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Cholesterol biosynthesis II (via 24,25-dihydrolanosterol)

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compartment	[Compartment:70101] cytosol
created	[InstanceEdit:1164949] Wu, Guanming, 2010-12-11
crossReference	[DatabaseIdentifier:1150129] RiceCyc:PWY66-3
dbId	1119286
displayName	Cholesterol biosynthesis II (via 24,25-dihydrolanosterol)
edited	[InstanceEdit:4719705] Amoss, Kindra, 2013-05-17 [InstanceEdit:4719712] Dharmawardhana, DP, 2013-07-31
eventOf	[Pathway:R-OSA-3906998] Fatty acid and lipid metabolism - Oryza sativa
goBiologicalProcess	[GO_BiologicalProcess:263550] cholesterol biosynthetic process via 24,25-dihydrolanosterol
hasDiagram	true
hasEvent	[Reaction:R-OSA-1119792] Farnesyl-diphosphate farnesyltransferase - Oryza sativa [Reaction:R-OSA-1120616] presqualene diphosphate+NAD(P)H->squalene+NAD(P)++diphosphate - Oryza sativa [Reaction:R-OSA-1120707] squalene+NADPH+O2->(S)-2,3-Epoxysqualene+NADP++H2O - Oryza sativa [Reaction:R-OSA-1121018] Lanosterol synthase - Oryza sativa [Reaction:R-OSA-1119897] lanosterol+NADPH->24,25-dihydrolanosterol+NADP+ - Oryza sativa [Reaction:R-OSA-1120103] 24,25-dihydrolanosterol+O2+NADPH->4,4-dimethyl-14alpha-hydroxymethyl-5alpha-cholesta-8-en-3beta-ol+H2O+NADP+ - Oryza sativa [Reaction:R-OSA-1121266] 4,4-dimethyl-14alpha-hydroxymethyl-5alpha-cholesta-8-en-3beta-ol+O2+NADPH->4,4-dimethyl-14alpha-formyl-5alpha-cholesta-8-en-3beta-ol+H2O+NADP+ - Oryza sativa [Reaction:R-OSA-1121462] 4,4-dimethyl-14alpha-formyl-5alpha-cholesta-8-en-3beta-ol+O2+NADPH->4,4-dimethyl-5-alpha-cholesta-8,14-dien-3-beta-ol+formate+H2O+NADP+ - Oryza sativa [Reaction:R-OSA-1120751] 4,4-dimethyl-5-alpha-cholesta-8,14-dien-3-beta-ol+NADPH->4,4-dimethyl-5alpha-cholesta-8-en-3-beta-ol+NADP+ - Oryza sativa [Reaction:R-OSA-1119717] 4,4-dimethyl-5alpha-cholesta-8-en-3-beta-ol+O2+NAD(P)H->4alpha-hydroxymethyl-4beta-methyl-5alpha-cholesta-8-en-3beta-ol+H2O+NAD(P)+ - Oryza sativa [Reaction:R-OSA-1119795] 4alpha-hydroxymethyl-4beta-methyl-5alpha-cholesta-8-en-3beta-ol+O2+NAD(P)H->4alpha-formyl-4beta-methyl-5alpha-cholesta-8-en-3beta-ol+H2O+NAD(P)+ - Oryza sativa [Reaction:R-OSA-1121434] 4alpha-formyl-4beta-methyl-5alpha-cholesta-8-en-3beta-ol+O2+NAD(P)H->4alpha-carboxy-4beta-methyl-5alpha-cholesta-8-en-3beta-ol+H2O+NAD(P)+ - Oryza sativa [Reaction:R-OSA-1120847] 4alpha-carboxy-4beta-methyl-5alpha-cholesta-8-en-3beta-ol+NAD(P)+->4alpha-methyl-5alpha-cholesta-8-en-3-one+NAD(P)H+CO2 - Oryza sativa [Reaction:R-OSA-1120331] 4alpha-methyl-5alpha-cholesta-8-en-3-one+NADPH->4alpha-methyl-cholesta-8-enol+NADP+ - Oryza sativa [Reaction:R-OSA-1119821] 4alpha-methyl-cholesta-8-enol+O2+NAD(P)H->4alpha-hydroxymethyl-5alpha-cholesta-8-en-3beta-ol+H2O+NAD(P)+ - Oryza sativa [Reaction:R-OSA-1120987] 4alpha-hydroxymethyl-5alpha-cholesta-8-en-3beta-ol+O2+NAD(P)H->4alpha-formyl-5alpha-cholesta-8-en-3beta-ol+H2O+NAD(P)+ - Oryza sativa [Reaction:R-OSA-1121645] 4alpha-formyl-5alpha-cholesta-8-en-3beta-ol+O2+NAD(P)H->4alpha-carboxy-5alpha-cholesta-8-en-3beta-ol+H2O+NAD(P)+ - Oryza sativa [Reaction:R-OSA-1119929] 4alpha-carboxy-5alpha-cholesta-8-en-3beta-ol+NAD(P)+->5alpha-cholesta-8-en-3-one+CO2+NAD(P)H - Oryza sativa [Reaction:R-OSA-1121395] 5alpha-cholesta-8-en-3-one+NADPH->zymostenol+NADP+ - Oryza sativa [Reaction:R-OSA-1120373] zymostenol->lathosterol - Oryza sativa [Reaction:R-OSA-1121497] lathosterol+NAD(P)H+O2->7-dehydro-cholesterol+NAD(P)++H2O - Oryza sativa [Reaction:R-OSA-1120741] 7-dehydro-cholesterol+NADPH->cholesterol+NADP+ - Oryza sativa
isInDisease	false
isInferred	false
literatureReference	[LiteratureReference:4719746] Biosynthesis of antinutritional alkaloids in solanaceous crops is mediated by clustered genes [LiteratureReference:4719709] Dual biosynthetic pathways to phytosterol via cycloartenol and lanosterol in Arabidopsis
modified	[InstanceEdit:11540225] Elser, Justin, 2025-03-27
name	Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) pathway100427
orthologousEvent	[Pathway:R-DEX-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Digitaria exilis [Pathway:R-QLO-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Quercus lobata [Pathway:R-ECG-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Echinochloa crus-galli [Pathway:R-AOF-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Asparagus officinalis [Pathway:R-CST-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Camelina sativa [Pathway:R-SMO-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Selaginella moellendorffii [Pathway:R-ATH-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Arabidopsis thaliana [Pathway:R-CSA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Cucumis sativus [Pathway:R-MTR-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Medicago truncatula [Pathway:R-TSP-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Triticum spelta subsp. spelta [Pathway:R-CME-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Cyanidioschyzon merolae [Pathway:R-SPC-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Synechocystis sp. PCC 6803 [Pathway:R-HVU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Hordeum vulgare [Pathway:R-AUM-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Aegilops umbellulata [Pathway:R-OPU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Oryza punctata [Pathway:R-ZMY-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Zea mays ver5 [Pathway:R-NAT-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Nicotiana attenuata [Pathway:R-CQI-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Chenopodium quinoa [Pathway:R-MDO-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Malus domestica [Pathway:R-OES-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Olea europaea var. sylvestris [Pathway:R-GRA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Gossypium raimondii [Pathway:R-PVE-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Pistacia vera [Pathway:R-TAE-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Triticum aestivum [Pathway:R-MAC-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Musa acuminata [Pathway:R-CCL-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Citrus clementina [Pathway:R-RCH-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Rosa chinensis [Pathway:R-PVU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Phaseolus vulgaris [Pathway:R-ADU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Arachis duranensis [Pathway:R-CRE-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Chlamydomonas reinhardtii [Pathway:R-NNU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Nelumbo nucifera [Pathway:R-LST-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Lathyrus sativus [Pathway:R-JCU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Jatropha curcas [Pathway:R-HAN-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Helianthus annuus [Pathway:R-CCA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Cajanus cajan [Pathway:R-OGR-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Oryza meyeriana var. granulata [Pathway:R-CSI-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Citrus sinensis [Pathway:R-VUN-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Vigna unguiculata [Pathway:R-PDA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Phoenix dactylifera [Pathway:R-HLP-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Humulus lupulus var. lupulus [Pathway:R-PTA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Pinus taeda [Pathway:R-AIP-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Arachis ipaensis [Pathway:R-AHA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Arabidopsis halleri [Pathway:R-LPE-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Leersia perrieri [Pathway:R-MPO-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Marchantia polymorpha [Pathway:R-TTU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Triticum turgidum [Pathway:R-CCD-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Cynara cardunculus var. scolymus [Pathway:R-ZMA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Zea mays [Pathway:R-FEX-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Fraxinus excelsior (European ash) [Pathway:R-MGU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Erythranthe guttata [Pathway:R-BVU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Beta vulgaris [Pathway:R-PTI-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Populus trichocarpa [Pathway:R-FCR-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Ficus carica [Pathway:R-CCR-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Chondrus crispus [Pathway:R-LAN-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Lupinus angustifolius [Pathway:R-BDI-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Brachypodium distachyon [Pathway:R-CAN-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Capsicum annuum [Pathway:R-OEU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Olea europaea [Pathway:R-OOF-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Oryza officinalis [Pathway:R-QSU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Quercus suber [Pathway:R-PED-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Phyllostachys edulis [Pathway:R-BOL-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Brassica oleracea [Pathway:R-CML-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Cucumis melo [Pathway:R-OLO-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Oryza longistaminata [Pathway:R-SVI-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Setaria viridis [Pathway:R-SCR-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Secale cereale [Pathway:R-DCA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Daucus carota [Pathway:R-ORU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Oryza rufipogon [Pathway:R-AHY-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Arachis hypogaea [Pathway:R-TUR-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Triticum urartu [Pathway:R-ACO-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Ananas comosus [Pathway:R-DRO-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Dioscorea rotundata [Pathway:R-CLA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Citrullus lanatus [Pathway:R-LSA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Lactuca sativa [Pathway:R-PPE-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Prunus persica [Pathway:R-SLY-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Solanum lycopersicum [Pathway:R-SHI-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Salvia hispanica [Pathway:R-OGL-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Oryza glaberrima [Pathway:R-COL-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Corchorus olitorius [Pathway:R-ZJA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Zoysia japonica [Pathway:R-OGU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Oryza glumaepatula [Pathway:R-CAR-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Cicer arietinum [Pathway:R-OBA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Oryza barthii [Pathway:R-ONI-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Oryza nivara [Pathway:R-GSO-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Glycine soja [Pathway:R-SBI-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Sorghum bicolor [Pathway:R-SIN-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Sesamum indicum [Pathway:R-OSI-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Oryza sativa Indica Group [Pathway:R-PAB-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Picea abies [Pathway:R-SSP-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Saccharum spontaneum [Pathway:R-TCA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Theobroma cacao [Pathway:R-ATR-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Amborella trichopoda [Pathway:R-CSK-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Cannabis sativa subsp. indica [Pathway:R-CCP-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Corchorus capsularis [Pathway:R-TDI-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Triticum dicoccoides [Pathway:R-FVE-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Fragaria vesca [Pathway:R-ETE-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Eragrostis tef [Pathway:R-BRA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Brassica rapa [Pathway:R-BNA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Brassica napus [Pathway:R-PAV-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Prunus avium [Pathway:R-BJU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Brassica juncea [Pathway:R-STU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Solanum tuberosum [Pathway:R-OME-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Oryza meridionalis [Pathway:R-MES-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Manihot esculenta [Pathway:R-CCN-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Coffea canephora [Pathway:R-TPR-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Trifolium pratense [Pathway:R-VRA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Vigna radiata [Pathway:R-OMI-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Oryza minuta [Pathway:R-GSU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Galdieria sulphuraria [Pathway:R-LPR-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Lolium perenne [Pathway:R-ITR-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Ipomoea triloba [Pathway:R-SIT-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Setaria italica [Pathway:R-GMA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Glycine max [Pathway:R-VAN-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Vigna angularis [Pathway:R-ALY-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Arabidopsis lyrata [Pathway:R-TTI-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Triticum timopheevii subsp. timopheevii [Pathway:R-VVN-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Vitis vinifera [Pathway:R-CRU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Capsella rubella [Pathway:R-PSA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Pisum sativum [Pathway:R-PHH-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Panicum hallii var. hallii ("HAL2") [Pathway:R-CBA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Chara braunii [Pathway:R-OLU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Ostreococcus lucimarinus [Pathway:R-PSO-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Papaver somniferum [Pathway:R-ECU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Eragrostis curvula [Pathway:R-ACH-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Actinidia chinensis [Pathway:R-PPA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Physcomitrella patens [Pathway:R-HLH-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Humulus lupulus [Pathway:R-OBR-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Oryza brachyantha [Pathway:R-NCO-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Nymphaea colorata [Pathway:R-JRE-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Juglans regia [Pathway:R-ASA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Avena sativa [Pathway:R-VFA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Vicia faba var. faba [Pathway:R-OAU-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Oryza australiensis [Pathway:R-EGR-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Eucalyptus grandis [Pathway:R-CSC-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Cannabis sativa [Pathway:R-ATA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Aegilops tauschii [Pathway:R-CCI-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Corymbia citriodora [Pathway:R-PHA-1119286] Cholesterol biosynthesis II (via 24,25-dihydrolanosterol) - Panicum hallii ("FIL2")
schemaClass	Pathway
species	[Species:186860] Oryza sativa
stId	R-OSA-1119286
summation	[Summation:4719731] cholesterol biosynthesis II (via 24,25-dihydrolanosterol) [Summation:2850146] Developed by Gramene.org

Referrals

(hasEvent)

[Pathway:R-OSA-3906998] Fatty acid and lipid metabolism - Oryza sativa