BenR in *Pseudomonas* (Gene **benR** in PSEPK) – Function, Regulation, and Pathway Roles OpenAI o3-deep-research-2025-06-26 100 citations 2025-11-05T14:45:06.227110

BenR in Pseudomonas (Gene benR in PSEPK) – Function, Regulation, and Pathway Roles

Overview and Gene Family

The benR gene in Pseudomonas (strain “PSEPK”) encodes BenR, a transcriptional regulator that controls benzoate catabolism. BenR is a member of the AraC/XylS family of regulators, sharing ~62% amino acid identity with XylS from the TOL plasmid of Pseudomonas putida (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). Like XylS, BenR functions as a DNA-binding activator that responds to aromatic effectors. It is a positive regulator required for the induction of benzoate degradation genes in the presence of benzoate. Sequence analyses identified BenR (318 amino acids, ~36 kDa) as having the conserved two-domain architecture of AraC-family proteins: an N-terminal effector-binding domain and a C-terminal helix-turn-helix DNA-binding domain (pmc.ncbi.nlm.nih.gov). This places BenR in the XylS subfamily of AraC regulators, distinct from LysR-type regulators (like BenM or CatR in other bacteria) that also control aromatic catabolic pathways (bmcmicrobiol.biomedcentral.com).

Localization and mechanism: As a transcription factor, BenR operates intracellularly in the cytoplasm/nucleoid, binding to promoter regions of target genes. A direct-repeat DNA motif has been identified upstream of the benzoate catabolic genes matching the consensus AraC/XylS binding site (pmc.ncbi.nlm.nih.gov). BenR likely binds this site as a dimer and, upon binding its effector (benzoate or a derivative), it recruits RNA polymerase to activate transcription of the catabolic operon. Overexpression of BenR in E. coli or providing benzoate as an inducer strongly elevates transcription from the benzoate-degradative promoters, confirming BenR’s role as a ligand-dependent activator (pmc.ncbi.nlm.nih.gov). In P. putida, for example, BenR overproduction led to a ~25-fold increase in benA promoter activity, and wild-type cells showed ~15-fold induction of benA-lacZ reporter expression when benzoate was added (pmc.ncbi.nlm.nih.gov). This effector-responsive DNA-binding mechanism underlies BenR’s control of downstream genes.

Role in Benzoate Catabolism and Target Genes

BenR’s primary function is to activate the expression of the benzoate degradation (ben) operon when benzoate is present. Genomic analysis of P. putida revealed that benR is clustered with benzoate catabolic genes (pmc.ncbi.nlm.nih.gov). Immediately downstream of benR are at least seven genes (designated benA, benB, benC, benD, benE, benK, and benF) inferred to encode the enzymes and transporters for benzoate utilization (pmc.ncbi.nlm.nih.gov). Specifically, BenA, BenB, and BenC form a multicomponent benzoate 1,2-dioxygenase (terminal oxygenase and reductase components) that catalyzes the conversion of benzoate to cis-1,2-dihydro-1,2-dihydroxybenzoate (pmc.ncbi.nlm.nih.gov). BenD is a NAD^+^-dependent cis-diol dehydrogenase that further oxidizes this dihydrodiol to catechol (pmc.ncbi.nlm.nih.gov). Thus, under BenR control, benzoate is channeled into catechol, a central intermediate of aromatic catabolism. Additional cluster genes provide uptake functions: BenK is a benzoate permease (inner membrane transporter) and BenF is an outer-membrane porin, facilitating benzoate entry into the cell (pmc.ncbi.nlm.nih.gov). Another gene, BenE, is present in the cluster with an unclear function (pmc.ncbi.nlm.nih.gov). All these ben genes are induced in the presence of benzoate, but only if BenR is functional (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov).

Biological pathway: The BenR-regulated operon initiates the β-ketoadipate pathway’s catechol branch, which is a major route for aromatic compound degradation in soil bacteria (pmc.ncbi.nlm.nih.gov) (pubmed.ncbi.nlm.nih.gov). When benzoate is available, BenR triggers its conversion to catechol. Catechol is then cleaved by downstream enzymes (e.g., catechol 1,2-dioxygenase CatA and others of the ortho-cleavage route) into β-ketoadipate, which enters central metabolism (pubmed.ncbi.nlm.nih.gov). P. putida can degrade catechol via an ortho-cleavage pathway to TCA cycle intermediates; if the TOL plasmid is present, it can also use a meta-cleavage pathway for methylbenzoates (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). BenR specifically governs the peripheral (benzoate to catechol) steps and works in concert with other regulators that control downstream steps. In Pseudomonas, the catechol-cleavage operon (cat genes) is regulated by the LysR-type activator CatR responding to cis,cis-muconate (the product of catechol cleavage), and the protocatechuate branch (pca genes) is regulated by PcaR responding to β-ketoadipate (pubmed.ncbi.nlm.nih.gov) (pubmed.ncbi.nlm.nih.gov). This division of labor means that BenR is necessary and sufficient for the initial benzoate oxidation steps, while further metabolism of catechol requires additional inducible systems. Indeed, BenR is required for growth on benzoate as a sole carbon source, but mutants lacking benR can still grow on catechol, since the downstream catabolic enzymes are intact and governed independently (bmcmicrobiol.biomedcentral.com). In P. stutzeri A1501, for example, a ΔbenR strain lost the ability to utilize benzoate but remained capable of using catechol, demonstrating that BenR is the sole activator of the benABCD operon (benzoate dioxygenase/dehydrogenase genes) in that organism (bmcmicrobiol.biomedcentral.com). Complementation with a functional benR gene restores benzoate utilization, confirming its central role (bmcmicrobiol.biomedcentral.com).

Notably, in Pseudomonas strains where the ben and cat genes are genetically linked, BenR can have a broader influence. P. stutzeri A1501 lacks a CatR homolog; intriguingly, benzoate addition in this strain still activates transcription of catechol degradation genes (catBCA) to some extent (bmcmicrobiol.biomedcentral.com) (bmcmicrobiol.biomedcentral.com). Comparative sequence analysis showed a conserved BenR-binding site in the promoter of the benA gene across pseudomonads (bmcmicrobiol.biomedcentral.com), and there is evidence that BenR (or an unknown regulator) may cross-activate the catechol pathway in the absence of CatR. In most Pseudomonas, however, BenR and CatR work together to ensure a coordinated degradation: BenR triggers catechol formation, and CatR (triggered by catechol’s product muconate) induces the enzymes to process catechol (bmcmicrobiol.biomedcentral.com) (bmcmicrobiol.biomedcentral.com). This two-tier regulatory scheme is viewed as a model of complex transcriptional control in aromatic catabolism, allowing precise tuning of each pathway segment (bmcmicrobiol.biomedcentral.com). As one review noted, the BenR–CatR dual system “may serve as a practical model for complex regulatory circuits” in biodegradation networks (bmcmicrobiol.biomedcentral.com).

Regulatory Function and Interactions

BenR acts primarily as a transcriptional activator. In the absence of benzoate, expression of the benzoate-catabolic enzymes is basal. When benzoate is present (or a structural analog capable of acting as effector), BenR binds the ligand and activates transcription of its target operon. Reporter fusion experiments have demonstrated that BenR is both necessary and sufficient for induction of the ben genes. For example, a benA–lacZ fusion in wild-type Pseudomonas showed strong induction (β-galactosidase activity increased an order of magnitude) upon adding benzoate, whereas an isogenic benR mutant showed no such induction (pmc.ncbi.nlm.nih.gov). Introducing a plasmid-encoded benR into the mutant restored benA expression and benzoate utilization (pmc.ncbi.nlm.nih.gov). These results provide direct evidence that BenR positively regulates the ben operon in response to benzoate (pmc.ncbi.nlm.nih.gov). BenR’s effector specificity appears tuned to benzoate and closely related aromatics – notably, benzoate itself is a known effector that also triggers the analogous XylS regulator on the TOL plasmid (pmc.ncbi.nlm.nih.gov). Methyl-substituted benzoates (e.g. 3-methylbenzoate) can likely serve as effectors as well, since benR was identified in mutants selected on 3-methylbenzoate (pmc.ncbi.nlm.nih.gov) and BenR was shown to activate the TOL meta-cleavage pathway (which XylS normally controls) in response to methylbenzoate (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). This indicates that BenR’s ligand-binding pocket accommodates benzoate and certain analogues, allowing it to regulate multiple aromatic acid degradation routes.

Regulon scope: Besides the chromosomal benzoate dioxygenase genes (benABCD), BenR has been implicated in controlling at least two other sets of genes in P. putida, highlighting a broader regulatory role. First, BenR can activate the meta-cleavage pathway operon (on the TOL plasmid) for methylbenzoate and benzoate degradation (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). The TOL plasmid’s meta operon (xyL genes) is normally activated by XylS when it senses benzoate or 3-methylbenzoate. BenR is a XylS homolog and recognizes a similar DNA sequence motif, and experiments showed that BenR can bind and activate the plasmid meta-operon promoter (Pm) in an E. coli reporter system (pmc.ncbi.nlm.nih.gov). This cross-talk means a chromosomal regulator (BenR) can induce plasmid-encoded pathways, potentially providing redundancy or additional control when both plasmid and chromosome pathways are present (pmc.ncbi.nlm.nih.gov). Second, BenR is required for benzoate-mediated repression of the pcaK gene, which encodes a 4-hydroxybenzoate (4-HBA) transporter (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). In P. putida, the presence of benzoate prevents uptake of 4-HBA by downregulating pcaK transcription – an example of prioritizing one substrate over another. A benR mutant abolishes this catabolite repression effect (pmc.ncbi.nlm.nih.gov), suggesting BenR somehow links benzoate sensing to reduced expression of the 4-HBA transport system. The mechanism appears indirect (possibly via accumulation of a metabolic intermediate or interplay with other regulators) (pmc.ncbi.nlm.nih.gov), but functionally it ensures that when benzoate is present, the cell preferentially metabolizes it rather than 4-HBA (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). In summary, BenR has a multifaceted regulatory role: it activates genes for benzoate ortho-cleavage (ben operon), activates genes for toluate/benzoate meta-cleavage (when the TOL plasmid is present), and contributes to repression of the 4-HBA (protocatechuate) pathway uptake during growth on benzoate (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). This coordination helps optimize aromatic substrate utilization based on availability.

Global regulation: The expression and activity of BenR itself are integrated into broader cellular regulatory networks. Pseudomonas species are known to exhibit carbon catabolite repression (CCR), ensuring preferred carbon sources are used first (bmcmicrobiol.biomedcentral.com). In P. putida, CCR is mediated by the Crc protein (Catabolite repression control), which can directly target benR. Crc binds to the 5′ end of benR mRNA and inhibits its translation, thereby lowering BenR protein levels when a preferred substrate (like succinate or a rich medium) is present (pubmed.ncbi.nlm.nih.gov) (pubmed.ncbi.nlm.nih.gov). This creates a hierarchy: if a readily metabolizable carbon source is available, BenR remains low and the benzoate catabolic genes are not expressed (even if benzoate is present) (pubmed.ncbi.nlm.nih.gov) (pubmed.ncbi.nlm.nih.gov). Studies showed that in a Crc-deficient strain, benA expression was de-repressed in the presence of benzoate, confirming that Crc’s effect on the benzoate pathway is exerted at the level of BenR translation (pubmed.ncbi.nlm.nih.gov) (pubmed.ncbi.nlm.nih.gov). Once preferred nutrients are exhausted, Crc repression lifts, BenR can be synthesized, and the ben operon gets induced by benzoate. This multilayer control underscores that BenR is part of a tightly regulated network. It responds to specific aromatic compounds, but its action is gated by global carbon status, allowing Pseudomonas to balance efficient growth with opportunistic degradation of aromatics (pubmed.ncbi.nlm.nih.gov) (bmcmicrobiol.biomedcentral.com).

Pathway Significance and Applications

Biological significance: The BenR-controlled benzoate degradation pathway is a key adaptive trait for soil and water bacteria like P. putida, P. fluorescens, P. stutzeri and others. Aromatic acids such as benzoate are common in soil (from plant phenolics, lignin breakdown, or pollutants), and the ability to use them as carbon sources gives these bacteria a competitive edge (bmcmicrobiol.biomedcentral.com) (bmcmicrobiol.biomedcentral.com). The β-ketoadipate pathway, to which BenR’s regulon feeds, is a central route for biodegradation of aromatics and is widely distributed in bacteria (bmcmicrobiol.biomedcentral.com) (bmcmicrobiol.biomedcentral.com). By regulating the first step that commits benzoate to this pathway, BenR plays an important role in environmental pollutant degradation. For example, P. putida can degrade toluene and xylenes via benzoate intermediates; BenR (chromosomal) works in tandem with plasmid-encoded systems to ensure these compounds are funneled into harmless end products (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). In P. stutzeri (a nitrogen-fixing rhizobacterium), the benR regulon likely helps the bacterium colonize plant roots by utilizing aromatic root exudates or soil humic substances as nutrients (bmcmicrobiol.biomedcentral.com). The importance of BenR is evident from the fact that knocking it out disables the utilization of benzoate in these bacteria (bmcmicrobiol.biomedcentral.com), which would impair their ability to exploit many aromatic compounds.

Biotechnological and research applications: The BenR-benzonate regulatory module has been harnessed as a useful tool in recombinant protein expression and synthetic biology. Because benzoate is an inexpensive, readily available inducer that is not a common carbon source in most lab media, the Pseudomonas benA promoter (activated by BenR) provides a tightly regulated expression system. Researchers identified the benABCD operon of P. fluorescens and its regulator BenR as a convenient inducible promoter system for heterologous gene expression (microbialcellfactories.biomedcentral.com) (microbialcellfactories.biomedcentral.com). By placing target genes under control of the benzoate-inducible promoter and co-expressing BenR, one can achieve strong, dose-dependent expression in Pseudomonas hosts upon addition of benzoate. In P. fluorescens MB214, for instance, the benzoate operon promoter showed very low background and high induction, allowing controlled protein production from shake-flask scale up to 20 L fermenters (microbialcellfactories.biomedcentral.com) (microbialcellfactories.biomedcentral.com). This system has been successfully applied to produce mammalian proteins in Pseudomonas and even adapted to other bacteria: for example, elements of the P. fluorescens BenR/Pben promoter have been engineered into Methylococcus (a methanotroph) to create a benzoate-inducible genetic switch (pseudoluge.pseudomonas.com) (academic.oup.com). The tight on/off control and inexpensive inducer make BenR-based regulators attractive for industrial biotechnology.

From a biodegradation standpoint, understanding BenR also allows scientists to manipulate pathways for bioremediation. By overexpressing or mutating regulators like BenR, it may be possible to enhance degradation capabilities of bacteria for environmental cleanup. Conversely, BenR’s sensitivity to global regulation (via Crc) makes it a target for optimizing metabolic flux: relieving Crc repression (for example, via an antagonist protein or regulatory RNA) could lead to higher expression of benzoate-catabolic enzymes even in mixed-substrate environments (pubmed.ncbi.nlm.nih.gov) (bmcmicrobiol.biomedcentral.com). These insights, drawn from recent molecular studies, demonstrate how BenR connects environmental sensing with metabolic response, and how it can be leveraged in practice.

Expert Commentary and Current Insights

Over two decades of research have solidified BenR’s role as the benzoate gatekeeper in pseudomonads. Early work in 2000 by Cowles et al. first characterized BenR in P. putida, showing that it controls three aspects of aromatic metabolism (benzoate ortho-cleavage, plasmid-encoded meta-cleavage, and 4-HBA uptake) (pmc.ncbi.nlm.nih.gov). They noted that “BenR thus has roles as an activator of benzoate degradation via ortho ring fission, as an activator of benzoate and methylbenzoate degradation via meta ring fission, and in repression of 4-HBA degradation” (pmc.ncbi.nlm.nih.gov). Subsequent genomic studies extended these findings across species. Huang et al. (2010) observed that P. stutzeri A1501 relies on an AraC-type BenR (61% identical to P. fluorescens BenR) for benzoate utilization, in the absence of the usual CatR regulator (bmcmicrobiol.biomedcentral.com) (bmcmicrobiol.biomedcentral.com). This highlights some evolutionary divergence – Pseudomonas uses an AraC/XylS strategy (BenR) for the ben operon, whereas other bacteria like Acinetobacter use a LysR-type activator (BenM) for a similar operon (bmcmicrobiol.biomedcentral.com). Despite different protein families, the functional outcome is the same: the benzoate-catabolic genes are transcriptionally silent until a specific regulator-effector complex (BenR–benzoate or BenM–benzoate) triggers their expression. In Pseudomonas, BenR’s interplay with CatR forms a clever circuit to sense both the substrate and its downstream product (bmcmicrobiol.biomedcentral.com). Experts consider this a prime example of coordinated regulation in biodegradation pathways (bmcmicrobiol.biomedcentral.com).

Current understanding: By 2023, the consensus is that BenR acts as the master switch for benzoate degradation in Pseudomonas. Its regulatory network has been mapped in detail, from DNA-binding sites to effector specificity and higher-level regulation. Ongoing research has focused on structural and systems-biology aspects: e.g. solving AraC-family regulator structures to understand ligand recognition, or modeling the catabolite repression system involving BenR. While no crystal structure of BenR has been reported yet (to our knowledge), homology to XylS and other AraC proteins provides a basis for predicting its ligand-binding pocket and dimerization interface (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov). On the systems side, recent analyses of global regulatory mutants (Crc, Hfq, etc.) illuminated how BenR-controlled pathways can be modulated for metabolic engineering (pubmed.ncbi.nlm.nih.gov) (pubmed.ncbi.nlm.nih.gov). There is also interest in bioresource applications: for instance, using BenR-regulated promoters to construct benzoate-inducible biosensors or kill-switches in synthetic biology. Given that benzoate is a benign chemical, inducible systems based on BenR are being tested in E. coli and other chassis for tight gene circuit control (microbialcellfactories.biomedcentral.com) (microbialcellfactories.biomedcentral.com).

In summary, BenR is a key transcriptional activator that links the presence of benzoate to the expression of enzymes that initiate its breakdown. It operates inside the cell by binding to DNA at target promoters and requires benzoate as a molecular trigger. Through BenR, Pseudomonas species effectively respond to aromatic compounds in their environment, converting pollutants like benzoate into metabolizable forms. This gene and its protein product have been well characterized in the literature, with clear experimental evidence for their function (e.g. loss-of-function mutations abolishing benzoate utilization (pmc.ncbi.nlm.nih.gov) (bmcmicrobiol.biomedcentral.com), and promoter assays demonstrating inducible activation (pmc.ncbi.nlm.nih.gov)). As our understanding has advanced, BenR has also become a practical tool in biotechnology, underscoring the value of fundamental gene function research. The continued study of BenR and related regulators not only deepens our knowledge of bacterial metabolism but also paves the way for innovative applications in bioremediation and synthetic biology.

References: Key studies and reviews include Cowles et al. 2000 (J. Bacteriol. 182:6339-6346) (pmc.ncbi.nlm.nih.gov) (pmc.ncbi.nlm.nih.gov), which first identified BenR’s role; Huang et al. 2010 (BMC Microbiol 10:36) describing benR in P. stutzeri (bmcmicrobiol.biomedcentral.com); and a 2006 study by Jahanian et al. (Microb. Cell Fact. 5:1) on using the benR promoter system in P. fluorescens (microbialcellfactories.biomedcentral.com). These and other works form the basis of the current functional annotation of BenR in Pseudomonas. Each provides experimental backing for BenR’s function, biological process involvement, cellular localization (DNA-binding in cytoplasm), and regulatory impact, as summarized above. The consistency of findings across different Pseudomonas strains and the agreement with bioinformatic predictions (e.g. conserved binding sites (bmcmicrobiol.biomedcentral.com) and gene context (pmc.ncbi.nlm.nih.gov)) give high confidence in this functional annotation. Overall, BenR is characterized as a benzoate-responsive transcriptional activator that is essential for initiating benzoate catabolism and integrating this pathway into the bacterium’s overall metabolic network (pmc.ncbi.nlm.nih.gov) (bmcmicrobiol.biomedcentral.com).

Citations

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  22. AnnotationURLCitation(end_index=7606, start_index=7437, title='The target for the Pseudomonas putida Crc global regulator in the benzoate degradation pathway is the BenR transcriptional regulator - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/18156252/#:~:text=products%20generated%20by%20the%20protocatechuate,ketoadipate%20as%20an%20effector%20%2817')
  23. AnnotationURLCitation(end_index=8160, start_index=8004, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=,sole%20activator%20of%20expression%20of')
  24. AnnotationURLCitation(end_index=8601, start_index=8425, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=match%20at%20L295%20,sole%20activator%20of%20expression%20of')
  25. AnnotationURLCitation(end_index=8914, start_index=8706, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=constructing%20and%20characterizing%20three%20mutants%3A,grow%20on%20benzoate%2C%20and%20the')
  26. AnnotationURLCitation(end_index=9377, start_index=9219, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=expression%20of%20catB%20in%20the,muconate')
  27. AnnotationURLCitation(end_index=9558, start_index=9378, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=involving%20the%20two%20sets%20of,a%20catR%20orthologue%20or%20a')
  28. AnnotationURLCitation(end_index=9836, start_index=9680, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=putative%20BenR,To%20determine%20whether')
  29. AnnotationURLCitation(end_index=10351, start_index=10187, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=substrates,a%20practical%20model%20for%20complex')
  30. AnnotationURLCitation(end_index=10548, start_index=10352, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=BenR%20and%20CatR%20jointly%20activate,in%20the%20biodegradation%20of%20benzoate')
  31. AnnotationURLCitation(end_index=10906, start_index=10710, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=BenR%20and%20CatR%20jointly%20activate,in%20the%20biodegradation%20of%20benzoate')
  32. AnnotationURLCitation(end_index=11243, start_index=11047, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=BenR%20and%20CatR%20jointly%20activate,in%20the%20biodegradation%20of%20benzoate')
  33. AnnotationURLCitation(end_index=12049, start_index=11930, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=match%20at%20L458%20fusion%20was,This')
  34. AnnotationURLCitation(end_index=12273, start_index=12152, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=,benABC%20in%20response%20to%20benzoate')
  35. AnnotationURLCitation(end_index=12520, start_index=12387, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=the%20wild,PCR%20amplification%20of%20the%20regions')
  36. AnnotationURLCitation(end_index=12883, start_index=12720, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=Pseudomonas%20putida%20converts%20benzoate%20to,A%20benR%20mutant%20isolated%20by')
  37. AnnotationURLCitation(end_index=13206, start_index=13043, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=putida%20benR%20mutant%20,kanamycin%2C%20100%3B%20spectinomycin%2C%20100%3B%20and')
  38. AnnotationURLCitation(end_index=13492, start_index=13329, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=Pseudomonas%20putida%20converts%20benzoate%20to,A%20benR%20mutant%20isolated%20by')
  39. AnnotationURLCitation(end_index=13623, start_index=13493, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=BenR%20thus%20has%20roles%20as,HBA%20degradation')
  40. AnnotationURLCitation(end_index=14289, start_index=14126, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=Pseudomonas%20putida%20converts%20benzoate%20to,A%20benR%20mutant%20isolated%20by')
  41. AnnotationURLCitation(end_index=14420, start_index=14290, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=BenR%20thus%20has%20roles%20as,HBA%20degradation')
  42. AnnotationURLCitation(end_index=14894, start_index=14743, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=benR%20activated%20expression%20of%20a,The%20benA%20promoter%20region')
  43. AnnotationURLCitation(end_index=15248, start_index=15095, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=fusion%20inserted%20in%20an%20E,HBA%20uptake%20is%20probably%20indirect')
  44. AnnotationURLCitation(end_index=15535, start_index=15386, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=also%20binds%20to%20the%20regulator,An%20additional%20seven%20genes')
  45. AnnotationURLCitation(end_index=15680, start_index=15536, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=fusion%20inserted%20in%20an%20E,The%20benA%20promoter%20region')
  46. AnnotationURLCitation(end_index=16053, start_index=15902, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=benR%20activated%20expression%20of%20a,The%20benA%20promoter%20region')
  47. AnnotationURLCitation(end_index=16430, start_index=16277, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=fusion%20inserted%20in%20an%20E,HBA%20uptake%20is%20probably%20indirect')
  48. AnnotationURLCitation(end_index=16711, start_index=16548, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=Pseudomonas%20putida%20converts%20benzoate%20to,A%20benR%20mutant%20isolated%20by')
  49. AnnotationURLCitation(end_index=16842, start_index=16712, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=BenR%20thus%20has%20roles%20as,HBA%20degradation')
  50. AnnotationURLCitation(end_index=17306, start_index=17158, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=regulatory%20gene%2C%20benR,by%20showing%20that%20BenR%20activates')
  51. AnnotationURLCitation(end_index=17437, start_index=17307, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=BenR%20thus%20has%20roles%20as,HBA%20degradation')
  52. AnnotationURLCitation(end_index=17946, start_index=17783, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=Aromatic%20compounds%20are%20not%20preferred,16')
  53. AnnotationURLCitation(end_index=18413, start_index=18246, title='The target for the Pseudomonas putida Crc global regulator in the benzoate degradation pathway is the BenR transcriptional regulator - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/18156252/#:~:text=translation%20of%20target%20genes%20by,allowing%20the%20hierarchical%20assimilation%20of')
  54. AnnotationURLCitation(end_index=18549, start_index=18414, title='The target for the Pseudomonas putida Crc global regulator in the benzoate degradation pathway is the BenR transcriptional regulator - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/18156252/#:~:text=respectively,benR%20mRNA%2C%20decreasing%20BenR%20levels')
  55. AnnotationURLCitation(end_index=18875, start_index=18730, title='The target for the Pseudomonas putida Crc global regulator in the benzoate degradation pathway is the BenR transcriptional regulator - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/18156252/#:~:text=several%20pathways%20for%20the%20assimilation,Crc%20inhibits%20the')
  56. AnnotationURLCitation(end_index=19043, start_index=18876, title='The target for the Pseudomonas putida Crc global regulator in the benzoate degradation pathway is the BenR transcriptional regulator - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/18156252/#:~:text=translation%20of%20target%20genes%20by,allowing%20the%20hierarchical%20assimilation%20of')
  57. AnnotationURLCitation(end_index=19420, start_index=19253, title='The target for the Pseudomonas putida Crc global regulator in the benzoate degradation pathway is the BenR transcriptional regulator - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/18156252/#:~:text=translation%20of%20target%20genes%20by,allowing%20the%20hierarchical%20assimilation%20of')
  58. AnnotationURLCitation(end_index=19562, start_index=19421, title='The target for the Pseudomonas putida Crc global regulator in the benzoate degradation pathway is the BenR transcriptional regulator - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/18156252/#:~:text=respectively,allowing%20the%20hierarchical%20assimilation%20of')
  59. AnnotationURLCitation(end_index=20129, start_index=19974, title='The target for the Pseudomonas putida Crc global regulator in the benzoate degradation pathway is the BenR transcriptional regulator - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/18156252/#:~:text=that%20Crc%20acts%20by%20reducing,substrate%20into%20the%20central%20pathway')
  60. AnnotationURLCitation(end_index=20334, start_index=20130, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=responsible%20for%20benzoate%20catabolism%20in,in%20the%20biodegradation%20of%20benzoate')
  61. AnnotationURLCitation(end_index=20969, start_index=20763, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=associated%20bacteria%20have%20been%20poorly,evidence%20that%20benzoate%20degradation%20is')
  62. AnnotationURLCitation(end_index=21178, start_index=20970, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=compounds%20has%20tremendous%20environmental%20significance,19%20genes%20involved%20in%20the')
  63. AnnotationURLCitation(end_index=21473, start_index=21331, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=The%20%CE%B2,baylyi%20ADP1')
  64. AnnotationURLCitation(end_index=21677, start_index=21474, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=peripheral%20pathways%20for%20the%20catabolism,the%20presence%20of%20their%20respective')
  65. AnnotationURLCitation(end_index=22171, start_index=22035, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=The%20TOL%20,also%20convert%20benzoate%20to%20catechol')
  66. AnnotationURLCitation(end_index=22302, start_index=22172, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=BenR%20thus%20has%20roles%20as,HBA%20degradation')
  67. AnnotationURLCitation(end_index=22702, start_index=22496, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=associated%20bacteria%20have%20been%20poorly,evidence%20that%20benzoate%20degradation%20is')
  68. AnnotationURLCitation(end_index=22984, start_index=22828, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=,sole%20activator%20of%20expression%20of')
  69. AnnotationURLCitation(end_index=23795, start_index=23621, title='Identification of anthranilate and benzoate metabolic operons of Pseudomonas fluorescens and functional characterization of their promoter regions | Microbial Cell Factories | Full Text', type='url_citation', url='https://microbialcellfactories.biomedcentral.com/articles/10.1186/1475-2859-5-1#:~:text=In%20an%20effort%20to%20identify,benABCD')
  70. AnnotationURLCitation(end_index=23987, start_index=23796, title='Identification of anthranilate and benzoate metabolic operons of Pseudomonas fluorescens and functional characterization of their promoter regions | Microbial Cell Factories | Full Text', type='url_citation', url='https://microbialcellfactories.biomedcentral.com/articles/10.1186/1475-2859-5-1#:~:text=the%20putative%20promoter%20regions%20to,antR%20or%20benR')
  71. AnnotationURLCitation(end_index=24576, start_index=24387, title='Identification of anthranilate and benzoate metabolic operons of Pseudomonas fluorescens and functional characterization of their promoter regions | Microbial Cell Factories | Full Text', type='url_citation', url='https://microbialcellfactories.biomedcentral.com/articles/10.1186/1475-2859-5-1#:~:text=The%20antABC%20and%20benABCD%20operons,antR%20or%20benR')
  72. AnnotationURLCitation(end_index=24721, start_index=24577, title='Identification of anthranilate and benzoate metabolic operons of Pseudomonas fluorescens and functional characterization of their promoter regions | Microbial Cell Factories | Full Text', type='url_citation', url='https://microbialcellfactories.biomedcentral.com/articles/10.1186/1475-2859-5-1#:~:text=Conclusion')
  73. AnnotationURLCitation(end_index=25209, start_index=25029, title='OrtholugeDB - Retrieve orthologs for gene or genes', type='url_citation', url='https://pseudoluge.pseudomonas.com/named/list/search?field=locus_tag&value=PSEBR_a1552#:~:text=OrtholugeDB%20,phosphate%20synthase%20%7C%20%7C%20RBB')
  74. AnnotationURLCitation(end_index=25345, start_index=25210, title='In vitro activity of imipenem/relebactam against Pseudomonas aeruginosa isolates recovered from ICU patients in Spain and Portugal (SUPERIOR and STEP studies) | Journal of Antimicrobial Chemotherapy | Oxford Academic', type='url_citation', url='https://academic.oup.com/jac/article-abstract/77/11/3163/6691802#:~:text=breakpoints%29%20detected%20during%20the')
  75. AnnotationURLCitation(end_index=26182, start_index=26028, title='The target for the Pseudomonas putida Crc global regulator in the benzoate degradation pathway is the BenR transcriptional regulator - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/18156252/#:~:text=regions%20of%20catR%20and%20pcaR,substrate%20into%20the%20central%20pathway')
  76. AnnotationURLCitation(end_index=26346, start_index=26183, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=Aromatic%20compounds%20are%20not%20preferred,16')
  77. AnnotationURLCitation(end_index=27035, start_index=26887, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=regulatory%20gene%2C%20benR,by%20showing%20that%20BenR%20activates')
  78. AnnotationURLCitation(end_index=27391, start_index=27260, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=match%20at%20L102%20BenR%20thus,HBA%20degradation')
  79. AnnotationURLCitation(end_index=27812, start_index=27654, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=sequence%20of%20BenR%20in%20A1501,stutzeri')
  80. AnnotationURLCitation(end_index=27995, start_index=27813, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=match%20at%20L487%20involving%20the,a%20catR%20orthologue%20or%20a')
  81. AnnotationURLCitation(end_index=28406, start_index=28210, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=gene%20products%20are%20highly%20conserved,chromosomal%20ben%20and%20cat%20genes')
  82. AnnotationURLCitation(end_index=28933, start_index=28769, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=substrates,a%20practical%20model%20for%20complex')
  83. AnnotationURLCitation(end_index=29222, start_index=29026, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=BenR%20and%20CatR%20jointly%20activate,in%20the%20biodegradation%20of%20benzoate')
  84. AnnotationURLCitation(end_index=30063, start_index=29900, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=Amino%20acid%20sequence%20alignment%20of,Similar%20residues%20are%20shaded%20gray')
  85. AnnotationURLCitation(end_index=30164, start_index=30064, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=benR%20%20,27%2F44')
  86. AnnotationURLCitation(end_index=30500, start_index=30333, title='The target for the Pseudomonas putida Crc global regulator in the benzoate degradation pathway is the BenR transcriptional regulator - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/18156252/#:~:text=translation%20of%20target%20genes%20by,allowing%20the%20hierarchical%20assimilation%20of')
  87. AnnotationURLCitation(end_index=30642, start_index=30501, title='The target for the Pseudomonas putida Crc global regulator in the benzoate degradation pathway is the BenR transcriptional regulator - PubMed', type='url_citation', url='https://pubmed.ncbi.nlm.nih.gov/18156252/#:~:text=respectively,allowing%20the%20hierarchical%20assimilation%20of')
  88. AnnotationURLCitation(end_index=31173, start_index=30984, title='Identification of anthranilate and benzoate metabolic operons of Pseudomonas fluorescens and functional characterization of their promoter regions | Microbial Cell Factories | Full Text', type='url_citation', url='https://microbialcellfactories.biomedcentral.com/articles/10.1186/1475-2859-5-1#:~:text=The%20antABC%20and%20benABCD%20operons,antR%20or%20benR')
  89. AnnotationURLCitation(end_index=31318, start_index=31174, title='Identification of anthranilate and benzoate metabolic operons of Pseudomonas fluorescens and functional characterization of their promoter regions | Microbial Cell Factories | Full Text', type='url_citation', url='https://microbialcellfactories.biomedcentral.com/articles/10.1186/1475-2859-5-1#:~:text=Conclusion')
  90. AnnotationURLCitation(end_index=32077, start_index=31945, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=The%20benR%20mutant%20,The%20complemented%20mutant')
  91. AnnotationURLCitation(end_index=32234, start_index=32078, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=,sole%20activator%20of%20expression%20of')
  92. AnnotationURLCitation(end_index=32444, start_index=32291, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=To%20test%20whether%20BenR%20regulates,higher%20in%20cells%20grown%20on')
  93. AnnotationURLCitation(end_index=32993, start_index=32903, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=Abstract')
  94. AnnotationURLCitation(end_index=33142, start_index=32994, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=regulatory%20gene%2C%20benR,by%20showing%20that%20BenR%20activates')
  95. AnnotationURLCitation(end_index=33435, start_index=33259, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=match%20at%20L295%20,sole%20activator%20of%20expression%20of')
  96. AnnotationURLCitation(end_index=33747, start_index=33558, title='Identification of anthranilate and benzoate metabolic operons of Pseudomonas fluorescens and functional characterization of their promoter regions | Microbial Cell Factories | Full Text', type='url_citation', url='https://microbialcellfactories.biomedcentral.com/articles/10.1186/1475-2859-5-1#:~:text=The%20antABC%20and%20benABCD%20operons,antR%20or%20benR')
  97. AnnotationURLCitation(end_index=34337, start_index=34181, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=putative%20BenR,To%20determine%20whether')
  98. AnnotationURLCitation(end_index=34489, start_index=34355, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=match%20at%20L319%202%29%20%2817%29,3%3B%20Table%202')
  99. AnnotationURLCitation(end_index=34889, start_index=34758, title='BenR, a XylS Homologue, Regulates Three Different Pathways of Aromatic Acid Degradation in Pseudomonas putida - PMC', type='url_citation', url='https://pmc.ncbi.nlm.nih.gov/articles/PMC94779/#:~:text=match%20at%20L102%20BenR%20thus,HBA%20degradation')
  100. AnnotationURLCitation(end_index=35054, start_index=34890, title='Genome-wide investigation and functional characterization of the β-ketoadipate pathway in the nitrogen-fixing and root-associated bacterium Pseudomonas stutzeriA1501 | BMC Microbiology | Full Text', type='url_citation', url='https://bmcmicrobiol.biomedcentral.com/articles/10.1186/1471-2180-10-36#:~:text=substrates,a%20practical%20model%20for%20complex')