黑僵菌属
黑僵菌属
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| 被黑僵菌杀死的蟑螂 | |
科学分类 
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| 界： | 真菌界 Fungi |
| 门： | 子囊菌门 Ascomycota |
| 亚门： | 盘菌亚门 Pezizomycotina |
| 纲： | 粪壳菌纲 Sordariomycetes |
| 目： | 肉座菌目 Hypocreales |
| 科： | 麦角菌科 Clavicipitaceae |
| 属： | 黑僵菌属 Metarhizium Sorokīn, 1879 |
| 模式种 | |
| 黑僵菌 M. anisopliae (Metchnikoff) Sorokīn
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| 种 | |
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黑僵菌属（学名：Metarhizium），又称绿僵菌属，是麦角菌科的一个属，其下物种为可感染昆虫的虫生真菌。本属物种过去多以外形区分，现代则以分子序列鉴定。许多本属物种可感染影响农作物的害虫，因此可用作生物农药以抑制害虫的生长。
分类
1879年，俄罗斯微生物学家伊利亚·梅契尼可夫首次于奥国赛丽金龟体内分离了黑僵菌，将其归入虫霉属中，命名为Entomophthora anisopliae，1883年，另一位俄罗斯生物学家N. Sorokin将其归入新属黑僵菌属（Metarhizium），后来许多文献将本属的属名误植为Metarrhizium[1]。在分子生物学技术于二十世纪末问世以前，黑僵菌属的物种多是以外形区分，其中分生孢子形态为重要的区分依据之一，较早被分出的物种包括黑僵菌、棕色绿僵菌、黄绿绿僵菌、柱孢绿僵菌、M. cicadinum与M. truncatum等。1976年玛格利特·塔洛克（Margaret Tulloch）整理本属的物种，只承认黑僵菌（M. anisopliae）与黄绿绿僵菌（M. flavoviride）两物种，将其他物种都视为异名或变种[2] 。1991年，有研究指出许多本属物种为虫草属物种的无性型[3]，2007年，本属的有性型与部分虫草属的物种一起被归入新属异虫草属中[4]。
DNA测序普及后，本属许多隐存种渐被发表。2000年，一篇分子种系发生学的研究发现黑僵菌与黄绿绿僵菌两物种皆分别由许多演化支组成[5]，2009年，另一篇分子种系发生学研究显示黑僵菌（M. anisopliae）为一复合种，其下可分为九个演化支，其中许多过去为黑僵菌的变种，皆被提升至独立物种[3]，随后亦有许多新种陆续被发表描述[6]。
| 黑僵菌（M. anisopliae）的复合种 |
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感染
本属真菌以分生孢子感染昆虫，分生孢子接触昆虫体表后，可萌发长出附着器，施加机械压力并分泌多种水解酶以穿透体表而进入血淋巴中。在血淋巴中，菌丝会出芽产生孢子（blastospores），又称芽孢体（hyphal bodies）[7][8]，可脱离菌丝而在血淋巴中游离生长、大量增殖，且其表面蛋白可抑制昆虫的吞噬细胞对其进行攻击[9]，菌丝亦可分泌多种黑僵菌素（destruxin）等多种毒素以帮助杀死宿主[1]，随后菌丝会长出体表并将分生孢子散播至环境中。感染本属真菌的昆虫死后数天身上即会呈现分生孢子的颜色，依物种而异可为白色、黄色、棕色或绿色[10]。
本属许多物种可感染影响农作物的害虫，因此可用作生物农药以抑制害虫的生长[11]，模式种黑僵菌还可感染孑孓，有助抑制疟疾的传播[12]。有些物种感染的宿主种类广泛，如罗伯茨绿僵菌（M. robertsii），有些物种感染宿主的种类则较专一，如蝗绿僵菌仅能感染直翅目的昆虫[11]，后者基因组编码的蛋白质种类较前者少[13]。另外部分物种可被整体病毒科与分体病毒科等双股RNA的真菌病毒感染，可能使其感染昆虫的能力降低[1]。
本属物种常可在土壤样本中被分离，但其于土壤中的生长形态仍不明，亦不清楚其可于土壤中生长，或仅处于休眠状态以等待感染宿主的机会[11][14]，有研究显示部分本属真菌在根际环境的存活率比在周边土壤中好[10][15]。
物种
以下为黑僵菌属的物种列表[注 1] （页面存档备份，存于互联网档案馆）。
- 蝗绿僵菌 Metarhizium acridum (Driver & Milner) J.F. Bisch., S.A. Rehner & Humber (2009)
- 黑僵菌 Metarhizium anisopliae (Metschn.) Sorokīn (1883)[18]，金龟子绿僵菌
- Metarhizium acridum (Driver & Milner) J.F.Bisch., S.A.Rehner & Humber[19]
- 暗绿绿僵菌 M. atrovirens (Kobayasi & Shimizu) Kepler, S.A. Rehner & Humber (2014)
- Metarhizium bibionidarum O. Nishi & H. Sato (2017)
- Metarhizium blattodeae C. Montalva, Humber, K. Collier & C. Luz (2016)
- Metarhizium brasiliense Kepler, S.A. Rehner & Humber (2014)
- 拟布里班克绿僵菌 M. brittlebankisoides (Zuo Y. Liu, Z.Q. Liang, Whalley, Y.J. Yao & A.Y. Liu) Kepler, S.A. Rehner & Humber (2014)
- 棕色绿僵菌 Metarhizium brunneum Petch (1935)，褐色绿僵菌
- 打铁虫绿僵菌 Metarhizium campsosterni (W.M. Zhang & T.H. Li) Kepler, S.A. Rehner & Humber (2014)
- Metarhizium carneum (Duché & R. Heim) Kepler, S.A. Rehner & Humber (2014)
- Metarhizium chaiyaphumense Tasan., Khons., Thanakitp., Mongkols. & Luangsa-ard (2017)
- Metarhizium cicadinum (Höhn.) Petch (1931)
- 柱孢绿僵菌 Metarhizium cylindrosporum Q.T. Chen & H.L. Guo (1986)
- 松毛虫绿僵菌 Metarhizium dendrolimatilis Z.Q. Liang, Wan H. Chen, Y.F. Han & X. Zou (2017)
- Metarhizium dianzhongense [20]
- 黄绿绿僵菌 Metarhizium flavoviride W. Gams & Rozsypal (1973)[18]
- 耐寒绿僵菌 Metarhizium frigidum J.F. Bisch. & S.A. Rehner (2007)[18]
- Metarhizium globosum J.F. Bisch., S.A. Rehner & Humber (2009)
- 贵州绿僵菌 Metarhizium guizhouense Q.T. Chen & H.L. Guo (1986)（戴氏绿僵菌 Metarhizium taii Z.Q. Liang & A.Y. Liu (1991)为其异名[3]）
- Metarhizium granulomatis (Sigler) Kepler, S.A. Rehner & Humber (2014)
- Metarhizium guniujiangense (C.R. Li, B. Huang, M.Z. Fan & Z.Z. Li) Kepler, S.A. Rehner & Humber (2014)
- Metarhizium indigoticum (Kobayasi & Shimizu) Kepler, S.A. Rehner & Humber (2014)
- Metarhizium kalasinense Tasan., Khons., Thanakitp., Mongkols. & Luangsa-ard (2017)
- Metarhizium khaoyaiense (Hywel-Jones) Kepler, S.A. Rehner & Humber (2014)
- Metarhizium koreanum Kepler, S.A. Rehner & Humber (2014)
- 草剃绿僵菌 Metarhizium kusanagiense (Kobayasi & Shimizu) Kepler, S.A. Rehner & Humber (2014)
- 鳞翅目绿僵菌 Metarhizium lepidiotae (Driver & Milner) J.F. Bisch., S.A. Rehner & Humber (2009)
- 大孢绿僵菌 Metarhizium majarosporae
- 大孢绿僵菌 Metarhizium majus (J.R. Johnst.) J.F. Bisch., S.A. Rehner & Humber (2009)
- Metarhizium marquandii (Massee) Kepler, S.A. Rehner & Humber (2014)
- Metarhizium megapomponiae Luangsa-ard, Khonsanit, Thanakitpipattana & Samson[21]
- Metarhizium minus (Rombach, Humber & D.W. Roberts) Kepler, S.A. Rehner & Humber (2014)
- Metarhizium megapomponiae Luangsa-ard, Khonsanit, Thanakitpipattana & Samson
- Metarhizium niveum Luangsa-ard, Khonsanit, Thanakitpipattana & Samson[21]
- 新西兰绿僵菌 Metarhizium novozealandicum (Driver & Milner) Kepler, S.A. Rehner & Humber (2014)
- Metarhizium ovoidosporum Luangsa-ard, Khonsanit, Thanakitpipattana & Samson[21]
- 尾张绿僵菌 Metarhizium owariense (Kobayasi) Kepler, S.A. Rehner & Humber (2014)
- 瘿绵蚜绿僵菌 Metarhizium pemphigi (Driver & Milner) Kepler, Humber & S.A. Rehner (2014)
- 平沙绿僵菌 Metarhizium pinghaense Q.T. Chen & H.L. Guo (1986)
- Metarhizium prachinense Tasan., Khons., Thanakitp., Mongkols. & Luangsa-ard (2017)
- 拟暗绿绿僵菌 Metarhizium pseudoatrovirens (Kobayasi & Shimizu) Kepler, S.A. Rehner & Humber (2014)
- Metarhizium purpureogenum O. Nishi, S. Shimizu & H. Sato (2017)
- Metarhizium reniforme (Samson & Evans) Luangsa-ard, Boucias & Hywel-Jones (2017)
- 莱氏绿僵菌 Metarhizium rileyi (Farl.) Kepler, S.A. Rehner & Humber (2014) （旧名Nomuraea rileyi）
- 罗伯茨绿僵菌 Metarhizium robertsii J.F. Bisch., S.A. Rehner & Humber (2009)
- Metarhizium samlanense Luangsa-ard, Thanakitp., Tasan., Mongkols. & Hywel-Jones (2017)
- 束梗绿僵菌 Metarhizium synnematis Z.Q.Liang, H.L.Chu & T.C.Wen, 2016
- Metarhizium takense Tasan., Thanakitp., Mongkols. & Luangsa-ard (2017)
- Metarhizium truncatum Petch (1931)
- Metarhizium viride (Segretain, Fromentin, Destombes, Brygoo & Dodin ex Samson) Kepler, S.A. Rehner & Humber (2014)
- Metarhizium viridulum (Tzean, L.S. Hsieh, J.L. Chen & W.J. Wu) B. Huang & Z.Z. Li (2004)
- Metarhizium yongmunense (G.H. Sung, J.M. Sung & Spatafora) Kepler, S.A. Rehner & Humber (2014)
已归入他属
注脚
参考资料
- ^ 1.0 1.1 1.2 Roberts, Donald W.; St. Leger, Raymond J. Metarhizium spp., Cosmopolitan Insect-Pathogenic Fungi: Mycological Aspects. Advances in Applied Microbiology. 2004, 54: 1–70. ISSN 0065-2164. doi:10.1016/S0065-2164(04)54001-7.
- ^ Tulloch, M. The genus Metarhizium. Transactions of the British Mycological Society. 1976, 66 (3): 407–411. ISSN 0007-1536. doi:10.1016/S0007-1536(76)80209-4.
- ^ 3.0 3.1 3.2 Bischoff J.F.; Rehner S.A. Humber R.A. A multilocus phylogeny of the Metarhizium anisopliae lineage. Mycologia. 2009, 101 (4): 512–530 [2019-07-05]. PMID 19623931. doi:10.3852/07-202. （原始内容存档于2020-08-19）.
- ^ Sung, G.-H.; Hywel-Jones, N.L.; Sung, J.-M.; Luangsa-ard, J.J.; Shrestha, B. & Spatafora, J.W. Phylogenetic classification of Cordyceps and the clavicipitaceous fungi. Studies in Mycology. 2007, 57: 5–59. PMC 2104736
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- ^ Driver, Felice; Milner, Richard J.; Trueman, John W.H. A taxonomic revision of Metarhizium based on a phylogenetic analysis of rDNA sequence data. Mycological Research. 2000, 104 (2): 134–150. ISSN 0953-7562. doi:10.1017/S0953756299001756.
- ^ 於Species fungorum搜索Metarhizium. [2019-07-05]. （原始内容存档于2008-05-11）.
- ^ Butt, T.M.; Coates, C.J.; Dubovskiy, I.M.; Ratcliffe, N.A. Entomopathogenic Fungi 94: 307–364. 2016. ISSN 0065-2660. doi:10.1016/bs.adgen.2016.01.006.
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- ^ Brian Lovett, Raymond St. Leger. Genetics and Molecular Biology of Entomopathogenic Fungi. Elsevier Science. 27 April 2016: 112–113 [2019-07-06]. ISBN 978-0-444-63723-9. （原始内容存档于2020-08-19）.
- ^ 10.0 10.1 Todd Ugine. Metarhizium (Order: Hypocreales, Family: Clavicipitaceae). College of Agriculture and Life Sciences, Cornell University. [2019-07-06]. （原始内容存档于2020-01-21）.
- ^ 11.0 11.1 11.2 Brunner-Mendoza, Carolina; Reyes-Montes, María del Rocío; Moonjely, Soumya; Bidochka, Michael J; Toriello, Conchita. A review on the genus Metarhizium as an entomopathogenic microbial biocontrol agent with emphasis on its use and utility in Mexico. Biocontrol Science and Technology. 2018, 29 (1): 83–102. ISSN 0958-3157. doi:10.1080/09583157.2018.1531111.
- ^ Bukhari T, Takken W, Koenraadt CJ. Development of Metarhizium anisopliae and Beauveria bassiana formulations for control of malaria mosquito larvae 4 (23). Parasites and Vectors. 2011. doi:10.1186/1756-3305-4-23.
- ^ Hu, Xiao; Xiao, Guohua; Zheng, Peng; Shang, Yanfang; Su, Yao; Zhang, Xinyu; Liu, Xingzhong; Zhan, Shuai; St. Leger, Raymond J.; Wang, Chengshu. Trajectory and genomic determinants of fungal-pathogen speciation and host adaptation. Proceedings of the National Academy of Sciences. 2014, 111 (47): 16796–16801. ISSN 0027-8424. doi:10.1073/pnas.1412662111.
- ^ G. Douglas Inglis, Mark S. Goettel, Tariq M. Butt And Hermann Strasser. Use of hyphomycetous fungi for managing insect pests. T.M. Butt, C. Jackson, N. Magan (编). Fungi as Biocontrol Agent: Progress, Problems and Potential (PDF). CABI Publishing. 2001: 23-70 [2019-07-05]. （原始内容存档 (PDF)于2019-02-20）.
- ^ Hu, G.; St. Leger, R. J. Field Studies Using a Recombinant Mycoinsecticide (Metarhizium anisopliae) Reveal that It Is Rhizosphere Competent. Applied and Environmental Microbiology. 2002, 68 (12): 6383–6387. ISSN 0099-2240. doi:10.1128/AEM.68.12.6383-6387.2002.
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- ^ 章西. 中国绿僵菌分子分类的研究. 安徽农业大学. 2015 [2019-07-05]. （原始内容存档于2020-08-19）.
- ^ 18.0 18.1 18.2 Joseph F. Bischoff, Stephen A. Rehner & Richard A. Humber (2006) Metarhizium frigidum sp. nov.: a cryptic species of M. anisopliae and a member of the M. flavoviride complex, Mycologia, 98:5, 737-745, DOI: 10.1080/15572536.2006.11832645
- ^ Metarhizium acridum. Parasite of the Day. [2024-03-30].
- ^ Wei, C.-Y.; Tang, M.; Xie, L.-Y.; Fan, Q.; Shen, S.-K.; Yang, Z.-L.; Deng, G.; Wang, Y.-B. Metarhizium dianzhongense sp. nov. and New Record of M. bibionidarum (Clavicipitaceae, Hyocreales) Attacking Insects from China. Diversity 2024, 16, 201. https://doi.org/10.3390/d16040201
- ^ 21.0 21.1 21.2 Suchada Mongkolsamrit, Artit Khonsanit, Donnaya Thanakitpipattana, K. Tasanathai, W. Noisripoom, S. Lamlertthon, W. Himaman, J. Houbraken, R. A. Samson and J. Luangsa-ard. 2020. Revisiting Metarhizium and the Description of New Species from Thailand. Studies in Mycology. 95; 171-251. DOI: 10.1016/j.simyco.2020.04.001
- ^ Gliocladium viridicolumnare. MycoBank. [2019-07-05]. （原始内容存档于2020-06-05）.
- ^ Metarhizium glutinosum. MycoBank. [2019-07-05]. （原始内容存档于2020-08-19）.
- ^ Luangsa-ard, J. Jennifer; Mongkolsamrit, Suchada; Thanakitpipattana, Donnaya; Khonsanit, Artit; Tasanathai, Kanoksri; Noisripoom, Wasana; Humber, Richard A. Clavicipitaceous entomopathogens: new species in Metarhizium and a new genus Nigelia. Mycological Progress. 2017, 16 (4): 369–391. ISSN 1617-416X. doi:10.1007/s11557-017-1277-1.
- ^ Nigelia martiale. MycoBank. [2019-07-05]. （原始内容存档于2020-08-19）.
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