Springer Nature is making SARS-CoV-2 and COVID-19 research free. View research | View latest news | Sign up for updates

Melanin biosynthesis in the desert-derived Aureobasidium melanogenum XJ5-1 is controlled mainly by the CWI signal pathway via a transcriptional activator Cmr1


The melanin produced by Aureobasidium melanogenum XJ5-1 obtained from the Taklimakan Desert can play an important role in adaptation of the yeast strain to various stress treatments. It is very important to know how the desert-derived yeast sense, respond and adapt to the harsh environments. However, it is still unclear how melanin is genetically controlled by signaling pathways and transcriptional factors. In this study, it was found that the mitogen-activated protein kinase (MAPK) Slt2 in the cell wall integrity (CWI) signal pathway could regulate activity of the transcriptional activator Swi4; in turn, the Swi4 could control the expression of the CMR1 gene. The melanin-specific transcriptional activator Cmr1 encoded by the CMR1 gene was specifically bound to the promoter with the sequence TTCTCTCCA of the PKS1 gene and strongly stimulated expression of the PKS1 gene and any other genes responsible for melanin biosynthesis, so that a large amount of melanin could be produced by A. melanogenum XJ5-1. Therefore, melanin biosynthesis in the desert-derived A. melanogenum XJ5-1 was controlled mainly by the CWI signal pathway among the cell wall-related signal pathways via a transcriptional activator Cmr and regulation of the melanin biosynthesis in A. melanogenum XJ5-1 was completely different from that of the melanin biosynthesis in any other fungi. This is the first time to show that melanin biosynthesis in the desert-derived A. melanogenum XJ5-1 is controlled mainly by the CWI signal pathway via a transcriptional activator Cmr1. This would provide the fundamentals for further research on the desert-derived yeast to sense, respond and adapt to the harsh environments.

This is a preview of subscription content, log in to check access.

Fig. 1
Fig. 2
Fig. 3
Fig. 4
Fig. 5
Fig. 6
Fig. 7
Fig. 8
Fig. 9


  1. Bahn YS, Kojima K, Cox GM (2006) A unique fungal two-component system regulates stress responses, drug sensitivity, sexual development, and virulence of Cryptococcus neoformans. Mol Biol Cell 17:3122–3135

  2. Bradford MM (1976) A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Anal Biochem 72:248–254

  3. Chen L, Tong Q, Zhang C, Ding K (2019) The transcription factor FgCrz1A is essential for fungal development, virulence, deoxynivalenol biosynthesis and stress responses in Fusarium graminearum. Curr Genet 65:153–166

  4. Chi Z, Wang XX, Ma ZC, Buzdar MA, Chi ZM (2012) The unique role of siderophore in marine-derived Aureobasidium pullulans HN6.2. Biometals 25:219–230

  5. Cho Y, Srivastava A, Ohm RA, Lawrence CB, Wang KH, Grigoriev IV, Sharadchandra P, Marahatta SP (2012) Transcription factor Amr1 induces melanin biosynthesis and suppresses virulence in Alternaria brassicicola. PLoS Pathog 8:e1002974

  6. Eliahu NA, Rose MS, Horwitz BA, Lev S (2007) Melanin biosynthesis in the maize pathogen Cochliobolus heterostrophus depends on two mitogen-activated protein kinases, Chk1 and Mps1, and the transcription factor Cmr1. Eukaryot Cell 6:421–429

  7. Engelberg D, Perlman R, Levitzki A (2014) Transmembrane signaling in Saccharomyces cerevisiae as a model for signaling in metazoans: state of the art after 25 years. Cell Signal 26:2865–2878

  8. Ganesh Kumar C, Mongolla P, Pombala S, Kamle A, Joseph J (2011) Physicochemical characterization and antioxidant activity of melanin from a novel strain of Aspergillus bridgeri ICTF-201. Lett Appl Microbiol 53:350–358

  9. Hellman LM, Fried MG (2007) Electrophoretic mobility shift assay (EMSA) for detecting protein-nucleic acid interactions. Nat Protoc 2:1849–1861

  10. Irniger S, Sarikaya-Bayramo O, Bayram O (2016) Fungal MAP-kinase-mediated regulatory pathways. In: The Mycota III D. Hoffmeister (ed) Biochemistry and molecular biology, 3rd edn. Springer, Cham, pp 97–116

  11. Jiang H, Liu NN, Liu GL, Chi Z, Wang JM, Zhang LL, Chi ZM (2016) Melanin production by a yeast strain XJ5-1 of Aureobasidium melanogenum isolated from the Taklimakan desert and its role in the yeast survival in stress environments. Extremophiles 20:567–577

  12. Jiang H, Liu GL, Chi Z, Wang JM, Zhang LL, Chi ZM (2017) Both a PKS and a PPTase are involved in melanin biosynthesis and regulation of Aureobasidium melanogenum XJ5-1 isolated from the Taklimakan desert. Gene 602:8–15

  13. Jiang H, Liu GL, Chi Z, Hu Z, Chi ZM (2018) Genetics of trehalose biosynthesis in desert-derived Aureobasidium melanogenum and role of trehalose in the adaptation of the yeast to extreme environments. Curr Genet 64:479–491

  14. Jiang H, Chen TJ, Chi Z, Hu Z, Liu GL, Chi ZM (2019) Macromolecular pullulan produced by Aureobasidium melanogenum 13-2 isolated from the Taklimakan desert and its crucial roles in resistance to the stress treatments. Int J Biol Macromol 132:429–436

  15. Laemmli UK (1970) Cleavage of structural proteins during the assembly of the head of bacteriophage T4. Nature 227:680–685

  16. Lenassi M, Gostincar C, Jackman S, Turk M, Sadowski I, Nislow C, Jones S, Inanc B, Gunde-Cimerman N, Plemenitas A (2013) Whole genome duplication and enrichment of metal cation transporters revealed by De Novo genome sequencing of extremely halotolerant black yeast Hortaea werneckii. PLoS ONE 8:e71328

  17. Li C, Lei J, Zhao Y, Xu X, Li S (2015) Effect of saline water irrigation on soil development and plant growth in the Taklimakan Desert Highway shelterbelt. Soil Tillage Res 146:99–107

  18. Liu GL, Wang DS, Wang LF, Zhao SF, Chi ZM (2011a) Mig1 is involved in mycelial formation and expression of the genes encoding extracellular enzymes in Saccharomycopsis fibuligera A11. Fungal Genet Biol 48:904–909

  19. Liu W, Soulie MC, Perrino C, Fillinger S (2011b) The osmosensing signal transduction pathway from Botrytis cinerea regulates cell wall integrity and MAP kinase pathways control melanin biosynthesis with influence of light. Fungal Genet Biol 48:377–387

  20. Ma ZC, Fu WJ, Liu GL, Wang ZP, Chi ZM (2014) High-level pullulan production by Aureobasidium pullulans var. melanogenum P16 isolated from mangrove system. Appl Microbiol Biotechnol 98:4865–4873

  21. Moriwaki A, Kubo E, Arase S, Kihara J (2006) Disruption of SRM1, a mitogen-activated protein kinase gene, affects sensitivity to osmotic and ultraviolet stressors in the phytopathogenic fungus Bipolaris oryzae. FEMS Microbiol Lett 257:253–261

  22. Motoyama T, Ochiai N, Morita M, Iida Y, Usami R, Kudo T (2008) Involvement of putative response regulator genes of the rice blast fungus Magnaporthe oryzae in osmotic stress response, fungicide action, and pathogenicity. Curr Genet 54:185–195

  23. Shelest E (2008) Transcription factors in fungi. FEMS Microbiol Lett 286:145–151

  24. Sterflinger K, Tesei D, Zakharova K (2012) Fungi in hot and cold deserts with particular reference to microcolonial fungi. Fungal Ecol 5:453–462

  25. Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins DG (1997) The Clustal X windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Res 24:4876–4882

  26. Toledo AV, Franco MEE, Lopez SMY, Troncozo MI, Saparrat MCN, Pedro PA, Balatti A (2017) Melanins in fungi: types, localization and putative biological roles. Physiol Mol Plant Pathol 99:2–6

  27. Tsuji G, Kenmochi Y, Takano Y, Sweigard J, Farrall L, Furusawa I, Horino O, Kubo Y (2000) Novel fungal transcriptional activators, Cmr1p of Colletotrichum lagenarium and pig1p of Magnaporthe grisea, contain Cys2His2 zinc finger and Zn(II)2Cys6 binuclear cluster DNA-binding motifs and regulate transcription of melanin biosynthesis genes in a developmentally specific manner. Mol Microbiol 38:940–954

  28. Valiante V, Baldin C, Chortschansky P, Jain R, Thywißen A, Straßburger M, Shelest E, Heinekamp T, Brakhage AA (2016) The Aspergillus fumigatus conidial melanin production is regulated by the bifunctional bHLH DevR and MADS-box RlmA transcription factors. Mol Microbiol 102:321–335

  29. Varghese G, Diwan AM (1983) Simultaneous staining of proteins during polyacrylamide gel electrophoresis in acidic gels by counter migration of Coomassie brilliant blue R-250. Anal Biochem 132:481–483

  30. Wang QQ, Lu Y, Ren ZY, Chi Z, Liu GL, Chi ZM (2017) CreA is directly involved in pullulan biosynthesis and regulation of Aureobasidium melanogenum P16. Curr Genet 63:471–485

  31. White RJ, Sharrocks AD (2010) Coordinated control of the gene expression machinery. Trends Genet 26:214–220

  32. Xu L, Wang M, Tang G, Ma Z, Shao W (2019) The endocytic cargo adaptor complex is required for cell-wall integrity via interacting with the sensor FgWsc2B in Fusarium graminearum. Curr. Genet.

  33. Xue SJ, Chen L, Jiang H, Liu GL, Chi ZM, Hu Z, Chi Z (2019) High pullulan biosynthesis from high concentration of glucose by a hyperosmotic resistant, yeast-like fungal strain isolated from a natural comb-honey. Food Chem 286:123–128

  34. Zhang Z, Lu Y, Chi Z, Liu GL, Jiang H, Hu Z, Chi ZM (2019) Genome editing of different strains of Aureobasidium melanogenum using an efficient Cre/loxp site-specific recombination system. Fungal Biol.

  35. Zhao T, Tian H, Yuxian Xia Y, Jin K (2019) MaPmt4, a protein O-mannosyltransferase, contributes to cell wall integrity, stress tolerance and virulence in Metarhizium acridum. Curr Genet.

  36. Zhou Y, Yang L, Wu M, Chen W, Li G, Zhang J (2017) A single-nucleotide deletion in the transcription factor gene bcsmr1 causes sclerotial-melanogenesis deficiency in Botrytis cinerea. Front Microbiol 8:1–14

Download references


This study was financially supported by National Natural Science Foundation of China (Grant nos. 31800090 and 31770061).

Author information

HJ carried out all the experiments; ZC and GLL designed some of the experiments; ZH wrote some parts of the manuscript; ZMC wrote the whole manuscript.

Correspondence to Zhen-Ming Chi.

Ethics declarations

Conflict of interest

The authors declare they have no conflict of interest.

Additional information

Publisher's Note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Communicated by M. Kupiec.

Electronic supplementary material

Rights and permissions

Reprints and Permissions

About this article

Verify currency and authenticity via CrossMark

Cite this article

Jiang, H., Chi, Z., Liu, G. et al. Melanin biosynthesis in the desert-derived Aureobasidium melanogenum XJ5-1 is controlled mainly by the CWI signal pathway via a transcriptional activator Cmr1. Curr Genet 66, 173–185 (2020).

Download citation


  • Taklimakan desert
  • Harsh environment
  • A. melanogenum
  • Transcriptional activator
  • Melanin biosynthesis