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Branching stipe morphologies have evolved multiple times across the kelp (Laminariales) lineage, creating morphological forms that drive the complexity of kelp forest habitats. Although branching is likely a complicated developmental process, it has evolved repeatedly through kelp evolution and the processes facilitating the emergence of branched forms from unbranched ancestors remain unclear. Here I report on abnormally branched individuals (n = 9) from five kelp species found in British Columbia, Canada that had atypical bifurcations in their stipes, creating a single dichotomous branch. One of these species generally lacks branching entirely (Laminaria ephemera) while the other four exhibit some branching but typically lack this stipe bifurcation (Alaria marginata, Laminaria setchellii, Nereocystis luetkeana, Pterygophora californica). These unusually branched individuals exhibited replicated morphological subunits distal to the stipe bifurcation, including more blades, pneumatocysts, and sporophylls than is typical. This suggests that unbranched species possess an inherent developmental capacity for modularity with autonomy in the development of individual modules that may have helped to facilitate the widespread emergence of branched morphologies. Given the role of kelp forests in coastal environments, branching may influence habitat characteristics, potentially influencing community dynamics, and is thus a trait of particular evolutionary interest. These findings highlight the need for experiments that manipulate kelp development to better characterise the ontogenetic processes of these globally important taxa.

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The phylogeny of brown algae (Phaeophyceae) has undergone extensive changes in the recent past due to regular new scientific insights. We used nuclear 18S rDNA with an extensive dataset, aiming to increase the accuracy and robustness of the reconstructed phylogenetic trees using a simultaneous sequence-structure approach. Individual secondary structures were generated for all 18S rDNA sequences. The sequence-structure information was encoded and used for an automated simultaneous sequence-structure alignment. Neighbor-joining and profile neighbor-joining trees were calculated based on 186 phaeophycean sequence-structure pairs. Additionally, sequence-structure neighbor-joining, maximum parsimony and maximum likelihood trees were reconstructed on a representative subset. Using a similar approach, ITS2 rDNA sequence-structure information was used to reconstruct a neighbor-joining tree including 604 sequence-structure pairs of the Laminariales. Our study results are in significant agreement with previous single marker 18S and ITS2 rDNA analyses. Moreover, the 18S results are in wide agreement with recent multi-marker analyses. The bootstrap support was significantly higher for our sequence-structure analysis in comparison to sequence-only analyses in this study and the available literature. This study supports the simultaneous inclusion of sequence-structure data at least for 18S to obtain more accurate and robust phylogenetic trees compared to sequence-only analyses.

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Kelp forests are highly productive and economically important ecosystems worldwide, especially in the North Pacific Ocean. However, current hypotheses for their evolutionary origins are reliant on a scant fossil record. Here, we report fossil hapteral kelp holdfasts from western Washington State, USA, indicating that kelp has existed in the northeastern Pacific Ocean since the earliest Oligocene. This is consistent with the proposed North Pacific origin of kelp associated with global cooling around the Eocene-Oligocene transition. These fossils also support the hypotheses that a hapteral holdfast, rather than a discoid holdfast, is the ancestral state in complex kelps and suggest that early kelps likely had a flexible rather than a stiff stipe. Early kelps were possibly grazed upon by mammals like desmostylians, but fossil evidence of the complex ecological interactions known from extant kelp forests is lacking. The fossil record further indicates that the present-day, multi-story kelp forest had developed at latest after the mid-Miocene climate optimum. In summary, the fossils signify a stepwise evolution of the kelp ecosystem in the North Pacific, likely enabled by changes in the ocean-climate system.

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BACKGROUND AND AIMS: Kelp forests are increasingly considered blue carbon habitats for ocean-based biological carbon dioxide removal, but knowledge gaps remain in our understanding of their carbon cycle. Of particular interest is the remineralization of detritus, which can remain photosynthetically active. Here, we study a widespread, thermotolerant kelp (Ecklonia radiata) to explore detrital photosynthesis as a mechanism underlying temperature and light as two key drivers of remineralization. METHODS: We used meta-analysis to constrain the thermal optimum (Topt) of E. radiata. Temperature and light were subsequently controlled over a 119-day ex situ decomposition experiment. Flow-through experimental tanks were kept in darkness at 15 °C or under a subcompensating maximal irradiance of 8 µmol photons m-2 s-1 at 15, 20 or 25 °C. Photosynthesis of laterals (analogues to leaves) was estimated using closed-chamber oxygen evolution in darkness and under a saturating irradiance of 420 µmol photons m-2 s-1. KEY RESULTS: T opt of E. radiata is 18 °C across performance variables (photosynthesis, growth, abundance, size, mass and fertility), life stages (gametophyte and sporophyte) and populations. Our models predict that a temperature of >15 °C reduces the potential for E. radiata detritus to be photosynthetically viable, hence detrital Topt ≤ 15 °C. Detritus is viable under subcompensating irradiance, where it performs better than in darkness. Comparison of net and gross photosynthesis indicates that elevated temperature primarily decreases detrital photosynthesis, whereas darkness primarily increases detrital respiration compared with optimal experimental conditions, in which detrital photosynthesis can persist for ≥119 days. CONCLUSIONS: T opt of kelp detritus is ≥3 °C colder than that of the intact plant. Given that E. radiata is one of the most temperature-tolerant kelps, this suggests that photosynthesis is generally more thermosensitive in the detrital phase, which partly explains the enhancing effect of temperature on remineralization. In contrast to darkness, even subcompensating irradiance maintains detrital viability, elucidating the accelerating effect of depth and its concomitant light reduction on remineralization to some extent. Detrital photosynthesis is a meaningful mechanism underlying at least two drivers of remineralization, even below the photoenvironment inhabited by the attached alga.

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Brown macroalgae are an important source of polysaccharides, mainly fucose-containing sulphated polysaccharides (FCSPs), associated with several biological activities. However, the structural diversity and structure-function relationships for their bioactivities are still undisclosed. Thus, the aim of this work was to characterize the chemical structure of water-soluble Saccharina latissima polysaccharides and evaluate their immunostimulatory and hypocholesterolemic activities, helping to pinpoint a structure-activity relationship. Alginate, laminarans (F1, neutral glucose-rich polysaccharides), and two fractions (F2 and F3) of FCSPs (negatively charged) were studied. Whereas F2 is rich in uronic acids (45 mol%) and fucose (29 mol%), F3 is rich in fucose (59 mol%) and galactose (21 mol%). These two fractions of FCSPs showed immunostimulatory activity on B lymphocytes, which could be associated with the presence of sulphate groups. Only F2 exhibited a significant effect in reductions in in vitro cholesterol's bioaccessibility attributed to the sequestration of bile salts. Therefore, S. latissima FCSPs were shown to have potential as immunostimulatory and hypocholesterolemic functional ingredients, where their content in uronic acids and sulphation seem to be relevant for the bioactive and healthy properties.

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The culturing and investigation of individual marine specimens in lab environments is crucial to further our understanding of this highly complex ecosystem. However, the obtained results and their relevance are often limited by a lack of suitable experimental setups enabling controlled specimen growth in a natural environment while allowing for precise monitoring and in-depth observations. In this work, we explore the viability of a microfluidic device for the investigation of the growth of the alga Saccharina latissima to enable high-resolution imaging by confining the samples, which usually grow in 3D, to a single 2D plane. We evaluate the specimen's health based on various factors such as its growth rate, cell shape, and major developmental steps with regard to the device's operating parameters and flow conditions before demonstrating its compatibility with state-of-the-art microscopy imaging technologies such as the skeletonisation of the specimen through calcofluor white-based vital staining of its cell contours as well as the immunolocalisation of the specimen's cell wall. Furthermore, by making use of the on-chip characterisation capabilities, we investigate the influence of altered environmental illuminations on the embryonic development using blue and red light. Finally, live tracking of fluorescent microspheres deposited on the surface of the embryo permits the quantitative characterisation of growth at various locations of the organism.

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Ploidy variants can be utilized to increase yield, introduce sterility, and modify specific traits with an economic impact. Despite economic importance of Saccharina species, their nuclear DNA content in different cell types and life stages remain unclear. The present research was initiated to determine the nuclear DNA content and intraindividual variation at different life cycle stages of the Laminarialean kelp Saccharina latissima. Nuclear DNA content in embryonic and mature sporophytes, released and unreleased zoospores, female, and male gametophytes from Sør-Trøndelag county in Norway were estimated by image analysis using the DNA-localizing fluorochrome DAPI and chicken's red blood cells as a standard. DNA content of a total of 6905 DAPI-stained nuclei was estimated. This is the first study of nuclear DNA content which covered the life cycle of kelp. The lowest level of DNA content (1C) was observed in zoospores with an average of 0.76 pg. Male and female single spore gametophyte cultures presented higher average DNA content, more than double that of zoospores, suggesting the presence of polyteny. Female gametophyte nuclei were slightly larger and more variable in size than those of male gametophytes. The DNA content observed in embryonic sporophytes and in meristoderm cells from older sporophytes (1.51 pg) was 2C as expected and in the range of previously published studies of sporophytes of S. latissima. Mature sporophytes showed intra-plant variation with DNA content values ranging from 2-16C. The main difference was between meristoderm cells (mostly 2C) and cortical and medullary cells (2-16C).

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Marine heatwaves threaten the persistence of kelp forests globally. However, the observed responses of kelp forests to these events have been highly variable on local scales. Here, we synthesize distribution data from an environmentally diverse region to examine spatial patterns of canopy kelp persistence through an unprecedented marine heatwave. We show that, although often overlooked, temperature variation occurring at fine spatial scales (i.e., a few kilometers or less) can be a critical driver of kelp forest persistence during these events. Specifically, though kelp forests nearly all persisted toward the cool outer coast, inshore areas were >3°C warmer at the surface and experienced extensive kelp loss. Although temperatures remained cool at depths below the thermocline, kelp persistence in these thermal refugia was strongly constrained by biotic interactions, specifically urchin populations that increased during the heatwave and drove transitions to urchin barrens in deeper rocky habitat. Urchins were, however, largely absent from mixed sand and cobble benthos, leading to an unexpected association between bottom substrate and kelp forest persistence at inshore sites with warm surface waters. Our findings demonstrate both that warm microclimates increase the risk of habitat loss during marine heatwaves and that biotic interactions modified by these events will modulate the capacity of cool microclimates to serve as thermal refugia.

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Kelp forest ecosystems and their associated ecosystem services are declining around the world. In response, marine managers are working to restore and counteract these declines. Kelp restoration first started in the 1700s in Japan and since then has spread across the globe. Restoration efforts, however, have been largely disconnected, with varying methodologies trialled by different actors in different countries. Moreover, a small subset of these efforts are 'afforestation', which focuses on creating new kelp habitat, as opposed to restoring kelp where it previously existed. To distil lessons learned over the last 300 years of kelp restoration, we review the history of kelp restoration (including afforestation) around the world and synthesise the results of 259 documented restoration attempts spanning from 1957 to 2020, across 16 countries, five languages, and multiple user groups. Our results show that kelp restoration projects have increased in frequency, have employed 10 different methodologies and targeted 17 different kelp genera. Of these projects, the majority have been led by academics (62%), have been conducted at sizes of less than 1 ha (80%) and took place over time spans of less than 2 years. We show that projects are most successful when they are located near existing kelp forests. Further, disturbance events such as sea-urchin grazing are identified as regular causes of project failure. Costs for restoration are historically high, averaging hundreds of thousands of dollars per hectare, therefore we explore avenues to reduce these costs and suggest financial and legal pathways for scaling up future restoration efforts. One key suggestion is the creation of a living database which serves as a platform for recording restoration projects, showcasing and/or re-analysing existing data, and providing updated information. Our work establishes the groundwork to provide adaptive and relevant recommendations on best practices for kelp restoration projects today and into the future.

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Undaria pinnatifida is the commercially second most important brown alga in the world. Its global annual yield has been more than two million tonnes since 2012. It is extensively cultivated in East Asia, mainly consumed as food but also used as feed for aquacultural animals and raw materials for extraction of chemicals applicable in pharmaceutics and cosmetics. Cultivar breeding, which is conducted on the basis of characteristics of the life history, plays a pivotal role in seaweed farming industry. The common basic life history shared by kelps determines that their cultivar breeding strategies are similar. Cultivar breeding and cultivation methods of U. pinnatifida have usually been learned or directly transferred from those of Saccharina japonica. However, recent studies have revealed certain peculiarity in the life history of U. pinnatifida. In this article, we review the studies relevant to cultivar breeding in this alga, including the peculiar component of the life history, and the genetics, transcriptomics and genomics tools available, as well as the main cultivar breeding methods. Then we discuss the prospects of cultivar breeding based on our understanding of this kelp and what we can learn from the model brown alga and land crops.

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Organellar genomes serve as useful models for genome evolution and contain some of the most widely used phylogenetic markers, but they are poorly characterized in many lineages. Here, we report 20 novel mitochondrial genomes and 16 novel plastid genomes from the brown algae. We focused our efforts on the orders Chordales and Laminariales but also provide the first plastid genomes (plastomes) from Desmarestiales and Sphacelariales, the first mitochondrial genome (mitome) from Ralfsiales and a nearly complete mitome from Sphacelariales. We then compared gene content, sequence evolution rates, shifts in genome structural arrangements, and intron distributions across lineages. We confirm that gene content is largely conserved in both organellar genomes across the brown algal tree of life, with few cases of gene gain or loss. We further show that substitution rates are generally lower in plastid than mitochondrial genes, but plastomes are more variable in gene arrangement, as mitomes tend to be colinear even among distantly related lineages (with exceptions). Patterns of intron distribution across organellar genomes are complex. In particular, the mitomes of several laminarialean species possess group II introns that have T7-like ORFs, found previously only in mitochondrial genomes of Pylaiella spp. (Ectocarpales). The distribution of these mitochondrial introns is inconsistent with vertical transmission and likely reflects invasion by horizontal gene transfer between lineages. In the most extreme case, the mitome of Hedophyllum nigripes is ∼40% larger than the mitomes of close relatives because of these introns. Our results provide substantial insight into organellar evolution across the brown algae.

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PREMISE: The resistance of macroalgae to hydrodynamic forces imposed by ambient water motion depends in part on the mechanical properties of their tissues. In wave-swept habitats, tissues are stretched (strained) at different rates as hydrodynamic forces change. Previous studies of mechanical properties of macroalgal tissues have used either a single strain rate or a small range of strain rates. Therefore, our knowledge of the mechanical properties of macroalgae is limited to a narrow fraction of the strain rates that can occur in nature. In addition, although mechanical properties of macroalgal tissues change with age, the effect of age on the strain-rate dependence of their mechanical behavior has not been documented. METHODS: Using the kelp Egregia menziesii, we measured how high strain rate (simulating wave impingement) and low strain rate (simulating wave surge) affected mechanical properties of frond tissues of various ages. RESULTS: Stiffness of tissues of all ages increased with strain rate, whereas extensibility was unaffected. Strength and toughness increased with strain rate for young tissue but were unaffected by strain rate for old tissue. CONCLUSIONS: Young tissue is weaker than old tissue and, therefore, the most susceptible to breakage from hydrodynamic forces. The increased strength of young tissue at high strain rates can help the frond resist breaking when pulled rapidly during wave impingement, when hydrodynamic forces are largest. Because breakage of young tissue can remove a frond's meristem and negatively impact the survival of the whole kelp, strain-rate dependence of the young tissue's strength can enhance kelp's survival.

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Kelps are canopy-forming brown seaweed sustaining critical ecosystem services in coastal habitats, including shelter, nursery grounds, and providing food resources to a myriad of associated species. This study modeled the fundamental niche of Laminaria abyssalis along the Brazilian continental margin, an endemic species of the South Atlantic, to anticipate potential distributional range shifts under two contrasting scenarios of future environmental changes (RCP2.6 and RCP8.5). The model for fundamental niche predictions considering the "present scenario" has shown a wider potential area than the realized niche (i.e., the area where the species actually occurs) along the Brazilian coast. In both future scenarios, the models have shown niche erosion on the northern portion of the Brazilian coast and niche gains towards the south. In both scenarios, L. abyssalis populations tend to shift to deeper regions of the reef. The restricted range of occurrence (33,000 km2), intense anthropic activities along these beds (e.g., trawling fisheries, oil/gas mining, or removal for agricultural purposes) acting synergically with global warming, may drive this ecosystem to collapse faster than kelp species' ability to adapt. We propose to classify L. abyssalis as Endangered - (EN) under IUCN criteria, and highlight that long-term monitoring of kelp beds is an urgent need to develop effective conservation initiatives to protect such rare and invaluable ecosystem.

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Kelps are colonized by a wide range of microbial symbionts. Among them, endophytic fungi remain poorly studied, but recent studies evidenced yet their high diversity and their central role in algal defense against various pathogens. Thus, studying the metabolic expressions of kelp endophytes under different conditions is important to have a better understanding of their impacts on host performance. In this context, fatty acid composition is essential to a given algae fitness and of interest to food web studies either to measure its nutritional quality or to infer about its contribution to consumers diets. In the present study, Paradendryphiella salina, a fungal endophyte was isolated from Saccharina latissima (L.) and Laminaria digitata (Hudson.) and its fatty acid composition was assessed at increasing salinity and temperature conditions. Results showed that fungal composition in terms of fatty acids displayed algal-dependent trajectories in response to temperature increase. This highlights that C18 unsaturated fatty acids are key components in the host-dependant acclimation of P. salina to salinity and temperature changes.

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ABSTRACT: Kelps (order Laminariales) are foundation species in temperate and arctic seas globally, but they are in decline in many places. Laminarian kelp have an alternation of generations and this poses challenges for experimental studies due to the difficulties in achieving zoospore release and gametophyte growth. Here, we review and synthesize the protocols that have been used to induce zoospore release in kelps to identify commonalities and provide guidance on best practices. We found 171 papers, where zoospore release was induced in four kelp families from 35 different ecoregions. The most commonly treated family was Laminariaceae, followed by Lessoniaceae and the most studied ecoregion was Central Chile, followed by the Southern California Bight. Zoospore release generally involved three steps: a pretreatment which included cleaning of the reproductive tissue to eliminate epiphytic organisms, followed by desiccation of the tissue, and finally a postdesiccation immersion of the reproductive material in a seawater medium for zoospore release. Despite these commonalities, there was a high degree of variation in the detail within each of these steps, even among studies within genera and from the same ecoregions. This suggests either that zoospore release may be relatively insensitive across the Laminariales or that little methods optimization has been undertaken. We suggest that greater attention to standardization of protocols and reporting of methodology and optimization would improve comparisons of kelp zoospore release across species and locations and facilitate a broader understanding of this key, but understudied life history stage. OPEN RESEARCH BADGES: This article has earned an Open Data Badge for making publicly available the digitally-shareable data necessary to reproduce the reported results. The data is available at https://doi.org/10.5061/dryad.0kh1f8j.

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Reconstructing phylogenetic topologies and divergence times is essential for inferring the timing of radiations, the appearance of adaptations, and the historical biogeography of key lineages. In temperate marine ecosystems, kelps (Laminariales) drive productivity and form essential habitat but an incomplete understanding of their phylogeny has limited our ability to infer their evolutionary origins and the spatial and temporal patterns of their diversification. Here, we reconstruct the diversification of habitat-forming kelps using a global genus-level phylogeny inferred primarily from organellar genome datasets, and investigate the timing of kelp radiation. We resolve several important phylogenetic features, including relationships among the morphologically simple kelp families and the broader radiation of complex kelps, demonstrating that the initial radiation of the latter resulted from an increase in speciation rate around the Eocene-Oligocene boundary. This burst in speciation rate is consistent with a possible role of recent climatic cooling in triggering the kelp radiation and pre-dates the origin of benthic-foraging carnivores. Historical biogeographical reconstructions point to a northeast Pacific origin of complex kelps, with subsequent colonization of new habitats likely playing an important role in driving their ecological diversification. We infer that complex morphologies associated with modern kelp forests (e.g. branching, pneumatocysts) evolved several times over the past 15-20 MY, highlighting the importance of morphological convergence in establishing modern upright kelp forests. Our phylogenomic findings provide new insights into the geographical and ecological proliferation of kelps and provide a timeline along which feedbacks between kelps and their food-webs could have shaped the structure of temperate ecosystems.

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Fucoidans are sulphated fucose-rich polysaccharides predominantly found in the cell walls of brown algae. The bioactive properties of fucoidans attract increasing interest from the medico-pharmaceutical industries and may drive an increase in demand of brown algae biomass. In nature, the biochemical composition of brown algae displays a seasonal fluctuation driven by environmental factors and endogenous rhythms. To cultivate and harvest kelps with high yields of fucoidans, knowledge is needed on seasonal variation and impact of environmental conditions on the fucoidan content of brown algae. The relations between the fucoidan content and key environmental factors (irradiance, nutrient availability, salinity and exposure) were examined by sampling natural populations of the common North Atlantic kelps, Saccharina latissima and Laminaria digitata, over a full year at Hanstholm in the North Sea and Aarhus in the Kattegat. In addition, laboratory experiments were carried out isolating the effects of the single factors. The results demonstrated that (1) seasonal variation alters the fucoidan content by a factor of 2-2.6; (2) interspecific differences exist in the concentrations of crude fucoidan (% of dry matter): L. digitata (11%) > S. latissima (6%); and (3) the effects of single environmental factors were not consistent between species or between different conspecific populations. The ambiguous response to single environmental factors complicates prospective directions for manipulating an increased content of fucoidan in a cultivation scenario and emphasizes the need for knowledge on performance of local kelp ecotypes.

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The absorption of anthropogenic CO2 by the oceans is causing a reduction in the pH of the surface waters termed ocean acidification (OA). This could have substantial effects on marine coastal environments where fleshy (non-calcareous) macroalgae are dominant primary producers and ecosystem engineers. Few OA studies have focused on the early life stages of large macroalgae such as kelps. This study evaluated the effects of seawater pH on the ontogenic development of meiospores of the native kelp Macrocystis pyrifera and the invasive kelp Undaria pinnatifida, in south-eastern New Zealand. Meiospores of both kelps were released into four seawater pH treatments (pHT 7.20, extreme OA predicted for 2300; pHT 7.65, OA predicted for 2100; pHT 8.01, ambient pH; and pHT 8.40, pre-industrial pH) and cultured for 15 d. Meiospore germination, germling growth rate, and gametophyte size and sex ratio were monitored and measured. Exposure to reduced pHT (7.20 and 7.65) had positive effects on germling growth rate and gametophyte size in both M. pyrifera and U. pinnatifida, whereas, higher pHT (8.01 and 8.40) reduced the gametophyte size in both kelps. Sex ratio of gametophytes of both kelps was biased toward females under all pHT treatments, except for U. pinnatifida at pHT 7.65. Germling growth rate under OA was significantly higher in M. pyrifera compared to U. pinnatifida but gametophyte development was equal for both kelps under all seawater pHT treatments, indicating that the microscopic stages of the native M. pyrifera and the invasive U. pinnatifida will respond similarly to OA.

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Brown algae have evolved complex multicellularity in the heterokont lineage. They are phylogenetically distant to land plants, fungi and animals. Especially, the members of Laminariales (so-called kelps) have developed highly differentiated tissues. Extracellular matrix (ECM) plays pivotal roles in a number of essential processes in multicellular organisms, such as cell adhesion, cell and tissue differentiations, cell-to-cell communication, and responses to environmental stimuli. In these processes, a set of extracellular secreted proteins called the secretome operates remodeling of the physicochemical nature of ECM and signal transduction by interacting with cell surface proteins and signaling molecules. Characterization of the secretome is a critical step to clarify the contributions of ECM to the multicellularity of brown algae. However, the identity of the brown algal secretome has been poorly understood. In order to reveal the repertory of the brown algal secretome and its involvement in the evolution of Laminariales, we conducted a genome-wide analysis of the brown algal secretome utilizing the published complete genome data of Ectocarpus siliculosus and Saccharina japonica as well as newly obtained RNA-seq data of seven laminarialean species (Agarum clathratum, Alaria crassifolia, Aureophycus aleuticus, Costaria costata, Pseudochorda nagaii, Saccharina angustata and Undaria pinnatifida) largely covering the laminarialean families. We established the in silico pipeline to systematically and accurately detect the secretome by combining multiple prediction algorithms for the N-terminal signal peptide and transmembrane domain within the protein sequence. From 16,189 proteins of E. siliculosus and 18,733 proteins of S. japonica, 552 and 964 proteins respectively were predicted to be classified as the secretome. Conserved domain analysis showed that the domain repertory were very similar to each other, and that of the brown algal secretome was partially common with that of the secretome of other multicellular organisms (land plants, fungi and animals). In the laminarialean species, it was estimated that the gene abundance and the domain architecture of putative ECM remodeling-related proteins were altered compared with those of E. siliculosus, and that the alteration started from the basal group of Laminariales. These results suggested that brown algae have developed their own secretome, and its functions became more elaborated in the more derived members in Laminariales.

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We confirmed the monophyly of the Agaraceae based on phylogenetic analyses of six mitochondrial and six chloroplast gene sequences from Agarum, Costaria, Dictyoneurum, and Thalassiophyllum species, as well as representative species from other laminarialean families. However, the genus Agarum was paraphyletic, comprising two independent clades, A. clathratum/A. turneri and A. fimbriatum/A. oharaense. The latter clade was genetically most closely related to Dictyoneurum spp., and morphologically, the species shared a flattened stipe bearing fimbriae (potential secondary haptera) in the mid- to upper portion. The phylogenetic position of Thalassiophyllum differed between the two datasets: in the chloroplast gene phylogeny, Thalassiophyllum was included in the A. clathratum/A. turneri clade, but in the mitochondrial gene phylogeny, it formed an independent clade at the base of the Agaraceae, the same position it took in the phylogeny when the data from both genomes were combined despite a larger number of bp being contributed by the chloroplast gene sequences. Considering the remarkable morphological differences between Thalassiophyllum and other Agaraceae, and the molecular support, we conclude that Thalassiophyllum should be reinstated as an independent genus. Dictyoneurum reticulatum was morphologically distinguishable from D. californicum due to its midrib, but because of their close genetic relationship, further investigations are needed to clarify species-level taxonomy. In summary, we propose the establishment of a new genus Neoagarum to accommodate A. fimbriatum and A. oharanese and the reinstatement of the genus Thalassiophyllum.

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