LCAT

Summary

LCAT (lecithin-cholesterol acyltransferase, HGNC:6522) is a protein-coding gene on chromosome 16q22.1, encoding Phosphatidylcholine-sterol acyltransferase (P04180). Central enzyme in the extracellular metabolism of plasma lipoproteins.

This gene encodes the extracellular cholesterol esterifying enzyme, lecithin-cholesterol acyltransferase. The esterification of cholesterol is required for cholesterol transport. Mutations in this gene have been found to cause fish-eye disease as well as LCAT deficiency.

Source: NCBI Gene 3931 — RefSeq curated summary.

At a glance

• Gene–disease (curated): LCAT deficiency (Definitive, GenCC) — +2 more curated relationships
• GWAS associations: 21
• Clinical variants (ClinVar): 352 total — 19 pathogenic, 15 likely-pathogenic
• Phenotypes (HPO): 21
• Druggable target: yes
• MANE Select transcript: NM_000229

Identifiers

Gene identifiers

Field	Value
HGNC ID	HGNC:6522
Approved symbol	LCAT
Name	lecithin-cholesterol acyltransferase
Location	16q22.1
Locus type	gene with protein product
Status	Approved
Ensembl gene	ENSG00000213398
Ensembl biotype	protein_coding
OMIM	606967
Entrez	3931

Gene structure

Transcript identifiers

Ensembl transcripts: 9 — 5 protein_coding, 2 nonsense_mediated_decay, 2 retained_intron

ENST00000264005, ENST00000570369, ENST00000570396, ENST00000570980, ENST00000573538, ENST00000573846, ENST00000575277, ENST00000575467, ENST00000576450

RefSeq mRNA: 1 — MANE Select: NM_000229 NM_000229

CCDS: CCDS10854

Canonical transcript exons

ENST00000264005 — 6 exons

Exon	Start	End
ENSE00000692168	67943056	67943212
ENSE00001308070	67939750	67940478
ENSE00003478042	67942671	67942766
ENSE00003564538	67942861	67942976
ENSE00003600601	67942363	67942587
ENSE00003893923	67943948	67944120

Expression profiles

Bgee: expression breadth ubiquitous, 194 present calls, max score 99.12.

FANTOM5 (CAGE): breadth broad, TPM avg 1.4337 / max 324.4416, expressed in 183 samples.

FANTOM5 promoters (1 alternative TSS)

Promoter ID	TPM avg	Samples expressed
157834	1.4337	183

Top tissues by expression

280 total, by Bgee expression score (0-100, higher = more expressed):

Tissue	Anatomy ID	Expression score	Quality
right lobe of liver	UBERON:0001114	99.12	gold quality
right hemisphere of cerebellum	UBERON:0014890	96.16	gold quality
tibial nerve	UBERON:0001323	96.14	gold quality
left lobe of thyroid gland	UBERON:0001120	96.11	gold quality
apex of heart	UBERON:0002098	96.05	gold quality
liver	UBERON:0002107	96.01	gold quality
right lobe of thyroid gland	UBERON:0001119	95.80	gold quality
ascending aorta	UBERON:0001496	95.68	gold quality
right coronary artery	UBERON:0001625	95.64	gold quality
thoracic aorta	UBERON:0001515	95.55	gold quality
endocervix	UBERON:0000458	95.54	gold quality
body of uterus	UBERON:0009853	95.49	gold quality
lower esophagus muscularis layer	UBERON:0035833	95.13	gold quality
lower esophagus	UBERON:0013473	95.10	gold quality
left coronary artery	UBERON:0001626	94.93	gold quality
esophagogastric junction muscularis propria	UBERON:0035841	94.92	gold quality
right lung	UBERON:0002167	94.81	gold quality
cerebellar hemisphere	UBERON:0002245	94.78	gold quality
descending thoracic aorta	UBERON:0002345	94.76	gold quality
upper lobe of left lung	UBERON:0008952	94.76	gold quality
cerebellar cortex	UBERON:0002129	94.66	gold quality
metanephros cortex	UBERON:0010533	94.49	gold quality
left uterine tube	UBERON:0001303	94.45	gold quality
aorta	UBERON:0000947	94.33	gold quality
thyroid gland	UBERON:0002046	94.30	gold quality
omental fat pad	UBERON:0010414	94.07	gold quality
coronary artery	UBERON:0001621	94.03	gold quality
peritoneum	UBERON:0002358	93.99	gold quality
muscle layer of sigmoid colon	UBERON:0035805	93.89	gold quality
mucosa of stomach	UBERON:0001199	93.83	gold quality

Single-cell (SCXA)

Detected in 1 experiment(s), a significant marker in 1.

Experiment	Marker?	Max mean expression
E-ANND-3	yes	3.57

Regulation

Is transcription factor: no

Upstream regulators (CollecTRI, top): PAX3, SP1, SP3, STAT3

Literature-anchored findings (GeneRIF, showing 40)

• The lecithin:cholesterol acyltransferase (LCAT) activity in cord blood was extremely low. (PMID:11882335)
• Results suggest that LCAT may take part in the detoxification of oxidants even after the loss of its cholesterol esterification function. (PMID:11966470)
• IL-6 induced activation of full-length LCAT promoter activity. A minimal IL-6 response element mapped within the distal promoter and was sufficient to mediate the IL-6 response (PMID:12032172)
• In conclusion, 12% of Hypoalphalipoproteinemia subjects were found to carry mutations in apo A-I, LCAT, or GBA genes (PMID:12048121)
• The results suggest that as follicle maturation progresses, Toc and Asc concentrations increase in follicular fluid, thus protecting LCAT from oxidative damage and loss of activity (PMID:12051518)
• Endotoxin-lipoprotein complex formation as a factor in atherogenesis: associations with hyperlipidemia and with lecithin:cholesterol acyltransferase activity (PMID:12139471)
• comparison of 5 natural point mutations of apo A-I illustrates that a specific sequence between amino acids 110 and 162 is required for LCAT activation (PMID:12573451)
• 608T polymorphism of LCAT gene was associated with higher plasma HDL-C level in Coronary Arteriosclerosis patients. (PMID:12673583)
• Familial lecithin:cholesterol acyltransferase (LCAT) deficiency, arising from mutation, is associated with complete absence of Lecithin:cholesterol acyltransferase (LCAT) activity. (PMID:12957688)
• Postprandial chylomicrons may play an important role in promoting reverse cholesterol transport in vivo by serving as the preferred ultimate vehicle for transporting cholesterol released from cell membranes to the liver via LCAT and CETP. (PMID:15102891)
• T13M mutation of LCAT gene causes fish-eye disease. (PMID:15115696)
• tested whether rare sequence variants of ABCA1, APOA1, and LCAT collectively contribute to variation in plasma levels of high density lipoprotein cholesterol; nonsynonymous sequence variants were significantly more common in individuals with low HDL-C (PMID:15297675)
• role of LCAT in estradiol-2 esterification and its involvement in antioxidant protection of HDL (PMID:15472210)
• Selective LCAT-mediated reactivity with pre-beta 1-high density lipoprotein represents a novel mechanism increasing the efficiency of reverse cholesterol transport. (PMID:15544352)
• Deficiency is manifested by fish eye disease. (PMID:15936482)
• the inheritance of a mutated LCAT genotype causes a gene-dose-dependent alteration in the plasma lipid/lipoprotein profile, which is remarkably similar between subjects classified as LCAT deficiency or fish eye disease (PMID:15994445)
• a novel G–>A mutation in exon 6 of LCAT gene, which resulted in an amino acid substitution of valine for methionine in familial LCAT deficiency (PMID:16051254)
• Mutation underlies increased carotid artery intima-media thickness, which suggests that LCAT protects against atherosclerosis. (PMID:16061733)
• analysis of mutations in the LCAT gene that result in an intermediate phenotype between LCAT-deficiency and fish-eye disease (case report) (PMID:16216249)
• Decreased LDL in LCAT-deficiency is attributable to increased catabolism caused by rapid catabolism of abnormal LDL and upregulation of LDL receptor pathway. Decreased catabolism of LpX contributes to its accumulation in LCAT-deficiency. (Lipoprotein X) (PMID:16543491)
• Redistribution of charged residues in mutant LCAT may be a major factor responsible for the dramatically reduced activity of the enzyme with HDL and reconstituted high density lipoprotein (rHDL). (PMID:16780378)
• The LCAT 608C/T polymorphism is possibly a predisposing factor in atherosclerotic cerebral infarction happening of Chinese Han population. (PMID:16883530)
• HDL particles generated by this pathway may account at least for some of the atheroprotective functions of apolipoprotein E (PMID:17206937)
• Apolipoprotein A=I activation affects the kinetics of the LCAT reaction (PMID:17216278)
• decreased cholesterol-total cholesterol ratio in the plasma of human apolipoprotein A-I transgenic Scavenger Receptors, Class B, Type I-/- mice is attributed to a reduction in LCAT reactivity with sphingomyelin-enriched HDL particles (PMID:17272829)
• analysis of mutations in LCAT in two patients with severe LCAT deficiency [case report] (PMID:17526537)
• The study establishes that apoA-I(Leu141Arg)Pisa and apoA-I(Leu159Arg)FIN inhibit an early step in the biogenesis of HDL due to inefficient esterification of the cholesterol of the prebeta1-HDL particles by the endogenous LCAT. (PMID:17711302)
• Multiple linear regression analysis confirmed the independent association of cellular cholesterol efflux to plasma with cholesteryl ester transfer protein genotype (PMID:18178167)
• Analysis of the LCAT gene showed two novel point mutations in exon 2 and exon 3, leading to amino acid substitution. As far, these mutations have not been reported in FLD or FED patients. (PMID:18397721)
• HbA1c provides an easy-to-assess, accurate measure of LCAT activity in type 2 diabetes. (PMID:18485513)
• oxidation of a single Met in apoA-I results in impaired LCAT activation (PMID:18719109)
• Plasma LCAT activity is elevated in metabolic syndrome and may be a marker of subclinical atherosclerosis (PMID:18782872)
• Functional LCAT is not required for macrophage cholesterol efflux to human serum. (PMID:18922527)
• Recent research findings from animal and human studies have revealed a potential beneficial role of LCAT in reducing atherosclerosis–REVIEW (PMID:19306528)
• Suggest that a LCAT-/- genotype associated with an APOE epsilon2 allele could be a novel mechanism leading to dysbetalipoproteinemia in familial LCAT deficient patients. (PMID:19515369)
• Low LCAT plasma levels are not associated with increased atherosclerosis in the general population. (PMID:19671930)
• Genetically determined low LCAT activity in Italian families is not associated with enhanced preclinical atherosclerosis despite low high-density lipoprotein cholesterol levels. (PMID:19687369)
• lecithin:cholesterol acyltransferase activity is not responsible for low incidence of cardiovascular events (PMID:19698944)
• Plasma lecithin: cholesterol acyltransferase activity modifies the inverse relationship of C-reactive protein with HDL cholesterol in nondiabetic men. (PMID:19800416)
• Novel N-terminal mutation of human apolipoprotein A-I reduces self-association and impairs LCAT activation (PMID:20884842)

Cross-species orthologs

3 orthologs

Organism	Symbol	Gene ID
danio_rerio	lcat	ENSDARG00000099424
mus_musculus	Lcat	ENSMUSG00000035237
rattus_norvegicus	Lcat	ENSRNOG00000019573

Paralogs (1): PLA2G15 (ENSG00000103066)

Protein

Protein identifiers

Phosphatidylcholine-sterol acyltransferase — P04180 (reviewed: P04180)

Alternative names: 1-alkyl-2-acetylglycerophosphocholine esterase, Lecithin-cholesterol acyltransferase, Phospholipid-cholesterol acyltransferase, Platelet-activating factor acetylhydrolase

All UniProt accessions (8): A0A140VK24, P04180, I3L0J6, I3L1Q6, I3L215, I3L3R0, J3QKT0, J3QSE5

UniProt curated annotations — full annotation on UniProt →

Function. Central enzyme in the extracellular metabolism of plasma lipoproteins. Synthesized mainly in the liver and secreted into plasma where it converts cholesterol and phosphatidylcholines (lecithins) to cholesteryl esters and lysophosphatidylcholines on the surface of high and low density lipoproteins (HDLs and LDLs). The cholesterol ester is then transported back to the liver. Has a preference for plasma 16:0-18:2 or 18:O-18:2 phosphatidylcholines. Also produced in the brain by primary astrocytes, and esterifies free cholesterol on nascent APOE-containing lipoproteins secreted from glia and influences cerebral spinal fluid (CSF) APOE- and APOA1 levels. Together with APOE and the cholesterol transporter ABCA1, plays a key role in the maturation of glial-derived, nascent lipoproteins. Required for remodeling high-density lipoprotein particles into their spherical forms. Catalyzes the hydrolysis of 1-O-alkyl-2-acetyl-sn-glycero-3-phosphocholine (platelet-activating factor or PAF) to 1-O-alkyl-sn-glycero-3-phosphocholine (lyso-PAF). Also catalyzes the transfer of the acetate group from PAF to 1-hexadecanoyl-sn-glycero-3-phosphocholine forming lyso-PAF. Catalyzes the esterification of (24S)-hydroxycholesterol (24(S)OH-C), also known as cerebrosterol to produce 24(S)OH-C monoesters.

Subcellular location. Secreted.

Tissue specificity. Detected in blood plasma. Detected in cerebral spinal fluid (at protein level). Detected in liver. Expressed mainly in brain, liver and testes.

Post-translational modifications. O- and N-glycosylated. O-glycosylation on Thr-431 and Ser-433 consists of sialylated galactose beta 1–>3N-acetylgalactosamine structures. N-glycosylated sites contain sialylated triantennary and/or biantennary complex structures.

Disease relevance. Lecithin-cholesterol acyltransferase deficiency (LCATD) [MIM:245900] A disorder of lipoprotein metabolism characterized by inadequate esterification of plasmatic cholesterol. Two clinical forms are recognized: complete LCAT deficiency and fish-eye disease. LCATD is generally referred to the complete form which is associated with absence of both alpha and beta LCAT activities resulting in esterification anomalies involving both HDL (alpha-LCAT activity) and LDL (beta-LCAT activity). It causes a typical triad of diffuse corneal opacities, target cell hemolytic anemia, and proteinuria with renal failure. The disease is caused by variants affecting the gene represented in this entry. Fish-eye disease (FED) [MIM:136120] A disorder of lipoprotein metabolism due to partial lecithin-cholesterol acyltransferase deficiency that affects only alpha-LCAT activity. FED is characterized by low plasma HDL and corneal opacities due to accumulation of cholesterol deposits in the cornea (‘fish-eye’). The disease is caused by variants affecting the gene represented in this entry.

Activity regulation. APOA1 is the most potent activator in plasma. Also activated by APOE, APOC1 and APOA4. Inhibited by haptoglobin and 5,5’-dithiobis-(2-nitrobenzoic acid) (DTNB).

Miscellaneous. Levels of LCAT activity correlates inversely with leptin levels as well as with obesity for a wide range of BMI values.

Similarity. Belongs to the AB hydrolase superfamily. Lipase family.

RefSeq proteins (1): NP_000220* (*=MANE)

Domains & families (InterPro)

ID	Name	Type
IPR003386	LACT/PDAT_acylTrfase	Family
IPR029058	AB_hydrolase_fold	Homologous_superfamily

Pfam: PF02450

Enzyme classification (BRENDA):

• EC 2.3.1.43 — phosphatidylcholine-sterol O-acyltransferase (BRENDA: 17 organisms, 163 substrates, 53 inhibitors, 25 Km, 0 kcat entries)

Substrate kinetics (BRENDA)

7 substrates with measured Km, best-characterized 7. Km ranges are aggregated across organisms/conditions.

Substrate	Km (mM)	Measurements
1,2-BIS-(1-PYRENEBUTANOYL)-SN-GLYCERO-3-PHOSPHOC	0.14–20.5	6
1-PALMITOYL-2-OLEOYL-SN-GLYCERO-3-PHOSPHOCHOLINE	0.0006–0.027	3
CHOLESTEROL	0.57–21.4	3
PHOSPHATIDYLCHOLINE	—	2
1,2-BIS[4-(1-PYRENO)BUTANOYL]-SN-GLYCERO-3-PHOSP	0.0006	1
APOLIPOPROTEIN-A-I	0.0001	1
DEHYDROERGOSTEROL	0.003	1

Catalyzed reactions (Rhea), 12 shown:

• a 1-O-alkyl-2-acetyl-sn-glycero-3-phosphocholine + H2O = a 1-O-alkyl-sn-glycero-3-phosphocholine + acetate + H(+) (RHEA:17777)
• a sterol + a 1,2-diacyl-sn-glycero-3-phosphocholine = a sterol ester + a 1-acyl-sn-glycero-3-phosphocholine (RHEA:21204)
• 1-hexadecanoyl-2-(5Z,8Z,11Z,14Z-eicosatetraenoyl)-sn-glycero-3-phosphocholine + H2O = 1-hexadecanoyl-sn-glycero-3-phosphocholine + (5Z,8Z,11Z,14Z)-eicosatetraenoate + H(+) (RHEA:40427)
• 1-hexadecanoyl-2-(9Z,12Z-octadecadienoyl)-sn-glycero-3-phosphocholine + H2O = (9Z,12Z)-octadecadienoate + 1-hexadecanoyl-sn-glycero-3-phosphocholine + H(+) (RHEA:40811)
• a 1-hexadecanoyl-2-acyl-sn-glycero-3-phosphocholine + (24S)-hydroxycholesterol = (24S)-24-hydroxycholesterol ester + 1-hexadecanoyl-sn-glycero-3-phosphocholine (RHEA:43216)
• (24S)-hydroxycholesterol + 1-hexadecanoyl-2-(9Z,12Z-octadecadienoyl)-sn-glycero-3-phosphocholine = (24S)-hydroxycholesterol 3-linoleoate + 1-hexadecanoyl-sn-glycero-3-phosphocholine (RHEA:43224)
• 1-hexadecanoyl-2-(5Z,8Z,11Z,14Z-eicosatetraenoyl)-sn-glycero-3-phosphocholine + cholesterol = cholesteryl (5Z,8Z,11Z,14Z)-eicosatetraenoate + 1-hexadecanoyl-sn-glycero-3-phosphocholine (RHEA:53448)
• 1-hexadecanoyl-2-(9Z-octadecenoyl)-sn-glycero-3-phosphocholine + cholesterol = cholesteryl (9Z-octadecenoate) + 1-hexadecanoyl-sn-glycero-3-phosphocholine (RHEA:53456)
• 1-hexadecanoyl-2-(5Z,8Z,11Z-eicosatrienoyl)-sn-glycero-3-phosphocholine + cholesterol = cholesteryl (5Z,8Z,11Z)-eicosatrienoate + 1-hexadecanoyl-sn-glycero-3-phosphocholine (RHEA:53460)
• 1-hexadecanoyl-2-(8Z,11Z,14Z-eicosatrienoyl)-sn-glycero-3-phosphocholine + cholesterol = cholesteryl (8Z,11Z,14Z)-eicosatrienoate + 1-hexadecanoyl-sn-glycero-3-phosphocholine (RHEA:53464)
• 1-hexadecanoyl-2-(5Z,8Z,11Z,14Z,17Z-eicosapentaenoyl)-sn-glycero-3-phosphocholine + cholesterol = (5Z,8Z,11Z,14Z,17Z-eicosapentaenoyl)-cholesterol + 1-hexadecanoyl-sn-glycero-3-phosphocholine (RHEA:53468)
• 1-hexadecanoyl-2-(9Z,12Z-octadecadienoyl)-sn-glycero-3-phosphocholine + cholesterol = cholesteryl (9Z,12Z)-octadecadienoate + 1-hexadecanoyl-sn-glycero-3-phosphocholine (RHEA:53472)

UniProt features (111 total): sequence variant 44, strand 19, helix 17, turn 11, glycosylation site 6, mutagenesis site 5, active site 3, disulfide bond 2, signal peptide 1, chain 1, site 1, sequence conflict 1

Structure

Experimental structures (PDB)

7 structures.

PDB	Method	Resolution (Å)
5BV7	X-RAY DIFFRACTION	2.45
4XWG	X-RAY DIFFRACTION	2.65
5TXF	X-RAY DIFFRACTION	3.1
6MVD	X-RAY DIFFRACTION	3.1
4XX1	X-RAY DIFFRACTION	3.6
4X96	X-RAY DIFFRACTION	8.69
9MXZ	ELECTRON MICROSCOPY	9.8

Predicted structure (AlphaFold)

Model	pLDDT	Fraction very-high
AF-P04180-F1	87.33	0.73

Antibody-complex structures (SAbDab): 3 — 4XWG, 4XX1, 5BV7

Functional residue map

Curated UniProt residues grouped by drug-discovery relevance — catalytic, ligand-binding, modification, and mutation-validated positions. Source: UniProtKB sequence features.

Catalytic / active sites (4): 205 (nucleophile); 369 (charge relay system); 401 (charge relay system); 173 (determinant for substrate specificity)

Disulfide bonds (2): 74–98, 337–380

Glycosylation sites (6): 431, 433, 44, 108, 296, 408

Mutagenesis-validated functional residues (5):

Position	Phenotype
173	increased activity towards papc. increased papc/popc activity ratio.
173	little change in enzyme specific activity nor in papc/popc activity ratio.
173	decreased enzyme specific activity. increased papc/popc activity ratio.

Function

Pathways and Gene Ontology

Reactome pathways

4 pathways

ID	Pathway
R-HSA-8964058	HDL remodeling
R-HSA-174824	Plasma lipoprotein assembly, remodeling, and clearance
R-HSA-382551	Transport of small molecules
R-HSA-8963899	Plasma lipoprotein remodeling

MSigDB gene sets: 239 (showing top): GOBP_PHOSPHOLIPID_METABOLIC_PROCESS, GOBP_PHOSPHATIDYLCHOLINE_METABOLIC_PROCESS, chr16q22, GOBP_STEROL_HOMEOSTASIS, GOBP_LIPOPROTEIN_METABOLIC_PROCESS, GOBP_PHOSPHATIDYLCHOLINE_BIOSYNTHETIC_PROCESS, GNF2_GSTM1, GOBP_RESPONSE_TO_CORTICOSTEROID, LFA1_Q6, GNF2_HPN, LINDGREN_BLADDER_CANCER_CLUSTER_3_DN, GOBP_ORGANOPHOSPHATE_METABOLIC_PROCESS, GOBP_RESPONSE_TO_COPPER_ION, GOBP_PROTEIN_LIPID_COMPLEX_ASSEMBLY, GOBP_REGULATION_OF_CELLULAR_COMPONENT_BIOGENESIS

GO Biological Process (17): lipid metabolic process (GO:0006629), phospholipid metabolic process (GO:0006644), phosphatidylcholine biosynthetic process (GO:0006656), cholesterol metabolic process (GO:0008203), cholesterol transport (GO:0030301), very-low-density lipoprotein particle remodeling (GO:0034372), high-density lipoprotein particle remodeling (GO:0034375), lipoprotein biosynthetic process (GO:0042158), cholesterol homeostasis (GO:0042632), reverse cholesterol transport (GO:0043691), aflatoxin metabolic process (GO:0046222), phosphatidylcholine metabolic process (GO:0046470), response to copper ion (GO:0046688), response to glucocorticoid (GO:0051384), regulation of high-density lipoprotein particle assembly (GO:0090107), steroid metabolic process (GO:0008202), lipoprotein metabolic process (GO:0042157)

GO Molecular Function (11): 1-alkyl-2-acetylglycerophosphocholine esterase activity (GO:0003847), phosphatidylcholine-sterol O-acyltransferase activity (GO:0004607), A2-type glycerophospholipase activity (GO:0004623), sterol ester esterase activity (GO:0004771), apolipoprotein A-I binding (GO:0034186), platelet-activating factor acetyltransferase activity (GO:0047179), protein binding (GO:0005515), obsolete O-acyltransferase activity (GO:0008374), transferase activity (GO:0016740), acyltransferase activity (GO:0016746), hydrolase activity (GO:0016787)

GO Cellular Component (5): extracellular region (GO:0005576), obsolete extracellular space (GO:0005615), high-density lipoprotein particle (GO:0034364), extracellular exosome (GO:0070062), membrane (GO:0016020)

Reactome top-level categories

Rollup of top-3 pathways:

Category	Pathways
Plasma lipoprotein remodeling	1
Transport of small molecules	1
Plasma lipoprotein assembly, remodeling, and clearance	1

GO top-level categories

Rollup of top GO terms by namespace:

Category	Terms
carboxylic ester hydrolase activity	3
lipid metabolic process	2
catalytic activity	2
cellular anatomical structure	2
primary metabolic process	1
organophosphate metabolic process	1
phosphatidylcholine metabolic process	1
glycerophospholipid biosynthetic process	1
sterol metabolic process	1
secondary alcohol metabolic process	1
sterol transport	1
triglyceride-rich lipoprotein particle remodeling	1
plasma lipoprotein particle remodeling	1
macromolecule biosynthetic process	1
lipoprotein metabolic process	1
sterol homeostasis	1
cholesterol transport	1
mycotoxin metabolic process	1
glycerophospholipid metabolic process	1
response to metal ion	1
response to corticosteroid	1
high-density lipoprotein particle assembly	1
regulation of protein-containing complex assembly	1
regulation of multicellular organismal process	1
protein metabolic process	1
acyltransferase activity, transferring groups other than amino-acyl groups	1
glycerophospholipase activity	1
lipase activity	1
apolipoprotein binding	1
acetyltransferase activity	1
binding	1
transferase activity	1
plasma lipoprotein particle	1
extracellular vesicle	1

Protein interactions and networks

STRING

1590 interactions, top by confidence (×1000):

Protein A	Protein B	Partner UniProt	Score
LCAT	APOA1	P02647	997
LCAT	CETP	P11597	955
LCAT	APOB	P04114	938
LCAT	APOA2	P02652	912
LCAT	ABCA1	O95477	911
LCAT	PON1	P27169	874
LCAT	APOD	P05090	856
LCAT	APOC3	P02656	849
LCAT	PLTP	P55058	828
LCAT	PSKH1	P11801	810
LCAT	APOE	P02649	800
LCAT	SCARB1	Q8WTV0	794
LCAT	LIPC	P11150	778
LCAT	APOC2	P02655	777
LCAT	LPL	P06858	777

IntAct

6 interactions, top by confidence:

A	B	Type	Score
APOA1	LCAT	psi-mi:“MI:0407”(direct interaction)	0.620
LCAT	WFS1	psi-mi:“MI:0915”(physical association)	0.560

BioGRID (13): APOA1 (Reconstituted Complex), PIKFYVE (Co-crystal Structure), LCAT (Affinity Capture-Western), LCAT (Reconstituted Complex), LCAT (Reconstituted Complex), LCAT (Reconstituted Complex), LCAT (Co-purification), LCAT (Co-purification), APOA1 (Biochemical Activity), APOA1 (Reconstituted Complex), APOA1 (Biochemical Activity), APOE (Biochemical Activity), LCAT (Two-hybrid)

ESM2 similar proteins: A0A3Q2HW92, A2AIG8, A4FV98, A5PK45, A6NDV4, A6QLK4, A6QQ24, D3YWP0, F1PZV2, O09009, O15303, O35790, O55240, P04180, P35349, Q08758, Q0P5C0, Q1JPJ0, Q3T0A0, Q3T9M1, Q3U481, Q501J2, Q5E9V4, Q5SX19, Q6AYG0, Q6NUT3, Q863I4, Q86VF5, Q86VU5, Q8BVM4, Q8K297, Q8NBJ5, Q8R2H9, Q8TCT0, Q8TCT1, Q8TCT7, Q8VCE6, Q92781, Q96AZ1, Q96FB5

Diamond homologs: O35502, O35573, O35724, O35840, P04180, P16301, P18424, P30930, P53760, P53761, Q08758, Q675A5, Q6XPZ3, Q8NCC3, Q8VEB4, Q8WMP9, Q9FZI8, P40345, Q10PI6, Q4VCM1, Q9FYC7, Q9FNA9

SIGNOR signaling

1 interactions.

A	Effect	B	Mechanism
APOA1	“up-regulates activity”	LCAT	binding

Disease & clinical

Clinical variants and AI predictions

ClinVar

352 variants total. Per-class counts are floors (≥ shown; pagination cap):

Classification	Count (floor)
Pathogenic	19
Likely pathogenic	15
Uncertain significance	143
Likely benign	131
Benign	8

Top pathogenic / likely-pathogenic (30)

Variant ID	HGVS	Classification
1969478	NM_000229.2(LCAT):c.23G>A (p.Trp8Ter)	Pathogenic
2736369	NM_000229.2(LCAT):c.803G>A (p.Arg268His)	Pathogenic
2879462	NM_000229.2(LCAT):c.849G>A (p.Trp283Ter)	Pathogenic
3581207	NM_000229.2(LCAT):c.511C>T (p.Arg171Trp)	Pathogenic
3656	NM_000229.2(LCAT):c.508T>C (p.Trp170Arg)	Pathogenic
3657	NM_000229.2(LCAT):c.951G>A (p.Met317Ile)	Pathogenic
3658	NM_000229.2(LCAT):c.491_493dup (p.Arg164_Ala165insGly)	Pathogenic
3659	NM_000229.2(LCAT):c.756C>A (p.Asn252Lys)	Pathogenic
3660	NM_000229.2(LCAT):c.440C>T (p.Thr147Ile)	Pathogenic
3661	NM_000229.2(LCAT):c.101dup (p.His35fs)	Pathogenic
3662	NM_000229.2(LCAT):c.101C>T (p.Pro34Leu)	Pathogenic
3665	NM_000229.2(LCAT):c.827T>A (p.Met276Lys)	Pathogenic
3666	NM_000229.2(LCAT):c.698T>C (p.Leu233Pro)	Pathogenic
3669	NM_000229.2(LCAT):c.1197dup (p.Gln400fs)	Pathogenic
3672	NM_000229.2(LCAT):c.969CCT[1] (p.Leu325del)	Pathogenic
3673	NM_000229.2(LCAT):c.463A>G (p.Asn155Asp)	Pathogenic
3674	NM_000229.2(LCAT):c.524-22T>C	Pathogenic
626357	NM_000229.2(LCAT):c.997G>A (p.Val333Met)	Pathogenic
626358	NM_000229.2(LCAT):c.1210A>G (p.Met404Val)	Pathogenic
1338728	NM_000229.2(LCAT):c.802C>T (p.Arg268Cys)	Likely pathogenic
1764448	NM_000229.2(LCAT):c.1267C>T (p.Arg423Cys)	Likely pathogenic
2585273	NM_000229.2(LCAT):c.428-2A>T	Likely pathogenic
2631093	NM_000229.2(LCAT):c.160G>A (p.Gly54Ser)	Likely pathogenic
3067749	NM_000229.2(LCAT):c.713del (p.Gly238fs)	Likely pathogenic
3581201	NM_000229.2(LCAT):c.799C>T (p.Gln267Ter)	Likely pathogenic
3581202	NM_000229.2(LCAT):c.794_795del (p.Glu265fs)	Likely pathogenic
3581209	NM_000229.2(LCAT):c.488_489del (p.Val163fs)	Likely pathogenic
3581214	NM_000229.2(LCAT):c.254G>A (p.Trp85Ter)	Likely pathogenic
3581217	NM_000229.2(LCAT):c.142_154+1dup	Likely pathogenic
3581220	NM_000229.2(LCAT):c.115A>T (p.Lys39Ter)	Likely pathogenic

SpliceAI

1035 predictions. Top by Δscore:

Variant	Effect	Δscore
16:67940480:T:G	acceptor_loss	1.0000
16:67942444:T:TA	donor_gain	1.0000
16:67942588:C:CC	acceptor_gain	1.0000
16:67942666:CTCA:C	donor_loss	1.0000
16:67942667:TCA:T	donor_loss	1.0000
16:67942668:CACCG:C	donor_loss	1.0000
16:67942669:A:AC	donor_gain	1.0000
16:67942669:A:T	donor_loss	1.0000
16:67942670:C:A	donor_loss	1.0000
16:67942670:C:CC	donor_gain	1.0000
16:67942670:CCGGG:C	donor_gain	1.0000
16:67942765:CC:C	acceptor_gain	1.0000
16:67942766:CC:C	acceptor_gain	1.0000
16:67942856:CAAA:C	donor_loss	1.0000
16:67942857:AAACC:A	donor_loss	1.0000
16:67942858:AAC:A	donor_loss	1.0000
16:67942859:A:T	donor_loss	1.0000
16:67942860:C:A	donor_loss	1.0000
16:67942972:CAACC:C	acceptor_gain	1.0000
16:67942975:CC:C	acceptor_gain	1.0000
16:67942975:CCCT:C	acceptor_loss	1.0000
16:67942976:CC:C	acceptor_gain	1.0000
16:67942976:CCT:C	acceptor_loss	1.0000
16:67942977:C:CC	acceptor_gain	1.0000
16:67942978:T:A	acceptor_loss	1.0000
16:67943053:TA:T	donor_loss	1.0000
16:67943054:A:AT	donor_loss	1.0000
16:67943055:CCT:C	donor_loss	1.0000
16:67943213:C:CC	acceptor_gain	1.0000
16:67940476:CAC:C	acceptor_gain	0.9900

AlphaMissense

2864 scored. Top likely-pathogenic:

Variant	Protein change	am_pathogenicity
16:67943074:C:G	C98S	0.998
16:67943075:A:T	C98S	0.998
16:67943146:C:G	C74S	0.998
16:67943147:A:T	C74S	0.998
16:67942497:C:A	S205I	0.997
16:67942739:A:G	L152P	0.996
16:67943074:C:T	C98Y	0.996
16:67943075:A:G	C98R	0.996
16:67943145:G:C	C74W	0.996
16:67943147:A:G	C74R	0.996
16:67943183:C:G	A62P	0.996
16:67940121:T:A	D369V	0.995
16:67942686:A:G	W170R	0.995
16:67942686:A:T	W170R	0.995
16:67943074:C:A	C98F	0.995
16:67943114:A:G	W85R	0.995
16:67943114:A:T	W85R	0.995
16:67943146:C:T	C74Y	0.995
16:67943193:A:C	N58K	0.995
16:67943193:A:T	N58K	0.995
16:67940122:C:G	D369H	0.994
16:67942419:A:T	I231N	0.994
16:67942431:A:T	I227N	0.994
16:67942442:C:A	K223N	0.994
16:67942442:C:G	K223N	0.994
16:67942703:C:G	R164P	0.994
16:67943073:G:C	C98W	0.994
16:67943146:C:A	C74F	0.994
16:67943188:A:G	L60P	0.994
16:67940099:G:C	S376R	0.993

dbSNP variants (sampled 300 via entrez): RS1001326689 (16:67939513 T>C), RS1001452642 (16:67942544 C>T), RS1001580962 (16:67945274 TC>T), RS1002920761 (16:67939502 C>T), RS1002935033 (16:67939686 G>A,T), RS1003101800 (16:67944195 G>A,T), RS1003139704 (16:67942571 G>A), RS1003444341 (16:67942772 G>A,C), RS1003769357 (16:67944193 A>G), RS1003800705 (16:67943911 T>C,G), RS1004727769 (16:67941827 T>C), RS1004923760 (16:67941342 C>T), RS1005551039 (16:67941574 G>A,T), RS1005597574 (16:67944230 G>A,T), RS1005931116 (16:67942791 G>A,C)

Disease associations

OMIM: gene MIM:606967 | disease phenotypes: MIM:245900, MIM:136120

GenCC curated gene-disease

Disease	Classification	Inheritance
LCAT deficiency	Definitive	Autosomal recessive
fish eye disease	Definitive	Autosomal recessive
Norum disease	Strong	Autosomal recessive

Mondo (3): Norum disease (MONDO:0009515), fish eye disease (MONDO:0007620), LCAT deficiency (MONDO:0018999)

Orphanet (3): Familial LCAT deficiency (Orphanet:79293), LCAT deficiency (Orphanet:650), Fish-eye disease (Orphanet:79292)

HPO phenotypes

21 total (21 of 21 shown, HPO-id order):

HPO	Term
HP:0000007	Autosomal recessive inheritance
HP:0000083	Renal insufficiency
HP:0000093	Proteinuria
HP:0000505	Visual impairment
HP:0001084	Corneal arcus
HP:0001681	Angina pectoris
HP:0001744	Splenomegaly
HP:0001878	Hemolytic anemia
HP:0001895	Normochromic anemia
HP:0002155	Hypertriglyceridemia
HP:0002240	Hepatomegaly
HP:0002621	Atherosclerosis
HP:0002716	Lymphadenopathy
HP:0003141	Increased LDL cholesterol concentration
HP:0003233	Decreased HDL cholesterol concentration
HP:0003362	Increased VLDL cholesterol concentration
HP:0003581	Adult onset
HP:0003651	Foam cells
HP:0007759	Opacification of the corneal stroma
HP:0007957	Corneal opacity
HP:0025433	Decreased lecithin cholesterol acyl transferase level

GWAS associations

21 associations (top):

Study	Trait	p-value
GCST000135_8	HDL cholesterol	1.000000e-07
GCST000284_3	HDL cholesterol	3.000000e-08
GCST000290_13	HDL cholesterol	9.000000e-13
GCST000755_31	HDL cholesterol	8.000000e-33
GCST000805_12	HDL cholesterol	2.000000e-07
GCST000974_7	HDL cholesterol	2.000000e-11
GCST002223_11	HDL cholesterol	8.000000e-54
GCST002539_84	Schizophrenia	2.000000e-08
GCST002899_33	HDL cholesterol	9.000000e-26
GCST004232_35	HDL cholesterol levels	3.000000e-09
GCST004232_4	HDL cholesterol levels	3.000000e-57
GCST004234_11	HDL cholesterol levels	4.000000e-10
GCST005446_33	Total cholesterol levels in HDL	6.000000e-10
GCST006614_16	Total cholesterol levels	1.000000e-14
GCST006803_42	Schizophrenia	4.000000e-08
GCST007850_6	HDL cholesterol	2.000000e-10
GCST008035_65	High density lipoprotein cholesterol levels	2.000000e-16
GCST008035_66	High density lipoprotein cholesterol levels	1.000000e-06
GCST010002_113	Refractive error	2.000000e-14
GCST90002397_246	Mean spheric corpuscular volume	2.000000e-32
GCST90002403_680	Red blood cell count	1.000000e-15

EFO canonical traits (3, from GWAS)

EFO ID	Trait name
EFO:0004612	high density lipoprotein cholesterol measurement
EFO:0004574	total cholesterol measurement
EFO:0004305	erythrocyte count

Drugs & pharmacology

Drug and pharmacology data

Is drug target: yes

ChEMBL targets (1): CHEMBL5942 (SINGLE PROTEIN)

PharmGKB: 1 entry (VIP=true, CPIC=false)

Binding affinities (BindingDB)

5 measured of 6 human assays (6 total across all organisms); most potent 5 below. Values come from heterogeneous assays and are not directly comparable.

Ligand	Measure	Value	Patent
(+)-4,5-Dihydroxy-4-(trifluoromethyl)-3-{1-[5-(trifluoromethyl)pyrazin-2-yl]piperidin-4-yl}-1,4,5,7-tetrahydro-6H-pyrazolo[3,4-b]pyridin-6-one	EC50	4 nM	US-10138240: Crystal of 5-hydroxy-4-(trifluoromethyl)pyrazolopyridine derivative
(+)-cis-5-Hydroxy-4-(trifluoromethyl)-3-{1-[2-(trifluoromethyl)pyrimidin-5-yl]piperidin-4-yl}-1,4,5,7-tetrahydro-6H-pyrazolo[3,4-b]pyridin-6-one	EC50	18 nM	US-10138240: Crystal of 5-hydroxy-4-(trifluoromethyl)pyrazolopyridine derivative
Difluoroketone phospholipid analogue, 3	IC50	28 nM
Difluoroketone phospholipid analogue, 2	IC50	60 nM
Difluoroketone phospholipid analogue, 4	IC50	60 nM

ChEMBL bioactivities

152 potent at pChembl≥5 of 152 total, top 50 by pChembl (potency: 10 = 0.1 nM, 6 = 1 µM).

pChembl	Type	Value	Unit	Molecule
8.40	EC50	4	nM	CHEMBL4571766
8.40	EC50	4	nM	CHEMBL4444702
8.15	EC50	7	nM	CHEMBL4543681
8.15	EC50	7	nM	CHEMBL4444091
8.15	EC50	7	nM	CHEMBL4567833
8.10	EC50	8	nM	CHEMBL4449961
8.05	EC50	9	nM	CHEMBL4438875
7.96	EC50	11	nM	CHEMBL4565710
7.92	EC50	12	nM	CHEMBL4565889
7.89	EC50	13	nM	CHEMBL4475628
7.89	EC50	13	nM	CHEMBL4449770
7.89	EC50	13	nM	CHEMBL4444770
7.82	EC50	15	nM	CHEMBL4462753
7.75	EC50	18	nM	CHEMBL4438084
7.75	EC50	18	nM	CHEMBL4435610
7.75	EC50	18	nM	CHEMBL4592989
7.75	EC50	18	nM	CHEMBL4463860
7.70	EC50	20	nM	CHEMBL4453452
7.68	EC50	21	nM	CHEMBL4438084
7.68	EC50	21	nM	CHEMBL4455643
7.68	EC50	21	nM	CHEMBL4445217
7.66	EC50	22	nM	CHEMBL4567833
7.64	EC50	23	nM	CHEMBL4443799
7.64	EC50	23	nM	CHEMBL4445235
7.62	EC50	24	nM	CHEMBL4456783
7.58	EC50	26	nM	CHEMBL4442211
7.57	EC50	27	nM	CHEMBL4445217
7.57	EC50	27	nM	CHEMBL4541184
7.57	EC50	27	nM	CHEMBL4435684
7.54	EC50	29	nM	CHEMBL4514555
7.52	EC50	30	nM	CHEMBL4536836
7.51	EC50	31	nM	CHEMBL4483570
7.51	EC50	31	nM	CHEMBL4563848
7.50	EC50	32	nM	CHEMBL4575561
7.50	EC50	32	nM	CHEMBL4461575
7.47	EC50	34	nM	CHEMBL4450717
7.46	EC50	35	nM	CHEMBL4515289
7.46	EC50	35	nM	CHEMBL4577479
7.46	EC50	35	nM	CHEMBL4466792
7.46	EC50	35	nM	CHEMBL4567145
7.43	EC50	37	nM	CHEMBL4461964
7.43	EC50	37	nM	CHEMBL4440456
7.43	EC50	37	nM	CHEMBL4448781
7.40	EC50	40	nM	CHEMBL4526869
7.39	EC50	41	nM	CHEMBL4436647
7.37	EC50	43	nM	CHEMBL4440507
7.35	EC50	45	nM	CHEMBL4435430
7.34	EC50	46	nM	CHEMBL4453731
7.32	EC50	48	nM	CHEMBL4567290
7.30	EC50	50	nM	CHEMBL4454322

CTD chemical–gene interactions

54 total (human), top 30 by PubMed support.

Chemical	Actions (top 5)	PubMed papers
Cyclosporine	decreases expression	3
bisphenol A	affects expression, affects cotreatment, decreases methylation	2
Hydrogen Peroxide	decreases expression, affects cotreatment, decreases activity, increases oxidation	2
Tobacco Smoke Pollution	decreases activity, decreases expression	2
Valproic Acid	decreases expression, increases methylation	2
Particulate Matter	decreases expression, increases abundance, increases expression	2
aristolochic acid I	increases expression	1
methylmercuric chloride	increases expression	1
triphenyl phosphate	affects expression	1
tris(2-butoxyethyl) phosphate	affects expression	1
tris(1,3-dichloro-2-propyl)phosphate	decreases expression	1
1,2-oleoylphosphatidylcholine	affects binding	1
4-hydroxy-2-nonenal	decreases expression	1
polyserine	decreases reaction, increases chemical synthesis, increases metabolic processing	1
27-hydroxycholesterol	increases metabolic processing	1
K 7174	decreases expression	1
obeticholic acid	decreases expression	1
abrine	increases expression	1
Decitabine	affects expression	1
Sunitinib	decreases expression	1
Fulvestrant	affects cotreatment, decreases methylation	1
Acetaminophen	increases expression	1
Air Pollutants	decreases expression, increases abundance	1
Benzo(a)pyrene	decreases expression	1
Cadmium	decreases expression, increases abundance	1
Chlorine	affects binding, decreases activity, increases reaction	1
Cholesterol	increases acylation	1
Cholesterol Esters	increases chemical synthesis, increases metabolic processing, decreases reaction	1
Cisplatin	affects expression	1
Copper	decreases activity	1

ChEMBL screening assays

5 unique, capped per target: 5 binding

Representative assays (with source publication via chembl_document):

Assay ID	Type	Description	Source paper
CHEMBL1015040	Binding	Inhibition of LCAT activity in human plasma assessed as change in cholesteryl ester/free cholesterol ratio by cell based assay	Design, synthesis, and biological evaluation of (2R,alphaS)-3,4-dihydro-2-[3-(1,1,2,2-tetrafluoroethoxy)phenyl]-5-[3-(trifluoromethoxy)-phenyl]-alpha-(trifluoromethyl)-1(2H)-quinolineethanol as potent and orally active cholesteryl ester transfer protein inhibitor. — J Med Chem

Clinical trials (associated diseases)

28 trials via MONDO — disease-level, not drug-specific.

Trial	Phase	Status	Title
NCT03368066	PHASE3	COMPLETED	Is Adrenal Insufficiency Under-diagnosed in Hospitalized Cirrhosis Patients?
NCT01283867	PHASE1	COMPLETED	Bioequivalence Study of Mycophenolate Mofetil 500 mg Tablets of Dr.Reddy’s Laboratories Limited Under Non-Fasting Conditions
NCT01283932	PHASE1	COMPLETED	Bioequivalence Study of Pantoprazole Sodium DR Tablets 40 mg of Dr. Reddy’s Laboratories Limited Under Fed Condition
NCT01314352	PHASE1	COMPLETED	Bioavailability Study of Desloratadine 5 mg Tablet of Dr. Reddy’s Under Fed Conditions
NCT01372306	PHASE1	COMPLETED	Galantamine Bioequivalence Study of Dr. Reddy’s Under Fed Condition
NCT01372358	PHASE1	COMPLETED	Bioavailability Study of Ciprofloxacin 1000 mg ER Tablets of Dr. Reddy’s Under Non-Fasting Conditions
NCT01503424	PHASE1	COMPLETED	Bioequivalence Study of Olanzapine Tablets, 5 mg Under Fed Study
NCT01503450	PHASE1	COMPLETED	Bioequivalence Study of Olanzapine Orally Disintegrating Tablets, 5 mg Under Fed Condition
NCT01505998	PHASE1	COMPLETED	Bioequivalence Study of Amlodipine Besylate/Benazepril HCl 10 mg/40 mg Capsules of Dr. Reddy’s Under Fed Conditions
NCT01567501	PHASE1	COMPLETED	Bioequivalence Study of Levocetirizine Dihydrochloride Tablets 5 mg Under Fed Condition
NCT01578265	PHASE1	COMPLETED	Bioequivalence Study of Ondansetron Tablets USP 8 mg Under Fed Condition
NCT01603186	PHASE1	COMPLETED	Bioequivalence Study of Quetiapine Fumarate Tablets 25 mg Under Fed Condition
NCT01618760	PHASE1	COMPLETED	Bioequivalence Study of Risperidone Tablet 1 mg Under Fed Condition
NCT01618825	PHASE1	COMPLETED	Bioequivalence Study of Lamotrigine Tablets 25 mg (2 x 25 mg Tablets) Under Fed Condition
NCT01722110	PHASE1	COMPLETED	Bioequivalence Study of Indomethacin Extended-Release Capsules 75 mg Under Fed Condition
NCT01735344	PHASE1	COMPLETED	Bioequivalence Study of Lisinopril Tablets 40 mg Under Fed Condition
NCT01735383	PHASE1	COMPLETED	Bioequivalence Study of Etodolac Tablet USP 500 mg Under Fed Condition
NCT01744873	PHASE1	COMPLETED	Bioequivalence Study of Bisoprolol Fumarate Tablet 10 mg Under Fed Condition
NCT01827878	PHASE1	COMPLETED	Bioequivalence Study of Lisinopril and Hydrochlorothiazide Tablets (20+25) mg Under Fed Conditions
NCT01884909	PHASE1	COMPLETED	Bioequivalence Study of Metoprolol Succinate Extended Release Tablets 200 mg Under Fed Conditions
NCT07396844	PHASE1	NOT_YET_RECRUITING	The Food Effect on Pharmacokinetics of Fixed Dose Combination of Gemigliptin/Dapagliflozin/Metformin in Healthy Volunteers
NCT01782027	Not specified	TERMINATED	Mendelian Reverse Cholesterol Transport Study
NCT04737720	Not specified	COMPLETED	Intravenous ACP-501 for Familial LCAT Deficiency (rhLCAT)
NCT06217588	Not specified	RECRUITING	LCAT (Lecithin Cholesterol Acyl Transferase) Natural History Study
NCT05995119	EARLY_PHASE1	COMPLETED	Comparative Bioavailability Study of TAH3311 5 mg Oral Dissolving Film vs ELIQUIS® 5 mg Tablet in Healthy Volunteers
NCT01721187	Not specified	COMPLETED	Satiety Effects on the Neural Valuation of Food
NCT03390881	Not specified	COMPLETED	Use of Breath Acetone aa a Marker of Energy Balance
NCT05837572	Not specified	UNKNOWN	Development of the Leeds Food Preference Questionnaire in Spanish