Upstream stimulatory factor gene transcriptions: Difference between revisions

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"The helix-loop-helix transcription factor USF (upstream stimulating factor) binds to a regulatory sequence of the human insulin gene enhancer."<ref name=Read/>
"The regulation of insulin gene expression is dependent on sequences located upstream of the transcription start site (Clark and Docherty, 1992). Two important ''cis''-acting elements, the insulin enhancer binding site 1 (IEBI) or NIR box and the IEB2 or FAR box, have been identified in the rat insulin I gene (Karlsson ''et al.'', 1987, 1989). Located at positions -104 (IEBI/NIR) and -233 (IEB2/FAR), these elements share an identical 8 bp sequence, GCCATCTG, which contains a consensus sequence, CANNTG, characteristic of E-box elements (Kingston, 1989). E boxes are present in enhancers from a variety of genes, including immunoglobulin and muscle-specific genes, where they interact with transcription factors containing a helix-loop-helix (HLH) dimerization domain (Murre ''et al.'', 1989)."<ref name=Read>{{ cite journal
|author=Martin L. Read, Andrew R. Clark and Kevin Docherty
|title=The helix-loop-helix transcription factor USF (upstream stimulating factor) binds to a regulatory sequence of the human insulin gene enhancer
|journal=Biochemical Journal
|date=1993
|volume=295
|issue=
|pages=233-237
|url=https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1134844/pdf/biochemj00102-0234.pdf
|arxiv=
|bibcode=
|doi=
|pmid=
|accessdate=14 August 2020 }}</ref>
"The IEB1 box is highly conserved among insulin genes, and is thus likely to play an important role in controlling transcription. The IEB2 site is not well conserved; in the rat insulin 2 gene the equivalent sequence is GCCACCCAGGAG, and in the human insulin gene the homologous sequence, which has been previously designated the GC2 box (Boam ''et al.'', 1990a), is GCCACCGG."<ref name=Read/>
"Confirmation that USF bound at the IEB2 site was obtained using an oligonucleotide containing the USF binding site from the adenovirus MLP."<ref name=Read/>


==Consensus sequences==
==Consensus sequences==

Revision as of 20:03, 20 August 2020

Associate Editor(s)-in-Chief: Henry A. Hoff

"A major environmental stress encountered by humans is solar UV light, which can cause skin inflammation, the induction of pro-inflammatory cytokines and skin ageing, as well as skin cancer, including the highly aggressive and increasingly common malignant melanoma (Elwood, 1996; Armstrong et al., 1997; Park and Gilchrest, 1999). The serious adverse effect of UV light on the skin has meant that humans have evolved an effective defence mechanism. In response to low levels of UV irradiation epidermal melanocytes increase the production of the pigment melanin in specialized organelles termed melanosomes (Jimbow et al., 1991). The melanosomes are transferred into surrounding keratinocytes where they act to protect against UV-induced DNA damage."[1]

"The Tyrosinase gene encodes the rate-limiting enzyme for the production of melanin and is absolutely required for pigmentation; the absence of a functional tyrosinase enzyme results in an albino phenotype. Although much of the tanning response comprises a post-translational activation of the melanosome, transcription of the Tyrosinase gene is UV responsive (Hara et al., 1994; Sturm et al., 1994; Imokawa et al., 1995, 1997; Ota et al., 1998). However, analysis of the Tyrosinase promoter (Bentley et al., 1994; Ganss et al., 1994) failed to reveal any classical UV or stress-response element. The human Tyrosinase promoter comprises an SP1 site and two E box motifs, one at the initiator, and a second, termed the M box (Lowings et al., 1992), located at –100 with respect to the transcription initiation site (Bentley et al., 1994). The E box motifs are essential for Tyrosinase promoter activity and are highly evolutionarily conserved. Numerous studies (Bentley et al., 1994; Ganss et al., 1994; Hemesath et al., 1994; Yasumoto et al., 1994; Yavuzer et al., 1995; Krylov et al., 1997) have demonstrated that the Tyrosinase initiator E box and M box elements are targets for the microphthalmia-associated basic helix–loop–helix-leucine zipper (bHLH-LZ) transcription factor Mitf (Hodgkinson et al., 1993; Hughes et al., 1993). In addition to its role in regulating pigmentation genes, [the microphthalmia-associated transcription factor] Mitf is also critically required for the development of the melanocyte (Steingrímsson et al., 1994; Opdecamp et al., 1997)."[1]

The "UV response is mediated by the ubiquitous bHLH-LZ transcription factor [upstream stimulatory factor] Usf-1, which, like Mitf, binds the conserved E box elements in the Tyrosinase promoter. The ability of Usf-1 to activate transcription is regulated by a signal transduction pathway that culminates in phosphorylation and activation of Usf-1 by the p38 stress-activated kinase."[1]

"Usf-1 and Usf-2 bind the Tyrosinase promoter in vivo."[1]

"USF comprises a combination of related ubiquitous bHLH-LZ transcription factors encoded by the Usf-1 and Usf-2 genes (Gregor et al., 1990; Sirito et al., 1992, 1994). Usf-1 and Usf-2 can form either homo- or heterodimers (Sirito et al., 1992; Viollet et al., 1996), and both are present in melanocytes and melanoma cell lines [...]."[1]

Upstream stimulating or stimulatory factors

The "expressions of [Homeobox transcript antisense intergenic RNA] HOTAIR and upstream stimulatory factor 1 (USF1) was up-regulated, but miR-148b-3p was down-regulated in glioma microvascular endothelial cells (GECs)."[2]

"Upstream stimulating factor (USF) includes upstream stimulating factor 1 (USF1) and upstream stimulating factor 2 (USF2), which belongs to the basic helix-loop-helix leucine zipper family of transcription factors. As an important regulatory factor, USF has highly conserved bHLH-LZ domain and binds to consensus sequence (CANNTG) of E-box to further regulate the transcription process of different proteins (Wu et al., 2013; Lupp et al., 2014). USF1 was reported to regulate the expression of IL-10 in glioma related microglia: inhibition of USF1 expression resulted in the up-regulation of IL-10 expression (Zhang et al., 2007). In liver cancer HepG2 cells, USF1 resisted oxygen sugar deprivation induced apoptosis by regulating miR-132 (Wang et al., 2014)."[2]

"The helix-loop-helix transcription factor USF (upstream stimulating factor) binds to a regulatory sequence of the human insulin gene enhancer."[3]

"The regulation of insulin gene expression is dependent on sequences located upstream of the transcription start site (Clark and Docherty, 1992). Two important cis-acting elements, the insulin enhancer binding site 1 (IEBI) or NIR box and the IEB2 or FAR box, have been identified in the rat insulin I gene (Karlsson et al., 1987, 1989). Located at positions -104 (IEBI/NIR) and -233 (IEB2/FAR), these elements share an identical 8 bp sequence, GCCATCTG, which contains a consensus sequence, CANNTG, characteristic of E-box elements (Kingston, 1989). E boxes are present in enhancers from a variety of genes, including immunoglobulin and muscle-specific genes, where they interact with transcription factors containing a helix-loop-helix (HLH) dimerization domain (Murre et al., 1989)."[3]

"The IEB1 box is highly conserved among insulin genes, and is thus likely to play an important role in controlling transcription. The IEB2 site is not well conserved; in the rat insulin 2 gene the equivalent sequence is GCCACCCAGGAG, and in the human insulin gene the homologous sequence, which has been previously designated the GC2 box (Boam et al., 1990a), is GCCACCGG."[3]

"Confirmation that USF bound at the IEB2 site was obtained using an oligonucleotide containing the USF binding site from the adenovirus MLP."[3]

Consensus sequences

A likely general USF box consensus sequence may be 3'-GCC(A/T)NN(C/G/T)(A/G)-5'.

Human genes

Gene ID: 7391 is USF1 upstream transcription factor 1 (aka upstream stimulatory factor 1). "This gene encodes a member of the basic helix-loop-helix leucine zipper family, and can function as a cellular transcription factor. The encoded protein can activate transcription through pyrimidine-rich initiator (Inr) elements and E-box motifs. This gene has been linked to familial combined hyperlipidemia (FCHL). Alternative splicing of this gene results in multiple transcript variants. A related pseudogene has been defined on chromosome 21."[4]

  1. NP_009053.1 upstream stimulatory factor 1 isoform 1 (variant 1).
  2. NP_996888.1 upstream stimulatory factor 1 isoform 2 (variant 2).
  3. NP_001263302.1 upstream stimulatory factor 1 isoform 1 (variant 3).

Gene ID: 7392 is USF2 upstream transcription factor 2, c-fos interacting. "This gene encodes a member of the basic helix-loop-helix leucine zipper family of transcription factors. The encoded protein can activate transcription through pyrimidine-rich initiator (Inr) elements and E-box motifs and is involved in regulating multiple cellular processes."[5]

  1. NP_003358.1 upstream stimulatory factor 2 isoform 1.
  2. NP_997174.1 upstream stimulatory factor 2 isoform 2.
  3. NP_001308079.1 upstream stimulatory factor 2 isoform 3.
  4. XP_024307452.1 upstream stimulatory factor 2 isoform X1.
  5. XP_005259254.1 upstream stimulatory factor 2 isoform X2.
  6. XP_024307453.1 upstream stimulatory factor 2 isoform X3.
  7. XP_016882688.1 upstream stimulatory factor 2 isoform X4.
  8. XP_011525562.1 upstream stimulatory factor 2 isoform X5.
  9. XP_011525563.1 upstream stimulatory factor 2 isoform X6.

Gene ID: 205717 is USF3 upstream transcription factor family member 3. "This gene encodes a large protein that contains a helix-loop-helix domain and a polyglutamine region. A deletion in the polyglutamine region was associated with risk for thyroid carcinoma."[6]

  1. NP_001009899.3 basic helix-loop-helix domain-containing protein USF3 (variant 1).
  2. NR_111981.1 RNA Sequence (non-coding, variant 2).
  3. XP_024309159.1 basic helix-loop-helix domain-containing protein USF3 isoform X1.
  4. XP_024309160.1 basic helix-loop-helix domain-containing protein USF3 isoform X2.
  5. XP_016861360.1 basic helix-loop-helix domain-containing protein USF3 isoform X1.
  6. XP_016861361.1 basic helix-loop-helix domain-containing protein USF3 isoform X2.
  7. XP_005247265.2 basic helix-loop-helix domain-containing protein USF3 isoform X2.

Gene ID: 100151645 is USF1P1 upstream transcription factor 1 pseudogene 1.

Hypotheses

  1. A1BG has no USF boxes in either promoter.

Samplings

Using the two versions of the consensus sequence and "⌘F" to locate these sequences in the nucleotides for A1BG listed in A1BG gene transcriptions revealed two occurrences (GCCTGGGA) and (GCCTTCCG) on the ZNF497 side.

For the Basic programs testing consensus sequence 3'-GCC(A/T)NN(C/G/T)(A/G)-5' (starting with SuccessablesUSFbox.bas) written to compare nucleotide sequences with the sequences on either the template strand (-), or coding strand (+), of the DNA, in the negative direction (-), or the positive direction (+), the programs are, are looking for, and found:

  1. negative strand in the negative direction (from ZSCAN22 to A1BG) is SuccessablesUSFbox--.bas, looking for 3'-GCC(A/T)NN(C/G/T)(A/G)-5', 0.
  2. negative strand in the positive direction (from ZNF497 to A1BG) is SuccessablesUSFbox-+.bas, looking for 3'-GCC(A/T)NN(C/G/T)(A/G)-5', 27 , 3'-GCCTGGGA-5', 288 , 3'-GCCAGCGG-5', 331 , 3'-GCCTTCCG-5', 678 , 3'-GCCTGCCG-5', 809 , 3'-GCCACACG-5', 886 , 3'-GCCTGCCG-5', 909 , 3'-GCCACACG-5', 986 , 3'-GCCTCATG-5', 1238 , 3'-GCCACCGG-5', 1294 , 3'-GCCTTCCG-5', 1334 , 3'-GCCTTCCG-5', 1434 , 3'-GCCACCGG-5', 1546 , 3'-GCCTTTCA-5', 1601 , 3'-GCCTTGGG-5', 1800 , 3'-GCCACCCA-5', 1853 , 3'-GCCTCCCA-5', 2396 , 3'-GCCTCCCA-5', 2532 , 3'-GCCAGGGA-5', 2576 , 3'-GCCAAAGG-5', 2828 , 3'-GCCTCTGG-5', 2883 , 3'-GCCAATGG-5', 2910 , 3'-GCCTGTGG-5', 3436 , 3'-GCCACATG-5', 3707 , 3'-GCCTGGGA-5', 3760 , 3'-GCCTCAGA-5', 4195 , 3'-GCCTTCCG-5', 4243 , 3'-GCCTCCTG-5', 4408.
  3. positive strand in the negative direction (from ZSCAN22 to A1BG) is SuccessablesUSFbox+-.bas, looking for 3'-GCC(A/T)NN(C/G/T)(A/G)-5', 11, 3'-GCCACCGA-5', 385, 3'-GCCTAGTG-5', 432, 3'-GCCACCGA-5', 658, 3'-GCCTAGTG-5', 705, 3'-GCCACTCG-5', 870, 3'-GCCACCGA-5', 1767, 3'-GCCTAGGG-5', 1814, 3'-GCCACCGA-5', 2202, 3'-GCCTCCGA-5', 2359, 3'-GCCACCGA-5', 2529, 3'-GCCTGACG-5', 4329.
  4. positive strand in the positive direction (from ZNF497 to A1BG) is SuccessablesUSFbox++.bas, looking for 3'-GCC(A/T)NN(C/G/T)(A/G)-5', 4 , 3'-GCCACCCG-5', 404, 3'-GCCTGACG-5', 748, 3'-GCCTCGTG-5', 1207, 3'-GCCAACGG-5', 3492.
  5. complement, negative strand, negative direction is SuccessablesUSFboxc--.bas, looking for 3'-CGG(A/T)NN(A/C/G)(C/T)-5', 13, 3'-CGGTATAC-5', 41, 3'-CGGTCCGT-5', 650, 3'-CGGTCCGT-5', 950, 3'-CGGAGGGC-5', 1508, 3'-CGGTTTCC-5', 1639, 3'-CGGTCCAC-5', 2079, 3'-CGGTCCGT-5', 2521, 3'-CGGTGGGT-5', 3135, 3'-CGGACAGT-5', 3202, 3'-CGGTCCAC-5', 3953, 3'-CGGTCCGT-5', 4104, 3'-CGGTCTGC-5', 4235, 3'-CGGACCCT-5', 4302.
  6. complement, negative strand, positive direction is SuccessablesUSFboxc-+.bas, looking for 3'-CGG(A/T)NN(A/C/G)(C/T)-5', 4, 3'-CGGTGGGC-5', 404, 3'-CGGACTGC-5', 748, 3'-CGGAGCAC-5', 1207, 3'-CGGTTGCC-5', 3492.
  7. complement, positive strand, negative direction is SuccessablesUSFboxc+-.bas, looking for 3'-CGG(A/T)NN(A/C/G)(C/T)-5', 11, 3'-CGGTGGCT-5', 385, 3'-CGGATCAC-5', 432, 3'-CGGTGGCT-5', 658, 3'-CGGATCAC-5', 705, 3'-CGGTGAGC-5', 870, 3'-CGGTGGCT-5', 1767, 3'-CGGATCCC-5', 1814, 3'-CGGTGGCT-5', 2202, 3'-CGGAGGCT-5', 2359, 3'-CGGTGGCT-5', 2529, 3'-CGGACTGC-5', 4329.
  8. complement, positive strand, positive direction is SuccessablesUSFboxc++.bas, looking for 3'-CGG(A/T)NN(A/C/G)(C/T)-5', 27 , 3'-CGGACCCT-5', 288 , 3'-CGGTCGCC-5', 331 , 3'-CGGAAGGC-5', 678 , 3'-CGGACGGC-5', 809 , 3'-CGGTGTGC-5', 886 , 3'-CGGACGGC-5', 909 , 3'-CGGTGTGC-5', 986 , 3'-CGGAGTAC-5', 1238 , 3'-CGGTGGCC-5', 1294 , 3'-CGGAAGGC-5', 1334 , 3'-CGGAAGGC-5', 1434 , 3'-CGGTGGCC-5', 1546 , 3'-CGGAAAGT-5', 1601 , 3'-CGGAACCC-5', 1800 , 3'-CGGTGGGT-5', 1853 , 3'-CGGAGGGT-5', 2396 , 3'-CGGAGGGT-5', 2532 , 3'-CGGTCCCT-5', 2576 , 3'-CGGTTTCC-5', 2828 , 3'-CGGAGACC-5', 2883 , 3'-CGGTTACC-5', 2910 , 3'-CGGACACC-5', 3436 , 3'-CGGTGTAC-5', 3707 , 3'-CGGACCCT-5', 3760 , 3'-CGGAGTCT-5', 4195 , 3'-CGGAAGGC-5', 4243 , 3'-CGGAGGAC-5', 4408.
  9. inverse complement, negative strand, negative direction is SuccessablesUSFboxci--.bas, looking for 3'-(C/T)(A/C/G)NN(A/T)GGC-5', 0.
  10. inverse complement, negative strand, positive direction is SuccessablesUSFboxci-+.bas, looking for 3'-(C/T)(A/C/G)NN(A/T)GGC-5', 22, 3'-CCTCAGGC-5', 91, 3'-TCACAGGC-5', 158, 3'-CAGGTGGC-5', 198, 3'-CGTGTGGC-5', 1023, 3'-TACCTGGC-5', 1200, 3'-CGAGAGGC-5', 1384, 3'-CGAGAGGC-5', 1484, 3'-CGAGAGGC-5', 1568, 3'-CGGCAGGC-5', 1906, 3'-TCTCTGGC-5', 1993, 3'-TCTCAGGC-5', 2116, 3'-TCTTAGGC-5', 2367, 3'-CACCTGGC-5', 2434, 3'-CACCTGGC-5', 2570, 3'-TGCAAGGC-5', 2694, 3'-CCTCTGGC-5', 2884, 3'-CCTCTGGC-5', 2984, 3'-CATCTGGC-5', 3406, 3'-CAACAGGC-5', 3637, 3'-CGGGAGGC-5', 3675, 3'-CAGCAGGC-5', 3695, 3'-CCTGAGGC-5', 4189.
  11. inverse complement, positive strand, negative direction is SuccessablesUSFboxci+-.bas, looking for 3'-(C/T)(A/C/G)NN(A/T)GGC-5', 29, 3'-TGGGAGGC-5', 415, 3'-TGCGAGGC-5', 451, 3'-TGGGAGGC-5', 552, 3'-TGGGAGGC-5', 688, 3'-CAGGAGGC-5', 854, 3'-CAGGAGGC-5', 988, 3'-TGGGAGGC-5', 1020, 3'-CAGGAGGC-5', 1279, 3'-TAGGAGGC-5', 1311, 3'-TGGGAGGC-5', 1797, 3'-TGGGAGGC-5', 1964, 3'-TGGGAGGC-5', 2106, 3'-TGGGAGGC-5', 2223, 3'-CCGGAGGC-5', 2358, 3'-TGGCAGGC-5', 2390, 3'-TGGAAGGC-5', 2559, 3'-CAGGAGGC-5', 2693, 3'-TAGATGGC-5', 2907, 3'-TGGGAGGC-5', 3082, 3'-CAGGAGGC-5', 3221, 3'-CATCAGGC-5', 3397, 3'-TAAAAGGC-5', 3442, 3'-CAGATGGC-5', 3629, 3'-TGGGAGGC-5', 3716, 3'-TAAGAGGC-5', 3896, 3'-TGGTTGGC-5', 3947, 3'-TGGGAGGC-5', 3991, 3'-CAGGAGGC-5', 4142, 3'-TGGGAGGC-5', 4273.
  12. inverse complement, positive strand, positive direction is SuccessablesUSFboxci++.bas, looking for 3'-(C/T)(A/C/G)NN(A/T)GGC-5', 15, 3'-TGGGTGGC-5', 74, 3'-TCCAAGGC-5', 307, 3'-CGTCTGGC-5', 441, 3'-CGGAAGGC-5', 678, 3'-CGAAAGGC-5', 1098, 3'-TCGGTGGC-5', 1293, 3'-CGGAAGGC-5', 1334, 3'-CGGAAGGC-5', 1434, 3'-TCGGTGGC-5', 1545, 3'-TCGGTGGC-5', 1608, 3'-CAGGTGGC-5', 1845, 3'-CCAGAGGC-5', 1961, 3'-TCGGTGGC-5', 2562, 3'-CCGTTGGC-5', 3912, 3'-CGGAAGGC-5', 4243.
  13. inverse negative strand, negative direction is SuccessablesUSFboxi--.bas, looking for 3'-(A/G)(C/G/T)NN(A/T)CCG-5', 29, 3'-ACCCTCCG-5', 415, 3'-ACGCTCCG-5', 451, 3'-ACCCTCCG-5', 552, 3'-ACCCTCCG-5', 688, 3'-GTCCTCCG-5', 854, 3'-GTCCTCCG-5', 988, 3'-ACCCTCCG-5', 1020, 3'-GTCCTCCG-5', 1279, 3'-ATCCTCCG-5', 1311, 3'-ACCCTCCG-5', 1797, 3'-ACCCTCCG-5', 1964, 3'-ACCCTCCG-5', 2106, 3'-ACCCTCCG-5', 2223, 3'-GGCCTCCG-5', 2358, 3'-ACCGTCCG-5', 2390, 3'-ACCTTCCG-5', 2559, 3'-GTCCTCCG-5', 2693, 3'-ATCTACCG-5', 2907, 3'-ACCCTCCG-5', 3082, 3'-GTCCTCCG-5', 3221, 3'-GTAGTCCG-5', 3397, 3'-ATTTTCCG-5', 3442, 3'-GTCTACCG-5', 3629, 3'-ACCCTCCG-5', 3716, 3'-ATTCTCCG-5', 3896, 3'-ACCAACCG-5', 3947, 3'-ACCCTCCG-5', 3991, 3'-GTCCTCCG-5', 4142, 3'-ACCCTCCG-5', 4273.
  14. inverse negative strand, positive direction is SuccessablesUSFboxi-+.bas, looking for 3'-(A/G)(C/G/T)NN(A/T)CCG-5', 15, 3'-ACCCACCG-5', 74, 3'-AGGTTCCG-5', 307, 3'-GCAGACCG-5', 441, 3'-GCCTTCCG-5', 678, 3'-GCTTTCCG-5', 1098, 3'-AGCCACCG-5', 1293, 3'-GCCTTCCG-5', 1334, 3'-GCCTTCCG-5', 1434, 3'-AGCCACCG-5', 1545, 3'-AGCCACCG-5', 1608, 3'-GTCCACCG-5', 1845, 3'-GGTCTCCG-5', 1961, 3'-AGCCACCG-5', 2562, 3'-GGCAACCG-5', 3912, 3'-GCCTTCCG-5', 4243.
  15. inverse positive strand, negative direction is SuccessablesUSFboxi+-.bas, looking for 3'-(A/G)(C/G/T)NN(A/T)CCG-5', 0.
  16. inverse positive strand, positive direction is SuccessablesUSFboxi++.bas, looking for 3'-(A/G)(C/G/T)NN(A/T)CCG-5', 22, 3'-GGAGTCCG-5', 91, 3'-AGTGTCCG-5', 158, 3'-GTCCACCG-5', 198, 3'-GCACACCG-5', 1023, 3'-ATGGACCG-5', 1200, 3'-GCTCTCCG-5', 1384, 3'-GCTCTCCG-5', 1484, 3'-GCTCTCCG-5', 1568, 3'-GCCGTCCG-5', 1906, 3'-AGAGACCG-5', 1993, 3'-AGAGTCCG-5', 2116, 3'-AGAATCCG-5', 2367, 3'-GTGGACCG-5', 2434, 3'-GTGGACCG-5', 2570, 3'-ACGTTCCG-5', 2694, 3'-GGAGACCG-5', 2884, 3'-GGAGACCG-5', 2984, 3'-GTAGACCG-5', 3406, 3'-GTTGTCCG-5', 3637, 3'-GCCCTCCG-5', 3675, 3'-GTCGTCCG-5', 3695, 3'-GGACTCCG-5', 4189.

See also

References

  1. 1.0 1.1 1.2 1.3 1.4 Marie-Dominique Galibert, Suzanne Carreira, and Colin R. Goding (2001 September 3). "The Usf-1 transcription factor is a novel target for the stress-responsive p38 kinase and mediates UV-induced Tyrosinase expression". EMBO Journal. 20 (17): 5022–5031. doi:10.1093/emboj/20.17.5022. Retrieved 7 December 2018. Check date values in: |date= (help)
  2. 2.0 2.1 Libo Sa, Yan Li, Lini Zhao1, Yunhui Liu, Ping Wang, Libo Liu, Zhen Li, Jun Ma, Heng Cai and Yixue Xue (28 June 2017). "The Role of HOTAIR/miR-148b-3p/USF1 on Regulating the Permeability of BTB". Frontiers in Molecular Neuroscience. 10 (194): 194. doi:10.3389/fnmol.2017.00194. Retrieved 20 August 2020.
  3. 3.0 3.1 3.2 3.3 Martin L. Read, Andrew R. Clark and Kevin Docherty (1993). "The helix-loop-helix transcription factor USF (upstream stimulating factor) binds to a regulatory sequence of the human insulin gene enhancer" (PDF). Biochemical Journal. 295: 233–237. Retrieved 14 August 2020.
  4. RefSeq (February 2013). "USF1 upstream transcription factor 1 [ Homo sapiens (human) ]". 8600 Rockville Pike, Bethesda MD, 20894 USA: National Center for Biotechnology Information, U.S. National Library of Medicine. Retrieved 10 December 2018.
  5. RefSeq (March 2016). "USF2 upstream transcription factor 2, c-fos interacting [ Homo sapiens (human) ]". 8600 Rockville Pike, Bethesda MD, 20894 USA: National Center for Biotechnology Information, U.S. National Library of Medicine. Retrieved 10 December 2018.
  6. RefSeq (May 2017). "USF3 upstream transcription factor family member 3 [ Homo sapiens (human) ]". 8600 Rockville Pike, Bethesda MD, 20894 USA: National Center for Biotechnology Information, U.S. National Library of Medicine. Retrieved 10 December 2018.

External links

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