接合菌蓝光受体蛋白研究进展
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摘要
光信号是一种重要的环境因子,尤其是蓝光可以调控真菌的生长发育、生理周期、基因表达及多种代谢活动。接合菌亚门的真菌对环境信号特别是对光刺激感应灵敏,具有明显的趋向性,是研究真菌信号感应的理想材料,而同时具有无性生殖和有性生殖的特点也使之成为研究性别决定的模型。接合菌处于真菌进化的早期,进化地位低,其多拷贝的蓝光受体蛋白在长期进化过程中形成了完善的光受体系统,能够感应不同波长、光强和方向的蓝光和近紫外光,在接合菌的趋光性和类胡萝卜素合成以及无性生殖和有性生殖等光响应过程中发挥承上启下的重要作用。以接合菌中的代表菌株为例,总结了目前在接合菌中报道的蓝光受体蛋白,从基因发现、功能鉴定、光化学特性和调控途径等方面进行了比较分析,进而从3个方面详细阐述了与高等植物不同的接合菌中光受体蛋白调控的特殊生理过程;最后对以后接合菌蓝光受体蛋白的研究重点和难点进行了思考和展望,旨为相关研究者系统了解接合菌蓝光受体蛋白的功能和发展趋势提供线索和参考。
Light is a key environmental signal. Light,most notably blue light,may regulate many aspects of the biology of fungi,including vegetative growth,development of fruiting body,the circadian rhythm,pigment formation and varied metabolic pathways.Zygomycete fungi are generally sensitive to environmental signals,especially light stimulation and have obvious tendency,and have served asmodel organisms to studying light signal transduction in fungi. Zygomycete fungi are at early stage of fungal evolution,i.e.,in low evolution.Their multi-copy blue photo-receptor proteins usually form perfect photoreceptor system during long-term evolution,which may sense blueand near ultraviolet lights at different wavelengths,intensities and directions,thus play a critical role in phototaxis and carotenoid synthesisof zygomyceta,as well as in the photo-response of asexual and sexual reproduction. In this paper,we took the representative strains ofZygomyceta as an example,summarized the blue-light photoreceptor proteins reported by several researchers,and made a comparative analysisfrom the aspects of gene discovery,function identification,photochemical characteristics,regulatory pathways,etc. Moreover,based onthe above three(four)aspects,we described special physiological process mediated and regulated by blue-photoreceptors thatwere entirelydifferent from it in higher plant. In the end,we suggested the future focuses and the confronting difficulties on the studies of blue-photoreceptorsin zygomycetes,expecting for providing direction and reference for the researchers in this field.
Light is a key environmental signal. Light,most notably blue light,may regulate many aspects of the biology of fungi,including vegetative growth,development of fruiting body,the circadian rhythm,pigment formation and varied metabolic pathways.Zygomycete fungi are generally sensitive to environmental signals,especially light stimulation and have obvious tendency,and have served asmodel organisms to studying light signal transduction in fungi. Zygomycete fungi are at early stage of fungal evolution,i.e.,in low evolution.Their multi-copy blue photo-receptor proteins usually form perfect photoreceptor system during long-term evolution,which may sense blueand near ultraviolet lights at different wavelengths,intensities and directions,thus play a critical role in phototaxis and carotenoid synthesisof zygomyceta,as well as in the photo-response of asexual and sexual reproduction. In this paper,we took the representative strains ofZygomyceta as an example,summarized the blue-light photoreceptor proteins reported by several researchers,and made a comparative analysisfrom the aspects of gene discovery,function identification,photochemical characteristics,regulatory pathways,etc. Moreover,based onthe above three(four)aspects,we described special physiological process mediated and regulated by blue-photoreceptors thatwere entirelydifferent from it in higher plant. In the end,we suggested the future focuses and the confronting difficulties on the studies of blue-photoreceptorsin zygomycetes,expecting for providing direction and reference for the researchers in this field.
引文
[1]Tisch D,Schmoll M.Light regulation of metabolic pathways infungi[J].Appl Microbiol Biotechnol,2010,5:1259-1277.
[2]Fuller KK,Loros JJ,Dunlap JC.Fungal photobiology:Visible lightas a signal for stress,space and time[J].Current Genetics,2015,61(3):275-288.
[3]Liu Y,He Q,Cheng P.Photoreception in Neurospora:A tale of twowhite collar proteins[J].Cellular and Molecular Life Sciences,2003,60(10):2131-2138.
[4]Corrochano LM,Garre V.Photobiology in the Zygomycota:Multiplephotoreceptor genes for complex responses to light[J].FungalGenet Biol,2010,47(11):893-899.
[5]Nyilasi I,Acs K,Papp T,et al.Agrobacterium tumefaciens-mediatedtransformation of Mucor circinelloides[J].Folia Microbiologica,2005,50(5):415-420.
[6]Ortiz-Alvarado R,Gonzalez-Hernandez GA,Torres-Guzman JC,et al.Transformation of Mucor circinelloides with autoreplicativevectors containing homologous and heterologous ars elements andthe dominant cbx(r)carboxine-resistance gene[J].CurrentMicrobiology,2006,52(3):178-181.
[7]Ballario P,Vittorioso P,Magrelli A,et al.White collar-1,a centralregulator of blue light responses in Neurospora,is a zinc fingerprotein[J].EMBO Journal,1996,15(7):1650-1657.
[8]Linden H,Macino G.White collar 2,a partner in blue-light signaltransduction,controlling expression of light-regulated genes inNeurospora crassa[J].EMBO Journal,1997,16(1):98-109.
[9]Linden H,Ballario P,Macino G.Blue light regulation in Neurospora crassa[J].Fungal Genet Biol,1997,22(3):141-150.
[10]Smith KM,Sancar G,Dekhang R,et al.Transcription factorsin light and circadian clock signaling networks revealed bygenomewide mapping of direct targets for Neurospora white collarcomplex[J].Eukaryot Cell,2010,9(10):1549-1556.
[11]Casas-Flores S,Rios-Momberg M,Bibbins M,et al.Blr-1 and blr-2,key regulatory elements of photoconidiation and mycelial growth inTrichoderma atroviride[J].Microbiology,2004,150(Pt 11):3561-3569.
[12]Terashima K,Yuki K,Muraguchi H,et al.The dst1 gene involvedin mushroom photomorphogenesis of Coprinus cinereus encodes aputative photoreceptor for blue light[J].Genetics,2005,171(1):101-108.
[13]Kuratani M,Tanaka K,Terashima K,et al.The dst2 gene essentialfor photomorphogenesis of Coprinopsis cinerea encodes a proteinwith a putative FAD-binding-4 domain[J].Fungal Genet Biol,2010,47(2):152-158.
[14]Tisch D,Schmoll M.Targets of light signalling in Trichoderma reesei[J].BMC Genomics,2013,14(1):1-18.
[15]Campuzano V,Galland P,Senger H,et al.Isolation andcharacterization of phototropism mutants of Phycomyces insensitiveto ultraviolet light[J].Current Genetics,1994,26(1):49-53.
[16]Sanz C,Rodriguez-Romero J,Idnurm A,et al.Phycomyces MADBinteracts with MADA to form the primary photoreceptor complex forfungal phototropism[J].Proc Natl Acad Sci USA,2009,106(17):7095-7100.
[17]Idnurm A,Rodriguez-Romero J,Corrochano LM,et al.ThePhycomyces mada gene encodes a blue-light photoreceptor forphototropism and other light responses[J].Proc Natl Acad SciUSA,2006,103(12):4546-4551.
[18]Silva F,Torres-Martinez S,Garre V.Distinct white collar-1 genescontrol specific light responses in Mucor circinelloides[J].MolMicrobiol,2006,61(4):1023-1037.
[19]Ma LJ,Ibrahim AS,Skory C,et al.Genomic analysis of thebasal lineage fungus Rhizopus oryzae reveals a whole-genomeduplication[J].PLo S Genet,2009,5(7):e1000549.
[20]Navarro E,Penaranda A,Hansberg W,et al.A white collar1-like protein mediates opposite regulatory functions in Mucor circinelloides[J].Fungal Genet Biol,2013,52(3):42-52.
[21]Lin C,Todo T.The cryptochromes[J].Genome Biol,2005,6(5):220.
[22]Mei Q,Dvornyk V.Evolutionary history of the photolyase/cryptochrome superfamily in eukaryotes[J].PLo S One,2015,10(9):e0135940.
[23]Froehlich AC,Chen CH,Belden WJ,et al.Genetic and molecularcharacterization of a cryptochrome from the filamentous fungusNeurospora crassa[J].Eukaryot Cell,2010,9(5):738-750.
[24]Guzman-Moreno J,Flores-Martinez A,Brieba LG,et al.TheTrichoderma reesei Cry1 protein is a member of the cryptochrome/photolyase family with 6-4 photoproduct repair activity[J].PLo SOne,2014,9(6):e100625.
[25]Castrillo M,Avalos J.The flavoproteins Cry D and Vvd Acooperate with the white collar protein wcoa in the control ofphotocarotenogenesis in Fusarium fujikuroi[J].PLo S One,2015,10(3):e0119785.
[26]Tagua VG,Pausch M,et al.Fungal cryptochrome with DNA repairactivity reveals an early stage in cryptochrome evolution[J].Proc Natl Acad Sci USA,2015,112(49):15130-15135.
[27]Chaves I,Pokorny R,Byrdin M,et al.The cryptochromes:Bluelight photoreceptors in plants and animals[J].Annu Rev PlantBiol,2011,62:335-364.
[28]Selby CP,Sancar A.A cryptochrome/photolyase class of enzymeswith single-stranded DNA-specific photolyase activity[J].ProcNatl Acad Sci USA,2006,103(47):17696-17700.
[29]Pokorny R,Klar T,et al.Recognition and repair of uv lesions inloop structures of duplex DNA by dash-type cryptochrome[J].Proc Natl Acad Sci USA,2008,105(52):21023-21027.
[30]Huang Y,Baxter R,Smith BS,et al.Crystal structure ofcryptochrome 3 from Arabidopsis thaliana and its implications forphotolyase activity[J].Proc Natl Acad Sci USA,2006,103(47):17701-17706.
[31]Galland P.Ultraviolet killing and photoreactivation of Phycomycesspores[J].Microbiological Research,1996,151(1):9-17.
[32]Shimura M,Ito Y,Ishii C,et al.Characterization of a Neurospora crassa photolyase-deficient mutant generated by repeat inducedpoint mutation of the phr gene[J].Fungal Genet Biol,1999,28(1):12-20.
[33]Galland P.Phototropism of the Phycomyces sporangiophore:Acomparison with higher plants[J].Photochem Photobiol,1990,52(1):233-248.
[34]Cerda-Olmedo E.Phycomyces and the biology of light andcolor[J].FEMS Microbiol Rev,2001,25(5):503-512.
[35]Galland P,Lipson ED.Blue-light reception in Phycomycesphototropism:Evidence for two photosystems operating in lowand high-intensity ranges[J].Proc Natl Acad Sci USA,1987,84(1):104-108.
[36]Martin-Rojas V,Greiner H,Wagner T,et al.Specific tropismcaused by ultraviolet c radiation in Phycomyces[J].Planta,1995,197(1):63-68.
[37]Galland P.Reception of far-ultraviolet light in Phycomyces:Antagonistic interaction with blue and red light[J].Planta,1998,205(2):269-276.
[38]Chen XY,Xiong YQ,Lipson ED.Action spectrum for subliminallight control of adaptation in Phycomyces phototropism[J].Photochem Photobiol,1993,58(3):425-431.
[39]Galland P,Amon S,Senger H,et al.Blue light reception inPhycomyces:Red light sensitization in mad C mutants[J].Botanica Acta,1995,108(4):344-350.
[40]Polaino S,Villalobos-Escobedo JM,et al.A Ras GTPase associatedprotein is involved in the phototropic and circadian photobiologyresponses in fungi[J].Sci Rep,2017,7:44790.
[41]Corrochano LM,Cerdá-Olmedo E.Photomorphogenesis inbehavioural and colour mutants of Phycomyces[J].Journal ofPhotochemistry and Photobiology B,1990,6(3):325-335.
[42]Page RM.Light and the asexual reproduction of Pilobolus[J].Science,1962,138(3546):1238-1245.
[43]Kubo H,Mihara H.Phototropic fluence-response curves forPilobolus crystallinus sporangiophore[J].Planta,1988,174(2):174-179.
[44]Kubo H,Mihara H.Effects of microbeam light on growth andphototropism of Pilobolus crystallinus sporangiophores[J].Mycoscience,1996,37(1):31-34.
[45]Ruiz-Hidalgo MJ,Benito EP,et al.The phytoene dehydrogenasegene of Phycomyces:Regulation of its expression by blue light andvitamin A[J].Mol Gen Genet,1997,253(6):734-744.
[46]Velayos A,Blasco JL,Alvarez MI,et al.Blue-light regulationof phytoene dehydrogenase(car B)gene expression in Mucor circinelloides[J].Planta,2000,210(6):938-946.
[47]Velayos A,Eslava AP,Iturriaga EA.A bifunctional enzyme withlycopene cyclase and phytoene synthase activities is encoded bythe car RP gene of Mucor circinelloides[J].Eur J Biochem,2000,267(17):5509-5519.
[48]Arrach N,Fernandez-Martin R,Cerda-Olmedo E,et al.A singlegene for lycopene cyclase,phytoene synthase,and regulation ofcarotene biosynthesis in Phycomyces[J].Proc Natl Acad SciUSA,2001,98(4):1687-1692.
[49]Rodriguez-Saiz M,et al.Blakeslea trispora genes for carotenebiosynthesis[J].Appl Environ Microbiol,2004,70(9):5589-5594.
[50]Bejarano ER,Avalos J,Lipson ED,et al.Photoinduced accumulation of carotene in Phycomyces[J].Planta,1991,183(1):1-9.
[51]Bergman K,et al.Mutants of Phycomyces with abnormalphototropism[J].Mol Gen Genet,1973,123(1):1-16.
[52]Quiles-Rosillo MD,Ruiz-Vazquez RM,Torres-Martinez S,et al.Light induction of the carotenoid biosynthesis pathway in Blakeslea trispora[J].Fungal Genet Biol,2005,42(2):141-153.
[53]Navarro E,et al.Overexpression of the crg A gene abolishes lightrequirement for carotenoid biosynthesis in Mucor circinelloides[J].Eur J Biochem,2000,267(3):800-807.
[54]Silva F,Navarro E,Penaranda A,et al.A RING-finger protein regulates carotenogenesis via proteolysis-independent ubiquitylation ofa White Collar-1-like activator[J].Mol Microbiol,2008,70(4):1026-1036.
[55]Navarro E,Lorca-Pascual JM,Quiles-Rosillo MD,et al.A negativeregulator of light-inducible carotenogenesis in Mucor circinelloides[J].Mol Genet Genomics,2001,3:463-470.
[56]Corrochano LM,Cerda-Olmedo E.Sex,light and carotenes:Thedevelopment of Phycomyces[J].Trends Genet,1992,8(8):268-274.
[57]Kubo H,Mihara H.Effects of light and temperature onsporangiophore initiation in Pilobolus crystallinus(wiggers)tode[J].Planta,1986,168(3):337-339.
[58]Quiles-Rosillo MD,Torres-Martinez S,Garre V.Cig A,a lightinducible gene involved in vegetative growth in Mucor circinelloidesis regulated by the carotenogenic repressor Crg A[J].FungalGenet Biol,2003,38(1):122-132.
[59]Nicolas FE,Calo S,et al.A RING-finger photocarotenogenic repressor involved in asexual sporulation in Mucor circinelloides[J].FEMS Microbiol Lett,2008,280(1):81-88.
[60]Yamazaki Y,Kataoka H,Miyazakl A,et al.Action spectrafor photoinhibition of sexual development in Phycomyces blakesleeanus[J].Photochem Photobiol,1996,2:389-392.
[61]Lee SC,Heitman J.Sex in the mucoralean fungi[J].Mycoses,2014,57(Suppl 3):18-24.
[62]Tagua VG,Medina HR,Martin-Dominguez R,et al.A gene forcarotene cleavage required for pheromone biosynthesis andcarotene regulation in the fungus Phycomyces blakesleeanus[J].Fungal Genet Biol,2012,49(5):398-404.
[63]Shakya VPS,Idnurm A.The inhibition of mating in Phycomyces blakesleeanus by light is dependent on the MADA-MADB complexthat acts in a sex-specific manner[J].Fungal Genet Biol,2017,101:20-30.
[2]Fuller KK,Loros JJ,Dunlap JC.Fungal photobiology:Visible lightas a signal for stress,space and time[J].Current Genetics,2015,61(3):275-288.
[3]Liu Y,He Q,Cheng P.Photoreception in Neurospora:A tale of twowhite collar proteins[J].Cellular and Molecular Life Sciences,2003,60(10):2131-2138.
[4]Corrochano LM,Garre V.Photobiology in the Zygomycota:Multiplephotoreceptor genes for complex responses to light[J].FungalGenet Biol,2010,47(11):893-899.
[5]Nyilasi I,Acs K,Papp T,et al.Agrobacterium tumefaciens-mediatedtransformation of Mucor circinelloides[J].Folia Microbiologica,2005,50(5):415-420.
[6]Ortiz-Alvarado R,Gonzalez-Hernandez GA,Torres-Guzman JC,et al.Transformation of Mucor circinelloides with autoreplicativevectors containing homologous and heterologous ars elements andthe dominant cbx(r)carboxine-resistance gene[J].CurrentMicrobiology,2006,52(3):178-181.
[7]Ballario P,Vittorioso P,Magrelli A,et al.White collar-1,a centralregulator of blue light responses in Neurospora,is a zinc fingerprotein[J].EMBO Journal,1996,15(7):1650-1657.
[8]Linden H,Macino G.White collar 2,a partner in blue-light signaltransduction,controlling expression of light-regulated genes inNeurospora crassa[J].EMBO Journal,1997,16(1):98-109.
[9]Linden H,Ballario P,Macino G.Blue light regulation in Neurospora crassa[J].Fungal Genet Biol,1997,22(3):141-150.
[10]Smith KM,Sancar G,Dekhang R,et al.Transcription factorsin light and circadian clock signaling networks revealed bygenomewide mapping of direct targets for Neurospora white collarcomplex[J].Eukaryot Cell,2010,9(10):1549-1556.
[11]Casas-Flores S,Rios-Momberg M,Bibbins M,et al.Blr-1 and blr-2,key regulatory elements of photoconidiation and mycelial growth inTrichoderma atroviride[J].Microbiology,2004,150(Pt 11):3561-3569.
[12]Terashima K,Yuki K,Muraguchi H,et al.The dst1 gene involvedin mushroom photomorphogenesis of Coprinus cinereus encodes aputative photoreceptor for blue light[J].Genetics,2005,171(1):101-108.
[13]Kuratani M,Tanaka K,Terashima K,et al.The dst2 gene essentialfor photomorphogenesis of Coprinopsis cinerea encodes a proteinwith a putative FAD-binding-4 domain[J].Fungal Genet Biol,2010,47(2):152-158.
[14]Tisch D,Schmoll M.Targets of light signalling in Trichoderma reesei[J].BMC Genomics,2013,14(1):1-18.
[15]Campuzano V,Galland P,Senger H,et al.Isolation andcharacterization of phototropism mutants of Phycomyces insensitiveto ultraviolet light[J].Current Genetics,1994,26(1):49-53.
[16]Sanz C,Rodriguez-Romero J,Idnurm A,et al.Phycomyces MADBinteracts with MADA to form the primary photoreceptor complex forfungal phototropism[J].Proc Natl Acad Sci USA,2009,106(17):7095-7100.
[17]Idnurm A,Rodriguez-Romero J,Corrochano LM,et al.ThePhycomyces mada gene encodes a blue-light photoreceptor forphototropism and other light responses[J].Proc Natl Acad SciUSA,2006,103(12):4546-4551.
[18]Silva F,Torres-Martinez S,Garre V.Distinct white collar-1 genescontrol specific light responses in Mucor circinelloides[J].MolMicrobiol,2006,61(4):1023-1037.
[19]Ma LJ,Ibrahim AS,Skory C,et al.Genomic analysis of thebasal lineage fungus Rhizopus oryzae reveals a whole-genomeduplication[J].PLo S Genet,2009,5(7):e1000549.
[20]Navarro E,Penaranda A,Hansberg W,et al.A white collar1-like protein mediates opposite regulatory functions in Mucor circinelloides[J].Fungal Genet Biol,2013,52(3):42-52.
[21]Lin C,Todo T.The cryptochromes[J].Genome Biol,2005,6(5):220.
[22]Mei Q,Dvornyk V.Evolutionary history of the photolyase/cryptochrome superfamily in eukaryotes[J].PLo S One,2015,10(9):e0135940.
[23]Froehlich AC,Chen CH,Belden WJ,et al.Genetic and molecularcharacterization of a cryptochrome from the filamentous fungusNeurospora crassa[J].Eukaryot Cell,2010,9(5):738-750.
[24]Guzman-Moreno J,Flores-Martinez A,Brieba LG,et al.TheTrichoderma reesei Cry1 protein is a member of the cryptochrome/photolyase family with 6-4 photoproduct repair activity[J].PLo SOne,2014,9(6):e100625.
[25]Castrillo M,Avalos J.The flavoproteins Cry D and Vvd Acooperate with the white collar protein wcoa in the control ofphotocarotenogenesis in Fusarium fujikuroi[J].PLo S One,2015,10(3):e0119785.
[26]Tagua VG,Pausch M,et al.Fungal cryptochrome with DNA repairactivity reveals an early stage in cryptochrome evolution[J].Proc Natl Acad Sci USA,2015,112(49):15130-15135.
[27]Chaves I,Pokorny R,Byrdin M,et al.The cryptochromes:Bluelight photoreceptors in plants and animals[J].Annu Rev PlantBiol,2011,62:335-364.
[28]Selby CP,Sancar A.A cryptochrome/photolyase class of enzymeswith single-stranded DNA-specific photolyase activity[J].ProcNatl Acad Sci USA,2006,103(47):17696-17700.
[29]Pokorny R,Klar T,et al.Recognition and repair of uv lesions inloop structures of duplex DNA by dash-type cryptochrome[J].Proc Natl Acad Sci USA,2008,105(52):21023-21027.
[30]Huang Y,Baxter R,Smith BS,et al.Crystal structure ofcryptochrome 3 from Arabidopsis thaliana and its implications forphotolyase activity[J].Proc Natl Acad Sci USA,2006,103(47):17701-17706.
[31]Galland P.Ultraviolet killing and photoreactivation of Phycomycesspores[J].Microbiological Research,1996,151(1):9-17.
[32]Shimura M,Ito Y,Ishii C,et al.Characterization of a Neurospora crassa photolyase-deficient mutant generated by repeat inducedpoint mutation of the phr gene[J].Fungal Genet Biol,1999,28(1):12-20.
[33]Galland P.Phototropism of the Phycomyces sporangiophore:Acomparison with higher plants[J].Photochem Photobiol,1990,52(1):233-248.
[34]Cerda-Olmedo E.Phycomyces and the biology of light andcolor[J].FEMS Microbiol Rev,2001,25(5):503-512.
[35]Galland P,Lipson ED.Blue-light reception in Phycomycesphototropism:Evidence for two photosystems operating in lowand high-intensity ranges[J].Proc Natl Acad Sci USA,1987,84(1):104-108.
[36]Martin-Rojas V,Greiner H,Wagner T,et al.Specific tropismcaused by ultraviolet c radiation in Phycomyces[J].Planta,1995,197(1):63-68.
[37]Galland P.Reception of far-ultraviolet light in Phycomyces:Antagonistic interaction with blue and red light[J].Planta,1998,205(2):269-276.
[38]Chen XY,Xiong YQ,Lipson ED.Action spectrum for subliminallight control of adaptation in Phycomyces phototropism[J].Photochem Photobiol,1993,58(3):425-431.
[39]Galland P,Amon S,Senger H,et al.Blue light reception inPhycomyces:Red light sensitization in mad C mutants[J].Botanica Acta,1995,108(4):344-350.
[40]Polaino S,Villalobos-Escobedo JM,et al.A Ras GTPase associatedprotein is involved in the phototropic and circadian photobiologyresponses in fungi[J].Sci Rep,2017,7:44790.
[41]Corrochano LM,Cerdá-Olmedo E.Photomorphogenesis inbehavioural and colour mutants of Phycomyces[J].Journal ofPhotochemistry and Photobiology B,1990,6(3):325-335.
[42]Page RM.Light and the asexual reproduction of Pilobolus[J].Science,1962,138(3546):1238-1245.
[43]Kubo H,Mihara H.Phototropic fluence-response curves forPilobolus crystallinus sporangiophore[J].Planta,1988,174(2):174-179.
[44]Kubo H,Mihara H.Effects of microbeam light on growth andphototropism of Pilobolus crystallinus sporangiophores[J].Mycoscience,1996,37(1):31-34.
[45]Ruiz-Hidalgo MJ,Benito EP,et al.The phytoene dehydrogenasegene of Phycomyces:Regulation of its expression by blue light andvitamin A[J].Mol Gen Genet,1997,253(6):734-744.
[46]Velayos A,Blasco JL,Alvarez MI,et al.Blue-light regulationof phytoene dehydrogenase(car B)gene expression in Mucor circinelloides[J].Planta,2000,210(6):938-946.
[47]Velayos A,Eslava AP,Iturriaga EA.A bifunctional enzyme withlycopene cyclase and phytoene synthase activities is encoded bythe car RP gene of Mucor circinelloides[J].Eur J Biochem,2000,267(17):5509-5519.
[48]Arrach N,Fernandez-Martin R,Cerda-Olmedo E,et al.A singlegene for lycopene cyclase,phytoene synthase,and regulation ofcarotene biosynthesis in Phycomyces[J].Proc Natl Acad SciUSA,2001,98(4):1687-1692.
[49]Rodriguez-Saiz M,et al.Blakeslea trispora genes for carotenebiosynthesis[J].Appl Environ Microbiol,2004,70(9):5589-5594.
[50]Bejarano ER,Avalos J,Lipson ED,et al.Photoinduced accumulation of carotene in Phycomyces[J].Planta,1991,183(1):1-9.
[51]Bergman K,et al.Mutants of Phycomyces with abnormalphototropism[J].Mol Gen Genet,1973,123(1):1-16.
[52]Quiles-Rosillo MD,Ruiz-Vazquez RM,Torres-Martinez S,et al.Light induction of the carotenoid biosynthesis pathway in Blakeslea trispora[J].Fungal Genet Biol,2005,42(2):141-153.
[53]Navarro E,et al.Overexpression of the crg A gene abolishes lightrequirement for carotenoid biosynthesis in Mucor circinelloides[J].Eur J Biochem,2000,267(3):800-807.
[54]Silva F,Navarro E,Penaranda A,et al.A RING-finger protein regulates carotenogenesis via proteolysis-independent ubiquitylation ofa White Collar-1-like activator[J].Mol Microbiol,2008,70(4):1026-1036.
[55]Navarro E,Lorca-Pascual JM,Quiles-Rosillo MD,et al.A negativeregulator of light-inducible carotenogenesis in Mucor circinelloides[J].Mol Genet Genomics,2001,3:463-470.
[56]Corrochano LM,Cerda-Olmedo E.Sex,light and carotenes:Thedevelopment of Phycomyces[J].Trends Genet,1992,8(8):268-274.
[57]Kubo H,Mihara H.Effects of light and temperature onsporangiophore initiation in Pilobolus crystallinus(wiggers)tode[J].Planta,1986,168(3):337-339.
[58]Quiles-Rosillo MD,Torres-Martinez S,Garre V.Cig A,a lightinducible gene involved in vegetative growth in Mucor circinelloidesis regulated by the carotenogenic repressor Crg A[J].FungalGenet Biol,2003,38(1):122-132.
[59]Nicolas FE,Calo S,et al.A RING-finger photocarotenogenic repressor involved in asexual sporulation in Mucor circinelloides[J].FEMS Microbiol Lett,2008,280(1):81-88.
[60]Yamazaki Y,Kataoka H,Miyazakl A,et al.Action spectrafor photoinhibition of sexual development in Phycomyces blakesleeanus[J].Photochem Photobiol,1996,2:389-392.
[61]Lee SC,Heitman J.Sex in the mucoralean fungi[J].Mycoses,2014,57(Suppl 3):18-24.
[62]Tagua VG,Medina HR,Martin-Dominguez R,et al.A gene forcarotene cleavage required for pheromone biosynthesis andcarotene regulation in the fungus Phycomyces blakesleeanus[J].Fungal Genet Biol,2012,49(5):398-404.
[63]Shakya VPS,Idnurm A.The inhibition of mating in Phycomyces blakesleeanus by light is dependent on the MADA-MADB complexthat acts in a sex-specific manner[J].Fungal Genet Biol,2017,101:20-30.
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