Zhu

Specialty type: Oncology

Provenance and peer review: Invited article; Externally peer reviewed.

Peer-review model: Single blind

Peer-review report’s scientific quality classification

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Grade B (Very good): B

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P-Reviewer: Shalaby MN, Egypt

Received: October 17, 2023 Peer-review started: October 17, 2023

First decision: December 5, 2023

Revised: December 10, 2023

Accepted: January 8, 2024

Article in press: January 8, 2024 Published online: February 15, 2024

Hong-Mei Wu, Yuan-Yuan Huang, Yu-Qiu Xu, Wei-Lai Xiang, Chang Yang, Ru-Yuan Liu, Di Li, Xue- Feng Guo, Zheng-Bao Zhang, Chun-Hua Bei, Sheng-Kui Tan, Xiao-Nian Zhu, Guangxi Key Laboratory of Environmental Exposomics and Entire Lifecycle Health, Guilin Medical University, Guilin 541199, Guangxi Zhuang Autonomous Region, China

Corresponding author: Xiao-Nian Zhu, MD, Professor, Guangxi Key Laboratory of Environmental Exposomics and Entire Lifecycle Health, Guilin Medical University, No. 1 Zhiyuan Road, Guilin 541199, Guangxi Zhuang Autonomous Region, China. zhuxiaonian0403@163.com

Abstract

BACKGROUND

B568 is a regulatory subunit of the serine/threonine protein phosphatase 2A, which is abnormally expressed in tumors and regulates various tumor cell functions. At present, the application of B568 in pan-cancer lacks a comprehensive analysis, and its role and mechanism in hepatocellular carcinoma (HCC) are still unclear.

AIM

To analyze B568 in pan-cancer, and explore its role and mechanism in HCC.

METHODS

The Cancer Genome Atlas, Genotype-Tissue Expression, Gene Expression Profiling Interactive Analysis, and Tumor Immune Estimation Resource databases were used to analyze B568 expression, prognostic mutations, somatic copy number alterations, and tumor immune characteristics in 33 tumors. The relationships between B568 expression levels and drug sensitivity, immuno- therapy, immune checkpoints, and human leukocyte antigen (HLA)-related genes were further analyzed. Gene Set Enrichment Analysis (GSEA) was performed to reveal the role of B568 in HCC. The Cell Counting Kit-8, plate cloning, wound healing, and transwell assays were conducted to assess the effects of B568 interference on the malignant behavior of HCC cells.

RESULTS

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In most tumors, B56 expression was upregulated, and high B568 expression was a risk factor for adrenocortical cancer, HCC, pancreatic adenocarcinoma, and pheochromocytoma and paraganglioma (all P < 0.05). B568 expression levels were correlated with a variety of immune cells, such as T helper 17 cells, B cells, and macro- phages. There was a positive correlation between B568 expression levels with immune checkpoint genes and HLA- related genes (all P < 0.05). The expression of B568 was negatively correlated with the sensitivity of most chemotherapy drugs, but a small number showed a positive correlation (all P < 0.05). GSEA analysis showed that B568 expression was related to the cancer pathway, p53 downstream pathway, and interleukin-mediated signaling in HCC. Knockdown of B568 expression in HCC cells inhibited the proliferation, migration, and invasion capacity of tumor cells.

CONCLUSION

B568 is associated with the microenvironment, immune evasion, and immune cell infiltration of multiple tumors. B568 plays an important role in HCC progression, supporting it as a prognostic marker and potential therapeutic target for HCC.

Key Words: B568; Prognosis; Tumor microenvironment; Immune infiltration; Immunotherapy; Hepatocellular carcinoma

@The Author(s) 2024. Published by Baishideng Publishing Group Inc. All rights reserved.

Core Tip: The expression of protein phosphatase 2A (PP2A) subunit B568 is up-regulated in most tumors, and its high expression is a risk factor for adrenocortical cancer, hepatocellular carcinoma (HCC), pancreatic adenocarcinoma, and pheochromocytoma and paraganglioma. B568 expression levels correlate with immune cells, immune checkpoint genes, human leukocyte antigen-related genes, and the sensitivity of chemotherapy drugs. In HCC, B568 expression is related to the cancer pathway. Knockdown of B568 expression in HCC cells can inhibit the proliferation, migration and invasion capacity of tumor cells. Our study supports PP2A subunit B568 as a prognostic marker and potential therapeutic target for HCC.

Citation: Wu HM, Huang YY, Xu YQ, Xiang WL, Yang C, Liu RY, Li D, Guo XF, Zhang ZB, Bei CH, Tan SK, Zhu XN. Compre- hensive analysis of the protein phosphatase 2A regulatory subunit B568 in pan-cancer and its role and mechanism in hepatocellular carcinoma. World J Gastrointest Oncol 2024; 16(2): 475-492 URL: https://www.wjgnet.com/1948-5204/full/v16/12/475.htm

DOI: https://dx.doi.org/10.4251/wjgo.v16.i2.475

INTRODUCTION

Hepatocellular carcinoma (HCC) is a global health challenge with a rising incidence. In the 2020 Global Cancer Statistics, HCC ranked sixth in the incidence and third in the mortality of cancers worldwide[1]. As the main histological type of liver cancer, HCC is the cause of the vast majority of liver cancer diagnoses and deaths[2,3]. The incidence of HCC varies by geography, with about 72% of cases occurring in Asia[4]. Its onset and progression are a multistep process associated with multiple risk factors, such as hepatitis B virus (HBV), hepatitis C virus, and the environment[5]. It is also controlled by genetic and epigenetic changes that inactivate tumor suppressor genes or activate oncogenes, ultimately leading to the occurrence of HCC[6]. Although after decades of exploration, we have some understanding of the molecular mechanism by which HCC occurs, the detailed pathogenesis is still poorly understood. Given the increasing rate of mortality from HCC worldwide, it is important to improve our understanding of the molecular mechanisms underlying the pathogenesis of HCC. Moreover, new diagnostic, prognostic biomarkers and therapeutic strategies are urgently needed to address this major public health issue.

Protein phosphatase 2A (PP2A) is a major serine/threonine phosphatase. It is involved in the feedback of multiple signaling pathways, affecting cell cycle progression, proliferation, transcription, and translation[7]. The PP2A holoenzyme complex contains scaffold subunit “A”, regulatory subunit “B”, and catalytic subunit “C”. The A and C subunits constitute the core enzyme, while the B subunit is responsible for regulating substrate specificity, cell loca- lization, and enzyme activity of PP2A holoenzyme trimers[8]. Studies have shown that PP2A can participate in the occurrence and development of HCC[9,10]. The PP2A B56 subfamily has five different subtypes a, B, y, 8, and ¿[11].

Recently, the B56 protein has become widely recognized and valued because of its role in the development of a variety of tumors such as melanoma[12], breast cancer[13], and prostate cancer[14]. The B56a (PPP2R5A), B56ß (PPP2R5B), and B568 (PPP2R5E) of the B56 subfamily have nuclear output signals at the C-terminus, resulting in the migration of the PP2A complex into the cytoplasm. B56y (PPP2R5C) and B568 (PPP2R5D) are mainly found in the nucleus because they lack the signal sequence at the C-terminus. One study shows that knockdown of B56y could promote xenograft tumor growth and HBV-mediated migration and invasion of HCC cells in vivo[15]. Mice lacking B568 spontaneously develop HCC, which is associated with increased carcinogenicity of c-Myc[16].

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As a member of the regulatory subunit B56 subfamily, the role of B568 in tumors has also attracted much attention. In a study of acute myeloid leukemia (AML), B568 was shown to induce caspase-dependent apoptosis by impairing cell prolif- eration, affecting the activation state of AKT and reducing the colony formation capacity of leukemia cells[17]. A study also showed that B568 can inhibit the growth of gastric cancer cells and induce cell apoptosis[18]. These results are consistent with its tumor suppressor properties in breast cancer[19] and human tongue squamous cell carcinoma[20]. Although B56 has been studied in some cancers, the mechanism of action of B56 in HCC is poorly understood.

To further explore the role and mechanism of B56 in the development and progression of HCC, we conducted a combination of bioinformatics analyses and cell experiment validation in this study. The Cancer Genome Atlas (TCGA), Genotype-Tissue Expression (GTEx), Gene Expression Profiling Interactive Analysis (GEPIA), and Tumor Immune Estimation Resource (TIMER) databases were used to analyze B568 expression, prognostic mutations, somatic copy number alterations (sCNAs), and tumor immune characteristics in 33 tumors. The relationships between B568 expression levels and drug sensitivity, immunotherapy, immune checkpoints, and human leukocyte antigen (HLA)-related genes were further analyzed. Gene Set Enrichment Analysis (GSEA) was performed to reveal the role of B568 in HCC. The Cell Counting Kit-8 (CCK-8), plate cloning, wound healing, and transwell assays were conducted to show the effects of B568 interference on the malignant behaviors of HCC cells. The findings from our study will provide insight into the potential value of B568 in the diagnosis, prognosis and treatment of HCC.

MATERIALS AND METHODS

Acquisition of data

RNA sequencing (RNA-seq) data in Trusted Platform Module (TPM) format for TCGA and GTEx were downloaded from the online website UCSC XENA (https://xenabrowser.net/datapages/), which was uniformly processed by the Toil process. Data corresponding to TCGA for 33 tumors and normal tissue data corresponding to GTEx were extracted. The 33 tumors included adrenocortical carcinoma (ACC), bladder urothelial carcinoma, breast invasive carcinoma (BRCA), cervical squamous cell carcinoma (CESC), cholangiocarcinoma (CHOL), colon adenocarcinoma (COAD), lymphoid neoplasm diffuse large B cell lymphoma (DLBC), esophageal carcinoma (ESCA), glioblastoma (GBM), head and neck squamous cell carcinoma, kidney chromophobe (KICH), kidney clear cell carcinoma (KIRC), kidney renal papillary cell carcinoma (KIRP), AML, low-grade glioma (LGG), HCC, lung adenocarcinoma (LUAD), lung squamous cell carcinoma (LUSC), mesothelioma, ovarian serous cystadenoma (OV), pancreatic adenocarcinoma (PAAD), pheochromocytoma and paraganglioma (PCPG), prostate adenocarcinoma (PRAD), rectal adenocarcinoma, sarcoma, skin cutaneous melanoma (SKCM), stomach adenocarcinoma (STAD), testicular germ cell tumor (TGCT), thyroid carcinoma (THCA), thymic carcinoma (THYM), endometrial cancer (UCEC), uterine carcinosarcoma (UCS), and uveal melanoma (UVM). Statistical analyses were performed with R software (version 4.2.1).

The RNA-seq data of the STAR process of the TCGA-Liver HCC (LIHC) project was downloaded and collated from TCGA database (https://portal.gdc.cancer.gov) and extracted in TPM format. Data from the paracancerous and carcinoma samples with corresponding number pairs were extracted and statistical analysis was performed with R.

Differential analysis of B56& expression in different tumors

The expression of B56 was analyzed in 33 tumors by the TIMER database (https://cistrome.shinyapps.io/timer/). RNA- seq expression data were statistically analyzed and visualized using R packages, ggplot2 (3.3.6), stats (4.2.1), and car. Expression profiles of B568 protein levels in HCC and corresponding immunohistochemical images were obtained through an online Human Protein Atlas (HPA) (http://www.proteinatlas.org/) database.

Analysis of the prognostic value of B56E in human cancer

The correlation between B568 expression and survival in pan-carcinoma (http://dna1.bio.kyutech.ac.jp/PrognoScan/ index.html) was analyzed. Univariate survival analysis was used to calculate the hazard ratio (HR) and 95% confidence interval of B568 in 33 tumors, and the results are shown as forest plots. The prognostic value of B568 expression in HCC was analyzed using GEPIA (http://gepia.cancer-pku.cn/). GEPIA is an interactive online platform that provides information on tumor samples from TCGA as well as normal sample information from TCGA and GTEx projects. The expression levels of B568 in cancerous and non-cancerous tissues were divided into a B568 high expression group and B568 low expression group according to the median.

Relationship between B56& and tumor mutation/immunity

The association of B568 expression with mutations and sCNAs in 33 tumors was first analyzed through the TIMER (http:/ / cistrome.org/TIMER/) database. Second, the expression of B568 was analyzed in six types of immune-infiltrating cells, namely B cells, CD4+ T cells, CD8+ T cells, neutrophils, macrophages, and dendritic cells (DCs). The relationship between B568 expression levels and tumor purity was also determined. After a general analysis of immune cell types, the correlation between B568 expression and multiple immunomarkers was analyzed to identify the potential subtypes of infiltrating immune cells. In addition, the relationship between B568 expression and immune checkpoint genes/HLA- associated genes were determined. Single-sample GSEA (ssGSEA), TIMER, and ESTIMATE were used to analyze the differences between tumor infiltrating immune cells and B56 expression levels. Drug susceptibility data were obtained from CellMiner with a screening criterion of P < 0.05 and correlation analysis was conducted using the Pearson’s test. The above data were statistically analyzed and the results were visualized in the form of box plots, heat maps, scatter plots,

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violin plots, or stick charts using the ggplot2 (3.3.6), stats (4.2.1), and car packages. The role of B568 expression in HCC immunity was explored through the TISIDB (http://cis.hku.hk/TISIDB) website.

GSEA

Based on “c2.cp.all.v2022.1.Hs.symbols.gmt [All Canonical Pathways] (3050)”, GSEA was performed from both Gene Ontology (GO) and Kyoto Encyclopedia of Genes and Genomes (KEGG) dimensions to explore the potential biological function of B568 expression in HCC. The molecules of the input data were wrapped with R “org. Hs.eg.db” after ID conversion, then the “clusterProfile” package was used for GSEA analysis, and finally the “ggplot2” package was used to visualize the results.

Cell lines and cell culture

The human hepatic cell line L02 and two types of human HCC cell lines (Hep3B and SK-Hep-1) obtained from the Cell Bank of the Chinese Academy of Sciences (Shanghai, China) were maintained in RPMI 1640, Dulbecco’s Modified Eagle Medium, and Minimum Essential Medium (Gibco, El Paso, TX, United States) with 10% fetal bovine serum (FBS) (OriCell, Guangzhou, China). Cells were cultured in a humidified incubator at 37 ℃ and 5% CO2.

Cell transfection

Hep3B and SK-Hep-1 cells were digested and seeded in 6-well plates after the logarithmic growth phase was reached. When the degree of cell confluency reached 80%, transfection was performed using Lipofectamine®2000 (Invitrogen, Carlsbad, CA, United States) according to the manufacturer’s protocol.

Quantitative polymerase chain reaction

The total RNA of cells was extracted with Trizol reagent and reverse transcribed into cDNA using MonScript™ RTIII All- in One Mix (Monad, Shanghai, China) with dsDNase. GAPDH served as the internal reference control to quantify the relative expression of the B568, which was calculated using the 2-44℃ method. The B568 primer sequences were as follows: Forward primer of 5’-GACTTCATGGACACGCTAT-3’ and reverse primer of 5’-CATTCTAACTACTTCAGGGTAA-3’. The GAPDH primer sequences were as follows: forward primer of 5’-ACAACTTTGGTATCGTGGAAGG-3’ and reverse primer of 5’- GCCATCACGCCACAGTTTC-3’.

Western blot analysis

Total cell protein was extracted using lysis buffer [10 µL PMSF, 1 mL RIPA Histiocytes Lysate Buffer (strong), 10 µL protein phosphatase inhibitor cocktail]. The protein concentration was measured using a BCA protein quantitation kit (Epizyme, Shanghai, China). The following primary antibodies were used: B568 primary antibody (1:500, PA5-118186; Invitrogen); and GAPDH primary antibody (1:10000, PR30011; Proteintech, Shanghai, China).

Cell proliferation

The CCK-8 (MedChemExpress, Shanghai, China) was used to measure cell proliferation. Cells were digested and seeded in 96-well plates. After 12 h, 24 h, 48 h, and 72 h, 10 µL CCK-8 was added to each well and cultured in the incubator for 2 h. Then, a microplate reader was used to detect the optical density value at 450 nm.

Plate cloning assay

A total of 3000 cells were seeded in 6-well plate for plate cloning and cultured for 10-15 d. When the size of the cell colony was suitable, the cells were fixed in 4% neutral paraformaldehyde solution, stained with 1% crystal violet, and finally photographed for calculating.

Wound healing assay

After cell transfection for 8 h in 6-well plates, cells were scratched vertically with a 10 uL tip, washed with phosphate- buffered saline, and then covered in serum-free medium for culturing. Pictures were taken at 0 h and 48 h with a light microscope, and the documented wound-healing areas were analyzed using ImageJ software.

Cell migration and invasion assays

HCC cell migration and invasion were examined using the transwell chambers (jetbiofil, Guangzhou, China). Cells were digested 48 h after transfection, counted, and prepared in cell suspension with serum-free medium. 600 µL of 10% FBS medium was added to a 24-well plate, and then the chamber was put in the plate with 200 uL cells. After 48 h, the cells were fixed with 4% formaldehyde and stained with 1% crystal violet. Five random fields were taken under the microscope and cells passing through the chamber were calculated.

Statistical analyses

SPSS 26.0 statistical software (IBM Corp., Armonk, NY, United States) was used to analyze the data. R software (version 4.2.1) and GraphPad Prism 9 software were used for drawing graphics. The Wilcoxon test was used for immune checkpoint gene and HLA-related gene analyses. Drug sensitivity was analyzed using Pearson’s correlation coefficient. P < 0.05 was considered statistically significant.

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RESULTS

Risk assessment of B56& expression in pan-cancer

The expression of B568 in pan-cancer was analyzed using the TIMER database. For tumors such as BRCA, CHOL, ESCA, HNSC, LIHC, LUSC, LUAD, and STAD, B56 was expressed higher in the tumor tissues than in the normal tissues (Figure 1A). Data analysis combining TCGA and GTEx databases showed that B568 was not only highly expressed in the above eight tumor types but also in sixteen other tumor types, including ACC, CESC, DLBC, GBM, KICH, LAML, LGG, OV, PAAD, PCPG, PRAD, READ, SKCM, THCA, THYM, and UCEC (Figure 1B). The PrognoScan online website was used to analyze the relationship between B56 expression and prognosis of tumor patients in the Gene Expression Omnibus dataset. As shown in Figure 1C, the high expression of B568 was associated with a poor prognosis in patients with brain or breast cancer, while the low expression of B568 was associated with a poor prognosis in patients with colorectal, lung, skin, or ovarian cancer. Univariate analysis was used to analyze the prognostic significance of B568 expression in 33 tumors. The results of the forest plot showed that the high expression of B568 was a risk factor for ACC, LIHC, PAAD, and PCPG (all HR > 1, P < 0.05), but for KIRC and THYM, the high expression of B568 was a protective factor (all HR < 1, P < 0.05) (Figure 1D). These results indicate that B568 has a deregulated expression in tumors and its expression is correlated with the prognosis of tumor patients.

B56& expression is associated with mutation, sCNAs, and immunity in different tumors

To understand the role of B568 expression in tumors, we first analyzed the association of B568 expression with mutations and sCNAs in 33 tumors through the TIMER database. It was found that diploid, arm-level deletion and arm-level gain were common mutation types in most cancers, accounting for a high proportion. The top four tumors with the highest mutation ratio were UCEC, SKCM, COAD, and UCS (Figures 2A and B). Second, the relationship between B568 expression and immune status showed that B568 expression was negatively correlated with estimated score, immune score, and stromal score (Figure 2C). In TGCT, the correlations between B568 expression and immune score (r = - 0.537) and estimated score (r = - 0.494) were the strongest (all P < 0.001). In ACC, the associations between B568 expression and stromal score (r = - 0.338), immune score (r = - 0.455) and estimated score (r = - 0.423) were also relatively strong (all P < 0.001). Finally, more than 30 common immune checkpoint genes were collected to analyze the correlation with B568 expression. As shown in Figure 2D, there was a positive relationship between the expression levels of B568 and several immune checkpoint genes in some tumors such as KICH, KIRP, LIHC, PAAD, and UVM. These results suggest that B568 expression might play a role in tumors by regulating tumor mutation, sCNAs, or immunity.

Prognostic significance of B56& expression in HCC

To identify the expression of B568 in HCC tissues, we downloaded liver cancer-related data from TCGA database for bioinformatics analysis (tumor = 374, normal = 50). It was found that B568 was highly expressed in HCC tissues compared to normal liver tissues (P < 0.001; Figures 3A and B). TCGA database combined with GTEx database was used to further analyze the expression of B568 in HCC tissues (tumor = 371, normal = 160). As shown in Figure 3C, it was consistent with the results of TCGA database analysis. The prognostic significance of B568 in HCC patients was analyzed using the GEPIA database. The results showed that the expression of B568 was negatively correlated with the overall survival (OS) of patients with HCC, indicating that the higher the expression level of B568, the worse the prognosis of HCC patients (P < 0.05; Figure 3D).

Further analysis of the differential protein expression of B568 between normal liver tissues and HCC tissues was conducted in the HPA database. It was found that B568 stained weakly in normal liver tissue samples, but had deeper staining in HCC tissues (Figure 3E). Univariate analysis was used to analyze the association of common pathological features with OS in HCC patients. The results showed that the differential protein expression of B568 was statistically significant with patient OS (P < 0.05). Further inclusion of variables in the multivariate regression model analysis found that the correlation was not significant (Table 1). These results indicate that B568 is highly expressed in HCC tissues and correlates with a poor prognosis in HCC patients.

B56& expression is correlated with HCC immunity

The role of B568 expression in HCC immune subtypes was explored using the TISIDB website. Immune subtypes were divided into the following six types: C1 (wound healing), C2 (interferon gamma dominant), C3 (inflammatory), C4 (lymphocyte depleted), C5 (immunologically quiet), and C6 (transforming growth factor ß dominant). As can be seen from Figure 4A, B568 exhibited high expression in the C3 type and lowest expression in the C2 type. The immune-related function and immune infiltration of B568 in HCC were further analyzed by the ssGSEA algorithm. The box plot in Figure 4B shows that the expression levels of macrophages, T helper (Th) cells, central memory T cells (TCM), effector memory cells (TEM), natural killer (NK) cells, and Th2 cells were significantly higher in the B568 high expression group than in the B568 low expression group. The opposite was true for Th17 cells, gamma delta T cells (Tgd), plasmacytoid DCs (pDCs), DCs, and cytotoxic cells. Figure 4C shows that a variety of immune cells were associated with B568 expression including Th17 cells (r = - 0.264, P < 0.001), pDCs (r = - 0.203, P < 0.001), DCs (r = - 0.175, P < 0.001), cytotoxic cells (r = - 0.141, P < 0.01), Tgd (r = - 0.103, P < 0.05), Th cells (r = 0.388, P < 0.001), TCM (r = 0.264, P < 0.001), TEM (r = 0.263, P < 0.001), Th2 cells (r=0.258, P<0.001), macrophages (r=0.195, P <0.001), and NK cells (r=0.165, P <0.01).

The relationship between B56 expression and the tumor microenvironment (TME) was further analyzed by ESTIMATE, immune, and stromal score algorithms. As shown in Figure 4D, the ESTIMATE score and stromal score were significantly higher in the B568 high expression group than in the B568 low expression group (P < 0.05). The relationship between B568 expression and six common immune cells were analyzed using the TIMER database. Figure 4E shows that

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Table 1 Correlation between B56 expression and hepatocellular carcinoma prognosis
VariablesnUnivariate analysis Multivariate analysis
HR (95%CI)P valueHR (95%CI)P value
Gender3731.261 (0.885-1.796)0.2041.052 (0.632-1.752)0.845
Age3731.205 (0.850-1.708)0.2931.238 (0.768-1.997)0.381
T3702.598 (1.826-3.697)< 0.001ª8.023 (0.461-139.715)0.153
N2582.029 (0.497-8.281)0.3754.476 (0.583-34.348)0.149
M2724.077 (1.281-12.973)0.0501.865 (0.546-6.367)0.320
Stage3492.504 (1.727-3.631)< 0.001ª0.357 (0.020-6.488)0.487
Grade3681.091 (0.761-1.564)0.6371.177 (0.735-1.885)0.498
B56₴3731.587 (1.120-2.249)0.009ª1.337 (0.829-2.156)0.234

ªp < 0.05.

CI: Confidence interval; HR: Hazard ratio.

Table 2 Correlation analysis of B56& expression with chemotherapy drugs
DrugsCorrelationP value
Allopurinol0.2550.049
Fluorouracil-0.3240.011
Methylprednisolone0.2790.031
Chelerythrine0.3340.009
Ergenyl-0.3570.005
Ribavirin0.3070.017
Claritin-0.3120.015
RAF-265-0.2930.023
Nelarabine0.2640.042
PLX-4720-0.2710.036
Econazole nitrate0.3160.014
Rabusertib-0.2760.033
Vemurafenib-0.3040.018
Vertex ATR inhibitor Cpd 45-0.2650.041
Dabrafenib-0.2630.042

B568 expression was positively correlated with B cells, CD8+ T cells, CD4+ T cells, macrophages, neutrophils, and DCs (all P < 0.05). These results suggest that the expression of B568 may have functions in HCC immunity.

To further explore the potential mechanism of B568 in HCC immunity, we evaluated the expression and correlations of 34 common immune checkpoints and 21 HLA-associated genes in different B568 expression groups. Excluding the adenosine A2A receptor gene, butyrophilin-like protein 2, and indoleamine 2,3-dioxygenase 2, the remaining 31 immune checkpoints were significantly correlated with the differential expression of B568, and all were positively correlated (Figure 5A). As shown in Figure 5B, with the exception of the HLA-F and HLA-G genes, the remaining 19 HLA- associated genes were significantly correlated with the differential expression of 568, and all were positively correlated.

Data related to cancer drugs were downloaded through the CellMiner database for analysis. It was found that the expression of B568 was negatively correlated with the sensitivity of most chemotherapy drugs, and a small number showed a positive correlation (Table 2). As shown in Figure 5C, four drugs with the strongest negative correlation and their differential expression were statistically significant in drug sensitivity, including Ergenyl (r = - 0.357, P < 0.01), fluorouracil (r = - 0.324, P < 0.05), Claritin (r = - 0.312, P < 0.05), and vemurafenib (r = - 0.303, P < 0.05). The remaining three drugs were statistically significant in the differential expression of B568 except Claritin (all P < 0.05). Furthermore, three

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Figure 1 Analysis of B56E expression levels and prognostic value in 33 types of cancers. A: B56E expression profile in pan-cancer by the Tumor Immune Estimation Resource database; B: B56E expression in pan-carcinoma was analyzed based on The Cancer Genome Atlas (TCGA) and Genotype-Tissue Expression databases; C: Relationship between B56E expression and tumor survival in the Gene Expression Omnibus dataset; D: Prognostic value of B56€ in pan- cancer based on TCGA database. ªP < 0.05; bP < 0.01; P < 0.001. OS: Overall survival; CI: Confidence interval; HR: Hazard ratio; ACC: Adrenocortical carcinoma; BLCA: Bladder urothelial carcinoma; BRCA: Breast invasive carcinoma; CESC: Cervical squamous cell carcinoma; CHOL: Cholangiocarcinoma; COAD: Colon adenocarcinoma; DLBC: Lymphoid neoplasm diffuse large B cell lymphoma; ESCA: Esophageal carcinoma; GBM: Glioblastoma; HNSC: Head and neck squamous cell carcinoma; KICH: Kidney chromophobe; KIRP: Kidney renal papillary cell carcinoma; KIRC: Kidney clear cell carcinoma; LGG: Low-grade glioma; LIHC: The Cancer Genome Atlas-Liver hepatocellular carcinoma; LUAD: Lung adenocarcinoma; LUSC: Lung squamous cell carcinoma; MESO: Mesothelioma; OV: Ovarian serous cystadenoma; PAAD: Pancreatic adenocarcinoma; PCPG: Pheochromocytoma and paraganglioma; PRAD: Prostate adenocarcinoma; READ: Rectal adenocarcinoma; SARC: Sarcoma; SKCM: Skin cutaneous melanoma; STAD: Stomach adenocarcinoma; TGCT: Testicular germ cell tumor; THCA: Thyroid carcinoma; THYM: Thymic carcinoma; UCEC: Endometrial cancer; UCS: Uterine carcinosarcoma; UVM: Uveal melanoma.

A

a

2

2)

0

0

0

0

0

0

0

0

0

0

PPP2R5E expression level (log2 TPM)

9

+

2

ACC.Tumor (n=79)

BLCA.Tumor (n=408).

BLCA.Normal (n=19)- BRCA.Tumor (n=1093)-

BRCA.Normal (n=112)

BRCA-Basal. Tumor (n=190)

BRCA-Her2.Tumor (n=82)

BRCA-LumA.Tumor (n=564)

BRCA-LumB. Tumor (n=217)

CESC.Tumor (n=304)

CESC.Normal (n=3)

CHOL.Tumor (n=36)

CHOL.Normal (n=9)

COAD.Tumor (n=457)

COAD.Normal (n=41)

DLBC.Tumor (n=48)

ESCA.Tumor (n=184)

ESCA.Normal (n=11)

GBM.Tumor (n=153)

GBM.Normal (n=5)

HNSC.Tumor (n=520)

HNSC.Normal (n=44)

HNSC-HPV+.Tumor (n=97)

HNSC-HPV -. Tumor (n=421)

KICH.Tumor (n=66)

KICH.Normal (n=25)

KIRC.Tumor (n=533)

KIRC.Normal (n=72)

KIRP.Tumor (n=290)

KIRP.Normal (n=32)

LAML. Tumor (n=173)

LGG.Tumor (n=516))

LIHC. Tumor (n=371)

LIHC.Normal (n=50)

LUAD.Tumor (n=515)

LUAD.Normal (n=59)

LUSC.Tumor (n=501)

LUSC.Normal (n=51)

MESO.Tumor (n=87)

OV.tumor (n=303)

PAAD.Tumor (n=178)

PAAD.Normal (n=4)

PCPG.Tumor (n=179)

PCPG.Normal (n=3)

PRAD.Tumor (n=497)

PRAD.Normal (n=52)

READ.Tumor (n=166)

READ.Normal (n=10)

SARC.Tumor (n=259)

SKCM.Tumor (n=103)

SKCM.Metastasis (n=368)

STAD.Tumor (n=415))

STAD.Normal (n=35)

TGCT.Tumor (n=150)

THCA. Tumor (n=501)

THCA.Normal (n=59)

THYM.Tumor (n=120)

UCEC.Tumor (n=545)

UCEC.Normal (n=35)

UCS.Tumor (n=57)

UCM.Tumor (n=80)

B

10

30

PPP2R5E expression Log2(TPM+1)

Jo

9

Ja

10

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10

10

10

10

jo

10

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10

10

10

10

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2

0

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COAD

DLBC

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GBM

HNSC

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KIRC

KIRP

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LIHC

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PCPG

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SARC

SKCM

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TGCT

THCA

THYM

UCEC

UCS

UVM

C

Brain cancer

GSE4412-GPL97

Breast cancer GSE9893

D

HRs (95%CI)

100

ACC

OS (%)

p=0.018

100

BLCA

3.317 (1.570-7.009)

80

BRCA

1.297 (0.969-1.736)

60

1.137 (0.827-1.564)

40

High (n = 32)

-

80

60

High (n = 26)

CESC

0.762 (0.478-1.214)

Low (n = 42)

40

Low (n = 129)

CHOL

0.781 (0.309-1.977)

20

0

20

p<0.001

COAD

DLBC

0.694 (0.472-1.022)

0.769 (0.202-2.928)

0

500

1000 1500 2000 2500

0

0

50

100

ESCA

0.903 (0.558-1.464)

Time (month)

150

GBM

Time (day)

HNSC

0.886 (0.633-1.239)

KICH

1.117 (0.856-1.457)

3.577 (0.968-13.213)

Colorectal cancer GSE17537

Lung cancer GSE11117

KIRC

KIRP

0.533 (0.396-0.717)

1.001 (0.554-1.808)

100

LAML

100

1.241 (0.815-1.890)

0

80

- High (n = 40)

2

80

LGG

LIHC

0.838 (0.597-1.176)

60

60

- High (n = 24)

LUAD

1.600 (1.120-2.290)

1.119 (0.841-1.490)

5

40

- Low (n = 15)

5

40

-Low (n = 17)

LUSC

1.056 (0.807-1.383)

20

p=0.047

20

p=0.042

MESO

OV

0.967 (0.611-1.531)

0

0

0.960 (0.742-1.241)

0

20 40 60 80 100

0

400

800

1200

PAAD

Time (month)

Time (day)

PCPG

1.649 (1.097-2.478)

PRAD

7.219 (1.805-28.869)

2.708 (0.781-9.387)

Skin cancer GSE19234

Ovarian cancer DUKE-OC

READ

SARC

1.087 (0.501-2.356)

SKCM

1.433 (0.966-2.126)

1.097 (0.839-1.434)

100

100

STAD

TGCT

1.167 (0.842-1.618)

80

de

80

60

p<0.001

60

- High (n = 18)

THCA

2.147 (0.294-15.667)

- High (n = 98)

1.207 (0.453-3.218)

40

THYM

0.223 (0.060-0.831)

20

p=0.044

- Low (n = 20)

40

- Low (n = 35)

20

UCEC

0.988 (0.659-1.482)

0

0

UCS

1.114 (0.574-2161)

0

200 400 600 8001000

0

50

100

150

UVM

1.188 (0.521-2.710)

Time (day)

Time (month)

0

1

2

3

4

Hazard ratio

DOI: 10.4251/wjgo.v16.i2.475 Copyright @The Author(s) 2024.

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% of samples with PPP2R5E mutation>>

0.06

25/531

0.04

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8/406

0.02

1/57

1/79

5/411

1/92

3/282

3/291

4/400

5/517

4/439

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1/140

3/485

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2/408

2/498

1/365

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2/1026

1/519

1/525

0

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1.0

0.5

0

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Estimate score

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bb
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Figure 2 Association analysis of B56E expression with tumor mutations, somatic copy number alterations, and immunity in pan-cancer. A: Percentage of samples with B56E mutation; B: Somatic copy number alterations by the Tumor Immune Estimation Resource database; C: Associations between B56& expression and estimate score, immune score, and stromal score in different cancers; D: Correlation analysis between immune checkpoint genes and the expression of B56€. ªP < 0.05; bP < 0.01; P < 0.001. SCNA: Somatic copy number alteration; ACC: Adrenocortical carcinoma; BLCA: Bladder urothelial carcinoma; BRCA: Breast invasive carcinoma; CESC: Cervical squamous cell carcinoma; CHOL: Cholangiocarcinoma; COAD: Colon adenocarcinoma; DLBC: Lymphoid neoplasm diffuse large B cell lymphoma; ESCA: Esophageal carcinoma; GBM: Glioblastoma; HNSC: Head and neck squamous cell carcinoma; KICH: Kidney chromophobe; KIRP: Kidney renal papillary cell carcinoma; KIRC: Kidney clear cell carcinoma; LGG: Low-grade glioma; LIHC: The Cancer Genome Atlas-Liver hepatocellular carcinoma; LUAD: Lung adenocarcinoma; LUSC: Lung squamous cell carcinoma; MESO: Mesothelioma; OV: Ovarian serous cystadenoma; PAAD: Pancreatic adenocarcinoma; PCPG: Pheochromocytoma and paraganglioma; PRAD: Prostate adenocarcinoma; READ: Rectal adenocarcinoma; SARC: Sarcoma; SKCM: Skin cutaneous melanoma; STAD: Stomach adenocarcinoma; TGCT: Testicular germ cell tumor; THCA: Thyroid carcinoma; THYM: Thymic carcinoma; UCEC: Endometrial cancer; UCS: Uterine carcinosarcoma; UVM: Uveal melanoma.

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I

DOI: 10.4251/wjgo.v16.i2.475 Copyright @The Author(s) 2024.

m Relative expression of PPP2R5E

Relative expression of PPP2R5E Log2(TPM+1)

C

D

4.0

c

C

Relative expression of PPP2R5E Log2(FPKM+1)

Log-rank Test

5.0

c

:

100

P = 0.021

3.5

Log2(FPKM+1)

4.0

3.0

4.0

Overall survival (%)

75

2.5

3.0

3.0

50

2.0

1.5

2.0

2.0

25

High B56£

1.0

1.0

0

Low B56&

.

Normal

Tumor

Normal

Tumor

Normal

Tumor

0

30

60

90

120

B56& expression in HPA database

Time (month)

Normal liver tissue

HCC tissue

DOI: 10.4251/wjgo.v16.i2.475 Copyright @The Author(s) 2024.

Figure 3 Expression and prognosis value of B56E in hepatocellular carcinoma. A: Analysis of the B56E expression difference between hepatocellular carcinoma (HCC) tissues and normal liver tissues in The Cancer Genome Atlas (TCGA); B: Pairwise difference comparison of B56E expression level between HCC tissues and normal liver tissues in the TCGA database; C: TCGA combined with Genotype-Tissue Expression databases to analyze the expression of B56E in HCC tissues and normal liver tissues; D: Gene Expression Profiling Interactive Analysis was utilized for analyzing the relationship between B56E expression and HCC overall survival time; E: Expression of B56E protein in normal liver tissue and HCC tissue by the Human Protein Atlas database. "P < 0.001. HCC: Hepatocellular carcinoma.

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@Low B56E

a

High B56£

b

ESTIMATE score

Stromal score

A

C

Β56€

D

T helper cells

Tcm

O

LIHC

Tem

Expression (Log2CPM)

Th2 cells

6

Macrophages

NK cells

4000

ns

Th1 cells

IDC

·

TFH

P value

Enrichment score

4

!

aDC-

0.75

i

H

8

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0.50

2000

NK CD56bright cells

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0.25

2

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0

0

P = 6.7e-03

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n= 22 C2

n= 45 C3

n = 135 C4

n = 159 C6

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n=1

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0

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Immune score

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0

Enrichment score

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a

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Eosinophils

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Tem

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Tgd

Th1 cells

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Purity

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CD4+ T cells

cor -0.067

partial.cor = 0.297

. p = 2.16e-01

p = 1.90e-08

:

partial.cor = 0.261

p = 9.72e-07

partial.cor = 0.43

p = 6.53e-17

4

2

Expression (Log2TPM)

0

0.25

0.50

0.75

1.00

0.1

0.2

0.3

0.4

0.2

0.4

0.6

0

0.1

0.2

0.3

0.4

Infiltration level

Macrophage

Neutrophil

Dendritic cells

partial.cor = 0.442

P= 9.10e-18

partial.cor = 0.514

·partial.cor = 0.395

p = 1.09e-24

· = 3.98e-14

0

0.1

0.2

0.3

0.05

0.10

0.15

0.20

0.25

0.50

0.75

1.00

Infiltration level

DOI: 10.4251/wjgo.v16.i2.475 Copyright @The Author(s) 2024.

Figure 4 Correlation analysis of B56E expression with immune subtype and immune cells. A: Analysis of B56E expression in different immune subtypes of hepatocellular carcinoma (HCC); B: Differential expression of immune cells in the B56E high and low expression groups; C: Correlation analysis between immune cells and the expression of B56E; D: Analysis of differences in tumor microenvironment scores between groups with a high and low B56E expression; E: Association of six common types of immune cells and B56E expression in HCC by Tumor Immune Estimation Resource. ªP < 0.05; bP < 0.01; cP < 0.001. aDC: Activate dendritic cell; iDC: Immature dendritic cell; NK: Natural killer; pDC: Plasmacytoid dendritic cell; TFH: Follicular helper T cell; Th: T helper; Treg: Regulatory T cell.

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drugs with the strongest positive correlation with their differential expression were statistically significant in drug sensitivity (Figure 5D), including chelerythrine (r = 0.334, P < 0.01), econazole nitrate (r = 0.316, P < 0.05), and ribravirin (r = 0.307, P < 0.05). The remaining two drugs were not statistically significant in the differential expression of B568 except chelerythrine (all P > 0.05). These results indicate that B568 expression is positively correlated with most of immune checkpoints and negatively correlated with the sensitivity of most chemotherapy drugs.

GSEA associated with B56 expression

The characteristics of GO and KEGG in cells at different expression levels of B568 were further explored by GSEA. As shown in Figure 6A, GO analysis indicated that B568 was mainly involved in histone modification, cell response to environmental stimulation, response regulation to DNA damage stimuli, the ERBB signaling pathway, epidermal growth factor receptor signaling pathway, and other processes in the biological process category. In terms of the cellular component, it was mainly related to transcriptional regulatory complexes, nuclear spots, DNA damage sites, and transcription factor TFIID complexes. In terms of molecular function, it was mainly related to protein serine/threonine kinase activity, transcriptional co-regulatory activity, histone binding, p53 binding, and tau protein kinase activity. When the expression level of B568 was elevated, it was mainly related to the cancer pathway, p53 downstream pathway, and interleukin-mediated signaling (Figure 6B). However, when the expression level of B568 was decreased, it was mainly related to translation, the peroxisome proliferator-activated receptor signaling pathway, and adipocytokine signaling pathway (Figure 6C). These results show that the expression level of B568 is correlated with different signaling pathways in HCC.

Functions of B56E in HCC cells

To further explore the role of B568 in HCC progression, we knocked down B56 in HCC cells. We first detected the expression of B568 in hepatic cells L02 and different HCC cell lines by quantitative polymerase chain reaction (qPCR). As shown in Figure 7A, B568 was more highly expressed in SK-Hep-1 cells and Hep3B cells than in L02 cells. B568 expression was verified by qPCR and western blotting after its knockdown in SK-Hep-1 cells and Hep3B cells (Figure 7B). Moreover, the proliferative ability of HCC cells was significantly weakened after B568 knockdown and detected by the CCK-8 assay (Figure 7C). Plate cloning experiments also found that HCC cells proliferated more slowly after B568 knockdown (Figure 7D). The migration and invasive ability of HCC cells after B568 knockdown was further detected by wound healing and transwell assays. The wound-healing area was smaller in B568 knockdown HCC cells than in control cells, indicating that the cell migration capacity was significantly decreased after B568 knockdown (Figure 7E, P < 0.05). The same results were found in the transwell assay, which showed that HCC cell invasion and migration capacity were significantly reduced in B568 knockdown SK-Hep-1 cells and Hep3B cells (Figure 7F). These results suggest that B568 may promote HCC cell proliferation and metastasis.

DISCUSSION

Herein, we first verified the tumor-promoting effect of B568 by comprehensive bioinformatics analysis and related ex- periments. Then, the correlation between B568 expression levels and immune infiltration, immune checkpoint molecules, and immune function were explored. Furthermore, the potential value of B568 as an immunomodulator was revealed in the evaluation of immunotherapy for HCC. Our study’s findings collectively suggest that B568 is a potential biomarker for pan-cancer prognosis and an immune target for HCC treatment.

Some studies have reported that PP2A is often inactivated in human cancers and is considered a tumor suppressor[21, 22]. Paradoxically, inhibition of PP2A also has the potential to be a therapeutic target for a variety of cancers. Decreased PR55a expression inhibits the migration and invasion of pancreatic cancer cells[23]. Another study showed that B55B overexpression markedly suppressed cell migration and invasion in HCC cells[24]. B568 is one member of the PP2A B56 regulatory subunit; its role in tumors has garnered much attention and is a subject of debate. B568 has a nuclear output signal at the C-terminus, which can lead to migration of the PP2A complex into the cytoplasm[25]. Through the TCGA, GTEx, PrognoScan, and TIMER databases, we found that B568 expression levels were up-regulated in most tumors, and the high expression of B568 was associated with a poor prognosis in patients with brain tumor or breast cancer. Moreover, we identified B568 as a high-risk prognostic factor in ACC, LIHC, PAAD, and PCPG.

After determining the expression characteristics of B568 in pan-cancer, we further investigated the key role of B568 in HCC. The results showed that the expression level of B568 in HCC tissues was higher than that of normal tissues, and HCC patients with high B568 expression had a poor prognosis. Additional cell function assays showed that downreg- ulated B568 can inhibit HCC cell proliferation, invasion, and migration. Consistent with our results, a recent study showed that decreased B568 can promote gastric cancer cell apoptosis to suppress cell growth[18]. Finally, GSEA was used to analyze the biological function of B568 in HCC. GO results showed that B568 expression was related to immune response and histone modification. When B568 was highly expressed, it was mainly enriched in the cancer pathway, p53 downstream pathway, interleukin-mediated signaling and other related pathways, revealing the potential mechanism of B568 in the malignant biological behavior of HCC cells.

The high incidence and mortality rate of HCC is a serious health problem worldwide[26]. Due to the insidious onset of HCC in the early stage, patients present in the intermediate to advanced stages, and the OS remains poor due to high rates of intrahepatic and extrahepatic metastasis and recurrence[27,28]. With the development of immunotherapy, the treatment of HCC has been further improved. Based on the application of cytotoxic T-lymphocyte associated protein 4 and programmed cell death protein 1/programmed death-ligand 1 monoclonal antibodies, T cell immune checkpoint

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Figure 5 Correlation analysis of B56E expression with immune checkpoint genes, human leukocyte antigen-associated genes and drug sensitivity. A: Correlation analysis between immune checkpoint genes and the expression of B56E; B: Correlation analysis between human leukocyte antigen- associated genes and the expression of B56E; C and D: Association between anticancer drug response and B56 expression in the CellMiner database. ªP < 0.05; bp < 0.01; P < 0.001. HLA: Human leukocyte antigen; NS: Not significant.

A

PPP2R5E

Log2(TPM+1)

C

6

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R =- 0.36

P=0.005

0

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a

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0

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2

R =- 0.32

2

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P=0.011

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1

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0

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-1

-2.

B

PPP2R5E Log2(TPM+1)

1

2

3

4

High

Low

6

B56& expression

A

2

High

0

Low

4.

a

B2Mc

HLA-Aª

Activity z-scores of Vemurafenib

3

R =- 0.30

HLA-Bª

HLA-Cc

2

P=0.018

2-

HLA-DMAª

HLA-DMBc

1

0

0

HLA-DOAc

B

HLA-DOBb

HLA-DPA1c

Z-score

-1

2.5

-2.

HLA-DPB1c

HLA-DQA1c

1

2

3

4

High

Low

HLA-DQA2ª

0

B56& expression

HLA-DQB1b

HLA-DRAc

HLA-DRB1c

2.5

HLA-Ec

HLA-Fns

HLA-Gns

TAP1c

TAP2€

TAPBPc

D

Activity z-scores of Chelerythrine

a

R=0.33

Activity z-scores of Econazole Nitrate

R=0.32

NS

Activity z-scores of Ribavirin

4.

R=0.31

6

NS

P=0.009

2

P=0.014

4

2

P=0.017

2.5

2

4.

0

0

0

:

2

0

2-

:

-2

2.5

-2

-2.

0

0-

I

I

-2

1

2

3

4

High

Low

1

2

3

4

High

Low

1

2

3

4

High

Low

B56& expression

B56& expression

B56& expression

DOI: 10.4251/wjgo.v16.i2.475 Copyright @The Author(s) 2024.

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WJGO

Figure 6 Gene Ontology and Kyoto Encyclopedia of Genes and Genomes analyses for samples with high and low B56E expression. A: Gene Ontology enrichment analysis was performed on B56€ differential expression samples; B: Kyoto Encyclopedia of Genes and Genomes (KEGG) signaling pathway in the high B56E expression group; C: KEGG signaling pathway in the low B56E expression group. BP: Biological process; CC: Cellular component; MF: Molecular function.

A

histone modification

cellular response to environmental stimulus

response to transforming growth factor beta

regulation of response to DNA damage stimulus

BP

ERBB signaling pathway

epidermal growth factor receptor signaling pathway

P adj

transcription regulator complex

0.075

nuclear speck

0.050

focal adhesion

0.025

nuclear membrane

CC

Counts

site of DNA damage

0 10

transcription factor TFIID complex

20

30

40

protein serine/threonine kinase activity

transcription coregulator activity

DNA-binding transcription factor binding

histone binding

MF

p53 binding

tau-protein kinase activity

0.02 0.04 0.06 0.08

Gene ratio

B

C

@ 0.5

Enrichment score

8 0.4-

Ranked list metric Enrichment score

0

0.3

0.2-

-0.2-

0.1-

Reactome] Leishmania infection

[Reactome] Translation

Reactome] Metabolism of amino acids and derivative’s

0

Beactome] Signaling by interleukins

-0.4.

PID] F53 downstream pathway

[WikiPathways] Ppar signaling pathway

KEGG] Calcium

[KEGG] Adipocytokine signaling pathway

0.1.

ŻEGĞİ Jak stat signaling pathway

KEGGI Pathway in cancer

-0.6-

-

[PID] Hnf3b pathway

KEGG] Parkinsons disease

Ranked list metric

4

4

2

0

-2

-2

0

10000

20000

30000

0

10000

20000

30000

Rank in ordered dataset

Rank in ordered dataset

DOI: 10.4251/wjgo.v16.i2.475 Copyright @The Author(s) 2024.

inhibitors (ICIs) bring a new clinical breakthrough in tumor immunotherapy[29-33]. Especially, the immunotherapy of several solid tumor and hematological tumors has achieved satisfactory efficacy, and sheds light on immune-based HCC therapy[34-38]. ICIs have been shown to eliminate tumor cells using an efficient immune response, including non-small cell lung cancer[39], melanoma[40], and HCC.

Our study found that B568 expression varied in different HCC immune subtypes. Moreover, the expression of B568 was correlated with a variety of immune cells, especially tumor-infiltrating lymphocytes (TILs) and played a vital role in TME. For example, B568 expression was significantly correlated with CD8+ T cells, macrophages, and DCs. The prognosis and immunotherapy efficacy can be predicted by TILs in the TME of cancer patients[41,42]. As a universal component of TME, macrophages have been shown to aid in immune evasion and suppression[43]. Some studies have suggested the presence of antitumor immunity in HCC patients. For example, tumor-associated antigen-specific CD8+ T-cell responses were found correlated with HCC prognosis[44]. The intratumoral density of activated cytotoxic T cells (CTLs) had a correlation with the OS of HCC patients, and the intratumoral balance between CTLs and regulatory T cells also affected the OS and disease-free survival[45]. These observations suggest that the immunogenic potential of HCC can be controlled through optimized immunotherapy.

This study had some limitations. The sample was limited to the patient information contained in TCGA dataset requires more clinical case validation. The potential mechanism of B568 tumor-promoting and immunomodulatory effects in HCC also needs to be further verified in clinical practice.

CONCLUSION

In summary, we demonstrate that B568 can be used as a prognostic biomarker for a variety of tumors and may modulate tumor immune cell infiltration and immune response. B56 can promote the proliferation, invasion, and migration of HCC. Our results can provide theoretical support and new ideas for HCC treatment.

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WJGO https://www.wjgnet.com

DOI: 10.4251/wjgo.v16.i2.475 Copyright @The Author(s) 2024. Figure 7 Effects of B56E downregulation on the proliferation, invasion, and migration of hepatocellular carcinoma cells. A: Quantitative polymerase chain reaction (qPCR) detected the expression of B56E in human hepatocellular carcinoma (HCC) cell lines; B: qPCR and western blotting verified B56€ knockdown efficiency; C: Downregulation of B56E expression on the proliferation of HCC cells was determined by the Cell Counting Kit-8 assay; D: Downregulation of B56E expression on the proliferation of HCC cells was determined by the colony formation assay; E and F: Wound healing assay (E) and transwell assay (F) showed the ability of cells to invade and migrate after downregulation of B56€. ªP < 0.05; bP < 0.01; "P < 0.001. NC: Negative vector control; KD-1: B56E knockdown vector 1;

Relative expression of B56£

B

SK-Hep-1

15

Hep3B

c

Relative expression of B56&

1.5

Β56€

GAPDH

Relative expression of B56&

1.5

B56€

10

GAPDH

1.0

0.62 0.43

1.0

T

0.75 0.69

5

c

C

C

0.5

c

c

c

C

ns

0.5

0

L02-

HepG2-

Hep3B-

SMMC7721

Huh7

SK-Hep-1-

MHCC97H-

0

0

NC

KD-1

KD-2

NC

KD-1

KD-2

C

SK-Hep-1

Hep3B

OD at 450 nm (CCK8)

0.5

NC

OD at 450 nm (CCK8)

0.4

NC

0.4

KD-1

KD-2

0.3

KD-1

KD-2

a

0.3

a

0.2

0.2

0.1

0.1

0

0

0

12

24

36

48

60

72

0

12

24

36

48

60

72

Time (h)

Time (h)

D

E

0 h

48 h

SK-Hep-1

Hep3B

160

a

a

NC

Wound area at 48h (%)

120

200

300

SK-Hep-1

80

KD-1

40

Colony number

150

Colony number

0

200

KD-2

NC

KD-1

KD-2

100

40

a

a

100

NC

Wound area at 48h (%)

120

a

a

50

C

Hep3B

80

KD-1

40

0

0

0

NC

KD-1

KD-2

NC

KD-1

KD-2

KD-2

NC

KD-1

KD-2

F

SK-Hep-1

NC

KD-1

KD-2

500

400

200

Migration

Migration cells

Invasion cells

300

150

200

b

a

100

b

b

Invasion

100

50

0

0

NC

KD-1

KD-2

NC

KD-1

KD-2

Hep3B

NC

KD-1

KD-2

30

20

10

Migration

Migration cells

Invasion cells

15

20

b

10

10

5

a

Invasion

HO

0

0

NC

KD-1

KD-2

NC

KD-1

KD-2

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KD-2: B56E knockdown vector 2; CCK-8: Cell Counting Kit-8.

ARTICLE HIGHLIGHTS

Research background

B568 is a regulatory subunit of the protein phosphatase 2A, which is abnormally expressed in tumors and regulates various tumor cell functions.

Research motivation

At present, the application of B568 in pan-carcinoma lacks a comprehensive analysis, and its role and mechanism in hepatocellular carcinoma (HCC) are still unclear.

Research objectives

The study aims to analyze B568 in pan-cancer, and explore its role and mechanism in HCC.

Research methods

The Cancer Genome Atlas (TCGA), Genotype-Tissue Expression, Gene Expression Profiling Interactive Analysis, and Tumor Immune Estimation Resource databases were used to analyze B568 expression, prognostic mutations, somatic copy number alterations, and tumor immune characteristics in 33 tumors. The relationship between B568 expression levels and drug sensitivity, immunotherapy, immune checkpoints, and human leukocyte antigen (HLA)-related genes were further analyzed. Gene Set Enrichment Analysis (GSEA) was performed to reveal the role of B568 in HCC. Cell Counting Kit-8, plate cloning, wound healing, and transwell experiments were conducted to show the effects of B568 interference on the malignant behaviors of HCC cells.

Research results

In most tumors, B568 expression was upregulated, and B568 high expression was a risk factor in adrenocortical cancer, HCC, pancreatic adenocarcinoma, and pheochromocytoma and paraganglioma (all P < 0.05). B568 expression levels were correlated with a variety of immune cells, such as T helper 17 cells, B cells, and macrophages. There was a positive correlation between B568 expression levels with immune checkpoint genes and HLA-related genes (all P < 0.05). The expression of B568 was negatively correlated with the sensitivity of most chemotherapy drugs, but a small number showed a positive correlation (all P < 0.05). GSEA showed that B568 expression was related to the cancer pathway, p53 downstream pathway, and interleukin-mediated signaling in HCC. Knockdown of B568 expression in HCC cells inhibited the proliferation, migration, and invasion capacity of tumor cells.

Research conclusions

B568 may regulate the microenvironment, immune evasion, and immune cell infiltration of multiple tumors. Moreover, B568 plays an important role in HCC progression. Our study supports B568 as a prognostic marker and potential therapeutic target for HCC.

Research perspectives

The patient information contained in TCGA dataset and requires more clinical case validation. In the future, the potential mechanism of B568 tumor-promoting and immunomodulatory effects in HCC also needs to be further verified in clinical practice.

FOOTNOTES

Co-first authors: Hong-Mei Wu and Yuan-Yuan Huang.

Author contributions: Wu HM, Huang YY, and Zhu XN conceived, designed, and wrote the original draft; Wu HM, Xu YQ, Xiang WL, Yang C, Liu RY, Li D, and Guo XF performed the formal analysis; Zhang ZB, Bei CH, and Tan SK conducted the methodology; Bei CH, Tan SK, and Zhu XN were responsible for the conceptualization, writing, review and editing; and all authors read and approved the final manuscript.

Supported by National Natural Science Foundation of China, No. 82060621, 82060607, and 82260664; Natural Science Foundation of Guangxi Province, No. 2020GXNSFDA297010 and 2020GXNSFAA297142; and Key Science and Technology Research and Development Program Project of Guangxi, No. AB22035017.

Institutional review board statement: The study was reviewed and approved by the Ethics Committee of Guilin Medical University (No. GLMC2020045).

Conflict-of-interest statement: All the authors report no relevant conflicts of interest for this article.

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Data sharing statement: Publicly available datasets were analyzed in this study. No additional data are available.

Open-Access: This article is an open-access article that was selected by an in-house editor and fully peer-reviewed by external reviewers. It is distributed in accordance with the Creative Commons Attribution NonCommercial (CC BY-NC 4.0) license, which permits others to distribute, remix, adapt, build upon this work non-commercially, and license their derivative works on different terms, provided the original work is properly cited and the use is non-commercial. See: https://creativecommons.org/Licenses/by-nc/4.0/

Country/Territory of origin: China

ORCID number: Sheng-Kui Tan 0000-0002-1955-0868; Xiao-Nian Zhu 0000-0002-6175-3204.

S-Editor: Wang JJ

L-Editor: A

P-Editor: Zhang XD

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