The AP-2 family of transcription factors: critical regulators of human development and cancer
Yi-Liu Yang1, Lin Yong Zhao2*
1West China School of Medicine, West China Hospital, Sichuan University, Chengdu, Chiny
2Gastrointestinal kirurgisk klinik og gastrisk kræftlaboratorium, National Key Biotherapy Laboratory, West China Hospital, Sichuan University and Biotherapy Innovation Center, Chengdu, Kina
The AP-2 family of transcription factors consists of DNA-binding proteins: AP-2α to AP-2ε. Members and homologues of this family are also known in frogs, fish and invertebrates. These proteins share the same central core region and the helix-span-helix dimerization motif required for DNA dimerization and binding. This family has been shown to influence the development of the face, limbs and kidneys in embryogenesis, while regulating differentiation and apoptosis. These proteins are also involved in the regulation of endocrine processes. In addition to their effects on growth and development, this family has been reported to correlate with carcinogenesis and cancer development. Currently, these families are related to ovarian, melanoma, lung, nasopharyngeal, breast, glioblastoma, neuroblastoma, colon, etc. They regulate the expression of many cancer-related genes and influence the occurrence, development, invasiveness and therapeutic response of cancers. Different levels of AP-2 expression are also associated with different survival rates. These results could bring a new idea to the diagnosis, classification, treatment and prognosis of cancer.
The AP-2 family of transcription factors (AP-2 family) includes five DNA-binding proteins: AP-2α to AP-2ε, encoded byTFAP2ADoTFAP2Eappropriate. Frogs, fish and invertebrates also have members or homologues of this family. With the exception of AP-2δ, the other four in the family are encoded by seven exons and share the same central core region and a helix-span-helix dimerization motif required for DNA dimerization and binding1. In addition to highly similar DNA-binding and dimerizing domains with other family members, AP-2δ has a unique sequence specificity not observed in the other four proteins. This may be useful for regulating target gene activation2.
The AP-2 family plays an important role in the regulation of cell differentiation and apoptosis. Found to affect body development in embryogenesis, including the formation of the face, limbs, kidneys, retina, central nervous system, and heart3,4. Mutations or defectsTFAP2AITFAP2Blead to development errors5,6. These genes are also involved in the regulation of endocrine processes. E.g,TFAP2Bassociated with insulin resistance and diabetes7, one secondTFAP2Cplays a key role in the regulation of estrogen signaling genes8.
In addition to their effects on cell fate and development, the AP-2 family has also been reported to be involved in tumorigenesis and cancer development. So far, this family has been found to be closely related to various cancers, including ovarian cancer, melanoma, lung cancer, nasopharyngeal cancer, breast cancer, glioblastoma, neuroblastoma, gastric cancer, colon cancer, etc.9-17. They regulate the expression of many cancer-related genes, especially in breast cancer. In addition, they coordinate with the occurrence, development, invasiveness and therapeutic response of cancers. Different levels of AP-2 expression are also associated with different survival rates.
AP-2α encoded byTFAP2Aregulates cell growth and tissue differentiation. Its expression has been observed in epithelial and neural crest cell lines at the early stage of mouse embryogenesis3. MutationsTFAP2AIt has been shown to cause Branchio-oculo-facial syndrome (BOFS), a rare orofacial cleft syndrome that includes skin, ocular, renal and ectodermal abnormalities along with a characteristic facial appearance5.
Different levels of AP-2a expression in tumor cells have also been reported.
AP-2α overexpression has been found in ovarian, nasopharyngeal, and lung cancers, and increased expression may promote tumorigenesis and lead to worse cancer outcomes. In epithelial cells of a normal ovary, AP-2α is expressed only in the cytoplasm. However, in malignant epithelial ovarian tumors, AP-2α is expressed both in the nucleus and in the cytoplasm. The level of AP-2α expression in the testis is associated with an increased risk of death9. High expression of AP-2α has also been reported in nasopharyngeal carcinoma cells, which promotes tumor growth, while downregulation of AP-2α suppresses cell viability and inhibits tumor growth along with microvascular density14. Increased expression levels are also found in lung cancer, which is strongly associated with poor prognosis12. This result is consistent with another study in whichTFAP2Aupregulates expressionKRT16, an independent prognostic predictor associated with poor survival in lung cancer18.
However, in some other cancers, AP-2α expression is reduced, which is associated with tumor progression. For example, suppressed AP-2a expression in breast cancer appears to be more common in invasive breast tumors than in ductal carcinoma in situ.19. Similarly, AP-2a expression inversely correlates with glioblastoma staging, which may suggest its direct role in glioblastoma carcinogenicity.11. The association between decreased AP-2α expression and increased carcinogenicity is also seen in colon cancer cells17and gastric adenocarcinoma20.
AP-2β plays a key role in the development of the ductus arteriosus and the shaping of the limbs21.TFAP2Bmutation leads to nonsyndromic patent ductus arteriosus and Char's syndrome, which is characterized by patent ductus arteriosus, facial dysmorphia, and fifth finger abnormalities6,22. In addition, AP-2β is also involved in glucose and fat metabolism. Gene variantsTFAP2Bis associated with insulin resistance and type 2 diabetes7. It has also been reported to affect conditions related to obesity and intrauterine growth23.
Similar to AP-2α, decreased or increasedTFAP2Bexpression has also been described in human cancers. Increased expression of AP-2β has been reported in lobular carcinoma in situ (LCIS) and invasive lobular carcinoma of the breast24. Furthermore, AP-2β overexpression is associated with poor prognosis in lung adenocarcinoma25and papillary thyroid cancer26while reduced expression appears to be correlated with poor prognosis and poor outcome in neuroblastoma15and endometrial cancer27.
Research suggests a link between AP-2γ and lung cancer. AP-2γ acts as an oncogenic factor that promotes lung cancer28and plays a key role in the development of lung cancer29. In addition, increased expression of AP-2γ is detected in lung cancer cells12which inhibit expressionGADD45BIPMAIP1, then promotes cell proliferation and motility in non-small cell lung cancer30.
Interestingly, AP-2γ has been reported to have the opposite effect in breast cancer: it first delays tumor initiation but then promotes tumor progression13.TFAP2CIt has been shown to influence the development of the luminal cell type during mammary development and to act as a critical transcriptional regulator that maintains the luminal phenotype31.TFAP2Cregulates the expression of many genes in breast cancer.TFAP2Cit was first reported to induce high expressionERBB2(Her2)IESR1 (ERA)which affects the hormonal response in breast cancer cells8. Furthermore, research suggestsTFAP2Ccoordinates the expression of some other major target genes, includingFOXA1, WWOX, GREB1, CDH2, HPSE, IGSF11itp32,33. Expressive levelTFAP2Cit also coordinates with the treatment response and survival rate of breast cancer patients. High expressionTFAP2Cwas reported for suppressionCD44, a gene associated with breast cancer, and leads to a higher pathologic complete response rate after neoadjuvant chemotherapy34. But other research suggests overexpressionTFAP2Cis associated with a shorter survival time of more than 10 years from diagnosis35.
Research on AP-2δ iTFAP2Dthey are relatively rare. Compared to other members of the AP-2 family, AP-2δ appears to affect mammalian development in a different way. AP-2δ expression has been reported in the retina, central nervous system, and developing heart, while the neural crest, facial mesenchyme, and limbs show almost no expression4. AP-2δ expression in ganglion cells promotes fine-tuning of axonal growth in the developing retina36. Additionally, loss of AP-2δ correlates with reduced axonal projections to the superior colliculus37.
AP-2δ has also been found in prostate tissue38. In addition, AP-2δ upregulation is demonstrated in the aggressive tumor phenotype of prostate cancer39.
Many studies of AP-2ε focus on its predictive value regarding patients' response to chemotherapy and treatment outcome for colorectal cancer. However, this issue is still controversial. Elbert's study indicates hypermethylationTFAP2Ecorrelates with resistance to chemotherapy in colorectal cancer40while other studies show that response to chemotherapy cannot be predicted by methylation levels41. Regarding the value of predicting prognosis, there is also no consensus. Some studies show that hypermethylation is associated with a survival advantage42an association with poorer overall and disease-free survival was also noted43.
In addition, AP-2ε is also associated with human neuroblastoma. It has been reported that AP-2ε is involved in the regulation of the DNA damage response in neuroblastoma cells44.
The AP-2 family of transcription factors play indispensable roles in embryogenesis, body formation, and development.TFAP2AITFAP2Bthey also participate in the regulation of endocrine processes. Furthermore, the family has been shown to be involved in tumorigenesis, development and prognosis of various human cancers. And now they are still being studied extensively for cancer in humans. Considering the influence of the AP-2 family on tumor type, therapeutic response and prognosis, especially as shown in breast and colon cancer, these studies may bring new ideas to the diagnosis, classification, treatment and prognosis of cancer. Meanwhile, there are still many controversial issues, including the predictive value of AP-2ε on patients' response to chemotherapy and colorectal cancer outcomes, to be investigated.
This work was supported by the Institute of Science and Technology of Sichuan Province, No. 2021YFS0111. The views expressed are those of the authors and are not necessarily those of the Department of Science and Technology of Sichuan Province. We apologize for the inability to cite all publications related to this topic due to journal space limitations.
TFAP: proteins that activate transcription factors.
KRT16: keratingen 16, type I cytokeratin
GADD45B: growth arrest gene and DNA damage induced beta
PMAIP1: phorbol 12-myristate-13-acetate-induced protein gene 1
ERBB2: V-Erb-B2 avian erythroblastic leukemia viral oncogene homolog 2
Co2: human epidermal growth factor-2 receptor gene
ESR1: estrogen receptor gene 1
Era: estrogen receptor α gene
FOXA1:forkhead box A1 proteingen
WWOX:an oxidoreductase gene containing a WW domain
GRIP1:estrogen growth regulator gene in breast cancer 1
IGSF11:gen for immunoglobulinsuperfamiliemedlem 11
CD44:differentiation cluster gen 44
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The AP-2 family of transcription factors consists of five different proteins in humans and mice: AP-2alpha, AP-2beta, AP-2gamma, AP-2delta and AP-2epsilon. Frogs and fish have known orthologs of some but not all of these proteins, and homologs of the family are also found in protochordates, insects and nematodes.What is the AP2 domain transcription factor? ›
The AP2 DNA binding domain characterizes the large AP2/ERF family of transcription factors in plants. In contrast with other important DNA binding domains, such as basic/helix-loop-helix, MYB, and homeodomain, which are conserved in many branches of the tree of life, the AP2 domain has been considered plant specific.Is AP2 a transcription factor? ›
AP-2a is an important transcription factor of DEK expression, which is correlated with the methylation level of the DEK core promoter in hepatocellular carcinoma . Hepatitis B virus X protein is able to elevate the expression of SPHK1 in hepatoma cells by upregulating transcription factor AP2 alpha.What are the families of transcription factors? ›
Transcription factors constitute a large functional family of proteins directly regulating the activity of genes. Most of them are sequence-specific DNA-binding proteins, thus reading out the information encoded in cis-regulatory DNA elements of promoters, enhancers and other regulatory regions of a genome.What is the AP 2 gene called? ›
Normal Function. The TFAP2A gene provides instructions for making a protein called transcription factor AP-2 alpha (AP-2α). As its name suggests, this protein is a transcription factor, which means it attaches (binds) to specific regions of DNA and helps control the activity of particular genes.What does transcription factor 2 do? ›
Transcription factor TFIIB is an essential component of the RNA polymerase II initiation complex. TFIIB carries out at least two functions: it interacts directly with the TATA-binding protein (TBP) and helps to recruit RNA polymerase II into the initiation complex.What is transcription factor like 2? ›
Transcription factor 7-like 2 (TCF7L2) is the main susceptibility gene for type 2 diabetes, predominantly by affecting insulin secretion of pancreatic β cells.What two domains are most important for transcription factor function? ›
- DNA-binding domain (DBD), which attaches to specific sequences of DNA (enhancer or promoter. ...
- Activation domain (AD), which contains binding sites for other proteins such as transcription coregulators.
Activator protein 1 (AP-1) is a transcription factor that regulates gene expression in response to a variety of stimuli, including cytokines, growth factors, stress, and bacterial and viral infections. AP-1 controls a number of cellular processes including differentiation, proliferation, and apoptosis.What are the subunits of AP2? ›
AP2 comprises two large subunits of 100 kDa (α- and β2-adaptin), a medium 50 kDa subunit (μ2) and a small 17 kDa subunit (σ2). The 100 kDa α- and β2-adaptins comprise an N-terminal head domain linked by a flexible hinge to 'ear' or 'appendage' domains.
JUN Gene - Jun Proto-Oncogene, AP-1 Transcription Factor Subunit.What is AP genetic? ›
Activator protein 1 (AP-1) is a heterodimer typically consisting of the oncogenic transcription factors Jun (often called c-Jun) and Fos (c-Fos) but can also include other less common AP-1 subfamily members. AP-1 binds DNA at conserved response element sequences via a leucine zipper domain (Abate and Curran, 1990).Why are transcription factors important? ›
Transcription factors are vital molecules in the control of gene expression, directly controling when, where and the degree to which genes are expressed. They bind to specific sequences of DNA and control the transcription of DNA into mRNA.What are the 4 classes of transcription factors? ›
TFClass is a classification of eukaryotic transcription factors based on the characteristics of their DNA-binding domains. It comprises four general levels (superclass, class, family, subfamily) and two levels of instantiation (genus and molecular species). Two of them (subfamily and factor species) are optional.How do transcription factors affect gene expression? ›
Transcription factors are proteins that help turn specific genes "on" or "off" by binding to nearby DNA. Transcription factors that are activators boost a gene's transcription. Repressors decrease transcription.What gene is mutated in aps? ›
APS-1 is caused by biallelic mutations of the autoimmune regulator (AIRE) gene located on chromosome 21q22.What is APC gene mutation? ›
In 1997, a Johns Hopkins research team found an inherited genetic mutation called APC I1307K. Adenomatous polyposis coli (APC) is a gene that suppresses tumor growth. If the APC gene is defective, it makes the gene unstable and more susceptible to additional changes that may lead to colon and rectal cancers.Where is the RP2 gene located? ›
RP2 Q158P located in the TBCC domain and destabilized RP2 protein in ARPE-19 cells. four frameshift mutations including three novel mutations of c.What are the 2 most important transcription factors in the inflammatory response? ›
The transcription factors about which there is most information in immune and inflammatory responses are nuclear factor-kappa (κ)B (NF-κB) and activator protein 1 (AP-1).What is the role of RNA polymerase 2 in the process of transcription in eukaryotes? ›
RNA polymerase II (Pol II) is an essential, multi-subunit, DNA-dependent, nucleotidyltransferase. In eukaryotes, Pol II is the one of three nuclear RNA polymerases. It is responsible for unspooling the genetic program in the form of protein-coding mRNAs and some small non-coding RNAs.
transcription factor / transcription factors
Transcription factors are proteins involved in the process of converting, or transcribing, DNA into RNA. Transcription factors include a wide number of proteins, excluding RNA polymerase, that initiate and regulate the transcription of genes.
transcription factor, molecule that controls the activity of a gene by determining whether the gene's DNA (deoxyribonucleic acid) is transcribed into RNA (ribonucleic acid). The enzyme RNA polymerase catalyzes the chemical reactions that synthesize RNA, using the gene's DNA as a template.How do transcription factors work? ›
Under the effect of transcription factors, the various cells of the body can function differently though they have the same genome. Transcription factors bind to one or more sequence sites, which are called transcription factor binding sites (TFBSs), attaching to specific DNA sequences of the genes they regulate .What are the 2 general types of gene specific transcription factors? ›
Distinguishing between General, Specific & Regulatory Transcription Factors. General transcription factors are protein-based factors that bind to the DNA at regions called promoters; on the other hand, specific transcription factors involve DNA sequences called enhancers or promoters.What do regulatory transcription factors do? ›
Transcription factors (TFs) are regulatory proteins whose function is to activate (or more rarely, to inhibit) transcription of DNA by binding to specific DNA sequences. TFs have defined DNA-binding domains with up to 106-fold higher affinity for their target sequences than for the remainder of the DNA strand.What transcription factors are important in development? ›
Zf-C2H2, HMG, and MYB Are Three Dominant Transcription Factor Families in Early Embryo Development.What is AP2 in endocytosis? ›
The AP2 adaptor complex is a multimeric protein that works on the cell membrane to internalize cargo in clathrin-mediated endocytosis. It is a stable complex of four adaptins which give rise to a structure that has a core domain and two appendage domains attached to the core domain by polypeptide linkers.What is made of two subunits and are composed of RNA and protein? ›
A ribosome is an intercellular structure made of both RNA and protein, and it is the site of protein synthesis in the cell.What are two subunits and are composed of RNA and protein? ›
The ribosome is a very large and complex structure, composed of two-thirds RNA and one-third protein. The determination, in 2000, of the entire three-dimensional structure of its large and small subunits is a major triumph of modern structural biology.Is BRCA1 a transcription factor? ›
BRCA1 is known to be involved in transcriptional regulation in that it acts in concert with the RNA Pol II holoenzyme, transcription factors, acetylases, and deacetylases and their associated proteins CBPp300 and CtIP (reviewed in ref. 17). Overexpression of BRCA1 induces genes in the apoptotic pathway (18, 19).
The E2F transcription factor family is known to play a key role in the timely expression of genes required for cell cycle progression and proliferation, but only a few E2F target genes have been identified.What is AP-1 and NFKB? ›
Nuclear factor κB (NF-κB) and activator protein 1 (AP-1) are key transcription factors that orchestrate expression of many genes involved in inflammation, embryonic development, lymphoid differentiation, oncogenesis, and apoptosis (48, 62).What does AP mean in biology? ›
The advanced-placement (AP) biology course sponsored by the College Entrance Examination Board (College Board) is a national program that provides an opportunity for high-school students to pursue and receive credit for college-level biology coursework.What chromosome is APS on? ›
Human prostate-specific antigen (APS) is a member of the glandular kallikrein gene family at 19q13. Cytogenet Cell Genet.What does AP stand for in biology? ›
Advanced Placement (AP) Biology (also known as AP Bio) is an Advanced Placement biology course and exam offered by the College Board in the United States.What is the transcription factor AP-1? ›
AP-1 (activating protein-1) is a collective term referring to dimeric transcription factors composed of Jun, Fos or ATF (activating transcription factor) subunits that bind to a common DNA site, the AP-1-binding site.What is AP in genetics? ›
Activator protein 1 (AP-1) is a heterodimer typically consisting of the oncogenic transcription factors Jun (often called c-Jun) and Fos (c-Fos) but can also include other less common AP-1 subfamily members.What does AP-1 transcription factor do? ›
Activator protein 1 (AP-1) is a transcription factor that regulates gene expression in response to a variety of stimuli, including cytokines, growth factors, stress, and bacterial and viral infections. AP-1 controls a number of cellular processes including differentiation, proliferation, and apoptosis.