fleQ

UniProt ID: Q88ET0
Organism: Pseudomonas putida KT2440
Review Status: COMPLETE
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Gene Description

FleQ is an NtrC-family, sigma-54-associated AAA+ transcriptional regulator that acts as the master controller of flagellar biogenesis in Pseudomonas putida KT2440. Under low c-di-GMP it activates the flagellar transcriptional cascade through sigma-N-dependent promoters, whereas c-di-GMP-responsive promoter control redirects FleQ activity toward adhesin and exopolysaccharide loci such as lapA, lapE, and bcs to coordinate the transition from motility to surface attachment and biofilm growth. ChIP-seq and targeted promoter studies support a broader regulon that includes additional adhesion, exopolysaccharide, iron-homeostasis, and type VI secretion genes, but the clearest core functions remain flagellum regulation and c-di-GMP-responsive control of biofilm-associated transcription.

Proposed New Ontology Terms

sigma-54-dependent DNA-binding transcription activator activity

Definition: Catalysis of transcriptional activation by a bacterial enhancer-binding protein that binds specific DNA sequences and uses ATP-dependent remodeling to activate RNA polymerase containing sigma-54 at target promoters.

Justification: FleQ and related enhancer-binding proteins are currently forced into generic DNA-binding transcription activator activity and ATP binding terms, which do not capture the sigma-54-specific promoter class or ATP-dependent remodeling mechanism.

Parent term: DNA-binding transcription activator activity

Existing Annotations Review

GO Term Evidence Action Reason
GO:0005524 ATP binding
IEA
GO_REF:0000002
ACCEPT
Summary: Accurate and core. FleQ is an AAA+ enhancer-binding transcriptional regulator with a central ATPase/sigma-54 interaction domain, and ATP binding is mechanistically required for its transcriptional control of flagellar promoters.
Reason: Although generic, ATP binding is part of the core mechanism of a sigma-54 bacterial enhancer-binding protein rather than an incidental property.
Supporting Evidence:
file:PSEPK/fleQ/fleQ-uniprot.txt
UniProt and InterPro annotate FleQ, AAA+ ATPase, and sigma-54 interaction domains in Q88ET0
PMID:30177764
central AAA+/ATPase Οƒ54 (RpoN)-interaction domain
file:PSEPK/fleQ/fleQ-deep-research-falcon.md
**FleN is required for full c-di-GMP-dependent activation** at certain matrix promoters and that **ATP hydrolysis is required for flagellar gene expression**, but not necessarily for matrix gene regulation.
GO:0006355 regulation of DNA-templated transcription
IEA
GO_REF:0000002
KEEP AS NON CORE
Summary: Correct but overly broad for FleQ. The gene product is a global transcriptional regulator with separable flagellar-activation and biofilm-control branches, so this parent term is best retained as non-core while more informative child terms are added below.
Reason: FleQ clearly regulates transcription, but this term is too unspecific to represent the biologically informative functions of the protein.
Supporting Evidence:
PMID:27636892
coordinate regulation of multiple functions related to motility and surface colonization in Pseudomonas putida
PMID:30177764
Putative genes regulated by FleQ included, as expected, flagellar and motility-related genes
file:PSEPK/fleQ/fleQ-deep-research-openai.md
master activator of flagellar biogenesis in pseudomonads
file:PSEPK/fleQ/fleQ-deep-research-falcon.md
FleQ acts as a **bifunctional regulator**: typically activating motility/adhesion genes and repressing some EPS genes.
GO:0043565 sequence-specific DNA binding
IEA
GO_REF:0000002
ACCEPT
Summary: Accurate and core. FleQ directly binds specific promoters in the flagellar region and at biofilm-associated loci including lapA, bcs, lapE, and cyaA, establishing sequence-specific promoter recognition as a central molecular function.
Reason: Direct DNA-binding evidence exists in KT2440 for multiple named targets, and this function underlies both the motility and biofilm branches of FleQ regulation.
Supporting Evidence:
PMID:27120564
In vitro experiments show that FleQ binds to lapA and bcs promoter DNA
PMID:33975969
Both in vitro and in vivo assays show that FleQ binds directly to promoters of the five genes
file:PSEPK/fleQ/fleQ-deep-research-falcon.md
promoter fragments predicted to contain FleQ boxes shifted upon incubation with purified FleQ, while control fragments lacking predicted motifs did not.
GO:0001216 DNA-binding transcription activator activity
IDA
PMID:27636892
Complex Interplay between FleQ, Cyclic Diguanylate and Multi...
NEW
Summary: FleQ is a bona fide DNA-binding transcription activator for the flagellar cascade. Under low c-di-GMP it activates sigma-N-dependent promoters in the flagellar regulon, and multiple KT2440 studies describe FleQ as the master activator of flagellar gene expression.
Reason: The seeded IEA set captures DNA binding and generic transcription regulation but misses FleQ's specific activator role.
Supporting Evidence:
PMID:27636892
Low cyclic diguanylate levels favor FleQ interaction with ΟƒN-dependent promoters to activate the flagellar cascade
PMID:30177764
It has been described that FleQ is able to activate the expression of genes involved in flagellar export
file:PSEPK/fleQ/fleQ-deep-research-falcon.md
FleQ directly activated a subset of Οƒ54-class promoters, including **PflgB, PflhA, and PflgA**
GO:1902210 positive regulation of bacterial-type flagellum assembly
IMP
PMID:30177764
Genome-wide analysis of the FleQ direct regulon in Pseudomon...
NEW
Summary: FleQ positively regulates flagellum assembly by driving transcription of flagellar biogenesis genes. fleQ mutants are non-motile and aflagellated, and ChIP-seq/promoter analyses place FleQ directly upstream of multiple flagellar loci required for assembly.
Reason: This term captures the specific biological outcome of FleQ's best-established regulatory role better than generic motility or transcription terms.
Supporting Evidence:
PMID:30177764
mutations in the fleQ gene result in non-motile, aflagellated bacteria
PMID:28968917
FleQ positively regulates flagella biosynthesis but negatively regulates the type VI secretion system in P. putida
file:PSEPK/fleQ/fleQ-deep-research-falcon.md
In KT2440-focused work, FleQ activates multiple flagellar promoters and sits at the top of a multi-tier regulatory cascade.
GO:1900190 regulation of single-species biofilm formation
IMP
PMID:27636892
Complex Interplay between FleQ, Cyclic Diguanylate and Multi...
NEW
Summary: FleQ directly regulates the transition into the biofilm state by controlling transcription of adhesin and matrix-associated loci such as lapA, lapE, and bcs in a c-di-GMP-responsive manner. The literature shows clear FleQ dependence for biofilm formation, but because FleQ can activate some matrix genes and repress others, the broader regulation term is safer than forcing a purely positive label.
Reason: Biofilm control is a major experimentally supported branch of FleQ function, but the directionality is promoter- and c-di-GMP-state dependent.
Supporting Evidence:
PMID:27636892
Disruption of fleQ caused strong defects in flagellar motility, biofilm formation and surface attachment
PMID:27120564
FleQ acts as an activator of lapA, but a repressor of bcs
PMID:33975969
These results demonstrated that FleQ was essential for c-di-GMP-mediated biofilm formation and LapA secretion
file:PSEPK/fleQ/fleQ-deep-research-falcon.md
Deletion of fleQ caused a **~2-fold drop in lapA mRNA** and a **~32-fold increase in bcsD mRNA**.
GO:0035438 cyclic-di-GMP binding
IDA
PMID:27503246
FleQ of Pseudomonas putida KT2440 is a multimeric cyclic dig...
NEW
Summary: FleQ of P. putida KT2440 directly binds the second messenger c-di-GMP, as demonstrated by isothermal titration calorimetry. c-di-GMP binding to the AAA+ ATPase module is the molecular switch that redirects FleQ between flagellar activation and biofilm/matrix gene control, making this a core molecular function distinct from generic ATP binding.
Reason: The seeded IEA set captures ATP binding but misses FleQ's experimentally demonstrated direct c-di-GMP binding, which is the key ligand-sensing function underlying its lifestyle-switch behavior.
Supporting Evidence:
PMID:27503246
we show in this work that FleQ of P. putida interacts with c-di-GMP and directly binds the promoter regions of flagellar and EPS genes
file:PSEPK/fleQ/fleQ-deep-research-falcon.md
FleQ is also described as a **c-di-GMP-binding** multimeric regulator, consistent with AAA+_ATPase, FleQ, and CheY-like/receiver-like domain annotations.
file:PSEPK/fleQ/fleQ-deep-research-falcon.md
**c-di-GMP** binds FleQ’s AAA+ region/Walker A-associated ATPase module and **inhibits ATPase activity**, shifting FleQ from flagellar activation toward biofilm-associated regulation.

Core Functions

ATP-dependent DNA-binding transcription activator that initiates the sigma-54-dependent flagellar transcriptional cascade and thereby promotes bacterial-type flagellum assembly

Supporting Evidence:
  • PMID:27636892
    Low cyclic diguanylate levels favor FleQ interaction with ΟƒN-dependent promoters to activate the flagellar cascade
  • PMID:30177764
    It has been described that FleQ is able to activate the expression of genes involved in flagellar export

c-di-GMP-responsive promoter-binding regulator that coordinates surface attachment and matrix-gene expression, especially through the lapA-lapE and bcs branches, during the transition to single-species biofilm growth

Supporting Evidence:
  • PMID:27120564
    In vitro experiments show that FleQ binds to lapA and bcs promoter DNA
  • PMID:33975969
    These results demonstrated that FleQ was essential for c-di-GMP-mediated biofilm formation and LapA secretion

Direct cyclic-di-GMP binding at the AAA+ ATPase module, which allosterically inhibits FleQ ATPase activity and acts as the second-messenger switch that redirects FleQ between flagellar activation and biofilm/matrix gene control

Supporting Evidence:
  • PMID:27503246
    we show in this work that FleQ of P. putida interacts with c-di-GMP and directly binds the promoter regions of flagellar and EPS genes
  • file:PSEPK/fleQ/fleQ-deep-research-falcon.md
    A 2024 authoritative review synthesizes structural and biochemical evidence (primarily from *P. aeruginosa* but presented as conserved across *Pseudomonas*) that **c-di-GMP binds the AAA+ ATPase domain of FleQ** and allosterically **inhibits ATPase activity**, enabling FleQ to switch from flagellar activation to matrix gene activation/repression at a single promoter without dissociation.

References

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Suggested Questions for Experts

Q: Which FleQ-regulated iron-homeostasis genes are direct physiological targets in KT2440 under rhizosphere-relevant iron limitation, and are these roles core or context-specific?

Q: What promoter or cofactor features determine whether c-di-GMP-bound FleQ activates versus represses a given surface-associated operon in P. putida?

Q: How much of the reported type VI secretion phenotype reflects direct FleQ control versus secondary remodeling of the motility-biofilm regulatory network?

Suggested Experiments

Experiment: ChIP-seq across low- and high-c-di-GMP states with matched RNA-seq in KT2440 to separate constitutive FleQ binding from c-di-GMP-dependent transcriptional output

Hypothesis: FleQ occupancy will remain at many promoters while c-di-GMP changes the transcriptional outcome and cofactor usage rather than simple DNA binding

Type: Differential ChIP-seq plus transcriptomics

Experiment: Promoter mutagenesis and in vitro transcription assays for fleS-fliF, flhA, lapA, bcs, and lapE promoters using purified FleQ with ATP, c-di-GMP, and FleN

Hypothesis: Distinct promoter architectures encode the activator-versus-repressor logic of FleQ and explain the opposite responses of lapA and bcs

Type: Promoter mutagenesis, EMSA, and in vitro transcription

Experiment: Separation-of-function FleQ mutants targeting ATPase versus c-di-GMP-binding surfaces tested for motility, attachment, biofilm, and root colonization phenotypes

Hypothesis: ATPase-deficient alleles will preferentially collapse the flagellar branch, whereas c-di-GMP-response mutants will uncouple motility from biofilm regulation

Type: Allelic replacement with phenotypic profiling

Deep Research

Falcon

(fleQ-deep-research-falcon.md)

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OpenAI

(fleQ-deep-research-openai.md)

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πŸ“š Additional Documentation

Notes

(fleQ-notes.md)

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