Table 1.

Terpenoid production using different organelle compartmentalization strategies

Organelle	Yeast species	Products	Titer or yield	Major engineering strategies	References
Endoplasmic reticulum	S. cerevisiae	β-amyrin	N.A.	Knock out PAH1	[33]
		Aedicagenic-28-O-glucoside	27.1 mg/L	Knock out PAH1	[33]
		Artemisinic acid	N.A.	Knock out PAH1	[33]
		Squalene	634 mg/L	Overexpression of INO2	[35]
		Ginsenoside	12.1 mg/L	Overexpression of INO2	[35]
Lipid droplet	Y. Lipolytica	Lycopene	16 mg/g	Strengthen the isoprenoid biosynthesis pathway and block the β-oxidation pathway	[47]
		Squalene	731.18 mg/L	Co-overexpression of tHMG1 and DGA1	[53]
	S. cerevisiae	Squalene	445.6 mg/L	Co-overexpression of tHMG1 and DGA1	[54]
		Lycopene	2.37 g/L (73.3 mg/g)	Strengthen the TAG pathway and modulate TAG fatty acyl composition	[49]
		Ginsenoside	5 g/L	Target protopanaxadiol synthase to LDs and strengthen the TAG pathway	[44]
		α-amyrin	1107.9 mg/L	Semi-rational design of MdOSC1, strengthen the MVA pathway and overexpress DGA1	[48]
Peroxisome	P. pastoris	Lycopene	73.9 mg/L	Target heterologous carotenogenic enzymes to peroxisomes	[74]
		α-humulene	3.2 g/L	Introduce the α-humulene synthesis pathway to peroxisomes	[77]
	S. cerevisiae	Squalene	11 g/L	Hybridization of the cytoplasm- and peroxisome-engineered strain	[63]
		Geraniol	2.75 mg/L	Deletion of PEX30 and PEX32 and introduce the geraniol synthesis pathway into peroxisomes	[72]
			5.5 g/L	Introduce a complete MVA pathway in peroxisomes	[75]
		(R)-(+)-limonene	2.6 g/L	Introduce a complete MVA pathway in peroxisomes	[75]
		Protopanaxadiol	N.A.	Knock out PEX11, PEX34, and ATG36	[73]
		α-humulene	1726.78 mg/L	Introduce the α-humulene biosynthesis pathway into peroxisomes and block the expression of ERG9	[76]
		β-Amyrin	2.6 g/L	Introduce the MVA pathway into peroxisomes	[78]
Mitochondrion	S. cerevisiae	Valencene	1.5 mg/L	Co-overexpression of tHMG1, mitochondrion-targeted heterologous FDP synthase and amorphadiene synthase	[84]
		Amorphadiene	20 mg/L	Co-overexpression of tHMG1, mitochondrion-targeted heterologous FDP synthase and amorphadiene synthase	[84]
		Amorpha-4,11-diene	427 mg/L	Introduce the amorpha-4,11-diene biosynthesis pathway to mitochondria	[86]
		Linalool	21 mg/L	Dual mevalonate pathways in mitochondria and cytoplasm	[88]
		Geraniol	43.3 mg/L	Introduce the geraniol biosynthetic pathway into mitochondria	[89]
		Patchoulol	19.24 mg/L	Introduce the DMAPP pathway into mitochondria	[90]
		Isoprene	2527 mg/L	Introduce the complete MVA pathway together with isoprene synthase (ISPS) into mitochondria	[91]
			11.9 g/L	Dual regulation of cytoplasmic and mitochondrial acetyl-CoA utilization	[92]
Plasma membrane	S. cerevisiae	β-Ionone	184 mg/L (32 mg/g)	Target the β-carotene cleavage dioxygenase to the membrane	[95]
	E. coli	Astaxanthin	N.A.	Target CrtW and CrtZ to the membrane via a GlpF protein	[94]
		Squalene	612 mg/L	Overexpression of TSR to expand membrane volume	[93]
		β-carotene	44.2 mg/g DCW	Overexpression of ALMGS and PLSB/PLSC to increase membrane surface area and enhance membrane synthesis	[24]