Informationsaustausch in der Ektomykorrhiza

To see the wood for the trees: Communication in ectomycorrhizal symbiosis The mutual symbiosis of ectomycorrhiza has been established in a co‐evolution that depends on a specific communication between the woody plant and the fungus. The exchange of inorganic nutrients and water (delivered by the fungus) for sugar (supplied by the host tree) provides the basis for the symbiosis. The interaction is initiated with signals that can be associated with root exudates and volatiles in the soil matrix. After recognition, the fungus is able to modulate plant response functions that usually suppress pathogens by excretion of effector molecules, which allows entry into the root. Within the root, specific cell wall proteins of the fungus like hydrophobins are important for host specificity. Signals in the mycorrhizal root like the auxin indole‐acetic acid modify the morphology of both partners resulting in the intimate interactions of fully established mycorrhiza. The soil hyphae of the fungus, at the same time, respond to other bacteria and fungi in the mycorrhizosphere.     

Medicago truncatula mtpt4 mutants reveal a role for nitrogen in the regulation of arbuscule degeneration in arbuscular mycorrhizal symbiosis

Plants acquire essential mineral nutrients such as phosphorus (P) and nitrogen (N) directly from the soil, but the majority of the vascular plants also gain access to these mineral nutrients through endosymbiotic associations with arbuscular mycorrhizal (AM) fungi. In AM symbiosis, the fungi deliver P and N to the root through branched hyphae called arbuscules. Previously we identified MtPT4, a Medicago truncatula phosphate transporter located in the periarbuscular membrane that is essential for symbiotic phosphate transport and for maintenance of the symbiosis. In mtpt4 mutants arbuscule degeneration occurs prematurely and symbiosis fails. Here, we show that premature arbuscule degeneration occurs in mtpt4 mutants even when the fungus has access to carbon from a nurse plant. Thus, carbon limitation is unlikely to be the primary cause of fungal death. Surprisingly, premature arbuscule degeneration is suppressed if mtpt4 mutants are deprived of nitrogen. In mtpt4 mutants with a low N status, arbuscule lifespan does not differ from that of the wild type, colonization of the mtpt4 root system occurs as in the wild type and the fungus completes its life cycle. Sulphur is another essential macronutrient delivered to the plant by the AM fungus; however, suppression of premature arbuscule degeneration does not occur in sulphur-deprived mtpt4 plants. The mtpt4 arbuscule phenotype is strongly correlated with shoot N levels. Analyses of an mtpt4-2 sunn-1 double mutant indicates that SUNN, required for N-mediated autoregulation of nodulation, is not involved. Together, the data reveal an unexpected role for N in the regulation of arbuscule lifespan in AM symbiosis.     

Tit for tat? A mycorrhizal fungus accumulates phosphorus under low plant carbon availability

The exchange of carbohydrates and mineral nutrients in the arbuscular mycorrhizal (AM) symbiosis must be controlled by both partners in order to sustain an evolutionarily stable mutualism. Plants downregulate their carbon (C) flow to the fungus when nutrient levels are sufficient, while the mechanism controlling fungal nutrient transfer is unknown. Here, we show that the fungus accumulates nutrients when connected to a host that is of less benefit to the fungus, indicating a potential of the fungus to control the transfer of nutrients. We used a monoxenic in vitro model of root organ cultures associated with Glomus intraradices, in which we manipulated the C availability to the plant. We found that G. intraradices accumulated up to seven times more nutrients in its spores, and up to nine times more in its hyphae, when the C pool available to the associated roots was halved. The strongest effect was found for phosphorus (P), considered to be the most important nutrient in the AM symbiosis. Other elements such as potassium and chorine were also accumulated, but to a lesser extent, while no accumulation of iron or manganese was found. Our results suggest a functional linkage between C and P exchange.     

Pathways of Phosphorus Absorption and Early Signaling between the Mycorrhizal Fungi and Plants

This review highlights the key role that mycorrhizal fungi play in making phosphorus (Pi) more available to plants, including pathways of phosphorus absorption, phosphate transporters and plant-mycorrhizal fungus symbiosis, especially in conditions where the level of inorganic phosphorus (Pi) in the soil is low. Mycorrhizal fungi colonization involves a series of signaling where the plant root exudates strigolactones, while the mycorrhizal fungi release a mixture of chito-oligosaccharides and liposaccharides, that activate the symbiosis process through gene signaling pathways, and contact between the hyphae and the root. Once the symbiosis is established, the extraradical mycelium acts as an extension of the roots and increases the absorption of nutrients, particularly phosphorus by the phosphate transporters. Pi then moves along the hyphae to the plant root/fungus interface. The transfer of Pi occurs in the apoplectic space; in the case of arbuscular mycorrhizal fungi, Pi is discharged from the arbuscular to the plant’s root symplasm, in the membrane that surrounds the arbuscule. Pi is then absorbed through the plant periarbuscular membrane by plant phosphate transporters. Furthermore, plants can acquire Pi from soil as a direct absorption pathway. As a result of this review, several genes that codify for high-affinity Pi transporters were identified. In plants, the main family is Pht1 although it is possible to find others such as Pht2, Pht3, Pho1 and Pho2. As in plants, mycorrhizal fungi have genes belonging to the Pht1 subfamily. In arbuscular mycorrhizal fungi we found L1PT1, GiPT, MtPT1, MtPT2, MtPT4, HvPT8, ZmPht1, TaPTH1.2, GmosPT and LYCes. HcPT1, HcPT2 and BePT have been characterized in ectomycorrhizal fungi. Each gene has a different way of expressing itself. In this review, we present diagrams of the symbiotic relationship between mycorrhizal fungi and the plant. This knowledge allows us to design solutions to regional problems such as food production in soils with low levels of Pi.

     

Localization and speciation of arsenic in Glomus intraradices by synchrotron radiation spectroscopic analysis

The protective mechanisms employed by arbuscular mycorrhizal fungi (AMF) to reduce the toxic effects of arsenic on host plants remain partially unknown. The goal of this research was identifying the in situ localization and speciation of arsenic (As) in the AM fungus Rhizophagus intraradices [formerly named Glomus intraradices] exposed to arsenate [As(V)]. By using a two-compartment in vitro fungal cultures of R. intraradices-transformed carrot roots, microspectroscopic X-ray fluorescence (μ-XRF), and microspectroscopic X-ray absorption near edge structure (μ-XANES), we observed that As(V) is absorbed after 1 h in the hyphae of AMF. Three hours after exposure a decrease in the concentration of As was noticed and after 24 and 72 h no detectable As concentrations were perceived suggesting that As taken up was pumped out from the hyphae. No As was detected within the roots or hyphae in the root compartment zone three or 45 h after exposure. This suggests a dual protective mechanism to the plant by rapidly excluding As from the fungus and preventing As translocation to the plant root. μ-XANES data showed that gradual As(V) reduction occurred in the AM hyphae between 1 and 3 h after arsenic exposure and was completed after 6 h. Principal component analysis (PCA) and linear combination fitting (LCF) of μ-XANES data showed that the dominant species after reduction of As(V) by R. intraradices extra-radical hyphal was As(III) complexed with a reduced iron(II) carbonate compound. The second most abundant As species present was As(V)–iron hydroxides. The remaining As(III) compounds identified by the LCF analyses suggested these molecules were made of reduced As and S. These results increase our knowledge on the mechanism of As transport in AMF and validate our hypotheses that R. intraradices directly participates in arsenic detoxification. These fungal mechanisms may help AMF colonized plants to increase their tolerance to As at contaminated sites.     

At the Root of the Wood Wide Web: Self Recognition and Non-Self Incompatibility in Mycorrhizal Networks

Arbuscular mycorrhizal (AM) fungi are mutualistic symbionts living in the roots of 80% of land plant species, and developing extensive, below-ground extraradical hyphae fundamental for the uptake of soil nutrients and their transfer to host plants. Since AM fungi have a wide host range, they are able to colonize and interconnect contiguous plants by means of hyphae extending from one root system to another. Such hyphae may fuse due to the widespread occurrence of anastomoses, whose formation depends on a highly regulated mechanism of self recognition. Here, we examine evidences of self recognition and non-self incompatibility in hyphal networks formed by AM fungi and discuss recent results showing that the root systems of plants belonging to different species, genera and families may be connected by means of anastomosis formation between extraradical mycorrhizal networks, which can create indefinitely large numbers of belowground fungal linkages within plant communities.Key Words: arbuscular mycorrhizal symbiosis, extraradical mycelium, anastomosis, plant interconnectedness, self recognition, non-self incompatibility, mycorrhizal networks     

Anatomy and mycotrophy of the achlorophyllous Afrothismia winkleri

Afrothismia winkleri develops fleshy rhizomes, densely covered with small root tubercles, narrowing to filiform roots with age. The exclusively intracellular mycorrhizal fungus has distinct morphologies in different tissues of the plant. In the filiform root the hyphae grow straight and vesicles are borne on short hyphal stalks. The straight hyphae are present in the epidermis of the root tubercles, but change to loosely coiled and swollen hyphae in the rhizome tissue. No penetration from epidermis to root cortex was found. From the rhizome, a separating cell layer permits only one or rarely two hyphal penetrations into the cortex of each root tubercle. The hyphae proceed apically within the root hypodermis in a spiral row of distinctively coiled hyphae, branches of which colonize the inner root cortex. In the inner root cortex the hyphal coils degenerate to amorphous clumps. In older roots the cortex itself also deteriorates, but epidermis, hypodermis, endodermis and central cylinder persist. The mycorrhizal pattern in A. winkleri is interpreted as an elaborate exploitation system whereby the fungus provides carbon and nutrients to the plant and, simultaneously but spatially distinct, its hyphae are used to translocate and store the matter within the plant. Several features indicate that the endophyte is an arbuscular mycorrhizal fungus.     

Hyphosphere interactions between Rhizophagus irregularis and Rahnella aquatilis promote carbon–phosphorus exchange at the peri-arbuscular space in Medicago truncatula

Arbuscular mycorrhizal (AM) fungi form a continuum between roots and soil. One end of this continuum is comprised of the highly intimate plant–fungus interface with intracellular organelles for nutrient exchange, while on the other end the fungus interacts with bacteria to compensate for the AM fungus' inability to take up organic nutrients from soil. How both interfaces communicate in this highly complex tripartite mutualism is widely unknown. Here, the effects of phosphate-solubilizing bacteria (PSB) Rahnella aquatilis dwelling at the surface of the extraradical hyphae of Rhizophagus irregularis was analysed based on the expression of genes involved in C-P exchange at the peri-arbuscular space (PAS) in Medicago truncatula. The interaction between AM fungus and PSB resulted in an increase in uptake and transport of Pi along the extraradical hyphae and its transfer from AM fungus to plant. In return, this was remunerated by a transfer of C from plant to AM fungus, improving the C-P exchange at the PAS. These results demonstrated that a microorganism (i.e., a PSB) developing at the hyphosphere interface can affect the C-P exchange at the PAS between plant and AM fungus, suggesting a fine-tuned communication operated between three organisms via two distantly connected interfaces.     

Transformation of Gaeumannomyces graminis with β‐Glucuronidase Reporter and Hygromycin Resistance Genes

Take‐all disease is caused by Gaeumannomyces graminis, (Sacc.) Arx & D. Olivier, a soil‐borne fungus, which colonizes the root and crown tissue of many members of the Poaceae plant family. This fungus is able to grow along the surface of roots as darkly pigmented runner hyphae, which has the ability to penetrate the root. Here, we describe a genetic transformation of G. graminis var. graminis by using polyethylene glycol (PEG)‐based protoplast transformation. Fungus cells were transformed with a plasmid, pHPG, containing the gusA reporter gene that codes for β‐glucuronidase (GUS) and the hph gene for hygromycin resistance as the selectable marker. A de novo transformant selection assay was developed to identify the putative transformants that were expressing the hph gene. In addition, the transformed cells maintained the ability to infect the plant tissues. The GUS‐expressing fungus can be used to study fungal infection processes including fungal penetration, colonization and the role(s) of melanin during pathogenesis. Thus, this study is the first report of G. graminis var. graminis transformed with a visibly detectable reporter gene that provides a useful tool to a better understanding of host–Gaeumannomyces interactions.     

Influence of arbuscular mycorrhiza on organic solutes in maize leaves under salt stress

A pot experiment was conducted to examine the effect of the arbuscular mycorrhizal (AM) fungus, Glomus mosseae, on plant biomass and organic solute accumulation in maize leaves. Maize plants were grown in sand and soil mixture with three NaCl levels (0, 0.5, and 1.0 g kg⁻¹ dry substrate) for 55 days, after 15 days of establishment under non-saline conditions. At all salinity levels, mycorrhizal plants had higher biomass and higher accumulation of organic solutes in leaves, which were dominated by soluble sugars, reducing sugars, soluble protein, and organic acids in both mycorrhizal and non-mycorrhizal plants. The relative abundance of free amino acids and proline in total organic solutes was lower in mycorrhizal than in non-mycorrhizal plants, while that of reducing sugars was higher. In addition, the AM symbiosis raised the concentrations of soluble sugars, reducing sugars, soluble protein, total organic acids, oxalic acid, fumaric acid, acetic acid, malic acid, and citric acid and decreased the concentrations of total free amino acids, proline, formic acid, and succinic acid in maize leaves. In mycorrhizal plants, the dominant organic acid was oxalic acid, while in non-mycorrhizal plants, the dominant organic acid was succinic acid. All the results presented here indicate that the accumulation of organic solutes in leaves is a specific physiological response of maize plants to the AM symbiosis, which could mitigate the negative impact of soil salinity on plant productivity.     

The promoting role of an isolate of dark-septate fungus on its host plant Saussurea involucrata Kar. et Kir.

A dark-septate endophytic (DSE) fungus EF-37 was isolated from the roots of Saussurea involucrata Kar. et Kir., an endangered Chinese medicinal plant. The molecular identification of the fungus was based on internal transcribed spacer regions and the result showed that EF-37 was congeneric to Mycocentrospora. This study was conducted to clarify the influence of the root endophyte EF-37 on the host plant S. involucrata using material grown in a sterile culture bottle. After cultivation for 40 days, fungal hyphae were found to be branching repeatedly and forming “hyphae nets” in the epidermal layers. Significant differences were detected between the study groups in plant dry weight, plant height, root dry weight, shoot dry weight, and the number of hair root tips. There was a positive effect of endophyte EF-37 on plant root development, with results showing that cortical cells dissolved and formed aerate structures. There was a positive effect of endophyte EF-37 on plant growth, but chlorophyll fluorescence analysis showed that there were no significant differences between the study groups. In addition, analysis of the chemical composition of seedlings showed that the level of rutin was higher in plants cultivated with the EF-37 fungus compared to the controls. This study helps to establish a basis for germplasm conservation and for further investigation of the interaction between dark-septate fungi and this alpine plant.     

Laser‐ablation electrospray ionization mass spectrometry with ion mobility separation reveals metabolites in the symbiotic interactions of soybean roots and rhizobia

Technologies enabling in situ metabolic profiling of living plant systems are invaluable for understanding physiological processes and could be used for rapid phenotypic screening (e.g., to produce plants with superior biological nitrogen‐fixing ability). The symbiotic interaction between legumes and nitrogen‐fixing soil bacteria results in a specialized plant organ (i.e., root nodule) where the exchange of nutrients between host and endosymbiont occurs. Laser‐ablation electrospray ionization mass spectrometry (LAESI‐MS) is a method that can be performed under ambient conditions requiring minimal sample preparation. Here, we employed LAESI‐MS to explore the well characterized symbiosis between soybean (Glycine max L. Merr.) and its compatible symbiont, Bradyrhizobium japonicum. The utilization of ion mobility separation (IMS) improved the molecular coverage, selectivity, and identification of the detected biomolecules. Specifically, incorporation of IMS resulted in an increase of 153 differentially abundant spectral features in the nodule samples. The data presented demonstrate the advantages of using LAESI–IMS–MS for the rapid analysis of intact root nodules, uninfected root segments, and free‐living rhizobia. Untargeted pathway analysis revealed several metabolic processes within the nodule (e.g., zeatin, riboflavin, and purine synthesis). Compounds specific to the uninfected root and bacteria were also detected. Lastly, we performed depth profiling of intact nodules to reveal the location of metabolites to the cortex and inside the infected region, and lateral profiling of sectioned nodules confirmed these molecular distributions. Our results established the feasibility of LAESI–IMS–MS for the analysis and spatial mapping of plant tissues, with its specific demonstration to improve our understanding of the soybean‐rhizobial symbiosis.     

Effects of prometryn and acetochlor on arbuscular mycorrhizal fungi and symbiotic system

Prometryn and acetochlor are common herbicides widely used to control weeds in agricultural systems. The impacts of the two herbicides on spore germination, hyphal elongation, the biomass and malondialdehyde content of carrot hairy roots were investigated using a strict in vitro cultivation system associating the Ri T‐DNA‐transferred carrot hairy roots with Glomus etunicatum. Alternatively, root colonization, daughter spore production and the proportion of hyphae with succinate dehydrogenase (SDH) and alkaline phosphatase (ALP) activities were also investigated. No significant impact on spore germination was noted in the presence of acetochlor at all three concentrations tested, while a significant decrease was observed with prometryn only at the highest concentration. Moreover, an inverse correlation was identified between herbicides concentrations and G. etunicatum root colonization and spore production as well as hyphal SDH and ALP activity, with a positive correlation identified among these four factors. Both herbicides exerted negative effects on the arbuscular mycorrhizal (AM) fungus and symbiosis at increasing concentrations, with prometryn apparently more toxic than acetochlor. Furthermore, the AM symbiotic system was shown to improve biomass, reduce malondialdehyde accumulation and ease lipid peroxidation in carrot hairy roots and decrease damage in host plants, thus enhancing plant tolerance to adverse conditions.

Significance and Impact of the Study

In this study, the effect of prometryn and acetochlor on the physiology and metabolic activities of the AM fungus Glomus etunicatum were investigated. Our findings demonstrate for the first time, the impact of the two herbicides at three concentrations (0·1, 1 and 10 mg l^?1) on transformed carrot hairy roots/AM fungus association under strict in vitro culture conditions, which may guide the application of the two herbicides in modern agriculture.     

High resolution characterization of ectomycorrhizal fungal-mineral interactions in axenic microcosm experiments

Microcosms with Pinus sylvestris seedlings in symbiosis with the fungus mycorrhizal Paxillus involutus were established, and atomic force microscopy (AFM) was used to characterise plant photosynthate-driven fungal interactions with mineral surfaces. Comparison of images of the same area of the minerals before and after mycorrhizal fungal colonization showed extensive growth of hyphae on three different mineral surfaces – hornblende, biotite and chlorite. A layer of biological exudate, or biolayer, covered the entire mineral surface and was composed of globular features of diameter 10–80 nm, and the morphology of the biolayer differed among mineral types. Similar-sized components were found on the fungal hyphae, but with a more elongated profile. Biolayer and hyphae surfaces both appeared to be hydrophobic with the hyphal surfaces yielding higher maximal adhesive interactions and a wider range of values: the mean (± SE) adhesive forces were 2.63 ± 0.03 and 3.46 ± 0.18 nN for biolayer and hypha, respectively. The highest adhesion forces are preferentially localized at the hyphal surface above the Spitzenkörper region and close to the tip, with a mean interaction force in this locality of 5.24 ± 0.49 nN. Biolayer thickness was between 10 and 40 nm. The underlying mineral was easily broken up by the tip, in contrast to the native mineral. These observations of mineral surfaces colonised by mycorrhizal fungus demonstrate how fungal hyphae are able to form a layer of organic exudates, or biolayer, and its role in hyphal attachment and potential weathering of ferromagnesian silicates, which may supply nutrients to the plant.     

Mechanisms of nutrient transport across interfaces in arbuscular mycorrhizas

Bidirectional nutrient transfer between the plant and the fungus is a key feature of arbuscular mycorrhizal symbiosis. The major nutrients exchanged between the symbiotic partners are reduced carbon, assimilated through the plant photosynthesis and phosphate, taken up by the fungal hyphae exploring soil microhabitats. This nutrient exchange takes place across the symbiotic interfaces which are bordered by the plant and fungal plasma membranes. This review provides an overview of the current knowledge of the mechanisms underlying nutrient transport processes in the symbiosis, with special emphasis on recent developments in the molecular biology of the plant and fungal primary (H⁺-ATPases) and secondary transporters.     

Carbon Uptake and the Metabolism and Transport of Lipids in an Arbuscular Mycorrhiza

Both the plant and the fungus benefit nutritionally in the arbuscular mycorrhizal symbiosis: The host plant enjoys enhanced mineral uptake and the fungus receives fixed carbon. In this exchange the uptake, metabolism, and translocation of carbon by the fungal partner are poorly understood. We therefore analyzed the fate of isotopically labeled substrates in an arbuscular mycorrhiza (in vitro cultures of Ri T-DNA-transformed carrot [Daucus carota] roots colonized by Glomus intraradices) using nuclear magnetic resonance spectroscopy. Labeling patterns observed in lipids and carbohydrates after substrates were supplied to the mycorrhizal roots or the extraradical mycelium indicated that: (a) ¹³C-labeled glucose and fructose (but not mannitol or succinate) are effectively taken up by the fungus within the root and are metabolized to yield labeled carbohydrates and lipids; (b) the extraradical mycelium does not use exogenous sugars for catabolism, storage, or transfer to the host; (c) the fungus converts sugars taken up in the root compartment into lipids that are then translocated to the extraradical mycelium (there being little or no lipid synthesis in the external mycelium); and (d) hexose in fungal tissue undergoes substantially higher fluxes through an oxidative pentose phosphate pathway than does hexose in the host plant.     

Growth-promotion of strawberry plants inoculated with Azospirillum brasilense

Azospirillum brasilense (strains REC3, RLC1, PEC5) were root inoculated in strawberry plants of the cultivars ‘Milsei’, ‘Selva’ and ‘Camarosa’ to assess plant growth-promoting effects. The bacteria were able to promote plant growth (expressed as root length, root area, and dry weight of root and shoot), depending on the genotypes of plants and bacteria used, whereas the stolon production (3–4) depended only on the strawberry cultivar. To explain whether root exudates plays any role on the growth-promotion observed herein, total protein and sugar were determined, and chemotaxis properties were evaluated. The strains showed positive chemotaxis toward the root exudates, being influenced by the total sugars content, suggesting that the latter plays an important role in the chemotaxis effect and may contribute to enhance the root capacity to recruit azospirilla from rhizosphere, thus improving the growth-promoting effect exerted by these bacteria.     

Sugar for my honey: carbohydrate partitioning in ectomycorrhizal symbiosis

Simple, readily utilizable carbohydrates, necessary for growth and maintenance of large numbers of microbes are rare in forest soils. Among other types of mutualistic interactions, the formation of ectomycorrhizas, a symbiosis between tree roots and certain soil fungi, is a way to overcome nutrient and carbohydrate limitations typical for many forest ecosystems. Ectomycorrhiza formation is typical for trees in boreal and temperate forests of the northern hemisphere and alpine regions world-wide. The main function of this symbiosis is the exchange of fungus-derived nutrients for plant-derived carbohydrates, enabling the colonization of mineral nutrient-poor environments. In ectomycorrhizal symbiosis up to 1/3 of plant photoassimilates could be transferred toward the fungal partner. The creation of such a strong sink is directly related to the efficiency of fungal hexose uptake at the plant/fungus interface, a modulated fungal carbohydrate metabolism in the ectomycorrhiza, and the export of carbohydrates towards soil growing hyphae. However, not only the fungus but also the plant partner increase its expression of hexose importer genes at the plant/fungus interface. This increase in hexose uptake capacity of plant roots in combination with an increase in photosynthesis may explain how the plant deals with the growing fungal carbohydrate demand in symbiosis and how it can restrict this loss of carbohydrates under certain conditions to avoid fungal parasitism.     

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.     

Reflections on the role of environmentally-governed reproductive versatility in the adaptation of plant populations

Summary

Mycorrhizal associations vary widely in structure and function, but the commonest interaction is the Arbuscular Mycorrhizal (AM) symbiosis which forms between the roots of over 80% of all terrestrial plant species and Zygomycete fungi of the Order Glomales. These are obligate symbionts which colonise plant root cells. This symbiosis confers benefits directly to the host plants through the acquisition of phosphate and other mineral nutrients from the soil by the fungus while the fungus receives a carbon source from the host. In addition, the symbiosis may also enhance the plants resistance to biotic and abiotic stresses. The beneficial effects of AM symbioses occur as a result of a complex molecular dialogue between the two symbiotic partners. Identifying the molecules and genes involved in the dialogue is necessary for a greater understanding of the symbiosis. This paper reviews the process of AM fungal colonisation of plant roots and the underlying molecular mechanisms associated with the formation and functioning of an AM symbiosis.