Sistotrema is a genus with ectomycorrhizal species − confirmation of what sequence studies already suggested

The ectomycorrhizal status of Sistotrema sp. is shown by morphological–anatomical and molecular identification, confirming earlier reports about Sistotrema DNA in ectomycorrhizae (ECM). For molecular identification of the ECM nuclear rDNA ITS sequences obtained from mycorrhizal root tips and fruitbodies of Sistotrema sp. were compared. Blast searches using the Sistotrema sp. sequences as query were performed in GenBank and UNITE for comparison with previously published Sistotrema sequences. The morphological–anatomical characterization of the ECM used well-established protocols for the examination of all mantle parts and rhizomorphs in different sections and views including detailed illustrations. The ECM are irregularly monopodial–pyramidal, whitish ochre to yellow ochre, and woolly. Older ones become more greyish and silvery at some patches. Diagnostic anatomical characteristics are irregularly inflated emanating and rhizomorph hyphae, ampullately inflated clamps, and the occurrence of yellow drops within the hyphae. The plectenchymatous mantle shows ring-like arranged hyphae, and a slightly gelatinous matrix. The ECM of Sistotrema sp. are compared to those of other species that form distinctly ampullate hyphae in rhizomorphs, too. The anatomically most similar ECM to those of this Sistotrema specimen are those of Hydnum repandum.     

The ectomycorrhiza <Emphasis Type="Italic">Piceirhiza internicrassihyphis</Emphasis>: A weak competitor of <Emphasis Type="Italic">Cortinarius obtusus</Emphasis>?

The ectomycorrhiza (ECM) Piceirhiza internicrassihyphis on Picea abies is described in detail. It is yellowish brown to brown with a mantle of a transitional type between plectenchymatous and pseudoparenchymatous. The inner mantle layers show some thick-walled hyphae, often with a few of them growing in parallel. These thick-walled hyphae contain particles which turn brownish in Melzer’s reagent, and their septa and walls are partially amyloid. Rhizomorphs occur infrequently and are undifferentiated, and are composed of rather loosely woven hyphae of uniform diameter. This combination of characteristics has not yet been identified in any other ECM. The lack of cystidia and the presence of amyloidy suggest that this ECM is formed by a species of the family Thelephoraceae. Piceirhiza internicrassihyphis can be found associated with ECM of Cortinarius obtusus, and the hyphae of the latter species often grow on the surface of P. internicrassihyphis and can even cover the whole tip. Although there is a close association and possibly a penetration between these two fungi, it can be excluded that the thick-walled hyphae of the inner mantle layers originate from C. obtusus. Some emanating hyphae of C. obtusus ECM seem to interact with those of P. internicrassihyphis, as indicated by short, hyphal outgrowths which were found attached to the foreign hyphae.     

Fungal relationships and structural identity of their ectomycorrhizae

Aproximately 5,000–6,000 fungal species form ectomyorrhizae (ECM), the symbiotic organs with roots of predominantly trees. The contributing fungi are not evenly distributed over the system of fungi. Within Basidiomycota exclusively Hymenomycetes and within Ascomycota exclusively Ascomycetes contribute to the symbiosis. Hymenomycetes play a big part, Ascomycetes a minor role; Zygomycetes only form exceptionally ECM. Responsible for ascomycetous ECM are mostly Pezizales with their hypogeous derivatives, whereas Boletales, Gomphales, Thelephorales, Amanitaceae, Cantharellaceae, Cortinariaceae, Russulaceae, and Tricholomataceae are the most important ectomycorrhizal relationships within Hymenomycetes. ECM, as transmitting organs between soil and roots, are transporting carbohydrates for growth of mycelium and fruitbodies from roots and have to satisfy the tree’s demand for water and nutrients. The latter task particularly influences the structure of ECM as nutrients are patchily distributed in the soil and saprotrophic as well as ectomycorrhizal fungi can act as strong competitors for nutrients. In focusing these requirements, ECM developed variously structured hyphal sheaths around the roots, the so-called mantles, and differently organized mycelium that emanates from the mantle, the so-called extramatrical mycelium. The mantles can be plectenchymatous consisting of loosely woven, differently arranged hyphae or they are densely packed, forming a pseudoparenchyma similar to the epidermis of leaves. The extramatrical mycelium grows either as simple scattered hyphae from the mantle into the soil or it can be united to undifferentiated rhizomorphs with a small reach or to highly organized root-like organs with vessel-like hyphae for efficient water and nutrient transport from distances of decimeters. Cystidia, sterile and variously shaped hyphal ends, possibly appropriate for preventing animal attack, in addition, can cover mantles and rhizomorphs. Although only a limited number of species could be considered, some general conclusions are possible.The genus Tuber forms needle-shaped cystidia and lacks rhizomorphs and clamps. Gomphales ECM are identified by rhizomorphs with ampullate inflations at septa of some hyphae and by oleoacanthocystidia or/and oleoacanthohyphae. Thelephoraceae reveal a great diversity of mantle structures and of extramatrical mycelium, with some additional optional characters, i.e., dark brown color, cystidia, blue granules, amyloid hyphae, or amyloid septa. Bankeraceae are mostly characterized by plectenchymatous mantles with star-like pattern and chlamydospores. Russulaceae possess smooth and hydrophilic ECM. Russula forms plectenchymatous mantles with knob-bearing cystidia, so-called russuloid cystidia, or pseudoparenchymatous mantles without cystidia. Lactarius lacks cystidia and shows laticifers within plectenchymatous or within pseudoparenchymatous mantles. The Boletales families Boletaceae, Gyroporaceae, Melanogastraceae, Paxillaceae, Rhizopogonaceae, Sclerodermataceae, and Suillaceae have the most advanced rhizomorph type, the so-called boletoid rhizomorphs, and reveal generally plectenchymatous mantles, frequently with ring-like patterns. Gomphidiaceae and Albatrellaceae provide cystidia, plectenchymatous mantles, and amyloidy; Gomphidiaceae are generally growing in ECM of Suillaceae and Rhizopogonaceae. Cortinariaceae reveal plectenchymatous mantles and undifferentiated or differentiated rhizomorphs or lack rhizomorphs at all. Cortinarius and Dermocybe are distinct by irregularly shaped, bent to tortuous ECM with many rhizomorphs, some growing over the mycorrhizal tip into the soil. Inocybe lacks rhizomorphs and its emanating hyphae are furnished by many secondary septa and prominent clamps with a hole. Rozites lacks rhizomorphs, too, and reveals a distinctly amyloid gelatinous mantle matrix. Descolea and Descomyces are covered by bolbitioid cystidia. Lastly, the genus Tricholoma forms plectenchymatous mantles and a high diversity of rhizomorphs. Some of the ectomycorrhizal features are used to hypothesize relationships at different taxonomic levels. These conclusions are compared with recently developed molecular hypotheses. Correspondence between the two types of hypotheses are evident, while some conflicts wait for a settlement.     

Two sebacinoid ectomycorrhizae on Chinese pine

Sebacinoid fungi show a broad mycorrhizal capacity; therefore, they play a very important role in natural systems. Worldwide, fungi of Sebacinales are present under different environmental conditions and associate with diverse plant hosts, however, are hitherto poorly studied in China. Two sebacinoid ectomycorrhizae (ECM), Pinirhiza multifurcata and Pinirhiza nondextrinoidea, are described in detail morphologically and anatomically in the present study. They share a plectenchymatous outer mantle with multiply ramified hyphae in a gelatinous matrix, clampless, thin, thick-walled emanating hyphae with mostly Y-shaped ramifications and triangular inflations at the point of ramification. P. multifurcata and P. nondextrinoidea can be distinguished by thick cells in mantle layers, the ramification of emanating hyphae, the presence or absence of rhizomorphs, as well as the differing color reaction in Melzer's reagent. The putative molecular phylogenetic relationships of P. multifurcata and P. nondextrinoidea were inferred by analyses of the partial large subunit nuclear rDNA (nLSU); however, an affiliation to fungal species was not possible. This is the first report of sebacinoid ECM on Chinese pine.     

Four ectomycorrhizae of Pyronemataceae (Pezizomycetes) on Chinese Pine (<Emphasis Type="Italic">Pinus tabulaeformis</Emphasis>): morpho-anatomical and molecular-phylogenetic analyses

Morphological and anatomical characters of four ectomycorrhizae with affinities to the genera Humaria, Geopora, and Trichophaea of Pyronemataceae (Pezizomycetes, Ascomycota) on Chinese Pine (Pinus tabulaeformis) are described. The ectomycorrhizae are yellowish brown to brown, and have pseudoparenchymatous outer mantle layers and partially warty emanating hyphae with thick walls and without clamps. Intrahyphal hyphae are present, and no rhizomorphs are formed. The four ectomycorrhizae are distinguishable by differences in cell shape of outer mantle layers and the presence of cystidia. Ectomycorrhizae of a possible Humaria species (Pinirhiza humarioides) lack cystidia and have irregularly inflated cells on the outer mantle layer that are connected with thin septa. The two ectomycorrhizae showing probable affinities to Geopora species (“P. daqingensis” and “P. geoporoides”) possess row-like arranged cells in the outer mantle layer and cell heaps, and differ by the presence or absence of cystidia as well as by the structure of the inner mantle layers. Ectomycorrhizae likely having been formed by a Trichophaea species (“P. trichophaeoides”) have oval to polygonal cells and no cystidia. The possible taxa affiliations were assessed by molecular-phylogenetic analyses of the internal transcribed spacer (ITS) and partial large subunit (LSU) nrDNA. Morphological and anatomical characters are discussed against the background of the LSU phylogeny.     

Ectomycorrhizas of Cortinarius helodes and Gyrodon monticola with Alnus acuminata from Argentina

Field ectomycorrhizas of Cortinarius helodes Moser, Matheny & Daniele (sp. nov) and Gyrodon monticola Sing. on Alnus acuminata Kunth (Andean alder, aliso del cerro) are described based on morphological and anatomical features. Ectomycorrhizal roots were sampled beneath fruitbodies of C. helodes and G. monticola from two homogeneous A. acuminata forest sites located in Tucumán and Catamarca Provinces in Argentina. C. helodes ectomycorrhizas showed a thick white to beige mantle exuding a milky juice when injured, were bluish toward the apex, and had hyphal strands in the mantle. G. monticola ectomycorrhizas showed some conspicuous features like highly differentiated rhizomorphs, inflated brown cells on the mantle surface, and hyaline and brown emanating hyphae with dolipores. Restriction fragment length polymorphism analysis of the nuclear rDNA internal transcribed spacer provided a distinctive profile for each of the collections of fruitbodies and the mycorrhizal morphotypes.     

Ectomycorrhizae between <Emphasis Type="Italic">Alnus acuminata</Emphasis> H.B.K. and <Emphasis Type="Italic">Naucoria escharoides</Emphasis> (Fr.:Fr.) Kummer from Argentina

Field ectomycorrhizae of Naucoria escharoides on Alnus acuminata ("andean alder", "aliso del cerro") are described in detail for the first time. Naturally occurring ectomycorrhizal roots were sampled beneath sporocarps of N. escharoides. The samples were taken from four natural forest plots at two homogeneous A. acuminata sites (Tucumán and Catamarca Provinces, Argentina). The ectomycorrhizae were characterized morphologically and compared by means of PCR/RFLP analysis of the internal transcribed spacer region of the nuclear rDNA. The most important morphological features of the ectomycorrhizae are a white to pale yellow mantle, simple to monopodial branches, hyaline emanating hyphae, abundant hyphal bundles emerging more or less perpendicularly from a plectenchymatous mantle, and an acute or rounded apex with or without a mantle. N. escharoides fruitbodies have white basal mycelium with emanating hyphae similar to those of andean alder ectomycorrhizae. The RFLP profiles of sporocarps and mycorrhizae were the same.     

Scanning Electron Microscope Studies of Interactions between Agaricus bisporus (Lang) Sing Hyphae and Bacteria in Casing Soil

Relationships between the hyphae of Agaricus bisporus (Lang) Sing and bacteria from the mushroom bed casing layer were examined with a scanning electron microscope. Hyphae growing in the casing layer differed morphologically from compost-grown hyphae. Whereas the compost contained thin single hyphae surrounded by calcium oxalate crystals, the casing layer contained mainly wide hyphae or mycelial strands without crystals. The bacterial population in the hyphal environment consisted of several types, some attached to the hyphae with filamentlike structures. This attachment may be important in stimulation of pinhead initiation.     

Characterization and identification of field ectomycorrhizae of Boletus edulis and Cistus ladanifer

Field ectomycorrhizae sampled under Boletus edulis and Cistus ladanifer have been characterized and described in detail based on standard morphological and anatomical characters. The described ectomycorrhiza has traits typical of Boletales: whitish with three differentiated plectenchymatous layers in the mantle in plan view forming ring-like structures and rhizomorphs with highly differentiated hyphae. The inflated, smooth cystidia-like clavate end cells on the surface of the rhizomorphs and their slightly twisted external hyphae are additional characterizing features. The Hartig net occupies 1 1/2 rows of cortical cells, partly reaching the endodermis. Not all hyphae have clamps. The identification of the fungal symbiont as B. edulis was confirmed by ITS rDNA sequence comparison between mycorrhizas and sporocarps. The singularity of this symbiotic association, as well as its ecological and practical implications, are discussed.     

Ectomycorrhizae of Tomentella albomarginata (Thelephoraceae) on Scots pine

 The ectomycorrhizae of Tomentella albomarginata are comprehensively described and compared to ectomycorrhizae of other Tomentella species and to ectomycorrhizae of some members of Thelephoraceae ss. Stalpers and Bankeraceae ss. Stalpers. The ectomycorrhizae of T. albomarginata are characterized by a hyphal net lying on a pseudoparenchymatous mantle surface, by tubular outgrowths of irregularly angular mantle cells, and by clamp-bearing emanating hyphae. Accepted: 15 July 1995     

Morphology,Anatomy and Histochemistry of Fagus grandifolia Ehrh. (North American Beech) Ectomycorrhizas

Abstract: Five distinct ectomycorrhizal morphotypes were recognized on root systems of Fagus grandifolia collected in a maple-beech woodlot. Three morphotypes showed extensive root branching whereas two had limited root branching. One morphotype, a bright orange, smooth type, was studied in detail anatomically. A compact mantle with few emanating extraradical hyphae covered the root apex of each mycorrhizal lateral. A Hartig net extended around epidermal and hypodermal cells. Roots were mostly diarch and a single-layered endodermis and a biseriate hypodermis with the inner hypodermis having suberized walls were present in all specimens examined. The mantle, and to a lesser extent the Hartig net, stored lipids, polysaccharides and proteins.     

Asexual reproduction of <Emphasis Type="Italic">Hygrophorus olivaceoalbus</Emphasis> by intracellular microsclerotia in root cells of <Emphasis Type="Italic">Picea abies</Emphasis> – A winner of ozone stress?

Hygrophorus olivaceoalbus has long been known as an ectomycorrhizal fungus, formerly designated with the artificial binomen Piceirhiza gelatinosa. Recently, it has been found to be abundant and very frequent under double ambient ozone free air fumigation of mature Norway spruce trees. As it has already been reported that this fungus can form intercellular hyphae within the root meristem, a more detailed study was performed to clarify its type of root colonization. The present study not only revealed intercellular hyphae within the meristematic zone but also intracellular hyphae within root cortex cells which grow towards and into large tannin droplets—phenolic compounds usually deposited as defensive aids—to finally fill them completely. The hyphal assemblages become globular which at first sit as separated hyphal balls within cells still containing cytoplasm. Later on, they are apparently released from the root cells, presumably as microsclerotia for dispersal of the species. Old ectomycorrhizae (ECM) show an apical pore, and later a large orifice of a tube-like cylinder formed by the thick, persistent hyphal mantle. The root tissue is progressively degrading towards proximal parts. Disintegrating root cells apparently liberate the microsclerotia through the orifice. Further studies have to find out the mechanism by which the microsclerotia are liberated and whether they operate as asexual propagules and lastly how and by whom they are propagated. As these ECM are found under ozone stress, and with identical features at higher altitudes, stress impact on trees might be the causative agent for the high frequency and abundance of Hygrophorus olivaceoalbus ECM.     

Matrix valency regulates integrin-mediated lymphoid adhesion via Syk kinase

Lymphocytes accumulate within the extracellular matrix (ECM) of tumor, wound, or inflammatory tissues. These tissues are largely comprised of polymerized adhesion proteins such as fibrin and fibronectin or their fragments. Nonactivated lymphoid cells attach preferentially to polymerized ECM proteins yet are unable to attach to monomeric forms or fragments of these proteins without previous activation. This adhesion event depends on the appropriate spacing of integrin adhesion sites. Adhesion of nonactivated lymphoid cells to polymeric ECM components results in activation of the antigen receptor-associated Syk kinase that accumulates in adhesion-promoting podosomes. In fact, activation of Syk by antigen or agonists, as well as expression of an activated Syk mutant in lymphoid cells, facilitates their adhesion to monomeric ECM proteins or their fragments. These results reveal a cooperative interaction between signals emanating from integrins and antigen receptors that can serve to regulate stable lymphoid cell adhesion and retention within a remodeling ECM.     

Hyphal interference by Trametes hispida

The antagonistic activity of Trametes hispida Bagl. in dual culture with Hirschioporus species and other non-basidiomycetous fungi is interpreted as hyphal interference. Hyphae of T. hispida grow into and over colonies of sensitive fungi. Contact with hyphae of T. hispida results in cessation of growth and rapid necrosis of affected cells. Ultrastructural studies of the affected hyphae of Hirschioporus pargamenus (Fr.) Bond. & Sing. showed an early formation of extraplasmalemmal wall deposits, disruption of membrane systems, coagulation of cytoplasm, localized dissolution of walls, and loss of cell contents without penetration by the antagonist.     

Morphological-anatomical characterization and identification of Tomentella ectomycorrhizas

Over the last two decades, much information has been gathered on the ectomycorrhizal fungus community composition of plant associations of boreal, temperate, and tropical regions. Worldwide, Tomentella ectomycorrhizas (ECM) are often common and dominant in the mycorrhizosphere of coniferous and deciduous forests. They are present under different environmental conditions and associate with diverse plant hosts. Tomentella sporocarps, however, are rarely found aboveground, so Tomentella species are often missing from fungus community studies based on fruit-body presence. Tomentella is a resupinate genus of Thelephoraceae (Basidiomycota) forming black-brown, brown, yellow, or ochre ECM on the roots of gymnosperm and angiosperm trees, distinguished by typical morphological-anatomical characteristics (clamped hyphae, angular mantle, surface network, special rhizomorphs and cystidia). In this paper, we review the taxonomic position and morphological-anatomical characteristics of Tomentella ECM. A short summary of the microscopic features used for distinguishing tomentelloids during morphotyping and identification is presented in order to support molecular and ecological studies of ectomycorrhizal fungus communities.     

New Neotropical Sebacinales Species from a Pakaraimaea dipterocarpacea Forest in the Guayana Region,Southern Venezuela: Structural Diversity and Phylogeography

Pakaraimaea dipterocarpacea, a member of the Dipterocarpaceae endemic in the Guayana region, is associated with a diverse community of ectomycorrhizal (ECM) fungi. Amongst the 41 ECM fungal species detected in a 400 m² P. dipterocarpacea ssp. nitida plot in Southern Venezuela, three species belonged to the Sebacinales. We tested whether ECM anatomotype characterization can be used as a feasible element in an integrative taxonomy in this diverse fungal group, where the relevance of fruitbody morphology for species delimitation seems limited. Using a combination of ECM morpho-anatomical characterizations and phylogenetic analyses based on nuclear ITS and LSU sequences, we report three new species. The main distinguishing features of Sebacina guayanensis are the yellowish cell walls together with conspicuous undifferentiated, uniform compact (type B) rhizomorphs. Staghorn-like hyphae are characteristic of S. tomentosa. The combination of clusters of thick-walled emanating hyphae, including hyphae similar to awl-shaped cystidia with basal dichotomous or trichotomous ramifications, and the presence of type B rhizomorphs were characteristic of a third, yet unnamed species. The three species belong to three different, possibly specifically tropical clades in Sebacinales Group A. The geographic distribution of phylogenetically related strains was wide, including a Dicymbe forest in Guyana and an Ecuadorian rainforest with Coccoloba species. We show that ECM morpho-anatomy can be used, in combination with other analyses, to delineate species within Sebacinales Group A. In addition to phylogenetic information, type B rhizomorphs observed in different Sebacinales clades have important ecological implications for this fungal group. The phylogeography of Sebacinales suggests that dispersion and host jump are important radiation mechanisms that shaped P. dipterocarpacea ECM fungal community. This study emphasizes the need for more sequence data to evaluate the hypothesis that phylogeographic relationships between neo- and paleotropical ECM fungal species could be attributed to the vicariance of cross-continental hosts such as the Dipterocarpacae.     

Ecological and anatomical characterization of some Pinus patula ectomycorrhizas from Mpumalanga, South Africa

 Ectomycorrhizal (ECM) fungi are an important component of the Pinus patula Schlechdt. et Cham. forest ecosystem in Mpumalanga, South Africa. ECM roots are intimately associated with accumulated litter on the forest floor and four ECM isolates where examined to determine whether they provide plant access to inorganic and organic sources of nitrogen (N) and inorganic, complexed inorganic and organic sources of phosphorus (P). In in vitro studies, all isolates were found to utilize most of the organic N compounds and organic or complexed inorganic P compounds supplied. Thus, ECM fungi could play a significant role in providing N and P to P. patula, especially from sources to which the host plant would not normally have access. Temperature sensitivities and pH optima of the four isolates differed. Of the ECM isolates WITS 01 and WITS 06 were collected from a high-litter site; WITS 01 mycorrhizas, identified as Scleroderma citrinum, were white, smooth and dichotomously branched with smooth, pale yellow, differentiated rhizomorphs. The mantle was plectenchymatous with outer and inner layers showing ring-like arrangements of hyphal bundles. The Hartig net had a palmetti shape. The WITS 02 (not identified) mycorrhizas were brown with lighter coloured root tips, with simple to dichotomous branching, smooth with no distinct mantle and sparse hyphae occurred on the root surface. The Hartig net was palmetti type with lobed haustoria. The results are discussed in relation to ECM distribution and function in nutrient cycling. Accepted: 7 January 1999     

Scanning electron microscopic study of clamp connections in higherBasidiomycetes

Clamp connections on vegetative hyphae of six species belonging to different groups of higherBasidiomycetes were studied by scanning electron microscopy. Different types of clamps, short and long, high and low, small and large, gently or abruptly curved, medallion-type and without a slit, single, coupled and in whorls, as well as clamps occurring on anastomoses were observed. No characteristic features usable for the species-specific differentiation of the vegetative mycelia of higherBasidiomycetes were found. Greater variability in the type of clamps, especially in their distribution on hyphae, found in species belonging toAphyllophorales contributes to the theory of their relative primitivism in comparison withAgaricales.     

Wirtelschnallen bei homo- und heterothallischenCoprinus-Arten (Agaricales)

Within agaric fungi whorled (multiple) clamps have been described so far only for the homothallicCoprinus myceliocephalus Lange. The new speciesCoprinus verticillatus Schulz-Weddigen (ined.) develops fertile mycelia which form septae without or with 1–5 clamps. Stronger hyphae tend to branch dichotomously. Most cells of the vegetative system contain many nuclei. Though single spore cultures remain sterile, they can develop multiple clamps, too. Moreover, very young mycelia produce a special type of oidiophores. According to its mating interactions a bipolar pattern characterizes the heterothallicC. verticillatus.