Αr14 RNA: Difference between revisions
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==Discovery and Structure== |
==Discovery and Structure== |
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Smr14C2 sRNA was described by del Val et al.,<ref name="Val07">{{cite journal | vauthors = del Val C, Rivas E, Torres-Quesada O, Toro N, Jiménez-Zurdo JI | title = Identification of differentially expressed small non-coding RNAs in the legume endosymbiont Sinorhizobium meliloti by comparative genomics | journal = Molecular Microbiology | volume = 66 | issue = 5 | pages = 1080–91 | date = December 2007 | pmid = 17971083 | pmc = 2780559 | doi = 10.1111/j.1365-2958.2007.05978.x }}</ref> as a result of a computational comparative genomic approach consisting in the integration of complementary strategies, designed to search for novel sRNA-encoding genes in the intergenic regions (IGRs) of the reference ''S. meliloti'' 1021 strain (http://iant.toulouse.inra.fr/bacteria/annotation/cgi/rhime.cgi) . Northern hybridization experiments confirmed that the predicted ''smr14C2'' locus did express a single transcript of the expected size, which accumulated differentially in free-living and endosymbiotic bacteria. TAP-based 5’-RACE experiments mapped the transcription start site (TSS) of the full-length Smr14C transcript to the 1,667,613 nt position in the S. meliloti 1021 genome (http://iant.toulouse.inra.fr/bacteria/annotation/cgi/rhime.cgi) whereas the 3’-end was assumed to be located at the 1,667,491 nt position matching the last residue of the consecutive stretch of Us of a bona fide Rho-independent terminator (Figure 5). Parallel and later studies<ref name="ulve">{{cite journal | vauthors = Ulvé VM, Sevin EW, Chéron A, Barloy-Hubler F | title = Identification of chromosomal alpha-proteobacterial small RNAs by comparative genome analysis and detection in Sinorhizobium meliloti strain 1021 | journal = BMC Genomics | volume = 8 | issue = 467 | pages = 467 | date = December 2007 | pmid = 18093320 | pmc = 2245857 | doi = 10.1186/1471-2164-8-467 }}</ref><ref name="valverde">{{cite journal | vauthors = Valverde C, Livny J, Schlüter JP, Reinkensmeier J, Becker A, Parisi G | title = Prediction of Sinorhizobium meliloti sRNA genes and experimental detection in strain 2011 | journal = BMC Genomics | volume = 9 | issue = 406 | pages = 416 | date = September 2008 | pmid = 18793445 | doi = 10.1186/1471-2164-9-416 }}</ref> in which Smr14C2 transcript is referred to as sra38 or Sm7', independently confirmed the expression this sRNA in ''S. melilloti'' and in its closely related strain 2011. Recent deep sequencing-based characterization of the small RNA fraction (50-350 nt) of ''S. meliloti'' 2011 further confirmed the expression of Smr14C2 (here referred to as SmelC397), and mapped the 5’- and 3´-ends of the full-length transcript to the same position in the ''S. meliloti'' 1021 genome.<ref name="schluter">{{cite journal | vauthors = Córdoba JM, Chavarro C, Schlueter JA, Jackson SA, Blair MW | title = Integration of physical and genetic maps of common bean through BAC-derived microsatellite markers | journal = BMC Genomics | volume = 11 | issue = 245 | pages = 436 | date = July 2010 | pmid = 20637113 | doi = 10.1186/1471-2164-11-436 }}</ref> |
Smr14C2 sRNA was described by del Val et al.,<ref name="Val07">{{cite journal | vauthors = del Val C, Rivas E, Torres-Quesada O, Toro N, Jiménez-Zurdo JI | title = Identification of differentially expressed small non-coding RNAs in the legume endosymbiont Sinorhizobium meliloti by comparative genomics | journal = Molecular Microbiology | volume = 66 | issue = 5 | pages = 1080–91 | date = December 2007 | pmid = 17971083 | pmc = 2780559 | doi = 10.1111/j.1365-2958.2007.05978.x }}</ref> as a result of a computational comparative genomic approach consisting in the integration of complementary strategies, designed to search for novel sRNA-encoding genes in the intergenic regions (IGRs) of the reference ''S. meliloti'' 1021 strain (http://iant.toulouse.inra.fr/bacteria/annotation/cgi/rhime.cgi) . Northern hybridization experiments confirmed that the predicted ''smr14C2'' locus did express a single transcript of the expected size, which accumulated differentially in free-living and endosymbiotic bacteria. TAP-based 5’-RACE experiments mapped the transcription start site (TSS) of the full-length Smr14C transcript to the 1,667,613 nt position in the S. meliloti 1021 genome (http://iant.toulouse.inra.fr/bacteria/annotation/cgi/rhime.cgi) whereas the 3’-end was assumed to be located at the 1,667,491 nt position matching the last residue of the consecutive stretch of Us of a bona fide Rho-independent terminator (Figure 5). Parallel and later studies<ref name="ulve">{{cite journal | vauthors = Ulvé VM, Sevin EW, Chéron A, Barloy-Hubler F | title = Identification of chromosomal alpha-proteobacterial small RNAs by comparative genome analysis and detection in Sinorhizobium meliloti strain 1021 | journal = BMC Genomics | volume = 8 | issue = 467 | pages = 467 | date = December 2007 | pmid = 18093320 | pmc = 2245857 | doi = 10.1186/1471-2164-8-467 }}</ref><ref name="valverde">{{cite journal | vauthors = Valverde C, Livny J, Schlüter JP, Reinkensmeier J, Becker A, Parisi G | title = Prediction of Sinorhizobium meliloti sRNA genes and experimental detection in strain 2011 | journal = BMC Genomics | volume = 9 | issue = 406 | pages = 416 | date = September 2008 | pmid = 18793445 | pmc = 2573895 | doi = 10.1186/1471-2164-9-416 }}</ref> in which Smr14C2 transcript is referred to as sra38 or Sm7', independently confirmed the expression this sRNA in ''S. melilloti'' and in its closely related strain 2011. Recent deep sequencing-based characterization of the small RNA fraction (50-350 nt) of ''S. meliloti'' 2011 further confirmed the expression of Smr14C2 (here referred to as SmelC397), and mapped the 5’- and 3´-ends of the full-length transcript to the same position in the ''S. meliloti'' 1021 genome.<ref name="schluter">{{cite journal | vauthors = Córdoba JM, Chavarro C, Schlueter JA, Jackson SA, Blair MW | title = Integration of physical and genetic maps of common bean through BAC-derived microsatellite markers | journal = BMC Genomics | volume = 11 | issue = 245 | pages = 436 | date = July 2010 | pmid = 20637113 | pmc = 3091635 | doi = 10.1186/1471-2164-11-436 }}</ref> |
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[[File:C14 annotated alignment structure conservation.png|thumb|center|1001px|Figure 1: Consensus secondary structure of Smr14C2 and the ar14 family predicted by RNA<ref name="RNAfold">{{cite journal|author=I. L. Hofacker, W. Fontana, P. F. Stadler, L. S. Bonhoeffer, M. Tacker and P. Schuster|title=Fast folding and comparison of RNA secondary structures| volume=125|issue=2|pages=167–188|doi=10.1007/BF00818163|journal= |
[[File:C14 annotated alignment structure conservation.png|thumb|center|1001px|Figure 1: Consensus secondary structure of Smr14C2 and the ar14 family predicted by RNA<ref name="RNAfold">{{cite journal|author=I. L. Hofacker, W. Fontana, P. F. Stadler, L. S. Bonhoeffer, M. Tacker and P. Schuster|title=Fast folding and comparison of RNA secondary structures| volume=125|issue=2|pages=167–188|doi=10.1007/BF00818163|journal=Monatshefte für Chem|year=1994}}</ref> and RNAalifold.<ref name="RNAalifold">{{cite journal | vauthors = Bernhart SH, Hofacker IL, Will S, Gruber AR, Stadler PF | title = RNAalifold: improved consensus structure prediction for RNA alignments | journal = BMC Bioinformatics | volume = 9 | issue = 474 | pages = 474 | date = November 2008 | pmid = 19014431 | pmc = 2621365 | doi = 10.1186/1471-2105-9-474 }}</ref> The color scheme represent the base pair probabilities.]] |
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The nucleotide sequence of Smr14C2 was initially used as query to search against the Rfam database (version 10.0;http://www.sanger.ac.uk/Software/Rfam). This homology search rendered no matches to known bacterial sRNA in this database. Smr14C2 was next BLASTed with default parameters against all the currently available bacterial genomes (1,615 sequences at 20 April 2011; https://www.ncbi.nlm.nih.gov;). The regions exhibiting significant homology to the query sequence (78-89% similarity) were extracted to create a Covariance Model (CM) from a seed alignment using Infernal (version1.0)<ref name="infernal">{{cite journal | vauthors = Nawrocki EP, Kolbe DL, Eddy SR | title = Infernal 1.0: inference of RNA alignments | journal = Bioinformatics | volume = 25 | issue = 10 | pages = 1335–7 | date = May 2009 | pmid = 19307242 | pmc = 2732312 | doi = 10.1093/bioinformatics/btp157 }}</ref> (Figure 2). This CM was used in a further search for new members of the αr9 family in the existing bacterial genomic databases and resumed in the following Table 2: |
The nucleotide sequence of Smr14C2 was initially used as query to search against the Rfam database (version 10.0;http://www.sanger.ac.uk/Software/Rfam). This homology search rendered no matches to known bacterial sRNA in this database. Smr14C2 was next BLASTed with default parameters against all the currently available bacterial genomes (1,615 sequences at 20 April 2011; https://www.ncbi.nlm.nih.gov;). The regions exhibiting significant homology to the query sequence (78-89% similarity) were extracted to create a Covariance Model (CM) from a seed alignment using Infernal (version1.0)<ref name="infernal">{{cite journal | vauthors = Nawrocki EP, Kolbe DL, Eddy SR | title = Infernal 1.0: inference of RNA alignments | journal = Bioinformatics | volume = 25 | issue = 10 | pages = 1335–7 | date = May 2009 | pmid = 19307242 | pmc = 2732312 | doi = 10.1093/bioinformatics/btp157 }}</ref> (Figure 2). This CM was used in a further search for new members of the αr9 family in the existing bacterial genomic databases and resumed in the following Table 2: |
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The results were manually inspected to deduce a consensus secondary structure for the family (Figure 1 and Figure 2). The consensus structure was also independently predicted with the program locARNATE<ref name="locarnate">{{cite journal |authors=Will S, Reiche K, Hofacker IL, Stadler PF, Backofen R|title=Inferring Noncoding RNA Families and Classes by Means of Genome-Scale Structure-Based Clustering|journal=PLoS Comput Biol|volume=4|issue=65|doi= |
The results were manually inspected to deduce a consensus secondary structure for the family (Figure 1 and Figure 2). The consensus structure was also independently predicted with the program locARNATE<ref name="locarnate">{{cite journal |authors=Will S, Reiche K, Hofacker IL, Stadler PF, Backofen R|title=Inferring Noncoding RNA Families and Classes by Means of Genome-Scale Structure-Based Clustering|journal=PLoS Comput Biol|volume=4|issue=65|pages=e65|doi=10.1371/journal.pcbi.0030065|pmid=17432929|pmc=1851984|year=2007}}</ref> with very similar predictions. The manual inspection of the sequences found with the CM using Infernal allowed finding 101 homolog sequences. The rhizobial species encoding the 36 closer homologs to Smr14C2 were: ''S. medicae'' and ''S. fredii'', two ''R. leguminosarum'' trifolii strains (WSM2304 and WSM1325), two ''R. etli'' strains CFN 42 and CIAT 652, the reference ''R. leguminosarum'' bv. viciae 3841 strain, and the ''Agrobacterium'' species ''A. vitis'',''A. tumefaciens'', ''A. radiobacter'' and ''A. H13''. All these sequences showed significant Infernal E-values (5.63E-29 - 8.16E-18) and bit-scores. The rest of the sequences found with the model showed high E-values between (1.33E-17 and 8.79E-03) but lower bit-scores and are encoded by ''Brucella'' species (''B. ovis'', ''B. canis'', ''B. abortus'', ''B. microtis'', and several biobars of ''B. melitensis''), ''Ochrobactrum anthropi'' and the ''Mesorhizobum'' species ''loti'', ''M. ciceri'' and ''M.'' BNC. |
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The model identified five additional copies in the ''S. meliloti'' genome, locating two of them in tandem in the same IGR (Smr14C2, Smr14C3). The same results were obtained for ''S. medicae'' (Smedr14C1, Smedr14C2), ''S. fredii'' (Sfr14C1, Sfr14C2), ''M. loti'' (Mlr14C1, Mlr14C2) and ''M. ciceri'' (Mcr14C1, Mcr14C2). |
The model identified five additional copies in the ''S. meliloti'' genome, locating two of them in tandem in the same IGR (Smr14C2, Smr14C3). The same results were obtained for ''S. medicae'' (Smedr14C1, Smedr14C2), ''S. fredii'' (Sfr14C1, Sfr14C2), ''M. loti'' (Mlr14C1, Mlr14C2) and ''M. ciceri'' (Mcr14C1, Mcr14C2). |
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==Promoter Analysis== |
==Promoter Analysis== |
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All the promoter regions of the αr14 family members examined so far are very conserved in a sequence stretch extending up to 120 bp upstream of the transcription start site of the sRNA. All closest homolog loci have recognizable σ70-dependent promoters showing a -35/-10 consensus motif CTTAGAC-n17-CTATAT, which has been previously shown to be widely conserved among several other genera in the α-subgroup of proteobacteria.<ref name="6r6">{{cite journal | vauthors = MacLellan SR, MacLean AM, Finan TM | title = Promoter prediction in the rhizobia | journal = Microbiology | volume = 152 | issue = Pt 6 | pages = 1751–63 | date = June 2006 | pmid = 16735738 | doi = 10.1099/mic.0.28743-0 }}</ref> To identify binding sites for other known transcription factors we used the fasta sequences provided by RegPredict<ref name="regpredict">{{cite journal | vauthors = Novichkov PS, Rodionov DA, Stavrovskaya ED, Novichkova ES, Kazakov AE, Gelfand MS, Arkin AP, Mironov AA, Dubchak I | title = RegPredict: an integrated system for regulon inference in prokaryotes by comparative genomics approach | journal = Nucleic Acids Research | volume = 38 | issue = Web Server issue | pages = |
All the promoter regions of the αr14 family members examined so far are very conserved in a sequence stretch extending up to 120 bp upstream of the transcription start site of the sRNA. All closest homolog loci have recognizable σ70-dependent promoters showing a -35/-10 consensus motif CTTAGAC-n17-CTATAT, which has been previously shown to be widely conserved among several other genera in the α-subgroup of proteobacteria.<ref name="6r6">{{cite journal | vauthors = MacLellan SR, MacLean AM, Finan TM | title = Promoter prediction in the rhizobia | journal = Microbiology | volume = 152 | issue = Pt 6 | pages = 1751–63 | date = June 2006 | pmid = 16735738 | doi = 10.1099/mic.0.28743-0 }}</ref> To identify binding sites for other known transcription factors we used the fasta sequences provided by RegPredict<ref name="regpredict">{{cite journal | vauthors = Novichkov PS, Rodionov DA, Stavrovskaya ED, Novichkova ES, Kazakov AE, Gelfand MS, Arkin AP, Mironov AA, Dubchak I | title = RegPredict: an integrated system for regulon inference in prokaryotes by comparative genomics approach | journal = Nucleic Acids Research | volume = 38 | issue = Web Server issue | pages = W299–307 | date = July 2010 | pmid = 20542910 | pmc = 2896116 | doi = 10.1093/nar/gkq531 }}</ref>(http://regpredict.lbl.gov/regpredict/help.html), and used those position weight matrices (PSWM) provided by RegulonDB<ref name="regulondb">{{cite journal | vauthors = Gama-Castro S, Salgado H, Peralta-Gil M, Santos-Zavaleta A, Muñiz-Rascado L, Solano-Lira H, Jimenez-Jacinto V, Weiss V, García-Sotelo JS, López-Fuentes A, Porrón-Sotelo L, Alquicira-Hernández S, Medina-Rivera A, Martínez-Flores I, Alquicira-Hernández K, Martínez-Adame R, Bonavides-Martínez C, Miranda-Ríos J, Huerta AM, Mendoza-Vargas A, Collado-Torres L, Taboada B, Vega-Alvarado L, Olvera M, Olvera L, Grande R, Morett E, Collado-Vides J | title = RegulonDB version 7.0: transcriptional regulation of Escherichia coli K-12 integrated within genetic sensory response units (Gensor Units) | journal = Nucleic Acids Research | volume = 39 | issue = Database issue | pages = D98–105 | date = January 2011 | pmid = 21051347 | pmc = 3013702 | doi = 10.1093/nar/gkq1110 }}</ref> (http://regulondb.ccg.unam.mx). We built PSWM for each transcription factor from the RegPredict sequences using the Consensus/Patser program, choosing the best final matrix for motif lengths between 14–30 {{Not a typo|bps}} a threshold average E-value < 10E-10 for each matrix was established, (see "Thresholded consensus" in http://gps-tools2.its.yale.edu). Moreover, we searched for conserved unknown motifs using MEME<ref name="meme">{{cite journal |vauthors=Bailey TL, Elkan C |title=Fitting a mixture model by expectation maximization to discover motifs in biopolymers|journal=Proceedings of the Second International Conference on Intelligent Systems for Molecular Biology| pages=28–36|publisher=AAAI Press, Menlo Park, California|year=1994}}</ref> (http://meme.sdsc.edu/meme4_6_1/intro.html) and used relaxed regular expressions (i.e. pattern matching) over all Smr14C2 homologs promoters. |
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This studies revealed two well defined groups of loci, the first one represented by the closest homologs (Figure 5) that presented a 26 bp long region very conserved between positions -40 and -75, marked as conserved MEME motif in (Figure 5), but no significant similarity to known transcription factor binding sites matrices could be established. A group of not so closely related members of the αr14 family constituted the second group of conserved promoters (Figure6). They presented a different promoter region, very well conserved across all members and an additional unknown 20 bp motif. |
This studies revealed two well defined groups of loci, the first one represented by the closest homologs (Figure 5) that presented a 26 bp long region very conserved between positions -40 and -75, marked as conserved MEME motif in (Figure 5), but no significant similarity to known transcription factor binding sites matrices could be established. A group of not so closely related members of the αr14 family constituted the second group of conserved promoters (Figure6). They presented a different promoter region, very well conserved across all members and an additional unknown 20 bp motif. |
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[[File:Promoters14C.png|thumb|center|1001px|Figure 5: Graphic representation of the αr14 seed members' promoter region. All members presented putative σ70 promoters with -30 and -10 boxes marked in green and red respectively.]] |
[[File:Promoters14C.png|thumb|center|1001px|Figure 5: Graphic representation of the αr14 seed members' promoter region. All members presented putative σ70 promoters with -30 and -10 boxes marked in green and red respectively.]] |
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==Genomic Context== |
==Genomic Context== |
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Most of the members of the αr14 family are trans-encoded sRNAs transcribed from independent promoters in chromosomal IGRs. Many of the neighboring genes of the seed alignment’s members were not annotated and thus were further manually curated.<ref name="gopet">{{cite journal | vauthors = Vinayagam A, del Val C, Schubert F, Eils R, Glatting KH, Suhai S, König R | title = GOPET: a tool for automated predictions of Gene Ontology terms | journal = BMC Bioinformatics | volume = 7 | pages = 161 | date = March 2006 | pmid = 16549020 | pmc = 1434778 | doi = 10.1186/1471-2105-7-161 }}</ref><ref name="blast2go">{{cite journal | vauthors = Conesa A, Götz S, García-Gómez JM, Terol J, Talón M, Robles M | title = Blast2GO: a universal tool for annotation, visualization and analysis in functional genomics research | journal = Bioinformatics | volume = 21 | issue = 18 | pages = 3674–6 | date = September 2005 | pmid = 16081474 | doi = 10.1093/bioinformatics/bti610 }}</ref><ref name="domainsweep">{{cite journal | vauthors = del Val C, Ernst P, Falkenhahn M, Fladerer C, Glatting KH, Suhai S, Hotz-Wagenblatt A | title = ProtSweep, 2Dsweep and DomainSweep: protein analysis suite at DKFZ | journal = Nucleic Acids Research | volume = 35 | issue = Web Server issue | pages = |
Most of the members of the αr14 family are trans-encoded sRNAs transcribed from independent promoters in chromosomal IGRs. Many of the neighboring genes of the seed alignment’s members were not annotated and thus were further manually curated.<ref name="gopet">{{cite journal | vauthors = Vinayagam A, del Val C, Schubert F, Eils R, Glatting KH, Suhai S, König R | title = GOPET: a tool for automated predictions of Gene Ontology terms | journal = BMC Bioinformatics | volume = 7 | pages = 161 | date = March 2006 | pmid = 16549020 | pmc = 1434778 | doi = 10.1186/1471-2105-7-161 }}</ref><ref name="blast2go">{{cite journal | vauthors = Conesa A, Götz S, García-Gómez JM, Terol J, Talón M, Robles M | title = Blast2GO: a universal tool for annotation, visualization and analysis in functional genomics research | journal = Bioinformatics | volume = 21 | issue = 18 | pages = 3674–6 | date = September 2005 | pmid = 16081474 | doi = 10.1093/bioinformatics/bti610 }}</ref><ref name="domainsweep">{{cite journal | vauthors = del Val C, Ernst P, Falkenhahn M, Fladerer C, Glatting KH, Suhai S, Hotz-Wagenblatt A | title = ProtSweep, 2Dsweep and DomainSweep: protein analysis suite at DKFZ | journal = Nucleic Acids Research | volume = 35 | issue = Web Server issue | pages = W444–50 | date = July 2007 | pmid = 17526514 | pmc = 1933246 | doi = 10.1093/nar/gkm364 }}</ref> |
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The tandem copies in the Sinorhizobium group’s genomes (Smr14C2, Smr14C3, Smedr14C1, Smedr14C2, Sfr14C1, Sfr14C2) presented a conserved genomic context. The gene upstream coded for an unknown protein containing the domain DUF1127 and the gene downstream coded for a trigger factor. |
The tandem copies in the Sinorhizobium group’s genomes (Smr14C2, Smr14C3, Smedr14C1, Smedr14C2, Sfr14C1, Sfr14C2) presented a conserved genomic context. The gene upstream coded for an unknown protein containing the domain DUF1127 and the gene downstream coded for a trigger factor. |
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Revision as of 09:25, 19 February 2019
αr14 is a family of bacterial small non-coding RNAs with representatives in a broad group of α-proteobacteria. The first member of this family (Smr14C2) was found in a Sinorhizobium meliloti 1021 locus located in the chromosome (C). It was later renamed NfeR1 (Nodule Formation Efficiency RNA) and shown to be highly expressed in salt stress and during the symbiotic interaction on legume roots.[1] Further homology and structure conservation analysis identified 2 other chromosomal copies and 3 plasmidic ones. Moreover, full-length Smr14C homologs have been identified in several nitrogen-fixing symbiotic rhizobia (i.e. R. leguminosarum bv.viciae, R. leguminosarum bv. trifolii, R. etli, and several Mesorhizobium species), in the plant pathogens belonging to Agrobacterium species (i.e. A. tumefaciens, A. vitis, A. radiobacter, and Agrobacterium H13) as well as in a broad spectrum of Brucella species (B. ovis, B. canis, B. abortus and B. microtis, and several viobars of B. melitensis). αr14C RNA species are 115-125 nt long (Table 1) and share a well defined common secondary structure (Figure 1). Most of the αr14 transcripts can be catalogued as trans-acting sRNAs expressed from well-defined promoter regions of independent transcription units within intergenic regions (IGRs) of the α-proteobacterial genomes (Figure 5).
CM model | Name | GI accession number | begin | end | strand | %GC | length | Organism |
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αr14 | Smr14C1 | gi|15963753|ref|NC_003047.1| | 206862 | 206980 | - | 63 | 119 | Sinorhizobium meliloti 1021 |
αr14 | Smr14C2 | gi|15963753|ref|NC_003047.1| | 1667491 | 1667613 | - | 65 | 123 | Sinorhizobium meliloti 1021 |
αr14 | Smr14C3 | gi|15963753|ref|NC_003047.1| | 1667769 | 1667891 | - | 62 | 123 | Sinorhizobium meliloti 1021 |
αr14 | Smr14A1 | gi|16262453|ref|NC_003037.1| | 1220690 | 1220808 | + | 62 | 119 | Sinorhizobium meliloti 1021 plasmid pSymA |
αr14 | Smr14A2 | gi|16262453|ref|NC_003037.1| | 1328176 | 1328301 | - | 60 | 126 | Sinorhizobium meliloti 1021 plasmid pSymA |
αr14 | Smr14B | gi|16263748|ref|NC_003078.1| | 1605826 | 1605943 | + | 64 | 118 | Sinorhizobium meliloti 1021 plasmid pSymB |
αr14 | Smedr14C1 | gi|150395228|ref|NC_009636.1| | 1258038 | 1258160 | - | 63 | 123 | Sinorhizobium medicae WSM419 chromosome |
αr14 | Smedr14C2 | gi|150395228|ref|NC_009636.1| | 1258311 | 1258431 | - | 65 | 121 | Sinorhizobium medicae WSM419 chromosome |
αr14 | Smedr14C3 | gi|150395228|ref|NC_009636.1| | 3590299 | 3590417 | - | 63 | 119 | Sinorhizobium medicae WSM419 chromosome |
αr14 | Smedr14p01 | gi|150375726|ref|NC_009620.1| | 679043 | 679160 | - | 66 | 118 | Sinorhizobium medicae WSM419 plasmid pSMED01 |
αr14 | Smedr14p021 | gi|150377168|ref|NC_009621.1| | 316304 | 316428 | + | 64 | 125 | Sinorhizobium medicae WSM419 plasmid pSMED02 |
αr14 | Smedr14p022 | gi|150377168|ref|NC_009622.1| | 338866 | 338990 | + | 63 | 125 | Sinorhizobium medicae WSM419 plasmid pSMED02 |
αr14 | Sfr14C1 | gi|227820587|ref|NC_012587.1| | 1434171 | 1434293 | - | 64 | 123 | Sinorhizobium fredii NGR234 chromosome |
αr14 | Sfr14C2 | gi|227820587|ref|NC_012587.1| | 1434463 | 1434586 | - | 60 | 124 | Sinorhizobium fredii NGR234 chromosome |
αr14 | Sfr14C3 | gi|227820587|ref|NC_012587.1| | 3720579 | 3720700 | + | 66 | 122 | Sinorhizobium fredii NGR234 chromosome |
αr14 | Sfr14b | gi|227818258|ref|NC_012586.1| | 2205311 | 2205428 | + | 64 | 118 | Sinorhizobium fredii NGR234 plasmid pNGR234b |
αr14 | Atr14C1 | gi|159184118|ref|NC_003062.2| | 109473 | 109601 | + | 59 | 129 | Agrobacterium tumefaciens str. C58 chromosome circular |
αr14 | Atr14C2 | gi|159185562|ref|NC_003063.2| | 1230297 | 1230426 | + | 60 | 130 | Agrobacterium tumefaciens str. C58 chromosome linear |
αr14 | Atr14C3 | gi|159185562|ref|NC_003063.2| | 1831483 | 1831614 | + | 57 | 132 | Agrobacterium tumefaciens str. C58 chromosome linear |
αr14 | AH13r14C1 | gi|325291453|ref|NC_015183.1| | 109372 | 109501 | + | 58 | 130 | Agrobacterium sp. H13-3 chromosome |
αr14 | ReCIATr14C1 | gi|190889639|ref|NC_010994.1| | 513993 | 514114 | + | 60 | 122 | Rhizobium etli CIAT 652 |
αr14 | ReCIATr14C2 | gi|190889639|ref|NC_010994.1| | 1173512 | 1173639 | + | 59 | 128 | Rhizobium etli CIAT 652 |
αr14 | ReCIATr14C3 | gi|190889639|ref|NC_010994.1| | 1912478 | 1912597 | - | 64 | 120 | Rhizobium etli CIAT 652 |
αr14 | ReCIATr14C4 | gi|190889639|ref|NC_010994.1| | 4444894 | 4445016 | - | 55 | 123 | Rhizobium etli CIAT 652 |
αr14 | ReCIATr14A | gi|190895317|ref|NC_010998.1| | 345900 | 346021 | - | 63 | 122 | Rhizobium etli CIAT 652 plasmid pA |
αr14 | ReCIATr14B | gi|190893983|ref|NC_010996.1| | 112292 | 112425 | + | 61 | 134 | Rhizobium etli CIAT 652 plasmid pB |
αr14 | Arr14CI1 | gi|222084201|ref|NC_011985.1| | 597903 | 598030 | + | 61 | 128 | Agrobacterium radiobacter K84 chromosome 1 |
αr14 | Arr14CI2 | gi|222084201|ref|NC_011985.1| | 1693278 | 1693409 | - | 60 | 132 | Agrobacterium radiobacter K84 chromosome 1 |
αr14 | Arr14CI3 | gi|222084201|ref|NC_011985.1| | 3945511 | 3945639 | - | 59 | 129 | Agrobacterium radiobacter K84 chromosome 1 |
αr14 | Arr14CII1 | gi|222080781|ref|NC_011983.1| | 957687 | 957817 | + | 58 | 131 | Agrobacterium radiobacter K84 chromosome 2 |
αr14 | Arr14CII2 | gi|222080781|ref|NC_011983.1| | 1921110 | 1921242 | - | 61 | 133 | Agrobacterium radiobacter K84 chromosome 2 |
αr14 | Arr14Atc | gi|222101962|ref|NC_011987.1| | 138495 | 138625 | - | 60 | 131 | Agrobacterium radiobacter K84 plasmid pAtK84c |
αr14 | Rlt2304r14C1 | gi|209547612|ref|NC_011369.1| | 98050 | 98171 | + | 60 | 122 | Rhizobium leguminosarum bv. trifolii WSM2304 chromosome |
αr14 | Rlt2304r14C2 | gi|209547612|ref|NC_011369.1| | 1534501 | 1534621 | - | 63 | 121 | Rhizobium leguminosarum bv. trifolii WSM2304 chromosome |
αr14 | Rlt2304r14C3 | gi|209547612|ref|NC_011369.1| | 4019326 | 4019446 | - | 57 | 121 | Rhizobium leguminosarum bv. trifolii WSM2304 chromosome |
αr14 | Rlt2304r14p011 | gi|209546450|ref|NC_011368.1| | 306339 | 306454 | - | 70 | 116 | Rhizobium leguminosarum bv trifolii WSM2304 plasmid pRLG201 |
αr14 | Rlt2304r14p012 | gi|209546450|ref|NC_011368.1| | 607636 | 607759 | + | 62 | 124 | Rhizobium leguminosarum bv trifolii WSM2304 plasmid pRLG201 |
αr14 | Rlt2304r14p02 | gi|209545999|ref|NC_011366.1| | 50814 | 50936 | + | 63 | 123 | Rhizobium leguminosarum bv trifolii WSM2304 plasmid pRLG202 |
αr14 | Avr14CI1 | gi|222147015|ref|NC_011989.1| | 115126 | 115257 | + | 58 | 132 | Agrobacterium vitis S4 chromosome 1 |
αr14 | Avr14CI2 | gi|222147015|ref|NC_011989.1| | 2043913 | 2044044 | - | 64 | 132 | Agrobacterium vitis S4 chromosome 1 |
αr14 | Avr14CI3 | gi|222147015|ref|NC_011989.1| | 2793803 | 2793939 | + | 63 | 137 | Agrobacterium vitis S4 chromosome 1 |
αr14 | Avr14CII1 | gi|222106127|ref|NC_011988.1| | 598426 | 598556 | + | 62 | 131 | Agrobacterium vitis S4 chromosome 2 |
αr14 | Avr14CII2 | gi|222106127|ref|NC_011988.1| | 901035 | 901159 | - | 63 | 125 | Agrobacterium vitis S4 chromosome 2 |
αr14 | Avr14Atc | gi|222083145|ref|NC_011984.1| | 7318 | 7444 | + | 61 | 127 | Agrobacterium vitis S4 plasmid pAtS4c |
αr14 | Rlvr14C1 | gi|116249766|ref|NC_008380.1| | 512658 | 512779 | + | 59 | 122 | Rhizobium leguminosarum bv. viciae 3841 |
αr14 | Rlvr14C2 | gi|116249766|ref|NC_008380.1| | 2174956 | 2175074 | - | 65 | 119 | Rhizobium leguminosarum bv. viciae 3841 |
αr14 | Rlvr14C3 | gi|116249766|ref|NC_008380.1| | 4988030 | 4988158 | - | 58 | 129 | Rhizobium leguminosarum bv. viciae 3841 |
αr14 | Rlvr14p11 | gi|116255200|ref|NC_008384.1| | 574471 | 574593 | - | 59 | 123 | Rhizobium leguminosarum bv. viciae 384 plasmid pRL11 |
αr14 | Rlvr14p12 | gi|116248676|ref|NC_008378.1| | 407202 | 407316 | + | 65 | 115 | Rhizobium leguminosarum bv. viciae 384 plasmid pRL12 |
αr14 | Rlt1325r14C1 | gi|241202755|ref|NC_012850.1| | 107063 | 107184 | + | 58 | 122 | Rhizobium leguminosarum bv. trifolii WSM1325 |
αr14 | Rlt1325r14C2 | gi|241202755|ref|NC_012850.1| | 1697038 | 1697156 | - | 63 | 119 | Rhizobium leguminosarum bv. trifolii WSM1325 |
αr14 | Rlt1325r14C3 | gi|241202755|ref|NC_012850.1| | 4296632 | 4296749 | - | 58 | 118 | Rhizobium leguminosarum bv. trifolii WSM1325 |
αr14 | Rlt1325r14p012 | gi|241113003|ref|NC_012848.1| | 265945 | 266069 | + | 62 | 125 | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132501 |
αr14 | Rlt1325r14p011 | gi|241113003|ref|NC_012848.1| | 112604 | 112728 | + | 66 | 125 | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132501 |
αr14 | Rlt1325r14p02 | gi|241666492|ref|NC_012858.1| | 28646 | 28767 | + | 60 | 122 | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132502 |
αr14 | Rlt1325r14p04 | gi|241258599|ref|NC_012852.1| | 119314 | 119443 | + | 59 | 130 | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132504 |
αr14 | Rlt1325r14p05 | gi|241554070|ref|NC_012854.1| | 2286 | 2409 | - | 65 | 124 | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132505 |
αr14 | ReCFNr14C1 | gi|86355669|ref|NC_007761.1| | 462566 | 462687 | - | 58 | 122 | Rhizobium etli CFN 42 |
αr14 | ReCFNr14C2 | gi|86355669|ref|NC_007761.1| | 1926239 | 1926358 | + | 65 | 120 | Rhizobium etli CFN 42 |
αr14 | ReCFNr14C3 | gi|86355669|ref|NC_007761.1| | 4314384 | 4314505 | + | 58 | 122 | Rhizobium etli CFN 42 |
αr14 | ReCFNr14e | gi|86360278|ref|NC_007765.1| | 443001 | 443129 | + | 64 | 129 | Rhizobium etli CFN 42 plasmid p42e |
αr14 | ReCFNr14f | gi|86360734|ref|NC_007766.1| | 22818 | 22935 | - | 64 | 118 | Rhizobium etli CFN 42 plasmid p42f |
αr14 | ReCFNr14d | gi|89255298|ref|NC_004041.2| | 77668 | 77794 | - | 61 | 127 | Rhizobium etli CFN 42 symbiotic plasmid p42d |
αr14 | Mlr14C1 | gi|57165207|ref|NC_002678.2| | 890582 | 890702 | + | 68 | 121 | Mesorhizobium loti MAFF303099 chromosome |
αr14 | Mlr14C2 | gi|57165207|ref|NC_002678.2| | 890760 | 890880 | + | 64 | 121 | Mesorhizobium loti MAFF303099 chromosome |
αr14 | Mlr14C3 | gi|57165207|ref|NC_002678.2| | 890941 | 891061 | + | 63 | 121 | Mesorhizobium loti MAFF303099 chromosome |
αr14 | Mlr14C4 | gi|57165207|ref|NC_002678.2| | 5350767 | 5350882 | + | 65 | 116 | Mesorhizobium loti MAFF303099 chromosome |
αr14 | Mlr14C5 | gi|57165207|ref|NC_002678.2| | 6092423 | 6092540 | - | 64 | 118 | Mesorhizobium loti MAFF303099 chromosome |
αr14 | Mcr14C1 | gi|319779749|ref|NC_014923.1| | 3674843 | 3674963 | - | 64 | 121 | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome |
αr14 | Mcr14C2 | gi|319779749|ref|NC_014923.1| | 3675022 | 3675142 | - | 67 | 121 | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome |
αr14 | Mcr14C3 | gi|319779749|ref|NC_014923.1| | 3675197 | 3675318 | - | 65 | 122 | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome |
αr14 | Mcr14C4 | gi|319779749|ref|NC_014923.1| | 2545673 | 2545788 | + | 64 | 116 | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome |
αr14 | Mcr14C5 | gi|319779749|ref|NC_014923.1| | 5265891 | 5266009 | + | 63 | 119 | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome |
αr14 | Mcr14p01 | gi|319777789|ref|NC_014918.1| | 241412 | 241531 | - | 63 | 120 | Mesorhizobium ciceri biovar biserrulae WSM1271 plasmid pMESCI01 |
αr14 | Bcr14CI1 | gi|161617991|ref|NC_010103.1| | 338021 | 338128 | + | 56 | 108 | Brucella canis ATCC 23365 chromosome I |
αr14 | Bcr14CI2 | gi|161617991|ref|NC_010103.1| | 867217 | 867335 | - | 60 | 119 | Brucella canis ATCC 23365 chromosome I |
αr14 | Bs23445r14CI1 | gi|163842277|ref|NC_010169.1| | 355557 | 355664 | + | 57 | 108 | Brucella suis ATCC 23445 chromosome I |
αr14 | Bs23445r14CI2 | gi|163842277|ref|NC_010169.1| | 888713 | 888831 | - | 60 | 119 | Brucella suis ATCC 23445 chromosome I |
αr14 | Bm16Mr14CI1 | gi|17986284|ref|NC_003317.1| | 1116661 | 1116787 | + | 59 | 127 | Brucella melitensis bv. 1 str. 16M chromosome |
αr14 | Bm16Mr14CI2 | gi|17986284|ref|NC_003317.1| | 1644167 | 1644273 | - | 56 | 107 | Brucella melitensis bv. 1 str. 16M chromosome |
αr14 | BaS19r14CI1 | gi|189023268|ref|NC_010742.1| | 358310 | 358417 | + | 56 | 108 | Brucella abortus S19 chromosome 1 |
αr14 | BaS19r14CI2 | gi|189023268|ref|NC_010742.1| | 888877 | 888995 | - | 59 | 119 | Brucella abortus S19 chromosome 1 |
αr14 | Bm23457r14CI2 | gi|225851546|ref|NC_012441.1| | 890343 | 890461 | - | 59 | 119 | Brucella melitensis ATCC 23457 chromosome I |
αr14 | Bm23457r14CI1 | gi|225851546|ref|NC_012441.1| | 358976 | 359082 | + | 56 | 107 | Brucella melitensis ATCC 23457 chromosome I |
αr14 | Bs1330r14CI1 | gi|56968325|ref|NC_004310.3| | 338032 | 338139 | + | 56 | 108 | Brucella suis 1330 chromosome I |
αr14 | Bs1330r14CI2 | gi|56968325|ref|NC_004310.3| | 868847 | 868965 | - | 60 | 119 | Brucella suis 1330 chromosome I |
αr14 | Ba19941r14CI1 | gi|62288991|ref|NC_006932.1| | 359925 | 360032 | + | 56 | 108 | Brucella abortus bv. 1 str. 9-941 chromosome I |
αr14 | Ba19941r14CI2 | gi|62288991|ref|NC_006932.1| | 890576 | 890694 | - | 59 | 119 | Brucella abortus bv. 1 str. 9-941 chromosome I |
αr14 | Bmar14CI2 | gi|82698932|ref|NC_007618.1| | 886854 | 886972 | - | 59 | 119 | Brucella melitensis biovar Abortus 2308 chromosome I |
αr14 | Bmar14CI1 | gi|82698932|ref|NC_007618.1| | 356293 | 356400 | + | 56 | 108 | Brucella melitensis biovar Abortus 2308 chromosome I |
αr14 | Bor14CI1 | gi|148558820|ref|NC_009505.1| | 359682 | 359789 | + | 56 | 108 | Brucella ovis ATCC 25840 chromosome I |
αr14 | Bor14CI2 | gi|148558820|ref|NC_009505.1| | 895749 | 895867 | - | 60 | 119 | Brucella ovis ATCC 25840 chromosome I |
αr14 | Bmir14CI1 | gi|256368465|ref|NC_013119.1| | 339699 | 339806 | + | 57 | 108 | Brucella microti CCM 4915 chromosome 1 |
αr14 | Bmir14CI2 | gi|256368465|ref|NC_013119.1| | 873049 | 873167 | - | 60 | 119 | Brucella microti CCM 4915 chromosome 1 |
αr14 | Oar14CII1 | gi|153010078|ref|NC_009668.1| | 274684 | 274807 | + | 55 | 124 | Ochrobactrum anthropi ATCC 49188 chromosome 2 |
αr14 | Oar14CII2 | gi|153010078|ref|NC_009668.1| | 1275649 | 1275772 | + | 58 | 124 | Ochrobactrum anthropi ATCC 49188 chromosome 2 |
αr14 | Oar14CI1 | gi|153007346|ref|NC_009667.1| | 448505 | 448614 | + | 58 | 110 | Ochrobactrum anthropi ATCC 49188 chromosome 1 |
αr14 | Oar14CI2 | gi|153007346|ref|NC_009667.1| | 2453973 | 2454100 | + | 56 | 128 | Ochrobactrum anthropi ATCC 49188 chromosome 1 |
αr14 | MsBCNr14C1 | gi|110632362|ref|NC_008254.1| | 1916694 | 1916817 | + | 60 | 124 | Mesorhizobium sp. BNC1 |
αr14 | MsBCNr14C2 | gi|110632362|ref|NC_008254.1| | 3977086 | 3977208 | - | 60 | 123 | Mesorhizobium sp. BNC1 |
αr14 | MsBCNr14p1 | gi|110346917|ref|NC_008242.1| | 135270 | 135386 | - | 66 | 117 | Mesorhizobium sp. BNC1 plasmid 1 |
Discovery and Structure
Smr14C2 sRNA was described by del Val et al.,[2] as a result of a computational comparative genomic approach consisting in the integration of complementary strategies, designed to search for novel sRNA-encoding genes in the intergenic regions (IGRs) of the reference S. meliloti 1021 strain (http://iant.toulouse.inra.fr/bacteria/annotation/cgi/rhime.cgi) . Northern hybridization experiments confirmed that the predicted smr14C2 locus did express a single transcript of the expected size, which accumulated differentially in free-living and endosymbiotic bacteria. TAP-based 5’-RACE experiments mapped the transcription start site (TSS) of the full-length Smr14C transcript to the 1,667,613 nt position in the S. meliloti 1021 genome (http://iant.toulouse.inra.fr/bacteria/annotation/cgi/rhime.cgi) whereas the 3’-end was assumed to be located at the 1,667,491 nt position matching the last residue of the consecutive stretch of Us of a bona fide Rho-independent terminator (Figure 5). Parallel and later studies[3][4] in which Smr14C2 transcript is referred to as sra38 or Sm7', independently confirmed the expression this sRNA in S. melilloti and in its closely related strain 2011. Recent deep sequencing-based characterization of the small RNA fraction (50-350 nt) of S. meliloti 2011 further confirmed the expression of Smr14C2 (here referred to as SmelC397), and mapped the 5’- and 3´-ends of the full-length transcript to the same position in the S. meliloti 1021 genome.[5]
The nucleotide sequence of Smr14C2 was initially used as query to search against the Rfam database (version 10.0;http://www.sanger.ac.uk/Software/Rfam). This homology search rendered no matches to known bacterial sRNA in this database. Smr14C2 was next BLASTed with default parameters against all the currently available bacterial genomes (1,615 sequences at 20 April 2011; https://www.ncbi.nlm.nih.gov;). The regions exhibiting significant homology to the query sequence (78-89% similarity) were extracted to create a Covariance Model (CM) from a seed alignment using Infernal (version1.0)[8] (Figure 2). This CM was used in a further search for new members of the αr9 family in the existing bacterial genomic databases and resumed in the following Table 2:
Organism | Nr. of αr14 chromosomal homologs | Nr. of αr14 plasmids homologs | Total nr Nr. of αr14 copies |
---|---|---|---|
Sinorhizobium meliloti 1021 | 3 | 3 | 6 |
Sinorhizobium medicae WSM419 | 3 | 3 | 6 |
Sinorhizobium fredii NGR234 | 3 | 3 | 6 |
Rhizobium etli CIAT 652 | 3 | 3 | 6 |
Rhizobium etli CFN 42 | 3 | 3 | 6 |
Rhizobium leguminosarum bv. trifolii WSM2304 | 3 | 2 | 5 |
Rhizobium leguminosarum bv. trifolii WSM1325 | 3 | 5 | 8 |
Rhizobium leguminosarum bv. viciae 3841 | 4 | 2 | 6 |
Agrobacterium sp. H13-3 | 2 | ||
Agrobacterium vitis S4 | 3 CI / 2 CII | 1 | 6 |
Agrobacterium radiobacter K84 | 3 CI / 2 CII | 1 | 6 |
Agrobacterium tumefaciens str. C58 | 3 | 3 | |
Brucella suis ATCC 23445 | 2 | 2 | |
Brucella microti CCM 4915 | 2 | 2 | |
Brucella ovis ATCC 25840 | 2 | 2 | |
Brucella canis ATCC 23365 | 2 | 2 | |
Brucella melitensis ATCC 23457 | 2 | 2 | |
Brucella abortus S19 | 2 | 2 | |
Brucella abortus bv 1 str 9-941 | 2 | 2 | |
Brucella melitensis biovar Abortus 2308 | 2 | 2 | |
Brucella suis 1330 | 2 | 2 | |
Mesorhizobium loti MAFF303099 | 5 | 5 | |
Mesorhizobium sp. BNC1 | 2 | 1 | 3 |
Mesorhizobium ciceri biovar biserrulae WSM127 | 4 | 1 | 5 |
Ochrobactrum anthropi ATCC 49188 | 2 CI / 2 CII | 4 |
The results were manually inspected to deduce a consensus secondary structure for the family (Figure 1 and Figure 2). The consensus structure was also independently predicted with the program locARNATE[9] with very similar predictions. The manual inspection of the sequences found with the CM using Infernal allowed finding 101 homolog sequences. The rhizobial species encoding the 36 closer homologs to Smr14C2 were: S. medicae and S. fredii, two R. leguminosarum trifolii strains (WSM2304 and WSM1325), two R. etli strains CFN 42 and CIAT 652, the reference R. leguminosarum bv. viciae 3841 strain, and the Agrobacterium species A. vitis,A. tumefaciens, A. radiobacter and A. H13. All these sequences showed significant Infernal E-values (5.63E-29 - 8.16E-18) and bit-scores. The rest of the sequences found with the model showed high E-values between (1.33E-17 and 8.79E-03) but lower bit-scores and are encoded by Brucella species (B. ovis, B. canis, B. abortus, B. microtis, and several biobars of B. melitensis), Ochrobactrum anthropi and the Mesorhizobum species loti, M. ciceri and M. BNC.
The model identified five additional copies in the S. meliloti genome, locating two of them in tandem in the same IGR (Smr14C2, Smr14C3). The same results were obtained for S. medicae (Smedr14C1, Smedr14C2), S. fredii (Sfr14C1, Sfr14C2), M. loti (Mlr14C1, Mlr14C2) and M. ciceri (Mcr14C1, Mcr14C2).
Concerning the additional copies found in the S. meliloti genome, the expression of four out of the five copies have been independently confirmed in recent studies:
- Smr14C3 referred as: sra38,[3] Sm7[4] or SmelC398[5]
- Smr14A1 referred as: sma8[4] or SmelA075[5]
- Smr14A2 referred as: SmelA099[5]
- Smr14B referred as: SmelB161[5]
There are no experimental evidences up to now for the predicted copy Smr14C1.
Expression information
Parallel studies assessed Smr14C expression in S. meliloti 1021 under different biological conditions; i.e. bacterial growth in TY, minimal medium (MM) and luteolin-MM broth and endosymbiotic bacteria (i.e. mature symbiotic alfalfa nodules)[2] and high salt stress, oxidative stress and cold and hot shock stresses.[4] Expression of Smr14C2 in free-living bacteria was found to be growth-dependent, being the gene strongly down-regulated when bacteria entered the stationary phase. Expression of Smr14C2 increased ~5-fold in nodules when compared with free-living bacteria (log phase TY or MM cultures), suggesting the induction of these sRNAs during bacterial infection and/or bacteroid differentiation.[2] Recent deep sequencing data[5] found differential expression of the plasmic copies. Smr14A1 showed differential expression conditions, with a very low expression level in complex medium and in the same medium at decreased temperature. However, it was strongly up-regulated by heat-shock stress.[5] Smr14B showed an increase of its expression in the stationary phase greater that 8 fold. Moreover, also showed a week upregulation (<8 fold) upon acidic, basic and oxidative stress[5]
Promoter Analysis
All the promoter regions of the αr14 family members examined so far are very conserved in a sequence stretch extending up to 120 bp upstream of the transcription start site of the sRNA. All closest homolog loci have recognizable σ70-dependent promoters showing a -35/-10 consensus motif CTTAGAC-n17-CTATAT, which has been previously shown to be widely conserved among several other genera in the α-subgroup of proteobacteria.[10] To identify binding sites for other known transcription factors we used the fasta sequences provided by RegPredict[11](http://regpredict.lbl.gov/regpredict/help.html), and used those position weight matrices (PSWM) provided by RegulonDB[12] (http://regulondb.ccg.unam.mx). We built PSWM for each transcription factor from the RegPredict sequences using the Consensus/Patser program, choosing the best final matrix for motif lengths between 14–30 bps a threshold average E-value < 10E-10 for each matrix was established, (see "Thresholded consensus" in http://gps-tools2.its.yale.edu). Moreover, we searched for conserved unknown motifs using MEME[13] (http://meme.sdsc.edu/meme4_6_1/intro.html) and used relaxed regular expressions (i.e. pattern matching) over all Smr14C2 homologs promoters. This studies revealed two well defined groups of loci, the first one represented by the closest homologs (Figure 5) that presented a 26 bp long region very conserved between positions -40 and -75, marked as conserved MEME motif in (Figure 5), but no significant similarity to known transcription factor binding sites matrices could be established. A group of not so closely related members of the αr14 family constituted the second group of conserved promoters (Figure6). They presented a different promoter region, very well conserved across all members and an additional unknown 20 bp motif.
Genomic Context
Most of the members of the αr14 family are trans-encoded sRNAs transcribed from independent promoters in chromosomal IGRs. Many of the neighboring genes of the seed alignment’s members were not annotated and thus were further manually curated.[14][15][16] The tandem copies in the Sinorhizobium group’s genomes (Smr14C2, Smr14C3, Smedr14C1, Smedr14C2, Sfr14C1, Sfr14C2) presented a conserved genomic context. The gene upstream coded for an unknown protein containing the domain DUF1127 and the gene downstream coded for a trigger factor.
Partially conserved genomic context, DUF1127 protein upstream and 5-keto-4-deoxyuronate isomerase downstream, was found for the plasmidic copies of the Rhizobium group Rlt1325r14p02, Rlvr14p11, Rlt2304r14p02, and ReCIATr14A.
A similar case of partial genomic conservation, DUF1127 protein upstream and tRNA (uracil-5-)-methyltransferase downstream, was found in the case of the Mesorhizobium species, which also presented tandem copies M. loti (Mlr14C1, Mlr14C2) and M. ciceri (Mcr14C1, Mcr14C2). The same context was found for the second copy in the chromosome II of Ochrobactrum anthropii (Oar14CII2) and the second copy in chromosome I in the Brucella group (Bor14CI2, Bcr14CI2, Bmir14CI2, Bs1330r14CI2, Bm16Mr14CI1, BaS19r14CI2, Bm23457r14CI1, BMEA_A0934, BruAb1_0906, Ba19941r14CI2). In all these cases the sRNAs did appear as single copy in their IGR. Bs23445r14CI2, Bmar14CI2, and Rlvr14C2 shared the gene coding for tRNA (uracil-5-)-methyltransferase downstream, but did not shared the genes coding upstream.
The results showed two other well-conserved genomic contexts. One with a gene coding for nicotinic acid mononucleotide adenylyltransferase upstream and a gene coding for a molybdenum ABC transporter downstream, which is shared by the third copy of ar14 family in the A. radiobacter genome (Arr14CI3) and in the Rhizobium group genomes (ReCFNr14C3, Rlt1325r14C3, Rlvr14C3, Rlt2304r14C3). The second group is formed by the first copy in the Chromosome I in the Brucellas genomes that share the gene upstream coding for an LrgB membrane protein and the gene downstream a coding for an alpha/beta hydrolase.
Additional files:
- Additional file 1
- Additional file 2
- Additional file 3
- Additional file 4
- Additional file 5
- Additional file 6
- Additional file 7
- Additional file 8
- Additional file 9
- Additional file 10
- Additional file 11
Family | Feature | Name | Strand | Begin | End | Protein name | Annotation | Organism |
---|---|---|---|---|---|---|---|---|
αr14 | gene | SMc02051 | R | 1666735 | 1666878 | NP_385651.1 | DUF1127 | Sinorhizobium meliloti 1021 (NC_003047) |
αr14 | sRNA | Smr14C2 | R | 1667491 | 1667613 | - | sRNA | Sinorhizobium meliloti 1021 (NC_003047) |
αr14 | sRNA | Smr14C3 | R | 1667769 | 1667891 | - | sRNA | Sinorhizobium meliloti 1021 (NC_003047) |
αr14 | gene | SMc02050 | D | 1668357 | 1668441 | YP_001326866.1 | trigger factor | Sinorhizobium meliloti 1021 (NC_003047) |
αr14 | gene | Smed_1180 | R | 1257614 | 1257814 | YP_001326866.1 | DUF1127 | Sinorhizobium medicae WSM419 (NC_009636) |
αr14 | sRNA | Smedr14C1 | R | 1258038 | 1258160 | - | sRNA | Sinorhizobium medicae WSM419 chromosome (NC_009636) |
αr14 | sRNA | Smedr14C2 | R | 1258311 | 1258431 | - | sRNA | Sinorhizobium medicae WSM419 chromosome (NC_009636) |
αr14 | gene | Smed_1181 | D | 1258759 | 1258843 | YP_001326867.1 | trigger factor | Sinorhizobium medicae WSM419 chromosome (NC_009636) |
αr14 | gene | RHECIAT_CH0001925 | D | 1911937 | 1912446 | YP_001978070.1 | hypothetical protein | Rhizobium etli CIAT 652 (NC_010994) |
αr14 | sRNA | ReCIATr14C3 | R | 1912478 | 1912597 | - | sRNA | Rhizobium etli CIAT 652 (NC_010994) |
αr14 | gene | RHECIAT_CH0001926 | D | 1912923 | 1914026 | YP_001978071.1 | membrane protein | Rhizobium etli CIAT 652 (NC_010994) |
αr14 | gene | Atu4110 | D | 1229427 | 1230176 | NP_356536.1 | serine dehydrogenase | Agrobacterium tumefaciens str. C58 chromosome linear (NC_003063) |
αr14 | sRNA | Atr14C2 | D | 1230297 | 1230426 | - | sRNA | Agrobacterium tumefaciens str. C58 chromosome linear (NC_003063) |
αr14 | gene | Atu4112 | D | 1230719 | 1231906 | NP_356534.2 | proline dipeptidase-metalloexopeptidase | Agrobacterium tumefaciens str. C58 chromosome linear (NC_003063) |
αr14 | gene | Rleg_1700 | R | 1696657 | 1696803 | YP_002975526.1 | DUF1127 | Rhizobium leguminosarum bv. trifolii WSM1325 (NC_012850) |
αr14 | sRNA | Rlt1325r14C2 | R | 1697038 | 1697156 | - | sRNA | Rhizobium leguminosarum bv. trifolii WSM1325 (NC_012850) |
αr14 | gene | Rleg_1701 | D | 1697482 | 1698585 | YP_002975527.1 | membrane protein | Rhizobium leguminosarum bv. trifolii WSM1325 (NC_012850) |
αr14 | gene | RHE_CH01838 | D | 1925699 | 1926187 | YP_469358.1 | hypothetical protein | Rhizobium etli CFN 42 (NC_007761) |
αr14 | sRNA | ReCFNr14C2 | R | 1926239 | 1926358 | - | sRNA | Rhizobium etli CFN 42 (NC_007761) |
αr14 | gene | RHE_CH01839 | D | 1926684 | 1927778 | YP_469359.1 | membrane protein | Rhizobium etli CFN 42 (NC_007761) |
αr14 | gene | Rleg2_1502 | R | 1533929 | 1534075 | YP_002281020.1 | DUF1127 | Rhizobium leguminosarum bv. trifolii WSM2304 chromosome (NC_011369) |
αr14 | sRNA | Rlt2304r14C2 | R | 1534501 | 1534621 | - | sRNA | Rhizobium leguminosarum bv. trifolii WSM2304 chromosome (NC_011369) |
αr14 | gene | Rleg2_1503 | D | 1534947 | 1536050 | YP_002281021.1 | membrane protein | Rhizobium leguminosarum bv. trifolii WSM2304 chromosome (NC_011369) |
αr14 | gene | AGROH133_02958 | R | 108794 | 109252 | YP_004277412.1 | aminoacyl-tRNA ligase | Agrobacterium sp. H13-3 chromosome (NC_015183) |
αr14 | sRNA | AH13r14C1 | D | 109372 | 109501 | - | sRNA | Agrobacterium sp. H13-3 chromosome (NC_015183) |
αr14 | gene | AGROH133_02961 | D | 109733 | 109942 | YP_004277414.1 | cold shock protein | Agrobacterium sp. H13-3 chromosome (NC_015183) |
αr14 | gene | NGR_c13890 | R | 1433785 | 1433985 | YP_002825922.1 | DUF1127 | Sinorhizobium fredii NGR234 chromosome (NC_012587) |
αr14 | sRNA | Sfr14C2 | R | 1434463 | 1434586 | - | sRNA | Sinorhizobium fredii NGR234 chromosome (NC_012587) |
αr14 | sRNA | Sfr14C1 | R | 1434171 | 1434293 | - | sRNA | Sinorhizobium fredii NGR234 chromosome (NC_012587) |
αr14 | gene | NGR_c13920 | R | 1434843 | 1434925 | YP_002825924.1 | trigger factor | Sinorhizobium fredii NGR234 chromosome (NC_012587) |
αr14 | gene | Rleg2_0104 | D | 96796 | 97677 | YP_002279632.1 | aldo/keto reductase | Rhizobium leguminosarum bv. trifolii WSM2304 chromosome (NC_011369) |
αr14 | sRNA | Rlt2304r14C1 | D | 98050 | 98171 | - | sRNA | Rhizobium leguminosarum bv. trifolii WSM2304 chromosome (NC_011369) |
αr14 | gene | Rleg2_0105 | D | 98282 | 99559 | YP_002279633.1 | DUF900, alpha/beta hydrolase | Rhizobium leguminosarum bv. trifolii WSM2304 chromosome (NC_011369) |
αr14 | gene | Atu0104 | R | 108895 | 109353 | NP_353139.1 | aminoacyl-tRNA ligase | Agrobacterium tumefaciens str. C58 chromosome circular (NC_003062) |
αr14 | sRNA | Atr14C1 | D | 109473 | 109601 | - | sRNA | Agrobacterium tumefaciens str. C58 chromosome circular (NC_003062) |
αr14 | gene | Atu0106 | D | 109823 | 110032 | NP_353140.2 | cold shock protein | Agrobacterium tumefaciens str. C58 chromosome circular (NC_003062) |
αr14 | gene | RHECIAT_CH0000515 | D | 512732 | 513613 | YP_001976686.1 | oxidoreductase | Rhizobium etli CIAT 652 (NC_010994) |
αr14 | sRNA | ReCIATr14C1 | D | 513993 | 514114 | - | sRNA | Rhizobium etli CIAT 652 (NC_010994) |
αr14 | gene | RHECIAT_CH0000517 | D | 514221 | 515498 | YP_001976688.1 | DUF900, alpha/beta hydrolase | Rhizobium etli CIAT 652 (NC_010994) |
αr14 | gene | RL0474 | D | 511376 | 512272 | YP_766083.1 | aldo/keto reductase | Rhizobium leguminosarum bv. viciae 3841 (NC_008380) |
αr14 | sRNA | Rlvr14C1 | D | 512658 | 512779 | - | sRNA | Rhizobium leguminosarum bv. viciae 3841 (NC_008380) |
αr14 | gene | RL0475 | R | 512816 | 513202 | YP_766084.1 | hypothetical protein | Rhizobium leguminosarum bv. viciae 3841 (NC_008380) |
αr14 | gene | RHE_CH00449 | D | 461301 | 462182 | YP_467997.1 | oxidoreductase | Rhizobium etli CFN 42 (NC_007761) |
αr14 | sRNA | ReCFNr14C1 | D | 462566 | 462687 | - | sRNA | Rhizobium etli CFN 42 (NC_007761) |
αr14 | gene | RHE_CH00450 | D | 462794 | 464071 | YP_467998.1 | DUF900, alpha/beta hydrolase | Rhizobium etli CFN 42 (NC_007761) |
αr14 | gene | Avi_0122 | R | 113190 | 114716 | YP_002548072.1 | sulfatase | Agrobacterium vitis S4 chromosome 1 chromosome 1 (NC_011989) |
αr14 | sRNA | Avr14CI1 | D | 115126 | 115257 | - | sRNA | Agrobacterium vitis S4 chromosome 1 chromosome 1 (NC_011989) |
αr14 | gene | Avi_0126 | D | 115395 | 116840 | YP_002548074.1 | phosphomannomutase | Agrobacterium vitis S4 chromosome 1 chromosome 1 (NC_011989) |
αr14 | gene | Arad_2144 | R | 1692677 | 1692823 | YP_002544330.1 | DUF1127 | Agrobacterium radiobacter K84 chromosome 1 (NC_011985) |
αr14 | sRNA | Arr14CI2 | R | 1693278 | 1693409 | - | sRNA | Agrobacterium radiobacter K84 chromosome 1 (NC_011985) |
αr14 | gene | Arad_2145 | D | 1693731 | 1694870 | YP_002544331.1 | membrane protein | Agrobacterium radiobacter K84 chromosome 1 (NC_011985) |
αr14 | gene | RHECIAT_CH0004359 | D | 4444289 | 4444867 | YP_001980464.1 | nicotinic acid mononucleotide adenylyltransferase | Rhizobium etli CIAT 652 (NC_010994) |
αr14 | sRNA | ReCIATr14C4 | R | 4444894 | 4445016 | - | sRNA | Rhizobium etli CIAT 652 (NC_010994) |
αr14 | gene | RHECIAT_CH0004360 | R | 4445118 | 4445327 | YP_001980465.1 | hypothetical protein | Rhizobium etli CIAT 652 (NC_010994) |
αr14 | gene | Rleg_0114 | D | 105825 | 106706 | YP_002973964.1 | aldo/keto reductase | Rhizobium leguminosarum bv. trifolii WSM1325 (NC_012850) |
αr14 | sRNA | Rlt1325r14C1 | D | 107063 | 107184 | - | sRNA | Rhizobium leguminosarum bv. trifolii WSM1325 (NC_012850) |
αr14 | gene | Rleg_0115 | D | 107377 | 108654 | YP_002973965.1 | DUF900, alpha/beta hydrolase | Rhizobium leguminosarum bv. trifolii WSM1325 (NC_012850) |
αr14 | gene | Avi_2480 | R | 2042548 | 2043660 | YP_002549780.1 | membrane protein | Agrobacterium vitis S4 chromosome 1 chromosome 1 (NC_011989) |
αr14 | sRNA | Avr14CI2 | D | 2043913 | 2044044 | - | sRNA | Agrobacterium vitis S4 chromosome 1 chromosome 1 (NC_011989) |
αr14 | gene | Avi_2481 | D | 2044292 | 2044441 | YP_002549781.1 | DUF1127 | Agrobacterium vitis S4 chromosome 1 chromosome 1 (NC_011989) |
αr14 | gene | NGR_b21490 | R | 2204845 | 2205168 | YP_002824347.1 | acylphosphatase | Sinorhizobium fredii NGR234 plasmid pNGR234b (NC_012586) |
αr14 | sRNA | Sfr14b | D | 2205311 | 2205428 | - | sRNA | Sinorhizobium fredii NGR234 plasmid pNGR234b (NC_012586) |
αr14 | gene | NGR_b21500 | D | 2205588 | 2206316 | YP_002824348.1 | RNA polymerase sigma factor | Sinorhizobium fredii NGR234 plasmid pNGR234b (NC_012586) |
αr14 | gene | Arad_4880 | D | 3944888 | 3945469 | YP_002546417.1 | nicotinic acid mononucleotide adenylyltransferase | Agrobacterium radiobacter K84 chromosome 1 (NC_011985) |
αr14 | sRNA | Arr14CI3 | R | 3945511 | 3945639 | - | sRNA | Agrobacterium radiobacter K84 chromosome 1 (NC_011985) |
αr14 | gene | Arad_4883 | D | 3946044 | 3946844 | YP_002546419.1 | molybdenum ABC transporter | Agrobacterium radiobacter K84 chromosome 1 (NC_011985) |
αr14 | gene | RHE_CH04070 | D | 4313781 | 4314359 | YP_471540.1 | nicotinic acid mononucleotide adenylyltransferase | Rhizobium etli CFN 42 (NC_007761) |
αr14 | sRNA | ReCFNr14C3 | R | 4314384 | 4314505 | - | sRNA | Rhizobium etli CFN 42 (NC_007761) |
αr14 | gene | RHE_CH04071 | D | 4314798 | 4315589 | YP_471541.1 | molybdenum ABC transporter | Rhizobium etli CFN 42 (NC_007761) |
αr14 | gene | Rleg_4201 | D | 4295976 | 4296575 | YP_002977981.1 | nicotinic acid mononucleotide adenylyltransferase | Rhizobium leguminosarum bv. trifolii WSM1325 (NC_012850) |
αr14 | sRNA | Rlt1325r14C3 | R | 4296632 | 4296749 | - | sRNA | Rhizobium leguminosarum bv. trifolii WSM1325 (NC_012850) |
αr14 | gene | Rleg_4202 | D | 4297098 | 4297889 | YP_002977982.1 | molybdenum ABC transporter | Rhizobium leguminosarum bv. trifolii WSM1325 (NC_012850) |
αr14 | gene | RL4684 | D | 4987413 | 4988012 | YP_770246.1 | nicotinic acid mononucleotide adenylyltransferase | Rhizobium leguminosarum bv. viciae 3841 (NC_008380) |
αr14 | sRNA | Rlvr14C3 | R | 4988030 | 4988158 | - | sRNA | Rhizobium leguminosarum bv. viciae 3841 (NC_008380) |
αr14 | gene | RL4685 | D | 4988507 | 4989298 | YP_770247.1 | molybdenum ABC transporter | Rhizobium leguminosarum bv. viciae 3841 (NC_008380) |
αr14 | gene | Rleg2_3877 | D | 4018721 | 4019299 | YP_002283365.1 | nicotinic acid mononucleotide adenylyltransferase | Rhizobium leguminosarum bv. trifolii WSM2304 chromosome (NC_011369) |
αr14 | sRNA | Rlt2304r14C3 | R | 4019326 | 4019446 | - | sRNA | Rhizobium leguminosarum bv. trifolii WSM2304 chromosome (NC_011369) |
αr14 | gene | Rleg2_3878 | D | 4019781 | 4020572 | YP_002283366.1 | molybdenum ABC transporter | Rhizobium leguminosarum bv. trifolii WSM2304 chromosome (NC_011369) |
αr14 | gene | Arad_12190 | D | 137950 | 138255 | YP_002546688.1 | FAD dependent oxidoreductase | Agrobacterium radiobacter K84 plasmid pAtK84c (NC_011987) |
αr14 | sRNA | Arr14Atc | R | 138495 | 138625 | - | sRNA | Agrobacterium radiobacter K84 plasmid pAtK84c (NC_011987) |
αr14 | gene | Arad_12193 | R | 139226 | 140683 | YP_002546689.1 | Cytochrome c2C mono- and diheme oxidoreductase | Agrobacterium radiobacter K84 plasmid pAtK84c (NC_011987) |
αr14 | gene | Arad_8084 | D | 957315 | 957461 | YP_002541036.1 | DUF1127 | Agrobacterium radiobacter K84 chromosome 2 (NC_011983) |
αr14 | sRNA | Arr14CII1 | D | 957686 | 957817 | - | sRNA | Agrobacterium radiobacter K84 chromosome 2 (NC_011983) |
αr14 | gene | Arad_8086 | R | 957909 | 958754 | YP_002541037.1 | DUF900, alpha/beta hydrolase | Agrobacterium radiobacter K84 chromosome 2 (NC_011983) |
αr14 | gene | SM_b20590 | R | 1605382 | 1605705 | NP_438042.1 | acylphosphatase | Sinorhizobium meliloti 1021 plasmid pSymB (NC_003078) |
αr14 | sRNA | Smr14B | D | 1605826 | 1605943 | - | sRNA | Sinorhizobium meliloti 1021 plasmid pSymB (NC_003078) |
αr14 | gene | SM_b20592 | D | 1606089 | 1606772 | NP_438044.1 | RNA polymerase sigma factor | Sinorhizobium meliloti 1021 plasmid pSymB (NC_003078) |
αr14 | gene | Avi_5586 | D | 597555 | 598229 | YP_002547399.1 | hypothetical protein | Agrobacterium vitis S4 chromosome 2 (NC_011988) |
αr14 | sRNA | Avr14CII1 | R | 598426 | 598556 | - | sRNA | Agrobacterium vitis S4 chromosome 2 (NC_011988) |
αr14 | gene | Avi_5587 | R | 598779 | 598925 | YP_002547400.1 | DUF1127 | Agrobacterium vitis S4 chromosome 2 (NC_011988) |
αr14 | gene | Smed_4197 | R | 678214 | 678885 | YP_001312935.1 | RNA polymerase sigma factor | Sinorhizobium medicae WSM419 plasmid pSMED01 (NC_009620) |
αr14 | sRNA | Smedr14p01 | R | 679043 | 679160 | - | sRNA | Sinorhizobium medicae WSM419 plasmid pSMED01 (NC_009620) |
αr14 | gene | Smed_4198 | D | 679315 | 679599 | YP_001312936.1 | acylphosphatase | Sinorhizobium medicae WSM419 plasmid pSMED01 (NC_009620) |
αr14 | gene | Rleg_6288 | D | 1484 | 2256 | YP_002979285.1 | hypothetical protein | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132505 (NC_012854) |
αr14 | sRNA | Rlt1325r14p05 | R | 2286 | 2409 | - | sRNA | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132505 (NC_012854) |
αr14 | gene | Rleg_6290 | D | 3237 | 3734 | YP_002979286.1 | hypothetical protein | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132505 (NC_012854) |
αr14 | gene | Rleg_6602 | D | 28280 | 28426 | YP_002984605.1 | DUF1127 | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132502 (NC_012858) |
αr14 | sRNA | Rlt1325r14p02 | D | 28646 | 28767 | - | sRNA | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132502(NC_012858) |
αr14 | gene | Rleg_6603 | D | 29055 | 29891 | YP_002984606.1 | 5-keto-4-deoxyuronate isomerase | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132502 (NC_012858) |
αr14 | gene | Smed_5370 | D | 338535 | 338681 | YP_001314073.1 | DUF1127 | Sinorhizobium medicae WSM419 plasmid pSMED02 (NC_009621) |
αr14 | sRNA | Smedr14p022 | D | 338866 | 338990 | - | sRNA | Sinorhizobium medicae WSM419 plasmid pSMED02 (NC_009621) |
αr14 | gene | Smed_5371 | R | 339017 | 339226 | YP_001314074.1 | hypothetical protein | Sinorhizobium medicae WSM419 plasmid pSMED02 (NC_009621) |
αr14 | gene | pRL110530 | R | 573348 | 574184 | YP_771564.1 | 5-keto-4-deoxyuronate isomerase | Rhizobium leguminosarum bv. viciae 3841 plasmid pRL11 (NC_008384) |
αr14 | sRNA | Rlvr14p11 | R | 574471 | 574593 | - | sRNA | Rhizobium leguminosarum bv. viciae 3841 plasmid pRL11 (NC_008384) |
αr14 | gene | pRL110532 | R | 574811 | 574957 | YP_771566.1 | DUF1127 | Rhizobium leguminosarum bv. viciae 3841 plasmid pRL11 (NC_008384) |
αr14 | gene | Arad_9216 | R | 1920052 | 1920927 | YP_002541901.1 | glutathione S-transferase Ygh | Agrobacterium radiobacter K84 chromosome 2 (NC_011983) |
αr14 | sRNA | Arr14CII2 | R | 1921110 | 1921242 | - | sRNA | Agrobacterium radiobacter K84 chromosome 2 (NC_011983) |
αr14 | gene | Arad_9217 | R | 1921485 | 1921631 | YP_002541902.1 | DUF1127 | Agrobacterium radiobacter K84 chromosome 2 (NC_011983) |
αr14 | gene | Arad_0725 | D | 596785 | 597666 | YP_002543276.1 | oxidoreductase | Agrobacterium radiobacter K84 chromosome 1 (NC_011985) |
αr14 | sRNA | Arr14CI1 | D | 597903 | 598030 | - | sRNA | Agrobacterium radiobacter K84 chromosome 1 (NC_011985) |
αr14 | gene | Arad_0727 | D | 598187 | 599503 | YP_002543277.1 | DUF900, alpha/beta hydrolase | Agrobacterium radiobacter K84 chromosome 1 (NC_011985) |
αr14 | gene | RHE_PE00400 | R | 441962 | 442798 | YP_472562.1 | 5-keto-4-deoxyuronate isomerase | Rhizobium etli CFN 42 plasmid p42e (NC_007765) |
αr14 | sRNA | ReCFNr14e | R | 443001 | 443129 | - | sRNA | Rhizobium etli CFN 42 plasmid p42e (NC_007765) |
αr14 | gene | RHE_PE00401 | R | 443336 | 443482 | YP_472563.1 | hypothetical protein | Rhizobium etli CFN 42 plasmid p42e (NC_007765) |
αr14 | gene | Rleg_4483 | D | 265481 | 265806 | - | pseudogen | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132501 (NC_012848) |
αr14 | sRNA | Rlt1325r14p012 | D | 265945 | 266069 | - | sRNA | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132501 (NC_012848) |
αr14 | gene | Rleg_4484 | R | 266933 | 267917 | - | pseudogen | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132501 (NC_012848) |
αr14 | gene | Rleg2_5661 | D | 50447 | 50593 | YP_002277936.1 | DUF1127 | Rhizobium leguminosarum bv. trifolii WSM2304 plasmid pR132501 (NC_011366) |
αr14 | sRNA | Rlt2304r14p02 | D | 50814 | 50936 | - | sRNA | Rhizobium leguminosarum bv. trifolii WSM2304 plasmid pR132501 (NC_011366) |
αr14 | gene | Rleg2_5662 | D | 51199 | 52035 | YP_002277937.1 | 5-keto-4-deoxyuronate isomerase | Rhizobium leguminosarum bv. trifolii WSM2304 plasmid pR132501 (NC_011366) |
αr14 | gene | Rleg2_4962 | R | 606812 | 607117 | YP_002278931.1 | hypothetical protein | Rhizobium leguminosarum bv. trifolii WSM2304 plasmid pR132501 (NC_011368) |
αr14 | sRNA | Rlt2304r14p012 | D | 607636 | 607759 | - | sRNA | Rhizobium leguminosarum bv. trifolii WSM2304 plasmid pR132501 (NC_011368) |
αr14 | gene | Rleg2_4963 | R | 607786 | 608067 | - | pseudogen | Rhizobium leguminosarum bv. trifolii WSM2304 plasmid pR132501 (NC_011368) |
αr14 | gene | Rleg_6084 | D | 118943 | 119089 | YP_002978590.1 | DUF1127 | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132504 (NC_012852) |
αr14 | sRNA | Rlt1325r14p04 | D | 119314 | 119443 | - | sRNA | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132504 (NC_012852) |
αr14 | gene | Rleg_6085 | R | 119664 | 120056 | YP_002978591.1 | Cell wall assembly/cell proliferation coordinating protein, KNR4-like | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132504 (NC_012852) |
αr14 | gene | Avi_3374 | D | 2792636 | 2793793 | YP_002550427.1 | DNA polymerase I | Agrobacterium vitis S4 chromosome 1 (NC_011989) |
αr14 | sRNA | Avr14CI3 | R | 2793803 | 2793939 | - | sRNA | Agrobacterium vitis S4 chromosome 1 (NC_011989) |
αr14 | gene | Avi_3375 | R | 2794154 | 2794300 | YP_002550428.1 | DUF1127 | Agrobacterium vitis S4 chromosome 1 (NC_011989) |
αr14 | gene | RHECIAT_PA0000319 | R | 344808 | 345644 | YP_001985926.1 | 5-keto-4-deoxyuronate isomerase | Rhizobium etli CIAT 652 plasmid pA (NC_010998) |
αr14 | sRNA | ReCIATr14A | R | 345900 | 346021 | - | sRNA | Rhizobium etli CIAT 652 plasmid pA (NC_010998) |
αr14 | gene | RHECIAT_PA0000320 | R | 346130 | 346366 | YP_001985927.1 | DUF1127 | Rhizobium etli CIAT 652 plasmid pA (NC_010998) |
αr14 | gene | Meso_1791 | D | 1916546 | 1916686 | YP_674350.1 | DUF1127 | Mesorhizobium sp. BNC1 (NC_008254) |
αr14 | sRNA | MsBCNr14C1 | D | 1916694 | 1916817 | - | sRNA | Mesorhizobium sp. BNC1 (NC_008254) |
αr14 | gene | Meso_1793 | R | 1918307 | 1920064 | YP_674352.1 | hypothetical protein | Mesorhizobium sp. BNC1 (NC_008254) |
αr14 | gene | Avi_5890 | D | 899428 | 900909 | YP_002547646.1 | sulfate permease | Agrobacterium vitis S4 chromosome 2 (NC_011988) |
αr14 | sRNA | Avr14CII2 | D | 901035 | 901159 | - | sRNA | Agrobacterium vitis S4 chromosome 2 (NC_011988) |
αr14 | gene | Avi_5891 | R | 901178 | 902659 | YP_002547647.1 | siroheme synthase | Agrobacterium vitis S4 chromosome 2 (NC_011988) |
αr14 | gene | Avi_9007 | D | 6113 | 7192 | YP_002542553.1 | two component sensor kinase | Agrobacterium vitis S4 plasmid pAtS4c (NC_011984) |
αr14 | sRNA | Avr14Atc | D | 7318 | 7444 | - | sRNA | Agrobacterium vitis S4 plasmid pAtS4c (NC_011984) |
αr14 | gene | Avi_9008 | R | 7500 | 8633 | YP_002542554.1 | phage integrase family protein | Agrobacterium vitis S4 plasmid pAtS4c (NC_011984) |
αr14 | gene | RHECIAT_CH0001166 | R | 1172125 | 1173279 | YP_001977325.1 | polyhydroxybutyrate depolymerase protein | Rhizobium etli CIAT 652 (NC_010994) |
αr14 | sRNA | ReCIATr14C2 | D | 1173512 | 1173639 | - | sRNA | Rhizobium etli CIAT 652 (NC_010994) |
αr14 | gene | RHECIAT_CH0001168 | D | 1173948 | 1174214 | YP_001977327.1 | DUF2277 | Rhizobium etli CIAT 652 (NC_010994) |
αr14 | gene | Atu4670 | R | 1831052 | 1831294 | NP_535148.1 | hypothetical protein | Agrobacterium tumefaciens str. C58 chromosome linear (NC_003063) |
αr14 | sRNA | Atr14C3 | D | 1831483 | 1831614 | - | sRNA | Agrobacterium tumefaciens str. C58 chromosome linear (NC_003063) |
αr14 | gene | Atu4671 | R | 1831642 | 1832085 | NP_355992.2 | response regulator | Agrobacterium tumefaciens str. C58 chromosome linear (NC_003063) |
αr14 | gene | RHECIAT_PB0000101 | R | 111975 | 112217 | YP_001984364.1 | hypothetical protein | Rhizobium etli CIAT 652 plasmid pB (NC_010996) |
αr14 | sRNA | ReCIATr14B | D | 112292 | 112425 | - | sRNA | Rhizobium etli CIAT 652 plasmid pB (NC_010996) |
αr14 | gene | RHECIAT_PB0000102 | D | 112536 | 113108 | YP_001984365.1 | hypothetical protein | Rhizobium etli CIAT 652 plasmid pB (NC_010996) |
αr14 | gene | RHE_PD00068 | D | 76637 | 77354 | NP_659968.2 | hypothetical protein | Rhizobium etli CFN 42 symbiotic plasmid p42d (NC_004041) |
αr14 | sRNA | ReCFNr14d | D | 77668 | 77794 | - | sRNA | Rhizobium etli CFN 42 symbiotic plasmid p42d (NC_004041) |
αr14 | gene | RHE_PD00069 | D | 77912 | 78484 | NP_659967.1 | hydrolase | Rhizobium etli CFN 42 symbiotic plasmid p42d (NC_004041) |
αr14 | gene | NGR_c35080 | R | 3718015 | 3720336 | YP_002827985.1 | penicillin-binding | Sinorhizobium fredii NGR234 chromosome (NC_012587) |
αr14 | sRNA | Sfr14C3 | D | 3720579 | 3720700 | - | sRNA | Sinorhizobium fredii NGR234 chromosome (NC_012587) |
αr14 | gene | NGR_c35100 | D | 3721900 | 3722382 | YP_002827987.1 | hypothetical protein | Sinorhizobium fredii NGR234 chromosome (NC_012587) |
αr14 | gene | Smed_3386 | R | 3589318 | 3590205 | YP_001329042.1 | membrane protein | Sinorhizobium medicae WSM419 chromosome (NC_009636) |
αr14 | sRNA | Smedr14C3 | R | 3590299 | 3590417 | - | sRNA | Sinorhizobium medicae WSM419 chromosome (NC_009636) |
αr14 | gene | Smed_3387 | R | 3590494 | 3590925 | YP_001329043.1 | pyrimidine dimer DNA glycosylase | Sinorhizobium medicae WSM419 chromosome (NC_009636) |
αr14 | gene | SMc02855 | R | 205860 | 206783 | NP_384286.1 | membrane protein | Sinorhizobium meliloti 1021 (NC_003047) |
αr14 | sRNA | Smr14C1 | R | 206862 | 206980 | - | sRNA | Sinorhizobium meliloti 1021 (NC_003047) |
αr14 | gene | SMc02856 | D | 207244 | 209547 | NP_384287.1 | penicillin-binding protein | Sinorhizobium meliloti 1021 (NC_003047) |
αr14 | gene | msr1062 | D | 890180 | 890389 | NP_102735.1 | DUF1127 | Mesorhizobium loti MAFF303099 chromosome (NC_002678) |
αr14 | sRNA | Mlr14C1 | D | 890582 | 890702 | - | sRNA | Mesorhizobium loti MAFF303099 chromosome (NC_002678) |
αr14 | sRNA | Mlr14C2 | D | 890760 | 890880 | - | sRNA | Mesorhizobium loti MAFF303099 chromosome (NC_002678) |
αr14 | gene | mlr1064 | D | 891150 | 892568 | NP_102736.1 | tRNA (uracil-5-)-methyltransferase | Mesorhizobium loti MAFF303099 chromosome (NC_002678) |
αr14 | gene | SMa2165 | D | 1219615 | 1220355 | NP_436422.1 | oxidoreductase | Sinorhizobium meliloti 1021 plasmid pSymA (NC_003037) |
αr14 | sRNA | Smr14A1 | D | 1220690 | 1220808 | - | sRNA | Sinorhizobium meliloti 1021 plasmid pSymA (NC_003037) |
αr14 | gene | SMa2167 | D | 1221480 | 1221947 | NP_436423.1 | transcription factor regulator | Sinorhizobium meliloti 1021 plasmid pSymA (NC_003037) |
αr14 | gene | pRL120374 | R | 403573 | 404652 | YP_764884.1 | ribonuclease domain-containing protein | Rhizobium leguminosarum bv. viciae 3841 plasmid pRL12 (NC_008378) |
αr14 | sRNA | Rlvr14p12 | D | 407202 | 407316 | - | sRNA | Rhizobium leguminosarum bv. viciae 3841 plasmid pRL12 (NC_008378) |
αr14 | gene | pRL120378 | R | 408099 | 409010 | YP_764885.1 | L-isoaspartate O-methyltransferase | Rhizobium leguminosarum bv. viciae 3841 plasmid pRL12 (NC_008378) |
αr14 | gene | Mesci_3513 | R | 3673344 | 3674753 | YP_004142683.1 | tRNA (uracil-5-)-methyltransferase | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome (NC_014923) |
αr14 | sRNA | Mcr14C1 | R | 3674843 | 3674963 | - | sRNA | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome (NC_014923) |
αr14 | sRNA | Mcr14C2 | R | 3675022 | 3675142 | - | sRNA | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome (NC_014923) |
αr14 | gene | Mesci_3514 | R | 3675321 | 3675461 | YP_004142684.1 | DUF1127 | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome (NC_014923) |
αr14 | gene | Oant_0422 | D | 447656 | 448351 | YP_001368982.1 | LrgB membrane protein /hydrolase | Ochrobactrum anthropi ATCC 49188 chromosome 1 (NC_009667) |
αr14 | sRNA | Oar14CI1 | D | 448505 | 448614 | - | sRNA | Ochrobactrum anthropi ATCC 49188 chromosome 1 (NC_009667) |
αr14 | gene | Oant_0423 | D | 448748 | 450301 | YP_001368983.1 | glyoxalase bleomycin resistance protein dioxygenase/hydrolase | Ochrobactrum anthropi ATCC 49188 chromosome 1 (NC_009667) |
αr14 | gene | Oant_2969 | R | 272174 | 273850 | YP_001371506.1 | hypothetical protein | Ochrobactrum anthropi ATCC 49188 chromosome 2 (NC_009668) |
αr14 | sRNA | Oar14CII1 | D | 274684 | 274807 | - | sRNA | Ochrobactrum anthropi ATCC 49188 chromosome 2 (NC_009668) |
αr14 | gene | Oant_2970 | D | 275652 | 276380 | YP_001371507.1 | septum formation inhibitor | Ochrobactrum anthropi ATCC 49188 chromosome 2 (NC_009668) |
αr14 | gene | SMa2355 | D | 1327042 | 1328151 | NP_436520.4 | DNA polymerase I | Sinorhizobium meliloti 1021 plasmid pSymA (NC_003037) |
αr14 | sRNA | Smr14A2 | R | 1328176 | 1328301 | - | sRNA | Sinorhizobium meliloti 1021 plasmid pSymA (NC_003037) |
αr14 | gene | SMa2357 | D | 1329978 | 1330670 | NP_436521.2 | adenylate guanylate cyclase | Sinorhizobium meliloti 1021 plasmid pSymA (NC_003037) |
αr14 | gene | Oant_2332 | D | 2453758 | 2453904 | YP_001370875.1 | DUF1127 | Ochrobactrum anthropi ATCC 49188 chromosome 1 (NC_009667) |
αr14 | sRNA | Oar14CI2 | D | 2453973 | 2454100 | - | sRNA | Ochrobactrum anthropi ATCC 49188 chromosome 1 (NC_009667) |
αr14 | gene | Oant_2333 | D | 2454171 | 2455601 | YP_001370876.1 | tRNA (uracil-5-)-methyltransferase | Ochrobactrum anthropi ATCC 49188 chromosome 1 (NC_009667) |
αr14 | gene | Rleg_4742 | D | 111695 | 112336 | YP_002972933.1 | hypothetical protein | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132501 (NC_012848) |
αr14 | sRNA | Rlt1325r14p011 | D | 112604 | 112728 | - | sRNA | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132501 (NC_012848) |
αr14 | gene | Rleg_4743 | R | 112925 | 113309 | - | hypothetical protein | Rhizobium leguminosarum bv. trifolii WSM1325 plasmid pR132501 (NC_012848) |
αr14 | gene | Meso_3687 | R | 3975340 | 3976962 | YP_676220.1 | alpha amylase-C hydrolase | Mesorhizobium sp. BNC1 (NC_008254) |
αr14 | sRNA | MsBCNr14C2 | R | 3977086 | 3977208 | - | sRNA | Mesorhizobium sp. BNC1 (NC_008254) |
αr14 | gene | Meso_3688 | D | 3977363 | 3978682 | YP_676221.1 | homoserine dehydrogenase-C NAD-binding | Mesorhizobium sp. BNC1 (NC_008254) |
αr14 | gene | BSUIS_A0355 | D | 354629 | 355336 | YP_001627017.1 | LrgB membrane protein /hydrolase | Brucella suis ATCC 23445 chromosome I (NC_010169) |
αr14 | sRNA | Bs23445r14CI1 | D | 355557 | 355664 | - | sRNA | Brucella suis ATCC 23445 chromosome I (NC_010169) |
αr14 | gene | BSUIS_A0356 | D | 355877 | 357124 | YP_001627018.1 | hypothetical protein | Brucella suis ATCC 23445 chromosome I (NC_010169) |
αr14 | gene | BMI_I332 | D | 338771 | 339478 | YP_003106291.1 | LrgB membrane protein /hydrolase | Brucella microti CCM 4915 chromosome I (NC_013119) |
αr14 | sRNA | Bmir14CI1 | D | 339699 | 339806 | - | sRNA | Brucella microti CCM 4915 chromosome I (NC_013119) |
αr14 | gene | BMI_I334 | D | 340025 | 341266 | YP_003106293.1 | DUF900, alpha/beta hydrolase | Brucella microti CCM 4915 chromosome I (NC_013119) |
αr14 | gene | BOV_0342 | D | 358754 | 359461 | YP_001258364.1 | hydrolase | Brucella ovis ATCC 25840 chromosome I (NC_009505) |
αr14 | sRNA | Bor14CI1 | D | 359682 | 359789 | - | sRNA | Brucella ovis ATCC 25840 chromosome I (NC_009505) |
αr14 | gene | BOV_0343 | D | 360008 | 361250 | - | pseudogen | Brucella ovis ATCC 25840 chromosome I (NC_009505) |
αr14 | gene | BCAN_A0335 | D | 337093 | 337800 | YP_001592196.1 | LrgB membrane protein /hydrolase | Brucella canis ATCC 23365 chromosome I (NC_010103) |
αr14 | sRNA | Bcr14CI1 | D | 338021 | 338128 | - | sRNA | Brucella canis ATCC 23365 chromosome I (NC_010103) |
αr14 | gene | BCAN_A0337 | D | 338347 | 339588 | YP_001592198.1 | DUF900, alpha/beta hydrolase | Brucella canis ATCC 23365 chromosome I (NC_010103) |
αr14 | gene | BMEA_A0364 | D | 358048 | 358755 | YP_002732104.1 | LrgB membrane protein /hydrolase | Brucella melitensis ATCC 23457 chromosome I (NC_012441) |
αr14 | sRNA | Bm23457r14CI1 | D | 358976 | 359082 | - | sRNA | Brucella melitensis ATCC 23457 chromosome I (NC_012441) |
αr14 | gene | BMEA_A0366 | D | 359302 | 360543 | YP_002732106.1 | DUF900, alpha/beta hydrolase | Brucella melitensis ATCC 23457 chromosome I (NC_012441) |
αr14 | gene | BAbS19_I03260 | D | 357382 | 358089 | YP_001934341.1 | LrgB membrane protein /hydrolase | Brucella abortus S19 chromosome I (NC_010742) |
αr14 | sRNA | BaS19r14CI1 | D | 358310 | 358417 | - | sRNA | Brucella abortus S19 chromosome I (NC_010742) |
αr14 | gene | BAbS19_I03270 | D | 358635 | 359876 | YP_001934342.1 | DUF900, alpha/beta hydrolase | Brucella abortus S19 chromosome I (NC_010742) |
αr14 | gene | BruAb1_0352 | D | 358997 | 359704 | YP_221113.1 | membrane protein | Brucella abortus bv. 1 str. 9-941 chromosome I (NC_006932) |
αr14 | sRNA | Ba19941r14CI1 | D | 359925 | 360032 | - | sRNA | Brucella abortus bv. 1 str. 9-941 chromosome I (NC_006932) |
αr14 | gene | BruAb1_0355 | D | 361578 | 362459 | YP_221115.1 | pantoate—beta-alanine ligase | Brucella abortus bv. 1 str. 9-941 chromosome I (NC_006932) |
αr14 | gene | BAB1_0356 | D | 355365 | 356072 | YP_413824.1 | LrgB membrane protein /hydrolase | Brucella melitensis biovar Abortus 2308 chromosome I (NC_007618) |
αr14 | sRNA | Bmar14CI1 | D | 356293 | 356400 | - | sRNA | Brucella melitensis biovar Abortus 2308 chromosome I (NC_007618) |
αr14 | gene | BAB1_0358 | D | 356618 | 357859 | YP_413825.1 | DUF900, alpha/beta hydrolase | Brucella melitensis biovar Abortus 2308 chromosome I (NC_007618) |
αr14 | gene | BR0326 | D | 337104 | 337811 | NP_697360.1 | DUF900, alpha/beta hydrolase | Brucella suis 1330 chromosome I (NC_004310) |
αr14 | sRNA | Bs1330r14CI1 | D | 338032 | 338139 | - | sRNA | Brucella suis 1330 chromosome I (NC_004310) |
αr14 | gene | BR0328 | D | 338358 | 339599 | NP_697362.1 | tRNA (uracil-5-)-methyltransferase | Brucella suis 1330 chromosome I (NC_004310) |
αr14 | gene | Smed_5352 | R | 313859 | 314596 | YP_001314057.1 | adenylate guanylate cyclase | Sinorhizobium medicae WSM419 plasmid pSMED02 (NC_009621) |
αr14 | sRNA | Smedr14p021 | D | 316304 | 316428 | - | sRNA | Sinorhizobium medicae WSM419 plasmid pSMED02 (NC_009621) |
αr14 | gene | Smed_5353 | R | 316485 | 317621 | YP_001314058.1 | DNA polymerase I | Sinorhizobium medicae WSM419 plasmid pSMED02 (NC_009621) |
αr14 | gene | RHE_PF00022 | R | 22159 | 22494 | YP_472643.1 | hypothetical protein | Rhizobium etli CFN 42 plasmid p42f (NC_007766) |
αr14 | sRNA | ReCFNr14f | D | 22818 | 22935 | - | sRNA | Rhizobium etli CFN 42 plasmid p42f (NC_007766) |
αr14 | gene | RHE_PF00023 | D | 23286 | 23726 | YP_472644.1 | carbon-sulfur lyase | Rhizobium etli CFN 42 plasmid p42f (NC_007766) |
αr14 | gene | Oant_3865 | D | 1275426 | 1275572 | YP_001372399.1 | DUF1127 | Ochrobactrum anthropi ATCC 49188 chromosome 2 (NC_009668) |
αr14 | sRNA | Oar14CII2 | D | 1275649 | 1275772 | - | sRNA | Ochrobactrum anthropi ATCC 49188 chromosome 2 (NC_009668) |
αr14 | gene | Oant_3866 | R | 1275787 | 1277010 | YP_001372400.1 | membrane protein | Ochrobactrum anthropi ATCC 49188 chromosome 2 (NC_009668) |
αr14 | gene | Rleg2_4668 | D | 305380 | 306273 | YP_002278660.1 | L-isoaspartate O-methyltransferase | Rhizobium leguminosarum bv. trifolii WSM2304 plasmid pRLG201 (NC_011368) |
αr14 | sRNA | Rlt2304r14p011 | R | 306339 | 306454 | - | sRNA | Rhizobium leguminosarum bv. trifolii WSM2304 plasmid pRLG201 (NC_011368) |
αr14 | gene | Rleg2_4670 | R | 306976 | 307923 | YP_002278662.1 | transposase IS110 family protein | Rhizobium leguminosarum bv. trifolii WSM2304 plasmid pRLG201 (NC_011368) |
αr14 | gene | Mesci_6302 | D | 240660 | 240910 | YP_004134462.1 | hypothetical protein | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome I (NC_014918) |
αr14 | sRNA | Mcr14p01 | R | 241412 | 241531 | - | sRNA | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome I (NC_014918) |
αr14 | gene | Mesci_6303 | R | 241784 | 242047 | YP_004134463.1 | hypothetical protein | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome I (NC_014918) |
αr14 | gene | Mesci_2426 | D | 2545134 | 2545592 | YP_004141622.1 | phasin | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome I (NC_014923) |
αr14 | sRNA | Mcr14CI4 | D | 2545673 | 2545788 | - | sRNA | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome I (NC_014923) |
αr14 | gene | Mesci_2427 | D | 2545962 | 2546038 | YP_004141623 | FAD dependent oxidoreductase | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome I (NC_014923) |
αr14 | gene | Mesci_5119 | D | 5264750 | 5265772 | YP_004144269.1 | LacI family transcription regulator | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome I (NC_014923) |
αr14 | sRNA | Mcr14CI5 | D | 5265891 | 5266009 | - | sRNA | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome I (NC_014923) |
αr14 | gene | Mesci_5121 | R | 5266103 | 5266693 | YP_004144271.1 | TetR family transcription regulator | Mesorhizobium ciceri biovar biserrulae WSM1271 chromosome I (NC_014923) |
αr14 | gene | BOV_0890 | R | 894271 | 895671 | YP_001258873.1 | tRNA (uracil-5-)-methyltransferase | Brucella ovis ATCC 25840 chromosome I (NC_009505) |
αr14 | sRNA | Bor14CI2 | R | 895749 | 895867 | - | sRNA | Brucella ovis ATCC 25840 chromosome I (NC_009505) |
αr14 | gene | BOV_0891 | R | 895931 | 896077 | YP_001258874.1 | DUF1127 | Brucella ovis ATCC 25840 chromosome I (NC_009505) |
αr14 | gene | BCAN_A0908 | R | 865739 | 867139 | YP_001592741.1 | tRNA (uracil-5-)-methyltransferase | Brucella canis ATCC 23365 chromosome I (NC_010103) |
αr14 | sRNA | Bcr14CI2 | R | 867217 | 867335 | - | sRNA | Brucella canis ATCC 23365 chromosome I (NC_010103) |
αr14 | gene | BCAN_A0909 | R | 867399 | 867545 | YP_001592742.1 | DUF1127 | Brucella canis ATCC 23365 chromosome I (NC_010103) |
αr14 | gene | BSUIS_A0933 | R | 887235 | 888635 | YP_001627570.1 | tRNA (uracil-5-)-methyltransferase | Brucella suis ATCC 23445 chromosome I (NC_010169) |
αr14 | sRNA | Bs23445r14CI2 | R | 888713 | 888831 | - | sRNA | Brucella suis ATCC 23445 chromosome I (NC_010169) |
αr14 | gene | BSUIS_A0934 | R | 888895 | 889041 | YP_001627571.1 | hypothetical protein | Brucella suis ATCC 23445 chromosome I (NC_010169) |
αr14 | gene | BMI_I892 | R | 871571 | 872971 | YP_003106830.1 | tRNA (uracil-5-)-methyltransferase | Brucella microti CCM 4915 chromosome 1 (NC_013119) |
αr14 | sRNA | Bmir14CI2 | R | 873049 | 873167 | - | sRNA | Brucella microti CCM 4915 chromosome 1 (NC_013119) |
αr14 | gene | BMI_I893 | R | 873231 | 873377 | YP_003106831.1 | DUF1127 | Brucella microti CCM 4915 chromosome 1 (NC_013119) |
αr14 | gene | BR0894 | R | 867369 | 868769 | NP_697907.1 | tRNA (uracil-5-)-methyltransferase | Brucella suis 1330 chromosome I (NC_004310) |
αr14 | sRNA | Bs1330r14CI2 | R | 868847 | 868965 | - | sRNA | Brucella suis 1330 chromosome I (NC_004310) |
αr14 | gene | BR0895 | R | 869029 | 869175 | NP_697908.1 | DUF1127 | Brucella suis 1330 chromosome I (NC_004310) |
αr14 | gene | mlr7359 | D | 6091731 | 6092324 | NP_107697.1 | transcription factor regulator | Mesorhizobium loti MAFF303099 chromosome (NC_002678) |
αr14 | sRNA | Mlr14C5 | R | 6092423 | 6092540 | - | sRNA | Mesorhizobium loti MAFF303099 chromosome (NC_002678) |
αr14 | gene | mll7360 | R | 6092659 | 6093681 | NP_107698.1 | transcription factor regulator | Mesorhizobium loti MAFF303099 chromosome (NC_002678) |
αr14 | gene | BMEI1072 | D | 1116469 | 1116615 | NP_539989.1 | DUF1127 | Brucella melitensis bv. 1 str. 16M chromosome (NC_003317) |
αr14 | sRNA | Bm16Mr14CI1 | D | 1116661 | 1116787 | - | sRNA | Brucella melitensis bv. 1 str. 16M chromosome (NC_003317) |
αr14 | gene | BMEI1073 | D | 1116821 | 1118257 | NP_539990.1 | tRNA (uracil-5-)-methyltransferase | Brucella melitensis bv. 1 str. 16M chromosome (NC_003317) |
αr14 | gene | BAbS19_I08530 | R | 887399 | 888799 | YP_001934848.1 | tRNA (uracil-5-)-methyltransferase | Brucella abortus S19 chromosome 1 (NC_010742) |
αr14 | sRNA | BaS19r14CI2 | R | 888877 | 888995 | - | sRNA | Brucella abortus S19 chromosome 1 (NC_010742) |
αr14 | gene | BAbS19_I08540 | R | 889041 | 889187 | YP_001934849.1 | DUF1127 | Brucella abortus S19 chromosome 1 (NC_010742) |
αr14 | gene | BMEA_A0933 | R | 888865 | 890265 | YP_002732640.1 | tRNA (uracil-5-)-methyltransferase | Brucella melitensis ATCC 23457 chromosome I (NC_012441) |
αr14 | sRNA | Bm23457r14CI2 | R | 890343 | 890461 | - | sRNA | Brucella melitensis ATCC 23457 chromosome I (NC_012441) |
αr14 | gene | BMEA_A0934 | R | 890507 | 890653 | YP_002732641.1 | DUF1127 | Brucella melitensis ATCC 23457 chromosome I (NC_012441) |
αr14 | gene | BruAb1_0906 | R | 889098 | 890498 | YP_221632.1 | tRNA (uracil-5-)-methyltransferase | Brucella abortus bv. 1 str. 9-941 chromosome I (NC_006932) |
αr14 | sRNA | Ba19941r14CI2 | R | 890576 | 890694 | - | sRNA | Brucella abortus bv. 1 str. 9-941 chromosome I (NC_006932) |
αr14 | gene | BruAb1_0907 | R | 890740 | 890886 | YP_221633.1 | DUF1127 | Brucella abortus bv. 1 str. 9-941 chromosome I (NC_006932) |
αr14 | gene | BAB1_0913 | R | 885376 | 886776 | YP_414338.1 | tRNA (uracil-5-)-methyltransferase | Brucella melitensis biovar Abortus 2308 chromosome I (NC_007618) |
αr14 | sRNA | Bmar14CI2 | R | 886854 | 886972 | - | sRNA | Brucella melitensis biovar Abortus 2308 chromosome I (NC_007618) |
αr14 | gene | BAB1_0914 | R | 887018 | 887164 | YP_414339.1 | oxidoreductase | Brucella melitensis biovar Abortus 2308 chromosome I (NC_007618) |
αr14 | gene | Meso_4235 | R | 134555 | 135097 | YP_665858.1 | hypothetical protein | Mesorhizobium sp. BNC1 plasmid 1 (NC_008242) |
αr14 | sRNA | MsBCNr14p1 | R | 135270 | 135386 | - | sRNA | Mesorhizobium sp. BNC1 plasmid 1 (NC_008242) |
αr14 | gene | Meso_4236 | R | 135539 | 136099 | YP_665859.1 | hypothetical protein | Mesorhizobium sp. BNC1 plasmid 1 (NC_008242) |
αr14 | gene | mlr6541 | D | 5350228 | 5350686 | NP_107027.1 | phasin | Mesorhizobium loti MAFF303099 chromosome (NC_002678) |
αr14 | sRNA | Mlr14C4 | D | 5350767 | 5350882 | - | sRNA | Mesorhizobium loti MAFF303099 chromosome (NC_002678) |
αr14 | gene | mlr6543 | D | 5351042 | 5351115 | - | two component regulator | Mesorhizobium loti MAFF303099 chromosome (NC_002678) |
αr14 | gene | RL2059 | R | 2172416 | 2173849 | YP_767657.1 | tRNA (uracil-5-)-methyltransferase | Rhizobium leguminosarum bv. viciae 3841 (NC_008380) |
αr14 | sRNA | Rlvr14C2 | R | 2174956 | 2175074 | - | sRNA | Rhizobium leguminosarum bv. viciae 3841 (NC_008380) |
αr14 | gene | RL2060 | D | 2175400 | 2176503 | YP_767658.1 | membrane protein | Rhizobium leguminosarum bv. viciae 3841 (NC_008380) |
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