Multiple-pathways light modulation in Pleurosigma strigosum bi-raphid diatom.

Ordered, quasi-ordered, and even disordered nanostructures can be identified as constituent components of several protists, plants and animals, making possible an efficient manipulation of light for intra- and inter- species communication, camouflage, or for the enhancement of primary production. Diatoms are ubiquitous unicellular microalgae inhabiting all the aquatic environments on Earth. They developed, through tens of millions of years of evolution, ultrastructured silica cell walls, the frustules, able to handle optical radiation through multiple diffractive, refractive, and wave-guiding processes, possibly at the basis of their high photosynthetic efficiency. In this study, we employed a range of imaging, spectroscopic and numerical techniques (including transmission imaging, digital holography, photoluminescence spectroscopy, and numerical simulations based on wide-angle beam propagation method) to identify and describe different mechanisms by which Pleurosigma strigosum frustules can modulate optical radiation of different spectral content. Finally, we correlated the optical response of the frustule to the interaction with light in living, individual cells within their aquatic environment following various irradiation treatments. The obtained results demonstrate the favorable transmission of photosynthetic active radiation inside the cell compared to potentially detrimental ultraviolet radiation.

Underwater Light Manipulation by the Benthic Diatom Ctenophora pulchella: From PAR Efficient Collection to UVR Screening.

Several species of diatoms, unicellular microalgae which constitute the main component of phytoplankton, are characterized by an impressive photosynthetic efficiency while presenting a noticeable tolerance versus exposure to detrimental UV radiation (UVR). In particular, the growth rate of the araphid diatom Ctenophora pulchella is not significantly affected by harsh treatments with UVR, even in absence of detectable, specific UV-absorbing pigments and even if it is not able to avoid high UV exposure by motility. In this work we applied a multi-disciplinary approach involving numerical computation, photonics, and biological parameters in order to investigate the possible role of the frustule, micro- and nano-patterned silica shell which encloses the cell, in the ability of C. pulchella to efficiently collect photosynthetic active radiation (PAR) and to simultaneously screen the protoplasm from UVR. The characterization of the photonic properties of the frustule has been accompanied by in vivo experiments conducted in water in order to investigate its function as optical coupler between light and plastids.

Effects of seawater scrubbing on a microplanktonic community during a summer-bloom in the Baltic Sea.

The International Maritime Organization (IMO) has gradually applied stricter regulations on the maximum sulphur content permitted in marine fuels and from January 1, 2020, the global fuel sulphur limit was reduced from 3.5% to 0.5%. An attractive option for shipowners is to install exhaust gas cleaning systems, also known as scrubbers, and continue to use high sulphur fuel oil. In the scrubber, the exhausts are led through a fine spray of water, in which sulphur oxides are easily dissolved. The process results in large volumes of acidic discharge water, but while regulations are focused on sulphur oxides removal and acidification, other pollutants e.g. polycyclic aromatic hydrocarbons, metals and nitrogen oxides can be transferred from the exhausts to the washwater and discharged to the marine environment. The aim of the current study was to investigate how different treatments of scrubber discharge water (1, 3 and 10%) affect a natural Baltic Sea summer microplanktonic community. To resolve potential contribution of acidification from the total effect of the scrubber discharge water, "pH controls" were included where the pH of natural sea water was reduced to match the scrubber treatments. Biological effects (e.g. microplankton species composition, biovolume and primary productivity) and chemical parameters (e.g. pH and alkalinity) were monitored and analysed during 14 days of exposure. Significant effects were observed in the 3% scrubber treatment, with more than 20% increase in total biovolume of microplankton compared to the control group, and an even greater effect in the 10% scrubber treatment. Group-specific impacts were recorded where diatoms, flagellates incertae sedis, chlorophytes and ciliates increased in biovolume with increasing concentrations of scrubber water while no effect was recorded for cyanobacteria. In contrast, these effects was not observed in the "pH controls", a suggestion that other parameters/stressors in the scrubber water were responsible for the observed effects.

A reflection on four impactful Ambio papers: The biotic perspective : This article belongs to Ambio's 50th Anniversary Collection. Theme: Climate change impacts.

Climate change represents one of the most pressing societal and scientific challenges of our time. While much of the current research on climate change focuses on future prediction, some of the strongest signals of warming can already be seen in Arctic and alpine areas, where temperatures are rising faster than the global average, and in the oceans, where the combination of rising temperatures and acidification due to increased CO2 concentrations has had catastrophic consequences for sensitive marine organisms inhabiting coral reefs. The scientific papers highlighted as part of this anniversary issue represent some of the most impactful advances in our understanding of the consequences of anthropogenic climate change. Here, we reflect on the legacy of these papers from the biotic perspective.

Cultivation and Photophysiological Characteristics of Desmids in Moderately Saline Aquaculture Wastewater.

Although desmids typically inhabit freshwater environments characterized by low amounts of nutrients and low salinity, several desmid species have been recorded in eutrophic waters, indicating their adaptation to elevated pollution and conductivity. This study aimed to determine whether desmids could be used for remediation of moderately saline aquaculture wastewater (AWW) from a fish farm situated in the southeast of Sweden. Fourteen desmid strains isolated from different climates (tropical to polar) and trophic conditions (oligotrophic to eutrophic) were cultivated in diluted AWW and we estimated their growth rates, biomass, nutrient removal efficiency, chlorophyll fluorescence parameters and cellular C, N and P quotas. Despite being grown at moderate salinity, unfavourable N:P ratio, and relatively low light/temperature regime the eutrophic strains, Cosmarium humile, Cosmarium laeve and a meso-oligotrophic species Cosmarium impressulum, completely absorbed nitrate and phosphate from AWW media after 7 d, indicating their potential for remediation of fish effluents in colder climates. These species, along with the typical eutrophic species, Cosmarium meneghinii and Staurastrum chaetoceras, had biomass in the range 0.45-1.19 g · L-1 while maximum growth rates ranged from 0.36 to 0.51 · d-1 , similar to published rates for several fast-growing green microalgae cultivated in various AWW types. Tropical desmids had distinctly high values of saturating irradiance (Ik  > 1,000 µmol photons · m-2  · s-1 ), and, along with eutrophic desmids, had high potential electron transport (rETRmax  > 155 rel. units). Hence, the desmids studied demonstrated inherent photophysiological responses corresponding to their climate and trophic origin under the suboptimal growth conditions.

Glacial melt disturbance shifts community metabolism of an Antarctic seafloor ecosystem from net autotrophy to heterotrophy.

Climate change-induced glacial melt affects benthic ecosystems along the West Antarctic Peninsula, but current understanding of the effects on benthic primary production and respiration is limited. Here we demonstrate with a series of in situ community metabolism measurements that climate-related glacial melt disturbance shifts benthic communities from net autotrophy to heterotrophy. With little glacial melt disturbance (during cold El Niño spring 2015), clear waters enabled high benthic microalgal production, resulting in net autotrophic benthic communities. In contrast, water column turbidity caused by increased glacial melt run-off (summer 2015 and warm La Niña spring 2016) limited benthic microalgal production and turned the benthic communities net heterotrophic. Ongoing accelerations in glacial melt and run-off may steer shallow Antarctic seafloor ecosystems towards net heterotrophy, altering the metabolic balance of benthic communities and potentially impacting the carbon balance and food webs at the Antarctic seafloor.

Future HAB science: Directions and challenges in a changing climate.

There is increasing concern that accelerating environmental change attributed to human-induced warming of the planet may substantially alter the patterns, distribution and intensity of Harmful Algal Blooms (HABs). Changes in temperature, ocean acidification, precipitation, nutrient stress or availability, and the physical structure of the water column all influence the productivity, composition, and global range of phytoplankton assemblages, but large uncertainty remains about how integration of these climate drivers might shape future HABs. Presented here are the collective deliberations from a symposium on HABs and climate change where the research challenges to understanding potential linkages between HABs and climate were considered, along with new research directions to better define these linkages. In addition to the likely effects of physical (temperature, salinity, stratification, light, changing storm intensity), chemical (nutrients, ocean acidification), and biological (grazer) drivers on microalgae (senso lato), symposium participants explored more broadly the subjects of cyanobacterial HABs, benthic HABs, HAB effects on fisheries, HAB modelling challenges, and the contributions that molecular approaches can bring to HAB studies. There was consensus that alongside traditional research, HAB scientists must set new courses of research and practices to deliver the conceptual and quantitative advances required to forecast future HAB trends. These different practices encompass laboratory and field studies, long-term observational programs, retrospectives, as well as the study of socioeconomic drivers and linkages with aquaculture and fisheries. In anticipation of growing HAB problems, research on potential mitigation strategies should be a priority. It is recommended that a substantial portion of HAB research among laboratories be directed collectively at a small sub-set of HAB species and questions in order to fast-track advances in our understanding. Climate-driven changes in coastal oceanographic and ecological systems are becoming substantial, in some cases exacerbated by localized human activities. That, combined with the slow pace of decreasing global carbon emissions, signals the urgency for HAB scientists to accelerate efforts across disciplines to provide society with the necessary insights regarding future HAB trends.

Fatty acids as chemotaxonomic and ecophysiological traits in green microalgae (desmids, Zygnematophyceae, Streptophyta): A discriminant analysis approach.

Desmids (Zygnematophyceae) are a group of poorly studied green microalgae. The aim of the present study was to identify fatty acids (FAs) that could be used as biomarkers in desmids in general, and to determine FAs as traits within different ecophysiological desmid groups. FA profiles of 29 desmid strains were determined and analysed with respect to their geographic origin, trophic preference and age of cultivation. It appeared that merely FAs present in relatively large proportions such as palmitic, linoleic, α-linolenic and hexadecatrienoic acids could be used as biomarkers for reliable categorization of this microalgal group. Linear discriminant analysis applied to three a priori defined groups of desmids, revealed clear strain-specific characteristics regarding FA distribution, influenced by climate and trophic conditions at the source sites as well as by the age of culture and growth phase. Accordingly, when considering FAs for the determination of lower taxonomic ranks we recommend using the term "trait" instead of "biomarker", as the latter designates unchangeable "fingerprint" of a specific taxon. Furthermore, despite that desmids were regarded as microalgae having stable genomes, long-term cultivation appeared to cause modifications in FA metabolic pathways, evident as a larger proportion of stearidonic acid in desmid strains cultivated over extensive time periods (>35 years).

Diatom frustules enhancing the efficiency of gel polymer electrolyte based dye-sensitized solar cells with multilayer photoelectrodes.

The incorporation of nanostructures that improve light scattering and dye adsorption has been suggested for dye-sensitized solar cells (DSSCs), but the manufacture of photonic and nanostructured materials with the desired properties is not an easy task. In nature, however, the process of light-harvesting for photosynthesis has, in some cases, evolved structures with remarkable wavelength-sensitive light-trapping properties. The present work is focused on enhancing the efficiency of quasi solid-state DSSCs by capitalizing on the light trapping properties of diatom frustules since they provide complex 3-dimensional structures for scattering and trapping light. This study reports a promising approach to prepare TiO2 nanocrystal (14 nm) based photo-electrodes by utilizing the waveguiding and photon localization effects of nanostructured diatom frustules for enhancing light harvesting without deteriorating the electron conduction. Single and double-layered photo-electrodes were prepared with different frustule/nanocrystal combinations and conformations on transparent conductive oxide substrates. This study clearly reports impressive efficiency and short circuit current density enhancements of about 35% and 39%, respectively, due to the incorporation of diatom frustules extracted from a ubiquitous species. The SEM images obtained in this work reveal that the produced thin films had a remarkable surface coverage of evenly distributed frustules within the TiO2 nanoparticle layer. To the best of our knowledge, this study reports the first quasi solid-state DSSC based on a photo-electrode with incorporated bio-formed nanostructures.

Effects of scrubber washwater discharge on microplankton in the Baltic Sea.

In 2020, the global cap of maximum allowable sulphur content in marine fuel will be reduced from the current 3.5% to 0.5%. Another way to reduce the sulphur emissions is to install a seawater scrubber that cleans exhausts but instead release acidic water containing nutrients and contaminants back to the marine environment. In the current study, scrubber washwater was tested on a Baltic Sea microplankton community. A significant increase in chlorophyll a, particulate organic phosphorus (POP), carbon (POC) and nitrogen (PON) were observed when the community was exposed to 10% scrubber washwater for 13 days as compared to the control. A laboratory experiment with the filamentous cyanobacteria Nodularia spumigena and the chain-forming diatom Melosira cf. arctica showed negative responses in photosynthetic activity (EC10 = 8.6% for N. spumigena) and increased primary productivity (EC10 = 5.5% for M. cf. arctica), implying species-specific responses to scrubber washwater discharge.

Microalgal photophysiology and macronutrient distribution in summer sea ice in the Amundsen and Ross Seas, Antarctica.

Our study addresses how environmental variables, such as macronutrients concentrations, snow cover, carbonate chemistry and salinity affect the photophysiology and biomass of Antarctic sea-ice algae. We have measured vertical profiles of inorganic macronutrients (phosphate, nitrite + nitrate and silicic acid) in summer sea ice and photophysiology of ice algal assemblages in the poorly studied Amundsen and Ross Seas sectors of the Southern Ocean. Brine-scaled bacterial abundance, chl a and macronutrient concentrations were often high in the ice and positively correlated with each other. Analysis of photosystem II rapid light curves showed that microalgal cells in samples with high phosphate and nitrite + nitrate concentrations had reduced maximum relative electron transport rate and photosynthetic efficiency. We also observed strong couplings of PSII parameters to snow depth, ice thickness and brine salinity, which highlights a wide range of photoacclimation in Antarctic pack-ice algae. It is likely that the pack ice was in a post-bloom situation during the late sea-ice season, with low photosynthetic efficiency and a high degree of nutrient accumulation occurring in the ice. In order to predict how key biogeochemical processes are affected by future changes in sea ice cover, such as in situ photosynthesis and nutrient cycling, we need to understand how physicochemical properties of sea ice affect the microbial community. Our results support existing hypothesis about sea-ice algal photophysiology, and provide additional observations on high nutrient concentrations in sea ice that could influence the planktonic communities as the ice is retreating.

Ocean acidification and desalination: climate-driven change in a Baltic Sea summer microplanktonic community.

Helcom scenario modelling suggests that the Baltic Sea, one of the largest brackish-water bodies in the world, could expect increased precipitation (decreased salinity) and increased concentration of atmospheric CO2 over the next 100 years. These changes are expected to affect the microplanktonic food web, and thereby nutrient and carbon cycling, in a complex and possibly synergistic manner. In the Baltic Proper, the extensive summer blooms dominated by the filamentous cyanobacteria Aphanizomenon sp., Dolichospermum spp. and the toxic Nodularia spumigena contribute up to 30% of the yearly new nitrogen and carbon exported to the sediment. In a 12 days outdoor microcosm experiment, we tested the combined effects of decreased salinity (from 6 to 3) and elevated CO2 concentrations (380 and 960 µatm) on a natural summer microplanktonic community, focusing on diazotrophic filamentous cyanobacteria. Elevated pCO2 had no significant effects on the natural microplanktonic community except for higher biovolume of Dolichospermum spp. and lower biomass of heterotrophic bacteria. At the end of the experimental period, heterotrophic bacterial abundance was correlated to the biovolume of N. spumigena. Lower salinity significantly affected cyanobacteria together with biovolumes of dinoflagellates, diatoms, ciliates and heterotrophic bacteria, with higher biovolume of Dolichospermum spp. and lower biovolume of N. spumigena, dinoflagellates, diatoms, ciliates and heterotrophic bacteria in reduced salinity. Although the salinity effects on diatoms were apparent, they could not clearly be separated from the influence of inorganic nutrients. We found a clear diurnal cycle in photosynthetic activity and pH, but without significant treatment effects. The same diurnal pattern was also observed in situ (pCO2, pH). Thus, considering the Baltic Proper, we do not expect any dramatic effects of increased pCO2 in combination with decreased salinity on the microplanktonic food web. However, long-term effects of the experimental treatments need to be further studied, and indirect effects of the lower salinity treatments could not be ruled out. Our study adds one piece to the complicated puzzle to reveal the combined effects of increased pCO2 and reduced salinity levels on the Baltic microplanktonic community.

Diatom frustules protect DNA from ultraviolet light.

The evolutionary causes for generation of nano and microstructured silica by photosynthetic algae are not yet deciphered. Diatoms are single photosynthetic algal cells populating the oceans and waters around the globe. They generate a considerable fraction (20-30%) of all oxygen from photosynthesis, and 45% of total primary production of organic material in the sea. There are more than 100,000 species of diatoms, classified by the shape of the glass cage in which they live, and which they build during algal growth. These glass structures have accumulated for the last 100 million of years, and left rich deposits of nano/microstructured silicon oxide in the form of diatomaceous earth around the globe. Here we show that reflection of ultraviolet light by nanostructured silica can protect the deoxyribonucleic acid (DNA) in the algal cells, and that this may be an evolutionary cause for the formation of glass cages.

Long-term acclimation to elevated pCO2 alters carbon metabolism and reduces growth in the Antarctic diatom Nitzschia lecointei.

Increasing atmospheric CO2 levels are driving changes in the seawater carbonate system, resulting in higher pCO2 and reduced pH (ocean acidification). Many studies on marine organisms have focused on short-term physiological responses to increased pCO2, and few on slow-growing polar organisms with a relative low adaptation potential. In order to recognize the consequences of climate change in biological systems, acclimation and adaptation to new environments are crucial to address. In this study, physiological responses to long-term acclimation (194 days, approx. 60 asexual generations) of three pCO2 levels (280, 390 and 960 µatm) were investigated in the psychrophilic sea ice diatom Nitzschia lecointei. After 147 days, a small reduction in growth was detected at 960 µatm pCO2. Previous short-term experiments have failed to detect altered growth in N. lecointei at high pCO2, which illustrates the importance of experimental duration in studies of climate change. In addition, carbon metabolism was significantly affected by the long-term treatments, resulting in higher cellular release of dissolved organic carbon (DOC). In turn, the release of labile organic carbon stimulated bacterial productivity in this system. We conclude that long-term acclimation to ocean acidification is important for N. lecointei and that carbon overconsumption and DOC exudation may increase in a high-CO2 world.

Physicochemical control of bacterial and protist community composition and diversity in Antarctic sea ice.

Due to climate change, sea ice experiences changes in terms of extent and physical properties. In order to understand how sea ice microbial communities are affected by changes in physicochemical properties of the ice, we used 454-sequencing of 16S and 18S rRNA genes to examine environmental control of microbial diversity and composition in Antarctic sea ice. We observed a high diversity and richness of bacteria, which were strongly negatively correlated with temperature and positively with brine salinity. We suggest that bacterial diversity in sea ice is mainly controlled by physicochemical properties of the ice, such as temperature and salinity, and that sea ice bacterial communities are sensitive to seasonal and environmental changes. For the first time in Antarctic interior sea ice, we observed a strong eukaryotic dominance of the dinoflagellate phylotype SL163A10, comprising 63% of the total sequences. This phylotype is known to be kleptoplastic and could be a significant primary producer in sea ice. We conclude that mixotrophic flagellates may play a greater role in the sea ice microbial ecosystem than previously believed, and not only during the polar night but also during summer when potential food sources are abundant.

Harmful algal blooms and climate change: Learning from the past and present to forecast the future.

Climate change pressures will influence marine planktonic systems globally, and it is conceivable that harmful algal blooms may increase in frequency and severity. These pressures will be manifest as alterations in temperature, stratification, light, ocean acidification, precipitation-induced nutrient inputs, and grazing, but absence of fundamental knowledge of the mechanisms driving harmful algal blooms frustrates most hope of forecasting their future prevalence. Summarized here is the consensus of a recent workshop held to address what currently is known and not known about the environmental conditions that favor initiation and maintenance of harmful algal blooms. There is expectation that harmful algal bloom (HAB) geographical domains should expand in some cases, as will seasonal windows of opportunity for harmful algal blooms at higher latitudes. Nonetheless there is only basic information to speculate upon which regions or habitats HAB species may be the most resilient or susceptible. Moreover, current research strategies are not well suited to inform these fundamental linkages. There is a critical absence of tenable hypotheses for how climate pressures mechanistically affect HAB species, and the lack of uniform experimental protocols limits the quantitative cross-investigation comparisons essential to advancement. A HAB "best practices" manual would help foster more uniform research strategies and protocols, and selection of a small target list of model HAB species or isolates for study would greatly promote the accumulation of knowledge. Despite the need to focus on keystone species, more studies need to address strain variability within species, their responses under multifactorial conditions, and the retrospective analyses of long-term plankton and cyst core data; research topics that are departures from the norm. Examples of some fundamental unknowns include how larger and more frequent extreme weather events may break down natural biogeographic barriers, how stratification may enhance or diminish HAB events, how trace nutrients (metals, vitamins) influence cell toxicity, and how grazing pressures may leverage, or mitigate HAB development. There is an absence of high quality time-series data in most regions currently experiencing HAB outbreaks, and little if any data from regions expected to develop HAB events in the future. A subset of observer sites is recommended to help develop stronger linkages among global, national, and regional climate change and HAB observation programs, providing fundamental datasets for investigating global changes in the prevalence of harmful algal blooms. Forecasting changes in HAB patterns over the next few decades will depend critically upon considering harmful algal blooms within the competitive context of plankton communities, and linking these insights to ecosystem, oceanographic and climate models. From a broader perspective, the nexus of HAB science and the social sciences of harmful algal blooms is inadequate and prevents quantitative assessment of impacts of future HAB changes on human well-being. These and other fundamental changes in HAB research will be necessary if HAB science is to obtain compelling evidence that climate change has caused alterations in HAB distributions, prevalence or character, and to develop the theoretical, experimental, and empirical evidence explaining the mechanisms underpinning these ecological shifts.

Inheritance of mitochondrial DNA in the Pennate diatom Haslea ostrearia (Naviculaceae) during auxosporulation suggests a uniparental transmission.

We present the first study examining mtDNA transmission in diatoms, using sexual progeny of the pennate species Haslea ostrearia (Naviculaceae). A fragment of the cytochrome oxidase subunit I gene (cox1) with 7 nucleic substitutions between parental clones was used as a parental tracer in 16 F1 clones obtained from two pairs of mating crosses. Each cross involved a parental clone isolated from France (Bay of Bourgneuf) and Sweden (Kattegat Bay). We determined that all progeny possessed only one cox1 parental haplotype. These results suggest that the mitochondrial DNA transmission in H. ostrearia is uniparental. Implications and new topics of investigation are discussed.

UVR defense mechanisms in eurytopic and invasive Gracilaria vermiculophylla (Gracilariales, Rhodophyta).

The invasive success of Gracilaria vermiculophylla has been attributed to its wide tolerance range to different abiotic factors, but its response to ultraviolet radiation (UVR) is yet to be investigated. In the laboratory, carpospores and vegetative thalli of an Atlantic population were exposed to different radiation treatments consisting of high PAR (photosynthetically active radiation) only (P), PAR+UV-A (PA) and PAR+UV-A+UV-B (PAB). Photosynthesis of carpospores was photoinhibited under different radiation treatments but photosystem II (PSII) function was restored after 12 h under dim white light. Growth of vegetative thalli was significantly higher under radiation supplemented with UVR. Decrease in chlorophyll a (Chl a) under daily continuous 16-h exposure to 300 µmol photons m(-2) s(-1) of PAR suggests preventive accumulation of excited chlorophyll molecules within the antennae to minimize the generation of dangerous reactive oxygen species. Moreover, an increase in total carotenoids and xanthophyll cycle pigments (i.e. violaxanthin, antheraxanthin and zeaxanthin) further suggests effective photoprotection under UVR. The presence of the ketocarotenoid ß-cryptoxanthin also indicates protection against UVR and oxidative stress. The initial concentration of total mycosporine-like amino acids (MAAs) in freshly-released spores increased approximately four times after 8-h laboratory radiation treatments. On the other hand, initial specific MAAs in vegetative thalli changed in composition after 7-day exposure to laboratory radiation conditions without affecting the total concentration. The above responses suggest that G. vermiculophylla have multiple UVR defense mechanisms to cope with the dynamic variation in light quantity and quality encountered in its habitat. Beside being eurytopic, the UVR photoprotective mechanisms likely contribute to the current invasive success of the species in shallow lagoons and estuaries exposed to high solar radiation.