Function
Locations
Acylate glycerol-3-phosphate at sn-1 to LPA.
Triacylglycerol (TAG) is the principal energy-storage lipid of eukaryotic cells. It is assembled from a three-carbon glycerol backbone and fatty acyl-CoAs, and the pathway's intermediates phosphatidic acid (PA) and diacylglycerol (DAG) are also the branch points to the membrane glycerophospholipids. Lysophosphatidic acid (LPA) is formed either by sn-1 acylation of glycerol-3-phosphate (glycerol-3-phosphate acyltransferase, GPAT) or by acylation of dihydroxyacetone phosphate followed by NADPH-dependent reduction of 1-acyl-DHAP; in fungi the same GPAT enzymes accept both acceptors, whereas the DHAP acyltransferase of animals belongs to peroxisomal ether-lipid synthesis. LPA is acylated at sn-2 to PA by 1-acylglycerol-3-phosphate O-acyltransferases (LPAATs) of the AGPAT family and, in fungi, also by MBOAT-family and lipid-droplet acyltransferases. A Mg2+-dependent lipin-family phosphatidate phosphatase, a soluble enzyme that acts on membranes, dephosphorylates PA to DAG. DAG is finally esterified at sn-3 either with an acyl-CoA by the diacylglycerol O-acyltransferases (the MBOAT-family DGAT1 and the DGAT2-family enzymes) or, independently of acyl-CoA, by a phospholipid:diacylglycerol acyltransferase (PDAT) that transfers an acyl chain from a glycerophospholipid. Synthesis takes place on the endoplasmic reticulum and on lipid droplets, into which TAG is packaged; in animals the first acylation is also performed at the mitochondrial outer membrane. Inherited defects in the mammalian enzymes cause metabolic disease (AGPAT2 congenital generalized lipodystrophy, LPIN1 recurrent rhabdomyolysis, DGAT1 congenital diarrhoea).
GENERALIZE (YeastPathways project, 2026-10). The former human-only module (GPAM -> AGPAT2 -> LPIN1 -> DGAT1/DGAT2) was broadened to eukaryotic glycerolipid/TAG synthesis. All human leaves, families and gene-review evidence are kept unchanged. Structural changes: (1) LPA formation is now a ONE_OR_MORE variant set of the glycerol-3-phosphate route and the DHAP route. The G3P-acylation node holds two GPAT annotons because animal GPAM (PANTHER PTHR12563, mitochondrial outer membrane) and fungal GPT2/SCT1 (PTHR31605, ER and lipid droplets) are not in the same PANTHER family; the fungal enzymes are dual G3P/DHAP acyltransferases (PMID:11544256), so they also ground the DHAP-acylation step. The animal DHAP acyltransferase GNPAT acts in peroxisomal ether-lipid synthesis (ether_lipid_plasmalogen_biosynthesis) and is not used here. (2) LPA acylation gained fungal exemplars: SLC1 and S. pombe SPAC1851.02 join AGPAT2 in PTHR10434, and two further annotons were added for the MBOAT-family ALE1/LPT1 (PTHR13906:SF4) and the lipid-droplet LOA1/VPS66 (PTHR23063:SF60). ICT1 (Q12385), reported as an LPAAT, was not added because its contribution to de novo PA synthesis is not established in the cached literature. (3) PAH1 and S. pombe ned1 were added to the lipin family node. (4) DAG acylation is a ONE_OR_MORE variant set of acyl-CoA-dependent DGAT (DGAT1 MBOAT; DGAT2 family with yeast DGA1 and S. pombe dga1) and acyl-CoA-independent PDAT (LRO1, S. pombe plh1). The sterol acyltransferase ARE2 (PTHR10408, the DGAT1 family) contributes a minor DGAT activity in yeast (PMID:11751875) but is not used as an exemplar because its principal function is steryl-ester synthesis. YeastCyc corrections: DPP1 and LPP1, listed as PA phosphatases in TRIGLSYN-PWY, are broad-specificity lipid phosphate phosphatases that are not involved in de novo TAG synthesis (PMID:22345606) and were not added; ALE1 and LOA1 are missing from YeastCyc and were added; RXN-12383 (DAG:DAG transacylase) has no yeast gene and is not modelled; DGK1 (RXN-12959) runs the reverse reaction (DAG -> PA with CTP, PMID:18458076) and is noted as a regulator of the PA/DAG balance outside the module; GPD1/GPD2 glycerol-3-phosphate supply is in glycerol_metabolism. Compartments: YeastCyc-derived GO-CAMs put every activity in cytosol, but the yeast GPATs and SLC1 are ER and lipid-droplet enzymes, AYR1 is on lipid droplets and ER, DGA1 is enriched on lipid droplets, and PAH1 is a soluble enzyme that acts on the nuclear/ER membrane (PMID:22345606, PMID:10617610, PMID:11751830, PMID:16467296). GO-CAM disagreements: (a) all four YeastPathways models place activities in cytosol; (b) YeastPathways_TRIGLSYN-PWY includes DPP1 and LPP1 as PA phosphatases and DGK1 as part of TAG biosynthesis; (c) the PomBase model 6796b94c00005663 also contains SPAC6G10.03c as an LPA acyltransferase (UniProt names it a probable mitochondrial cardiolipin-specific deacylase), SPAC977.08/ayr2 as a second 1-acyl-DHAP reductase (UniProt: uncharacterized oxidoreductase), ptp4/dgk1, pld1, gpd1/gpd2 and ept1, which are either unverified for this activity or outside this module. Plant seed-oil assembly is modelled separately in seed_triacylglycerol_biosynthesis; the CDP-diacylglycerol and Kennedy branches that consume PA and DAG are cdp_dag_phospholipid_synthesis and kennedy_pathway_phospholipid_synthesis.
All recommended fields populated.
✗ none found
No MODULE:triacylglycerol_biosynthesis deep-research report alongside the module YAML.
✓ every leaf node grounds to a representative protein.
✓ every declared conforms_to bundle matches its template motif.
✓ every PRECEDES step chains, or its break is acknowledged via chaining_status.
22 gene-function check(s) reflected in core functions · 0 conflict(s) · 2 evidence or coverage gap(s) (40 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 |
|---|---|---|---|
| gpam_activity | GPAM / GPAT1 (human) UniProtKB:Q9HCL2 Family Representative |
glycerol-3-phosphate O-acyltransferase activity GO:0004366 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| fungal_gpat_activity | GPT2 / GAT1 (Saccharomyces cerevisiae) UniProtKB:P36148 Family Representative |
glycerol-3-phosphate O-acyltransferase activity GO:0004366 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| fungal_gpat_activity | SCT1 / GAT2 (Saccharomyces cerevisiae) UniProtKB:P32784 Family Representative |
glycerol-3-phosphate O-acyltransferase activity GO:0004366 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| fungal_gpat_activity | sct1 / SPBC1718.04 (Schizosaccharomyces pombe) UniProtKB:Q9P7P0 Family Representative |
glycerol-3-phosphate O-acyltransferase activity GO:0004366 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| fungal_dhapat_activity | GPT2 / GAT1 (Saccharomyces cerevisiae) UniProtKB:P36148 Family Representative |
glycerone-phosphate O-acyltransferase activity GO:0016287 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| fungal_dhapat_activity | SCT1 / GAT2 (Saccharomyces cerevisiae) UniProtKB:P32784 Family Representative |
glycerone-phosphate O-acyltransferase activity GO:0016287 |
Annotation Supported
A retained annotation or curated replacement supports this function, but core_functions does not yet reflect it. |
| fungal_dhapat_activity | sct1 / SPBC1718.04 (Schizosaccharomyces pombe) UniProtKB:Q9P7P0 Family Representative |
glycerone-phosphate O-acyltransferase activity GO:0016287 |
Non Core Supported
The review explicitly retains this function as non-core; promotion into core_functions is not implied. |
| ayr1_activity | AYR1 (Saccharomyces cerevisiae) UniProtKB:P40471 Family Representative |
acylglycerone-phosphate reductase (NADP+) activity GO:0000140 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| ayr1_activity | ayr1 (Schizosaccharomyces pombe) UniProtKB:Q09851 Family Representative |
acylglycerone-phosphate reductase (NADP+) activity GO:0000140 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| agpat2_activity | AGPAT2 / LPAAT-beta (human) UniProtKB:O15120 Family Representative |
1-acylglycerol-3-phosphate O-acyltransferase activity GO:0003841 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| agpat2_activity | SLC1 (Saccharomyces cerevisiae) UniProtKB:P33333 Family Representative |
1-acylglycerol-3-phosphate O-acyltransferase activity GO:0003841 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| agpat2_activity | slc1 / SPAC1851.02 (Schizosaccharomyces pombe) UniProtKB:Q9US20 Family Representative |
1-acylglycerol-3-phosphate O-acyltransferase activity GO:0003841 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| mboat_lpaat_activity | ALE1 / LPT1 (Saccharomyces cerevisiae) UniProtKB:Q08548 Family Representative |
1-acylglycerol-3-phosphate O-acyltransferase activity GO:0003841 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| mboat_lpaat_activity | ale1 (Schizosaccharomyces pombe) UniProtKB:O42916 Family Representative |
1-acylglycerol-3-phosphate O-acyltransferase activity GO:0003841 |
Non Core Supported
The review explicitly retains this function as non-core; promotion into core_functions is not implied. |
| loa1_lpaat_activity | LOA1 / VPS66 (Saccharomyces cerevisiae) UniProtKB:Q06508 Family Representative |
1-acylglycerol-3-phosphate O-acyltransferase activity GO:0003841 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| loa1_lpaat_activity | vps66 (Schizosaccharomyces pombe) UniProtKB:Q9US27 Family Representative |
1-acylglycerol-3-phosphate O-acyltransferase activity GO:0003841 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| lpin1_activity | LPIN1 / lipin-1 (human) UniProtKB:Q14693 Family Representative |
phosphatidate phosphatase activity GO:0008195 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| lpin1_activity | PAH1 (Saccharomyces cerevisiae) UniProtKB:P32567 Family Representative |
phosphatidate phosphatase activity GO:0008195 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| lpin1_activity | ned1 (Schizosaccharomyces pombe) UniProtKB:Q9UUJ6 Family Representative |
phosphatidate phosphatase activity GO:0008195 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| dgat1_activity | DGAT1 (human) UniProtKB:O75907 Family Representative |
diacylglycerol O-acyltransferase activity GO:0004144 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| dgat2_activity | DGAT2 (human) UniProtKB:Q96PD7 Family Representative |
diacylglycerol O-acyltransferase activity GO:0004144 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| dgat2_activity | DGA1 (Saccharomyces cerevisiae) UniProtKB:Q08650 Family Representative |
diacylglycerol O-acyltransferase activity GO:0004144 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| dgat2_activity | dga1 (Schizosaccharomyces pombe) UniProtKB:O74850 Family Representative |
diacylglycerol O-acyltransferase activity GO:0004144 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| pdat_activity | LRO1 (Saccharomyces cerevisiae) UniProtKB:P40345 Family Representative |
phospholipid:diacylglycerol acyltransferase activity GO:0046027 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| pdat_activity | plh1 (Schizosaccharomyces pombe) UniProtKB:O94680 Family Representative |
phospholipid:diacylglycerol acyltransferase activity GO:0046027 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| gpam_activity | GPAT / glycerophosphate O-acyltransferase family PANTHER:PTHR12563 Family |
glycerol-3-phosphate O-acyltransferase activity GO:0004366 |
Family Function Unreviewed
Family review leaves GO:0004366 UNRESOLVED. The GPAM/GPAT1 branch in SF16 has directly demonstrated glycerol-3-phosphate acyltransferase activity. The family also contains GPAT2 and bacterial PlsB proteins, so this activity is not exclusive to SF16. DAPAT proteins introduce a different acceptor specificity. The exhaustive set of substrate-compatible branches remains unresolved; the GPAT1 experiment establishes a positive example rather than an exclusion of other GPAT enzymes. |
| fungal_gpat_activity | fungal GPT2/SCT1 glycerol-3-phosphate/DHAP acyltransferase family PANTHER:PTHR31605 Family |
glycerol-3-phosphate O-acyltransferase activity GO:0004366 |
Family Supported
Family review supports GO:0004366 FAMILY_WIDE. Asserted by triacylglycerol_biosynthesis; accepted by GPT2, SCT1 and SPBC1718.04. |
| fungal_dhapat_activity | fungal GPT2/SCT1 glycerol-3-phosphate/DHAP acyltransferase family PANTHER:PTHR31605 Family |
glycerone-phosphate O-acyltransferase activity GO:0016287 |
Family Supported
Family review supports GO:0016287 FAMILY_WIDE. DHAP acyltransferase activity; accepted by GPT2 and SCT1, kept as non-core by S. pombe. |
| ayr1_activity | AYR1 1-acyldihydroxyacetone-phosphate reductase family (SDR) PANTHER:PTHR44169:SF6 Family |
acylglycerone-phosphate reductase (NADP+) activity GO:0000140 |
Family Supported
Family review supports GO:0000140 for PANTHER:PTHR44169:SF6. Asserted by triacylglycerol_biosynthesis; accepted by AYR1 and ayr1. |
| agpat2_activity | AGPAT / 1-acylglycerol-3-phosphate O-acyltransferase family PANTHER:PTHR10434 Family |
1-acylglycerol-3-phosphate O-acyltransferase activity GO:0003841 |
Family Supported
Family review supports GO:0003841 FAMILY_WIDE. Asserted by triacylglycerol_biosynthesis; accepted by S. cerevisiae SLC1, S. pombe SPAC1851.02 and human AGPAT2 in two subfamilies. |
| mboat_lpaat_activity | ALE1 / LPLAT MBOAT family PANTHER:PTHR13906:SF4 Family |
1-acylglycerol-3-phosphate O-acyltransferase activity GO:0003841 |
Family Supported
Family review supports GO:0003841 for PANTHER:PTHR13906:SF4. ALE1-type broad-specificity LPLATs (SF4), as triacylglycerol_biosynthesis selects; accepted by ALE1, kept as non-core by ale1. |
| loa1_lpaat_activity | LOA1 glycerolipid acyltransferase subfamily PANTHER:PTHR23063:SF60 Family |
1-acylglycerol-3-phosphate O-acyltransferase activity GO:0003841 |
Family Supported
Family review supports GO:0003841 for PANTHER:PTHR23063:SF60. LOA1 (SF60), selected by triacylglycerol_biosynthesis, and the LPCAT1/2 and GPAT3 subfamilies whose reviews keep it as a secondary activity. |
| lpin1_activity | Lipin / phosphatidate phosphatase family PANTHER:PTHR12181 Family |
phosphatidate phosphatase activity GO:0008195 |
Family Supported
Family review supports GO:0008195 FAMILY_WIDE. Asserted by triacylglycerol_biosynthesis; accepted by PAH1, ned1 and LPIN1-3. |
| dgat1_activity | DGAT1 / MBOAT acyltransferase family PANTHER:PTHR10408:SF7 Family |
diacylglycerol O-acyltransferase activity GO:0004144 |
Family Supported
Family review supports GO:0004144 for PANTHER:PTHR10408:SF7. DGAT1 subfamilies only (SF7, SF12, SF22). triacylglycerol_biosynthesis selects the base family for DGAT1, which also takes in the sterol acyltransferases. |
| dgat1_activity | DGAT1 / MBOAT acyltransferase family PANTHER:PTHR10408:SF12 Family |
diacylglycerol O-acyltransferase activity GO:0004144 |
Family Supported
Family review supports GO:0004144 for PANTHER:PTHR10408:SF12. DGAT1 subfamilies only (SF7, SF12, SF22). triacylglycerol_biosynthesis selects the base family for DGAT1, which also takes in the sterol acyltransferases. |
| dgat1_activity | DGAT1 / MBOAT acyltransferase family PANTHER:PTHR10408:SF22 Family |
diacylglycerol O-acyltransferase activity GO:0004144 |
Family Supported
Family review supports GO:0004144 for PANTHER:PTHR10408:SF22. DGAT1 subfamilies only (SF7, SF12, SF22). triacylglycerol_biosynthesis selects the base family for DGAT1, which also takes in the sterol acyltransferases. |
| dgat2_activity | DGAT2 / MOGAT-DGAT2 acyltransferase family PANTHER:PTHR12317:SF0 Family |
diacylglycerol O-acyltransferase activity GO:0004144 |
Family Supported
Family review supports GO:0004144 for PANTHER:PTHR12317:SF0. DGAT2 subfamilies; triacylglycerol_biosynthesis selects the base family, which also takes in MOGAT and AWAT. |
| dgat2_activity | DGAT2 / MOGAT-DGAT2 acyltransferase family PANTHER:PTHR12317:SF14 Family |
diacylglycerol O-acyltransferase activity GO:0004144 |
Family Supported
Family review supports GO:0004144 for PANTHER:PTHR12317:SF14. DGAT2 subfamilies; triacylglycerol_biosynthesis selects the base family, which also takes in MOGAT and AWAT. |
| dgat2_activity | DGAT2 / MOGAT-DGAT2 acyltransferase family PANTHER:PTHR12317:SF63 Family |
diacylglycerol O-acyltransferase activity GO:0004144 |
Family Supported
Family review supports GO:0004144 for PANTHER:PTHR12317:SF63. DGAT2 subfamilies; triacylglycerol_biosynthesis selects the base family, which also takes in MOGAT and AWAT. |
| pdat_activity | PDAT (LCAT-related) subfamily PANTHER:PTHR11440:SF105 Family |
phospholipid:diacylglycerol acyltransferase activity GO:0046027 |
Family Supported
Family review supports GO:0046027 for PANTHER:PTHR11440:SF105. PDAT (SF105), as triacylglycerol_biosynthesis selects. |
21 complete review(s) · 10 with deep research · 0 missing review · 12 reviewed but lacking deep research
| Gene | Review | Complete | Deep research |
|---|---|---|---|
| AGPAT2 O15120 | ✓ | 25/31 | ✗ |
| ale1 O42916 | ✓ | ✓ | ✗ |
| ALE1 Q08548 | ✓ | ✓ | ✓ |
| AYR1 P40471 | ✓ | ✓ | ✓ |
| ayr1 Q09851 | ✓ | ✓ | ✗ |
| dga1 O74850 | ✓ | ✓ | ✗ |
| DGA1 Q08650 | ✓ | ✓ | ✓ |
| DGAT1 O75907 | ✓ | ✓ | ✗ |
| DGAT2 Q96PD7 | ✓ | ✓ | ✗ |
| GPAM Q9HCL2 | ✓ | ✓ | ✓ |
| GPT2 P36148 | ✓ | ✓ | ✓ |
| LOA1 Q06508 | ✓ | ✓ | ✓ |
| LPIN1 Q14693 | ✓ | ✓ | ✗ |
| LRO1 P40345 | ✓ | ✓ | ✓ |
| ned1 Q9UUJ6 | ✓ | ✓ | ✗ |
| PAH1 P32567 | ✓ | ✓ | ✓ |
| plh1 O94680 | ✓ | ✓ | ✗ |
| SCT1 P32784 | ✓ | ✓ | ✓ |
| SLC1 P33333 | ✓ | ✓ | ✓ |
| SPAC1851.02 Q9US20 | ✓ | ✓ | ✗ |
| SPBC1718.04 Q9P7P0 | ✓ | ✓ | ✗ |
| vps66 Q9US27 | ✓ | ✓ | ✗ |
sn-1 acylation of glycerol-3-phosphate gives LPA directly; acylation of DHAP gives 1-acyl-DHAP, which is reduced with NADPH to LPA. Both routes operate in yeast and in fungi are served by the same dual-substrate GPATs.
Acylate glycerol-3-phosphate at sn-1 to LPA.
Acylate glycerol-3-phosphate at sn-1 to LPA (fungal ER/lipid-droplet GPAT; also uses DHAP).
DHAP is acylated at sn-1 and the 1-acyl-DHAP is reduced with NADPH to LPA. In yeast both steps occur on lipid droplets and in the ER; in labelling experiments DHAP and G3P both contribute to the glycerolipid backbone (PMID:10049376).
Acylate DHAP at sn-1 (the same fungal enzymes as the G3P acylation).
Reduce 1-acyl-DHAP to LPA with NADPH.
sn-2 acylation of LPA. The AGPAT-family enzyme (AGPAT2 in animals, SLC1 in yeast) is the main activity; fungi add the MBOAT-family lysophospholipid acyltransferase ALE1 and the lipid-droplet LOA1.
Acylate LPA at sn-2 to phosphatidic acid.
Secondary ER LPAAT that also remodels other lysophospholipids (MBOAT family).
Lipid-droplet LPAAT that channels PA toward TAG.
Dephosphorylate PA to DAG on the ER membrane (soluble enzyme recruited to membranes).
Esterify DAG at sn-3 to TAG (MBOAT family; absent as a dedicated DGAT from budding yeast).
Esterify DAG at sn-3 to TAG (DGAT2 family; ER/lipid droplet).
An LCAT-related enzyme transfers the sn-2 acyl chain of a glycerophospholipid (mainly phosphatidylcholine) to DAG, giving TAG and a lysophospholipid. Present in fungi and plants (PMID:10829075); not found in animals.
Acyl-CoA-independent TAG synthesis from phospholipid acyl donors (fungi, plants).