van Rossum HM, Kozak BU, Niemeijer MS, Dykstra JC,
Luttik MAH, Daran J-MG, van Maris AJA, Pronk
JT (2016c) Requirements for carnitine shuttlemediated translocation of mitochondrial acetyl
moieties to the yeast cytosol. MBio 7:517
Viegas C (1997) Effects of low temperatures (9–33
C)
and pH (3.3–5.7) in the loss of Saccharomyces
cerevisiae viability by combining lethal concentrations of ethanol with octanoic and decanoic acids.
Int J Food Microbiol 34:267–277
Viegas CA, Sa ´-Correia I (1995) Toxicity of octanoic acid
in Saccharomyces cerevisiae at temperatures
between 8.5 and 30
C. Enzym Microb Technol
17:826–831
Viegas CA, Rosa MF, Sa ´-Correia I, Novais JM (1989)
Inhibition of yeast growth by octanoic and
decanoic acids produced during ethanolic fermentation. Appl Environ Microbiol 55:21–28
Viegas CA, Almeida PF, Cavaco M, Sa ´-Correia I (1998)
The H(+)-ATPase in the plasma membrane of
Saccharomyces cerevisiae is activated during
growth latency in octanoic acid-supplemented
medium accompanying the decrease in intracellular pH and cell viability. Appl Environ Microbiol
64:779–783
Vorapreeda T, Thammarongtham C, Cheevadhanarak
S, Laoteng K (2012) Alternative routes of acetylCoA synthesis identified by comparative genomic
analysis: involvement in the lipid production of
oleaginous yeast and fungi. Microbiology
(Reading, England) 158:217–228
Waks Z, Silver PA (2009) Engineering a synthetic dualorganism system for hydrogen production. Appl
Environ Microbiol 75:1867–1875
Wang BL, Ghaderi A, Zhou H, Agresti J, Weitz DA, Fink
GR, Stephanopoulos G (2014a) Microfluidic highthroughput culturing of single cells for selection
based on extracellular metabolite production or
consumption. Nat Biotechnol 32:473–478
Wang Y, Chen H, Yu O (2014b) A plant malonyl-CoA
synthetase enhances lipid content and polyketide
yield in yeast cells. Appl Microbiol Biotechnol
98:5435–5447
Wang M, Li S, Zhao H (2016) Design and engineering of
intracellular-metabolite-sensing/regulation gene
circuits in Saccharomyces cerevisiae. Biotechnol
Bioeng 113:206–215
Wattanachaisaereekul S, Lantz AE, Nielsen ML, Nielsen
J (2008) Production of the polyketide 6-MSA in
yeast engineered for increased malonyl-CoA supply. Metab Eng 10:246–254
Welch JW, Burlingame AL (1973) Very long-chain fatty
acids in yeast. J Bacteriol 115:464–466
Wenning L, Yu T, David F, Nielsen J, Siewers V (2017)
Establishing very long-chain fatty alcohol and wax
ester biosynthesis in Saccharomyces cerevisiae.
Biotechnol Bioeng 114:1025–1035
Williams TC, Xu X, Ostrowski M, Pretorius IS, Paulsen
IT (2017) Positive-feedback, ratiometric biosensor
expression improves high-throughput metaboliteproducer screening efficiency in yeast. Synth Biol
2:1–13
Wu C-C, Tsai Y-Y, Ohashi T, Misaki R, Limtong S,
Fujiyama K (2018) Isolation of a thermotolerant
Rhodosporidium toruloides DMKU3-TK16 mutant
and its fatty acid profile at high temperature.
FEMS Microbiol Lett 365(21). https://doi.org/
10.1093/femsle/fny203
Xu P, Qiao K, Ahn WS, Stephanopoulos G (2016) Engineering Yarrowia lipolytica as a platform for synthesis of drop-in transportation fuels and
oleochemicals. Proc Natl Acad Sci U S A
113:10848–10853
Xue S-J, Chi Z, Zhang Y, Li Y-F, Liu G-L, Jiang H, Hu Z,
Chi Z-M (2018) Fatty acids from oleaginous yeasts
and yeast-like fungi and their potential applications. Crit Rev Biotechnol 38:1049–1060
Yang H, Bard M, Bruner DA, Gleeson A, Deckelbaum
RJ, Aljinovic G, Pohl TM, Rothstein R, Sturley SL
(1996) Sterol esterification in yeast: a two-gene
process. Science (New York, NY) 272:1353–1356
Yoshikawa K, Tanaka T, Furusawa C, Nagahisa K, Hirasawa T, Shimizu H (2009) Comprehensive phenotypic analysis for identification of genes affecting
growth under ethanol stress in Saccharomyces cerevisiae. FEMS Yeast Res 9:32–44
Yu A-Q, Pratomo Juwono NK, Leong SSJ, Chang MW
(2014) Production of fatty acid-derived valuable
chemicals in synthetic microbes. Front Bioeng
Biotechnol 2:1219
Yu A-Q, Pratomo Juwono NK, Foo JL, Leong SSJ, Chang
MW (2016) Metabolic engineering of Saccharomyces cerevisiae for the overproduction of short
branched-chain fatty acids. Metab Eng 34:36–43
Yu T, Zhou YJ, Wenning L, Liu Q, Krivoruchko A,
Siewers V, Nielsen J, David F (2017) Metabolic
engineering of Saccharomyces cerevisiae for production of very long chain fatty acid-derived chemicals. Nat Commun 8:15587
Yu T, Zhou YJ, Huang M, Liu Q, Pereira R, David F,
Nielsen J (2018) Reprogramming yeast metabolism from alcoholic fermentation to lipogenesis.
Cell 174:1549–1558.e14
Zhang F, Carothers JM, Keasling JD (2012) Design of a
dynamic sensor-regulator system for production
of chemicals and fuels derived from fatty acids.
Nat Biotechnol 30:354–359
Zhang Y, Nielsen J, Liu Z (2018) Metabolic engineering
of Saccharomyces cerevisiae for production of fatty
acid-derived hydrocarbons. Biotechnol Bioeng
110:87
Zhou YJ, Buijs NA, Zhu Z, Go ´mez DO, Boonsombuti A,
Siewers V, Nielsen J (2016a) Harnessing yeast peroxisomes for biosynthesis of fatty-acid-derived
biofuels and chemicals with relieved side-pathway
competition. J Am Chem Soc 138:15368–15377
Zhou YJ, Buijs NA, Zhu Z, Qin J, Siewers V, Nielsen J
(2016b) Production of fatty acid-derived oleo14 Engineering Saccharomyces cerevisiae for Production of Fatty Acids and Their Derivatives
367
Luttik MAH, Daran J-MG, van Maris AJA, Pronk
JT (2016c) Requirements for carnitine shuttlemediated translocation of mitochondrial acetyl
moieties to the yeast cytosol. MBio 7:517
Viegas C (1997) Effects of low temperatures (9–33
C)
and pH (3.3–5.7) in the loss of Saccharomyces
cerevisiae viability by combining lethal concentrations of ethanol with octanoic and decanoic acids.
Int J Food Microbiol 34:267–277
Viegas CA, Sa ´-Correia I (1995) Toxicity of octanoic acid
in Saccharomyces cerevisiae at temperatures
between 8.5 and 30
C. Enzym Microb Technol
17:826–831
Viegas CA, Rosa MF, Sa ´-Correia I, Novais JM (1989)
Inhibition of yeast growth by octanoic and
decanoic acids produced during ethanolic fermentation. Appl Environ Microbiol 55:21–28
Viegas CA, Almeida PF, Cavaco M, Sa ´-Correia I (1998)
The H(+)-ATPase in the plasma membrane of
Saccharomyces cerevisiae is activated during
growth latency in octanoic acid-supplemented
medium accompanying the decrease in intracellular pH and cell viability. Appl Environ Microbiol
64:779–783
Vorapreeda T, Thammarongtham C, Cheevadhanarak
S, Laoteng K (2012) Alternative routes of acetylCoA synthesis identified by comparative genomic
analysis: involvement in the lipid production of
oleaginous yeast and fungi. Microbiology
(Reading, England) 158:217–228
Waks Z, Silver PA (2009) Engineering a synthetic dualorganism system for hydrogen production. Appl
Environ Microbiol 75:1867–1875
Wang BL, Ghaderi A, Zhou H, Agresti J, Weitz DA, Fink
GR, Stephanopoulos G (2014a) Microfluidic highthroughput culturing of single cells for selection
based on extracellular metabolite production or
consumption. Nat Biotechnol 32:473–478
Wang Y, Chen H, Yu O (2014b) A plant malonyl-CoA
synthetase enhances lipid content and polyketide
yield in yeast cells. Appl Microbiol Biotechnol
98:5435–5447
Wang M, Li S, Zhao H (2016) Design and engineering of
intracellular-metabolite-sensing/regulation gene
circuits in Saccharomyces cerevisiae. Biotechnol
Bioeng 113:206–215
Wattanachaisaereekul S, Lantz AE, Nielsen ML, Nielsen
J (2008) Production of the polyketide 6-MSA in
yeast engineered for increased malonyl-CoA supply. Metab Eng 10:246–254
Welch JW, Burlingame AL (1973) Very long-chain fatty
acids in yeast. J Bacteriol 115:464–466
Wenning L, Yu T, David F, Nielsen J, Siewers V (2017)
Establishing very long-chain fatty alcohol and wax
ester biosynthesis in Saccharomyces cerevisiae.
Biotechnol Bioeng 114:1025–1035
Williams TC, Xu X, Ostrowski M, Pretorius IS, Paulsen
IT (2017) Positive-feedback, ratiometric biosensor
expression improves high-throughput metaboliteproducer screening efficiency in yeast. Synth Biol
2:1–13
Wu C-C, Tsai Y-Y, Ohashi T, Misaki R, Limtong S,
Fujiyama K (2018) Isolation of a thermotolerant
Rhodosporidium toruloides DMKU3-TK16 mutant
and its fatty acid profile at high temperature.
FEMS Microbiol Lett 365(21). https://doi.org/
10.1093/femsle/fny203
Xu P, Qiao K, Ahn WS, Stephanopoulos G (2016) Engineering Yarrowia lipolytica as a platform for synthesis of drop-in transportation fuels and
oleochemicals. Proc Natl Acad Sci U S A
113:10848–10853
Xue S-J, Chi Z, Zhang Y, Li Y-F, Liu G-L, Jiang H, Hu Z,
Chi Z-M (2018) Fatty acids from oleaginous yeasts
and yeast-like fungi and their potential applications. Crit Rev Biotechnol 38:1049–1060
Yang H, Bard M, Bruner DA, Gleeson A, Deckelbaum
RJ, Aljinovic G, Pohl TM, Rothstein R, Sturley SL
(1996) Sterol esterification in yeast: a two-gene
process. Science (New York, NY) 272:1353–1356
Yoshikawa K, Tanaka T, Furusawa C, Nagahisa K, Hirasawa T, Shimizu H (2009) Comprehensive phenotypic analysis for identification of genes affecting
growth under ethanol stress in Saccharomyces cerevisiae. FEMS Yeast Res 9:32–44
Yu A-Q, Pratomo Juwono NK, Leong SSJ, Chang MW
(2014) Production of fatty acid-derived valuable
chemicals in synthetic microbes. Front Bioeng
Biotechnol 2:1219
Yu A-Q, Pratomo Juwono NK, Foo JL, Leong SSJ, Chang
MW (2016) Metabolic engineering of Saccharomyces cerevisiae for the overproduction of short
branched-chain fatty acids. Metab Eng 34:36–43
Yu T, Zhou YJ, Wenning L, Liu Q, Krivoruchko A,
Siewers V, Nielsen J, David F (2017) Metabolic
engineering of Saccharomyces cerevisiae for production of very long chain fatty acid-derived chemicals. Nat Commun 8:15587
Yu T, Zhou YJ, Huang M, Liu Q, Pereira R, David F,
Nielsen J (2018) Reprogramming yeast metabolism from alcoholic fermentation to lipogenesis.
Cell 174:1549–1558.e14
Zhang F, Carothers JM, Keasling JD (2012) Design of a
dynamic sensor-regulator system for production
of chemicals and fuels derived from fatty acids.
Nat Biotechnol 30:354–359
Zhang Y, Nielsen J, Liu Z (2018) Metabolic engineering
of Saccharomyces cerevisiae for production of fatty
acid-derived hydrocarbons. Biotechnol Bioeng
110:87
Zhou YJ, Buijs NA, Zhu Z, Go ´mez DO, Boonsombuti A,
Siewers V, Nielsen J (2016a) Harnessing yeast peroxisomes for biosynthesis of fatty-acid-derived
biofuels and chemicals with relieved side-pathway
competition. J Am Chem Soc 138:15368–15377
Zhou YJ, Buijs NA, Zhu Z, Qin J, Siewers V, Nielsen J
(2016b) Production of fatty acid-derived oleo14 Engineering Saccharomyces cerevisiae for Production of Fatty Acids and Their Derivatives
367