Biological Roles and Mechanisms of Circular RNA in Human Cancers

Figures & data

Figure 1 circRNAs involve in cancer progression by acting as “miRNA sponge” or protein scaffolding. (A) circRNAs competitively bind to miRNA with the participation of Ago2 protein by acting as “miRNA sponge,” consequently release the target gene of miRNA. (B) circRNAs directly interact with RNA-binding proteins (RBPs), or mediate the interaction between proteins by serving as protein scaffolding to regulate the expression of downstream targets.

Figure 2 circRNAs involve in cancer progression by regulation of gene splicing, transcription, translation and cell autophagy. (A) circRNAs influence the progression of human cancers by regulating gene splicing, transcription and translation via interaction with U1 small nuclear RNA (snRNA), RNA polymerase II (Pol II), and alternative splicing regulator. Moreover, circRNA containing AUG sites and open reading frame (ORF) driven by the internal ribosome entry site (IRES) may translate into the functional protein. (B) circRNAs are associated with the tumor cell autophagy through regulating the autophagy-related signaling pathways.

Table 1 Expression and Functional Characterization of circRNAs in Cancers

Type of Cancers	Name of circRNAs	Levels	Target Genes	Mode of Functions	Functional Phenotypes	References
Lung cancer	Circ-UBR5	Down	QK/NOVA1/U1 snRNA	Regulate RNA splicing	Correlate with tumor differentiation	[^Citation18]
	CircNOL10	Down	SCML1	Regulate transcription	Inhibit cell proliferation and promote cell apoptosis	[^Citation38]
	CircPTPRA	Down	miR-96-5p	MiRNA sponge	Suppress EMT and invasion	[^Citation39]
	Circ_0012673	Up	miR-22	MiRNA sponge	Promote cell proliferation	[^Citation53]
	Circ_0007766	Up	Cyclin	Cell cycle regulator	Induce the cell proliferation and migration	[^Citation59]
Breast cancer	CircRNA-MTO1	Down	TRAF4/Eg5	Regulate gene translation	Suppress cell viability and reverse monastrol resistance	[^Citation67]
	CircTADA2A-E6	Down	miR-203a-3p	MiRNA sponge	Suppress breast cancer progression and metastasis	[^Citation68]
	CircPLK1	Up	miR-296-5p	MiRNA sponge	Promote cell growth and invasion	[^Citation70]
	CircBMPR2	Down	miR-553	MiRNA sponge	Inhibit cell proliferation migration and invasion	[^Citation72]
Colorectal cancer	CircITGA7	Down	ITGA7	Regulate gene transcription	Inhibit cell growth and metastasis	[^Citation17]
	Circ_0009361	Down	miR-582	MiRNA sponge	Suppress colorectal cancer progression	[^Citation81]
	Circ_001569	Up	miR-145	MiRNA sponge	Promote cell proliferation and invasion	[^Citation88]
Esophageal Squamous Cell Carcinoma	Circ-PRKCI	Up	miR-3680-3p	MiRNA sponge	Stimulate cell migration and proliferation	[^Citation95]
	CiRS-7	Up	miR-7	MiRNA sponge	Promote growth and metastasis	[^Citation98]
	Circ_0000337	Up	miR-670-5p	MiRNA sponge	Promote cell proliferation, migration and invasion	[^Citation99]
Gastric cancer	CircYAP1	Down	miR-367-5p	MiRNA sponge	Suppress cell proliferation and invasion	[^Citation105]
	Circ-NOTCH1	Up	miR-637	MiRNA sponge	Enhance cell proliferation and invasion;reduce apoptosis;	[^Citation108]
	Circ-DONSON	Up	NURF complex	Protein scaffolding	Facilitate gastric cancer growth and invasion	[^Citation119]
	CircAGO2	Up	HuR protein	Interact with RBPs	Promote the tumorigenesis and aggressiveness	[^Citation120]
Hepatocellular carcinoma	Circ_0101432	Up	miR-1258/miR-622	MiRNA sponge	Inhibit cell apoptosis, promote cell proliferation and invasive	[^Citation126]
	CircZNF652	Up	miR-203/miR-502-5p	MiRNA sponge	Promote cell metastasis	[^Citation127]
	Circ_101280	Up	miR-375	MiRNA sponge	Promote cell proliferation and suppress apoptosis	[^Citation134]
	CircSLC3A2	Up	miR-490-3p	MiRNA sponge	Promote cell proliferation and invasion;	[^Citation135]
	CircARSP91	Down	ULBP1	Immune-association	Increase cell susceptibility to NK cell cytotoxicity;	[^Citation137]
Bladder cancer	CircBCRC-3	Down	miR-182-5p	MiRNA sponge	Suppress cell proliferation	[^Citation150]
	CircUVRAG	Up	miR-223	MiRNA sponge	Inhibit cell growth and metastasis	[^Citation151]
	CircGprc5a	Up	Gprc5a-Peptide	Peptide-dependent manner	Promote Bladder Oncogenesis and Metastasis	[^Citation156]
Pancreatic cancer	Circ_0006215	Up	miR-378a-3p	MiRNA sponge	Promote cell viability, apoptosis and migration	[^Citation161]
	CircRNA_100782	Up	miR-124	MiRNA sponge	Promote cell proliferation	[^Citation162]
	CircZMYM2	Up	miR-335-5p	MiRNA sponge	Promote cell proliferation and induce cell apoptosis	[^Citation163]
	CircRHOT1	Up	miR-26b/miR-125a/miR-330	MiRNA sponge	Promote cell proliferation and cell invasion	[^Citation164]
Papillary thyroid carcinoma	CircBACH2	Up	miR-139-5p	MiRNA sponge	Promote cell proliferation, migration and invasion	[^Citation172]
	CircRAPGEF5	Up	miR-198	MiRNA sponge	Promote cell proliferation, migration and invasion	[^Citation173]
	CircRNA_102171	Up	CTNNBIP1	Protein scaffolding	Promote cell proliferation, migration and invasion;promote cell apoptosis.	[^Citation175]
	Circ-ITCH	Down	miR-22-3p	MiRNA sponge	Suppress cell proliferation and invasion;Promote cell apoptosis	[^Citation176]
Osteosarcoma	CircFAT1	Up	miR-375	MiRNA sponge	Promote cell migration, invasion and tumorigenesis	[^Citation186]
	CircTADA2A	Up	miR-203a-3p	MiRNA sponge	Promotes tumor progression and metastasis	[^Citation188]
Glioblastoma	Circ_0001946	Down	miR-671-5p	MiRNA sponge	Reduce cell proliferation, migration	[^Citation191]
	CircNT5E	Up	miR-422a	MiRNA sponge	Promote cell proliferation, migration and invasion	[^Citation193]
	Circ-SHPRH	Down	SHPRH-146aa	Protein translation	Suppress tumor progression and tumorigenesis	[^Citation196]
Ovarian cancer	circPUM1	Up	miR-615-5p and miR-6753-5p	MiRNA sponge	Promote cell proliferation, migration and invasion and inhibited cell apoptosis	[^Citation199]
	circWHSC1	Up	miR-145 and miR-1182	MiRNA sponge	Promote cell proliferation, migration and invasion and inhibited cell apoptosis	[^Citation200]
	circ_0061140	Up	miR-370	MiRNA sponge	Promote cell proliferation, migration and the EMT	[^Citation207]
Prostate cancer	circUCK2	Down	miR-767-5p	MiRNA sponge	Inhibit cell invasion and proliferation	[^Citation217]
	circFOXO3	Up	miR‐29a‐3p	MiRNA sponge	Promote cell cycle, proliferation and inhibit cell apoptosis	[^Citation218]
	circHIPK3	Up	miRNA-338-3p	MiRNA sponge	Inhibit the proliferative and invasive	[^Citation220]
Myeloid leukemia	circ_0009910	Up	miR-20a-5p	MiRNA sponge	Promote cell proliferation and induced apoptosis	[^Citation229]
	circ_100290	Up	MiR-203	MiRNA sponge	Promote cell proliferation and inhibit apoptosis	[^Citation230]

Note: The expression, molecular targets, functional phenotypes of cirRNA in different cancers were summarized.

Table 2 The Roles in Clinical Relevance and Prognosis of circRNAs in Human Cancers

Type of Cancers	Name of circRNAs	Levels	Clinical Relevancies	References
Lung cancer	Circ_0016760	Up	Associated with advanced TNM stages, lymph node metastasis and adverse prognosis	[^Citation42]
	CircFADS2	Up	Correlated with advanced TNM stage, lymph node metastasis, poor differentiation, tumor size and shorter overall survival	[^Citation46]
	Circ-FOXM1	Up	Associated with lymph node invasion, higher TNM stage and unfavorable prognosis.	[^Citation48]
	CircMTO1	Down	Associated with malignant features and prognosis	[^Citation50]
	Circ_003645	Up	Related to advanced TNM stages, positive lymph node invasion and unfavorable prognosis	[^Citation51]
	Circ_0020123	Up	Correlated with positive lymph node metastasis, advanced TNM stages, and adverse prognosis	[^Citation54]
	CircVANGL1	Up	Associated with tumor size, TNM stage and overall survival	[^Citation60]
Breast cancer	CircANKS1B	Up	Associated with lymph node metastasis and advanced clinical stage	[^Citation65]
	CircKIF4A	Up	Associated with tumor size, lymph node metastasis and TNM Stage	[^Citation69]
	Circ-UBE2D2	Up	Associated with tumor size, lymph node metastasis and TNM Stage and differentiation	[^Citation71]
	Circ_0025202	Down	Correlated with lymphatic metastasis and histological grade	[^Citation73]
	Circ_001783	Up	Correlated with tumor size, lymph node status, TNM stage, ER status, PR status, molecular subtype and Ki-67 index	[^Citation76]
Colorectal cancer	CircVAPA	Up	Correlated with tumor size, Lymphovascular invasion, Differentiation, Distant metastasis lymph node metastasis and TNM stage	[^Citation82]
	Circ_0026344	Down	Correlated with CRC advance and lymphoid node metastasis	[^Citation86]
	CircHIPK3	Up	Correlated with Pathological T category, Lymph node metastasis, Distant metastasis and TNM stage	[^Citation87]
Esophageal Squamous Cell Carcinoma	Circ-TTC17	Up	Associated with TNM stage and Lymph node metastasis	[^Citation94]
	Circ_0006168	Up	Associated with lymph node metastasis and TNM stage	[^Citation96]
	Circ-DLG1	Up	Associated with TNM stage	[^Citation97]
Gastric cancer	CircNRIP1	Up	Associated with Lymphatic invasion and tumor size	[^Citation106]
	Circ_006100	Up	Associated with TNM stage, poor cell differentiation and lymphnode metastasis	[^Citation107]
	Circ-DCAF6	Up	Associated with deeper tumor invasion, positive lymph node metastasis and higher TNM stages	[^Citation109]
Hepatocellular carcinoma	Circ-ZEB1.33	Up	Associated with HBV infection, TNM stages and tumor size	[^Citation124]
	CircDAMTS13	Down	Associated with cirrhosis, tumor size and stage	[^Citation132]
	CircTRIM33-12	Down	Associated with tumor size, encapsulation invasion, vascular invasion and tumor number	[^Citation136]
Bladder cancer	CircCEP128	Up	Associated with tumor size, TNM stage and Lymphatic metastasis	[^Citation149]
	Circ-BPTF	Up	Associated with histological grade and prognosis	[^Citation152]
	Circ-cTFRC	Up	Associated with tumor stage, grade, number of tumors and Lymphatic metastasis	[^Citation154]
	CircUBXN7	Down	Associated with tumor grade and Pathology stage	[^Citation155]
Pancreatic cancer	Circ-PDE8A	Up	Correlated with lymphatic invasion, TNM stage and survival rate	[^Citation165]
Papillary thyroid carcinoma	CircZFR	Up	Correlated with TNM stage and overall survival	[^Citation177]
Osteosarcoma	CircCDR1	Up	Correlated with tumor size, localization, stage and metastasis	[^Citation187]
Glioblastoma	Circ_0029426	Up	Correlated with tumor grade	[^Citation194]
Ovarian cancer	circSETDB1	Up	Associated with advanced clinical stage, lymph node metastasis and increased chemoresistance	[^Citation209]
Prostate cancer	CircITCH	Down	Related with preoperative PSA, tumor stage and Gleason score	[^Citation216]
	circABCC4	Up	Correlated with advanced stage, metastasis and overall survival	[^Citation219]
Myeloid leukemia	Circ-Foxo3	Down	Correlated with overall survival	[^Citation233]
	circ_100053	Up	Associated with clinical stage, BCR/ABL mutant status and imatinib resistance	[^Citation236]

Note: The role of cirNA in clinical relevance and prognosis in different cancers was summarized.

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