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1.  Persistent Polar Depletion of Stratospheric Ozone and Emergent Mechanisms of Ultraviolet Radiation-Mediated Health Dysregulation 
Reviews on environmental health  2012;27(0):103-116.
Year 2011 noted the first definable ozone “hole” in the Arctic region, serving as an indicator to the continued threat of dangerous ultraviolet radiation (UVR) exposure caused by the deterioration of stratospheric ozone in the northern hemisphere. Despite mandates of the Montreal Protocol to phase out the production of ozone depleting chemicals (ODCs), the relative stability of ODCs validates popular notions of persistent stratospheric ozone for several decades. Moreover, increased UVR exposure through stratospheric ozone depletion is occurring within a larger context of physiological stress and climate change across the biosphere. In this review, we provide commentaries on stratospheric ozone depletion with relative comparisons between the well-known Antarctic ozone hole and the newly defined ozone hole in the Arctic. Compared to the Antarctic region, increased UVR exposure in the Northern Hemisphere poses a threat to denser human populations across North America, Europe and Asia. In this context, we discuss emerging targets of UVR exposure that can potentially offset normal biological rhythms in terms of taxonomically conserved photoperiod dependent seasonal signaling and entrainment of circadian clocks. Consequences of seasonal shifts during critical life history stages can alter the fitness and condition, while circadian disruption is increasingly becoming associated as a causal link to increased carcinogenesis. We further review the significance of genomic alterations via UVR induced modulations of phase I and phase II transcription factors, the aryl hydrocarbon receptor (AhR) and the Nuclear factor (erythroid-derived 2)-like 2 (Nrf2), with emphasis on mechanism that can lead to metabolic shifts and cancer. While concern for adverse health consequences due to increased UVR exposure are longstanding, recent advances in biochemical research suggest that AhR and Nrf2 transcriptional regulators are likely targets for UVR mediated dysregulations of rhymicity and homeostasis among animals, including humans.
PMCID: PMC3768272  PMID: 23023879
biological rhythmicity; seasonal cycling; circadian rhythms; AhR& Nrf2 transcription factors; carcinogenicity; climate change
2.  Predicted changes in vegetation structure affect the susceptibility to invasion of bryophyte-dominated subarctic heath 
Annals of Botany  2011;108(1):177-183.
Background and Aims
A meta-analysis of global change experiments in arctic tundra sites suggests that plant productivity and the cover of shrubs, grasses and dead plant material (i.e. litter) will increase and the cover of bryophytes will decrease in response to higher air temperatures. However, little is known about which effects these changes in vegetation structure will have on seedling recruitment of species and invasibility of arctic ecosystems.
A field experiment was done in a bryophyte-dominated, species-rich subarctic heath by manipulating the cover of bryophytes and litter in a factorial design. Three phases of seedling recruitment (seedling emergence, summer seedling survival, first-year recruitment) of the grass Anthoxanthum alpinum and the shrub Betula nana were analysed after they were sown into the experimental plots.
Key Results
Bryophyte and litter removal significantly increased seedling emergence of both species but the effects of manipulations of vegetation structure varied strongly for the later phases of recruitment. Summer survival and first-year recruitment were significantly higher in Anthoxanthum. Although bryophyte removal generally increased summer survival and recruitment, seedlings of Betula showed high mortality in early August on plots where bryophytes had been removed.
Large species-specific variation and significant effects of experimental manipulations on seedling recruitment suggest that changes in vegetation structure as a consequence of global warming will affect the abundance of grasses and shrubs, the species composition and the susceptibility to invasion of subarctic heath vegetation.
PMCID: PMC3119609  PMID: 21624960
Anthoxanthum alpinum; Betula nana; bryophytes; deciduous shrubs; global warming; graminoids; invasibility; litter; recruitment; seedling emergence
3.  Arctic Climate Tipping Points 
Ambio  2012;41(1):10-22.
There is widespread concern that anthropogenic global warming will trigger Arctic climate tipping points. The Arctic has a long history of natural, abrupt climate changes, which together with current observations and model projections, can help us to identify which parts of the Arctic climate system might pass future tipping points. Here the climate tipping points are defined, noting that not all of them involve bifurcations leading to irreversible change. Past abrupt climate changes in the Arctic are briefly reviewed. Then, the current behaviour of a range of Arctic systems is summarised. Looking ahead, a range of potential tipping phenomena are described. This leads to a revised and expanded list of potential Arctic climate tipping elements, whose likelihood is assessed, in terms of how much warming will be required to tip them. Finally, the available responses are considered, especially the prospects for avoiding Arctic climate tipping points.
PMCID: PMC3357822  PMID: 22270703
Arctic; Tipping points; Sea-ice; Greenland ice sheet; Atlantic thermohaline circulation; Boreal forest
4.  Changes in Phenolic Compounds and Cellular Ultrastructure of Arctic and Antarctic Strains of Zygnema (Zygnematophyceae, Streptophyta) after Exposure to Experimentally Enhanced UV to PAR Ratio 
Microbial Ecology  2012;65(1):68-83.
Ultraviolet (UV) radiation has become an important stress factor in polar regions due to anthropogenically induced ozone depletion. Although extensive research has been conducted on adaptations of polar organisms to this stress factor, few studies have focused on semi-terrestrial algae so far, in spite of their apparent vulnerability. This study investigates the effect of UV on two semi-terrestrial arctic strains (B, G) and one Antarctic strain (E) of the green alga Zygnema, isolated from Arctic and Antarctic habitats. Isolates of Zygnema were exposed to experimentally enhanced UV A and B (predominant UV A) to photosynthetic active radiation (PAR) ratio. The pigment content, photosynthetic performance and ultrastructure were studied by means of high-performance liquid chromatography (HPLC), chlorophyll a fluorescence and transmission electron microscopy (TEM). In addition, phylogenetic relationships of the investigated strains were characterised using rbcL sequences, which determined that the Antarctic isolate (E) and one of the Arctic isolates (B) were closely related, while G is a distinct lineage. The production of protective phenolic compounds was confirmed in all of the tested strains by HPLC analysis for both controls and UV-exposed samples. Moreover, in strain E, the content of phenolics increased significantly (p = 0.001) after UV treatment. Simultaneously, the maximum quantum yield of photosystem II photochemistry significantly decreased in UV-exposed strains E and G (p < 0.001), showing a clear stress response. The phenolics were most probably stored at the cell periphery in vacuoles and cytoplasmic bodies that appear as electron-dense particles when observed by TEM after high-pressure freeze fixation. While two strains reacted moderately on UV exposure in their ultrastructure, in strain G, damage was found in chloroplasts and mitochondria. Plastidal pigments and xanthophyll cycle pigments were investigated by HPLC analysis; UV A- and UV B-exposed samples had a higher deepoxidation state as controls, particularly evident in strain B. The results indicate that phenolics are involved in UV protection of Zygnema and also revealed different responses to UV stress across the three strains, suggesting that other protection mechanisms may be involved in these organisms.
PMCID: PMC3541927  PMID: 22903087
5.  The response of Arctic vegetation and soils following an unusually severe tundra fire 
Fire causes dramatic short-term changes in vegetation and ecosystem function, and may promote rapid vegetation change by creating recruitment opportunities. Climate warming likely will increase the frequency of wildfire in the Arctic, where it is not common now. In 2007, the unusually severe Anaktuvuk River fire burned 1039 km2 of tundra on Alaska's North Slope. Four years later, we harvested plant biomass and soils across a gradient of burn severity, to assess recovery. In burned areas, above-ground net primary productivity of vascular plants equalled that in unburned areas, though total live biomass was less. Graminoid biomass had recovered to unburned levels, but shrubs had not. Virtually all vascular plant biomass had resprouted from surviving underground parts; no non-native species were seen. However, bryophytes were mostly disturbance-adapted species, and non-vascular biomass had recovered less than vascular plant biomass. Soil nitrogen availability did not differ between burned and unburned sites. Graminoids showed allocation changes consistent with nitrogen stress. These patterns are similar to those seen following other, smaller tundra fires. Soil nitrogen limitation and the persistence of resprouters will likely lead to recovery of mixed shrub–sedge tussock tundra, unless permafrost thaws, as climate warms, more extensively than has yet occurred.
PMCID: PMC3720061  PMID: 23836794
Alaskan tussock tundra; fire; vegetation recovery; permafrost; climate change; soil N availability
6.  Interactions between herbivory and warming in aboveground biomass production of arctic vegetation 
BMC Ecology  2008;8:17.
Many studies investigating the ecosystem effects of global climate change have focused on arctic ecosystems because the Arctic is expected to undergo the earliest and most pronounced changes in response to increasing global temperatures, and arctic ecosystems are considerably limited by low temperatures and permafrost. In these nutrient limited systems, a warmer climate is expected to increase plant biomass production, primarily through increases in shrubs over graminoids and forbs. But, the influence of vertebrate and invertebrate herbivores has been largely absent in studies investigating the effects of vegetation responses to climate change, despite the fact that herbivory can have a major influence on plant community composition, biomass and nutrient cycling. Here, we present results from a multi-annual field experiment investigating the effects of vertebrate herbivory on plant biomass response to simulated climate warming in arctic Greenland.
The results after four years of treatments did not give any clear evidence of increased biomass of shrubs in response climate warming. Nor did our study indicate that vertebrate grazing mediated any increased domination of shrubs over other functional plant groups in response to warming. However, our results indicate an important role of insect outbreaks on aboveground biomass. Intense caterpillar foraging from a two-year outbreak of the moth Eurois occulta during two growing seasons may have concealed any treatment effects. However, there was some evidence suggesting that vertebrate herbivores constrain the biomass production of shrubs over graminoids and forbs.
Although inconclusive, our results were likely constrained by the overwhelming influence of an unexpected caterpillar outbreak on aboveground biomass. It is likely that the role of large vertebrate herbivores in vegetation response to warming will become more evident as this experiment proceeds and the plant community recovers from the caterpillar outbreak. Due to the greater influence of invertebrate herbivory in this study, it is advisable to consider both the effect of invertebrate and vertebrate herbivores in studies investigating climate change effects on plant communities.
PMCID: PMC2576048  PMID: 18945359
7.  MabCent: Arctic marine bioprospecting in Norway 
Phytochemistry Reviews  2012;12:567-578.
The deep waters surrounding the coastline of the northern parts of Norway represent an exciting biotope for marine exploration. Dark and cold Arctic water generates a hostile environment where the ability to adapt is crucial to survival. These waters are nonetheless bountiful and a diverse plethora of marine organisms thrive in these extreme conditions, many with the help of specialised chemical compounds. In comparison to warmer, perhaps more inviting shallower tropical waters, the Arctic region has not been as thoroughly investigated. MabCent is a Norwegian initiative based in Tromsø that aims to change this. Since 2007, scientists within MabCent have focussed their efforts on the study of marine organisms inhabiting the Arctic waters with the long term goal of novel drug discovery and development. The activities of MabCent are diverse and range from sampling the Arctic ice shelf to the chemical synthesis of promising secondary metabolites discovered during the screening process. The current review will present the MabCent pipeline from isolation to identification of new bioactive marine compounds via an extensive screening process. An overview of the main activities will be given with particular focus on isolation strategies, bioactivity screening and structure determination. Pitfalls, hard earned lessons and the results so far are also discussed.
PMCID: PMC3777186  PMID: 24078803
Pharmacognosy; Arctic; Cold adaption; Psychrophile; Screening; Bioprospecting; Norway; MabCent
8.  Major transitions in the evolution of early land plants: a bryological perspective 
Annals of Botany  2012;109(5):851-871.
Molecular phylogeny has resolved the liverworts as the earliest-divergent clade of land plants and mosses as the sister group to hornworts plus tracheophytes, with alternative topologies resolving the hornworts as sister to mosses plus tracheophytes less well supported. The tracheophytes plus fossil plants putatively lacking lignified vascular tissue form the polysporangiophyte clade.
This paper reviews phylogenetic, developmental, anatomical, genetic and paleontological data with the aim of reconstructing the succession of events that shaped major land plant lineages.
Fundamental land plant characters primarily evolved in the bryophyte grade, and hence the key to a better understanding of the early evolution of land plants is in bryophytes. The last common ancestor of land plants was probably a leafless axial gametophyte bearing simple unisporangiate sporophytes. Water-conducting tissue, if present, was restricted to the gametophyte and presumably consisted of perforate cells similar to those in the early-divergent bryophytes Haplomitrium and Takakia. Stomata were a sporophyte innovation with the possible ancestral functions of producing a transpiration-driven flow of water and solutes from the parental gametophyte and facilitating spore separation before release. Stomata in mosses, hornworts and polysporangiophytes are viewed as homologous, and hence these three lineages are collectively referred to as the ‘stomatophytes’. An indeterminate sporophyte body (the sporophyte shoot) developing from an apical meristem was the key innovation in polysporangiophytes. Poikilohydry is the ancestral condition in land plants; homoiohydry evolved in the sporophyte of polysporangiophytes. Fungal symbiotic associations ancestral to modern arbuscular mycorrhizas evolved in the gametophytic generation before the separation of major present-living lineages. Hydroids are imperforate water-conducting cells specific to advanced mosses. Xylem vascular cells in polysporangiophytes arose either from perforate cells or de novo. Food-conducting cells were a very early innovation in land plant evolution. The inferences presented here await testing by molecular genetics.
PMCID: PMC3310499  PMID: 22356739
Bryophytes; cuticle; homoiohydry; mycorrhizas; phylogeny; poikilohydry; polysporangiophytes; shoot apical meristem (SAM); stomata; stomatophytes; tracheophytes; vascular tissues
9.  Dividing without centrioles: innovative plant microtubule organizing centres organize mitotic spindles in bryophytes, the earliest extant lineages of land plants 
AoB Plants  2011;2011:plr028.
Triple staining of γ-tubulin, microtubules, and nuclei reveal that three types of MTOCs initiate spindles in bryophytes. Polar organizers in liverworts and plastid MTOCs in hornworts are unique and nuclear envelope MTOCs in mosses appear like those in seed plants.
Background and aims
As remnants of the earliest land plants, the bryophytes (liverworts, mosses and hornworts) are important in understanding microtubule organization in plant cells. Land plants have an anastral mitotic spindle that forms in the absence of centrosomes, and a cytokinetic apparatus comprised of a predictive preprophase band (PPB) before mitosis and a phragmoplast after mitosis. These microtubule arrays have no counterpart in animal cells and the nature of the plant microtubule organizing centre (MTOC) remained an enigma for many years until antibodies to γ-tubulin, an essential component of the MTOC in all eukaryotes, became available for tracing the origin of microtubule arrays.
We used immunofluorescence techniques to colocalize γ-tubulin, microtubules and chromosomes in mitotic cells of a representative liverwort, moss and hornwort to study the organization of microtubules during mitotic cell division.
Principal results
The future division site is marked by a PPB in all taxa but the MTOCs initially generating the half spindles differ: polar organizers in the liverwort, plastid MTOCs in the hornwort, and nuclear envelope-associated MTOCs in the moss. By mid-prophase, the forming spindles become more similar as γ-tubulin begins to spread around the polar regions of the nuclear envelope.
Regardless of origin, mature metaphase spindles are identical and indistinguishable from the typical anastral spindle of higher plants with broad polar regions consisting of numerous subsets of converging microtubules. A curious phenomenon of plant spindles, true of bryophytes as well as higher plants, is the movement of γ-tubulin into the metaphase spindle itself. The bipolar arrays of phragmoplast microtubules are organized by diffuse γ-tubulin located at proximal surfaces of reforming nuclear envelopes. Phragmoplast development appears similar in the three taxa and to vascular plants as well.
PMCID: PMC3240993  PMID: 22476498
10.  UVB Radiation as a Potential Selective Factor Favoring Microcystin Producing Bloom Forming Cyanobacteria 
PLoS ONE  2013;8(9):e73919.
Due to the stratospheric ozone depletion, several organisms will become exposed to increased biologically active UVB (280–320 nm) radiation, not only at polar but also at temperate and tropical latitudes. Bloom forming cyanobacteria are exposed to UVB radiation on a mass scale, particularly during the surface bloom and scum formation that can persist for long periods of time. All buoyant species of cyanobacteria are at least periodically exposed to higher irradiation during their vertical migration to the surface that usually occurs several times a day. The aim of this study is to assess the influence on cyanobacteria of UVB radiation at realistic environmental intensities. The effects of two UVB intensities of 0.5 and 0.99 W/m2 in up to 0.5 cm water depth were studied in vitro on Microcystis aeruginosa strains, two microcystin producing and one non-producing. After UVB exposure their ability to proliferate was estimated by cell counting, while cell fitness and integrity were evaluated using light microscopy, autofluorescence and immunofluorescence. Gene damage was assessed by TUNEL assay and SYBR Green staining of the nucleoide area. We conclude that UVB exposure causes damage to the genetic material, cytoskeletal elements, higher sedimentation rates and consequent cell death. In contrast to microcystin producers (PCC7806 and FACHB905), the microcystin non-producing strain PCC7005 is more susceptible to the deleterious effects of radiation, with weak recovery ability. The ecological relevance of the results is discussed using data from eleven years’ continuous UVB radiation measurements within the area of Ljubljana city (Slovenia, Central Europe). Our results suggest that increased solar radiation in temperate latitudes can have its strongest effect during cyanobacterial bloom formation in spring and early summer. UVB radiation in this period may significantly influence strain composition of cyanobacterial blooms in favor of microcystin producers.
PMCID: PMC3772863  PMID: 24058503
11.  Levoglucosan indicates high levels of biomass burning aerosols over oceans from the Arctic to Antarctic 
Scientific Reports  2013;3:3119.
Biomass burning is known to affect air quality, global carbon cycle, and climate. However, the extent to which biomass burning gases/aerosols are present on a global scale, especially in the marine atmosphere, is poorly understood. Here we report the molecular tracer levoglucosan concentrations in marine air from the Arctic Ocean through the North and South Pacific Ocean to Antarctica during burning season. Levoglucosan was found to be present in all regions at ng/m3 levels with the highest atmospheric loadings present in the mid-latitudes (30°–60° N and S), intermediate loadings in the Arctic, and lowest loadings in the Antarctic and equatorial latitudes. As a whole, levoglucosan concentrations in the Southern Hemisphere were comparable to those in the Northern Hemisphere. Biomass burning has a significant impact on atmospheric Hg and water-soluble organic carbon (WSOC) from pole-to-pole, with more contribution to WSOC in the Northern Hemisphere than in the Southern Hemisphere.
PMCID: PMC3814847  PMID: 24176935
12.  Long-Term Effects of Grazing and Global Warming on the Composition and Carrying Capacity of Graminoid Marshes for Moulting Geese in East Greenland 
Ambio  2011;40(6):638-649.
Greening of the Arctic due to climate warming may provide herbivores with richer food supplies, resulting in higher herbivore densities. In turn, this may cause changes in vegetation composition and ecosystem function. In 1982–1984, we studied the ecology of non-breeding moulting geese in Jameson Land, low Arctic East Greenland. By then, geese consumed most of the graminoid production in available moss fens, and it appeared that the geese had filled up the available habitat. In 2008, we revisited the area and found that the number of moulting geese and the temperature sum for June–July had tripled, while the above-ground biomass in a moss fen ungrazed by geese had more than doubled. In a goose-grazed fen, the overall plant composition was unchanged, but the frequency of graminoids had decreased and the area with dead vegetation and open spots had increased. We suggest that climate warming has lead to increased productivity, allowing for higher numbers of moulting geese. However, the reduction of vegetation cover by grazing may have longer term negative consequences for the number of geese the habitat can sustain.
PMCID: PMC3357869  PMID: 21954726
Barnacle geese; Carex; Climate change; Grazing impact; Herbivory; Pink-footed geese
13.  Altitudinal changes in temperature responses of net photosynthesis and dark respiration in tropical bryophytes 
Annals of Botany  2012;111(3):455-465.
Background and Aims
There is a conspicuous increase of poikilohydric organisms (mosses, liverworts and macrolichens) with altitude in the tropics. This study addresses the hypothesis that the lack of bryophytes in the lowlands is due to high-temperature effects on the carbon balance. In particular, it is tested experimentally whether temperature responses of CO2-exchange rates would lead to higher respiratory carbon losses at night, relative to potential daily gains, in lowland compared with lower montane forests.
Gas-exchange measurements were used to determine water-, light-, CO2- and temperature-response curves of net photosynthesis and dark respiration of 18 tropical bryophyte species from three altitudes (sea level, 500 m and 1200 m) in Panama.
Key Results
Optimum temperatures of net photosynthesis were closely related to mean temperatures in the habitats in which the species grew at the different altitudes. The ratio of dark respiration to net photosynthesis at mean ambient night and day temperatures did not, as expected, decrease with altitude. Water-, light- and CO2-responses varied between species but not systematically with altitude.
Drivers other than temperature-dependent metabolic rates must be more important in explaining the altitudinal gradient in bryophyte abundance. This does not discard near-zero carbon balances as a major problem for lowland species, but the main effect of temperature probably lies in increasing evaporation rates, thus restricting the time available for photosynthetic carbon gain, rather than in increasing nightly respiration rates. Since optimum temperatures for photosynthesis were so fine tuned to habitat temperatures we analysed published temperature responses of bryophyte species worldwide and found the same pattern on the large scale as we found along the tropical mountain slope we studied.
PMCID: PMC3579435  PMID: 23258418
Altitudinal gradient; bryophytes; carbon balance; dark respiration; gas-exchange measurements; hepatics; liverworts; mosses; net photosynthesis; photosynthesis-response curves; temperature; tropical rain forest
14.  One mechanism contributing to co-variability of the Atlantic inflow branches to the Arctic 
Nature Communications  2013;4:1488-.
The two-branched inflow of warm and saline Atlantic Water to the Arctic is the major contributor of oceanic heat to the Arctic climate system. However, while the Atlantic Water entering the Arctic through the Fram Strait retains a large part of its heat as it flows along the Arctic continental slope, the branch flowing through the shallow Barents Sea releases a substantial amount of heat to the atmosphere. Hence, the pathway of the Atlantic Water into the Arctic to a large degree determines the short term fate of its heat. Here we show events in which the relative strengths of the two branches are affected by wind-induced Ekman-transport off the northern Barents Sea shelf. The resulting decrease in sea surface height induces a cyclonic circulation anomaly along the slope encircling the northern Barents Sea shelf area, which enhances the flow through the Barents Sea while weakening the branch flowing along the Arctic continental slope.
The branched inflow of warm Atlantic Water to the Arctic has been known for more than a hundred years, yet what controls the relative strengths of the two pathways remains poorly understood. Here, the authors identify the role of atmospheric circulation over the northern Barents Sea in controlling inflow.
PMCID: PMC3586715  PMID: 23403588
15.  Tipping Elements in the Arctic Marine Ecosystem 
Ambio  2012;41(1):44-55.
The Arctic marine ecosystem contains multiple elements that present alternative states. The most obvious of which is an Arctic Ocean largely covered by an ice sheet in summer versus one largely devoid of such cover. Ecosystems under pressure typically shift between such alternative states in an abrupt, rather than smooth manner, with the level of forcing required for shifting this status termed threshold or tipping point. Loss of Arctic ice due to anthropogenic climate change is accelerating, with the extent of Arctic sea ice displaying increased variance at present, a leading indicator of the proximity of a possible tipping point. Reduced ice extent is expected, in turn, to trigger a number of additional tipping elements, physical, chemical, and biological, in motion, with potentially large impacts on the Arctic marine ecosystem.
PMCID: PMC3357823  PMID: 22270704
Arctic; Tipping points; Ecosystem; Non-linearity; Ice; Plankton
16.  Altitude affects the reproductive performance in monoicous and dioicous bryophytes: examples from a Brazilian Atlantic rainforest 
AoB Plants  2012;2012:pls016.
Species traits, such as breeding system, phylum and growth form and habitat characteristics are shown to influence reproductive performance of liverworts and mosses in the Brazilian Atlantic Rainforest, and drive life-history differentiation among species and populations.
Background and aims
Short life cycles and trade-offs linked to breeding systems make bryophytes good models for the study of plant reproductive strategies. Our aim was to test if differences in sexual reproductive performance of bryophytes in tropical rainforests are driven by the breeding system of the species (monoicous or dioicous) or are mainly affected by the habitat.
The reproductive performance (sexual branches, gametangia (sex organs), fertilization and sporophyte production) of 11 species was repeatedly monitored and analysed from populations at sea-level and montane sites of a Brazilian Atlantic rainforest over 15 months.
Principal results
Monoicous species had the highest reproductive performance, particularly for sexual branches, fertilized gametangia and sporophyte production. Species at the sea-level site produced more sexual branches and had more female-biased sex ratios of gametangia than species in the montane site. Fertilizations were more frequent at the montane site, but sporophyte frequency was similar between the two sites. Fertilization tended to occur mostly in the periods of heavy rain (October to December).
Breeding system is not the only major influence on the reproductive performance of bryophytes. We show that habitat is also an important factor determining life-history differentiation. Female-biased sex ratios and low rates of fertilization are seen to be compensated for by high production of reproductive structures at the initial phases of the reproductive cycle.
PMCID: PMC3401027  PMID: 22822422
17.  Comparative Cryptogam Ecology: A Review of Bryophyte and Lichen Traits that Drive Biogeochemistry 
Annals of Botany  2007;99(5):987-1001.
Recent decades have seen a major surge in the study of interspecific variation in functional traits in comparative plant ecology, as a tool to understanding and predicting ecosystem functions and their responses to environmental change. However, this research has been biased almost exclusively towards vascular plants. Very little is known about the role and applicability of functional traits of non-vascular cryptogams, particularly bryophytes and lichens, with respect to biogeochemical cycling. Yet these organisms are paramount determinants of biogeochemistry in several biomes, particularly cold biomes and tropical rainforests, where they: (1) contribute substantially to above-ground biomass (lichens, bryophytes); (2) host nitrogen-fixing bacteria, providing major soil N input (lichens, bryophytes); (3) control soil chemistry and nutrition through the accumulation of recalcitrant polyphenols (bryophytes) and through their control over soil and vegetation hydrology and temperatures; (4) both promote erosion (rock weathering by lichens) and prevent it (biological crusts in deserts); (5) provide a staple food to mammals such as reindeer (lichens) and arthropodes, with important feedbacks to soils and biota; and (6) both facilitate and compete with vascular plants.
Here we review current knowledge about interspecific variation in cryptogam traits with respect to biogeochemical cycling and discuss to what extent traits and measuring protocols needed for bryophytes and lichens correspond with those applied to vascular plants. We also propose and discuss several new or recently introduced traits that may help us understand and predict the control of cryptogams over several aspects of the biogeochemistry of ecosystems.
Whilst many methodological challenges lie ahead, comparative cryptogam ecology has the potential to meet some of the important challenges of understanding and predicting the biogeochemical and climate consequences of large-scale environmental changes driving shifts in the cryptogam components of vegetation composition.
PMCID: PMC2802918  PMID: 17353205
Biogeochemical processes; carbon; cryptogam; decomposition; defence; functional trait; growth rate; interspecific variation; moss; lichen; liverwort; nutrients
18.  Medical Consequences of Stratospheric Ozone Depletion 
Canadian Family Physician  1989;35:2283-2355.
The absorption of ultraviolet (UV) light by stratospheric ozone is crucial to the provision of an environment suitable for terrestrial life. Ultraviolet radiation is the part of the solar spectrum with wavelengths between 240 and 400 nm. Photons at wavelengths below about 320 nm can produce photochemical damage to the cells of living organisms. Data suggesting a causative role for UV exposure in squamous cell carcinoma, basal cell carcinoma, melanoma, cataract formation, and (as yet) poorly defined immune suppression comes from epidemiologic studies and, in some cases, clinical observation and experimental work. The author concentrates on the direct health effects of UV radiation, although the factors that lead to stratospheric ozone depletion also exert other powerful influences on the biosphere that will have less predictable direct and indirect effects on human health.
PMCID: PMC2280294  PMID: 21248921
carcinoma; cataracts; environmental health; melanoma; oncology; ozone; sunglasses; UV radiation
19.  Biogeography of Deep-Sea Benthic Bacteria at Regional Scale (LTER HAUSGARTEN, Fram Strait, Arctic) 
PLoS ONE  2013;8(9):e72779.
Knowledge on spatial scales of the distribution of deep-sea life is still sparse, but highly relevant to the understanding of dispersal, habitat ranges and ecological processes. We examined regional spatial distribution patterns of the benthic bacterial community and covarying environmental parameters such as water depth, biomass and energy availability at the Arctic Long-Term Ecological Research (LTER) site HAUSGARTEN (Eastern Fram Strait). Samples from 13 stations were retrieved from a bathymetric (1,284–3,535 m water depth, 54 km in length) and a latitudinal transect (∼ 2,500 m water depth; 123 km in length). 454 massively parallel tag sequencing (MPTS) and automated ribosomal intergenic spacer analysis (ARISA) were combined to describe both abundant and rare types shaping the bacterial community. This spatial sampling scheme allowed detection of up to 99% of the estimated richness on phylum and class levels. At the resolution of operational taxonomic units (97% sequence identity; OTU3%) only 36% of the Chao1 estimated richness was recovered, indicating a high diversity, mostly due to rare types (62% of all OTU3%). Accordingly, a high turnover of the bacterial community was also observed between any two sampling stations (average replacement of 79% of OTU3%), yet no direct correlation with spatial distance was observed within the region. Bacterial community composition and structure differed significantly with increasing water depth along the bathymetric transect. The relative sequence abundance of Verrucomicrobia and Planctomycetes decreased significantly with water depth, and that of Deferribacteres increased. Energy availability, estimated from phytodetrital pigment concentrations in the sediments, partly explained the variation in community structure. Overall, this study indicates a high proportion of unique bacterial types on relatively small spatial scales (tens of kilometers), and supports the sampling design of the LTER site HAUSGARTEN to study bacterial community shifts in this rapidly changing area of the world’s oceans.
PMCID: PMC3759371  PMID: 24023770
20.  Sensitivity of Rice to Ultraviolet-B Radiation 
Annals of Botany  2006;97(6):933-942.
• Background Depletion of the stratospheric ozone layer leads to an increase in ultraviolet-B (UVB: 280–320 nm) radiation reaching the earth's surface, and the enhanced solar UVB radiation predicted by atmospheric models will result in reduction of growth and yield of crops in the future. Over the last two decades, extensive studies of the physiological, biochemical and morphological effects of UVB in plants, as well as the mechanisms of UVB resistance, have been carried out.
• Scope In this review, we describe recent research into the mechanisms of UVB resistance in higher plants, with an emphasis on rice (Oryza sativa), one of the world's most important staple food crops. Recent studies have brought to light the following remarkable findings. UV-absorbing compounds accumulating in the epidermal cell layers have traditionally been considered to function as UV filters, and to play an important role in countering the damaging effects of UVB radiation. Although these compounds are effective in reducing cyclobutane pyrimidine dimer (CPD) induction in plants exposed to a challenge exposure to UVB, certain levels of CPD are maintained constitutively in light conditions containing UVB, regardless of the quantity or presence of visible light. These findings imply that the systems for repairing DNA damage and scavenging reactive oxygen species (ROS) are essential for plants to grow in light conditions containing UVB.
• Conclusion CPD photolyase activity is a crucial factor determining the differences in UVB sensitivity between rice cultivars. The substitution of one or two bases in the CPD photolyase gene can alter the activity of the enzyme, and the associated resistance of the plant to UVB radiation. These findings open up the possibility, in the near future, of increasing the resistance of rice to UVB radiation, by selective breeding or bioengineering of the genes encoding CPD photolyase.
PMCID: PMC2803405  PMID: 16520342
Ultraviolet-B radiation (UVB: 280–320 nm); rice (Oryza sativa); cyclobutane pyrimidine dimer (CPD); CPD photolyase; reactive oxygen species (ROS); UV-absorbing compounds; UVB resistance; UVB sensitivity; photorepair; dark repair; bioengineering; selective breeding
21.  Impacts of Ocean Acidification on Sediment Processes in Shallow Waters of the Arctic Ocean 
PLoS ONE  2014;9(4):e94068.
Despite the important roles of shallow-water sediments in global biogeochemical cycling, the effects of ocean acidification on sedimentary processes have received relatively little attention. As high-latitude cold waters can absorb more CO2 and usually have a lower buffering capacity than warmer waters, acidification rates in these areas are faster than those in sub-tropical regions. The present study investigates the effects of ocean acidification on sediment composition, processes and sediment-water fluxes in an Arctic coastal system. Undisturbed sediment cores, exempt of large dwelling organisms, were collected, incubated for a period of 14 days, and subject to a gradient of pCO2 covering the range of values projected for the end of the century. On five occasions during the experimental period, the sediment cores were isolated for flux measurements (oxygen, alkalinity, dissolved inorganic carbon, ammonium, nitrate, nitrite, phosphate and silicate). At the end of the experimental period, denitrification rates were measured and sediment samples were taken at several depth intervals for solid-phase analyses. Most of the parameters and processes (i.e. mineralization, denitrification) investigated showed no relationship with the overlying seawater pH, suggesting that ocean acidification will have limited impacts on the microbial activity and associated sediment-water fluxes on Arctic shelves, in the absence of active bio-irrigating organisms. Only following a pH decrease of 1 pH unit, not foreseen in the coming 300 years, significant enhancements of calcium carbonate dissolution and anammox rates were observed. Longer-term experiments on different sediment types are still required to confirm the limited impact of ocean acidification on shallow Arctic sediment processes as observed in this study.
PMCID: PMC3981760  PMID: 24718610
22.  Two Decades of Experimental Manipulations of Heaths and Forest Understory in the Subarctic 
Ambio  2012;41(Suppl 3):218-230.
Current atmospheric warming due to increase of greenhouse gases will have severe consequences for the structure and functioning of arctic ecosystems with changes that, in turn, may feed back on the global-scale composition of the atmosphere. During more than two decades, environmental controls on biological and biogeochemical processes and possible atmospheric feedbacks have been intensely investigated at Abisko, Sweden, by long-term ecosystem manipulations. The research has addressed questions like environmental regulation of plant and microbial community structure and biomass, carbon and nutrient pools and element cycling, including exchange of greenhouse gases and volatile organic compounds, with focus on fundamental processes in the interface between plants, soil and root-associated and free-living soil microorganisms. The ultimate goal has been to infer from these multi-decadal experiments how subarctic and arctic ecosystems will respond to likely environmental changes in the future. Here we give an overview of some of the experiments and main results.
PMCID: PMC3535062  PMID: 22864696
Tundra; Warming; Field experiments; Plant–microbe interactions; Carbon and nitrogen cycling
23.  Rapid transcriptome responses of maize (Zea mays) to UV-B in irradiated and shielded tissues 
Genome Biology  2004;5(3):R16.
Profiling the transcriptional response of maize tissues to UV-B irradiation suggests that a signal is transmitted from irradiated to shielded tissue. The transcriptional response occurs rapidly, even in shielded tissue.
Depletion of stratospheric ozone has raised terrestrial levels of ultraviolet-B radiation (UV-B), an environmental change linked to an increased risk of skin cancer and with potentially deleterious consequences for plants. To better understand the processes of UV-B acclimation that result in altered plant morphology and physiology, we investigated gene expression in different organs of maize at several UV-B fluence rates and exposure times.
Microarray hybridization was used to assess UV-B responses in directly exposed maize organs and organs shielded by a plastic that absorbs UV-B. After 8 hours of high UV-B, the abundance of 347 transcripts was altered: 285 were increased significantly in at least one organ and 80 were downregulated. More transcript changes occurred in directly exposed than in shielded organs, and the levels of more transcripts were changed in adult compared to seedling tissues. The time course of transcript abundance changes indicated that the response kinetics to UV-B is very rapid, as some transcript levels were altered within 1 hour of exposure.
Most of the UV-B regulated genes are organ-specific. Because shielded tissues, including roots, immature ears, and leaves, displayed altered transcriptome profiles after exposure of the plant to UV-B, some signal(s) must be transmitted from irradiated to shielded tissues. These results indicate that there are integrated responses to UV-B radiation above normal levels. As the same total UV-B irradiation dose applied at three intensities elicited different transcript profiles, the transcriptome changes exhibit threshold effects rather than a reciprocal dose-effect response. Transcriptome profiling highlights possible signaling pathways and molecules for future research.
PMCID: PMC395766  PMID: 15003119
24.  Hormonal regulation in green plant lineage families 
The patterns of phytohormones distribution, their native function and possible origin of hormonal regulation across the green plant lineages (chlorophytes, charophytes, bryophytes and tracheophytes) are discussed. The five classical phytohormones — auxins, cytokinins, gibberellins (GA), abscisic acid (ABA) and ethylene occur ubiquitously in green plants. They are produced as secondary metabolites by microorganisms. Some of the bacterial species use phytohormones to interact with the plant as a part of their colonization strategy. Phytohormone biosynthetic pathways in plants seem to be of microbial origin and furthermore, the origin of high affinity perception mechanism could have preceded the recruitment of a metabolite as a hormone. The bryophytes represent the earliest land plants which respond to the phytohormones with the exception of gibberellins. The regulation by auxin and ABA may have evolved before the separation of green algal lineage. Auxin enhances rhizoid and caulonemal differentiation while cytokinins enhance shoot bud formation in mosses. Ethylene retards cell division but seems to promote cell elongation. The presence of responses specific to cytokinins and ethylene strongly suggest the origin of their regulation in bryophytes. The hormonal role of GAs could have evolved in some of the ferns where antheridiogens (compounds related to GAs) and GAs themselves regulate the formation of antheridia.
During migration of life forms to land, the tolerance to desiccation may have evolved and is now observed in some of the microorganisms, animals and plants. Besides plants, sequences coding for late embryogenesis abundant-like proteins occur in the genomes of other anhydrobiotic species of microorganisms and nematodes. ABA acts as a stress signal and increases rapidly upon desiccation or in response to some of the abiotic stresses in green plants. As the salt stress also increases ABA release in the culture medium of cyanobacterium Trichormus variabilis, the recruitment of ABA in the regulation of stress responses could have been derived from prokaryotes and present at the level of common ancestor of green plants. The overall hormonal action mechanisms in mosses are remarkably similar to that of the higher plants. As plants are thought to be monophyletic in origin, the existence of remarkably similar hormonal mechanisms in the mosses and higher plants, suggests that some of the basic elements of regulation cascade could have also evolved at the level of common ancestor of plants. The networking of various steps in a cascade or the crosstalk between different cascades is variable and reflects the dynamic interaction between a species and its specific environment.
PMCID: PMC3550668  PMID: 23572871
Green plant families; Bryophytes; Abscisic acid; Auxin; Cytokinin; Ethylene; Antheridiogens; miRNA; Origin of hormonal regulation
25.  Desiccation Tolerance in the Moss Polytrichum formosum: Physiological and Fine-structural Changes during Desiccation and Recovery 
Annals of Botany  2006;99(1):75-93.
Background and Aims
This study explores basic physiological features and time relations of recovery of photosynthetic activity and CO2 uptake following rehydration of a desiccation-tolerant moss in relation to the full temporal sequence of cytological changes associated with recovery to the normal hydrated state. It seeks reconciliation of the apparently conflicting published physiological and cytological evidence on recovery from desiccation in bryophytes.
Observations were made of water-stress responses and recovery using infrared gas analysis and modulated chlorophyll fluorescence, and of structural and ultrastructural changes by light and transmission electron microscopy.
Key Results
Net CO2 uptake fell to zero at approx. 40 % RWC, paralleling the fluorescence parameter ΦPSII at 200 µmol m–2 s–1 PPFD. On re-wetting the moss after 9–18 d desiccation, the initially negative net CO2 uptake became positive 10–30 min after re-wetting, restoring a net carbon balance after approx. 0·3–1 h. The parameter Fv/Fm reached approx. 80 % of its pre-desiccation value within approx. 10 min of re-wetting. In the presence of the protein-synthesis inhibitors chloramphenicol and cycloheximide, recovery of Fv/Fm (and CO2 exchange) proceeded normally in the dark, but declined rapidly in the light. Though initial recovery was rapid, both net CO2 uptake and Fv/Fm required approx. 24 h to recover completely to pre-desiccation values. The fixation protocols produced neither swelling of tissues nor plasmolysis. Thylakoids, grana and mitochondrial cristae remained intact throughout the drying–re-wetting cycle, but there were striking changes in the form of the organelles, especially the chloroplasts, which had prominent lobes and lamellar extensions in the normally hydrated state, but rounded off when desiccated, returning slowly to their normal state within approx. 24 h of re-wetting. Sub-cellular events during desiccation and re-wetting were generally similar to those seen in published data from the pteridophyte Selaginella lepidophylla.
Initial recovery of respiration and photosynthesis (as of protein synthesis) is very rapid, and independent of protein synthesis, suggesting physical reactivation of systems conserved intact through desiccation and rehydration, but full recovery takes approx. 24 h. This is consistent with the cytological evidence, which shows the thylakoids and cristae remaining intact through the whole course of dehydration and rehydration. Substantial and co-ordinated changes in other cell components, which must affect spatial relationships of organelles and metabolic systems, return to normal on a time span similar to full recovery of photosynthesis. Comparison of the present data with recently published results suggests a significant role for the cytoskeleton in desiccation responses.
PMCID: PMC2802982  PMID: 17158142
Bryophyta; chlorophyll fluorescence; chloroplasts; CO2 exchange desiccation tolerance; electron microscopy; metabolic inhibitors; mosses; Polytrichum formosum

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