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We studied the effects of different radiation treatments on the physiology and UV-absorbing compounds of the model liverwort Marchantia polymorpha subsp. ruderalis. Starting from gemmae, samples were exposed to five radiation treatments: low photosynthetically active radiation (PAR), low PAR+ UV-A, low PAR + UV-B, low PAR + UV-A + UV-B, and high PAR. After 35 days, the maximum quantum yield of photosystem II was similar between treatments, which suggested comparable photoinhibition and physiological vitality, also supported by results showing an unchanged chlorophyll a/b ratio and only slight changes in growth. However, the total contents of both chlorophylls and carotenoids decreased in the UV radiation treatments and, more strongly, in the high-PAR samples, suggesting mainly PAR-dependent damage to the photosynthetic pigments. The xanthophyll index (antheraxanthin + zeaxanthin)/(violaxanthin + antheraxanthin + zeaxanthin) was only increased in the high-PAR samples, indicating an increase in photoprotection through nonphotochemical dissipation of the excess energy. The sclerophylly index (the ratio between the thallus dry mass and surface area) was increased in the UV-B-exposed samples, suggesting a UV-induced structural protection. Only the UV-B-exposed samples showed DNA damage. Several apigenin and luteolin derivatives were found in the methanol-soluble vacuolar fraction of the liverwort and p-coumaric and ferulic acids in the methanol-insoluble cell wall-bound fraction. Most individual soluble compounds, the bulk level of soluble compounds, and chalcone synthase expression increased in UV-B-exposed samples, whereas individual insoluble compounds increased in the samples exposed to only PAR. Principal components analysis summarized these responses, showing the strong influence of both UV-B and PAR levels on the physiology and UV protection of the samples.

The liverwort Marchantia polymorpha subsp. ruderalis is an emerging model plant, and some data are available on its responses to ultraviolet (UV) radiation. However, it is unknown if the developmental stage of the thalli modulates the effects of UV radiation on the contents of potentially protecting phenolic compounds. To fill this gap, liverwort samples were exposed or non-exposed to UV radiation for 38 days under controlled conditions, using three developmental stages: gemmae (G), one-month thalli (T1), and two-month thalli (T2). Then, the bulk level of methanol-soluble UV-absorbing compounds and the contents of six flavones (apigenin and luteolin derivatives) were measured. The UV responsiveness decreased with thallus age: G and T1 plants were the most UV-responsive and showed a strong increase in all the variables, with G plants more responsive than T1 plants. In UV-exposed T2 plants, only apigenin derivatives increased and more modestly, probably due to a lower acclimation capacity. Nevertheless, the thalli became progressively tougher due to a decreasing water content, representing a possible structural protection against UV. In UV-exposed plants, the temporal patterns of the accumulation of phenolic compounds were compound-specific. Most compounds decreased with thallus age, but di-glucuronide derivatives showed a bell-shaped pattern, with T1 plants showing the highest contents. A Principal Components Analysis (PCA) ordination of the different samples summarized the results found. The patterns described above should be taken into account to select thalli of an adequate developmental stage for experiments investigating the induction of phenolic compounds by UV radiation.

Bryophytes are small and structurally simple plants consisting of around 20,000 species distributed in three evolutionary lineages (mosses, liverworts, and hornworts). Bryophytes usually occupy harsh environments where other photosynthetic organisms are little competitive. This capacity is based on the tolerance of bryophytes to adverse environmental factors (particularly desiccation) and their economy in production, which in turn are found on diverse physiological mechanisms and ecological adaptations. One of these mechanisms is the outstanding capacity of bryophytes to produce bioactive compounds with diverse biological functions. In particular, liverworts can synthesize a great variety of terpenoids, and the three bryophyte lineages can produce phenolic derivatives (from simple cinnamic acids to complex flavonoids), alkaloids, and lipids. These compounds, many of them specific, contribute to the development of special smells and flavors and may have both positive (antibiotic, fungicidal, deterrent, herbicidal, antioxidant, cytotoxic, antidiabetic) and negative (allergenic) properties for human beings. Bisbibenzyls and sesquiterpenoid derivatives in liverworts and the diterpenoid derivatives momilactones in mosses are the most relevant compounds. Of particular interest is the recent possibility to use model bryophytes (the moss Physcomitrella patens and the liverwort Marchantia polymorpha) as biotechnological tools (in bioreactors) to produce pharmaceutical substances. Overall, and despite the advances experienced in the past decades, biotechnological substances have only been studied in a relatively small number of bryophyte species, and thus, this discipline has a strong research potential for the future.

Ultraviolet (UV) radiation strongly influences grape composition, but only a few studies have focused on how this influence is conserved in the resulting wines. Here we analyzed to what extent the changes induced by exposing Tempranillo grapes to UV radiation from budbreak to harvest were conserved in wine. By using different cut-off filters and lamps, we differentiated the effects of ambient levels of UV-A and UV-B wavelengths, as well as the effects of a realistic UV-B enhancement associated with climate change. Among phenolic compounds, the most consistent responses to UV were those of flavonols (particularly quercetin-, kaempferol-, isorhamnetin- and myricetin-glycosides), which significantly increased in wines whose grapes had been exposed to a synergic combination of UV-A and UV-B radiation. This confirms that flavonols are the phenolic compounds most reliably conserved from UV-exposed grapes to wine, despite the possible influence of the winemaking process. Flavonols are important compounds because they contribute to wine co-pigmentation by stabilizing anthocyanins, and they are interesting antioxidants and nutraceuticals. Hydroxycinnamic acids also increased under the same UV combination or under UV-A alone. Wine VOCs were much less reactive to the UV received by grapes than phenolic compounds, and only esters showed significantly higher values under (mainly) UV-A alone. This was surprising because (1) UV-A has been considered to be less important than UV-B to induce metabolic changes in plants, and (2) esters are produced during winemaking. Esters are relevant due to their contribution to the fruity aroma in wines. In general, the remaining phenolic compounds (stilbenes, flavanols, hydroxybenzoic acids, and anthocyanins) and VOCs (alcohols, hydrocarbons, and fatty acids), together with wine color and antioxidant capacity, showed inconsistent or non-significant responses to UV radiation. These results were summarized by a multivariate analysis. Our study opens up new possibilities to artificially manipulate UV radiation in grapevine cultivation to improve both grape and wine quality.

We studied the acclimation modalities of bryophytes to sun and shade under ambient or close-to-ambient conditions, measuring variables usually influenced by photosynthetically active (PAR) and ultraviolet (UV) radiations. Our aim was to elucidate to what extent the responses to changing radiations were influenced by PAR and UV wavelengths. For this aim, we used three taxonomically and structurally different species: the thalloid liverwort Marchantia polymorpha subsp. polymorpha, the leafy liverwort Jungermannia exsertifolia subsp. cordifolia, and the moss Fontinalis antipyretica. In the field, liverworts were more radiation-responsive than the moss, and the thalloid liverwort was more responsive than the leafy liverwort. Sun plants of M. polymorpha showed, in comparison to shade plants, higher sclerophylly, lower Chl a + b contents, higher Chl a/b ratios, higher (antheraxanthin + zeaxanthin)/(violaxanthin + antheraxanthin + zeaxanthin) ratios (xanthophyll index), lower Fv/Fm values, higher contents of methanol-soluble vacuolar UV-absorbing compounds (soluble UVACs), higher values of the ratio between the contents of methanol-insoluble cell wall-bound UVACs (insoluble UVACs) and soluble UVACs, higher contents of soluble luteolin and apigenin derivatives and riccionidin A, and higher contents of insoluble p-coumaric and ferulic acids. Overall, these responses reduced light absorption, alleviated overexcitation, increased photoprotection through non-photochemical energy dissipation, increased UV protection through UV screening and antioxidant capacity, and denoted photoinhibition. J. exsertifolia showed moderate differences between sun and shade plants, while responses of F. antipyretica were rather diffuse. The increase in the xanthophyll index was the most consistent response to sun conditions, occurring in the three species studied. The responses of soluble UVACs were generally clearer than those of insoluble UVACs, probably because insoluble UVACs are relatively immobilized in the cell wall. These modalities of radiation acclimation were reliably summarized by principal components analysis. Using the most radiation-responsive species in the field (M. polymorpha), we found, under close-to-ambient greenhouse conditions, that sclerophylly and Chl a + b content were only influenced by PAR, Fv/Fm, and luteolin and apigenin derivatives were only determined by UV, and xanthophyll index was influenced by both radiation types. Thus, responses of bryophytes to radiation can be better interpreted considering the influence of both PAR and UV radiation.

In the present study we assessed the effects of ambient solar UV exclusion on leaf physiology, and leaf and berry skin phenolic composition, of a major grapevine cultivar (Tempranillo) grown under typically Mediterranean field conditions over an entire season. In general, the effects of time were stronger than those of UV radiation. Ambient UV caused a little stressing effect (eustress) on leaf physiology, with decreasing net photosynthesis rates and stomatal conductances. However, it was not accompanied by alterations in F/F or photosynthetic pigments, and was partially counterbalanced by the UV-induced accumulation of protective flavonols. Consequently, Tempranillo leaves are notably adapted to current UV levels. The responses of berry skin phenolic compounds were diverse, moderate, and mostly transitory. At harvest, the clearest response in UV-exposed berries was again flavonol accumulation, together with a decrease in the flavonol hydroxylation level. Contrarily, responses of anthocyanins, flavanols, stilbenes and hydroxycinnamic derivatives were much more subtle or nonexistent. Kaempferols were the only compounds whose leaf and berry skin contents were correlated, which suggests a mostly different regulation of phenolic metabolism for each organ. Interestingly, the dose of biologically effective UV radiation (UV) was correlated with the leaf and berry skin contents of quercetins and kaempferols; relationships were linear except for the exponential relationship between UV dose and berry skin kaempferols. This opens management possibilities to modify kaempferol and quercetin contents in grapevine through UV manipulation., The authors wish to thank the Ministerio de Economía y Competitividad of Spain (MINECO) and FEDER (Project CGL2014-54127-P) for financial support. MADCA and GS benefited from grants of the Universidad de La Rioja (Plan Propio 2014, 2013, respectively).

Background This paper aims to investigate the correlations between the concentrations of nine heavy metals in moss and atmospheric deposition within ecological land classes covering Europe. Additionally, it is examined to what extent the statistical relations are affected by the land use around the moss sampling sites. Based on moss data collected in 2010/2011 throughout Europe and data on total atmospheric deposition modelled by two chemical transport models (EMEP MSC-E, LOTOS-EUROS), correlation coefficients between concentrations of heavy metals in moss and in modelled atmospheric deposition were specified for spatial subsamples defined by ecological land classes of Europe (ELCE) as a spatial reference system. Linear discriminant analysis (LDA) and logistic regression (LR) were then used to separate moss sampling sites regarding their contribution to the strength of correlation considering the areal percentage of urban, agricultural and forestry land use around the sampling location. After verification LDA models by LR, LDA models were used to transform spatial information on the land use to maps of potential correlation levels, applicable for future network planning in the European Moss Survey. Results Correlations between concentrations of heavy metals in moss and in modelled atmospheric deposition were found to be specific for elements and ELCE units. Land use around the sampling sites mainly influences the correlation level. Small radiuses around the sampling sites examined (5 km) are more relevant for Cd, Cu, Ni, and Zn, while the areal percentage of urban and agricultural land use within large radiuses (75–100 km) is more relevant for As, Cr, Hg, Pb, and V. Most valid LDA models pattern with error rates of

Aquatic bryophytes show the same basic characteristics of bryophytes, but certain structural and ecophysiological features make them peculiar among bryophytes. Terrestrial, emergent and facultative aquatic bryophytes mostly respond to the consequences of climate change occurring in the atmospheric environment, because they are exposed to air temperature, atmospheric CO2 and ambient sunlight. Aquatic bryophytes are adapted to low temperatures, and show optimum photosynthesis and growth at around 15°C. It can be observed that bryophytes are ideal plants for climate change bioindication, due to their sensitivity to ecological changes. Rheophilous and limnophilous bryophytes may show different structural and physiological adaptations. Within bryophytes, obligate aquatic bryophytes (OAB) inhabit fragile ecosystems that are severely threatened nowadays by climate change factors. Therefore, OAB are relevant tools to monitor and predict the effects of climate change on aquatic ecosystems. Adverse factors for OAB in lakes are waterlogging, cold water, low irradiances in the deep zones, high hydrostatic pressure, and abrasion along the shores.

To study the potential quick responses to ultraviolet (UV) radiation of bryophyte phenolic compounds, we cultivated two thalloid liverworts, two leafy liverworts, and two mosses under three moderate realistic UV levels in the laboratory for 22 days. At the end of the daylight period on the first and last culture days, we measured the bulk levels and individual contents of phenolic UV-absorbing compounds (UVACs) of each species, differentiating in both cases the UVACs located in the methanol-soluble (mainly vacuolar) and -insoluble (cell wall-bound) fractions (SUVACs and IUVACs, respectively). The bulk levels of SUVACs and IUVACs mostly showed linear or hyperbolic relationships with the UV dose applied. Thirteen flavones (apigenin and luteolin derivatives) and two hydroxycinnamic acids (p-coumaric and ferulic acids) were identified in the soluble and insoluble fractions, respectively. Only two compounds (p-coumaric and ferulic acids) from the insoluble fraction of the leafy liverwort Plagiochila asplenioides showed a significant quick accumulation in response to UV radiation in the first day of culture, whereas six UVACs (mainly soluble apigenin and luteolin derivatives) from different species (mainly liverworts) were significantly accumulated at the end of the culture. In conclusion, the responses of bryophyte UVACs to UV radiation were influenced by the specific compound considered, the fraction in which each UVAC was located, the global or individual way of UVACs quantification, the bryophyte species and evolutionary lineage, and the experimental conditions used. Particularly, SUVACs were more UV-responsive than IUVACs and liverworts than mosses, and responses were not especially quick.

Aquatic bryophytes are ecologically important in river phytobenthos, especially in mountain streams, despite the adverse environmental conditions. Other hostile aquatic habitats, such as deep lakes with dramatic light starvation, are also successfully inhabited by bryophytes. Some future trends in the investigation of the ecophysiology of pigments in aquatic bryophytes, such as the functioning of the xanthophyll cycle and the use of chlorophyll fluorescence as a vitality index, are outlined. The core complex of PSII contains only chl-a and beta-carotene as photosynthetic pigments. The close correlation that has been reported in aquatic bryophytes between the spectrophotometrically-determined concentrations of chl-b and phaeophytin-a probably derives from the characteristic presence of both pigments in PSII. The genetic influence on pigment composition is clearly observed in the colour of the different species, which is sometimes used as a taxonomical tool. In higher plants, pigment variations are diagnostic symptoms for several nutrient deficiencies, toxicities or antagonisms.

Excessive exposure to ultraviolet (UV) radiation can be harmful to photosynthetic organisms, that most frequently respond with the accumulation of protective UV-absorbing compounds (UVACs). UVACs location in different cell compartments can influence their preferential protective role as antioxidants and/or UV screens. However, the phylogenetic, ecological and evolutionary implications of UVACs compartmentation has been little studied, particularly in bryophytes. We analysed UVACs in the methanol-soluble and -insoluble fractions (SUVACs and IUVACs respectively) in extracts of 87 bryophytes belonging to their three evolutionary lineages: 22 liverworts, 64 mosses and one hornwort. Assuming that the cell wall-bound IUVACs are more effective UV screens than the mainly vacuolar SUVACs, thus conferring a higher UV tolerance, we evaluated whether UVACs levels and compartmentation were related to: (1) the bryophyte phylogeny down to the Order level; and (2) the bryophyte ecological attributes, including sun exposure and Ellenberg indicator values (numerical system classifying species' habitat along gradients of environmental factors). A similar phylogenetic and ecological analysis was conducted on the sclerophylly index (the ratio between the dry mass and the surface area of the bryophyte shoot). Mosses showed lower SUVACs but higher IUVACs and total UVACs, together with higher IUVAC/SUVAC ratios, than liverworts. Thus, mosses would better tolerate UV radiation than liverworts, which matches well with their general ecological preferences. As bryophytes were the earliest diverging land plants, we could infer that the different UVACs compartmentation between mosses and liverworts could have influenced their ecological segregation upon plant land colonization. UVACs compartmentation also differed between the two major moss lineages (acrocarpous and pleurocarpous), while the anatomically peculiar Sphagnales were the best characterized Order. There were not solid relationships between UVACs and the ecological attributes considered. Hence, UVACs might be mainly constitutive in bryophytes, depending more on the species phylogeny than on the habitat occupied in nature. Liverworts were less sclerophyllous than mosses, and, as in tracheophytes, water restrictions and high sun exposures increased sclerophylly. In conclusion, UVACs compartmentation represents an ecophysiological trait useful to understand the bryophyte ecophylogeny, because different compartmentation modalities seem to imply different UV adaptations and tolerance. A plain language summary is available for this article. © 2018 British Ecological Society.

Hornworts are the least species-rich bryophyte lineage, but represent a key group to understand the evolution of plants because, together with the remaining bryophyte lineages (mosses and liverworts), they constitute the earliest diverging land plants. The responses of hornworts to ultraviolet (UV) radiation are unknown, but they may be important to infer how primitive hornworts (and bryophytes in general) coped with UV upon land colonization. In this context, our aim was to show the first data on the effects of UV radiation on the accumulation patterns of phenolic UV-absorbing compounds (UVACs) in the emerging model hornwort Anthoceros agrestis. Thalli of 52 days age were exposed to photosynthetically active radiation (PAR) alone (P regime) and to a combination of PAR + UV-A + UV-B radiation (PAB regime) for 21 days, using realistic UV doses (equivalent to the natural ambient doses received in summer at mid-latitudes). At the end of the culture period, we measured the bulk levels and individual contents of phenolic UV-absorbing compounds (UVACs), differentiating in both cases the UVACs located in the methanol-soluble (mainly vacuolar) and -insoluble (cell wall-bound) fractions (SUVACs and IUVACs, respectively). Three soluble and one insoluble compounds were identified, among which the soluble rosmarinic and anthocerotonic acids are not present in any other bryophyte lineage. The bulk levels of SUVACs were higher than those of IUVACs, a physiological trait more typical of liverworts than of mosses. None of the variables measured responded significantly to UV exposure, but all of them showed an increasing trend under the PAB regime. Given that UV responsiveness of phenolic compounds depends on the UV levels used and the thallus age (with decreasing responsiveness as age increases), further research using higher UV levels and younger thalli should be conducted to more reliably establish the UV reactiveness of Anthoceros agrestis. © 2018 Adac. Tous droits réservés.

The spatial variability of ultraviolet-absorbing compounds (UVACs) in the freshwater liverwort Jungermannia exsertifolia subsp. cordifolia was studied in mid-latitudes (the Atlantic-Mediterranean transition) across a wide lati-altitudinal gradient, with the aim of testing the usefulness of UVACs as biomarkers of current ambient levels of UV radiation. We analysed 17 samples from streams located in the main mountain ranges of the Iberian Peninsula, differentiating methanol-soluble (SUVACs, mainly located in the vacuoles) and methanol-insoluble (IUVACs, bound to cell walls) compounds, since they represent different manners to cope with UV radiation. In both fractions, the bulk level of UVACs and the concentrations of several individual compounds were measured. In addition, we measured Fv/Fm, DNA damage and sclerophylly index (SI) as possible additional UV biomarkers. UVACs showed a high variability, probably due not only to the gradients of macroenvironmental factors (UV radiation, PAR, and water temperature), but also to microenvironmental factors inherent to the dynamic nature of mountain streams. Two soluble coumarins were positively correlated with UV levels and could be used for ambient UV biomonitoring in the spatial scale. In contrast to the variability in UVACs, the relatively homogeneous values of Fv/Fm and the lack of any DNA damage made these variables useless for ambient UV biomonitoring, but suggested a strong acclimation capacity of this liverwort to changing environmental conditions (in particular, to UV levels). Finally, UVACs of fresh samples of the liverwort were compared to those of herbarium samples collected in the same lati-altitudinal gradient. SUVACs were significantly higher in fresh samples, whereas IUVACs generally showed the contrary. Thus, IUVACs were more stable than SUVACs and hence more adequate for retrospective UV biomonitoring. In conclusion, UVAC compartmentation should be taken into account for bryophyte-based UV biomonitoring in future studies.

Ultraviolet-absorbing compounds (UVACs) were analyzed in 90 herbarium samples of the liverwort Jungermannia exsertifolia subsp. cordifolia collected throughout Spain in the period 1913–2006, in order to evaluate their usefulness in UV radiation biomonitoring. This was assessed, for the first time in bryophytes, differentiating the soluble (mainly vacuolar) and insoluble (cell wall-bound) UVACs (respectively, SUVACs and IUVACs), which may represent different mechanisms to cope with UV radiation. The bulk levels of SUVACs and IUVACs, and the concentrations of seven individual compounds (five SUVACs and two IUVACs) were analyzed, and several environmental variables of the sampling points were obtained: longitude, latitude, altitude, ozone column, and modelled UV levels. Diverse positive relationships of UVAC variables with UV radiation levels were identified in both the temporal and spatial scales. The soluble compound p -coumaroylmalic acid showed significantly higher concentrations in the years after the onset of stratospheric ozone degradation (around 1975) with respect to the previous years. The bulk level and the concentrations of three individual SUVACs were higher in the Spanish samples analyzed in the present study than in the northern European samples analyzed in a previous study in which IUVACs were not considered. The bulk level of IUVACs was the biological variable best spatially correlated with the UV exposure of the samples and with important factors determining it, such as altitude and latitude. Furthermore, the bulk level of IUVACs was the biological variable selected by a stepwise multiple linear regression analysis to reconstruct the past levels of UV-B radiation. The reconstructed UV-B did not show a clear trend in the period 1913–2006, which was coincident with other climatic and biological UV reconstructions. Both SUVACs and IUVACs were significant loading factors differentiating summer and autumn samples in the Principal Components Analysis conducted using both UVACs and the environmental variables, but again the bulk level of IUVACs was the main loading factor differentiating the samples collected from the highest and most UV-exposed localities (Sierra Nevada). In general, these findings suggest that UVACs were stimulated by increased UV levels in time or space, and that IUVACs would be better UV protectants than SUVACs, probably due to their more efficient screening capacity. Overall, UVACs of J. exsertifolia subsp. cordifolia were good UV biomarkers, and their location in the soluble or cell wall-bound fractions determined their usefulness in the different aspects of UV biomonitoring, including the assessment of the UV-B increase associated to anthropogenic ozone degradation.

The available data on the effects of UVB radiation (UVBR) are scarce for submerged macrophytes, particularly charophytes. We studied the effects of UVBR on Chara baltica, Chara hispida, Chara vulgaris, Nitella hyalina and Myriophyllum spicatum, collected from shallow Mediterranean waterbodies. In a short-term laboratory experiment, we subjected these species to three different UVBR treatments corresponding to daily biologically effective integrated doses of 0, 1 and 6 kJ m−2. The analysed response variables were DNA damage, UV-absorbing compounds (both the methanol-soluble–SUVACs- and, for the first time in charophytes, the methanol-insoluble cell wall-bound fraction–WUVACs-), the stoichiometric composition, photosynthetic pigments, metabolic and growth rates and morphological architecture. UVBR induced harmful effects on DNA in all species, but M. spicatum appeared to be the most sensitive. The observed DNA damage only appeared to be UVB-dose dependent for C. vulgaris. The charophytes produced higher WUVACs levels and the angiosperm higher SUVACs levels. Higher chlorophyll and carotenoid concentrations were detected in the presence of UVBR, but their ratio was not modified by the applied treatments. The Chl a/b ratio decreased under UVBR in two charophyte species, whereas it increased in the angiosperm, which also exhibited a greater photosynthetic response. Relative growth rates were negatively influenced by UVBR, although the degree of this influence depending on the dose was different among species. UVBR modified growth patterns, particularly in those species exhibiting apical growth. We observed more species-specific differences than commonalities in the effects of and responses to UVBR across the investigated species.

Premise of the Study: Ultraviolet (UV) radiation influences the viability of algal spores and seed-plant pollen depending on the species, the dose, and the wavelength. In bryophytes, one of the dominant groups of plants in many habitats, UV radiation could determine their spore dispersal strategy, and such data are critical for reconstructing the ancestral state in plants and for determining the distribution range and persistence of bryophyte species. Methods: Spores of four bryophyte species of the moss genus Orthotrichum that were either hygrochastic or xerochastic (spores dispersed under wet or dry conditions, respectively) were exposed to realistic doses of UV radiation under laboratory conditions. Spore viability was evaluated through germination experiments and, for the first time in bryophytes, ultrastructural observations. Given that the UV-B doses used were relatively higher than the UV-A doses, the UV effect was probably due more to UV-B than UV-A wavelengths. Key Results: All four species reduced their spore germination capacity in a UV dose-dependent manner, concomitantly increasing spore ultrastructural damage (cytoplasmic and plastid alterations). Most spores eventually died when exposed to the highest UV dose. Interestingly, spores of hygrochastic species were much more UV-sensitive than those of xerochastic species. Conclusions: UV tolerance determines moss spore viability, as indicated by germination capacity and ultrastructural damage, and differs between spores of species with different dispersal strategies. Specifically, the higher UV tolerance of xerochastic spores may enable them to be dispersed to longer distances than hygrochastic spores, thus extending more efficiently the distribution range of the corresponding species.

The ultraviolet-B (UV-B) photoreceptor UV RESISTANCE LOCUS 8 (UVR8) mediates photomorphogenic responses to UV-B in Arabidopsis through differential gene expression, but little is known about UVR8 in other species. Bryophyte lineages were the earliest diverging embryophytes, thus being the first plants facing the UV-B regime typical of land. We therefore examined whether liverwort and moss species have functional UVR8 proteins and whether they are regulated similarly to Arabidopsis UVR8. We examined the expression, dimer/monomer status, cellular localisation and function of Marchantia polymorpha and Physcomitrella patens UVR8 in experiments with bryophyte tissue and expression of green fluorescent protein (GFP)–UVR8 fusions in Nicotiana leaves and transgenic Arabidopsis. P. patens expresses two UVR8 genes that encode functional proteins, whereas the single M. polymorpha UVR8 gene expresses two transcripts by alternative splicing that encode functional UVR8 variants. P. patens UVR8 proteins form dimers that monomerise and accumulate in the nucleus following UV-B exposure, similar to Arabidopsis UVR8, but M. polymorpha UVR8 has weaker dimers and the proteins appear more constitutively nuclear. We conclude that liverwort and moss species produce functional UVR8 proteins. Although there are differences in expression and regulation of P. patens and M. polymorpha UVR8, the mechanism of UVR8 action is strongly conserved in evolution. © 2017 The Authors. New Phytologist © 2017 New Phytologist Trust

International audience; AbstractKey messageMoss surveys provide spatially dense data on environmental concentrations of heavy metals and nitrogen which, together with other biomonitoring and modelling data, can be used for indicating deposition to terrestrial ecosystems and related effects across time and areas of different spatial extension.ContextFor enhancing the spatial resolution of measuring and mapping atmospheric deposition by technical devices and by modelling, moss is used complementarily as bio-monitor.AimsThis paper investigated whether nitrogen and heavy metal concentrations derived by biomonitoring of atmospheric deposition are statistically meaningful in terms of compliance with minimum sample size across several spatial levels (objective 1), whether this is also true in terms of geostatistical criteria such as spatial auto-correlation and, by this, estimated values for unsampled locations (objective 2) and whether moss indicates atmospheric deposition in a similar way as modelled deposition, tree foliage and natural surface soil at the European and country level, and whether they indicate site-specific variance due to canopy drip (objective 3).MethodsData from modelling and biomonitoring atmospheric deposition were statistically analysed by means of minimum sample size calculation, by geostatistics as well as by bivariate correlation analyses and by multivariate correlation analyses using the Classification and Regression Tree approach and the Random Forests method.ResultsIt was found that the compliance of measurements with the minimum sample size varies by spatial scale and element measured. For unsampled locations, estimation could be derived. Statistically significant correlations between concentrations of heavy metals and nitrogen in moss and modelled atmospheric deposition, and concentrations in leaves, needles and soil were found. Significant influence of canopy drip on nitrogen concentration in moss was proven.ConclusionMoss surveys should complement modelled atmospheric deposition data as well as other biomonitoring approaches and offer a great potential for various terrestrial monitoring programmes dealing with exposure and effects.

A 2-year study explored metabolic and phenotypic plasticity of sun-acclimated Vitis vinifera cv. Pinot noir leaves collected from 12 locations across a 36.69-49.98°N latitudinal gradient. Leaf morphological and biochemical parameters were analysed in the context of meteorological parameters and the latitudinal gradient. We found that leaf fresh weight and area were negatively correlated with both global and ultraviolet (UV) radiation, cumulated global radiation being a stronger correlator. Cumulative UV radiation (sumUVR) was the strongest correlator with most leaf metabolites and pigments. Leaf UV-absorbing pigments, total antioxidant capacities, and phenolic compounds increased with increasing sumUVR, whereas total carotenoids and xanthophylls decreased. Despite of this reallocation of metabolic resources from carotenoids to phenolics, an increase in xanthophyll-cycle pigments (the sum of the amounts of three xanthophylls: violaxanthin, antheraxanthin, and zeaxanthin) with increasing sumUVR indicates active, dynamic protection for the photosynthetic apparatus. In addition, increased amounts of flavonoids (quercetin glycosides) and constitutive β-carotene and α-tocopherol pools provide antioxidant protection against reactive oxygen species. However, rather than a continuum of plant acclimation responses, principal component analysis indicates clusters of metabolic states across the explored 1,500-km-long latitudinal gradient. This study emphasizes the physiological component of plant responses to latitudinal gradients and reveals the physiological plasticity that may act to complement genetic adaptations.

The accumulation of ultraviolet-absorbing compounds (UVACs) within the cell walls of bryophytes provides a spatially uniform filter of ultraviolet-B (UV-B) radiation, and thus represents an important protective mechanism against its adverse effects. Here, the abilities of several cell-wall-degrading enzymes and of alkali (NaOH) in extracting cell-wall-bound UVACs from the aquatic liverwort Jungermannia exsertifolia Steph. subsp. cordifolia (Dumort.) Vaňa were compared, in order to select the most appropriate extraction method and to achieve the most reliable assessment of the degree of protection afforded by UVACs against UV-B. We analysed both the overall level of UVACs and the concentrations of two hydroxycinnamic acids (p-coumaric acid and ferulic acid). The most effective extraction (statistically significant) of cell-wall-bound UVACs, in terms both of their overall levels and of the concentrations of individual compounds, was achieved using alkaline digestion, which was more efficient than any of the...

BACKGROUND: Ultraviolet (UV) radiation induces adaptive responses that can be used for plant production improvement. The aim of this study was to assess the effect of solar UV exclusion on the physiology and phenolic compounds of leaves and berry skins of Vitis vinifera L. cv. Graciano under field conditions. Phenolic compounds were analyzed globally and individually in both the vacuolar fraction and, for the first time in grapevine, the cell wall-bound fraction. These different locations may represent diverse modalities of phenolic response to and protection against UV. RESULTS: UV exclusion led to a decrease in Fv/Fm in leaves, revealing that solar UV is needed for adequate photoprotection. Only p-caffeoyl-tartaric acid from the soluble fraction of leaves and myricetin-3-O-glucoside from the soluble fraction of berry skins were significantly higher in the presence of UV radiation, and thus they could play a role in UV protection. Other hydroxycinnamic acids, flavonols, flavanols and stilbenes did not respond to UV exclusion. CONCLUSION: UV exclusion led to subtle changes in leaves and berry skins of Graciano cultivar, which would be well adapted to current UV levels. This may help support decision-making on viticultural practices modifying UV exposure of leaves and berries, which could improve grape and wine quality. © 2014 Society of Chemical Industry

We studied the among- and within-genus variability of UV protection in 23 mosses belonging to the genera Andreaea, Grimmia and Racomitrium, which were collected from sun-exposed rocks in northern and central Portugal mountains. We explored the influence of genus, species and environmental factors on their UV-absorption capacity, differentiating the soluble UV-absorbing compounds (SUVAC, mainly located in the vacuoles) and the insoluble ones (WUVAC, bound to the cell wall). All the physiological variables were significantly affected by both the genus and the species, which clearly reflects the influence of genetics on the levels and compartmentation of UV-absorbing compounds (UVAC). The physiological variables analyzed were also significantly influenced by both micro- (for example, substrate and orientation) and macro-scale (chain of mountains) variables. In addition, this study showed the interaction between genetics and the environmental factors known to enhance UV exposure (for example, higher ...

We studied the relationships between environmental variables and the physiology of two (semi-)aquatic mosses (Bryum pseudotriquetrum and Fontinalis antipyretica) in an unforested headwater stream over a three year period. Neither environmental or physiological variables showed significant interannual variations. Most environmental variables (water temperature, stratospheric ozone, and photosynthetic, UV-A and UV-B radiation) showed distinct seasonal variations, but only a few physiological ones did. In both species, photoprotection variables (the activity of the xanthophyll cycle and the bulk UV absorbance of the methanol-extractable UV-absorbing compounds, MEUVAC) varied more seasonally than variables related to physiological activity, such as the sclerophylly index and chlorophyll fluorescence parameters (Fv/Fm and ΦPSII). Changes in physiological activity would be attenuated by the buffering capacity of water with respect to the influence of environmental factors, and dynamic variables like Fv...

We analyzed bulk UV absorbance of methanolic extracts and levels of five UV-absorbing compounds (hydroxycinnamic acid derivatives) in 135 herbarium samples of the liverwort Jungermannia exsertifolia subsp. cordifolia from northern Europe. Samples had been collected in 1850-2006 (96% in June-August). Both UV absorbance and compound levels were correlated positively with collection year. p-Coumaroylmalic acid (C1) was the only compound showing a significant (and negative) correlation with stratospheric ozone and UV irradiance in the period that real data of these variables existed. Stratospheric ozone reconstruction (1850-2006) based on C1 showed higher values in June than in July and August, which coincides with the normal monthly variation of ozone. Combining all the data, there was no long-term temporal trend from 1850 to 2006. Reconstructed UV showed higher values in June-July than in August, but again no temporal trend was detected in 1918-2006 using the joint data. This agrees with previous UV reconstructions.

We analyzed the form of the UV-absorption spectra and calculated the bulk UV-absorption capacity of the methanolic extracts (BUVACME) in 15 bryophytes (five liverworts and 10 mosses) from unshaded aquatic habitats of Tierra del Fuego (Argentina). BUVACME was obtained as the area under the absorbance curve (AUC) in several ranges of the UV band to allow comparison with the literature data. Both the spectra form and the AUC values depended on the species considered. The spectra showed either no, one or two defined peaks. The peaks were probably due to phenolic derivatives, which could act as both screening compounds and antioxidants. The different AUCs calculated were highly and significantly correlated, and thus it may be unimportant which of these is used. The AUC values of most liverworts were higher than those of most mosses and in particular, the liverworts Noteroclada confluens and Triandrophyllum subtrifidum showed much higher BUVACMEs than those analyzed in any other bryophyte. Thus, the acc...

The interaction between UV radiation and mineral availability has not been investigated in bryophytes, despite the important peculiarities of their mineral nutrition. The question under investigation was whether an improvement in phosphorus (P) availability, and the consequent increase in tissue P concentration, modified the responses to enhanced UV radiation in two aquatic bryophytes (the liverwort Jungermannia exsertifolia subsp. cordifolia and the moss Fontinalis antipyretica) under laboratory conditions. These responses were evaluated in terms of photosynthetic pigment composition, chlorophyll fluorescence, photosynthesis and respiration rates and the accumulation of protecting UV-absorbing compounds (both the commonly used bulk UV-absorbance of methanol extracts in both species and the concentrations of five hydroxycinnamic acid derivatives in the liverwort). The culture period (36 days) affected significantly most variables of both species. However, no variable in either species showed sign...

We examined, under laboratory conditions, the physiological responses of the aquatic liverwort Jungermannia exsertifolia subsp. cordifolia to artificially enhanced ultraviolet (UV) radiation for 82 days, especially considering the responses of five hydroxycinnamic acid derivatives. This species lives in mountain streams, where it is exposed to low temperatures and high UV levels, and this combination is believed to increase the adverse effects of UV. Enhanced UV radiation hardly caused any change in several physiological variables indicative of vitality, such as Fv/Fm and chlorophylls/phaeopigments ratio (OD430/OD410). Thus, this liverwort seemed to be tolerant to UV radiation, probably due to the accumulation of three UV-absorbing hydroxycinnamic acid derivatives: p-coumaroylmalic acid, 5''-(7'',8''-dihydroxycoumaroyl)-2-caffeoylmalic acid, and 5''-(7'',8''-dihydroxy-7-O-beta-glucosyl-coumaroyl)-2-caffeoylmalic acid. These compounds might serve as bioindicators of enhanced UV radiation.

The aquatic liverwort Jungermannia exsertifolia subsp. cordifolia was cultivated for 15 d under controlled conditions to study the single and combined effects of cadmium and enhanced ultraviolet (UV) radiation. Both cadmium and UV radiation caused chlorophyll degradation and a decrease in the maximum quantum yield of photosystem II (PSII), together with an increase in the mechanisms of non-photochemical dissipation of energy (increase in the xanthophyll index). Cadmium was more stressing than UV radiation, since the metal also influenced photosynthesis globally and caused a decrease in net photosynthetic rates, in the effective quantum yield of photosynthetic energy conversion of PSII, and in the maximal apparent electron transport rate through PSII. Ultraviolet radiation increased the level of trans-p-coumaroylmalic acid and cadmium increased trans-phaselic and feruloylmalic acids. The increase in these compounds was probably related to both a more efficient absorption of harmful UV radiation and an enhanced protection against oxidative stress. DNA damage was specifically caused by UV-B radiation, but was intensified under the presence of cadmium, probably because the metal impairs the DNA enzymatic repair mechanisms. Ultraviolet radiation and cadmium seemed to operate additively on some physiological processes, while other responses were probably due to either factor alone.

Two kinds of parasitic fungal infections are described in the rhizoidal systems of both wild and cultured materials of Funaria hygrometrica, Bryum capillare and B. pseudotriquetrum. Protonemal and rhizoidal cells of Funaria produce pegs of host wall materialaroundsites of hyphal penetration. Similar structures appear to be much more widespread in hepatics and in Monoclea have been misinterpreted as rudimentary pegs, the characteristic feature ofpeggedrhizoids.Swollen side branch initials and the tip cells of major rhizoidal axescontaina single rozellopsidalean (Oomycota formerly Chytridiomycota) zoosporangium in Bryum pseudotriquetrum and up to three similar structures in B. capillare. Each zoosporangium produces a single exit tube. Whereas the infected cells die the subapical cells remain alive and,afterzoospore discharge, produce side branches whose tip cells also become infected. Restriction of the parasites to sidebranchinitials and rhizoid tip cells is perhaps because these are the only cells...

We examined, under laboratory conditions, the influence of temperature (2 °C vs. 10 °C) on the physiological responses of two aquatic bryophytes from a mountain stream to artificially enhanced UV-B radiation for 82 d. These organisms may be exposed naturally to relatively low temperatures and high levels of UV-B radiation, and this combination is believed to increase the adverse effects of UV-B radiation. In the moss Fontinalis antipyretica, UV-B-treated samples showed severe physiological damages, including significant decreases in chlorophyll (Chl) and carotenoid (Car) contents, Chl a/b and Chl/phaeopigment ratios, Chl a fluorescence parameters Fv/Fm and ΦPS2, electron transport rate (ETRmax), and growth. In the liverwort Jungermannia cordifolia, UV-B radiation hardly caused any physiological change except for growth reduction. Thus, this liverwort seemed to be more tolerant to UV-B radiation than the moss under the specific experimental conditions used, maybe partly due to the accumulation of UV-B absorbing compounds. The influence of temperature on the effects of UV-B radiation depended on the species: the higher the UV-B tolerance, the lower the influence of temperature. Also, different physiological variables showed varied responses to this influence. Particularly, the lower temperature used in our study enhanced the adverse effects of UV-B radiation on important physiological variables such as Fv/Fm, growth, and Chl/phaeopigment ratios in the UV-B-sensitive F. antipyretica, but not in the more UV-B-tolerant J. cordifolia. Thus, the adverse effects of cold and UV-B radiation were apparently additive in the moss, but this additiveness was lacking in the liverwort. The Principal Components Analyses (PCA) conducted for both species with the physiological data obtained after 36 and 82 d of culture confirmed the above results. Under natural conditions, the relatively high water temperatures in summer might facilitate the acclimation of aquatic bryophytes from mountain streams to high levels of UV-B radiation. This may be relevant to predict the consequences of concomitant global warming and increasing UV-B radiation.

Two aquatic bryophytes, the moss Fontinalis antipyretica Hedw. and the foliose liverwort Jungermannia exsertifolia Steph. subsp. cordifolia (Dumort.) Vana were analysed with regard to their ultraviolet-B (UV-B) tolerance. The plants were cultivated in the laboratory for 36 days under three different radiation regimes which were set up using appropriate lamps and filters: P (PAR alone, control), PA (PAR + UV-A) and PAB (PAR + UV-A + UV-B). The two bryophytes responded differently to the enhancement in UV-B radiation, while UV-A radiation had only a slight biological effect. The samples of the moss which were irradiated with UV-B showed, with respect to the control, decreases in the chlorophyll and carotenoids concentration, the chlorophyll a/b quotient, the chlorophylls/phaeopigments ratios, the net photosynthesis rates, the light saturation point, the maximum quantum yield of photosystem II (Fv/Fm) and the apparent electron transport rate (ETR). They also showed increases in the sclerophylly index...

The effects of four increasing levels of KH2PO4 on the physiology of the aquatic liverwort Jungermannia exsertifolia subsp. cordifolia were analyzed in the laboratory in the short term (15 d). The accumulation of P and K in the liverwort tissues was influenced (ANOVA) by the level of KH2PO4-enrichment and was significantly higher in the more enriched culture solutions. However, only the concentration of P was influenced by the effect of time (ANOVA), and the gradual P accumulation throughout the culture period contrasted with the fluctuations observed in K accumulation; these were presumably due to the higher liability of K to be leaked from the cells. Our results suggest that the analysis of P in transplants of J. cordifolia may be a useful bioindicator of short term water eutrophication both in the spatial and the temporal scales, although the highest PO43− concentration used in this study (20 mg liter−1) may induce a P saturation in J. cordifolia tissue (0.53% DM). The rates of net photosynthe...

12 Pags.- 4 Tab.- 6 Fig. - Available online 2 July 2014. - The definitive version is available at: http://www.sciencedirect.com/science/article/pii/S009884721400166X, Climate models predict an increase in ultraviolet (UV) radiation and a reduction in precipitation in the Mediterranean region in the coming decades. High levels of UV radiation and water shortage can both cause photo-oxidative stress in plants. The aim of this study was to investigate the effects of enhanced UV radiation and its interaction with low water availability on seedling performance, biomass production, and photoprotective mechanisms of the sclerophyllous evergreen species Laurus nobilis L. (laurel). To achieve this goal, one-year-old seedlings of L. nobilis were grown outdoors under three UV conditions (ambient UV, enhanced UV-A, and enhanced UV-A + UV-B) and under two watering regimes (watered to field capacity and reduced water supply). The results show that plants produced more biomass when exposed to above ambient levels of UV-A or UV-A + UV-B radiation, especially under low water availability. This was probably related to a UV-induced increase in leaf relative water content and in leaf water use efficiency under water shortage. Even though our results suggest that UV-A supplementation may play an important role in the stimulation of biomass production, plants grown under enhanced UV-A plots showed higher levels of energy dissipation as heat (measured as NPQ) and a higher de-epoxidation state of the violaxanthin cycle. This suggests a greater excess of light energy under UV-A supplementation, in accordance with the observed reduction in the foliar content of light-absorbing pigments in these plants. Strikingly, the addition of UV-B radiation mitigated these effects. In conclusion, UV enhancement might benefit water status and growth of L. nobilis seedlings, especially under low water availability. The results also indicate the activation of different plant response mechanisms to UV-A and UV-B radiation, which would interact to produce the overall plant response., This research was supported by the following projects: CGL2007-64583, CGC2010-17172, Consolider Ingenio Montes (CSD2008-00040) and CGL2011-26977 funded by the Spanish government and by the SGR 2009-458 project funded by the Catalan government.

Ultraviolet (UV) radiation induces adaptive responses that can be used for plant production improvement. The aim of this study was to assess the effect of solar UV exclusion on the physiology and phenolic compounds of leaves and berry skins of Vitis vinifera L. cv. Graciano under field conditions. Phenolic compounds were analyzed globally and individually in both the vacuolar fraction and, for the first time in grapevine, the cell wall-bound fraction. These different locations may represent diverse modalities of phenolic response to and protection against UV.UV exclusion led to a decrease in Fv /Fm in leaves, revealing that solar UV is needed for adequate photoprotection. Only p-caffeoyl-tartaric acid from the soluble fraction of leaves and myricetin-3-O-glucoside from the soluble fraction of berry skins were significantly higher in the presence of UV radiation, and thus they could play a role in UV protection. Other hydroxycinnamic acids, flavonols, flavanols and stilbenes did not respond to UV exclusion.UV exclusion led to subtle changes in leaves and berry skins of Graciano cultivar, which would be well adapted to current UV levels. This may help support decision-making on viticultural practices modifying UV exposure of leaves and berries, which could improve grape and wine quality.