The Arctic picoeukaryote Micromonas pusilla benefits synergistically from warming and ocean acidification

In the Arctic Ocean, climate change effects such as warming and ocean acidification (OA) are manifesting faster than in other regions. Yet, we are lacking a mechanistic understanding of the interactive effects of these drivers on Arctic primary producers. In the current study, one of the most abundant species of the Arctic Ocean, the prasinophyte Micromonas pusilla, was exposed to a range of different pCO2 levels at two temperatures representing realistic current and future scenarios for nutrient-replete conditions. We observed that warming and OA synergistically increased growth rates at intermediate to high pCO2 levels. Furthermore, elevated temperatures shifted the pCO2 optimum of biomass production to higher levels. Based on changes in cellular composition and photophysiology, we hypothesise that the observed synergies can be explained by beneficial effects of warming on carbon fixation in combination with facilitated carbon acquisition under OA. Our findings help to understand the higher abundances of picoeukaryotes such as M. pusilla under OA, as has been observed in many mesocosm studies

Resilience by diversity: Large intraspecific variation in climate change responses of an Arctic diatom

Primary productivity in the Arctic Ocean is vastly driven by single-celled phytoplankton. Within a comparably short growing season, they provide most of the carbon and energy for higher trophic levels and simultaneously influence biogeochemical cycles and the global climate. With the Arctic Ocean being one of the few regions that are expected to increase their productivity with future global change, it is especially important to understand how phytoplankton there is going to adapt. The potential for this adaptation to future climate change is often extrapolated from studies using single strains of a representative species that were cultured in laboratories for years. Since several decades, however, it is known that phytoplankton species and even local populations can exhibit large intraspecific diversity. During a field campaign on Svalbard, we isolated different strains of the Arctic diatom Thalassiosira hyalina from community incubations, which resembled present and future climate conditions. We then exposed these freshly isolated monocultures again to a matrix of CO and temperature. The results revealed that even within a single species, response patterns can differ greatly, comparable to variations seen between diatom species. Moreover, while only minor reactions of the communities were observed, the strain responses corresponded strongly with the previous selection environment. A strain isolated from future-like treatments, for instance, had its growth optimum at a higher temperature and pCO than another strain isolated from more present-like conditions. This suggests that intraspecific variability and the selection between coexisting ecotypes may be an underestimated source of species’ plasticity under changing environmental conditions and could ‘buffer’ functional species shifts. Adaptation of phytoplankton assemblages may therefore occur also by selection within rather than only between species, and species-wide inferences from single strain experiments should be handled with great care

The Arctic picoeukaryote Micromonas pusilla benefits from ocean acidification under constant and dynamic light

Compared to the rest of the globe, the Arctic Ocean is affected disproportionately by climate change. Despite these fast environmental changes, we currently know little about the effects of ocean acidification (OA) on marine key species in this area. Moreover, the existing studies typically test the effects of OA under constant, hence artificial, light fields. In this study, the abundant Arctic picoeukaryote Micromonas pusilla was acclimated to current (400 µatm) and future (1000 µatm) pCO2 levels under a constant as well as a dynamic light, simulating more realistic light fields as experienced in the upper mixed layer. To describe and understand the responses to these drivers, growth, particulate organic carbon (POC) production, elemental composition, photophysiology and reactive oxygen species (ROS) production were analysed. M. pusilla was able to benefit from OA on various scales, ranging from an increase in growth rates to enhanced photosynthetic capacity, irrespective of the light regime. These beneficial effects were, however, not reflected in the POC production rates, which can be explained by energy partitioning towards cell division rather than biomass build-up. In the dynamic light regime, M. pusilla was able to optimize its photophysiology for effective light usage during both low- and high-light periods. This photoacclimative response, which was achieved by modifications to photosys-tem II (PSII), imposed high metabolic costs leading to a reduction in growth and POC production rates when compared to constant light. There were no significant interactions observed between dynamic light and OA, indicating that M. pusilla is able to maintain effective photoacclimation without increased photoinactivation under high pCO2. Based on these findings, M. pusilla is likely to cope well with future conditions in the Arctic Ocean

Pelagic and ice-associated microalgae under elevated light and pCO2: Contrasting physiological strategies in two Arctic diatoms

Sea ice retreat, changing stratification and ocean acidification are fundamentally changing the light availability and physico-chemical conditions for primary producers in the Arctic ocean. However, detailed studies on ecophysiological strategies and performance of key species in the pelagic and ice-associated habitat remain scarce. We therefore investigated the acclimated responses of the diatoms Thalassiosira hyalina and Melosira arctica towards elevated irradiance and CO2 partial pressures. Next to growth, elemental composition and biomass production, we assessed detailed photophysiological responses through fluorometry and gas-flux measurements, including respiration and carbon acquisition. In the pelagic T. hyalina, growth rates remained high in all treatments and biomass production increased strongly with light. Even under low irradiances cells maintained a high-light acclimated state, allowing them to opportunistically utilize high irradiances by means of a highly plastic photosynthetic machinery and carbon uptake. The ice-associated M. arctica proved to be less plastic and more specialized on low-light. Its acclimation to high irradiances was characterized by minimizing photon harvest and photosynthetic efficiency, which led to lowered growth. Comparably low growth rates and strong silification advocate a strategy of persistence rather than of fast proliferation, which is also in line with the observed formation of resting stages under low-light conditions. In both species, responses to elevated pCO2 were comparably minor. Although both diatom species persisted under the applied conditions, their competitive abilities and strategies differ strongly. With the anticipated extension of Arctic pelagic habitats, flexible high-light specialists like T. hyalina seem to face a brighter future

Southern Ocean phytoplankton under multiple stressors: The modulation of Ocean Acidification effects by iron and light

The Southern Ocean (SO) contributes significantly to the sequestration of anthropogenic CO2 and is furthermore especially prone to Ocean Acidification (OA). The aim of this thesis was to investigate how key environmental factors influence the manifestation of OA effects on phytoplankton physiology and ecology. Publication I describes systematically occurring inconsistencies between pCO2 values calculated from different pairs of input parameters and their implications for the OA research field. Publication II presents the results of an experiment, in which interactive effects of OA and iron availability on SO primary production and species composition were observed. In publication III, dynamic light was found to strongly alter the effects of OA on the diatom Chaetoceros debilis. The aim of publication IV was to understand how iron, light and other factors control natural SO phytoplankton blooms. In conclusion, there is no universal phytoplankton response to OA. The effects of OA will always be modulated by the respective set of environmental conditions prevailing in the ecosystem of interest, which themselves may be subject to change

Resilience and adaptive mechanisms of Arctic phytoplankton under heatwaves: Acclimation, microevolution and community resilience

Trait adjustments of phytoplankton communities to changing environmental conditions can take place through responses on several fundamental ecological levels. These include physiological acclimation of single genotypes, evolution through sorting among genotypes of the same species, and selection within the entire multi-species community. Which of these different levels responds to environmental change can have large ecological and biogeochemical implications, but especially in protists, these levels are extremely difficult to disentangle. Arctic phytoplankton at base of the foodweb in one of the most rapidly warming regions on the planet, are faced with especially large changes, but often show high resilience. Among these changes are more frequent and intense heatwaves, which expose organisms to vast temperature fluctuations. In dedicated experimental setups of different ecological complexity, we investigated how phytoplankton responds and adjusts to heatwaves, and on which of the mentioned levels shifts can be observed. We resolved not only physiological features and productivity, but also composition on the species as well as the intraspecific level, using a novel molecular approach to efficiently examine the composition of protist populations in diverse contexts. This setup provides a comprehensive approach to investigate how phytoplankton communities respond to stable and fluctuating temperature scenarios, physiologically and ecologically

The hidden flows within species: Phytoplankton population dynamics in Arctic assemblages

Progressing climate change and concurrent alterations of environmental conditions pose challenges of adaptation on organisms and ecosystems, especially in rapidly changing places like the Arctic. While more diverse systems are usually considered to be more resilient, biodiversity does not only describe the number of species, but can also consist of diverse individuals within a species. Especially in protists, with large census sizes and fast proliferation, intraspecific lineage sorting can be an important mechanism of plasticity and trait adjustment. For phytoplankton communities at the base of the foodweb, physiological acclimation and species shifts are frequently described, but intraspecific composition and diversity are methodologically still difficult to resolve, especially in diverse natural contexts and at temporal resolution. Therefore, our knowledge on the functioning and importance of intraspecific selection dynamics in phytoplankton is still limited. In recent years, we have developed and applied a new, high throughput methodology for phytoplankton population composition, which can make temporal and spatial population dynamics visible that were before extremely difficult to resolve. Next to experiments with natural phytoplankton communities and artificial populations under controlled settings, a time-series of Arctic spring blooms has been investigated towards the year-to year composition of species but also of intraspecific populations of a dominant diatom. Datasets emerging now thanks to such novel technologies can offer new, more comprehensive perspectives on our understanding of the mechanisms and results of microevolution and local adaptation, and can reveal formerly hidden patterns of species’ strategies of persistence and development

The response of the red mangrove rhizophora mucronata lam, to changes in salinity, inundation and light : predictions for future climate change

Mangrove forests are subjected to many environmental factors which influence species distribution, zonation patterns as well as succession. Important driving factors in these forests are salinity, water level fluctuations and available light. This study investigated the response of red mangrove (Rhizophora mucronata Lam.) seedlings to these factors in controlled laboratory experiments. Increase in salinity and prolonged inundation within estuaries are predicted impacts resulting from sea level rise due to climate change. The study investigated the effect of five salinity treatments (0, 8, 18, 35 and 45 ppt) with a semi-diurnal tidal cycle on seedling growth. In a separate experiment the effect of different inundation treatments: no inundation, 3, 6, 9 hour tidal cycles and continuous inundation (24 h) were investigated. Both morphological and physiological responses of R. mucronata seedlings were measured. There was a decrease in growth (plant height, biomass and leaf production) with increasing salinity. Seedlings in the seawater, hypersaline and no inundation treatments showed symptoms of stress, having increased leaf necrosis ("burn marks"). The highest growth occurred in the low salinity (8 ppt) treatment, but the highest photosynthetic performance and stomatal conductance occurred in the freshwater treatment (0 ppt). The typical response of stem elongation with increasing inundation was observed in the 24 hr inundation treatment. In the light and salinity combination study there were ten different treatments of five different light treatments (unshaded, 20 percent, 50 percent, 80 percent and 90 percent shade) combined with two salinity concentrations (18 and 35 ppt). In this study the seedling growth: plant height, biomass, leaf surface area and leaf production were higher in the moderate salinity (18 ppt) treatments compared to the seawater (35 ppt) treatments. Biomass in the 35 ppt experiment decreased with increasing shade as well as in the unshaded treatments. Photosynthetic performance and stomatal conductance were lower for the unshaded treatment in both 18 and 35 ppt salinity compared to all other treatments with the same salinity. This suggests that R. mucronata more shade than sun tolerant, but overall it can be concluded that the species has a broad tolerance range. The results may be relevant in mangrove rehabilitation and predicting responses to climate change. This is important as mangrove ecosystems may adapt to changing sea levels and in order to restore areas it will be necessary to choose the mangrove species which will grow best. The results may also help to increase the protection of existing mangrove habitats

Response of mangroves in South Africa to anthropogenic and natural impacts

The total mangrove area cover in South Africa is 1631.7 ha, with the largest area cover in a few estuaries in the KwaZulu-Natal Province (1391.1 ha) and the remainder recorded in the Eastern Cape Province with 240.6 ha. This represents 0.05 percent of Africa‟s mangrove area cover and although small adds irreplaceable value to the biodiversity of South Africa. Mangroves are threatened by over-utilization through harvesting for firewood and building materials as well as excessive browsing and trampling by livestock. The objective of this study was to investigate the response of mangroves to different stressors from natural change as well as anthropogenic pressures. This was done by identifying pressures, measuring area cover, population structure and environmental parameters such as sediment characteristics. Mangroves in 17 estuaries along the east coast were investigated. Population structure and the area covered by mangroves in 2011/2012 were compared with data from the same area for 1999. Detailed studies were conducted in St. Lucia Estuary to investigate the response of mangroves to reduced tidal flooding; mangrove expansion at a latitudinal limit in a protected area at Nahoon Estuary was studied and the effect of cattle browsing on mangroves was measured at Nxaxo Estuary. The St. Lucia Estuary (28°S; 32°E) represented a unique study site as the mouth has been closed to the sea since 2002 and the mangrove habitats have been non-tidal. St. Lucia Estuary is both a Ramsar and World Heritage site and therefore understanding the response of mangroves to changes in the environment is important. In 2010 sediment characteristics and mangrove population structure were measured at four sites which were chosen to represent different salinity and water level conditions. The site fringing the main channel had the highest density of mangrove seedlings and saplings. The dry site had a lower density of mangroves with mostly only tall adult trees and few saplings. Mangrove tree height and density increased at sites with high sediment moisture and low surface sediment salinity. Few seedlings and saplings were found at sites with dry surface sediment and high salinity. Long term data are needed to assess the influence of mouth closure on recruitment and survival of the mangrove forest at St. Lucia Estuary; however this study has shown that sediment characteristics are unfavourable for mangrove growth at sites now characterized by a lack of tidal flooding. It is not known when exactly the mangroves were planted in Nahoon Estuary (32°S; 27° E), East London, but it is suspected that this was in the early 1970s. Avicennia marina (Forrsk.)Vierh. was planted first, followed a few years later by the planting of Bruguiera gymnorrhiza (L.) Lam. and Rhizophora mucronata (L.) among the larger A. marina trees. Surprisingly the mangrove population appears to be thriving and this study tested the hypothesis that mangroves have expanded and replaced salt marsh over a 33 year period. This study provides important information on mangroves growing at higher latitudes, where they were thought to not occur naturally due to lower annual average temperatures. It further provides insights on future scenarios of possible shifts in vegetation types due to climate change at one of the most southerly distribution sites worldwide. The expansion of mangroves was measured over a 33 year period (1978 - 2011) using past aerial photographs and Esri ArcGIS Desktop 10 software. In addition, field surveys were completed in 2011 to determine the population structure of the present mangrove forest and relate this to environmental conditions. The study showed that mangrove area cover increased linearly at a rate of 0.06 ha-1 expanding over a bare mudflat area, while the salt marsh area cover also increased (0.09 ha-1) but was found to be variable over time. The mangrove area is still small ( 70 percent). It was observed that browsing on trees resulted in a clear browse-line and browsing on propagules mainly by goats resulted in reduced seedling establishment in most of the estuaries except those in protected areas. Mangroves had re-established in estuaries where they had been previously lost but mouth closure due to drought and sea storms resulted in the mass die back of mangroves in the Kobonqaba Estuary. There was a total loss of 31.5 ha in mangrove area cover in the last 30 years and this was a total reduction of 10.5 ha (11 percent) for every decade. This is high considering that the present total mangrove area cover is only 240.6 ha for all the Transkei estuaries. In this study it was concluded that the anthropogenic impacts such as livestock browsing and trampling as well as harvesting in these estuaries contributed most to the mangrove degradation as these are continuous pressures occurring over long periods and are expected to increase in future with increasing human population. Natural changes such as sea storms occur less frequently but could result in large scale destruction over shorter periods. Examples of these are mouth closure that result in mangrove mass mortality as well as strong floods which destroy forest by scouring of the banks

Response of mangroves in South Africa to anthropogenic and natural impacts

The total mangrove area cover in South Africa is 1631.7 ha, with the largest area cover in a few estuaries in the KwaZulu-Natal Province (1391.1 ha) and the remainder recorded in the Eastern Cape Province with 240.6 ha. This represents 0.05 percent of Africa‟s mangrove area cover and although small adds irreplaceable value to the biodiversity of South Africa. Mangroves are threatened by over-utilization through harvesting for firewood and building materials as well as excessive browsing and trampling by livestock. The objective of this study was to investigate the response of mangroves to different stressors from natural change as well as anthropogenic pressures. This was done by identifying pressures, measuring area cover, population structure and environmental parameters such as sediment characteristics. Mangroves in 17 estuaries along the east coast were investigated. Population structure and the area covered by mangroves in 2011/2012 were compared with data from the same area for 1999. Detailed studies were conducted in St. Lucia Estuary to investigate the response of mangroves to reduced tidal flooding; mangrove expansion at a latitudinal limit in a protected area at Nahoon Estuary was studied and the effect of cattle browsing on mangroves was measured at Nxaxo Estuary. The St. Lucia Estuary (28°S; 32°E) represented a unique study site as the mouth has been closed to the sea since 2002 and the mangrove habitats have been non-tidal. St. Lucia Estuary is both a Ramsar and World Heritage site and therefore understanding the response of mangroves to changes in the environment is important. In 2010 sediment characteristics and mangrove population structure were measured at four sites which were chosen to represent different salinity and water level conditions. The site fringing the main channel had the highest density of mangrove seedlings and saplings. The dry site had a lower density of mangroves with mostly only tall adult trees and few saplings. Mangrove tree height and density increased at sites with high sediment moisture and low surface sediment salinity. Few seedlings and saplings were found at sites with dry surface sediment and high salinity. Long term data are needed to assess the influence of mouth closure on recruitment and survival of the mangrove forest at St. Lucia Estuary; however this study has shown that sediment characteristics are unfavourable for mangrove growth at sites now characterized by a lack of tidal flooding. It is not known when exactly the mangroves were planted in Nahoon Estuary (32°S; 27° E), East London, but it is suspected that this was in the early 1970s. Avicennia marina (Forrsk.)Vierh. was planted first, followed a few years later by the planting of Bruguiera gymnorrhiza (L.) Lam. and Rhizophora mucronata (L.) among the larger A. marina trees. Surprisingly the mangrove population appears to be thriving and this study tested the hypothesis that mangroves have expanded and replaced salt marsh over a 33 year period. This study provides important information on mangroves growing at higher latitudes, where they were thought to not occur naturally due to lower annual average temperatures. It further provides insights on future scenarios of possible shifts in vegetation types due to climate change at one of the most southerly distribution sites worldwide. The expansion of mangroves was measured over a 33 year period (1978 - 2011) using past aerial photographs and Esri ArcGIS Desktop 10 software. In addition, field surveys were completed in 2011 to determine the population structure of the present mangrove forest and relate this to environmental conditions. The study showed that mangrove area cover increased linearly at a rate of 0.06 ha-1 expanding over a bare mudflat area, while the salt marsh area cover also increased (0.09 ha-1) but was found to be variable over time. The mangrove area is still small ( 70 percent). It was observed that browsing on trees resulted in a clear browse-line and browsing on propagules mainly by goats resulted in reduced seedling establishment in most of the estuaries except those in protected areas. Mangroves had re-established in estuaries where they had been previously lost but mouth closure due to drought and sea storms resulted in the mass die back of mangroves in the Kobonqaba Estuary. There was a total loss of 31.5 ha in mangrove area cover in the last 30 years and this was a total reduction of 10.5 ha (11 percent) for every decade. This is high considering that the present total mangrove area cover is only 240.6 ha for all the Transkei estuaries. In this study it was concluded that the anthropogenic impacts such as livestock browsing and trampling as well as harvesting in these estuaries contributed most to the mangrove degradation as these are continuous pressures occurring over long periods and are expected to increase in future with increasing human population. Natural changes such as sea storms occur less frequently but could result in large scale destruction over shorter periods. Examples of these are mouth closure that result in mangrove mass mortality as well as strong floods which destroy forest by scouring of the banks