Catalyst

Reference

Evidence

CompoundsSubstances with curated findings
OutcomesMeasured endpoints
ProteinsTransporters and enzymes
InteractionsPairs that share biology
PapersEvery paper a finding cites
ArticlesReadings of the evidence

Chemistry

ReactionsRhea and Human-GEM, by subsystem
ReachWhat a compound can become

Reference layer

PathwaysReactions grouped under one name
ProcessesWhat proteins are annotated as doing
AnatomyTissues, by what is expressed there
Cell typesCells, by what is expressed there
OrganismsSpecies a compound is prepared from
MaterialsSubstances that are not one molecule
AssessmentsRegulatory determinations, transcribed
DiseasesFeatures, genes, and where each sits

Atlas Graph Twin Access

Search the graph

Search ?

Reference

Evidence

Compounds
Outcomes
Proteins
Interactions
Papers
Articles

Chemistry

Reactions
Reach

Reference layer

Pathways
Processes
Anatomy
Cell types
Organisms
Materials
Assessments
Diseases

Atlas Graph Twin Access

Loading the page.

Catalyst

Computational infrastructure for human biology, built on a provenance-aware knowledge graph. Findings retain their sources; research and simulation are the next layers.

The graph

Compounds
Outcomes
Proteins
Atlas
Interactions
Papers
Articles
Graph
Search

Reference layer

Pathways
Processes
Anatomy
Cell types
Organisms
Materials
Assessments
Diseases
Reactions
Reach

How it works

Mission
Method
Twin
How to read the evidence
Sources and licences

Build on it

Access
In your assistant
For agents
Query the graph
Bulk export

A reference for what the literature reports. Not medical advice.

Reach

a cholesterol ester

362,296 routes · scope human · up to 3 steps · built 5 October 2026 · Reactions · API

Inferred

Every route here is a hypothesis drawn from reference reactions (Rhea and Human-GEM), not a finding. Currency species such as water, ATP and NAD(H) never form a step. Evidence key

204 compounds within 3 steps: 8 in 1, 47 in 2, 149 in 3

Does it reach a compound?

Two steps

(9Z,12Z,15Z-octadecatrienoyl)-cholesterolReach from here →
1. a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
2. (9Z,12Z,15Z)-octadecatrienoyl-CoA + cholesterol = (9Z,12Z,15Z-octadecatrienoyl)-cholesterol + CoA
Catalysed by CES1, LIPN, NCEH1, LDAH and 1 moreOne of 2 routes
4beta-hydroxycholesterolReach from here →
1. a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
2. cholesterol + reduced [NADPH--hemoprotein reductase] + O2 = 4beta-hydroxycholesterol + oxidized [NADPH--hemoprotein reductase] + H2O + H(+)
Catalysed by CES1, LIPN, NCEH1, LDAH and 1 more
cholesteryl (5Z,8Z,11Z)-eicosatrienoateReach from here →
1. a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
2. 1-hexadecanoyl-2-(5Z,8Z,11Z-eicosatrienoyl)-sn-glycero-3-phosphocholine + cholesterol = cholesteryl (5Z,8Z,11Z)-eicosatrienoate + 1-hexadecanoyl-sn-glycero-3-phosphocholine
Catalysed by CES1, LIPN, NCEH1, LDAH and 1 more
(6Z,9Z,12Z-octadecatrienoyl)-cholesterolReach from here →
1. a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
2. 1-hexadecanoyl-2-(6Z,9Z,12Z-octadecatrienoyl)-sn-glycero-3-phosphocholine + cholesterol = (6Z,9Z,12Z-octadecatrienoyl)-cholesterol + 1-hexadecanoyl-sn-glycero-3-phosphocholine
Catalysed by CES1, LIPN, NCEH1, LDAH and 1 more
cholesteryl (11Z)-octadecenoateReach from here →
1. a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
2. (11Z)-octadecenoyl-CoA + cholesterol = cholesteryl (11Z)-octadecenoate + CoA
Catalysed by CES1, LIPN, NCEH1, LDAH and 1 more

Three steps

(S)-carnitineReach from here →
Catalysed by CEL, LIPAlysosome, endoplasmic reticulum, cytosolOne of 7 routes
CHEBI:1146900Reach from here →
Catalysed by CEL, LIPA

Page 2 of 5

, SOAT2

, SOAT1

lysosome, endoplasmic reticulum

One of 61 routes

(20R,22R)-20,22-dihydroxycholesterolReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
2 reduced [adrenodoxin] + cholesterol + O2 + 2 H(+) = (22R)-hydroxycholesterol + 2 oxidized [adrenodoxin] + H2O
(22R)-hydroxycholesterol + 2 reduced [adrenodoxin] + O2 + 2 H(+) = (20R,22R)-20,22-dihydroxycholesterol + 2 oxidized [adrenodoxin] + H2O

Catalysed by CES1, LIPN, NCEH1, LDAH and 1 more

CHEBI:12947Reach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
a beta-D-glucosyl-(1<->1')-N-acylsphing-4-enine + cholesterol = cholesteryl 3-beta-D-glucoside + an N-acylsphing-4-enine
UDP-alpha-D-galactose:N-acylsphingosine D-galactosyltransferase

Catalysed by CES1, LIPN, NCEH1, LDAH and 3 morecytosolOne of 5 routes

(22S)-22-hydroxycholesterolReach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
cytochrome P450 (cholesterol)

Catalysed by CEL, LIPA, CYP11A1, CYP1B1lysosome, mitochondrial matrix

(18S)-hydroxy-(5Z,8Z,11Z,14Z,16E)-eicosapentaenoateReach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
(5Z,8Z,11Z,14Z,17Z)-eicosapentaenoate + AH2 + O2 = (18S)-hydroxy-(5Z,8Z,11Z,14Z,16E)-eicosapentaenoate + A + H2O

Catalysed by CEL, LIPA, PTGS2lysosome

(15S)-hydroxy-(5Z,8Z,11Z,13E,17Z)-eicosapentaenoateReach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
(5Z,8Z,11Z,14Z,17Z)-eicosapentaenoate + AH2 + O2 = (15S)-hydroxy-(5Z,8Z,11Z,13E,17Z)-eicosapentaenoate + A + H2O

Catalysed by CEL, LIPA, PTGS2lysosome

pregnenolone sulfateReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
6 reduced [adrenodoxin] + cholesterol + 3 O2 + 6 H(+) = 4-methylpentanal + pregnenolone + 6 oxidized [adrenodoxin] + 4 H2O
3'-phosphoadenylyl-sulfate:pregnenolone sulfotransferase

Catalysed by CES1, LIPN, NCEH1, LDAH and 2 morecytosolOne of 4 routes

prostaglandin G1Reach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
(8Z,11Z,14Z)-eicosatrienoate + 2 O2 = prostaglandin G1

Catalysed by CEL, LIPA, PTGS2lysosome

prostaglandin G3Reach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
(5Z,8Z,11Z,14Z,17Z)-eicosapentaenoate + 2 O2 = prostaglandin G3

Catalysed by CEL, LIPA, PTGS2lysosome

N-(9Z-octadecenoyl)-L-phenylalanineReach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
L-phenylalanine + (9Z)-octadecenoate = N-(9Z-octadecenoyl)-L-phenylalanine + H2O

Catalysed by CEL, LIPA, PM20D1lysosome

N-(9Z-octadecenoyl)-L-leucineReach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
N-(9Z-octadecenoyl)-L-leucine + H2O = L-leucine + (9Z)-octadecenoate

Catalysed by CEL, LIPA, PM20D1lysosome

8,9-epoxy-(5Z,11Z,14Z,17Z)-eicosatetraenoateReach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
(5Z,8Z,11Z,14Z,17Z)-eicosapentaenoate + reduced [NADPH--hemoprotein reductase] + O2 = 8,9-epoxy-(5Z,11Z,14Z,17Z)-eicosatetraenoate + oxidized [NADPH--hemoprotein reductase] + H2O + H(+)

Catalysed by CEL, LIPA, CYP2C9lysosome

11,12-epoxy-(5Z,8Z,14Z,17Z)-eicosatetraenoateReach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
(5Z,8Z,11Z,14Z,17Z)-eicosapentaenoate + reduced [NADPH--hemoprotein reductase] + O2 = 11,12-epoxy-(5Z,8Z,14Z,17Z)-eicosatetraenoate + oxidized [NADPH--hemoprotein reductase] + H2O + H(+)

Catalysed by CEL, LIPA, CYP2C8, CYP2C9lysosome

14,15-epoxy-(5Z,8Z,11Z,17Z)-eicosatetraenoateReach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
(5Z,8Z,11Z,14Z,17Z)-eicosapentaenoate + reduced [NADPH--hemoprotein reductase] + O2 = 14,15-epoxy-(5Z,8Z,11Z,17Z)-eicosatetraenoate + oxidized [NADPH--hemoprotein reductase] + H2O + H(+)

Catalysed by CEL, LIPA, CYP2C8, CYP2C9lysosome

(24S)-hydroxycholesterol 24-sulfateReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
cholesterol + reduced [NADPH--hemoprotein reductase] + O2 = (24S)-hydroxycholesterol + oxidized [NADPH--hemoprotein reductase] + H2O + H(+)
(24S)-hydroxycholesterol + 3'-phosphoadenylyl sulfate = (24S)-hydroxycholesterol 24-sulfate + adenosine 3',5'-bisphosphate + H(+)

Catalysed by CES1, LIPN, NCEH1, LDAH and 2 more

(24S)-hydroxycholesterol 3-sulfateReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
cholesterol + reduced [NADPH--hemoprotein reductase] + O2 = (24S)-hydroxycholesterol + oxidized [NADPH--hemoprotein reductase] + H2O + H(+)
(24S)-hydroxycholesterol + 3'-phosphoadenylyl sulfate = (24S)-hydroxycholesterol 3-sulfate + adenosine 3',5'-bisphosphate + H(+)

Catalysed by CES1, LIPN, NCEH1, LDAH and 4 more

beta-D-glucosyl-N-(9Z-octadecenoyl)-sphing-4E-enineReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
a beta-D-glucosyl-(1<->1')-N-acylsphing-4-enine + cholesterol = cholesteryl 3-beta-D-glucoside + an N-acylsphing-4-enine
beta-D-glucosyl-N-(9Z-octadecenoyl)-sphing-4E-enine + cholesterol = N-(9Z-octadecenoyl)-sphing-4-enine + cholesteryl 3-beta-D-glucoside

Catalysed by CES1, LIPN, NCEH1, LDAH and 2 moreOne of 2 routes

an N-acyl-sphinga-4E,14Z-dienineReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
a beta-D-glucosyl-(1<->1')-N-acylsphing-4-enine + cholesterol = cholesteryl 3-beta-D-glucoside + an N-acylsphing-4-enine
an N-acylsphing-4-enine + 2 Fe(II)-[cytochrome b5] + O2 + 2 H(+) = an N-acyl-sphinga-4E,14Z-dienine + 2 Fe(III)-[cytochrome b5] + 2 H2O

Catalysed by CES1, LIPN, NCEH1, LDAH and 3 more

a 3-hydroxyacyl derivative of bacterial toxinReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
a 3-(acyloxy)acyl derivative of bacterial toxin + H2O = a 3-hydroxyacyl derivative of bacterial toxin + a fatty acid + H(+)
a 3-(acyloxy)acyl derivative of bacterial toxin + H2O = a 3-hydroxyacyl derivative of bacterial toxin + a fatty acid + H(+)

Catalysed by CES1, LIPN, NCEH1, LDAH and 1 more

sn-glycero-3-phosphoethanolamineReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
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
1-hexadecanoyl-sn-glycero-3-phosphoethanolamine + 1-hexadecanoyl-sn-glycero-3-phosphocholine = sn-glycero-3-phosphoethanolamine + 1,2-dihexadecanoyl-sn-glycero-3-phosphocholine

Catalysed by CES1, LIPN, NCEH1, LDAH and 2 more

a 2-acyl-6-alpha-D-glucosaminyl-1-(1,2-diacyl-sn-glycero-3-phospho)-1D-myo-inositolReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
a fatty acid + ATP + CoA = a fatty acyl-CoA + AMP + diphosphate
a 6-(alpha-D-glucosaminyl)-1-(1,2-diacyl-sn-glycero-3-phospho)-1D-myo-inositol + a fatty acyl-CoA = a 2-acyl-6-alpha-D-glucosaminyl-1-(1,2-diacyl-sn-glycero-3-phospho)-1D-myo-inositol + CoA

Catalysed by CES1, LIPN, NCEH1, LDAH and 2 more

an N-(fatty acyl)-glycineReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
a fatty acid + ATP + CoA = a fatty acyl-CoA + AMP + diphosphate
a fatty acyl-CoA + glycine = an N-(fatty acyl)-glycine + CoA + H(+)

Catalysed by CES1, LIPN, NCEH1, LDAH and 2 more

cholineReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
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
1-hexadecanoyl-sn-glycero-3-phosphocholine + H2O = 1-hexadecanoyl-sn-glycero-3-phosphate + choline + H(+)

Catalysed by CES1, LIPN, NCEH1, LDAH and 4 more

palmitoyl-CoAReach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
Palmitate:CoA ligase (AMP-forming)

Catalysed by CEL, LIPA, SLC27A2, ACSL4 and 6 morelysosome, cytosolOne of 3 routes

oleoyl-CoAReach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
long-chain-fatty-acid-CoA ligase (oleate)

Catalysed by CEL, LIPA, SLC27A2, ACSL4 and 6 morelysosome, cytosolOne of 3 routes

betaine aldehydeReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
cholesterol + AH2 + O2 = 25-hydroxycholesterol + A + H2O
choline + A = betaine aldehyde + AH2

Catalysed by CES1, LIPN, NCEH1, LDAH and 2 more

sulfateReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
cholesterol + 3'-phosphoadenylyl sulfate = cholesterol sulfate + adenosine 3',5'-bisphosphate + H(+)
Cholesterol-sulfate sulfohydrolase

Catalysed by CES1, LIPN, NCEH1, LDAH and 3 morecytosolOne of 4 routes

11-cis-retinolReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
an all-trans-retinyl ester + H2O = 11-cis-retinol + a fatty acid + H(+)
all-trans-retinyl ester acylhydrolase, 11-cis-retinol forming

Catalysed by CES1, LIPN, NCEH1, LDAH and 1 morecytosolOne of 2 routes

oxaloacetateReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
cholesterol + AH2 + O2 = 25-hydroxycholesterol + A + H2O
(R)-malate + A = oxaloacetate + AH2

Catalysed by CES1, LIPN, NCEH1, LDAH and 2 more

1-phosphatidyl-1D-myo-inositolReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
a 1,2-diacyl-sn-glycero-3-phospho-(1D-myo-inositol 4-phosphate)(out) + cholesterol(in) = a 1,2-diacyl-sn-glycero-3-phospho-(1D-myo-inositol 4-phosphate)(in) + cholesterol(out)
ATP:1-Phosphatidyl-1D-myo-inositol 4-phosphotransferase

Catalysed by CES1, LIPN, NCEH1, LDAH and 14 moreinner side of the membrane, outer side of the membrane, cytosolOne of 2 routes

formaldehydeReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
cholesterol + AH2 + O2 = 25-hydroxycholesterol + A + H2O
N(6),N(6)-dimethyl-L-lysyl(4)-[histone H3] + 2 A + 2 H2O = L-lysyl(4)-[histone H3] + 2 formaldehyde + 2 AH2

Catalysed by CES1, LIPN, NCEH1, LDAH and 3 moreOne of 3 routes

sn-glycerol 3-phosphocholineReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
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
1-hexadecanoyl-sn-glycero-3-phosphocholine + H2O = sn-glycerol 3-phosphocholine + hexadecanoate + H(+)

Catalysed by CES1, LIPN, NCEH1, LDAH and 18 moreOne of 6 routes

1-acyl-sn-glycerol 3-phosphateReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
a monoacylglycerol + H2O = glycerol + a fatty acid + H(+)
ATP:acylglycerol 3-phosphotransferase

Catalysed by CES1, LIPN, NCEH1, LDAH and 2 morecytosolOne of 6 routes

progesteroneReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
6 reduced [adrenodoxin] + cholesterol + 3 O2 + 6 H(+) = 4-methylpentanal + pregnenolone + 6 oxidized [adrenodoxin] + 4 H2O
Pregnenolone:NAD+ 3-oxidoreductase

Catalysed by CES1, LIPN, NCEH1, LDAH and 3 morecytosolOne of 2 routes

octadecanalReach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
octadecanal + NAD(+) + H2O = octadecanoate + NADH + 2 H(+)

Catalysed by CEL, LIPA, ALDH3A2lysosome

all-trans-retinolReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
cholesterol + AH2 + O2 = 25-hydroxycholesterol + A + H2O
all-trans-13,14-dihydroretinol + A = all-trans-retinol + AH2

Catalysed by CES1, LIPN, NCEH1, LDAH and 2 moreOne of 3 routes

AH2Reach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
cholesterol + AH2 + O2 = 25-hydroxycholesterol + A + H2O
A + NADH + H(+) = AH2 + NAD(+)

Catalysed by CES1, LIPN, NCEH1, LDAH and 3 moreOne of 19 routes

1-O-acyl-sn-glycero-3-phosphocholine(1+)Reach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
Phosphatidylcholine:sterol O-acyltransferase

Catalysed by CEL, LIPA, LCATlysosome, cytosolOne of 2 routes

L-cysteineReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
cholesterol + AH2 + O2 = 25-hydroxycholesterol + A + H2O
Mo-molybdopterin + L-cysteine + AH2 = thio-Mo-molybdopterin + L-alanine + A + H2O

Catalysed by CES1, LIPN, NCEH1, LDAH and 2 more

hexadecanalReach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
2 H+[c] + NADH[c] + palmitate[c] = H2O[c] + hexadecanal[c] + NAD+[c]

Catalysed by CEL, LIPA, ALDH3B1, ALDH3B2 and 1 morelysosome, cytosolOne of 2 routes

a sphingomyelinReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
a beta-D-glucosyl-(1<->1')-N-acylsphing-4-enine + cholesterol = cholesteryl 3-beta-D-glucoside + an N-acylsphing-4-enine
ceramide:phosphatidylcholine cholinephosphotransferase

Catalysed by CES1, LIPN, NCEH1, LDAH and 4 moreGolgi apparatusOne of 8 routes

desmosterolReach from here →

cholesterol-ester plasma pool to cholesterol-ester-10,13,16,19-docosa conversion
Steryl-ester acylhydrolase
3-hydroxyacyl-CoA dehydrogenase (desmosterol)

Catalysed by CEL, LIPAlysosome, nucleus

glycerolReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
a monoacylglycerol + H2O = glycerol + a fatty acid + H(+)
a monoacylglycerol + H2O = glycerol + a fatty acid + H(+)

Catalysed by CES1, LIPN, NCEH1, LDAH and 1 more

a 1,2-diacyl-sn-glycerolReach from here →

a cholesterol ester + H2O = cholesterol + a fatty acid + H(+)
a beta-D-glucosyl-(1<->1')-N-acylsphing-4-enine + cholesterol = cholesteryl 3-beta-D-glucoside + an N-acylsphing-4-enine
ceramide:phosphatidylcholine cholinephosphotransferase

Catalysed by CES1, LIPN, NCEH1, LDAH and 4 moreGolgi apparatusOne of 9 routes