proteorhodopsin phototrophy

traitmech:000036 · CLASS · REVIEWED

A light-harvesting metabolism in which a retinal-containing membrane protein (proteorhodopsin) acts as a light-driven proton pump, generating proton motive force without chlorophyll-based reaction centers. Widespread among marine bacterioplankton.

Trait evidence (2)

  • DOI:10.1126/science.289.5486.1902

    Béjà et al. identified proteorhodopsin, a retinal-binding light-driven proton pump in an uncultivated marine bacterium, as evidence for a new type of phototrophy in the sea.

  • DOI:10.1038/35081051

    Béjà et al., "Proteorhodopsin phototrophy in the ocean", supports proteorhodopsin as a widespread, spectrally tuned light-energy capture system in marine bacteria.

Proteorhodopsin pumps protons under illumination, generating PMF

Evidence-backed causal sketch linking retinal-based proteorhodopsin light-driven proton pumping to proton motive force without chlorophyll-based reaction centers.

MECHANISTIC · Represents the directly demonstrated retinal-dependent light-driven proton pump and downstream proton-motive-force branch; the exemplar is the original EBAC31A08 proteorhodopsin.

Proteorhodopsin pumps protons under illumination, generating PMF Interactive directed graph showing evidence-backed causal relationships for proteorhodopsin phototrophy.

Edge evidence

  • proteorhodopsin confers proteorhodopsin phototrophy METPO:2007700

    The retinal-binding pump realizes proteorhodopsin phototrophy.

    • DOI:10.1126/science.289.5486.1902 bound retinal to form an active, light-driven proton pump Béjà et al. identified proteorhodopsin as a retinal-binding light-driven proton pump.
  • proteorhodopsin phototrophy contributes to proton motive force RO:0002326

    Light-driven proton pumping by proteorhodopsin generates PMF.

    • DOI:10.1073/pnas.0611035104 Illumination of these cells with light coinciding with PR's absorption spectrum creates a proton motive force (pmf) A primary physiological study directly measured light-dependent PMF generation in proteorhodopsin-expressing cells.
  • light-driven proton pumping generates proton motive force biolink:produces

    Light-driven proton pumping by proteorhodopsin drives PMF generation.

    • DOI:10.1073/pnas.0611035104 Illumination of these cells with light coinciding with PR's absorption spectrum creates a proton motive force (pmf) The measured light-dependent PMF is the direct output of the represented proton-pumping process.
  • proton motive force enables ATP production RO:0002327

    PR-mediated proton transport generates a membrane potential sufficient for ATP production.

    • DOI:10.1073/pnas.0611035104 Sinks include rotation of the flagellar motor and ATP synthesis. The primary study's PMF model identifies ATP synthesis as a sink for the proteorhodopsin-generated motive force.
  • proteorhodopsin phototrophy does not produce NAD(P)H METPO:2007801

    Proteorhodopsin phototrophy cannot generate NAD(P)H for anabolic metabolism; ATP-only energy module.

    • DOI:10.4014/jmb.2410.10034 the PR function of strain IMCC1322, which supplies only ATP. Oh 2024: "PR can never be harnessed to generate NAD(P)H for anabolic metabolism".
  • all-trans retinal required for proteorhodopsin

    Functional proteorhodopsin requires all-trans retinal bound via a Schiff base to a central lysine.

    • DOI:10.1038/s41467-024-50960-3 all-trans retinal needs to be incorporated by forming a Schiff base with a central lysine in the chromophore binding pocket. Hirschi 2024: "Functional proteorhodopsin requires insertion of all-trans retinal by formation of a Schiff base with a central lysine".
  • beta-carotene cleaved to produce all-trans retinal

    Beta-carotene is cleaved by dioxygenase into retinal, an essential element of functional PR.

    • DOI:10.1128/MMBR.69.1.51-78.2005 converts β,β-carotene into retinal Jing 2022: "beta-carotene can be cleaved by dioxygenase into retinal, which is an important element in functional PR".

Protein and taxon examples

Graph nodeProteinTaxonUniProt statusRole and evidence
proteorhodopsin UniProtKB:Q9F7P4
Green-light absorbing proteorhodopsin
gamma proteobacterium EBAC31A08
NCBITaxon:133804
REVIEWED
retrieved 2026-08-23 · entry v97 · sequence v1

Retinal-binding light-driven proton pump that establishes the core energy-transduction mechanism represented in this graph.

  • DOI:10.1126/science.289.5486.1902 primarily as a light-driven proton pump The discovery study functionally characterized proteorhodopsin from environmental clone EBAC31A08; UniProtKB Q9F7P4 verifies the reviewed protein and source taxon.

Provenance

Identifier source
TraitMech local identifier
Definition source
DOI:10.1126/science.289.5486.1902

Parent traits (1)

Synonyms (1)

  • rhodopsin-based phototrophy RELATED_SYNONYM · DOI:10.1038/35081051

kg-microbe context

Matched 1 kg-microbe node via parent_proxy.

  • METPO:1000060 [-1.052, -1.766, -1.194, +0.291, …]

512-dim DeepWalkSkipGramEnsmallen embedding from kg-microbe (2026-04-25).

Nearest neighbors in embedding space

Top-8 cosine-similar METPO traits from the 2026-04-25 deepwalk (512-D).

Deep research

Generated by just research-trait; source: research/traits/metabolism/proteorhodopsin_phototrophy-deep-research-falcon.md

Unreviewed literature output — not curated TraitMech content Ontology identifiers suggested below have not been resolved against their ontologies, and some are known to be wrong. Check any CURIE against the source before using it.
# Curation report: proteorhodopsin phototrophy

## Trait record and scope

- **Trait label:** proteorhodopsin phototrophy
- **Trait identifier:** **`traitmech:000036`**
- **Category / kind / status:** METABOLISM / CLASS / REVIEWED
- **Parent:** `traitmech:000037`
- **Synonym:** rhodopsin-based phototrophy

### Recommended operational definition

Proteorhodopsin phototrophy is the physiological capacity to use a **retinal-bound, outward proton-pumping proteorhodopsin in the cytoplasmic membrane** to convert absorbed light into a transmembrane electrochemical proton gradient. That proton-motive force can support ATP synthesis and, depending on organism and conditions, transport, motility, maintenance, growth, or survival. It is generally an **energy-supplementing photoheterotrophic mechanism**, not carbon-fixing photosynthesis: proteorhodopsin supplies energy but does not itself provide a reaction center, electron-transfer chain, reducing power, or carbon-fixation pathway. PR-bearing SAR86 genomes, for example, lack autotrophic carbon-fixation pathways despite having PR-based energetic potential. (feng2023isproteorhodopsina pages 17-23, dupont2012genomicinsightsto pages 8-9)

The minimum mechanistic phenotype should require evidence for functional light-driven proton translocation, or a sufficiently complete genotype plus functional validation. A proteorhodopsin-like sequence alone is not enough: retinal availability, membrane expression, pump direction, and ion specificity can determine whether the phenotype exists. SAR86 illustrates this distinction because some genomes carry PR but apparently lack retinal biosynthesis and may have to acquire the chromophore externally. (dupont2012genomicinsightsto pages 8-9)

### Boundary cases

**Include:** outward H⁺-pumping bacterial proteorhodopsins that bind retinal and generate proton motive force under illumination, including systems that principally improve maintenance or stress survival rather than measurable exponential growth.

**Exclude or model separately:**

1. **Sensory rhodopsins**, light-gated channels, inward proton pumps, and chloride/sodium pumps; sequence homology to microbial rhodopsins does not establish PR phototrophy.
2. **Chlorophyll-based photosynthesis and aerobic anoxygenic phototrophy**, which use reaction centers and electron-transfer machinery absent from the minimal PR system.
3. **Retinal biosynthesis alone** or an unexpressed PR gene.
4. **Viral or eukaryotic proton-pumping rhodopsins** unless TraitMech explicitly intends the trait to extend beyond cellular bacterial PR systems. Viral rhodopsins can bind retinal and pump protons, but are evolutionarily and host-contextually distinct. (needham2019adistinctlineage pages 10-10)
5. **Autotrophy as an intrinsic consequence.** A 2024 culture study found no significant light–dark difference in inorganic-carbon assimilation, showing that PR-dependent ATP generation does not automatically imply carbon fixation.

| Subject | Predicate | Object | Confidence | Key evidence |
|---|---|---|---|---|
| light | activates | retinal-bound proteorhodopsin | High | PR is a "light-driven proton pump" whose mechanism depends on retinal/photoisomerization; foundational identification in marine bacteria and later mechanistic summaries support light activation (DOI:10.1126/science.289.5486.1902; DOI:10.1038/35081051; DOI:10.25959/23241740) (feng2023isproteorhodopsina pages 17-23, feng2023isproteorhodopsina pages 28-34) |
| retinal-bound proteorhodopsin | exports | protons (H+) across the membrane | High | Direct functional evidence from illuminated cells and heterologous expression shows outward proton pumping / external pH change; recent work summarizes PR as bacterial proton pumps (DOI:10.1128/AEM.02425-09; DOI:10.25959/23241740; DOI:10.1126/sciadv.adu5303) (johnson2010enhancementofsurvival pages 1-2, feng2023isproteorhodopsina pages 89-95, bukhdruker2025proteorhodopsininsightsinto pages 18-19) |
| proton export by proteorhodopsin | generates | proton motive force | High | Multiple sources explicitly state PR translocates protons to create membrane potential/PMF; direct in engineered Shewanella, established across PR literature (DOI:10.1128/AEM.02425-09; DOI:10.25959/23241740) (johnson2010enhancementofsurvival pages 1-2, feng2023isproteorhodopsina pages 28-34, feng2023isproteorhodopsina pages 17-23) |
| proton motive force | drives | F-type ATP synthase | Medium-High | Established bioenergetic coupling rather than directly reconstituted in every PR study: sources state PR-generated PMF drives ATP synthesis via F0F1-ATPase / proton-translocating ATPase (DOI:10.25959/23241740; DOI:10.1128/AEM.02425-09) (feng2023isproteorhodopsina pages 28-34, feng2023isproteorhodopsina pages 17-23, feng2023isproteorhodopsina pages 34-38, johnson2010enhancementofsurvival pages 1-2) |
| F-type ATP synthase | produces | ATP | High | Established ATP synthase function; PR studies cite ATP increases or ATP synthesis supported by PR-generated PMF in heterologous/native systems (DOI:10.1128/AEM.02425-09; DOI:10.25959/23241740) (johnson2010enhancementofsurvival pages 1-2, feng2023isproteorhodopsina pages 28-34) |
| beta-carotene cleavage / retinal supply | enables | functional proteorhodopsin | High | Genome and biochemical evidence link PR function to retinal availability: PR-containing marine genomes carry retinal biosynthesis genes including blh and carotenoid genes; PR requires retinal/retinol for functionality (DOI:10.1073/pnas.0712027105; DOI:10.1038/ismej.2011.189) (dupont2012genomicinsightsto pages 8-9) |


*Table: Compact curation table of the strongest mechanistic edges for proteorhodopsin phototrophy, from light activation and proton export to PMF-coupled ATP production, plus retinal supply. It distinguishes directly shown PR steps from the broader established ATP synthase coupling used in TraitMech curation.*

## Candidate graph nodes

Identifiers below are conservative suggestions; exact ontology labels and releases should be checked during repository validation. No identifier is proposed for a strain-specific PR protein without a verified accession.

### Genes, proteins, and complexes

| Candidate node | Role | Suggested grounding |
|---|---|---|
| proteorhodopsin gene (`pr`) | Encodes the retinal-binding outward H⁺ pump | Label-only unless a taxon-specific gene/protein accession is used |
| proteorhodopsin holoprotein | Light-activated integral-membrane proton pump | GO:0015078, proton transmembrane transporter activity; optionally label-only for protein identity |
| proteorhodopsin apoprotein | PR before retinal incorporation | Label-only |
| retinal–proteorhodopsin Schiff-base complex | Photoactive holoprotein state | Label-only |
| conserved retinal-binding lysine | Forms the protonated Schiff base with retinal | Label-only; residue numbering is protein-specific (reported as Lys230 in one PR context) (feng2023isproteorhodopsina pages 17-23) |
| `blh` β-carotene 15,15′-dioxygenase | Cleaves β-carotene to supply retinal | EC:1.13.11.63; use a verified UniProt accession only for a specified organism |
| `crtE`, `crtB`, `crtI`, `crtY` | Carotenoid/β-carotene precursor synthesis | Gene-label nodes unless exact enzyme reaction and taxon are specified |
| F-type H⁺-transporting ATP synthase | Uses PMF to phosphorylate ADP | GO:0015986, ATP synthesis coupled proton transport |
| carotenoid–PR antenna complex | Expanded light harvesting in certain Bacteroidota PRs | Label-only; recent and taxon-restricted |

Polaribacter sp. MED152 contains PR together with carotenoid/retinal synthesis genes `crtEBIY` and `blh`, supporting inclusion of a retinal-supply module, although not every PR-bearing genome contains that complete module. (dupont2012genomicinsightsto pages 8-9)

Showing the first 60 of 227 lines of findings; the linked file also carries the run's front matter and the prompt it was given — read the full report.

Canonical examples (2)

Organisms cited as exemplars of this trait. Taxon ids are NCBITaxon and link out to the NCBI record.

  • Dokdonia sp. MED134 NCBITaxon:313590 PMID:17215843 Canonical proteorhodopsin phototroph: light stimulates growth under oligotrophic conditions (retinal-inhibition causally proven). Ca. Pelagibacter ubique uses PR for starvation maintenance.
  • gamma proteobacterium EBAC31A08 NCBITaxon:133804 DOI:10.1126/science.289.5486.1902 Source organism for the first functionally characterized green-light-absorbing proteorhodopsin sequence.

Curation history

  1. · CURATE_PROTEIN_TAXON_EXAMPLE · codex

    Reviewed the graph as mechanistic and added DOI-backed, taxon-paired UniProtKB Q9F7P4 as the EBAC31A08 proteorhodopsin exemplar.

  2. · PROPOSED_FROM_RESEARCH · claude

    Proposed candidate METABOLISM trait (proteorhodopsin phototrophy) from literature research to fill the light-driven-energy gap.

  3. · CURATED_CAUSAL_GRAPH · claude

    Added evidence-backed causal graph (proteorhodopsin light-driven proton pump) with METPO node grounding and RO predicate groundings; promoted PROPOSED to REVIEWED.

  4. · GROUND_CAUSAL_NODES · claude

    Grounded 1 causal-node grounding field(s) via mappings/node_grounding.tsv (UniProtKB:A0A1D7QWW9×1).

  5. · ENRICH_CAUSAL_GRAPH · claude

    Added 5 evidence-backed generic edges (5 new nodes) from the deep-research report.

  6. · GROUND_CAUSAL_PREDICATES · claude

    Grounded 2 causal-edge predicate_id field(s) via mappings/predicate_grounding.tsv (biolink:produces×1, RO:0002327×1).

  7. · GROUND_CAUSAL_NODES · claude

    Grounded 2 causal-node grounding field(s) via mappings/node_grounding.tsv (CHEBI:13392×1, CHEBI:17579×1).

  8. · GROUND_CAUSAL_PREDICATES · claude

    Grounded 1 causal-edge predicate_id field(s) via mappings/predicate_grounding.tsv (METPO:2000222×1).

  9. · MIGRATE_ENABLES_TRAIT_EDGES · claude

    Migrated 1 causal edge(s) off enables/RO:0002327 with a TRAIT object (1 to confers), issue 302. RO:0002327 has range 'biological process or activity', which a trait (a disposition) cannot satisfy, so the previous form entailed trait is-a BiologicalProcessOrActivity. The replacements are proposed in proposals/metpo_traitmech_v8 and are placeholder ids until METPO mints them.

  10. · MIGRATE_MICROBE_DOMAIN_EDGES_PART2 · claude

    Re-grounded 1 causal edge(s) off microbe-domain METPO predicates onto their causal-graph counterparts (1 to does not produce), issue 301 part 2. The previous predicates are transitively rdfs:subPropertyOf METPO:2000001, whose rdfs:domain is METPO:1000525 (microbe), so a causal-graph subject entailed that the subject IS a microbe; CausalNodeTypeEnum has no organism member, so no such edge could ever satisfy the domain. Each replacement is a 1:1 mirror of its source predicate that changes only the domain, so the claim each edge makes is unchanged and directions are unchanged. The replacements are proposed in proposals/metpo_traitmech_v9 and are placeholder ids until METPO mints them.

  11. · REGROUND_CAUSAL_EDGE · claude

    Re-grounded the edge from enables/RO:0002327 to RO:0002326 (contributes to), issue 334. biolink declares enables range 'biological process or activity' and the object is a STATE, which does not satisfy it. Unlike the tolerance/capacity nodes in this same pass, this object is a genuine state rather than a mis-typed disposition - a gradient, a community composition, an internal environment - so retyping it to TRAIT would be wrong. contributes to fits because the subject genuinely contributes to the OCCURRENCE OR GENERATION of the object, which is biolink's definition; that is the same test the motive-force edges FAILED in issue 341, where the subject powers a machine it does not generate. RO:0002326 declares no rdfs:domain or rdfs:range.

  12. · REVIEW_UNIPROT_INSTANCE_GROUNDINGS · codex

    Reviewed 1 organism-specific UniProtKB grounding(s): replaced 1 with taxon-agnostic GO/InterPro terms and retracted 0 to label-only where no exact semantic term was supported (docs/GROUNDING_POLICY.md).

  13. · REVIEW_EVIDENCE_REFERENCE_CHURN · codex

    Offline review for issue 520 retained 4 evidence-reference replacement(s) that PR 511 made on surviving causal edges outside its stated protein-taxon scope. The pre-tranche evidence entries had references but no snippets; the retained entries supply edge-specific snippets and explanatory notes. Reverting would discard that claim-level provenance, so the scope defect is resolved by documenting the decision instead. This audit changed no causal claim or evidence field. Reviewed replacements: DOI:10.1007/s12275-024-00125-0 -> DOI:10.1073/pnas.0611035104 (1 edge); DOI:10.1038/35081051 -> DOI:10.1073/pnas.0611035104 (1 edge); DOI:10.34133/2022/9782712 -> DOI:10.1128/MMBR.69.1.51-78.2005 (1 edge); DOI:10.4014/jmb.2410.10034 -> DOI:10.1073/pnas.0611035104 (1 edge).