Research progress in promoting symbiotic germination mechanism of Orchidaceae plant seeds and its industrial application

doi:10.16178/j.issn.0528-9017.20230904

Abstract

Abstract:

Most Orchidaceae plants have bright and highly ornamental flowers, some of which are often used for medicinal purposes, and therefore have important social, economic, and medicinal value. However, its seeds are small and have no endosperm, so it is difficult to germinate and grow normally under natural conditions. Most of them are bred rapidly through vegetative reproduction or artificial tissue culture. However, in special circumstances, some orchid plants can also achieve seed reproduction under natural conditions, where the symbiosis of endophyte helps the seeds to grow normally. During this process, endophyte provide nutrients, hormones, and enhance the plant's resistance to environmental stress for the seeds, scientifically promote the establishment of such a good collaborative relationship, which is conducive to the rapid and efficient cultivation of high-quality Orchidaceae plant seedlings. Therefore, this article reviews the mechanism of seed symbiotic germination of Orchidaceae plants through research and analysis of their interactions with endophyte. Taking Orchidaceae plants such as Gastrodia elata Bl,Bletilla striata and Dendrobium officinale as examples, the practical production and application value of symbiotic germination technology in industrialization is explained, achieving the protection of Orchidaceae plant germplasm resources and promoting their scientific development and utilization.

Key words: endophyte, Orchidaceae plants, symbiotic germination, industrial application

CLC Number:

S682.31

YE Yueyu, LI Xiaomei, CHEN Man, WANG Xia, JIANG Fusheng, ZHANG Chunchun, CHEN Haimin, LI Shiqing. Research progress in promoting symbiotic germination mechanism of Orchidaceae plant seeds and its industrial application[J]. Journal of Zhejiang Agricultural Sciences, 2024, 65(12): 2944-2953.

Figures/Tables 1

References 76

[1]	丁绍武, 张鹏, 刘梦铭. 植物内生菌对植物生长的影响研究进展[J]. 现代农业科技, 2020, 21(11): 132-134.
[2]	CHRISTENHUSZ M J M, BYNG J W. The number of known plants species in the world and its annual increase[J]. Phytotaxa, 2016, 261(3): 201.
[3]	蔡媛, 刘浩, 孔文平, 等. 多花黄精内生菌群落结构多样性及其与有效成分含量相关性研究[J]. 中草药, 2021, 52(13): 4023-4030.
[4]	陈金阳, 陆儒涵, 王玲, 等. 药用植物内生菌抗氧化活性研究进展[J]. 中草药, 2016, 47(20): 3720-3727.
[5]	杨建文, 陈晓梅, 孟志霞, 等. 胶膜菌属真菌S7(Tulasnella sp.) 及提取物对铁皮石斛种子萌发的影响[J]. 中国科学(生命科学), 2020, 50(5): 559-570.
[6]	SMITH S E, READ D J. Mycorrhizal symbiosis[J]. Quarterly Review of Biology, 2008, 3(3): 273-281.
[7]	涂然, 武坤毅, 冯正平, 等. 同一生境下白及与黄花白及内生菌群差异的初步研究[J]. 广西植物, 2021, 41(7): 1173-1180.
[8]	郑超文, 肖娅萍. 兰科菌根真菌研究方法的概述[J]. 微生物学杂志, 2014, 34(4): 85-89.
[9]	张镇梁, 王美娜, 李健, 等. 管叶槽舌兰新鲜及硅胶干燥根样内生菌多样性研究[J]. 广西植物, 2023, 43(6): 991-1005.
[10]	高阳. 金钗石斛内生菌多样性与石斛碱含量的相关性研究[D]. 贵阳: 贵州师范大学, 2017.
[11]	DEARNALEY J D W, MARTOS F, SELOSSE M A. 12 orchid mycorrhizas: molecular ecology, physiology, evolution and conservation aspects[M]//HOCK B. Fungal associations. Berlin, Heidelberg: Springer, 2012: 207-230.
[12]	RASMUSSEN H N, DIXON K W, JERSÁKOVÁ J, et al. Germination and seedling establishment in orchids: a complex of requirements[J]. Annals of Botany, 2015, 116(3): 391-402.
[13]	李佳瑶, 赵泽宇, 高越, 等. 药用植物绶草(兰科)的菌根真菌群落组成分析[J]. 菌物学报, 2021, 40(6): 1317-1327.
[14]	孙晓颖. 五种野生兜兰植物菌根真菌多样性研究[D]. 北京: 北京林业大学, 2014.
[15]	高越, 郭顺星, 邢晓科. 兰科植物种子共生萌发真菌多样性及共生萌发机制研究进展[J]. 菌物学报, 2019, 38(11): 1808-1825.
[16]	MCCORMICK M K, WHIGHAM D F, O'NEILL J. Mycorrhizal diversity in photosynthetic terrestrial orchids[J]. New Phytologist, 2004, 163(2): 425-438.
[17]	李孟凯, 牛昱龙, 杨文娟, 等. 西藏野生兰科植物内生真菌多样性与共生萌发研究[J]. 高原农业, 2020, 4(6): 580-584, 653.
[18]	WILKINSON K G, DIXON K W, SIVASITHAMPARAM K, et al. Effect of IAA on symbiotic germination of an Australian orchid and its production by orchid-associated bacteria[J]. Plant and Soil, 1994, 159(2): 291-295.
[19]	WILKINSON K G, SIVASITHAMPARAM K, DIXON K W, et al. Identification and characterisation of bacteria associated with Western Australian orchids[J]. Soil Biology and Biochemistry, 1994, 26(1): 137-142.
[20]	WILKINSON K G, DIXON K W, SIVASITHAMPARAM K. Interaction of soil bacteria, mycorrhizal fungi and orchid seed in relation to germination of Australian orchids[J]. New Phytologist, 1989, 112(3): 429-435.
[21]	TSAVKELOVA E A, CHERDYNTSEVA T A, KLIMOVA S Y, et al. Orchid-associated bacteria produce indole-3-acetic acid, promote seed germination, and increase their microbial yield in response to exogenous auxin[J]. Archives of Microbiology, 2007, 188(6): 655-664.
[22]	赵银. 白及优质种苗繁育技术及其促生内生菌的筛选和鉴定[D]. 杭州: 浙江大学, 2020.
[23]	潘丽晶, 陈继敏, 张妙彬, 等. 蝴蝶兰根内生细菌的分离及可分泌IAA细菌的筛选[J]. 中国农学通报, 2014, 30(16): 148-152.
[24]	张芳芳. 五唇兰根部内生细菌筛选及其促生效应研究[D]. 海口: 海南大学, 2015.
[25]	WU Y S, XIAO S H, QI J S, et al. Pseudomonas fluorescens BsEB-1: an endophytic bacterium isolated from the root of Bletilla striata that can promote its growth[J]. Plant Signaling & Behavior, 2022, 17(1): 2100626.
[26]	陈耀丽, 俞龙春, 钱悦, 等. 大尖囊蝴蝶兰内生真菌和细菌的分离与鉴定[J]. 热带生物学报, 2019, 10(4): 372-379.
[27]	曾旭, 杨建文, 凌鸿, 等. 天麻种子与真菌共生萌发的蛋白组学研究[J]. 菌物学报, 2018, 37(1): 64-72.
[28]	FAN J, CHEN C X, YU Q B, et al. Comparative iTRAQ proteome and transcriptome analyses of sweet orange infected by “Candidatus Liberibacter asiaticus”[J]. Physiologia Plantarum, 2011, 143(3): 235-245.
[29]	WARCUP J H. Specificity of mycorrhizal association in some Australian terrestrial orchids[J]. New Phytologist, 1971, 70(1): 41-46.
[30]	YOUM J Y, HAN H K, CHUNG J M, et al. Identification of orchid mycorrhizal fungi isolated from five species of terrestrial orchids in Korea[J]. The Korean Journal of Mycology, 2012, 40(3): 132-135.
[31]	ZI X M, SHENG C L, GOODALE U M, et al. In situ seed baiting to isolate germination-enhancing fungi for an epiphytic orchid, Dendrobium aphyllum (Orchidaceae)[J]. Mycorrhiza, 2014, 24(7): 487-499.
[32]	HUANG H, ZI X M, LIN H, et al. Host-specificity of symbiotic mycorrhizal fungi for enhancing seed germination, protocorm formation and seedling development of over-collected medicinal orchid, Dendrobium devonianum[J]. Journal of Microbiology, 2018, 56(1): 42-48.
[33]	许璐, 田佳妮, 王涛, 等. 胶膜菌属真菌与兰科植物的共生体系建立[J]. 核农学报, 2017, 31(5): 876-883.
[34]	徐玲玲, 张焱, 赵明阳, 等. 菌根真菌对白及种子萌发和幼苗生根的作用[J]. 菌物学报, 2019, 38(9): 1440-1449.
[35]	ILLYÉS Z, RUDNÓY S, BRATEK Z. Aspects of in situ, in vitro germination and mycorrhizal partners of Liparis loeselii[J]. Acta Biologica Szegediensis, 2005, 49(1-2): 137-139.
[36]	BONNARDEAUX Y, BRUNDRETT M, BATTY A, et al. Diversity of mycorrhizal fungi of terrestrial orchids: compatibility webs, brief encounters, lasting relationships and alien invasions[J]. Mycological Research, 2007, 111(Pt 1): 51-61.
[37]	ROCHE S A, CARTER R J, PEAKALL R, et al. A narrow group of monophyletic Tulasnella (Tulasnellaceae) symbiont lineages are associated with multiple species of Chiloglottis (Orchidaceae): implications for orchid diversity[J]. American Journal of Botany, 2010, 97(8): 1313-1327.
[38]	RAFTER M, YOKOYA K, SCHOFIELD E J, et al. Non-specific symbiotic germination of Cynorkis purpurea (Thouars) Kraezl., a habitat-specific terrestrial orchid from the Central Highlands of Madagascar[J]. Mycorrhiza, 2016, 26(6): 541-552.
[39]	STÖCKEL M, TěŠITELOVÁ T, JERSÁKOVÁ J, et al. Carbon and nitrogen gain during the growth of orchid seedlings in nature[J]. New Phytologist, 2014, 202(2): 606-615.
[40]	MCKENDRICK S L, LEAKE J R, TAYLOR D L, et al. Symbiotic germination and development of myco-heterotrophic plants in nature: ontogeny of Corallorhiza trifida and characterization of its mycorrhizal fungi[J]. New Phytologist, 2000, 145(3): 523-537.
[41]	FRACCHIA S, ARANDA-RICKERT A, FLACHSLAND E, et al. Mycorrhizal compatibility and symbiotic reproduction of Gavilea australis, an endangered terrestrial orchid from south Patagonia[J]. Mycorrhiza, 2014, 24(8): 627-634.
[42]	MCKENDRICK S L, LEAKE J R, TAYLOR D L, et al. Symbiotic germination and development of the myco-heterotrophic orchid Neottia nidus-avis in nature and its requirement for locally distributed Sebacina spp[J]. New Phytologist, 2002, 154(1): 233-247.
[43]	GAO Y Y, JI J, ZHANG Y J, et al. Biochemical and transcriptomic analyses of the symbiotic interaction between Cremastra appendiculata and the mycorrhizal fungus Coprinellus disseminatus[J]. BMC Plant Biology, 2022, 22(1): 15.
[44]	TIAN F, LIAO X F, WANG L H, et al. Isolation and identification of beneficial orchid mycorrhizal fungi in Paphiopedilum barbigerum (Orchidaceae)[J]. Plant Signaling & Behavior, 2022, 17(1): 2005882.
[45]	CHUTIMA R, DELL B, VESSABUTR S, et al. Endophytic fungi from Pecteilis susannae (L.) Rafin (Orchidaceae), a threatened terrestrial orchid in Thailand[J]. Mycorrhiza, 2011, 21(3): 221-229.
[46]	ATHIPUNYAKOM P, MANOCH L, PILUEK C, et al. Mycorrhizal fungi from Spathoglottis plicata and the use of these fungi to germinate seeds of S. plicata in vitro[J]. Agriculture and Natural Resources, 2004, 38(1): 83-93.
[47]	STEWART S L, KANE M E. Symbiotic seed germination of Habenaria macroceratitis (Orchidaceae), a rare Florida terrestrial orchid[J]. Plant Cell, Tissue and Organ Culture, 2006, 86(2): 159-167.
[48]	CHEN J, WANG H, LIU S S, et al. Ultrastructure of symbiotic germination of the orchid Dendrobium officinale with its mycobiont, Sebacina sp[J]. Australian Journal of Botany, 2014, 62(3): 229.
[49]	范黎, 郭顺星, 肖培根. 密花石斛等六种兰科植物菌根的显微结构研究[J]. 植物学通报, 2000, 17 (1): 73.
[50]	SHUBHA J, SRINIVAS C. Diversity and extracellular enzymes of endophytic fungi associated with Cymbidium aloifolium L.[J]. Academic Journals, 2017, 16(48): 2248-2258.
[51]	唐燕静. 药用石斛种子与菌根真菌共生萌发专一性及其作用机制初探[D]. 北京: 北京协和医学院, 2021.
[52]	RICARDO M J, ALVAREZ M R. Ultrastructural changes associated with utilization of metabolite reserves and trichome differentiation in the protocorm of Vanda[J]. American Journal of Botany, 1971, 58(3): 229.
[53]	FOCHI V, CHITARRA W, KOHLER A, et al. Fungal and plant gene expression in the Tulasnella calospora-Serapias vomeracea symbiosis provides clues about nitrogen pathways in orchid mycorrhizas[J]. New Phytologist, 2017, 213(1): 365-379.
[54]	KUGA Y, SAKAMOTO N, YURIMOTO H. Stable isotope cellular imaging reveals that both live and degenerating fungal pelotons transfer carbon and nitrogen to orchid protocorms[J]. New Phytologist, 2014, 202(2): 594-605.
[55]	BOUGOURE J J, BRUNDRETT M C, GRIERSON P F. Carbon and nitrogen supply to the underground orchid, Rhizanthella gardneri[J]. New Phytologist, 2010, 186(4): 947-956.
[56]	BARRA-BUCAREI L, GONZÁLEZ M G, IGLESIAS A F, et al. Beauveria bassiana multifunction as an endophyte: growth promotion and biologic control of Trialeurodes vaporariorum, (Westwood) (Hemiptera: Aleyrodidae) in tomato[J]. Insects, 2020, 11(9): 591.
[57]	童文君. 美花石斛内生菌多样性分析及促生潜力研究[D]. 南京: 南京师范大学, 2014.
[58]	HUREK T, HANDLEY L L, REINHOLD-HUREK B, et al. Azoarcus grass endophytes contribute fixed nitrogen to the plant in an unculturable state[J]. Molecular Plant-Microbe Interactions, 2002, 15(3): 233-242.
[59]	孙磊, 邵红, 刘琳, 等. 可产生铁载体的春兰根内生细菌多样性[J]. 微生物学报, 2011, 51(2): 189-195.
[60]	张萍, 宋希强. 兰科植物内生细菌物种多样性及其促生机理研究进展[J]. 热带亚热带植物学报, 2012, 20(1): 92-98.
[61]	CHEN X M, DONG H L, HU K X, et al. Diversity and antimicrobial and plant-growth-promoting activities of endophytic fungi in Dendrobium loddigesii Rolfe[J]. Journal of Plant Growth Regulation, 2010, 29(3): 328-337.
[62]	裴东方. 盘龙参内生菌多样性及一株内生细菌的抑菌活性研究[D]. 荆州: 长江大学, 2020.
[63]	程萍, 郑燕玲, 黎永坚, 等. 石斛兰镰刀菌叶斑病的生物防治研究[J]. 中国农学通报, 2008, 24(9): 357-361.
[64]	何劲, 雷帮星, 康冀川, 等. 石斛抗真菌内生细菌的筛选及其抗菌特性的初步研究[J]. 西南大学学报(自然科学版), 2009, 31(6): 92-96.
[65]	VALADARES R B S, PEROTTO S, SANTOS E C, et al. Proteome changes in Oncidium sphacelatum (Orchidaceae) at different trophic stages of symbiotic germination[J]. Mycorrhiza, 2014, 24(5): 349-360.
[66]	国家药典委员会. 中华人民共和国药典-一部: 2020年版[M]. 北京: 中国医药科技出版社, 2020.
[67]	张照宇, 孙建华, 陈士林, 等. 天麻种质资源及其与双菌共生分子机制研究[J]. 世界中医药, 2022, 17(13): 1819-1826.
[68]	刘华, 谭德仁, 曾祥福, 等. 菌材伴栽与天麻高产栽培技术研究[J]. 湖北林业科技, 2006, 35(4): 14-16.
[69]	郭顺星, 王秋颖. 促进天麻种子萌发的石斛小菇优良菌株特性及作用[J]. 菌物系统, 2001, 20(3): 408-412.
[70]	王秋颖, 郭顺星, 关凤斌. 不同来源蜜环菌对天麻产量影响的研究[J]. 中草药, 2001, 32(9): 839-841.
[71]	王永, 王彩云, 侯俊, 等. 不同蜜环菌菌株对红天麻农艺性状、产量及质量的影响[J]. 微生物学杂志, 2020, 40(6): 59-65.
[72]	赵麒鸣, 吴鹏, 刘鸿高, 等. 蜜环菌与天麻的共生关系研究进展[J]. 云南农业科技, 2022(2): 56-58.
[73]	张升明, 王绍柏. 从市场需求角度谈天麻产业发展趋势与建议[J]. 食药用菌, 2022, 30(1): 7-10.
[74]	王兆春. 铁皮石斛接种枝孢霉菌的响应机制及其种子萌发关键环境因子研究[D]. 重庆: 西南大学, 2019.
[75]	杨社峰. 金沙江石斛传粉生物学及种子共生萌发研究[D]. 昆明: 云南大学, 2018.
[76]	字肖萌, 高江云. 不同真菌对2种药用石斛种子共生萌发的效应[J]. 中国中药杂志, 2014, 39(17): 3238-3244.

内生菌	促萌发内生细菌种属	兰科植物种属	文献
内生细菌	恶臭假单胞菌Pseudomonas putida	翅柱兰属Pterostylis vittata	[18]
	嗜麦芽黄单胞菌Xanthomonas maltophilia		[18]
	蜡样芽孢杆菌Bacillus cereus		[20]
	戴氏菌属Dyella	蝴蝶兰Phalaenopsis amabilis	[23]
	鞘氨醇菌属Sphingomonas	杓唇石斛Dendrobium moschatum(Buch.-Ham.)Sw.	[21]
	分枝菌属Mycobacterium		[21]
	贝莱斯芽孢杆菌Bacillus velezensis	白及Bletilla striata	[21]
	芽孢杆菌属Bacillus megaterium	五唇兰Phalaenopsis pulcherrima (Lindl) J.J. Sm.	[24]
	芽孢杆菌属Bacillus mycoides
	芽孢杆菌属Bacillus cereus
	荧光假单胞菌Pseudomonas fluorescens	白及Bletilla striata	[25]
内生真菌	镰刀菌属Fusarium Link ex Fr.	铁皮石斛Dendrobium officinale Kimura et Migo	[26]
	木霉属Trichoderma	华石斛Dendrobium sinense	[26]
	紫箕小菇Mycena osmundicola J.E.Lange	天麻Gastrodia elata	[27]
	石斛小菇Mycena dendrobii L.Fan et S.X.Guo		[27]
	兰小菇Mycena orchidicola Fan et Guo		[27]
	兰小菇Mycena orchidicola Fan et Guo	报春石斛Dendrobium polyanthum Wall. ex Lindl.	[28]
		长苏石斛Dendrobium brymerianum Rchb. f.	[28]
		鼓槌石斛Dendrobium chrysotoxum	[28]
		密花石斛Dendrobium densiflorum Lindl.	[28]
	胶膜菌属Tulasnella	驴兰Diuris	[29]
		虾脊兰属Calanthe	[30]
		兜唇石斛Dendrobium aphyllum	[31]
		硬叶兰Cymbidium mannii	[32]
		树兰Epidnedium	[33]
		白及Bletilla striata	[34]
		羊耳蒜属Liparis lilifolia	[16]
		羊耳蒜属Liparis loeselii	[35]
		萼距兰属Disa bracteata	[36]
		飞鸟兰属Chiloglottis	[37]
		Cynorkis purpurea	[38]
		长药兰属Serapias parviflora	[39]
		珊瑚兰属Corallorhiza trifida	[40]
	角担菌属Ceratobasidium	羽柱鹦喙兰Gavilea australis	[41]
	蜡壳菌属Sebacina	鸟巢兰Neottia nidus-avis	[42]
		Cynorkis purpurea	[38]
		白及Bletilla striata	[34]
	小假鬼伞Coprinellus disseminatus	杜鹃兰Cremastra appendiculata	[43]
	毛霉属Mucor sp;	细茎石斛Dendrobium moniliforme(L.) Sw.	[17]
	镰孢菌属Fusarium		[17]
	轮枝孢菌属Verticilli		[17]
	瘤菌根菌属Epulorhiza sp.	小叶兜兰Paphiopedilum barbigerum (Orchidaceae)	[44]
		陆生兰花Habenaria macroceratitis	[45]
		龙头兰Pecteilis susannae (L.) Rafin	[45]
		紫苞舌兰Spathoglottis plicata	[46]
	丝核菌属Rhizoctonia globularis	紫苞舌兰Spathoglottis plicata	[46]
	蜡壳菌属Serendipita sp.	白及Bletilla striata	[34]
		大蜘蛛兰Caladenia huegelii	[47]

内生菌	促萌发内生细菌种属	兰科植物种属	文献
内生细菌	恶臭假单胞菌Pseudomonas putida	翅柱兰属Pterostylis vittata	[18]
	嗜麦芽黄单胞菌Xanthomonas maltophilia		[18]
	蜡样芽孢杆菌Bacillus cereus		[20]
	戴氏菌属Dyella	蝴蝶兰Phalaenopsis amabilis	[23]
	鞘氨醇菌属Sphingomonas	杓唇石斛Dendrobium moschatum(Buch.-Ham.)Sw.	[21]
	分枝菌属Mycobacterium		[21]
	贝莱斯芽孢杆菌Bacillus velezensis	白及Bletilla striata	[21]
	芽孢杆菌属Bacillus megaterium	五唇兰Phalaenopsis pulcherrima (Lindl) J.J. Sm.	[24]
	芽孢杆菌属Bacillus mycoides
	芽孢杆菌属Bacillus cereus
	荧光假单胞菌Pseudomonas fluorescens	白及Bletilla striata	[25]
内生真菌	镰刀菌属Fusarium Link ex Fr.	铁皮石斛Dendrobium officinale Kimura et Migo	[26]
	木霉属Trichoderma	华石斛Dendrobium sinense	[26]
	紫箕小菇Mycena osmundicola J.E.Lange	天麻Gastrodia elata	[27]
	石斛小菇Mycena dendrobii L.Fan et S.X.Guo		[27]
	兰小菇Mycena orchidicola Fan et Guo		[27]
	兰小菇Mycena orchidicola Fan et Guo	报春石斛Dendrobium polyanthum Wall. ex Lindl.	[28]
		长苏石斛Dendrobium brymerianum Rchb. f.	[28]
		鼓槌石斛Dendrobium chrysotoxum	[28]
		密花石斛Dendrobium densiflorum Lindl.	[28]
	胶膜菌属Tulasnella	驴兰Diuris	[29]
		虾脊兰属Calanthe	[30]
		兜唇石斛Dendrobium aphyllum	[31]
		硬叶兰Cymbidium mannii	[32]
		树兰Epidnedium	[33]
		白及Bletilla striata	[34]
		羊耳蒜属Liparis lilifolia	[16]
		羊耳蒜属Liparis loeselii	[35]
		萼距兰属Disa bracteata	[36]
		飞鸟兰属Chiloglottis	[37]
		Cynorkis purpurea	[38]
		长药兰属Serapias parviflora	[39]
		珊瑚兰属Corallorhiza trifida	[40]
	角担菌属Ceratobasidium	羽柱鹦喙兰Gavilea australis	[41]
	蜡壳菌属Sebacina	鸟巢兰Neottia nidus-avis	[42]
		Cynorkis purpurea	[38]
		白及Bletilla striata	[34]
	小假鬼伞Coprinellus disseminatus	杜鹃兰Cremastra appendiculata	[43]
	毛霉属Mucor sp;	细茎石斛Dendrobium moniliforme(L.) Sw.	[17]
	镰孢菌属Fusarium		[17]
	轮枝孢菌属Verticilli		[17]
	瘤菌根菌属Epulorhiza sp.	小叶兜兰Paphiopedilum barbigerum (Orchidaceae)	[44]
		陆生兰花Habenaria macroceratitis	[45]
		龙头兰Pecteilis susannae (L.) Rafin	[45]
		紫苞舌兰Spathoglottis plicata	[46]
	丝核菌属Rhizoctonia globularis	紫苞舌兰Spathoglottis plicata	[46]
	蜡壳菌属Serendipita sp.	白及Bletilla striata	[34]
		大蜘蛛兰Caladenia huegelii	[47]