野外模拟酸雨胁迫下接种外生菌根真菌对马尾松幼苗的缓解作用

外生菌根真菌能够提高宿主植物对外界环境胁迫的抵抗力。主要探讨野外条件下外生菌根真菌对酸雨胁迫下马尾松(Pinus massoniana)幼苗生长、养分元素以及表层土壤的影响,以期为酸雨严重区马尾松林恢复提供科学依据。以2年生马尾松幼苗为材料,采用原位试验,共设置6个处理:p H5.6(对照)处理未接种、对照处理接种、p H4.5酸雨处理未接种、p H4.5酸雨处理接种、p H3.5酸雨处理未接种、p H3.5酸雨处理接种。研究表明:(1)酸雨处理与对照处理相比显著降低了非菌根苗总生物量及各部位生物量(根、茎、叶),对株高无显著影响,接种外生菌根真菌可以缓解酸雨对马尾松幼苗生长的不利影响;(2)与对照处理相比,酸雨处理的非菌根苗的针叶中N、P、Ca含量升高,Mg含量降低,根系中N、P、Ca含量降低,Mg含量随p H的降低先升高后降低。接种外生菌根真菌显著提高了p H3.5酸雨处理的马尾松幼苗根系中N、P、Ca、Mg含量,而对针叶中N、P、Ca、Mg含量无显著影响。(3)在非菌根土壤中,p H3.5酸雨处理与对照处理相比显著降低了土壤中有机质、速效磷、速效钾、可溶性碳、可溶性氮、铵态氮、硝态氮含量,而接种外生菌根真菌显著提高了上述指标。酸雨对土壤阳离子交换量无显著影响。总而言之,接种外生菌根真菌促进了酸雨处理的马尾松幼苗生长、缓解了酸雨对马尾松幼苗养分元素和表层土壤的不利影响,由此可见接种外生菌根真菌是减轻酸雨对马尾松危害的一个重要途径。     

外生菌根菌与森林树木的相互关系

生态系统的每个过程都伴随着各种微生物的活动,其中最重要的功能群之一是菌根真菌(菌根菌)。一般认为,菌根菌是自然界多数植物生存最基本的组成部分,陆地上约90％以上的高等植物都具有菌根菌。这些菌类的菌丝体与植物根系结合形成菌根,使植物生长成为可能,使不同种类植物的根系联在一起。根据菌根菌入侵植物根系的方式及菌根的形态特征,菌根可分为外生菌根、内生菌根和内外生菌根3组共7种类型。外生菌根主要出现在松科、桦木科、壳斗科等树种的森林生态系统中,在根系表面形成菌丝鞘,部分菌丝进入根系皮层细胞间隙形成哈氏网表面。菌根菌剂在森林经营中得到广泛地应用。外生菌根菌对森林树木的作用可归纳为：1)促进造林或育苗成活与生长;2)提高森林生态系统中植物的多样性、稳定性和生产力;3)对森林生态系统的综合效应,主要表现在增加植物一土壤联结,改善土壤结构,促进土壤微生物,增强植物器官的功能;4)抗拮植物根部病害病原菌等。树木与菌根菌相互关系研究主要包括：1)菌根共生的机理;2)菌根菌在退化森林生态系统恢复与改造中的作用;3)菌根菌的分布格局与森林生态系统服务功能的关系;4)菌根菌对森林生态系统的综合效应,如菌根菌与森林植物群落结构、物种多样性以及森林系统稳定性和生产力的研究。     

模拟酸雨和接种外生菌根真菌对马尾松土壤养分、土壤团聚体及有机碳组分的影响

以2年生马尾松(Pinus massoniana)幼苗为试验对象,采用原位试验,研究了模拟酸雨和接种外生菌根真菌对土壤有机碳含量、土壤团聚体组成及土壤养分(N、P、K)含量的影响。设置3种强度(p H5.6(CK),4.5,3.5)的模拟酸雨条件,每个强度酸雨下分别设置接种外生菌根真菌和不接种外生菌根真菌,共计6个处理。结果表明:p H3.5酸雨处理显著降低了土壤中速效磷、总磷、速效钾、铵态氮、硝态氮及总氮含量,而外接外生菌根真菌则有效缓解了这一情况;与对照比,酸雨处理提高了微团聚体(0.053 mm,0.053～0.25 mm)含量,降低了大团聚体(0.25～2 mm)含量; pH3.5处理显著降低各粒级尤其是0.25～2 mm粒级团聚体有机碳含量,而接种外生菌根真菌显著增加了有机碳含量;未接种外生菌根真菌土壤中,酸雨处理显著降低了土壤轻组、重组有机碳含量,而接种外生菌根真菌显著提高了土壤轻组、重组有机碳含量;接种外生菌根真菌在一定程度上可以缓解酸雨胁迫对土壤质量的不利影响。     

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。     

氮沉降对外生菌根真菌的影响

综述了国外氮沉降对外生菌根真菌的影响研究现状 ,主要从菌根形成、形态 (菌丝体、菌根 )变化、子实体生产力和外生菌根真菌群落结构等方面对氮沉降的响应进行了综述 ,并初步探讨了氮饱和的临界负荷。研究表明 ,过量氮沉降会给外生菌根真菌在以下几个方面带来负影响 :(1)影响外生菌根真菌与寄主植物之间的养分分配和循环 ;(2 )降低子实体生产力 ;(3)减少菌丝 ;(4 )降低菌根量及其活力 ;(5 )降低外生菌根真菌丰富度 ;(6 )改变外生菌根真菌群落结构组成 ;(7)降低外生菌根真菌群落功能。还指出了未来该方面研究重点和方向     

中国菌根研究60年:过去、现在和将来

菌根(真菌根系)存在于大约90%的植物中,在促进土壤结构、植物养分与生长、元素生物地球化学循环和陆地生态系统结构与功能等方面具有重要作用.过去60年尤其是近30年,中国菌根研究成果举世瞩目,如共鉴定出20种新种与120余种新记录种丛枝菌根真菌、30种新种与800余种新记录种外生菌根真菌以及10种新种与100余种新记录种兰花菌根真菌.同时,在菌根真菌菌种丰富度与遗传结构、菌种生态分布与植物种群,植物养分摄取与生长、植物修复与土地复垦、植物抗病性和与其他土壤微生物相互作用、菌根植物酶学性质及大气CO2和O3浓度升高对丛枝菌根多样性的影响等方面也取得重大进展.本文选介中国菌根主要研究成就,进行研究前景展望,以促进我国菌根研究的深入开展.     

外生菌根与植物抗重金属胁迫机理

外生菌根是外生菌根真菌和植物营养根形成的共生体,能够增加植物对污染胁迫的抵抗能力。本文综述了20多年来国内外研究外生菌根增加植物抗重金属毒害的成果,指出了外生菌根在植物抗重金属毒害中的积极作用,并概括其抗性的主要机理为：外延菌丝的吸收作用;菌根分泌物的调节与螯合作用;菌根菌套或哈蒂氏网吸收过滤有毒金属;菌根菌套的疏水性作用。在研究外生菌根抗重金属毒害机理的基础上,提出了该领域今后的研究前景。     

钾与信号抑制剂对外生菌根真菌分泌乙酸的调控作用

外生菌根真菌是森林生态系统中的重要成分,参与树木养分的吸收利用。试验液体培养外生菌根真菌,设置不同供钾水平,添加钙信号抑制剂,研究了它们的有机酸和氢离子分泌,以及乙酸分泌对供钾和信号抑制剂的响应。结果表明,供试菌株的生长,氮、磷、钾含量和吸收量因菌株不同而异,生物量变化于52.91—121.72 mg/瓶之间,相差1倍以上。外生菌根真菌吸收养分的差异可能与它们对土壤养分环境的长期适应、进化、选择有关。在外生菌根真菌的培养液中,分别检测出草酸、乙酸、苹果酸、柠檬酸和丁二酸等,以及大量的氢离子,说明菌根真菌能分解土壤含钾矿物,释放钾离子,改善寄主植物的钾营养。其中,乙酸分泌量较大,具有普遍性,低钾刺激分泌乙酸,高钾时分泌减少,其分泌速率与供钾浓度和菌丝吸钾量之间呈显著负相关(r=-0.734,r=-0.617,n=60)。钾对菌根真菌分泌乙酸的调控作用具有改善森林钾素营养,防止土壤养分淋失的生理和生态意义。此外,在低钾条件下,阴离子通道和钙信号抑制剂抑制外生菌根真菌分泌乙酸。说明钙信号和阴离子通道参与了乙酸分泌,缺钾可能是刺激乙酸分泌的原初信号,通过信号转导和一系列级联反应促进乙酸分泌。     

外生菌根真菌对矿物风化作用研究进展

外生菌根真菌是生态系统的重要组成部分,其对岩石和矿物的风化作用可对生态系统的物质循环过程产生影响,并在一定程度上提高土壤养分的生物有效性,有利于改善植物营养状况。本文回顾了外生菌根真菌对矿物风化作用的研究进展,并对未来的研究方向进行了展望。研究表明,外生菌根真菌可以通过向矿物表面施加物理压力、酸化根际环境以及络合矿物中的金属离子等途径加快包括磷灰石、云母、蛭石等矿物在内的风化进程,促使养分从矿物中析出,并使自身及与其共生的植物受益。但外生菌根真菌对矿物的风化作用受到诸如真菌与矿物的类型、土壤有效养分水平、氮素类型与水平、菌根真菌的生长条件以及根际其他生物的活动等多种因素的影响。目前,通过对矿质元素转移的定量计算、对真菌分泌物的测定分析、对菌丝与矿物颗粒接触面的直接观察分析以及对能量流动的监测等一系列途径,已经对外生菌根真菌矿物风化作用的现象和机理有了一定的认识,但针对这种现象所带来影响的定量计算及其生态意义还有待进一步探究。     

外生菌根真菌生物地理学研究进展

外生菌根真菌对植物矿质营养、生态系统物质循环、物种演化进程等具有十分重要的作用,但其生物地理学研究长期滞后于动植物。扩散和隔离是解释外生菌根真菌生物地理分布格局的重要理论。古地质、古气候和宿主植物是外生菌根真菌地理分布格局形成的重要推动因子。基因组学、生物信息学技术与生物地理学方法相互结合和补充,可以用来研究一些复杂的真菌生物地理学问题。文中详细阐述了外生菌根真菌生物地理学研究的重要过程和现状,如真菌物种界定、常用数据分析方法、起源和演化规律及其在生物多样性保护方面的应用等,并初步探讨了其未来的发展方向。     

接种彩色豆马勃对模拟酸沉降下马尾松幼苗生物量的影响

外生菌根能够提高宿主植物对外界环境胁迫的抵抗力,促进植物的生长,本文试图揭示外生菌根对酸雨胁迫下马尾松生长的保护作用。本研究采用盆栽试验,共设置四个处理：酸雨对照处理（Control check (CK), 约pH值5.5）不接种,酸雨对照处理接种,酸雨pH值3.5处理不接种,酸雨pH值3.5处理接种。pH值3.5的酸雨处理降低马尾松的生物量,在试验前期降低根冠比,试验中后期则提高根冠比,在试验初期增加了叶面积,但中后期显著降低了叶面积。接种外生菌根菌有利于马尾松幼苗的生长,pH值3.5处理下接种外生菌根菌能提高马尾松幼苗的生物量,外生菌根菌对生物量分配和叶面积的影响与酸雨胁迫的影响是相反的,即外生菌根菌抵消了酸雨胁迫对马尾松的影响。     

Ectomycorrhizal fungi inoculation alleviates simulated acid rain effects on soil ammonia oxidizers and denitrifiers in Masson pine forest

Acid rain can cause severe effects on soil biota and nutrient biogeochemical cycles in the forest ecosystem, but how plant-symbiotic ectomycorrhizal fungi will modulate the effects remains unknown. Here, we conducted a full factorial field experiment in a Masson pine forest by simultaneously controlling the acidity of the simulated rain (pH 5.6 vs. pH 3.5) and the ectomycorrhizal fungi Pisolithus tinctorius inoculation (non-inoculation vs. inoculation), to investigate the effects on ammonia oxidizers and denitrifiers. After 10 months, compared with the control (rain pH 5.6, and non-inoculation), simulated acid rain (pH 3.5) reduced soil nutrient content, decreased archaeal amoA gene abundance and inhibited denitrification enzyme activity. Also, simulated acid rain altered the community compositions of all the examined functional genes (archaeal amoA, bacterial amoA, nirK, nirS and nosZ). However, inoculation with ectomycorrhizal fungi under acid rain stress recovered soil nutrient content, archaeal amoA gene abundance and denitrification enzyme activity to levels comparable to the control, suggesting that ectomycorrhizal fungi inoculation ameliorates simulated acid rain effects. Taken together, ectomycorrhizal fungi inoculation – potentially through improving soil substrate availability – could alleviate the deleterious effects of acid rain on nitrogen cycling microbes in forest soils.     

Ectomycorrhizal fungi are shared between seedlings and adults in a monodominant Gilbertiodendron dewevrei rain forest in Cameroon

Ectomycorrhizal networks may facilitate the establishment and survival of seedlings regenerating under the canopies of tropical forests and are often invoked as a potential contributor to monodominance. We identified ectomycorrhizal fungi in a monodominant Gilbertiodendron dewevrei (Fabaceae) rain forest in Cameroon, using sporocarps and ectomycorrhizae of three age categories (seedlings, intermediate trees, and large trees) and tentatively revealed nutrient transfer through ectomycorrhizal networks by measuring spontaneous isotopic (¹³C and ¹⁵N) abundances in seedlings. Sporocarp surveys revealed fewer ectomycorrhizal fungal taxa (59 species from 1030 sporocarps) than molecular barcoding of ectomycorrhizal roots (75 operational taxonomic units from 828 ectomycorrhizae). Our observations suggest that ectomycorrhizal fungal diversity is similar to that in other mixed tropical forests and provide the first report of the Tuber‐Helvella lineage in a tropical forest. Despite some differences, all age categories of G. dewevrei had overlapping ectomycorrhizal fungal communities, with families belonging to Thelephoraceae, Russulaceae, Sebacinaceae, Boletaceae, and Clavulinaceae. Of the 49 operational taxonomic units shared by the three age categories (65.3% of the ectomycorrhizal fungal community), 19 were the most abundant on root tips of all categories (38.7% of the shared taxa), supporting the likelihood of ectomycorrhizal networks. However, we obtained no evidence for nutrient transfer from trees to seedlings. We discuss the composition of the ectomycorrhizal fungal community among the G. dewevrei age categories and the possible role of common ectomycorrhizal networks in this rain forest.     

The Multifunctional Role of Ectomycorrhizal Associations in Forest Ecosystem Processes

Belowground biological interactions that occur among plant roots, microorganisms and animals are dynamic and substantially influence ecosystem processes. Among these interactions, the ectomycorrhizal (ECM) symbiosis is remarkable but unfortunately these associations have mainly been considered within the rather narrow perspective of their effects on the uptake of dissolved mineral nutrients by individual plants. More recent research has placed emphasis on a wider, multifunctional perspective, including the effects of ectomycorrhizal symbiosis on plant and microbial communities, and on ecosystem processes. This includes mobilization of N and P from organic polymers, release of nutrients from mineral particles or rock surfaces via weathering, effects on carbon cycling, interactions with mycoheterotrophic plants, mediation of plant responses to stress factors such as drought, soil acidification, toxic metals, and plant pathogens, rehabilitation and regeneration of degraded forest ecosystems, as well as a range of possible interactions with groups of other soil microorganisms. Ectomycorrhizas are almost invariably characterized by a Hartig net composed of highly branched hyphae which entirely surround the outer root cortical cells. The Hartig net is the place of massive bidirectional exchanges of nutrients between the host and the fungus. Through these branched hyphae ectomycorrhizal fungi connect their plant hosts to the heterogeneously distributed nutrients required for their growth, enabling the flow of energy-rich compounds required for nutrient mobilization whilst simultaneously providing conduits for the translocation of mobilized products back to their hosts. In addition to increasing the nutrient absorptive surface area of their host plant root systems, the extraradical mycelium of ectomycorrhizal fungi provides a direct pathway for translocation of photosynthetically derived carbon from their hosts to microsites in the soil and a large surface area for interaction with other soil micro-organisms. The detailed functioning and regulation of these mycorrhizosphere processes is still poorly understood and needs detailed molecular approach to study these mycorrhizosphere processes but recent progress in ectomycorrhizal associations is reviewed and potential benefits of improved understanding of mycorrhizosphere interactions are discussed.     

外生菌根真菌的共生互作和宿主选择机制研究进展

外生菌根真菌作为树木的共生伙伴,是森林生态系统重要组成部分,在森林天然更新、植物抗逆性形成、协助植物吸收限制性营养等方面扮演重要角色。真菌和植物跨界共生具有复杂的分子互作过程,在共生的不同阶段有不同的分子互作机制,其调控反馈网络还有许多未知。基因组与转录组研究技术和方法的进步,为一些新的信号分子、效应蛋白以及相关通路的发现提供了可能。真菌与宿主植物之间营养转移调控对共生的影响也逐渐受到关注,营养相关的转运蛋白对共生的建立和维持提供了物质基础。真菌的宿主选择机制是值得重点关注的领域,由于外生菌根真菌的多谱系起源和演化史中存在多次宿主转换事件,真菌演化出多样的应对机制用来区分相容性宿主、不相容性宿主和非宿主。通过对不同真菌与宿主植物的组学研究,宿主选择机制研究取得了一定进展。本文对近十年来国内外的研究报道进行梳理与总结,并对未来在该领域的探索方向做出展望。     

Host plants and edaphic factors influence the distribution and diversity of ectomycorrhizal fungal fruiting bodies within rainforests from Tshopo,Democratic Republic of the Congo

Ectomycorrhizal fungi constitute an important component of forest ecosystems that enhances plant nutrition and resistance against stresses. Diversity of ectomycorrhizal (EcM) fungi is, however, affected by host plant diversity and soil heterogeneity. This study provides information about the influence of host plants and soil resources on the diversity of ectomycorrhizal fungal fruiting bodies from rainforests of the Democratic Republic of the Congo. Based on the presence of fungal fruiting bodies, significant differences in the number of ectomycorrhizal fungi species existed between forest stand types (p < 0.001). The most ectomycorrhizal species‐rich forest was the Gilbertiodendron dewevrei‐dominated forest (61 species). Of all 93 species of ectomycorrhizal fungi, 19 demonstrated a significant indicator value for particular forest stand types. Of all analysed edaphic factors, the percentage of silt particles was the most important parameter influencing EcM fungi host plant tree distribution. Both host trees and edaphic factors strongly affected the distribution and diversity of EcM fungi. EcM fungi may have developed differently their ability to successfully colonise root systems in relation to the availability of nutrients.     

酸雨引起森林生态系统钙流失研究进展

酸雨作为一个全球性环境问题,随着我国经济的快速发展,其危害也日益严重。诸多长期观测研究认为酸雨导致森林生态系统营养流失,特别是Ca的流失是酸雨导致森林衰退的一个重要原因。长期酸雨影响下,土壤中的Ca大量流失,在红壤等酸性土壤中表现得尤为严重;此外酸雨也会直接淋溶出植物叶片中的Ca,造成林冠中Ca不同程度的淋溶流失,并可能造成叶片Ca含量下降。且酸雨会导致叶片细胞内Ca稳态的失衡,影响植物的正常生理响应。对于整个森林生态系统而言,酸雨改变了生态系统中Ca生物地球化学循环的特征,大量的Ca从森林生态系统中以地表径流的形式流出系统,导致可利用Ca库的下降和周转速率的上升。酸雨对森林土壤、植物及生态系统各层面中的Ca均造成了长期广泛的影响,深入研究其影响程度与植物的应对机制,对我国酸雨危害区的酸雨防治和森林保育具有重大意义。     

Ectomycorrhizal specificity patterns in a mixed Pinus contorta and Picea engelmannii forest in Yellowstone National Park

We used molecular genetic methods to test two hypotheses, (i) that host plant specificity among ectomycorrhizal fungi would be common in a closed-canopy, mixed Pinus contorta-Picea engelmannii forest in Yellowstone National Park and (ii) that specificity would be more common in the early successional tree species, P. contorta, than in the invader, P. engelmannii. We identified 28 ectomycorrhizal fungal species collected from 27 soil cores. The proportion of P. engelmannii to P. contorta ectomycorrhizae was nearly equal (52 and 48%, respectively). Of the 28 fungal species, 18 composed greater than 95% of the fungal community. No species was associated exclusively with P. contorta, but four species, each found in only one core, and one species found in two cores were associated exclusively with P. engelmannii. These fungi composed less than 5% of the total ectomycorrhizae. Thus, neither hypothesis was supported, and hypothesized benefits of ectomycorrhizal specificity to both trees and fungi probably do not exist in this system.     

Ectomycorrhizal communities in a productive Tuber aestivum Vittad. orchard: composition, host influence and species replacement

Truffles (Tuber spp.) and other ectomycorrhizal species form species-rich assemblages in the wild as well as in cultivated ecosystems. We aimed to investigate the ectomycorrhizal communities of hazels and hornbeams that are growing in a 24-year-old Tuber aestivum orchard. We demonstrated that the ectomycorrhizal communities included numerous species and were phylogenetically diverse. Twenty-nine ectomycorrhizal taxa were identified. Tuber aestivum ectomycorrhizae were abundant (9.3%), only those of Tricholoma scalpturatum were more so (21.4%), and were detected in both plant symbionts with a variation in distribution and abundance between the two different hosts. The Thelephoraceae family was the most diverse, being represented by 12 taxa. The overall observed diversity represented 85% of the potential one as determined by a jackknife estimation of richness and was significantly higher in hazel than in hornbeam. The ectomycorrhizal communities of hornbeam trees were closely related phylogenetically, whereas no clear distribution pattern was observed for the communities in hazel. Uniform site characteristics indicated that ectomycorrhizal relationships were host mediated, but not host specific. Despite the fact that different plant species hosted diverse ectomycorrhizal communities and that the abundance of T. aestivum differed among sites, no difference was detected in the production of fruiting bodies.