苔藓-蓝藻共生体关系与固氮能力研究进展

doi:10.17521/cjpe.2017.0191

摘要/Abstract

摘要：

苔藓-蓝藻共生体(BCS)能固氮, 是养分贫瘠地区森林氮输入的不可忽视的来源。BCS关系与固氮能力研究为科学认识生态系统氮输入与氮循环过程和机理提供了新的视角和有效途径, 具有重要的理论价值。然而, BCS关系、固氮作用与机理的研究迄今未受到足够关注, 报道较少, 认识仍然是零星而片段化的。基于系统查阅的相关文献, 该文综述了BCS的种类组成与共生关系类型、固氮能力及所固定氮的去向及其影响因素和作用机理, 指出了存在的问题及需要深入关注和亟待突破的4个研究方向。

关键词: 苔藓-蓝藻共生体, 固氮作用, 生态系统氮输入, 氮循环, 影响因素

Abstract:

Bryophyte-cyanobacteria symbiosis (BCS) is a key source of nitrogen input into ecosystems in nutrient-poor regions. Investigating BCS relationships and the nitrogen fixation capacity can be a new pathway and window to explore the process and mechanism of nitrogen input and nitrogen cycling. However, BCS relationships and nitrogen fixation/cycling processes and mechanisms remain poorly studied, and most of these studies have only focused on the boreal forest, with no report from Chinese forests. Based on systematic literature search and analysis, this review provides a summary on BCS relationships, the nitrogen fixation capability of BCS, the fate of fixed nitrogen, as well as the environmental factors and driving mechanisms of BCS. Firstly, we synthesized different types of BCS, the mechanisms by which the fixed nitrogen is transferred to and used by other plants within the forest, the rate of fixed nitrogen, the factors influencing the rate of nitrogen fixation. Moreover we point out the existing problems that need to pay close attention to and at least four research directions need to break through. Furthermore, the theoretical basis of BCS is provided for further research, promote and deepen the cognition of BCS and nitrogen-fixing research.

Key words: bryophyte-cyanobacteria symbiosis, nitrogen fixation, ecosystem nitrogen inputs, nitrogen cycle, driving factor

皮春燕, 刘鑫, 王喆, 包维楷. 苔藓-蓝藻共生体关系与固氮能力研究进展. 植物生态学报, 2018, 42(4): 407-418. DOI: 10.17521/cjpe.2017.0191

Chun-Yan PI, Xin LIU, Zhe WANG, Wei-Kai BAO. Bryophyte-cyanobacteria symbioses and their nitrogen fixation capacity—A review. Chinese Journal of Plant Ecology, 2018, 42(4): 407-418. DOI: 10.17521/cjpe.2017.0191

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https://www.plant-ecology.com/CN/Y2018/V42/I4/407

图/表 3

参考文献 80

1	Adams DG , Duggan PS ( 2008). Cyanobacteria-bryophyte symbioses. Journal of Experimental Botany, 59, 1047- 1058.
2	Adams DG , Duggan PS ( 2012). Signalling in Cyanobacteria-?Plant Symbioses. Signaling and Communication in Plant Symbiosis. Springer,Berlin. 11, 93- 121.
3	Arróniz-Crespo M , Pérez-Ortega S , De los Ríos A , Green TGA , Ochoa-Hueso R , Casermeiro Má , de la Cruz MT , Pintado A , Palacios D , Rozzi R , Tysklind N , Sancho LG ( 2014). Bryophyte-cyanobacteria associations during primary succession in recently deglaciated areas of Tierra del Fuego (Chile). PLOS ONE, 9, e96081. DOI: 10.1371/journal. pone.0096081. DOI URL PMID
4	Basilier K , Granhall U , Stenstr?m TA ( 1978). Nitrogen fixation in wet minerotrophic moss communities of a subarctic mire. Oikos, 31, 236- 246. DOI URL
5	Bay G , Nahar N , Oubre M , Whitehouse MJ , Wardle DA , Zackrisson O , Nilsson MC , Rasmussen U ( 2013). Boreal feather mosses secrete chemical signals to gain nitrogen. New Phytology, 200, 54- 60. DOI URL PMID
6	Berg A , Danielsson ? , Svensson BH ( 2012). Transfer of fixed-N from N₂-fixing cyanobacteria associated with the moss Sphagnum riparium results in enhanced growth of the moss. Plant and Soil, 362, 271- 278. DOI URL
7	Blundon DJ , Dale MRT ( 1990). Dinitrogen fixation (acetylene reduction) in primary succession near Mount Robson, British Columbia, Canada. Arctic Antarctic and Alpine Research, 22, 255- 263. DOI URL
8	Carleton TJ , Read DJ ( 1991). Ectomycorrhizas and nutrient transfer in conifer-feather moss ecosystems. Canadian Journal of Botany, 69, 778- 785. DOI URL
9	Chapin DM , Bliss LC , Bledsoe LJ ( 1991). Environment-?regulation of nitrogen-fixation in a high arctic lowland ecosystem. Canadian Journal of Botany, 69, 2744- 2755.
10	Chen Y , Fang DW ( 1983). Light-regulation of nitrogen fixation in the blue-green algae Anabaena 7120. Acta Phytophysiologia Sinica, 9, 51- 59.
	[ 陈因, 方大惟 ( 1983). 蓝藻Anabaena 7120固氮的光调节. 植物生理学报, 9, 51- 59.]
11	Cleveland CC , Townsend AR , Schimel DS , Fisher H , Howarth RW , Hedin LO , Perakis SS , Latty EF , Von Fischer JC , Elseroad A , Wasson MF ( 1999). Global patterns of terrestrial biological nitrogen (N₂) fixation in natural ecosystems. Global Biogeochemical Cycles, 13, 623- 645. DOI URL
12	Cornelissen JHC , Lang SI , Soudzilovskaia NA , During HJ ( 2007). Comparative cryptogam ecology: A review of bryophyte and lichen traits that drive biogeochemistry. Annals of Botany, 99, 987- 1001.
13	Coxson DS ( 1991). Nutrient release from epiphytic bryophytes in tropical montane rain forest (Guadeloupe). Canadian Journal of Botany, 69, 2122- 2129.
14	Davey A , Marchant HJ ( 1983). Seasonal variation in nitrogen fixation by Nostoc commune Vaucher at the Vestfold Hills, Antarctica. Phycologia, 22, 377- 385. DOI URL
15	Deluca T , Zackrisson O , Gundale M , Nilsson MC ( 2008). Ecosystem feedbacks and nitrogen fixation in boreal forests. Science, 320, 1181. DOI URL PMID
16	Deluca TH , Nilsson MC , Zackrisson O ( 2002a). Nitrogen mineralization and phenol accumulation along a fire chronosequence in northern Sweden. Oecologia, 133, 206- 214. DOI URL PMID
17	Deluca TH , Zackrisson O , Gentili F , Sellstedt A , Nilsson MC ( 2007). Ecosystem controls on nitrogen fixation in boreal feather moss communities. Oecologia, 152, 121- 130.
18	Deluca TH , Zackrisson O , Nilsson MC , Sellstedt A ( 2002b). Quantifying nitrogen-fixation in feather moss carpets of boreal forest. Nature, 419, 917- 920. DOI URL PMID
19	Du EZ , Liu XY , Fang JY ( 2014). Effects of nitrogen additions on biomass, stoichiometry and nutrient pools of moss Rhytidium rugosum in a boreal forest in northeast China. Environmental Pollution, 188, 166- 171. DOI URL PMID
20	Gavazov KS , Soudzilovskaia NA , van Logtestijn RSP , Braster M , Cornelissen JHC ( 2010). Isotopic analysis of cyanobacterial nitrogen fixation associated with subarctic lichen and bryophyte species. Plant and Soil, 333, 507- 517. DOI URL
21	Gentili F , Nilsson MC , Zackrisson O , Deluca TH , Sellstedt A ( 2005). Physiological and molecular diversity of feather moss associative N₂-fixing cyanobacteria. Journal of Experimental Botany, 56, 3121- 3127. DOI URL PMID
22	Gerber S , Hedin LO , Oppenheimer M , Pacala SW , Shevliakova E ( 2010). Nitrogen cycling and feedbacks in a global dynamic land model. Global Biogeochemical Cycles, 24, 723- 738. DOI URL
23	Granhall U , Basilier K ( 1973). Nitrogen fixation in tundra moss communities. Progress Report, Swedish IBP. Tundra Biome Project Technical Report, 14, 174- 190.
24	Granhall U , Selander H ( 1973). Nitrogen fixation in a subarctic mire. Oikos, 24, 8- 15.
25	Gundale MJ , Bach LH , Nordin A ( 2013). The impact of simulated chronic nitrogen deposition on the biomass and N₂-fixation activity of two boreal feather moss-cyanobacteria associations. Biology Letters, 9, 1- 4. DOI URL PMID
26	Gundale MJ , Deluca TH , Nordin A ( 2011). Bryophytes attenuate anthropogenic nitrogen inputs in boreal forests. Global Change Biology, 17, 2743- 2753.
27	Gundale MJ , Gustafsson H , Nilsson MC ( 2009). The sensitivity of nitrogen fixation by a feathermoss-cyanobacteria association to litter and moisture variability in young and old boreal forests. Canadian Journal of Forest Research, 39, 2542- 2549. DOI URL
28	Gundale MJ , Nilsson M , Bansal S , Jaderlund A ( 2012a). The interactive effects of temperature and light on biological nitrogen fixation in boreal forests. New Phytologist, 194, 453- 463. DOI URL PMID
29	Gundale MJ , Wardle DA , Nilsson MC ( 2012b). The effect of altered macroclimate on N-fixation by boreal feather mosses. Biology Letters, 8, 805- 808. DOI URL PMID
30	Han B , Zou XM , Kong JJ , Sha LQ , Gong HD , Yu Z , Cao T ( 2010). Nitrogen fixation of epiphytic plants enwrapping trees in Ailao Mountain cloud forests, Yunnan, China. Protoplasma, 247, 103- 110. DOI URL PMID
31	Houle D , Bilodeau GS , Paquet S , Planas D , Warren A ( 2006). Identification of two genera of N₂-fixing cyanobacteria growing on three feather moss species in boreal forests of Quebec, Canada. Canadian Journal of Botany, 84, 1025- 1029. DOI URL
32	Houlton BZ , Wang YP , Vitousek PM , Field CB ( 2008). A unifying framework for dinitrogen fixation in the terrestrial biosphere. Nature, 454, 327- 330. DOI URL PMID
33	Jackson BG , Martin P , Nilsson MC , Wardle DA ( 2011). Response of feather moss associated N₂ fixation and litter decomposition to variations in simulated rainfall intensity and frequency. Oikos, 120, 570- 581. DOI URL
34	Jones MLM , Wallace HL , Norris D , Brittain SA , Haria S , Jones RE , Rhind PM , Reynolds BR , Emmett BA ( 2004). Changes in vegetation and soil characteristics in coastal sand dunes along a gradient of atmospheric nitrogen deposition. Plant Biology, 6, 598- 605.
35	Lagerstrom A , Nilsson MC , Zackrisson O , Wardle DA ( 2007). Ecosystem input of nitrogen through biological ?xation in feather mosses during ecosystem retrogression. Functional Ecology, 21, 1027- 1033. DOI URL
36	Li YJ , Li J , Chen J , Huang GW ( 2004). Responses of Dolichomitriopsis diveirsiformis photosynthesis rate to light, air temperature and plant water content. Chinese Journal of Applied Ecology, 15, 391- 395.
	[ 李佑稷, 李菁, 陈军, 黄国文 ( 2004). 不同含水量下尖叶拟船叶藓光合速率对光温的响应及其模型. 应用生态学报, 15, 391- 395.]
37	Lindo Z , Whiteley JA ( 2011). Old trees contribute bio-available nitrogen through canopy bryophytes. Plant and Soil, 342, 141- 148. DOI URL
38	Liu JH , Bao WK , Li FL ( 2005). Major bryophyte patch structures and their relationships with environmental factors under a coniferous forest of eastern Tibetan plateau. Ecology and Environment, 14, 735- 741. DOI URL
	[ 刘俊华, 包维楷, 李芳兰 ( 2005). 青藏高原东部原始林下地表主要苔藓斑块特征及其影响因素. 生态环境学报, 14, 735- 741.] DOI URL
39	Liu X , Bao WK ( 2014). Understory plant assemblages present distinct short-term responses to the clear-cutting of an old-growth spruce forest near an alpine timberline. Canadian Journal of Forest Research, 44, 562- 571. DOI URL
40	Luo W , Shen JY ( 2007). Responses of growth of two Cyanobacteria species to different pH environments. Journal of Shanghai Jiaotong University (Agriculture Science), 25, 566- 569.
	[ 罗伟, 沈健英 ( 2007). 2种蓝藻的生长对不同环境pH的响应. 上海交通大学学报(农业科学版), 25, 566- 569.]
41	Markham JH ( 2009). Variation in moss-associated nitrogen fixation in boreal forest stands. Oecologia, 161, 353- 359. DOI URL PMID
42	Meeks JC ( 1990). Cyanobacterial-Bryophyte Associations. Chemical Rubber Company Press, Boca Raton.
43	Meeks JC ( 2007). Physiological adaptations in nitrogen-fixing Nostoc-plant symbiotic associations. In: Pawlowski K ed. Prokaryotic symbionts in plants. Microbiology Monographs. Vol. 8. Springer, Berlin.
44	Newmaster SG , Belland RJ , Arsenault A , Vitt DH ( 2003). Patterns of bryophyte diversity in humid coastal and inland cedar-hemlock forests of British Columbia. Environmental Reviews, 11, 159- 185.
45	Proctor M ( 2001). Patterns of desiccation tolerance and recovery in bryophytes. Plant Growth Regulation, 35, 147- 156.
46	Rai AN , Bergman B , Rasmussen U ( 2002). Cyanobacteria in Symbiosis. Springer,Berlin.
47	Rai AN , Trumbore SE , Davison EA , Harden JW , Veldhuis H ( 2000). Cyanobacterium-plant symbioses. New Phytologist, 174, 449- 481.
48	Reddy GB , Giddens J ( 1981). Nitrogen fixation by moss-algal associations in grassland. Soil Biology and Biochemistry, 13, 537- 538. DOI URL
49	Reed SC , Cleveland CC , Townsend AR ( 2011). Functional ecology of free-living nitrogen fixation: A contemporary perspective. Annual Review of Ecology, Evolution, and Systematics, 42, 489- 512. DOI URL
50	Rippka R , Deruelles J , Waterbury JB , Herdman M , Stanier RY ( 1979). Generic assignments, strain histories and properties of pure cultures of cyanobacteria. Journal of General Microbiology, 11, 1- 61.
51	Rosén K , Lindberg T ( 1980). Biological nitrogen fixation in coniferous forest watershed areas in central Sweden. Ecography, 3, 137- 140. DOI URL
52	Rousk K , Degboe J , Michelsen A , Bradley R , Bellenger JP ( 2017a). Molybdenum and phosphorus limitation of moss- associated nitrogen fixation in boreal ecosystems. New Phytologist, 214, 97- 107. DOI URL PMID
53	Rousk K , Deluca TH , Rousk J ( 2013a). The cyanobacterial role in the resistance of feather mosses to decomposition—?Toward a new hypothesis. PLOS ONE, 8, e62058. DOI: 10.1371/journal.pone.0062058. DOI URL PMID
54	Rousk K , Jones DL , Deluca TH ( 2013b). Exposure to nitrogen does not eliminate N₂ fixation in the feather moss Pleurozium schreberi (Brid.) Mitt. Plant and Soil, 374, 513- 521. DOI URL
55	Rousk K , Jones DL , Deluca TH ( 2013c). Moss-cyanobacteria associations as biogenic sources of nitrogen in boreal forest ecosystems. Frontiers in Microbiology, 4, 150. DOI URL PMID
56	Rousk K , Jones DL , Deluca TH ( 2013d). The resilience of nitrogen fixation in feather moss (Pleurozium schreberi)- cyanobacteria associations after a drying and rewetting cycle. Plant and Soil, 377, 159- 167. DOI URL
57	Rousk K , Michelsen A ( 2016). The sensitivity of moss- associated nitrogen fixation towards repeated nitrogen input. PLOS ONE, 11, e0146655. DOI: 10.1371/journal. pone.0146655. DOI URL PMID
58	Rousk K , Pedersen PA , Dyrnum K , Michelsen A ( 2017b). The interactive effects of temperature and moisture on nitrogen fixation in two temperate-arctic mosses. Theoretical and Experimental Plant Physiology, 29, 25- 36. DOI URL
59	Rousk K , Rousk J , Jones DL , Zackrisson O , Deluca TH ( 2013e). Feather moss nitrogen acquisition across natural fertility gradients in boreal forests. Soil Biology and Biochemistry, 61, 86- 95. DOI URL
60	Rousk K , Sorensen PL , Michelsen A ( 2016f). Nitrogen transfer from four nitrogen-fixer associations to plants and soils. Ecosystems, 19, 1491- 1504. DOI URL
61	Sanna ML , Antti JR , Marja T ( 2015). Nitrogen fixation in Sphagnum mosses is affected by moss species and water table level. Plant and Soil, 389, 185- 196. DOI URL
62	Smith VR ( 1984). Effects of abiotic factors on acetylene reduction by cyanobacteria epiphytic on moss at a subantarctic island. Applied and Environmental Microbiology, 48, 594- 600. DOI URL PMID
63	Sorensen PL , Michelsen A ( 2011). Long-term warming and litter addition affects nitrogen fixation in a subarctic heath. Global Change Biology, 17, 528- 537. DOI URL
64	Stewart KJ , Coxson D , Grogan P ( 2011a). Nitrogen inputs by associative cyanobacteria across a low arctic tundra landscape. Arctic Antarctic and Alpine Research, 43, 267- 278.
65	Stewart KJ , Lamb EG , Coxson DS , Siciliano SD ( 2011b). Bryophyte-cyanobacterial associations as a key factor in N₂-fixation across the Canadian Arctic. Plant and Soil, 344, 335- 346. DOI URL
66	Stewart WDP ( 1966). Nitrogen Fixation in Plants. Athlone Press, London.
67	Stuiver BM , Gundale MJ , Wardle DA , Nilsson MC ( 2015). Nitrogen fixation rates associated with the feather mosses Pleurozium schreberi and Hylocomium splendens during forest stand development following clear-cutting. Forest Ecology and Management, 347, 130- 139. DOI URL
68	Su YG , Li XR , Zhao X , Li AX , Chen YW ( 2011). The nitrogenase activity of biological soil crusts and their responses to environmental factors. Advances in Earth Science, 26, 332- 338.
	[ 苏延桂, 李新荣, 赵昕, 李爱霞, 陈应武 ( 2011). 不同类型生物土壤结皮固氮活性及对环境因子的响应研究. 地球科学进展, 26, 332- 338.]
69	Tamm CO ( 1991). Nitrogen in Terrestrial Ecosystems. Springer, Berlin.
70	Turetsky MR , Crow SE , Evans RJ , Vitt DH , Wieder RK ( 2008). Trade-offs in resource allocation among moss species control decomposition in boreal peatlands. Journal of Ecology, 96, 1297- 1305. DOI URL
71	Vitousek PM , Cassman K , Cleveland C ( 2002). The Nitrogen Cycle at Regional to Global Scales. Springer, Berlin. 1- 45. DOI URL
72	West NJ , Adams DG ( 1997). Phenotypic and genotypic comparison of symbiotic and free-living cyanobacteria from a single field site. Applied and Environmental Microbiology, 63, 4479- 4484. DOI URL PMID
73	Wilson JA , Coxson DS ( 1999). Carbon flux in a subalpine spruce-fir forest: Pulse release from Hylocomium splendens feather-moss mats. Canadian Journal of Botany, 77, 564- 569.
74	Yang L , Deng CC , Chen YM , He RL , Zhang J , Liu Y ( 2015). Relationships between decomposition rate of leaf litter and initial quality across the alpine timberline ecotone in Western Sichuan, China. Chinese Journal of Applied Ecology, 26, 3602- 3610.
	[ 杨林, 邓长春, 陈亚梅, 和润莲, 张健, 刘洋 ( 2015). 川西高山林线交错带凋落叶分解速率与初始质量的关系. 应用生态学报, 26, 3602- 3610.]
75	Zackrisson O , DeLluca TH , Gentili F , Sellstedt A , Jaderlund A ( 2009). Nitrogen fixation in mixed Hylocomium splendens moss communities. Oecologia, 160, 309- 319. DOI URL PMID
76	Zackrisson O , Deluca TH , Nilsson MC , Sellstedt A , Berglund LM ( 2004). Nitrogen fixation increases with successional age in boreal forests. Ecology, 85, 3327- 3334. DOI URL
77	Zhang Y , Guo LD ( 2007). Arbuscular mycorrhizal structure and fungi associated with mosses. Mycorrhiz, 17, 319- 325. DOI URL PMID
78	Zheng YP , Zhao JC , Zhang BC , Xu M , Liang HZ , Li M ( 2010). Influences of different physicochemical-chemical factors on three Cyanobacteria separated from biological soil crusts. Agriculture Research in the Arid Area, 28, 206- 211.
	[ 郑云普, 赵建成, 张丙昌, 徐明, 梁红柱, 李敏 ( 2010). 不同理化因子对荒漠生物结皮中三种蓝藻生长的影响. 干旱地区农业研究, 28, 206- 211.]
79	Zielke M , Ekker A , Olsen R , Spjelkavik S , Solheim B ( 2002). The influence of abiotic factors on biological nitrogen fixation in different types of vegetation in the high Arctic, Svalbard. Arctic Antarctic and Alpine Research, 34, 293- 299. DOI URL
80	Zielke M , Solheim B , Spjelkavik S ( 2005). Nitrogen fixation in the high arctic: Role of vegetation and environmental conditions. Arctic Antarctic and Alpine Research, 37, 372- 378. DOI URL

类型 Type	苔藓种类及与蓝藻关系 Bbryophyte and the relationship with cyanobacteria (En or Ep)	蓝藻种类 Cyanobacteria	参考文献 Reference
角苔-藻共生体 Hornwort-cyanobacteria symbiosis	角苔属 Anthoceros sp. (En 1) 树角苔属 Dendroceros sp. (En 1) 短角苔属 Notothylas sp. (En 1) 黄角苔属 Phaeoceros sp. (En 1)	1 念珠藻属 Nostoc sp.	Rippka et al., 1979; Meeks, 1990; West & Adams, 1997; Houle et al., 2006; Arróniz-Crespo et al., 2014
苔-藻共生体 Liverwort-cyanobacteria symbiosis	Anastrophyllum involutifolium (Ep1) 壶苞苔属 Blasia sp. (Ep 1, 2, 4; En 1) Cavicularia sp. (En 1） Chiloscyphus leptanthus (Ep 1) 地钱属 Marchantia sp. (Ep 1) 光萼苔 Porella sp. (Ep 1)	1 念珠藻属 Nostoc sp. 2 眉藻属 Calothrix sp. 3 真枝藻属 Stigonema sp. 4 Chlorogloeopsis sp.	Rippka et al., 1979; Meeks, 1990; West & Adams, 1997; Houle et al., 2006; Arróniz-Crespo et al., 2014
藓-藻共生体 Moss-cyanobacteria symbiosis	Acroschisma wilsonii (Ep 1) Andreaea alpine (Ep 1) Andreaea laxifolia (Ep 1) 皱缩藓 Aulacomnium palustre (Ep 1) Blepharidophyllum densifolium (Ep 1) 真藓属 Bryum sp. (Ep 1, 5, 6 7) Calliergon richardsonii (Ep 1) 角齿藓 Ceratodon purpureus (Ep 5, 6, 7) Clasmatocolea humilis (Ep 1) Cryptochila grandiflora (Ep 1) Dendroligotrichum squamosum (Ep 1) Dicranoloma chilense (Ep1） Ditrichum cylindricarpum (Ep 1) 镰刀藓属 Drepanocladus sp. (Ep 1, 2, 4) Drepanocladus exannulatus (Ep 1) Dupontia sp. (Ep 1) 紫萼藓属 Grimmia sp. (Ep 1) Heteroscyphus magellanicus (Ep 1) 塔藓 Hylocomium splendens (Ep 1, 3) 沼寒藓 Paludella squarrosa (Ep 1) 赤茎藓 Pleurozium schreberi (Ep 1, 3) 毛梳藓 Ptilium sp. (Ep 1, 3) Racomitrium Subcrispipilum (Ep 1) Racomitrium laevigatum (Ep 1) 白毛砂藓 Racomitrium lanuginosum (Ep 1) Racomitrium didymium (Ep 1) 三洋藓 Sanionia uncinata (EP 1) Sphagnum lindebergii (Ep 1; En 1.1) 岸生泥炭藓 Sphagnum riparium (Ep 1; En 1.1) 毛青藓 Tomentypnum nitens (Ep 1) 小石藓 Weisia controversa (Ep 5, 6, 7)	1 念珠藻属 Nostoc sp. 1.1 灰色念珠藻 Nostoc muscorum 2 眉藻属 Calothrix sp. 3 真枝藻属 Stigonema sp. 4 伪枝藻属 Scytonema sp. 5 Anobena sp. 6 颤藻属Oscillatoria sp. 7 鞘丝藻属 Lyngbya sp.	Granhall & Selander, 1973b; Rippka et al., 1979; Reddy & Giddens, 1981; Davey & Marchant, 1983; Meeks, 1990; West & Adams, 1997; Gentili et al., 2005; Houle et al., 2006; Gavazov et al., 2010; Arróniz-Crespo et al., 2014
小计 Total	41	8

类型 Type	苔藓种类及与蓝藻关系 Bbryophyte and the relationship with cyanobacteria (En or Ep)	蓝藻种类 Cyanobacteria	参考文献 Reference
角苔-藻共生体 Hornwort-cyanobacteria symbiosis	角苔属 Anthoceros sp. (En 1) 树角苔属 Dendroceros sp. (En 1) 短角苔属 Notothylas sp. (En 1) 黄角苔属 Phaeoceros sp. (En 1)	1 念珠藻属 Nostoc sp.	Rippka et al., 1979; Meeks, 1990; West & Adams, 1997; Houle et al., 2006; Arróniz-Crespo et al., 2014
苔-藻共生体 Liverwort-cyanobacteria symbiosis	Anastrophyllum involutifolium (Ep1) 壶苞苔属 Blasia sp. (Ep 1, 2, 4; En 1) Cavicularia sp. (En 1） Chiloscyphus leptanthus (Ep 1) 地钱属 Marchantia sp. (Ep 1) 光萼苔 Porella sp. (Ep 1)	1 念珠藻属 Nostoc sp. 2 眉藻属 Calothrix sp. 3 真枝藻属 Stigonema sp. 4 Chlorogloeopsis sp.	Rippka et al., 1979; Meeks, 1990; West & Adams, 1997; Houle et al., 2006; Arróniz-Crespo et al., 2014
藓-藻共生体 Moss-cyanobacteria symbiosis	Acroschisma wilsonii (Ep 1) Andreaea alpine (Ep 1) Andreaea laxifolia (Ep 1) 皱缩藓 Aulacomnium palustre (Ep 1) Blepharidophyllum densifolium (Ep 1) 真藓属 Bryum sp. (Ep 1, 5, 6 7) Calliergon richardsonii (Ep 1) 角齿藓 Ceratodon purpureus (Ep 5, 6, 7) Clasmatocolea humilis (Ep 1) Cryptochila grandiflora (Ep 1) Dendroligotrichum squamosum (Ep 1) Dicranoloma chilense (Ep1） Ditrichum cylindricarpum (Ep 1) 镰刀藓属 Drepanocladus sp. (Ep 1, 2, 4) Drepanocladus exannulatus (Ep 1) Dupontia sp. (Ep 1) 紫萼藓属 Grimmia sp. (Ep 1) Heteroscyphus magellanicus (Ep 1) 塔藓 Hylocomium splendens (Ep 1, 3) 沼寒藓 Paludella squarrosa (Ep 1) 赤茎藓 Pleurozium schreberi (Ep 1, 3) 毛梳藓 Ptilium sp. (Ep 1, 3) Racomitrium Subcrispipilum (Ep 1) Racomitrium laevigatum (Ep 1) 白毛砂藓 Racomitrium lanuginosum (Ep 1) Racomitrium didymium (Ep 1) 三洋藓 Sanionia uncinata (EP 1) Sphagnum lindebergii (Ep 1; En 1.1) 岸生泥炭藓 Sphagnum riparium (Ep 1; En 1.1) 毛青藓 Tomentypnum nitens (Ep 1) 小石藓 Weisia controversa (Ep 5, 6, 7)	1 念珠藻属 Nostoc sp. 1.1 灰色念珠藻 Nostoc muscorum 2 眉藻属 Calothrix sp. 3 真枝藻属 Stigonema sp. 4 伪枝藻属 Scytonema sp. 5 Anobena sp. 6 颤藻属Oscillatoria sp. 7 鞘丝藻属 Lyngbya sp.	Granhall & Selander, 1973b; Rippka et al., 1979; Reddy & Giddens, 1981; Davey & Marchant, 1983; Meeks, 1990; West & Adams, 1997; Gentili et al., 2005; Houle et al., 2006; Gavazov et al., 2010; Arróniz-Crespo et al., 2014
小计 Total	41	8

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