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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 (K d ) 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 K d 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., (© 2023 The Authors. Journal of Phycology published by Wiley Periodicals LLC on behalf of Phycological Society of America.)

The Arctic Ocean is a unique ecosystem hosting a biodiversity that has not yet been elucidated in full detail. There is increasing evidence that there are more kelp species constricted to Arctic/sub-Arctic habitats hitherto not well investigated, such as Hedophyllum nigripes, which is morphologically very similar to cold-temperate Laminaria digitata. Hedophyllum nigripes was originally described as L. nigripes by Agardh from Spitsbergen but has often been misidentified as L. digitata in the European Arctic. We initiated a systematic algal survey along a depth gradient (0-7.5 m) in Kongsfjorden (Spitsbergen) in June and July 2015 and thereby confirmed a predominant presence of H. nigripes (73%). Hedophyllum nigripes is occurring between 0 and 7.5 m while L. digitata was most abundant at 2.5 m depth. Hedophyllum nigripes individuals were generally younger (2.3 vs. 3.6 years) and stipe and blade length shorter (31 vs. 54 cm and 76 vs. 96 cm, respectively) compared to L. digitata. A combination of molecular (COI-5P) and morpho-anatomical tools (presence of sori and mucilage ducts in the stipe) was used to differentiate specimens of H. nigripes and L. digitata. Both kelp species were indistinguishable in most cases by external blade and stipe morphology. The different blade shapes represented different ontogenetic stages rather than phenotypic plasticity. The presence of mucilage ducts in the stipe was correlated with H. nigripes and changed with depth from 17%, 36%, and 85% at 2.5, 5, and 7.5 m, respectively. In addition, all summer fertile specimens were L. digitata., (© 2020 Phycological Society of America.)

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., Competing Interests: The authors have declared that no competing interests exist.

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]

This text provides a summary of the life and work of Klaus Lüning, a prominent researcher in the field of seaweed biology. Lüning's career spanned several decades, during which he made significant contributions to the study of seaweed communities and their environmental requirements. He conducted research on the thermal requirements of seaweeds and investigated the circadian rhythms of growth in kelp species. Lüning also explored the presence of the hormone melatonin in seaweeds. Throughout his career, he supervised numerous students and collaborated with researchers from around the world. After retiring, Lüning established an algal farm and continued his research on Ulva. Unfortunately, he passed away in 2023 after battling cancer. Lüning's pioneering work and dedication to science left a lasting impact on the field of macroalgal research. [Extracted from the article]

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]

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 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]

Kelps are important providers and constituents of marine ecological niches, the coastal kelp forests. Kelp species have differing distribution ranges, but mainly thrive in temperate and arctic regions. Although the principal factors determining biogeographic distribution ranges are known, genomics could provide additional answers to this question. We sequenced DNA from two Laminaria species with contrasting distribution ranges, Laminaria digitata and Laminaria solidungula. Laminaria digitata is found in the Northern Atlantic with a southern boundary in Brittany (France) or Massachusetts (USA) and a northern boundary in the Arctic, whereas L. solidungula is endemic to the Arctic only. From the raw reads of DNA, we reconstructed both chloroplast genomes and annotated them. A concatenated data set of all available brown algae chloroplast sequences was used for the calculation of a robust phylogeny, and sequence variations were analyzed. The two Laminaria chloroplast genomes are collinear to previously analyzed kelp chloroplast genomes with important exceptions. Rearrangements at the inverted repeat regions led to the pseudogenization of ycf37 in L. solidungula, a gene possibly required under high light conditions. This defunct gene might be one of the reasons why the habitat range of L. solidungula is restricted to lowlight sublittoral sites in the Arctic. The inheritance pattern of single nucleotide polymorphisms suggests incomplete lineage sorting of chloroplast genomes in kelp species. Our analysis of kelp chloroplast genomes shows that not only evolutionary information could be gleaned from sequence data. Concomitantly, those sequences can also tell us something about the ecological conditions which are required for species well‐being. [ABSTRACT FROM AUTHOR]

To study the effect of different radiation conditions on sporogenesis of Laminaria digitata (Huds.) J. V. Lamour., excised disks were induced to form sporangia under PAR (P), PAR + ultraviolet-A (UVA) (PA), and PAR + UVA + ultraviolet-B (UVB) (PAB) conditions in the laboratory. Vitality of meiospores, released from sori induced under different radiation conditions in the laboratory and from sori of wild sporophytes acclimated to in situ solar radiation in the presence and absence of ultraviolet radiation (UVR), was measured in terms of their germination capacity. Sorus induction in disks of laboratory-grown sporophytes was not hampered under light supplemented with UVR, and sorus area was not significantly different among P, PA, and PAB. Vitality and germination rate of meiospores released from sori induced under different radiation treatments was comparable. Likewise, screening of UVR of the natural solar radiation did not promote higher germination rates of meiospores released from wild sporophytes. Germination rates were, however, higher in meiospores released from laboratory-induced sori compared to sori of wild sporophytes. Higher DNA damage (formation of cyclobutane pyrimidine dimers, CPDs) was observed in laboratory-grown nonsorus compared to sorus tissue, while CPDs were nondetectable in both sorus and nonsorus tissue of wild sporophytes. To explain the apparent protection of developing meiospores and the unexpected UV resistance of soral tissue, concurrent anatomical investigations of sporogenic tissue were performed. We observed the previously unreported existence of two types of sterile paraphysis cells. One type of paraphysis cells, the most frequent type, contained several red-fluorescing plastids. The other type, less frequently occurring, was completely filled with substances emitting blue fluorescence under violet excitation, presumably brown algal phenolic compounds (phlorotannins). Cells of this type were irregularly scattered within the sorus and did not contain red-fluorescing plastids. Meiospore-containing sporangia were positioned embedded between both types of paraphysis cells. In vegetative tissue, blue autofluorescence was observed only in injured parts of the blade. Results of our study suggest that the sorus structure with phlorotannins localized in the specialized paraphysis cells may be able to screen harmful UVR and protect UV-sensitive meiospores inside the sporangia. [ABSTRACT FROM AUTHOR]

This review about the genus Laminaria sensu lato summarizes the extensive literature that has been published since the overview of the genus given by Kain in 1979. The recent proposal to divide the genus into the two genera Laminaria and Saccharina is acknowledged, but the published data are discussed under a 'sensu lato' concept, introduced here. This includes all species which have been considered to be 'Laminaria' before the division of the genus. In detail, after an introduction the review covers recent insights into phylogeny and taxonomy, and discusses morphotypes, ecotypes, population genetics and demography. It describes growth and photosynthetic performance of sporophytes with special paragraphs on the regulation of sporogenesis, regulation by endogenous rhythms, nutrient metabolism, storage products, and salinity tolerance. The biology of microstages is discussed separately. The ecology of these kelps is described with a focus on stress defence against abiotic and biotic factors and the role of Laminaria as habitat, its trophic interactions and its competition is discussed. Finally, recent developments in aquaculture are summarized. In conclusion to each section, as a perspective and guide to future research, we draw attention to the remaining gaps in the knowledge about the genus and kelps in general. [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]

Kelp species (Laminariales, Phaeophyceae) are globally widespread along temperate to Polar rocky coastal lines. Here we analyse the mitochondrial and chloroplast genomes of Laminaria rodriguezii , in comparison to the organellar genomes of other kelp species. We also provide the complete mitochondrial genome sequence of another endemic kelp species from a Polar habitat, the Arctic Laminaria solidungula. We compare phylogenetic trees derived from twenty complete mitochondrial and seven complete chloroplast kelp genomes. Interestingly, we found a stretch of more than 700 bp in the mitochondrial genome of L.rodriguezii , which is not present in any other yet sequenced member of the Phaeophyceae. This stretch matches a protein coding region in the mitochondrial genome from Desmarestia viridis , another brown seaweed. Their high similarity suggests that these sequences originated through independent introduction into the two species. Their origin could have been by infection by yet unknown similar mitoviruses, currently only known from fungi and plants. • We present three newly sequenced organellar genomes from the kelp species Laminaria rodriguezii and Laminaria solidungula. • Their specific features and phylogenetic information content is analysed and compared to other kelp organellar genomes. • Collinearity of the organellar genomes within the kelps make them apt as robust phylogenetic markers for the entire order. • We found an independent introduction of homologous nucleotide sequences into two brown algal mitochondrial genomes. • Their potential mitovirus origin is unprecedented and should be the beginning of endeavours to search for similar sequences. [ABSTRACT FROM AUTHOR]