Abstract
Carotenoids are widespread pigments in nature, obtained by direct synthesis or by ingestion in all taxonomic groups, and playing a large diversity of biological functions. Carotenoid biosynthesis is frequently found in fungi, and the amenability of some producing species to research studies has made them ideal models to investigate the genes and enzymes involved in the biosynthesis and its regulation. Best known examples are those for the production of β-carotene by the mucorales Phycomyces blakesleeanus, Mucor circinelloides, and Blakeslea trispora, neurosporaxanthin by the ascomycetes Neurospora crassa and Fusarium fujikuroi, and astaxanthin by the basidiomycete yeast Xanthophyllomyces dendrorhous, formerly Phaffia rhodozyma. Because of their coloring and health-promoting properties, some carotenoids have biotechnological applications, usually as food or feed additives. In the case of the fungi, the biotechnological studies have been mostly centered on the productions of β-carotene or its red precursor lycopene by B. trispora and astaxanthin by X. dendrorhous, extended to the heterologous expression of the relevant genes in non-carotenogenic yeasts as potentially favorable industrial producers. Less attention has been addressed to the synthesis of other carotenoids, with the only exception of torularhodin, produced by Rhodotorula and other related basidiomycete yeasts, but the genetics of its biosynthesis has not been investigated.
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References
Aasen AJ, Jensen SL (1965) Fungal carotenoids II. The structure of the carotenoid acid neurosporaxanthin. Acta Chem Scand 19:1843–1853
Acheampong EA, Martin AM (1995) Kinetic studies on the yeast Phaffia rhodozyma. J Basic Microbiol 35:147–155
Ádám AL, García-Martínez J, Szücs EP et al (2011) The MAT1-2-1 mating-type gene upregulates photo-inducible carotenoid biosynthesis in Fusarium verticillioides. FEMS Microbiol Lett 318:76–83
Ahrazem O, Gómez-Gómez L, Rodrigo M et al (2016) Carotenoid cleavage oxygenases from microbes and photosynthetic organisms: features and functions. Int J Mol Sci 17:1781
Aksu Z, Eren AT (2005) Carotenoids production by the yeast Rhodotorula mucilaginosa: use of agricultural wastes as a carbon source. Process Biochem 40:2985–2991
Aksu Z, Eren AT (2007) Production of carotenoids by the isolated yeast of Rhodotorula glutinis. Biochem Eng J 35:107–113
Alcaíno J, Barahona S, Carmona M et al (2008) Cloning of the cytochrome p450 reductase (crtR) gene and its involvement in the astaxanthin biosynthesis of Xanthophyllomyces dendrorhous. BMC Microbiol 8:169
Alcaíno J, Fuentealba M, Cabrera R et al (2012) Modeling the interfacial interactions between CrtS and CrtR from Xanthophyllomyces dendrorhous, a P450 system involved in astaxanthin production. J Agric Food Chem 60:8640–8647
Alcaino J, Baeza M, Cifuentes V (2014) Astaxanthin and related xanthophylls. In: Martín J-F, García-Estrada C, Zeilinger S (eds) Biosynthesis and molecular genetics of fungal secondary metabolites. Springer, New York, NY, pp 187–208
Alcaíno J, Romero I, Niklitschek M et al (2014) Functional characterization of the Xanthophyllomyces dendrorhous farnesyl pyrophosphate synthase and geranylgeranyl pyrophosphate synthase encoding genes that are involved in the synthesis of isoprenoid precursors. PLoS One 9:e96626
Alcaíno J, Bravo N, Córdova P et al (2016) The involvement of Mig1 from Xanthophyllomyces dendrorhous in catabolic repression: an active mechanism contributing to the regulation of carotenoid production. PLoS One 11:e0162838
Almeida ER, Cerdá-Olmedo E (2008) Gene expression in the regulation of carotene biosynthesis in Phycomyces. Curr Genet 53:129–137
Álvarez V, Rodríguez-Sáiz M, de la Fuente JL et al (2006) The crtS gene of Xanthophyllomyces dendrorhous encodes a novel cytochrome-P450 hydroxylase involved in the conversion of β-carotene into astaxanthin and other xanthophylls. Fungal Genet Biol 43:261–272
Amado IR, Vázquez JA (2015) Mussel processing wastewater: a low-cost substrate for the production of astaxanthin by Xanthophyllomyces dendrorhous. Microb Cell Factories 14:177
Ambati RR, Phang SM, Ravi S, Aswathanarayana RG (2014) Astaxanthin: sources, extraction, stability, biological activities and its commercial applications—a review. Mar Drugs 12:128–152
An GH (1997) Photosensitization of the yeast Phaffia rhodozyma at a low temperature for screening carotenoid hyperproducing mutants. Appl Biochem Biotechnol 66:263–268
An GH, Johnson EA (1990) Influence of light on growth and pigmentation of the yeast Phaffia rhodozyma. Antonie Van Leeuwenhoek 57:191–203
An GH, Schuman DB, Johnson EA (1989) Isolation of Phaffia rhodozyma mutants with increased astaxanthin content. Appl Environ Microbiol 55:116–124
An GH, Bielich J, Auerbach R, Johnson EA (1991) Isolation and characterization of carotenoid hyperproducing mutants of yeast by flow cytometry and cell sorting. Biotechnology (N Y) 9:70–73
An GH, Cho MH, Johnson EA (1999) Monocyclic carotenoid biosynthetic pathway in the yeast Phaffia rhodozyma (Xanthophyllomyces dendrorhous). J Biosci Bioeng 88:189–193
An GH, Jang BG, Cho MH (2001) Cultivation of the carotenoid-hyperproducing mutant 2A2N of the red yeast Xanthophyllomyces dendrorhous (Phaffia rhodozyma) with molasses. J Biosci Bioeng 92:121–125
Andrewes AG, Phaff HJ, Starr MP (1976) Carotenoids of Phaffia rhodozyma, a red-pigmented fermenting yeast. Phytochemistry 15:1003–1007
Aragón CM, Murillo FJ, de la Guardia MD, Cerdá-Olmedo E (1976) An enzyme complex for the dehydrogenation of phytoene in Phycomyces. Eur J Biochem FEBS 63:71–75
Araya-Garay JM, Ageitos JM, Vallejo JA et al (2012a) Construction of a novel Pichia pastoris strain for production of xanthophylls. AMB Express 2:24
Araya-Garay JM, Feijoo-Siota L, Rosa-dos-Santos F et al (2012b) Construction of new Pichia pastoris X-33 strains for production of lycopene and β-carotene. Appl Microbiol Biotechnol 93:2483–2492
Arpaia G, Carattoli A, Macino G (1995) Light and development regulate the expression of the albino-3 gene in Neurospora crassa. Dev Biol 170:626–635
Arpin N, Liaaen-Jensen S (1967) Chemotaxonomic research on fungi. Fungal carotenoids. IV. Carotenoids of Phillipsia carminea (Pat.) Le Gal; isolation and identification of a new natural xanthophyll. Bull Soc Chim Biol (Paris) 49:527–536
Arrach N, Fernández-Martín R, Cerdá-Olmedo E, Avalos J (2001) A single gene for lycopene cyclase, phytoene synthase, and regulation of carotene biosynthesis in Phycomyces. Proc Natl Acad Sci U S A 98:1687–1692
Arrach N, Schmidhauser TJ, Avalos J (2002) Mutants of the carotene cyclase domain of al-2 from Neurospora crassa. Mol Gen Genomics 266:914–921
Austin DG, Bu’Lock JD, Winstanley DJ (1969) Trisporic acid biosynthesis and carotenogenesis in Blakesleea trispora. Biochem J 113:34P
Austin DJ, Bu’Lock JD, Drake D (1970) The biosynthesis of trisporic acids from β-carotene via retinal and trisporol. Experientia 26:348–349
Avalos J, Cerdá-Olmedo E (1987) Carotenoid mutants of Gibberella fujikuroi. Curr Genet 25:1837–1841
Avalos J, Cerdá-Olmedo E (2004) Fungal carotenoid production. In: Arora DK (ed) Handbook of fungal biotechnology, 2nd edn. Marcel Dekker, New York, pp 367–378
Avalos J, Corrochano LM (2013) Carotenoid biosynthesis in Neurospora. In: Kasbekar DP, McCluskey K (eds) Neurospora: genomics and molecular biology. Caister Academic Press, Norfolk, pp 227–241
Avalos J, Estrada AF (2010) Regulation by light in Fusarium. Fungal Genet Biol 47:930–938
Avalos J, Schrott EL (1990) Photoinduction of carotenoid biosynthesis in Gibberella fujikuroi. FEMS Microbiol Lett 66:295–298
Avalos J, Mackenzie A, Nelki DS, Bramley PM (1988) Terpenoid biosynthesis in cell-extracts of wild type and mutant strains of Gibberella fujikuroi. Biochim Biophys Acta 966:257–265
Avalos J, Díaz-Sánchez V, García-Martínez J et al (2014a) Carotenoids. In: Martín JF, García-Estrada C, Zeilinger S (eds) Biosynthesis and molecular genetics of fungal secondary metabolites. Springer, New York, pp 149–185
Baima S, Macino G, Morelli G (1991) Photoregulation of the albino-3 gene in Neurospora crassa. J Photochem Photobiol B 11:107–115
Barbachano-Torres A, Castelblanco-Matiz LM, Ramos-Valdivia AC et al (2014) Analysis of proteomic changes in colored mutants of Xanthophyllomyces dendrorhous (Phaffia rhodozyma). Arch Microbiol 196:411–421
Barba-Ostria C, Lledias F, Georgellis D (2011) The Neurospora crassa DCC-1 protein, a putative histidine kinase, is required for normal sexual and asexual development and carotenogenesis. Eukaryot Cell 10:1733–1739
Barbato C, Calissano M, Pickford A et al (1996) Mild RIP-an alternative method for in vivo mutagenesis of the albino-3 gene in Neurospora crassa. Mol Gen Genet 252:353–361
Barredo J-L (ed) (2012) Microbial carotenoids from fungi: methods and protocols. Humana Press, New York
Barrero AF, Herrador MM, Arteaga P et al (2011) New apocarotenoids and β-carotene cleavage in Blakeslea trispora. Org Biomol Chem 9:7190–7195
Bartley GE, Schmidhauser TJ, Yanofsky C, Scolnik PA (1990) Carotenoid desaturases from Rhodobacter capsulatus and Neurospora crassa are structurally and functionally conserved and contain domains homologous to flavoprotein disulfide oxidoreductases. J Biol Chem 265:16020–16024
Beekwilder J, van Rossum HM, Koopman F et al (2014) Polycistronic expression of a β-carotene biosynthetic pathway in Saccharomyces cerevisiae coupled to β-ionone production. J Biotechnol 192(Pt B):383–392
Bejarano ER, Cerdá-Olmedo E (1989) Inhibition of phytoene dehydrogenation and activation of carotenogenesis in Phycomyces. Phytochemistry 28:1623–1626
Bejarano ER, Avalos J, Lipson ED, Cerdá-Olmedo E (1991) Photoinduced accumulation of carotene in Phycomyces. Planta 183:1–9
Bejarano ER, Cerdá-Olmedo E (1992) Independence of the carotene and sterol pathways of Phycomyces. FEBS Lett 306:209–212
Bejarano ER, Govind NS, Cerdá-Olmedo E (1987) ξ-carotene and other carotenes in a Phycomyces mutant. Phytochemistry 26:2251–2254
Bejarano ER, Parra F, Murillo FJ, Cerdá-Olmedo E (1988) End-product regulation of carotenogenesis in Phycomyces. Arch Microbiol 150:209–214
Bergman K, Eslava AP, Cerdá-Olmedo E (1973) Mutants of Phycomyces with abnormal phototropism. Mol Gen Genet 123:1–16
Bieszke JA, Braun EL, Bean LE et al (1999a) The nop-1 gene of Neurospora crassa encodes a seven transmembrane helix retinal-binding protein homologous to archaeal rhodopsins. Proc Natl Acad Sci U S A 96:8034–8039
Bieszke JA, Spudich EN, Scott KL et al (1999b) A eukaryotic protein, NOP-1, binds retinal to form an archaeal rhodopsin-like photochemically reactive pigment. Biochemistry (Mosc) 38:14138–14145
Bindl E, Lang W, Rau W (1970) Untersuchungen über die lichtabhängige Carotinoidsynthese. VI Zeitlicher Verlauf der Synthese der einzelnen Carotinoide bei Fusarium aquaeductuum unter verschiedenen Induktionsbedingungen. Planta 94:156–174
Blasco JL, Roeßink D, Iturriaga EA et al (2001) Photocarotenogenesis in phycomyces: expression of the carB gene encoding phytoene dehydrogenase. J Plant Res 114:25–31
Blomhoff R, Blomhoff HK (2006) Overview of retinoid metabolism and function. J Neurobiol 66:606–630
Böhme K, Richter C, Pätz R (2006) New insights into mechanisms of growth and β-carotene production in Blakeslea trispora. Biotechnol J 1:1080–1084
Bramley PM (1985) The in vitro biosynthesis of carotenoids. Adv Lipid Res 21:243–279
Brehm-Stecher BF, Johnson EA (2012) Isolation of carotenoid hyperproducing mutants of Xanthophyllomyces dendrorhous (Phaffia rhodozyma) by flow cytometry and cell sorting. Methods Mol Biol 898:207–217
Breitenbach J, Fraser PD, Sandmann G (2012) Carotenoid synthesis and phytoene synthase activity during mating of Blakeslea trispora. Phytochemistry 76:40–45
Breitenbach J, Visser H, Verdoes JC et al (2011) Engineering of geranylgeranyl pyrophosphate synthase levels and physiological conditions for enhanced carotenoid and astaxanthin synthesis in Xanthophyllomyces dendrorhous. Biotechnol Lett 33:755–761
Britton G, Liaaen-Jensen S, Pfander H (1998) Carotenoids. Birkhäuser Verlag, Basel
Britton G, Liaaen-Jensen S, Pfander H (2004) Carotenoids: handbook. Birkhauser, Boston
Burgeff H (1924) Untersuchungen über Sexualität und Parasitismus bei Mucorineen. Gustav Fischer, Jena
Burmester A, Richter M, Schultze K et al (2007) Cleavage of β-carotene as the first step in sexual hormone synthesis in zygomycetes is mediated by a trisporic acid regulated β-carotene oxygenase. Fungal Genet Biol 44:1096–1108
Buzzini P, Innocenti M, Turchetti B et al (2007) Carotenoid profiles of yeasts belonging to the genera Rhodotorula, Rhodosporidium, Sporobolomyces, and Sporidiobolus. Can J Microbiol 53:1024–1031
Buzzini P, Martini A (1999) Production of carotenoids by strains of Rhodotorula glutinis cultured in raw materials of agro-industrial origin. Bioresour Technol 71:41–44
Calo P, de Miguel T, Velázquez JB, Villa TG (1995) Mevalonic acid increases trans-astaxanthin and carotenoid biosynthesis in Phaffia rhodozyma. Biotechnol Lett 17:575–578
Candau R, Avalos J, Cerdá-Olmedo E (1991a) Gibberellins and carotenoids in the wild type and mutants of Gibberella fujikuroi. Appl Environ Microbiol 57:3378–3382
Candau R, Bejarano ER, Cerdá-Olmedo E (1991b) In vivo channeling of substrates in an enzyme aggregate for β-carotene biosynthesis. Proc Natl Acad Sci U S A 88:4936–4940
Carattoli A, Cogoni C, Morelli G, Macino G (1994) Molecular characterization of upstream regulatory sequences controlling the photoinduced expression of the albino-3 gene of Neurospora crassa. Mol Microbiol 13:787–795
Carattoli A, Romano N, Ballario P et al (1991) The Neurospora crassa carotenoid biosynthetic gene (albino 3) reveals highly conserved regions among prenyltransferases. J Biol Chem 266:5854–5859
Cardoso LA, Jäckel S, Karp SG et al (2016) Improvement of Sporobolomyces ruberrimus carotenoids production by the use of raw glycerol. Bioresour Technol 200:374–379
Castelblanco-Matiz LM, Barbachano-Torres A, Ponce-Noyola T et al (2015) Carotenoid production and gene expression in an astaxanthin-overproducing Xanthophyllomyces dendrorhous mutant strain. Arch Microbiol 197:1129–1139
Castrillo M, Avalos J (2015) The flavoproteins CryD and VvdA cooperate with the white collar protein WcoA in the control of photocarotenogenesis in Fusarium fujikuroi. PLoS One 10:e0119785
Castrillo M, Bernhardt A, Avalos J et al (2015) Biochemical characterization of the DASH-type cryptochrome CryD from Fusarium fujikuroi. Photochem Photobiol 91:1356–1367
Castrillo M, García-Martínez J, Avalos J (2013) Light-dependent functions of the Fusarium fujikuroi CryD DASH cryptochrome in development and secondary metabolism. Appl Environ Microbiol 79:2777–2788
Cederberg E, Neujahr HY (1969) Activation of β-carotene synthesis in Blakeslea trispora by certain terpenes. Acta Chem Scand 23:957–961
Cerdá-Olmedo E (1987) Carotene. In: Cerdá-Olmedo E, Lipson ED (eds) Phycomyces. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, pp 199–222
Cerdá-Olmedo E (1985) Carotene mutants of Phycomyces. Methods Enzymol 110:220–243
Cerdá-Olmedo E (1989) Production of carotenoids with fungi. In: Vandamme E (ed) Biotechnology of vitamin, growth factor and pigment production. Elsevier Applied Science, London, pp 27–42
Cerdá-Olmedo E, Hüttermann A (1986) Förderung und Hemmung der Carotinsynthese bei Phycomyces durch Aromaten. Angew Bot 60:59–70
Cerdá-Olmedo E, Mehta BJ (2012) Isolation of mutants and construction of intersexual heterokaryons of Blakeslea trispora. Methods Mol Biol 898:75–84
Chang J-J, Thia C, Lin H-Y et al (2015) Integrating an algal β-carotene hydroxylase gene into a designed carotenoid-biosynthesis pathway increases carotenoid production in yeast. Bioresour Technol 184:2–8
Chen CH, Dunlap JC, Loros JJ (2010) Neurospora illuminates fungal photoreception. Fungal Genet Biol 47:922–929
Chi S, He Y, Ren J et al (2015) Overexpression of a bifunctional enzyme, CrtS, enhances astaxanthin synthesis through two pathways in Phaffia rhodozyma. Microb Cell Factories 14:90
Choudhari S, Singhal R (2008) Media optimization for the production of β-carotene by Blakeslea trispora: a statistical approach. Bioresour Technol 99:722–730
Choudhari SM, Ananthanarayan L, Singhal RS (2008) Use of metabolic stimulators and inhibitors for enhanced production of β-carotene and lycopene by Blakeslea trispora NRRL 2895 and 2896. Bioresour Technol 99:3166–3173
Chumpolkulwong N, Kakizono T, Nagai S, Nishio N (1997) Increased astaxanthin production by Phaffia rhodozyma mutants isolated as resistant to diphenylamine. J Ferment Bioeng 83:429–434
Corrochano LM, Cerdá-Olmedo E (1992) Sex, light and carotenes: the development of Phycomyces. Trends Genet TIG 8:268–274
Corrochano LM, Cerdá-Olmedo E (1990) Photomorphogenesis in behavioural and colour mutants of Phycomyces. J Photochem Photobiol B 6:325–335
Corrochano LM, Garre V (2010) Photobiology in the Zygomycota: multiple photoreceptor genes for complex responses to light. Fungal Genet Biol 47:893–899
Csernetics A, Nagy G, Iturriaga EA et al (2011) Expression of three isoprenoid biosynthesis genes and their effects on the carotenoid production of the zygomycete Mucor circinelloides. Fungal Genet Biol 48:696–703
Csernetics Á, Tóth E, Farkas A et al (2015) Expression of Xanthophyllomyces dendrorhous cytochrome-P450 hydroxylase and reductase in Mucor circinelloides. World J Microbiol Biotechnol 31:321–336
Daub ME, Payne GA (1989) The role of carotenoids in resistance of fungi to cercosporin. Phytopathology 79:180–185
de Fabo EC, Harding RW, Shropshire W Jr (1976) Action spectrum between 260 and 800 nanometers for the photoinduction of carotenoid biosynthesis in Neurospora crassa. Plant Physiol 57:440–445
de la Fuente JL, Rodríguez-Sáiz M, Schleissner C et al (2010) High-titer production of astaxanthin by the semi-industrial fermentation of Xanthophyllomyces dendrorhous. J Biotechnol 148:144–146
De la Guardia MD, Aragón CM, Murillo FJ, Cerdá-Olmedo E (1971) A carotenogenic enzyme aggregate in Phycomyces: evidence from quantitive complementation. Proc Natl Acad Sci U S A 68:2012–2015
de Miguel T, Calo P, Díaz A, Villa TG (1997) The genus Rhodosporidium: a potential source of β-carotene. Microbiologia 13:67–70
DeBose-Boyd RA (2008) Feedback regulation of cholesterol synthesis: sterol-accelerated ubiquitination and degradation of HMG CoA reductase. Cell Res 18:609–621
Degli-Innocenti F, Russo VE (1984) Isolation of new white collar mutants of Neurospora crassa and studies on their behavior in the blue light-induced formation of protoperithecia. J Bacteriol 159:757–761
Díaz-Sánchez V, Estrada AF, Limón MC et al (2013) The oxygenase CAO-1 of Neurospora crassa is a resveratrol cleavage enzyme. Eukaryot Cell 12:1305–1314
Díaz-Sánchez V, Estrada AF, Trautmann D et al (2011a) Analysis of al-2 mutations in Neurospora. PLoS One 6:e21948
Díaz-Sánchez V, Estrada AF, Trautmann D et al (2011b) The gene carD encodes the aldehyde dehydrogenase responsible for neurosporaxanthin biosynthesis in Fusarium fujikuroi. FEBS J 278:3164–3176
Dimitrova S, Pavlova K, Lukanov L et al (2013) Production of metabolites with antioxidant and emulsifying properties by antarctic strain Sporobolomyces salmonicolor AL1. Appl Biochem Biotechnol 169:301–311
Domenech CE, Giordano W, Avalos J, Cerdá-Olmedo E (1996) Separate compartments for the production of sterols, carotenoids and gibberellins in Gibberella fujikuroi. Eur J Biochem 239:720–725
Domínguez-Bocanegra AR, Torres-Muñoz JA (2004) Astaxanthin hyperproduction by Phaffia rhodozyma (now Xanthophyllomyces dendrorhous) with raw coconut milk as sole source of energy. Appl Microbiol Biotechnol 66:249–252
Domonkos I, Kis M, Gombos Z, Ughy B (2013) Carotenoids, versatile components of oxygenic photosynthesis. Prog Lipid Res 52:539–561
Ducrey Sanpietro LM, Kula MR (1998) Studies of astaxanthin biosynthesis in Xanthophyllomyces dendrorhous (Phaffia rhodozyma). Effect of inhibitors and low temperature. Yeast Chichester Engl 14:1007–1016
Echavarri-Erasun C, Johnson EA (2004) Stimulation of astaxanthin formation in the yeast Xanthophyllomyces dendrorhous by the fungus Epicoccum nigrum. FEMS Yeast Res 4:511–519
Edgar RC (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Res 32:1792–1797
Ehrenshaft M, Daub ME (1994) Isolation, sequence, and characterization of the Cercospora nicotianae phytoene dehydrogenase gene. Appl Environ Microbiol 60:2766–2771
Elahi M, Chichester CO, Simpson KL (1973a) Biosynthesis of carotenoids by Phycomyces blakesleeanus mutants in the presence of nitrogenous heterocyclic compounds. Phytochemistry 12:1627–1632
Elahi M, Lee TH, Simpson KL, Chichester CO (1973b) Effect of CPTA and cycocel on the biosynthesis of carotenoids by Phycomyces blakesleeanus mutants. Phytochemistry 12:1633–1639
El-Jack M, Mackenzie A, Bramley PM (1988) The photoregulation of carotenoid biosynthesis in Aspergillus giganteus mut. alba. Planta 174:59–66
Eslava AP, Alvarez MI, Cerdá-Olmedo E (1974) Regulation of carotene biosynthesis in Phycomyces by vitamin A and β-ionone. Eur J Biochem 48:617–623
Estrada AF, Avalos J (2009) Regulation and targeted mutation of opsA, coding for the NOP-1 opsin orthologue in Fusarium fujikuroi. J Mol Biol 387:59–73
Estrada AF, Avalos J (2008) The White Collar protein WcoA of Fusarium fujikuroi is not essential for photocarotenogenesis, but is involved in the regulation of secondary metabolism and conidiation. Fungal Genet Biol 45:705–718
Estrada AF, Brefort T, Mengel C et al (2010) Ustilago maydis accumulates β-carotene at levels determined by a retinal-forming carotenoid oxygenase. Fungal Genet Biol 46:803–813
Estrada AF, Maier D, Scherzinger D et al (2008a) Novel apocarotenoid intermediates in Neurospora crassa mutants imply a new biosynthetic reaction sequence leading to neurosporaxanthin formation. Fungal Genet Biol 45:1497–1505
Estrada AF, Youssar L, Scherzinger D et al (2008b) The ylo-1 gene encodes an aldehyde dehydrogenase responsible for the last reaction in the Neurospora carotenoid pathway. Mol Microbiol 69:1207–1220
Fang TJ, Cheng YS (1992) Isolation of astaxanthin over-producing mutants of Phaffia rhodozyma and their fermentation kinetics. Chin J Microbiol Inmunol 25:209–222
Fernández-Martín R, Cerdá-Olmedo E, Avalos J (2000) Homologous recombination and allele replacement in transformants of Fusarium fujikuroi. Mol Gen Genet 263:838–845
Flores R, Cerdá-Olmedo E, Corrochano LM (1998) Separate sensory pathways for photomorphogenesis in Phycomyces. Photochem Photobiol 67:467–472
Flores-Cotera LB, Martín R, Sánchez S (2001) Citrate, a possible precursor of astaxanthin in Phaffia rhodozyma: influence of varying levels of ammonium, phosphate and citrate in a chemically defined medium. Appl Microbiol Biotechnol 55:341–347
Fontana JD, Guimarães MF, Martins NT et al (1996) Culture of the astaxanthinogenic yeast Phaffia rhodozyma in low-cost media. Appl Biochem Biotechnol 57–58:413–422
Fraser PD, Bramley PM (1994) The purification of phytoene dehydrogenase from Phycomyces blakesleeanus. Biochim Biophys Acta 1212:59–66
Fraser PD, Miura Y, Misawa N (1997) In vitro characterization of astaxanthin biosynthetic enzymes. J Biol Chem 272:6128–6135
Fraser PD, Ruiz-Hidalgo MJ, López-Matas MA et al (1996) Carotenoid biosynthesis in wild type and mutant strains of Mucor circinelloides. Biochim Biophys Acta 1289:203–208
Freitas C, Parreira TM, Roseiro J et al (2014) Selecting low-cost carbon sources for carotenoid and lipid production by the pink yeast Rhodosporidium toruloides> NCYC 921 using flow cytometry. Bioresour Technol 158:355–359
Frengova G, Simova E, Beshkova D (2004a) Use of whey ultrafiltrate as a substrate for production of carotenoids by the yeast Rhodotorula rubra. Appl Biochem Biotechnol 112:133–141
Frengova G, Simova E, Pavlova K et al (1994) Formation of carotenoids by Rhodotorula glutinis in whey ultrafiltrate. Biotechnol Bioeng 44:888–894
Frengova GI, Beshkova DM (2009) Carotenoids from Rhodotorula and Phaffia: yeasts of biotechnological importance. J Ind Microbiol Biotechnol 36:163–180
Frengova GI, Simova ED, Beshkova DM (2004b) Improvement of carotenoid-synthesizing yeast Rhodotorula rubra by chemical mutagenesis. Z Naturforschung C J Biosci 59:99–103
Garbayo I, Vílchez C, Nava-Saucedo JE, Barbotin JN (2003) Nitrogen, carbon and light-mediated regulation studies of carotenoid biosynthesis in immobilized mycelia of Gibberella fujikuroi. Enzym Microb Technol 33:629–634
García-Martínez J, Ádám AL, Avalos J (2012) Adenylyl cyclase plays a regulatory role in development, stress resistance and secondary metabolism in Fusarium fujikuroi. PLoS One 7:e28849
García-Martínez J, Brunk M, Avalos J, Terpitz U (2015) The CarO rhodopsin of the fungus Fusarium fujikuroi is a light-driven proton pump that retards spore germination. Sci Rep 5:7798
Gassel S, Breitenbach J, Sandmann G (2014) Genetic engineering of the complete carotenoid pathway towards enhanced astaxanthin formation in Xanthophyllomyces dendrorhous starting from a high-yield mutant. Appl Microbiol Biotechnol 98:345–350
Gassel S, Schewe H, Schmidt I et al (2013) Multiple improvement of astaxanthin biosynthesis in Xanthophyllomyces dendrorhous by a combination of conventional mutagenesis and metabolic pathway engineering. Biotechnol Lett 35:565–569
Georgiou CD, Tairis N, Polycratis A (2001a) Production of β-carotene by Sclerotinia sclerotiorum and its role in sclerotium differentiation. Mycol Res 105:1110–1115
Georgiou CD, Zervoudakis G, Tairis N, Kornaros M (2001b) β-Carotene production and its role in sclerotial differentiation of Sclerotium rolfsii. Fungal Genet Biol 34:11–20
Gessler NN, Sokolov AV, Bykhovsky VY, Belozerskaya TA (2002) Superoxide dismutase and catalase activities in carotenoid-synthesizing fungi Blakeslea trispora and Neurospora crassa in oxidative stress. Appl Biochem Microbiol 38:205–209
Girard P, Falconnier B, Bricout J, Vladescu B (1994) β-Carotene producing mutants of Phaffia rhodozyma. Appl Microbiol Biotechnol 41:183–191
Goksungur Y, Mantzouridou F, Roukas T, Kotzekidou P (2004) Production of β-carotene from beet molasses by Blakeslea trispora in stirred-tank and bubble column reactors: development of a mathematical modeling. Appl Biochem Biotechnol 112:37–54
Goldie AH, Subden RE (1973) The neutral carotenoids of wild-type and mutant strains of Neurospora crassa. Biochem Genet 10:275–284
Goodwin TW (1980) The biochemistry of the carotenoids, 2nd edn. Chapman & Hall, London
Govind NS, Cerdá-Olmedo E (1986) Sexual activation of carotenogenesis in Phycomyces blakesleeanus. J Gen Microbiol 132:2775–2780
Gu WL, An GH, Johnson EA (1997) Ethanol increases carotenoid production in Phaffia rhodozyma. J Ind Microbiol Biotechnol 19:114–117
Han JR, Zhao WJ, Gao YY, Yuan JM (2005) Effect of oxidative stress and exogenous β-carotene on sclerotial differentiation and carotenoid yield of Penicillium sp. PT95. Lett Appl Microbiol 40:412–417
Han M, He Q, Zhang WG (2012) Carotenoids production in different culture conditions by Sporidiobolus pararoseus. Prep Biochem Biotechnol 42:293–303
Hara KY, Morita T, Mochizuki M et al (2014) Development of a multi-gene expression system in Xanthophyllomyces dendrorhous. Microb Cell Factories 13:175
Harding RW (1974) The effect of temperature on photo-induced carotenoid biosynthesis in Neurospora crassa. Plant Physiol 54:142–147
Harding RW (1973) Inhibition of conidiation and photoinduced carotenoid biosynthesis by cyclic AMP. Neurospora Newsl 20:20–21
Harding RW, Huang PC, Mitchell HK (1969) Photochemical studies of the carotenoid biosynthetic pathway in Neurospora crassa. Arch Biochem Biophys 129:696–707
Harding RW, Philip DQ, Drozdowicz BZ, Williams NP (1984) A Neurospora crassa mutant which overaccumulates carotenoid pigments. Neurospora Newsl 31:23–25
Harding RW, Turner RV (1981) Photoregulation of the carotenoid biosynthetic pathway in albino and white collar mutants of Neurospora crassa. Plant Physiol 68:745–749
Hausmann A, Sandmann G (2000) A single five-step desaturase is involved in the carotenoid biosynthesis pathway to β-carotene and torulene in Neurospora crassa. Fungal Genet Biol 30:147–153
Haxo F (1950) Carotenoids of the mushroom Cantharellus cinnabarinus. Bot Gaz 112:228–232
Hayman GT, Mannarelli BM, Leathers TD (1995) Production of carotenoids by Phaffia rhodozyma grown on media composed of corn wet-milling co-products. J Ind Microbiol 14:389–395
He Q, Liu Y (2005) Molecular mechanism of light responses in Neurospora: from light-induced transcription to photoadaptation. Genes Dev 19:2888–2899
Hernández-Almanza A, Cesar Montanez J, Aguilar-González MA et al (2014a) Rhodotorula glutinis as source of pigments and metabolites for food industry. Food Biosci 5:64–72
Hernández-Almanza A, Montañez-Sáenz J, Martínez-Ávila C et al (2014b) Carotenoid production by Rhodotorula glutinis YB-252 in solid-state fermentation. Food Biosci 7:31–36
Herz S, Weber RW, Anke H et al (2007) Intermediates in the oxidative pathway from torulene to torularhodin in the red yeasts Cystofilobasidium infirmominiatum and C. capitatum (Heterobasidiomycetes, Fungi). Phytochemistry 68:2503–2511
Higuera-Ciapara I, Félix-Valenzuela L, Goycoolea FM (2006) Astaxanthin: a review of its chemistry and applications. Crit Rev Food Sci Nutr 46:185–196
Hsu W-J, Poling SM, Yokoyama H (1974) Effects of amines on the carotenogenesis in Blakeslea trispora. Phytochemistry 13:415–419
Hsu WJ, Yokoyama H, Coggins CW (1972) Carotenoid biosynthesis in Blakeslea trispora. Phytochemistry 11:2985–2990
Hu X, Li H, Tang P et al (2013a) GC-MS-based metabolomics study of the responses to arachidonic acid in Blakeslea trispora. Fungal Genet Biol 57:33–41
Hu X, Ma X, Tang P, Yuan Q (2013b) Improved β-carotene production by oxidative stress in Blakeslea trispora induced by liquid paraffin. Biotechnol Lett 35:559–563
Hu X, Sun J, Yuan Q (2012) Improved β-carotene biosynthesis and gene transcription in Blakeslea trispora with arachidonic acid. Biotechnol Lett 34:2107–2111
Huang PC (1964) Recombination and complementation of albino mutants in Neurospora. Genetics 49:453–469
Idnurm A, Rodríguez-Romero J, Corrochano LM et al (2006) The Phycomyces madA gene encodes a blue-light photoreceptor for phototropism and other light responses. Proc Natl Acad Sci U S A 103:4546–4551
Iurkov AM, Vustin MM, Tiaglov BV et al (2008) Pigmented basidiomycete yeasts are a promising source of carotenoids and ubiquinone Q10. Mikrobiologiia 77:5–10
Jayaram M, Leutwiler L, Delbrück M (1980) Light-induced carotene synthesis in mutants of Phycomyces with abnormal phototropism. Photochem Photobiol 32:241–245
Jeong JC, Lee IY, Kim SW, Park YH (1999) Stimulation of β-carotene synthesis by hydrogen peroxide in Blakeslea trispora. Biotechnol Lett 21:683–686
Jin JM, Lee J, Lee YW (2010) Characterization of carotenoid biosynthetic genes in the ascomycete Gibberella zeae. FEMS Microbiol Lett 302:197–202
Jirasripongpun K, Pewlong W, Kitraksa P, Krudngern C (2008) Carotenoid production by Xanthophyllomyces dendrorhous: use of pineapple juice as a production medium. Lett Appl Microbiol 47:112–116
Johnson EA (2003) Phaffia rhodozyma: colorful odyssey. Int Microbiol 6:169–174
Johnson EA, Lewis MJ (1979) Astaxanthin formation by the yeast Phaffia rhodozyma. J Gen Microbiol 115:173–183
Kajiwara S, Fraser PD, Kondo K, Misawa N (1997) Expression of an exogenous isopentenyl diphosphate isomerase gene enhances isoprenoid biosynthesis in Escherichia coli. Biochem J 324:421–426
Kim S-K, Lee J-H, Lee C-H, Yoon Y-C (2007) Increased carotenoid production in Xanthophyllomyces dendrorhous G276 using plant extracts. J Microbiol Seoul Korea 45:128–132
Kirti K, Amita S, Priti S et al (2014) Colorful world of microbes: carotenoids and their applications. Adv Biol 2014:1–13
Krinsky NI, Johnson EJ (2005) Carotenoid actions and their relation to health and disease. Mol Asp Med 26:459–516
Kritsky MS, Sokolovsky VY, Belozerskaya TA, Chernysheva EK (1982) Relationship between cyclic AMP level and accumulation of carotenoid pigments in Neurospora crassa. Arch Microbiol 133:206–208
Kucsera J, Pfeiffer I, Ferenczy L (1998) Homothallic life cycle in the diploid red yeast Xanthophyllomyces dendrorhous (Phaffia rhodozyma). Antonie Van Leeuwenhoek 73:163–168
Kuzina V, Cerdá-Olmedo E (2006) Modification of sexual development and carotene production by acetate and other small carboxylic acids in Blakeslea trispora and Phycomyces blakesleeanus. Appl Environ Microbiol 72:4917–4922
Kuzina V, Domenech C, Cerdá-Olmedo E (2006) Relationships among the biosyntheses of ubiquinone, carotene, sterols, and triacylglycerols in Zygomycetes. Arch Microbiol 186:485–493
Kuzina V, Ramírez-Medina H, Visser H et al (2008) Genes involved in carotene synthesis and mating in Blakeslea trispora. Curr Genet 54:143–152
Lampila LE, Wallen SE, Bullerman LB (1985) A review of factors affecting biosynthesis of carotenoids by the order Mucorales. Mycopathologia 90:65–80
Lange N, Steinbüchel A (2011) β-carotene production by Saccharomyces cerevisiae with regard to plasmid stability and culture media. Appl Microbiol Biotechnol 91:1611–1622
Ledetzky N, Osawa A, Iki K et al (2014) Multiple transformation with the crtYB gene of the limiting enzyme increased carotenoid synthesis and generated novel derivatives in Xanthophyllomyces dendrorhous. Arch Biochem Biophys 545:141–147
Lee JJL, Chen L, Cao B, Chen WN (2016) Engineering Rhodosporidium toruloides with a membrane transporter facilitates production and separation of carotenoids and lipids in a bi-phasic culture. Appl Microbiol Biotechnol 100:869–877
Lewis MJ, Ragot N, Berlant MC, Miranda M (1990) Selection of astaxanthin-overproducing mutants of Phaffia rhodozyma with β-Ionone. Appl Environ Microbiol 56:2944–2945
Li C, Schmidhauser TJ (1995) Developmental and photoregulation of al-1 and al-2, structural genes for two enzymes essential for carotenoid biosynthesis in Neurospora. Dev Biol 169:90–95
Li Q, Sun Z, Li J, Zhang Y (2013) Enhancing beta-carotene production in Saccharomyces cerevisiae by metabolic engineering. FEMS Microbiol Lett 345:94–101
Linden H, Rodríguez-Franco M, Macino G (1997) Mutants of Neurospora crassa defective in regulation of blue light perception. Mol Gen Genet 254:111–118
Linnemannstöns P, Prado MM, Fernández-Martín R et al (2002) A carotenoid biosynthesis gene cluster in Fusarium fujikuroi: the genes carB and carRA. Mol Gen Genomics 267:593–602
Liu X-J, Liu R-S, Li H-M, Tang Y-J (2012) Lycopene production from synthetic medium by Blakeslea trispora NRRL 2895 (+) and 2896 (-) in a stirred-tank fermenter. Bioprocess Biosyst Eng 35:739–749
Liu YS, Wu JY (2006a) Use of n-hexadecane as an oxygen vector to improve Phaffia rhodozyma growth and carotenoid production in shake-flask cultures. J Appl Microbiol 101:1033–1038
Liu YS, Wu JY (2006b) Hydrogen peroxide-induced astaxanthin biosynthesis and catalase activity in Xanthophyllomyces dendrorhous. Appl Microbiol Biotechnol 73:663–668
Liu YS, Wu JY (2007) Perfusion culture process plus H2O2 stimulation for efficient astaxanthin production by Xanthophyllomyces dendrorhous. Biotechnol Bioeng 97:568–573
Liu Z-Q, Zhang J-F, Zheng Y-G, Shen Y-C (2008) Improvement of astaxanthin production by a newly isolated Phaffia rhodozyma mutant with low-energy ion beam implantation. J Appl Microbiol 104:861–872
Lodato P, Alcaino J, Barahona S et al (2003) Alternative splicing of transcripts from crtI and crtYB genes of Xanthophyllomyces dendrorhous. Appl Environ Microbiol 69:4676–4682
Lodato P, Alcaíno J, Barahona S et al (2007) Expression of the carotenoid biosynthesis genes in Xanthophyllomyces dendrorhous. Biol Res 40:73–84
Lodato P, Alcaíno J, Barahona S et al (2004) Study of the expression of carotenoid biosynthesis genes in wild-type and deregulated strains of Xanthophyllomyces dendrorhous (Ex.: Phaffia rhodozyma). Biol Res 37:83–93
López J, Essus K, Kim I et al (2015) Production of β-ionone by combined expression of carotenogenic and plant CCD1 genes in Saccharomyces cerevisiae. Microb Cell Factories 14:84
López-Díaz I, Cerdá-Olmedo E (1980) Relationship of photocarotenogenesis to other behavioural and regulatory responses in Phycomyces. Planta 150:134–139
López-Nieto MJ, Costa J, Peiro E et al (2004) Biotechnological lycopene production by mated fermentation of Blakeslea trispora. Appl Microbiol Biotechnol 66:153–159
Lorca-Pascual JM, Murcia-Flores L, Garre V et al (2004) The RING-finger domain of the fungal repressor crgA is essential for accurate light regulation of carotenogenesis. Mol Microbiol 52:1463–1474
Loto I, Gutiérrez MS, Barahona S et al (2012) Enhancement of carotenoid production by disrupting the C22-sterol desaturase gene (CYP61) in Xanthophyllomyces dendrorhous. BMC Microbiol 12:235
Madhour A, Anke H, Mucci A et al (2005) Biosynthesis of the xanthophyll plectaniaxanthin as a stress response in the red yeast Dioszegia (Tremellales, Heterobasidiomycetes, Fungi). Phytochemistry 66:2617–2626
Mantzouridou F, Naziri E, Tsimidou MZ (2008) Industrial glycerol as a supplementary carbon source in the production of β-carotene by Blakeslea trispora. J Agric Food Chem 56:2668–2675
Mantzouridou F, Roukasa T, Kotzekidoua P, Liakopoulou M (2002) Optimization of β-carotene production from synthetic medium by Blakeslea trispora: a mathematical modeling. Appl Biochem Biotechnol 101:153–175
Mantzouridou F, Tsimidou MZ (2008) Lycopene formation in Blakeslea trispora. Chemical aspects of a bioprocess. Trends Food Sci Technol 19:363–371
Mantzouridou F, Tsimidou MZ, Roukas T (2006) Performance of crude olive pomace oil and soybean oil during carotenoid production by Blakeslea trispora in submerged fermentation. J Agric Food Chem 54:2575–2581
Marcoleta A, Niklitschek M, Wozniak A et al (2011) Glucose and ethanol-dependent transcriptional regulation of the astaxanthin biosynthesis pathway in Xanthophyllomyces dendrorhous. BMC Microbiol 11:190
Marova I, Carnecka M, Halienova A et al (2012) Use of several waste substrates for carotenoid-rich yeast biomass production. J Environ Manag 95:S338–S342
Martín JF, Gudiña E, Barredo JL (2008) Conversion of β-carotene into astaxanthin: two separate enzymes or a bifunctional hydroxylase-ketolase protein? Microb Cell Factories 7:3
Martínez C, Hermosilla G, León R et al (1998) Genetic transformation of astaxanthin mutants of Phaffia rhodozyma. Antonie Van Leeuwenhoek 73:147–153
Martinez-Moya P, Niehaus K, Alcaíno J et al (2015) Proteomic and metabolomic analysis of the carotenogenic yeast Xanthophyllomyces dendrorhous using different carbon sources. BMC Genomics 16:289
Martinez-Moya P, Watt SA, Niehaus K et al (2011) Proteomic analysis of the carotenogenic yeast Xanthophyllomyces dendrorhous. BMC Microbiol 11:131
Mata-Gómez LC, Montañez JC, Méndez-Zavala A, Aguilar CN (2014) Biotechnological production of carotenoids by yeasts: an overview. Microb Cell Factories 13:12
Medina HR, Cerdá-Olmedo E, Al-Babili S (2011) Cleavage oxygenases for the biosynthesis of trisporoids and other apocarotenoids in Phycomyces. Mol Microbiol 82:199–208
Mehta BJ, Cerdá-Olmedo E (1995) Mutants of carotene production in Blakeslea trispora. Appl Microbiol Biotechnol 42:836–838
Mehta BJ, Cerdá-Olmedo E (1999) Lycopene cyclization in Blakeslea trispora. Mycoscience 40:307–310
Mehta BJ, Cerdá-Olmedo E (2001) Intersexual partial diploids of Phycomyces. Genetics 158:635–641
Mehta BJ, Obraztsova IN, Cerdá-Olmedo E (2003) Mutants and intersexual heterokaryons of Blakeslea trispora for production of β-carotene and lycopene. Appl Environ Microbiol 69:4043–4048
Mehta BJ, Salgado LM, Bejarano ER, Cerdá-Olmedo E (1997) New mutants of Phycomyces blakesleeanus for β-carotene production. Appl Environ Microbiol 63:3657–3661
Mende K, Homann V, Tudzynski B (1997) The geranylgeranyl diphosphate synthase gene of Gibberella fujikuroi: isolation and expression. Mol Gen Genet 255:96–105
Meyer PS, Du Preez JC (1994) Photo-regulated astaxanthin production by Phaffia rhodozyma mutants. Syst Appl Microbiol 17:24–31
Meyer PS, du Preez JC, Kilian SG (1993) Selection and evaluation of astaxanthin-overproducing mutants of Phaffia rhodozyma. World J Microbiol Biotechnol 9:514–520
Miao L, Chi S, Tang Y et al (2011) Astaxanthin biosynthesis is enhanced by high carotenogenic gene expression and decrease of fatty acids and ergosterol in a Phaffia rhodozyma mutant strain. FEMS Yeast Res 11:192–201
Miao L, Wang Y, Chi S et al (2010) Reduction of fatty acid flux results in enhancement of astaxanthin synthesis in a mutant strain of Phaffia rhodozyma. J Ind Microbiol Biotechnol 37:595–602
Mitzka U, Rau W (1977) Composition and photoinduced biosynthesis of the carotenoids of a protoplast-like Neurospora crassa “slime” mutant. Arch Microbiol 111:261–263
Mitzka-Schnabel U (1985) Carotenogenic enzymes from Neurospora. Pure Appl Chem 57:667–669
Mitzka-Schnabel U, Rau W (1980) The subcellular distribution of carotenoids in Neurospora crassa. Phytochemistry 19:1409–1413
Miura Y, Kondo K, Saito T et al (1998) Production of the carotenoids lycopene, β-carotene, and astaxanthin in the food yeast Candida utilis. Appl Environ Microbiol 64:1226–1229
Moliné M, Flores MR, Libkind D et al (2010) Photoprotection by carotenoid pigments in the yeast Rhodotorula mucilaginosa: the role of torularhodin. Photochem Photobiol Sci 9:1145–1151
Moliné M, Libkind D, van Broock M (2012) Production of torularhodin, torulene, and β-carotene by Rhodotorula yeasts. Methods Mol Biol 898:275–283
Montanti J, Nghiem NP, Johnston DB (2011) Production of astaxanthin from cellulosic biomass sugars by mutants of the yeast Phaffia rhodozyma. Appl Biochem Biotechnol 164:655–665
Moore MM, Breedveld MW, Autor AP (1989) The role of carotenoids in preventing oxidative damage in the pigmented yeast, Rhodotorula mucilaginosa. Arch Biochem Biophys 270:419–431
Moran NA, Jarvik T (2010) Lateral transfer of genes from fungi underlies carotenoid production in aphids. Science 328:624–627
Murcia-Flores L, Lorca-Pascual JM, Garre V et al (2007) Non-AUG translation initiation of a fungal RING finger repressor involved in photocarotenogenesis. J Biol Chem 282:15394–15403
Murillo FJ, Calderón IL, López-Díaz I, Cerdá-Olmedo E (1978) Carotene-superproducing strains of Phycomyces. Appl Environ Microbiol 36:639–642
Murillo FJ, Cerdá-Olmedo E (1976) Regulation of carotene synthesis in Phycomyces. Mol Gen Genet 148:19–24
Murillo FJ, Torres-Martínez S, Aragón CM, Cerdá-Olmedo E (1981) Substrate transfer in carotene biosynthesis in Phycomyces. Eur J Biochem 119:511–516
Nanou K, Roukas T (2011) Stimulation of the biosynthesis of carotenes by oxidative stress in Blakeslea trispora induced by elevated dissolved oxygen levels in the culture medium. Bioresour Technol 102:8159–8164
Nanou K, Roukas T (2010) Oxidative stress response and morphological changes of Blakeslea trispora induced by butylated hydroxytoluene during carotene production. Appl Biochem Biotechnol 160:2415–2423
Nanou K, Roukas T (2013) Oxidative stress response of Blakeslea trispora induced by iron ions during carotene production in shake flask culture. Appl Biochem Biotechnol 169:2281–2289
Nanou K, Roukas T (2016) Waste cooking oil: a new substrate for carotene production by Blakeslea trispora in submerged fermentation. Bioresour Technol 203:198–203
Navarro E, Lorca-Pascual JM, Quiles-Rosillo MD et al (2001) A negative regulator of light-inducible carotenogenesis in Mucor circinelloides. Mol Gen Genomics 266:463–470
Navarro E, Ruiz-Pérez VL, Torres-Martínez S (2000) Overexpression of the crgA gene abolishes light requirement for carotenoid biosynthesis in Mucor circinelloides. Eur J Biochem 267:800–807
Navarro E, Sandmann G, Torres-Martínez S (1995) Mutants of the carotenoid biosynthetic pathway of Mucor circinelloides. Exp Mycol 19:186–190
Navarro-Sampedro L, Yanofsky C, Corrochano LM (2008) A genetic selection for Neurospora crassa mutants altered in their light regulation of transcription. Genetics 178:171–183
Nelson MA, Morelli G, Carattoli A et al (1989) Molecular cloning of a Neurospora crassa carotenoid biosynthetic gene (albino-3) regulated by blue light and the products of the white collar genes. Mol Cell Biol 9:1271–1276
Neupert W, Ludwig GD (1971) Sites of biosynthesis of outer and inner membrane proteins of Neurospora crassa mitochondria. Eur J Biochem 19:523–532
Niklitschek M, Alcaíno J, Barahona S et al (2008) Genomic organization of the structural genes controlling the astaxanthin biosynthesis pathway of Xanthophyllomyces dendrorhous. Biol Res 41:93–108
Ninet L, Renaut J, Tissier R (1969) Activation of the biosynthesis of carotenoids by Blakeslea trispora. Biotechnol Bioeng 11:1195–1210
Ojima K, Breitenbach J, Visser H et al (2006) Cloning of the astaxanthin synthase gene from Xanthophyllomyces dendrorhous (Phaffia rhodozyma) and its assignment as a β-carotene 3-hydroxylase/4-ketolase. Mol Gen Genomics 275:148–158
Ootaki T, Crafts-Lighty A, Delbrück M, Hsu WJ (1973) Complementation between mutants of Phycomyces deficient with respect to carotenogenesis. Mol Gen Genet 121:57–70
Paietta J, Sargent ML (1981) Photoreception in Neurospora crassa: correlation of reduced light sensitivity with flavin deficiency. Proc Natl Acad Sci U S A 78:5573–5577
Paietta J, Sargent ML (1983) Modification of blue light photoresponses by riboflavin analogs in Neurospora crassa. Plant Physiol 72:764–766
Papaioannou EH, Liakopoulou-Kyriakides M (2012) Agro-food wastes utilization by Blakeslea trispora for carotenoids production. Acta Biochim Pol 59:151–153
Papp T, Csernetics A, Nagy G et al (2013) Canthaxanthin production with modified Mucor circinelloides strains. Appl Microbiol Biotechnol 97:4937–4950
Papp T, Velayos A, Bartok T et al (2006) Heterologous expression of astaxanthin biosynthesis genes in Mucor circinelloides. Appl Microbiol Biotechnol 69:526–531
Parajo JC, Santos VV, Vazquez M (1998) Production of carotenoids by Phaffia rhodozyma growing on media made from hemicellulosic hydrolysates of Eucalyptus globulus wood. Biotechnol Bioeng 59:501–506
Pegklidou K, Mantzouridou F, Tsimidou MZ (2008) Lycopene production using Blakeslea trispora in the presence of 2-methyl imidazole: yield, selectivity, and safety aspects. J Agric Food Chem 56:4482–4490
Polaino S, Herrador MM, Cerdá-Olmedo E, Barrero AF (2010) Splitting of β-carotene in the sexual interaction of Phycomyces. Org Biomol Chem 8:4229–4231
Prado MM, Prado-Cabrero A, Fernández-Martín R, Avalos J (2004) A gene of the opsin family in the carotenoid gene cluster of Fusarium fujikuroi. Curr Genet 46:47–58
Prado-Cabrero A, Estrada AF, Al-Babili S, Avalos J (2007) Identification and biochemical characterization of a novel carotenoid oxygenase: elucidation of the cleavage step in the Fusarium carotenoid pathway. Mol Microbiol 64:448–460
Prado-Cabrero A, Schaub P, Díaz-Sánchez V et al (2009) Deviation of the neurosporaxanthin pathway towards β-carotene biosynthesis in Fusarium fujikuroi by a point mutation in the phytoene desaturase gene. FEBS J 276:4582–4597
Qiang W, Ling-ran F, Luo W et al (2014) Mutation breeding of lycopene-producing strain Blakeslea trispora by a novel atmospheric and room temperature plasma (ARTP). Appl Biochem Biotechnol 174:452–460
Quiles-Rosillo MD, Ruiz-Vázquez RM, Torres-Martínez S, Garre V (2005) Light induction of the carotenoid biosynthesis pathway in Blakeslea trispora. Fungal Genet Biol 42:141–153
Quiles-Rosillo MD, Torres-Martínez S, Garre V (2003) cigA, a light-inducible gene involved in vegetative growth in Mucor circinelloides is regulated by the carotenogenic repressor crgA. Fungal Genet Biol 38:122–132
Ramírez J, Gutierrez H, Gschaedler A (2001) Optimization of astaxanthin production by Phaffia rhodozyma through factorial design and response surface methodology. J Biotechnol 88:259–268
Rao AV, Rao LG (2007) Carotenoids and human health. Pharmacol Res 55:207–216
Rau W (1962) Über den Einfluss der Temperatur auf die lichtabhängige Carotinoidbildung von Fusarium aquaeductuum. Planta 59:123–137
Rau W (1967) Untersuchungen über die lichtabhängige Carotinoidsynthese. I Das Wirkungsspektrum von Fusarium aquaeductuum. Planta 72:14–28
Rau W, Lindemann I, Rau-Hund A (1968) Untersuchungen über die lichtabhängige Carotinoidsynthese. III Die Farbstoffbildung von Neurospora crassa in Submerskultur. Planta 80:309–316
Razani SH, Mousavi SM, Yeganeh HM, Marc I (2007) Fatty acid and carotenoid production by Sporobolomyces ruberrimus when using technical glycerol and ammonium sulfate. J Microbiol Biotechnol 17:1591–1597
Revuelta JL, Eslava AP (1983) A new gene (carC) involved in the regulation of carotenogenesis in Phycomyces. Mol Gen Genet 192:225–229
Ribeiro BD, Barreto DW, Coelho MAZ (2011) Technological aspects of β-carotene production. Food Bioprocess Technol 4:693–701
Rodríguez-Ortiz R, Limón MC, Avalos J (2009) Regulation of carotenogenesis and secondary metabolism by nitrogen in wild-type Fusarium fujikuroi and carotenoid-overproducing mutants. Appl Environ Microbiol 75:405–413
Rodríguez-Ortiz R, Limón MC, Avalos J (2013) Functional analysis of the carS gene of Fusarium fujikuroi. Mol Gen Genomics 288:157–173
Rodríguez-Ortiz R, Michielse C, Rep M et al (2012) Genetic basis of carotenoid overproduction in Fusarium oxysporum. Fungal Genet Biol 49:684–696
Rodríguez-Sáiz M, de la Fuente JL, Barredo JL (2010) Xanthophyllomyces dendrorhous for the industrial production of astaxanthin. Appl Microbiol Biotechnol 88:645–658
Rodríguez-Saiz M, Paz B, De La Fuente JL et al (2004) Blakeslea trispora genes for carotene biosynthesis. Appl Environ Microbiol 70:5589–5594
Rohmer M, Knani M, Simonin P et al (1993) Isoprenoid biosynthesis in bacteria: a novel pathway for the early steps leading to isopentenyl diphosphate. Biochem J 15:517–524
Roukas T (2016) The role of oxidative stress on carotene production by Blakeslea trispora in submerged fermentation. Crit Rev Biotechnol 36:424–433
Roukas T, Varzakakou M, Kotzekidou P (2015) From cheese whey to carotenes by Blakeslea trispora in a bubble column reactor. Appl Biochem Biotechnol 175:182–193
Ruiz-Hidalgo MJ, Benito EP, Sandmann G, Eslava AP (1997a) The phytoene dehydrogenase gene of Phycomyces: regulation of its expression by blue light and vitamin A. Mol Gen Genet 253:734–744
Ruiz-Roldán MC, Garre V, Guarro J et al (2008) Role of the white collar 1 photoreceptor in carotenogenesis, UV resistance, hydrophobicity, and virulence of Fusarium oxysporum. Eukaryot Cell 7:1227–1230
Saelices L, Youssar L, Holdermann I et al (2007) Identification of the gene responsible for torulene cleavage in the Neurospora carotenoid pathway. Mol Gen Genomics 278:527–537
Saenge C, Cheirsilp B, Suksaroge TT, Bourtoom T (2011) Potential use of oleaginous red yeast Rhodotorula glutinis for the bioconversion of crude glycerol from biodiesel plant to lipids and carotenoids. Process Biochem 46:210–218
Sahadevan Y, Richter-Fecken M, Kaerger K et al (2013) Early and late trisporoids differentially regulate β-carotene production and gene transcript levels in the mucoralean fungi Blakeslea trispora and Mucor mucedo. Appl Environ Microbiol 79:7466–7475
Sakaki H, Kaneno H, Sumiya Y et al (2002a) A new carotenoid glycosyl ester isolated from a marine microorganism, Fusarium strain T-1. J Nat Prod 65:1683–1684
Sakaki H, Nakanishi T, Satonaka K et al (2000) Properties of a high-torularhodin-producing mutant of Rhodotorula glutinis cultivated under oxidative stress. J Biosci Bioeng 89:203–205
Sakaki H, Nakanishi T, Tada A et al (2001) Activation of torularhodin production by Rhodotorula glutinis using weak white light irradiation. J Biosci Bioeng 92:294–297
Sakaki H, Nochide H, Komemushi S, Miki W (2002b) Effect of active oxygen species on the productivity of torularhodin by Rhodotorula glutinis No. 21. J Biosci Bioeng 93:338–340
Salgado LM, Avalos J, Bejarano ER, Cerdá-Olmedo E (1991) Correlation between in vivo and in vitro carotenogenesis in Phycomyces. Phytochemistry 30:2587–2591
Salgado LM, Bejarano ER, Cerdá-Olmedo E (1989) Carotene-superproducing mutants of Phycomyces blakesleeanus. Exp Mycol 13:332–336
Salgado LM, Cerdá-Olmedo E (1992) Genetic interactions in the regulation of carotenogenesis in Phycomyces. Curr Genet 21:67–71
Sandmann G (2002) Combinatorial biosynthesis of carotenoids in a heterologous host: a powerful approach for the biosynthesis of novel structures. Chembiochem Eur J Chem Biol 3:629–635
Sandmann G, Misawa N (2002) Fungal carotenoids. In: Osiewacz HD (ed) The mycota X. Industrial applications. Springer, Berlin, pp 247–262
Sandmann G, Misawa N, Wiedemann M et al (1993) Functional identification of al-3 from Neurospora crassa as the gene for geranylgeranyl pyrophosphate synthase by complementation with crt genes, in vitro characterization of the gene product and mutant analysis. J Photochem Photobiol B 18:245–251
Sandmann G, Takaichi S, Fraser PD (2008) C(35)-apocarotenoids in the yellow mutant Neurospora crassa YLO. Phytochemistry 69:2886–2890
Sandmann G, Zhu C, Krubasik P, Fraser PD (2006) The biotechnological potential of the al-2 gene from Neurospora crassa for the production of monocyclic keto hydroxy carotenoids. Biochim Biophys Acta 1761:1085–1092
Sanz C, Alvarez MI, Orejas M et al (2002) Interallelic complementation provides genetic evidence for the multimeric organization of the Phycomyces blakesleeanus phytoene dehydrogenase. Eur J Biochem 269:902–908
Sanz C, Benito EP, Orejas M et al (2010) Protein-DNA interactions in the promoter region of the Phycomyces carB and carRA genes correlate with the kinetics of their mRNA accumulation in response to light. Fungal Genet Biol 47:773–781
Sanz C, Rodríguez-Romero J, Idnurm A et al (2009) Phycomyces MADB interacts with MADA to form the primary photoreceptor complex for fungal phototropism. Proc Natl Acad Sci U S A 106:7095–7100
Sanz C, Velayos A, Alvarez MI et al (2011) Functional analysis of the Phycomyces carRA gene encoding the enzymes phytoene synthase and lycopene cyclase. PLoS One 6:e23102
Schachtschabel D, David A, Menzel K-D et al (2008) Cooperative biosynthesis of trisporoids by the (+) and (-) mating types of the zygomycete Blakeslea trispora. Chembiochem Eur J Chem Biol 9:3004–3012
Schmidhauser TJ, Lauter FR, Russo VE, Yanofsky C (1990) Cloning, sequence, and photoregulation of al-1, a carotenoid biosynthetic gene of Neurospora crassa. Mol Cell Biol 10:5064–5070
Schmidhauser TJ, Lauter FR, Schumacher M et al (1994) Characterization of al-2, the phytoene synthase gene of Neurospora crassa. Cloning, sequence analysis, and photoregulation. J Biol Chem 269:12060–12066
Schmidt AD, Heinekamp T, Matuschek M et al (2005) Analysis of mating-dependent transcription of Blakeslea trispora carotenoid biosynthesis genes carB and carRA by quantitative real-time PCR. Appl Microbiol Biotechnol 67:549–555
Schmidt I, Schewe H, Gassel S et al (2011) Biotechnological production of astaxanthin with Phaffia rhodozyma/Xanthophyllomyces dendrorhous. Appl Microbiol Biotechnol 89:555–571
Schmidt-Dannert C (2000) Engineering novel carotenoids in microorganisms. Curr Opin Biotechnol 11:255–261
Schneider T, Graeff-Hönninger S, French WT et al (2013) Lipid and carotenoid production by oleaginous red yeast Rhodotorula glutinis cultivated on brewery effluents. Energy 61:34–43
Schroeder WA, Johnson EA (1995a) Singlet oxygen and peroxyl radicals regulate carotenoid biosynthesis in Phaffia rhodozyma. J Biol Chem 270:18374–18379
Schroeder WA, Johnson EA (1993) Antioxidant role of carotenoids in Phaffia rhodozyma. J Gen Microbiol 139:907–912
Schroeder WA, Johnson EA (1995b) Carotenoids protect Phaffia rhodozyma against singlet oxygen damage. J Ind Microbiol 14:502–507
Schrott EL (1980) Fluence response relationship of carotenogenesis in Neurospora crassa. Planta 150:174–179
Schrott EL (1981) The biphasic fluence response of carotenogenesis in Neurospora crassa: temporary insensitivity of the photoreceptor system. Planta 151:371–374
Schwerdtfeger C, Linden H (2003) VIVID is a flavoprotein and serves as a fungal blue light photoreceptor for photoadaptation. EMBO J 22:4846–4855
Shi F, Zhan W, Li Y, Wang X (2014) Temperature influences β-carotene production in recombinant Saccharomyces cerevisiae expressing carotenogenic genes from Phaffia rhodozyma. World J Microbiol Biotechnol 30:125–133
Shi Y, Xin X, Yuan Q (2012) Improved lycopene production by Blakeslea trispora with isopentenyl compounds and metabolic precursors. Biotechnol Lett 34:849–852
Shimada H, Kondo K, Fraser PD et al (1998) Increased carotenoid production by the food yeast Candida utilis through metabolic engineering of the isoprenoid pathway. Appl Environ Microbiol 64:2676–2680
Shrode LB, Lewis ZA, White LD et al (2001) vvd is required for light adaptation of conidiation-specific genes of Neurospora crassa, but not circadian conidiation. Fungal Genet Biol 32:169–181
Sieiro C, Poza M, de Miguel T, Villa TG (2003) Genetic basis of microbial carotenogenesis. Int Microbiol 6:11–16
Silva F, Navarro E, Peñaranda A et al (2008) A RING-finger protein regulates carotenogenesis via proteolysis-independent ubiquitylation of a White Collar-1-like activator. Mol Microbiol 70:1026–1036
Silva F, Torres-Martínez S, Garre V (2006) Distinct white collar-1 genes control specific light responses in Mucor circinelloides. Mol Microbiol 61:1023–1037
Simpson KL, Nakayama TO, Chichester CO (1964) Biosynthesis of yeast carotenoids. J Bacteriol 88:1688–1694
Sokolovsky VY, Lauter FR, Müller-Röber B et al (1992) Nitrogen regulation of blue light-inducible genes in Neurospora crassa. J Gen Microbiol 138:2045–2049
Sperstad S, Lutnaes BF, Stormo SK et al (2006) Torularhodin and torulene are the major contributors to the carotenoid pool of marine Rhodosporidium babjevae (Golubev). J Ind Microbiol Biotechnol 33:269–273
Spurgeon SL, Turner RV, Harding RW (1979) Biosynthesis of phytoene from isopentenyl pyrophosphate by a Neurospora enzyme system. Arch Biochem Biophys 195:23–29
Stachowiak B (2013) Efficiency of selected mutagens in generating Xanthophyllomyces dendrorhous strains hyperproducing astaxanthin. Pol J Microbiol 62:67–72
Stahl W, Sies H (2005) Bioactivity and protective effects of natural carotenoids. Biochim Biophys Acta 1740:101–107
Stahl W, Sies H (2003) Antioxidant activity of carotenoids. Mol Asp Med 24:345–351
Strobel I, Breitenbach J, Scheckhuber CQ et al (2009) Carotenoids and carotenogenic genes in Podospora anserina: engineering of the carotenoid composition extends the life span of the mycelium. Curr Genet 55:175–184
Sun J, Li H, Yuan Q (2012) Metabolic regulation of trisporic acid on Blakeslea trispora revealed by a GC-MS-based metabolomic approach. PLoS One 7:e46110
Sun N, Lee S, Song KB (2004) Characterization of a carotenoid-hyperproducing yeast mutant isolated by low-dose gamma irradiation. Int J Food Microbiol 94:263–267
Sutter RP (1970) Effect of light on β-carotene accumulation in Blakeslea trispora. J Gen Microbiol 64:215–221
Sutter RP (1987) Sexual development. In: Cerdá-Olmedo E, Lipson ED (eds) Phycomyces. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, pp 317–336
Tagua VG, Medina HR, Martín-Domínguez R et al (2012) A gene for carotene cleavage required for pheromone biosynthesis and carotene regulation in the fungus Phycomyces blakesleeanus. Fungal Genet Biol 49:398–404
Tang Q, Li Y, Yuan QP (2008) Effects of an ergosterol synthesis inhibitor on gene transcription of terpenoid biosynthesis in Blakeslea trispora. Curr Microbiol 57:527–531
Tapiero H, Townsend DM, Tew KD (2004) The role of carotenoids in the prevention of human pathologies. Biomed Pharmacother Biomedecine Pharmacother 58:100–110
Thewes S, Prado-Cabrero A, Prado MM et al (2005) Characterization of a gene in the car cluster of Fusarium fujikuroi that codes for a protein of the carotenoid oxygenase family. Mol Gen Genomics 274:217–228
Tinoi J, Rakariyatham N, Deming RL (2006) Utilization of mustard waste isolates for improved production of astaxanthin by Xanthophyllomyces dendrorhous. J Ind Microbiol Biotechnol 33:309–314
Tinoi J, Rakariyatham N, Deming RL (2005) Simplex optimization of carotenoid production by Rhodotorula glutinis using hydrolyzed mung bean waste flour as substrate. Process Biochem 40:2551–2557
Torres-Martínez S, Murillo FJ, Cerdá-Olmedo E (1980) Genetics of lycopene cyclization and substrate transfer in β-carotene biosynthesis in Phycomyces. Genet Res 36:299–309
Torres-Martínez S, Ruiz-Vázquez RM, Garre V et al (2012) Molecular tools for carotenogenesis analysis in the zygomycete Mucor circinelloides. Methods Mol Biol 898:85–107
Tsolakis G, Parashi E, Galland P, Kotzabasis K (1999) Blue light signaling chains in Phycomyces: phototransduction of carotenogenesis and morphogenesis involves distinct protein kinase/phosphatase elements. Fungal Genet Biol 28:201–213
Tudzynski B (2005) Gibberellin biosynthesis in fungi: genes, enzymes, evolution, and impact on biotechnology. Appl Microbiol Biotechnol 66:597–611
Ukibe K, Hashida K, Yoshida N, Takagi H (2009) Metabolic engineering of Saccharomyces cerevisiae for astaxanthin production and oxidative stress tolerance. Appl Environ Microbiol 75:7205–7211
Valadon LRG, Cooke RC (1963) Carotenoid pigments of the nematode-trapping hyphomycete Arthrobotrys oligospora (fres.) Phytochemistry 2:103–105
Valadon LRG, Mummery RS (1969) Biosynthesis of neurosporaxanthin. Microbios 1A:3–8
Valadon LRG, Mummery RS (1977) Natural β-apo-4′-carotenoic acid methyl ester in the fungus Verticillium agaricinum. Phytochemistry 16:613–614
Valduga E, Rausch Ribeiro AH, Cence K et al (2014) Carotenoids production from a newly isolated Sporidiobolus pararoseus strain using agroindustrial substrates. Biocatal Agric Biotechnol 3:207–213
Valduga E, Valerio A, Treichel H et al (2009) Kinetic and stoichiometric parameters in the production of carotenoids by Sporidiobolus salmonicolor (CBS 2636) in synthetic and agroindustrial media. Appl Biochem Biotechnol 157:61–69
van Eijk GW, Mummery RS, Roeymans HJ, Valadon LR (1979) A comparative study of carotenoids of Aschersonia aleyroides and Aspergillus giganteus. Antonie Van Leeuwenhoek 45:417–422
Varzakakou M, Roukas T (2010) Identification of carotenoids produced from cheese whey by Blakeslea trispora in submerged fermentation. Prep Biochem Biotechnol 40:76–82
Varzakakou M, Roukas T, Papaioannou E et al (2011) Autolysis of Blakeslea trispora during carotene production from cheese whey in an airlift reactor. Prep Biochem Biotechnol 41:7–21
Vázquez M, Martin AM (1998) Optimization of Phaffia rhodozyma continuous culture through response surface methodology. Biotechnol Bioeng 57:314–320
Velayos A, Blasco JL, Álvarez MI et al (2000a) Blue-light regulation of phytoene dehydrogenase (carB) gene expression in Mucor circinelloides. Planta 210:938–946
Velayos A, Eslava AP, Iturriaga EA (2000b) A bifunctional enzyme with lycopene cyclase and phytoene synthase activities is encoded by the carRP gene of Mucor circinelloides. Eur J Biochem 267:5509–5519
Velayos A, López-Matas MA, Ruiz-Hidalgo MJ, Eslava AP (1997) Complementation analysis of carotenogenic mutants of Mucor circinelloides. Fungal Genet Biol 22:19–27
Velayos A, Papp T, Aguilar-Elena R et al (2003) Expression of the carG gene, encoding geranylgeranyl pyrophosphate synthase, is up-regulated by blue light in Mucor circinelloides. Curr Genet 43:112–120
Verdoes JC, Krubasik P, Sandmann G, van Ooyen AJJ (1999a) Isolation and functional characterisation of a novel type of carotenoid biosynthetic gene from Xanthophyllomyces dendrorhous. Mol Gen Genet 262:453–461
Verdoes JC, Misawa N, van Ooyen AJJ (1999b) Cloning and characterization of the astaxanthin biosynthetic gene encoding phytoene desaturase of Xanthophyllomyces dendrorhous. Biotechnol Bioeng 63:750–755
Verdoes JC, Sandmann G, Visser H et al (2003) Metabolic engineering of the carotenoid biosynthetic pathway in the yeast Xanthophyllomyces dendrorhous (Phaffia rhodozyma). Appl Environ Microbiol 69:3728–3738
Verwaal R, Jiang Y, Wang J et al (2010) Heterologous carotenoid production in Saccharomyces cerevisiae induces the pleiotropic drug resistance stress response. Yeast Chichester Engl 27:983–998
Verwaal R, Wang J, Meijnen J-P et al (2007) High-level production of beta-carotene in Saccharomyces cerevisiae by successive transformation with carotenogenic genes from Xanthophyllomyces dendrorhous. Appl Environ Microbiol 73:4342–4350
Visser H, van Ooyen AJJ, Verdoes JC (2003) Metabolic engineering of the astaxanthin-biosynthetic pathway of Xanthophyllomyces dendrorhous. FEMS Yeast Res 4:221–231
Vittorioso P, Carattoli A, Londei P, Macino G (1994) Internal translational initiation in the mRNA from the Neurospora crassa albino-3 gene. J Biol Chem 269:26650–26654
Wang GY, Keasling JD (2002) Amplification of HMG-CoA reductase production enhances carotenoid accumulation in Neurospora crassa. Metab Eng 4:193–201
Wang H-B, Luo J, Huang X-Y et al (2014) Oxidative stress response of Blakeslea trispora induced by H2O2 during β-carotene biosynthesis. J Ind Microbiol Biotechnol 41:555–561
Wang Q, Luo W, Gu Q-Y et al (2013) Enhanced lycopene content in Blakeslea trispora by effective mutation-screening method. Appl Biochem Biotechnol 171:1692–1700
Wang W, Yu L, Zhou P (2006) Effects of different fungal elicitors on growth, total carotenoids and astaxanthin formation by Xanthophyllomyces dendrorhous. Bioresour Technol 97:26–31
Wang Y, Chen X, Hong X et al (2016) Cyclase inhibitor tripropylamine significantly enhanced lycopene accumulation in Blakeslea trispora. J Biosci Bioeng 122(5):570–576
Will OH, Ruddat M, Garber ED, Kezdy FJ (1984) Characterization of carotene accumulation in Ustilago violacea using high-performance liquid chromatography. Curr Microbiol 10:57–63
Will OH, Ruddat M, Newland NA (1985) Characterization of carotene accumulation in species of the fungal genus Ustilago using high-performance liquid chromatography. Bot Gaz 146:204–207
Woodside JV, McGrath AJ, Lyner N, McKinley MC (2015) Carotenoids and health in older people. Maturitas 80:63–68
Wozniak A, Lozano C, Barahona S et al (2011) Differential carotenoid production and gene expression in Xanthophyllomyces dendrorhous grown in a nonfermentable carbon source. FEMS Yeast Res 11:252–262
Wu W, Lu M, Yu L (2011) Expression of carotenogenic genes and astaxanthin production in Xanthophyllomyces dendrorhous as a function of oxygen tension. Z Naturforschung C J Biosci 66:283–286
Xie W, Lv X, Ye L et al (2015) Construction of lycopene-overproducing Saccharomyces cerevisiae by combining directed evolution and metabolic engineering. Metab Eng 30:69–78
Xu F, Yuan Q-P, Zhu Y (2007) Improved production of lycopene and β-carotene by Blakeslea trispora with oxygen-vectors. Process Biochem 42:289–293
Yamamoto K, Hara KY, Morita T et al (2016) Enhancement of astaxanthin production in Xanthophyllomyces dendrorhous by efficient method for the complete deletion of genes. Microb Cell Factories 15:155
Yamane Y, Higashida K, Nakashimada Y et al (1997) Influence of oxygen and glucose on primary metabolism and astaxanthin production by Phaffia rhodozyma in batch and fed-batch cultures: kinetic and stoichiometric analysis. Appl Environ Microbiol 63:4471–4478
Yamano S, Ishii T, Nakagawa M et al (1994) Metabolic engineering for production of β-carotene and lycopene in Saccharomyces cerevisiae. Biosci Biotechnol Biochem 58:1112–1114
Yang Q, Borkovich KA (1999) Mutational activation of a Gαi causes uncontrolled proliferation of aerial hyphae and increased sensitivity to heat and oxidative stress in Neurospora crassa. Genetics 151:107–117
Yen H-W, Chang J-T (2015) Growth of oleaginous Rhodotorula glutinis in an internal-loop airlift bioreactor by using lignocellulosic biomass hydrolysate as the carbon source. J Biosci Bioeng 119:580–584
Youssar L, Avalos J (2007) Genetic basis of the ovc phenotype of Neurospora: identification and analysis of a 77 kb deletion. Curr Genet 51:19–30
Youssar L, Schmidhauser TJ, Avalos J (2005) The Neurospora crassa gene responsible for the cut and ovc phenotypes encodes a protein of the haloacid dehalogenase family. Mol Microbiol 55:828–838
Zahra Bathaie S, Ashrafi M, Azizian M, Tamanoi F (2016) Mevalonate pathway and human cancers. Curr Mol Pharmacol 10:77–85
Zalokar M (1954) Studies on biosynthesis of carotenoids in Neurospora crassa. Arch Biochem Biophys 50:71–80
Zalokar M (1957) Isolation of an acidic pigment in Neurospora. Arch Biochem Biophys 70:568–571
Zalokar M (1955) Biosynthesis of carotenoids in Neurospora. Action spectrum of photoactivation. Arch Biochem Biophys 56:318–325
Zhang Z, Zhang X, Tan T (2014) Lipid and carotenoid production by Rhodotorula glutinis under irradiation/high-temperature and dark/low-temperature cultivation. Bioresour Technol 157:149–153
Zoz L, Carvalho JC, Soccol VT et al (2015) Torularhodin and torulene: bioproduction, properties and prospective applications in food and cosmetics—a review. Braz Arch Biol Technol 58:278–288
Acknowledgments
We thank the Spanish Government (projects BIO2012-39716 and BIO2015-69613-R), and Andalusian Government (project CTS-6638) for funding support. The grants include support from the European Union (European Regional Development Fund [ERDF]).
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Avalos, J., Nordzieke, S., Parra, O., Pardo-Medina, J., Carmen Limón, M. (2017). Carotenoid Production by Filamentous Fungi and Yeasts. In: Sibirny, A. (eds) Biotechnology of Yeasts and Filamentous Fungi. Springer, Cham. https://doi.org/10.1007/978-3-319-58829-2_8
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