丛枝菌根真菌脂类代谢对共生信号调控的响应和反馈

丛枝菌根(AM)真菌是自然生态系统中分布最为广泛的真菌之一,在自然界物质循环和能量流动中发挥着重要作用。经过长期的协同进化,AM真菌和宿主植物之间形成了完美的互惠互利的共生关系,而真菌的脂类代谢可能是揭示共生秘密的关键所在。本文综述了AM真菌脂类代谢在共生关系建立和维持中关键作用的最新研究进展,重点探讨了AM真菌脂类代谢对共生信号调控的响应和反馈机制,主要包括:AM真菌脂类存储和释放对共生和非共生状态的响应,以及脂类代谢产物变化与共生营养传递之间的关系;脂类分解过程在共生建立初期对信号分子调控发生的响应,以及相应的物质转化和能量代谢;菌根共生互惠互利关系维持中,真菌脂类代谢与信号分子交流通道的相互渗透和影响。本文对于理解菌根共生机制,促进菌根在生产中的应用具有促进作用。     

Competitive interactions are mediated in a sex‐specific manner by arbuscular mycorrhiza in Antennaria dioica

• Plants usually interact with other plants, and the outcome of such interaction ranges from facilitation to competition depending on the identity of the plants, including their sexual expression. Arbuscular mycorrhizal (AM) fungi have been shown to modify competitive interactions in plants. However, few studies have evaluated how AM fungi influence plant intraspecific and interspecific interactions in dioecious species.
• The competitive abilities of female and male plants of Antennaria dioica were examined in a greenhouse experiment. Females and males were grown in the following competitive settings: (i) without competition, (ii) with intrasexual competition, (iii) with intersexual competition, and (iv) with interspecific competition by Hieracium pilosella – a plant with similar characteristics to A. dioica. Half of the pots were grown with Claroideoglomus claroideum, an AM fungus isolated from the same habitat as the plant material. We evaluated plant survival, growth, flowering phenology, and production of AM fungal structures.
• Plant survival was unaffected by competition or AM fungi. Competition and the presence of AM fungi reduced plant biomass. However, the sexes responded differently to the interaction between fungal and competition treatments. Both intra‐ and interspecific competition results were sex‐specific, and in general, female performance was reduced by AM colonization. Plant competition or sex did not affect the intraradical structures, extraradical hyphae, or spore production of the AM fungus.
• These findings suggest that plant sexual differences affect fundamental processes such as competitive ability and symbiotic relationships with AM fungi.

     

Developmental genetics and evolution of symbiotic structures in nitrogen-fixing nodules and arbuscular mycorrhiza.

Genetic and molecular mechanisms of development are compared for two major plant-microbe endosymbioses: N(2)-fixing nodules (with rhizobia or actinomycetes Frankia) and arbuscular mycorrhiza (with Glomales fungi). Development from the primordia formed de novo in root tissues is common for all known types of N(2)-fixing nodules. However, their structure varies greatly with respect to: (i) tissue topology (location of vascular bundles is peripherical in legumes or central in non-legumes); (ii) position of nodule primordium (inner or outer cortex in legumes, pericycle in non-legumes); (iii) stability of apical meristem (persistent in the indeterminate nodules, transient in the determinate ones). In addition, legumes vary in ability to form compartments harboring endosymbiotic rhizobia and located intercellularly (infection threads) and intracellularly (symbiosomes). Using pea (Pisum sativum) symbiotic mutants, the nodule developmental program is dissected into a range of spatially and temporarily differentiated steps comprising four sub-programs (development of endosymbiotic compartments; nodule histogenesis; autoregulation of nodulation; bacteroid differentiation). The developmental mutations are suggested in some cases to reverse the endosymbiotic system into the morphologically simpler forms some of which may correspond to the ancestral stages of nodule evolution. The origin of legume-rhizobial and actinorhizal symbioses is suggested to be based on a set of preadaptations many of which had been evolved in angiosperms during coevolution with arbuscular mycorrhizal fungi (e.g., inter- and intracellular maintenance of symbionts, their control via defence-like reactions and recognition of chitin-like molecules). An analysis of parallel morphological variation in symbiotic mutants and wild-growing legume species enables us to reconstruct the major stages of evolution for N(2)-fixing symbioses.     

丛枝菌根结构与功能研究进展

丛枝菌根(arbuscular mycorrhiza,AM)是陆地生态系统中分布最广泛、最重要的互惠共生体之一,对提高植物抗逆性、修复污染生境、保持生态系统稳定与可持续生产力的作用显著.AM结构特征是判断菌根形成的主要指标,与其功能密切相关.本文总结了AM丛枝结构、泡囊结构、菌丝结构和侵入点结构等发育特征;分析了A型丛枝结构、P型丛枝结构、泡囊结构和根外菌丝结构与促进寄主植物养分吸收和生长、提高植物抗旱性、耐涝性、耐盐性、抗高温、拮抗病原物、提高植物抗病性、抗重金属毒性、分解有毒有机物、修复污染与退化土壤等功能的关系,及其所发挥的重要作用;探讨了影响AM结构与功能的因子,以及基于AM不同结构所发挥功能的作用机制.旨在为系统研究AM真菌发育特征、AM真菌效能机制,以及评价和筛选AM真菌高效菌种提供依据.     

Fungal genes related to calcium homeostasis and signalling are upregulated in symbiotic arbuscular mycorrhiza interactions

Fluctuations in intracellular calcium levels generate signalling events and regulate different cellular processes. Whilst the implication of Ca²⁺ in plant responses during arbuscular mycorrhiza (AM) interactions is well documented, nothing is known about the regulation or role of this secondary messenger in the fungal symbiont. The spatio-temporal expression pattern of putatively Ca²⁺-related genes of Glomus intraradices BEG141 encoding five proteins involved in membrane transport and one nuclear protein kinase, was investigated during the AM symbiosis. Expression profiles related to successful colonization of host roots were observed in interactions of G. intraradices with roots of wild-type Medicago truncatula (line J5) compared to the mycorrhiza-defective mutant dmi3/Mtsym13. Symbiotic fungal activity was monitored using stearoyl-CoA desaturase and phosphate transporter genes. Laser microdissection based-mapping of fungal gene expression in mycorrhizal root tissues indicated that the Ca²⁺-related genes were differentially upregulated in arbuscules and/or in intercellular hyphae. The spatio-temporal variations in gene expression suggest that the encoded proteins may have different functions in fungal development or function during symbiosis development. Full-length cDNA obtained for two genes with interesting expression profiles confirmed a close similarity with an endoplasmic reticulum P-type ATPase and a Vcx1-like vacuolar Ca²⁺ ion transporter functionally characterized in other fungi and involved in the regulation of cell calcium pools. Possible mechanisms are discussed in which Ca²⁺-related proteins G. intraradices BEG141 may play a role in mobilization and perception of the intracellular messenger by the AM fungus during symbiotic interactions with host roots.     

环境污染物对丛枝菌根(AM)形成及功能的影响

丛枝菌根(AM)具有植物和微生物的双重特性,在污染土壤修复中受到越来越多的重视.AM在修复污染土壤的同时,也深受污染物毒害的影响,从而降低AM在污染土壤修复中的作用.如何减少环境污染物对AM的不利影响,是AM应用中需要考虑的问题.从有机和无机污染物角度, 综述了不同污染物对AM形成及功能的影响,并分析了可能的影响机理.大量研究表明,无论是有机污染物还是无机污染物,都会对AM的结构、形成和功能产生破坏性影响,主要表现在孢子萌发、侵染率、菌丝伸长受抑制等.有机污染物可能通过影响光合产物向AM真菌的分配, 间接影响AM真菌的活性,而重金属则通过抑制AM真菌活性, 直接对其产生影响.     

Microbial ecology of the arbuscular mycorrhiza

Arbuscular mycorrhizal (AM) fungi interact with a wide variety of organisms during all stages of their life. Some of these interactions such as grazing of the external mycelium are detrimental, while others including interactions with plant growth promoting rhizobacteria (PG PR) promote mycorrhizal functioning. Following mycorrhizal colonisation the functions of the root become modified, with consequences for the rhizosphere community which is extended into the mycorrhizosphere due to the presence of the AM external mycelium. However, we still know relatively little of the ecology of AM fungi and, in particular, the mycelium network under natural conditions. This area merits attention in the future with emphasis on the fungal partner in the association rather than the plant which has been the focus in the past.     

Temperature‐dependence of minimum resource requirements alters competitive hierarchies in phytoplankton

Resource competition theory is a conceptual framework that provides mechanistic insights into competition and community assembly of species with different resource requirements. However, there has been little exploration of how resource requirements depend on other environmental factors, including temperature. Changes in resource requirements as influenced by environmental temperature would imply that climate warming can alter the outcomes of competition and community assembly. We experimentally demonstrate that environmental temperature alters the minimum light and nitrogen requirements – as well as other growth parameters – of six widespread phytoplankton species from distinct taxonomic groups. We found that species require the most nitrogen at the highest temperatures while light requirements tend to be lowest at intermediate temperatures, although there are substantial interspecific differences in the exact shape of this relationship. We also experimentally parameterize two competition models, which we use to illustrate how temperature, through its effects on species’ traits, alters competitive hierarchies in multispecies assemblages, determining community dynamics. Developing a mechanistic understanding of how temperature influences the ability to compete for limiting resources is a critical step towards improving forecasts of community dynamics under climate warming.     

Soil-mediated local adaptation alters seedling survival and performance

Background and aims

Soils can act as agents of natural selection, causing differential fitness among genotypes and/or families of the same plant species, especially when soils have extreme physical or chemical properties. More subtle changes in soils, such as variation in microbial communities, may also act as agents of selection. We hypothesized that variation in soil properties within a single river drainage can be a selective gradient, driving local adaptation in plants.

Methods

Using seeds collected from individual genotypes of Populus angustifolia James and soils collected from underneath the same trees, we use a reciprocal transplant design to test whether seedlings would be locally adapted to their parental soil type.

Results

We found three patterns: 1. Soils from beneath individual genotypes varied in pH, soil texture, nutrient content, microbial biomass and the physiological status of microorganisms. 2. Seedlings grown in local soils experienced 2.5-fold greater survival than seedlings planted in non-local soils. 3. Using a composite of height, number of leaves and leaf area to measure plant growth, seedlings grew ～17.5% larger in their local soil than in non-local soil.

Conclusions

These data support the hypothesis that variation in soils across subtle gradients can act as an important selective agent, causing differential fitness and local adaptation in plants.     

Photon-harvesting efficiency and arbuscular mycorrhiza in amphibious plants

The amphibious plant species of intermittent aquatic habitats thrive both submerged and emerged. In order to outline the adaptive characters of these two life forms photochemical efficiency of photosystem 2, leaf contents of chlorophyll (Chl) a and b, carotenoids (Car), anthocyanins (Ant), and UV-B absorbing compounds (UV-B abs), and root aerenchyma and arbuscular mycorrhizal (AM) colonisation were studied in Glyceria fluitans, Gratiola officinalis, Ranunculus lingua, Teucrium scordium, Sium latifolium, Sparganium emersum, and Veronica anagallis-aquatica. Water level fluctuations did not exert a severe effect on photon harvesting efficiency. Submerged specimens had higher contents of Car and Ant whereas higher contents of UV-B abs were found in emerged specimens indicating efficient protection against the harmful effects of solar radiation. Roots of all species studied had extensive aerenchyma and were colonised by AM fungi, which were significantly more abundant in emerged specimens. This is the first report on AM symbiosis in S. latifolium and S. emersum.     

植物激素调控丛枝菌根发育的作用机制研究进展

丛枝菌根（arbuscular mycorrhiza,AM）是土壤中AM真菌和绝大多数维管植物根系长期进化过程中相互识别、相互作用形成的互利共生体。AM的发育与功能效应依赖AM真菌-寄主植物之间精准的“分子对话”,同时受到环境条件特别是土壤养分水平、干旱和盐渍化的制约。植物激素作为低浓度的小分子有机物,是参与调控AM共生过程的重要信号分子。其中,主要有9种植物激素参与AM发育过程且分工各有不同：独脚金内酯（strigolactones,SLs）参与AM真菌-寄主植物之间最初的共生识别,脱落酸（abscisic acid,ABA）和油菜素内酯（brassinosteroid,BR）促进前期的菌丝入侵,但水杨酸（salicylic acid,SA）和乙烯（ethylene,ET）抑制前期的菌丝入侵,生长素（auxin,Aux）、ABA和BR促进随后的丛枝形成而ET和赤霉素（gibberellin,GA）的作用则相反,茉莉酸（jasmonic acid,JA）对菌丝入侵与丛枝形成均可能存在正调控或负调控作用。目前细胞分裂素（cytokinin,CTK）在AM发育中的作用尚不明确。更为复杂的是,通常植物激素信号之间的交叉互作决定AM的发育进程。本文针对AM发育过程总结了不同植物激素的调控作用特点和不同植物激素信号之间的互作（协同或拮抗）,以及胁迫条件下不同植物激素信号的可能调控机制。深入研究和系统阐明植物激素调控AM真菌-寄主植物共生的生理/分子机制,将有助于促进生物共生学理论研究及菌根技术的应用。     

Neighbouring weeds influence the formation of arbuscular mycorrhiza in grapevine

Grapevine (Vitis vinifera L.) and two selected weeds from Mediterranean Croatian vineyards (Plantago lanceolata L. and Tanacetum cinerariifolium (Trevir.) Sch.Bip.) were examined in pot culture experiments, individually or when combined, to see whether multiple hosts influenced the formation of the symbiosis with arbuscular mycorrhizal fungi (AMF). The results after six-month period showed that plant identity and density significantly influenced development of mycorrhizal intra- and extraradical mycelium and/or sporulation. Grapevine and T. cinerariifolium individually and in combination resulted in a greater development of arbuscular mycorrhizae in terms of spore production, extraradical mycelium length and root colonization compared with pots containing P. lanceolata. Herbaceous weed species seemed to promote a different set of dominant AMF, potentially providing a wider spectrum of AMF for colonising grapevine roots. This indicates the value of encouraging host plant diversity in vineyards. AMF sequences obtained in this study are the first data reported for soils in Croatia.     

丛枝菌根共生建成的信号识别机制

丛枝菌根(Arbuscular mycorrhiza,AM)共生是自然界中普遍存在的一种互惠共生现象,对促进土壤生态系统物质循环及维持生态系统稳定具有重要的意义。AM共生体的建立需要AM真菌和宿主植物间一系列复杂的信号识别、交换和传导。本文总结近年来相关文献,从AM共生体形成前期及AM共生体形成期两个阶段,分别综述了信号物质的生物合成过程、调控过程及其作用机制,希望有助于进一步认识AM共生体建成过程,同时通过分析当前研究工作的不足及未来研究动向,期望推动相关研究工作。     

Cadmium accumulation in sunflower plants influenced by arbuscular mycorrhiza

In order to investigate the cadmium (Cd) accumulation patterns and possible alleviation of Cd stress by mycorrhization, sunflower plants (Helianthus annuus L.) were grown in the presence or absence of Cd (20 micromol L(-1)) and inoculated or not inoculated with the arbuscular mycorrhizal fungus (AMF) Glomus intraradices. No visual symptoms of Cd phytotoxicity were observed; nevertheless, in non-mycorrhizal plants the presence of Cd decreased plant growth. The addition of Cd had no significant effect on either mycorrhizal colonization or the amount of extra-radical mycelia that was produced by the AMF. Cd accumulated mainly in roots; only 22% of the total Cd absorbed was translocated to the shoots, where it accumulated to an average of 228 mg Cd kg(-1). Although the shoot-to-root ratio of Cd was similar in both the AMF inoculated and non-inoculated plants, the total absorbed Cd was 23% higher in mycorrhizal plants. Cd concentration in AMF extra-radical mycelium was 728 microg g(-1) dry weight. Despite the greater absorption of Cd, mycorrhizal plants showed higher photosynthetic pigment concentrations and shoot P contents. Cd also influenced mineral nutrition, leading to decreased Ca and Cu shoot concentrations; N, Fe and Cu shoot contents; and increased S and K shoot concentrations. Cd induced guaiacol peroxidase activity in roots in both mycorrhizal and non-mycorrhizal plants, but this increase was much more accentuated in non-mycorrhizal roots. In conclusion, sunflower plants associated with G. intraradices were less sensitive to Cd stress than non-mycorrhizal plants. Mycorrhizal sunflowers showed enhanced Cd accumulation and some tolerance to excessive Cd concentrations in plant tissues.     

Fast moves in arbuscular mycorrhizal symbiotic signalling

Exciting research looking at early events in arbuscular mycorrhizal symbioses has shown how the fungus and plant get together. Kohki Akiyama et al. have demonstrated that strigolactones in root exudates are fungal germ tube branching factors, and Arnaud Besserer et al. found that these compounds rapidly induce fungal mitochondrial activity. Andrea Genre et al. have shown that subsequent development of appressoria on host roots induces construction of a transient prepenetration apparatus inside epidermal cells that is reminiscent of nodulation infection.     

Genetic evidence for auxin involvement in arbuscular mycorrhiza initiation

? Formation of arbuscular mycorrhiza (AM) is controlled by a host of small, diffusible signaling molecules, including phytohormones. To test the hypothesis that the plant hormone auxin controls mycorrhiza development, we assessed mycorrhiza formation in two mutants of tomato (Solanum lycopersicum): diageotropica (dgt), an auxin-resistant mutant, and polycotyledon (pct), a mutant with hyperactive polar auxin transport. ? Mutant and wild-type (WT) roots were inoculated with spores of the AM fungus Glomus intraradices. Presymbiotic root-fungus interactions were observed in root organ culture (ROC) and internal fungal colonization was quantified both in ROC and in intact seedlings. ? In ROC, G. intraradices stimulated presymbiotic root branching in pct but not in dgt roots. pct roots stimulated production of hyphal fans indicative of appressorium formation and were colonized more rapidly than WT roots. By contrast, approaching hyphae reversed direction to grow away from cultured dgt roots and failed to colonize them. In intact seedlings, pct and dgt roots were colonized poorly, but development of hyphae, arbuscules, and vesicles was morphologically normal within roots of both mutants. ? We conclude that auxin signaling within host roots is required for the early stages of AM formation, including during presymbiotic signal exchange.     

The incidence of arbuscular mycorrhiza in two submerged Isoëtes species

We investigated the occurrence of arbuscular mycorrhizal fungi in the roots of Isoëtes lacustris and I. echinospora. These submerged lycopsids are the only macrophyte species inhabiting the bottom of two acidified glacial lakes in the Czech Republic. Arbuscular mycorrhizal (AM) fungi were detected in the roots of both species but the percentage of root colonization was both low and variable. Nevertheless, planting Littorella uniflora in the sediments from Isoëtes rhizosphere revealed high levels of viable AM propagules in both lakes. While AM colonization of Isoëtes roots did not exceed 25%, the average colonization of Littorella roots amounted to more than 80%. Although colonization of quillwort roots by AM fungi is evident, the taxonomic identity and role of these AM fungi in plant growth remain unclear. In addition to AM fungi, root-colonizing dark septate endophytic fungi were observed in both Isoëtes species.