Algal Lichens Research Articles

Genome Analysis of Two Lichen Bacteriobionts, <i>Lichenibacterium</i> <i>ramalinae</i> and <i>Lichenibacterium minor</i>: Factors of Virulence and Adaptation

Successful invasion and adaptation of bacteria into lichen symbiosis requires the involvement of a number of physiological mechanisms that regulate interactions between bacteriobionts, mycobionts and algobionts and ensure the optimization of thallus growth and development. The genomes of two lichen bacteriobionts Lichenibacterium ramalinae and L. minor were analysed to identify those factors. No nodulation genes traditional for rhizobia were found. TIGR02302 protein has been suggested to have a similar function. Genes encoding proteins associated with the implementation of planktonic and biofilm phenotypes – flagellins and pilins – have been found. These genes have a high level of similarity with the genes of plant symbiont bacteria. Genetic determinants of virulence factors such as locus B invasion proteins, Integration host factor (IHF), sensory histidine kinases, siderophore receptor and transport proteins and monooxygenase of antibiotic synthesis (Abm) were found in the genomes of the studied bacteria. The listed genes have similarities with those in organisms forming associations of different degrees of association with plants. Based on the data obtained, the assumption of a priority association of bacteria of the Lichenibacteriaceae family with green lichen algae was suggested.

Chemical properties of key metabolites determine the global distribution of lichens

In lichen symbioses, fungal secondary metabolites provide UV protection on which lichen algae such as trebouxiophycean green algae-the most prominent group of photobionts in lichen symbioses-sensitively depend. These metabolites differ in their UV absorbance capability and solvability, and thus vary in their propensity of being leached from the lichen body in humid and warm environments, with still unknown implications for the global distribution of lichens. In this study covering more than 10,000 lichenised fungal species, we show that the occurrence of fungal-derived metabolites in combination with their UV absorbance capability and their probability of being leached in warm and humid environments are important eco-evolutionary drivers of global lichen distribution. Fungal-derived UV protection seems to represent an indirect environmental adaptation in which the lichen fungus invests to protect the trebouxiophycean photobiont from high UV radiation in warm and humid climates and, by doing this, secures its carbon source.

Lichen algae: the photosynthetic partners in lichen symbioses

AbstractA review of algal (including cyanobacterial) symbionts associated with lichen-forming fungi is presented. General aspects of their biology relevant to lichen symbioses are summarized. The genera of algae currently believed to include lichen symbionts are outlined; approximately 50 can be recognized at present. References reporting algal taxa in lichen symbiosis are tabulated, with emphasis on those published since the 1988 review by Tschermak-Woess, and particularly those providing molecular evidence for their identifications. This review is dedicated in honour of Austrian phycologist Elisabeth Tschermak-Woess (1917–2001), for her numerous and significant contributions to our knowledge of lichen algae (some published under the names Elisabeth Tschermak and Liesl Tschermak).

Survey of the occurrence of desiccation-induced quenching of basal fluorescence in 28 species of green microalgae.

Desiccation-induced chlorophyll fluorescence quenching seems to be an indispensable part of desiccation resistance in the surveyed 28 green microalgal species. Lichens are desiccation tolerant meta-organisms. In the desiccated state photosynthesis is inhibited rendering the photobionts potentially sensitive to photoinhibition. As a photoprotective mechanism, strong non-radiative dissipation of absorbed light leading to quenching of chlorophyll fluorescence has been proposed. Desiccation-induced quenching affects not only variable fluorescence, but also the so-called basal fluorescence, F0. This phenomenon is well-known for intact lichens and some free living aero-terrestrial algae, but it was often absent in isolated lichen algae. Therefore, a thorough screening for the appearance of desiccation-induced quenching was undertaken with 13 different aero-terrestrial microalgal species and lichen photobionts. They were compared with 15 aquatic green microalgal species, among them also three marine species. We asked the following questions: Do isolated lichen algae show desiccation-induced quenching? Are aero-terrestrial algae different in this respect to aquatic algae and is the potential for desiccation-induced quenching coupled to desiccation tolerance? How variable is desiccation-induced quenching among species? Most of the aero-terrestrial algae, including all lichen photobionts, showed desiccation-induced quenching, although highly variable in extent, whereas most of the aquatic algae did not. All algae displaying quenching were also desiccation tolerant, whereas all algae unable to perform desiccation-induced quenching were desiccation intolerant. Desiccation-induced fluorescence quenching seems to be an indispensable part of desiccation resistance in the investigated species.

Origin and evolution of chloroplast group I introns in lichen algae.

The history of group I introns is characterized by repeated horizontal transfers, even among phylogenetically distant species. The symbiogenetic thalli of lichens are good candidates for the horizontal transfer of genetic material among distantly related organisms, such as fungi and green algae. The main goal of this study was to determine whether there were different trends in intron distribution and properties among Chlorophyte algae based on their phylogenetic relationships and living conditions. Therefore, we investigated the occurrence, distribution and properties of group I introns within the chloroplast LSU rDNA in 87 Chlorophyte algae including lichen and free-living Trebouxiophyceae compared to free-living non-Trebouxiophyceae species. Overall, our findings showed that there was high diversity of group I introns and homing endonucleases (HEs) between Trebouxiophyceae and non-Trebouxiophyceae Chlorophyte algae, with divergence in their distribution patterns, frequencies and properties. However, the differences between lichen Trebouxiophyceae and free-living Trebouxiophyceae were smaller. An exception was the cL2449 intron, which was closely related to ω elements in yeasts. Such introns seem to occur more frequently in lichen Trebouxiophyceae compared to free-living Trebouxiophyceae. Our data suggest that lichenization and maintenance of lichen symbiosis for millions of years of evolution may have facilitated horizontal transfers of specific introns/HEs between symbionts. The data also suggest that sequencing of more chloroplast genes harboring group I introns in diverse algal groups may help us to understand the group I intron/HE transmission process within these organisms.

Algal carbohydrates affect polyketide synthesis of the lichen-forming fungus Cladonia rangiferina

Lichen secondary metabolites (polyketides) are produced by the fungal partner, but the role of algal carbohydrates in polyketide biosynthesis is not clear. This study examined whether the type and concentration of algal carbohydrate explained differences in polyketide production and gene transcription by a lichen fungus (Cladonia rangiferina). The carbohydrates identified from a free-living cyanobacterium (Spirulina platensis; glucose), a lichen-forming alga (Diplosphaera chodatii; sorbitol) and the lichen alga that associates with C. rangiferina (Asterochloris sp.; ribitol) were used in each of 1%, 5% and 10% concentrations to enrich malt yeast extract media for culturing the mycobiont. Polyketides were determined by high performance liquid chromatography (HPLC), and polyketide synthase (PKS) gene transcription was measured by quantitative PCR of the ketosynthase domain of four PKS genes. The lower concentrations of carbohydrates induced the PKS gene expression where ribitol up-regulated CrPKS1 and CrPKS16 gene transcription and sorbitol up-regulated CrPKS3 and CrPKS7 gene transcription. The HPLC results revealed that lower concentrations of carbon sources increased polyketide production for three carbohydrates. One polyketide from the natural lichen thallus (fumarprotocetraric acid) also was produced by the fungal culture in ribitol supplemented media only. This study provides a better understanding of the role of the type and concentration of the carbon source in fungal polyketide biosynthesis in the lichen Cladonia rangiferina.

Identification of Phymatolithon lamii (Me. Lemoine) Y.M. Chamberlain 1991 following bleaching in the upper intertidal zone of the Minas Basin, Nova Scotia, Canada

Objective: To identify an unusual encrusting calcareous organism found growing on upper intertidal rocks along the coast of the Minas Basin, Nova Scotia, Canada, which was thought to potentially represent a new, perhaps invasive species to the area. Methods: Observations were made of the organism growing in situ and of intact and dissected specimens under microscopes in the laboratory. Taxonomic keys were used to determine probable identity of the organism as a tunicate, sponge, bryozoan, marine lichen, or coralline red alga. These keys were then further used to identify samples to species. Results: The organism was determined to be Phymatolithon lamii (Me. Lemoine) Y.M. Chamberlain 1991, an encrusting coralline red alga native to the study region. This species normally occurs in the subtidal zone, so its occurrence in the high intertidal zone was unusual and resulted in the alga becoming bleached and taking on an unusual appearance. Conclusions: The sample did not represent a new or invasive species to the study area. However, occurrence of this species on the high intertidal zone was novel and may have resulted from unusual climatic and/or oceanographic conditions in the study area in 2015 that allowed this alga to settle and grow in the intertidal zone for a time before death and bleaching occurred. Events such as this resulting from climate change may negatively impact recruitment to populations of coastal marine species and should be investigated further.

Effects of short-term low temperature stress on chlorophyll fluorescence transients in Antarctic lichen species

Chlorophyll fluorescence is an effective tool for investigating characteristics of any photosynthesizing organisms and its responses due to different stressors. Here, we have studied a short-term temperature response on two Antarctic green algal lichen species: Umbilicaria antarctica, and Physconia muscigena. We measured slow chlorophyll fluorescence transients in the species during slow a cooling of thallus temperature from 20°C to 5°C with a 10 min. acclimation at each temperature in dark. The measurements were supplemented with saturation pulses for the analysis of chlorophyll fluorescence parameters: maximum yield of PS II photochemistry (FV/FM), effective quantum yield of PS II photochemistry (FPSII) and non-photochemical quenching (NPQ). In response to decreasing thallus temperature, we observed species-specific changes in chlorophyll fluorescence levels P, S, M, T reached during chlorophyll fluorescence transient as well as in the shape of the chlorophyll fluorescence transients. With a decrease in temperature, the time at which M and T chlorophyll fluorescence levels were reached, increased. These changes were attributed to redox state of plastoquinon pool, changes in Calvin-Benson cycle activity, non-photochemical quenching components, state transition in particular. In this study, we present some chlorophyll fluorescence ratios (P/M, M/T, P/T) and chlorophyll fluorescence increase rates (FR1, i.e. O to P, and FR2 - i.e. S to M) as the parameters reflecting direct temperature effects on chloroplastic apparatus of lichen alga sensitively. We proposed that species-specific changes in the slow phase of chlorophyll fluorescence transients could be potentially used as indicators of low temperature effects in photosynthetic apparatus of lichen algal photobionts. Interspecific differences in response to low temperature might be evaluated using the approach as well.

Lichen photobionts of the rocky outcrops of Karadag massif (Crimean Peninsula)

Despite the fact that lichen algae have been explored for over100 years, their diversity, taxonomy and ecology still remains understudied. In present study we analyzed 114 specimens representing 72 lichen taxa collected on the Crimean Peninsula, culturing and sequencing their algal partners. In total we identified 26 taxa of photobionts representing 10 algal genera (Asterochloris, Chloroidium, Dilabifilum, Diplosphaera, Gloeocapsa, Myrmecia, Nostoc, Printzina, Stichococcus, Trebouxia). As most of the studied lichens (more than 80 % of studied lichen species) contained Trebouxia, for which morphology-based identification is often difficult and insufficient, we additionally used molecular methods. According to the molecular phylogeny of internal transcribed spacer (ITS) rDNA, Trebouxia species were related to three main clades: “Arboricola” (88.9 % of the investigated Trebouxia-containing lichens), “Impressa” (4.4 %) and “Simplex” (6.7 %). Based on results of molecular phylogeny and morphology three new species of Trebouxia, namely T. solaris, T. vagua and T. cretacea were described. In several lichens, two coexisting Trebouxia species were revealed in the same thallus.

Comparison of Lichen-Forming Cyanobacterial and Green Algal Photobionts with Free-Living Algae

Cyanobacteria of the genus Nostoc and green algae in the Chlorophyceae are widespread in nature and may occur in symbiotic associations with lichen-forming ascomycetes or as free-living cyanobacteria. Recent findings for some groups of lichens suggest that special lichen-forming photobiont lineages may exist independent of the free-living lineages, but few comparisons on photobiont growth, pigment, and polysaccharide production have been made. The goal of this study was to isolate photobionts, confirm their identity, and characterize their growth, pigment and polysaccharide production relative to free-living lineages. Algal growth, pigment contents, and polysaccharide concentration was measured using standard methods. The identification of Nostoc species was determined using transfer RNA for Leucine (trnL) nucleotide sequences and green algae using the internal transcribed spacer 1 of ribosomal DNA (ITS rDNA) sequences. An additional heptanucleotide repeat was present in the trnL gene of the Nostoc strain that associates with Leptogium rivulare. The biomass of pigment and polysaccharide production was highest in the lichenized Diplosphaera chodatii but the specific growth rate was highest in the free-living green alga, Chlorella vulgaris. The specific growth of the free-living Nostoc was higher than the lichenized Nostoc but pigment production was similar and polysaccharide production was lower than some of the lichenized Nostoc isolates. It was further hypothesized that the rates of growth, polysaccharide and pigment production may be key factors in compatibility of lichen algae with the fungus.

Production of UV-B screens and changes in photosynthetic efficiency in Antarctic Nostoc commune colonies and a lichen Xanthoria elegans depend on a dose and duration of UV-B stress

The survival of non-vascular autotrophs in the extreme polar conditions and the principles of their high tolerance to extreme physical factors have been intriguing scientists in last decades. Therefore, this study focuses on the capacity of production of UV-B screening pigments in two model Antarctic species, one algal lichen, and colony of a cyanobacterium. Dose-dependent activation of protective mechanisms of Antarctic cyanobacterium (Nostoc commune) and algal lichen (Xanthoria elegans), synthesis of UV-B screening compounds in particular, were studied together with the changes in photosynthetic efficiency induced by a background photosynthetically active radiation (PAR) supplemented with UV-B radiation. The samples were exposed to different doses of UV-B (280–320 nm), low (0.7 W m-2), medium (1.5 W m-2) and high (3.0 W m-2) for 5 days. Untreated samples (control) were shielded from UV-B radiation during experiment. Chlorophyll fluorescence parameters and secondary UV-B protective metabolites were analysed in the intervals of 24 h, 48 h and 120 h. Amount of UV-B screening pigments was measured spectrophotometrically using several specific wavelengths in UV-B absorption range. Results showed that if exposed to a low dose of UV-B radiation or a short-term treatment, both species exhibited an increase in UV-B screening pigments to protect the lichen photobiont against UV-B damage. However, if exposed to a high dose of UV-B radiation or a long-term treatment, a decrease of UV-B screening compounds occured. This implies that Antarctic lichen and cyanobacterium can protect themselves against an increase of stress factors ranging within physiological limits, like e.g. increased synthesis of UV-B screening compounds thanks to a thinning of the ozone layer and consequent increase in UV radiation doses incident on Antarctic terrestrial ecosystems. Nevertheless, the likely increased UV-B radiation due to more intense depletion of stratospheric ozone layer may lead to alterations in UV-B tolerance in Antarctic lichens in future.

Evidence of ancient horizontal gene transfer between fungi and the terrestrial alga Trebouxia

Intimate and long-lasting relationships of fungi and algae have been known for centuries by scientists, and these ancient symbioses might have provided excellent opportunities for horizontal gene transfer (HGT) of protein encoding genes between the two organismal partners. In this study, we sequenced and assembled 451 Mbp of novel genomic DNA from Trebouxia decolorans (Trebouxiaceae, Chlorophyta), the green algal photobiont of the lichen Xanthoria parietina (Teloschistaceae, Lecanoromycetes, Ascomycota). This alga also occurs as a free-living terrestrial organism. The aim of our work was to search for candidate genes pointing to HGT between lichenized fungi and lichen algae. We found evidence for three putative HGT events of fungal genes into the Trebouxia genome, but these are likely more ancient (over 600 mya) than the origin of lichenization within the fungal Ascomycetes. The three transferred genes are part of gene groups that in other species encode a tellurite-resistance dicarboxylate transporter (TDT) family protein, a class-1 nitrilase/cyanide hydratase (CH), and an oxidoreductase/retinol dehydrogenase. In each case, our phylogenomic analyses show orthologs from Trebouxia as sister to orthologs from all fungi or basally placed within Ascomycetes, while the orthologs from green algae and land plants form separate, independent evolutionary lineages. Alternative hypothesis tests significantly support these HGT events. The presence of these genes in Trebouxia was validated by PCR amplification of separately isolated Trebouxia DNA. The ancient incorporation of fungal genes in the genomes of these particular green algae are intriguing and could be early evidence for symbiotic and co-evolutionary relationships among the major eukaryotic branches of algae and fungi present in early terrestrial life on Earth. These genes could have played a pre-disposition role for some fungi and algae in the origin of lichen symbiosis, but further studies are needed to evaluate this in detail.

Photobiont selectivity leads to ecological tolerance and evolutionary divergence in a polymorphic complex of lichenized fungi

The integrity and evolution of lichen symbioses depend on a fine-tuned combination of algal and fungal genotypes. Geographically widespread species complexes of lichenized fungi can occur in habitats with slightly varying ecological conditions, and it remains unclear how this variation correlates with symbiont selectivity patterns in lichens. In an attempt to address this question, >300 samples were taken of the globally distributed and ecologically variable lichen-forming species complex Tephromela atra, together with closely allied species, in order to study genetic diversity and the selectivity patterns of their photobionts. Lichen thalli of T. atra and of closely related species T. grumosa, T. nashii and T. atrocaesia were collected from six continents, across 24 countries and 62 localities representing a wide range of habitats. Analyses of genetic diversity and phylogenetic relationships were carried out both for photobionts amplified directly from the lichen thalli and from those isolated in axenic cultures. Morphological and anatomical traits were studied with light and transmission electron microscopy in the isolated algal strains. Tephromela fungal species were found to associate with 12 lineages of Trebouxia. Five new clades demonstrate the still-unrecognized genetic diversity of lichen algae. Culturable, undescribed lineages were also characterized by phenotypic traits. Strong selectivity of the mycobionts for the photobionts was observed in six monophyletic Tephromela clades. Seven Trebouxia lineages were detected in the poorly resolved lineage T. atra sensu lato, where co-occurrence of multiple photobiont lineages in single thalli was repeatedly observed. Low selectivity apparently allows widespread lichen-forming fungi to establish successful symbioses with locally adapted photobionts in a broader range of habitats. This flexibility might correlate with both lower phylogenetic resolution and evolutionary divergence in species complexes of crustose lichen-forming fungi.

The earliest records of internally stratified cyanobacterial and algal lichens from the Lower Devonian of the Welsh Borderland

Lichenization is assumed to be a very ancient mode of fungal nutrition, but fossil records are rare. Here we describe two fragments of exceptionally preserved, probably charred, lichen thalli with internal stratification. Cyanolichenomycites devonicus has a cyanobacterial and Chlorolichenomycites salopensis a unicellular, presumably green algal photobiont. Fruiting bodies are missing. Cyanolichenomycites devonicus forms asexual spores in a pycnidium. All specimens were examined with scanning electron microscopy techniques. The fossils were extracted by maceration. Extant lichens and free-living cyanobacteria were either experimentally charcoalified for comparison or conventionally prepared. Based on their septate hyphal structure, both specimens are tentatively interpreted as representatives of the Pezizomycotina (Ascomycota). Their presence in 415 million yr (Myr) old rocks from the Welsh Borderland predates existing Late Cretaceous records of pycnidial conidiomata by some 325 Myr and Triassic records of lichens with broadly similar organization by some 195 Myr. These fossils represent the oldest known record of lichens with symbionts and anatomy as typically found in morphologically advanced taxa today. The latter does not apply to Winfrenatia reticulata, the enigmatic crustose lichen fossil from the Lower Devonian, nor to presumed lichen-like organisms such as the Cambrian Farghera robusta or to the Lower Devonian Spongiophyton minutissimum.

Desiccation-induced non-radiative dissipation in isolated green lichen algae

Lichens are able to tolerate almost complete desiccation and can quickly resume metabolic activity after rehydration. In the desiccated state, photosynthesis is completely blocked and absorbed excitation energy cannot be used for electron transport, leading to a potential strong vulnerability for high light damage. Although desiccation and high insolation often occur simultaneously and many lichens colonize exposed habitats, these organisms show surprisingly little photodamage. In the desiccated state, variable chlorophyll fluorescence is lost, indicating a suspension of charge separation in photosystem II. At the same time, basal fluorescence (F (0)) is strongly quenched, which has been interpreted as an indication for high photoprotective non-radiative dissipation (NRD) of absorbed excitation energy. In an attempt to provide evidence for a photoprotective function of NRD in the desiccated state, isolated green lichen algae of the species Coccomyxa sp. and Trebouxia asymmetrica were used as experimental system. In contrast to experiments with intact lichens this system provided high reproducibility of the data without major optical artifacts on desiccation. The presence of 5mM trehalose during desiccation had no effect but culture of the algae in seawater enhanced F (0) quenching in T. asymmetrica together with a reduced depression of F (V)/F (M) after high light treatment. While this effect could not be induced using artificial seawater medium lacking trace elements, the addition of ZnCl(2) and NaI in small amounts to the normal growth medium led to qualitatively and quantitatively identical results as with pure seawater. It is concluded that NRD indicated by F (0) quenching is photoprotective. The formation of NRD in lichen algae is apparently partially dependent on the presence of specific micronutrients.

Recent literature on lichens—224

Alpsoy, L., A. Asian, E. Kotan, G. Agar & M. Anar. 2011. Protective role of two lichens in human lymphocytes in vitro. Fresenius Environmental Bulletin 20(7): 1661–1666. Aptroot, A. 2011. Mossen en korstmossen op aangevoerde olijfbomen en geimporteerde stenige substraten [Mosses and lichens on introduced olive trees with notes on further introduced mosses]. Buxbaumiella 90: 31–37. [In Dutch with English abstract.] Armaleo, D., F. Lutzoni, F. Collart, S. Baker, J. Magnuson, O. Andresson, D. Auberry, D. Culley, F. Dietrich, I. Grigoriev, B. Hodkinson, S. Karpowicz, A. Kuo, P. Larsen, F. Martin, T. McDonald, S. Merchant, E. Morin, O. Mueller, E. Panisko, M. V. Sanchez, I. Small & B. B. Xavier. 2011. Systems Biology of Lichen Systems: Pure Cultures of Cladonia grayi and Lichen-Dominated Biological Soil Crusts. Pages 158–159. In DOE Genomic Science: Systems Biology for Energy and Environment. Oak Ridge National Lab, Oak Ridge, TN. ———, X. Sun & C. Culberson. 2011. Insights from the first putative biosynthetic gene cluster for a lichen depside and depsidone. Mycologia 103(4): 741–754. Armstrong, R. A. 2010. Lobe formation and division in the foliose lichen Xanthoparmelia conspersa. Symbiosis 51(3): 227–232. Arslan, B., S. Ozturk & S. Oran. 2011. Lecanora, Phaeophyscia and Rinodina species new to Turkey. Mycotaxon 116: 49–52. Asplund, J. 2011. Chemical races of Lobaria pulmonaria differ in palatability to gastropods. The Lichenologist 43(5): 491–494. ———. 2011. Snails avoid the medulla of Lobaria pulmonaria and L. scrobiculata due to presence of secondary compounds. Fungal Ecology 4(5): 356–358. Aydin, E. & H. Turkez. 2011. Antioxidant and genotoxicity screening of aqueous extracts of four lichens collected from north east Anatolia. Fresenius Environmental Bulletin 20(8A): 2085–2091. Azizuddin & M. I. Choudhary. 2011. Radical scavenging and lipoxygenase inhibition studies of the compounds isolated from a medicinal lichen, Usnea longissima. Chemistry of Natural Compounds 47(3): 481–484. Ballantyne, K. & E. Nol. 2011. Nesting habitat selection and hatching success of whimbrels near Churchill, Manitoba, Canada. Waterbirds 34(2): 151–159. Beeching, S. Q. 2009. ‘I Like You but What Can You Do, Can You be a Bird?’: Adventures in the Lichen Trade. Public Domain, Inc., Atlanta, GA. 185 pp. ———. 2011. The Licheniad. Squamules Unlimited, New York. 28 verses. [Available at http://squamules.blogspot.com/ 2011/02/licheniad.html.] ———. 2011. Three rhymes on the lichen symbiosis occasioned by a lichen video’s narrator having said that the lichen fungus eats the lichen algae. Squamules Unlimited, New York. 3 verses. [Available at http://squamules. blogspot.com/2011/01/poemules.html.] 1 Corresponding author: brendan.hodkinson@gmail.com The cumulative database for this series is available in searchable form on the World Wide Web at http:// www.nhm.uio.no/botanisk/lav/RLL/RLL.HTM; for more recent entries, full abstracts of electronically available articles will be databased along with the information published here. We owe thanks to Einar Timdal for his work on the RLL database, to James Lendemer for proofreading this manuscript, to Bill Buck for alerting us to some of the more obscure sources, to Jim Bennett for sharing his Scopus alerts, and to the cooperating authors who send reprints or electronic versions of their works to us for inclusion. DOI: 10.1639/007-2745-115.1.183

Ascomycete fungi on dimension stone of the “Burg Gleichen”, Thuringia

In the present study, the diversity of ascomycete fungi was investigated on two wall areas of the “Burg Gleichen”, Thuringia (Germany), made of various types of sandstones, travertine and Grenzdolomit. From a W-exposed, shaded wall area, free-living ascomycetes (mainly “black fungi”) and green algae could be retrieved from sandstone lithologies. Sandstone from an ESE-exposed area was mainly colonized by lichen ascomycetes and the lichen alga Trebouxia. Both areas share a small number of generalist species, related to the ascomycete black fungi Sarcinomyces petricola, Phaeococcomyces chersonesos and Stichococcus mirabilis. Free-living black fungi were isolated and characterized with respect to cell wall morphology and melanin content. A remarkably rigid melanin layer, incorporated in the cell wall of a Cladosporium isolate is presented in detail.

Evolutionary inferences based on ITS rDNA and actin sequences reveal extensive diversity of the common lichen alga Asterochloris (Trebouxiophyceae, Chlorophyta)

The genus Asterochloris is one of the most common lichen photobionts. We present a molecular investigation of 41 cultured strains, for which nuclear-encoded ITS rDNA and partial actin I sequences were determined. The loci studied revealed considerable differences in their evolutionary dynamics as well as sequence variation. As compared to ITS data, the actin sequences show much greater variation, and the phylogenies yield strong resolution and support of many internal branches. The partitioning of ITS dataset into several regions yielded better node resolution. We recognized 16 well-supported monophyletic lineages, of which one represents the type species of the genus (Asterochloris phycobiontica), and six correspond to species previously classified to the genus Trebouxia (T. erici, T. excentrica, T. glomerata, T. irregularis, T. italiana and T. magna). Only 15% of isolated photobionts considered in our study could be assigned with certainty to previously described species, emphasizing amazing cryptic variability in Asterochloris. Concurrently with the formal delimitation of the genus Asterochloris, we propose new combinations for the former Trebouxia species; furthermore, T. pyriformis is reduced to a synonym of A. glomerata. The present knowledge of global diversity of Asterochloris algae is discussed.

Erratum to: Dehydration rate and time of desiccation affect recovery of the lichen alga Trebouxia erici: alternative and classical protective mechanisms

Erratum to: Dehydration rate and time of desiccation affect recovery of the lichen alga Trebouxia erici: alternative and classical protective mechanisms

Evolution of ITS Ribosomal RNA Secondary Structures in Fungal and Algal Symbionts of Selected Species of Cladonia sect. Cladonia (Cladoniaceae, Ascomycotina)

Evolutionary studies in lichen associations follow that of the fungal symbiont (mycobiont), which is the symbiont after which the lichen is named and forms the majority of the thallus. However, evolution of the algal partner (photobiont) is important to maintain compatibility between symbionts and to optimize productivity of the lichen association. The internal transcribed spacer (ITS) regions of the nuclear ribosomal DNA (rDNA) were examined for primary DNA sequence patterns and for patterns in the secondary structure of the rRNA transcripts in both symbionts of the genus Cladonia. Fungal and algal symbionts show opposite trends in rates of evolution and fragment lengths. Both symbionts showed stronger conservation of the ITS2 structure than the ITS1 structure. Homology was evident in the secondary structures between the two highly divergent chlorophyte and ascomycete taxonomic groups. Most fungal species and all species complexes were polyphyletic. The ITS rDNA of the natural lichen algae from Manitoba and four known algal species is highly similar. The natural lichen algae segregate into highly supported clades by environmental features, suggesting that algae that are already adapted to the environment may associate with germinating fungal propagules in the genus Cladonia. Fungal plasticity may allow the mycobiont to adapt to the environment of the photobiont producing variation in lichen morphology. This might explain the incongruence of phylogenetic patterns between the algal and fungal partners tested and the polyphyly of the fungal species.