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EDITED BY Feng Jiang, Fudan University, China REVIEWED BY Xiaojing Chang, Second Hospital of Hebei Medical University, China Sammed Mandape, University of North Texas Health Science Center, United States

*CORRESPONDENCE Yali Zhang, zhangyl_2013@sina.com

“These authors have contributed equally to this work

SPECIALTY SECTION This article was submitted to Cancer Genetics and Oncogenomics, a section of the journal Frontiers in Genetics

RECEIVED 17 July 2022 ACCEPTED 29 November 2022 PUBLISHED 04 January 2023

CITATION Lu K, Yuan X, Zhao L, Wang B and Zhang Y (2023), Comprehensive pan- cancer analysis and the regulatory mechanism of AURKA, a gene associated with prognosis of ferroptosis of adrenal cortical carcinoma in the tumor micro-environment. Front. Genet. 13:996180. doi: 10.3389/fgene.2022.996180

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@ 2023 Lu, Yuan, Zhao, Wang and Zhang. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

Comprehensive pan-cancer analysis and the regulatory mechanism of AURKA, a gene associated with prognosis of ferroptosis of adrenal cortical carcinoma in the tumor micro-environment

Keqiang Lut, Xingxing Yuan1, Lingling Zhao, Bingyu Wang and Yali Zhang*

Department of Gastroenterology, Heilongjiang Academy of Traditional Chinese Medicine, Harbin, China

Background: The only curative option for patients with locally or locally advanced adrenocortical carcinoma is primary tumor curative sexual resection (ACC). However, overall survival remains low, with most deaths occurring within the first 2 years following surgery. The 5-year survival rate after surgery is less than 30%. As a result, more accurate prognosis-related predictive biomarkers must be investigated urgently to detect patients’ disease status after surgery.

Methods: Data from FerrDb were obtained to identify ferroptosis-related genes, and ACC gene expression profiles were collected from the GEO database to find differentially expressed ACC ferroptosis-related genes using differential expression analysis. The DEFGs were subjected to Gene Ontology gene enrichment analysis and KEGG signaling pathway enrichment analysis. PPI network building and predictive analysis were used to filter core genes. The expression of critical genes in ACC pathological stage and pan-cancer was then investigated. In recent years, immune-related factors, DNA repair genes, and methyltransferase genes have been employed in diagnosing and prognosis of different malignancies. Cancer cells are mutated due to DNA repair genes, and highly expressed DNA repair genes promote cancer. Dysregulation of methyltransferase genes and Immune-related factors, which are shown to be significantly expressed in numerous malignancies, also plays a crucial role in cancer. As a result, we investigated the relationship of AURKA with immunological checkpoints, DNA repair genes, and methyltransferases in pan-cancer.

Result: The DEGs found in the GEO database were crossed with ferroptosis- related genes, yielding 42 differentially expressed ferroptosis-related genes. Six of these 42 genes, particularly AURKA, are linked to the prognosis of ACC. AURKA expression was significantly correlated with poor prognosis in patients

with multiple cancers, and there was a significant positive correlation with Th2 cells. Furthermore, AURKA expression was positively associated with tumor immune infiltration in Lung adenocarcinoma (LUAD), Liver hepatocellular carcinoma (LIHC), Sarcoma (SARC), Esophageal carcinoma (ESCA), and Stomach adenocarcinoma (STAD), but negatively correlated with the immune score, matrix score, and calculated score in these tumors. Further investigation into the relationship between AURKA expression and immune examination gene expression revealed that AURKA could control the tumor- resistant pattern in most tumors by regulating the expression level of specific immune examination genes.

Conclusion: AURKA may be an independent prognostic marker for predicting ACC patient prognosis. AURKA may play an essential role in the tumor microenvironment and tumor immunity, according to a pan-cancer analysis, and it has the potential to be a predictive biomarker for multiple cancers.

KEYWORDS

AURKA, pan-cancer analysis, tumor micro-environment, regulatory mechanism, ferroptosis

Introduction

Adrenal cortical carcinoma (ACC) is a rare malignant tumor with an annual incidence of one to two per million that can occur at any age and is more common in women (Cheng et al., 2021; Faron et al., 2022; Pitsava et al., 2022). It is an incidental adrenal tumor and one of the most common reasons for adrenalectomy, accounting for 14% of all spontaneous adrenal tumors (Alyateem and Nilubol, 2021). Although radical resection is the only option for the majority of ACC patients, postoperative survival remains low. As a result, understanding the molecular mechanism of ACC and identifying key target molecules can help predict tumor prognosis.

Currently, ACC is diagnosed using hormone detection and imaging, which plays a vital role in the initial diagnosis and prognostic detection and necessitates repeated detection (Mete et al., 2022). Efforts have been made for decades to discover new reliable, usable diagnostic and prognostic factors. Despite these achievements, 5-year mortality remains higher than 50% (Mizdrak et al., 2021). Accordingly, it is critical to discover new biomarkers that can predict patient outcomes and provide new treatment options.

Ferroptosis, a distinct mechanism of cell death caused by iron-dependent phospholipid peroxidation, has been shown to damage treatment-resistant cancer cells, particularly those in mesenchymal condition and prone to metastasis (Jiang et al., 2021). Correlative research has demonstrated that ferroptosis- related genes are linked to prognosis in various malignancies, including uveal melanoma, glioma, and adrenocortical tumors (Chen et al., 2021a; Luo and Ma, 2021; Zheng et al., 2021).

Aurora kinase A (AURKA) is a serine/threonine kinase family member, and its activation has been linked to several malignancies. Several studies have shown that highly expressed

AURKA can be used as a prognostic marker in various malignancies, including ACC (Du et al., 2021; Tang et al., 2021; Zhang et al., 2022).

Tumor samples from GEO databases were combined with standard models in this study. Differential expression analysis and ACC predictive analysis revealed significantly correlated genes. Pan-cancer analysis was used to study the expression of target genes in 40 different types of cancer. Then the correlations between target gene expression and tumor immune microenvironment, immune checkpoints, DNA repair genes, and methyltransferase were discovered.

Materials and methods

Data source

The GEO database (https://www.ncbi.nlm.nih.gov/geo) was used to download the RNA expression data for ACC from accession numbers GSE12368, GSE19750, and GSE75415, which contained 17 regular and 74 tumor tissues. All data were quantile normalized using a log2-scale transformation. The gene symbols found in multiple probes were calculated using their mean expression levels.

The “Limma” package of R software was used to investigate the differential expression genes (DEGs) of ACC (version: 3.42.2). p-values were adjusted to account for false-positive results. The number of highly expressed molecules in groups 1 (tumor) and 2 (standard control) that met the |log2(FC)|

>1&p. Adj0.05 threshold was counted. The DEGs were also visualized using the “ComplexHeatmap” and “ggplot2” packages. The DEGs and ferroptosis-related genes were then intersected to obtain ferroptosis-related genes with differential expression (DEFGs).

Functional analysis

Metascape Online (https://metascape.org/gp/index.html#/ main/step1) was used for available analysis. Metascape was used to perform functional analysis and build a PPI network using the ferroptosis-related genes. MCODE was used to reveal more densely connected regions.

Construction and prognostic value of IRSS

Univariate (Wei et al., 2022) Cox regression model is a semi- parametric regression model. The model’s dependent variables are survival results and survival time. It may examine the impact of several variables on survival time simultaneously. It does not require estimated data and can evaluate data with suppressed survival time. The least absolute shrinkage and selection operator (LASSO) is an L1-regularized linear regression approach. Using L1-regularization, part of the learned feature weights will be set to zero, achieving the goal of sparsity and feature selection (Tian et al., 2022). Univariate Cox regression analysis of DEFGs was used to identify significant prognosis-related genes, followed by LASSO regression analysis to obtain independent genes. A multivariate Cox regression analysis was also performed to obtain regression coefficients for independent prognostic factors. Finally, an immune risk score signature (IRSS) based on the Cox regression coefficient beta value was developed.

Survival analysis

One-way Cox was used to analyze the association of ACC expression with patient survival, and Xian Tao Academic created a forest plot of the correlation of overall survival and disease- specific survival of ARUKA in pan-cancer (https://www.xiantao. love).

Immune correlation analysis

The TIMER database was used to download data from multiple immune-infiltrating cells in 40 cancers, and the correlation between target gene expression and immune cell scores was examined separately. A lollipop graph of the correlation of target genes with immune cells in the cancer microenvironment and a diagram of the correlation of target

genes with immune scores, stromal scores, and computational scores in five cancers were drawn using Xian Tao Academic (https://www.xiantao.love).

Correlation analysis of DNA repair genes and methyltransferases

Using the TCGA expression profiling data, the correlation of DNA repair genes with target gene expression was assessed. The relationship between methyltransferases and the target gene was also investigated. Xian Tao Academic (https://www.xiantao.love) was used to create heat maps, with red dots indicating significant correlations.

Results

Results of DEGs screening in ACC

The information on the GEO database used is listed in Table 1. A total of 2,311 differentially expressed genes were identified following differential gene analysis: in GSE12368, the total number of molecules after filtering was 21,655, of which 849 IDs met the |log2(FC)|>1&p. Adj0.05 threshold. There were 170 highly expressed (logFC is positive) individuals in the standard group and 679 highly expressed (logFC is negative) individuals in the tumor group. The number of molecules in GSE19750 after filtering is 21,655, and 849 IDs meet the |log2(FC)| >1&p. Adj0.05 threshold.

Under this threshold, the regular group has a high expression (logFC is positive). The number was 170, with 679 having a high face (logFC is negative) in the tumor group. The number of molecules in GSE19750 after filtering is 21,655, and 849 IDs meet the |log2(FC)| >1&p. Adj0.05 threshold. Under this threshold, the usual group has a high expression (logFC is positive). The number was 170, with 679 highly expressed (logFC is negative) in the tumor group. After filtering in GSE75415, 12,548 molecules were obtained, of which 660 dysregulated genes satisfy |log2(FC)|>1&p. Adj0.05; under this threshold, the number of highly expressed (logFC is positive) genes in the standard group is equal to the number of highly expressed (logFC is positive) genes in the standard group. There were 258 in the tumor group, with 402 being highly expressed (logFC is negative) (Figure 1A). DEFGs was created by intersecting DEGs from GEO databases and ferroptosis- related genes (Figure 1B).

We used Metascape Online to perform a functional analysis to investigate ACC’s underlying mechanisms of ferroptosis signatures. The Gene Ontology (GO) analysis results show that these DEFGs were primarily enriched in response to

TABLE 1 The information of datasets from the GEO database.
Accession numberPlatformSamplesExperiment type
GSE75415GPL9625expression profiling by array
GSE12368GPL57018expression profiling by array
GSE19750GPL57048expression profiling by array

A

KONJ5

NCAPG TOPZA BACS

PMAIP1

Apelan

GRINZC

NPYIR

GRIA2

CCNB1 RRM2

SOLE

TCEAL2

MND1

IL32

OBH

FOXMI

CHGA

Group

PEG3-AS1

Group

BUB1

HOMER1

SST

GTF21

Group

SCNN1A

group1

CBXS

ZNF460

group1

COL11A1

ERICH3

SMCy HIST1H4E

TACC3

group2

UBEZS ASPM

group1

ASUS

CASA

SUR

PARK

group2

KIFZDA

group2

CYP1182

2

HEMES

BODILI

2

NEKS

BPPHI

PRI

2

BOTH

KLAA 1004

1

LUZPI

TMEFF2

1

TOP2A

TPX2

PTMS

SLC2AB

1

MUMIL1

RPS4Y1

0

0

ANGPT2

CENPK

CYP11B1

HSD382

CYP1182

0

LE

KIFZDA DEPDC1

-1

FAM1668

KONQ1

AADAC

-1

LMOD1

HOPX

-1

E

CDC25C

S100A8

11

1PX2

-2

FARBA

Ppare ADH18

-2

NOW

WNT4

DEPDC1B

HSD382

-2

AADIRE

FOXM!

ANIN

FMOD

ASPM

eyes

HIRZB

PIGDS

CSOCZ

HACE

L

CCNB2

OTL

CYP1182

COKI

OPR98

PDGFRA

SCG2

OFBP5

ZG16B

CALB!

CARTPT

LSP1

PNLIPRP3

PENK

CHGA

FATE1

TH

RARRE52

C3

MIR210HG

CHGA

PNMT

NR442

FOSB

GSE12368

GSE19750

GSE75415

B

C

GO:0034599: cellular response to oxidative stress

GO:0051348: negative regulation of transferase activity

GSE19750

GSE75415

R-HSA-8953897: Cellular responses to stimuli

GO:0034097: response to cytokine

GO:0007346: regulation of mitotic cell cycle

WP236: Adipogenesis

GSE12368

588

378

FerrDB

GO:0050727: regulation of inflammatory response

GO:1901652: response to peptide

hsa05167: Kaposi sarcoma-associated herpesvirus infection

118

49

17

R-HSA-9648895: Response of EIF2AK1 (HRI) to heme deficiency WP4313: Ferroptosis

GO:0002520: immune system development

77

2

GO:0080135: regulation of cellular response to stress

WP5044: Kynurenine pathway and links to cell senescence

462

217

R-HSA-9663891: Selective autophagy

3

WP2882: Nuclear receptors meta-pathway

GO:0048871: multicellular organismal homeostasis

132

10

M14: PID AURORA B PATHWAY

GO:0018958: phenol-containing compound metabolic process

2

2

WP231: TNF-alpha signaling pathway

GO:0031331: positive regulation of cellular catabolic process

R-HSA-556833: Metabolism of lipids

6

GO:0001889: liver development

GO:0006575: cellular modified amino acid metabolic process

WP5115: Network map of SARS-COV-2 signaling pathway

GO:0022411: cellular component disassembly

0

2

4

6

8

10

-log10(P)

D

E

TMNI

cellular response to oxidative stress

negative regulation of transferase activity

Cellular responses 10

response to cytokine

regulation of mitotic cell cycle

DOKONZA

Adipogenesis

regulation of inflammatory response

response to peptide

Kaposi sarcoma-associated herpesvirus infection

Response of EIF2AK1 (HRI) to heme deficiency Ferroptosis

KURKA

COKNIA

immune system development

regulation of cellular response to stress

Kynurenine pathway and links to cell senescence

Selective autophagy

Nuclear receptors meta-pathway

multicellular organismal homeostasis

PID AURORA B PATHWAY

phenol-containing compound metabolic process

ANPICS

TNF-alpha signaling pathway

NFAP3

FIGURE 1 Access to key genes. (A) Heatmap of differentially expressed genes in three GEO databases. (B) Venn diagram of differential ferroptosis genes. (C,D) Graph showing the GO and KEGG analysis based on the Metascape Online, bar plot, and network showing the distribution and relationship of the different functions. (E) PPI network and MCODE reveal hub genes in differential ferroptosis gene sets.

A

1.0

HELLS

1.0

FANCD2

1.0

SLC40A1

Low

High

Low

Low

0.8

High

High

Survival probability

0.8

Survival probability

0.8

Survival probability

0.6

0.6

0.6

0.4

0.4

0.4

0.2

Overall Survival HR = 8.83 (3.29-23.68)

0.2

0.2

Overall Survival HR = 5.10 (2.14-12.17)

Overall Survival HR = 0.23 (0.10-0.54)

0.0

P < 0.001

0.0

P < 0.001

0.0

P = 0.001

0

50

100

150

0

50

100

150

0

50

100

150

Time (months)

Time (months)

Time (months)

1.0

TNFAIP3

1.0

STMN1

Low

1.0

AURKA

Low

High

High

Low

High

Survival probability

0.8

Survival probability

0.8

Survival probability

0.8

0.6

0.6

0.6

0.4

0.4

0.4

0.2

Overall Survival HR = 5.74 (2.24-14.72)

0.2

Overall Survival HR = 10.62 (3.63-31.08)

0.2

Overall Survival HR = 6.55 (2.62-16.37)

0.0

P < 0.001

0.0

P < 0.001

0.0

P < 0.001

0

50

100

150

0

50

100

150

0

50

100

150

B

Time (months)

Time (months)

Time (months)

0

10

20

30

D

E

4

Risk score

3

66666666666666666666666666655533333322211110

6

6

6

3

0

Risk group

2

Low

0.6-

High

1

Coefficients

0.4-

Survival time

4000

0.2.

3000

Status

2000

1

0

0.0-

1000

-0.2-

TNFAIP3

-5

-4

-3

-2

-1

-5

-4

-3

-2

-1

Log (

Log (1)

AURKA

HELLS

-2

-1

0

1

2

CharacteristicsTotal(N)HR(95% CI) Univariate analysisP value Univariate analysis
HELLS(High vs. Low)798.829 (3.292-23.675) 1<0.001
FANCD2(High vs. Low)795.098 (2.136-12.166) 1<0.001
SLC40A1(High vs. Low)790.229 (0.097-0.542)<0.001
TNFAIP3(High vs. Low)795.742 (2.241-14.717) 1<0.001
STMN1(High vs. Low)7910.622 (3.631-31.076) I 1<0.001
AURKA(High vs. Low)796.545 (2.617-16.369) 1<0.001

C

Partial Likelihood Deviance

9.5

9.0

8.5-

8.0-

FIGURE 2 Establishment of ACC ferroptosis-related prognostic model. (A) six Significantly Differential Gene Survival Analysis Survival Chart. (B) Forest plot showing the results of a univariate Cox regression analysis. (C) Ten-fold cross-validation plot. (D) LASSO coefficient trajectory diagram. (E) The risk score, survival status and heat map of three key genes in patients.

A

B

1.0

C

1.0

1.0

Low

High

0.8

0.8

Survival probability

0.8

Sensitivity (TPR)

Sensitivity (TPR)

0.6

0.6

0.6

0.4

0.4

0.4

0.2

0.2

0.2

AURKA

HR = 11.63 (3.96-34.12)

3-Year (AUC = 0.918)

5-Year (AUC = 0.902)

2-Year (AUC = 0.906)

3-Year (AUC = 0.920)

0.0

P < 0.001

0.0

7-Year (AUC = 0.853)

0.0

5-Year (AUC = 0.818)

0

1000

2000

3000

4000

0.0

0.2

0.4

0.6

0.8

1.0

0.0

0.2

0.4

0.6

0.8

1.0

Time

1-Specificity (FPR)

1-Specificity (FPR)

D

E

6

10

ns

ns

The expression of AURKA Log2 (TPM+1)

5

The expression of AURKA Log2 (TPM+1)

8

ns

4

ns

6

3

2

4

1

2

0

T

Stage I

Stage II Stage III Stage IV Pathologic stage

Normal

Tumor

F

8

ns

The expression of AURKA Log2 (FPKM+1)

. .

6

Normal

Tumor

4

:

-

·

.

2

. …

-…

..

.

:

-

0

DE

ACC

BLCA

BRCA

CESC

CHOL

COAD

DLBC

ESCA

GBM

HNSC

KICH

KIRC

KIRP

LAML

LGG

LIHC

LUAD

LUSC

MESO

OV

PAAD

PCPG

PRAD

READ

SARC

SKCM

STAD

TGCT

THCA

THYM

UCEC

UCS

UVM

FIGURE 3 Validation of the model. (A) Survival map of high and low risk patients. (B) 3-Gene time-dependent ROC plot. (C) Single-gene time-dependent ROC plot. (D) Expression of AURKA in normal population and ACC patients. (E) Expression of AURKA in ACC patients at different stages. (F) AURKA expression in a wide range of cancers.

stimuli, oxidative stress responses, immune system processes, and negative regulators of transferase activity, as shown in Figures 1C, D. According to the Kyoto Encyclopedia of Genes and Genomes (KEGG) pathway analysis, these d DEFGs were primarily enriched in ferroptosis, cellular responses to stimuli, selective autophagy, and EIE2AKI response to heme deficiency. As a result of these findings, we decided to investigate the

relationship between the ferroptosis-gene set and the tumor immune microenvironment. Furthermore, the MCODE plugin and the MetascapeOnline-based protein-protein interaction (PPI) network identified necessary modules in these filiform genes (Figure 1D). STMN1, CDKN2A, CDKN1A, MAP3K5, TNFAIP3, and AURKA are involved in seven edges and six nodes.

A

B

TumorHR (95%CI)p valueTumorHR (95%CI)p value
ACC5.48 (2.28-13.17)<0.001ACC5.21 (2.14-12.69)<0.001
BLCA1.31 (0.98-1.76)0.069BLCA1.51 (1.06-2.16)0.024
BRCA1.24 (0.90-1.71)0.184BRCA1.41 (0.92-2.18)0.114
CESC1.17 (0.74-1.86)0.505CESC1.25 (0.74-2.13)0.409
CHOL1.73 (0.65-4.65)0.275CHOL1.92 (0.66-5.56)0.231
COAD0.78 (0.53-1.15)0.205
COAD0.73 (0.44-1.20)0.21
COADREAD0.77 (0.54-1.08)0.133
DLBC0.80 (0.18-3.59)0.766COADREAD0.82 (0.52-1.28)0.373
ESAD1.85 (0.96-3.53)0.064DLBC1.14 (0.16-8.23)0.9
ESCA0.96 (0.59-1.58)0.886ESAD1.79 (0.84-3.81)0.13
ESCC0.46 (0.20-1.10)0.08ESCA0.97 (0.54-1.74)0.925
GBM1.20 (0.85-1.69)0.295ESCC0.68 (0.26-1.80)0.439
GBMLGG4.90 (3.68-6.54)<0.001GBM1.24 (0.86-1.79)0.252
HNSC1.31 (1.00-1.71)0.05GBMLGG5.22 (3.84-7.10)<0.001
KICH8.39 (1.05-67.12)0.045HNSC1.46 (1.03-2.07)0.033
KIRC1.61 (1.19-2.19)0.002KICH6.38 (0.77-52.97)0.086
KIRP2.73 (1.46-5.11)0.002KIRC2.28 (1.52-3.42)<0.001
LAML0.98 (0.64-1.49)0.914KIRP7.40 (2.56-21.35)<0.001
LGG2.60 (1.78-3.82)<0.0012.72 (1.81-4.07)
LGG<0.001
LIHC1.83 (1.29-2.59)0.001
LIHC2.24 (1.41-3.54)0.001
LUAD1.36 (1.02-1.81)0.037
LUADLUSC1.12 (0.92-1.36)0.264LUAD1.45 (1.01-2.10)0.045
LUSC0.96 (0.73-1.26)0.779LUADLUSC1.23 (0.93-1.62)0.147
MESO3.46 (2.10-5.71)<0.001LUSC1.41 (0.92-2.16)0.118
OS0.83 (0.45-1.54)0.557MESO3.77 (2.00-7.12)<0.001
OSCC1.49 (1.08-2.06)0.016OSCC1.56 (1.03-2.36)0.034
OV0.98 (0.76-1.27)0.887OV0.92 (0.70-1.22)0.56
PAAD1.83 (1.20-2.78)0.005PAAD1.75 (1.09-2.81)0.02
PCPG7.53 (0.92-61.38)0.059PCPG5.50 (0.64-47.25)0.12
PRAD1.96 (0.50-7.61)0.331PRAD3.29 (0.37-29.58)0.289
READ0.97 (0.45-2.11)0.945READ1.85 (0.62-5.54)0.269
SARC1.30 (0.88-1.94)0.191SARC1.31 (0.85-2.03)0.225
SKCM1.29 (0.98-1.68)0.068
STAD0.93 (0.67-1.29)0.654SKCM1.29 (0.97-1.72)0.082
TGCT2.81 (0.29-27.06)0.371STAD0.89 (0.59-1.35 ...0.59
THCA1.31 (0.49-3.53)0.589TGCT1.83 (0.17-20.24 ...0.62
THYM0.17 (0.04-0.86)0.032THYM0.55 (0.07-4.19)0.566
UCEC2.13 (1.39-3.27)0.001UCEC3.07 (1.76-5.36)<0.001
UCS1.16 (0.59-2.28)0.671UCS1.26 (0.60-2.63)0.542
UVM3.09 (1.26-7.61)0.014UVM4.11 (1.49-11.29)0.006

0.0

2.5

5.0

7.5

10.0

0

2

4

6

8

FIGURE 4 Prognostic analysis of AURKA in pan-cancer. (A) Forest plot of overall survival prognostic analysis of AURKA in pan-cancer. (B) Disease-specific survival prognostic analysis of AURKA in pan-cancer.

Construction and prognostic value of IRSS

The associations of 42 DEFGs with overall survival in ACC were calculated separately using univariate survival analysis. Six

genes were significantly related to ACC prognosis, including AURKA, TNFAIP, HELLS, STMN1, FANCD2, and SLC4OA1. The high expression of the six genes associated with poor prognosis in ACC, as shown in Figure 2A, greatly impacted

the overall survival of ACC patients and was followed by LASSO regression analysis.

LASSO regression can improve model accuracy and interpretability while also eliminating the issue of collinearity between independent variables (Yu et al., 2021). The results of Figures 2C, D determined that the model fit best when the penalty coefficient was 3, and the corresponding three immune genes, TNFAIP3, AURKA, and HELLS, were included in the model (Figure 2E).

Each patient’s risk score was calculated as previously described (Peng et al., 2022). Furthermore, the risk score of each ACC patient was directly computed using the above formula. The samples were then divided into high- and low-risk groups, which were then grouped based on the median. The KM curve results showed that the high-risk group had a worse prognosis than the low-risk group (Figure 3A, log-rank p 0.001; HR = 11.63% CI = 3.9634,12).

The area under the receiver operating characteristic (ROC) curve (AUC) was used to assess IRSS’s prognostic predictive value in ACC patients. The receiver operating characteristic curves are referred to as ROC curves, with sensitivity as the ordinate and 1-specificity as the abscissa (DeLong et al., 1988). The AUC is a probability value ranging from 0.5 to -1 that is used to evaluate the accuracy of the model prediction; a more extensive area indicates higher accuracy. In the current study, the greater its value, the greater the agreement between predicted and actual overall survival.

The area under the curve (AUC) was 0.918 (3-year OS), 0.902 (5-year OS), and 0.853 (7-year OS), as shown in Figure 3B, indicating that the prediction model was well established. We also created ROC curves for the effect of AURKA alone on survival time in ACC patients, with AUCs of 0.906 (2-year OS), 0.920 (3-year OS), and 0.818 (5-year OS) (Figure 3C). The above results demonstrated the model’s robustness and accuracy in predicting patient prognosis. Simultaneously, we discovered that AURKA’s single-gene and polygenic prognostic models have similar prediction results. AURKA is a common intersection of ferroptosis- related genes and three differentially expressed gene sets in the GEO database. As a result, we make the bold assumption that AURKA is a crucial gene associated with ferroptosis prognosis in ACC. Then, we looked at AURKA’s pan-cancer expression and its relationship to ACC pathological stage.

Expression of AURKA in pan-cancer

The expression level of AURKA was higher in ACC tissue (Figure 3D), and the expression level of AURKA in different stages of ACC was shown in Figure 3E, indicating that the expression level of AURK increased with the progression of ACC. We then investigated AURKA expression in pan- cancer, and the results show that AURKA was highly expressed in all 31 tumors except PCPG and THCA (Figure 3F).

Prognostic analysis of AURKA expression in ACC and other cancers

The correlation of AURKA expression with overall survival and disease-specific survival in 40 TCGA tumors was calculated using univariate survival analysis. AURKA expression, as shown in Figure 4A, significantly impacted overall survival in multiple cancers.

In addition to COAD, COADREAD, DLBC, ESCC, and THYM, forest plot results revealed that high AURKA expression was associated with poor patient prognosis. Figure 4B depicts the correlation of AURKA expression with disease-specific survival, demonstrating that in ACC, GBMLGG, KICH, KIRC, KIRP, and LGG, patients with high AURKA expression had significantly lower disease-specific survival than patients with common AURKA expression. Overall, the findings suggest that AURKA could be used to predict the prognosis of ACC and other cancers.

Correlation of AURKA with immune cells in the pan-cancer microenvironment

It has been studied whether AURKA expression correlates with immune infiltration in ACC or other types of cancer. The findings revealed that AURKA expression is associated with the level of immune infiltration in various tumors. Particularly Th2 cells. AURKA was significantly positively correlated with Th2 cells in all 40 cancers studied, and it was the first positive correlation. We also chose 12 cancers to map the relationship between AURKA and immune cells in these cancer microenvironments (GBM, LUSC, LUAD, TGCT, CESC, COADREAD, SARC, ACC, KICH, ESAD, STAD, READ). Figure 5 shows that, in addition to Th2 cells, many other immune cells were negatively correlated with AURKA. The killer CD8+T regulated by Th1 was the main focus of the previous immunotherapy study for AURKA. Perhaps Th1- executing B Cells will have an unanticipated effect on AURKA targeted therapy. AURKA may also inhibit other immune cells in the tumor microenvironment, though the specific mechanism is unknown.

Xiantao Academic then created a correlation chart of AURKA expression levels in LUAD, SARC, ACC, ESCA, STAD, immune score, matrix score, and calculation score, which were all negatively correlated (Figure 6).

Correlation of AURKA expression with immune checkpoints

More than 40 common immune checkpoint genes were analyzed, as was the relationship between AURKA expression and immune checkpoint gene expression. Figure 7 depicts the results. AURKA was

GBM

LUSC

LUAD

Th2 cells

Th2 oulis

Th2 cells

T helper cells

T helper cells

Tgd

Tom

Tgđ

NK CD55dim cells

Tgd

Tem

T helper cells

NK cells

Tem

aDC

CD8 T cels

aDC

TReg

TReg

P value

Th17 cells

P value

Cytotoxic cells

P value

Tem

0.75

NK CD56dim cells

PDC

0.50

Eosinophils

0.2

Tem

0.4

Neutrophils -

0.3

NK CD56dim cells

0.26

NK CD56bright cells

0.1

Thi cells

02

B cells

0.00

TReg

Tom -

0.1

0.0

NK CD56bright cells

Correlation

Cytotoxic cells

Correlation

NK CD55bright cells

Correlation

Thi cells

0.2

T cells

0.1

T cells -

0.2

TFH

0.4

CDB T cells

0.2

Th17 cells B cells

0.4

T cells

0.6

TFH

0.3

0.6

Mast cells

Neutrophils

Macrophages CD8 T cells

aDC

Th1 cells

DC

NK cells

pDC

Cytotoxic cells

Macrophages

NK cells -

Th17 cels

B cells

TFH-

iDC

PDC

DC

Neutrophils

DC

Eosinophils

Macrophages

iDC

iDC

Eosinophils

Mast cells

Mast cells

-0.4

-0.2

0.0

0.2

0.4

0.6

0.8

-0.4

-0.2

0.0

0.2

-0.6

-0.4

-0.2

0.0

0.2

0.4

0.6

0.8

Correlation

Correlation

Correlation

TGCT

CESC

COADREAD

Th2 cells

Th2 cells

Th2 cells

NK CD56bright cels

T helper cells

T helper cells

Tom

Tcm

Th17 cells

NK cells

Tgd

NK CD66dim cells

T helper cells

Eosinophils

aDC

Tgd

P value

Tem

Tgd -

TReg

0.75

Th17 cells

P value

Tem

aDC

0.50

NK cells

0.8

Neutrophils

P value

Macrophages Mast cells

NK CD56bright cells

0.6

0.25

0.4

Eosinophils

0.75

aDC

0.2

TReg-

0.50

DC

Macrophages

B cells

0.25

NK CD56dim cells

Correlation

0.1

NK CD56dim cells

Correlation

DC

T cells

02

Thi cells

0.1

Tem -

Correlation

Neutrophils

0.3

Neutrophils

0.2

Th1 cells-

0.1

Thi cells

0.4

TFH

0.3

Macrophages CD8 T cells-

0.2

Th17 cells

0.5

TReg

Eosinophils

T cells

Mast cells

iDC

DC

Cytotoxic cells

Cytotoxic cells

CDB T cells

T cells

B cells

IDC

TFH -

CDB T cells

Cytotoxic cells

PDC

TFH

B cells

NK CD56bright cells

Tem

Mast cells

NK cells

PDC

PDC

IDC

-0.4

-0.2

0.0

0.2

-0.3

-0.2

-0.1

0.0

0.1

0.2

0.3

-0.3

-0.2

-0.1

0.0

0.1

0.2

0.3

Correlation

Correlation

Correlation

SARC

ACC

KICH

Th2 cells

Th2 cells

Th2 cells

T helper cells

aDC

aDC

TReg

T helper o

T helper cells

Neutrophils

Tgd

Tcm

Th1 cells

TReg

Macrophages

NK CD56bright cels

Tcm

Eosinophils

aDC

P value

NK cells

P value

Neutrophils

P value

Macrophages

0.8

0.6

Tem

0,6

NK cells

0.8

NK CD56dim cels

0.8

0.4

IDC

0.4

B cells

0.4

Th17 cells

0.2

DC

0.2

TReg

0.2

Tgd

0.0

Neutrophils

0.0

TFH

T cells

Correlation

NK CD56bright cells Macrophages

Correlation

Tgd

Correlation

DC

0.2

0.2

NK CD56dim cells

0.1

IDC

0.4

PDC

0.4

Mast cells

0.2

Tom

0.6

NK CD56dim cells

0.6

Th1 cells

0.3

TFH

Th17 cell

T cells

0.4

B cells

Eosinophils

PDC -

Tem

Th1 cells

Tem -

Cytotoxic cells

T cells

NK CD55bright cells

Mast cells

B cells

CD8 T cells

CD8 T cells

TFH

iDC

Eosinophils

CDB T cells Mast cells

Th17 cells

NK cells

DC

PDC

Cytotoxic cells

Cytotoxic cells

-0.4

-0.2

0.0

0.2

0.4

0.6

0.8

-0.6

-0.4

-0.2

0.0

0.2

0.4

0.6

0,8

-0,4

-0.2

0.0

0.2

0.4

Correlation

Correlation

Correlation

ESAD

STAD

READ

Th2 cells

Th2 cells

Th2 cells

Tgd

Th17 cells

T helper cells

Th17 cells

NK CD56bright cells

aDC-

T helper cels

T helper cells

Tom -

NK CD56bright cells

NK CD56dim cells

Tgd -

NK cels

aDC

Macrophages NK CD56dim cells

aDC

P value

Neutrophils

P value

P value

Neutrophils

0.6

0,75

Thi cells

0.75

DC-

0.4

Tgd

TReg

0.50

0.25

TReg

0.50

Tem

0.2

Th1 cells

0.25

Eosinophils

Macrophages

0.00

Th17 cells

Correlation

Correlation

B cells

Thi cells

Tem -

Comelation

Macrophages

0.1

Eosinophils

0,1

0.1

Tom

0.2

Tcm

0.2

Neutrophils

02

iDC

0.3

iDC

T cells

0.3

0.4

Tem

0.3

0.4

PDC

TReg

TFH

DC

0.4

PDC

Cytotoxic cells

Mast cells

TFH

DC

CD8 T cells

T cells

T cells

Cytotoxic cells

NK CD56dim cells

B cells

TFH -

B cels

NK cells

Eosinophils

Cytotoxic cells

CDB T cells

NK CD56bright cells-

CDB T cells

Mast cells

NIK cells

Mast cells

PDC

iDC

-0.4

-0.2

0.0

0.2

0.4

-0.4

-0.2

0.0

0.2

0.4

-0.4

-0.2

0.0

0.2

0.4

Correlation

Correlation

Correlation

FIGURE 5 Lollipop plot of AURKA’s association with immune cells in 12 cancers.

FIGURE 6 Correlation of AURKA expression level with immune score, stroma score, calculated score in LUAD (A), SARC (B), ACC (C), ESCA (D), STAD (E).

A

2000

4000

3000

StromalScore

1000

ESTIMATEScore

ImmuneScore

2000

2000

.

0

1000

0

-1000

Spearman

Spearman

0

Dearman

r= - 0.224

r =- 0.219

-2000

P < 0.001

-2000

o

r= - 0.182

P < 0.001

-1000

P <0.001

2

4

6

8

2

4

6

8

2

4

6

8

The expression of AURKA Log2 (TPM+1)

The expression of AURKA Log2 (TPM+1)

The expression of AURKA Log2 (TPM+1)

B

6000

4000

2000

.

StromalScore

ESTIMATEScore

4000

3000

.

ImmuneScore

1000

2000

2000

1000

0

0

0

Spearman

-2000

Spearman

-1000

Spearman

-1000

r= - 0.284

-0.199

r= - 0.119

P < 0.001

P = 0.001

2

-2000

P = 0.053

3

4

5

6

7

2

3

4

5

6

7

2

3

4

5

6

7

The expression of AURKA Log2 (TPM+1)

The expression of AURKA Log2 (TPM+1)

The expression of AURKA Log2 (TPM+1)

C

.

4000

2000

.

1000

StromalScore

ESTIMATEScore

2000

ImmuneScore

1000

0

0

.

0

.

-1000

.

00

.

Spearman

®

F-0.258

-2000

Spearman

Spearman

P = 0.022

= - 0.270

-1000

= - 0.292

-2000

P = 0.017

.

P = 0.009

2

4

6

2

4

6

2

4

6

The expression of AURKA Log2 (TPM+1)

The expression of AURKA Log2 (TPM+1)

The expression of AURKA Log2 (TPM+1)

D

2000

.

6000

3000

1000

4000

StromalScore

ESTIMATEScore

ImmuneScore

2000

0

2000

1000

-1000

0

.

.

0

Spearman

®

-2000

Spearman

·

-2000

0.188

9-0.214

Spearman

-1000

45-0.217

P = 0.017

-4000

P = 0.006

P = 0.006

3

4

5

6

7

3

4

5

6

7

3

4

5

6

7

The expression of AURKA Log2 (TPM+1)

The expression of AURKA Log2 (TPM+1)

The expression of AURKA Log2 (TPM+1)

E

2000

3000

.

4000

StromalScore

1000

ESTIMATEScore

.

ImmuneScore

2000

2000

.

0

1000

®

0

-1000

0

Spearmar

Spearman

Spearman

T= - 0,422

= - 0.421

r= - 0,334

-2000

P < 0.001

-2000

··· P < 0.001

-1000

P < 0.001

3

5

7

9

3

5

7

9

3

5

7

9

The expression of AURKA Log2 (TPM+1)

The expression of AURKA Log2 (TPM+1)

The expression of AURKA Log2 (TPM+1)

BTLA

CD200

TNFRSF14

NRP1

LAIR1

TNFSF4

CD244

LAG3

ICOS

CD40LG

CTLA4

CD48

CD28

CD200R1

HAVCR2

ADORA2A

CD276

KIR3DL1

CD80

PDCD1

Correlation

1.0

LGALS9

0.5

CD160

0.0

TNFSF14

-0.5

IDO2

-1.0

ICOSLG

TMIGD2

VTCN1

IDO1

PDCD1LG2

HHLA2

TNFSF18

BTNL2

CD70

TNFSF9

TNFRSF8

CD27

TNFRSF25

VSIR

TNFRSF4

CD40

TNFRSF18

TNFSF15

TIGIT

CD274

CD86

CD44

TNFRSF9

ACC

p1 LIHC

p2

COAD

p3

THYM

p4

E

p5

PAAD

p6

LUSC

p7

GBMLGG

p8

BRCA

p9

PRAD

p10

KIRC

1

ESCA

p12

LUAD

p13

UCEC

p14

HNSC

p15

THCA

p16

COADREAD

p17

BLCA

p18

FIGURE 7 Heatmap of AURKA’s association with immune checkpoints in a broad range of cancers.

A

MSH2

MSH3

MSH6

Correlation

1.0

MLH1

0.5

PMS2

0.0

EPCAM

-0.5

MGMT

-1.0

ALKBH2

ALKBH3

ACC

p1

LIHC

p2

COAD

p3

THYM

p4

STAD

p5

PAAD

p6

LUSC

p7

GBMLGG

p8

BRCA

p9

PRAD

p10

KIRC

p11

ESCA

p12

LUAD

p13

UCEC

p14

HNSC

p15

THCA

p16

COADREAD

p17

BLCA

p18

B

Correlation

DNMT1

1.0

0.5

DNMT3A

0.0

DNMT3B

-0.5

ACC

p1

LIHC

p2

COAD

p3

THYM

p4

STAD

p5

PAAD

p6

LUSC

p7

GBMLGG

p8

BRCA

p9

PRAD

p10

KIRC

p11

ESCA

p12

LUAD

p13

UCEC

p14

HNSC

p15

THCA

p16

COADREAD

p17

BLCA

p18

-1.0

FIGURE 8 The relationship between AURKA expression and DNA repair gene and methyltransferase expression. (A) Heatmap of correlations between AURKA and DNA repair genes. (B) Heatmap of correlation between AURKA and methyltransferase genes.

positively correlated with the presentation of immune checkpoint genes in many cancers, which supports our findings in Figure 5. Meanwhile, we discovered that AURKA was significantly negatively associated with most checkpoint genes in thymic carcinoma. The thymus is the site of T cell maturation and a mechanism that inhibits the AURKA-mediated increase in immune checkpoint expression, protecting T cells in the thymus.

The relationship between AURKA expression and DNA repair gene and methyltransferase expression

AURKA was found to be associated with DNA repair genes as well as methyltransferase genes in several common cancers, as shown in Figures 8A, B. AURKA may have an indirect effect on cancer development and progression by modulating epigenetic status.

Discussion

Although the incidence of adrenal cortical carcinoma is very low, it is one of the most aggressive solid tumors with a poor

prognosis (Yeoh et al., 2022). Furthermore, the recurrence of ACC patients after surgery is still common. As a result, more biomarkers are required for more accurate predictive detection in ACC patients to improve the detection of postoperative risk. The discovery of predictive cancer biomarkers can aid in predicting each patient’s prognosis (Mizdrak et al., 2021; Lippert et al., 2022; Waszut and Taylor, 2022). Using robust rank analysis and a PPI network, XiaoH et al. identified five genes (TOP2A, NDC80, CEP55, CDKN3, and CDK1) that could predict the prognosis of ACC (Xiao et al., 2018). Giordano et al.’s laid the groundwork for ACC molecular classification and prediction, as well as a rich source of potential diagnostic and prognostic markers (Xu et al., 2019).

Ferroptosis is a new iron-dependent programmed cell death method discovered that can induce cell death by promoting cellular lipid peroxidation. It is involved in the occurrence and development of many diseases and plays an essential regulatory role in disease processes. Related studies have shown that ferroptosis plays a role in the progression of various cancers. For example, inhibiting glutathione synthesis in ccRCC in clear cell renal cell carcinoma can induce ferroptosis and inhibit tumor growth (Miess et al., 2018); According to other research (Liu et al., 2020; Chen et al., 2021b; Lu et al., 2022),

ferroptosis attenuates the viability of glioma cells, and activation of ferroptosis inhibits glioma cell proliferation. Inhibition of ferroptosis accelerates glioma proliferation and metastasis and promotes angiogenesis and malignant transformation of gliomas. One study discovered that ferroptosis sensitivity was significantly increased in adrenocortical carcinoma and proposed ferroptosis induction as a treatment option for ACC (Belavgeni et al., 2019).

We obtained six critical genes in this study by crossing the up-regulated genes in ACC with the genes associated with overall survival in ACC. Three of them were chosen to build a polygenic model. The AURKA prediction model and the polygenic model produced very similar results. Meanwhile, AURKA is the point of convergence for the ferroptosis-related gene set and the GEO databases. As a result, we boldly identified AURKA as a critical gene in ACC ferroptosis. We then looked at AURKA expression in ACC and other cancers to see if it had any predictive value. The findings revealed that AURKA was highly expressed in ACC and most cancers and that its expression level increased as ACC progressed. It is consistent with previous research findings (Naso et al., 2021; Sankhe et al., 2021; Ng et al., 2022; Wang et al., 2022). Related studies have also shown high levels of AURKA as an indicator of poor prognosis in bladder cancer. It is also associated with the development and prognosis of rectal cancer, hepatocellular carcinoma, and head and neck cancer (Lu et al., 2021; Tsepenko et al., 2021; Guo et al., 2022; Huang et al., 2022). This research discovered that high AURKA was related to a bad prognosis in various malignancies by creating a deep forest graph and feeding back the association between AURKA, overall survival, and disease-specific survival. It gives compelling evidence that ARUKA may be used to predict the prognosis of ACC and other malignancies.

In addition, we investigated the relationship between AURKA and immune cells in the pan-cancer microenvironment. We discovered that AURKA had a substantial positive link with Th2 cells in all 40 malignancies studied, and these were all the first positive correlations. We next chose 12 malignancies to investigate the association between AURKA and immune cells in them, finding that all immune cells except Th2 cells were adversely connected with AURKA. Previous research (Bustos- Moran et al., 2019; Sun et al., 2021; Long and Zhang, 2022) has shown that AURKA may impact T cells, reshape the immunosuppressive tumor microenvironment, apoptosis, and hypoxia and hence contribute to immunological control, particularly CD8+ T cells that govern Th1 regulation. For example, studies (Han et al., 2020) suggest that decreasing Aurora-A activity or deleting the AURKA gene might boost IL10-induced infiltration and growth of CD8+ T cells in malignancies. Th1-executing B Cells may have unanticipated impacts on AURKA targeted treatment. AURKA may also block other immune cells in the tumor microenvironment, albeit the particular mechanism is unknown. We next looked at the relationship between AURKA expression level and immunological score, stromal score, and computational score in

five malignancies (ACC, SARC, LUAD, ESCA, and STAD), which were all shown to be negatively linked. AURKA has also been identified to affect tumor immunological patterns in diverse malignancies by controlling the expression of particular immune checkpoint genes, according to subsequent research. AURKA was shown to be favorably connected with the indication of immune checkpoint genes, which supports our prior results from a pan-cancer immunological correlation study. The discovery of immunological checkpoints opens up new avenues for tumor therapy. Immune checkpoint inhibitors have been employed in treating many tumors recently, and their effectiveness and safety have been objectively validated (Cai et al., 2022; Minegishi et al., 2022). In addition, we discovered an intriguing phenomenon. AURKA was strongly inversely related to most checkpoint genes in thymic cancer. The thymus is the location of T cell maturation. Thymic cancer has a mechanism that blocks the AURKA-mediated rise in immune checkpoint expression, safeguarding T cells in the thymus.

DNA repair capacity, which is primarily determined by repair gene expression levels, is the first line of defense against genotoxic stress, which causes metabolic changes, inflammation, and cancer, and is also required for maintaining genome stability and protecting cells from endogenous and exogenous DNA traumatic injuring (Shao et al., 2022; Zuo et al., 2022).

This study looked at nine DNA repair genes: MSH2, MSH3, MSH6, MLH1, PMS2, EPCAM, MGMT, ALKBH2, and ALKBH3. In most malignancies, AURKA expression was strongly positively linked with DNA repair genes, according to the findings. Furthermore, the results of this study revealed that the levels of ARUKA and the methyltransferase gene expression exhibited a substantial positive link in a range of malignancies.

Conclusion

To summarize, we did differential expression analysis on the GEO database data, obtaining DEFGs by intersecting with ferroptosis-related genes and exploring some information from them. The significant result is that AURKA is a critical gene for the prognosis of ferroptosis in ACC and can be exploited as an ACC biomarker. The expression of ARUKA is connected with the tumor microenvironment and the number of immune cells in the pan-cancer study, which can impact cancer growth by controlling the level of immune cells, DNA repair, and DNA methylation. This result can only be reached from bioinformatics research, and thus further biological tests are required to demonstrate ARUKA’s probable relevant activities, action mechanisms, and signaling pathways in ACC ferroptosis. It is believed that this work would aid in related research while providing additional biological information about the mechanism of AURKA in tumor immunity and the tumor microenvironment in future research.

Data availability statement

The original contributions presented in the study are included in the article/Supplementary Materials, further inquiries can be directed to the corresponding author.

Author contributions

KL and YZ designed the study; KL and XY wrote the initial manuscript; XY, BW, and YZ collected data; KL. YZ, XY, BW, and YZ analyzed data and contributed in writing the revised manuscript.

Funding

This research was supported by Natural Science Foundation of Heilongjiang Province (LH2019H095) and Outstanding Young Talents Project of Heilongjiang Natural Science Foundation (YQ2022H015).

References

Alyateem, G., and Nilubol, N. (2021). Current status and future targeted therapy in adrenocortical cancer. Front. Endocrinol. 12, 613248. doi:10.3389/fendo.2021. 613248

Belavgeni, A., Bornstein, S. R., von Massenhausen, A., Tonnus, W., Stumpf, J., Meyer, C., et al. (2019). Exquisite sensitivity of adrenocortical carcinomas to induction of ferroptosis. Proc. Natl. Acad. Sci. U. S. A. 116, 22269-22274. doi:10.1073/pnas.1912700116

Bustos-Moran, E., Blas-Rus, N., Alcaraz-Serna, A., Iborra, S., Gonzalez-Martinez, J., Malumbres, M., et al. (2019). Aurora A controls CD8(+) T cell cytotoxic activity and antiviral response. Sci. Rep. 9, 2211. doi:10.1038/s41598-019-38647-y

Cai, Z., Zhan, P., Song, Y., Liu, H., and Lv, T. (2022). Safety and efficacy of retreatment with immune checkpoint inhibitors in non-small cell lung cancer: A systematic review and meta-analysis. Transl. Lung Cancer Res. 11, 1555-1566. doi:10.21037/tlcr-22-140

Chen, Q., Wang, W., Wu, Z., Chen, S., Chen, X., Zhuang, S., et al. (2021). Over- expression of IncRNA TMEM161B-AS1 promotes the malignant biological behavior of glioma cells and the resistance to temozolomide via up-regulating the expression of multiple ferroptosis-related genes by sponging hsa-miR-27a-3p. Cell Death Discov. 7, 311. doi:10.1038/s41420-021-00709-4

Chen, X., Yan, L., Jiang, F., Lu, Y., Zeng, N., Yang, S., et al. (2021). Identification of a ferroptosis-related signature associated with prognosis and immune infiltration in adrenocortical carcinoma. Int. J. Endocrinol. 2021, 4654302. doi:10.1155/2021/4654302

Cheng, Y., Kou, W., Zhu, D., Yu, X., and Zhu, Y. (2021). Future directions in diagnosis, prognosis and disease monitoring of adrenocortical carcinoma: Novel non-invasive biomarkers. Front. Endocrinol. 12, 811293. doi:10.3389/fendo.2021.811293

Du, R., Huang, C., Liu, K., Li, X., and Dong, Z. (2021). Targeting AURKA in cancer: Molecular mechanisms and opportunities for cancer therapy. Mol. Cancer 20, 15. doi:10.1186/s12943-020-01305-3

Faron, M., Lamartina, L., Hescot, S., Moog, S., Deschamps, F., Roux, C., et al. (2022). New endpoints in adrenocortical carcinoma studies: A mini review. Endocrine 77, 419-424. doi:10.1007/s12020-022-03128-2

Guo, J., Li, W., Cheng, L., and Gao, X. (2022). Identification and validation of hub genes with poor prognosis in hepatocellular carcinoma by integrated bioinformatical analysis. Int. J. Gen. Med. 15, 3933-3941. doi:10.2147/IJGM.S353708

Han, J., Jiang, Z., Wang, C., Chen, X., Li, R., Sun, N., et al. (2020). Inhibition of aurora-A promotes CD8(+) T-cell infiltration by mediating IL10 production in cancer cells. Mol. Cancer Res. 18, 1589-1602. doi:10.1158/1541-7786.MCR-19-1226

Conflict of interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

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Supplementary material

The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fgene. 2022.996180/full#supplementary-material

Huang, C., Cheng, Y., Li, W., Huang, Y., Luo, H., Zhong, C., et al. (2022). Examining the mechanisms of huachansu injection on liver cancer through integrated bioinformatics analysis. Recent Pat. anticancer. Drug Discov. 17. doi:10.2174/1574892817666220511162046

Jiang, X., Stockwell, B. R., and Conrad, M. (2021). Ferroptosis: Mechanisms, biology and role in disease. Nat. Rev. Mol. Cell Biol. 22, 266-282. doi:10.1038/s41580-020-00324-8

Lippert, J., Fassnacht, M., and Ronchi, C. L. (2022). The role of molecular profiling in adrenocortical carcinoma. Clin. Endocrinol. 97, 460-472. doi:10.1111/ cen.14629

Liu, H. J., Hu, H. M., Li, G. Z., Zhang, Y., Wu, F., Liu, X., et al. (2020). Ferroptosis- related gene signature predicts glioma cell death and glioma patient progression. Front. Cell Dev. Biol. 8, 538. doi:10.3389/fcell.2020.00538

Long, S., and Zhang, X. F. (2022). AURKA is a prognostic potential therapeutic target in skin cutaneous melanoma modulating the tumor microenvironment, apoptosis, and hypoxia. J. Cancer Res. Clin. Oncol. doi:10.1007/s00432-022-04164-1

Lu, H., Li, L., Sun, D., Duan, Y., Yue, K., Wu, Y., et al. (2021). Identification of novel hub genes associated with lymph node metastasis of head and neck squamous cell carcinoma by completive bioinformatics analysis. Ann. Transl. Med. 9, 1678. doi:10.21037/atm-21-5704

Lu, M., Zhou, Y., Sun, L., Shafi, S., Ahmad, N., Sun, M., et al. (2022). The molecular mechanisms of ferroptosis and its role in glioma progression and treatment. Front. Oncol. 12, 917537. doi:10.3389/fonc.2022.917537

Luo, H., and Ma, C. (2021). A novel ferroptosis-associated gene signature to predict prognosis in patients with uveal melanoma. Diagn. (Basel) 11, 219. doi:10. 3390/diagnostics11020219

Mete, O., Erickson, L. A., Juhlin, C. C., de Krijger, R. R., Sasano, H., Volante, M., et al. (2022). Overview of the 2022 WHO classification of adrenal cortical tumors. Endocr. Pathol. 33, 155-196. doi:10.1007/s12022-022-09710-8

Miess, H., Dankworth, B., Gouw, A. M., Rosenfeldt, M., Schmitz, W., Jiang, M., et al. (2018). The glutathione redox system is essential to prevent ferroptosis caused by impaired lipid metabolism in clear cell renal cell carcinoma. Oncogene 37, 5435-5450. doi:10.1038/s41388-018-0315-z

Minegishi, S., Kinguchi, S., Horita, N., Namkoong, H., Briasoulis, A., Ishigami, T., et al. (2022). Immune checkpoint inhibitors do not increase short-term risk of hypertension in cancer patients: A systematic literature review and meta- analysis. Hypertension 79, 2611-2621. doi:10.1161/HYPERTENSIONAHA.122. 19865

Mizdrak, M., Ticinovic Kurir, T., and Bozic, J. (2021). The role of biomarkers in adrenocortical carcinoma: A review of current evidence and future perspectives. Biomedicines 9, 174. doi:10.3390/biomedicines9020174

Naso, F. D., Boi, D., Ascanelli, C., Pamfil, G., Lindon, C., Paiardini, A., et al. (2021). Nuclear localisation of aurora-A: Its regulation and significance for aurora- A functions in cancer. Oncogene 40, 3917-3928. doi:10.1038/s41388-021-01766-w

Ng, C. K. Y., Dazert, E., Boldanova, T., Coto-Llerena, M., Nuciforo, S., Ercan, C., et al. (2022). Integrative proteogenomic characterization of hepatocellular carcinoma across etiologies and stages. Nat. Commun. 13, 2436. doi:10.1038/ s41467-022-29960-8

Peng, J., Lu, Y., Chen, L., Qiu, K., Chen, F., Liu, J., et al. (2022). The prognostic value of machine learning techniques versus cox regression model for head and neck cancer. Methods 205, 123-132. doi:10.1016/j.ymeth.2022.07.001

Pitsava, G., Maria, A. G., and Faucz, F. R. (2022). Disorders of the adrenal cortex: Genetic and molecular aspects. Front. Endocrinol. 13, 931389. doi:10.3389/fendo.2022.931389

Sankhe, K., Prabhu, A., and Khan, T. (2021). Design strategies, SAR, and mechanistic insight of Aurora kinase inhibitors in cancer. Chem. Biol. Drug Des. 98, 73-93. doi:10.1111/cbdd.13850

Shao, X., Yang, X., Liu, Y., Song, Q., Pan, X., Chen, W., et al. (2022). Genetic polymorphisms in DNA repair genes and their association with risk of cervical cancer: A systematic review and meta-analysis. J. Obstet. Gynaecol. Res. 48, 2405-2418. doi:10.1111/jog.15325

Sun, S., Zhou, W., Li, X., Peng, F., Yan, M., Zhan, Y., et al. (2021). Nuclear Aurora kinase A triggers programmed death-ligand 1-mediated immune suppression by activating MYC transcription in triple-negative breast cancer. Cancer Commun. 41, 851-866. doi:10.1002/cac2.12190

Tang, J., Yang, L., Li, Y., Ning, X., Chaulagain, A., Wang, T., et al. (2021). ARID3A promotes the development of colorectal cancer by upregulating AURKA. Carcinogenesis 42, 578-586. doi:10.1093/carcin/bgaa118

Tian, L., Li, X., Zheng, H., Wang, L., Qin, Y., and Cai, J. (2022). Fisher discriminant model based on LASSO logistic regression for computed tomography imaging diagnosis of pelvic rhabdomyosarcoma in children. Sci. Rep. 12, 15631. doi:10.1038/s41598-022-20051-8

Tsepenko, V. V., Shkavrova, T. G., Cherkesov, V. N., Golub, E. V., and Mikhailova, G. F. (2021). Asynchronous DNA replication of biallelically expressed genes in human peripheral blood lymphocytes as a prognostic sign of cancer. Sovrem. Tekhnologii Med. 13, 33-38. doi:10.17691/stm2021.13.3.04

Wang, F., Zhang, H., Wang, H., Qiu, T., He, B., and Yang, Q. (2022). Combination of AURKA inhibitor and HSP90 inhibitor to treat breast cancer with AURKA overexpression and TP53 mutations. Med. Oncol. 39, 180. doi:10. 1007/s12032-022-01777-x

Waszut, U., and Taylor, N. F. (2022). Use of dissected paraffin block tissue as a source of mRNA for transcriptional profiling and biomarker identification: A review, with preliminary findings in adrenocortical carcinoma tissue. Acta Biochim. Pol. 69, 273-281. doi:10.18388/abp.2020_5611

Wei, S., Zhang, J., Shi, R., Yu, Z., Chen, X., and Wang, H. (2022). Identification of an integrated kinase-related prognostic gene signature associated with tumor immune microenvironment in human uterine corpus endometrial carcinoma. Front. Oncol. 12, 944000. doi:10.3389/fonc.2022.944000

Xiao, H., Xu, D., Chen, P., Zeng, G., Wang, X., and Zhang, X. (2018). Identification of five genes as a potential biomarker for predicting progress and prognosis in adrenocortical carcinoma. J. Cancer 9, 4484-4495. doi:10.7150/jca. 26698

Xu, W. H., Wu, J., Wang, J., Wan, F. N., Wang, H. K., Cao, D. L., et al. (2019). Screening and identification of potential prognostic biomarkers in adrenocortical carcinoma. Front. Genet. 10, 821. doi:10.3389/fgene.2019.00821

Yeoh, P., Czuber-Dochan, W., Aylwin, S., and Sturt, J. (2022). Lived experience of people with adrenocortical carcinoma and associated adrenal insufficiency. Endocrinol. Diabetes Metab. 5, e341. doi:10.1002/edm2.341

Yu, S. H., Cai, J. H., Chen, D. L., Liao, S. H., Lin, Y. Z., Chung, Y. T., et al. (2021). LASSO and bioinformatics analysis in the identification of key genes for prognostic genes of gynecologic cancer. J. Pers. Med. 11, 1177. doi:10.3390/ jpm11111177

Zhang, P., Chen, X., Zhang, L., Cao, D., Chen, Y., Guo, Z., et al. (2022). POLE2 facilitates the malignant phenotypes of glioblastoma through promoting AURKA-mediated stabilization of FOXM1. Cell Death Dis. 13, 61. doi:10.1038/ s41419-021-04498-7

Zheng, Y., Ji, Q., Xie, L., Wang, C., Yu, C. N., Wang, Y. L., et al. (2021). Ferroptosis-related gene signature as a prognostic marker for lower-grade gliomas. J. Cell. Mol. Med. 25, 3080-3090. doi:10.1111/jcmm.16368

Zuo, C., Lv, X., Liu, T., Yang, L., Yang, Z., Yu, C., et al. (2022). Polymorphisms in ERCC4 and ERCC5 and risk of cancers: Systematic research synopsis, meta- analysis, and epidemiological evidence. Front. Oncol. 12, 951193. doi:10.3389/ fonc.2022.951193