SciCombinator

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Concept: Moss

756

Recent climate change on the Antarctic Peninsula is well documented [1-5], with warming, alongside increases in precipitation, wind strength, and melt season length [1, 6, 7], driving environmental change [8, 9]. However, meteorological records mostly began in the 1950s, and paleoenvironmental datasets that provide a longer-term context to recent climate change are limited in number and often from single sites [7] and/or discontinuous in time [10, 11]. Here we use moss bank cores from a 600-km transect from Green Island (65.3°S) to Elephant Island (61.1°S) as paleoclimate archives sensitive to regional temperature change, moderated by water availability and surface microclimate [12, 13]. Mosses grow slowly, but cold temperatures minimize decomposition, facilitating multi-proxy analysis of preserved peat [14]. Carbon isotope discrimination (Δ(13)C) in cellulose indicates the favorability of conditions for photosynthesis [15]. Testate amoebae are representative heterotrophs in peatlands [16-18], so their populations are an indicator of microbial productivity [14]. Moss growth and mass accumulation rates represent the balance between growth and decomposition [19]. Analyzing these proxies in five cores at three sites over 150 years reveals increased biological activity over the past ca. 50 years, in response to climate change. We identified significant changepoints in all sites and proxies, suggesting fundamental and widespread changes in the terrestrial biosphere. The regional sensitivity of moss growth to past temperature rises suggests that terrestrial ecosystems will alter rapidly under future warming, leading to major changes in the biology and landscape of this iconic region-an Antarctic greening to parallel well-established observations in the Arctic [20].

Concepts: Plant, Climate, Weather, Temperature, Antarctica, Cold, Peat, Moss

373

Establishing the timescale of early land plant evolution is essential for testing hypotheses on the coevolution of land plants and Earth’s System. The sparseness of early land plant megafossils and stratigraphic controls on their distribution make the fossil record an unreliable guide, leaving only the molecular clock. However, the application of molecular clock methodology is challenged by the current impasse in attempts to resolve the evolutionary relationships among the living bryophytes and tracheophytes. Here, we establish a timescale for early land plant evolution that integrates over topological uncertainty by exploring the impact of competing hypotheses on bryophyte-tracheophyte relationships, among other variables, on divergence time estimation. We codify 37 fossil calibrations for Viridiplantae following best practice. We apply these calibrations in a Bayesian relaxed molecular clock analysis of a phylogenomic dataset encompassing the diversity of Embryophyta and their relatives within Viridiplantae. Topology and dataset sizes have little impact on age estimates, with greater differences among alternative clock models and calibration strategies. For all analyses, a Cambrian origin of Embryophyta is recovered with highest probability. The estimated ages for crown tracheophytes range from Late Ordovician to late Silurian. This timescale implies an early establishment of terrestrial ecosystems by land plants that is in close accord with recent estimates for the origin of terrestrial animal lineages. Biogeochemical models that are constrained by the fossil record of early land plants, or attempt to explain their impact, must consider the implications of a much earlier, middle Cambrian-Early Ordovician, origin.

Concepts: Evolution, Plant, Plants, Bryophyte, Embryophyte, Moss, Marchantiophyta, Hornwort

174

A hydrophobic cuticle consisting of waxes and the polyester cutin covers the aerial epidermis of all land plants, providing essential protection from desiccation and other stresses. We have determined the enzymatic basis of cutin polymerization through characterization of a tomato extracellular acyltransferase, CD1, and its substrate, 2-mono(10,16-dihydroxyhexadecanoyl)glycerol. CD1 has in vitro polyester synthesis activity and is required for cutin accumulation in vivo, indicating that it is a cutin synthase.

Concepts: Plant, In vivo, In vitro, Annual plant, Epidermis, Bryophyte, Moss, Plant cuticle

168

Moss species Physcomitrella patens has been used as a model system in plant science for several years, because it has a short life cycle and is easy to be handled. With the completion of its genome sequencing, more and more proteomic analyses were conducted to study the mechanisms of P. patens abiotic stress resistance. It can be concluded from these studies that abiotic stresses could lead to the repression of photosynthesis and enhancement of respiration in P. patens, although different stresses could also result in specific responses. Comparative analysis showed that the responses to drought and salinity were very similar to that of abscisic acid, while the response to cold was quite different from these three. Based on previous studies, it is proposed that sub-proteomic studies on organelles or protein modifications, as well as functional characterization of those candidate proteins identified from proteomic studies will help us to further understand the mechanisms of abiotic stress resistance in P. patens.

Concepts: DNA, Photosynthesis, Protein, Genome, Plant, Model organism, Moss, Physcomitrella patens

148

Mosses are among the earliest branching embryophytes and probably originated not later than the early Ordovician when atmospheric CO2 was higher and O2 was lower than today. The C3 biochemistry and physiology of their photosynthesis suggests, by analogy with tracheophytes, that growth of extant bryophytes in high CO2 approximating Ordovician values would increase the growth rate. This occurs for many mosses, including Physcomitrella patens in suspension culture, although recently published transcriptomic data on this species at high CO2 and present-day CO2 show down-regulation of the transcription of several genes related to photosynthesis. It would be useful if transcriptomic (and proteomic) data comparing growth conditions are linked to measurements of growth and physiology on the same, or parallel, cultures. Mosses (like later-originating embryophytes) have been subject to changes in bulk atmospheric CO2 and O2 throughout their existence, with evidence, albeit limited, for positive selection of moss Rubisco. Extant mosses are subject to a large range of CO2 and O2 concentrations in their immediate environments, especially aquatic mosses, and mosses are particularly influenced by CO2 generated by, and O2 consumed by, soil chemoorganotrophy from organic C produced by tracheophytes (if present) and bryophytes.

Concepts: Carbon dioxide, Plant, Bryophyte, Embryophyte, Moss, Marchantiophyta, Physcomitrella patens, Mosses

147

Metals deposited into ecosystems are non-degradable and become one of the major toxic agents which accumulate in habitats. Thus, their concentration requires precise monitoring. To evaluate pollution around a chlor-alkali plant, a glass smelter, two power plants and a ceramic and porcelain factory, we selected terrestrial mosses with different life forms: the orthotropic and endohydric Polytrichum commune and plagiotropic and ectohydric Pleurozium schreberi. Metal concentrations were determined in both species growing together at sites situated at various distances approximately 0.75, 1.5, 3 and 6 km from polluters. MARS analysis evaluated different tendencies of both species for Cd, Co and Pb accumulation depending on the distance from the emitter. In P. schreberi, the concentration of these metals diminished relatively rapidly with an increasing distance from the emitter up to 3000 m and then stabilised. For P. commune, a steady decrease could be observed with increasing the distance up to 6000 m. PCCA ordination explained that both species from the vicinity of the chlor-alkali plant were correlated with the highest Co, Cr, Cu, Fe and Pb as well as Mn and Ni concentrations in their tissues. The mosses from sites closest to both power plants were correlated with the highest Cd and Zn concentrations. P. commune contained significantly higher Cd, Cr, Ni, Pb and Zn concentrations compared to P. schreberi. This may be caused by the lamellae found in the leaves of P. commune which increase the surface area of the possible aerial absorption of contaminants. Soil may also be an additional source of metals, and it affects the uptake in endohydric P. commune more than in ectohydric P. schreberi. However, the precise explanation of these relations needs further investigation.

Concepts: Iron, Plant, Metal, Copper, Pollution, Moss, Mosses, Polytrichum commune

146

Momilactones, which are diterpenoid phytoalexins with antimicrobial and allelopathic functions, have been found only in rice and the moss Hypnum plumaeforme. Although these two evolutionarily distinct plant species are thought to produce momilactones as a chemical defence, the momilactone biosynthetic pathway in H. plumaeforme has been unclear. Here, we identified a gene encoding syn-pimara-7,15-diene synthase (HpDTC1) responsible for the first step of momilactone biosynthesis in the moss. HpDTC1 is a bifunctional diterpene cyclase that catalyses a two-step cyclization reaction of geranylgeranyl diphosphate to syn-pimara-7,15-diene. HpDTC1 transcription was up-regulated in response to abiotic and biotic stress treatments. HpDTC1 promoter-GUS analysis in transgenic Physcomitrella patens showed similar transcriptional responses as H. plumaeforme to the stresses, suggesting that a common response system to stress exists in mosses. Jasmonic acid (JA), a potent signalling molecule for inducing plant defences, could not activate HpDTC1 expression. In contrast, 12-oxo-phytodienoic acid, an oxylipin precursor of JA in vascular plants, enhanced HpDTC1 expression and momilactone accumulation, implying that as-yet-unknown oxylipins could regulate momilactone biosynthesis in H. plumaeforme. These results demonstrate the existence of an evolutionarily conserved chemical defence system utilizing momilactones and suggest the molecular basis of the regulation for inductive production of momilactones in H. plumaeforme.

Concepts: DNA, Gene, Gene expression, Amino acid, Molecular biology, Plant, Moss, Mosses

49

Mosses, dominant elements in the vegetation of polar and alpine regions, have well-developed stress tolerance features permitting cryptobiosis. However, direct regeneration after longer periods of cryptobiosis has been demonstrated only from herbarium and frozen material preserved for 20 years at most [1]. Recent field observations of new moss growth on the surface of small moss clumps re-exposed from a cold-based glacier after about 400 years of ice cover have been accompanied by regeneration in culture from homogenised material [2], but there are no reported instances of regrowth occurring directly from older preserved material.

Concepts: Glacier, English-language films, Antarctica, Moss, Glaciology, Mosses

45

Lignin, one of the most abundant biopolymers on Earth, derives from the plant phenolic metabolism. It appeared upon terrestrialization and is thought critical for plant colonization of land. Early diverging land plants do not form lignin, but already have elements of its biosynthetic machinery. Here we delete in a moss the P450 oxygenase that defines the entry point in angiosperm lignin metabolism, and find that its pre-lignin pathway is essential for development. This pathway does not involve biochemical regulation via shikimate coupling, but instead is coupled with ascorbate catabolism, and controls the synthesis of the moss cuticle, which prevents desiccation and organ fusion. These cuticles share common features with lignin, cutin and suberin, and may represent the extant representative of a common ancestor. Our results demonstrate a critical role for the ancestral phenolic metabolism in moss erect growth and cuticle permeability, consistent with importance in plant adaptation to terrestrial conditions.

Concepts: Amino acid, Metabolism, Plant, Cell wall, Bryophyte, Embryophyte, Moss, Marchantiophyta

32

The evolutionary emergence of land plant body plans transformed the planet. However, our understanding of this formative episode is mired in the uncertainty associated with the phylogenetic relationships among bryophytes (hornworts, liverworts, and mosses) and tracheophytes (vascular plants). Here we attempt to clarify this problem by analyzing a large transcriptomic dataset with models that allow for compositional heterogeneity between sites. Zygnematophyceae is resolved as sister to land plants, but we obtain several distinct relationships between bryophytes and tracheophytes. Concatenated sequence analyses that can explicitly accommodate site-specific compositional heterogeneity give more support for a mosses-liverworts clade, “Setaphyta,” as the sister to all other land plants, and weak support for hornworts as the sister to all other land plants. Bryophyte monophyly is supported by gene concatenation analyses using models explicitly accommodating lineage-specific compositional heterogeneity and analyses of gene trees. Both maximum-likelihood analyses that compare the fit of each gene tree to proposed species trees and Bayesian supertree estimation based on gene trees support bryophyte monophyly. Of the 15 distinct rooted relationships for embryophytes, we reject all but three hypotheses, which differ only in the position of hornworts. Our results imply that the ancestral embryophyte was more complex than has been envisaged based on topologies recognizing liverworts as the sister lineage to all other embryophytes. This requires many phenotypic character losses and transformations in the liverwort lineage, diminishes inconsistency between phylogeny and the fossil record, and prompts re-evaluation of the phylogenetic affinity of early land plant fossils, the majority of which are considered stem tracheophytes.

Concepts: Evolution, Plant, Phylogenetics, Bryophyte, Embryophyte, Moss, Marchantiophyta, Hornwort