Function
Processes
Locations
Entry reaction of the cycle; removes 2-phosphoglycolate, which inhibits Calvin-Benson cycle enzymes.
The photorespiratory C2 cycle salvages 2-phosphoglycolate, the two-carbon product of the oxygenase reaction of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), and returns three of every four carbons to the Calvin-Benson cycle as 3-phosphoglycerate. The pathway runs through three organelles. In the chloroplast, 2-phosphoglycolate phosphatase dephosphorylates 2-phosphoglycolate to glycolate, which the inner-envelope glycolate/glycerate translocator exports. In the peroxisome, glycolate oxidase oxidizes glycolate to glyoxylate with O2, generating H2O2 that catalase removes, and glyoxylate is transaminated to glycine, mainly by glutamate:glyoxylate aminotransferase. In the mitochondrion, the glycine decarboxylase (glycine cleavage) complex converts one glycine to CO2, NH3 and 5,10-methylenetetrahydrofolate, and serine hydroxymethyltransferase uses the one-carbon unit and a second glycine to make serine. Back in the peroxisome, serine:glyoxylate aminotransferase converts serine to hydroxypyruvate while aminating a second glyoxylate, and hydroxypyruvate reductase reduces hydroxypyruvate to glycerate; a cytosolic hydroxypyruvate reductase provides a bypass. In the chloroplast, D-glycerate 3-kinase phosphorylates glycerate to 3-phosphoglycerate. The NH3 released in the mitochondrion is refixed in the chloroplast by plastidic glutamine synthetase and ferredoxin-dependent glutamate synthase, which return the glutamate used by the glyoxylate transamination. Loss of most core steps makes plants unable to grow in normal air but viable at elevated CO2. The Rubisco oxygenase reaction itself, the Calvin-Benson cycle, the transporters for the other intermediates (largely unidentified), supply of NADH to the peroxisome, alternative glycolate routes in algae and cyanobacteria, and C4 or CAM carbon-concentrating mechanisms are outside this module's boundary.
Arabidopsis thaliana proteins are used as representative members; the module itself is phrased for land plants. Every Arabidopsis representative, and the glycine decarboxylase subunits named in step 5 (GLDP1, GLDP2, GDCH, GDCST, mtLPD1, mtLPD2), has a gene review under genes/ARATH/, and each annoton molecular function matches a core function of the corresponding review. Photorespiration (GO:0009853) is the process term for every step except catalase, matching the GOA experimental rows and the gene reviews' core functions; GO:0009854 oxidative photosynthetic carbon pathway, which is part_of it, is accepted in the reviews where GOA carries it but is not used here, so that the carbon-cycle steps and ammonia refixation share one term. Catalase removes the H2O2 that glycolate oxidase produces but does no carbon chemistry of the cycle, and no catalase in GO carries photorespiration, so its annoton uses GO:0042744 hydrogen peroxide catabolic process. GO:0008891 glycolate oxidase activity is obsolete, so the glycolate oxidase step uses its parent GO:0003973 (S)-2-hydroxy-acid oxidase activity, which is the term UniProt assigns to GLO1 and GLO2. The glycine decarboxylase step reuses the glycine_cleavage_system module by reference; its Arabidopsis subunits are named in that node's description. No PANTHER family ids are asserted: they have not been checked against interpro/panther/panther.obo for these families. Paralogs not listed (GLU2, the other PGLP, GOX and CAT genes) are not modelled. No production GO-CAM in gocams/index.tsv contains these Arabidopsis gene products.
knowledge_gaps[0].provenance[0] · reference_section_type
(0/1)knowledge_gaps[0].provenance[1] · reference_section_type
(0/1)✗ none found
No MODULE:photorespiration_c2_cycle deep-research report alongside the module YAML.
✓ representative grounding skipped for abstract module.
✓ every declared conforms_to bundle matches its template motif.
✓ every PRECEDES step chains, or its break is acknowledged via chaining_status.
14 gene-function check(s) reflected in core functions · 0 conflict(s) · 12 evidence or coverage gap(s) (26 comparison(s))
Retained noncore functions are reported separately; they do not require promotion to core functions. Support from representative genes does not establish conservation across the whole family. Missing reviews or identifiers leave a claim unverified.
| Annoton | Participant | Function | Assessment |
|---|---|---|---|
| pglp_activity | PGLP1 (Arabidopsis; At5g36700) UniProtKB:P0DKC3 Family Representative |
phosphoglycolate phosphatase activity GO:0008967 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| plgg1_transport | PLGG1 (Arabidopsis; At1g32080) UniProtKB:Q9FVQ4 Family Representative |
glycolate transmembrane transporter activity GO:0043879 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| gox_activity | GLO1/GOX1 (Arabidopsis; At3g14420) UniProtKB:Q9LRR9 Family Representative |
(S)-2-hydroxy-acid oxidase activity GO:0003973 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| gox_activity | GLO2/GOX2 (Arabidopsis; At3g14415) UniProtKB:Q9LRS0 Family Representative |
(S)-2-hydroxy-acid oxidase activity GO:0003973 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| cat2_activity | CAT2 (Arabidopsis; At4g35090) UniProtKB:P25819 Family Representative |
catalase activity GO:0004096 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| ggat_activity | GGAT1/AOAT1 (Arabidopsis; At1g23310) UniProtKB:Q9LR30 Family Representative |
glycine:2-oxoglutarate transaminase activity GO:0047958 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| ggat_activity | GGAT2/AOAT2 (Arabidopsis; At1g70580) UniProtKB:Q9S7E9 Family Representative |
glycine:2-oxoglutarate transaminase activity GO:0047958 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| shm1_activity | SHM1/SHMT1 (Arabidopsis; At4g37930) UniProtKB:Q9SZJ5 Family Representative |
glycine hydroxymethyltransferase activity GO:0004372 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| sgat_activity | AGT1/SGAT (Arabidopsis; At2g13360) UniProtKB:Q56YA5 Family Representative |
L-serine:glyoxylate transaminase activity GO:0050281 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| hpr1_activity | HPR1 (Arabidopsis; At1g68010) UniProtKB:Q9C9W5 Family Representative |
hydroxypyruvate reductase (NADH) activity GO:0008465 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| hpr2_activity | HPR2 (Arabidopsis; At1g79870) UniProtKB:Q9CA90 Family Representative |
hydroxypyruvate reductase [NAD(P)H] activity GO:0016618 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| glyk_activity | GLYK (Arabidopsis; At1g80380) UniProtKB:Q944I4 Family Representative |
glycerate kinase activity GO:0008887 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| gs2_activity | GLN2/GS2 (Arabidopsis; At5g35630) UniProtKB:Q43127 Family Representative |
glutamine synthetase activity GO:0004356 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| fd_gogat_activity | GLU1/Fd-GOGAT1 (Arabidopsis; At5g04140) UniProtKB:Q9ZNZ7 Family Representative |
glutamate synthase (ferredoxin) activity GO:0016041 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| pglp_activity | plastid photorespiratory 2-phosphoglycolate phosphatase Family |
phosphoglycolate phosphatase activity GO:0008967 |
Family Id Missing
The family has no identifier linking it to a curated family review. |
| plgg1_transport | LrgB-like plastidic glycolate/glycerate translocator Family |
glycolate transmembrane transporter activity GO:0043879 |
Family Id Missing
The family has no identifier linking it to a curated family review. |
| gox_activity | peroxisomal glycolate oxidase Family |
(S)-2-hydroxy-acid oxidase activity GO:0003973 |
Family Id Missing
The family has no identifier linking it to a curated family review. |
| cat2_activity | peroxisomal monofunctional catalase Family |
catalase activity GO:0004096 |
Family Id Missing
The family has no identifier linking it to a curated family review. |
| ggat_activity | peroxisomal alanine aminotransferase-like GGAT Family |
glycine:2-oxoglutarate transaminase activity GO:0047958 |
Family Id Missing
The family has no identifier linking it to a curated family review. |
| shm1_activity | mitochondrial serine hydroxymethyltransferase Family |
glycine hydroxymethyltransferase activity GO:0004372 |
Family Id Missing
The family has no identifier linking it to a curated family review. |
| sgat_activity | class V aminotransferase serine:glyoxylate aminotransferase Family |
L-serine:glyoxylate transaminase activity GO:0050281 |
Family Id Missing
The family has no identifier linking it to a curated family review. |
| hpr1_activity | peroxisomal hydroxypyruvate reductase Family |
hydroxypyruvate reductase (NADH) activity GO:0008465 |
Family Id Missing
The family has no identifier linking it to a curated family review. |
| hpr2_activity | cytosolic glyoxylate/hydroxypyruvate reductase Family |
hydroxypyruvate reductase [NAD(P)H] activity GO:0016618 |
Family Id Missing
The family has no identifier linking it to a curated family review. |
| glyk_activity | plant-type D-glycerate 3-kinase Family |
glycerate kinase activity GO:0008887 |
Family Id Missing
The family has no identifier linking it to a curated family review. |
| gs2_activity | plastidic glutamine synthetase (GS2) Family |
glutamine synthetase activity GO:0004356 |
Family Id Missing
The family has no identifier linking it to a curated family review. |
| fd_gogat_activity | ferredoxin-dependent glutamate synthase Family |
glutamate synthase (ferredoxin) activity GO:0016041 |
Family Id Missing
The family has no identifier linking it to a curated family review. |
11 complete review(s) · 0 with deep research · 0 missing review · 14 reviewed but lacking deep research
| Gene | Review | Complete | Deep research |
|---|---|---|---|
| AGT1 Q56YA5 | ✓ | ✓ | ✗ |
| CAT2 P25819 | ✓ | 37/38 | ✗ |
| GGAT1 Q9LR30 | ✓ | ✓ | ✗ |
| GGAT2 Q9S7E9 | ✓ | ✓ | ✗ |
| GLN2 Q43127 | ✓ | ✓ | ✗ |
| GLO1 Q9LRR9 | ✓ | ✓ | ✗ |
| GLO2 Q9LRS0 | ✓ | 24/25 | ✗ |
| GLU1 Q9ZNZ7 | ✓ | ✓ | ✗ |
| GLYK Q944I4 | ✓ | ✓ | ✗ |
| HPR Q9C9W5 | ✓ | 22/23 | ✗ |
| HPR2 Q9CA90 | ✓ | ✓ | ✗ |
| PGLP1A P0DKC3 | ✓ | ✓ | ✗ |
| PLGG1 Q9FVQ4 | ✓ | ✓ | ✗ |
| SHM1 Q9SZJ5 | ✓ | ✓ | ✗ |
Entry reaction of the cycle; removes 2-phosphoglycolate, which inhibits Calvin-Benson cycle enzymes.
The same translocator imports glycerate for the final kinase step (glycerate transmembrane transporter activity, GO:1901974). The dicarboxylate translocators DiT1/DiT2 that exchange 2-oxoglutarate and glutamate for the ammonia refixation step are not modelled.
Glycolate oxidase, an FMN-dependent (S)-2-hydroxy-acid oxidase, uses O2 to oxidize glycolate to glyoxylate and produces H2O2; peroxisomal catalase disproportionates the H2O2. In Arabidopsis two co-regulated leaf genes, GOX1 and GOX2, are redundant enough that no glycolate oxidase mutant was found in the classical photorespiratory screens.
GO:0008891 glycolate oxidase activity is obsolete; the parent (S)-2-hydroxy-acid oxidase term is used.
Ancillary detoxification step; cat2 mutants show high-CO2 reversible growth and redox phenotypes in air.
GGAT1 knockouts have a weak photorespiratory phenotype, with residual activity from GGAT2; serine:glyoxylate aminotransferase (step 7) aminates the other half of the glyoxylate.
Glycine is oxidatively decarboxylated and deaminated to CO2, NH3, NADH and 5,10-methylenetetrahydrofolate. In Arabidopsis the P protein is encoded by GLDP1 (UniProtKB:Q94B78) and GLDP2 (UniProtKB:O80988), which are redundant; the H and T proteins include GDCH (UniProtKB:P25855) and GDCST (UniProtKB:O65396). The leaf L protein is mainly mitochondrial LPD1 (mtLPD1, At1g48030, UniProtKB:Q9M5K3); the paralog mtLPD2 (At3g17240, UniProtKB:Q9M5K2) is a minor but real L subunit of the leaf complex, at about a quarter of the mtLPD1 level, and mutant analysis shows that both mtLPDs contribute to photorespiration, mtLPD1 predominantly. mtLPD2 was first proposed to serve chiefly the 2-oxoacid dehydrogenase complexes from its expression pattern. Loss of both P-protein genes is lethal even at high CO2, because the complex is also required for one-carbon metabolism.
Realized by module
glycine_cleavage_system
Supplies the second amino group for glycine formation, which keeps the nitrogen stoichiometry of the cycle closed.
Single hpr1 mutants are only mildly affected in air, whereas hpr1 hpr2 double mutants are air-sensitive, so the peroxisomal and cytosolic reductases act in parallel.
Concluding step; returns 3-phosphoglycerate to the Calvin-Benson cycle. GLYK is unrelated in structure to bacterial and animal glycerate kinases.
NH3 released by glycine decarboxylase is reassimilated in the chloroplast by the plastidic glutamine synthetase (GS2) and ferredoxin-dependent glutamate synthase (Fd-GOGAT), producing the glutamate that the peroxisomal glyoxylate transamination consumes. Of the two Arabidopsis Fd-GOGAT genes only GLU1 mutants (gls) are photorespiratory.