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The carbonate chemistry of sea ice is known to play a role in global carbon cycles, but its importance is uncertain in part due to disparities in reported results. Variability in physical and biological drivers is usually invoked to explain differences between studies. In the Canadian Arctic Archipelago, “invisible polynyas” – areas of strong currents, thin ice, and potentially high biological productivity – are examples of extreme spatial variability. We used an invisible polynya as a natural laboratory to study the effects of inferred initial ice formation conditions, ice growth rate, and algal biomass on the distribution of carbonate species by collecting enough cores to perform a statistical comparison between sites located within, and just outside of, a polynya near Iqaluktuttiaq (Cambridge Bay, Nunavut, Canada). At both sites, the uppermost 10 cm ice horizon showed evidence of CO2 off-gassing, while carbonate distributions in the middle and bottommost 10 cm horizons largely followed the salinity distribution. In the polynya, the upper ice horizon had significantly higher bulk total inorganic carbon (TIC), total alkalinity (TA), and salinity potentially due to freeze-up conditions that favoured frazil ice production. The middle ice horizons were statistically indistinguishable between sites, suggesting that ice growth rate is not an important factor for the carbonate distribution under mid-winter conditions. The thicker (non-polynya) site experienced higher algal biomass, TIC, and TA in the bottom horizon. Carbonate chemistry in the bottom horizon could largely be explained by the salinity distribution, with the strong currents at the polynya site potentially playing a role in desalinization; biology appeared to exert only a minor control, with some evidence that the ice algae community was net heterotrophic. We did see evidence of calcium carbonate precipitation but with little impact on the TIC:TA ratio and little difference between sites. Because differences were constrained to relatively thin layers at the top and bottom, vertically averaged values of TIC, TA, and especially the TIC:TA ratio were not meaningfully different between sites. This provides some justification for using a single bulk value for each parameter when modelling sea ice effects on ocean chemistry at coarse resolution. Exactly what value to use (particularly for the TIC:TA ratio) likely varies by region but could potentially be approximated from knowledge of the source seawater and sea ice salinity. Further insights await a rigorous intercomparison of existing data.

International audience; One of the most striking ecological divides on Earth is between marine and nearby freshwater environments, as relatively few taxa can move between the two. Microbial eukaryotes contribute to biogeochemical and energy cycling in both fresh and marine waters, with little species overlap between the two ecosystems. Arctic and sub-Arctic marine systems are relatively fresh compared to tropical and temperate systems, but details of microbial eukaryote communities along river-to-sea transitions are poorly known. To bridge this knowledge gap, we investigated three river-to-sea transitions (Nelson, Churchill, and Great Whale Rivers) in sub-Arctic Hudson Bay through 18S rRNA amplicon sequencing to identify microbial eukaryotes along the salinity and biogeochemical gradients. Salinity acted as the principal dispersal barrier preventing freshwater microorganisms from colonizing marine coastal waters, with microbial eukaryote communities of the three rivers clustering together. Just offshore, communities clustered by coastal regions associated with nutrient concentrations. Analysis of indicator species revealed that communities in the nitrate-depleted coastal water off the Churchill and Great Whale Rivers were dominated by heterotrophic taxa and small photosynthetic protists. In contrast, the Nelson offshore community was characterized by a high proportion of the diatom Rhizosolenia. A distinct community of heterotrophic protists was identified in the three estuarine transition zones, suggesting specialized estuarine communities. Such specialization was most marked in the Nelson River system that was sampled more intensely and showed estuarine circulation.The autochthonous community was composed of the bacterial grazers Katablepharis, Mataza, and Cryothecomonas, as well as brackish species of the diatoms Skeletonema and Thalassiosira. These findings suggest that flow regulation on the Nelson River that modifies estuarine circulation would affect estuarine community composition and distribution in the transition zone.

The abundance of diatoms and heterotrophic bacteria in sea ice rapidly increases during the spring. However, the number and activity of these microorganisms vary with changing environmental conditions and potentially the taxonomic composition of the algal community during this time. In this study, we assessed the spring bottom-ice community composition in Dease Strait, Nunavut, and investigated potential controls of chlorophyll a (chl a), particulate organic carbon (POC), cell abundance, and production from early March until early June. We found that using flow cytometry to estimate photosynthetic nanoeukaryote (2–20 μm) abundance gave results very similar to light microscopy counts, except when pennate diatoms with lengths close to 20 μm, the maximum size detected by flow cytometry, were abundant. Using the average abundance of nanoeukaryotes from the two methods, we documented a change in the size of cells comprising the ice algal community over the spring, from largely pico- (

During the Norwegian young sea ICE expedition (N-ICE2015) from January to June 2015 the pack ice in the Arctic Ocean north of Svalbard was studied during four drifts between 83° and 80° N. This pack ice consisted of a mix of second-year, first-year and young ice. The physical properties and ice algal community composition was investigated in the three different ice types during the winter-spring-summer transition. Our results indicate that algae remaining in sea ice that survived the summer melt season are subsequently trapped in the upper layers of the ice column during winter and may function as an algal seed repository. Once the connectivity in the entire ice column is established, as a result of temperature-driven increase in ice porosity during spring, algae in the upper parts of the ice are able to migrate towards the bottom and initiate the ice-algal spring bloom. Furthermore, this algal repository might seed the bloom in younger ice formed in adjacent leads. This mechanism was studied in detail for the often dominating ice diatom Nitzschia frigida.The proposed seeding mechanism may be compromised due to the disappearance of older ice in the anticipated regime shift towards a seasonally ice-free Arctic Ocean.

In spring 2015, we observed an extensive phytoplankton bloom of Phaeocystis pouchetii, with chlorophyll a concentrations up to 7.5 mg m−3, under compact snow-covered Arctic sea ice at 80-81˚N during the Norwegian young sea ICE (N-ICE2015) expedition. We investigated the influence of the under-ice bloom on inherent optical properties (IOPs) of the upper ocean. Absorption and scattering in the upper 20 m of the water column at visible wavebands increased threefold and tenfold, respectively, relative to pre-bloom conditions. The scattering-to-absorption ratio during the Phaeocystis under-ice bloom was higher than in previous Arctic studies investigating diatom blooms. During the bloom, absorption by colored dissolved organic matter (at 375 nm), seemingly of autochthonous origin, doubled. Total absorption by particles (at 440 nm), dominated by phytoplankton (> 90%), increased tenfold. Measured absorption and scattering in the water were used as inputs for a 1D coupled atmosphere-ice-ocean radiative transfer model (AccuRT) to investigate effects of altered IOPs on the under-ice light field. Multiple scattering between sea ice and phytoplankton in the ocean led to an increase in scalar irradiance in the photosynthetically active radiation range (Eo(PAR)) at the ice-ocean interface by 6–7% compared to pre-bloom situation. This increase could have a positive feedback on ice-algal and under-ice phytoplankton productivity. The ratio between Eo(PAR) and downwelling planar irradiance (Ed(PAR)) below sea ice reached 1.85. Therefore, the use of Ed(PAR) might significantly underestimate the amount of PAR available for photosynthesis underneath sea ice. Our findings could help to improve light parameterizations in primary production models.

A number of common sterols were identified in sea ice diatoms from the Arctic and the Antarctic. The main sterols in floating sea ice algal aggregates collected from Resolute Passage (Canadian Arctic) and Nansen Basin (North Svalbard) in 2012 were 22E-dehydrocholesterol, cholesterol, epi-brassicasterol, 24-methylenecholesterol and 24-ethylcholesterol, although the distribution varied between the two locations, likely reflecting compositional differences in diatom taxa. The three major sterols in cells of Berkeleya adeliensis picked from a melted sea ice core collected from Ryder Bay in the Antarctic Peninsula in 2014, were 24-ethylcholesterol, cholesterol and 22E-dehydrocholesterol. We suggest that certain sea ice diatoms may thus contribute to the sedimentary budget of common sterols in seasonally sea ice-covered locations following ice melt.

Light is a major environmental cue affecting various physiological and metabolic processes in plants. Although plant photoreceptors are well characterized, the mechanisms by which light regulates downstream responses are less clear. In Arabidopsis thaliana, the accumulation of photoprotective anthocyanin pigments is light dependent, and the R2R3 MYB transcription factor MYB75/PAP1 regulates anthocyanin accumulation. Here, we report that MYB75 interacts with and is phosphorylated by MAP KINASE4 (MPK4). Their interaction is dependent on MPK4 kinase activity and is required for full function of MYB75. MPK4 can be activated in response to light and is involved in the light-induced accumulation of anthocyanins. We show that MPK4 phosphorylation of MYB75 increases its stability and is essential for light-induced anthocyanin accumulation. Our findings reveal an important role for a MAPK pathway in light signal transduction.

Genetics studies the structure/function of genes via the characterization of their mutant phenotypes. In plants, a readily scorable mutant phenotype comprises macroscopic lesions symptomatic of disease in the absence of pathogens. Such mutants therefore exhibit autoimmune phenotypes. Many of these mutants are considered to be associated with immunity and the corresponding genes have been described as 'negative regulators' of immunity and/or cell death. Pathogens deliver effectors into host cells to increase infectivity by modifying or removing host proteins. Plants detect effectors via nucleotide-binding, leucine-rich repeat (NLR) immune receptors, which monitor host effector targets. In response to effector-mediated target tampering, NLR proteins potentiate immunity. The guard hypothesis proposes that NLRs 'guard' host 'guardees' targeted by pathogen effectors. An obvious corollary to this guard model is that forms of plant autoimmunity are a result of inappropriate NLR protein activation. In this review, we discuss what is known about some of the 'negative regulators' of immunity, and propose simple strategies that may help to characterize autoimmune mutants.

Marine ecosystems at high latitudes are characterized by extreme seasonal changes in light conditions, as well as a limited period of high primary production during spring and early summer. As light returns at the end of winter to Arctic ice-covered seas, a first algal bloom takes place in the bottom layer of the sea ice. This bottom ice algae community develops through three distinct phases in the transition from winter to spring, starting with phase I, a predominantly net heterotroph community that has limited interaction with the pelagic or benthic realms. Phase II begins in the spring once light for photosynthesis becomes available at the ice bottom, although interaction with the water column and benthos remains limited. The transition to the final phase III is then mainly driven by a balance of atmospheric and oceanographic forcing that induce structural changes in the sea ice and ultimately the removal of algal biomass from the ice. Due to limited data availability an incomplete understanding exists of all the processes determining ice algal bloom phenology and the considerable geographic differences in sympagic algal standing stocks and primary production. We present here the first pan-Arctic compilation of available time-series data on vernal sea ice algal bloom development and identify the most important factors controlling its development and termination. Using data from the area surrounding Resolute Bay (Nunavut, Canada) as an example, we support previous investigations that snow cover on top of the ice influences sea ice algal phenology, with highest biomass development, but also earliest termination of blooms, under low snow cover. We also provide a pan-Arctic overview of sea ice algae standing stocks and primary production, and discuss the pertinent processes behind the geographic differences we observed. Finally, we assess potential future changes in vernal algal bloom phenology as a consequence of climate change, including their importance to different groups of grazers.

The Marginal Ice Zone (MIZ) of the Arctic Ocean is changing rapidly due to a warming Arctic climate with commensurate reductions in sea ice extent and thickness. This Pan-Arctic review summarizes the main changes in the Arctic ocean–sea ice–atmosphere (OSA) interface, with implications for primary- and secondary producers in the ice and the underlying water column. Changes in the Arctic MIZ were interpreted for the period 1979–2010, based on best-fit regressions for each month. Trends of increasingly open water were statistically significant for each month, with quadratic fit for August–November, illustrating particularly strong seasonal feedbacks in sea-ice formation and decay. Geographic interpretations of physical and biological changes were based on comparison of regions with significant changes in sea ice: (1) The Pacific Sector of the Arctic Ocean including the Canada Basin and the Beaufort, Chukchi and East Siberian seas; (2) The Canadian Arctic Archipelago; (3) Baffin Bay and Hudson Bay; and (4) the Barents and Kara seas. Changes in ice conditions in the Barents sea/Kara sea region appear to be primarily forced by ocean heat fluxes during winter, whereas changes in the other sectors appear to be more summer–autumn related and primarily atmospherically forced. Effects of seasonal and regional changes in OSA-system with regard to increased open water were summarized for photosynthetically available radiation, nutrient delivery to the euphotic zone, primary production of ice algae and phytoplankton, ice-associated fauna and zooplankton, and gas exchange of CO2. Changes in the physical factors varied amongst regions, and showed direct effects on organisms linked to sea ice. Zooplankton species appear to be more flexible and likely able to adapt to variability in the onset of primary production. The major changes identified for the ice-associated ecosystem are with regard to production timing and abundance or biomass of ice flora and fauna, which are related to regional changes in sea-ice conditions.

The importance of newly formed sea ice in spring is likely to increase with formation of leads in a more dynamic Arctic icescape. We followed the ice algal species succession in young ice (≤ 0.27 m) in spring at high temporal resolution (sampling every second day for 1 month in May–June 2015) in the Arctic Ocean north of Svalbard. We document the early development of the ice algal community based on species abundance and chemotaxonomic marker pigments, and relate the young-ice algal community to the communities in the under-ice water column and the surrounding older ice. The seeding source seemed to vary between algal groups. Dinoflagellates were concluded to originate from the water column and diatoms from the surrounding older ice, which emphasizes the importance of older ice as a seeding source over deep oceanic regions and in early spring when algal abundance in the water column is low. In total, 120 taxa (80 identified to species or genus level) were recorded in the young ice. The protist community developed over the study period from a ciliate, flagellate, and dinoflagellate dominated community to one dominated by pennate diatoms. Environmental variables such as light were not a strong driver for the community composition, based on statistical analysis and comparison to the surrounding thicker ice with low light transmission. The photoprotective carotenoids to Chl a ratio increased over time to levels found in other high-light habitats, which shows that the algae were able to acclimate to the light levels of the thin ice. The development into a pennate diatom-dominated community, similar to the older ice, suggests that successional patterns tend toward ice-associated algae fairly independent of environmental conditions like light availability, season or ice type, and that biological traits, including morphological and physiological specialization to the sea ice habitat, play an important role in colonization of the sea ice environment. However, recruitment of ice-associated algae could be negatively affected by the ongoing loss of older ice, which acts as a seeding repository. Copyright © 2018 Kauko, Olsen, Duarte, Peeken, Granskog, Johnsen, Fernández-Méndez, Pavlov, Mundy and Assmy. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

The presence of algal pigments in sea ice alters under-ice irradiance spectra, and the relationship between these variables can be used as a non-invasive means for estimating ice- associated algal biomass on ecologically relevant spatial and temporal scales. While the influence of snow cover and ice algal biomass on spectra transmitted through the snow-ice matrix has been examined for the Arctic, it has not been tested for Antarctic sea ice at regional scales. We used paired measurements of sea ice core chla concentrations and hyperspectral-transmitted under-ice irradiances from 59 sites sampled off East Antarctica and in the Weddell Sea to develop algorithms for estimating algal biomass in Antarctic pack ice. We compared 4 approaches that have been used in various bio-optical studies for marine systems: normalised difference indices, ratios of spectral irradiance, scaled band area and empirical orthogonal functions. The percentage of vari- ance explained by these models ranged from 38 to 79%, with the best-performing approach being normalised difference indices. Given the low concentrations of integrated chl a observed in our study compared with previous studies, our statistical models performed surprisingly well in explaining variability in these concentrations. Our findings provide a basis for future work to develop methods for non-invasive time series measurements and medium- to large-scale spatial mapping of Antarctic ice algal biomass using instrumented underwater vehicles.

The bottom ice algae chlorophyll a (chl a)–snow depth (HS) relationship was investigated for first-year sea ice in Allen Bay, Nunavut, from 27 April to 13 June 2011. A transmitted irradiance technique was used to estimate ice algae chl a throughout the period at time series locations covered and cleared of snow. Furthermore, chl a was estimated along transects perpendicular to dominant snowdrift orientation, and at short-term snow clear experimental sites. The association between chl a and most snow depths was characterized by four phases over the spring; light limitation (negative relationship), a transitional period (no relationship), chl a decline associated with higher transmitted irradiance (positive relationship), and a final phase of chl a decline independent from HS (no relationship). Algal chl a under areas cleared of snow was lower, reached zero chl a earlier and declined faster than snow-covered control sites. Results indicated that snow removal caused these chl a responses through photoinhibition, as well as ice melt later in the spring. Based on this research we propose that weather events that can rapidly melt the snowpack could significantly deplete bottom ice chl a and cause early termination of the bloom if they occur late in the spring.

We present the results of a 6 week time series of carbonate system and stable isotope measurements investigating the effects of sea ice on air-sea CO2 exchange during the early melt period in the Canadian Arctic Archipelago. Our observations revealed significant changes in sea ice and sackhole brine carbonate system parameters that were associated with increasing temperatures and the buildup of chlorophyll a in bottom ice. The warming sea-ice column could be separated into distinct geochemical zones where biotic and abiotic processes exerted different influences on inorganic carbon and pCO2 distributions. In the bottom ice, biological carbon uptake maintained undersaturated pCO2 conditions throughout the time series, while pCO2 was supersaturated in the upper ice. Low CO2 permeability of the sea ice matrix and snow cover effectively impeded CO2 efflux to the atmosphere, despite a strong pCO2 gradient. Throughout the middle of the ice column, brine pCO2 decreased significantly with time and was tightly controlled by solubility, as sea ice temperature and in situ melt dilution increased. Once the influence of melt dilution was accounted for, both CaCO3 dissolution and seawater mixing were found to contribute alkalinity and dissolved inorganic carbon to brines, with the CaCO3 contribution driving brine pCO2 to values lower than predicted from melt-water dilution alone. This field study reveals a dynamic carbon system within the rapidly warming sea ice, prior to snow melt. We suggest that the early spring period drives the ice column toward pCO2 undersaturation, contributing to a weak atmospheric CO2 sink as the melt period advances.

This study reports on the temporal variations in algal and bacterial metabolism of dissolved dimethylsulfoniopropionate (DMSPd) in Arctic ice-covered waters in response to the release of organic matter (OM) from the sea ice and the onset of under-ice phytoplankton growth. Sampling took place between 21 May and 21 June 2012 at a station located in Resolute Passage. A snow and ice melt event was accompanied by an important release of OM and total DMSP from the bottom ice to the water column. This input of OM coincided with increases in DMSPd and DMSPd loss rate constant at the ice-water interface and, 2 days later, with increases in DMSPd and bacter- ial dimethylsulfide (DMS) yields from DMSPd at 0.5 m under the ice. The different microbial re - sponses suggest that DMSPd-rich brines were released first, followed by the release of sympagic algae due to ice melt. In both cases, the changes in DMSPd metabolism resulted in an increase in gross DMS production from 0.15 to 1.9 nmol l �1 d �1 . The initiation of phytoplankton growth resulted in increases in bacterial abundance, DMSPd loss-rate constant and DMSP-sulfur assimilation. In contrast, DMS yield remained low during the onset of phytoplankton growth, indicating that bac- teria used DMSP as a carbon and sulfur source. These results show that ice DMSPd can be rapidly (

A recent upgrade to the optics configuration of a thermal source FTIR microscope equipped with a focal plane array detector has enabled rapid acquisition of high magnification spectrochemical images, in transmission, with an effective geometric pixel size of ∼1 × 1 μm(2) at the sample plane. Examples, including standard imaging targets for scale and accuracy, as well as biomedical tissues and microorganisms, have been imaged with the new system and contrasted with data acquired at normal magnification and with a high magnification multi-beam synchrotron instrument. With this optics upgrade, one can now conduct rapid biodiagnostic ex vivo tissue imaging in-house, with images collected over larger areas, in less time (minutes) and with comparable quality and resolution to the best synchrotron source FTIR imaging capabilities.

Plant mitogen-activated protein kinases (MPKs) transduce signals required for the induction of immunity triggered by host recognition of pathogen-associated molecular patterns. We isolated a full-length cDNA of a group B MPK (PgMPK4) from pearl millet. Autophosphorylation assay of recombinant PgMPK4 produced in Escherichia coli confirmed it as a kinase. Differential accumulation of PgMPK4 mRNA and kinase activity was observed between pearl millet cultivars 852B and IP18292 in response to inoculation with the downy mildew oomycete pathogen Sclerospora graminicola. This increased accumulation of PgMPK4 mRNA, kinase activity as well as nuclear-localization of PgMPK protein(s) was only detected in the S. graminicola resistant cultivar IP18292 with a ~tenfold peak at 9 h post inoculation. In the susceptible cultivar 852B, PgMPK4 mRNA and immuno-detectable nuclear PgMPK could be induced by application of the chemical elicitor β-amino butyric acid, the non-pathogenic bacteria Pseudomonas fluorescens, or by the phytohormones jasmonic acid (JA) or salicylic acid (SA). Furthermore, kinase inhibitor treatments indicated that PgMPK4 is involved in the JA- and SA-mediated expression of three defense genes, lipoxygenase, catalase 3 and polygalacturonase-inhibitor protein. These findings indicate that PgMPK/s contribute to pearl millet defense against the downy mildew pathogen by activating the expression of defense proteins.

We measured the concentrations of selected lipids and some of their degradation products in a time series of sea ice samples collected from Resolute Bay in the Canadian Arctic in 2012. The identification of specific tracers of photo- and autoxidation reactions provided evidence of both abiotic processes acting on organic matter, although none of the lipids appeared sensitive to significant biodegradation. Some differences in lipid reactivity were observed between the upper (3–10 cm) and lower (0–3 cm) sections of the cores, possibly as a result of increased exchange at the ice–water boundary for the latter. In terms of photodegradation, the phytyl side chain of chlorophyll and a tri-unsaturated highly branched isoprenoid (HBI) alkene were most affected, 24-methylenecholesterol (from diatoms) was more susceptible to photodegradation than brassicasterol (from diatoms and prymnesiophytes), while sitosterol and cholesterol, likely from a range of sources, were largely unaffected. With respect to autoxidation, the reactivity trend for the sterols was reversed, with significant autoxidation of sitosterol and cholesterol, but not of 24-methylenecholesterol or brassicasterol. The phytyl side chain of chlorophyll and a highly branched isoprenoid triene were also particularly susceptible to autoxidation. In contrast, the diatom fatty acid, C16:1, was not substantially altered by any degradation pathway. By measuring temporal changes in the proportions of specific tracers of each degradation process, we provide evidence that such abiotic reactions take place via sequential photooxidation and autoxidation, with homolytic cleavage of photochemically produced hydroperoxides leading to autoxidation of initial substrates and subsequent oxidation products. The observation of significant abiotic degradation of several lipids appears to take place despite the low temperature in sea ice (ca. 0 °C) and should be considered carefully alongside other removal processes involving organic matter in marine systems.

A MAP kinase pathway below a chitin receptor in the moss Physcomitrella patens induces immune responses including rapid growth inhibition, a novel fluorescence burst, and cell wall depositions. The molecular mechanisms producing these three responses are currently unknown but warrant further investigation in this simple model system. Here we describe qualitative, time-lapse, and quantitative assays to monitor and measure these responses.

The assays described here are pertinent to protein kinase studies in any plant. They include an immunoblot phosphorylation/activation assay and an in-gel activity assay for MAP kinases (MPKs) using the general protein kinase substrate myelin basic protein. They also include a novel in-gel peptide substrate assay for Snf1-related kinase family 2 members (SnRK2s). This kinase family-specific assay overcomes some limitations of in-gel assays and permits the identification of different types of kinase activities in total protein extracts.

We have used thermal source Fourier Transform Infrared (FTIR) microtomographic imaging to compare sea ice diatoms growing under different light conditions. A prototype tomography accessory was designed to have sufficient degrees of freedom to align any tilted cylindrical sample relative to the axis of rotation, minimizing the off-axis path traced during rotation. The lightweight device rests on the motorized stage to position the sample in the field-of-view and enable mosaic imaging. Reconstruction routines were tested with simulated and real phantoms, to assess limitations in the Radon back-projection method employed. The distribution and abundance of biochemicals is analysed for targets larger than a single FPA tile. Two and three dimensional (2D and 3D) FTIR spectrochemical images were obtained with a Focal Plane Array (FPA, nominal 1.1 μm pixel edges) for phantoms (polystyrene beads in polyvinyl alcohol matrix) and diatom cells harvested from land fast, first-year ice sites in Resolute Passage (74 43.628'N; 95 33.330'W) and Dease Strait (69° 1.11'N; 105° 21.29'W), Nunavut, Canada. The analysis of relative concentrations of organic matter within the encapsulating silica frustules of diatoms is important for a better understanding of both the physiological state and the individual cellular response to environmental pressures. Analysis of 3D FTIR images of Nitzschia frigida collected from beneath high (17-19 cm) and low (3-7 cm) snow depth revealed higher concentrations of lipids in diatoms collected under low snow cover, uniquely based on spectroscopically determined total 3D cell volume and biochemical content.

A MAP kinase pathway below a chitin receptor in the moss Physcomitrella patens induces immune responses including rapid growth inhibition, a novel fluorescence burst, and cell wall depositions. The molecular mechanisms producing these three responses are currently unknown but warrant further investigation in this simple model system. Here we describe qualitative, time-lapse, and quantitative assays to monitor and measure these responses.

The assays described here are pertinent to protein kinase studies in any plant. They include an immunoblot phosphorylation/activation assay and an in-gel activity assay for MAP kinases (MPKs) using the general protein kinase substrate myelin basic protein. They also include a novel in-gel peptide substrate assay for Snf1-related kinase family 2 members (SnRK2s). This kinase family-specific assay overcomes some limitations of in-gel assays and permits the identification of different types of kinase activities in total protein extracts.

Relatively few sea ice energy balance studies have successfully captured the transition season of warming, snowmelt, and melt pond formation. In this paper, we report a surface energy budget for landfast sea ice that captures this important period. The study was conducted in the Canadian Arctic Archipelago from 10 May to 20 June 2010. Over the first 20 days of the study, we found that short periods (1–3 days) of increased net radiation associated with low longwave loss provided most of the energy required to warm the snowpack from winter conditions. An extended period of low longwave loss (5 days) combined with the seasonal increase in incoming shortwave radiation then triggered snowmelt onset. Melt progressed with a rapid reduction in albedo and attendant increases in shortwave energy absorption, resulting in melt pond formation 8 days later. The key role of longwave radiation in initiating melt onset supports past findings, and confirms the importance of clouds and water vapor associated with synoptic weather systems. However, we also observed a period of strong turbulent energy exchange associated with the passage of a cyclone. The cyclone event occurred shortly after melt pond formation, but it delivered enough energy to significantly hasten melt onset had it occurred earlier in the season. Changes in the frequency, duration, and timing of synoptic-scale weather events that deliver clouds and/or strong turbulent heat fluxes may be important in explaining observed changes in sea ice melt onset timing.

This study presents temporal variations in concentrations of chlorophyll a (Chl a), particulate and dissolved dimethylsulfoniopropionate (DMSPp and DMSPd) in the sea ice and underlying water column in the Canadian Arctic Archipelago during the spring of 2010 and 2011. During both years, bottom ice Chl a, DMSPp and DMSPd concentrations were high (up to 1328 µg L−1, 15,082 nmol L−1, and 6110 nmol L−1, respectively) in May and decreased thereafter. The release of bottom ice algae and DMSPp in the water column was gradual in 2010 and rapid (8 days) in 2011. Bottom brine drainage during the presnowmelt period in 2010 and a rapid loss of the snow cover in 2011 coinciding with rain events explain most of the difference between the 2 years. During both years, less than 13% of the DMSPd lost from the ice was detected in the water column, suggesting a rapid microbial consumption. An under-ice diatom bloom developed in both years. In 2010, the bloom was dominated by centric diatoms while in 2011 pennates dominated, likely reflecting seeding by ice algae following the faster snowmelt progression induced by rainfall events in 2011. Both under-ice blooms were associated with high DMSPp concentrations (up to 185 nmol L−1), but pennate diatoms showed DMSPp/Chl a ratios twice higher than centrics. These results highlight the key role of snowmelt and precipitation on the temporal pattern of ice-DMSP release to the water column and on the timing, taxonomic composition, and DMSP content of phytoplankton under-ice blooms in the Arctic.

It has been common practice in scientific studies to assume negligible phytoplankton production when the ocean is ice-covered, due to the strong light attenuation properties of snow, sea ice, and ice algae. Recent observations of massive under-ice blooms in the Arctic challenge this concept and call for a re-evaluation of light conditions prevailing under ice during the melt period. Using hydrographic data collected under landfast ice cover in Resolute Passage, Nunavut, Canada between 9 May and 21 June 2010, we documented the exponential growth phase of a substantial under-ice phytoplankton bloom. Numerous factors appeared to influence bloom initi- ation: (1) transmitted light increased with the onset of snowmelt and termination of the ice algal bloom; (2) initial phytoplankton growth resulted in the accumulation of biomass below the devel- oping surface melt layer where nutrient concentrations were high and turbulent mixing was rela- tively low; and (3) melt pond formation rapidly increased light transmission, while spring-tidal energy helped form a surface mixed layer influenced by ice melt — both were believed to influ- ence the final rapid increase in phytoplankton growth. By the end of the study, nitrate+nitrite was depleted in the upper 10 m of the water column and the under-ice bloom had accumulated 508 mg chl a m �2 with a new production estimate of 17.5 g C m �2 over the upper 50 m of the water column. The timing of bloom initiation with melt onset suggests a strong link to climate change where sea ice is both thinning and melting earlier, the implication being an earlier and more ubiquitous phytoplankton bloom in Arctic ice-covered regions.

Summary The accelerated cell death 11 ( acd11 ) mutant of Arabidopsis provides a genetic model for studying immune response activation and localized cellular suicide that halt pathogen spread during infection in plants. Here, we elucidate ACD11 structure and function and show that acd11 disruption dramatically alters the in vivo balance of sphingolipid mediators that regulate eukaryotic-programmed cell death. In acd11 mutants, normally low ceramide-1-phosphate (C1P) levels become elevated, but the relatively abundant cell death inducer phytoceramide rises acutely. ACD11 exhibits selective intermembrane transfer of C1P and phyto-C1P. Crystal structures establish C1P binding via a surface-localized, phosphate headgroup recognition center connected to an interior hydrophobic pocket that adaptively ensheaths lipid chains via a cleft-like gating mechanism. Point mutation mapping confirms functional involvement of binding site residues. A π helix (π bulge) near the lipid binding cleft distinguishes apo-ACD11 from other GLTP folds. The global two-layer, α-helically dominated, "sandwich" topology displaying C1P-selective binding identifies ACD11 as the plant prototype of a GLTP fold subfamily.

The lipid content of sea ice samples collected in 2011 and 2012 from Resolute Passage in the Canadian Arctic Archipelago was measured and compared with related samples obtained from the Amundsen Gulf in 2008. The highly branched isoprenoid (HBI) sea ice biomarker, IP 25 , was found in sea ice samples from each study, consistent with its formation by diatoms during the spring bloom. Our analysis also revealed the occurrence of a number of sterols in Arctic sea ice and these were rigorously identified and quantified for the first time. Concentrations of IP 25 and sterols exhibited some variability between sampling studies, with somewhat higher values in samples from Resolute in 2012 than for the other two datasets, consistent with a general increase in biomass; however, major differences in biomarker concentration between sampling studies were not observed. An estimate of the proportion of Arctic sea ice diatoms that produce IP 25 (ca. 1–5%) was obtained by comparison of the concentration of IP 25 in the samples with those in laboratory cultures of known HBI-producing diatoms and cell enumeration in selected sea ice samples. The estimate is similar to the proportion of Haslea spp. in the same samples, providing further support to the suggestion that at least some species of the Haslea genus may be responsible for the biosynthesis of IP 25 and related HBI diatom lipids in Arctic sea ice and that IP 25 is made by a relatively small proportion of sea ice diatoms. In contrast, median sterol/IP 25 values were all substantially higher than those in cultures of HBI-producing diatoms, suggesting that sterols are made by the majority of sea ice diatoms. The sterol/IP 25 ratio was quite variable between locations and samples, likely as a result of differences in diatom assemblage; however, a comparison of individual and median sterol/IP 25 values in sea ice with those from surface sediments from different Arctic regions demonstrated that sterols from sea ice diatoms may, in some cases, have a significant impact on the sedimentary budget. This should be considered carefully for quantitative estimates of palaeo sea ice reconstruction using methods such as the PIP 25 index, which are based on the relative concentrations of IP 25 and sterols in Arctic marine sediment cores.

During summer 2008, as part of the Circumpolar Flaw Lead system study, we measured phytoplankton photosynthetic parameters to understand regional patterns in primary productivity, including the degree and timescale of photoacclimation and how variability in environmental conditions influences this response. Photosynthesis–irradiance measurements were taken at 15 sites primarily from the depth of the subsurface chlorophyll a (Chl a) maximum (SCM) within the Beaufort Sea flaw lead polynya. The physiological response of phytoplankton to a range of light levels was used to assess maximum rates of carbon (C) fixation (P m * ), photosynthetic efficiency (α *), photoacclimation (E k), and photoinhibition (β *). SCM samples taken along a transect from under ice into open water exhibited a >3-fold increase in α * and P m * , showing these parameters can vary substantially over relatively small spatial scales, primarily in response to changes in the ambient light field. Algae were able to maintain relatively high rates of C fixation despite low light at the SCM, particularly in the large (>5 μm) size fraction at open water sites. This may substantially impact biogenic C drawdown if species composition shifts in response to future climate change. Our results suggest that phytoplankton in this region are well acclimated to existing environmental conditions, including sea ice cover, low light, and nutrient pulses. Furthermore, this photoacclimatory response can be rapid and keep pace with a developing SCM, as phytoplankton maintain photosynthetic rates and efficiencies in a narrow “shade-acclimated” range.

Biological characteristics of ice-associated algal communities were studied in Darnley Bay (western Canadian Arctic) during a 2-week period in July 2008 when the landfast ice cover had reached an advanced stage of melt. We found two distinct and separate algal communities: (1) an interior ice community confined to brine channel networks beneath white ice covers; and (2) an ice melt water community in the brackish waters of both surface melt ponds and the layer immediately below the ice cover. Both communities reached maximum chlorophyll a concentrations of about 2.5 mg m−3, but with diatoms dominating the interior ice while flagellates dominated the melt water community. The microflora of each community was diverse, containing both unique and shared algal species, the latter suggesting an initial seeding of the ice melt water by the bottom ice community. Absorption characteristics of the algae indicated the presence of mycosporine-like amino acids (MAAs) and carotenoid pigments as a photoprotective strategy against being confined to high-light near-surface layers. Although likely not contributing substantially to total annual primary production, these ice-associated communities may play an important ecological role in the Arctic marine ecosystem, supplying an accessible and stable food source to higher trophic levels during the period of ice melt.

Programmed cell death (PCD) is an integral part of plant development and of responses to abiotic stress or pathogens. Although the morphology of plant PCD is, in some cases, well characterised and molecular mechanisms controlling plant PCD are beginning to emerge, there is still confusion about the classification of PCD in plants. Here we suggest a classification based on morphological criteria. According to this classification, the use of the term 'apoptosis' is not justified in plants, but at least two classes of PCD can be distinguished: vacuolar cell death and necrosis. During vacuolar cell death, the cell contents are removed by a combination of autophagy-like process and release of hydrolases from collapsed lytic vacuoles. Necrosis is characterised by early rupture of the plasma membrane, shrinkage of the protoplast and absence of vacuolar cell death features. Vacuolar cell death is common during tissue and organ formation and elimination, whereas necrosis is typically found under abiotic stress. Some examples of plant PCD cannot be ascribed to either major class and are therefore classified as separate modalities. These are PCD associated with the hypersensitive response to biotrophic pathogens, which can express features of both necrosis and vacuolar cell death, PCD in starchy cereal endosperm and during self-incompatibility. The present classification is not static, but will be subject to further revision, especially when specific biochemical pathways are better defined.

Early summer in the Arctic with extensive ice melt and break-up represents a dramatic change for sympagic–pelagic fauna below seasonal sea ice. As part of the International Polar Year-Circumpolar Flaw Lead system study (IPY-CFL), this investigation quantified zooplankton in the meltwater layer below landfast ice and remaining ice fauna below melting ice during June (2008) in Franklin Bay and Darnley Bay, Amundsen Gulf, Canada. The ice was in a state of advanced melt, with fully developed melt ponds. Intense melting resulted in a 0.3- to 0.5-m-thick meltwater layer below the ice, with a strong halocline to the Arctic water below. Zooplankton under the ice, in and below the meltwater layer, was sampled by SCUBA divers. Dense concentrations (max. 1,400 ind. m−3) of Calanus glacialis were associated with the meltwater layer, with dominant copepodid stages CIV and CV and high abundance of nauplii. Less abundant species included Pseudocalanus spp., Oithona similis and C. hyperboreus. The copepods were likely feeding on phytoplankton (0.5–2.3 mg Chl-a m−3) in the meltwater layer. Ice amphipods were present at low abundance (

Variations in the concentrations of the sea ice diatom biomarker, IP25 (Ice Proxy with 25 carbon atoms), were measured in the bottom 10 cm of sea ice collected from the eastern Beaufort Sea and Amundsen Gulf from January to June 2008, as part of the International Polar Year–Circumpolar Flaw Lead system study. Temporal and vertical changes in IP25 concentrations were compared against other biomarkers and indicators of ice algal production. IP25 was not detected in sea ice samples collected from mid-winter to early spring, likely as a result of light-limiting conditions for algal growth and accumulation. From early March to mid-June, IP25 concentrations correlated well with those of fatty acids (r = 0.79; P 5% which is consistent with the hypothesis that brine channel connectivity limits the internal colonisation of sea ice by diatoms. Maximum IP25 concentrations occurred at 1–3 cm from the ice–water interface providing further evidence for a selective sea ice diatom origin for this biomarker. In contrast, vertical concentration profiles for fatty acids and sterols indicated mixed sources for these biomarkers.

SummaryAutophagy has been implicated as a prosurvival mechanism to restrict programmed cell death (PCD) associated with the pathogen-triggered hypersensitive response (HR) during plant innate immunity. This model is based on the observation that HR lesions spread in plants with reduced autophagy gene expression. Here, we examined receptor-mediated HR PCD responses in autophagy-deficient Arabidopsis knockout mutants (atg), and show that infection-induced lesions are contained in atg mutants. We also provide evidence that HR cell death initiated via Toll/Interleukin-1 (TIR)-type immune receptors through the defense regulator EDS1 is suppressed in atg mutants. Furthermore, we demonstrate that PCD triggered by coiled-coil (CC)-type immune receptors via NDR1 is either autophagy-independent or engages autophagic components with cathepsins and other unidentified cell death mediators. Thus, autophagic cell death contributes to HR PCD and can function in parallel with other prodeath pathways.

The Arabidopsis acd11 mutant exhibits runaway, programmed cell death due to the loss of a putative sphingosine transfer protein (ACD11) with homology to mammalian GLTP. We demonstrate that transgenic expression in Arabidopsis thaliana of human GLTP partially suppressed the phenotype of the acd11 null mutant, resulting in delayed programmed cell death development and plant survival. Surprisingly, a GLTP mutant form impaired in glycolipid transfer activity also complemented the acd11 mutants. To understand the relationship between functional complementarity and transfer activity, we generated site-specific mutants in ACD11 based on homologous GLTP residues required for glycolipid transfer. We show that these ACD11 mutant forms are impaired in their in vitro transfer activity of sphingolipids. However, transgenic expression of these mutant forms fully complemented acd11 mutant cell death, and transgenic plants showed normal induction of hypersensitive cell death upon infection with avirulent strains of Pseudomonas syringae. The significance of these findings with respect to the function(s) of ACD11 in sphingolipid transport and cell death regulation is discussed.

Plant and animal perception of microbes through pathogen surveillance proteins leads to MAP kinase signalling and the expression of defence genes. However, little is known about how plant MAP kinases regulate specific gene expression. We report that, in the absence of pathogens, Arabidopsis MAP kinase 4 (MPK4) exists in nuclear complexes with the WRKY33 transcription factor. This complex depends on the MPK4 substrate MKS1. Challenge with Pseudomonas syringae or flagellin leads to the activation of MPK4 and phosphorylation of MKS1. Subsequently, complexes with MKS1 and WRKY33 are released from MPK4, and WRKY33 targets the promoter of PHYTOALEXIN DEFICIENT3 (PAD3) encoding an enzyme required for the synthesis of antimicrobial camalexin. Hence, wrky33 mutants are impaired in the accumulation of PAD3 mRNA and camalexin production upon infection. That WRKY33 is an effector of MPK4 is further supported by the suppression of PAD3 expression in mpk4-wrky33 double mutant backgrounds. Our data establish direct links between MPK4 and innate immunity and provide an example of how a plant MAP kinase can regulate gene expression by releasing transcription factors in the nucleus upon activation.

The Arabidopsis (Arabidopsis thaliana) MKK1 and MKK2 mitogen-activated protein kinase kinases have been implicated in biotic and abiotic stress responses as part of a signaling cascade including MEKK1 and MPK4. Here, the double loss-of-function mutant (mkk1/2) of MKK1 and MKK2 is shown to have marked phenotypes in development and disease resistance similar to those of the single mekk1 and mpk4 mutants. Because mkk1 or mkk2 single mutants appear wild type, basal levels of MPK4 activity are not impaired in them, and MKK1 and MKK2 are in part functionally redundant in unchallenged plants. These findings are confirmed and extended by biochemical and molecular analyses implicating the kinases in jasmonate- and salicylate-dependent defense responses, mediated in part via the MPK4 substrate MKS1. In addition, transcriptome analyses delineate overlapping and specific effects of the kinases on global gene expression patterns demonstrating both redundant and unique functions for MKK1 and MKK2.

The monopoly enjoyed by the BBC Television Service in the UK prior to the introduction of ITV in 1955 led to accusations that the BBC was unprepared for competition and was reluctant to provide popular programming of the kind transmitted by its commercial rival. Using an archival case study of one popular entertainer, the comedian Norman Wisdom, this paper argues that sections of the BBC Television Service, notably Variety and Light Entertainment, worked hard from 1947 onwards to provide programming that relied on notions of ‘celebrity’ and ‘stardom’ in ways which anticipated the later success of ITV. Though hampered by financial constraints, entrenched institutional values and technical limitations, as well as competition from live variety, the film industry and radio, key personnel within the BBC Television Service did much to establish television as a popular medium even before the introduction of commercial broadcasting in 1955.

The characteristics of chromophoric dissolved organic matter (CDOM) were studied in Hudson Bay and Hudson Strait in the Canadian Arctic. Hudson Bay receives a disproportionately large influx of river runoff. With high dissolved organic matter (DOM) concentrations in Arctic rivers the influence of CDOM on coastal and ocean systems can be significant, yet the distribution, characteristics and potential consequences of CDOM in these waters remain unknown. We collected 470 discrete water samples in offshore, coastal, estuarine and river waters in the region during September and October 2005. Mixing of CDOM appeared conservative with salinity, although regional differences exist due to variable DOM composition in the rivers discharging to the Bay and the presence of sea-ice melt, which has low CDOM concentrations and low salinity. There were higher concentrations of CDOM in Hudson Bay, especially in coastal waters with salinities 28 , due to river runoff. Using CDOM composition of water masses as a tracer for the freshwater components revealed that river runoff is largely constrained to nearshore waters in Hudson Bay, while sea-ice melt is distributed more evenly in the Bay. Strong inshore–offshore gradients in the bio-optical properties of the surface waters in the Hudson Bay cause large variation in penetration of ultraviolet radiation and the photic depth within the bay, potentially controlling the vertical distribution of biomass and occurrence of deep chlorophyll maxima which are prevalent only in the more transparent offshore waters of the bay. The CDOM distribution and associated photoprocesses may influence the thermodynamics and stratification of the coastal waters, through trapping of radiant heating within the top few meters of the water column. Photoproduction of biologically labile substrates from CDOM could potentially stimulate the growth of biomass in Hudson Bay coastal waters. Further studies are needed to investigate the importance of terrestrial DOM in the Hudson Bay region, and the impact of hydroelectric development and climate change on these processes.

Programmed cell death (PCD) occurs during vegetative and reproductive plant growth, as typified by autumnal leaf senescence and the terminal differentiation of the endosperm of cereals which provide our major source of food. PCD also occurs in response to environmental stress and pathogen attack, and these inducible PCD forms are intensively studied due their experimental tractability. In general, evidence exists for plant cell death pathways which have similarities to the apoptotic, autophagic and necrotic forms described in yeast and metazoans. Recent research aiming to understand these pathways and their molecular components in plants are reviewed here.

Microscale photographs were taken of the ice bottom to examine linkages of algal chlorophyll a (chl a) biomass distribution with bottom ice features in thick Arctic first-year sea ice during a spring field program which took place from May 5 to 21, 2003. The photographic technique developed in this paper has resulted in the first in situ observations of microscale variability in bottom ice algae distribution in Arctic first-year sea ice in relation to ice morphology. Observations of brine channel diameter (1.65–2.68 mm) and number density (5.33–10.35 per 100 cm2) showed that the number of these channels at the bottom of thick first-year sea ice may be greater than previously measured on extracted ice samples. A variogram analysis showed that over areas of low chl a biomass (≤20.7 mg chl a m−2), patchiness in bottom ice chl a biomass was at the scale of brine layer spacing and small brine channels (∼1–3 mm). Over areas of high chl a biomass (≥34.6 mg chl a m−2), patchiness in biomass was related to the spacing of larger brine channels on the ice bottom (∼10–26 mm). Brine layers and channels are thought to provide microscale maxima of light, nutrient replenishment and space availability which would explain the small scale patchiness over areas of low algal biomass. However, ice melt and erosion near brine channels may play a more important role in areas with high algal biomass and low snow cover.

The geophysical, thermodynamic and dielectric properties of snow are important state variables that are known to be sensitive to Arctic climate variability and change. Given recent observations of changes in the Arctic physical system (Arctic Climate Impact Assessment, 2004), it is important to focus on the processes that give rise to variability in the horizontal, vertical and temporal dimensions of the life-history of snow on sea ice. The objectives in this study are to present these ‘state’ variables and to investigate the processes that govern variability in the vertical, horizontal and temporal dimension by using a case study over land-fast first-year sea ice for the period December 2003 to June 2004. Results from two sampling areas (thin and thick snowpacks) show that differences in snowpack thickness can substantially change the vertical and temporal evolution of snow properties. During the late fall and early winter (cooling period) we measured no significant changes in the physical properties, except for thin snow-cover salinity, which decreased throughout the period. Fall-snow desalination was only observed under thin snowpacks with a rate of −0·12 ppt day−1. Significant changes occurred in the late winter and early spring (warming period), especially for snow grain size. Snow grain kinetic growth of 0·25–0·48 mm·day−1 was measured coincidently with increasing salinity and wetness for both thin and thick snowpacks. Copyright © 2006 John Wiley & Sons, Ltd.

MAP kinase (MPK) cascades in Arabidopsis thaliana and other vascular plants are activated by developmental cues, abiotic stress, and pathogen infection. Much less is known of MPK functions in nonvascular land plants such as the moss Physcomitrella patens. Here, we provide evidence for a signaling pathway in P. patens required for immunity triggered by pathogen associated molecular patterns (PAMPs). This pathway induces rapid growth inhibition, a novel fluorescence burst, cell wall depositions, and accumulation of defense-related transcripts. Two P. patens MPKs (MPK4a and MPK4b) are phosphorylated and activated in response to PAMPs. This activation in response to the fungal PAMP chitin requires a chitin receptor and one or more MAP kinase kinase kinases and MAP kinase kinases. Knockout lines of MPK4a appear wild type but have increased susceptibility to the pathogenic fungi Botrytis cinerea and Alternaria brassisicola. Both PAMPs and osmotic stress activate some of the same MPKs in Arabidopsis. In contrast, abscisic acid treatment or osmotic stress of P. patens does not activate MPK4a or any other MPK, but activates at least one SnRK2 kinase. Signaling via MPK4a may therefore be specific to immunity, and the moss relies on other pathways to respond to osmotic stress.