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Mariaestrella Legaz - One of the best experts on this subject based on the ideXlab platform.
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Sugarcane glycoproteins control dynamics of cytoskeleton during Teliospore germination of Sporisorium scitamineum
Mycological Progress, 2019Co-Authors: Elena Sánchez-elordi, Carlos Vicente, Frantisek Baluska, Mariaestrella LegazAbstract:Sporisorium scitamineum Teliospores possess an organized cytoskeleton involved in important developmental and physiological processes. It has been described that microtubules appear to be fundamental for nucleus translocation during germination and hyphal growth, whereas actin polymerization is necessary for the formation of invaginations during Teliospore displacement. Here, a global vision of the actin cytoskeleton organization throughout the life cycle of S. scitamineum cells is shown, providing evidence that a perfectly structured F-actin network is necessary to trigger smut pathogenicity. Moreover, although myosin presence in Teliospores had been previously described, herein actin and myosin co-locations are demonstrated by confocal microscopy during both invaginations formation and germination. In turn, F-actin and microtubules (MTs) interact, jointly participating in the establishment of cell polarity. The resistant sugarcane cultivar Mayari 55-14 produces high molecular mass glycoproteins (HMMG) that differently affect F-actin organization at different stages of fungal development. HMMG first supported F-actin to induce the movement of Teliospores towards the cytoagglutination points. At later stages of fungal development, HMMG disorganized F-actin which prevented the protrusion of germinative tube. A continuous exposure to HMMG provoked apoptosis in pathogenic, diploid cells and a delay in sporidia conjugation that could be crucial for plant resistance.
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sugar cane arginase competes with the same fungal enzyme as a false quorum signal against smut Teliospores
Phytochemistry Letters, 2015Co-Authors: Elena Sanchezelordi, Laura Morales De Los Rios, Carlos Vicente, Mariaestrella LegazAbstract:Sugar cane cultivars resistant to smut produce a pool of glycoproteins after experimental infection, one of them develops arginine activity. This arginase induces cytoagglutination of smut Teliospores but impedes germination. Teliospores also secrete a fungal arginase that accelerates their own germination. This fungal arginase binds to Teliospore cell walls. The affinity of this arginase for the ligand shows to be higher than that found for plant arginase. In fact, fungal arginase removes sugar cane arginase previously bound to their ligands in the cell wall whereas the inverse process is practically negligible. The enzymatic activity is required to the binding of the protein to Teliospore cell wall but it is not related to the germination process. Thus, it can be concluded that the binding of fungal arginase to smut Teliospores activates a signal transduction cascade that enhances germination.
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in vitro effects of caffeic acid upon growth of the fungi sporisorium scitamineum
Journal of Plant Interactions, 2010Co-Authors: Rocio Santiago, Carlos Vicente, Roberto De Armas, Maria Blanch, Mariaestrella LegazAbstract:Abstract Caffeic acid (CA) sometimes behaves as a potent phytotoxin affecting plant and fungi growth and physiology. The aim of the present study was to investigate whether CA at a concentration range similar to that found in sugarcane leaves, had any effect against different phases of Sporisorium scitamineum growth cycle. Leaf CA concentration from two different sugarcane cultivars, Mayari (My) 55-14, resistant, and Barbados (B) 42231, susceptible to smut, was chromatographically quantified by HPLC. A smut elicitor promoted an increase of CA concentration in the resistant cv. while no effect was produced in the susceptible one. The effect of CA upon S. scitamineum growth cycle showed to be dependent of both concentration and time. At 5.0 µg ml−1, CA produced and inhibition of Teliospore germination, haploid sporidia production and dikaryotic mycelium appearance. At 30 µg ml−1, CA produced similar effects to these just described. Inhibition was more evident after 24 h or 28 h incubation of Teliospores in ...
Franz Oberwinkler - One of the best experts on this subject based on the ideXlab platform.
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Teliosporogenesis of Uleiella chilensis as seen by transmission electron microscopy.
2016Co-Authors: Kai Riess, Matthias Lutz, Franz Oberwinkler, Max E. Schön, Heinz Butin, Sigisfredo GarnicaAbstract:Material from a–e was prepared from a herbarium specimen. (a) Section through a sorus showing external Teliospores (one is indicated by an arrow). (b) Teliospore wall with a sheath (arrowhead), an exosporium with ornaments (small arrow) and an endosporium (large arrow). (c) Section through a young Teliospore with ornaments showing the beginning of septation (arrow). (d) Section through a Teliospore showing one complete septum (arrow). (e) Section through a mature Teliospore with two more or less rounded segments. (f) Section through a germinating Teliospore, showing the multicellular content. Scale bar = 10 μm in (a), 0.2 μm in (b–c) and 0.5 μm in (d–f).
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Sections through galls of Entorrhiza casparyana on Juncus articulatus, as seen by SEM.
2015Co-Authors: Robert Bauer, Franz Oberwinkler, Sigisfredo Garnica, Kai Riess, Michael Weiß, Dominik BegerowAbstract:(a) Section showing intracellular Teliospore packets (one is indicated by an arrow). (b) Teliospore packet in detail, showing Teliospores and hyphal spirals (one is indicated by an arrow). Scale bar is 1 mm in (a) and 20 μm in (b).
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Teliospore formation and germination of Entorrhiza casparyana, as seen by TEM (a-c) or LM (d).
2015Co-Authors: Robert Bauer, Franz Oberwinkler, Sigisfredo Garnica, Kai Riess, Michael Weiß, Dominik BegerowAbstract:(a) Section through an intracellular hyphal coil showing a terminal Teliospore initial with two nuclei (n). (b) Section through a Teliospore tetrad showing the cruciform septation. (c) Section through a Teliospore tetrad showing the germination of one of the four Teliospore compartments. One nucleus is visible at the arrow, probably migrating into the germination hypha. (d) Light microscopic micrograph of a germinating Teliospore illustrated to show the synchronous development of the germination hyphae and the terminal sigmoid propagules (one is indicated by an arrow). Scale bar is 2 μm in (a-b), 3 μm in (c), and 40 μm in (d).
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spore liberation and dispersal in smut fungi
Botanica Acta, 1998Co-Authors: Meike Piepenbring, G Hagedorn, Franz OberwinklerAbstract:: Teliospores are the most important diaspores of smut fungi, albeit not the only ones. The role of basidla, basidiospores, secondary spores, yeast cells, and infected parts of the host for dispersal has often been neglected. Many smut species have soral structures like galls, peridia, and elaters, which cause Teliospores to be liberated over prolonged periods. This increases the chance that at least some spores are released under favourable wet climatic conditions and while host plants are susceptible. In this review, the diversity of dispersal units as well as vectors of smut fungi are presented. The importance of timing of diaspore liberation, flexibility in dispersal strategies, and the genetic and evolutionary implications of dispersal strategies of smut fungi are discussed. The general considerations are complemented by examples based on original field and laboratory observations: peridia of Farysia corniculata and certain species of Sporisorium expose the spore mass by hygroscopic movement under wet conditions (hygrochasy) favourable for Teliospore germination and infection of a host plant. Basidia with firmly attached basidiospores liberated from spore balls of Doassansiopsis deformans, branched basidiospores of Rhamphospora nymphaeae, needle-shaped basidiospores of species of Entyloma, folded basidiospores of Mycosyrinx cissi, and stellate groups of yeast cells of Trichocintractia utriculicola show enlarged surfaces, which are advantageous for dispersal in water. Galls filled with spore balls of Doassansiopsis limnocharidis and witches' brooms formed by spikelets infected by Cintractia standleyana separate from the host and fall into water where they are dispersed.
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Teliospores of smut fungi Teliospore walls and the development of ornamentation studied by electron microscopy
Protoplasma, 1998Co-Authors: M. Piepenbring, R. Bauer, Franz OberwinklerAbstract:The walls of mature Teliospores and the development of ornamentation, as seen by transmission electron microscopy, are described for 37 genera of smut fungi, based on observations of ca. 120 species and on literature. Structural diversity of mature Teliospore walls is due to differences in spore wall layers forming the spore wall (endosporium, middle layer, exosporium, ornamentation) and to different elements forming the ornamentation (exosporium, ornaments, sheath, hyphal wall, adjacent fungal cells, material of the host). During teliosporogenesis the outer layers are usually deposited first. At the beginning of the formation of the ornamentation the plasma membrane may be smooth or undulated carrying the developing ornaments on its tips or in its depressions. The ornamentation of some genera appears similar when seen by scanning electron microscopy, but can be the product of different developmental patterns (e.g., warts of species of Farysia, Tilletia , and Ustilago ), however, warty and reticulate ornamentation can both be produced by similar developmental processes (shown, e.g., for species of Cintractia and Tilletia ). Typical structures of the mature Teliospore wall and developmental patterns based on homologous similarities are described for the following groups of genera or species: Macalpinomyces, Melanopsichium, Sporisorium , and Ustilago infecting members of the family Poaceae; Kuntzeomyces, Testicularia , and Trichocintractia; Anthracoidea, Cintractia, Heterotolyposporium piluliforme , and Tolyposporium junci; Glomosporium, Sorosporium , and Thecaphora; Conidiosporomyces, Erratomyces, Ingoldiomyces, Neovossia, Oberwinkleria , and Tilletia; Entyloma , and genera of the Doassansia group; Liroa, Microbotryum, Sphacelotheca, Ustilago infecting dicotyledons, and Zundeliomyces; Aurantiosporium, Fulvisporium , and Ustilentyloma . Special characteristics of the Teliospore wall were observed for the genera Dermatosorus, Doassinga, Entorrhha, Farysia, Mycosyrinx, Rhamphospora , and some species of Tolyposporium .
M R Bonde - One of the best experts on this subject based on the ideXlab platform.
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survival of Teliospores of tilletia indica in arizona field soils
Plant Disease, 2004Co-Authors: M R Bonde, S E Nester, Mary W Olsen, D. K. BernerAbstract:The survival of Teliospores of the Karnal bunt of wheat pathogen, Tilletia indica, was determined in field plots in Tucson, AZ. Two methods were used to test viability during a 48-month period in which 21-μm-pore-size polyester mesh bags of Teliospore-infested soil were buried in irrigated and nonirrigated field plots at two sites. One method determined the total number of viable Teliospores in a soil sample, regardless of whether or not they could be extracted from the soil using a sucrose centrifugation technique. The total number of viable Teliospores declined over time in both irrigated and nonirrigated field plots and in the same soils in the laboratory. Based on nonlinear regressions, total number of viable Teliospores decreased from 55.7% at time zero to 9.7 and 6.7% for nonirrigated and irrigated field soils, respectively, in 48 months. Total number of viable Teliospores in soil in the laboratory decreased from 55.7 to 34.0% after 48 months. The second method determined germination percentages of Teliospores extracted from the soil samples by means of a sucrose centrifugation technique. Based on linear regressions of transformed data, germination of Teliospores extracted from irrigated and nonirrigated field soils, and control (laboratory) soil, significantly decreased over time. The rate of decrease in germination was significantly greater for Teliospores from irrigated field plots than from nonirrigated plots and the laboratory soil. At time zero, 55.7% of Teliospores germinated, and by 48 months, average germination of Teliospores extracted from soil in nonirrigated plots had decreased to 13.6% compared with 4.4% in irrigated plots and 36.8% for Teliospores in the laboratory control. Regression over time of total number of viable Teliospores accounted for more of the overall variability than did regression over time of germination percentages of extracted Teliospores. Neither field site nor soil depth had any effect on total number of viable Teliospores or on Teliospore germination percentages.
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survival of tilletia indica Teliospores in different soils
Plant Disease, 2004Co-Authors: M R Bonde, D. K. Berner, S E Nester, Mary W Olsen, Gary L Peterson, B M Cunfer, T SimAbstract:ABSTRACT To determine the potential for Tilletia indica, cause of Karnal bunt of wheat, to survive and become established in new areas, a Teliospore longevity study was initiated in Kansas, Maryland, Georgia, and Arizona. Soil from each location was infested with T. indica Teliospores and placed in polyester mesh bags. The bags were placed within soil from the same location within polyvinyl chloride pipes. Pipes were buried in the respective plots such that the bags were at 5-, 10-, and 25-cm depths. Each pipe was open at the ends to allow interaction with the outside environment, however fitted with screens preventing possibility of Teliospore escape. In the Karnal bunt-quarantine area of Arizona, bags of infested soil also were placed outside the pipes. Teliospore-infested soil from each location was maintained dry in a laboratory. During the first 2 years, viability declined more rapidly in pipes than outside pipes, and more rapidly in fields in Kansas and Maryland than in Georgia or Arizona. After 2 y...
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survival of Teliospores of tilletia indica in soil
Plant Disease, 2004Co-Authors: M Babadoost, D E Mathre, R H Johnston, M R BondeAbstract:Babadoost, M., Mathre, D. E., Johnston, R. H., and Bonde, M. R. 2004. Survival of Teliospores of Tilletia indica in soil. Plant Dis. 88:56-62. This study was conducted to assess survival of Tilletia indica Teliospores in a location in the northern United States. Soils differing in texture and other characteristics were collected from four locations, equilibrated to –0.3 MPa, and infested with Teliospores of T. indica to give a density of 10 3 Teliospores per gram of dry soil. Samples (22 g) of the infested soil were placed in 20-µm mesh polyester bags, which were sealed and placed at 2-, 10-, and 25-cm depths in polyvinyl chloride tubes containing the same field soil as the infested bags. Tubes were buried vertically in the ground at Bozeman, MT, in October 1997. Soil samples were assayed for recovery and germination of T. indica Teliospores 1 day and 8, 20, and 32 months after incorporation of Teliospores into soil. The rates of Teliospores recovered from soil samples were 90.2, 18.7, 16.1, and 13.3% after 1 day and 8, 20, and 32 months after incorporation of Teliospores into soil, respectively, and was significantly (P < 0.01) affected by soil source. The percentage of Teliospore recovery from soil was the greatest in loam soil and lowest from a silt loam soil. The rate of Teliospores recovered from soil was not significantly affected by depth of burial and the soil source–depth interaction during the 32-month period. The percentage of germination of Teliospores was significantly (P < 0.01) affected by soil source and depth of burial over the 32month period. The mean percentage of Teliospore germination at 1 day, and 8, 20, and 32 months after incorporation into soils was 51.3, 15.1, 16.4, and 16.5%, respectively. In another experiment, samples of silty clay loam soil with 5 × 10 3 Teliospores of T. indica per gram of soil were stored at different temperatures in the laboratory. After 37 months of incubation at 22, 4, –5, and –18°C, the rates of Teliospore recovered from soil were 1.6, 2.0, 5.7, and 11.3%, respectively. The percentage of spore germination from soil samples was highest at –5°C. Microscopy studies revealed that disintegration of Teliospores begin after breakdown of the sheath-covering Teliospore. The results of this study showed that Teliospores of T. indica can survive in Montana for more than 32 months and remain viable. Additional keywords: Neovossia indica, Triticum aestivum
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size selective sieving for detecting Teliospores of tilletia indica in wheat seed samples
Plant Disease, 2000Co-Authors: Gary L Peterson, M R Bonde, J G PhillipsAbstract:A method was developed to isolate Teliospores of Tilletia indica from infested grain. The technique was evaluated to determine its sensitivity for detection and quantification of Teliospores, the time required to conduct an individual test, and its utility for the detection and identification of the pathogen for phytosanitary regulation and seed certification. A seed wash of a 50-g grain sample was washed through 53-μm and 20-μm pore size nylon screens to remove unwanted debris and to concentrate and isolate Teliospores. The material retained in the 20-μm screen was suspended for direct microscopic examination or plated on water agar for Teliospore germination and identification by polymerase chain reaction (PCR) utilizing two pairs of T. indica-specific primers. The reliability of detection for both light microscopy and PCR are 100% at an infestation of five Teliospores per 50-g sample. The proportion of Teliospores recovered from grain samples artificially infested with T. indica was 0, 82, 88, 81, and 82%, respectively, at infestation levels of 0, 1, 2, 5, and 10 Teliospores per 50-g wheat sample. Extraction efficiency was comparable to the centrifuge seed-wash method currently used by most seed health laboratories. Sample analysis using size-selective sieving was more than 83% faster than the standard centrifuge seed wash.
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comparison of effects of acidic electrolyzed water and naocl on tilletia indica Teliospore germination
Plant Disease, 1999Co-Authors: M R Bonde, S E Nester, A Khayat, Joseph L Smilanick, Reid D Frederick, Norman W SchaadAbstract:Bonde, M. R., Nester, S. E., Khayat, A., Smilanick, J. L., Frederick, R. D., and Schaad, N. W. 1999. Comparison of effects of acidic electrolyzed water and NaOCl on Tilletia indica Teliospore germination. Plant Dis. 83:627-632. Definitive identification of free Teliospores of Tilletia indica, causal agent of Karnal bunt of wheat, requires polymerase chain reaction (PCR)-based diagnostic tests. Since direct PCR amplification from Teliospores has not been reliable, Teliospores first must be germinated in order to obtain adequate DNA. We have routinely surface-sterilized Teliospores for 2 min with 0.4% (vol/vol) sodium hypochlorite (NaOCl) to stimulate germination and produce axenic cultures. However, we observed that some spores were killed even with a 2-min NaOCl treatment, the shortest feasible duration. Decreasing the NaOCl concentration in our study from 0.4% to 0.3 and 0.2%, respectively, increased Teliospore germination, but treatment times longer than 2 min still progressively reduced the germination percentages. In testing alternative met hods, we found “acidic electrolyzed water” (AEW), generated by electrolysis of a weak solution of sodium chloride, also surface-sterilized and increased the rate of T. indica Teliospore germination. In a representative experiment comparing the two methods, NaOCl (0.4%) for 2 min and AEW for 30 min increased germination from 19% (control) to 41 and 54%, respectively, by 7 days after treatment. Because Teliospores can be treated with AEW for up to 2 h with little, if any, loss of viability, compared with 1 to 2 min for NaOCl, treatment with AEW has certain advantages over NaOCl for surface sterilizing and increasing germination of Teliospores of suspect T. indica.
James W Kronstad - One of the best experts on this subject based on the ideXlab platform.
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the hgl1 gene is required for dimorphism and Teliospore formation in the fungal pathogen ustilago maydis
Molecular Microbiology, 2001Co-Authors: Franz Durrenberger, R D Laidlaw, James W KronstadAbstract:The fungal pathogen Ustilago maydis causes a dramatic disease in maize involving the induction of tumours and the formation of masses of black Teliospores. In this fungus, mating between haploid, budding cells results in the formation of the infectious, filamentous cell type that invades host tissue. Mating and filamentous growth are governed by the mating-type loci and by cAMP signalling, perhaps in response to signals from maize. To dissect the involvement of cAMP signalling further, the constitutive filamentous phenotype of a mutant with a defect in the catalytic subunit of protein kinase A was used to isolate suppressor mutations that restore budding growth. One such mutation identified the hgl1 gene, which is shown to be required for both the switch between budding and filamentous growth and Teliospore formation during infection. In addition, the hgl1 gene product may be a target of phosphorylation by protein kinase A, and transcript levels for the gene are elevated during mating. Thus, the hgl1 gene provides a connection between mating, cAMP signalling and two important aspects of virulence: filamentous growth and the formation of Teliospores.
R. Bauer - One of the best experts on this subject based on the ideXlab platform.
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Teliospores of smut fungi Teliospore walls and the development of ornamentation studied by electron microscopy
Protoplasma, 1998Co-Authors: M. Piepenbring, R. Bauer, Franz OberwinklerAbstract:The walls of mature Teliospores and the development of ornamentation, as seen by transmission electron microscopy, are described for 37 genera of smut fungi, based on observations of ca. 120 species and on literature. Structural diversity of mature Teliospore walls is due to differences in spore wall layers forming the spore wall (endosporium, middle layer, exosporium, ornamentation) and to different elements forming the ornamentation (exosporium, ornaments, sheath, hyphal wall, adjacent fungal cells, material of the host). During teliosporogenesis the outer layers are usually deposited first. At the beginning of the formation of the ornamentation the plasma membrane may be smooth or undulated carrying the developing ornaments on its tips or in its depressions. The ornamentation of some genera appears similar when seen by scanning electron microscopy, but can be the product of different developmental patterns (e.g., warts of species of Farysia, Tilletia , and Ustilago ), however, warty and reticulate ornamentation can both be produced by similar developmental processes (shown, e.g., for species of Cintractia and Tilletia ). Typical structures of the mature Teliospore wall and developmental patterns based on homologous similarities are described for the following groups of genera or species: Macalpinomyces, Melanopsichium, Sporisorium , and Ustilago infecting members of the family Poaceae; Kuntzeomyces, Testicularia , and Trichocintractia; Anthracoidea, Cintractia, Heterotolyposporium piluliforme , and Tolyposporium junci; Glomosporium, Sorosporium , and Thecaphora; Conidiosporomyces, Erratomyces, Ingoldiomyces, Neovossia, Oberwinkleria , and Tilletia; Entyloma , and genera of the Doassansia group; Liroa, Microbotryum, Sphacelotheca, Ustilago infecting dicotyledons, and Zundeliomyces; Aurantiosporium, Fulvisporium , and Ustilentyloma . Special characteristics of the Teliospore wall were observed for the genera Dermatosorus, Doassinga, Entorrhha, Farysia, Mycosyrinx, Rhamphospora , and some species of Tolyposporium .
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Teliospores of smut fungi general aspects of Teliospore walls and sporogenesis
Protoplasma, 1998Co-Authors: R. BauerAbstract:The concept and nomenclature for the elements of Teliospore walls in smut fungi are presented and a survey of teliosporogenesis is given, as seen by light and transmission electron microscopy. Four developmental types are distinguished: the Ustilago, Microbotryum, Tilletia, and Entorrhiza type. In the Ustilago type, sporogenous hyphae are completely segmented into Teliospore initials which are embedded in a hyaline matrix formed by gelatinised hyphal walls (found in species of Anthracoidea, Cintractia, Heterotolyposporium, Kuntzeomyces, Macalpinomyces, Melanopsichium, Sporisorium, Testicularia, Tolyposporium junci, Trichocintractia , and species of Ustilago infecting members of the family Poaceae). In the Microbotryum type, septate sporogenous hyphae are also completely segmented into Teliospore initials, however, they are not surrounded by a hyaline matrix ( Microbotryum, Sphacelotheca, Ustilago spp. infecting dicotyledons). A yeast-like budding of teliosporogenic cells is observed for some species of Microbotryum, Sphacelotheca , and Ustilago infecting dicotyledons. In the Tilletia type, Teliospores differentiate locally in the sporogenous hyphae, in an apical or intercalary position, without a hyaline matrix ( Conidiosporomyces, Doassinga, Entyloma, Erratomyces, Ingoldiomyces, Neovossia, Oberwinkleria, Rhamphospora, Tilletia ). In all these types, the Teliospore initials first develop a hyaline sheath under which the ornamentation, the exosporium, sometimes a middle layer, and the endosporium are successively deposited by the fungal cell. In the Entorrhiza type, the Teliospores develop inside vital host cells with the wall of the sporogenous hypha included into the Teliospore wall. The fungus develops a middle layer and an electron-transparent endosporium inside the hyphal wall while a layer forming the ornamentation is deposited onto the hyphal wall, probably by vesicles of dictyosomes of the host cell.
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Teliospores of smut fungi Teliospore connections, appendages, and germ pores studied by electron microscopy; phylogenetic discussion of characteristics of Teliospores
Protoplasma, 1998Co-Authors: M. Piepenbring, R. Bauer, Franz OberwinklerAbstract:Special features of Teliospores in smut fungi are described, including Teliospore connections, appendages, and germ pores. Balls of Teliospores in species of many different genera cohere by remnants of hyphal walls, sheaths, and sometimes interlocking ornamentation. Teliospores are connected in pairs in species of Mycosyrinx and Geminago by special local structures. Appendages can be formed locally by persistent material from the sheath ( Cintractia, Anthracoidea, Sphacelotheca ), thickened parts of the spore wall (e.g., Georgefischeria, Jamesdicksonia, Rhamphospora, Tolyposporella ), or persistent walls of sporogenous hyphae ( Rhamphospora , genera of the Tilletia relationship). Species of Georgefischeria, Jamesdicksonia , and Tolyposporella have Teliospore walls composed of more than three layers of different electron density. “Germ areas” corresponding to thinner parts of the spore wall are known, e.g., for species of Anthracoidea, Cintractia , and Ustilago infecting members of the family Poaceae, while distinct germ pores, one per Teliospore, are found in some species of Thecaphora, “Tolyposporium” , and Sporisorium . Teliospores of Mycosyrinx cissi have a germination ring. Characteristics of Teliospores are used to discuss the phylogeny of smut fungi. A phylogenetic tree in accordance with Teliospore characteristics is compared to those obtained from ultrastructural characteristics of host-parasite interaction, of septal pores, and from sequence data. Aspects of Teliospore development help to define taxa at a high systematic level (Entorrhizales, Ustilaginales, Tilletiales/Entylomatales, Microbotryaceae), while details of ornamentation ontogeny delimit groups of genera (e.g., genera related to Ustilago on members of the Poaceae and Sporisorium, Cintractia and Anthracoidea, Tilletia ) or single genera (e.g., Melanopsichium, Dermatosorus, Mycosyrinx, Doassinga, Rhamphospora ). Types of ornamentation (warty, reticulate), middle layers, Teliospore balls, and germ pores evolved repeatedly by convergence. The smut Teliospore itself probably evolved independently at least twice, or perhaps three (or more) times, in the Microbotryales, in the Entorrhizales, and in a common ancestor of the remainder of the Ustilaginomycetes.
