块菌属

块菌属
科学分类
界：	真菌界 Fungi
门：	子囊菌门 Ascomycota
纲：	盘菌纲 Pezizomycetes
目：	盘菌目 Pezizales
科：	块菌科 Tuberaceae
属：	块菌属 Tuber; P.Micheli（英语：P.Micheli） ex F.H.Wigg.（英语：F.H.Wigg.） (1780)
模式种
	Tuber aestivum; Vittad.（英语：Vittad.） (1831)
异名
	Tuber P.Micheli (1729); Tuberium Raf. (1815); Aschion Wallr.（英语：Wallr.） (1833); Ensaluta Zobel (1854); Lespiaultinia Zobel (1854); Oogaster Corda (1854); Tuber subgen. Vittadinion Zobel (1854); Vittadinion Zobel (1854); Terfeziopsis Harkn.（英语：Harkn.） (1899); Delastreopsis Mattir.（英语：Oreste Mattirolo） (1905); Mukagomyces S.Imai (1940)

块菌属（学名：Tuber），又称西洋松露属，为子囊菌门块菌科的一属，本属目前已知222种，其中有几种可以烹调，且价位极高^[2]。

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子囊果地下生（英语：Hypogeal），不具囊盖（inoperculate），形态上成球状、瘤块状，且略不规则。外表随种类不同而有不同表现，可能布满金字塔型或圆锥形的突起鳞片，如黑松露（Tuber melanosporum）；也可能如意大利白松露（Tuber magnatum）呈平滑表面。颜色从黑色、紫黑色、红棕色、黄褐色至灰白色不等^[3]。子囊果的巨观构造可以大致分为最外层的包被（perido）和内部的产孢腔（gleba）^[4]。包被外层由外囊盘被（ectal excipulum）所构成，周边皮质则由致密的囊盘被髓质（medullary excipulum）构成。皮质常为假薄壁（pseudoparenchymatous）组织形态，有些则为长轴（prosenchymatous）组织形态^[3]。

产孢腔切面则可见因子囊盘的重叠压缩所造成的迷宫样致密曲折纹路，这些纹路称为脉（venae）。脉又分为外脉（venae externae）^{[注 1]}及颜色较白内脉（venae internae）^[3]^{[注 2]}。外脉为子实层，负责产生子实，组织内含有空气；内脉不具生殖功能，在发育初期颜色呈白色，随着个体逐渐成熟逐渐变深，其材质较为致密且不含气体^[3]。

子囊内含一至八颗孢子。本属的孢子多有复杂纹饰，可分为两类：一类有许多的棘状突起，此类以黑孢松露为代表；另一类则具有蜂巢般的格纹，意大利白松露属于此型^[4]。

一般盘菌的子囊前端有囊盖（operculate），成熟时囊盖会打开并释出孢子。但块菌生长于地下，又无法弹射孢子，因此会散发浓郁气味，吸引哺乳动物挖掘食用，借此散布孢子^[5]。会食用块菌的动物包括熊及猪，因此前人常使用猪的嗅觉来寻找块菌，在母猪嗅闻到块菌的位置后，再将松露掘出。但后来因为猪的力气较大不易控制，因此改用狗来寻找块菌^[5]。

种间关系

块菌属属于菌根真菌，常会与其他植物产生菌根^[5]。欧洲最常见的共生乔木包含橡树和榛树。在亚洲则主要与主要与壳斗科、桦木科及松科等树种的根系共生^[5]^[6]。

过往认为块菌无法单独生存，必须与其他植物形成共生性的外生菌根，以吸收孢外间质才能生存^[7]。但后来发现某些块菌（如黑冬松露（英语：Tuber melanosporum））也会摄取环境的资源，甚至会使一些植物的根部退化，借此吸收其残留下来的营养和矿物质。因此块菌与植物间的种间关系并不仅有互利共生，可能也会出现片利共生、寄生，及死体营养（necrotrophy）等交互关系^[7]。目前研究支持块菌属于辅助性腐生（auxiliary saprotrophic nutrition）生物，当植物的根系密集时，块菌采取腐生模式，吸收孢外间质的养分；根系较稀疏时，块菌会开始掠夺周边的资源。然而，自由生长的块菌由于缺乏营养供给，因此无法形成子囊果，且未必能与其他物种竞争^[7]。

除乔木的外生菌根之外，某些块菌也被发现可以与兰科植物形成兰菌根，也能与杜鹃花类植物形成浆果鹃类菌根（arbutoid mycorrhizas）^[8]。

本属包括以下物种：^[9]

夏块菌 Tuber aestivum (Wulfen) Spreng.
白脐块菌 Tuber alboumbilicum Y. Wang & Shu H. Li
Tuber anniae W. Colgan & Trappe
Tuber argenteum Gilkey
Tuber argentinum Speg.
Tuber arnoldianum Healy, Zurier & Bonito
阿萨块菌 Tuber asa-foetida
Tuber aztecorum G. Guevara, Bonito & M.E. Sm.
Tuber badium S.P. Wan
Tuber bellisporum Bonito & Trappe
Tuber bernardinii Gori
Tuber besseyi Gilkey
Tuber beyerlei Trappe, Bonito & G. Guevara
Tuber bisporum Gilkey
Tuber bituminatum Berk. & Broome
波密块菌 Tuber bomiense K.M. Su & W.P. Xiong
Tuber bonitoi G. Guevara & Trappe
Tuber bonnetii Roum.
勃氏块菌 Tuber borchii Vittad.，波氏块菌
勃氏块菌球孢变种 Tuber borchii var. sphaerosperma Malençon
Tuber brennemanii A. Grupe, Healy & M.E. Sm.
冬块菌 Tuber brumale Vittad.
Tuber brunneum G. Guevara, Bonito & Trappe
加州块菌 Tuber californicum
Tuber calosporum S.P. Wan
Tuber canaliculatum Gilkey
Tuber caoi L. Fan
Tuber castellanoi Bonito & Trappe
Tuber castilloi G. Guevara, Bonito & Trappe
Tuber cistophilum P. Alvarado, G. Moreno, Manjón, Gelpi & Jaime Muñoz
Tuber clarei Gilkey
Tuber crassitunicatum L. Fan & X.Y. Yan
Tuber cryptobrumale Merényi, T. Varga & Bratek
Tuber decipiens Bozac, Širić & Kos
Tuber depressum S.P. Wan
Tuber donnagotto Bozac, Širić & Kos
喜栎块菌 Tuber dryophilum Tul. & C. Tul.
深脉松露 Tuber elevatireticulatum K.F. Wong & H.T. Li
凹陷块菌 Tuber excavatum Vittad.
Tuber ferrugineum Vittad.
Tuber flavidosporum Hir. Sasaki, A. Kinosh. & Nara
Tuber floridanum A. Grupe, Sulzbacher & M.E. Sm.
臭块菌 Tuber foetidum Vittad.
台湾块菌 Tuber formosanum H.T. Hu & Y. Wang
粉状块菌 Tuber furfuraceum H.T. Hu & Y.I. Wang
Tuber gardneri Gilkey
Tuber gennadii (Chatin) Pat.
Tuber giganteum Gilkey
巨孢块菌 Tuber gigantosporum Y. Wang & Z.P. Li
光巨孢块菌 Tuber glabrum L. Fan & S. Feng
Tuber guevarae Bonito & Trappe
Tuber guzmanii Trappe & Cázares
Tuber harknessii Gilkey
喜马拉雅块菌 Tuber himalayense B.C. Zhang & Minter
Tuber hiromichii (S. Imai) Trappe
湖北块菌 Tuber hubeiense L. Fan
会东块菌 Tuber huidongense Y. Wang
会泽块菌 Tuber huizeanum L. Fan & C.L. Hou
Tuber incognitum Piña Páez, Bonito, Guevara & Castellano
印度块菌 Tuber indicum Cooke & Massee，异名：Tuber sinense K. Tao& B. Liu)
Tuber intermedium Bucholtz
Tuber irradians Gilkey
Tuber itzcuinzapotl de la Fuente & Rosales-Rosales, 2024 ^[10]
日本块菌 Tuber japonicum Hir. Sasaki, A. Kinosh. & Nara
金沙江块菌 Tuber jinshajiangense L. Fan
朝鲜块菌 Tuber koreanum H.Park & A.H.Eom, 2021^[11]
Tuber lacunosum Mattir.
Tuber lannaense Suwannar. & Lumyong
宽孢块菌 Tuber latisporum Juan Chen & P.G. Liu，阔孢块菌
Tuber lauryi Trappe, Bonito & G. Guevara
Tuber levissimum Gilkey
辽东块菌 Tuber liaotongense Y. Wang
Tuber lignarium (Harkn.) Gilkey
丽江块菌 Tuber lijiangense L. Fan & J.Z. Cao
Tuber linsdalei Gilkey
Tuber lishanense L. Fan & X.Y. Yan
小西氏石栎松露 Tuber lithocarpii K.F. Wong & H.T. Li
刘氏块菌 Tuber liui A-S. Xu
李玉块菌 Tuber liyuanum L. Fan & J.Z. Cao
Tuber longisporum Gilkey
Tuber lucentum Bordallo
莱氏块菌 Tuber lyonii Butters
Tuber macrosporum Vittad.
斑点块菌 Tuber maculatum Vittad.
Tuber magentipunctatum Merényi, I. Nagy, Stielow & Bratek
白松露 Tuber magnatum Picco, 1788
Tuber malacodermum E. Fisch.
Tuber malenconii Donadini, Riousset, G. Riousset & G. Chev.
Tuber maresa Font Quer
黑松露 Tuber melanosporum Vittad.
Tuber melosporum (G. Moreno, J. Díez & Manjón) P. Alvarado, G. Moreno, Manjón & J. Díez
膜皱块菌 Tuber mesentericum Vittad.
Tuber mexiusanum G. Guevara, Bonito & Cázares
Tuber michailowskianum Bucholtz
小孢块菌 Tuber microspermum L. Fan & J.Z. Cao
小球孢块菌 Tuber microsphaerosporum L. Fan & Y. Li
密网孢块菌 Tuber microspiculatum L. Fan & J.Z. Cao
细疣块菌 Tuber microverrucosum L. Fan & C.L. Hou
Tuber miquihuanense G. Guevara, Bonito & Cázares
Tuber monosporum (Mattir.) Vizzini
山地块菌 Tuber montanum Chatin, 1892
Tuber moravicum Velen.
Tuber moretii Maire
Tuber multimaculatum Parladé, Trappe & I.F. Alvarez^[12]
网孢凹陷块菌 Tuber neoexcavatum L. Fan & Yu Li
光果块菌 Tuber nitidum Vittad.
少孢块菌 Tuber oligospermum (Tul. & C. Tul.) Trappe
Tuber oregonense Trappe, Bonito & P. Rawl.
Tuber pacificum Trappe, Castellano & Bushnell
攀枝花块菌 Tuber panzhihuanense X.J. Deng & Y. Wang
小块菌 Tuber parvomurphium L. Fan
Tuber petrophilum Milenković, P. Jovan., Grebenc, Ivančević & Marković
Tuber phlebodermum (Gilkey) Trappe
Tuber piceatum L. Fan, X.Y. Yan & M.S. Song
Tuber polymorphosporum S.P. Wan
多孢块菌 Tuber polyspermum L. Fan & J.Z. Cao
假柄块菌 Tuber pseudobrumale Y. Wang & Shu H. Li
假凹陷块菌 Tuber pseudoexcavatum Y. Wang, G. Moreno, Riousset, Manjón & G. Riousset
假喜马拉雅块菌 Tuber pseudohimalayense G. Moreno, Manjón, J. Díez & García-Mont.
拟白块菌 Tuber pseudomagnatum L. Fan
Tuber pseudoseparans G. Guevara, Bonito & Trappe
拟球孢块菌 Tuber pseudosphaerosporum L. Fan
短毛块菌 Tuber puberulum Berk. & Broome
Tuber pustulatum M. Leonardi, A. Paz, G. Guevara & Pacioni
Tuber queletianum Bataille
Tuber quercicola J.L. Frank, D. Southw. & Trappe
曲靖块菌 Tuber qujingense S.P. Wan
芜菁味块菌 Tuber rapaeodorum Tul. & C. Tul.
Tuber regianum Montecchi & Lazzari
Tuber regimontanum G. Guevara, Bonito & Julio Rodr.
棕红块菌 Tuber rufum Pollini
Tuber scleroneuron Berk. & Broome
Tuber separans Gilkey
歇尔氏块菌 Tuber shearii Harkn.
Tuber shidianense S.P. Wan & F.Q. Yu
Tuber shii L. Fan & Y.W. Wang
中国块菌 Tuber sinense K. Tao & B. Liu
中华夏块菌 Tuber sinoaestivum J.P. Zhang & P.G. Liu
中华白块菌 Tuber sinoalbidum L. Fan & J.Z. Cao
中华凹陷块菌 Tuber sinoexcavatum L. Fan & Yu Li
中华巨孢块菌 Tuber sinomonosporum J.Z. Cao & L. Fan
Tuber sinoniveum S.P. Wan
中华短毛块菌 Tuber sinopuberulum L. Fan & J.Z. Cao
中华球孢块菌 Tuber sinosphaerosporum L. Fan, J.Z. Cao & Yu Li
松露块菌 Tuber songlu S.P. Wan
球孢块菌 Tuber sphaerospermum (Malençon) P. Roux, Guy García & M.C. Roux
Tuber sphaerosporum Gilkey
Tuber spinoreticulatum Uecker & Burds.
Tuber suave Pacioni & M. Leonardi, 2021^[13]
Tuber suaveolens Ant. Rodr. & Morte 2021
亚球孢块菌 Tuber subglobosum L. Fan & C.L. Hou
太原块菌 Tuber taiyuanense B. Liu
Tuber tequilanum G. Guevara, Bonito & Trappe
德州块菌 Tuber texense Heimsch
泰国块菌 Tuber thailandicum Suwannar., Kumla & Lumyong
Tuber theleascum M. Leonardi, A. Paz, G. Guevara & Pacioni
姜黄块菌 Tuber turmericum L. Fan
脐状块菌 Tuber umbilicatum Juan Chen & P.G. Liu
钩块菌 Tuber uncinatum Chatin, 1888
单色块菌 Tuber unicolor Gilkey
Tuber vacini Velen.
Tuber verii Pacioni & Lalli
Tuber verrucosivolvum S.P. Wan
囊被块菌 Tuber vesicoperidium L. Fan
Tuber walkeri Healy, Bonito & G. Guevara
Tuber wanglangense L. Fan
汶川块菌 Tuber wenchuanense L. Fan & J.Z. Cao
Tuber whetstonense J.L. Frank, D. Southw. & Trappe
乌蒙块菌 Tuber wumengense L. Fan
黄孢块菌 Tuber xanthomonosporum Qing & Yun Wang bis
西藏块菌 Tuber xizangense A-S. Xu
宣化块菌 Tuber xuanhuaense L. Fan
Tuber yigongense L. Fan & W.P. Xiong
中甸块菌 Tuber zhongdianense X.Y. He, Hai M. Li & Y. Wang

[注 1]
其他别称还有：venae lymphaticae、veines aquiferes of Tulasne、venae internae of Vittadini。
[注 2]
其他别称还有：venae lymphaticae、veines aquiferes of Tulasne、venae internae of Vittadini。

[1]
Tuber P. Micheli ex F.H. Wigg. 1780. MycoBank. International Mycological Association. [2011-07-13].
[2]
Kirk PM, Cannon PF, Minter DW, Stalpers JA. Dictionary of the Fungi 10th. Wallingford, UK: CABI. 2008: 708. ISBN 978-0-85199-826-8.
[3]
Hanlin, Richard T. Illustrated genera of ascomycetes. TUBER P. Mich. ex A. Wigg.. St. Paul, Minn.: APS Press. 1990: 126. ISBN 0890541078. OCLC 21300121.
[4]
ROOT TIP: the terminal portion of a root of a tree.ASCUS: the spore bearing sac.. www.tuber.it. [2018-08-12]. （原始内容存档于2018-08-12）.
[5]
国立自然科学博物馆. 菌類學習資源. [2018-08-13]. （原始内容存档于2016-03-24）（中文（台湾））.
[6]
歷時5年調查世界新種「深脈松露」發表揭開台灣真菌多樣性珍寶. 环境资讯中心、行政院农业委员会林务局. 2019-03-06 [2023-07-04]. （原始内容存档于2023-07-05）.
[7]
Gryndler, Milan. True Truffle Host Diversity. True Truffle (Tuber spp.) in the World. Cham: Springer International Publishing. 2016: 267–281 [2018-08-13]. ISBN 9783319314341. doi:10.1007/978-3-319-31436-5_16. （原始内容存档于2018-08-13）（英语）.
[8]
Taschen, Elisa; Sauve, Mathieu; Taudiere, Adrien; Parlade, Javier; Selosse, Marc-André; Richard, Franck. Whose truffle is this? Distribution patterns of ectomycorrhizal fungal diversity inTuber melanosporumbrûlés developed in multi-host Mediterranean plant communities. Environmental Microbiology. 2015-02-14, 17 (8): 2747–2761. ISSN 1462-2912. doi:10.1111/1462-2920.12741.
[9]
Tuber P.Micheli ex F.H.Wigg.. GBIF. [2023-01-25]. （原始内容存档于2023-01-25）.
[10]
Javier Isaac de la Fuente, Wendy Rosales-Rosales, César Ramiro Martínez-González, Magdalena Martínez-Reyes, Andrea Carolina Elizondo-Salas and Jesus Perez-Moreno. 2024. Tuber itzcuinzapotl sp. nov. (Tuberaceae, Pezizomycetes), the First Edible Truffle reported from Mexico with Traditional Biocultural Importance. Phytotaxa. 635(3); 206-216. DOI: 10.11646/phytotaxa.635.3.2
[11]
Hyeok Park, Ju-Hui Gwon, Jong-Chul Lee & Ahn-Heum Eom (2021) Report on a New Truffle Species, Tuber koreanum sp. nov., from Korea, Mycobiology, 49:6, 527-533, DOI: 10.1080/12298093.2021.1992089
[12]
Isabel F. Alvarez, Javier Parladé & James M. Trappe (1992) Loculotuber Gennadii Gen. Et Comb. Nov. and Tuber Multimaculatum Sp. Nov., Mycologia, 84:6, 926-929, DOI: 10.1080/00275514.1992.12026228
[13]
Leonardi, M.; Salvi, D.; Iotti, M.; Rana, G.L.; Paz-Conde, A.; Pacioni, G. Multilocus Phylogeography of the Tuber mesentericum Complex Unearths Three Highly Divergent Cryptic Species. J. Fungi 2021, 7, 1090. https://doi.org/10.3390/jof7121090

块菌属在Index Fungorum中的资讯
The Lightning and the Truffle （页面存档备份，存于互联网档案馆）

[5] [注 1]
其他别称还有：venae lymphaticae、veines aquiferes of Tulasne、venae internae of Vittadini。

[6] [注 2]
其他别称还有：venae lymphaticae、veines aquiferes of Tulasne、venae internae of Vittadini。

[urlMycoBank:_Tuber-1] [1]
Tuber P. Micheli ex F.H. Wigg. 1780. MycoBank. International Mycological Association. [2011-07-13].

[Kirk2008-2] [2]
Kirk PM, Cannon PF, Minter DW, Stalpers JA. Dictionary of the Fungi 10th. Wallingford, UK: CABI. 2008: 708. ISBN 978-0-85199-826-8.

[:0-3] [3]
Hanlin, Richard T. Illustrated genera of ascomycetes. TUBER P. Mich. ex A. Wigg.. St. Paul, Minn.: APS Press. 1990: 126. ISBN 0890541078. OCLC 21300121.

[:1-4] [4]
ROOT TIP: the terminal portion of a root of a tree.ASCUS: the spore bearing sac.. www.tuber.it. [2018-08-12]. （原始内容存档于2018-08-12）.

[:2-7] [5]
国立自然科学博物馆. 菌類學習資源. [2018-08-13]. （原始内容存档于2016-03-24）（中文（台湾））.

[:02-8] [6]
歷時5年調查世界新種「深脈松露」發表揭開台灣真菌多樣性珍寶. 环境资讯中心、行政院农业委员会林务局. 2019-03-06 [2023-07-04]. （原始内容存档于2023-07-05）.

[:3-9] [7]
Gryndler, Milan. True Truffle Host Diversity. True Truffle (Tuber spp.) in the World. Cham: Springer International Publishing. 2016: 267–281 [2018-08-13]. ISBN 9783319314341. doi:10.1007/978-3-319-31436-5_16. （原始内容存档于2018-08-13）（英语）.

[10] [8]
Taschen, Elisa; Sauve, Mathieu; Taudiere, Adrien; Parlade, Javier; Selosse, Marc-André; Richard, Franck. Whose truffle is this? Distribution patterns of ectomycorrhizal fungal diversity inTuber melanosporumbrûlés developed in multi-host Mediterranean plant communities. Environmental Microbiology. 2015-02-14, 17 (8): 2747–2761. ISSN 1462-2912. doi:10.1111/1462-2920.12741.

[11] [9]
Tuber P.Micheli ex F.H.Wigg.. GBIF. [2023-01-25]. （原始内容存档于2023-01-25）.

[12] [10]
Javier Isaac de la Fuente, Wendy Rosales-Rosales, César Ramiro Martínez-González, Magdalena Martínez-Reyes, Andrea Carolina Elizondo-Salas and Jesus Perez-Moreno. 2024. Tuber itzcuinzapotl sp. nov. (Tuberaceae, Pezizomycetes), the First Edible Truffle reported from Mexico with Traditional Biocultural Importance. Phytotaxa. 635(3); 206-216. DOI: 10.11646/phytotaxa.635.3.2

[13] [11]
Hyeok Park, Ju-Hui Gwon, Jong-Chul Lee & Ahn-Heum Eom (2021) Report on a New Truffle Species, Tuber koreanum sp. nov., from Korea, Mycobiology, 49:6, 527-533, DOI: 10.1080/12298093.2021.1992089

[14] [12]
Isabel F. Alvarez, Javier Parladé & James M. Trappe (1992) Loculotuber Gennadii Gen. Et Comb. Nov. and Tuber Multimaculatum Sp. Nov., Mycologia, 84:6, 926-929, DOI: 10.1080/00275514.1992.12026228

[15] [13]
Leonardi, M.; Salvi, D.; Iotti, M.; Rana, G.L.; Paz-Conde, A.; Pacioni, G. Multilocus Phylogeography of the Tuber mesentericum Complex Unearths Three Highly Divergent Cryptic Species. J. Fungi 2021, 7, 1090. https://doi.org/10.3390/jof7121090

[2]

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[3]

[4]

[注 1]

[注 2]

[5]

[6]

[7]

[8]

[9]

[10]

[11]

[12]

[13]

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生物学

下属物种

参见

注释

参考文献

外部链接