The Transporter Classification Database (TCDB): 2021 update
Open accessAbstract
The Transporter Classification Database (TCDB; tcdb.org) is a freely accessible reference resource, which provides functional, structural, mechanistic, medical and biotechnological information about transporters from organisms of all types. TCDB is the only transport protein classification database adopted by the International Union of Biochemistry and Molecular Biology (IUBMB) and now (October 1, 2020) consists of 20 653 proteins classified in 15 528 non-redundant transport systems with 1567 tabulated 3D structures, 18 336 reference citations describing 1536 transporter families, of which 26% are members of 82 recognized superfamilies. Overall, this is an increase of over 50% since the last published update of the database in 2016. This comprehensive update of the database contents and features include (i) adoption of a chemical ontology for substrates of transporters, (ii) inclusion of new superfamilies, (iii) a domain-based characterization of transporter families for the identification of new members as well as functional and evolutionary relationships between families, (iv) development of novel software to facilitate curation and use of the database, (v) addition of new subclasses of transport systems including 11 novel types of channels and 3 types of group translocators and (vi) the inclusion of many man-made (artificial) transmembrane pores/channels and carriers.
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Operon 36 encodes a 4.A.3 family PTS, a GH1 6-phospho-beta-glucosidase (bglA), and a gntR/yydK-like regulator.
“Likewise, operon 36 encodes a 4.A.3 family PTS [90] , a GH1 6-phospho-beta-glucosidase ( bgl A), and a gnt R/ yyd K-like regulator [Figure 6B].”
From Genome-scale insights into the metabolic landscape and evolutionary development of Bifidobacterium bifidum (2026) · cites this paper as [90]
The citation is attached to the identification of the operon’s PTS as family 4.A.3, not to the later gene descriptions. The Introduction describes TCDB’s transporter-classification system, but searches for “4.A.3” and “phosphotransferase system PTS” found no passage in our copy identifying that family or discussing operon 36. (LLM, conf=0.72)
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Cites
Gapped BLAST and PSI-BLAST: a new generation of protein database search programs
Altschul, S. (1997) · Nucleic Acids Research · doi:10.1093/nar/25.17.3389 · Cited by 73,302 · see citing works
MUSCLE: multiple sequence alignment with high accuracy and high throughput
Edgar, R. C. (2004) · Nucleic Acids Research · doi:10.1093/nar/gkh340 · Cited by 44,662 · see citing works
Accelerated Profile HMM Searches
Eddy, S. R. (2011) · PLoS Computational Biology · doi:10.1371/journal.pcbi.1002195 · Cited by 6,975 · see citing works
The Pfam protein families database in 2019
El-Gebali, S., Mistry, J., Bateman, A., Eddy, S. R., Luciani, A., Potter, S. C., Qureshi, M., Richardson, L. J., Salazar, G. A., Smart, A., et al. (2019) · Nucleic Acids Research · doi:10.1093/nar/gky995 · Cited by 4,602 · see citing works
CDD/SPARCLE: the conserved domain database in 2020
Lu, S., Wang, J., Chitsaz, F., Derbyshire, M. K., Geer, R. C., Gonzales, N. R., Gwadz, M., Hurwitz, D. I., Marchler, G. H., Song, J. S., et al. (2019) · Nucleic Acids Research · doi:10.1093/nar/gkz991 · Cited by 3,049 · see citing works
Self-citationA new view of the tree of life
Hug, L. A., Baker, B. J., Anantharaman, K., Brown, C. T., Probst, A. J., Castelle, C. J., Butterfield, C. N., Hernsdorf, A. W., Amano, Y., Ise, K., et al. (2016) · Nature Microbiology · doi:10.1038/nmicrobiol.2016.48 · Cited by 2,039 · see citing works
Asgard archaea illuminate the origin of eukaryotic cellular complexity
Zaremba-Niedzwiedzka, K., Caceres, E. F., Saw, J. H., Bäckström, D., Juzokaite, L., Vancaester, E., Seitz, K. W., Anantharaman, K., Starnawski, P., Kjeldsen, K. U., et al. (2017) · Nature · doi:10.1038/nature21031 · Cited by 1,226 · see citing works
ChEBI in 2016: Improved services and an expanding collection of metabolites
Hastings, J., Owen, G., Dekker, A., Ennis, M., Kale, N., Muthukrishnan, V., Turner, S., Swainston, N., Mendes, P., & Steinbeck, C. (2016) · Nucleic Acids Research · doi:10.1093/nar/gkv1031 · Cited by 1,083 · see citing works
The Transporter Classification Database (TCDB): recent advances
Saier, M. H., Reddy, V. S., Tsu, B. V., Ahmed, M. S., Li, C., & Moreno‐Hagelsieb, G. (2015) · Nucleic Acids Research · doi:10.1093/nar/gkv1103 · Cited by 989 · see citing works
Self-citationA major superfamily of transmembrane facilitators that catalyse uniport, symport and antiport
Marger, M. D., & Saier, M. H. (1993) · Trends in Biochemical Sciences · doi:10.1016/0968-0004(93)90081-w · Cited by 954 · see citing works
Self-citationSaier, M. H. (2005) · Nucleic Acids Research · doi:10.1093/nar/gkj001 · Cited by 848 · see citing works
Self-citationMolecular Properties of Bacterial Multidrug Transporters
Putman, M., van Veen, H. W., & Konings, W. N. (2000) · Microbiology and Molecular Biology Reviews · doi:10.1128/mmbr.64.4.672-693.2000 · Cited by 792 · see citing works
The Transporter Classification Database
Saier, M. H., Reddy, V. S., Tamang, D. G., & Västermark, Å. (2013) · Nucleic Acids Research · doi:10.1093/nar/gkt1097 · Cited by 493 · see citing works
Self-citationThe Transporter Classification Database: recent advances
Saier, M. H., Yen, M. R., Noto, K., Tamang, D. G., & Elkan, C. (2008) · Nucleic Acids Research · doi:10.1093/nar/gkn862 · Cited by 414 · see citing works
Self-citationBurley, S. K., Berman, H. M., Christie, C. H., Duarte, J. M., Feng, Z., Westbrook, J., Young, J. Y., & Zardecki, C. (2017) · Protein Science · doi:10.1002/pro.3331 · Cited by 296 · see citing works
The Transporter Classification (TC) System, 2002
Busch, W., & Saier, M. H. (2002) · Critical Reviews in Biochemistry and Molecular Biology · doi:10.1080/10409230290771528 · Cited by 259 · see citing works
Self-citationErnst, C. M., & Peschel, A. (2011) · Molecular Microbiology · doi:10.1111/j.1365-2958.2011.07576.x · Cited by 238 · see citing works
Bacterial Multidrug Efflux Transporters
Delmar, J. A., Su, C., & Yu, E. W. (2014) · Annual Review of Biophysics · doi:10.1146/annurev-biophys-051013-022855 · Cited by 187 · see citing works
Opsin Is a Phospholipid Flippase
Menon, I. A., Huber, T., Sanyal, S., Banerjee, S., Barré, P., Canis, S., Warren, J. D., Hwa, J., Sakmar, T. P., & Menon, A. K. (2011) · Current Biology · doi:10.1016/j.cub.2010.12.031 · Cited by 159 · see citing works
Tracing pathways of transport protein evolution
Saier, M. H. (2003) · Molecular Microbiology · doi:10.1046/j.1365-2958.2003.03499.x · Cited by 154 · see citing works
Self-citationEnergetics of Pathogenic Bacteria and Opportunities for Drug Development
Cook, G. M., Greening, C., Hards, K., & Berney, M. (2014) · Advances in microbial physiology/Advances in Microbial Physiology · doi:10.1016/bs.ampbs.2014.08.001 · Cited by 126 · see citing works
Maity, K., Heumann, J. M., McGrath, A. P., Kopcho, N. J., Hsu, P., Lee, C., Mapes, J., Garza, D., Krishnan, S., Morgan, G., et al. (2019) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1900774116 · Cited by 125 · see citing works
Self-citationConstitutive phospholipid scramblase activity of a G protein-coupled receptor
Goren, M. A., Morizumi, T., Menon, I. A., Joseph, J. S., Dittman, J. S., Cherezov, V., Stevens, R. C., Ernst, O. P., & Menon, A. K. (2014) · Nature Communications · doi:10.1038/ncomms6115 · Cited by 122 · see citing works
Saier, M. H., & Reddy, B. L. (2014) · Journal of Bacteriology · doi:10.1128/jb.02046-14 · Cited by 122 · see citing works
Self-citationCastelle, C., Brown, C., Thomas, B., Williams, K., & Banfield, J. (2017) · Scientific Reports · doi:10.1038/srep40101 · Cited by 96 · see citing works
Sondén, B., Kocíncová, D., Deshayes, C., Euphrasie, D., Rhayat, L., Laval, F., Fréhel, C., Daffé, M., Etienne, G., & Reyrat, J. (2005) · Molecular Microbiology · doi:10.1111/j.1365-2958.2005.04847.x · Cited by 88 · see citing works
Membrane Porters of ATP-Binding Cassette Transport Systems Are Polyphyletic
Wang, B., Dukarevich, M., Sun, E. I., Yen, M. R., & Saier, M. H. (2009) · The Journal of Membrane Biology · doi:10.1007/s00232-009-9200-6 · Cited by 85 · see citing works
Self-citationVrljic, M., Garg, J., Bellmann, A., et al. (1999) · PubMed · Cited by 83 · see citing works
Topological and phylogenetic analyses of bacterial holin families and superfamilies
Reddy, B. L., & Saier, M. H. (2013) · Biochimica et Biophysica Acta (BBA) - Biomembranes · doi:10.1016/j.bbamem.2013.07.004 · Cited by 80 · see citing works
Self-citationThe IUBMB-Endorsed Transporter Classification System
Busch, W., & Saier, M. H. (2004) · Molecular Biotechnology · doi:10.1385/mb:27:3:253 · Cited by 80 · see citing works
Self-citationWang, S. C., Davejan, P. M., Hendargo, K. J., Javadi-Razaz, I., Chou, A., Yee, D. C., Ghazi, F., Lam, K. J. K., Conn, A. M., Madrigal, A. A., et al. (2020) · Biochimica et Biophysica Acta (BBA) - Biomembranes · doi:10.1016/j.bbamem.2020.183277 · Cited by 69 · see citing works
Self-citationStructure of the rice hyperosmolality-gated ion channel OSCA1.2
Maity, K., Heumann, J. M., McGrath, A. P., Chang, G. A., & Stowell, M. H. B. (2019) · Worldwide Protein Data Bank · doi:10.2210/pdb6oce/pdb · Cited by 63 · see citing works
Medrano-Soto, A., Moreno‐Hagelsieb, G., McLaughlin, D. J., Ye, Z. S., Hendargo, K. J., & Saier, M. H. (2018) · PLoS ONE · doi:10.1371/journal.pone.0192851 · Cited by 57 · see citing works
Self-citationTransport protein evolution deduced from analysis of sequence, topology and structure
Saier, M. H. (2016) · Current Opinion in Structural Biology · doi:10.1016/j.sbi.2016.05.001 · Cited by 55 · see citing works
Self-citationYen, M. R., Choi, J., & Saier, M. H. (2009) · Microbial Physiology · doi:10.1159/000239667 · Cited by 49 · see citing works
Self-citationThe Membrane Attack Complex/Perforin Superfamily
Moreno‐Hagelsieb, G., Vitug, B., Medrano-Soto, A., et al. (2017) · Microbial Physiology · doi:10.1159/000481286 · Cited by 46 · see citing works
Self-citationPhospholipid scrambling by rhodopsin
Ernst, O. P., & Menon, A. K. (2015) · Photochemical & Photobiological Sciences · doi:10.1039/c5pp00195a · Cited by 41 · see citing works
Comparative genomics of transport proteins in seven Bacteroides species
Zafar, H., & Saier, M. H. (2018) · PLoS ONE · doi:10.1371/journal.pone.0208151 · Cited by 39 · see citing works
Self-citationThe LysE Superfamily of Transport Proteins Involved in Cell Physiology and Pathogenesis
Tsu, B. V., & Saier, M. H. (2015) · PLoS ONE · doi:10.1371/journal.pone.0137184 · Cited by 37 · see citing works
Self-citationStructure and multistate function of the transmembrane electron transporter CcdA
Williamson, J. A., Cho, S.-H., Ye, J., Collet, J., Beckwith, J. R., & Chou, J. J. (2015) · Nature Structural & Molecular Biology · doi:10.1038/nsmb.3099 · Cited by 32 · see citing works
The IUBMB-Endorsed Transporter Classification System
Busch, W., & Saier, M. H. (2003) · Humana Press eBooks · doi:10.1385/1-59259-387-9:21 · Cited by 31 · see citing works
Self-citationDo, J., Zafar, H., & Saier, M. H. (2017) · Microbial Pathogenesis · doi:10.1016/j.micpath.2017.03.022 · Cited by 28 · see citing works
Self-citationReddy, B. L., & Saier, M. H. (2016) · PLoS ONE · doi:10.1371/journal.pone.0152733 · Cited by 27 · see citing works
Self-citationlibChEBI: an API for accessing the ChEBI database
Swainston, N., Hastings, J., Dekker, A., Muthukrishnan, V., May, J. W., Steinbeck, C., & Mendes, P. (2016) · Journal of Cheminformatics · doi:10.1186/s13321-016-0123-9 · Cited by 25 · see citing works
Comparative genomics of the transportome of Ten Treponema species
Buyuktimkin, B., Zafar, H., & Saier, M. H. (2019) · Microbial Pathogenesis · doi:10.1016/j.micpath.2019.04.034 · Cited by 23 · see citing works
Self-citationSolution structure and elevator mechanism of the membrane electron transporter CcdA
Zhou, Y., & Bushweller, J. H. (2018) · Nature Structural & Molecular Biology · doi:10.1038/s41594-018-0022-z · Cited by 19 · see citing works
Medrano-Soto, A., Ghazi, F., Hendargo, K. J., Moreno‐Hagelsieb, G., Myers, S., & Saier, M. H. (2020) · PLoS ONE · doi:10.1371/journal.pone.0231085 · Cited by 18 · see citing works
Self-citationComparative analyses of transport proteins encoded within the genomes of Leptospira species
Buyuktimkin, B., & Saier, M. H. (2016) · Microbial Pathogenesis · doi:10.1016/j.micpath.2016.06.013 · Cited by 11 · see citing works
Self-citationLee, J., Ghosh, S., & Saier, M. H. (2017) · Journal of Phycology · doi:10.1111/jpy.12534 · Cited by 10 · see citing works
Self-citationTajabadi, F. H., Medrano-Soto, A., Ahmadzadeh, M., Jouzani, G. S., & Saier, M. H. (2017) · Microbial Physiology · doi:10.1159/000484563 · Cited by 9 · see citing works
Self-citationA Riboflavin Transporter in Bdellovibrio exovorous JSS
Rodionova, I. A., Tajabadi, F. H., Zhang, Z., et al. (2019) · Microbial Physiology · doi:10.1159/000501354 · Cited by 4 · see citing works
Characterization of the Tetraspan Junctional Complex (4JC) superfamily
Chou, A., Lee, A. D. W., Hendargo, K. J., et al. (2016) · Biochimica et Biophysica Acta (BBA) - Biomembranes · doi:10.1016/j.bbamem.2016.11.015 · Cited by 3 · see citing works
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Wang, J., Chitsaz, F., Derbyshire, M. K., Gonzales, N. R., Gwadz, M., Lu, S., Marchler, G. H., Song, J. S., Thanki, N., Yamashita, R. A., et al. (2023) · Nucleic Acids Research · doi:10.1093/nar/gkac1096 · Cited by 1,277 · see citing works
Self-citationIns and outs of AlphaFold2 transmembrane protein structure predictions
Hegedűs, T., Geisler, M., Lukács, G. L., & Farkas, B. (2022) · Cellular and Molecular Life Sciences · doi:10.1007/s00018-021-04112-1 · Cited by 127 · see citing works
mGWAS-Explorer: Linking SNPs, Genes, Metabolites, and Diseases for Functional Insights
Chang, L., Zhou, G., Ou, H., & Xia, J. (2022) · Metabolites · doi:10.3390/metabo12060526 · Cited by 15 · see citing works
Selleri, E., Longhi, G., Tarracchini, C., Carolis, E. D., Petraro, S., Mancabelli, L., Milani, C., Turroni, F., Ventura, M., & Lugli, G. A. (2026) · Microbiome Research Reports · doi:10.20517/mrr.2026.19
- Authors
- Saier, M. H., Reddy, V. S., Moreno-Hagelsieb, G., Hendargo, K. J., Zhang, Y., Iddamsetty, V., Lam, K. J. K., Tian, N., Russum, S., Wang, J., & Medrano-Soto, A.
- Affiliations
- University of California San Diego (US)
- Department of Biology, Wilfrid Laurier University, Waterloo, Ontario N2L 3C5, Canada
- Department of Molecular Biology, Division of Biological Sciences, University of California at San Diego, La Jolla, CA 92093-0116, USA
- Corresponding author
- Saier, Milton H
- Medrano-Soto, Arturo
- Venue
- Nucleic Acids Research
- Publisher
- Oxford University Press (OUP)
- Published
- 2020-11-10
- Volume
- 49
- Issue
- D1
- Pages
- D461-D467
- ISSN
- 0305-1048 (ISSN-L)
- 1362-4962
- PMC
- PMC7778945
- OpenAlex
- W3099496190
- Cited by
- 467
- References
- 52
- Citations by year
- 2026: 72 · 2025: 112 · 2024: 118 · 2023: 81 · 2022: 61 · 2021: 23
- FWCI
- 14.90
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- Locations
- Nucleic Acids Research (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- Funders
- National Institutes of Health (GM077402)
- Topics
- Health Sciences › Medicine › Oncology › Drug Transport and Resistance Mechanisms
- Physical Sciences › Materials Science › Materials Chemistry › Enzyme Structure and Function
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Protein Structure and Dynamics
- Keywords
- transporter classification, membrane transporters, transporter families, transport channels
- MeSH terms
- Protein Domains, Software, Substrate Specificity, Membrane Transport Proteins, Databases, Protein, Metagenomics
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