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Epipogium aphyllum is a Eurasian achlorophyllous, mycoheterotrophic forest orchid. Due to its rarity, it is often protected, and its biology is poorly known. The identity and pattern of colonization of fungal associates providing carbon to this orchid have not been studied previously. Using samples from 34 individuals from Free fuck buddy in Kukes populations in Japan, Russia and France, the following were investigated: Microscopic investigations revealed that thick rhizomes were densely colonized by fungi bearing clamp-connections and dolipores, i.

Additionally, Carbon sexual encounters in Nagano, probably endophytes or parasites, were sometimes present. Since basidiomycetes colonizing the orchid are ectomycorrhizal, surrounding trees are probably the ultimate carbon source. Conversely, long, thin stolons bearing bulbils indicated active asexual multiplication, but these propagules were not colonized by fungi.

These findings are discussed in the framework of ecology and evolution of mycoheterotrophy. Orchids depend on their fungal symbionts at germination since their seeds are devoid of food reserves. Soil fungi that colonize orchid seeds provide carbon and mineral resources and allow their development into a heterotrophic, underground protocorm.

Adult orchids are often autotrophic and still harbour fungi in their roots, forming typical mycorrhizal associations Smith and Read, and perhaps reversing the carbon flow toward fungi Cameron et al. Recent research on MH orchids has used molecular techniques to identify fungal symbionts that are often unculturable and thus unidentifiable from their morphology in vitro Taylor et al.

MH fungal symbionts turned out to differ from the usual rhizoctonias, both at taxonomic and ecological levels. Most MH orchids associate with basidiomycetes that also form so-called ectomycorrhizae ECM on roots of trees and shrubs Smith and Read, Russulaceae were Carbon sexual encounters in Nagano in Corallorhiza maculata Taylor and Bruns, C.

Sebacinales were found Carbon sexual encounters in Nagano Neottia nidus-avis McKendrick et al. Using 14 C-labelling, McKendrick et al. Some rhizoctonias belonging to Ceratobasidiaceae also associate with the MH Rhizanthella gardneri Warcup, and Chamaegastrodia sikokiana Yagame et al. However, there are also some reports of saprobic basidiomycetes in MH orchids, such as Armillaria species in Galeola septentrionalis Cha and Igarashi, and Gastrodia elata Lan et Carbon sexual encounters in Nagano. In fact, Armillaria mellea colonizes and induces growth of seedlings of Galeola septentrionalis Terashita,and Erythromyces crocicreas induces germination of Galeola Carbon sexual encounters in Nagano Umata, Campbell isolated several saprobic fungi from MH orchids.

However, these saprobic fungi were identified after in vitro isolation, and saprobic contaminants may have overgrown the true mycorrhizal fungi that are often slow growing or unculturable Taylor et al. Indeed, Carbon sexual encounters in Nagano isolated by Campbell were not confirmed by recent molecular studies. Molecular methods are also sensitive to contamination by DNA from endophytic fungi or spores, so that direct observations of Adult sex dating in Luuq e.

Recently, fungi belonging to Coprinaceae, a Carbon sexual encounters in Nagano of saprobic basidiomycetes, were found in the Asian MH Epipogium roseum Yamato et al. Epipogium occurs throughout Eurasia and Africa Pridgeon et al. Alternatively, the other Epipogium species associate with ECM fungi, in which case Epipogium would illustrate a hitherto unknown variability in ecology of fungal partners.

This study focuses on E. Questions raised by previous studies on E. Epipogium aphyllum has a complex vegetative morphology, forming two kinds of rhizomes Fig. The latter presumably contribute to asexual reproduction. Therefore it is also questioned here c whether the anatomy of hypogeous plant parts has an impact on fungal colonization strategies and d how transmission to asexual offspring occurs.

These data are of relevance to future in situ or ex situ conservation activities, especially because this species is rare over its range. For example, it has been considered extinct in the United Kingdom since Harrap and Harrap, ; Kull and Hitchings, and is highly protected in France Danton and Baffray, ; in Japan, it is listed as an endangered species in the Red Data Book Environment Agency of Japan, Rhizomes were sampled in August in three countries where official authorizations were obtained France, Russia and Japanusing a protocol that allows plant survival Carbon sexual encounters in Nagano not shown.

One to ten independent rhizome fragments were harvested by digging about 20 cm away from shoots and then carefully approaching underground parts of the plant from one side; after sampling, the hole was refilled with the same soil. In the same population, rhizome fragments were sampled from seven plants to address intrapopulation diversity.

Samples were carefully washed with water to eliminate Carbon sexual encounters in Nagano soil particles. The rhizome was rinsed and cut with a sterilized scalpel into three pieces, each placed in 5 mL of sterilized distilled water in a Petri dish 9 cm in diameter and crushed with a sterilized glass rod to disperse the intracellular hyphal pelotons.

Twenty fungal pelotons per rhizome piece were harvested and pooled. Identification of fungal sequences retrieved from the investigated E. Superscript numbers indicate the number of times a clone or a direct sequence was obtained. ITS fragments amplified from coralloid rhizomes were directly sequenced as in Selosse et al. Whenever direct sequencing was not possible, PCR products were cloned as in Selosse et al. RFLP were carried out as in Selosse et al. Further, to ensure absence of usual rhizoctonia orchid symbionts with highly derived rDNA sequences, i.

To ensure relatedness of the E. Edited sequences or consensus sequences for similar clones were deposited in GenBank. Embedded samples were processed for ultramicrotomy: Stolon samples were embedded in paraffin before sectioning.

To look for fungal colonization of stolons, thin Carbon sexual encounters in Nagano and gently crushed stolons were stained with Trypan Blue Koske and Gemma, After alignment using Bioedit v7.

Genetic distances Carbon sexual encounters in Nagano estimated by maximum likelihood using a general time-reversible model Lanave et al. This model of DNA substitution was chosen using a series of hierarchical likelihood-ratio test in Modeltest 3.

Base frequencies were estimated before running the analysis, and bootstrap replicates were performed. Percentages of sequence identity between fungal ITS sequences were measured with Bioedit and compared with geographical distances and elevations by a Mantel test using XLstat; Addinsoft, Paris, France to test whether these two factors were correlated to any differences in fungal associates. The below-ground portion of E. Inflorescence buds were filled with starch not shown.

These coralloid rhizomes proved to be densely colonized by fungi, with the exception of the meristematic zone at their apex, characterized by a whitish colour Fig. Transverse sections showed that the outer cell layers were usually not colonized Fig. Some isolated hyphae running from soil to the more internal cortical cells were occasionally observed not shownbut Carbon sexual encounters in Nagano route of penetration could not be followed.

The vascular bundle and the air-filled intercellular spaces Fig. The inner cortical cells were filled with pelotons that were often elongated in one direction; these push the enlarged cell nucleus towards the periphery Fig. Direction of elongation varied from one cell to another, allowing some linear hyphae to be longitudinally cut Fig. TEM investigations confirmed that hyphae occurred in living cells with intact organelles but without starch not shown. Hyphae were surrounded by the host plasma membrane arrowed in Fig.

They consistently showed dolipores between cells Carbon sexual encounters in Nagano. On the bases of these cytological features, the fungus is confirmed as a basidiomycete. Morphology and fungal colonization of coralloid rhizomes: A fragment of a coralloid dichotomously branching rhizome; B Carbon sexual encounters in Nagano of the underground parts with bases of inflorescence shoots isthe coralloid rhizome cr sometimes giving rise to stolon s bearing bulbils sb ; C section at low magnification with pelotons p in the internal cell layers, and absence of hyphae in cortical cells cc is, intercellular spaces; n, orchid cell nucleus ; D magnification detail Carbon sexual encounters in Nagano infected cells with view of elongated fungal pelotons, cut transversally in P1 and longitudinally in P2 C and D optical microscopy stained with Trypan Blue ; E TEM of hyphae h in a peloton cut transversally as in P1with Carbon sexual encounters in Nagano dolipore d in one hypha; F TEM view of a dolipore with parenthesome p in a fungal hypha; Chat privat online erotic TEM detail of a cell with longitudinal cut of the peloton as in P2showing a clamp connection c-c.

Note the host membrane surrounding the hyphae arrows in E and G. This structure is loose and fragile, and separation from Married female wanting black male in Algeria mother plant occurred easily with disturbance.

The oldest bulbils were covered with rhizoids Fig. Sections illustrated the contrasting features of cortical cells of stolons elongated, often empty, separated by air-filled intercellular spaces What to sayemail on dating sites those of Carbon sexual encounters in Nagano densely filled with starch and stacked together, Fig.

Xylem Fuck my wife in Slovakia poorly differentiated in vascular bundles not shown. In bulbils, cell nuclei were central, and smaller cells formed an apical meristem Fig. Morphology and absence of fungal colonization of stolons and lateral bulbils b: A apex of Swinger parties in Puttalan stolon with bulbils and scaly sheathing leaf l and vascular bundle vb in transparent stolon tissues; B detail of the sheathing leaf protecting a bulbil; C sheathing leaf pushed aside to show rhizoids r covering a bulbil; D transverse section of a stolon with large, empty Carbon sexual encounters in Nagano and its lateral bulbil with starch-filled cells bc ; E contact between stolon and bulbil n, cell nucleus ; F longitudinal section showing the smaller cells in bulbils; G detailed view of the meristematic zone m at the bulbil apex.

Direct sequences were obtained for rhizome fragments from 21 plants and 13 peloton pools from Japanese plants. Two plants from one Japanese population exhibited a fungus related to Hebeloma Fig. Phylogenetic position of the E. Russian samples for Carbon sexual encounters in Nagano fungal ITSs were not sequenced are excluded. To investigate fungal diversity in rhizome fragments for which ITS was not directly sequenced, 12 ITS amplification products were cloned.

At the population level, only one sequence was shared between two plants NB1 and NC1. In an analysis of Inocybethe various sequences retrieved clustered into seven well-supported clades Fig. Five clades clustered with identified species two-thirds of the sequences clustered with I. No significant correlation with elevation was found not shown. Compared with other orchids, E. As expected from the literature Irmisch, ; Ziegenspeck,the plant encompasses two kinds of specialized underground shoots: Since numerous inflorescence buds were found on some flowering plants, the common idea that ramets die after fruiting monocarpic development, e.

Ziegenspeck, ; Rasmussen, is questionable. Instead, periods of underground growth could explain the irregular appearance of inflorescences for each plant Summerhayes, ; Soyrinki, Carbon sexual encounters in Nagano Robin, Ramet survival thus deserves further studies. Coralloid rhizomes are densely colonized by fungi Fig.

Fungal colonization closely resembles that in other orchids Scannerini and Bonfante, ; Smith and Read,with a final lysis of pelotons not shown and no colonization of meristems, inflorescence buds and central cylinder.

Lack of colonization of outer rhizome cells Fig. Another similarity with N. There is a sharp separation between infection and storage zone, i.

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