Table 1.
Overview of metabolic engineering strategies for terpenoid production in Y. lipolytica.
| Compound | Carbon Source | Parental Strain | Modifications related to terpenoid biosynthesis | Titer | Reference |
|---|---|---|---|---|---|
| Hemiterpenoids: | |||||
| Isoprene | glucose | Po1g | ↑HMG ↑ERG13 ↑IDI ↑PmISPS | 530.4 μg/L (sealed vials) | Shaikh and Odaneth (2021) |
| Monoterpenoids: | |||||
| Limonene | Glucose Pyruvic acid | Po1f | ↑TArLS ↑tSlNDPS1 ↑HMG ↑ERG12 | 23.6 mg/L (shakeflask) | Cao et al. (2016) |
| Limonene | Glucose | Po1g | ↑HMG (↑ClLS or ↑MsLS) | D-limonene: 11.7 mg/L or | Pang et al. (2019) |
| L-limonene: 11.1 mg/L, respectively (bioreactor) | |||||
| Limonene | Glycerol Citrate | Po1f | ↑TArLS ↑tSlNDPS1 ↑HMG ↑ERG12 | 165.3 mg/L (bioreactor) | Cheng et al. (2019) |
| Limonene | Glucose | ATCC 20460 | ↑HMG ↑ERG12 ↑ACL1 ↑SeACSL641P↑IDI ↑ERG20F88W–N119W↓SQS ↑PfLS | 35.9 mg/L (glass tube) | Arnesen et al. (2020) |
| Limonene | Lignocellulosic hydrolysate Citric acid | Po1f | ↑ssXR ↑ssXDH ↑XKS | 20.57 mg/L (shake flasks) | Yao et al. (2020) |
| ↑TArLS ↑SltNDPS1 ↑HMG ↑ERG12 | |||||
| Limonene | Waste cooking oil | Po1f | (↑ClLS or ↑MsLS) ↑HMG ↑IDI ↑tSlNDPS1 | 91.24 (D-limonene) and 83.06 (L-limonene) mg/L (shake flasks) | Li et al. (2022) |
| Linalool | Citrate Pyruvate | Po1f | ↑AaLIS ↑ERG20F88W–N119W↑HMG ↑IDI | 6.96 mg/L (shake flask) | Cao et al. (2017) |
| α-pinene | Waste cooking oil, soybean oil, or lignocellulosic hydrolysate medium | Po1f | ↑HMG ↑tSlNDPS1 ↑tPtPS ↑ERG8,12 ↑MBP-ERG12 ↑AMPD | 33.8, 36.6 or 36.1 mg/L for each carbon source, respectively (shake flasks) | Wei et al. (2021b) |
| Sesquiterpenoids: | |||||
| Abscisic acid | Glucose | ATCC 20460 | ↑HMG ↑ERG12 ↑ACL1 ↑SeACSL641P↑IDI ↑ERG20 ↓SQS ↑BcABA1-4 ↑BcCPR1 ↑POS5 ↑AtDTX50 | 263.5 mg/L (deepwell plate) | Arnesen et al. (2022) |
| Amorphadiene | Glucose | Po1g | ↑AaADS ↑HMG ↑ERG12 | 171.5 mg/L (shakeflask) | Marsafari and Xu (2020) |
| α-bisabolene | Waste cooking oil | Po1g | ↑HMG ↑GcABCG1 (↑AgαBS, ↑ZoβBS, or ↑HaγBS) | 973.1 mg/L, | Zhao et al. (2021) |
| β-bisabolene | 68.2 mg/L, or | ||||
| γ-bisabolene | 20.2 mg/L, respectively (culture tubes) | ||||
| (−)-α-bisabolol | Glucose | Po1f | ↑POT1 ↑MrBBS ↑tHMG ↑ERG20 ↓SQS | 364.23 mg/L (shake flasks) | (Yirong Ma et al., 2021) |
| α-farnesene | Glucose Fructose | Po1h | ↑tScHMG ↑IDI ↑MdFS-L-ERG20 | 260 mg/L (bioreactor) | Yang et al. (2016) |
| α-farnesene | Glucose | Po1f | ↑EcAtoB ↑BpHMG ↑ERG13 ↑MdFS-L-ERG20 ↑ERG12 ↑IDI ↑ERG8,19 ↑GPPS | 25.55 g/L (bioreactor) | (Y. Liu et al., 2019) |
| α-farnesene | Glucose Glycerol | Po1f | ↑FS-L-ERG20 ↑tScHMG1 ↑IDI ↑HMG ↑ERG19 | 2.57 g/L (bioreactor) | (S. C. Liu et al., 2020) |
| α-farnesene | Oleic acid | Po1f | ↑VHb ↑MdFS-L-ERG20 ↑ERG12 ↑IDI ↑ERG8,19 | 10.2 g/L (bioreactor) | (Y. Liu et al., 2021) |
| β-farnesene | Glucose | ATCC 20460 | ↑HMG ↑ERG12 ↑ACL1 ↑SeACSL641P↑IDI ↑ERG20 ↑AaBFS | 955 mg/L (glass tube) | Arnesen et al. (2020) |
| β-farnesene | Glucose | Po1f | ΔDGA1 ΔDGA2 ↑tHMG ↑BFS ↑ERG8,10,12, 13,19,20 IDI Δgut2 Δpox3,4,5,6 | 22.8 g/L (bioreactor) | (T. Shi et al., 2021) |
| α-humulene | Glucose | Po1f | ↑POT1 ↑AcACHS2PTS↑RpHMGPTS↑ANT1 ↑(ERG12,8,20,10,13,19)PTS↑IDIPTS | 3.2 g/L (bioreactor) | Guo et al. (2021) |
| α-santalene | Glucose | ATCC 201249 | ↑ClSTS ↑ERG8 ↑tHMG | 27.92 mg/L (bioreactor) | Jia et al. (2019) |
| Nootkatone | Glucose | ATCC 201249 | ↑CnVS ↑CnCYP706M1-tAtATR1 ↑tScHMG ↑ERG20 | Nootkatone: 978.2 μg/L | Guo et al. (2018) |
| Valencene | Valencene: 22.8 mg/L (shake flask) | ||||
| Valencene | Glucose | ATCC 20460 | ↑HMG ↑ERG12 ↑ACL1 ↑SeACSL641P↑IDI ↑ERG20 ↓SQS ↑CnVS | 113.9 mg/L (glass tube) | Arnesen et al. (2020) |
| Sterols: | |||||
| Campesterol | Sunflower seed oil | ATCC 201249 | Δerg5 ↑XlDHCR7 | 453 mg/L (bioreactor) | Du et al. (2016) |
| Campesterol | Sunflower seed oil | ATCC 201249 | Δerg5 ↑DrDHCR7 ↑POX2 | 942 mg/L (bioreactor) | Zhang et al. (2017) |
| Triterpenoids: | |||||
| Betulinic acid | Glycerol | ATCC 201249 | ↑tHMG1 ↑SQS ↑AtLUP1 ↑MtCYP716A12-tAtATR1 | 26.53 mg/L (shake flask) | Sun et al. (2019) |
| Betulinic acid | Glucose | ATCC 201249 | ↑RcLUS ↑BPLO ↑LjCPR ↑SQS ↑SQE ↑HMG ↑MFE1 | 204.89 mg/L total triterpenoid (shake flask) | Jin et al. (2019) |
| Ginsenoside K | Glucose | ATCC 201249 | ↑tHMG ↑ERG20 ↑SQS ↑PgDS ↑PgPPDS-L-tAtATR1 ↑PgUGT1 | 161.8 mg/L (bioreactor) | Li et al. (2019) |
| Lupeol | Glucose Pyruvic acid | ATCC 201249 | ↑RcLUS ↑HMG ↑SQS ↑SQE ↑OLE1 Δpah1 Δdgk1 | 411.72 (shake flasks) | (J.-L. Zhang et al., 2020) |
| Oleanolic acid | Glucose | ATCC 201249 | ↑tHMG ↑ERG20 ↑SQS ↑GgBAS ↑MtCYP716A12-L-tAtATR1 | 540.7 mg/L (bioreactor) | (D. Li et al., 2020) |
| 2,3-oxidosqualene | Glucose | ATCC 20460 | ↑HMG ↑ERG12 ↓ERG7 ↑ACL1 ↑SeACSL641P↑IDI ↑ERG20 ↑SQS ↑SQE | 22 mg/L (deepwell plate) | Arnesen et al. (2020) |
| Protopanaxadiol | Xylose | ATCC 201249 | ↑SsXR ↑SsXDH ↑XKS ↑PgDS ↑PgPPDS-L-AtATR1 ↑tHMG ↑ERG20 ↑SQS ↑TKL ↑TAL ↑TX Δpox1,2,3 | 300.63 mg/L (bioreactor) | Wu et al. (2019) |
| Squalene | Glucose Citrate | Po1f | ↑HMG ↑ACL1 ↑SeACSL641p | 10 mg/gDCW (shake flask) | Huang et al. (2018) |
| Squalene | Glucose | Po1f | ↑carB ↑carRP ↑ERG8,10,12,13,19,20 ↑tHMG ↑IDI Δgut2 Δpox3,4,5,6 | 531.6 mg/L | Gao et al. (2017b) |
| Squalene | Glucose | ATCC 20460 | ↑HMG ↑ERG12 ↓ERG7 ↑ACL1 ↑ SeACSL641P↑IDI ↑ERG20 ↑SQS | 402.4 mg/L (deepwell plate) | Arnesen et al. (2020) |
| Squalene | Glucose | Po1g | ↑SQS ↑HMG ↑MnDH2 | 502.7 mg/L (shake flask) | (H. Liu et al., 2020) |
| Squalene | Glucose | Po1f | Δpex10 Δure2 ↑HMG ↑DGA1 | 240.5 mg/L (shake flask) | Wei et al. (2021a) |
| Diterpenoids: | |||||
| Gibberellins | Glucose | GB20 | ↑tHMG ↑GGPPS ↓SQS ↑AtCPS ↑AtKS ↑AtKO ↑YlCyb5 ↑AtATR2 ↑SsGGPPS ↑GfCyb ↑GfCybRed ↑GfCPR ↑GfDES ↑GfP450-1 ↑GfP450-2 ↑GfP450-3 ↑tAtCPS ↑tAtKS ↑tAtKO | 12.81 mg/L GA3, 16.41 mg/L GA4, 0.79 mg/L GA7, and 4.70 mg/L GA9. | Kildegaard et al. (2021) |
| Tetraterpenoids: | |||||
| Astaxanthin | Glucose | GB20 | ↑XdcrtYB ↑XdcrtI ↑HMG ↓SQS ↑XdcrtE ↑PscrtW ↑PacrtZ | 54.6 mg/L (microtiterplate) | Kildegaard et al. (2017) |
| Astaxanthin | Glucose | GB20 | ↑XdcrtYB ↑XdcrtI ↑HMG ↓SQS ↑XdcrtE ↑SsGGPPS ↑HpBKT ↑HpCrtZ | 285 mg/L (bioreactor) | Tramontin et al. (2019) |
| Astaxanthin | Safflower oil | GB20 | ↑XdcrtYB ↑XdcrtI ↑HMG ↓SQS ↑XdcrtE ↑PscrtW ↑PacrtZ | 167 mg/L (bioreactor) | (N. Li et al., 2020) |
| Astaxanthin | Glucose | Not described | ↑PsCrtW-HpCrtZ-SKL ↑PsCrtW-HpCrtZ-oleosin ↑PsCrtW-HpCrtZ-KDEL ↑SaGGPPS ↑McCarRP ↑McCarB | 858 mg/L (shake flasks) | (Yongshuo Ma et al., 2021) |
| Cantaxanthin | glucose | Po1f | ↑BsCrtW ↑McCarB ↑McCarRP ↑GGPPS | 36.1 mg/L (shake flask) | Cui et al. (2021) |
| β-carotene | Glucose | Po1f | ↑McCarB ↑McCarRP ↑ERG8,10,12, 13,19,20 ↑GGPPS ↑tHMG ↑IDI Δpox3,4,5,6 | 4 g/L (bioreactor) | Gao et al. (2017b) |
| β-carotene | Glucose | Po1f | ↑GGPPS ↑SsCarS | 0.41 mg/gDCW | Gao et al. (2017a) |
| β-carotene | Glucose | ATCC 20460 | ↑McCarB ↑McCarRP ↑HMG ↑GGPPS ↑DGA2 ↑GPD1 Δpox1–6 Δtgl4 | 6.5 g/L (bioreactor) | Larroude et al. (2018) |
| β-carotene | Glucose | Po1g | ↑EcAtoB ↑ScERG13, ↑HMG ↑ERG8,12,19,20 ↑IDI ↑GGPPS ↑McCarB ↑McCarRP | 12.1 mg/gDCW | Cui et al. (2019) |
| β-carotene | Glucose | S11073 (in-house strain collection) | ↑McCarB ↑McCarRP | 75 mg/L (shake flask) | Bruder et al. (2020) |
| β-carotene | Glucose | ATCC 20460 | ↑HMG ↑ERG12 ↑ACL1 ↑ SeACSL641P↑IDI ↑GGPPS ↓SQS ↑XdcrtYB ↑XdcrtI | 164 mg/L (deepwell plate) | Arnesen et al. (2020) |
| β-carotene | Glucose | Po1f | ↑McCarB ↑McCarRP ↑GGPPS ↑HMG ↑ERG13 Δpox2,3 Δmfe | 4.5 g/L (bioreactor) | (X.-K. Zhang et al., 2020) |
| β-carotene | Glucose | Po1f | ↑tHMG ↑BtCarB ↑BtCarRA ↑GGPPS Δgut2 ↑ERG13 ↑Hxk | 2.4 g/L (bioreactor) | Qiang et al. (2020) |
| β-carotene | Glucose | Po1f | ↑tHMG ↑CarB ↑CarRP ↑GGPPS Δgut2 ↑DID2 | 2.01 g/L (bioreactor) | Lv et al. (2020) |
| β-carotene | Glucose | IMUFRJ 50682 | ↑McCarB ↑McCarRP ↑GGPPS | 50.1 mg/L (shake flask) | de Souza et al. (2020) |
| β-carotene | Glucose | Po1f | ↑tHMG ↑BtCarB ↑BtCarRA ↑GGPPS Δgut2 ↑DID2 | ∼2.6 g/L (bioreactor) | Yang et al. (2021) |
| β-carotene | Glucose | Po1f | ↑tHMG ↑GGPPS ↑BtCarRA ↑BtCarB | 1.7 g/L (bioreactor) | (L. Liu et al., 2021) |
| β-carotene | Glucose | Po1f | ↑McCarB ↑McCarRP Δndt80 | ∼12.5 mg/gDCW (96-well plate) | Liu et al. (2022) |
| β-carotene | Glucose | Po1f | ΔCLA4 ΔMHY1 ↑AfGGPS ↑IDI ↑ERG8,10,12,19,20 ↑VHb ↑MccarRP ↑GGPPS ↑McCarB ↑ScERG13 ↑HMG | 7.6 g/L (bioreactor) | (M. Liu et al., 2021) |
| β-carotene | Glucose | Po1f | ↑ScCK ↑AtIPK ↑ERG12 ↑tHMG ↑ERG20 ↑IDI ↑XdcrtE ↑McCarB ↑McCarRPY27Ror McCarRPE78K | 39.5 g/L β-carotene or 17.6 g/L lycopene, respectively (bioreactor) | Ma et al. (2022) |
| Lycopene | |||||
| Lycopene | Glucose Fructose | Po1f | ↑PaCrtE ↑PaCrtB ↑PaCrtI | 242 mg/L (bioreactor) | Nambou et al. (2015) |
| Lycopene | Glucose | H222 | ↑PaCrtB ↑PaCrtI ↑GGPPS ↑HMG Δpox1–6 Δgut2 | 16 mg/gDCW (bioreactor) | Matthäus et al. (2014) |
| Lycopene | Glucose | Po1f | ↑HMG ↑PaCrtE ↑PaCrtB ↑PaCrtI ↑ERG8,19 | 213 mg/L (bioreactor) | Schwartz et al. (2017) |
| Lycopene | Glucose | Po1f | ↑PaCrtE ↑PaCrtB ↑PaCrtI ↑AMPD | 745 mg/L (bioreactor) | Zhang et al. (2019) |
| Lycopene | Glucose Palmitic acid | Po1f | ↑PvIDI ↑LpCrtE ↑LpCrtB ↑LpCrtI ↑AtIPK ↑ScCHK ↑ERG20 | 4.2 g/L (bioreactor) | Luo et al. (2020) |
| Apocarotenoids: | |||||
| α-ionone | Glucose | Po1f | Proprietary information | 408 mg/L (bioreactor) | Czajka et al. (2020) |
| β-ionone | Glucose | Po1f | ↑McCarB ↑McCarRP ↑OfCCD1 ↑SsNphT7 ↑HpIDI ↑ERG8,10,12,13,19 ↑tHMG ↑GPPS ↑ERG20-GGPPS | 380 mg/L (bioreactor) | Czajka et al. (2018) |
| β-ionone | Glucose | Po1f | ↑McCarB ↑McCarRP ↑PhCCD1 ↑GGPPS ↑tHMG ↑ERG8,10,12,13,19,20 Δpox3,5 ↑IDI ↑bbPK ↑bsPTA | 0.98 g/L (bioreactor) | Lu et al. (2020) |
AaADS, Artimisia annua amorphadiene synthase. AaBFS, A. annua β-farnesene synthase. AaLIS, Actinidia arguta linalool synthase. AcACHS2, Aquilaria crassna α-humulene synthase. ACL1, ATP citrate lyase 1. AfGGPS, Archaeoglobus fulgidus geranylgeranyl diphosphate synthase. AgαBS, Abies grandis α-bisabolene synthase. AMPD, adenosine monophosphate deaminase. ANT1, peroxisomal adenine nucleotide transporter. AtATR1, Arabidopsis thaliana cytochrome P450 reductase 1. AtATR2, A. Thaliana cytochrome P450 reductase 2. AtCPS, A. Thaliana copalyl diphosphate synthase. AtDTX, A. thaliana abscisic acid exporter. AtIPK, A. thaliana isopentenyl phosphate kinase. AtKO, A. Thaliana ent-kaurene oxidase. AtKS, A. Thaliana ent-kaurene synthase. AtLUP1, A. thaliana lupeol synthase. BbPK, Bifidobacterium bifidum phosphoketolase. BcABA1, Botrytis cinerea cytochrome P450. BcABA2, B. cinerea cytochrome P450. BcABA3, B. cinerea α-ionylideneethane synthase. BcABA4, B. cinerea dehydrogenase. BcCPR1, B. cinerea cytochrome P450 reductase. BpHMG, Bordetella petrii 3-Hydroxy-3-methylglutaryl-CoA reductase. BpLO, Betula platyphylla lupeol C-28 oxidase. BsCrtW, Brevundimonas sp. β-carotene ketolase. BsPTA, Bacillus subtilis phosphotransacetylase. BtCarB, Blakeslea trispora phytoene dehydrogenase. BtCarRA, B. trispora phytoene synthase/lycopene cyclase. CarB, phytoene dehydrogenase from unknown species.CarRP, phytoene synthase/lycopene cyclase from unknown species.CLA4, protein kinase involved in hyphal development. ClLS, Citrus limon D-limonene synthase. ClSTS, Clausena lansium α-santalene synthase. CnCYP706M1-AtATR1, fusion of Callitropsis nootkatensis cytochrome P450 and AtATR1. CnVS, C. nootkatensis valencene synthase. Cyb5, Cytochrome b5. DGA1-2, diacylglycerol acyltransferase 1–2. DGK1, diacylglycerol kinase. DID2, endosomal sorting complex subunit. DrDHCR1, Danio rerio 7-dehydrocholesterol reductase. EcAtoB, Escherichia coli acetyl-CoA acetyltransferase. ERG10, acetyl-CoA acetyltransferase. ERG12, mevalonate kinase. ERG13, 3-hydroxy-3-methylglutaryl-CoA synthase. ERG19, mevalonate diphosphate decarboxylase. ERG20, farnesyl diphosphate synthase. ERG20F88W–N119W, geranyl diphosphate synthase. ERG20-GGPPS, fusion of ERG20p and geranylgeranyl diphosphate synthase. ERG5, C22-sterol desaturase. ERG7, lanosterol synthase. ERG8, phosphomevalonate kinase. FS-L-ERG20, α-farnesene synthase from unknown species fused to ERG20p. GcABCG1, Grosmania clavigera ABC-G1 efflux pump. GfCPR, Fusarium (Gibberella) fujikuroi cytochrome P450 reductase. GfCyb, F. fujikuroi cytochrome b5. GfCybRed, F. fujikuroi cytochrome b5 reductase. GfDES, F. fujikuroi desaturase. GfP450-1, F. fujikuroi cytochrome P450 1. GfP450-2, F. fujikuroi cytochrome P450 2. GfP450-3, F. fujikuroi cytochrome P450 3. GgBAS, Glycyrrhiza glabra β-amyrin synthase. GGPPS, geranylgeranyl diphosphate synthase. GPD1, glyceraldehyde-3-phosphate dehydrogenase. GPPS, geranyl diphosphate synthase. GUT2, glycerol-3-phosphate dehydrogenase 2. HaγBS, Helianthus annuus γ-bisabolene synthase. HMG, 3-Hydroxy-3-methylglutaryl-CoA reductase. HpBKT, Haematococcus pluvialis β-carotene ketolase. HpCrtZ, H. pluvialis β-carotene hydroxylase. HpIPI, H. pluvialis isopentyl diphosphate isomerase. Hxk, hexokinase. IDI, isopentyl diphosphate isomerase. LjCPR, Lotus japonicus cytochrome P450 reductase. LpCrtB, Lamprocystis purpurea phytoene synthase. LpCrtE, L. purpurea geranylgeranyl diphosphate syntase. LpCrtI, L. purpurea phytoene desaturase. MBP-ERG12, Maltose binding protein N-terminally fused to ERG12p. McCarB, Mucor circinelloides phytoene dehydrogenase. McCarRP, M. circinelloides phytoene synthase/lycopene cyclase. MdFS-L-ERG20, Malus domestica α-farnesene synthase linked to ERG20p. MFE1, multifunctional β-oxidation enzyme 1. MHY1, transcription factor involved in hyphal formation. MnDH2, Mannitol dehydrogenase. MrBBS, Matricaria recutita (−)-α-bisabolol synthase. MsLS, Mentha spicata L-limonene synthase. MtCYP716A12-L-tAtATR1, Medicago truncatula cytochrome P450 fused to truncated AtATR1p. MtCYP716A12-tAtATR1, M. truncatula cytochrome P450 directly linked to truncated AtATR1p. NDT80, transcription factor affecting lipid and ergosterol biosynthesis. OfCCD1, Osmanthus fragrans carotenoid cleavage dioxygenase 1. OLE1, Δ9-fatty acid desaturase. PaCrtB, Pantoea ananatis phytoene synthase. PaCrtE, P. ananatis geranylgeranyl diphosphate syntase. PaCrtI, P. ananatis phytoene desaturase. PacrtZ, P. ananatis β-carotene hydroxylase. PAH1, phosphatidic acid phosphatase. PfLS, Perilla frutescens limonene synthase. PgDS, Panax ginseng dammarenediol II synthase. PgPPDS, P. ginseng cytochrome P450 enzyme. PgPPDS-L-AtATR1, PgPPDSp linked to AtATR1p. PgPPDS-L-tAtATR1, PgPPDSp linked to truncated AtATR1p. PgUGT1, P. ginseng UDP-glycosyltransferase. PhCCD1, Petunia hybrida carotenoid cleavage dioxygenase. PmISPS, Pueraria montana isoprene synthase. POS5, Putative NAD + kinase. POT1, 3-ketoacyl-CoA thiolase. POX1-6, peroxisome acyl-CoA oxidase 1–6. PsCrtW, Paracoccus sp. β-carotene ketolase. PsCrtW-HpCrtZ-KDEL, fusion of PsCrtW-HpCrtZp with endoplasmic reticulum targeting sequence. PsCrtW-HpCrtZ-oleosin, fusion of PsCrtW-HpCrtZp with lipid body targeting signal. PsCrtW-HpCrtZ-SKL, fusion of PsCrtW-HpCrtZp with peroxisome targeting signal. PTS, C-terminal peroxisomal targeting signal. PvIDI, Pseudescherichia vulneris isopentyl diphosphate isomerase. RcLUS, Ricinus communis lupeol synthase. RpHMG1, Ruegeria Pomeroyi NADH-dependent HMGp. SaGGPPS, Sulfolobus acidocaldarius geranylgeranyl diphosphate synthase. ScCHK, Saccharomyces cerevisiae choline kinase. tScHMG, truncated S. cerevisiae HMGp. SeACSL641P, mutated Salmonella enterica acetyl-CoA synthetase. SQE, squalene epoxidase. SQS, squalene synthase. SsCarS, Multifunctional Schizochytrium sp. carotene synthase. SsGGPPS, Synechococcus sp. GGPPS. SsNphT7, Streptomyces sp. acetoacetyl CoA synthase. SsXDH, Scheffersomyces stipites xylose dehydrogenase. SsXR, S. stipites xylose reductase. TAL, transaldolase. tArLS, truncated Agastache rugosa limonene synthase. tAtCPS, truncated AtCPSp with plastidial targeting sequence removed. tAtKO, truncated AtKOp with plastidial targeting sequence removed. tAtKS, truncated AtKSp with plastidial targeting sequence removed. TGL3-4, triacylglycerol lipase 3–4. TKL, transketolase. tPtPS, truncated Pinus taeda α-pinene synthase without plastidial targeting sequence. tSlNDPS1, truncated Solanum lycopersicum neryl diphosphate synthase 1 without plastidial targeting sequence. TX, xylose transporter. URE2, transcriptional regulator involved in nitrogen catabolism repression. VHb, Vitreoscilla stercoraria hemoglobin. XdcrtE, Xanthophyllomyces dendrorhous geranylgeranyl diphosphate synthase. XdcrtI, X. dendrorhous phytoene desaturase. XdcrtYB, X. dendrorhous bi-functional phytoene synthase/lycopene cyclase. XKS, xylulose kinase. XlDHCR7, Xenapus laevis 7-dehydrocholesterol reductase. ZoβBS, Zingiber officinal β-bisabolene synthase. The table and accompanying text is expanded from (Arnesen et al., 2020).