Characterization of yeast groupings in the phyllosphere of <Emphasis Type="Italic">Sphagnum</Emphasis> mosses

Significant differences were revealed in the taxonomic structure of the epiphytic yeast communities formed on Sphagnum mosses and on the leaves of vascular plants. On mosses, low abundance of red yeasts was found (the most typical epiphytes on vascular plant leaves), along with a relatively high content and diversity of nonpigmented dimorphic basidiomycetes related to the order Leucosporidiales. The species composition of epiphytic yeasts from mosses is different from that of both forest and meadow grasses and of the parts of vascular plants submerged in the turf. The specific composition of the Sphagnum mosses yeast community is probably determined by the biochemical characteristics of this environment, rather than by the hydrothermal regime in the turf.     

Comparative desiccation tolerance of two Sphagnum mosses

Summary Sphagnum fallax (Klinggr.) Klinggr., a moss growing in hollows close to the water table, is more desiccation tolerant than S. nemoreum Scop., a hummock former distributed high above the hollows. Sphagnum fallax recovered to a greater proportion of its predesiccation photosynthetic rate after one and five days of tissue dryness. Further, a greater percentage of S. fallax plants survived five and ten day periods at low tissue water contents. Longer desiccated periods and lower water contents during these periods decreased both photosynthetic recovery and survival.Water contents measured in Bloomingdale Bog (Adirondack Mountains, NY, USA) showed that S. fallax probably dries more frequently and for longer periods than S. nemoreum. These results support previous findings that the greater ability of S. nemoreum to remain moist in the field is the most important character in its success as a hummock former. Greater tolerance of desiccation helps S. fallax to compensate for its greater tendency to become dry, and is a key physiological feature enabling it to dominate hollows.     

Mineral nutrient economy in competing species of Sphagnum mosses

Bog vegetation, which is dominated by Sphagnum mosses, depends exclusively on aerial deposition of mineral nutrients. We studied how the main mineral nutrients are distributed between intracellular and extracellular exchangeable fractions and along the vertical physiological gradient of shoot age in seven Sphagnum species occupying contrasting bog microhabitats. While the Sphagnum exchangeable cation content decreased generally in the order Ca²⁺ ≥ K⁺, Na⁺, Mg²⁺ > Al³⁺ > NH₄ ⁺, intracellular element content decreased in the order N > K > Na, Mg, P, Ca, Al. Calcium occurred mainly in the exchangeable form while Mg, Na and particularly K, Al and N occurred inside cells. Hummock species with a higher cation exchange capacity (CEC) accumulated more exchangeable Ca²⁺, while the hollow species with a lower CEC accumulated more exchangeable Na⁺, particularly in dead shoot segments. Intracellular N and P, but not metallic elements, were consistently lower in dead shoot segments, indicating the possibility of N and P reutilization from senescing segments. The greatest variation in tissue nutrient content and distribution was between species from contrasting microhabitats. The greatest variation within microhabitats was between the dissimilar species S. angustifolium and S. magellanicum. The latter species had the intracellular N content about 40% lower than other species, including even this species when grown alone. This indicates unequal competition for N, which can lead to outcompeting of S. magellanicum from mixed patches. We assume that efficient cation exchange enables Sphagnum vegetation to retain immediately the cationic nutrients from rainwater. This may represent an important mechanism of temporal extension of mineral nutrient availability to subsequent slow intracellular nutrient uptake.     

Notes on Guayana mosses with new information of Sphagnum ornatum

Mosses restricted to tepui summits in the Guayana Highland are relatively few.Sphagnum ornatum Crum, though, only recently described from Brazil, has been found subsequently in a number of localities in nearby Venezuela and Guyana. These additional collections make for a better understanding of the diagnostic features of a species characteristically found on the tops of the sandstone mesas of southern Venezuela and adjacent areas.     

Recombination and introgression of nuclear and chloroplast genomes between the peat mosses, Sphagnum capillifolium and Sphagnum quinquefarium

Haploid hybrid gametophytes are often present at low frequencies in sympatric populations of Sphagnum capillifolium and Sphagnum quinquefarium. We used intersimple sequence repeat (ISSR) markers and polymerase chain reaction-restriction fragment length polymorphism of the trnL(UAA) intron of the chloroplast genome to reveal the nuclear and chloroplast composition of mature hybrid gametophytes from natural populations and of gametophytes derived from spores of hybrid sporophytes collected in nature. Asymmetrical nuclear inheritance was found in the progeny of the hybrid sporophytes, indicating that only spores with a low level of recombination of parental genomes were viable. A similarly skewed nuclear composition was found among the naturally occurring hybrid gametophytes. All hybrid genomes contained a larger proportion of S. capillifolium ISSR markers, combined with only two to five S. quinquefarium markers together with a chloroplast haplotype derived from S. quinquefarium. In this way, a pattern resembling introgression is created within a single generation. Some individuals possessed nuclear genomes typical for S. capillifolium in combination with the chloroplast haplotype of S. quinquefarium, possibly indicating backcrossing. Our results indicate that hybridization between S. capillifolium and S. quinquefarium is relatively common, but the resistance of large parts of the genome against heterospecific genes maintains the genetic distinctness of the species. Further evolutionary and phylogenetic consequences of restricted interspecific gene exchange are discussed.     

Fatal interactions between Scots pine and Sphagnum mosses in bog ecosystems

In this study, we explore how Sphagnum mosses and Scots pine, Pinus sylvestris , interact on different spatial and temporal scales in a boreal bog ecosystem. We were particularly interested in relationships between the occurrence of Sphagnum- dominated habitats and the occurrence of Scots pines of different age and size.
Juvenile and adult pines occurred in different habitats. While juveniles mainly occurred in Sphagnum- dominated habitats, predominantly with Sphagnum rubellum , adult pines were found in habitats dominated by lichens, or with a sparse vegetation cover. Examination of surface peat cores sampled close to adult pines revealed that almost all pines (97%) had established in a Sphagnum -dominated environment and that the habitat had changed since pine establishment. Scots pine is thus capable of changing and exterminating the Sphagnum -dominated environment preferred for germination and establishment. Pines impede Sphagnum growth and peat accumulation significantly once they have reached a stem diameter of approximately 20 mm. It takes from 30 to 90 yr for a pine to reach that size.
Our results show the importance of interactions between Scots pine and Sphagnum mosses in bog ecosystems. We conclude that interactions between trees and Sphagnum mosses are important driving forces behind the vegetation change that has characterised boreal bogs during the Holocene.     

Effect of water content components on desiccation and recovery in Sphagnum mosses

BACKGROUND AND AIMS: The basic parameters of water relations were measured in Sphagnum mosses. The relationships of these parameters to the photosynthetic response to desiccation and the ecology of these mosses were then tested. METHODS: The water relations parameters of six Sphagnum species (mosses typical of wet habitats) and Atrichum androgynum (a moss more typical of mesophytic conditions) were calculated from pressure-volume isotherms. Photosynthetic properties during and after moderate desiccation were monitored by chlorophyll fluorescence. KEY RESULTS: When desiccated, the hummock-forming species S. fuscum and S. magellanicum lost more water before turgor started dropping than other sphagna inhabiting less exposed habitats (73 % compared with 56 % on average). Osmotic potentials at full turgor were similar in all species, with an average value of -1.1 MPa. Hummock sphagna had clearly more rigid cell walls than species of wet habitats (epsilon = 3 x 55 compared with 1 x 93 MPa). As a result, their chlorophyllous cells lost turgor at higher relative water contents (RWCs) than species of wet habitats (0 x 61 compared with 0 x 46) and at less negative osmotic potentials (-2 x 28 compared with -3 x 00 MPa). During drying, Phi(PSII) started declining earlier in hummock species (at an RWC of 0 x 65 compared with 0 x 44), and F(v)/F(m) behaved similarly. Compared with other species, hummock sphagna desiccated to -20 or -40 MPa recovered more completely after rehydration. Atrichum androgynum responded to desiccation similarly to hummock sphagna, suggesting that their desiccation tolerance may have a similar physiological basis. CONCLUSIONS: Assuming a fixed rate of desiccation, the higher water-holding capacities of hummock sphagna will allow them to continue metabolism for longer than other species. While this could be viewed as a form of 'desiccation avoidance', hummock species also recover faster than other species during rehydration, suggesting that they have higher inherent tolerance. This may help them to persist in drought-exposed hummocks. In contrast, species growing in wet habitats lack such strong avoidance and tolerance mechanisms. However, their turgor maintenance mechanisms, for example more elastic cell walls, enable them to continue metabolizing longer as their water contents fall to the turgor-loss point.     

Endophytic bacteria of Sphagnum mosses as promising objects of agricultural microbiology

Sphagnum mosses serve as a unique habitat for microorganisms, which play an important role both for the host plants and the peatland ecosystems. The aim of the present study was to isolate endophytic bacteria from the tissues of Sphagnum mosses and to screen them for strains promising for further application in agricultural microbiology. About 50 samples of Sphagnum fallax (H. Klinggr.) H. Klinggr. and Sphagnum magellanicum Brid. were collected in the Austrian Alps and the Lenindgrad Region of Russia in 2009–2010. Endophytic bacteria were detected inside the moss plants using fluorescent in situ hybridization (FISH) followed by confocal laser scanning microscopy (CLSM). Altogether, 283 isolates were obtained by cultivation on the nutrient media. Examination of the isolates for the antagonistic activity revealed that more than 50% of them could suppress the growth of phytopathogenic and toxigenic fungi. More than 30% of isolates showed some antagonistic activity against microbial phytopathogens. The isolated strains could colonize crops and promote their growth. Molecular-genetic identification coupled with physiological/biochemical characterization showed that the dominant endophytic groups belonged to the genera Burkholderia, Pseudomonas, Flavobacterium, Serratia and Collimonas. The isolated endophytes were shown to be promising objects for the development of effective growth-promoting and protective microbiological preparations to be used in agriculture.     

Investigations of the structure and function of bacterial communities associated with Sphagnum mosses

High acidity, low temperature and extremely low concentration of nutrients form Sphagnum bogs into extreme habitats for organisms. Little is known about the bacteria associated with living Sphagnum plantlets, especially about their function for the host. Therefore, we analysed the endo- and ectophytic bacterial populations associated with two widely distributed Sphagnum species, Sphagnum magellanicum and Sphagnum fallax , by a multiphasic approach. The screening of 1222 isolates for antagonistic activity resulted in 326 active isolates. The bacterial communities harboured a high proportion of antifungal (26%) but a low proportion of antibacterial isolates (0.4%). Members of the genus Burkholderia (38%) were found to be the most dominant group of antagonistic bacteria. The finding that a large proportion (89%) of the antagonistic bacteria produced antifungal compounds may provide an explanation for the well-known antimicrobial activity of certain Sphagnum species. The secondary metabolites of the Sphagnum species themselves were analysed by HPLC-PDA. The different spectra of detected compounds may not only explain the antifungal activity but also the species specificity of the microbial communities. The latter was analysed using cultivation-independent single-stranded conformation polymorphism (SSCP) analysis. Using Burkholderia -specific primers we found a high diversity of Burkholderia isolates in the endophytic and ectophytic habitats of Sphagnum . Furthermore, a high diversity of nitrogen–fixing bacteria was detected by using nifH-specific primers, especially inside Sphagnum mosses. In conclusion, this study provides evidence that both Sphagnum species were colonized by characteristic bacterial populations, which appear to be important for pathogen defence and nitrogen fixation.     

Three-genome mosses: complex double allopolyploid origins for triploid gametophytes in Sphagnum

This paper documents the occurrence of allotriploidy (having three differentiated genomes) in gametophytes of two Southern Hemisphere Sphagnum species ( S. australe, S . falcatulum ). The pattern of microsatellite alleles indicates that both species are composed of a complex of allodiploid and allotriploid gametophytes, with the latter resulting from two allopolyploidization events. No haploid ( n  =  x ) gametophytes were found for either species. The ploidal levels suggested by the pattern of microsatellite alleles were confirmed by flow cytometry and Feulgen DNA image densitometry. For both S. australe and S. falcatulum , the respective allodiploid plants (or their ancestors) are one of the parent species of the allotriploid plants. This is the first report of triploidy in Sphagnum gametophytes occurring in nature and also the first report of the presence of three differentiated genomes in any bryophyte. It is also the first report of intersectional allopolyploidy in Sphagnum , with S. australe appearing to have parental species from Sphagnum sections Rigida and Sphagnum, and S. falcatulum having parental species from Sphagnum sections Cuspidata and Subsecunda . In both species, the allotriploid cytotypes were the most prevalent cytotype on the South Island of New Zealand. The pattern of microsatellite alleles shows the presence of two genetically distinct populations of allodiploid S. australe , possibly indicating multiple origins of polyploidy for that allodiploid cytotype. Morphological evidence is also highly indicative of recurrent polyploidy in the allotriploid cytotype of S. falcatulum . Allopolyploidy has clearly played a major evolutionary role in these two Southern Hemisphere taxa. This study, in conjunction with other recent research, indicates that allopolyploidy is a common, if not the predominant, form of polyploidy in Sphagnum .     

Photoinhibition as a control on photosynthesis and production of Sphagnum mosses

The effect of high light intensity on photosynthesis and growth of Sphagnum moss species from Alaskan arctic tundra was studied under field and laboratory conditions. Field experiments consisted of experimental shading of mosses at sites normally exposed to full ambient irradiance, and removal of the vascular plant canopy from above mosses in tundra water track habitats. Moss growth was then monitored in the experimental plots and in adjacent control areas for 50 days from late June to early August 1988. In shaded plots total moss growth was 2–3 times higher than that measured in control plots, while significant reductions in moss growth were found in canopy removal plots. The possibility that photoinhibition of photosynthesis might occur under high-light conditions and affect growth was studied under controlled laboratory conditions with mosses collected from the arctic study site, as well as from a temperate location in the Sierra Nevada, California. After 2 days of high-light treatment (800 mol photons m^–2 s^–1) in a controlled environmental chamber, moss photosynthetic capacity was significantly lowered in both arctic and temperate samples, and did not recover during the 14-day experimental period. The observed decrease in photosynthetic capacity was correlated (r ²=0.735, P<0.001) with a decrease in the ratio of variable to maximum chlorophyll fluorescence (F _v/F _m) in arctic and temperate mosses. This relationship indicates photoinhibition of photosynthesis in both arctic and temperate mosses at even moderately high light intensities. It is suggested that susceptibility to photoinhibition and failure to photoacclimate to higher light intensities in Sphagnum spp. may be related to low tissue nitrogen levels in these exclusively ombrotrophic plants. Photoinhibition of photosynthesis leading to lowered annual carbon gain in Sphagnum mosses may be an important factor affecting CO₂ flux at the ecosystem level, given the abundance of these plants in Alaskan tussock tundra.     

Sphagnum mosses harbour highly specific bacterial diversity during their whole lifecycle

Knowledge about Sphagnum-associated microbial communities, their structure and their origin is important to understand and maintain climate-relevant Sphagnum-dominated bog ecosystems. We studied bacterial communities of two cosmopolitan Sphagnum species, which are well adapted to different abiotic parameters (Sphagnum magellanicum, which are strongly acidic and ombrotrophic, and Sphagnum fallax, which are weakly acidic and mesotrophic), in three Alpine bogs in Austria by a multifaceted approach. Great differences between bacterial fingerprints of both Sphagna were found independently from the site. This remarkable specificity was confirmed by a cloning and a deep sequencing approach. Besides the common Alphaproteobacteria, we found a discriminative spectrum of bacteria; although Gammaproteobacteria dominated S. magellanicum, S. fallax was mainly colonised by Verrucomicrobia and Planctomycetes. Using this information for fluorescent in situ hybridisation analyses, corresponding colonisation patterns for Alphaproteobacteria and Planctomycetes were detected. Bacterial colonies were found in high abundances inside the dead big hyalocytes, but they were always connected with the living chlorocytes. Using multivariate statistical analysis, the abiotic factors nutrient richness and pH were identified to modulate the composition of Sphagnum-specific bacterial communities. Interestingly, we found that the immense bacterial diversity was transferred via the sporophyte to the gametophyte, which can explain the high specificity of Sphagnum-associated bacteria over long distances. In contrast to higher plants, which acquire their bacteria mainly from the environment, mosses as the phylogenetically oldest land plants maintain their bacterial diversity within the whole lifecycle.     

Climatic modifiers of the response to nitrogen deposition in peat-forming Sphagnum mosses: a meta-analysis

Peatlands in the northern hemisphere have accumulated more atmospheric carbon (C) during the Holocene than any other terrestrial ecosystem, making peatlands long-term C sinks of global importance. Projected increases in nitrogen (N) deposition and temperature make future accumulation rates uncertain. Here, we assessed the impact of N deposition on peatland C sequestration potential by investigating the effects of experimental N addition on Sphagnum moss. We employed meta-regressions to the results of 107 field experiments, accounting for sampling dependence in the data. We found that high N loading (comprising N application rate, experiment duration, background N deposition) depressed Sphagnum production relative to untreated controls. The interactive effects of presence of competitive vascular plants and high tissue N concentrations indicated intensified biotic interactions and altered nutrient stochiometry as mechanisms underlying the detrimental N effects. Importantly, a higher summer temperature (mean for July) and increased annual precipitation intensified the negative effects of N. The temperature effect was comparable to an experimental application of almost 4 g?N m(-2) yr(-1) for each 1°C increase. Our results indicate that current rates of N deposition in a warmer environment will strongly inhibit C sequestration by Sphagnum-dominated vegetation.     

Detection, isolation, and characterization of acidophilic methanotrophs from Sphagnum mosses

Sphagnum peatlands are important ecosystems in the methane cycle. Methane-oxidizing bacteria in these ecosystems serve as a methane filter and limit methane emissions. Yet little is known about the diversity and identity of the methanotrophs present in and on Sphagnum mosses of peatlands, and only a few isolates are known. The methanotrophic community in Sphagnum mosses, originating from a Dutch peat bog, was investigated using a pmoA microarray. A high biodiversity of both gamma- and alphaproteobacterial methanotrophs was found. With Sphagnum mosses as the inoculum, alpha- and gammaproteobacterial acidophilic methanotrophs were isolated using established and newly designed media. The 16S rRNA, pmoA, pxmA, and mmoX gene sequences showed that the alphaproteobacterial isolates belonged to the Methylocystis and Methylosinus genera. The Methylosinus species isolated are the first acid-tolerant members of this genus. Of the acidophilic gammaproteobacterial strains isolated, strain M5 was affiliated with the Methylomonas genus, and the other strain, M200, may represent a novel genus, most closely related to the genera Methylosoma and Methylovulum. So far, no acidophilic or acid-tolerant methanotrophs in the Gammaproteobacteria class are known. All strains showed the typical features of either type I or II methanotrophs and are, to the best of our knowledge, the first isolated (acidophilic or acid-tolerant) methanotrophs from Sphagnum mosses.     

Nitrogen deposition interacts with climate in affecting production and decomposition rates in Sphagnum mosses

Increasing rates of atmospheric nitrogen (N) deposition may reduce growth and accelerate decomposition of Sphagnum mosses in bogs. Sphagnum growth and rates of Sphagnum litter decomposition may also vary because of climate change as both processes are controlled by climatic factors. The initial purpose of this study was to assess if growth and litter decomposition of hummock and lawn Sphagnum species varied with increasing N input in a factorial mid‐term (2002–2005) experiment of N and phosphorus (P) addition, in a bog on the southern Alps of Italy. However, as the experimental period was characterized by an exceptional heat wave in summer 2003, we also explored the interacting effects of fertilization and strongly varying climate on growth and decomposition rates of Sphagnum. The heat wave implied strong dehydration of the upper Sphagnum layer even if precipitation in summer 2003 did not differ appreciably from the overall mean. Sphagnum production was somewhat depressed by high levels (3 g m⁻² yr⁻¹) of N addition without concomitant addition of P presumably because of nutrient imbalance in the tissues, but production rates were much lower than the overall means in 2003, when no effect of nutrient addition could be observed. Adding N at high level also increased the potential decay of Sphagnum litter. Higher CO₂ emission from N‐fertilized litter was due to amelioration of litter chemistry showing lower C/N quotients in the N‐fertilized treatments. Rates of CO₂ emission from incubated litter also were more strongly affected by water content than by nutrient status, with practically no CO₂ emission detected when litter was dry. We conclude that higher rates of atmospheric N availability input may depress Sphagnum growth because of P, and presumably potassium, (co‐)limitation. Higher N availability is also expected to promote potential decay of Sphagnum litter by ameliorating litter chemistry. However, both effects are less pronounced if the growing Sphagnum apex and the underlying senescing tissues dry out.     

Range change evolution of peat mosses (Sphagnum) within and between climate zones

Peat mosses (Sphagnum) hold exceptional importance in the control of global carbon fluxes and climate because of the vast stores of carbon bound up in partially decomposed biomass (peat). This study tests the hypothesis that the early diversification of Sphagnum was in the Northern Hemisphere, with subsequent range expansions to tropical latitudes and the Southern Hemisphere. A phylogenetic analysis of 192 accessions representing the moss class Sphagnopsida based on four plastid loci was conducted in conjunction with biogeographic analyses using BioGeoBEARS to investigate the tempo and mode of geographic range evolution. Analyses support the hypothesis that the major intrageneric clades of peat‐forming species accounting for >90% of peat moss diversity originated and diversified at northern latitudes. The genus underwent multiple range expansions into tropical and Southern Hemisphere regions. Range evolution in peat mosses was most common within latitudinal zones, attesting to the relative difficulty of successfully invading new climate zones. Allopolyploidy in Sphagnum (inferred from microsatellite heterozygosity) does not appear to be biased with regard to geographic region nor intrageneric clade. The inference that Sphagnum diversified in cool‐or cold‐climate regions and repeatedly expanded its range into tropical regions makes the genus an excellent model for studying morphological, physiological, and genomic traits associated with adaptation to warming climates.