西北干旱区AM真菌多样性研究

为了探明我国新疆、甘肃河西走廊等西北干旱区丛枝菌根(AM)真菌的资源与分布, 我们调查了荒漠、草原、针阔叶混交林、草甸和绿洲农田5种植被类型中AM真菌类群分布特征, 以及不同生态因子对AM真菌多样性特征的影响。在西北干旱区的26科60种植物的根区土壤中, 共分离鉴定出AM真菌6属40种, 其中35个鉴定到种, 5个鉴定到属。在调查区域内球囊霉属(Glomus)为优势属, G. claroideum为优势菌种。AM真菌在不同植被类型中的发生和分布规律明显不同: 内养囊霉属(Entrophospora)只在荒漠中有分布, 类球囊霉属(Paraglomus)在绿洲农田和荒漠中有分布, 原囊霉属(Archaeospora)在草原、荒漠和绿洲农田中分布, 盾巨孢囊霉属(Scutellospora)在草甸、针阔叶混交林和草原中有分布。研究观察到在绿洲农田和草甸中AM真菌的物种多样性指数高于其他植被类型, 这反映出在干旱区土壤水分条件可能是AM真菌发生和分布的重要制约因素。     

泰山丛枝菌根真菌群落结构特征

2007年对泰山植被根围内丛枝菌根(arbuscular mycorrhiza,AM)真菌群落组成、数量、分布及其与植物多样性的关系进行了研究。从泰山傲徕峰、黑龙潭库区等样地共分离出4属16种AM真菌:球囊霉属Glomus 9种、无梗囊霉属Acaulospora 4种、巨孢囊霉属Gigaspora 2种和盾巨孢囊霉属Scutellospora1种。其中,球囊霉属Glomus及聚球囊霉Glomus fasciculatum的孢子密度、相对多度、分布频度和重要值均最高,分别为泰山植被区根围内AM真菌优势属和优势种。各样地之间Sorenson相似系数在0.60和0.85之间。植被数量与孢子密度(r=0.80,p0.01)、植物种的丰富度与AM真菌种的丰富度(r=0.77,p0.01)以及与孢子密度(r=0.59,p0.01)均呈极显著正相关关系。研究结果表明植物多样性对于提高AM真菌多样性发挥极为重要的作用。     

Arbuscular mycorrhizal fungal diversity and species dominance in a temperate soil with long-term conventional and low-input cropping systems

The aim of this work was to study the effect of long-term contrasting cropping systems on the indigenous arbuscular mycorrhizal fungal (AMF) spore populations in the soil of a field experiment located in western Finland. Conventional and low-input cropping systems were compared, each with two nutrient management regimes. The conventional cropping system with a non-leguminous 6-year crop rotation (barley–barley–rye–oat–potato–oat) was fertilized at either full (rotation A) or half (rotation B) the recommended rate. In the low-input cropping system, plant residues were returned to the plots either as such (rotation C) or composted (rotation D). In the rotation of this system, 1 year with barley was replaced by clover, and oat was cultivated mixed with pea. Thus, the 6-year rotation was barley–red clover–rye–oat + pea–potato–oat + pea. Each rotation was replicated three times, starting the 6-year rotation in three different years, these being designated point 1, point 2, and point 3, respectively. In the low-input system, biotite and rock phosphate were used to compensate for K and P in the harvested yield, while animal manure was applied at the start only. After 13 years, rotation points 1 and 3 were studied. Barley was the standing crop in all plots of rotation point 1, while oat and oat + pea were grown in rotations C and D, respectively. AMF spores were studied by direct extraction and by trapping, sampled on 15 June and 15 August. In addition, a special assay was designed for isolation of fast colonizing, dominating AMF. The cropping system did not significantly affect AMF spore densities, although the low-input cropping system with composted plant residues had the highest density with 44 spores on average and the conventional system with full fertilization 24 spores per 100 cm³ soil in the autumn samples. Species richness was low in the experimental area. Five Glomus spp., one Acaulospora, and one Scutellospora were identified at the species level. In addition to these, three unidentified Glomus spp. were found. Species richness was not affected by cropping system, rotation point, or their interactions. The Shannon–Wiener index of AMF spore distributions was significantly higher in the fully fertilized than in the half-fertilized conventional plots. Glomus claroideum was the most commonly identified single species in the experimental area. It occurred in all the cropping systems and their various rotation points, representing about 30% of the total number of identified spores. In August, G. claroideum accounted for as much as 45–55% of the total numbers of spores identified in the conventional system with halved fertilization. In contrast, Glomus mosseae occurred more commonly in June (26%) than in August (9%). A bioassay using roots as inoculum for isolation and culture of dominating AMF was successfully developed and yielded only G. claroideum. This indicates a high probability of being able to more generally identify, isolate, and culture fast colonizing generalist AMF for use as inoculants in agriculture and horticulture.     

长期施肥对砂姜黑土丛枝菌根真菌群落的影响

在农业生态系统中,丛枝菌根真菌(AM真菌)与很多作物的根系都存在互惠互利的共生关系,与作物的生长和健康密切相关,同时,这类特殊的真菌群落也会受到施肥等农业措施的影响.本研究依托长期定位试验4个试验处理(不施肥、单施化肥、化肥配施秸秆、化肥配施粪肥),研究砂姜黑土AM真菌群落对不同施肥措施的响应及其影响因素,探索不同处理AM真菌指示种的存在.结果表明: 砂姜黑土中的主要AM真菌类群为原囊霉科、多孢囊霉科、巨孢囊霉科、近明球囊霉科、球囊霉科和类球囊霉科;其中类球囊霉属在化肥和有机物料配施中具有显著指示作用.与对照相比,长期单施化肥显著改变AM真菌群落结构并降低其多样性,配施秸秆处理进一步降低AM真菌群落多样性,而配施粪肥明显缓解因施用化肥而造成的多样性减少现象.检验发现,导致AM真菌群落变化最主要的影响因素是土壤pH和可溶性碳.总之,长期不同有机物料和化肥配施对砂姜黑土AM真菌群落结构和多样性会产生不同影响,其中化肥配施粪肥更有利于土壤AM真菌群落多样性的维持.     

AM真菌物种多样性:生态功能、影响因素及维持机制

AM真菌物种多样性是土壤生态系统生物多样性的重要组分之一。尽管对AM真菌多样性已有多年研究,但是,已有研究绝大多数仅停留在对AM真菌群落种属解析层面上,对AM真菌物种多样性生态功能及维持机制方面的认识较浅。从生态功能、影响因素及维持机制3个方面系统地综述了近年来AM真菌多样性领域的研究进展。认为AM真菌多样性对植物群落生产力的调控机制及结合理论与实践解析AM真菌多样性维持机制是该领域未来的重点研究方向。     

长期施肥对麦田大型土壤动物群落结构的影响

于2007、2008年春季在中国科学院封丘农业生态实验站,研究长期定位施肥对麦田大型土壤动物群落结构的影响。调查共获得大型土壤动物3068头,隶属8纲,19目,28科。不同施肥处理条件下土壤动物类群数差异显著(F=2.51, P＜0.05, df=55),个体数差异极显著(F=8.99, P＜0.01, df=55)。结果显示,大型土壤动物的类群数和个体数在有机肥和营养均衡条件下较高,缺磷条件下较低。土壤动物多样性指数(H')以有机肥和营养均衡条件下显著高于营养不均衡和不施肥处理(P＜0.05)。主成分分析结果显示,第一主成分贡献值47.14%,第二主成分贡献值30.10%。经线性检验第一主成分与土壤动物个体数(y=0.335x-2.163,R²=0.51)和类群数(y=0.042x-1.25,R²=0.67)均呈线性关系。总之,长期施用有机肥和营养均衡大型土壤动物群落构成相似。土壤有机质和磷含量是影响土壤动物群落构主要生态因子。     

Arbuscular mycorrhizal fungal propagules in a salt marsh

The tolerance of indigenous arbuscular mycorrhizal fungi (AMF) to stressful soil conditions and the relative contribution of spores of these fungi to plant colonization were examined in a Portuguese salt marsh. Glomus geosporum is dominant in this salt marsh. Using tetrazolium as a vital stain, a high proportion of field-collected spores were found to be metabolically active at all sampling dates. Spore germination tests showed that salt marsh spores were not affected by increasing levels of salinity, in contrast to two non-marsh spore isolates, and had a significantly higher ability to germinate under increased levels of salinity (20) than in the absence of or at low salinity (10). Germination of salt marsh spores was not affected by soil water levels above field capacity, in contrast to one of the two non-marsh spore isolates. For the evaluation of infectivity, a bioassay was established with undisturbed soil cores (containing all types of AM fungal propagules) and soil cores containing only spores as AM fungal propagules. Different types of propagules were able to initiate and to expand the root colonization of a native plant species, but spores were slower than mycelium and/or root fragments in colonizing host roots. The AM fungal adaptation shown by this study may explain the maintenance of AMF in salt marshes.     

Preliminary assessment of arbuscular mycorrhizal fungal diversity and community structure in an urban ecosystem

Arbuscular mycorrhizal fungal (AMF) species richness, composition, spore density and diversity indices were evaluated in the Phoenix metropolitan area, Arizona, USA at 20 sampling sites selected to represent the four predominant land-use types found in the greater urban area: urban-residential, urban non-residential, agriculture and desert. AMF spores were extracted and identified from soil samples and from trap cultures established using soil collected at each site. Data were analyzed according to land use, land-use history, soil chemistry and vegetation characteristics at each site. Current agricultural sites were associated with decreased spore densities and historically agricultural sites with decreased species richness. Overall species composition was similar to that previously reported for the Sonoran desert, but composition at each sampling site was influenced by the vegetation from which samples were collected. Sites with the highest degrees of similarity in AMF species composition were also similar to each other in native plants or land use. Conversely, sites with the lowest similarity in AMF composition were those from which the majority of samples were collected from non-mycorrhizal plants, predominately ectomycorrhizal plants or bare soil. Spores of Glomus microggregatum were most abundant in urban sites, while those of G. eburneum were most abundant in desert and agricultural sites. Further studies are needed to determine the functional implications of shifts in AMF communities in urban ecosystems, including effects on plant primary productivity.     

Soil fungal community changes in response to long-term fire cessation and N fertilization in tallgrass prairie

In grasslands, fire management and fertilization are established drivers of plant community change, but associated soil fungal responses are less well defined. We predicted that soil fungal communities would change seasonally, that decades of fire cessation and nitrogen (N) fertilization would alter fungal distributions, and that plant and fungal community change would be correlated. Surface soils were sampled monthly for 1 y from a 30-y fire by fertilization experiment to evaluate fungal community dynamics and assess correlation with plant community heterogeneity. ITS gene community composition was seasonally stable, excepting increased arbuscular mycorrhizal fungal summer abundance in the burned, fertilized treatment. Long-term treatments affected soil fungal and plant communities, with correlated heterogeneity patterns. Despite woody encroachment in the fire cessation treatment, soil fungal communities did not resemble those of forests. This study provides evidence supporting the strength of feedbacks between fungal and plant community change in response to long-term grassland fire and N management changes.     

长期不同施肥方式对麦田杂草群落的影响

以太湖地区农业科学研究所31a的长期肥料定位试验田为材料,分别于2011—2012年小麦苗期、拔节期和收获期进行了杂草群落调查,研究杂草类型与密度的分布、杂草多样性指数的变化,并对杂草种群分布与土壤养分因子进行冗余分析。结果表明:小飞蓬、看麦娘、大巢菜、稻槎菜、通泉草是本地区小麦生长期的主要杂草类型;随着小麦的生长以及氮肥、有机肥的施入,杂草密度呈下降趋势;施入有机肥降低了麦季杂草的群落多样性指数,在小麦生长的不同时期均衡施肥的CNPK处理以及不施肥的C0处理的群落各项多样性指数能维持在一个较高的水平。RDA分析显示土壤氮含量以及有机质含量与第一排序轴相关性大,是对杂草分布影响最大的两个土壤养分因子。太湖地区稻麦两熟制条件下,长期有机无机肥料单一或配合投入可显著影响麦田杂草的群落组成,其中氮肥和有机肥的施入能显著降低杂草密度;土壤养分的差异影响田面杂草密度和优势种群,均衡施肥能降低优势杂草种群的优势地位,抑制其发生危害程度,提高农田生态系统的生产力及稳定性。     

Arbuscular mycorrhizal fungal community differs between a coexisting native shrub and introduced annual grass

Arbuscular mycorrhizal fungi (AMF) have been implicated in non-native plant invasion success and persistence. However, few studies have identified the AMF species associating directly with plant invaders, or how these associations differ from those of native plant species. Identifying changes to the AMF community due to plant invasion could yield key plant–AMF interactions necessary for the restoration of native plant communities. This research compared AMF associating with coexisting Bromus tectorum, an invasive annual grass, and Artemisia tridentata, the dominant native shrub in western North America. At three sites, soil and root samples from Bromus and Artemisia were collected. Sporulation was induced using trap cultures, and spores were identified using morphological characteristics. DNA was extracted from root and soil subsamples and amplified. Sequences obtained were aligned and analyzed to compare diversity, composition, and phylogenetic distance between hosts and sites. Richness of AMF species associated with Artemisia in cultures was higher than AMF species associated with Bromus. Gamma diversity was similar and beta diversity was higher in AMF associated with Bromus compared to Artemisia. AMF community composition differed between hosts in both cultures and roots. Two AMF species (Archaeospora trappei and Viscospora viscosum) associated more frequently with Artemisia than Bromus across multiple sites. AMF communities in Bromus roots were more phylogenetically dispersed than in Artemisia roots, indicating a greater competition for resources within the invasive grass. Bromus associated with an AMF community that differed from Artemisia in a number of ways, and these changes could restrict native plant establishment.     

中国丛枝菌根真菌分子多样性

【背景】随着分子生物学技术的发展,通过分子生物学方法鉴定丛枝菌根真菌(Arbuscular mycorrhizal fungi,AMF)多样性的研究越来越多。目前,AMF在全球的分子多样性及其分布已引起关注,但在我国关于AMF分子多样性的研究进展状况尚未见报道。【目的】探究我国AMF分子多样性及其分布,研究不同气候区域以及不同生态系统中AMF分子多样性并揭示其分布规律,为我国AMF在分子水平上的研究提供基础数据。【方法】利用Davison于2015年发表于"Science"上的一篇有关全球AMF分子多样性的数据库,选用中国AMF多样性的数据,并补充了近年来已发表的该数据库中未涉及的数据,建立中国AMF分子多样性的新数据库。【结果】共得到145个分子生物学鉴定出的虚拟种(Virtualtaxa,VT),隶属于8科12属。对于不同生态系统而言,草地生态系统中AMF的分子多样性最高,占到总VT数的60.7%,人为生态系统和森林生态系统也分别达到55.2%和43.4%。同样,AMF的VT数也随气候区域的不同而有所变化,温带气候区域所占比例最高(64.1%),亚热带气候区域次之(60.7%),极北气候区域最少(20.7%)。而对于根系和土壤样品中AMF的分子多样性而言,根系样品中发现的AMF分子多样性(80.0%)高于土壤中AMF的分子多样性(48.3%)。进一步对不同生态系统、气候区域和样品来源分别在属的水平上进行研究,同样也存在一定的差异。【结论】我国具有丰富的AMF分子多样性,且不同气候区域、生态系统以及样品来源都影响着AMF的分子多样性及其分布。     

Arbuscular mycorrhizal fungal community composition associated with Juniperus brevifolia in native Azorean forest

The communities of glomeromycotan fungi (arbuscular mycorrhizal fungi, AMF) under native Juniperus brevifolia forest from two Azorean islands, Terceira and São Miguel, were compared, mainly by spore morphology, and when possible, by molecular analysis. Thirty-nine morphotypes were detected from 12 genera. Glomeromycotan fungal richness was similar in Terceira and São Miguel, but significantly different among the four fragments of native forest. Spore diversity and community composition differed significantly between the two islands. The less degraded island, Terceira, showed 10 exclusive morphotypes including more rare types, whereas the more disturbed forest on São Miguel showed 13 morphs, mostly of common types. Forests from Terceira were dominated by Acaulosporaceae and Glomeraceae. Whereas members of Acaulosporaceae, Glomeraceae and Ambisporaceae were most frequent and abundant in those from São Miguel. Spore abundance was greatest on Terceira, and correlated with soil chemical properties (pH), average monthly temperature and relative humidity.     

长期肥料施用对华北大田玉米土壤丛枝菌根真菌群落结构的影响

华北平原是我国北方最重要的粮食主产区之一,过去30年间其粮食产量稳定提高的同时,肥料等生产资料的投入也大幅增加。丛枝菌根(Arbuscular mycorrhiza,AM)真菌作为一类重要的土壤功能性微生物,研究施肥措施对其群落、功能的影响,在挖掘农田有益微生物的潜在作用方面具有重要意义。本研究采用分子生物学方法测定了华北平原典型集约化农田体系中不同养分用量和投入模式对土壤AM真菌群落的影响。试验样地于2007年开始对冬小麦-夏玉米轮作体系进行施肥处理:有机肥(牛粪9 000kg/hm~2)、高量化肥(296.4kg N/hm~2、177.3kg P/hm~2、287.8kg K/hm~2,较当地常规化肥施用量上调25%)、低量化肥(74.1kg N/hm~2、44.3kg P/hm~2、71.9kg K/hm~2,较当地常规化肥施用量下调75%)和不施肥对照,每个处理3次重复,于2011年10月玉米收获季进行土壤取样。试验获得的31个OTUs主要属于Glomerales目(球囊霉目)(Glomus Group A/B)及Diversisporales目(多孢囊霉目),其中部分AM真菌类群如Glo12、Aca3、Scutellospora aurigloba(Scut.aurigloba全球盾巨孢囊霉)以及Div2仅在有机肥处理中被检测到。与其他处理相比,有机肥处理的116bp片段(Glo9/12/13/14;Fun1/2;Rhi1)相对丰度显著升高,而140bp片段(Uncultured Glomus)(球囊霉属)则主要受到化肥处理的促进。对于Diversisporales目AM真菌所对应的片段(141、142、169bp),施肥明显对其有促进作用。4年的肥料施用对土壤AM真菌物种多样性无显著影响,施肥处理0–20、20–40、40–60cm深度土壤AM真菌群落的多样性水平分别为1.14–1.78、1.61–1.84、1.23–2.07,丰富度分别为3.7–7.5、8.0–8.3、5.0–7.3。表明深层土壤是AM真菌重要的库,多样性水平并不低,对于维持土壤微生物多样性水平具有重要作用。本研究发现,p H值是影响表层土壤AM真菌群落分布的主要原因,土壤全氮、全碳、速效磷等肥力因子也是影响表层土壤AM真菌群落分布的重要因子,而深层土壤AM真菌的群落组成与固有的盐分特征(土壤电导率EC)有着较好的相关性。     

菌根真菌多样性与植物多样性的相互作用研究进展

菌根共生双方多样性影响着生态系统的过程与功能。菌根真菌-寄主植物之间的共生组合存在偏好性或特异性,这导致菌根真菌对寄主植物的效益差异和寄主植物对菌根真菌的利益差别：两者在互利共生过程中不仅相互选择,还存在相互促进与制约的关系(如互补与选择效应、竞争),从而在一定程度上决定生态系统的演化与发展。本文概述了植物多样性与菌根真菌多样性的相互影响,探讨了两者互作可能存在的调控因素与机制,对存在的问题和争议进行了总结,并提出了进一步研究的方向。深入阐明植物多样性与菌根真菌多样性之间的互作关系,将丰富生物共生学理论,增强菌根应用潜力及生物多样性的维持。     

Effects of long-term fertilization on AM fungal community structure and Glomalin-related soil protein in the Loess Plateau of China

Arbuscular mycorrhizal fungi (AMF) are crucial for ecosystem functioning, and thus have potential use for sustainable agriculture. In this study, we investigated the impact of organic and mineral fertilizers on the AMF community composition and content of Glomalin-related soil protein (GRSP) in a field experimental station which was established in 1979, in the Loess Plateau of China. Roots and soils were sampled three times during the growing period of winter wheat in 2008. The treatments including: N (inorganic N), NP (inorganic N and P), SNP (straw, inorganic N and P), M (farmyard manure), MNP (farmyard manure, inorganic N and P), and CK (no fertilization). AMF communities of root and soil samples were analyzed using PCR-DGGE, cloning and sequencing techniques; and GRSP content was determined by Bradford assay. Our results indicated that spore density, GRSP, and AMF community varied significantly in soils of long-term fertilization plots at three different wheat growing stages. The effects of wheat growing period on AMF community in roots were much more evident than fertilization regimes. However, the diversity of AMF was low in our study field. Up to five AMF phylotypes appeared in each sample, with the overwhelming dominance of a Glomus-like phylotype affiliated to G. mosseae. GRSP content was correlated positively with organic carbon, total phosphorus, available phosphorus, soil pH, and spore densities, but correlated negatively with soil C/N (P?<?0.05). The results of our study highlight that the richness of AMF in Loess Plateau agricultural region is low, and long-term fertilization, especially amendments with manure and straw, has beneficial effects on accumulation of soil organic carbon, spore density, GRSP content, and AMF diversity. Host phenology, edaphic factors (influenced by long-term fertilization), and habitats interacted to affect the AMF community and agoecosystem functioning. Additionally, soil moisture and pH make a greater contribution than other determined soil parameters to the AMF community dynamics in such a special semi-arid agroecosystem where crops rely greatly on rainfall.     

Soil mycorrhizal and nematode diversity vary in response to bioenergy crop identity and fertilization

The mandate by the Energy Independence and Security Act of 2007 to increase renewable fuel production in the USA has resulted in extensive research into the sustainability of perennial bioenergy crops such as switchgrass (Panicum virgatum) and miscanthus (Miscanthus× giganteus). Perennial grassland crops have been shown to support greater aboveground biodiversity and ecosystem function than annual crops. However, management considerations, such as what crop to plant or whether to use fertilizer, may alter belowground diversity and ecosystem functioning associated with these grasslands as well. In this study, we compared crop type (switchgrass or miscanthus) and nitrogen fertilization effects on arbuscular mycorrhizal fungal (AMF) and soil nematode abundance, activity, and diversity in a long‐term experiment. We quantified AMF root colonization, AMF extra‐radical hyphal length, soil glomalin concentrations, AMF richness and diversity, plant‐parasitic nematode abundance, and nematode family richness and diversity in each treatment. Mycorrhizal activity and diversity were higher with switchgrass than with miscanthus, leading to higher potential soil carbon contributions via increased hyphal growth and glomalin production. Plant‐parasitic nematode (PPN) abundance was 2.3 ×  higher in miscanthus plots compared to switchgrass, mostly due to increases in dagger nematodes (Xiphinema). The higher PPN abundance in miscanthus may be a consequence of lower AMF in this species, as AMF can provide protection against PPN through a variety of mechanisms. Nitrogen fertilization had minor negative effects on AMF and nematode diversity associated with these crops. Overall, we found that crop type and fertilizer application associated with perennial bioenergy cropping systems can have detectable effects on the diversity and composition of soil communities, which may have important consequences for the ecosystem services provided by these systems.