Growth physiology and chlorophyll fluorescence analysis of two moss species under different LED light qualities.

This study investigated the responses of Didymodon constrictus and Hypnum plumaeforme to different light qualities emitted by light-emitting diodes (LEDs), including white light (WL), red light (RL), blue light (BL), yellow light (YL), green light (GL), and a combination of red and blue light (R1B1L). The research analyzed the fluorescence imaging, photosynthetic pigments, coloration, and growth characteristics related to antioxidant enzymes in these two moss species. The results indicated that R1B1L significantly enhanced the content of photosynthetic pigments, maximum relative electron transport rate (rETRmax), saturation light intensity (IK), and the greenness of the moss. RL improved the maximum quantum yield (Fv/Fm), the light energy efficiency of H. plumaeforme and effective quantum yield in both moss species. In contrast, BL notably increased non-photochemical quenching (NPQ), photochemical quenching (qp), and the steady-state fluorescence decrease ratio (RFD) in H. plumaeforme. The application of GL significantly increases the maximum photon yield (Fv/Fm) in D. constrictus, as well as the light energy efficiency and elongation length, resulting in a shift in the color composition of both moss species towards yellow. Among the light treatments, R1B1L had the highest induction rate and promotional effect on the growth of both moss species. These mosses absorbed GL and RL effectively, while BL played a crucial role in the dissipation of heat and electron transfer in H. plumaeforme. This research provides valuable insights for the regulation of LED light environments and the physiological adaptability of moss in artificial cultivation.

Characteristics of spatial and temporal variability in the distribution of biological soil crusts on the Loess Plateau, China.

Biological soil crust (biocrust) is widely distributed on the Loess Plateau and plays multiple roles in regulating ecosystem stability and multifunctionality. Few reports are available on the distribution characteristics of biocrust in this region, which limits the assessment of its ecological functions. Based on 388 sampling points in different precipitation zones on the Loess Plateau from 2009 to 2020, we analyzed the coverage, composition, and influencing factors of biocrust across different durations since land abandonment, precipitation levels, topography (slope aspect and position), and utilization of abandoned slopelands (shrubland, forest, and grassland). On this base, with the assistance of machine learning and spatial modeling methods, we generated a distribution map of biocrust and its composition at a resolution of 250 m × 250 m, and analyzed the spatial distribution of biocrust on the Loess Plateau. The results showed that the average biocrust coverage in the woodlands and grasslands was 47.3%, of which cyanobacterial crust accounted for 25.5%, moss crust 19.7%, and lichen crust 2.1%. There were significant temporal and spatial variations. Temporally, the coverage of biocrust in specific regions fluctuated with the extension of the abandoned durations and coverage of cyanobacterial crust, while moss crust showed a reverse pattern. In addition, the coverage of biocrust in the wet season was slightly higher than that in the dry season within a year. Spatially, the coverage of biocrusts on the sandy lands area on the Loess Plateau was higher and dominated by cyanobacterial crusts, while the coverage was lower in the hilly and gully area. Precipitation and utilization of abandoned land were the major factors driving biocrust coverage and composition, while slope direction and position did not show obvious effect. In addition, soil organic carbon content, pH, and texture were related to the distribution of biocrust. This study uncovered the spatial and temporal variability of biocrust distribution, which might provide important data support for the research and management of biocrust in the Loess Plateau region.

Small but visible: Predicting rare bryophyte distribution and richness patterns using remote sensing-based ensembles of small models.

In Canadian boreal forests, bryophytes represent an essential component of biodiversity and play a significant role in ecosystem functioning. Despite their ecological importance and sensitivity to disturbances, bryophytes are overlooked in conservation strategies due to knowledge gaps on their distribution, which is known as the Wallacean shortfall. Rare species deserve priority attention in conservation as they are at a high risk of extinction. This study aims to elaborate predictive models of rare bryophyte species in Canadian boreal forests using remote sensing-derived predictors in an Ensemble of Small Models (ESMs) framework. We hypothesize that high ESMs-based prediction accuracy can be achieved for rare bryophyte species despite their low number of occurrences. We also assess if there is a spatial correspondence between rare and overall bryophyte richness patterns. The study area is located in western Quebec and covers 72,292 km2. We selected 52 bryophyte species with <30 occurrences from a presence-only database (214 species, 389 plots in total). ESMs were built from Random Forest and Maxent techniques using remote sensing-derived predictors related to topography and vegetation. Lee's L statistic was used to assess and map the spatial relationship between rare and overall bryophyte richness patterns. ESMs yielded poor to excellent prediction accuracy (AUC > 0.5) for 73% of the modeled species, with AUC values > 0.8 for 19 species, which confirmed our hypothesis. In fact, ESMs provided better predictions for the rarest bryophytes. Likewise, our study revealed a spatial concordance between rare and overall bryophyte richness patterns in different regions of the study area, which have important implications for conservation planning. This study demonstrates the potential of remote sensing for assessing and making predictions on inconspicuous and rare species across the landscape and lays the basis for the eventual inclusion of bryophytes into sustainable development planning.

Evolutionary insight of plant cuticle biosynthesis in bryophytes.

As an adaptive innovation in plant terrestrialization, cuticle covers the plant surface and greatly contributes to the development and stress tolerance in land plants. Although past decades have seen great progress in understanding the molecular mechanism of cuticle biosynthesis in flowering plants with the contribution of cuticle biosynthesis mutants and advanced cuticle composition profiling techniques, origins and evolution of cuticle biosynthesis are poorly understood. Recent chemical, phylogenomic, and molecular genetic studies on cuticle biosynthesis in early-diverging extant land plant lineages, the bryophytes, shed novel light on the origins and evolution of plant cuticle biosynthesis. In this mini-review, we highlighted these recent advances in the molecular biology of cuticle biosynthesis in bryophytes, and provided evolutionary insights into plant cuticle biosynthesis.

Quantitative cell biology of tip growth in moss.

KEY MESSAGE: Here we review, from a quantitative point of view, the cell biology of protonemal tip growth in the model moss Physcomitrium patens. We focus on the role of the cytoskeleton, vesicle trafficking, and cell wall mechanics, including reviewing some of the existing mathematical models of tip growth. We provide a primer for existing cell biological tools that can be applied to the future study of tip growth in moss. Polarized cell growth is a ubiquitous process throughout the plant kingdom in which the cell elongates in a self-similar manner. This process is important for nutrient uptake by root hairs, fertilization by pollen, and gametophyte development by the protonemata of bryophytes and ferns. In this review, we will focus on the tip growth of moss cells, emphasizing the role of cytoskeletal organization, cytoplasmic zonation, vesicle trafficking, cell wall composition, and dynamics. We compare some of the existing knowledge on tip growth in protonemata against what is known in pollen tubes and root hairs, which are better-studied tip growing cells. To fully understand how plant cells grow requires that we deepen our knowledge in a variety of forms of plant cell growth. We focus this review on the model plant Physcomitrium patens, which uses tip growth as the dominant form of growth at its protonemal stage. Because mosses and vascular plants shared a common ancestor more than 450 million years ago, we anticipate that both similarities and differences between tip growing plant cells will provide mechanistic information of tip growth as well as of plant cell growth in general. Towards this mechanistic understanding, we will also review some of the existing mathematical models of plant tip growth and their applicability to investigate protonemal morphogenesis. We attempt to integrate the conclusions and data across cell biology and physical modeling to our current state of knowledge of polarized cell growth in P. patens and highlight future directions in the field.

Assessment of cultured media for desert moss crust by physiological responses.

The physiological responses of desert moss crusts under four artificial media (Beneck, Part, BG11, and Hogland) were investigated to evaluate the function of culture media during different culture periods. The results showed that the value of malondialdehyde (MDA) was at a maximum at 11d, on the contrary, chlorophyll-a, soluble protein, and soluble sugar were at a minimum. As the time increased, the value of MDA and soluble protein decreased faster in the Hogland, while the value of chlorophyll-a and soluble sugar increased. At the end of the culture period, the value of chlorophyll-a and soluble sugar was at a maximum in the Hogland, while the value of MDA and soluble protein was at a minimum. The results suggested that the Hogland medium had a promoting effect on the growth of desert moss crusts. The selected artificial cultivation medium towards wider and larger scale field applications of cultural desert biocrust was widely anticipated.

Improvement of moss photosynthesis by humic acids from Antarctic tundra soil.

There have been several published reports regarding the growth promoting effect of humic acids (HA) on vascular plants; however, the effect of HA on bryophytes is still unknown. Due to the ecological importance of mosses, which dominate the Antarctic flora, we assessed the effectiveness of HA as a biostimulant using three moss species: Antarctic Ceratodon purpureus KMA5038, Arctic Bryum sp. KMR5045, and Physcomitrella patens which inhabits temperate regions. Natural HA (KS1-3_HA) were extracted through acidic precipitation of alkaline extracts from Antarctic tundra soil. Spectroscopic structural properties of KS1-3_HA were characterized and determined to possess several functional groups such as hydroxyl (R-OH) and carboxyl (R-COOH), implying they could have a growth-related biological function. For two polar mosses, increasing HA concentrations correlated with increased growth and photosynthesis. The efficiency for temperate moss increased at lower concentrations tested, but rather began to reduce at the highest HA concentration, indicating that effective concentrations of HA vary depending on the moss species and habitat. Based on these results, Antarctic HA may have ecological role in enhancing the growth and photosynthesis of Antarctic mosses. We believe this is the first study to establish a positive physiological effect of HA on mosses and hope it may serve as a basis for studying the role of HA in preserving the terrestrial ecosystem of Antarctica.

Spatial and Temporal Patterns of Endopolyploidy in Mosses.

Somatic polyploidy or endopolyploidy is common in the plant kingdom; it ensures growth and allows adaptation to the environment. It is present in the majority of plant groups, including mosses. Endopolyploidy had only been previously studied in about 65 moss species, which represents less than 1% of known mosses. We analyzed 11 selected moss species to determine the spatial and temporal distribution of endopolyploidy using flow cytometry to identify patterns in ploidy levels among gametophytes and sporophytes. All of the studied mosses possessed cells with various ploidy levels in gametophytes, and four of six species investigated in sporophytic stage had endopolyploid sporophytes. The proportion of endopolyploid cells varied among organs, parts of gametophytes and sporophytes, and ontogenetic stages. Higher ploidy levels were seen in basal parts of gametophytes and sporophytes than in apical parts. Slight changes in ploidy levels were observed during ontogenesis in cultivated mosses; the youngest (apical) parts of thalli tend to have lower levels of endopolyploidy. Differences between parts of cauloid and phylloids of Plagiomnium ellipticum and Polytrichum formosum were also documented; proximal parts had higher levels of endopolyploidy than distal parts. Endopolyploidy is spatially and temporally differentiated in the gametophytes of endopolyploid mosses and follows a pattern similar to that seen in angiosperms.

Light- and hormone-mediated development in non-flowering plants: An overview.

MAIN CONCLUSION: Light, hormones and their interaction regulate different aspects of development in non-flowering plants. They might have played a role in the evolution of different plant groups by conferring specific adaptive evolutionary changes. Plants are sessile organisms. Unlike animals, they lack the opportunity to abandon their habitat in unfavorable conditions. They respond to different environmental cues and adapt accordingly to control their growth and developmental pattern. While phytohormones are known to be internal regulators of plant development, light is a major environmental signal that shapes plant processes. It is plausible that light-hormone crosstalk might have played an important role in plant evolution. But how the crosstalk between light and phytohormone signaling pathways might have shaped the plant evolution is unclear. One of the possible reasons is that flowering plants have been studied extensively in context of plant development, which cannot serve the purpose of evolutionary comparisons. In order to elucidate the role of light, hormone and their crosstalk in the evolutionary adaptation in plant kingdom, one needs to understand various light- and hormone-mediated processes in diverse non-flowering plants. This review is an attempt to outline major light- and phytohormone-mediated responses in non-flowering plant groups such as algae, bryophytes, pteridophytes and gymnosperms.

Bryophytes are predicted to lag behind future climate change despite their high dispersal capacities.

The extent to which species can balance out the loss of suitable habitats due to climate warming by shifting their ranges is an area of controversy. Here, we assess whether highly efficient wind-dispersed organisms like bryophytes can keep-up with projected shifts in their areas of suitable climate. Using a hybrid statistical-mechanistic approach accounting for spatial and temporal variations in both climatic and wind conditions, we simulate future migrations across Europe for 40 bryophyte species until 2050. The median ratios between predicted range loss vs expansion by 2050 across species and climate change scenarios range from 1.6 to 3.3 when only shifts in climatic suitability were considered, but increase to 34.7-96.8 when species dispersal abilities are added to our models. This highlights the importance of accounting for dispersal restrictions when projecting future distribution ranges and suggests that even highly dispersive organisms like bryophytes are not equipped to fully track the rates of ongoing climate change in the course of the next decades.

Forest canopy-cover composition and landscape influence on bryophyte communities in Nothofagus forests of southern Patagonia.

Bryophytes (liverworts, mosses and hornworts) are one of the most diverse plant groups worldwide but one of the least studied in temperate forests from an ecological perspective. In comparison to vascular plants, bryophytes have a broader distribution and a longer altitudinal gradient, and their influence on the landscape is poorly understood. The objective was to evaluate environmental drivers that can influence bryophyte cover, richness, diversity, and nestedness in different forest canopy compositions in two typical landscapes across the natural distribution of bryophytes in Tierra del Fuego (Argentina). Three natural Nothofagus forest types (pure deciduous, pure evergreen, and mixed deciduous-evergreen) in two landscapes (coasts < 100 m.a.s.l. and mountains > 400 m.a.s.l.) were selected (N = 60 plots). In each plot, we established one transect (10 m length) to measure bryophyte cover (point-intercept method). Data were evaluated using generalized linear mixed models and multivariate analyses. The studied environmental drivers were mainly explained by the microclimate, with higher effective annual precipitation and relative air humidity in the coastal forests and higher soil moisture in the mountain forests. Greater liverwort richness was found in evergreen forests at the mountain (9 species) than at the coastal, while mosses showed higher richness in mixed deciduous-evergreen forests at the coastal (11 species) than at the mountain. However, the expected richness according to the rarefaction/extrapolation curves suggested that it is possible to record additional species, except for liverworts in pure deciduous forests on the coasts. Similarities and differences among the studied forest types and among plots of the same forest type and landscape were detected. These differences in the studied indexes (similarity that varied between 0 and 1) ranged from 0.09-0.48 for liverworts and 0.05-0.65 for mosses. Moreover, these results indicated that pure evergreen and mixed deciduous-evergreen forests presented higher moss cover (10.7% and 10.0%, respectively), mainly in the mountains than on the coast. These outputs highlight the need to explore differences at greater altitudinal ranges to achieve sustainability objectives conservation planning for bryophytes in southernmost forests.

Life under quartz: Hypolithic mosses in the Mojave Desert.

Several species of dryland cyanobacteria are known to occur as hypoliths under semi-translucent rocks. In the Mojave Desert, these organisms find refuge from intense solar radiation under milky quartz where moisture persists for a longer period of time than in adjacent soil surface habitat. Desert mosses, which are extremely desiccation-tolerant, can also occur in these hypolithic spaces, though little is known about this unique moss microhabitat and how species composition compares to that of adjacent soil surface communities. To address this question, we deployed microclimate dataloggers and collected moss samples from under and adjacent to 18 milky quartz rocks (quartz mean center thickness 26 ± 15 mm) in a western high elevation Mojave Desert site. Light transmission through Mojave quartz rocks may be as low as 1.2%, and data from microclimate loggers deployed for five months support the hypothesis that quartz provides thermal buffering and higher relative humidity compared to the soil surface. Of the 53 samples collected from hypolith and surface microhabitats, 68% were Syntrichia caninervis, the dominant bryophyte of the Mojave Desert biological soil crust. Tortula inermis accounted for 28% of the samples and 4% were Bryum argenteum. In a comparison of moss community composition, we found that S. caninervis was more likely to be on the soil surface, though it was abundant in both microhabitats, while T. inermis was more restricted to hypoliths, perhaps due to protection from temperature extremes. In our study site, the differences between hypolithic and surface microhabitats enable niche partitioning between T. inermis and S. caninervis, enhancing alpha diversity. This work points to the need to thoroughly consider microhabitats when assessing bryophyte species diversity and modelling species distributions. This focus is particularly important in extreme environments, where mosses may find refuge from the prevailing macroclimatic conditions in microhabitats such as hypoliths.

Cross-kingdom signalling regulates spore germination in the moss Physcomitrella patens.

Plants live in close association with microorganisms that can have beneficial or detrimental effects. The activity of bacteria in association with flowering plants has been extensively analysed. Bacteria use quorum-sensing as a way of monitoring their population density and interacting with their environment. A key group of quorum sensing molecules in Gram-negative bacteria are the N-acylhomoserine lactones (AHLs), which are known to affect the growth and development of both flowering plants, including crops, and marine algae. Thus, AHLs have potentially important roles in agriculture and aquaculture. Nothing is known about the effects of AHLs on the earliest-diverging land plants, thus the evolution of AHL-mediated bacterial-plant/algal interactions is unknown. In this paper, we show that AHLs can affect spore germination in a representative of the earliest plants on land, the Bryophyte moss Physcomitrella patens. Furthermore, we demonstrate that sporophytes of some wild isolates of Physcomitrella patens are associated with AHL-producing bacteria.

Bryophytes and the symbiotic microorganisms, the pioneers of vegetation restoration in karst rocky desertification areas in southwestern China.

In karst rocky desertification areas, bryophytes coexist with algae, bacteria, and fungi on exposed calcareous rocks to form a bryophyte crust, which plays an irreplaceable role in the restoration of karst degraded ecosystems. We investigated the biodiversity of crust bryophytes in karst rocky desertification areas from Guizhou Province, China. A total of 145 species in 22 families and 56 genera were identified. According to frequency and coverage, seven candidate dominant mosses were screened out, and five drought-resistant indexes of them were measured. Hypnum leptothallum, Racopilum cuspidigerum, and Hyophila involuta have high drought adaptability. We explored the interactions between two dominant mosses (H. leptothallum, H. involuta) and the structure of microbial communities in three karst rocky desertification types. Microbial diversity and function analysis showed that both moss species and karst rocky desertification types affect microbial communities. Moss species much more strongly affected the diversity and changed the community composition of these microbial groups. Bacteria were more sensitive in the microbiome as their communities changed strongly between mosses and drought resistance factors. Moreover, several species of fungi and bacteria could be significantly associated with three drought-resistant indexes: Pro (free proline content), SOD (superoxide dismutase activity), and POD (peroxidase activity), which were closely related to the drought adaptability of mosses. Our results enforced the potential role of moss-associated microbes that are important components involved in the related biological processes when bryophytes adapted to arid habitats, or as one kind of promoters in the distribution pattern of early mosses succession in karst rocky desertification areas.

When facilitation meets clonal integration in forest canopies.

Few studies have explored how - within the same system - clonality and positive plant-plant interactions might interact to regulate plant community composition. Canopy-dwelling epiphytes in species-rich forests provide an ideal system for studying this because many epiphytic vascular plants undertake clonal growth and because vascular epiphytes colonize canopy habitats after the formation of nonvascular epiphyte (i.e. bryophyte and lichen) mats. We investigated how clonal integration of seven dominant vascular epiphytes influenced inter-specific interactions between vascular epiphytes and nonvascular epiphytes in a subtropical montane moist forest in southwest China. Both clonal integration and environmental buffering from nonvascular epiphytes increased survival and growth of vascular epiphytes. The benefits of clonal integration for vascular epiphytes were higher when nonvascular epiphytes were removed. Similarly, facilitation from nonvascular epiphytes played a more important role when clonal integration of vascular epiphytes was eliminated. Overall, clonal integration had greater benefits than inter-specific facilitation. This study provides novel evidence for interactive effects of clonality and facilitation between vascular and nonvascular species, and has implications for our understanding of a wide range of ecosystems where both high levels of clonality and facilitation are expected to occur.

Species diversity patterns in managed Scots pine stands in ancient forest sites.

Continuity in forest habitats is crucial for species diversity and richness. Ancient Scots pine forests are usually under forest management, which disturbs vegetation and causes differentiation in terms of tree stand age. To date, vegetation variability in ancient Scots pine forests has not been examined based on tree stand age classes. In the present study the continuity of a large Scots pine forest complex was investigated, and a system of sampling plots established in five tree stand age classes: initiation stands (4-10 years), young stands (20-35 years), middle-aged stands (45-60 years), pre-mature stands (70-85 years) and mature stands (95-110 years). Species composition, including vascular plants, bryophytes and lichens, on soil, tree trunks, and coarse woody debris, was analyzed. Based on existing classifications systems, forest species and ancient forest species groups were distinguished. In the studied ancient Scots pine forests the species pool and richness were relatively low, and the vegetation consisted mostly of generalist species. Cryptogams, which can grow on diverse substrates, were the most abundant species. Moreover, most species could tolerate both forest and non-forest conditions. Age class forests provided different environmental niches for species. Initiation stands were optimal for terrestrial light-demanding species, and in terms of species composition, initiation stands were most specific. Young stands were most preferred by species on coarse woody debris, and at this stage of stand maturation epiphytic species re-appeared. The oldest stands were not rich in forest specialists, i.e. species of closed forest and ancient forest species. Cryptogams of closed forests inhabited different substrates, and they were not associated only with the oldest stands. The low number of forest specialists in the oldest stands may be a general feature of acidophilus pine forests. However, it may also be a result of the lack of species sources in the vicinity of maturing pine stands. In managed forests a frequent diversity pattern is an increase in a species pool and richness after clear-cut logging. In the present study we obtained higher species pools in initiation and young stands, but richness was similar in all tree stand age classes. This resulted from taking into account species of different substrates (terrestrial, epixylous and epiphytic species) which changed their participation in the vegetation of subsequent stages of tree stand development.

Can mowing restore boreal rich-fen vegetation in the face of climate change?

Low-frequency mowing has been proposed to be an effective strategy for the restoration and management of boreal fens after abandonment of traditional haymaking. This study investigates how mowing affects long-term vegetation change in both oceanic and continental boreal rich-fen vegetation. This will allow evaluation of the effectiveness of mowing as a management and restoration tool in this ecosystem in the face of climate change. At two nature reserves in Central Norway (Tågdalen, 63° 03' N, 9° 05 E, oceanic climate and Sølendet, 62° 40' N, 11° 50' E, continental climate), we used permanent plot data from the two sites to compare plant species composition from the late 1960s to the early 1980s with that recorded in 2012-2015 in abandoned and mown fens. Changes in species composition and frequency were analysed by multivariate and univariate methods in relation to environmental variables and modelled climate and groundwater data. Mowing resulted in a decline in shrub and Molinia caerulea cover at the continental and oceanic sites respectively, and the total cover of specialist fen species had increased to a significantly greater extent in the mown plots than the unmown at the continental site. However, mowing did not have an effect on the cover of specialist bryophyte species, and some specialist species declined regardless of mowing treatment. Temperature sums had increased at both sites, but precipitation had not changed significantly. Mowing was shown to be the most important determinant of plant community composition at both sites, with local environmental conditions being of secondary importance. In conclusion, the abandonment of traditional management practices results in the loss of characteristic fen species. In order to encourage the restoration of typical rich-fen vegetation, particularly in oceanic areas, additional management measures, such as more intensive mowing, may be required.

Small-scale Variation of Testate Amoeba Assemblages: the Effect of Site Heterogeneity and Empty Shell Inclusion.

Studies on testate amoeba species distribution at small scales (i.e., single peatland sites) are rare and mostly focus on bogs or mineral-poor Sphagnum fens, leaving spatial patterns within mineral-rich fens completely unexplored. In this study, two mineral-rich fen sites of contrasting groundwater chemistry and moss layer composition were selected for the analysis of testate amoeba compositional variance within a single site. At each study site, samples from 20 randomly chosen moss-dominated plots were collected with several environmental variables being measured at each sampling spot. We also distinguished between empty shells and living individuals to evaluate the effect of empty shell inclusion on recorded species distribution. At the heterogeneous-rich Sphagnum-fen, a clear composition turnover in testate amoebae between Sphagnum-dominated and brown moss-dominated samples was closely related to water pH, temperature and redox potential. We also found notable species composition variance within the homogeneous calcareous fen, yet it was not as high as for the former site and the likely drivers of community assembly remained unidentified. The exclusion of empty shells provided more accurate data on species distribution as well as their relationship with some environmental variables, particularly moisture. Small-scale variability in species composition of communities seems to be a worthwhile aspect in testate amoeba research and should be considered in future sampling strategies along with a possible empty shell bias for more precise understanding of testate amoeba ecology and paleoecology.