黑殭菌屬

黑殭菌屬（學名：Metarhizium），又稱绿僵菌属，是麥角菌科的一個屬，其下物種為可感染昆蟲的蟲生真菌（英语：entomopathogenic fungi）。本屬物種過去多以外形區分，現代則以分子序列鑑定。許多本屬物種可感染影響農作物的害蟲，因此可用作生物農藥以抑制害蟲的生長。 1879年，俄羅斯微生物學家伊利亞·梅契尼可夫首次於奧國賽麗金龜（英语：Anisoplia austriaca）體內分離了黑殭菌，將其歸入蟲霉屬（英语：Entomophthora）中，命名為Entomophthora anisopliae，1883年，另一位俄羅斯生物學家N. Sorokin將其歸入新屬黑殭菌屬（Metarhizium），後來許多文獻將本屬的屬名誤植為Metarrhizium。在分子生物學技術於二十世紀末問世以前，黑殭菌屬的物種多是以外形區分，其中分生孢子形態為重要的區分依據之一，較早被分出的物種包括黑殭菌、棕色绿僵菌（英语：Metarhizium brunneum）、黄绿绿僵菌（英语：Metarhizium flavoviride）、柱孢绿僵菌（瑞典语：Metarhizium cylindrosporum）、M. cicadinum與M. truncatum等。1976年瑪格利特·塔洛克（Margaret Tulloch）整理本屬的物種，只承認黑殭菌（M. anisopliae）與黄绿绿僵菌（M. flavoviride）兩物種，將其他物種都視為異名或變種 。1991年，有研究指出許多本屬物種為蟲草屬（英语：Cordyceps）物種的無性型（英语：Teleomorph, anamorph and holomorph），2007年，本屬的有性型與部分蟲草屬的物種一起被歸入新屬异虫草属（英语：Metacordyceps）中。 DNA測序普及後，本屬許多隱存種漸被發表。2000年，一篇分子種系發生學的研究發現黑殭菌與黄绿绿僵菌兩物種皆分別由許多演化支組成，2009年，另一篇分子種系發生學研究顯示黑殭菌（M. anisopliae）為一複合種，其下可分為九個演化支，其中許多過去為黑殭菌的變種，皆被提升至獨立物種，隨後亦有許多新種陸續被發表描述。 本屬真菌以分生孢子感染昆蟲，分生孢子接觸昆蟲體表後，可萌發長出附著器，施加機械壓力並分泌多種水解酶以穿透體表而進入血淋巴中。在血淋巴中，菌絲會出芽產生孢子（blastospores），又稱芽孢體（hyphal bodies），可脫離菌絲而在血淋巴中游離生長、大量增殖，且其表面蛋白可抑制昆蟲的吞噬細胞對其進行攻擊，菌絲亦可分泌多種黑殭菌素（destruxin）等多種毒素以幫助殺死宿主，隨後菌絲會長出體表並將分生孢子散播至環境中。感染本屬真菌的昆蟲死後數天身上即會呈現分生孢子的顏色，依物種而異可為白色、黃色、棕色或綠色。 本屬許多物種可感染影響農作物的害蟲，因此可用作生物農藥以抑制害蟲的生長，模式種黑殭菌還可感染孑孓，有助抑制瘧疾的傳播。有些物種感染的宿主種類廣泛，如罗伯茨绿僵菌（M. robertsii），有些物種感染宿主的種類則較專一，如蝗綠殭菌（英语：Metarhizium acridum）僅能感染直翅目的昆蟲，後者基因組編碼的蛋白質種類較前者少。另外部分物種可被整體病毒科與分體病毒科等雙股RNA（英语：Double-stranded RNA viruses）的真菌病毒感染，可能使其感染昆蟲的能力降低。 本屬物種常可在土壤樣本中被分離，但其於土壤中的生長形態仍不明，亦不清楚其可於土壤中生長，或僅處於休眠狀態以等待感染宿主的機會，有研究顯示部分本屬真菌在根際環境的存活率比在周邊土壤中好。 以下為黑殭菌屬的物種列表 （页面存档备份，存于互联网档案馆）。 蝗綠殭菌 Metarhizium acridum (Driver & Milner) J.F. Bisch., S.A. Rehner & Humber (2009) 黑殭菌 Metarhizium anisopliae (Metschn.) Sorokīn (1883)，金龟子绿僵菌 Metarhizium acridum (Driver & Milner) J.F.Bisch., S.A.Rehner & Humber 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 黄绿绿僵菌 Metarhizium flavoviride W. Gams & Rozsypal (1973) 耐寒绿僵菌 Metarhizium frigidum J.F. Bisch. & S.A. Rehner (2007) Metarhizium globosum J.F. Bisch., S.A. Rehner & Humber (2009) 貴州绿僵菌 Metarhizium guizhouense Q.T. Chen & H.L. Guo (1986)（戴氏绿僵菌 Metarhizium taii Z.Q. Liang & A.Y. Liu (1991)為其異名） 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 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 新西兰绿僵菌 Metarhizium novozealandicum (Driver & Milner) Kepler, S.A. Rehner & Humber (2014) Metarhizium ovoidosporum Luangsa-ard, Khonsanit, Thanakitpipattana & Samson 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) 平沙綠僵菌 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) 萊氏綠殭菌 Metarhizium rileyi (Farl.) Kepler, S.A. Rehner & Humber (2014) （舊名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) Metarhizium viridicolumnare (Matsush.) Matsush. (1993)：被認為與黏帚黴屬（英语：Gliocladium）物種Gliocladium viridicolumnare的異名。 Metarhizium glutinosum S.A. Pope (1944)：被認為是疣孢漆斑菌（英语：Myrothecium verrucaria）（Myrothecium verrucaria）的異名。 Metarhizium martiale (Spegazzini) Kepler, S.A. Rehner & Humber (2014)：已被歸入新屬Nigelia屬，與黑殭菌屬關係接近。 维基共享资源上的相关多媒体资源：黑殭菌屬 部分中文名稱參考： 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.  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]. 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