id: MODULE:oxygenic_photosynthesis
title: Oxygenic photosynthesis module
description: >-
  A taxon-neutral decomposition of oxygenic photosynthesis as a recursively
  decomposable module. The module separates the thylakoid light reactions
  (light harvesting, water oxidation at photosystem II, the cytochrome b6f
  complex, photosystem I, mobile electron carriers, ferredoxin-NADP+ reductase,
  and the ATP synthase) from carbon fixation by the Calvin-Benson-Bassham
  reductive pentose-phosphate cycle, and adds optional photoprotection/electron
  balancing, the inorganic carbon-concentrating mechanism, and chlorophyll
  supply. It is phrased as functions, complexes, and pathway segments rather
  than a fixed gene list so it can represent cyanobacterial, algal, and plant
  implementations. Anoxygenic photosynthesis (single reaction center, non-water
  electron donors) is explicitly out of scope.
status: DRAFT
evidence:
  - source_id: file:modules/photosynthesis-deep-research-falcon.md
    title: Photosynthesis module deep research (falcon)
    statement: >-
      Module-targeted falcon deep research (generated from this module's
      outline) used to cross-check component coverage, variants, and
      failure modes. Machine-generated citations are not yet manually adjudicated.
  - source_id: file:terms/photosynthesis/photosynthesis-deep-research-falcon.md
    title: Photosynthesis concept deep research (falcon)
    statement: >-
      Falcon deep-research synthesis used to set module boundaries and the
      module-by-module component breakdown across Arabidopsis, Synechocystis,
      and Chlamydomonas (core vs accessory vs regulatory factors).
  - source_id: file:terms/photosynthesis/photosynthesis-notes.md
    title: Photosynthesis concept notes
    statement: Manually authored, GO- and PubMed-verified concept notes underpinning this module.
  - source_id: GO:0015979
    title: photosynthesis
    statement: The module is grounded in the GO biological-process term for photosynthesis.
  - source_id: PMID:21499260
    title: "Crystal structure of oxygen-evolving photosystem II at a resolution of 1.9 Å."
    statement: >-
      Defines the PSII D1/D2 reaction center, CP43/CP47 inner antennae, and the
      Mn4CaO5 water-oxidation cluster stabilized by the extrinsic OEC proteins.
  - source_id: PMID:11418848
    title: "Three-dimensional structure of cyanobacterial photosystem I at 2.5 A resolution"
    statement: >-
      Defines the PsaA/PsaB reaction-center heterodimer and the terminal Fe-S
      clusters carried by PsaC.
  - source_id: PMID:18294858
    title: Structure and function of Rubisco
    statement: >-
      Supports Rubisco as the major carboxylating enzyme of the CBB cycle and
      its competing oxygenase side-reaction.
  - source_id: PMID:33761918
    title: A new type of flexible CP12 protein in the marine diatom Thalassiosira pseudonana
    statement: >-
      Supports CP12 as a redox-controlled scaffold that forms a PRK-GAPDH-CP12
      ternary complex regulating the CBB cycle.
  - source_id: PMID:37549282
    title: NTRC regulates CP12 to activate Calvin-Benson cycle during cold acclimation
    statement: >-
      Supports redox (NTRC/thioredoxin)-dependent assembly/disassembly of the
      PRK-CP12-GAPDH supracomplex gating CBB-cycle activity.
notes: >-
  Identifiers are grounded only where verified against the local GO term cache,
  GOlr, or UniProt; descriptors without a `term` are deliberate (no confident
  identifier yet) rather than oversights. Representative UniProt members are
  concrete orienting examples, not exhaustive or species-restricting claims.
module:
  id: oxygenic_photosynthesis
  label: Oxygenic photosynthesis
  module_type: BIOLOGICAL_PROCESS
  concepts:
    - preferred_term: oxygenic photosynthesis
      term:
        id: GO:0015979
        label: photosynthesis
      description: >-
        Light-driven oxidation of water and reduction of CO2 to organic carbon,
        releasing O2. Comprises coupled light reactions and carbon fixation.
  context:
    taxa:
      - preferred_term: cyanobacteria
      - preferred_term: eukaryotic algae
      - preferred_term: land plants
    cellular_components:
      - preferred_term: photosynthetic membrane
        term:
          id: GO:0034357
          label: photosynthetic membrane
  parts:
    - order: 1
      role: light reactions (electron transport and photophosphorylation)
      node:
        id: light_reactions
        label: Thylakoid light reactions
        module_type: BIOLOGICAL_PROCESS
        concepts:
          - preferred_term: photosynthesis, light reaction
            term:
              id: GO:0019684
              label: photosynthesis, light reaction
          - preferred_term: photosynthetic electron transport chain
            term:
              id: GO:0009767
              label: photosynthetic electron transport chain
        parts:
          - order: 1
            role: light harvesting / antenna
            node:
              id: light_harvesting
              label: Light-harvesting antenna
              module_type: MOLECULAR_FUNCTION
              annotons:
                - id: antenna_chlorophyll_binding
                  label: Antenna chlorophyll/carotenoid light capture
                  participant:
                    selector_type: FAMILY
                    family:
                      preferred_term: light-harvesting chlorophyll a/b binding (LHC) family
                      description: >-
                        Membrane antenna proteins (LHCII/LHCI in plants and algae;
                        phycobilisomes substitute in most cyanobacteria).
                  function:
                    preferred_term: chlorophyll binding for excitation energy transfer
                    term:
                      id: GO:0016168
                      label: chlorophyll binding
                    cofactors:
                      - preferred_term: chlorophyll a
                      - preferred_term: chlorophyll b
                      - preferred_term: carotenoid
                  locations:
                    - preferred_term: thylakoid membrane
                      term:
                        id: GO:0034357
                        label: photosynthetic membrane
                  role_description: >-
                    Increases absorption cross-section and funnels excitation to
                    the reaction centers; also a site of photoprotective quenching.
          - order: 2
            role: water oxidation (photosystem II)
            node:
              id: photosystem_II
              label: Photosystem II (water:plastoquinone oxidoreductase)
              module_type: PROTEIN_COMPLEX
              concepts:
                - preferred_term: photosystem II
                  term:
                    id: GO:0009523
                    label: photosystem II
              annotons:
                - id: psii_water_oxidation
                  label: Light-driven water oxidation / oxygen evolution
                  participant:
                    selector_type: PROTEIN_COMPLEX
                    protein_complex:
                      preferred_term: photosystem II complex
                      term:
                        id: GO:0009523
                        label: photosystem II
                      active_units:
                        - id: psii_d1
                          label: D1 reaction-center subunit (PsbA)
                          role: binds P680 and the Mn4CaO5 cluster; high-turnover repair substrate
                          participant:
                            selector_type: FAMILY
                            family:
                              preferred_term: PsbA / D1
                        - id: psii_d2
                          label: D2 reaction-center subunit (PsbD)
                          role: reaction-center partner of D1
                          participant:
                            selector_type: FAMILY
                            family:
                              preferred_term: PsbD / D2
                        - id: psii_cp43_cp47
                          label: Core antennae CP43/CP47 (PsbC/PsbB)
                          role: inner chlorophyll antennae feeding excitation to P680
                          participant:
                            selector_type: FAMILY
                            family:
                              preferred_term: PsbB/PsbC core antenna
                        - id: psii_oec
                          label: Oxygen-evolving complex extrinsic proteins (PsbO/PsbP/PsbQ)
                          role: stabilize the Mn4CaO5 water-oxidation cluster on the lumenal side
                          participant:
                            selector_type: FAMILY
                            family:
                              preferred_term: oxygen-evolving complex extrinsic proteins
                              representative_members:
                                - preferred_term: spinach PsbO (oxygen-evolving enhancer protein 1)
                                  term:
                                    id: UniProtKB:P12359
                                    label: Oxygen-evolving enhancer protein 1, chloroplastic
                  function:
                    preferred_term: oxygen evolving activity
                    term:
                      id: GO:0010242
                      label: oxygen evolving activity
                    substrates:
                      - preferred_term: water
                      - preferred_term: plastoquinone
                    products:
                      - preferred_term: dioxygen
                      - preferred_term: plastoquinol
                      - preferred_term: proton (lumen)
                  locations:
                    - preferred_term: thylakoid membrane
                      term:
                        id: GO:0042651
                        label: thylakoid membrane
                  role_description: >-
                    Uses light energy to oxidize water at the Mn4CaO5 cluster,
                    reducing plastoquinone and releasing O2 and lumenal protons.
                  evidence:
                    - source_id: PMID:21499260
                      statement: PSII structure defining the reaction center, antennae, and Mn4CaO5 cluster.
          - order: 3
            role: plastoquinol oxidation and proton translocation (cytochrome b6f)
            node:
              id: cytochrome_b6f
              label: Cytochrome b6f complex
              module_type: PROTEIN_COMPLEX
              concepts:
                - preferred_term: cytochrome b6f complex
                  term:
                    id: GO:0009512
                    label: cytochrome b6f complex
              annotons:
                - id: b6f_plastoquinol_plastocyanin_reductase
                  label: Plastoquinol--plastocyanin reductase (Q-cycle)
                  participant:
                    selector_type: PROTEIN_COMPLEX
                    protein_complex:
                      preferred_term: cytochrome b6f complex
                      term:
                        id: GO:0009512
                        label: cytochrome b6f complex
                      active_units:
                        - id: b6f_cytf
                          label: Cytochrome f (PetA)
                          role: high-potential heme; reduces plastocyanin/cyt c6
                          participant:
                            selector_type: FAMILY
                            family:
                              preferred_term: cytochrome f (PetA)
                              representative_members:
                                - preferred_term: Synechocystis cytochrome f (petA)
                                  term:
                                    id: UniProtKB:P26287
                                    label: Cytochrome f
                        - id: b6f_cytb6
                          label: Cytochrome b6 (PetB)
                          role: low-potential hemes; Q-cycle electron bifurcation
                          participant:
                            selector_type: FAMILY
                            family:
                              preferred_term: cytochrome b6 (PetB)
                        - id: b6f_rieske
                          label: Rieske iron-sulfur protein (PetC)
                          role: 2Fe-2S cluster; oxidizes plastoquinol
                          participant:
                            selector_type: FAMILY
                            family:
                              preferred_term: Rieske Fe-S protein (PetC)
                              representative_members:
                                - preferred_term: wheat Rieske Fe-S subunit (petC)
                                  term:
                                    id: UniProtKB:Q7X9A6
                                    label: Cytochrome b6-f complex iron-sulfur subunit, chloroplastic
                  function:
                    preferred_term: plastoquinol--plastocyanin reductase activity
                    term:
                      id: GO:0009496
                      label: plastoquinol--plastocyanin reductase activity
                    substrates:
                      - preferred_term: plastoquinol
                      - preferred_term: plastocyanin (oxidized)
                    products:
                      - preferred_term: plastoquinone
                      - preferred_term: plastocyanin (reduced)
                      - preferred_term: proton (lumen)
                  locations:
                    - preferred_term: thylakoid membrane
                      term:
                        id: GO:0009535
                        label: chloroplast thylakoid membrane
                  role_description: >-
                    Links PSII to PSI, oxidizing plastoquinol and reducing the
                    mobile carrier while translocating protons via the Q-cycle;
                    frequently rate-limiting for linear electron transport.
          - order: 4
            role: inter-photosystem mobile electron carrier
            node:
              id: mobile_carrier_b6f_to_psi
              label: Mobile carrier from b6f to PSI
              module_type: MOLECULAR_FUNCTION
              description: >-
                Soluble lumenal carrier shuttling electrons from cytochrome b6f
                to photosystem I; the carrier used is lineage- and
                metal-availability-dependent.
              variant_sets:
                - id: lumenal_carrier_variants
                  label: Plastocyanin vs cytochrome c6
                  axis: lineage / metal availability
                  selection: EXACTLY_ONE
                  variants:
                    - id: plastocyanin_variant
                      label: Plastocyanin (copper carrier)
                      module_type: MOLECULAR_FUNCTION
                      annotons:
                        - id: plastocyanin_electron_transfer
                          label: Plastocyanin electron transfer
                          participant:
                            selector_type: FAMILY
                            family:
                              preferred_term: plastocyanin
                          function:
                            preferred_term: electron transfer activity
                            term:
                              id: GO:0009055
                              label: electron transfer activity
                            cofactors:
                              - preferred_term: copper ion
                          locations:
                            - preferred_term: thylakoid lumen
                              term:
                                id: GO:0009543
                                label: chloroplast thylakoid lumen
                    - id: cytochrome_c6_variant
                      label: Cytochrome c6 (iron carrier)
                      module_type: MOLECULAR_FUNCTION
                      annotons:
                        - id: cytc6_electron_transfer
                          label: Cytochrome c6 electron transfer
                          participant:
                            selector_type: FAMILY
                            family:
                              preferred_term: cytochrome c6
                          function:
                            preferred_term: electron transfer activity
                            term:
                              id: GO:0009055
                              label: electron transfer activity
                            cofactors:
                              - preferred_term: heme c
                          locations:
                            - preferred_term: thylakoid lumen
                              term:
                                id: GO:0031977
                                label: thylakoid lumen
                  notes: >-
                    Many cyanobacteria and algae switch between plastocyanin and
                    cytochrome c6 depending on copper/iron availability.
          - order: 5
            role: P700 photo-oxidation and ferredoxin reduction (photosystem I)
            node:
              id: photosystem_I
              label: Photosystem I (plastocyanin:ferredoxin oxidoreductase)
              module_type: PROTEIN_COMPLEX
              concepts:
                - preferred_term: photosystem I
                  term:
                    id: GO:0009522
                    label: photosystem I
              annotons:
                - id: psi_ferredoxin_reduction
                  label: Light-driven ferredoxin reduction
                  participant:
                    selector_type: PROTEIN_COMPLEX
                    protein_complex:
                      preferred_term: photosystem I complex
                      term:
                        id: GO:0009522
                        label: photosystem I
                      active_units:
                        - id: psi_psaa_psab
                          label: PsaA/PsaB reaction-center heterodimer
                          role: bind P700 and early electron-transfer cofactors
                          participant:
                            selector_type: FAMILY
                            family:
                              preferred_term: PsaA/PsaB reaction-center
                        - id: psi_psac
                          label: PsaC terminal Fe-S clusters (FA/FB)
                          role: terminal 4Fe-4S clusters that reduce ferredoxin
                          participant:
                            selector_type: FAMILY
                            family:
                              preferred_term: PsaC
                              representative_members:
                                - preferred_term: Chlamydomonas psaC
                                  term:
                                    id: UniProtKB:Q00914
                                    label: Photosystem I iron-sulfur center
                  function:
                    preferred_term: light-driven plastocyanin:ferredoxin oxidoreduction
                    description: >-
                      Photo-oxidation of P700 drives electron transfer through
                      Fe-S clusters to reduce ferredoxin; no single exact GO MF
                      is asserted for the whole complex here.
                    substrates:
                      - preferred_term: plastocyanin (reduced) or cytochrome c6
                      - preferred_term: ferredoxin (oxidized)
                    products:
                      - preferred_term: ferredoxin (reduced)
                  processes:
                    - preferred_term: photosynthetic electron transport in photosystem I
                      term:
                        id: GO:0009773
                        label: photosynthetic electron transport in photosystem I
                  locations:
                    - preferred_term: thylakoid membrane
                      term:
                        id: GO:0009535
                        label: chloroplast thylakoid membrane
                  evidence:
                    - source_id: PMID:11418848
                      statement: PSI structure defining PsaA/PsaB and terminal Fe-S clusters.
          - order: 6
            role: NADPH production
            node:
              id: fnr_step
              label: Ferredoxin-NADP+ reductase
              module_type: REACTION
              annotons:
                - id: fnr_activity
                  label: Ferredoxin-NADP+ reductase
                  participant:
                    selector_type: ANY_WITH_FUNCTION
                    required_function:
                      preferred_term: ferredoxin-NADP+ reductase activity
                      term:
                        id: GO:0004324
                        label: ferredoxin-NADP+ reductase activity
                  function:
                    preferred_term: ferredoxin-NADP+ reductase activity
                    term:
                      id: GO:0004324
                      label: ferredoxin-NADP+ reductase activity
                    substrates:
                      - preferred_term: ferredoxin (reduced)
                      - preferred_term: NADP+
                    products:
                      - preferred_term: ferredoxin (oxidized)
                      - preferred_term: NADPH
          - order: 7
            role: ATP synthesis from the proton-motive force
            node:
              id: atp_synthase
              label: Thylakoid (chloroplast/cyanobacterial) ATP synthase
              module_type: PROTEIN_COMPLEX
              concepts:
                - preferred_term: proton-transporting ATP synthase complex
                  term:
                    id: GO:0045259
                    label: proton-transporting ATP synthase complex
              annotons:
                - id: photophosphorylation
                  label: Proton-motive-force-driven ATP synthesis
                  participant:
                    selector_type: PROTEIN_COMPLEX
                    protein_complex:
                      preferred_term: thylakoid ATP synthase (CF1FO)
                      term:
                        id: GO:0045259
                        label: proton-transporting ATP synthase complex
                  function:
                    preferred_term: "proton-transporting ATP synthase activity, rotational mechanism"
                    term:
                      id: GO:0046933
                      label: "proton-transporting ATP synthase activity, rotational mechanism"
                    substrates:
                      - preferred_term: ADP
                      - preferred_term: phosphate
                      - preferred_term: proton (lumen)
                    products:
                      - preferred_term: ATP
                  locations:
                    - preferred_term: thylakoid membrane
                      term:
                        id: GO:0009535
                        label: chloroplast thylakoid membrane
        connections:
          - source: photosystem_II
            target: cytochrome_b6f
            connection_type: PROVIDES_INPUT_FOR
            description: PSII reduces plastoquinone to plastoquinol, the substrate oxidized by b6f.
          - source: cytochrome_b6f
            target: mobile_carrier_b6f_to_psi
            connection_type: PROVIDES_INPUT_FOR
            description: b6f reduces the mobile lumenal carrier (plastocyanin or cyt c6).
          - source: mobile_carrier_b6f_to_psi
            target: photosystem_I
            connection_type: PROVIDES_INPUT_FOR
            description: The reduced carrier re-reduces photo-oxidized P700 in PSI.
          - source: photosystem_I
            target: fnr_step
            connection_type: PROVIDES_INPUT_FOR
            description: PSI-reduced ferredoxin is the electron donor for FNR.
          - source: fnr_step
            target: atp_synthase
            connection_type: PRECEDES
            description: >-
              Linear electron flow and water oxidation build the proton-motive
              force used by the ATP synthase (coupling, not a metabolite hand-off).
    - order: 2
      role: carbon fixation (Calvin-Benson-Bassham cycle)
      node:
        id: carbon_fixation
        label: Calvin-Benson-Bassham reductive pentose-phosphate cycle
        module_type: METABOLIC_PATHWAY
        concepts:
          - preferred_term: reductive pentose-phosphate cycle
            term:
              id: GO:0019253
              label: reductive pentose-phosphate cycle
          - preferred_term: carbon fixation
            term:
              id: GO:0015977
              label: carbon fixation
        parts:
          - order: 1
            role: carboxylation
            node:
              id: rubisco_carboxylation
              label: Rubisco carboxylation of RuBP
              module_type: REACTION
              annotons:
                - id: rubisco_activity
                  label: Ribulose-bisphosphate carboxylase
                  participant:
                    selector_type: ANY_WITH_FUNCTION
                    required_function:
                      preferred_term: ribulose-bisphosphate carboxylase activity
                      term:
                        id: GO:0016984
                        label: ribulose-bisphosphate carboxylase activity
                    description: >-
                      Form I Rubisco (L8S8, RbcL+RbcS) in plants, algae, and
                      cyanobacteria; form II (L2) in some bacteria/dinoflagellates.
                  function:
                    preferred_term: ribulose-bisphosphate carboxylase activity
                    term:
                      id: GO:0016984
                      label: ribulose-bisphosphate carboxylase activity
                    substrates:
                      - preferred_term: ribulose 1,5-bisphosphate
                      - preferred_term: CO2
                    products:
                      - preferred_term: 3-phosphoglycerate
                  locations:
                    - preferred_term: chloroplast stroma / cyanobacterial cytoplasm
                      term:
                        id: GO:0009570
                        label: chloroplast stroma
                  role_description: >-
                    Fixes CO2 onto RuBP. The competing oxygenase reaction with O2
                    produces 2-phosphoglycolate (the substrate of photorespiration).
                  evidence:
                    - source_id: PMID:18294858
                      statement: Rubisco as the principal carboxylating enzyme with a competing oxygenase activity.
          - order: 2
            role: reduction phase
            node:
              id: cbb_reduction_phase
              label: Reduction of 3-phosphoglycerate to triose phosphate
              module_type: METABOLIC_PATHWAY
              description: >-
                ATP- and NADPH-consuming reduction of 3-phosphoglycerate via
                1,3-bisphosphoglycerate to glyceraldehyde 3-phosphate
                (phosphoglycerate kinase + chloroplastic NADP-GAPDH).
              annotons:
                - id: cbb_gapdh
                  label: NADP-dependent glyceraldehyde-3-phosphate dehydrogenase
                  participant:
                    selector_type: FAMILY
                    family:
                      preferred_term: chloroplastic GAPDH (GapA/GapB)
                  function:
                    preferred_term: glyceraldehyde-3-phosphate dehydrogenase (NADP+) (phosphorylating) activity
                    description: NADPH-consuming reduction step; no confident exact GO MF id asserted here.
                    substrates:
                      - preferred_term: 1,3-bisphosphoglycerate
                      - preferred_term: NADPH
                    products:
                      - preferred_term: glyceraldehyde 3-phosphate
                      - preferred_term: NADP+
          - order: 3
            role: regeneration phase
            node:
              id: cbb_regeneration_phase
              label: Regeneration of ribulose 1,5-bisphosphate
              module_type: METABOLIC_PATHWAY
              description: >-
                Rearrangement of triose phosphates through transketolase,
                aldolase, fructose-1,6-bisphosphatase, and sedoheptulose-1,7-
                bisphosphatase, ending in phosphoribulokinase regeneration of RuBP.
              annotons:
                - id: cbb_prk
                  label: Phosphoribulokinase
                  participant:
                    selector_type: ANY_WITH_FUNCTION
                    required_function:
                      preferred_term: phosphoribulokinase activity
                      term:
                        id: GO:0008974
                        label: phosphoribulokinase activity
                  function:
                    preferred_term: phosphoribulokinase activity
                    term:
                      id: GO:0008974
                      label: phosphoribulokinase activity
                    substrates:
                      - preferred_term: ribulose 5-phosphate
                      - preferred_term: ATP
                    products:
                      - preferred_term: ribulose 1,5-bisphosphate
                      - preferred_term: ADP
                  role_description: Regenerates the Rubisco substrate; a major redox-regulated control point.
                - id: cbb_sbpase_fbpase
                  label: Bisphosphatase regeneration steps (FBPase, SBPase)
                  participant:
                    selector_type: FAMILY
                    family:
                      preferred_term: chloroplast FBPase / SBPase
                  function:
                    preferred_term: bisphosphatase regeneration steps
                    description: >-
                      Fructose-1,6-bisphosphatase and sedoheptulose-1,7-
                      bisphosphatase; SBPase is a frequent yield-engineering target.
          - order: 4
            role: Rubisco activation and inhibitor repair
            optional: true
            node:
              id: rubisco_activation
              label: Rubisco activase and inhibitor removal
              module_type: REGULATORY_STEP
              annotons:
                - id: rubisco_activase
                  label: Rubisco activase
                  participant:
                    selector_type: FAMILY
                    family:
                      preferred_term: Rubisco activase (Rca)
                  function:
                    preferred_term: ATP-dependent removal of inhibitory sugar phosphates from Rubisco
                    description: AAA+ chaperone-like remodeling of Rubisco active sites; thermolabile.
                  role_description: Restores catalytically competent Rubisco; major thermal-tolerance target.
          - order: 5
            role: redox gating of the CBB cycle
            optional: true
            node:
              id: cbb_redox_regulation
              label: CP12-mediated redox regulation
              module_type: REGULATORY_STEP
              annotons:
                - id: cp12_scaffold
                  label: CP12 redox scaffold
                  participant:
                    selector_type: FAMILY
                    family:
                      preferred_term: CP12
                      representative_members:
                        - preferred_term: Chlamydomonas CP12
                          term:
                            id: UniProtKB:A6Q0K5
                            label: Calvin cycle protein CP12
                  function:
                    preferred_term: redox-dependent scaffolding of the PRK-GAPDH-CP12 ternary complex
                    description: >-
                      In the dark/oxidizing conditions CP12 assembles an
                      autoinhibitory PRK-GAPDH complex; reduction (thioredoxin/
                      NTRC) dissociates it to activate the cycle.
                    targets:
                      - preferred_term: phosphoribulokinase
                      - preferred_term: glyceraldehyde-3-phosphate dehydrogenase
                  role_description: Couples CBB-cycle activity to the light/redox state of the chloroplast.
                  evidence:
                    - source_id: PMID:33761918
                      statement: CP12 forms a PRK-GAPDH ternary complex regulating the CBB cycle.
                    - source_id: PMID:37549282
                      statement: NTRC-dependent redox control of the PRK-CP12-GAPDH supracomplex.
              connections:
                - source: cp12_scaffold
                  target: cbb_prk
                  connection_type: NEGATIVELY_REGULATES
                  description: CP12 inhibits PRK in the dark/oxidized state.
                - source: cp12_scaffold
                  target: cbb_gapdh
                  connection_type: NEGATIVELY_REGULATES
                  description: CP12 inhibits GAPDH in the dark/oxidized state.
    - order: 3
      role: photoprotection and ATP/NADPH balancing
      optional: true
      node:
        id: photoprotection_balancing
        label: Photoprotection and electron/energy balancing
        module_type: REGULATORY_STEP
        description: >-
          Mechanisms that dissipate excess excitation, redistribute antennae,
          and tune the ATP:NADPH ratio under fluctuating light.
        variant_sets:
          - id: balancing_mechanisms
            label: Photoprotection and balancing mechanisms
            axis: regulatory mechanism
            selection: ZERO_OR_MORE
            variants:
              - id: npq_variant
                label: Non-photochemical quenching (qE)
                module_type: REGULATORY_STEP
                annotons:
                  - id: npq_psbs_xanthophyll
                    label: PsbS- and xanthophyll-cycle-dependent quenching
                    participant:
                      selector_type: FAMILY
                      family:
                        preferred_term: PsbS and xanthophyll-cycle enzymes (VDE/ZEP)
                    function:
                      preferred_term: heat dissipation of excess PSII excitation
                    role_description: Rapid, reversible thermal dissipation protecting PSII under high light.
              - id: state_transitions_variant
                label: State transitions
                module_type: REGULATORY_STEP
                annotons:
                  - id: state_transition_kinase
                    label: STN7/Stt7-dependent LHCII redistribution
                    participant:
                      selector_type: FAMILY
                      family:
                        preferred_term: state-transition kinase (STN7/Stt7) and TAP38/PPH1 phosphatase
                    function:
                      preferred_term: redox-controlled LHCII phosphorylation balancing PSII/PSI
                    role_description: Rebalances excitation between the photosystems via antenna migration.
              - id: cyclic_electron_flow_variant
                label: Cyclic electron flow around PSI
                module_type: REGULATORY_STEP
                annotons:
                  - id: cef_pgr5_ndh
                    label: PGR5/PGRL1- and NDH-dependent cyclic electron flow
                    participant:
                      selector_type: FAMILY
                      family:
                        preferred_term: PGR5/PGRL1 and NDH(-1) complex
                    function:
                      preferred_term: cyclic electron transfer generating extra proton-motive force
                    role_description: Raises ATP:NADPH and contributes to photoprotection.
    - order: 4
      role: inorganic carbon-concentrating mechanism (CCM)
      optional: true
      node:
        id: carbon_concentrating_mechanism
        label: Carbon-concentrating mechanism
        module_type: REGULATORY_STEP
        description: >-
          Active inorganic-carbon uptake and CO2 concentration around Rubisco to
          suppress the oxygenase reaction; prominent in cyanobacteria and algae.
        annotons:
          - id: ccm_carbonic_anhydrase
            label: Carbonic anhydrase (CO2/HCO3- interconversion)
            participant:
              selector_type: ANY_WITH_FUNCTION
              required_function:
                preferred_term: carbonate dehydratase activity
                term:
                  id: GO:0004089
                  label: carbonate dehydratase activity
            function:
              preferred_term: carbonate dehydratase activity
              term:
                id: GO:0004089
                label: carbonate dehydratase activity
              substrates:
                - preferred_term: bicarbonate
              products:
                - preferred_term: CO2
                - preferred_term: water
        variant_sets:
          - id: ccm_strategy
            label: CCM compartmentation strategy
            axis: lineage / compartment
            selection: EXACTLY_ONE
            variants:
              - id: carboxysome_ccm
                label: Cyanobacterial carboxysome CCM
                module_type: CELLULAR_COMPONENT
                annotons:
                  - id: carboxysome_unit
                    label: Carboxysome microcompartment with encapsulated Rubisco/CA
                    participant:
                      selector_type: FAMILY
                      family:
                        preferred_term: carboxysome shell and cargo (Ccm/Cso)
                    role_description: Bacterial microcompartment concentrating CO2 around Rubisco.
              - id: pyrenoid_ccm
                label: Algal pyrenoid CCM
                module_type: CELLULAR_COMPONENT
                annotons:
                  - id: pyrenoid_unit
                    label: Pyrenoid and bicarbonate transporter network
                    participant:
                      selector_type: FAMILY
                      family:
                        preferred_term: pyrenoid CCM components (HLA3, LCIA, LCI1, LCIB, CAH3, CCM1/CIA5)
                        representative_members:
                          - preferred_term: Chlamydomonas LCI5 (low-CO2-inducible protein)
                            term:
                              id: UniProtKB:Q94ET8
                              label: Low-CO2-inducible protein 5
                    role_description: Eukaryotic algal biophysical CCM organized around the pyrenoid.
    - order: 5
      role: chlorophyll supply (supporting context)
      optional: true
      node:
        id: pigment_biosynthesis
        label: Chlorophyll biosynthesis (supporting)
        module_type: METABOLIC_PATHWAY
        description: Supplies the chlorophyll cofactors required by antennae and reaction centers.
        annotons:
          - id: mg_chelatase
            label: Magnesium chelatase (committed step)
            participant:
              selector_type: FAMILY
              family:
                preferred_term: magnesium chelatase (CHLH/CHLD/CHLI)
            function:
              preferred_term: magnesium chelatase activity
              description: Inserts Mg2+ into protoporphyrin IX; committed step toward chlorophyll. No confident GO MF id asserted here.
          - id: por_reduction
            label: Protochlorophyllide reductase
            participant:
              selector_type: ANY_WITH_FUNCTION
              required_function:
                preferred_term: protochlorophyllide reductase activity
                term:
                  id: GO:0016630
                  label: protochlorophyllide reductase activity
            function:
              preferred_term: protochlorophyllide reductase activity
              term:
                id: GO:0016630
                label: protochlorophyllide reductase activity
              substrates:
                - preferred_term: protochlorophyllide
              products:
                - preferred_term: chlorophyllide
  connections:
    - source: light_reactions
      target: carbon_fixation
      connection_type: PROVIDES_INPUT_FOR
      description: >-
        The light reactions supply the ATP and NADPH consumed by the
        Calvin-Benson-Bassham cycle.
    - source: carbon_concentrating_mechanism
      target: rubisco_carboxylation
      connection_type: POSITIVELY_REGULATES
      description: The CCM raises CO2 around Rubisco, favoring carboxylation over oxygenation.
  notes: >-
    Concrete organism modules can specialize abstract participants with specific
    genes, complexes, compartments, and transporters (e.g. Synechocystis psbA/
    psaA, Chlamydomonas pyrenoid CCM, Arabidopsis PsbS/RbcS) without the generic
    module naming every protein.
