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木腐真菌

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干腐菌英语Serpula lacrymans引发的木材腐朽

木腐真菌是指一类能在树木上生长并侵入木材组织分解木材的真菌,大多数的木腐真菌只能在枯死的树木上进行生长,例如褐腐真菌,但仍有少数腐菌能攻击活树,如蜜环菌属。真菌在木材上生长和繁殖需要有高于纤维饱和点以上的水分[1] ,当真菌渗入木材的纤维结构后,能产生酶来攻击木材组织,分解复杂分子并将养分送回土壤[2] 。不同的木腐真菌所产生的酶不同,可以降解不同的木材成分,并产生不同的木材腐朽形态[3] ,并可借此将木材腐朽真菌大致区分为褐腐、软腐和白腐真菌。[4][5] 由不同真菌作用分解的残留物具有不同的酸碱度、溶解度和氧化还原电势,这些残留物最后会变成土壤中的沉积物,随着时间的累积,可能对该地区的环境产生显著影响。[3]目前全世界约有5,000余种木腐真菌[6]

褐腐真菌

一块橡木上的褐腐真菌

褐腐真菌会分解木材中的半纤维素纤维素。纤维素是借由半纤维素分解过程中产生的过氧化氢(H2O2)来分解。[4] 由于过氧化氢的分子小,因此能在木材中迅速扩散,影响超出真菌菌丝周围的木材。因为无法破坏木质素,所以植物细胞的形状不会被破坏,但分解后会导致木材收缩破裂,呈现褐色的块状(cubical)龟裂。

褐腐真菌主要为担子菌,大多在针叶树上生长,具有特殊经济意义的褐腐真菌包括Serpula lacrymans,Fibroporia vaillantii和Coniophora puteana,因为它们可能会侵蚀建筑物中的木材。[7]褐腐真菌分解的特征是将木质素广泛地去甲基化,而白腐菌则是只有产生少量的去甲基官能基团分子。[8]

热带气候温带地区的南方很少有褐腐真菌,褐腐真菌主要位于北回归线以北(北纬23.5度),并且大多数位于北纬35度以北,大致分布在北方。那些在北纬23.5°到35°之间的褐腐真菌,通常是在高海拔地区的松树林或温带针叶林落矶山脉喜马拉雅山脉中发现。[9]

软腐真菌

软腐真菌不同于其他木材腐朽真菌,需要较高的固定氮(约1%)才能合成酶,通常需要由木材本身或周围的环境来提供氮源。软腐真菌会从其菌丝分泌纤维素酶来分解木材中的纤维素[4] ,这会造常木材内部产生微小的空腔,有时会造成类似于褐腐的变色和开裂现象。[4][5] 引起软腐的真菌有 ChaetomiumCeratocystisKretzschmaria deusta

软腐真菌的分解效率不如白腐真菌[3],但能够在褐腐真菌或白腐真菌无法栖息的条件下生长,同时还可以分解具有高含量抗生物侵蚀化合物的木材。树皮作为植物保护内部脆弱组织的第一层防护[10],许多木本植物的树皮中含有高浓度的单宁,这很难使真菌分解,而木栓质更是可以有效阻挡微生物的屏障。[10]

白腐真菌

桦木上的白腐真菌
橡木上的白腐真菌

白腐真菌不只能分解纤维素和半纤维素,更能分解木质素[5]因此改变了木材的质地,使木材变成潮湿柔软的海绵状或丝状残骸,颜色则变成了白色或黄色(因为木材的颜色主要来至木质素及二次代谢物的累积)[11] 由于白腐真菌能够产生分解木质素和其他复杂有机分子所需的酶(如漆酶),因此已有科学家研究其应用于真菌修复的可行性[12]。白腐真菌的分解作用涉及多种酶,其中一些直接氧化木质素[13] ,木质素的的侧链丙基会因为分解作用而降低。[8] 据研究,秀珍菇平菇)会优先降解木质素而不是多糖[14] ,这点不同于其他的白腐真菌(如Phanerochaete chrysosporium),它们对木质纤维素没有选择性[14],而Honey mushroom (Armillaria spp.) 则是一种可攻击活树的白腐真菌。

平菇是一种普遍被培养的白腐真菌[12] ,但并不是活体寄生真菌,不会在活树上生长,除非树木已经死亡[15] 。其他白腐真菌如 turkey tail, artist's conk, 和 tinder fungus.[7]白腐真菌在世界各地都有种植,可以作为食物的来源,例如常见的香菇(shiitake mushroom)在2003年时约占菇总产量的25%。[15][16]

红腐真菌

感染红腐症之植株

为白腐真菌之子分类,得名于因造成木材损坏而在感染初期变色为红色,此症状因此称之为红腐症,如:棕黑蜜环菌英语Armillaria ostoyaeArmillaria ostoyae)和Heterobasidion annosum英语Heterobasidion annosum

参见

  • Snag (ecology)
  • Compartmentalization of decay in trees

参考资料

  1. ^ Harris, Samuel Y. Building Pathology: Deterioration, Diagnostics, and Intervention. John Wiley & Sons. 2001: 106 [2019-10-22]. ISBN 978-0-471-33172-8. (原始内容存档于2020-02-25). 
  2. ^ Wood Decay in Living and Dead Trees: A Pictorial Overview (PDF). [2018-02-28]. (原始内容存档 (PDF)于2018-07-29). 
  3. ^ 3.0 3.1 3.2 Vane, C. H., et al. (2005). "Decay of cultivated apricot wood (Prunus armeniaca) by the ascomycete Hypocrea sulphurea, using solid state 13C NMR and off-line TMAH thermochemolysis with GC–MS." International Biodeterioration & Biodegradation 55(3): 175-185.
  4. ^ 4.0 4.1 4.2 4.3 J. Deacon, Wood decay and wood-rotting fungi页面存档备份,存于互联网档案馆). University of Edinburgh (2005?).
  5. ^ 5.0 5.1 5.2 Microorganisms causing decay in trees and wood页面存档备份,存于互联网档案馆). University of Minnesota.
  6. ^ Tong, Li; Junning, Li; Yulian, Wei. Species diversity and distribution of wood-decaying fungi in Gutianshan National Nature Reserve. Biodiversity Science. 2019, 27 (1): 81–87. doi:10.17520/biods.2018156. 
  7. ^ 7.0 7.1 Stamets, Paul. Mycelium running: how mushrooms can help save the world. Random House, Inc. 2005: 83–84. ISBN 978-1-58008-579-3. 
  8. ^ 8.0 8.1 Vane, C. H., et al. (2001). "The effect of fungal decay (Agaricus bisporus) on wheat straw lignin using pyrolysis–GC–MS in the presence of tetramethylammonium hydroxide (TMAH)." Journal of Analytical and Applied Pyrolysis 60(1): 69-78.
  9. ^ Ryvarden, Leif. Tropical polypores. Isaac, Susan (编). Aspects of Tropical Mycology. British Mycological Society. Symposium. Cambridge University Press. 1993: 159 [2019-10-22]. ISBN 978-0-521-45050-8. (原始内容存档于2020-02-25). 
  10. ^ 10.0 10.1 Vane, C. H., et al. (2006). "Bark decay by the white-rot fungus Lentinula edodes: Polysaccharide loss, lignin resistance and the unmasking of suberin." International Biodeterioration & Biodegradation 57(1): 14-23.
  11. ^ Wood Decay Fungi in Landscape Trees Management Guidelines--UC IPM. www.ipm.ucdavis.edu. [24 February 2018]. (原始内容存档于2016-04-04). 
  12. ^ 12.0 12.1 Cohen, R.; Persky, L.; Hadar, Y. Biotechnological applications and potential of wood-degrading mushrooms of the genus Pleurotus (PDF). Applied Microbiology and Biotechnology. 2002, 58 (5): 582–94. PMID 11956739. doi:10.1007/s00253-002-0930-y. [永久失效链接]
  13. ^ Vane, C. H., et al. (2003). "Biodegradation of Oak (Quercus alba) Wood during Growth of the Shiitake Mushroom (Lentinula edodes): A Molecular Approach." Journal of Agricultural and Food Chemistry 51(4): 947–956.
  14. ^ 14.0 14.1 Vane, C. H., et al. (2001). "Degradation of Lignin in Wheat Straw during Growth of the Oyster Mushroom (Pleurotus ostreatus) Using Off-line Thermochemolysis with Tetramethylammonium Hydroxide and Solid-State 13C NMR." Journal of Agricultural and Food Chemistry 49(6): 2709–2716.
  15. ^ Stamets, Paul. Chapter 2: The Role of Mushrooms in Nature. Growing gourmet and medicinal mushrooms = [Shokuyo oyobi yakuyo kinoko no sabai] 3rd. Berkeley, California, USA: Ten Speed Press. 2000: 9–11. ISBN 978-1-58008-175-7. 
  16. ^ Vane, C. H. (2003). "Monitoring Decay of Black Gum Wood (Nyssa sylvatica) During Growth of the Shiitake Mushroom (Lentinula edodes) Using Diffuse Reflectance Infrared Spectroscopy." Applied Spectroscopy 57(5): 514–517.

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木腐真菌
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