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When studying the effects of climate change on eukaryotic organisms we often oversee a major ecological process: the interaction with microbes. Eukaryotic hosts and microbes form functional units, termed holobionts, where microbes play crucial roles in host functioning. Environmental stress may disturb these complex mutualistic relations. Macroalgae form the foundation of coastal ecosystems worldwide and provide important ecosystem services - services they could likely not provide without their microbial associates. Still, today we do not know how environmental stress will affect the macroalgal holobiont in an increasingly changing ocean. In this review, we provide a conceptual framework that contributes to understanding the different levels at which the holobiont and environment interact, and we suggest a manipulative experimental approach as a guideline for future research.

WOS:000569919500022; This study was focused on the phenolic content and associated cosmetic activities of seven brown marine macroalgae:Alaria esculenta,Ascophyllum nodosum,Bifurcaria bifurcata,Fucus serratus,Halidrys siliquosa,Himanthalia elongata, andLaminaria ochroleuca, selected for their abundance in Brittany and their phlorotannin production. Phlorotannins were extracted by maceration followed by a liquid/liquid purification to obtain fractions enriched in phlorotannins. Seasonal and yearly variations in phenolic contents were investigated thanks to the Folin-Ciocalteu assay to find the best harvesting season for each species.H-1 nuclear magnetic resonance (NMR) was used to validate the presence of phenolic compounds and to compare the species in terms of phenolic signals, as they produce different phlorotannin molecules. Activities required for cosmetic applications (i.e., radical scavenging, antioxidant, photoprotective, and anti-aging activities) were assessed by in vitro tests. Fucales, especiallyA. nodosum,F. serratus,H. elongata, andH. siliquosa, showed higher phenolic contents than Laminariales.Temporal variations were apparent for five seaweeds with the highest phenolic content, notably in autumn. The phenolic content was high regardless of the season and antioxidant and photoprotective activities were similar to those of commercial molecules, allowing year-round algal harvesting. The anti-aging activity ofA. nodosumandF. serratusfractions was higher than the active tea flavonol, epigallocatechin gallate. These results are promising for the use of abundant marine macroalgae as a source of natural active ingredients for cosmetic applications.

Seagrass and seaweed habitats constitute hotspots for diversity and ecosystem services in coastal ecosystems. These habitats are subject to anthropogenic pressures, of which eutrophication is one major stressor. Eutrophication favours fast-growing ephemeral algae over perennial macroalgae and seagrasses, causing habitat degradation. However, changes in top-down control, caused by, for example, overfishing, may also have negative impacts on such habitats by decreasing grazer control of ephemeral algae. Meanwhile, systematic analyses estimating top-down effects of predator manipulations across a wide range of studies are missing, limiting the potential use of top-down control measures in coastal management. Here, we review the literature on experiments that test top-down and bottom-up controls in seagrass Zostera marina and seaweed Fucus spp. food webs in the North Atlantic. Using meta-analysis and meta-regression, we compare effect sizes of consumer and nutrient manipulations on primary producers, grazers and mesopredators. Presence of mesopredators on average doubled the biomass of ephemeral algae through trophic cascades, mainly mediated via negative effects on amphipods and isopods. Of the grazers, gastropods had twice as strong a negative effect on ephemeral algae as amphipods/isopods, but responded weakly to both predators and fertilization. In accordance with theory, top-down effects became stronger with eutrophication. Across studies, top-down effects on ephemeral algae at all trophic levels are on par with eutrophication effects. However, the few studies manipulating piscivorous fish make estimates of their top-down effects uncertain.Synthesis and applications. Consistently strong top-down effects in coastal ecosystems call for an integrated ecosystem perspective. Management should consider measures to improve stocks of predatory fish and reduce mesopredators for restoration and conservation of essential seagrass and seaweed habitats, thereby increasing the long-term viability of ecosystem services from coastal habitats. Editor's Choice

Predictive species distribution models are mostly based on statistical dependence between environmental and distributional data and therefore may fail to account for physiological limits and biological interactions that are fundamental when modelling species distributions under future climate conditions. Here, we developed a state-of-the-art method integrating biological theory with survey and experimental data in a way that allows us to explicitly model both physical tolerance limits of species and inherent natural variability in regional conditions and thereby improve the reliability of species distribution predictions under future climate conditions. By using a macroalga-herbivore association (Fucus vesiculosus - Idotea balthica) as a case study, we illustrated how salinity reduction and temperature increase under future climate conditions may significantly reduce the occurrence and biomass of these important coastal species. Moreover, we showed that the reduction of herbivore occurrence is linked to reduction of their host macroalgae. Spatial predictive modelling and experimental biology have been traditionally seen as separate fields but stronger interlinkages between these disciplines can improve species distribution projections under climate change. Experiments enable qualitative prior knowledge to be defined and identify cause-effect relationships, and thereby better foresee alterations in ecosystem structure and functioning under future climate conditions that are not necessarily seen in projections based on non-causal statistical relationships alone.

When studying the effects of climate change on eukaryotic organisms we often oversee a major ecological process: the interaction with microbes. Eukaryotic hosts and microbes form functional units, termed holobionts, where microbes play crucial roles in host functioning. Environmental stress may disturb these complex mutualistic relations. Macroalgae form the foundation of coastal ecosystems worldwide and provide important ecosystem services - services they could likely not provide without their microbial associates. Still, today we do not know how environmental stress will affect the macroalgal holobiont in an increasingly changing ocean. In this review, we provide a conceptual framework that contributes to understanding the different levels at which the holobiont and environment interact, and we suggest a manipulative experimental approach as a guideline for future research.

Induced chemical defense reactions are widespread in marine brown algae. Despite the evidence that the biosynthesis of defense metabolites can be up-regulated upon herbivory, we do not know how this regulation of biosynthetic pathways to secondary metabolites is achieved in brown algae. In higher plants, the phytohormone jasmonic acid (JA) is crucial for the mediation of induced chemical defenses, and several findings of this metabolite from marine sources have been reported. We tested the hypothesis that JA or related metabolites play a role in induced brown algal defense. Quantification of oxylipins with a detection limit around 20 ng g-1 algal tissue did not reveal the presence of JA in the seven examined brown algal species Dictyota dichotoma, Colpomenia peregrina, Ectocarpus fasciculatus, Fucus vesiculosus, Himanthalia elongata, Saccharina latissima (formerly Laminaria saccharina), and Sargassum muticum. Moreover, treatment with ecologically relevant concentrations of JA and methyl jasmonate did not lead to a significant change in the profile of medium- and non-polar metabolites of the tested algae. Only when high concentrations of >= 500 mu g ml-1 medium of the phytohormones were applied that a metabolic response which could be attributed to unspecific stress was observed. Bioassays with D. dichotoma that focused on medium- and non-polar compounds confirmed the lack of a biological role of JA and methyl jasmonate in the induction of algal induced chemical defenses. The phytohormone-treated samples did not exhibit any increased defense potential towards the amphipod Ampithoe longimana and the isopod Paracerceis caudata. JA and related phytohormones, known to be active in higher plants, thus appear to play no role in brown algae for induction of the defense chemicals studied here.

The brown macroalga Cystoseira tamariscifolia is a foundation species along the northeastern Atlantic and western Mediterranean Sea. It occurs from lower intertidal rock pools to the shallow subtidal. Anthropogenic pollution and rising seawater temperatures can threaten its local distributions. In order to address impacts of historical and current environmental changes, to quantify effective dispersal and population connectivity, and to provide genetic tools for restoration and coastal management strategies, we have developed ten microsatellite markers, validated on 48 individuals from a single population. With 2-11 alleles per locus, the observed heterozygosity varied between 0.244 and 0.875. All of the developed microsatellites cross-amplified also on Cystoseira amentacea. The ten microsatellite loci developed here show high genetic diversity, making them useful for connectivity and population genetic studies aimed at small to large spatial scales, and provide essential insight for the development of conservation strategies for this important but threatened foundation species. Fundacao para a Ciencia e Tecnologia (FCT) [EXTANT-EXCL/AAG-GLO/0661/2012] [SFRH/PBD/107878/2015]

The biosorption capability of two marine macroalgae (green Ulva lactuca and brown Fucus vesiculosus) was evaluated in the removal of toxic metals (Hg, Cd and Pb) from saline waters, under realistic conditions. Results showed that, independently of the contamination scenario tested, both macroalgae have a remarkable capacity to biosorb Hg and Pb. In single-contaminant systems, by using only c.a. 500 mg of non-pre-treated algae biomass (size < 200 mu m) per litter, it was possible to achieve removal efficiencies between 96 and 99 % for Hg and up to 86 % for Pb. Despite the higher removal of Hg, equilibrium was reached more quickly for Pb (after 8 h). In multi-contaminant systems, macroalgae exhibited a similar selectivity toward the target metals: Hg > Pb > > Cd, although Pb removal by U. lactuca was more inhibited than that achieved by F. vesiculosus. Under the experimental conditions used, none of the macroalgae was effective to remove Cd (maximum removal of 20 %). In all cases, the kinetics of biosorption was mathematically described with success. Globally, it became clear that the studied macroalgae may be part of simple, efficient, and cost-effective water treatment technologies. Nevertheless, Fucus vesiculosus has greater potential, since it always presented higher initial sorption rates and higher removal efficiencies.

Pressurized liquids, PLE, and enzyme-assisted extraction, EAE, have been tested to improve the extraction of phlorotannins from the seaweed Sargassum muticum. Enzymatic treatment with proteases and carbohydrases, alkaline hydrolysis and PLE with ethanol:water as extracting solvent have been studied in terms of extraction yield, total phenolic content and antioxidant activity (TEAC assay). Results demonstrated that the application of PLE alone provided the highest yields and relevant antioxidant activity. An experimental design was employed to further optimize the PLE extraction conditions; optimum parameters included the use of 160 °C and 95% ethanol. Under these conditions, values of 21.9%, 94.0 mg gallic acid equivalents g−1, 5.018 mg phloroglucinol equivalents g−1 and 1.275 mmol trolox equivalents g−1 were obtained for extraction yield, total phenols, total phlorotannins and TEAC, respectively. A preliminary chemical characterization by liquid chromatography coupled to mass spectrometry provided insight in terms of the mechanisms involved in the different processes., A.P.S.C. thanks the Administrative Department of Science, Technology and Innovation COLCIENCIAS (Colombia) for her Ph.D. Scholarship. M.H. would like to thank MICINN (Spain) for his “Ramón y Cajal” research contract. The authors want to thank Projects AGL2011-29857-C03-01 (MINECO, Spain) and ALIBIRD-CM, S2013/ABI-2728 (Comunidad de Madrid) for the financial support.

Degradation of ecological resources by large-scale disturbances highlights the need to demonstrate biological properties that increase resistance to change and promote the resilience of ecosystem regimes. Coastal eutrophication is a global-scale disturbance that drives ecosystem change by increasing primary production and favouring ephemeral and bloom-forming life-forms. Recent synthesis indicates that consumption processes increase the resistance of coastal communities to nutrient loading by controlling the responses of ephemeral macroalgae. Here we suggest a similar ecological function for canopy cover by demonstrating that the presence of a canopy species modifies both resource and consumer control of bloom-forming algae associated with nutrient enrichment. We tested effects of canopy presence on the interaction between consumer and resource control, by field-manipulations of a dominant canopy forming seaweed (Fucus vesiculosus), grazer presence (dominated by the gastropod Littorina littorea) and nutrient enrichment (common agricultural NPK fertilizer). Canopy cover and grazers jointly controlled strong increases of ephemeral bloom-forming algae (dominated by Ulva spp) from nutrient enrichment; nutrients increased ephemeral recruitment almost 10-fold, but only in the absence of both grazers and canopy cover. Recruitment success of the canopy-forming seaweed itself decreased additively with 56.1, 71.3 and 50.5% from independent effects of canopy cover, grazers and nutrient enrichment, respectively. A meta-analysis of nine nutrient enrichment experiments including seaweed, seagrass and stream communities, showed that in the presence of canopies average nutrient effects were reduced by more than 90% compared to without canopies. This corroborates the generality of our finding that dominating canopy species are important for aquatic ecosystems by increasing community resistance to the propagation of nutrient effects.

Degradation of ecological resources by large-scale disturbances highlights the need to demonstrate biological properties that increase resistance to change and promote the resilience of ecosystem regimes. Coastal eutrophication is a global-scale disturbance that drives ecosystem change by increasing primary production and favouring ephemeral and bloom-forming life-forms. Recent synthesis indicates that consumption processes increase the resistance of coastal communities to nutrient loading by controlling the responses of ephemeral macroalgae. Here we suggest a similar ecological function for canopy cover by demonstrating that the presence of a canopy species modifies both resource and consumer control of bloom-forming algae associated with nutrient enrichment. We tested effects of canopy presence on the interaction between consumer and resource control, by field-manipulations of a dominant canopy forming seaweed (Fucus vesiculosus), grazer presence (dominated by the gastropod Littorina littorea) and nutrient enrichment (common agricultural NPK fertilizer). Canopy cover and grazers jointly controlled strong increases of ephemeral bloom-forming algae (dominated by Ulva spp) from nutrient enrichment; nutrients increased ephemeral recruitment almost 10-fold, but only in the absence of both grazers and canopy cover. Recruitment success of the canopy-forming seaweed itself decreased additively with 56.1, 71.3 and 50.5% from independent effects of canopy cover, grazers and nutrient enrichment, respectively. A meta-analysis of nine nutrient enrichment experiments including seaweed, seagrass and stream communities, showed that in the presence of canopies average nutrient effects were reduced by more than 90% compared to without canopies. This corroborates the generality of our finding that dominating canopy species are important for aquatic ecosystems by increasing community resistance to the propagation of nutrient effects.

The southern sea palm kelp Eisenia arborea produces wide, bullate (bumpy) blades in low-flow areas, whereas in adjacent high-flow areas blades are flat and narrow. Here we determine if morphological differences in these 2 closely associated populations are correlated with physical factors in the environment, and whether this response is a genetically fixed or plastic trait. Both phenotypes were subjected to morphometric analysis, field and laboratory transplant experiments, and genetic analysis (M13 DNA fingerprinting). Physical variables (water motion, temperature, and light) were monitored simultaneously in the field. Results indicate that the 2 populations, separated by

The global redistribution of biodiversity will intensify in the coming decades of climate change, making projections of species range shifts and of associated genetic losses important components of conservation planning. Highly-structured marine species, notably brown seaweeds, often harbor unique genetic variation at warmer low-latitude rear edges and thus are of particular concern. Here, a combination of Ecological Niche Models (ENMs) and molecular data is used to forecast the potential near-future impacts of climate change for a warm-temperate, canopy forming seaweed, Bifurcaria bifurcata. ENMs for B. bifurcata were developed using marine and terrestrial climatic variables, and its range projected for 2040-50 and 2090-2100 under two greenhouse emission scenarios. Geographical patterns of genetic diversity were assessed by screening 18 populations spawning the entire distribution for two organelle genes and 6 microsatellite markers. The southern limit of B. bifurcata was predicted to shift northwards to central Morocco by the mid-century. By 2090-2100, depending on the emission scenario, it could either retreat further north to western Iberia or be relocated back to Western Sahara. At the opposing margin, B. bifurcata was predicted to expand its range to Scotland or even Norway. Microsatellite diversity and endemism were highest in Morocco, where a unique and very restricted lineage was also identified. Our results imply that B. bifurcata will maintain a relatively broad latitudinal distribution. Although its persistence is not threatened, the predicted extirpation of a unique southern lineage or even the entire Moroccan diversity hotspot will erase a rich evolutionary legacy and shrink global diversity to current (low) European levels. NW Africa and similarly understudied southern regions should receive added attention if expected range changes and diversity loss of warm-temperate species is not to occur unnoticed. Portuguese FCT (Fundacao para a Ciencia e a Tecnologia) [PTDC/AAC-CLI/109108/2008, EXPL/BIA-BIC/1471/2012, EXCL/AAG-GLO/0661/2012]; [SFRH/BPD/88935/2012] info:eu-repo/semantics/publishedVersion

Here we describe a rapid method for extracting DNA from dried brown algae material using a microtitre plate system in conjunction with a milling instrument. The method allows the preparation of nuclear and organelle DNA of quality suitable for polymerase chain reaction amplification. It combines high throughput with low cost per sample: DNA from 192 samples can be extracted in c. 3 h for

Plants optimise their resistance to herbivores by regulating deterrent responses on demand. Induction of anti-herbivory defences can occur directly in grazed plants or from emission of risk cues to the environment, which modifies interactions of adjacent plants with, for instance, their consumers. This study confirmed the induction of anti-herbivory responses by water-borne risk cues between adjoining con-specific seaweeds and firstly examined whether plant-plant signalling also exists among adjacent hetero-specific seaweeds. Furthermore, differential abilities and geographic variation in plant-plant signalling by a non-indigenous seaweed as well as native seaweeds were assessed. Twelve-day induction experiments using the non-indigenous seaweed Sargassum muticum were conducted in the laboratory in Portugal and Germany with one local con-familiar (Portugal: Cystoseira humilis, Germany: Halidrys siliquosa) and hetero-familiar native species (Portugal: Fucus spiralis, Germany: F. vesiculosus). All seaweeds were grazed by a local isopod species (Portugal: Stenosoma nadejda, Germany: Idotea baltica) and were positioned upstream of con-and hetero-specific seaweeds. Grazing-induced modification in seaweed traits were tested in three-day feeding assays between cue-exposed and cue-free (= control) pieces of both fresh and reconstituted seaweeds. Both Fucus species reduced their palatability when positioned downstream of isopod-grazed con-specifics. Yet, the palatability of non-indigenous S. muticum remained constant in the presence of upstream grazed con-specifics and native hetero-specifics. In contrast, both con-familiar (but neither hetero-familiar) native species reduced palatability when located downstream of grazed S. muticum. Similar patterns of grazer-deterrent responses to water-borne cues were observed on both European shores, and were almost identical between assays using fresh and reconstituted seaweeds. Hence, seaweeds may use plant-plant signalling to optimise chemical resistance to consumers, though this ability appeared to be species-specific. Furthermore, this study suggests that native species may benefit more than a non-indigenous species from water-borne cue mediated reduction in consumption as only natives responded to signals emitted by hetero-specifics. Portuguese Science Foundation (Fundacao para a Ciencia e a Tecnologia, FCT) under Programa Operacional Ciencia e Inovacao do Quadro Comunitario de Apoio III e comparticipado pelo Fundo Comunitario Europeu FEDER [POCI/MAR/55377/2004]; FCT scholarships [SFRH/BPD/7153/2001, SFRH/BPD/63703/2009] info:eu-repo/semantics/publishedVersion

Presence of habitat-forming macroalgae is supposed to mitigate effects of altered resources on benthic microalgae and their consumers. In a field experiment in a microtidal area of the Western Baltic Sea, we tested the interactive effects of nutrient enrichment, artificial shading, and habitat complexity on microalgal biomass and diversity as well as invertebrate abundance and richness. Habitat complexity comprised three levels, the presence of macroalgal canopy of Fucus vesiculosus, the presence of macroalgal propagules, and the absence of both (=control). Microalgal biomass (and richness) was significantly reduced by canopy presence (-88%, compared to control) and shading (-42%), with the highest biomass in the absence of both canopy and macroalgal propagules at ambient light. Within the microalgal assemblage, higher biomass was related to lower evenness (higher dominance). Density of two main invertebrate groups (snails and amphipods) strongly increased with canopy presence (on average from 53 to 154 individuals m(-2) stone area for snails, and from 234 to 1203 individuals m-2 for amphipods) and so did invertebrate richness (from 4.3 to 10.3). Additionally, snail density doubled with increasing light availability. The snail responses to light and canopy were independent, the former relating to higher availability of microalgal prey, the latter to more structure. Microalgal taxon richness and biomass decreased with increasing invertebrate richness and with density of snails and amphipods. Our experiment thus showed that the presence of habitat-forming macroalgae alters biomass and diversity across trophic levels in benthic coastal communities as well as their response to resource manipulations. (C) 2011 Elsevier B.V. All rights reserved.

The unicellular red alga Rhodella maculata Evans is reported for the first time from the central Baltic Sea (habitat salinity 5 psu). To evaluate whether this Baltic isolate is locally adapted to the brackish environment, its salt stress response was compared with a marine isolate (33 psu) by measuring growth rates, photosynthetic performance and the concentration of the osmolyte mannitol as a function of changing salinity between 1 and 60 psu. The brackish Baltic isolate grew between 5 and 40 psu, a narrower salinity range than in the marine isolate (1-60 psu). The marine isolate of R. maculata showed adaptations to an intertidal habitat with fluctuating salinities rather than to stable salinity conditions. The cells of the marine isolate performed a pronounced regulation of primary photosynthesis via non-photochemical quenching (NPQ), and accumulation of mannitol was involved in osmotic acclimation as intracellular concentrations increased considerably with rising salinities from 1 to 180 mu mol g(-1) dry weight. At its habitat salinity (5 psu), the brackish isolate achieved only 39% of the growth rate measured at 33 psu; when exposed to higher salinities it lacked the ability to carry out substantial NPQ regulation, and it did not accumulate mannitol. However, when cultured at 33 psu for more than three months, the Baltic isolate showed a considerably improved salinity response in the higher salinity range (33-60 psu). Thus, the Baltic Sea isolate of R. maculata does not seem to be locally adapted to its brackish environment, and should be regarded as a stressed population at its distributional limit.

Here we describe a rapid method for extracting DNA from dried brown algae material using a microtitre plate system in conjunction with a milling instrument. The method allows the preparation of nuclear and organelle DNA of quality suitable for polymerase chain reaction amplification. It combines high throughput with low cost per sample: DNA from 192 samples can be extracted in c. 3 h for