# KLF4

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Protein-coding gene in the species Homo sapiens

KLF4 Available structures PDB Ortholog search: PDBe RCSB List of PDB id codes 2WBS, 2WBU, 4M9E Identifiers Aliases KLF4, EZF, GKLF, Kruppel-like factor 4 (gut), Kruppel like factor 4 External IDs OMIM: 602253; MGI: 1342287; HomoloGene: 3123; GeneCards: KLF4; OMA:KLF4 - orthologs Gene location (Human) Chr. Chromosome 9 (human)[1] Band 9q31.2 Start 107,484,852 bp[1] End 107,490,482 bp[1] Gene location (Mouse) Chr. Chromosome 4 (mouse)[2] Band 4 B3|4 29.76 cM Start 55,527,143 bp[2] End 55,532,466 bp[2] RNA expression pattern Bgee Human Mouse (ortholog) Top expressed in skin of thigh skin of hip human penis mucosa of sigmoid colon synovial joint skin of arm skin of abdomen vulva jejunal mucosa mucosa of pharynx Top expressed in epithelium of stomach mucous cell of stomach skin of external ear conjunctival fornix left colon pyloric antrum intestinal villus umbilical cord cervix stria vascularis More reference expression data BioGPS More reference expression data Gene ontology Molecular function DNA binding sequence-specific DNA binding beta-catenin binding phosphatidylinositol 3-kinase regulator activity DNA-binding transcription factor activity zinc ion binding DNA-binding transcription activator activity, RNA polymerase II-specific transcription factor binding cis-regulatory region sequence-specific DNA binding metal ion binding RNA polymerase II sequence-specific DNA-binding transcription factor recruiting activity protein binding nucleic acid binding double-stranded DNA binding promoter-specific chromatin binding RNA polymerase II cis-regulatory region sequence-specific DNA binding DNA-binding transcription factor activity, RNA polymerase II-specific transcription coregulator binding histone deacetylase binding Cellular component cytoplasm transcription regulator complex nucleoplasm chromatin nucleus Biological process negative regulation of chemokine (C-X-C motif) ligand 2 production epidermal cell differentiation cellular response to retinoic acid negative regulation of protein kinase B signaling negative regulation of muscle hyperplasia negative regulation of smooth muscle cell proliferation positive regulation of protein metabolic process negative regulation of cysteine-type endopeptidase activity involved in apoptotic process regulation of axon regeneration cellular response to peptide regulation of transcription, DNA-templated positive regulation of hemoglobin biosynthetic process response to retinoic acid somatic stem cell population maintenance cellular response to laminar fluid shear stress regulation of transcription by RNA polymerase II cell differentiation epidermis morphogenesis positive regulation of nitric oxide biosynthetic process cellular response to growth factor stimulus cellular response to organic cyclic compound post-embryonic hemopoiesis negative regulation of transcription by RNA polymerase II transcription by RNA polymerase II response to organic substance negative regulation of gene expression negative regulation of response to cytokine stimulus transcription, DNA-templated negative regulation of DNA-binding transcription factor activity negative regulation of phosphatidylinositol 3-kinase signaling stem cell population maintenance negative regulation of cell migration involved in sprouting angiogenesis positive regulation of gene expression negative regulation of cell migration regulation of cell population proliferation post-embryonic camera-type eye development regulation of phosphatidylinositol 3-kinase activity positive regulation of telomerase activity canonical Wnt signaling pathway negative regulation of ERK1 and ERK2 cascade regulation of cell differentiation mesodermal cell fate determination negative regulation of heterotypic cell-cell adhesion negative regulation of transcription, DNA-templated negative regulation of NF-kappaB transcription factor activity cellular response to cycloheximide negative regulation of inflammatory response fat cell differentiation positive regulation of transcription by RNA polymerase II negative regulation of cell population proliferation cellular response to hydrogen peroxide negative regulation of leukocyte adhesion to arterial endothelial cell positive regulation of core promoter binding cellular response to leukemia inhibitory factor negative regulation of angiogenesis pri-miRNA transcription by RNA polymerase II negative regulation of G1/S transition of mitotic cell cycle positive regulation of transcription, DNA-templated positive regulation of sprouting angiogenesis Sources:Amigo / QuickGO Orthologs Species Human Mouse Entrez 9314 16600 Ensembl ENSG00000136826 ENSMUSG00000003032 UniProt O43474 Q60793 RefSeq (mRNA) NM_001314052 NM_004235 NM_010637 RefSeq (protein) NP_001300981 NP_004226 NP_034767 Location (UCSC) Chr 9: 107.48 – 107.49 Mb Chr 4: 55.53 – 55.53 Mb PubMed search [3] [4] Wikidata View/Edit Human View/Edit Mouse

**Krüppel-like factor 4** (**KLF4**; gut-enriched Krüppel-like factor or **GKLF**) is a member of the [KLF family](/source/Kruppel-like_factors) of [zinc finger transcription factors](/source/Zinc_finger_transcription_factors), which belongs to the relatively large family of [SP1](/source/Sp1_transcription_factor)-like transcription factors.[5][6][7] KLF4 is involved in the regulation of [proliferation](/source/Cell_proliferation), [differentiation](/source/Cellular_differentiation), [apoptosis](/source/Apoptosis) and [somatic cell](/source/Somatic_cell) reprogramming. Evidence also suggests that KLF4 is a [tumor suppressor](/source/Tumor_suppressor) in certain [cancers](/source/Cancers), including [colorectal cancer](/source/Colorectal_cancer).[8] It has three Cys2His2-[zinc fingers](/source/Zinc_fingers) at its [carboxyl terminus](/source/Carboxyl_terminus) that are closely related to another KLF, [KLF2](/source/KLF2).[6] It has two [nuclear localization sequences](/source/Nuclear_localization_sequences) that signals it to localize to the nucleus.[9] In [embryonic stem cells](/source/Embryonic_stem_cells) (ESCs), KLF4 has been demonstrated to be a good indicator of stem-like capacity. It is suggested that the same is true in [mesenchymal stem cells](/source/Mesenchymal_stem_cells).

In humans, the [protein](/source/Protein) is 513 amino acids, with a predicted molecular weight of approximately 55kDa, and is encoded by the *KLF4* [gene](/source/Gene).[10] The KLF4 gene is conserved in chimpanzee, [rhesus monkey](/source/Rhesus_monkey), dog, cow, mouse, rat, chicken, [zebrafish](/source/Zebrafish), and frog.[11] KLF4 was first identified in 1996.[12]

## Interactions

KLF4 can activate transcription by interacting via its N-terminus with specific transcriptional co-activators, such as [p300-CBP coactivator family](/source/P300-CBP_coactivator_family).[13][14][15] Transcriptional repression by KLF4 is carried out by KLF4 competing with an activator for binding to a target DNA sequence (9-12).[16][17][18][19] KLF4 has been shown to [interact](/source/Protein-protein_interaction) with [CREB-binding protein](/source/CREB-binding_protein).[14] Furthermore, KLF4 has also been documented to play a role in long range chromatin interactions across the genome. KLF4 has been shown to play a role in methylation of chromatin thus contributing to higher order architectural changes across the genome. Specifically, KLF4 has been shown to bind to methylated gene motifs causing switching from repressive to active chromatin markers, and causing downstream enhanced transcription of KLF4 gene targets.[20] KLF4 has also been documented to interact with protein arginine methyltransferase 1 ([Prmt1](/source/PRMT1)), which is involved in [post-translational modifications](/source/Post-translational_modification) in regards to cell fate decisions via methylation of histones. It has been shown that KLF4 is methylated by Prmt1 and the methylation of KLF4 by Prmt1 prevents the emergence of primitive endoderm progenitor cells, thus showing the role in which KLF4 interactions play with cell development.[21]

## Function

KLF4 has diverse functions, and has gained attention because some of its functions are apparently contradictory, but mainly since the discovery of its integral role as one of four key factors that are essential for inducing [pluripotent stem cells](/source/Pluripotent_stem_cell).[22][23] KLF4 is highly expressed in non-dividing cells and its overexpression induces cell cycle arrest.[12][24][25][26][27] KLF4 is particularly important in preventing cell division when the DNA is damaged.[24][26][27][28] KLF4 is also important in regulating centrosome number and chromosome number (genetic stability),[8][29][30] and in promoting cell survival.[31][32][33][34][35][36] However, some studies have revealed that under certain conditions KLF4 may switch its role from pro-cell survival to pro-cell death.[35][37][38][39]

KLF4 is expressed in the cells that are non-dividing and are terminally differentiated in the [intestinal epithelium](/source/Intestinal_epithelium), where KLF4 is important in the regulation of intestinal epithelium homeostasis (terminal cell differentiation and proper localization of the different intestinal epithelium cell types).[40][41][42][43] In the intestinal epithelium, KLF4 is an important regulator of [Wnt signaling pathway](/source/Wnt_signaling_pathway) genes of genes regulating differentiation.[43]

KLF4 is expressed in a variety of tissues and organs such as: the [cornea](/source/Cornea) where it is required for epithelial barrier function[44][45] and is a regulator of genes required for corneal homeostasis;[46] the [skin](/source/Skin) where it is required for the development of skin permeability barrier function;[47][48][49] the [bone](/source/Bone) and teeth tissues where it regulates normal skeletal development;[50][51][52][53] epithelial cell of the mouse male and female reproductive tract[54] where in the males it is important for proper spermatogenesis;[55][56][57] vascular endothelial cells[58] where it is critical in preventing vascular leakage in response to inflammatory stimuli;[59] white blood cells where it mediates inflammatory responses cellular differentiation[60][61][62][63] and proliferation;[63][64] the kidneys where it is involved in the differentiation of embryonic stem cells and induced pluripotent stem (iPS) cells to renal lineage in vitro[65] and its dysregulation has been linked to some renal pathologies.[66][67][68]

## Roles in diseases

Several lines of evidence have shown that KLF4 role in disease is context dependent where under certain conditions it may play one role and under different conditions it may assume a complete opposite role.

KLF4 is an anti-tumorigenic factor and its expression is often lost in various human cancer types, such as [Colorectal cancer](/source/Colorectal_cancer),[69] [gastric cancer](/source/Gastric_cancer),[70] esophageal squamous cell carcinoma,[32] intestinal cancer,[71] [prostate cancer](/source/Prostate_cancer),[72] [bladder cancer](/source/Bladder_cancer)[73] and [lung cancer](/source/Lung_cancer).[74]

However, in some cancer types KLF4 may act as a tumor promoter where increased KLF4 expression has been reported, such as in oral squamous cell carcinoma[75] and in primary breast ductal carcinoma.[76] Also, overexpression of KLF4 in skin resulted in [hyperplasia](/source/Hyperplasia) and [dysplasia](/source/Dysplasia),[77] which lead to the development of squamous cell carcinoma.[78] Similar finding in esophageal epithelium was observed, where overexpression of KLF4 resulted in increased inflammation that eventually lead to the development of esophageal squamous cell cancer in mice.[79]

The role of KLF4 in [Epithelial–mesenchymal transition](/source/Epithelial%E2%80%93mesenchymal_transition) (EMT) is also controversial. It was shown to stimulate EMT in some systems by promoting/maintaining stemness of cancer cells, as is the case in [pancreatic cancer](/source/Pancreatic_cancer),[80][81][82] head and neck cancer,[83] [endometrial cancer](/source/Endometrial_cancer),[84] [nasopharyngeal cancer](/source/Nasopharyngeal_cancer),[85] prostate cancer[86] and non-small lung cancer.[87] Under conditions of TGFβ-induced EMT KLF4 was shown to suppress EMT in the same systems where it was shown to promote EMT, such as prostate cancer[88] and pancreatic cancer.[89] Additionally, KLF4 was shown to suppress EMT in epidermal cancer,[90] breast cancer,[35] lung cancer,[91] cisplatin-resistant nasopharyngeal carcinoma cells,[92] and in hepatocellular carcinoma cells.[*[citation needed](https://en.wikipedia.org/wiki/Wikipedia:Citation_needed)*]

KLF4 plays an important role in several vascular diseases where it was shown to regulate vascular inflammation by controlling macrophage polarization[93] and plaque formation in [atherosclerosis](/source/Atherosclerosis).[94][95][96] It up-regulates [Apolipoprotein E](/source/Apolipoprotein_E), which is an anti-atherosclerotic factor.[95] It is also involved in the regulation of [angiogenesis](/source/Angiogenesis). It may suppress angiogenesis by regulating [NOTCH1](/source/NOTCH1) activity,[97] while in the central nervous system its overexpression leads to vascular dysplasia.[98]

KLF4 may promote inflammation by mediating NF-κB-dependent inflammatory pathway such as in macrophages,[18] esophageal epithelium[79] and in chemically-induced acute colitis in mice.[99] Additionally, KLF-4 downregulates TNF-α-induced VCAM1 expression by targeting and blocking the binding site of NF-κB to the VCAM1 promoter.[100]

However, KLF4 may also suppress the activation of inflammatory signaling such as in endothelial cells in response to pro-inflammatory stimuli.[58]

KLF4 is essential for the cellular response to DNA damage. It is required for preventing cell cycle entry into mitosis following γ-irradiation-induced DNA damage,[26][27] in promoting DNA repair mechanisms (20) and in preventing the irradiated cell from undergoing programmed cell death (apoptosis) (23,25,26).[31][33][34] In one study, the in vivo importance of KLF4 in response to γ-irradiation-induced DNA damage was revealed where deletion of KLF4 specifically from the intestinal epithelium in mice lead to inability of the intestinal epithelium to regenerate and resulting in increased mortality of these mice.[34]

## Importance in stem cells

KLF4 in [post-translational modification](/source/Post-translational_modification) of [stem cells](/source/Stem_cell)

Research has proposed there to be four main factors that have the ability to reprogram differentiated cells back into an embryonic stem cell-like state through the introduction of the Yamanaka factors, Oct4, Sox2, c-Myc, and KLF4.[101] While not as frequently studied as Oct4 and Sox2, KLF4 has become of interest in regards to stem cells due to its complex abilities regarding cell fate and [cell potency](/source/Cell_potency). Specifically, it has been shown how KLF4 alone can affect a cell's level of potency, for instance, rapid KLF4 gene down regulation occurs during both serum and serum free differentiation conditions of embryoid bodies.[102] Furthermore, embryonic stem cells over expressing KLF4 demonstrated reduced differentiation, along with a greater capacity to self renew[103]

Since 2006 up to today, the work on clinically relevant research in stem cells and stem cell induction, has increased dramatically (more than 10,000 research articles, as compared to about 60 between years 1900 to 2005). In vivo functional studies on the role of KLF4 in stem cells are rare. Recently a group investigated the role of KLF4 in a particular population of intestinal stem cells, the Bmi1+ stem cells. This population of intestinal stem cells: are normally slow dividing, are known to be resistant to radiation injury, and are the ones responsible for intestinal epithelium regeneration following radiation injury.[104] The study showed that in the intestine, following γ-irradiation-induced DNA damage, KLF4 may regulate epithelial regeneration by modulating the fate of Bmi1+(([BMI1](/source/BMI1))) stem cells themselves, and consequently the development of Bmi1+ + intestinal stem cell-derived lineage.

## See also

- [Kruppel-like factors](/source/Kruppel-like_factors)

## Notes

The version of this article was updated by an external expert under a dual publication model. The corresponding academic peer reviewed article was published in Gene and can be cited as: Amr M Ghaleb, Vincent W Yang (22 February 2017). "Krüppel-like factor 4 (KLF4): What we currently know". Gene. Gene Wiki Review Series. 611: 27–37. doi:10.1016/J.GENE.2017.02.025. ISSN 0378-1119. PMC 5391259. PMID 28237823. Wikidata Q39151066.

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1. **[^](#cite_ref-98)** Cuttano R, Rudini N, Bravi L, Corada M, Giampietro C, Papa E, et al. (November 2015). ["KLF4 is a key determinant in the development and progression of cerebral cavernous malformations"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4718159). *EMBO Molecular Medicine*. **8** (1): 6–24. [doi](/source/Doi_(identifier)):[10.15252/emmm.201505433](https://doi.org/10.15252%2Femmm.201505433). [PMC](/source/PMC_(identifier)) [4718159](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4718159). [PMID](/source/PMID_(identifier)) [26612856](https://pubmed.ncbi.nlm.nih.gov/26612856).

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## Further reading

- Rowland BD, Peeper DS (January 2006). "KLF4, p21 and context-dependent opposing forces in cancer". *Nature Reviews. Cancer*. **6** (1): 11–23. [doi](/source/Doi_(identifier)):[10.1038/nrc1780](https://doi.org/10.1038%2Fnrc1780). [PMID](/source/PMID_(identifier)) [16372018](https://pubmed.ncbi.nlm.nih.gov/16372018). [S2CID](/source/S2CID_(identifier)) [41981721](https://api.semanticscholar.org/CorpusID:41981721).

- Shields JM, Christy RJ, Yang VW (August 1996). ["Identification and characterization of a gene encoding a gut-enriched Krüppel-like factor expressed during growth arrest"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2330254). *The Journal of Biological Chemistry*. **271** (33): 20009–20017. [doi](/source/Doi_(identifier)):[10.1074/jbc.271.33.20009](https://doi.org/10.1074%2Fjbc.271.33.20009). [PMC](/source/PMC_(identifier)) [2330254](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2330254). [PMID](/source/PMID_(identifier)) [8702718](https://pubmed.ncbi.nlm.nih.gov/8702718).

- Garrett-Sinha LA, Eberspaecher H, Seldin MF, de Crombrugghe B (December 1996). ["A gene for a novel zinc-finger protein expressed in differentiated epithelial cells and transiently in certain mesenchymal cells"](https://doi.org/10.1074%2Fjbc.271.49.31384). *The Journal of Biological Chemistry*. **271** (49): 31384–31390. [doi](/source/Doi_(identifier)):[10.1074/jbc.271.49.31384](https://doi.org/10.1074%2Fjbc.271.49.31384). [PMID](/source/PMID_(identifier)) [8940147](https://pubmed.ncbi.nlm.nih.gov/8940147).

- Yet SF, McA'Nulty MM, Folta SC, Yen HW, Yoshizumi M, Hsieh CM, et al. (January 1998). ["Human EZF, a Krüppel-like zinc finger protein, is expressed in vascular endothelial cells and contains transcriptional activation and repression domains"](https://doi.org/10.1074%2Fjbc.273.2.1026). *The Journal of Biological Chemistry*. **273** (2): 1026–1031. [doi](/source/Doi_(identifier)):[10.1074/jbc.273.2.1026](https://doi.org/10.1074%2Fjbc.273.2.1026). [PMID](/source/PMID_(identifier)) [9422764](https://pubmed.ncbi.nlm.nih.gov/9422764).

- Zhang W, Shields JM, Sogawa K, Fujii-Kuriyama Y, Yang VW (July 1998). ["The gut-enriched Krüppel-like factor suppresses the activity of the CYP1A1 promoter in an Sp1-dependent fashion"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2275057). *The Journal of Biological Chemistry*. **273** (28): 17917–17925. [doi](/source/Doi_(identifier)):[10.1074/jbc.273.28.17917](https://doi.org/10.1074%2Fjbc.273.28.17917). [PMC](/source/PMC_(identifier)) [2275057](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2275057). [PMID](/source/PMID_(identifier)) [9651398](https://pubmed.ncbi.nlm.nih.gov/9651398).

- Foster KW, Ren S, Louro ID, Lobo-Ruppert SM, McKie-Bell P, Grizzle W, et al. (June 1999). "Oncogene expression cloning by retroviral transduction of adenovirus E1A-immortalized rat kidney RK3E cells: transformation of a host with epithelial features by c-MYC and the zinc finger protein GKLF". *Cell Growth & Differentiation*. **10** (6): 423–434. [PMID](/source/PMID_(identifier)) [10392904](https://pubmed.ncbi.nlm.nih.gov/10392904).

- Segre JA, Bauer C, Fuchs E (August 1999). "Klf4 is a transcription factor required for establishing the barrier function of the skin". *Nature Genetics*. **22** (4): 356–360. [doi](/source/Doi_(identifier)):[10.1038/11926](https://doi.org/10.1038%2F11926). [PMID](/source/PMID_(identifier)) [10431239](https://pubmed.ncbi.nlm.nih.gov/10431239). [S2CID](/source/S2CID_(identifier)) [3014700](https://api.semanticscholar.org/CorpusID:3014700).

- Geiman DE, Ton-That H, Johnson JM, Yang VW (March 2000). ["Transactivation and growth suppression by the gut-enriched Krüppel-like factor (Krüppel-like factor 4) are dependent on acidic amino acid residues and protein-protein interaction"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC102607). *Nucleic Acids Research*. **28** (5): 1106–1113. [doi](/source/Doi_(identifier)):[10.1093/nar/28.5.1106](https://doi.org/10.1093%2Fnar%2F28.5.1106). [PMC](/source/PMC_(identifier)) [102607](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC102607). [PMID](/source/PMID_(identifier)) [10666450](https://pubmed.ncbi.nlm.nih.gov/10666450).

- Zhang W, Geiman DE, Shields JM, Dang DT, Mahatan CS, Kaestner KH, et al. (June 2000). ["The gut-enriched Kruppel-like factor (Kruppel-like factor 4) mediates the transactivating effect of p53 on the p21WAF1/Cip1 promoter"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2231805). *The Journal of Biological Chemistry*. **275** (24): 18391–18398. [doi](/source/Doi_(identifier)):[10.1074/jbc.C000062200](https://doi.org/10.1074%2Fjbc.C000062200). [PMC](/source/PMC_(identifier)) [2231805](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2231805). [PMID](/source/PMID_(identifier)) [10749849](https://pubmed.ncbi.nlm.nih.gov/10749849).

- Okano J, Opitz OG, Nakagawa H, Jenkins TD, Friedman SL, Rustgi AK (May 2000). ["The Krüppel-like transcriptional factors Zf9 and GKLF coactivate the human keratin 4 promoter and physically interact"](https://doi.org/10.1016%2FS0014-5793%2800%2901468-X). *FEBS Letters*. **473** (1): 95–100. [Bibcode](/source/Bibcode_(identifier)):[2000FEBSL.473...95O](https://ui.adsabs.harvard.edu/abs/2000FEBSL.473...95O). [doi](/source/Doi_(identifier)):[10.1016/S0014-5793(00)01468-X](https://doi.org/10.1016%2FS0014-5793%2800%2901468-X). [PMID](/source/PMID_(identifier)) [10802067](https://pubmed.ncbi.nlm.nih.gov/10802067). [S2CID](/source/S2CID_(identifier)) [34923598](https://api.semanticscholar.org/CorpusID:34923598).

- Higaki Y, Schullery D, Kawata Y, Shnyreva M, Abrass C, Bomsztyk K (June 2002). ["Synergistic activation of the rat laminin gamma1 chain promoter by the gut-enriched Kruppel-like factor (GKLF/KLF4) and Sp1"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC117209). *Nucleic Acids Research*. **30** (11): 2270–2279. [doi](/source/Doi_(identifier)):[10.1093/nar/30.11.2270](https://doi.org/10.1093%2Fnar%2F30.11.2270). [PMC](/source/PMC_(identifier)) [117209](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC117209). [PMID](/source/PMID_(identifier)) [12034813](https://pubmed.ncbi.nlm.nih.gov/12034813).

- Chen ZY, Shie JL, Tseng CC (November 2002). ["Gut-enriched Kruppel-like factor represses ornithine decarboxylase gene expression and functions as checkpoint regulator in colonic cancer cells"](https://doi.org/10.1074%2Fjbc.M204816200). *The Journal of Biological Chemistry*. **277** (48): 46831–46839. [doi](/source/Doi_(identifier)):[10.1074/jbc.M204816200](https://doi.org/10.1074%2Fjbc.M204816200). [PMID](/source/PMID_(identifier)) [12297499](https://pubmed.ncbi.nlm.nih.gov/12297499).

- Yoon HS, Chen X, Yang VW (January 2003). ["Kruppel-like factor 4 mediates p53-dependent G1/S cell cycle arrest in response to DNA damage"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2229830). *The Journal of Biological Chemistry*. **278** (4): 2101–2105. [doi](/source/Doi_(identifier)):[10.1074/jbc.M211027200](https://doi.org/10.1074%2Fjbc.M211027200). [PMC](/source/PMC_(identifier)) [2229830](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2229830). [PMID](/source/PMID_(identifier)) [12427745](https://pubmed.ncbi.nlm.nih.gov/12427745).

- Wang N, Liu ZH, Ding F, Wang XQ, Zhou CN, Wu M (December 2002). ["Down-regulation of gut-enriched Kruppel-like factor expression in esophageal cancer"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4656400). *World Journal of Gastroenterology*. **8** (6): 966–970. [doi](/source/Doi_(identifier)):[10.3748/wjg.v8.i6.966](https://doi.org/10.3748%2Fwjg.v8.i6.966). [PMC](/source/PMC_(identifier)) [4656400](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4656400). [PMID](/source/PMID_(identifier)) [12439907](https://pubmed.ncbi.nlm.nih.gov/12439907).

- Chen X, Whitney EM, Gao SY, Yang VW (February 2003). ["Transcriptional profiling of Krüppel-like factor 4 reveals a function in cell cycle regulation and epithelial differentiation"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2693487). *Journal of Molecular Biology*. **326** (3): 665–677. [doi](/source/Doi_(identifier)):[10.1016/S0022-2836(02)01449-3](https://doi.org/10.1016%2FS0022-2836%2802%2901449-3). [PMC](/source/PMC_(identifier)) [2693487](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2693487). [PMID](/source/PMID_(identifier)) [12581631](https://pubmed.ncbi.nlm.nih.gov/12581631).

- Dang DT, Chen X, Feng J, Torbenson M, Dang LH, Yang VW (May 2003). ["Overexpression of Krüppel-like factor 4 in the human colon cancer cell line RKO leads to reduced tumorigenecity"](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2275074). *Oncogene*. **22** (22): 3424–3430. [doi](/source/Doi_(identifier)):[10.1038/sj.onc.1206413](https://doi.org/10.1038%2Fsj.onc.1206413). [PMC](/source/PMC_(identifier)) [2275074](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2275074). [PMID](/source/PMID_(identifier)) [12776194](https://pubmed.ncbi.nlm.nih.gov/12776194).

- Mao Z, Song S, Zhu Y, Yi X, Zhang H, Shang Y, et al. (July 2003). ["Transcriptional regulation of A33 antigen expression by gut-enriched Krüppel-like factor"](https://doi.org/10.1038%2Fsj.onc.1206508). *Oncogene*. **22** (28): 4434–4443. [doi](/source/Doi_(identifier)):[10.1038/sj.onc.1206508](https://doi.org/10.1038%2Fsj.onc.1206508). [PMID](/source/PMID_(identifier)) [12853980](https://pubmed.ncbi.nlm.nih.gov/12853980).

- Ohnishi S, Ohnami S, Laub F, Aoki K, Suzuki K, Kanai Y, et al. (August 2003). "Downregulation and growth inhibitory effect of epithelial-type Krüppel-like transcription factor KLF4, but not KLF5, in bladder cancer". *Biochemical and Biophysical Research Communications*. **308** (2): 251–256. [Bibcode](/source/Bibcode_(identifier)):[2003BBRC..308..251O](https://ui.adsabs.harvard.edu/abs/2003BBRC..308..251O). [doi](/source/Doi_(identifier)):[10.1016/S0006-291X(03)01356-1](https://doi.org/10.1016%2FS0006-291X%2803%2901356-1). [PMID](/source/PMID_(identifier)) [12901861](https://pubmed.ncbi.nlm.nih.gov/12901861).

- Hinnebusch BF, Siddique A, Henderson JW, Malo MS, Zhang W, Athaide CP, et al. (January 2004). "Enterocyte differentiation marker intestinal alkaline phosphatase is a target gene of the gut-enriched Kruppel-like factor". *American Journal of Physiology. Gastrointestinal and Liver Physiology*. **286** (1): G23–G30. [doi](/source/Doi_(identifier)):[10.1152/ajpgi.00203.2003](https://doi.org/10.1152%2Fajpgi.00203.2003). [PMID](/source/PMID_(identifier)) [12919939](https://pubmed.ncbi.nlm.nih.gov/12919939).

## External links

- [KLF4+protein,+human](https://meshb.nlm.nih.gov/record/ui?name=KLF4+protein%2C+human) at the U.S. National Library of Medicine [Medical Subject Headings](/source/Medical_Subject_Headings) (MeSH)

- [KLF4 microarray expression results and literature](http://www.nextbio.com/b/home/home.nb?q=KLF4&id=34891&type=feature&name=KLF4&synonym=)

v t e Transcription factors and intracellular receptors (1) Basic domains (1.1) Basic leucine zipper (bZIP) Activating transcription factor AATF 1 2 3 4 5 6 7 AP-1 c-Fos FOSB FOSL1 FOSL2 JDP2 c-Jun JUNB JunD BACH 1 2 BATF BLZF1 C/EBP α β γ δ ε ζ CREB 1 3 L1 CREM DBP DDIT3 GABPA GCN4 HLF MAF B sMaf F G K NFE 2 L1 L2 L3 NFIL3 NRL NRF 1 2 3 XBP1 (1.2) Basic helix–loop–helix (bHLH) Group A AS-C ASCL1 ASCL2 ATOH1 HAND 1 2 MESP2 Myogenic regulatory factors MyoD Myogenin MYF5 MYF6 NeuroD 1 2 Neurogenins 1 2 3 OLIG 1 2 Paraxis TCF15 Scleraxis SLC LYL1 TAL 1 2 Twist Group B FIGLA Myc c-Myc l-Myc n-Myc MXD4 TCF4 Group C bHLH-PAS AhR AHRR ARNT ARNTL ARNTL2 CLOCK HIF 1A EPAS1 3A NPAS 1 2 3 PER 1 2 3 Period SIM 1 2 Group D DEC 1 2 BHLHA9 Pho4 ID 1 2 3 4 Group E HES 1 2 3 4 5 6 7 HEY 1 2 L Group F bHLH-COE EBF1 (1.3) bHLH-ZIP AP-4 MAX MXD1 MXD3 MITF MNT MLX MLXIPL MXI1 Myc SREBP 1 2 USF1 (1.4) NF-1 NFI A B C X SMAD R-SMAD 1 2 3 5 9 I-SMAD 6 7 4) (1.5) RF-X RFX 1 2 3 4 5 6 ANK (1.6) Basic helix-span-helix (bHSH) AP-2 α β γ δ ε (2) Zinc finger DNA-binding domains (2.1) Nuclear receptor (Cys4) subfamily 1 Thyroid hormone α β CAR FXR LXR α β PPAR α β/δ γ PXR RAR α β γ ROR α β γ Rev-ErbA α β VDR subfamily 2 COUP-TF (I II Ear-2 HNF4 α γ PNR RXR α β γ Testicular receptor 2 4 TLX subfamily 3 Steroid hormone Androgen Estrogen α β Glucocorticoid Mineralocorticoid Progesterone Estrogen related α β γ subfamily 4 NUR NGFIB NOR1 NURR1 subfamily 5 LRH-1 SF1 subfamily 6 GCNF subfamily 0 DAX1 SHP (2.2) Other Cys4 GATA 1 2 3 4 5 6 MTA 1 2 3 TRPS1 (2.3) Cys2His2 General transcription factors TFIIA TFIIB TFIID TFIIE 1 2 TFIIF 1 2 TFIIH 1 2 4 2I 3A 3C1 3C2 ATBF1 BCL 6 11A 11B CTCF E4F1 EGR 1 2 3 4 ERV3 GFI1 GLI family 1 2 3 REST S1 S2 YY1 HIC 1 2 HIVEP 1 2 3 IKZF 1 2 3 ILF 2 3 Sp/KLF family KLF 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 17 SP 1 2 4 7 8 MTF1 MYT1 OSR1 PRDM9 SALL 1 2 3 4 TSHZ3 WT1 Zbtb7 7A 7B ZBTB 11 16 17 20 21 32 33 40 zinc finger 3 7 9 10 19 22 24 33B 34 35 41 43 44 51 74 143 146 148 165 202 217 219 238 239 259 267 268 281 300 318 330 346 350 365 366 384 423 451 452 471 593 638 644 649 655 804A (2.4) Cys6 HIVEP1 (2.5) Alternating composition AIRE DIDO1 GRLF1 ING 1 2 4 JARID 1A 1B 1C 1D 2 JMJD1B (2.6) WRKY WRKY (3) Helix-turn-helix domains (3.1) Homeodomain Antennapedia ANTP class protoHOX Hox-like ParaHox Gsx 1 2 Xlox PDX1 Cdx 1 2 4 extended Hox: Evx1 Evx2 MEOX1 MEOX2 Homeobox A1 A2 A3 A4 A5 A7 A9 A10 A11 A13 B1 B2 B3 B4 B5 B6 B7 B8 B9 B13 C4 C5 C6 C8 C9 C10 C11 C12 C13 D1 D3 D4 D8 D9 D10 D11 D12 D13 GBX1 GBX2 MNX1 metaHOX NK-like BARHL1 BARHL2 BARX1 BARX2 BSX DBX 1 2 DLX 1 2 3 4 5 6 EMX 1 2 EN 1 2 HHEX HLX LBX1 LBX2 MSX 1 2 NANOG NKX 2-1 2-2 2-3 2-5 3-1 3-2 HMX1 HMX2 HMX3 6-1 6-2 NOTO TLX1 TLX2 TLX3 VAX1 VAX2 other ARX CRX CUTL1 FHL 1 2 3 HESX1 HOPX LMX 1A 1B NOBOX TALE IRX 1 2 3 4 5 6 MKX MEIS 1 2 PBX 1 2 3 PKNOX 1 2 SIX 1 2 3 4 5 PHF 1 3 6 8 10 16 17 20 21A POU domain PIT-1 BRN-3: A B C Octamer transcription factor: 1 2 3/4 6 7 11 SATB2 ZEB 1 2 (3.2) Paired box PAX 1 2 3 4 5 6 7 8 9 PRRX 1 2 PROP1 PHOX 2A 2B RAX SHOX SHOX2 VSX1 VSX2 Bicoid GSC BICD2 OTX 1 2 PITX 1 2 3 (3.3) Fork head / winged helix E2F 1 2 3 4 5 FOX proteins A1 A2 A3 B1 B2 C1 C2 D1 D2 D3 D4 D4L1 D4L3 D4L4 D4L5 D4L6 E1 E3 F1 F2 G1 H1 I1 I2 I3 J1 J2 J3 K1 K2 L1 L2 M1 N1 N2 N3 N4 O1 O3 O4 O6 P1 P2 P3 P4 Q1 R1 R2 S1 (3.4) Heat shock factors HSF 1 2 4 (3.5) Tryptophan clusters ELF 2 4 5 EHF ELK 1 3 4 ERF ETS 1 2 ERG SPIB ETV 1 4 5 6 FLI1 Interferon regulatory factors 1 2 3 4 5 6 7 8 MYB MYBL2 (3.6) TEA domain transcriptional enhancer factor 1 2 3 4 (4) β-Scaffold factors with minor groove contacts (4.1) Rel homology region NF-κB NFKB1 NFKB2 REL RELA RELB NFAT C1 C2 C3 C4 5 (4.2) STAT STAT 1 2 3 4 5 6 (4.3) p53-like p53 p63 p73 family p53 TP63 p73 TBX 1 2 3 5 19 21 22 TBR1 TBR2 TFT MYRF (4.4) MADS box Mef2 A B C D SRF (4.6) TATA-binding proteins TBP TBPL1 (4.7) High-mobility group BBX HMGB 1 2 3 4 HMGN 1 2 3 4 HNF 1A 1B SOX 1 2 3 4 5 6 8 9 10 11 12 13 14 15 17 18 21 SRY SSRP1 TCF/LEF TCF 1 3 4 LEF1 TOX 1 2 3 4 (4.9) Grainyhead TFCP2 (4.10) Cold-shock domain CSDA YBX1 (4.11) Runt CBF CBFA2T2 CBFA2T3 RUNX1 RUNX2 RUNX3 RUNX1T1 (0) Other transcription factors (0.2) HMGI(Y) HMGA 1 2 HBP1 (0.3) Pocket domain Rb RBL1 RBL2 (0.5) AP-2/EREBP-related factors Apetala 2 EREBP B3 (0.6) Miscellaneous ARID 1A 1B 2 3A 3B 4A CAP IFI 16 35 MLL 2 3 T1 MNDA NFY A B C Rho/Sigma see also transcription factor/coregulator deficiencies

*This article incorporates text from the [United States National Library of Medicine](/source/United_States_National_Library_of_Medicine), which is in the [public domain](/source/Public_domain).*

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Adapted from the Wikipedia article [KLF4](https://en.wikipedia.org/wiki/KLF4) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/KLF4?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.