Lichen Photobionts Research Articles

Secondary Metabolites from Australian Lichens Ramalina celastri and Stereocaulon ramulosum Affect Growth and Metabolism of Photobiont Asterochloris erici through Allelopathy.

In the present work, the phytotoxic effects of secondary metabolites extracted from lichen Ramalina celastri (usnic acid) and lichen Stereocaulon ramulosum (a naturally occurring mixture of atranorin and perlatolic acid, approx. 3:1) on cultures of the aposymbiotically grown lichen photobiont Asterochloris erici were evaluated. Algae were cultivated on the surface of glass microfiber disks with applied crystals of lichen extracts for 14 days. The toxicity of each extract was tested at the two selected doses in quantities of 0.01 mg/disk and 0.1 mg/disk. Cytotoxicity of lichen extracts was assessed using selected physiological parameters, such as growth (biomass production) of photobiont cultures, content of soluble proteins, chlorophyll a fluorescence, chlorophyll a integrity, contents of chlorophylls and total carotenoids, hydrogen peroxide, superoxide anion, TBARS, ascorbic acid (AsA), reduced (GSH) and oxidized (GSSG) glutathione, and composition of selected organic acids of the Krebs cycle. The application of both tested metabolic extracts decreased the growth of photobiont cells in a dose-dependent manner; however, a mixture of atranorin and perlatolic acid was more effective when compared to usnic acid at the same dose tested. A higher degree of cytotoxicity of extracts from lichen S. ramulosum when compared to identical doses of extracts from lichen R. celastri was also confirmed by a more pronounced decrease in chlorophyll a fluorescence and chlorophyll a integrity, decreased content of chlorophylls and total carotenoids, increased production of hydrogen peroxide and superoxide anion, peroxidation of membrane lipids (assessed as TBARS), and a strong decrease in non-enzymatic antioxidants such as AsA, GSH, and GSSG. The cytotoxicity of lichen compounds was confirmed by a strong alteration in the composition of selected organic acids included in the Krebs cycle. The increased ratio between pyruvic acid and citric acid was a very sensitive parameter of phytotoxicity of lichen secondary metabolites to the algal partner of symbiosis. Secondary metabolites of lichens are potent allelochemicals and play significant roles in maintaining the balance between mycobionts and photobionts, forming lichen thallus.

Diversity of lichen mycobionts and photobionts and their relationships in the Ny-Ålesund region (Svalbard, High Arctic).

Lichens are dual organisms, with one major mycobiont and one major photobiont in each lichen symbiosis, which can survive extreme environmental conditions in the Arctic. However, the diversity and distribution of lichen photobionts in the Arctic remain poorly understood compared to their mycobiont partners. This study explored the diversity of lichen mycobionts and photobionts in 197 lichen samples collected from the Ny-Ålesund region (Svalbard, High Arctic). The nuclear ribosomal internal transcribed spacer (ITS) regions were sequenced and phylogenetic analyses were performed. The relationships between mycobionts and photobionts, as well as the association patterns, were also investigated. A total of 48 species of lichen mycobionts (16 families, nine orders) and 31 species/lineages of photobionts were identified. These 31 photobiont species belonged to one class (Trebouxiophyceae) and five genera, including 22 species of Trebouxia, five species of Asterochloris, two species of Chloroidium, one species of Symbiochloris, and one species of Coccomyxa. The results indicated that most analyzed lichen mycobionts could associate with multiple photobiont species, and the photobionts also exhibited a similar pattern. The results provided an important reference dataset for characterizing the diversity of lichen mycobionts and photobionts in the High Arctic region.

The symbiotic alga Trebouxia fuels a coherent soil ecosystem on the landscape scale in the Atacama Desert

Biocrusts represent associations of lichens, green algae, cyanobacteria, fungi and other microorganisms, colonizing soils in varying proportions of principally arid biomes. The so-called grit crust represents a recently discovered type of biocrust situated in the Coastal Range of the Atacama Desert (Chile) made of microorganisms growing on and in granitoid pebbles, resulting in a checkerboard pattern visible to the naked eye on the landscape scale. This specific microbiome fulfills a broad range of ecosystem services, all probably driven by fog and dew-induced photosynthetic activity of mainly micro-lichens. To understand its biodiversity and impact, we applied a polyphasic approach on the phototrophic microbiome of this biocrust, combining isolation and characterization of the lichen photobionts, multi-gene phylogeny of the photobionts and mycobionts based on a direct sequencing and microphotography approach, metabarcoding and determination of chlorophylla+b contents. Metabarcoding showed that yet undescribed lichens within the Caliciaceae dominated the biocrust together with Trebouxia as the most abundant eukaryote in all plots. Together with high mean chlorophylla+b contents exceeding 410 mg m−2, this distinguished the symbiotic algae Trebouxia as the main driver of the grit crust ecosystem. The trebouxioid photobionts could be assigned to the I (T. impressa/gelatinosa) and A (T. arboricola) clades and represented several lineages containing five potential species candidates, which were identified based on the unique phylogenetic position, morphological features, and developmental cycles of the corresponding isolates. These results designate the grit crust as the only known coherent soil layer with significant landscape covering impact of at least 440 km2, predominantly ruled by a single symbiotic algal genus.

Isolation and Identification of Lichen Photobionts Collected from Different Environments in North of Portugal and Evaluation of Bioactivities of Their Extracts.

Lichens are organisms constituted by a symbiotic relationship between a fungus (mycobiont) and a photoautotrophic partner (photobiont). Lichens produce several bioactive compounds; however, the biotechnological exploitation of this organism is hampered by its slow growth. To start studying the possibility of exploiting lichens as alternative sources of bioactive compounds, eighteen lichens were collected in the north of Portugal in order to isolate and study the bioactivity of their photobionts. It was possible to isolate and cultivate only eight photobionts. Three of them, LFR1, LFA2 and LCF3, belong to the Coelastrella genus, the other two (LFA1 and LCF1) belong to the Chlorella genus and for the remaining three photobionts, LFS1, LCA1 and LCR1, it was impossible to isolate their microalgae. These only grow in consortium with bacteria and/or cyanobacteria. All extracts showed antioxidant activity, mainly at a concentration of 10 mg.mL-1. LFS1, a consortium extract, showed the highest antioxidant power, as well as the highest concentration of phenolic compounds (5.16 ± 0.53 mg of gallic acid equivalents (GAE).g-1). The extracts under study did not show significant antibacterial activity against Escherichia coli, Listeria or Salmonella. The Coelastrella sp. and LFA1 extracts showed the highest hyaluronidase inhibition. The LFR1 extract at a concentration of 5 mg.mL-1 showed the highest anti-inflammatory activity (79.77 ± 7.66%). The extracts of Coelastrella sp. and LFA1 also showed greater antidiabetic activity, demonstrating the high inhibitory power of α-amylase and α-glucosidase. LFR1 at a concentration of 5 mg.mL-1, due to its selective cytotoxicity inhibiting the growth of cancer cells (Caco-2 cells), is a promising anticancer agent.

A direct PCR approach with low-biomass insert opens new horizons for molecular sciences on cryptogam communities.

Molecular sequence data have transformed research on cryptogams (e.g., lichens, microalgae, fungi, and symbionts thereof) but methods are still strongly hampered by the small size and intermingled growth of the target organisms, poor cultivability and detrimental effects of their secondary metabolites. Here, we aim to showcase examples on which a modified direct PCR approach for diverse aspects of molecular work on environmental samples concerning biocrusts, biofilms, and cryptogams gives new options for the research community. Unlike traditional approaches, this methodology only requires biomass equivalent to colonies and fragments of 0.2 mm in diameter, which can be picked directly from the environmental sample, and includes a quick DNA lysis followed by a standardized PCR cycle that allows co-cycling of various organisms/target regions in the same run. We demonstrate that this modified method can (i) amplify the most widely used taxonomic gene regions and those used for applied and environmental sciences from single colonies and filaments of free-living cyanobacteria, bryophytes, fungi, and lichens, including their mycobionts, chlorobionts, and cyanobionts from both isolates and in situ material during co-cycling; (ii) act as a tool to confirm that the dominant lichen photobiont was isolated from the original sample; and (iii) optionally remove inhibitory secondary lichen substances. Our results represent examples which highlight the method's potential for future applications covering mycology, phycology, biocrusts, and lichenology, in particular.IMPORTANCECyanobacteria, green algae, lichens, and other cryptogams play crucial roles in complex microbial systems such as biological soil crusts of arid biomes or biofilms in caves. Molecular investigations on environmental samples or isolates of these microorganisms are often hampered by their dense aggregation, small size, or metabolism products which complicate DNA extraction and subsequent PCRs. Our work presents various examples of how a direct DNA extraction and PCR method relying on low biomass inserts can overcome these common problems and discusses additional applications of the workflow including adaptations.

Assessing above and belowground recovery from ammonium sulfate addition and wildfire in a lowland heath: mycorrhizal fungi as potential indicators

Atmospheric pollution containing soil‐nitrifying ammonium sulfate ((NH₄)₂SO₄) affects semi‐natural ecosystems worldwide. Long‐term additions of (NH₄)₂SO₄ to nitrogen (N)‐limited habitats, including heathlands, increase climate stress affecting recovery from wildfires. Although heathland vegetation largely depends on ericoid mycorrhizal fungi (ErM) to access soil N, we lack detailed understanding of how prolonged exposure to (NH₄)₂SO₄ may alter ErM community composition and host plants' reliance on fungal partners following wildfire and affect recovery. Simulation of atmospheric pollution ((NH₄)₂SO₄) occurred bi‐weekly for 5 years after a 2006 wildfire in a UK heathland. Ten years after treatments ceased, we measured vegetation structure, lichen and lichen photobiont composition, soil characteristics, ErM colonization, ErM diversity in roots and soil, and assessed ErM potential as novel recovery indicators. Heather height and density, and moss groundcover, were greater in N‐enriched plots. Lichen community indices showed significant treatment effects, but without differences in photobionts. Soil pH and Mg were significantly lower in treated plots while soil cation exchange capacity was significantly higher. There were no detectable differences in ErM composition and keystone ErM taxa between control and treated plots. Soil carbon stock measures were variable. Our results indicate atmospheric pollution following fire can have significant lingering effects above‐ and belowground. ErM diversity and root colonization were not assessed in the original N‐addition experiment; we advocate for their inclusion in future studies as an integral part of the recovery assessment toolkit. We show that mycorrhizal fungi diversity is a viable ecological tool and summarize key steps for ErM identification.

Non-photochemical quenching may contribute to the dominance of the pale mat-forming lichen Cladonia stellaris over the sympatric melanic Cetraria islandica.

The mat-forming fruticose lichens Cladonia stellaris and Cetraria islandica frequently co-occur on soils in sun-exposed boreal, subarctic, and alpine ecosystems. While the dominant reindeer lichen Cladonia lacks a cortex but produces the light-reflecting pale pigment usnic acid on its surface, the common but patchier Cetraria has a firm cortex sealed by the light-absorbing pigment melanin. By measuring reflectance spectra, high-light tolerance, photosynthetic responses, and chlorophyll fluorescence in sympatric populations of these lichens differing in fungal pigments, we aimed to study how they cope with high light while hydrated. Specimens of the two species tolerated high light equally well but with different protective mechanisms. The mycobiont of the melanic species efficiently absorbed excess light, consistent with a lower need for its photobiont to protect itself by non-photochemical quenching (NPQ). By contrast, usnic acid screened light at 450-700nm by reflectance and absorbed shorter wavelengths. The ecorticate usnic species with less efficient fungal light screening exhibited a consistently lower light compensation point and higher CO2 uptake rates than the melanic lichen. In both species, steady state NPQ rapidly increased at increasing light with no signs of light saturation. To compensate for less internal shading causing light fluctuations with a larger amplitude, the usnic lichen photobiont adjusted to changing light by faster induction and faster relaxation of NPQ rapidly transforming excess excitation energy to less damaging heat. The high and flexible NPQ tracking fluctuations in solar radiation probably contributes to the strong dominance of the usnic mat-forming Cladonia in open lichen-dominated heaths.

Allelopathic effects of three lichen secondary metabolites on cultures of aposymbiotically grown lichen photobionts and free-living alga Scenedesmus quadricauda

In the present work, we studied the influence of lichen secondary metabolites (evernic acid, orcinol, usnic acid) on selected species of algae: Trebouxia erici, photobiont of lichen Cladonia cristatella producing typical lichen secondary metabolites, Coccomyxa solorinae-saccatae, photobiont of lichen Solorina saccata which does not produce typical secondary metabolites of lichens, and free-living alga Scenedesmus quadricauda.For the determination of the cytotoxic effects of these compounds, algae were cultivated on the surface of disks composed of glass microfibers, in quantities of 0.1 mg/disk for each metabolite. After 14-days prolonged cultivation, we analysed selected physiological parameters: growth of algae, chlorophyll a fluorescence, the content of ascorbic acid, content of reduced and oxidized glutathione and contents of selected organic acids. The presence of secondary compounds typically inhibited the growth of tested algae and chlorophyll a fluorescence. In the presence of evernic acid and usnic acid, we observed a decrease of ascorbic acid, glutathione, and significant changes in the composition of organic acids.Lichen metabolites, except for their other biological and ecological roles, may act as phytotoxic substances, causing oxidative stress in photobiont cells. These metabolites may play an important role in the “controlled parasitism” of fungal partner on algal/cyanobacterial partner of symbiosis. Differences in the sensitivity of algal cells from different habitats to lichen compounds are, however, not understood sufficiently.

The pattern of photosynthetic response and adaptation to changing light conditions in lichens is linked to their ecological range

Epiphytic lichens constitute an important component of biodiversity in both deforested and forest ecosystems. Widespread occurrence is the domain of generalist lichens or those that prefer open areas. While, many stenoecious lichens find shelter only in a shaded interior of forests. Light is one of the factors known to be responsible for lichen distribution. Nevertheless, the effect of light intensity on photosynthesis of lichen photobionts remain largely unknown. We investigated photosynthesis in lichens with different ecological properties in relation to light as the only parameter modified during the experiments. The aim was to find links between this parameter and habitat requirements of a given lichen. We applied the methods based on a saturating light pulse and modulated light to perform comprehensive analyses of fast and slow chlorophyll fluorescence transient (OJIP and PSMT) combined with quenching analysis. We also examined the rate of CO2 assimilation. Common or generalist lichens, i.e. Hypogymnia physodes, Flavoparmelia caperata and Parmelia sulcata, are able to adapt to a wide range of light intensity. Moreover, the latter species, which prefers open areas, dissipates the excess energy most efficiently. Conversely, Cetrelia cetrarioides considered an old-growth forest indicator, demonstrates definitely lower range of energy dissipation than other species, although it assimilates CO2 efficiently both at low and high light. We conclude that functional plasticity of the thylakoid membranes of photobionts largely determines the dispersal abilities of lichens and light intensity is one of the most important factors determining the specificity of a species to a given habitat.

Alternative electron transport pathways contribute to tolerance to high light stress in lichenized algae.

The photosynthetic apparatus of lichen photobionts has been well-characterized by chlorophyll fluorescence analysis (e.g., by pulse amplitude modulation [PAM]), which provides a proxy of the activity of photosystem II (PSII) and its antenna. However, such kinetics are unable to directly characterize photosystem I (PSI) activity and the associated alternative electron pathways that may be involved in photoprotection. Instead, PSI can be probed in vivo by near-infrared absorption, measured at the same time as standard chlorophyll fluorescence (e.g., using the WALZ Dual PAM). Here, we used the Dual PAM to investigate cyclic electron flow and photoprotection in a range of mostly temperate lichens sampled from shaded to more open microhabitats. Sun species displayed lower acceptor side limitation of PSI (Y[NA]) early in illumination when compared to shade species, indicative of higher flavodiiron-mediated pseudocyclic electron flow. In response to high irradiance, some lichens accumulate melanin, and Y[NA] was lower and NAD(P)H dehydrogenase (NDH-2)-type cyclic flow was higher in melanised than pale forms. Furthermore, non-photochemical quenching (NPQ) was higher and faster relaxing in shade than sun species, while all lichens displayed high rates of photosynthetic cyclic electron flow. In conclusion, our data suggest that (1) low acceptor side limitation of PSI is important for sun-exposed lichens; (2) NPQ helps shade species tolerate brief exposure to high irradiance; and (3) cyclic electron flow is a prominent feature of lichens regardless of habitat, although NDH-2-type flow is associated with high light acclimation.

Lichen photobionts can be hardened to photoinhibition by pretreatment with light

Lichen photobionts can be hardened to photoinhibition by pretreatment with light

Adaptions of photosynthesis in sun and shade in populations of some Afromontane lichens

AbstractPhotosynthetic organisms have evolved a great variety of mechanisms to optimize their use of sunlight. Some of the clearest examples of adaptations can be seen by comparing photosynthesis in different species and in different individuals of the same species that grow under high and low light levels. While the adaptations of sun and shade higher plants have been relatively well studied, much less information is available on the photobionts of lichenized Ascomycetes. An important adaptation that can protect photosynthetic organisms from the potentially harmful effects of excess light is non-photochemical quenching (NPQ); NPQ can dissipate unused light energy as heat. Here we used chlorophyll fluorescence to compare the induction and relaxation of NPQ and the induction of electron transport (rETR) in collections of the same lichen species from exposed and from more shaded locations. All species have trebouxioid photobionts and normally grow in more exposed microhabitats but can also be readily collected from more shaded locations. Shade forms display generally higher NPQ, presumably to protect lichens from occasional rapid increases in light that occur during sunflecks. Furthermore, the NPQ of shade forms relaxes quickly when light levels are reduced, presumably to ensure efficient photosynthesis after a sunfleck has passed. The maximal relative electron transport rate is lower in shade than sun collections, probably reflecting a downregulation of photosynthetic capacity to reduce energy costs. We also compared collections of pale and melanized thalli from three species of shade lichens with Symbiochloris as their photobiont. Interestingly, NPQ in melanized thalli from slightly more exposed microhabitats induced and relaxed in a way that resembled shade rather than sun forms of the trebouxioid lichens. This might suggest that in some locations melanization induced during a temporary period of high light may be excessive and could potentially reduce photosynthesis later in the growing season. Taken together, the results suggest that lichen photobionts can flexibly adjust the amount and type of NPQ, and their levels of rETR in response to light availability.

Metabolite Profiling in Green Microalgae with Varying Degrees of Desiccation Tolerance.

Trebouxiophyceae are microalgae occupying even extreme environments such as polar regions or deserts, terrestrial or aquatic, and can occur free-living or as lichen photobionts. Yet, it is poorly understood how environmental factors shape their metabolism. Here, we report on responses to light and temperature, and metabolic adjustments to desiccation in Diplosphaera epiphytica, isolated from a lichen, and Edaphochlorella mirabilis, isolated from Tundra soil, assessed via growth and photosynthetic performance parameters. Metabolite profiling was conducted by GC–MS. A meta-analysis together with data from a terrestrial and an aquatic Chlorella vulgaris strain reflected elements of phylogenetic relationship, lifestyle, and relative desiccation tolerance of the four algal strains. For example, compatible solutes associated with desiccation tolerance were up-accumulated in D. epiphytica, but also sugars and sugar alcohols typically produced by lichen photobionts. The aquatic C. vulgaris, the most desiccation-sensitive strain, showed the greatest variation in metabolite accumulation after desiccation and rehydration, whereas the most desiccation-tolerant strain, D. epiphytica, showed the least, suggesting that it has a more efficient constitutive protection from desiccation and/or that desiccation disturbed the metabolic steady-state less than in the other three strains. The authors hope that this study will stimulate more research into desiccation tolerance mechanisms in these under-investigated microorganisms.

Photobiont Diversity in Lichen Symbioses From Extreme Environments.

Fungal–algal relationships—both across evolutionary and ecological scales—are finely modulated by the presence of the symbionts in the environments and by the degree of selectivity and specificity that either symbiont develop reciprocally. In lichens, the green algal genus Trebouxia Puymaly is one of the most frequently recovered chlorobionts. Trebouxia species-level lineages have been recognized on the basis of their morphological and phylogenetic diversity, while their ecological preferences and distribution are still only partially unknown. We selected two cosmopolitan species complexes of lichen-forming fungi as reference models, i.e., Rhizoplaca melanophthalma and Tephromela atra, to investigate the diversity of their associated Trebouxia spp. in montane habitats across their distributional range worldwide. The greatest diversity of Trebouxia species-level lineages was recovered in the altitudinal range 1,000–2,500 m a.s.l. A total of 10 distinct Trebouxia species-level lineages were found to associate with either mycobiont, for which new photobionts are reported. One previously unrecognized Trebouxia species-level lineage was identified and is here provisionally named Trebouxia “A52.” Analyses of cell morphology and ultrastructure were performed on axenically isolated strains to fully characterize the new Trebouxia “A52” and three other previously recognized lineages, i.e., Trebouxia “A02,” T. vagua “A04,” and T. vagua “A10,” which were successfully isolated in culture during this study. The species-level diversity of Trebouxia associating with the two lichen-forming fungi in extreme habitats helps elucidate the evolutionary pathways that this lichen photobiont genus traversed to occupy varied climatic and vegetative regimes.

Lichens from the littoral zone host diverse Ulvophycean photobionts.

Crustose Verrucariaceae lichens form a distinctive black belt on seashores all over the world. This lifestyle is apparently enabled by a specific set of photobionts. However, their diversity is understudied. We sampled these lichens from the northern Patagonian Pacific coast of Chile. Using molecular markers, we identified both mycobionts and photobionts. The lichens, belonging to the genus Hydropunctaria and to the Wahlenbergiella group, hosted solely Ulvophycean photobionts. Pseudendoclonium submarinum (Kornmanniaceae, Ulvales) was the most common, but representatives of other closely related, yet undescribed, lineages were also found. Undulifilum symbioticum gen. et sp. nov. is described within Kornmanniaceae based on culture morphology and DNA sequence data. Furthermore, the free-living macroscopic genus Urospora (Acrosiphoniaceae, Ulotrichales) is reported as a lichen photobiont for the first time and is the first of its kind in the order. These results indicate that undescribed algal diversity is waiting to be uncovered in seashore lichens.

Inhibition of Primary Photosynthesis in Desiccating Antarctic Lichens Differing in Their Photobionts, Thallus Morphology, and Spectral Properties.

Five macrolichens of different thallus morphology from Antarctica (King George Island) were used for this ecophysiological study. The effect of thallus desiccation on primary photosynthetic processes was examined. We investigated the lichens’ responses to the relative water content (RWC) in their thalli during the transition from a wet (RWC of 100%) to a dry state (RWC of 0%). The slow Kautsky kinetics of chlorophyll fluorescence (ChlF) that was recorded during controlled dehydration (RWC decreased from 100 to 0%) and supplemented with a quenching analysis revealed a polyphasic species-specific response of variable fluorescence. The changes in ChlF at a steady state (Fs), potential and effective quantum yields of photosystem II (FV/FM, ΦPSII), and nonphotochemical quenching (NPQ) reflected a desiccation-induced inhibition of the photosynthetic processes. The dehydration-dependent fall in FV/FM and ΦPSII was species-specific, starting at an RWC range of 22–32%. The critical RWC for ΦPSII was below 5%. The changes indicated the involvement of protective mechanisms in the chloroplastic apparatus of lichen photobionts at RWCs of below 20%. In both the wet and dry states, the spectral reflectance curves (SRC) (wavelength 400–800 nm) and indices (NDVI, PRI) of the studied lichen species were measured. Black Himantormia lugubris showed no difference in the SRCs between wet and dry state. Other lichens showed a higher reflectance in the dry state compared to the wet state. The lichen morphology and anatomy data, together with the ChlF and spectral reflectance data, are discussed in relation to its potential for ecophysiological studies in Antarctic lichens.

Photoprotection in lichens: adaptations of photobionts to high light

AbstractLichens often grow in microhabitats where they are exposed to severe abiotic stresses such as desiccation and temperature extremes. They are also often exposed to levels of light that are greater than lichen photobionts can use in carbon fixation. Unless regulated, excess energy absorbed by the photobionts can convert ground state oxygen to reactive oxygen species (ROS). These ROS can attack the photosynthetic apparatus, causing photoinhibition and photo-oxidative stress, reducing the ability of the photobionts to fix carbon. Here, we outline our current understanding of the effects of high light on lichens and the mechanisms they use to mitigate or tolerate this stress in hydrated and desiccated states. Tolerance to high light can be achieved first by lowering ROS formation, via synthesizing light screening pigments or by thermally dissipating the excess light energy absorbed; second, by scavenging ROS once formed; or third, by repairing ROS-induced damage. While the primary focus of this review is tolerance to high light in lichen photobionts, our knowledge is rather fragmentary, and therefore we also include recent findings in free-living relatives to stimulate new lines of research in the study of high light tolerance in lichens.

Global Biodiversity Patterns of the Photobionts Associated with the Genus Cladonia (Lecanorales, Ascomycota)

The diversity of lichen photobionts is not fully known. We studied here the diversity of the photobionts associated with Cladonia, a sub-cosmopolitan genus ecologically important, whose photobionts belong to the green algae genus Asterochloris. The genetic diversity of Asterochloris was screened by using the ITS rDNA and actin type I regions in 223 specimens and 135 species of Cladonia collected all over the world. These data, added to those available in GenBank, were compiled in a dataset of altogether 545 Asterochloris sequences occurring in 172 species of Cladonia. A high diversity of Asterochloris associated with Cladonia was found. The commonest photobiont lineages associated with this genus are A. glomerata, A. italiana, and A. mediterranea. Analyses of partitioned variation were carried out in order to elucidate the relative influence on the photobiont genetic variation of the following factors: mycobiont identity, geographic distribution, climate, and mycobiont phylogeny. The mycobiont identity and climate were found to be the main drivers for the genetic variation of Asterochloris. The geographical distribution of the different Asterochloris lineages was described. Some lineages showed a clear dominance in one or several climatic regions. In addition, the specificity and the selectivity were studied for 18 species of Cladonia. Potentially specialist and generalist species of Cladonia were identified. A correlation was found between the sexual reproduction frequency of the host and the frequency of certain Asterochloris OTUs. Some Asterochloris lineages co-occur with higher frequency than randomly expected in the Cladonia species.

Myco- and photobiont associations in crustose lichens in the McMurdo Dry Valleys (Antarctica) reveal high differentiation along an elevational gradient

Climatically extreme regions such as the polar deserts of the McMurdo Dry Valleys (78° S) in Continental Antarctica are key areas for a better understanding of changes in ecosystems. Therefore, it is particularly important to analyze and communicate current patterns of biodiversity in these sensitive areas, where precipitation mostly occurs in form of snow and liquid water is rare. Humidity provided by dew, clouds, and fog are the main water sources, especially for rock-dwelling crustose lichens as one of the most common vegetation-forming organisms. We investigated the diversity and interaction specificity of myco-/photobiont associations of 232 crustose lichen specimens, collected along an elevational gradient (171–959 m a.s.l.) within the McMurdo Dry Valleys. The mycobiont species and photobiont OTUs were identified by using three markers each (nrITS, mtSSU, RPB1, and nrITS, psbJ-L, COX2). Elevation, positively associated with water availability, turned out to be the key factor explaining most of the distribution patterns of the mycobionts. Pairwise comparisons showed Lecidea cancriformis and Rhizoplaca macleanii to be significantly more common at higher elevations and Carbonea vorticosa and Lecidea polypycnidophora at lower elevations. Lichen photobionts were dominated by the globally distributed Trebouxia OTU, Tr_A02 which occurred at all habitats. Network specialization resulting from myco-/photobiont bipartite network structure varied with elevation and associated abiotic factors. Along an elevational gradient, the spatial distribution, diversity, and genetic variability of the lichen symbionts appear to be mainly influenced by improved water relations at higher altitudes.

Lichens Bite the Dust - A Bioweathering Scenario in the Atacama Desert.

SummaryBioweathering mediated by microorganisms plays a significant role in biogeochemical cycles on global scales over geological timescales. Single processes induced by specific taxa have been described but could rarely be demonstrated for complex communities that dominate whole landscapes. The recently discovered grit crust of the coastal Atacama Desert, which is a transitional community between a cryptogamic ground cover and a rock-bound lithic assemblage, offers the unique chance to elucidate various bioweathering processes that occur simultaneously. Here, we present a bioweathering scenario of this biocenosis including processes such as penetration of the lithomatrix, microbial responses to wet-dry cycles, alkalinolysis, enzyme activity, and mineral re-localization. Frequently occurring fog, for example, led to a volume increase of microorganisms and the lithomatrix. This, together with pH shifts and dust accumulation, consequently results in biophysical breakdown and the formation of a terrestrial protopedon, an initial stage of pedogenesis fueled by the grit crust.