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The Arctic archipelago of Svalbard is a hotspot of global warming and many fjords experience a continuous increase in seawater temperature and glacial melt while sea‐ice cover declines. In 1996/1998, 2012–2014, and 2021 macroalgal biomass and species diversity were quantified at the study site Hansneset, Kongsfjorden (W‐Spitsbergen) in order to identify potential changes over time. In 2021, we repeated the earlier studies by stratified random sampling (1 × 1 m2, n = 3) along a sublittoral depth transect (0, 2.5, 5, 10, and 15 m) and investigated the lower depth limits of dominant brown algae between 3 and 19 m. The maximum fresh weight (FW) of all seaweeds was 11.5 kg m−2 at 2.5 m and to 99.9% constituted of kelp. Although biomass distribution along the depth transect in 2021 was not significantly different compared to 2012/2013, the digitate kelp community (Laminaria digitata/Hedophyllum nigripes) had transformed into an Alaria esculenta‐dominated kelp forest. Consequently, a pronounced shift in kelp forest structure occurred over time as we demonstrate that biomass allocation to thallus parts is kelp species‐specific. Over the past decade, kelp demography changed and in 2021 a balanced age structure of kelps (juveniles plus many older kelp individuals) was only apparent at 2.5 m. In addition, the abundances and lower depth limits of all dominant brown algae declined noticeably over the last 25 years while the red algal flora abundance remained unchanged at depth. We propose that the major factor driving the observed changes in the macroalgal community are alterations in underwater light climate, as in situ data showed increasing turbidity and decreasing irradiance since 2012 and 2017, respectively. As a consequence, the interplay between kelp forest retreat to lower depth levels caused by coastal darkening and potential macroalgal biomass gain with increasing temperatures will possibly intensify in the future with unforeseen consequences for melting Arctic coasts and fjord ecosystem services. [ABSTRACT FROM AUTHOR]

Coastal kelp forests produce substantial marine carbon due to high annual net primary production (NPP) rates, but upscaling of NPP estimates over time and space remains difficult. We investigated the impact of variable underwater photosynthetically active radiation (PAR) and photosynthetic parameters on photosynthetic oxygen production of Laminaria hyperborea, the dominant NE‐Atlantic kelp species, throughout summer 2014. Collection depth of kelp had no effect on chlorophyll a content, pointing to a high photoacclimation potential of L. hyperborea towards incident light. However, chlorophyll a and photosynthesis versus irradiance parameters differed significantly along the blade gradient when normalized to fresh mass, potentially introducing large uncertainties in NPP upscaling to whole thalli. Therefore, we recommend a normalization to kelp tissue area, which is stable over the blade gradient. Continuous PAR measurements revealed a highly variable underwater light climate at our study site (Helgoland, North Sea) in summer 2014, reflected by PAR attenuation coefficients (Kd) between 0.28 and 0.87 m−1. Our data highlight the importance of continuous underwater light measurements or representative average values using a weighted Kd to account for large PAR variability in NPP calculations. Strong winds in August increased turbidity, resulting in a negative carbon balance at depths >3–4 m over several weeks, considerably impacting kelp productivity. Estimated daily summer NPP over all four depths was 1.48 ± 0.97 g C · m−2 seafloor · d−1 for the Helgolandic kelp forest, which is in the range of other kelp forests along European coastlines. [ABSTRACT FROM AUTHOR]

Kelp forests are important habitats in the strongly environmentally and seasonally variable Arctic. There is a critical lack of knowledge about how seasonal conditions and climate change scenarios influence survival and reproduction of kelp early life stages. To better understand the regulation of kelp life cycle processes in this harsh environment we focused on the physiological performance and reproductive success of early life stages in Alaria esculenta and Laminaria digitata from Kongsfjorden, Spitsbergen. Gametophyte growth and survival during Arctic winter and subsequent sporophyte recruitment under spring conditions were investigated. Winter conditions (2°C, complete darkness) halted gametophyte growth and prevented the onset of gametogenesis in both species. The gametophytes of L. digitata but not A. esculenta became fertile after returning to spring conditions, suggesting that sporogenesis, sexual reproduction and recruitment in A. esculenta must occur successively during summer/autumn while in L. digitata a new generation of sporophytes could develop from over-wintering gametophytes. The effects of simulated canopy shading (offering protection against extreme irradiance stress, particularly as sea ice retreats), present-day and projected Arctic summer seawater temperatures, and nutrient levels on gametophyte survival, fertility and sporophyte recruitment success were also investigated in both species. A. esculenta gametophytes had greater survival and reproductive success than L. digitata, except under very low light (simulating dense canopy). In contrast, shading was required for reproductive success in L. digitata gametophytes. Predicted summer temperatures of 9°C reduced sexual reproduction in both species. Interactions observed between these environmental drivers probably reflect species-specific seasonal patterns of survival and reproduction. These differences between kelp species in response to abiotic factors and light levels (simulated canopy shading) suggest that climate change could alter community structure in the Arctic through effects on sexual reproduction and sporophyte recruitment success. HIGHLIGHTS• Gametophytes were able to endure long periods of darkness.• Parental kelp canopy is key for gametophyte survival and recruitment.• Climate change may alter kelp recruitment patterns. [ABSTRACT FROM AUTHOR]

Two morphologically similar digitate kelp species, Laminaria digitata and Hedophyllum nigripes, co-occur along a shallow sublittoral depth gradient in the Arctic but, in contrast to L. digitata, very few ecophysiological data exist for H. nigripes. We investigated growth, survival, photosynthetic characteristics and carbon:nitrogen ratios of juvenile sporophytes, and recruitment and survival of gametophytes in genetically verified Arctic isolates of both species along temperature gradients (0–25 °C) over 14 days. Laminaria digitata gametophytes survived 23–24 °C, while sporophytes survived 21–22 °C. Hedophyllum nigripes had lower temperature affinities. Gametophytes survived 19–21 °C, while sporophytes survived 18 °C. Male gametophytes were more heat-tolerant than female gametophytes in both species. The pronounced cold adaption of H. nigripes compared to L. digitata also became apparent in different sporophyte growth optima (L. digitata: 15 °C; H. nigripes: 10 °C) and gametogenesis optima (L. digitata: 5–15 °C; H. nigripes: 0–10 °C). Higher carbon:nitrogen ratios in H. nigripes suggest an adaptation to nutrient poor Arctic conditions. The overall temperature performance of H. nigripes possibly restricts the species to Arctic–Sub-Arctic regions, while Arctic L. digitata behaved similarly to cold-temperate populations. Our data suggest that a future increase in seawater temperatures may hamper the success of H. nigripes and favour L. digitata in Arctic environments. [ABSTRACT FROM AUTHOR]

SUMMARY The sugar kelp Saccharina latissima experiences a wide range of environmental conditions along its geographical and vertical distribution range. Temperature and salinity are two critical drivers influencing growth, photosynthesis and biochemical composition. Moreover, interactive effects might modify the results described for single effects. In shallow water coastal systems, exposure to rising temperatures and low salinity are expected as consequence of global warming, increased precipitation and coastal run‐off. To understand the acclimation mechanisms of S. latissima to changes in temperature and salinity and their interactions, we performed a mechanistic laboratory experiment in which juvenile sporophytes from Brittany, France were exposed to a combination of three temperatures (0, 8 and 15°C) and two salinity levels (20 and 30 psu (practical salinity units)). After a temperature acclimation of 7 days, sporophytes were exposed to low salinity (20 psu) for a period of 11 days. Growth, and maximal quantum yield of photosystem II (Fv/Fm), pigments, mannitol content and C:N ratio were measured over time. We report for the first time in S. latissima a fivefold increase in the osmolyte mannitol in response to low temperature (0°C) compared to 8 and 15°C that may have ecological and economic implications. Low temperatures significantly affected all parameters, mostly in a negative way. Chlorophyll a, the accessory pigment pool, growth and Fv/Fm were significantly lower at 0°C, while the de‐epoxidation state of the xanthophyll cycle was increased at both 0 and 8°C compared to 15°C. Mannitol content and growth decreased with decreased salinity; in contrast, pigment content and Fv/Fm were to a large extent irresponsive to salinity. In comparison to S. latissima originating from an Arctic population, despite some reported differences, this study reveals a remarkably similar impact of temperature and salinity variation, reflecting the large degree of adaptability in this species. [ABSTRACT FROM AUTHOR]

To understand the thermal plasticity of a coastal foundation species across its latitudinal distribution, we assess physiological responses to high temperature stress in the kelp Laminaria digitata in combination with population genetic characteristics and relate heat resilience to genetic features and phylogeography. We hypothesize that populations from Arctic and cold‐temperate locations are less heat resilient than populations from warm distributional edges. Using meristems of natural L. digitata populations from six locations ranging between Kongsfjorden, Spitsbergen (79°N), and Quiberon, France (47°N), we performed a common‐garden heat stress experiment applying 15°C to 23°C over eight days. We assessed growth, photosynthetic quantum yield, carbon and nitrogen storage, and xanthophyll pigment contents as response traits. Population connectivity and genetic diversity were analyzed with microsatellite markers. Results from the heat stress experiment suggest that the upper temperature limit of L. digitata is nearly identical across its distribution range, but subtle differences in growth and stress responses were revealed for three populations from the species' ecological range margins. Two populations at the species' warm distribution limit showed higher temperature tolerance compared to other populations in growth at 19°C and recovery from 21°C (Quiberon, France), and photosynthetic quantum yield and xanthophyll pigment responses at 23°C (Helgoland, Germany). In L. digitata from the northernmost population (Spitsbergen, Norway), quantum yield indicated the highest heat sensitivity. Microsatellite genotyping revealed all sampled populations to be genetically distinct, with a strong hierarchical structure between southern and northern clades. Genetic diversity was lowest in the isolated population of the North Sea island of Helgoland and highest in Roscoff in the English Channel. All together, these results support the hypothesis of moderate local differentiation across L. digitata's European distribution, whereas effects are likely too weak to ameliorate the species' capacity to withstand ocean warming and marine heatwaves at the southern range edge. [ABSTRACT FROM AUTHOR]

The plasticity of different kelp populations to heat stress has seldom been investigated excluding environmental effects due to thermal histories, by raising a generation under common garden conditions. Comparisons of populations in the absence of environmental effects allow unbiased quantification of the meta-population adaptive potential and resolution of population-specific differentiation. Following this approach, we tested the hypothesis that genetically distinct arctic and temperate kelp exhibit different thermal phenotypes, by comparing the capacity of their microscopic life stages to recover from elevated temperatures. Gametophytes of Laminaria digitata (Arctic and North Sea) grown at 15°C for 3 years were subjected to common garden conditions with static or dynamic (i.e., gradual) thermal treatments ranging between 15 and 25°C and also to darkness. Gametophyte growth and survival during thermal stress conditions, and subsequent sporophyte recruitment at two recovery temperatures (5 and 15°C), were investigated. Population-specific responses were apparent; North Sea gametophytes exhibited higher growth rates and greater sporophyte recruitment than those from the Arctic when recovering from high temperatures, revealing differential thermal adaptation. All gametophytes performed poorly after recovery from a static 8-day exposure at 22.5°C compared to the response under a dynamic thermal treatment with a peak temperature of 25°C, demonstrating the importance of gradual warming and/or acclimation time in modifying thermal limits. Recovery temperature markedly affected the capacity of gametophytes to reproduce following high temperatures, regardless of the population. Recovery at 5°C resulted in higher sporophyte production following a 15°C and 20°C static exposure, whereas recovery at 15°C was better for gametophyte exposures to static 22.5°C or dynamic heat stress to 25°C. The subtle performance differences between populations originating from sites with contrasting local in situ temperatures support our hypothesis that their thermal plasticity has diverged over evolutionary time scales. [ABSTRACT FROM AUTHOR]

The Arctic region is currently facing substantial environmental changes due to global warming. Melting glaciers cause reduced salinity environments in coastal Arctic habitats, which may be stressful for kelp beds. To investigate the responses of the kelp Saccharina latissima to the warming Arctic, we studied the transcriptomic changes of S. latissima from Kongsfjorden (Svalbard, Norway) over a 24‐hour exposure to two salinities (Absolute Salinity [SA] 20 and 30) after a 7‐day pre‐acclimation at three temperatures (0, 8 and 15°C). In addition, corresponding physiological data were assessed during an 11‐days salinity/temperature experiment. Growth and maximal quantum yield for photosystem II fluorescence were positively affected by increased temperature during acclimation, whereas hyposalinity caused negative effects at the last day of treatment. In contrast, hyposalinity induced marked changes on the transcriptomic level. Compared to the control (8°C – SA 30), the 8°C – SA 20 exhibited the highest number of differentially expressed genes (DEGs), followed by the 0°C – SA 20. Comparisons indicate that S. latissima tends to convert its energy from primary metabolism (e.g. photosynthesis) to antioxidant activity under hyposaline stress. The increase in physiological performance at 15°C shows that S. latissima in the Arctic region can adjust and might even benefit from increased temperatures. However, in Arctic fjord environments its performance might become impaired by decreased salinity as a result of ice melting. [ABSTRACT FROM AUTHOR]

Ocean warming can mediate species interactions and provoke changes in community structure worldwide. Species interactions vary along environmental gradients and life-history stages and increasing temperatures may change competitive dominance between species. Kelps, being marine foundation species, have a complex heteromorphic life cycle, with the early developmental stages being a bottleneck for successful establishment of the adult population. Here, we investigated how temperature influences interactions in early life-history stages of two kelp species with different thermal affinities (Alaria esculenta and Laminaria digitata from Spitsbergen) by cultivating them in mono- and co-culture and different temperatures. Irrespectively of cultivation treatment, spore germination, gametogenesis, and sporophyte development of both species were mostly positively stimulated by a temperature increase from mean ambient summer temperatures (4–5 °C) to a global warming scenario for the Arctic future (9–10 °C) but not at 15 °C which is the southern temperature limit of A. esculenta. At 15 °C gametogenesis and sporophyte formation of A. esculenta were greatly inhibited in monoculture but not so in L. digitata. On the other hand at 5 °C and 10 °C, gametogenesis and sporophyte growth were generally faster in A. esculenta than in L. digitata, leading to a competitive advantage of A. esculenta over L. digitata in the co-cultivation treatments. The interactive effects of co-cultivation and temperature were evident, where development of A. esculenta was accelerated in the presence of L. digitata at 9 °C but not at 4 °C. Although the mechanisms triggering interspecific interactions were not determined in this study, future global warming was found to give competitive advantage of A. esculenta over L. digitata, which could affect community structure and dominance in coastal environments. [ABSTRACT FROM AUTHOR]

Kelps are globally important bioengineering species with high ecological and economic value, but are increasingly threatened by climate-driven geographic range shifts. The inheritance of economically important traits from parents to offspring is poorly understood in kelps but it is of utmost interest to seaweed farmers wishing to select strains with superior performance and resilience to environmental change. For two allopatric kelp species (N-Atlantic Laminaria digitata and S-Atlantic L. pallida), we compared the speed of gametogenesis and reproductive success in parental gametophytes, and produced intraspecific and reciprocal interspecific crosses of female × male gametophyte parents isolated from the two species. We then compared the upper thermal resilience of microscopic and macroscopic sibling sporophytes in an exposure experiment over two weeks. The upper thermal limit of the sporophytes resulting from intraspecific crosses of the two species deviated by 1°C. In contrast, sporophytes from both interspecific hybrid crosses had a 2–3°C higher upper thermal tolerance than single species sporophytes, indicating heterosis for thermal tolerance. This phenotypic response appears partially sex-dependent in our study, with female parents being more important in determining the thermal-response phenotype than male parents. The presence of male gametophytes generally enhanced female reproductive success. Both gametogenesis rate and reproductive success differed among the types of reciprocal crosses. Although the interspecific crosses were artificial in an ecological sense, they may provide a tool for understanding the molecular basis of heterosis and thermal tolerance in kelps (e.g. by investigating species-specific gene expression), or for aquaculture breeding programmes against a background of rapid environmental change. [ABSTRACT FROM AUTHOR]

In haplodiplontic lineages, sexual reproduction occurs in haploid parents without meiosis. Although widespread in multicellular lineages such as brown algae (Phaeophyceae), haplodiplontic gametogenesis has been little studied at the molecular level. We addressed this by generating an annotated reference transcriptome for the gametophytic phase of the sugar kelp, Saccharina latissima. Transcriptional profiles of microscopic male and female gametophytes were analysed at four time points during the transition from vegetative growth to gametogenesis. Gametogenic signals resulting from a switch in culture irradiance from red to white light activated a core set of genes in a sex-independent manner, involving rapid activation of ribosome biogenesis, transcription and translation related pathways, with several acting at the post-transcriptional or post-translational level. Additional genes regulating nutrient acquisition and key carbohydrate-energy pathways were also identified. Candidate sex-biased genes under gametogenic conditions had potentially key roles in controlling female- and male-specific gametogenesis. Among these were several sex-biased or -specific E3 ubiquitin-protein ligases that may have important regulatory roles. Females specifically expressed several genes that coordinate gene expression and/or protein degradation, and the synthesis of inositol-containing compounds. Other female-biased genes supported parallels with oogenesis in divergent multicellular lineages, in particular reactive oxygen signalling via an NADPH-oxidase. Males specifically expressed the hypothesised brown algal sex-determining factor. Male-biased expression mainly involved upregulation of genes that control mitotic cell proliferation and spermatogenesis in other systems, as well as multiple flagella-related genes. Our data and results enhance genome-level understanding of gametogenesis in this ecologically and economically important multicellular lineage. [ABSTRACT FROM AUTHOR]

Spores represent the most vulnerable life history stage of kelps. While UV-induced inhibition of spore germination has been readily documented, the impact of in situ underwater radiation below kelp canopies has been largely overlooked. We determined spectral composition and intensity of underwater radiation along a density gradient in an Alaria esculenta kelp forest at 3 m depth in Kongsfjorden, Svalbard. Accordingly, we set up a laboratory experiment simulating five different radiation conditions corresponding to irradiances under very dense to no canopy cover on a cloudless summer day. Spore responses (photosynthetic quantum yield, pigment and phlorotannin contents, swimming activity, and germination success) were determined after 4, 8, 16, and 24 h of exposure. In situ spectral radiation composition differed strongly from conditions applied in previous studies, which underestimated photosynthetically active radiation and overestimated UV-radiation effects. Furthermore, spore solutions differed significantly in quantum yield, pigment, and phlorotannin contents upon release. Nevertheless, spores reacted dynamically to different radiation conditions and exposure times. Highest radiation (PAR 61.8 W m−2, 1.9 W m−2 UVA, 0.01 W m−2 UVB) caused photodamage after exposure for ≥ 8 h, while intermediate radiation led to photoinhibition. Lowest radiation (PAR 0.23 W m−2, 0 W m−2 UVA, 0 W m−2 UVB) caused inconsistent reactions. There was a reduction of absolute pigment content in all treatments, but reduction rates of photosynthetic pigments were significantly different between radiation treatments. Soluble phlorotannin content decreased under all conditions but was not significantly affected by experimental conditions. High radiation reduced swimming activity of spores, but experimental conditions had almost no effect on germination success. Consequently, it seems unlikely that in situ radiation conditions negatively affect spores in present and future radiation scenarios. [ABSTRACT FROM AUTHOR]

Submerged marine forests of macroalgae known as kelp are one of the key structures for coastal ecosystems worldwide. These communities are responding to climate driven habitat changes and are therefore appropriate indicators of ecosystem status and health. Hyperspectral remote sensing provides a tool for a spatial kelp habitat mapping. The difficulty in optical kelp mapping is the retrieval of a significant kelp signal through the water column. Detecting submerged kelp habitats is challenging, in particular in turbid coastal waters. We developed a fully automated simple feature detection processor to detect the presence of kelp in submerged habitats. We compared the performance of this new approach to a common maximum likelihood classification using hyperspectral AisaEAGLE data from the subtidal zones of Helgoland, Germany. The classification results of 13 flight stripes were validated with transect diving mappings. The feature detection showed a higher accuracy till a depth of 6 m (overall accuracy = 80.18%) than the accuracy of a maximum likelihood classification (overall accuracy = 57.66%). The feature detection processor turned out as a time-effective approach to assess and monitor submerged kelp at the limit of water visibility depth. [ABSTRACT FROM AUTHOR]

Kelp-derived detritus is a potential food item at the base level of marine food webs. A good knowledge of the factors that influence stable isotope composition in kelps is essential for reliable food web models. The variation of stable isotope signatures was analysed in three species of kelp in Kongsfjorden, Svalbard. Between 15 and 2.5 m depth, towards higher photon fluence rate, Alaria esculenta was gradually enriched in 13C while 15N did not change. The carbon/nitrogen ratio decreased with depth. 2- and 3-year-old kelp individuals had significantly higher δ15N values in their blades than 5- and 6-year-old ones. C/N ratio in blades remained statistically the same over the range of 2–6 years old plants. A two-way ANOVA did not show any interactive effects between depth and age group. A. esculenta as well as Saccharina latissima and Laminaria digitata exhibited enrichment in heavy carbon isotopes in the blades compared to the stipes, while within each species, δ15N values remained the same between blades and stipes. A more detailed analysis of kelp blades showed that independent of location in the fjord, year or season, and species, young tissue was more enriched in 13C than meristem and that decaying apical tissue was depleted relative to young tissue. The degree of this 13C depletion varied with kelp species. Meristem tissue appeared most uniform in the blades compared to young and decaying tissues and its δ13C signature was characteristic for each species with no significant difference in δ15N values. Decaying tissue showed no difference in δ13C between species. Carbon and nitrogen contents along the blades were species-specific and varied with growing location and year or season. Some advice is given on the temporal, spatial, and tissue-related choice of samples for different research objectives. [ABSTRACT FROM AUTHOR]