2.A.102 The 4-Toluene Sulfonate Uptake Permease (TSUP) Family The TSUP family is also referred to as the TauE/SafE/YfcA/DUF81/COG0730 Family. Although its members have not been rigorously characterized, evidence is available that at least some members function in the transport of sulfur containing organic compounds. These include sulfolactate which may be exported by the TauE protein of Cupriavidus necator (Weinitschke et al. 2007; Mayer et al. 2012) and sulfoacetate which may be exported by the SafE1 protein of Neptuniibacter caesariensis (Krejcík et al. 2008). Another member of the TSUP family, TsaS of Comamonas testosteroni, has been reported to function in the uptake of 4-toluene sulfonate (Mampel et al. 2004). None of these functional assignments can be considered to be certain. The TSUP family has been characterized using bioinformatic approaches. Shlykov et al. 2012; showed that prokaryotic members of the family outnumber the eukaryotic members substantially, and in prokaryotes, but not eukaryotes, extensive lateral gene transfer occurred during family evolution. Despite unequal representation, homologues from the three taxonomic domains of life share well-conserved motifs. A prototypical eight TMS topology apparently arose by intragenic duplication of a four transmembrane segment (TMS) unit. Possibly, a two TMS α-helical hairpin structure was the precursor of the 4 TMS repeat unit. Genome context analyses confirmed the proposal of a sulfur-based compound transport role for many TSUP homologues, but functional outliers appear to be prevalent as well (Shlykov et al. 2012). The TSUP is an established member of the Transporter/Opsin/G protein-coupled receptor (TOG) superfamily (Yee et al. 2013).
This family belongs to the Transporter-Opsin-G protein-coupled receptor (TOG) Superfamily.
References:
Deutschbauer, A., M.N. Price, K.M. Wetmore, W. Shao, J.K. Baumohl, Z. Xu, M. Nguyen, R. Tamse, R.W. Davis, and A.P. Arkin. (2011). Evidence-based annotation of gene function in Shewanella oneidensis MR-1 using genome-wide fitness profiling across 121 conditions. PLoS Genet 7: e1002385.
Gristwood, T., M.B. McNeil, J.S. Clulow, G.P. Salmond, and P.C. Fineran. (2011). PigS and PigP regulate prodigiosin biosynthesis in Serratia via differential control of divergent operons, which include predicted transporters of sulfur-containing molecules. J. Bacteriol. 193: 1076-1085.
Hug, L.A., B.J. Baker, K. Anantharaman, C.T. Brown, A.J. Probst, C.J. Castelle, C.N. Butterfield, A.W. Hernsdorf, Y. Amano, K. Ise, Y. Suzuki, N. Dudek, D.A. Relman, K.M. Finstad, R. Amundson, B.C. Thomas, and J.F. Banfield. (2016). A new view of the tree of life. Nat Microbiol 1: 16048.
Krejcík, Z., K. Denger, S. Weinitschke, K. Hollemeyer, V. Paces, A.M. Cook, and T.H. Smits. (2008). Sulfoacetate released during the assimilation of taurine-nitrogen by Neptuniibacter caesariensis: purification of sulfoacetaldehyde dehydrogenase. Arch. Microbiol. 190: 159-168.
Lin, J., T. Peng, L. Jiang, J.Z. Ni, Q. Liu, L. Chen, and Y. Zhang. (2015). Comparative genomics reveals new candidate genes involved in selenium metabolism in prokaryotes. Genome Biol Evol 7: 664-676.
Mampel, J., E. Maier, T. Tralau, J. Ruff, R. Benz, and A.M. Cook. (2004). A novel outer-membrane anion channel (porin) as part of a putatively two-component transport system for 4-toluenesulphonate in Comamonas testosteroni T-2. Biochem. J. 383: 91-99.
Mayer, J., K. Denger, K. Hollemeyer, D. Schleheck, and A.M. Cook. (2012). (R)-Cysteate-nitrogen assimilation by Cupriavidus necator H16 with excretion of 3-sulfolactate: a patchwork pathway. Arch. Microbiol. 194: 949-957.
Shlykov, M.A., W.H. Zheng, J.S. Chen, and M.H. Saier, Jr. (2012). Bioinformatic characterization of the 4-Toluene Sulfonate Uptake Permease (TSUP) family of transmembrane proteins. Biochim. Biophys. Acta. 1818: 703-717.
Weinitschke, S., K. Denger, A.M. Cook, and T.H. Smits. (2007). The DUF81 protein TauE in Cupriavidus necator H16, a sulfite exporter in the metabolism of C2 sulfonates. Microbiology 153: 3055-3060.
Yee, D.C., M.A. Shlykov, A. Västermark, V.S. Reddy, S. Arora, E.I. Sun, and M.H. Saier, Jr. (2013). The transporter-opsin-G protein-coupled receptor (TOG) superfamily. FEBS J. 280: 5780-5800.
Examples:
TC#	Name	Organismal Type	Example
2.A.102.1.1	The putative 4-toluenesulfonate uptake permease, TsaS	Proteobacteria	*TsaS of Comamonas testosteroni* (Q6A553)**

Examples:
TC#	Name	Organismal Type	Example
Examples:
TC#	Name	Organismal Type	Example
2.A.102.2.1	Sulfolactate (sulfite/organosulfonate) exporter,TauE of 256 aas with 8 TMSs in a 4 + 4 TMS arrangement. Catalyzes export of sulfolactate as the terminal step in the metabolism of cysteate (Weinitschke et al. 2007; Mayer et al. 2012).	Bacteria	*TauE of Cupriavidus necator (Ralstonia eutropha) (Q0K020)*

2.A.102.2.10	Uncharacterized protein of 250 aas nad 6 TMSs.		UP of Arthrobacter crystallopoietes

2.A.102.2.11	Putative sulfite exporter, the TauE/SafE family protein of 241 aas and 7 or 8 TMSs with a probable 4 + 4 TMS arrangement.		*TauE homolog of Adlercreutzia* sp.**

2.A.102.2.12	TauE/SafE family of putative sulfite exporters.		*TauE homolog of Hydrogenobaculum* sp. (metagenome)**

2.A.102.2.2	*The SafE1 sulfoacetate efflux pump (Krejcík et al., 2008).*	Bacteria	*SafE of Neptuniibacter caesariensis* (Q2BM66)**

2.A.102.2.3	Transporter of unknown function	Bacteria	Uncharacterized permease of Rhodopirellula baltica

2.A.102.2.4	Uncharacterized protein	Proteobacteria	Uncharacterized protein of Rhodopseudomonas palustris

2.A.102.2.5	Uncharacterized protein	Actinobacteria	Uncharacterized protein of Streptomyces coelicolor

2.A.102.2.6	Putative sulfite exporter, TauE or SafE, of 347 aas and 7 - 8 TMSs.	Spirochaetes	SafE of Leptospira kmetyi

2.A.102.2.7	Uncharacterized protein of 240 aas	Actinobacteria	UP of Mycobacterium smegmatis

2.A.102.2.8	Uncharacterized protein of 245 aas	Proteobacteria	UP of Achromobacter xylosoxidans (Achromobacter insuavis)

2.A.102.2.9	Uncharacterized protein of 291 aas and 8 TMSs. A member of the TSUP or TauE/SafE family.		UP of E. coli

Examples:
TC#	Name	Organismal Type	Example
2.A.102.3.1	Orf of 269 aas and 6 TMSs	Bacteria	*YfcA of E. coli* (P0AD30)**

2.A.102.3.2	Uncharacterized protein of 253 aas and 8 TMSs		UP of Gordonia amicalis

2.A.102.3.3	*Uncharacterized protein of 251 aas and 6 TMSs (Hug et al.* 2016).**		UP of Candidatus Peribacter riflensis

2.A.102.3.4	*Glycine transporter, YfcA, of 269 aas and 9 TMSs (Deutschbauer et al.* 2011).**		YfcA of Shewanella oneidensis

Examples:
TC#	Name	Organismal Type	Example
2.A.102.4.1	Orf of 253aas and 8 TMSs	Archaea	*Orf of Pyrococcus abyssi (Q9UYH7)*

2.A.102.4.10	Uncharacterized protein of 671 aas and 9 TMSs. TMS1 is N-terminal, followed by a peptidase domain of 350 aas, followed by a transmembrane domain with 8 putative TMSs, homologous to other members of this family.	Proteobacteria	UP of Magnetococcus sp.

2.A.102.4.11	Putative permease of 259 aas and 8 TMSs		PP of Bdellovibrio bacteriovorus

2.A.102.4.12	Uncharacterized protein of 245 aas and 8 TMSs.		UP of Lactobacillus rhamnosus

2.A.102.4.13	*Acetate transporter of 266 aas and 8 TMSs, SO_1333 (Deutschbauer et al.* 2011).**		SO_1333 of Shewanella oneidensis

2.A.102.4.14	Uncharacterized membrane protein of 339 aas and 10 TMSs. The gene encoding this protein is adjacent to the ABC exporter with TC# 3.A.1.148.1.		*UP of Microbispora* sp. ATCC PTA-5024**

2.A.102.4.15	Uncharacterized protein of 270 aas and 8 TMSs.		UP of candidatus Bathyarchaeota archaeon (marine sediment metagenome)

2.A.102.4.2	Possible organo-sulfur-containing compound transporter (Gristwood et al., 2011) (271aas; 8 TMSs; duplicated). May function with two 4 TMS proteins, PmpA and PmpB of the YedE/YeeE family (TC#9.B.102).	Bacteria	*PmpC of Serratia sp.* (E7BBJ3)**

2.A.102.4.3	Hypothetical protein	Euryarchaea	HP of Methanococcus maripaludis

2.A.102.4.4	Hypothetical protein	Firmicutes	HP of Streptococcus pneumoniae

2.A.102.4.5	Putative sulfite exporter of 304 aas and 9 TMSs, TauE/SafE.	Proteobacteria	*TauE homologue of Thalassiobium* sp.**

2.A.102.4.6	Uncharacterized protein of the TauE family of 380 aas and 8 TMSs.	Proteobacteria	TauE homologue of Bradyrhizobium japonicum

2.A.102.4.7	DUF81 protein of 330 aas and 8 TMSs	Archaea	DUF81 protein of Halobacterium salinarum

2.A.102.4.8	*The putative sulfate transporter, CysZ (Rückert et al., 2005).*	Bacteria	CysZ of Corynebacterium glutamicum (Q8NLX4)

2.A.102.4.9	TauE (DUF81) homologue of 322 aas and 9 putative TMSs	Red Algae	TauE homologue of Galdieria sulphuraria

Examples:
TC#	Name	Organismal Type	Example
2.A.102.5.1	Orf of 590 aas and 12 TMSs	Plants	*Orf of Oryza sativa* (Q5ZAL4)**

2.A.102.5.2	Uncharacterized protein of 480 aas and 8 TMSs.		UP of Marchantia polymorpha

2.A.102.5.3	Uncharacterized protein of 10 - 12 TMSs in a 3 - 5 + 2 + 5 TMS arrangement.		UP of Giardia intestinalis

2.A.102.5.4	Putative sulfite exporter,TauE/SafE family protein, of 476 aas and 10 TMSs.		TauE homolog of Arabidopsis thaliana

Examples:
TC#	Name	Organismal Type	Example
2.A.102.6.1	Putative permease of 208 aas and 6 TMSs		Putative permease of Methylococcaceae bacterium HT3

2.A.102.6.2	Uncharacterized protein of 174 aas and 4 TMSs		UP of Strongylus vulgaris

2.A.102.6.3	Putative sulfite exporter TauE/SafE family protein of 333 aas and 11 TM		TauE homologue of Desulfobacterium autotrophicum

Examples:
TC#	Name	Organismal Type	Example
2.A.102.7.1	Putative sulfite exporter TauE/SafE family protein of 331 aas and 9 TMSs.		TauE homolog of Candidatus Filomicrobium marinum

2.A.102.7.2	Uncharacterized protein of 318 aas and 9 TMSs.		UP of Pseudomaricurvus alcaniphilus

2.A.102.7.3	Putative sulfite exporter TauE/SafE family protein of 307 aas and 9 TM		TauE homologue of Litorilinea aerophila