Development of the arbuscular mycorrhizal symbiosis

The arbuscular mycorrhizal (AM) symbiosis formed between plant roots and fungi is one of the most widespread symbiotic associations found in plants, yet our understanding of events underlying its development are limited. The recent integration of biochemical, molecular and genetic approaches into analyses of the symbiosis is providing new insights into various aspects of its development. In the past year there have been advances in our understanding of the signals required for the formation of appressoria, the molecular changes in the root in response to colonisation, and components of the signal transduction pathways common to both the AM and Rhizobium symbioses.     

丛枝菌根真菌-豆科植物-根瘤菌共生体系的研究进展

丛枝菌根真菌（Arbuscular Mycorrhizal Fungi,AMF）-豆科植物-根瘤菌（Rhizobia）三者形成的共生体。是植物与微生物共生中的一种特殊类型。本文对这种共生体中微生物与植物之间的营养关系;AMF和根瘤菌双接种豆科植物的效应以及影响双接种效应的因素;AMF和根瘤菌在与豆科植物形成共生过程中的分子互作机制等进行了综述。同时对这种共生体还需进一步研究的问题及其在基础研究和实践应用方面的前景进行了讨论。     

Functional complementarity in the arbuscular mycorrhizal symbiosis

The causes and consequences of biodiversity are central themes in ecology. Perhaps one reason for much of the current interest in biodiversity is the belief that the loss of species (by extinction) or their gain (by invasion) will significantly influence ecosystem function. Arbuscular mycorrhizal (AM) fungi are components of most terrestrial ecosystems and, while many research programs have shown that variability among species or isolates of AM fungi does occur (Giovannetti & Gianinazzi-Pearson, 1994), the basis for this variability and its consequences to the function of communities and ecosystems remains largely unexplored. Smith et al . (pp. 357–366 in this issue) now show clearly that ecologically significant functional diversity exists among AM fungal species in the regions of the soil from which they absorb phosphate, and their results suggest that such diversity may have significant ecological consequences.     

Molecular genetics of the arbuscular mycorrhizal symbiosis

During arbuscular mycorrhiza (AM) development, fungal hyphae grow throughout root epidermal, exodermal and cortical cell layers to reach the inner cortex where the symbiosis' functional units, the arbuscles, develop. Three essential components of a plant signalling network, a receptor-like kinase, a predicted ion-channel and a calmodulin-dependent protein kinase have been identified. A detailed morphological study of symbiotic plant mutants revealed that different subsets of plant genes support the progress of fungal infection in successive root cell layers. Moreover, evidence of a diffusible fungal signalling factor that triggers gene activation in the root has recently been obtained.     

Plant hormones as signals in arbuscular mycorrhizal symbiosis

Abstract

Arbuscular mycorrhizal (AM) fungi are non-specific symbionts developing mutual and beneficial symbiosis with most terrestrial plants. Because of the obligatory nature of the symbiosis, the presence of the host plant during the onset and proceeding of symbiosis is necessary. However, AM fungal spores are able to germinate in the absence of the host plant. The fungi detect the presence of the host plant through some signal communications. Among the signal molecules, which can affect mycorrhizal symbiosis are plant hormones, which may positively or adversely affect the symbiosis. In this review article, some of the most recent findings regarding the signaling effects of plant hormones, on mycorrhizal fungal symbiosis are reviewed. This may be useful for the production of plants, which are more responsive to mycorrhizal symbiosis under stress.     

Alleviation of cadmium stress by arbuscular mycorrhizal symbiosis

Abstract

Owing to the realization of the harmful effect of cadmium on the environment and plants and as the plants are sessile organisms, they need to increase the protective mechanisms to cope with Cd stress. Inoculation the plant with soil microbes at the place of their growing is an important strategy to support the plants against stresses. In this study, trigonella plants were inoculated with arbuscular mycorrhizal (AM) fungi under different CdCl₂ concentrations (0, 2.25, and 6.25?mM). AM inoculation increased growth parameters, chlorophyll, and protein contents. Root colonization was significantly increased at low Cd concentration (2.25?mM) and decreased at high one (6.25?mM). Also, with AM fungal inoculation, the translocation factor of trigonella plants significantly decreased as compared to non-AM ones at both low and high Cd concentrations. In addition, it was clearly that malondialdehyde content of trigonella plants increased significantly at both Cd concentrations and with AM fungal inoculation its content decreased compared to those of non-AM ones. AM inoculation significantly increased antioxidant enzymes activities compared to non-AM ones. Consequently, this study showed a tolerance strategy of AM trigonella plants against Cd stress, thus mycorrhizal symbiosis becomes a promising and suitable as phytostabilizers of Cd stressed soil.     

The importance of integration and scale in the arbuscular mycorrhizal symbiosis

The arbuscular mycorrhizal (AM) fungus contributes to system processes and functions at various hierarchical organizational levels, through their establishment of linkages and feedbacks between whole-plants and nutrient cycles. Even though these fungal mediated feedbacks and linkages involve lower-organizational level processes (e.g. photo-assimilate partitioning, interfacial assimilate uptake and transport mechanisms, intraradical versus extraradical fungal growth), they influence higher-organizational scales that affect community and ecosystem behavior (e.g. whole-plant photosynthesis, biodiversity, nutrient and carbon cycling, soil structure). Hence, incorporating AM fungi into research directed at understanding many of the diverse environmental issues confronting society will require knowledge of how these fungi respond to or initiate changes in vegetation dynamics, soil fertility or both. Within the last few years, the rapid advancement in the development of analytical tools has increased the resolution by which we are able to quantify the mycorrhizal symbiosis. It is important that these tools are applied within a conceptual framework that is temporally and spatially relevant to fungus and host. Unfortunately, many of the studies being conducted on the mycorrhizal symbiosis at lower organizational scales are concerned with questions directed solely at understanding fungus or host without awareness of what the plant physiologist or ecologist needs for integrating the mycorrhizal association into larger organizational scales or process levels. We show by using the flow of C from plant-to-fungus-to-soil, that through thoughtful integration, we have the ability to bridge different organizational scales. Thus, an essential need of mycorrhizal research is not only to better integrate the various disciplines of mycorrhizal research, but also to identify those relevant links and scales needing further investigation for understanding the larger-organizational level responses. The U.S. Government's right to retain a non-exclusive, royalty-free licence in and to any copyright is acknowledged. This revised version was published online in June 2006 with corrections to the Cover Date.     

Programming good relations--development of the arbuscular mycorrhizal symbiosis

The majority of plants live in symbiotic associations with fungi or bacteria that improve their nutrition. Critical steps in a symbiosis are mutual recognition and subsequently the establishment of an intimate association, which involves the penetration of plant tissues and, in many cases, the invasion of individual host cells by the microbial symbiont. Recent advances revealed that in the arbuscular mycorrhizal symbiosis with soil fungi of the order Glomeromycota, plant-derived signals attract fungal hyphae and stimulate their growth. Upon physical attachment of the fungal symbiont to the root surface, an active plant developmental program prepares the epidermal cells for penetration by the fungus. Thus, plants actively help symbiotic fungi to colonize their roots rather than just tolerating them.     

Molecular and cell biology of arbuscular mycorrhizal symbiosis

The roots of most extant plants are able to become engaged in an interaction with a small group of fungi of the fungal order Glomales (Glomeromycota). This interaction—arbuscular mycorrhizal (AM) symbiosis—is the evolutionary precursor of most other mutualistic root-microbe associations. The molecular analysis of this interaction can elucidate basic principles regarding such associations. This review summarizes our present knowledge about cellular and molecular aspects of AM. Emphasis is placed on morphological changes in colonized cells, transfer of nutrients between both interacting partners, and plant defence responses. Similarities to and differences from other associations of plant and microorganisms are highlighted regarding defence reactions and signal perception.     

Signaling events during initiation of arbuscular mycorrhizal symbiosis

Under nutrient‐limiting conditions, plants will enter into symbiosis with arbuscular mycorrhizal(AM) fungi for the enhancement of mineral nutrient acquisition from the surrounding soil. AM fungi live in close, intracellular association with plant roots where they transfer phosphate and nitrogen to the plant in exchange for carbon. They are obligate fungi,relying on their host as their only carbon source. Much has been discovered in the last decade concerning the signaling events during initiation of the AM symbiosis, including the identification of signaling molecules generated by both partners. This signaling occurs through symbiosis‐specific gene products in the host plant, which are indispensable for normal AM development. At the same time, plants have adapted complex mechanisms for avoiding infection by pathogenic fungi, including an innate immune response to general microbial molecules, such as chitin present in fungal cell walls. How it is that AM fungal colonization is maintained without eliciting a defensive response from the host is still uncertain. In this review, we present a summary of the molecular signals and their elicited responses during initiation of the AM symbiosis, including plant immune responses and their suppression.     

Medicago truncatula Vapyrin is a novel protein required for arbuscular mycorrhizal symbiosis

Arbuscular mycorrhizal (AM) symbiosis is a widespread mutualism formed between vascular plants and fungi of the Glomeromycota. In this endosymbiosis, fungal hyphae enter the roots, growing through epidermal cells to the cortex where they establish differentiated hyphae called arbuscules in the cortical cells. Reprogramming of the plant epidermal and cortical cells occurs to enable intracellular growth of the fungal symbiont; however, the plant genes underlying this process are largely unknown. Here, through the use of RNAi, we demonstrate that the expression of a Medicago truncatula gene named Vapyrin is essential for arbuscule formation, and also for efficient epidermal penetration by AM fungi. Vapyrin is induced transiently in the epidermis coincident with hyphal penetration, and then in the cortex during arbuscule formation. The Vapyrin protein is cytoplasmic, and in cells containing AM fungal hyphae, the protein accumulates in small puncta that move through the cytoplasm. Vapyrin is a novel protein composed of two domains that mediate protein–protein interactions: an N‐terminal VAMP‐associated protein (VAP)/major sperm protein (MSP) domain and a C‐terminal ankyrin‐repeat domain. Putative Vapyrin orthologs exist widely in the plant kingdom, but not in Arabidopsis, or in non‐plant species. The data suggest a role for Vapyrin in cellular remodeling to support the intracellular development of fungal hyphae during AM symbiosis.     

The arbuscular mycorrhizal symbiosis links N mineralization to plant demand

Arbuscular mycorrhizal (AM) fungi facilitate inorganic N (NH₄ ⁺ or NO₃ ⁻) uptake by plants, but their role in N mobilization from organic sources is unclear. We hypothesized that arbuscular mycorrhizae enhance the ability of a plant to use organic residues (ORs) as a source of N. This was tested under controlled glasshouse conditions by burying a patch of OR in soil separated by 20-μm nylon mesh so that only fungal hyphae can pass through it. The fate of the N contained in the OR patch, as influenced by Glomus claroideum, Glomus clarum, or Glomus intraradices over 24 weeks, was determined using ¹⁵N as a tracer. AM fungal species enhanced N mineralization from OR to different levels. N recovery and translocation to Russian wild rye by hyphae reached 25% of mineralized N in G. clarum, which was most effective despite its smaller extraradical development in soil. Mobilization of N by G. clarum relieved plant N deficiency and enhanced plant growth. We show that AM hyphae modify soil functioning by linking plant growth to N mineralization from OR. AM species enhance N mineralization differentially leading to species-specific changes in the quality of the soil environment (soil C-to-N ratio) and structure of the soil microbial community.     

Fungal and plant gene expression in arbuscular mycorrhizal symbiosis

Arbuscular mycorrhizas (AMs) are a unique example of symbiosis between two eukaryotes, soil fungi and plants. This association induces important physiological changes in each partner that lead to reciprocal benefits, mainly in nutrient supply. The symbiosis results from modifications in plant and fungal cell organization caused by specific changes in gene expression. Recently, much effort has gone into studying these gene expression patterns to identify a wider spectrum of genes involved. We aim in this review to describe AM symbiosis in terms of current knowledge on plant and fungal gene expression profiles.An erratum to this article can be found at     

Dating in the dark: how roots respond to fungal signals to establish arbuscular mycorrhizal symbiosis

The arbuscular mycorrhizal symbiosis that involves most plants and Glomeromycota fungi is the result of a complex exchange of molecular information, which commences before the partners are in physical contact. On the one hand, plants release soluble factors, including strigolactones that activate both the metabolism and branching of the fungal partners. On the other hand, fungi use compounds that trigger the signaling transduction pathways that are required for the symbiotic modus of plant cells. Here we describe some of the recent discoveries regarding the fungal molecules involved in rhizospheric conversation, and the way in which they are perceived by their hosts. We conclude that similar signaling molecules may have different meanings, depending on the context. However, at the end, specificity must be maintained to ensure appropriate partners enter symbiosis.     

Regulation of the plant defence response in arbuscular mycorrhizal symbiosis

The response of plants to arbuscular mycorrhizal fungi involves a temporal and spatial activation of different defence mechanisms. The activation and regulation of these defences have been proposed to play a role in the maintenance of the mutualistic status of the association, however, how these defences affect the functioning and development of arbuscular mycorrhiza remains unclear. A number of regulatory mechanisms of plant defence response have been described during the establishment of the arbuscular mycorrhizal symbiosis, including elicitor degradation, modulation of second messenger concentration, nutritional and hormonal plant defence regulation, and activation of regulatory symbiotic gene expression. The functional characterization of these regulatory mechanisms on arbuscular mycorrhiza, including cross-talk between them, will be the aim and objective of future work on this topic.     

Preference,specificity and cheating in the arbuscular mycorrhizal symbiosis

Arbuscular mycorrhizal symbioses are mutualistic interactions between fungi and most plants. There is considerable interest in this symbiosis because of the strong nutritional benefits conferred to plants and its influence on plant diversity. Until recently, the symbiosis was assumed to be unspecific. However, two studies have now revealed that although it can be largely unspecific with the fungal community composition changing seasonally, in certain ecosystems it can also be highly specific and might potentially allow plants to cheat the arbuscular mycorrhizal network that connects plants below ground.