Effect of plant species on communities of arbuscular mycorrhizal fungi in the Mongolian steppe

Communities of arbuscular mycorrhizal (AM) fungi were investigated in Stipa krylovii, Leymus chinensis (Poaceae), Allium bidentatum (Liliaceae), and Astragalus brevifolius (Fabaceae) in the Mongolian steppe to examine the effect of plant species on the communities in this study. The AM fungal communities were examined by molecular analysis based on the partial sequences of a small subunit of the ribosomal RNA gene. The sequences obtained were divided into 23 phylotypes by the sequence similarity >98%. Many of the AM fungal phylotypes included AM fungi previously detected in high-altitude regions in the Tibet and Loes plateaus, which suggested that these AM fungi may have wide distribution with stressful conditions of aridity and coldness. Among the 23 phylotypes, 12 phylotypes were found in all four plants, and 87.4% of the all obtained sequences were affiliated into these 12 types. For the distribution of the AM fungal phylotypes, overlapping of the phylotypes among the four plant species were significantly higher than that simulated by random chance. These results suggested that AM fungal communities were less diversified among the examined plant species.     

西藏高原针茅草地土壤因子对丛枝菌根真菌物种多样性的影响

在对西藏高原北部针茅草地根围土壤中的丛枝菌根(AM)真菌种类分离鉴定基础上,研究了藏北针茅草地的土壤质地、pH、有机质和有效磷含量对AM真菌孢子密度、分离频度、相对多度、重要值、物种多样性指数和均匀度的影响.结果表明: 针茅草地根围土壤中共分离鉴定出AM真菌3属15种,其中,球囊霉属9种、无梗囊霉属6种、盾巨孢囊霉属1种.球囊霉属和无梗囊霉属为藏北针茅草地AM真菌的优势属;近明球囊霉和光壁无梗囊霉为藏北高寒草原针茅属植物根围AM真菌的优势种.不同质地土壤中AM真菌孢子密度、分离频度、相对多度和重要值均表现出球囊霉属＞无梗囊霉属＞盾巨孢囊霉属的趋势;土壤pH值对AM真菌种群组成无明显影响,球囊霉属和无梗囊霉属真菌分离频度、相对多度和重要值随土壤pH升高而增加,盾巨孢囊霉属则呈现相反趋势;不同土壤有机质含量范围内,AM真菌孢子密度等各项指标均呈球囊霉属＞无梗囊霉属＞盾巨孢囊霉属,而AM真菌属的分布没有明显规律;土壤有效磷含量对AM真菌种丰度和孢子密度影响较小.研究区域内AM真菌物种多样性指数和均匀度随着土壤有效磷含量升高而增加.     

Communities, populations and individuals of arbuscular mycorrhizal fungi

Arbuscular mycorrhizal fungi in the phylum Glomeromycota are found globally in most vegetation types, where they form a mutualistic symbiosis with plant roots. Despite their wide distribution, only relatively few species are described. The taxonomy is based on morphological characters of the asexual resting spores, but molecular approaches to community ecology have revealed a considerable unknown diversity from colonized roots. Although the lack of genetic recombination is not unique in the fungal kingdom, arbuscular mycorrhizal fungi are probably ancient asexuals. The long asexual evolution of the fungi has resulted in considerable genetic diversity within morphologically recognizable species, and challenges our concepts of individuals and populations. This review critically examines the concepts of species, communities, populations and individuals of arbuscular mycorrhizal fungi.     

不同放牧强度下克氏针茅(Stipa krylovii)草原的根系特征

以呼伦贝尔克氏针茅草原不同放牧强度下的演替群落为对象,开展群落及其群落建群种的地下生物量和根系形态特征研究.结果表明:从轻度放牧到重度放牧,群落种类组成和根系功能群类型趋于简单化;群落地下生物量的空间分布形态呈“T”型;不同放牧强度下草原群落的建群种出现了明显替代现象,轻度放牧样地群落建群种为密丛型根系的克氏针茅,中度放牧为疏丛型根系的糙隐子草(Cleistogenes squarrosa,重度放牧为鳞茎型根系的碱韭(Allium polyrhizum);随着放牧强度的增大,群落建群种根冠比逐渐增加,分别为0.47、1.0、4.1,并且群落建群种根系数量、根系体积、根系生物量、比根长及根长密度等各指标均发生了明显变化.另外,3种放牧强度样地群落建群种根冠比、根长密度均与土壤速效氮含量呈现显著正相关(P＜0.05).     

Ancestral lineages of arbuscular mycorrhizal fungi (Glomales)

Using new and existing 18S rRNA sequence data, we show that at least five species of glomalean fungi lie outside the previously defined families and diverged very early in the evolution of that group. These five fungi would have been missed by many previous ecological studies because their sequences are not well matched to available taxon-specific primers and they do not stain well with the standard reagents used for morphological analysis. Based upon spore morphology, these species are currently assigned to Glomus and Acaulospora, and two of the species are dimorphic, exhibiting spore stages of both genera. This suggests that dimorphic spores are the ancestral state for the order and that one or the other morphology was lost in various lineages. Our analyses also show that Geosiphon pyriforme, a symbiont with cyanobacteria, is not necessarily a sister group of the Glomales; instead, it may be derived from mycorrhizal ancestors.     

The relationship between the diversity of arbuscular mycorrhizal fungi and grazing in a meadow steppe

Aims

To study the relationship between changes in soil properties and plant community characters produced by grazing in a meadow steppe grassland and the composition and diversity of spore-producing arbuscular mycorrhizal fungi (AMF).

Methods

A field survey was carried out in a meadow steppe area with a gradient of grazing pressures (a site with four grazing intensities and a reserve closed to grazing). The AMF community composition (characterized by spore abundance) and diversity, the vegetation characters and soil properties were measured, and root colonization by AMF was assessed.

Results

AMF diversity (richness and evenness) was higher under light to moderate grazing pressure and declined under intense grazing pressures. Results of multiple regressions indicated that soil electrical conductivity was highly associated with AMF diversity. The variation in AMF diversity was partially associated to the density of tillers of the dominant grass (Leymus chinensis), the above and below-ground biomass and the richness of the plant community.

Conclusions

We propose that the relationship between plants and AMF is altered by environmental stress (salinity) which is in turn influenced by animal grazing. Direct and indirect interactions between vegetation, soil properties, and AMF community need to be elucidated to improve our ability to manage these communities.     

内蒙古锡林郭勒克氏针茅退化草原土壤种子库特征

以内蒙古锡林郭勒地区克氏针茅(Stipa krylovii)草原不同退化等级群落为对象,研究克氏针茅退化草原可萌发土壤种子库特征.结果表明,随着草原退化程度的增加,不论是土壤总种子库还是持久土壤种子库,组成和密度均明显下降, 重度和极度退化草原土壤总种子库密度下降至仅为轻度退化草原的46.8%和11.1%.代表土壤总种子库的4月份取样, 轻度、中度、重度和极度退化草原各样地0～9 cm土壤种子库密度分别为2800、1278、1311和311粒·m-2;代表持久土壤种子库的6月底取样,4个样地土壤种子库密度分别为1667、967、334和167粒·m-2.多数植物土壤种子库主要分布在0～6 cm土层,各样地种子库密度随土壤深度的增加而减少,轻度、中度、重度和极度退化草原4月份0～6 cm土层种子库分别占总种子库(0～9 cm)的98.4、96.5、95.8 和85.7%.不同退化等级草原地上植被与土壤种子库的Sorensen相似性指数介于0.24～0.48.     

Genetic processes in arbuscular mycorrhizal fungi

Arbuscular mycorrhizal (AM) fungi (Glomeromycota) colonize roots of the majority of land plants and facilitate their mineral nutrient uptake. Consequently, AM fungi play an important role in terrestrial ecosystems and are becoming a component of sustainable land management practices. The absence of sexual reproductive structures in modern Glomeromycota combined with their long evolutionary history suggest that these fungi may represent an ancient asexual lineage of great potential interest to evolutionary biology. However, many aspects of basic AM fungal biology, including genome structure, within-individual genetic variation, and reproductive mode are poorly understood. These knowledge gaps hinder research on the mechanisms of AM fungal interactions with individual plants and plant communities, and utilization of AM fungi in agricultural practices. I present here the current state of research on the reproduction in AM fungi and indicate what new findings can be expected in the future.     

Communities of arbuscular mycorrhizal fungi in arable soils are not necessarily low in diversity

Communities of arbuscular mycorrhizal fungi (AMF) in five agricultural field sites of different management intensities were studied. Variable regions of the ribosomal RNA genes were used to detect and identify AMF directly within colonized roots. Roots from a continuous maize monoculture showed low AMF diversity, in agreement with previous reports on molecular diversity of AMF in agricultural soils. In contrast, a substantially higher diversity of AMF was found throughout the long term 'DOK' field experiment, where organic and conventional agricultural practices have been compared side by side since 1978. In this experiment, a 7-year crop rotation is performed under lower levels of inorganic fertilizer input and chemical pest control. These results are in good agreement with analyses of the spore community previously conducted in these field sites. In a third site, an organically managed leek field with soil of very high phosphate content reflecting the highly intensive conventional field history and intensive tillage, we detected a low-diversity community comparable to the maize monoculture. In addition to fungi from Glomus group A, which have previously been reported to dominate arable soils, we regularly found members of the genera Scutellospora, Paraglomus and Acaulospora. The genus Acaulospora was shown to occur more frequently early in the growing season, suggesting that the life history strategy of AMF may influence the active community at a given time. These data show that the diversity of AMF is not always low in arable soils. Furthermore, low-input agriculture involving crop rotation may provide better conditions to preserve AMF diversity, by preventing the selection for the few AMF taxa tolerating high nutrient levels.     

内蒙古克氏针茅草原生态系统-大气通量交换特征

基于内蒙古克氏针茅草原生态系统的涡动相关观测资料和小气候梯度系统观测资料,对研究区碳通量和水热通量的日、季动态进行了研究.结果表明：克氏针茅草原生态系统生长季的碳通量日动态呈U型曲线,即日出前释放CO₂,日出后开始吸收CO₂,正午前后达峰值,午后吸收CO₂减弱,日落后重新转为释放CO₂;9月白天CO₂吸收最为强烈,8月次之,10月最低.克氏针茅草原的感热和潜热通量的日动态均呈倒U型曲线,与碳通量日动态相反,即白天感热和潜热通量多为正值,夜间感热为负值,潜热接近于零;感热通量以5月最高,潜热通量以9月最高.冬季草地为弱碳源,CO₂通量较小,夏季表现为明显的碳汇.     

丛枝菌根真菌生物地理学研究进展

丛枝菌根真菌(arbuscular mycorrhizal fungi,AMF)普遍存在于陆地生态系统中,能与绝大多数高等植物形成菌根共生体系。AMF能够促进植物对矿质养分的吸收,增强植物的抗逆能力,在维持生态系统稳定性和生产力中发挥着重要作用。AMF生物地理学主要研究AMF的生物地理分布格局和群落构建机制,对于理解AMF在不同生态系统中的重要性至关重要。总结了AMF生物地理学最新研究进展及研究方法,提出了AMF生物地理学研究理论框架。认为AMF在自然界中并非简单随机分布,宿主植物、地理因子、气候因子和土壤因子共同决定AMF的群落结构,不同尺度下的AMF群落构建符合生态位-中性连续统假说,但在不同尺度下这些驱动因子的相对重要性不同。在全球尺度和区域尺度下,AMF的地理分布格局主要受地理距离和气候因子的影响,中性理论的作用大于生态位理论。随着空间尺度的缩减,宿主植物和环境因子对AMF群落的影响胜过地理距离和扩散限制的作用,生态位理论取代中性理论在AMF群落构建中的主导地位。此外,很多研究发现,同一生境中AMF的群落构建机制并非一成不变,会随环境的变化而发生改变。在未来的研究中,应在野外调查和公共数据库的基础上加强整合分析和数据挖掘工作,从而进一步丰富和完善AMF生物地理学理论。     

Cd-tolerant arbuscular mycorrhizal (AM) fungi from heavy-metal polluted soils

Spores of arbuscular mycorrhizal (AM) fungi were isolated from two heavy-metal polluted soils in France via trap culture with leek (Allium porrum L.). Preliminary identification showed that the predominant spore type of both cultures (P2 and Cd40) belongs to the Glomus mosseae group. Their sensitivity to cadmium was compared to a laboratory reference strain (G. mosseae) by in vitro germination tests with cadmium nitrate solutions at a range of concentrations (0 to 100 mg L^–1) as well as extracts from a metal-polluted and unpolluted soils. Both cultures of AM fungi from heavy-metal polluted soils were more tolerant to cadmium than the G. mosseae reference strain. The graphically estimated EC₅₀ was 0.8 mg L^–1 Cd (concentration added to the test device) for G. mosseae and 7 mg L^–1 for P2 culture, corresponding to effective Cd concentrations of approximately 50–70 g L^–1 and 200–500 g L^–1, respectively. The extract of the metal-polluted soil P2 decreased germination of spores from the reference G. mosseae but not from P2 culture. However, the extracts of two unpolluted soils with different physico-chemical characteristics did not affect G. mosseae, whereas germination of P2 spores was markedly decreased in the presence of one of the extracts. These results indicate a potential adaptation of AM fungi to elevated metal concentrations in soil. The tested spores may be considered as metal-tolerant ecotypes. Spore germination results in presence of soil extracts show the difficulty of assessing the ecotoxic effect of metals on AM fungi without considering other soil factors that may interfere in spore germination and hyphal extension.     

Biolistic transformation of arbuscular mycorrhizal fungi

Gene transfer systems have proved effective for the transformation of a range of organisms for both fundamental and applied studies. Biolistic transformation is a powerful method for the gene transfer into various organisms and tissues that have proved recalcitrant to more conventional means. For fungi, the biolistic approach is particularly effective where protoplasts are difficult to obtain and/or the organisms are difficult to culture. This is particularly applicable to arbuscular mycorrhizal (AM) fungi, being as they are obligate symbionts that can only be propagated in association with intact plants or root explants. Furthermore, these fungi are aseptate and protoplasts cannot be released. Recent advancements in gene transformation systems have enabled the use of biolistic technology to introduce foreign DNA linked to molecular markers into these fungi. In this review we discuss the development of transformation strategies for AM fungi by biolistics and highlight the areas of this technology which require further development for the stable transformation of these elusive organisms.     

Sex-specific variation in the interaction between Distichlis spicata (Poaceae) and mycorrhizal fungi

Associations between mycorrhizal fungi and plants can influence intraspecific competition and shape plant population structure. While variation in plant genotypes is known to affect mycorrhizal colonization in crop systems, little is known about how genotypes affect colonization in natural plant populations or how plant sex might influence colonization with mycorrhizal fungi in plant species with dimorphic sexual systems. In this study, we analyzed mycorrhizal colonization in males and females of the wetland dioecious grass Distichlis spicata, which has spatially segregated sexes. Our results suggest that D. spicata males and females interact with mycorrhizal fungi differently. We discuss the implications for the role of this sex-specific symbiotic interaction in the maintenance of the within-population sex ratio bias of D. spicata.     

An evidence-based consensus for the classification of arbuscular mycorrhizal fungi (Glomeromycota)

The publication of a large number of taxon names at all levels within the arbuscular mycorrhizal fungi (Glomeromycota) has resulted in conflicting systematic schemes and generated considerable confusion among biologists working with these important plant symbionts. A group of biologists with more than a century of collective experience in the systematics of Glomeromycota examined all available molecular–phylogenetic evidence within the framework of phylogenetic hypotheses, incorporating morphological characters when they were congruent. This study is the outcome, wherein the classification of Glomeromycota is revised by rejecting some new names on the grounds that they are founded in error and by synonymizing others that, while validly published, are not evidence-based. The proposed “consensus” will provide a framework for additional original research aimed at clarifying the evolutionary history of this important group of symbiotic fungi.     

围栏封育和自由放牧对克氏针茅草原植被空间异质性的影响

群落组成结构的异质性是群落动态的主要属性。本研究采用样线、样方调查方法,基于物种频度与幂函数法则,分析了内蒙古克氏针茅草原围栏封育和自由放牧条件下群落物种结构与空间异质性的变化。结果表明:在围栏外和围栏内不同利用方式下,物种均呈现集中分布;从水分生态型、生活型、生育型、种子传播型和植物科属等不同功能群类别来看,旱生植物和中旱生植物广泛分布,生活型以多年生草本植物为主,依靠重力传播种子的植物占有相对较大的比例,达到60%以上,禾本科、百合科、菊科和豆科等适牧植物具有相对优势;在围栏保护状态下,物种多样性增加,群落异质性降低,放牧退化草原开始向群落状况良好的草原类型演替。     

Enkianthus campanulatus (Ericaceae) is commonly associated with arbuscular mycorrhizal fungi

Enkianthus is the most basal extant genus in the phylogeny of ericaceous plants. Its members harbor arbuscular mycorrhiza (AM)-like hyphal structures in their roots but, as yet, no study has surveyed the AM fungal species component. Roots from six species of Enkianthus were collected from five distantly located sites in Japan. Intracellular hyphal coils were observed in the root cortical cells of all species. Fungal DNA sequences of the small subunit ribosomal RNA gene were obtained from 73 of 75 segments of Enkianthus campanulatus roots by PCR using either AML2 or NS31/AM1primer pairs. Results indicated that all E. campanulatus trees were extensively associated with Glomus spp. A phylogenetic analysis showed that 71 root segments harbored fungi belonging to Glomus group A. Among eight delineated clades, seven did not nest with any known AM fungal species. One clade was detected in all roots at all sites at relatively high frequencies, but the rest were detected sporadically at each site. The placement of sequences from distantly located sites into a single clade without known AM fungal species suggests the common association of E. campanulatus with particular AM fungal taxa.     

Specific arbuscular mycorrhizal fungi associated with non-photosynthetic Petrosavia sakuraii (Petrosaviaceae)

Mycorrhizal fungi in roots of the achlorophyllous Petrosavia sakuraii (Petrosaviaceae) were identified by molecular methods. Habitats examined were plantations of the Japanese cypress Chamaecyparis obtusa in Honshu, an evergreen broad-leaved forest in Amami Island in Japan and a mixed deciduous and evergreen forest in China. Aseptate hyphal coils were observed in root cortical cells of P. sakuraii, suggesting Paris-type arbuscular mycorrhiza (AM). Furthermore, hyphal coils that had degenerated to amorphous clumps were found in various layers of the root cortex. Despite extensive sampling of P. sakuraii from various sites in Japan and China, most of the obtained AM fungal sequences of the nuclear small subunit ribosomal RNA gene were nearly identical and phylogenetic analysis revealed that they formed a single clade in the Glomus group A lineage. This suggests that the symbiotic relationship is highly specific. AM fungi of P. sakuraii were phylogenetically different from those previously detected in the roots of some mycoheterotrophic plants. In a habitat in C. obtusa plantation, approximately half of the AM fungi detected in roots of C. obtusa surrounding P. sakuraii belonged to the same clade as that of P. sakuraii. This indicates that particular AM fungi are selected by P. sakuraii from diverse indigenous AM fungi. The same AM fungi can colonize both plant species, and photosynthates of C. obtusa may be supplied to P. sakuraii through a shared AM fungal mycelial network. Although C. obtusa plantations are widely distributed throughout Japan, P. petrosavia is a rare plant species, probably because of its high specificity towards particular AM fungi.