臺灣博碩士論文加值系統

冬蟲夏草為真菌中珍貴的藥用真菌，屬於子囊菌亞門，核菌綱，球殼菌目，麥角菌科，蟲草屬。於蟲草屬(Cordyceps)中又有282種。其中蛹蟲草(Cordyceps militaris)為本屬之代表種，並於1950年中發現蟲草素(cordycepin)，並曾利用在抗腫瘤的治療。近年又將蟲草素利用於抗病毒的研究也發現有其價值。故本論文希望藉由育種的技術，找到具有高產量蟲草素的品系。為找出含有高產量的品系，並了解蛹蟲草的交配型態，期能首先將子座上產生的有性孢子收集。使用PDA(Potato Dextrose Agar)分別進行單一孢子培養，發現外觀菌絲的顏色可分為三類,鮮黃色,淡黃色及白色。也觀察到其生長的情形似乎也與菌絲的顏色有關聯性，其中白色生長速度最慢，鮮黃色最快。從子座上取下的菌絲生長速度最快。經玻片培養並以HCl-Giemsa stain後觀察細胞核的型態,從染色的型態學上觀察並無法區分，觀察到均為單一的細胞核。過去研究子囊菌均無法由交配反應判斷性別因子。本論文使用流式細胞儀，單染Propidium Iodide (PI) 以及Acridine
Orange(AO)藉由DNA含量不同，來確定單倍體或是二倍體；再經由不同單孢的交配，收集交配的菌絲，使用流式細胞儀分析是否成為二倍體，另外使用鏈鎖聚合酶反應，經由放大MAT基因確定性別的控制因子為2個基因，分別為MAT1-1-1及MAT1-2-1。因此證明了C. militaris為heterothallism的性別遺傳模式。C. militaris液態培養中蟲草素的含量，經層液相色層分析法(TLC)分析，結果顯示菌液中確實有蟲草素的成分。並發現在不同的天數產生蟲草素的產量有差異，進ㄧ步使用High Performance Liquid Chromatography (HPLC)分析菌液中蟲草素的差異，顯示單性菌株蟲草素之產量高於交配後之兩性菌株，因此，在選種上可採用單子囊孢子菌株，進行篩選。

Many species in the genus of Cordyceps were used as traditional medicine for several hundred years. Among them, Cordyceps militaris was recently developed as functional food in Taiwan. C. militaris was characterizes with the production of cordycepin that was found to be functional in antitumor and antivirus. The purpose of the present study is to breed a high cordycepin production strain through mating of mono-ascosporoes cultures and the mating types of C. militaris were detected in the same time. Ascospores discharged from cultivated stroma were also isolated individually from the surface of agar plate and grown in PDA slants at 25℃. The pigmentation of the monosporous strains can be separated into three types as white, light-yellow and deep-yellow. The growth rate were also different among these types, that indicated the meiotic segregation of the strains. HCl-Giemsa stain revealed to the hypha of slide culture revealed that all the strains before and after a all possible mating were single nucleated in hypha and phialoconidia. Flow cytometric analysis on the conidia suspension stained by acridine orange indicated that the mating occurred for some mating and the intensity of DNA demonstrated in two form which were suggested to be diploid and haploid strains. To confirm the mating type of C. militaris, primer of MAT1-1-1and MAT1-2-1 were design for PCR. TLC and HPLC were employed to analysis the production of cordycepin of the strains before and after mating of the monoascosporous strains. However, all the mated strain produced less cordycepin in the liquid culture. This result indicated that high cordycepin production strain could be screened from monosporous isolation.

目錄

摘要...............................................................
壹.緒論............................................................1
ㄧ.Cordycepes..................................................1
二.真菌性別遺傳模式之研究......................................5
三.確定子囊菌性別之方法.......................................11
四.蛹蟲草液態發酵的影響因子...................................12
五.蟲草素.....................................................15
六.真菌二次代謝物之合成調控...................................17
九.研究動機...................................................19
貳.實驗架構.......................................................20
参.實驗材料與方法.................................................21
ㄧ.實驗材料...................................................21
二.實驗方法...................................................24
肆.結果...........................................................30
伍.討論...........................................................37
陸.結論...........................................................43
柒.參考文獻.......................................................44
圖................................................................57





圖表目錄

Fig. 1 蟲草素及腺苷酸構式………………………..…………..….. 57
Fig. 2 菌種照片……………………………..………………..…… 58
Fig. 3 單孢品系………………………………………..…………..… 59
Fig. 4 單孢品系生長情形圖………………………..……….…….. 60
Fig. 5 交配反應……………………………………………………..… 61
Fig. 6A；6B 單胞品系染色圖………………………..…………..….. 62
Fig. 7交配後之菌絲染色圖…………………………………..…….… 63
Fig. 8 S. cereviceae麵包酵母圖…………………………………... 64
Fig. 9 S. cereviceae利用以PI染色後使用流式細胞儀分析…... 65
Fig. 10 C. militaris之分生孢子以PI染色使用流式細胞儀分析... 66
Fig. 11 C. militaris之分生孢子以AO染色使用流式細胞儀分析.. 67
Fig. 12 有性單孢品系經廣用引子増幅之PCR圖…………….….. 68
Fig. 13 C. militaris單孢及二倍體菌株之MAT基因圖…………… 69
Fig. 14 C. militaris與C. sinensis之MAT基因圖…………………. 70
Fig. 15 AT1-1-1基因定序圖………………………………………….. 71
Fig. 16 AT1-21基因定序圖…………………………………………… 72
Fig. 17 利用固態培養基培養C. militaris………………………….. 73
Fig. 18 單孢品系產生之子座……………………………………….. 74
Fig. 19A；19B 交配後菌種產生之子座……………………………. 75
Fig. 20A；20B 無性孢子…………………………………………….. 76
Fig. 21A；21B 無性孢子品系………………………………………. 77
Fig. 22 C. militaris無性單孢品系之MAT基因圖（一）………… 78
Fig. 23 C. militaris無性單孢品系之MAT基因圖（二）………… 79
Fig. 24 子座頂端……………………………………………………… 80
Fig. 25 子座經廣用引子増幅之PCR圖……………………………. 81
Fig. 26 C.militaris 子座之MAT 基因圖…………………………… 82
Fig. 27 子座性別因子示意圖……………………………………….. 83
Fig. 28 C.militaris 之生活史………………………………………… 84
Fig. 29 TLC分析蟲草素………………………………………………. 85
Fig. 30 蟲草素標準品檢量線……………………………………….. 86
Fig. 31 HPLC分析菌液中之蟲草素…………………………………. 87
Fig. 32 液態培養天數之分析圖…………………………………….. 88
Fig. 33 單孢品系中蟲草素之含量含量分析圖……………………. 89
Fig. 34 交配前後蟲草素含量比較圖（一）……………………….. 90
Fig. 35 交配前後蟲草素含量比較圖（二）……………………….. 91
Fig. 36 交配前後蟲草素含量比較圖（三）……………………….. 92
Table.1 單孢品系之遺傳性別基因…………………………………. 93

蘇慶華. (1985). 新屬植生蟲草形態遺傳及生化之研究。國立台灣大學農業化學研究所。
Ahn, Y. J., S. J. Park, et al. (2000). "Cordycepin: selective growth inhibitor derived from liquid culture of Cordyceps militaris against Clostridium spp." J Agric Food Chem 48(7): 2744-8.
Asami, K., K. Takahashi, et al. (2000). "Progression of cell cycle monitored by dielectric spectroscopy and flow-cytometric analysis of DNA content." Yeast 16(15): 1359-63.
Aspegren, A. and E. Bridge (2002). "Release of snRNP and RNA from transcription sites in adenovirus-infected cells." Exp Cell Res 276(2): 273-83.
Astell, C. R., L. Ahlstrom-Jonasson, et al. (1981). "The sequence of the DNAs coding for the mating-type loci of Saccharomyces cerevisiae." Cell 27(1 Pt 2): 15-23.
Bistis, G. N. (1998). "Physiological Heterothallism and Sexuality in Euascomycetes: A Partial History." Fungal Genet Biol 23(3): 213-22.
Calado, A. and M. Carmo-Fonseca (2000). "Localization of poly(A)-binding protein 2 (PABP2) in nuclear speckles is independent of import into the nucleus and requires binding to poly(A) RNA." J Cell Sci 113 ( Pt 12): 2309-18.
Carmeliet, P. and R. K. Jain (2000). "Angiogenesis in cancer and other diseases." Nature 407(6801): 249-57.
Carter, B. L., A. T. Bull, et al. (1971). "Relationship between energy substrate utilization and specific growth rate in Aspergillus nidulans." J Bacteriol 108(1): 309-13.
Cases-Gonzalez, C. E., M. Gutierrez-Rivas, et al. (2000). "Coupling ribose selection to fidelity of DNA synthesis. The role of Tyr-115 of human immunodeficiency virus type 1 reverse transcriptase." J Biol Chem 275(26): 19759-67.
Chassy, B. M. and R. J. Suhadolnik (1969). "Nucleoside antibiotics. IV. Metabolic fate of adenosine and cordycepin by Cordyceps militaris during cordycepin biosynthesis." Biochim Biophys Acta 182(2): 307-15.
Choi, S. B., C. H. Park, et al. (2004). "Improvement of insulin resistance and insulin secretion by water extracts of Cordyceps militaris, Phellinus linteus, and Paecilomyces tenuipes in 90% pancreatectomized rats." Biosci Biotechnol Biochem 68(11): 2257-64.
Christiansen, S. K., O. C. Yoder, et al. (1997). "Efficient Cloning of Ascomycete Mating Type Genes by PCR Amplification of the Conserved MAT HMG Box." Fungal Genet Biol 21(1): 118-30.
Chung, E. J., K. Choi, et al. (2003). "Analysis of cell cycle gene expression responding to acetoxyscirpendiol isolated from Paecilomyces tenuipes." Biol Pharm Bull 26(1): 32-6.
Coppin, E. and R. Debuchy (2000). "Co-expression of the mating-type genes involved in internuclear recognition is lethal in Podospora anserina." Genetics 155(2): 657-69.
Coppin, E., R. Debuchy, et al. (1997). "Mating types and sexual development in filamentous ascomycetes." Microbiol Mol Biol Rev 61(4): 411-28.
Cory, J. G., R. J. Suhadolnik, et al. (1965). "Incorporation of cordycepin (3'-deoxyadenosine) into ribonucleic acid and deoxyribonucleic acid of human tumor cells." Biochim Biophys Acta 103(4): 646-53.
Covert, S. F., A. Briley, et al. (1999). "Partial MAT-2 gene structure and the influence of temperature on mating success in Gibberella circinata." Fungal Genet Biol 28(1): 43-54.
Cunningham, K. G., W. Manson, et al. (1950). "Cordycepin, a metabolic product isolated from cultures of Cordyceps militaris (Linn.) Link." Nature 166(4231): 949.
Darnell, J. E., L. Philipson, et al. (1971). "Polyadenylic acid sequences: role in conversion of nuclear RNA into messenger RNA." Science 174(8): 507-10.
Dixon, N. M. and D. B. Kell (1989). "The inhibition by CO2 of the growth and metabolism of micro-organisms." J Appl Bacteriol 67(2): 109-36.
Dolan, J. W. and S. Fields (1991). "Cell-type-specific transcription in yeast." Biochim Biophys Acta 1088(2): 155-69.
Dyer, P. S., P. A. Furneaux, et al. (2001). "A multiplex PCR test for determination of mating type applied to the plant pathogens Tapesia yallundae and Tapesia acuformis." Fungal Genet Biol 33(3): 173-80.
Ferreira, A. V., Z. An, et al. (1998). "Characterization of mat A-2, mat A-3 and deltamatA mating-type mutants of Neurospora crassa." Genetics 148(3): 1069-79.
Fiume, L., I. Nardi, et al. (1972). "Effects of cordycepin on morphology and RNA synthesis of amphibian lampbrush chromosomes." Exp Cell Res 75(1): 11-4.
Frederiksen, S., H. Malling, et al. (1965). "Isolation of 3'-Deoxyadenosine (Cordycepin) from the Liquid Medium of Cordyceps Militaris (L. Ex Fr.) Link." Biochim Biophys Acta 95: 189-93.
Glass, N. L., J. Grotelueschen, et al. (1990). "Neurospora crassa A mating-type region." Proc Natl Acad Sci U S A 87(13): 4912-6.
Glass, N. L. and M. L. Smith (1994). "Structure and function of a mating-type gene from the homothallic species Neurospora africana." Mol Gen Genet 244(4): 401-9.
Glass, N. L., S. J. Vollmer, et al. (1988). "DNAs of the two mating-type alleles of Neurospora crassa are highly dissimilar." Science 241(4865): 570-3.
Haase, S. B. and D. J. Lew (1997). "Flow cytometric analysis of DNA content in budding yeast." Methods Enzymol 283: 322-32.
Haase, S. B. and S. I. Reed (2002). "Improved flow cytometric analysis of the budding yeast cell cycle." Cell Cycle 1(2): 132-6.
Herskowitz, I. (1989). "A regulatory hierarchy for cell specialization in yeast." Nature 342(6251): 749-57.
Hiscock, S. J. and U. Kues (1999). "Cellular and molecular mechanisms of sexual incompatibility in plants and fungi." Int Rev Cytol 193: 165-295.
Hotz, M. A., J. Gong, et al. (1994). "Flow cytometric detection of apoptosis: comparison of the assays of in situ DNA degradation and chromatin changes." Cytometry 15(3): 237-44.
Huang, L. F., Y. Z. Liang, et al. (2003). "Simultaneous separation and determination of active components in Cordyceps sinensis and Cordyceps militarris by LC/ESI-MS." J Pharm Biomed Anal 33(5): 1155-62.
Isegawa, Y., J. Sheng, et al. (1992). "Selective amplification of cDNA sequence from total RNA by cassette-ligation mediated polymerase chain reaction (PCR): application to sequencing 6.5 kb genome segment of hantavirus strain B-1." Mol Cell Probes 6(6): 467-75.
Jagger, D. V., N. M. Kredich, et al. (1961). "Inhibition of Ehrlich mouse ascites tumor growth by cordycepin." Cancer Res 21: 216-20.
Johnson, A. D. (1995). "Molecular mechanisms of cell-type determination in budding yeast." Curr Opin Genet Dev 5(5): 552-8.
Kaczka, E. A., N. R. Trenner, et al. (1964). "Identification of cordycepin, a metabolite of Cordyceps militaris, as 3'-deoxyadenosine." Biochem Biophys Res Commun 14: 456-7.
Kerenyi, Z., K. Zeller, et al. (1999). "Molecular standardization of mating type terminology in the Gibberella fujikuroi species complex." Appl Environ Microbiol 65(9): 4071-6.
Kim, G. Y., W. S. Ko, et al. (2006). "Water extract of Cordyceps militaris enhances maturation of murine bone marrow-derived dendritic cells in vitro." Biol Pharm Bull 29(2): 354-60.
Kim, J. R., S. H. Yeon, et al. (2002). "Larvicidal activity against Plutella xylostella of cordycepin from the fruiting body of Cordyceps militaris." Pest Manag Sci 58(7): 713-7.
Kim, S. W., H. J. Hwang, et al. (2003). "Optimization of submerged culture process for the production of mycelial biomass and exo-polysaccharides by Cordyceps militaris C738." J Appl Microbiol 94(1): 120-6.
Kim, S. W., C. P. Xu, et al. (2003). "Production and characterization of exopolysaccharides from an enthomopathogenic fungus Cordyceps militaris NG3." Biotechnol Prog 19(2): 428-35.
Kodama, E. N., R. P. McCaffrey, et al. (2000). "Antileukemic activity and mechanism of action of cordycepin against terminal deoxynucleotidyl transferase-positive (TdT+) leukemic cells." Biochem Pharmacol 59(3): 273-81.
Kornblihtt, A. R., M. de la Mata, et al. (2004). "Multiple links between transcription and splicing." Rna 10(10): 1489-98.
Kronstad, J. W. and C. Staben (1997). "Mating type in filamentous fungi." Annu Rev Genet 31: 245-76.
Kubicek, C. P. and M. Rohr (1980). "Regulation of citrate synthase from the citric acid-accumulating fungus, Aspergillus niger." Biochim Biophys Acta 615(2): 449-57.
Kuo, H. C., Y. L. Su, et al. (2005). "Identification of Chinese medicinal fungus Cordyceps sinensis by PCR-single-stranded conformation polymorphism and phylogenetic relationship." J Agric Food Chem 53(10): 3963-8.
Lallas, G. C., N. Courtis, et al. (2004). "K562 cell sensitization to 5-fluorouracil- or interferon-alpha-induced apoptosis via cordycepin (3'-deoxyadenosine): fine control of cell apoptosis via poly(A) polymerase upregulation." Int J Biol Markers 19(1): 58-66.
Laudet, V., D. Stehelin, et al. (1993). "Ancestry and diversity of the HMG box superfamily." Nucleic Acids Res 21(10): 2493-501.
Lee, H., Y. J. Kim, et al. (2006). "Induction of apoptosis by Cordyceps militaris through activation of caspase-3 in leukemia HL-60 cells." Biol Pharm Bull 29(4): 670-4.
Leuchtmann, A. (1992). "Systematics, distribution, and host specificity of grass endophytes." Nat Toxins 1(3): 150-62.
Luo, G., R. K. Hamatake, et al. (2000). "De novo initiation of RNA synthesis by the RNA-dependent RNA polymerase (NS5B) of hepatitis C virus." J Virol 74(2): 851-63.
Mainelis, G., R. Gorny, et al. (2005). "Rapid counting of liquid-borne microorganisms by light scattering spectrometry." Ann Agric Environ Med 12(1): 141-8.
Mao, X. B. and J. J. Zhong (2004). "Hyperproduction of cordycepin by two-stage dissolved oxygen control in submerged cultivation of medicinal mushroom Cordyceps militaris in bioreactors." Biotechnol Prog 20(5): 1408-13.
Mazumder, A., M. Gupta, et al. (1994). "Methylphosphonodiester substitution near the conserved CA dinucleotide in the HIV LTR alters both extent of 3'-processing and choice of nucleophile by HIV-1 integrase." Nucleic Acids Res 22(21): 4441-8.
McGuire, I. C., R. E. Marra, et al. (2001). "Analysis of mating-type genes in the chestnut blight fungus, Cryphonectria parasitica." Fungal Genet Biol 34(2): 131-44.
Melling, J., F. C. Belton, et al. (1972). "Production of pure cordycepin (3'-deoxyadenosine) from Cordyceps militaris." J Pharm Pharmacol 24: Suppl:125P.
Metzenberg, R. L. and N. L. Glass (1990). "Mating type and mating strategies in Neurospora." Bioessays 12(2): 53-9.
Molenaar, C., A. Abdulle, et al. (2004). "Poly(A)+ RNAs roam the cell nucleus and pass through speckle domains in transcriptionally active and inactive cells." J Cell Biol 165(2): 191-202.
Montefiori, D. C., R. W. Sobol, Jr., et al. (1989). "Phosphorothioate and cordycepin analogues of 2',5'-oligoadenylate: inhibition of human immunodeficiency virus type 1 reverse transcriptase and infection in vitro." Proc Natl Acad Sci U S A 86(18): 7191-4.
Muller, W. E., B. E. Weiler, et al. (1991). "Cordycepin analogues of 2',5'-oligoadenylate inhibit human immunodeficiency virus infection via inhibition of reverse transcriptase." Biochemistry 30(8): 2027-33.
Oh, G. S., K. H. Hong, et al. (2001). "4-Acetyl-12,13-epoxyl-9-trichothecene-3,15-diol isolated from the fruiting bodies of Isariajaponica Yasuda induces apoptosis of human leukemia cells (HL-60)." Biol Pharm Bull 24(7): 785-9.
Park, J. P., S. W. Kim, et al. (2001). "Optimization of submerged culture conditions for the mycelial growth and exo-biopolymer production by Cordyceps militaris." Lett Appl Microbiol 33(1): 76-81.
Park, J. P., Y. M. Kim, et al. (2002). "Effect of agitation intensity on the exo-biopolymer production and mycelial morphology in Cordyceps militaris." Lett Appl Microbiol 34(6): 433-8.
Penman, S., M. Rosbash, et al. (1970). "Messenger and heterogeneous nuclear RNA in HeLa cells: differential inhibition by cordycepin." Proc Natl Acad Sci U S A 67(4): 1878-85.
Philipson, L., R. Wall, et al. (1971). "Addition of polyadenylate sequences to virus-specific RNA during adenovirus replication." Proc Natl Acad Sci U S A 68(11): 2806-9.
Philley, M. L. and C. Staben (1994). "Functional analyses of the Neurospora crassa MT a-1 mating type polypeptide." Genetics 137(3): 715-22.
Piga, A., K. Ganeshaguru, et al. (1989). "Selective toxicity of purine nucleosides to human leukaemic cells." Adv Exp Med Biol 253B: 291-8.
Plagemann, P. G. (1971). "Effects of 3'deoxyadenosine (cordycepin) and 2'deoxyadenosine on nucleoside transport, macromolecular synthesis, and replication of cultured Novikoff hepatoma cells." Arch Biochem Biophys 144(1): 401-12.
Poggeler, S. (1999). "Phylogenetic relationships between mating-type sequences from homothallic and heterothallic ascomycetes." Curr Genet 36(4): 222-31.
Poggeler, S. and U. Kuck (2000). "Comparative analysis of the mating-type loci from Neurospora crassa and Sordaria macrospora: identification of novel transcribed ORFs." Mol Gen Genet 263(2): 292-301.
Poggeler, S., S. Risch, et al. (1997). "Mating-type genes from the homothallic fungus Sordaria macrospora are functionally expressed in a heterothallic ascomycete." Genetics 147(2): 567-80.
Rosowsky, A., H. Lazarus, et al. (1976). "Nucleosides. 1. 9-(3'-Alkyl-3'-deoxy-beta-D-ribofuranosyl)adenines as lipophilic analogues of cordycepin. Synthesis and preliminary biological studies." J Med Chem 19(11): 1265-70.
Rottman, F. and A. J. Guarino (1964). "The Inhibition of Purine Biosynthesis De Novo in Bacillus Subtilis by Cordycepin." Biochim Biophys Acta 80: 640-7.
Rottman, F. and A. J. Guarino (1964). "Studies on the Inhibition of Bacillus Subtilis Growth by Cordycepin." Biochim Biophys Acta 80: 632-9.
Rudensky, B., E. Broidie, et al. (2005). "Rapid flow-cytometric susceptibility testing of Candida species." J Antimicrob Chemother 55(1): 106-9.
Sanchez-Murillo, R. I., M. de la Torre-Martinez, et al. (2004). "Light-regulated asexual reproduction in Paecilomyces fumosoroseus." Microbiology 150(Pt 2): 311-9.
Shiu, P. K. and N. L. Glass (2000). "Cell and nuclear recognition mechanisms mediated by mating type in filamentous ascomycetes." Curr Opin Microbiol 3(2): 183-8.
Siev, M., R. Weinberg, et al. (1969). "The selective interruption of nucleolar RNA synthesis in HeLa cells by cordycepin." J Cell Biol 41(2): 510-20.
Singh, G. and A. M. Ashby (1998). "Cloning of the mating type loci from Pyrenopeziza brassicae reveals the presence of a novel mating type gene within a discomycete MAT 1-2 locus encoding a putative metallothionein-like protein." Mol Microbiol 30(4): 799-806.
Staben, C. and C. Yanofsky (1990). "Neurospora crassa a mating-type region." Proc Natl Acad Sci U S A 87(13): 4917-21.
Stein, P., P. Svoboda, et al. (2003). "RNAi: mammalian oocytes do it without RNA-dependent RNA polymerase." Rna 9(2): 187-92.
Tabak, H. H. and W. B. Cooke (1968). "Growth and metabolism of fungi in an atmosphere of nitrogen." Mycologia 60(1): 115-40.
Taylor, J., D. Jacobson, et al. (1999). "THE EVOLUTION OF ASEXUAL FUNGI: Reproduction, Speciation and Classification." Annu Rev Phytopathol 37: 197-246.
Thomadaki, H., C. M. Tsiapalis, et al. (2005). "Polyadenylate polymerase modulations in human epithelioid cervix and breast cancer cell lines, treated with etoposide or cordycepin, follow cell cycle rather than apoptosis induction." Biol Chem 386(5): 471-80.
Tosetti, F., N. Ferrari, et al. (2002). "Angioprevention': angiogenesis is a common and key target for cancer chemopreventive agents." Faseb J 16(1): 2-14.
Turgeon, B. G. (1998). "Application of mating type gene technology to problems in fungal biology." Annu Rev Phytopathol 36: 115-37.
Turgeon, B. G., H. Bohlmann, et al. (1993). "Cloning and analysis of the mating type genes from Cochliobolus heterostrophus." Mol Gen Genet 238(1-2): 270-84.
Turgeon, B. G. and O. C. Yoder (2000). "Proposed nomenclature for mating type genes of filamentous ascomycetes." Fungal Genet Biol 31(1): 1-5.
Weill, L., D. Louis, et al. (2004). "Selection and evolution of NTP-specific aptamers." Nucleic Acids Res 32(17): 5045-58.
Witthuhn, R. C., T. C. Harrington, et al. (2000). "Deletion of the MAT-2 mating-type gene during uni-directional mating-type switching in Ceratocystis." Curr Genet 38(1): 48-52.
Won, S. Y. and E. H. Park (2005). "Anti-inflammatory and related pharmacological activities of cultured mycelia and fruiting bodies of Cordyceps militaris." J Ethnopharmacol 96(3): 555-61.
Wu, A. M., R. C. Ting, et al. (1972). "Cordycepin inhibits induction of murine leukovirus production by 5-iodo-2'-deoxyuridine." Proc Natl Acad Sci U S A 69(12): 3820-4.
Yokoyama, E., K. Yamagishi, et al. (2003). "Structures of the mating-type loci of Cordyceps takaomontana." Appl Environ Microbiol 69(8): 5019-22.
Yokoyama, E., K. Yamagishi, et al. (2004). "Development of a PCR-based mating-type assay for Clavicipitaceae." FEMS Microbiol Lett 237(2): 205-12.
Yokoyama, E., K. Yamagishi, et al. (2005). "Heterothallism in Cordyceps takaomontana." FEMS Microbiol Lett 250(1): 145-50.
Yoo, H. S., J. W. Shin, et al. (2004). "Effects of Cordyceps militaris extract on angiogenesis and tumor growth." Acta Pharmacol Sin 25(5): 657-65.
Yu, H. M., B. S. Wang, et al. (2006). "Comparison of protective effects between cultured Cordyceps militaris and natural Cordyceps sinensis against oxidative damage." J Agric Food Chem 54(8): 3132-8.
Yun, S. H., T. Arie, et al. (2000). "Molecular organization of mating type loci in heterothallic, homothallic, and asexual Gibberella/Fusarium species." Fungal Genet Biol 31(1): 7-20.
Zhang, G., Y. Huang, et al. (2006). "Hypoglycemic activity of the fungi Cordyceps militaris, Cordyceps sinensis, Tricholoma mongolicum, and Omphalia lapidescens in streptozotocin-induced diabetic rats." Appl Microbiol Biotechnol.
Zhou, X., C. U. Meyer, et al. (2002). "Effect of cordycepin on interleukin-10 production of human peripheral blood mononuclear cells." Eur J Pharmacol 453(2-3): 309-17.
Zickler, D., S. Arnaise, et al. (1995). "Altered mating-type identity in the fungus Podospora anserina leads to selfish nuclei, uniparental progeny, and haploid meiosis." Genetics 140(2): 493-503.
Zylber, E. A., S. Perlman, et al. (1971). "Mitochondrial RNA turnover in the presence of cordycepin." Biochim Biophys Acta 240(4): 588-93.