Recent advances in exploring physiology and biodiversity of ectomycorrhizas highlight the functioning of these symbioses in ecosystems

Ectomycorrhizas, the dominating mycorrhizal symbiosis in boreal, temperate and some tropical forests, are formed by 5000-6000 species of the asco- and basidiomycetes. This high diversity of fungal partners allows optimal foraging and mobilisation of various nitrogen and phosphorus forms from organic soil layers. In this review, two approaches to study the functioning of this multitude of symbiotic associations are presented. On selected culture models, physiological and molecular investigations have shown that the supply of hexoses has a key function in controlling the plant^fungus interaction via partner-specific regulation of gene expression. Environmental factors which affect fungal carbon supply, such as increased nitrogen availability, also affect mycorrhiza formation. Based on such laboratory results, the adaptative capability of ectomycorrhizas to changing field conditions is discussed. The second approach consists of analysing the distribution of mycorrhizas in ecosystem compartments and to relate distribution patterns to variations of ecological factors. Recent advances in identification of fungal partners in ectomycorrhizas by analysing the internal transcribed spacer of ribosomal DNA are presented, which can help to resolve sampling problems in field studies. The limits of the laboratory and the field approaches are discussed. Despite some problems, this combined approach is the most promising. Direct investigation of gene expression, which has been introduced for soil bacteria, will be difficult in the case of mycorrhizal fungi which constitute organisms with functionally varying structure

Análise de uma região de DNA de Araucaria angustifolia que pode estar envolvida com resistência ao frio

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Harnessing ectomycorrhizal genomics for ecological insights

International audienceThe ectomycorrhizal (ECM) symbiosis is increasingly seen as a crucial component for nutrient cycling in sustainable forest ecosystems. To date, the complete genome sequence of only a single symbiotic fungus, the ECM basidiomycete Laccaria bicolor, has been released. Its analysis revealed unexpected features, including numerous transposons, secreted effectorlike proteins and a lack of carbohydrate-hydrolysing enzymes acting on plant cell walls. Genome-wide transcript profilings showed that most symbiosis-induced transcripts code for proteins with unknown function and identifying their role will be challenging. However, these studies confirmed the key role of membrane transporters in the symbiosis. The analysis of additional genomes from ECM fungi will provide further insights into the evolution, development and ecological role of the mycorrhizal symbiosis

Investigation of horizontal gene transfer in poplar/

Fine roots of forest trees form together with certain soil fungi symbiotic structures (ectomycorrhizas), where fungal hyphae are in intimate contact with plant cells. Due to root cell degeneration, plant DNA is released and could be taken up by the fungus. The possibility that horizontal gene transfer might result in a risk for the environment should be evaluated before a massive release of genetically engineered trees into nature occurs, even though only a few convincing examples of horizontal gene transfer are known. Transgenic poplars containing a construct of the Streptomyces hygroscopicus bar gene under the control of the Cochliobolus heterostrophus GPD (glyceraldehyde-3-phosphate dehydrogenase) promoter were generated by Agrobacterium-mediated transformation. The functionality of this construct in the ectomycorrhizal model fungus Amanita muscaria was previously verified by protoplast-based fungal transformation. 35 000 ectomycorrhizas, formed between transgenic poplars and non-transgenic A. muscaria hyphae, were isolated and transferred to selective agar plates. Putative herbicide-resistant fungal colonies were obtained after the first round of selection. However, none of these colonies survived a transfer onto fresh selection medium, nor did they contain the bar gene, indicating that no horizontal gene transfer from poplar to A. muscaria occurred during symbiosis under axenic conditions. However, since ectomycorrhizas are associated under natural conditions with viruses, bacteria and other fungi, these additional associations should be evaluated in future

Sugar for my honey: Carbohydrate partitioning in ectomycorrhizal symbiosis

Nehls U, Grunze N, Willmann M, Reich M, Kuester H. Sugar for my honey: Carbohydrate partitioning in ectomycorrhizal symbiosis. PHYTOCHEMISTRY. 2007;68(1):82-91.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. (c) 2006 Elsevier Ltd. All rights reserved

Expression of major intrinsic protein genes in Sorghum bicolor roots under water deficit depends on arbuscular mycorrhizal fungal species

Drought is a limiting factor for crop plant production, especially in arid and semi-arid climates. In this study, sorghum (Sorghum bicolor) was inoculated with two arbuscular mycorrhizal fungi, either the standard Rhizophagus irregularis or the desert-adapted Rhizophagus arabicus, and grown in experimental microcosms under well-watered or drought conditions. We investigated gene expression of selected major intrinsic proteins (MIPs) of sorghum in these mycorrhizal plants, compared to non-inoculated, well-watered sorghum (control). Colonization with R. irregularis induced the MIPs SbPIP2.2 and SbPIP2.5, regardless of whether sorghum plants were well watered or not. Root colonization with R. arabicus, however, caused an exclusive, strong reduction in the transcript levels of three MIP genes (SbTIP2.1, SbNIP1.2, SbNIP2.2) under drought conditions. We also studied water transport properties of mycorrhiza-regulated MIPs. One particular MIP, SbPIP2.8, was associated with high water permeability of roots. Expression of this gene was strongly repressed in all sorghum plants (mycorrhizal and non-inoculated) that experienced drought conditions

A novel class of ectomycorrhiza-regulated cell wall polypeptides in Pisolithus tinctorius

Development of the ectomycorrhizal symbiosis leads to the aggregation of fungal hyphae to form the mantle, To identify cell surface proteins involved in this developmental step, changes in the biosynthesis of fungal cell wall proteins were examined in Eucalyptus globulus-Pisolithus tinctorius ectomycorrhizas by two-dimensional polyacrylamide gel electrophoresis. Enhanced synthesis of several immunologically related fungal 31- and 32-kDa polypeptides, so-called symbiosis-regulated acidic polypeptides (SRAPs), was observed. Peptide sequences of SRAP(32d) were obtained after trypsin digestion. These peptides were found in the predicted sequence of six closely related fungal cDNAs coding for ectomycorrhiza up-regulated transcripts. The PtSRAP32 cDNAs represented about 10% of the differentially expressed cDNAs in ectomycorrhiza and are predicted to encode alanine-rich proteins of 28.2 kDa, There are no sequence homologies between SRAPs and previously identified proteins, but they contain the Arg-Gly-Asp (RGD) motif found in cell-adhesion proteins. SRAPs were observed on the hyphal surface by immunoelectron microscopy, They were also found in the host cell wall when P. tinctorius attached to the root surface. RNA blot analysis showed that the steady-state level of PtSRAP32 transcripts exhibited a drastic up-regulation when fungal hyphae form the mantle. These results suggest that SRAPs may form part of a cell-cell adhesion system needed for aggregation of hyphae in ectomycorrhizas