丛枝菌根真菌在地球圈层中的功能多样性

1、石石 兆兆 勇勇2008年年12月月26日日生态讲坛生态讲坛北京大学生态系北京大学生态系 地下结构地下结构 地下菌丝体的分布地下菌丝体的分布互惠关系互惠关系宿主宿主- -菌根真菌菌根真菌（Smith & Read, 1997）70%的磷的磷（George et al.,1995）30%的氮的氮（Frey et al.,1993） 光合产物总量光合产物总量的的4%-20%（Peng et al.,1993） 丛枝菌根（丛枝菌根（AM菌根）菌根）Kinden & Brown, 1975Brundrett, 1984Shi et al. , 2007Shi et al. , 2007

2、(Read 1991; Read et al. 2004).Gai et al., 2006.Cyperaceae丛枝菌根在地球上的分布丛枝菌根在地球上的分布丛枝菌根丛枝菌根地球最广泛的共生体地球最广泛的共生体（Brachmann & Parniske 2006; Brachmann & Parniske 2006; 石兆勇石兆勇等，等，20072007） 具有丛枝菌根的植物占具有丛枝菌根的植物占The majority of plants, strictly speaking,do not have roots; they have mycorrhizas. 地球上最广泛的共

3、生体地球上最广泛的共生体丛枝菌根丛枝菌根The study of plants without their mycorrhizasis the study of artefacts. 地下生态学研究地下生态学研究菌根菌根http:/www.kent.ac.uk/Light microscope picture of a fossil fungal spore with attached hypha from the Ordovician, 460 million years ago. (c)Science Magazine CREDIT: Dirk Redecker/UC BerkeleyIn

4、 addition, mycorrhizal fungi have been found in fossils as old as 400 million years, in company with later, more evolved vascular plants. Light microscope picture of a fossil fungal spore with attached hypha from the Ordovician, 460 million years ago. (c)Science Magazine CREDIT: Dirk Redecker/UC Ber

5、keleyLight microscope picture of a fossil fungal spore with attached hypha from the Ordovician, 460 million years ago. (c)Science Magazine CREDIT: Dirk Redecker/UC Berkeley内生菌根内生菌根 外生菌根外生菌根登陆：帮助生物体由海洋登上陆地登陆：帮助生物体由海洋登上陆地扩大养分的吸收面积帮助植物吸收 养分帮助植物获取食物生存Contribution of AMF to plant nutrient uptake Macronut

6、rients:P30 92 % (Kothari et al 1991, Yao et al 2004, Sally et al 2005)N20 42 % (Ames et al. 1983, Marschner & Dell 1994, Mder et al. 2000)K0 -10 % (?)(Smith et al. 1981, George et al. 1992, Smith & Read 1997) Micronutrients:Zn30 80 % ( Li et al 1991, Brkert & Robson 1994)Cu 20 62 % (Li e

7、t al. 1991, Lambert & Weidensaul 1991)Mn, Fe, B 0 -70 % (Lambert & Weidensaul 1991, Kothari et al. 1991)AM真菌对宿主植物养分吸收的贡献真菌对宿主植物养分吸收的贡献From Li X L菌根侵染诱导植物根细胞专一性菌根侵染诱导植物根细胞专一性P转运子转运子mycorrhizalnon-mycorrhizalBased on papers by Rosewarne et al 1999; Harrison and van Buuren 1995; Rausch et al. 2

8、002; Maldonado-Mendoza et al. 2002; Paszkowski et al. 2002；S Smith 2005Glomus intraradices, GiPT Harrison and Van Buuren,1995 NatureGlomus versiforme, GvPT Maldonado-Mendozaet al., 2001 MPMIGlomus mosseae GmosPT Benedetto et al., 2005 MycorrhizalSolanum tuberosum StPT3 StPT4 Rausch et al., 2001 Natu

9、re; Karandashov et al., 2004 PNASMedicago truncatula MtPT4 Harrison et al., 2002 Plant Cell; Liu, et al., 1998 MPMIOryza sativa ORYsa;Pht1;1 Paszkowski et al., 2002 PNASLotus japonicus, LjPT3 Maeda et al., 2006 Plant Cell Physiol.CFitter et al. 1998NPBethlenfalvay et al. 1991物质的传递物质的传递联盟互为帮助联盟互为帮助资源

10、共享资源共享菌丝桥菌丝桥Marschner 植物间物质和信号的传递地下菌丝网络 提供防御：帮助植物战胜逆境在菌丝际在菌丝际与病原菌与病原菌进行战斗，进行战斗，距病菌于距病菌于根际之外根际之外植物王国植物王国的远征部的远征部队队类别名称真菌病害枯萎病、根腐病、黑水病原生动物病害线虫病害细菌病害猝倒病、根癌病、斑点病提高植物的抗/耐病性+AM1 +AM2 -AM枯萎病枯萎病Wang et al. 2004+ AM- AM杀死线虫的功能提高植物对重金属的抗性提高植物对重金属的抗性CKAM0 10 50 100 200As的浓度（的浓度（mg/kg）Xu et al. 2004(Feng et al.

11、 2002; Tian et al. 2004)抗抗/耐盐性耐盐性抗抗/耐旱性耐旱性Shi et al. unpublishedNieves Goicoechea et al.，2004t：毛状体s:气孔w：上表皮蜡状物抗抗/耐耐旱旱机机制制不不接接种种AM接接种种AMvan der Heijden, M.G.A., Klironomos, J.N., Ursic, M., Moutoglis, P., Streitwolf-Engel, R., Boller, T., Wiemken, A., Sanders, I.R., 1998. Nature 396, 6972.决定植物生态系统的生物

12、多样性、变异性和生产力Mycorrhizal fungal diversity determines plant biodiversity, ecosystem variability and productivityArbuscular mycorrhizal fungi promote plant diversity and productivityVANDERHEIJDENetal.，Nature1998CommentarybyDavidRead:Plantsontheweb.From Zhu改善土壤的团粒结构 -M G.m G.v菌根分泌物对土壤团聚体的影响菌根分泌物对土壤团聚体的影响

13、尼龙网上粘结土壤重量尼龙网上粘结土壤重量(g)3.1 4.3 3.6From Li X L菌根菌对土壤有机质的贡献 投入土壤有机质 0.860 g kg-1 y-1菌丝碳 0.049 g kg-1 y-1菌根素碳 0.005 g kg-1 y-1菌根菌总碳 0.051 g kg-1 y-1菌根菌碳占土壤总碳：5.9 % 滞留时间 (y)土壤有机碳 111 细根 6.1菌丝碳 2.4 快-菌根素碳 15慢-菌根素碳 200提高土壤的有机质提高土壤的有机质加速岩石的矿质化加速岩石的矿质化(Masanori and Takuya 2002)Mt. Fugendake, Japan.Pang &

14、; Paul, 1980; Paul & Kucey, 1981; Kucey & Paul, 1982; Snellgrove et al., 1982; Koch & Johnson, 1984; Harris et al., 1985; Douds et al., 1988; Wang et al., 1989; Jakobsen& Rosendahl, 1990; Pearson & Jakobsen, 1993; Peng et al., 1993. Bago et al., 2000; Douds et al., 2002; Graham, 2004The C cost of AM symbiosis as between 4% and 20% of the plants C ph