footprintDB has been developed by Alvaro Sebastian Yague and Bruno Contreras Moreira and it is maintained at the Laboratory of Computational Biology (Zaragoza, Spain), which is supported by ARAID and CSIC.

footprintDB is part of the project Novel Knowledge Based Abiotic Stress Regulators (STREG) funded by the the 2008 Euro Research Program [EUI2008-03612].

If you use this resource we ask you to cite the following paper:

and the original data sources from footprintDB:
  • Matys V, Fricke E, Geffers R, Gössling E, Haubrock M, Hehl R, Hornischer K, Karas D, Kel A.E, Kel-Margoulis O.V, Kloos D.U, Land S, Lewicki-Potapov B, Michael H, Münch R, Reuter I, Rotert S, Saxel H, Scheer M, Thiele S, Wingender E. TRANSFAC: transcriptional regulation, from patterns to profiles. Nucleic acids research 31:374-8 (2003). [Pubmed]
  • Salgado H, Peralta-Gil M, Gama-Castro S, Santos-Zavaleta A, Muniz-Rascado L, Garcia-Sotelo JS, Weiss V, Solano-Lira H, Martinez-Flores I, Medina-Rivera A, Salgado-Osorio G, Alquicira-Hernandez S, Alquicira-Hernandez K, Lopez-Fuentes A, Porron-Sotelo L, Huerta AM, Bonavides-Martinez C, Balderas-Martinez YI, Pannier L, Olvera M, Labastida A, Jimenez-Jacinto V, Vega-Alvarado L, Del Moral-Chavez V, Hernandez-Alvarez A, Morett E, Collado-Vides J. RegulonDB v8.0: omics data sets, evolutionary conservation, regulatory phrases, cross-validated gold standards and more. Nucleic Acids Res. 2013 Jan 1;41(D1):D203-D213. [Pubmed]
  • Contreras-Moreira B. 3D-footprint: a database for the structural analysis of protein-DNA complexes. Nucleic acids research 38:D91-7 (2010). [Pubmed]
  • Hume MA, Barrera LA, Gisselbrecht SS, Bulyk ML. UniPROBE, update 2015: new tools and content for the online database of protein-binding microarray data on protein-DNA interactions. Nucleic Acids Res : (2015). [Pubmed]
  • Down T.A, Bergman C.M, Su J, Hubbard T.J. Large-scale discovery of promoter motifs in Drosophila melanogaster. PLoS computational biology 3:e7 (2007). [Pubmed]
  • Bülow L, Engelmann S, Schindler M, Hehl R. AthaMap, integrating transcriptional and post-transcriptional data. Nucleic acids research 37:D983-6 (2009). [Pubmed]
  • Sierro N, Makita Y, de Hoon M, Nakai K. DBTBS: a database of transcriptional regulation in Bacillus subtilis containing upstream intergenic conservation information. Nucleic acids research 36:D93-6 (2008). [Pubmed]
  • Jolma A, Yan J, Whitington T, Toivonen J, Nitta KR, Rastas P, Morgunova E, Enge M, Taipale M, Wei G, Palin K, Vaquerizas JM, Vincentelli R, Luscombe NM, Hughes TR, Lemaire P, Ukkonen E, Kivioja T, Taipale J. DNA-Binding Specificities of Human Transcription Factors. Cell. 2013 Jan 17;152(1-2):327-39. [Pubmed]
  • Kulakovskiy IV, Vorontsov IE, Yevshin IS, Soboleva AV, Kasianov AS, Ashoor H, Ba-Alawi W, Bajic VB, Medvedeva YA, Kolpakov FA, Makeev VJ. HOCOMOCO: expansion and enhancement of the collection of transcription factor binding sites models. Nucleic Acids Res : (2016). [Pubmed]
  • Franco-Zorrilla J.M, López-Vidriero I, Carrasco J.L, Godoy M, Vera P, Solano R. DNA-binding specificities of plant transcription factors and their potential to define target genes. Proceedings of the National Academy of Sciences of the United States of America : (2014). [Pubmed]
  • Enuameh MS et al (2013) Global analysis of Drosophila Cys2-His2 zinc finger proteins reveals a multitude of novel recognition motifs and binding determinants. Genome Res. 23(6):928-40. doi: 10.1101/gr.151472.112 [Pubmed]
  • Kelemen Z., Sebastian A., Xu W., Grain D., Salsac F., Avon A., Berger N., Tran J., Dubrecq B., Lurin C., Lepiniec L., Contreras-Moreira B., Dubos C. Analysis of the DNA-Binding Activities of the Arabidopsis R2R3-MYB Transcription Factor Family by One-Hybrid Experiments in Yeast. PLoS One 10, e0141044 (2015). [Pubmed]
  • Weirauch MT, Yang A, Albu M, Cote AG, Montenegro-Montero A, Drewe P, Najafabadi HS, Lambert SA, Mann I, Cook K, Zheng H, Goity A, van Bakel H, Lozano JC, Galli M, Lewsey MG, Huang E, Mukherjee T, Chen X, Reece-Hoyes JS, Govindarajan S, Shaulsky G, et al. Determination and inference of eukaryotic transcription factor sequence specificity. Cell. 2014 Sep 11;158(6):1431-43. doi: 10.1016/j.cell.2014.08.009. [Pubmed]
  • Jolma A, Yin Y, Nitta KR, Dave K, Popov A, Taipale M, Enge M, Kivioja T, Morgunova E, Taipale J. DNA-dependent formation of transcription factor pairs alters their binding specificity. Nature 527:384-8 (2015). [Pubmed]
  • O'Malley RC, Huang SS, Song L, Lewsey MG, Bartlett A, Nery JR, Galli M, Gallavotti A, Ecker JR. Cistrome and Epicistrome Features Shape the Regulatory DNA Landscape. Cell 165:1280-92 (2016). [Pubmed]
  • Isakova A, Groux R, Imbeault M, Rainer P, Alpern D, Dainese R, Ambrosini G, Trono D, Bucher P, Deplancke B. SMiLE-seq identifies binding motifs of single and dimeric transcription factors. Nat Methods 14:316-322 (2017). [Pubmed]
  • Yin Y, Morgunova E, Jolma A, Kaasinen E, Sahu B, Khund-Sayeed S, Das PK, Kivioja T, Dave K, Zhong F, Nitta KR, Taipale M, Popov A, Ginno PA, Domcke S, Yan J, Schübeler D, Vinson C, Taipale J. Impact of cytosine methylation on DNA binding specificities of human transcription factors. Science : (2017). [Pubmed]
  • Najafabadi HS, Mnaimneh S, Schmitges FW, Garton M, Lam KN, Yang A, Albu M, Weirauch MT, Radovani E, Kim PM, Greenblatt J, Frey BJ, Hughes TR. C2H2 zinc finger proteins greatly expand the human regulatory lexicon. Nat Biotechnol 33:555-62 (2015). [Pubmed]
  • [Pubmed]

You can also consider citing other relevant papers:
  • Contreras-Moreira B. 3D-footprint: a database for the structural analysis of protein-DNA complexes. Nucleic Acids Res. 2010 Jan;38(Database issue):D91-7. Epub 2009 Sep 18. [Pubmed]
  • Contreras-Moreira B, Branger PA, Collado-Vides J. TFmodeller: comparative modelling of protein-DNA complexes. Bioinformatics. 2007 Jul 1;23(13):1694-6. Epub 2007 Apr 25. [Pubmed]
  • Angarica VE, Pérez AG, Vasconcelos AT, Collado-Vides J, Contreras-Moreira B. Prediction of TF target sites based on atomistic models of protein-DNA complexes. BMC Bioinformatics. 2008 Oct 16;9:436. [Pubmed]
  • Morozov AV, Siggia ED. Connecting protein structure with predictions of regulatory sites. Proc Natl Acad Sci U S A. 2007 Apr 24;104(17):7068-73. Epub 2007 Apr 16. [Pubmed]
  • Contreras-Moreira B, Sancho J, Angarica VE. Comparison of DNA binding across protein superfamilies. Proteins. 2010 Jan;78(1):52-62. [Pubmed]

footprintDB relies on several valuable resources like the Protein Data Bank, SCOP, Superfamily, Pfam, BLAST, HMMER, MAMMOTH, STAMP, WEBLOGO, TFmodeller, PubMed, R and RSA-tools.

Should you have any questions regarding footprintDB, please get in touch at


These data are available AS IS and at your own risk. The EEAD/CSIC do not give any representation or warranty nor assume any liability or responsibility for the data nor the results posted (whether as to their accuracy, completeness, quality or otherwise). Access to these data is available free of charge for ordinary use in the course of research.