以杨树为例论述林木共生菌系统及其作用机制

杨树(Populus)是重要造林树种,也是研究林木基础生物学性状的模式材料。不仅如此,杨树可与多种细菌(内生细菌、内生固氮菌和根际促生菌)和真菌(外生菌根真菌、丛枝菌根真菌和内生真菌)类群建立共生关系,为揭示树木和微生物之间的互惠共生机制提供了理想模型。这些共生菌能积极调控林木生长发育、营养吸收和生理生态过程。目前在杨树-双色蜡蘑(Laccaria bicolor)形成的外生菌根发育、提高杨树耐盐、耐重金属的生理与分子机制、叶片内生真菌群落结构与病害发生、菌根辅助细菌和菌丝内共生细菌-真菌-杨树形成的三重跨界共生等方面取得多项突破。近年来,一批模式草本植物微生物组(microbiome)计划相继实施,对共生菌群落结构和功能的认识有了革命性的进步。以美洲黑杨、毛果杨和胶杨为代表的林木微生物组研究也已启动,表明宿主基因型和环境因子可显著影响共生菌群落结构与物种组成;在根际(rhizosphere)和内生(endosphere)环境存在结构和功能迥异的菌群。另一方面,以根系为诱饵,通过宿主表型来推测菌群功能的反向“钓鱼”策略将推动林木根际微生物工程研究,为揭示杨树-微生物群落的相互关系、菌群进化搭建了研究模型。总之,深入认识多元微生物对林木表型和生理代谢的表观遗传学调控机制将为今后创制新型菌剂并用于高效育苗和抗性育种提供新的思路,具有重要的科学意义和应用价值。     

以杨树为例论述林木共生菌系统及其作用机制

杨树(Populus)是重要造林树种,也是研究林木基础生物学性状的模式材料。不仅如此,杨树可与多种细菌(内生细菌、内生固氮菌和根际促生菌)和真菌(外生菌根真菌、丛枝菌根真菌和内生真菌)类群建立共生关系,为揭示树木和微生物之间的互惠共生机制提供了理想模型。这些共生菌能积极调控林木生长发育、营养吸收和生理生态过程。目前在杨树-双色蜡蘑(Laccaria bicolor)形成的外生菌根发育、提高杨树耐盐、耐重金属的生理与分子机制、叶片内生真菌群落结构与病害发生、菌根辅助细菌和菌丝内共生细菌-真菌-杨树形成的三重跨界共生等方面取得多项突破。近年来,一批模式草本植物微生物组(microbiome)计划相继实施,对共生菌群落结构和功能的认识有了革命性的进步。以美洲黑杨、毛果杨和胶杨为代表的林木微生物组研究也已启动,表明宿主基因型和环境因子可显著影响共生菌群落结构与物种组成;在根际(rhizosphere)和内生(endosphere)环境存在结构和功能迥异的菌群。另一方面,以根系为诱饵,通过宿主表型来推测菌群功能的反向“钓鱼”策略将推动林木根际微生物工程研究,为揭示杨树-微生物群落的相互关系、菌群进化搭建了研究模型。总之,深入认识多元微生物对林木表型和生理代谢的表观遗传学调控机制将为今后创制新型菌剂并用于高效育苗和抗性育种提供新的思路,具有重要的科学意义和应用价值。     

外生菌根真菌与内生细菌共生互作的研究进展

外生菌根真菌能与很多高等植物共生,广泛存在于自然界,在促进植物生长和养分吸收、增强宿主抗逆性及维持森林生态系统稳定等方面发挥着重要作用。除与寄主植物密切联系外,外生菌根真菌,在其生命周期中与细菌群落进行物理和代谢相互作用常形成共生关系。这些细菌对外生菌根真菌菌丝生长、生物量增加及子实体的形成具有积极影响。本文阐述了外生菌根真菌与内生细菌共生现象的发现、共生关系的建立、内生细菌促进外生菌根真菌生长和发育及宿主与微生物组的研究方法等,以期更好地巩固外生菌根真菌的生物学及生态学等基础性知识,并利用细菌与真菌的相互作用为可食用外生菌根真菌的生物防治、菌肥研究、人工驯化及栽培提供思路。     

内生真菌对氮添加羽茅根际土壤特性和微生物群落的影响

内生真菌不仅能改变与其共生植物的生理和生长指标,还可通过宿主植物间接对土壤的理化性质和微生物群落结构产生影响。以天然禾草——羽茅(Achnatherum sibiricum)为研究材料,探究在不同施氮水平下,内生真菌的种类对不同基因型的宿主植物根际土壤理化性质和微生物群落产生何种影响。结果表明,内生真菌侵染显著提高了羽茅根际土壤的pH值和微生物总量,但降低了土壤中真菌与细菌的比值。同时,土壤的pH值还受到了内生真菌种类的影响,其中感染Epichlo3 sibirica的羽茅根际土壤pH显著高于感染Epichlo3 gansuense-1的羽茅,而感染Epichlo3 gansuense-2的羽茅根际土壤pH与感染E.gansuensis-1、E.sibirica菌的羽茅相比没有显著差异。另外,内生真菌感染与否、内生真菌种类、施氮量以及宿主植物基因型对土壤总碳、总氮、微生物及碳矿化能力均无显著影响。     

禾草内生真菌对其它微生物的影响研究进展

禾草内生真菌在宿主植物的茎叶等地上组织中普遍存在,不仅能够提高禾草对生物与非生物逆境的抗性,而且能够对周围环境中的不同微生物类群产生影响。主要总结了禾草Neotyphodium/Epichlo内生真菌对病原真菌、丛枝菌根真菌和土壤微生物的影响及其作用机理。发现禾草内生真菌普遍存在对病原真菌的抑制作用,而对丛枝菌根真菌存在不对称的竞争作用,且因种类而异。禾草内生真菌对土壤微生物群落的作用则会随着土壤类型和时间等外界因素发生变化。禾草内生真菌对不同类群微生物的影响机制主要包括:通过生态位竞争、抑菌物质分泌、诱导抗病性等对病原真菌造成影响;通过根系化学物质释放、营养元素调节、侵染条件差异等对丛枝菌根真菌造成影响;通过根际沉积物和凋落物等对土壤微生物群落造成影响。禾草内生真菌产生的生物碱能提高宿主植物对包括昆虫在内草食动物采食的抗性,影响病原菌的侵入、定殖和扩展;根组织分泌物中包含次生代谢产物能够抑制菌根真菌、土传病原真菌及其它土壤微生物的侵染与群落组成;也可能通过次生代谢物影响禾草的其它抗性。因此,禾草内生真菌在植物-微生物系统中的作用应该给予更多的关注和深入研究。     

植物与内生真菌互作的生理与分子机制研究进展

在自然生态系统中,植物组织可作为许多微生物定居的生态位.内生真菌普遍存在于植物组织内,与宿主建立复杂的相互作用(互惠、拮抗和中性之间的相互转化),并且存在不同的传播方式(垂直和水平传播).内生真菌通过多样化途径来增强植物体的营养生理和抗性机能.但这种生理功能的实现有赖于双方精细的调控机制,表明宿主和真菌双方都进化形成特有的分子调控机制来维持这种互惠共生关系.环境因子(如气候、土壤性质等)、宿主种类和生理状态、真菌基因型的变化都将改变互作结果.此外,菌根真菌和真菌病毒等也可能普遍参与植物-内生真菌共生体,形成三重互作体系,最终影响宿主的表型.研究试图从形态、生理和分子水平阐述内生真菌与植物互作的基础.     

喀斯特石漠化治理区钙果种植年限对根内细菌群落的影响

存在于健康植物根中的内生细菌不但能够与宿主植物建立共生关系,而且还具有促进植物生长、提升植物对营养元素摄取能力等功能,从而对维持陆地生态平衡、提升喀斯特石漠化综合治理成效具有重要的意义。【目的】探究宿主植物根中的内生细菌群落结构,为深入认识宿主植物-内生细菌的互作机制提供理论依据。【方法】以被引种到喀斯特断陷盆地石漠化综合治理区不同种植年限钙果根中的内生细菌和根际土为研究对象,分析根内生细菌群落特征和根际土理化性质。【结果】钙果种植年限对研究区土壤质量整体有着直接、显著的影响,并间接影响根内生细菌群落。根内生细菌群落以共生互作为主,通过共网络结构识别出种植第一年和第三年的前三位优势属为链霉菌属(Streptomyces)、卡氏伯克霍尔德菌属(Burkholderia-Caballeronia- Paraburkholderia)和噬几丁质菌属(Chitinophaga),种植第五年的前三位优势属为Streptomyces、Chitinophaga和海无柄孢囊黏细菌属(Haliangium)。根内生细菌群落的形成主要由随机性过程中的生态漂变所主导。【结论】不同生长阶段钙果根内生细菌群落结构的差异是由于随机性过程赋予了微生物物种多样化。内生细菌群落的共生互作关系以及优势菌具有生防功能,可增强钙果定殖能力和生长,进而提升钙果在喀斯特断陷盆地石漠化综合治理区的生态效益和经济效益。     

药用植物刺山柑不同部位细菌群落结构及其多样性

【目的】研究传统药用植物刺山柑（Capparis spinosa L.）不同部位细菌群落结构、物种组成和多样性特征,为药用植物微生物资源的开发及微生物与宿主互作提供理论依据。【方法】本研究以刺山柑地上部植物组织（果实、茎）和地下部土壤（根际土壤、非根际土壤）为研究材料,采用高通量测序技术分析刺山柑不同部位细菌的16S rRNA基因多样性,比较其细菌群落结构和物种组成特征。【结果】刺山柑4种样本共获得的3 649个操作分类单元（operational taxonomic unit,OTU）,属于34门、88纲、248目、464科和1 051属。土壤样本的细菌多样性大于植物组织,细菌群落多样性以根际土壤、非根际土壤、茎和果实的顺序逐渐降低,果实内生细菌群落多样性始终最低,显著低于根际土壤。不同部位相对丰度较高的细菌门如下：植物组织中以变形菌门为主,根际土壤中为变形菌门和放线菌门,非根际土中为厚壁菌门和放线菌门。无色杆菌属（Achromobacter）、欧文氏菌属（Erwinia）、肠球菌属（Enterococcus）、微小杆菌属（Exiguobacterium）、乳杆菌属（Lactobacillus）和克雷伯氏菌属（Klebsiella）主要存在于刺山柑植物组织中。游动球菌属（Planomicrobium）、库克菌属（Kocuria）、类芽孢杆菌属（Paenibacillus）、链霉菌属（Streptomyces）、微枝形杆菌属（Microvirga）和节杆菌属（Arthrobacter）主要分布于土壤中。β多样性分析结果表明,刺山柑植物组织和土壤的细菌群落结构具有显著差异,同类型样本的细菌群落结构相似。【结论】刺山柑土壤样本中细菌群落的多样性和丰富度均高于植物组织,刺山柑不同部位的细菌群落组成不同。本研究对刺山柑不同部位细菌群落结构进行了初步分析,鉴定了各部位细菌群落中的核心菌群,为以后挖掘刺山柑的功能研究和利用提供了准确的微生物信息。     

管叶槽舌兰新鲜及硅胶干燥根样内生菌多样性研究

管叶槽舌兰(Holcoglossum kimballianum)是一种珍稀濒危兰科植物,其野生种群亟需保护。内生菌对兰科植物的生长发育至关重要,为评估管叶槽舌兰内生菌的多样性、分析采样方式对其内生菌的影响,该文采用高通量测序技术对迁地保育状态下新鲜与硅胶干燥的管叶槽舌兰根内生菌进行研究。结果表明:(1)新鲜及干燥管叶槽舌兰根内生菌物种组成明显不同,管叶槽舌兰内生真菌注释到6门46科51属,内生细菌注释到15门105科178属; 干燥后管叶槽舌兰根内生真菌注释到6门88科116属,内生细菌注释到21门154科336属。(2)迁地保育状态的管叶槽舌兰根样内生菌具有丰富的多样性,并且内生细菌群落丰富度和多样性远比内生真菌群落高; 经硅胶干燥后,内生真菌α多样性指数升高、β多样性指数降低,而内生细菌的α多样性指数降低、β多样性指数则升高。(3)差异显著性真菌黄盖小脆柄菇(Psathyrella candolleana)和刺盘孢属(Colletotrichum)的C. tofieldiae只存在于新鲜根样中,新鲜管叶槽舌兰差异显著性细菌是马赛菌属(Massilia),干燥根样中差异显著性细菌类群包括拜叶林克氏菌科(Beijerinckiaceae)、黄色杆菌科(Xanthobacteraceae)及慢生根瘤菌属(Bradyrhizobium)。(4)共发生网络分析显示,经干燥后管叶槽舌兰根样内生菌群落中占互作主导地位的优势物种和互作模式都发生了改变。综上认为,不同采样处理会影响管叶槽舌兰根内生菌的群落结构,在研究兰科植物根样内生菌时宜使用新鲜的根样。该研究结果为管叶槽舌兰野生种群保护及人工栽培提供了内生菌数据基础,也为兰科植物内生微生物采样方法提供了参考。     

土地利用驱动的土壤性状变化影响微生物群落结构和功能

微生物在调节陆地生态系统地球化学循环过程中具有重要作用。土地利用方式改变显著影响土壤微生物群落结构和功能,但对土地利用驱动的土壤性状变化与微生物群落结构和功能关系的研究相对匮乏。依托长期定位监测试验（始于1984年）,通过16S rRNA基因片段和ITS高通量测序,研究了土地利用方式（裸地、农田、草地）驱动的土壤碳氮变化对微生物群落结构和功能的影响。结果表明：对于细菌群落而言,裸地中α-多样性最高、其次是草地、农田中最低,农田和草地中细菌优势菌群变形菌（Proteobacteria）和放线菌门（Actinobacteria）相对丰度较裸地低4.5%、3.9%和5.5%、3.8%;对于真菌群落而言,裸地子囊菌门（Ascomycota）相对丰度最高、农田次之、草地最低;化能异养型、好氧化能异养型细菌相对丰度裸地显著高于农田和草地（P<0.05）,而硝化型和好氧氨氧化型细菌裸地显著低于农田和草地（P<0.05）;腐生型真菌相对丰度大小排序为：裸地>农田>草地。细菌群落变化主要与土壤容重、全氮、矿质氮、C ： N比和微生物量碳有关,而真菌群落与土壤矿质氮有关。细菌和真菌功能菌群主要受土壤容重、土壤有机碳、土壤全氮、C ： N比和微生物量碳影响。因此,土壤容重、土壤全氮、土壤有机碳、C ： N比、微生物量碳、矿质氮差异可能是影响不同土地利用方式中微生物群落和功能变化的主要因素。     

Continental‐scale nitrogen pollution is shifting forest mycorrhizal associations and soil carbon stocks

Most tree roots on Earth form a symbiosis with either ecto‐ or arbuscular mycorrhizal fungi. Nitrogen fertilization is hypothesized to favor arbuscular mycorrhizal tree species at the expense of ectomycorrhizal species due to differences in fungal nitrogen acquisition strategies, and this may alter soil carbon balance, as differences in forest mycorrhizal associations are linked to differences in soil carbon pools. Combining nitrogen deposition data with continental‐scale US forest data, we show that nitrogen pollution is spatially associated with a decline in ectomycorrhizal vs. arbuscular mycorrhizal trees. Furthermore, nitrogen deposition has contrasting effects on arbuscular vs. ectomycorrhizal demographic processes, favoring arbuscular mycorrhizal trees at the expense of ectomycorrhizal trees, and is spatially correlated with reduced soil carbon stocks. This implies future changes in nitrogen deposition may alter the capacity of forests to sequester carbon and offset climate change via interactions with the forest microbiome.     

植物病害防治相关微生物组研究进展与展望

微生物是人类活动过程中重要的生物资源。植物及其根围土壤中生存着大量多种多样的微生物,这些微生物与植物健康之间存在着密不可分的关系。近年来,基因测序技术的快速发展为植物微生物组结构和功能的研究提供了极大的便利,多种植物相关的微生物组得到了解析。同时更多研究者聚焦于植物病害相关的微生物组研究,通过差异分析,发现了一些特定的有益于植物健康的微生物菌群。此外,植物根围或根内微生物塑造的内在原理也得到了进一步的揭示。一系列植物微生物组研究为植物病害防治和新的微生物资源的挖掘提供了更多思路。     

Mycorrhizal and dark septate endophytic fungi of <Emphasis Type="Italic">Pedicularis</Emphasis> species from northwest of Yunnan Province,China

Colonization of mycorrhizal and root endophytic fungi in 14 Pedicularis species from northwest of Yunnan Province, southwest China, was examined. These species included: Pedicularis gracilis Wall., Pedicularis longipes Maxim., Pedicularis axillaris Franch., Pedicularis cephalantha Franch., Pedicularis tenuisecta Franch., Pedicularis tapaoensis Tsoong, Pedicularis likiangensis Franch., Pedicularis dichotoma Bonati, Pedicularis yui Li, Pedicularis rhinanthoides Schrenk, Pedicularis rex C.B. Clarke, Pedicularis longiflora Rudolph., Pedicularis siphonantha Don, and Pedicularis oxycarpa Franch., among which nine are endemic to China (one to Yunnan). Three types of potentially beneficial fungi associated with roots of these species were observed, namely, arbuscular mycorrhizal fungi, ectomycorrhizal fungi, and dark septate endophytic fungi (DSEF), with DSEF as the most common colonizers. An unexpected high colonization level was detected in this hemiparasitic genus. Of the 19 sampling sites examined, 10 gave colonization frequency of above 50% and 6 showed a colonization index of above 50. Heavy colonization suggested a significant ecological role of these fungi and their potential to be applied to successful cultivation of these intractable plants.     

The mycorrhizal status and colonization of 26 tree species growing in urban and rural environments

Urban environments are highly disturbed and fragmented ecosystems that commonly have lower mycorrhizal fungal species richness and diversity compared to rural or natural ecosystems. In this study, we assessed whether the mycorrhizal status and colonization of trees are influenced by the overall environment (rural vs. urban) they are growing in. Soil cores were collected from the rhizosphere of trees growing in urban and rural environments around southern Ontario. Roots were extracted from the soil cores to determine whether the trees were colonized by arbuscular mycorrhizal fungi, ectomycorrhizal fungi, or both, and to quantify the percent colonization of each type of mycorrhizal fungi. All 26 tree species were colonized by arbuscular mycorrhizal fungi, and seven tree species were dually colonized by arbuscular mycorrhizal and ectomycorrhizal fungi. Overall, arbuscular mycorrhizal and ectomycorrhizal fungal colonization was significantly (p < 0.001) lower in trees growing in urban compared to rural environments. It is not clear what ‘urban’ factors are responsible for the reduction in mycorrhizal fungal colonization; more research is needed to determine whether inoculating urban trees with mycorrhizal fungi would increase colonization levels and growth of the trees.     

黄精根际及药用部位内生真菌群落组成和生态功能分析

为寻找促进药用植物活性代谢产物合成的微生物,该文以黄精为研究对象,利用高通量测序技术和生态功能预测平台,测定根际土真菌、根茎和根内生真菌的ITS序列,分析其真菌多样性和群落组成,并预测根茎内生真菌的生态功能.结果表明:(1)测序得到1023个可操作分类单元(OTUs),根际、根茎和根真菌OTU数分别为703、128和1...     

A Multifactor Analysis of Fungal and Bacterial Community Structure in the Root Microbiome of Mature Populus deltoides Trees

Bacterial and fungal communities associated with plant roots are central to the host health, survival and growth. However, a robust understanding of the root-microbiome and the factors that drive host associated microbial community structure have remained elusive, especially in mature perennial plants from natural settings. Here, we investigated relationships of bacterial and fungal communities in the rhizosphere and root endosphere of the riparian tree species Populus deltoides, and the influence of soil parameters, environmental properties (host phenotype and aboveground environmental settings), host plant genotype (Simple Sequence Repeat (SSR) markers), season (Spring vs. Fall) and geographic setting (at scales from regional watersheds to local riparian zones) on microbial community structure. Each of the trees sampled displayed unique aspects to its associated community structure with high numbers of Operational Taxonomic Units (OTUs) specific to an individual trees (bacteria >90%, fungi >60%). Over the diverse conditions surveyed only a small number of OTUs were common to all samples within rhizosphere (35 bacterial and 4 fungal) and endosphere (1 bacterial and 1 fungal) microbiomes. As expected, Proteobacteria and Ascomycota were dominant in root communities (>50%) while other higher-level phylogenetic groups (Chytridiomycota, Acidobacteria) displayed greatly reduced abundance in endosphere compared to the rhizosphere. Variance partitioning partially explained differences in microbiome composition between all sampled roots on the basis of seasonal and soil properties (4% to 23%). While most variation remains unattributed, we observed significant differences in the microbiota between watersheds (Tennessee vs. North Carolina) and seasons (Spring vs. Fall). SSR markers clearly delineated two host populations associated with the samples taken in TN vs. NC, but overall host genotypic distances did not have a significant effect on corresponding communities that could be separated from other measured effects.     

Relationship between genotype and soil environment during colonization of poplar roots by mycorrhizal and endophytic fungi

Poplars are among the few tree genera that can develop both ectomycorrhizal (ECM) and arbuscular (AM) associations; however, variable ratios of ECM/AM in dual mycorrhizal colonizations were observed in the roots of a variety of poplar species and hybrids. The objective of our study was to analyze the effect of internal and external factors on growth and dual AM and ECM colonization of poplar roots in three 12–15-year-old common gardens in Poland. We also analyzed the abundance of nonmycorrhizal fungal endophytes in the poplar roots. The Populus clones comprised black poplars (Populus deltoides and P. deltoides × Populus nigra), balsam poplars (Populus maximowiczii × Populus trichocarpa), and a hybrid of black and balsam poplars (P. deltoides × P. trichocarpa). Of the three sites that we studied, one was located in the vicinity of a copper smelter, where soil was contaminated with copper and lead. Poplar root tip abundance, mycorrhizal colonization, and soil fungi biomass were lower at this heavily polluted site. The total mycorrhizal colonization and the ratio of ECM and AM colonization differed among the study sites and according to soil depth. The influence of Populus genotype was significantly pronounced only within the individual study sites. The contribution of nonmycorrhizal fungal endophytes differed among the poplar clones and was higher at the polluted site than at the sites free of pollution. Our results indicate that poplar fine root abundance and AM and ECM symbiosis are influenced by environmental conditions. Further studies of different site conditions are required to characterize the utility of poplars for purposes such as the phytoremediation of polluted sites.     

荒漠植物黑沙蒿嫩枝与成熟枝内生菌群落结构差异

[背景]黑沙蒿是我国北方沙漠地区分布广泛、抗旱性能优良的固沙灌木,对稳定沙漠地区生态系统有至关重要的作用.[目的]内生菌在植物生命过程中扮演着重要角色,认识植物生长发育阶段幼嫩和成熟组织内生菌群的结构变化,对于理解菌群间的相互作用及保护宿主植物抵御生物和非生物胁迫具有积极意义.[方法]以宁夏拉巴湖林场黑沙蒿为研究对象,...     

辽宁丹东地区柞树菌根真菌生态分布的研究

通过5年连续调查研究,发现辽宁丹东地区天然次生柞林中及部分柞树经济林中柞树菌根菌种类共有36种,隶属于7个科13个属,在此基础上研究分析了柞树菌根菌的生态分布与共生树种、树龄及季节的关系,结果表明,与蒙古栎共生的菌根菌种类最多,达29种;红菇属种类与柞树共生的树龄范围最广,鹅膏属种类偏重于与树龄较大的柞树共生;夏季的7、8月份为菌根菌发生的高峰期,75％的种类在7月份发生。100％的种类在8月份发生,8月中下旬菌根菌发生的相对密度最高,分别为19．5％和18．4％。     

Waiting for fungi: the ectomycorrhizal invasion of lowland heathlands

1.  In England, the loss of lowland heathland, a habitat of global conservation importance, is primarily due to the invasion of birch and pine. This encroachment has been researched in depth from a plant perspective but little is known about the role of mycorrhizal fungi. In lowland heathlands the resident dwarf shrubs form ericoid mycorrhizas whereas invading trees form ectomycorrhizas. Therefore, tree encroachment into heathlands can be regarded as the replacement of a resident mycorrhizal community by an invading one.
2 . This study examined how fungi form mycorrhizas with Betula and Pinus in lowland heathlands. We addressed the question of whether there are mycorrhizal fungi that mediate invasion using a molecular ecology approach to compare the mycorrhizal inoculum potential of soil at three levels of invasion (uninvaded heathland, invaded heathland and woodland) and the fungi forming mycorrhizas on tree seedlings and trees across diverse sites.
3.  We show that in lowland heathlands: (i) seedlings have severely limited access to ectomycorrhizal fungi relative to woodlands, (ii) there are few keystone spore-dispersed ectomycorrhizal fungi that can mediate tree invasion, (iii) tree seedlings can remain non-mycorrhizal for at least one year when no inoculum is present, even near saplings, and (iv) mycorrhizal seedlings achieve greater biomass than non-mycorrhizal seedlings. Within uninvaded heathland we detected only Rhizopogon luteolus , Suillus variegatus , S. bovinus ( Pinus symbionts) and Laccaria proxima (primarily a Betula symbiont).
4. Synthesis . Overall, ectomycorrhizal inoculum in lowland heathlands is rare; most tree seedlings growing in heathland soil are not mycorrhizal due to limited spore dispersal, poorly developed spore banks and weak common mycorrhizal networks. These seedlings can persist awaiting mycorrhization to boost their growth.