Rv2492 Resolved · medium

H37Rv Rv2492 · MTBC0 mtbc0_002654 · 250 aa · 2830168–2830920 (+) · RefSeq NP_217008.1

Annotation: from legacy to revised

Legacy (H37Rv / Mycobrowser)hypothetical protein
MTBC0 PGAP re-annotationhypothetical protein
Revised (this work)Thymidylate-synthase (TS)-superfamily (hydroxymethyl)transferase acting on a deoxyuridylate-type nucleotide, a distinct non-essential locus separate from the essential thymidylate synthases ThyA/ThyX. RefSeq leaves it 'hypothetical protein'. The model superposes on MilA (a CMP hydroxymethyltransferase, PDB 5b6d) with a complete catalytic constellation at the nucleotide (nucleophile Cys151, phosphate-binding Arg175, base-reading Asp178, ribose-binding His222/Tyr224); HHpred places it 100% in the thymidylate-synthase / dCMP-hydroxymethylase superfamily. The specificity residues reassign the substrate away from the template's cytidylate: Asn186 (vs MilA Asp186) and Ser176 (deoxy-type) point to a (deoxy)uridylate (dUMP-type) substrate with one-carbon chemistry on the pyrimidine C5. Vertically conserved through the human and animal MTBC and M. canettii (an earlier anti-phage-defence hypothesis is rejected). A structural prediction; substrate narrowed, not fixed.

Curated reference (UniProt)

UniProt I6YDJ7 TrEMBL · unreviewed · Predicted
UniProt nameThymidylate synthase

Conservation & selection (intra-MTBC, 145 209 strains)

pN/pS 0.705 · relaxed/neutral
Polymorphic sites (≥ 0.1% of strains) 4 synonymous, 10 missense, 0 nonsense, 0 frameshift

pN/pS from segregating SNPs (singletons removed) normalised by possible sites. Low pN/pS = purifying selection (a strong signal that a "hypothetical" is a real, constrained gene). A high pN/pS is ambiguous: relaxed constraint or positive selection (drug resistance, antigenic variation) inflate it; e.g. rpoB/katG/pncA score high here for resistance, not loss of function. A clonal disruption (one allele over a clade) suggests lineage pseudogenisation; a convergent one (many independent alleles) is typical of resistance loss-of-function.

Domains (Pfam, hmmscan --cut_ga)

No Pfam-A domain above the gathering threshold (or not yet scanned).

Structural neighbours (Foldseek on the ESMFold model, exploratory)

ESMFold model confidence: mean pLDDT 86.1 (confident). A confident model makes the fold comparison meaningful.

Best matches against the PDB, ranked by Foldseek homology probability. A high probability / TM-score suggests a shared fold; unless flagged sig (E < 0.01) these are fold hypotheses, not assignments.

TargetProbTME-valueDescription
5jnh-assembly1_B 1.00 0.73 3.7e-11 sig 5jnh-assembly1_B Crystal Structure of cytidine monophosphate hydroxymethylase MilA
5b6d-assembly1_A 1.00 0.71 4.2e-11 sig 5b6d-assembly1_A Crystal Structure of cytidine monophosphate hydroxymethylase MilA with CMP
5jnh-assembly1_A 1.00 0.63 2.3e-12 sig 5jnh-assembly1_A Crystal Structure of cytidine monophosphate hydroxymethylase MilA
5b6e-assembly1_A 1.00 0.70 6.3e-11 sig 5b6e-assembly1_A Crystal Structure of cytidine monophosphate hydroxymethylase MilA with hmCMP
5jp9-assembly1_A 1.00 0.65 1.0e-10 sig 5jp9-assembly1_A Crystal Structure of cytidine monophosphate hydroxymethylase MilA with dCMP
5j7w-assembly2_B 1.00 0.64 1.2e-10 sig 5j7w-assembly2_B Enterococcus faecalis thymidylate synthase complex with methotrexate
4xsd-assembly1_A 1.00 0.64 7.6e-11 sig 4xsd-assembly1_A Complex structure of thymidylate synthase from varicella zoster virus with a dUMP
4fog-assembly2_B 1.00 0.68 4.0e-10 sig 4fog-assembly2_B Crystal Structure of Mtb ThyA in Complex with 5-Fluoro-dUMP and 5-methyltetrahydrofolic acid

Functional interaction network (STRING v12, guilt-by-association)

Closest characterised functional partner: vapC38 (ribonuclease VapC38), medium confidence from genomic context alone (score 619 excluding text-mining). This association is the citable seed of a function hypothesis for this hypothetical protein.

PartnerProductScoreNo text-miningChannels (≥400)
Rv2491 hyp hypothetical protein 967 967 ctx neighborhood:882 coexpression:731
Rv3109 moaA1 cyclic pyranopterin monophosphate synthase 778 778 coexpression:778
Rv3114 hyp hypothetical protein 754 739 coexpression:730
Rv2763c dfrA dihydrofolate reductase 749 715 coexpression:699
Rv1407 fmu 16S rRNA m5C967 methyltransferase 720 708 coexpression:667
Rv2902c rnhB ribonuclease HII 717 705 coexpression:649
Rv0570 nrdZ vitamin B12-dependent ribonucleoside-diphosphate reductase 688 659 coexpression:647
Rv3051c nrdE ribonucleoside-diphosphate reductase subunit alpha 687 658 coexpression:646
Rv2116 lppK lipoprotein LppK 675 655 coexpression:598
Rv0002 dnaN DNA polymerase III subunit beta 675 654 coexpression:597
Rv2494 vapC38 ribonuclease VapC38 619 619 ctx neighborhood:619
Rv2493 vapB38 antitoxin VapB38 619 619 ctx neighborhood:619
Rv2697c dut deoxyuridine 5'-triphosphate nucleotidohydrolase 587 531 coexpression:443
Rv1389 gmk guanylate kinase 536 480 coexpression:479
Rv0038 hyp hypothetical protein 494 467 coexpression:467

STRING combines evidence channels (neighborhood, fusion, cooccurrence, coexpression, experimental, database, text-mining) into a 0–1000 score. The ctx badge marks edges carried by the genomic-context channels (conserved neighborhood, fusion, phylogenetic co-occurrence), which are independent of orthology and structure and the strongest signal for an unknown gene. The no text-mining column recomputes the score from data alone, so a link that does not depend on the literature is visible. Association is a function hypothesis, not proof: corroborate with the operon context and the primary literature before assigning a function.

Evidence

  • RefSeq: hypothetical protein
  • Superposition on MilA+CMP (5b6d); HHpred 100% thymidylate-synthase / dCMP-hydroxymethylase superfamily
  • Competent catalytic core: Cys151, Arg175, Asp178, His222, Tyr224 at H-bonding distance of the nucleotide
  • Specificity residues Asn186 + Ser176 -> (deoxy)uridylate (dUMP-type), not the template CMP
  • Distinct, non-essential locus (25% id); not redundant with ThyA/ThyX
  • Curated against the companion dark-enzymes re-annotation (Guyeux 2026)

Sources

  • Ancestral sequence & coordinates: Harrison LB et al. (2024), An imputed ancestral reference genome for the MTBC, doi:10.1101/2023.09.07.556366
  • Product annotation: NCBI PGAP on MTBC0; legacy from H37Rv NC_000962.3 (RefSeq NP_217008.1)
  • Domains: Pfam-A via hmmscan --cut_ga — none above threshold
  • Sequence-level signal: ESM Atlas (EvolutionaryScale × BioHub) — exploratory
  • Curated reference: UniProt I6YDJ7 (TrEMBL, unreviewed; Predicted)
  • Intra-MTBC selection: pN/pS and disruption from SPDI variants of 145 209 MTBC strains (this work, local collection vs H37Rv NC_000962.3)
  • Model confidence: ESMFold per-residue pLDDT (mean 86.1, confident)
  • Interaction network: STRING v12.0 (Szklarczyk et al. 2023, doi:10.1093/nar/gkac1000), taxon 83332, CC-BY 4.0 — 35 functional partner(s); context anchor vapC38
  • Primary literature: Guyeux C (2026). Structure-guided functional hypotheses for uncharacterised enzymes of Mycobacterium tuberculosis in preparation. doi:10.5281/zenodo.20571950

Ancestral MTBC0 protein sequence

>mtbc0_002654|Rv2492|
MSRRIINEFGVQIYGATIGDTWAGLVRAVLDLGSQCFDEDRERIALSNVRIKSSVQNYPDLTIEEHCNSDQLKAMLDFMFNTDTMEDIDVVKSFSRGAKSYHRRIKEGRMIEFVIERLSLIPESKKAVVVFPTYEDYAAVMRNHRDDYLPCLVSIQFRLLPDGKDYVFHTTFYSRSMDAWQKGHGNLLSIAKLSDWVRENVSARIGRKIMLGPLDGMICDVHIYKETYAEACKRLANLDLRRTQFDAVRN