Sheng J, Stevens J, Feng X (2016) Pathway compartmentalization in peroxisome of Saccharomyces
cerevisiae to produce versatile medium chain
fatty alcohols. Sci Rep 6:177
Shi S, Valle-Rodrı ´guez JO, Khoomrung S, Siewers V,
Nielsen J (2012) Functional expression and characterization of five wax ester synthases in Saccharomyces cerevisiae and their utility for biodiesel
production. Biotechnol Biofuels 5:7
Shi S, Valle-Rodrı ´guez JO, Siewers V, Nielsen J (2014a)
Engineering of chromosomal wax ester synthase
integrated Saccharomyces cerevisiae mutants for
improved biosynthesis of fatty acid ethyl esters.
Biotechnol Bioeng 111:1740–1747
Shi S, Chen Y, Siewers V, Nielsen J (2014b) Improving
production of malonyl coenzyme A-derived metabolites by abolishing Snf1-dependent regulation of
Acc1. MBio 5:321
Shi S, Ang EL, Zhao H (2018) In vivo biosensors:
mechanisms, development, and applications. J
Ind Microbiol Biotechnol 45:491–516
Shiba Y, Paradise EM, Kirby J, Ro D-K, Keasling JD
(2007) Engineering of the pyruvate dehydrogenase
bypass in Saccharomyces cerevisiae for high-level
production of isoprenoids. Metab Eng 9:160–168
Shin G-H, Veen M, Stahl U, Lang C (2012) Overexpression of genes of the fatty acid biosynthetic pathway
leads to accumulation of sterols in Saccharomyces
cerevisiae. Yeast 29:371–383
Skjoedt ML, Snoek T, Kildegaard KR, Arsovska D,
Eichenberger M, Goedecke TJ, Rajkumar AS,
Zhang J, Kristensen M, Lehka BJ, Siedler S, Borodina I, Jensen MK, Keasling JD (2016) Engineering
prokaryotic transcriptional activators as metabolite biosensors in yeast. Nat Chem Biol 12:951–958
Smidt O, Du Preez JC, Albertyn J (2012) Molecular and
physiological aspects of alcohol dehydrogenases in
the ethanol metabolism of Saccharomyces cerevisiae. FEMS Yeast Res 12:33–47
Stincone A, Prigione A, Cramer T, Wamelink MMC,
Campbell K, Cheung E, Olin-Sandoval V, Gru ¨ning
N-M, Kru ¨ger A, Tauqeer Alam M, Keller MA, Breitenbach M, Brindle KM, Rabinowitz JD, Ralser M
(2015) The return of metabolism: biochemistry
and physiology of the pentose phosphate pathway.
Biol Rev Camb Philos Soc 90:927–963
Sumper M, Oesterhelt D, Riepertinger C, Lynen F
(1969) Die Synthese verschiedener Carbonsa ¨uren
durch den Multienzymkomplex der Fettsa ¨uresynthese aus Hefe und die Erkla ¨rung ihrer Bildung. Eur J Biochem 10:377–387
Suomalainen H, Kera ¨nen AJA (1963) The effect of biotin deficiency on the synthesis of fatty acids by
yeast. Biochim Biophys Acta 70:493–503
Tang X, Chen WN (2015) Enhanced production of fatty
alcohols by engineering the TAGs synthesis pathway in Saccharomyces cerevisiae. Biotechnol
Bioeng 112:386–392
Tang X, Feng H, Chen WN (2013) Metabolic engineering for enhanced fatty acids synthesis in Saccharomyces cerevisiae. Metab Eng 16:95–102
Tang X, Feng L, Chen L, Chen WN (2017) Engineering
Saccharomyces cerevisiae for efficient biosynthesis
of fatty alcohols based on enhanced supply of free
fatty acids. ACS Omega 2:3284–3290
Tehlivets O, Scheuringer K, Kohlwein SD (2007) Fatty
acid synthesis and elongation in yeast. Biochim
Biophys Acta 1771:255–270
Teixeira PG, Ferreira R, Zhou YJ, Siewers V, Nielsen J
(2017) Dynamic regulation of fatty acid pools for
improved production of fatty alcohols in Saccharomyces cerevisiae. Microb Cell Factories 16:1185
Teo WS, Hee KS, Chang MW (2013) Bacterial FadR and
synthetic promoters function as modular fatty acid
sensor- regulators in Saccharomyces cerevisiae.
Eng Life Sci 13:456–463
Thompson RA, Trinh CT (2014) Enhancing fatty acid
ethyl ester production in Saccharomyces cerevisiae
through metabolic engineering and medium optimization. Biotechnol Bioeng 111:2200–2208
Toke DA, Martin CE (1996) Isolation and characterization of a gene affecting fatty acid elongation in
Saccharomyces cerevisiae. J Biol Chem 271:18413–
18422
Valle-Rodrı ´guez JO, Shi S, Siewers V, Nielsen J (2014)
Metabolic engineering of Saccharomyces cerevisiae
for production of fatty acid ethyl esters, an
advanced biofuel, by eliminating non-essential
fatty acid utilization pathways. Appl Energy
115:226–232
van Roermund CW, Tabak HF, van den Berg M, Wanders RJ, Hettema EH (2000) Pex11p plays a primary role in medium-chain fatty acid oxidation, a
process that affects peroxisome number and size
in Saccharomyces cerevisiae. J Cell Biol 150:489–
498
van Roermund CW, Drissen R, van den Berg M, Ijlst L,
Hettema EH, Tabak HF, Waterham HR, Wanders
RJ (2001) Identification of a peroxisomal ATP carrier required for medium-chain fatty acid b-oxidation and normal peroxisome proliferation in
Saccharomyces cerevisiae. Mol Cell Biol 21:4321–
4329
van Roermund CWT, Waterham HR, Ijlst L, Wanders
RJA (2003) Fatty acid metabolism in Saccharomyces cerevisiae. Cell Mol Life Sci 60:1838–1851
van Rossum HM, Kozak BU, Niemeijer MS, Duine HJ,
Luttik MAH, Boer VM, Ko ¨tter P, Daran J-MG, van
Maris AJA, Pronk JT (2016a) Alternative reactions
at the interface of glycolysis and citric acid cycle in
Saccharomyces cerevisiae. FEMS Yeast Res 16:
fow017
van Rossum HM, Kozak BU, Pronk JT, van Maris AJA
(2016b) Engineering cytosolic acetyl-coenzyme A
supply in Saccharomyces cerevisiae: pathway stoichiometry, free-energy conservation and redoxcofactor balancing. Metab Eng 36:99–115
366
L. Baumann et al.
cerevisiae to produce versatile medium chain
fatty alcohols. Sci Rep 6:177
Shi S, Valle-Rodrı ´guez JO, Khoomrung S, Siewers V,
Nielsen J (2012) Functional expression and characterization of five wax ester synthases in Saccharomyces cerevisiae and their utility for biodiesel
production. Biotechnol Biofuels 5:7
Shi S, Valle-Rodrı ´guez JO, Siewers V, Nielsen J (2014a)
Engineering of chromosomal wax ester synthase
integrated Saccharomyces cerevisiae mutants for
improved biosynthesis of fatty acid ethyl esters.
Biotechnol Bioeng 111:1740–1747
Shi S, Chen Y, Siewers V, Nielsen J (2014b) Improving
production of malonyl coenzyme A-derived metabolites by abolishing Snf1-dependent regulation of
Acc1. MBio 5:321
Shi S, Ang EL, Zhao H (2018) In vivo biosensors:
mechanisms, development, and applications. J
Ind Microbiol Biotechnol 45:491–516
Shiba Y, Paradise EM, Kirby J, Ro D-K, Keasling JD
(2007) Engineering of the pyruvate dehydrogenase
bypass in Saccharomyces cerevisiae for high-level
production of isoprenoids. Metab Eng 9:160–168
Shin G-H, Veen M, Stahl U, Lang C (2012) Overexpression of genes of the fatty acid biosynthetic pathway
leads to accumulation of sterols in Saccharomyces
cerevisiae. Yeast 29:371–383
Skjoedt ML, Snoek T, Kildegaard KR, Arsovska D,
Eichenberger M, Goedecke TJ, Rajkumar AS,
Zhang J, Kristensen M, Lehka BJ, Siedler S, Borodina I, Jensen MK, Keasling JD (2016) Engineering
prokaryotic transcriptional activators as metabolite biosensors in yeast. Nat Chem Biol 12:951–958
Smidt O, Du Preez JC, Albertyn J (2012) Molecular and
physiological aspects of alcohol dehydrogenases in
the ethanol metabolism of Saccharomyces cerevisiae. FEMS Yeast Res 12:33–47
Stincone A, Prigione A, Cramer T, Wamelink MMC,
Campbell K, Cheung E, Olin-Sandoval V, Gru ¨ning
N-M, Kru ¨ger A, Tauqeer Alam M, Keller MA, Breitenbach M, Brindle KM, Rabinowitz JD, Ralser M
(2015) The return of metabolism: biochemistry
and physiology of the pentose phosphate pathway.
Biol Rev Camb Philos Soc 90:927–963
Sumper M, Oesterhelt D, Riepertinger C, Lynen F
(1969) Die Synthese verschiedener Carbonsa ¨uren
durch den Multienzymkomplex der Fettsa ¨uresynthese aus Hefe und die Erkla ¨rung ihrer Bildung. Eur J Biochem 10:377–387
Suomalainen H, Kera ¨nen AJA (1963) The effect of biotin deficiency on the synthesis of fatty acids by
yeast. Biochim Biophys Acta 70:493–503
Tang X, Chen WN (2015) Enhanced production of fatty
alcohols by engineering the TAGs synthesis pathway in Saccharomyces cerevisiae. Biotechnol
Bioeng 112:386–392
Tang X, Feng H, Chen WN (2013) Metabolic engineering for enhanced fatty acids synthesis in Saccharomyces cerevisiae. Metab Eng 16:95–102
Tang X, Feng L, Chen L, Chen WN (2017) Engineering
Saccharomyces cerevisiae for efficient biosynthesis
of fatty alcohols based on enhanced supply of free
fatty acids. ACS Omega 2:3284–3290
Tehlivets O, Scheuringer K, Kohlwein SD (2007) Fatty
acid synthesis and elongation in yeast. Biochim
Biophys Acta 1771:255–270
Teixeira PG, Ferreira R, Zhou YJ, Siewers V, Nielsen J
(2017) Dynamic regulation of fatty acid pools for
improved production of fatty alcohols in Saccharomyces cerevisiae. Microb Cell Factories 16:1185
Teo WS, Hee KS, Chang MW (2013) Bacterial FadR and
synthetic promoters function as modular fatty acid
sensor- regulators in Saccharomyces cerevisiae.
Eng Life Sci 13:456–463
Thompson RA, Trinh CT (2014) Enhancing fatty acid
ethyl ester production in Saccharomyces cerevisiae
through metabolic engineering and medium optimization. Biotechnol Bioeng 111:2200–2208
Toke DA, Martin CE (1996) Isolation and characterization of a gene affecting fatty acid elongation in
Saccharomyces cerevisiae. J Biol Chem 271:18413–
18422
Valle-Rodrı ´guez JO, Shi S, Siewers V, Nielsen J (2014)
Metabolic engineering of Saccharomyces cerevisiae
for production of fatty acid ethyl esters, an
advanced biofuel, by eliminating non-essential
fatty acid utilization pathways. Appl Energy
115:226–232
van Roermund CW, Tabak HF, van den Berg M, Wanders RJ, Hettema EH (2000) Pex11p plays a primary role in medium-chain fatty acid oxidation, a
process that affects peroxisome number and size
in Saccharomyces cerevisiae. J Cell Biol 150:489–
498
van Roermund CW, Drissen R, van den Berg M, Ijlst L,
Hettema EH, Tabak HF, Waterham HR, Wanders
RJ (2001) Identification of a peroxisomal ATP carrier required for medium-chain fatty acid b-oxidation and normal peroxisome proliferation in
Saccharomyces cerevisiae. Mol Cell Biol 21:4321–
4329
van Roermund CWT, Waterham HR, Ijlst L, Wanders
RJA (2003) Fatty acid metabolism in Saccharomyces cerevisiae. Cell Mol Life Sci 60:1838–1851
van Rossum HM, Kozak BU, Niemeijer MS, Duine HJ,
Luttik MAH, Boer VM, Ko ¨tter P, Daran J-MG, van
Maris AJA, Pronk JT (2016a) Alternative reactions
at the interface of glycolysis and citric acid cycle in
Saccharomyces cerevisiae. FEMS Yeast Res 16:
fow017
van Rossum HM, Kozak BU, Pronk JT, van Maris AJA
(2016b) Engineering cytosolic acetyl-coenzyme A
supply in Saccharomyces cerevisiae: pathway stoichiometry, free-energy conservation and redoxcofactor balancing. Metab Eng 36:99–115
366
L. Baumann et al.