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Background: Rockweeds are among the most important foundation species of temperate rocky littoral shores. In the Baltic Sea, the rockweed Fucus vesiculosus is distributed along a decreasing salinity gradient from the North Atlantic entrance to the low-salinity regions in the north-eastern margins, thus, demonstrating a remarkable tolerance to hyposalinity. The underlying mechanisms for this tolerance are still poorly understood. Here, we exposed F. vesiculosus from two range-margin populations to the hyposaline (2.5 PSU - practical salinity unit) conditions that are projected to occur in the region by the end of this century as a result of climate change. We used transcriptome analysis (RNA-seq) to determine the gene expression patterns associated with hyposalinity acclimation, and examined the variation in these patterns between the sampled populations., Results: Hyposalinity induced different responses in the two populations: in one, only 26 genes were differentially expressed between salinity treatments, while the other population demonstrated up- or downregulation in 3072 genes. In the latter population, the projected future hyposalinity induced an acute response in terms of antioxidant production. Genes associated with membrane composition and structure were also heavily involved, with the upregulation of fatty acid and actin production, and the downregulation of ion channels and alginate pathways. Changes in gene expression patterns clearly indicated an inhibition of the photosynthetic machinery, with a consequent downregulation of carbohydrate production. Simultaneously, energy consumption increased, as revealed by the upregulation of genes associated with respiration and ATP synthesis. Overall, the genes that demonstrated the largest increase in expression were ribosomal proteins involved in translation pathways. The fixation rate of SNP:s was higher within genes responding to hyposalinity than elsewhere in the transcriptome., Conclusions: The high fixation rate in the genes coding for salinity acclimation mechanisms implies strong selection for them. The among-population differentiation that we observed in the transcriptomic response to hyposalinity stress suggests that populations of F. vesiculosus may differ in their tolerance to future desalination, possibly as a result of local adaptation to salinity conditions within the Baltic Sea. These results emphasise the importance of considering interspecific genetic variation when evaluating the consequences of environmental change.

Due to increased sea transport and offshore gas and oil exploration, the Arctic is facing an unprecedented risk of marine oil spills. Although beached oil spills can lead to acute and chronic impacts on intertidal ecosystems, the effects of oil spills on macro-algae in Arctic ecosystems is lacking. Here, we assessed the effect and response of the tidal macro-algae Fucus distichus to oiling, i.e. self-cleaning potential by seawater wash and photosynthetic activity. Oiling with four oil types (ANS, Grane, IFO30 and MGO) was simulated by exposing F. distichus tips to oil. Oil removal half-times ranged between 0.8 - 4.5 days, indicating that oiling of macro-algae with the tested oils was short-term. Further, Grane oil mostly inhibited photosynthetic activity whereas oil from ANS, IFO30 and MGO stimulated it. The photosynthetic activity of F. distichus continued to be affected (inhibited or stimulated), even after oil on the tip surface was washed off., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

The viability and physiological state of brown macroalgae Fucus vesiculosus and its associated epiphytic bacteria exposed to diesel water-accommodated fraction (WAF), as well as the capacity of this association to deplete petroleum hydrocarbons (HCs) were experimentally tested. After a 6-day exposure treatment, the algal-surface associated bacteria were identified as primarily hydrocarbon-oxidising bacteria (HOB), and the algal-HOB association was able to deplete petroleum hydrocarbons from the diesel WAF by 80%. The HOB density on the algal surface exposed to diesel WAF was 350% higher compared to the control (i.e. HOB density on the algal surface exposed to ambient seawater), which suggest that they actively proliferated in the presence of hydrocarbons and most likely consumed hydrocarbons as their primary organic substrate. Exposure to diesel WAF did not affect the metabolic activity of F. vesiculosus. Higher lipid peroxidation was observed in F. vesiculosus exposed to diesel WAF while catalase concentration decreased only during the first day of exposure. Results suggest F. vesiculosus is tolerant to oil pollution and the algal-HOB association can efficiently deplete petroleum hydrocarbons in oil-contaminated seas., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Ecological impact of global change is generated by multiple synchronous or asynchronous drivers which interact with each other and with intraspecific variability of sensitivities. In three near-natural experiments, we explored response correlations of full-sibling germling families of the seaweed Fucus vesiculosus towards four global change drivers: elevated CO 2 (ocean acidification, OA), ocean warming (OW), combined OA and warming (OAW), nutrient enrichment and hypoxic upwelling. Among families, performance responses to OA and OW as well as to OAW and nutrient enrichment correlated positively whereas performance responses to OAW and hypoxia anti-correlated. This indicates (i) that families robust to one of the three drivers (OA, OW, nutrients) will also not suffer from the two other shifts, and vice versa and (ii) families benefitting from OAW will more easily succumb to hypoxia. Our results may imply that selection under either OA, OW or eutrophication would enhance performance under the other two drivers but simultaneously render the population more susceptible to hypoxia. We conclude that intraspecific response correlations have a high potential to boost or hinder adaptation to multifactorial global change scenarios.

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.

To predict the effects of climate change, we first need information on both the current tolerance ranges of species and their future adaptive potential. Adaptive responses may originate either in genetic variation or in phenotypic plasticity, but the relative importance of these factors is poorly understood. Here, we tested the tolerance of Fucus radicans to the combination of hyposalinity and warming projected by climate models for 2070-2099. We measured the growth and survival responses of thalli in both current and future conditions, focusing on variations in tolerance among and within different clonal lineages. Survival was 32% lower in future than in current conditions, but the weight and length of the thalli which survived was respectively 267% and 178% higher when exposed to future conditions. The relatively high tolerance to the future conditions suggests that F. radicans is likely to persist in its current distributional range, which is limited to the Gulf of Bothia and Estonian coast in the Baltic Sea. Furthermore, this species may be able to expand its distribution southward and replace its congener F. vesiculosus, which, in previous studies, has not tolerated the future conditions as well. In addition, we discovered variation in tolerance to future conditions within one of the clonal lineages, which have been hitherto presumed to lack adaptive variation. The discovery of intra-clonal phenotypic plasticity means that this alga has the potential for adaptive responses to climate change, which may be the key to the future persistence of F. radicans in the Baltic Sea., (© 2018 Phycological Society of America.)

Coexistence of highly similar species is at odds with ecological theory of competition; coexistence, then, requires stabilizing mechanisms such as differences in ecological niche. In the bacterivore nematode Litoditis marina species complex, which occurs associated with macro-algae, four cryptic lineages (Pm I-IV) co-occur in the field along the south-western coast and estuaries of The Netherlands. Here we investigate the temporal and/or spatial niche differentiation in their natural environment using a qPCR-based detection and relative quantification method. We collected different algal species (i.e. two Fucus species and Ulva sp.) and separated algal structures (i.e. receptacula, thalli, non-fertile tips and bladders) at different sampling months and times (i.e. twice per sampling month), to examine differences in microhabitat use between coexisting L. marina species. Results demonstrate that the cryptic species composition varied among different algal species and algal structures, which was also subject to temporal shifts. Pm I dominated on Fucus spp., Pm II showed dominance on Ulva sp., while Pm III overall had the lowest frequencies. Microhabitat partitioning was most pronounced between the two cryptic species which had similar microbiomes (Pm I and Pm II), and less so between the two species which had significantly different microbiomes (Pm I and Pm III), suggesting that species which share the same microhabitats may avoid competition through resource partitioning. The interplay of microhabitat differentiation and temporal dynamics among the cryptic species of L. marina implies that there is a complex interaction between biotic components and abiotic factors which contributes to their coexistence in the field., Competing Interests: The authors have declared that no competing interests exist.

Poor physiological acclimatization to climate change has led to shifts in the distributional ranges of various species and to biodiversity loss. However, evidence also suggests the relevance of non-climatic physical factors, such as light, and biotic factors, which may act in interactive or additive way. We used a mechanistic approach to evaluate the ecophysiological responses of four seaweed species (three dominant intertidal fucoids, Fucus serratus, Ascophyllum nodosum, Bifurcaria bifurcata, and the invasive Sargassum muticum) to different conditions of grazing, light irradiance and ultraviolet (UV) radiation. We performed a large-scale mesocosm experiment with a total of 800 individual thalli of macroalgae. The factorial experimental design included major algal traits, photoacclimation, nutrient stoichiometry and chemical defence as response variables. Few significant effects of the factors acting alone or in combination were observed, suggesting a good capacity for acclimatization in all four species. The significant effects were generally additive and there were no potentially deleterious synergistic effects between factors. Fucus serratus, a species currently undergoing a drastic contraction of its southern distribution limit in Europe, was the most strongly affected species, showing overall lower photosynthetic efficiency than the other species. The growth rate of F. serratus decreased when UV radiation was filtered out, but only in the presence of grazers. Moreover, more individuals of this species tended to reach maturity in the absence of grazers, and the nitrogen content of tissues decreased under full-spectrum light. Only the phlorotannin content of tissues of B. bifurcata and of exudates of A. nodosum, both slow-growing species, were positively affected by respectively removal of UVB radiation and the presence of grazers. The findings for S. muticum, a well-established invasive seaweed across European coasts, suggested similar physiological response of this fast-growing species to different levels of grazing activity and light quality/intensity. As expected, this species grew faster than the other species. Bifurcaria bifurcata and A. nodosum only showed minor effects of light quality and grazing on phlorotannins content, which suggests good resistance of these two long-lived species to the experimental conditions. Mechanistic approaches that are designed to analyse interactive effects of physical and biotic factors provide an understanding of physiological responses of species and help to improve the confidence of predictive distribution models., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Empirical work on the dynamics of range limits of species distributions often lack replications of edge populations. We compared the local environment and performance of two groups of geographically peripheral populations of the foundation intertidal alga Fucus serratus L. at its southern range boundary in the NW Iberian Peninsula. Two populations were located on the Western Galician coast in large embayments or rias, and the other two on a Northern open coastal stretch in Lugo province. Sharp differences were detected in the local environment and performance of the two groups of populations. While recruitment was quite consistent throughout the year in rias, it was very limited and variable in Lugo. Furthermore, thalli from rias were severely damaged following their transplantation in Lugo, and poor conditions of local adult plants were detected there in subsequent years. These results suggest a higher vulnerability of Lugo populations under new climate conditions, while western rias, strongly influenced by upwelling events, feasibly act as contemporary refugia for this species. If sustained over time, these refugia may mitigate the retreat of the seaweed' rear-edge predicted by large spatial scale models., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Climate change is threating species' persistence worldwide. To predict species responses to climate change we need information not just on their environmental tolerance but also on its adaptive potential. We tested how the foundation species of rocky littoral habitats, Fucus vesiculosus, responds to combined hyposalinity and warming projected to the Baltic Sea by 2070-2099. We quantified responses of replicated populations originating from the entrance, central, and marginal Baltic regions. Using replicated individuals, we tested for the presence of within-population tolerance variation. Future conditions hampered growth and survival of the central and marginal populations whereas the entrance populations fared well. Further, both the among- and within-population variation in responses to climate change indicated existence of genetic variation in tolerance. Such standing genetic variation provides the raw material necessary for adaptation to a changing environment, which may eventually ensure the persistence of the species in the inner Baltic Sea., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Competition and fouling defence are important traits that may facilitate invasions by non-indigenous species. The 'novel weapons hypothesis' (NWH) predicts that the invasive success of exotic species is closely linked to the possession of chemical defence compounds that the recipient community in the new range is not adapted to. In order to assess whether chemical defence traits contribute to invasion success, anti-bacterial, anti-quorum sensing, anti-diatom, anti-larval and anti-algal properties were investigated for the following algae: a) the invasive brown alga Sargassum muticum from both, its native (Japan) and invasive (Germany) range, b) the two non- or weak invasive species Sargassum fusiforme and Sargassum horneri from Japan, and c) Fucus vesiculosus, a native brown alga from Germany. Crude and surface extracts and lipid fractions of active extracts were tested against common fouling organisms and zygotes of a dominant competing brown alga. Extracts of the native brown alga F. vesiculosus inhibited more bacterial strains (75%) than any of the Sargassum spp. (17 to 29%). However, Sargassum spp. from Japan exhibited the strongest settlement inhibition against the diatom Cylindrotheca closterium, larvae of the bryozoan Bugula neritina and zygotes of the brown alga F. vesiculosus. Overall, extracts of S. muticum from the invasive range were less active compared to those of the native range suggesting an adaptation to lower fouling pressure and competition in the new range resulting in a shift of resource allocation from costly chemical defence to reproduction and growth. Non-invasive Sargassum spp. from Japan was equally defended against fouling and competitors like S. muticum from Japan indicating a necessity to include these species in European monitoring programs. The variable antifouling activity of surface and crude extracts highlights the importance to use both for an initial screening for antifouling activity.

In vivo variable chlorophyll fluorescence measurements of photosystem II (PSII) quantum yields in optically dense systems are complicated by steep tissue light gradients due to scattering and absorption. Consequently, externally measured effective PSII quantum yields may be composed of signals derived from cells differentially exposed to actinic light, where cells located deeper inside tissues receive lower irradiance than cells closer to the surface and can display distinct photophysiological status. We demonstrate how measured distributions of PSII quantum yields in plant tissue change under natural tissue light gradients as compared with conventionally measured quantum yields with even exposure to actinic light. This was achieved by applying actinic irradiance perpendicular to one side of thallus cross sections of the aquatic macrophyte Fucus vesiculosus with laser light sheets of defined spectral composition, while imaging variable chlorophyll fluorescence from cross sections with a microscope-mounted pulse amplitude-modulated imaging system. We show that quantum yields are highly affected by light gradients and that traditional surface-based variable chlorophyll fluorescence measurements result in substantial underestimations and/or overestimations, depending on incident actinic irradiance. We present a method for using chlorophyll fluorescence profiles in combination with integrating sphere measurements of reflectance and transmittance to calculate depth-resolved photon absorption profiles, which can be used to correct apparent PSII electron transport rates to photons absorbed by PSII. Absorption profiles of the investigated aquatic macrophyte were different in shape from what is typically observed in terrestrial leaves, and based on this finding, we discuss strategies for optimizing photon absorption via modulation of the structural organization of phytoelements according to in situ light environments., (© 2017 American Society of Plant Biologists. All Rights Reserved.)

Nitrogen stable isotope ratios (δ 15 N) in macroalgae are often used to identify sources of nitrogenous pollution in fluvial and estuarine settings. This approach assumes that the macroalgal δ 15 N is representative of the sources of the pollution averaged over a timespan in the order of days to weeks. The preferential uptake of a particular nitrogen compound or potential for fractionation in the water column or during uptake and assimilation by the macroalgae could make this assumption invalid. Laboratory studies were therefore performed to investigate the uptake and assimilation of both nitrate and ammonium at a variety of concentrations using the vegetative (non-fertile) tips of the brown macroalgae, Fucus vesiculosus. Nitrate appeared to fractionate at high concentrations, and was found to be taken up more rapidly than ammonia; within 13 days, the macroalgae tips were in isotopic equilibrium with the nitrate solution at 500 μM. These experiments were complemented by an investigation involving the translocation of macroalgae collected from a site enriched in 15 N relative to natural levels (Staithes, UK), to the River Tees, Middlesbrough (UK), a site depleted in 15 N relative to natural levels. The nitrogen isotope signature shifted by ∼50% within 7 days, with samples deployed nearer the surface subject to greater change. These findings suggest that the translocation of macroalgae with isotopically distinct signatures can be used as a rapid, cost-efficient method for nitrogen biomonitoring in estuarine environments., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Background: Establishing populations in ecologically marginal habitats may require substantial phenotypic changes that come about through phenotypic plasticity, local adaptation, or both. West-Eberhard's "plasticity-first" model suggests that plasticity allows for rapid colonisation of a new environment, followed by directional selection that develops local adaptation. Two predictions from this model are that (i) individuals of the original population have high enough plasticity to survive and reproduce in the marginal environment, and (ii) individuals of the marginal population show evidence of local adaptation. Individuals of the macroalga Fucus vesiculosus from the North Sea colonised the hyposaline (≥2-3‰) Baltic Sea less than 8000 years ago. The colonisation involved a switch from fully sexual to facultative asexual recruitment with release of adventitious branches that grow rhizoids and attach to the substratum. To test the predictions from the plasticity-first model we reciprocally transplanted F. vesiculosus from the original population (ambient salinity 24‰) and from the marginal population inside the Baltic Sea (ambient salinity 4‰). We also transplanted individuals of the Baltic endemic sister species F. radicans from 4 to 24‰. We assessed the degree of plasticity and local adaptation in growth and reproductive traits after 6 months by comparing the performance of individuals in 4 and 24‰., Results: Branches of all individuals survived the 6 months period in both salinities, but grew better in their native salinity. Baltic Sea individuals more frequently developed asexual traits while North Sea individuals initiated formation of receptacles for sexual reproduction., Conclusions: Marine individuals of F. vesiculosus are highly plastic with respect to salinity and North Sea populations can survive the extreme hyposaline conditions of the Baltic Sea without selective mortality. Plasticity alone would thus allow for an initial establishment of this species inside the postglacial Baltic Sea at salinities where reproduction remains functional. Since establishment, the Baltic Sea populations have evolved adaptations to extreme hyposaline waters and have in addition evolved asexual recruitment that, however, tends to impede local adaptation. Overall, our results support the "plasticity-first" model for the initial colonisation of the Baltic Sea by Fucus vesiculosus.

The role of hyaline hairs on the thallus of brown algae in the genus Fucus is long debated and several functions have been proposed. We used a novel motorized set-up for two-dimensional and three-dimensional mapping with O 2 microsensors to investigate the spatial heterogeneity of the diffusive boundary layer (DBL) and O 2 flux around single and multiple tufts of hyaline hairs on the thallus of Fucus vesiculosus. Flow was a major determinant of DBL thickness, where higher flow decreased DBL thickness and increased O 2 flux between the algal thallus and the surrounding seawater. However, the topography of the DBL varied and did not directly follow the contour of the underlying thallus. Areas around single tufts of hyaline hairs exhibited a more complex mass-transfer boundary layer, showing both increased and decreased thickness when compared with areas over smooth thallus surfaces. Over thallus areas with several hyaline hair tufts, the overall effect was an apparent increase in the boundary layer thickness. We also found indications for advective O 2 transport driven by pressure gradients or vortex shedding downstream from dense tufts of hyaline hairs that could alleviate local mass-transfer resistances. Mass-transfer dynamics around hyaline hair tufts are thus more complex than hitherto assumed and may have important implications for algal physiology and plant-microbe interactions., (© 2017 The Author(s).)

Human-induced ocean warming and acidification have received increasing attention over the past decade and are considered to have substantial consequences for a broad range of marine species and their interactions. Understanding how these interactions shift in response to climate change is particularly important with regard to foundation species, such as the brown alga Fucus vesiculosus. This macroalga represents the dominant habitat former on coastal rocky substrata of the Baltic Sea, fulfilling functions essential for the entire benthic community. Its ability to withstand extensive fouling and herbivory regulates the associated community and ecosystem dynamics. This study tested the interactive effects of future warming, acidification, and seasonality on the interactions of a marine macroalga with potential foulers and consumers. F. vesiculosus rockweeds were exposed to different combinations of conditions predicted regionally for the year 2100 (+∆5°C, +∆700 μatm CO 2 ) using multifactorial long-term experiments in novel outdoor benthic mesocosms ("Benthocosms") over 9-12-week periods in four seasons. Possible shifts in the macroalgal susceptibility to fouling and consumption were tested using consecutive bioassays. Algal susceptibility to fouling and grazing varied substantially among seasons and between treatments. In all seasons, warming predominantly affected anti-fouling and anti-herbivory interactions while acidification had a subtle nonsignificant influence. Interestingly, anti-microfouling activity was highest during winter under warming, while anti-macrofouling and anti-herbivory activities were highest in the summer under warming. These contrasting findings indicate that seasonal changes in anti-fouling and anti-herbivory traits may interact with ocean warming in altering F. vesiculosus community composition in the future., (© 2016 Phycological Society of America.)

In oogamous species of brown algae such as Saccharina japonica and Fucus distichus, the sperm possess an unusual long posterior flagellum, which oscillates actively and produces a propulsive force during swimming. In this study, we quantitatively analyzed the effect of chemotactic responses on sperm swimming and flagellar waveforms by high-speed video recordings. We found that the thigmotactic response to the chemo-attractant was not enhanced during chemotactic swimming and that the swimming velocity of sperm did not decrease. As concentration of the chemo-attractant decreased, the sperm performed drastic U-turn movements, which was caused by a rapid and large bend of the posterior flagellum. Unilateral bending of the posterior flagellum when sensing a decrease in the concentration of the chemo-attractant may be a common response in male gametes during fertilization of brown algae both oogamous and isogamous species.

Intertidal seaweeds must cope with a suite of stressors imposed by aerial exposure at low tide, including nutrient limitation due to emersion. Seaweeds can access nutrients only when submerged, so individuals living higher compared to lower on the shore may have adaptations allowing them to acquire sufficient amounts of nutrients to survive and maintain growth. Using a combination of observations and experiments, we aimed to identify intraspecific variation in nitrate uptake rates across the intertidal distribution of F. vesiculosus, as well as test for acclimation in response to a change in tide height. We replicated our study at sites spanning nearly the entire Gulf of Maine coastline, to examine how local environmental variability may alter intraspecific variation in nitrate uptake. We found that average nitrate uptake rates were ~18% higher in upper compared to lower intertidal Fucus vesiculosus. Furthermore, we found evidence for both acclimation and adaptation to tide height during a transplant experiment. F. vesiculosus transplanted from the lower to the upper intertidal zone was characterized by increased nitrate uptake, but individuals transplanted from the upper to the lower intertidal zone retained high uptake rates. Our observations differed among Gulf of Maine regions and among time points of our study. Importantly, these differences may reflect associations between nitrate uptake rates and abiotic environmental conditions and seaweed nutrient status. Our study highlights the importance of long-term variation in ambient nutrient supply in driving intraspecific variation of seaweeds across the intertidal gradient and local and seasonal variation in ambient nutrient levels in mediating intraspecific differences., (© 2016 Phycological Society of America.)

Prezygotic isolating mechanisms act to limit hybridization and maintain the genetic identity of closely-related species. While synchronous intraspecific spawning is a common phenomenon amongst marine organisms and plays an important role in reproductive success, asynchronous spawning between potentially hybridizing lineages may also be important in maintaining species boundaries. We tested this hypothesis by comparing reproductive synchrony over daily to hourly timescales in a sympatric assemblage of intertidal fucoid algae containing selfing hermaphroditic (Fucus spiralis and Fucus guiryi) and dioecious (Fucus vesiculosus and Fucus serratus) species. Our results confirm that gametes are released on semi-lunar cycles in all species. However, sister species with different mating systems showed asynchronous spawning at finer circadian timescales, thus providing evidence for a partial reproductive barrier between hermaphroditic and dioecious species. Finally, our data also emphasize the ecological, developmental, and/or physiological constraints that operate to restrict reproduction to narrow temporal windows of opportunity in the intertidal zone and more generally the role of ecological factors in marine speciation.

The timing and synchrony of gamete release in broadcast spawners have important implications for fertilization success, recruitment and to explain differences in reproductive success under distinct reproductive modes in sympatry. Our objective was to compare the reproductive timing and investment for sister species with contrasting mating systems; Fucus guiryi (selfing hermaphroditic) and Fucus vesiculosus (dioecious) in habitats with different wave exposures (exposed shore and estuary). Over two months, daily gamete release, recruitment and population structure were recorded. Our results show spawning synchrony between species and habitats, but release events in hermaphrodites occupied broader temporal windows in estuarine than exposed shore habitats. On the exposed shore both species increased the synchrony of release and amount of eggs. In the estuary, hermaphrodites relied on broader temporal spawning windows and a larger canopy, and the dioecious species had higher recruitment success, important factors determining persistence., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

Expected increases in the frequency and intensity of storm surges and river flooding may greatly affect the relative salinity of estuarine environments over the coming decades. In this experiment we used detritus from three contrasting environments (marine Fucus vesiculosus; estuarine Spartina anglica; terrestrial Quercus robur) to test the prediction that the decomposition of the different types of litter would be highest in the environment with which they are associated. Patterns of decomposition broadly fitted our prediction: Quercus detritus decomposed more rapidly in freshwater compared with saline conditions while Fucus showed the opposite trend; Spartina showed an intermediate response. Variation in macro-invertebrate assemblages was detected along the salinity gradient but with different patterns between estuaries, suggesting that breakdown rates may be linked in part to local invertebrate assemblages. Nonetheless, our results suggest that perturbation of salinity gradients through climate change could affect the process of litter decomposition and thus alter nutrient cycling in estuarine transition zones. Understanding the vulnerability of estuaries to changes in local abiotic conditions is important given the need to better integrate coastal proceses into a wider management framework at a time when coastlines are increasingly threatened by human activities., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

Macroalgae live in an ever-changing light environment affected by wave motion, self-shading and light-scattering effects, and on the thallus scale, gradients of light and chemical parameters influence algal photosynthesis. However, the thallus microenvironment and internal gradients remain underexplored. In this study, microsensors were used to quantify gradients of light, O2 concentration, variable chlorophyll fluorescence, photosynthesis and O2 consumption as a function of irradiance in the cortex and medulla layers of Fucus serratus. The two cortex layers showed more efficient light utilization compared to the medulla, calculated both from electron transport rates through photosystem II and from photosynthesis-irradiance curves. At moderate irradiance, the upper cortex exhibited onset of photosynthetic saturation, whereas lower thallus layers exhibited net O2 consumption. O2 consumption rates in light varied with depth and irradiance and were more than two-fold higher than dark respiration. We show that the thallus microenvironment of F. serratus exhibits a highly stratified balance of production and consumption of O2 , and when the frond was held in a fixed position, high incident irradiance levels on the upper cortex did not saturate photosynthesis in the lower thallus layers. We discuss possible photoadaptive responses and consequences for optimizing photosynthetic activity on the basis of vertical differences in light attenuation coefficients., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

A role as UV sunscreens has been suggested for phlorotannins, the phenolic compounds that accumulate in brown algae in response to a number of external stimuli and take part in cell wall structure. After exposure of the intertidal brown alga Fucus vesiculosus to artificial UV-B radiation, we examined its physiological responses by following the transcript level of the pksIII gene encoding a phloroglucinol synthase, likely to be involved in the first step of phlorotannins biosynthesis. We also monitored the expression of three targeted genes, encoding a heat shock protein (hsp70), which is involved in global stress responses, an aryl sulfotransferase (ast), which could be involved in the sulfation of phlorotannins, and a vanadium bromoperoxidase (vbpo), which can potentially participate in the scavenging of Reactive Oxygen Species (ROS) and in the cross-linking and condensation of phlorotannins. We investigated whether transcriptional regulation of these genes is correlated with an induction of phlorotannin accumulation by establishing metabolite profiling of purified fractions of low molecular weight phlorotannins. Our findings demonstrated that a high dose of UV-B radiation induced a significant overexpression of hsp70 after 12 and 24 hours following the exposure to the UV-B treatment, compared to control treatment. The physiological performance of algae quantified by the photosynthetic efficiency (Fv/Fm) was slightly reduced. However UV-B treatment did not induce the accumulation of soluble phlorotannins in F. vesiculosus during the kinetics of four weeks, a result that may be related to the lack of induction of the pksIII gene expression. Taken together these results suggest a constitutive accumulation of phlorotannins occurring during the development of F.vesiculosus, rather than inducible processes. Gene expression studies and phlorotannin profiling provide here complementary approaches to global quantifications currently used in studies of phenolic compounds in brown algae.

Macroalgae, especially perennial species, are exposed to a seasonally variable fouling pressure. It was hypothesized that macroalgae regulate their antifouling defense to fouling pressure. Over one year, the macrofouling pressure and the chemical anti-macrofouling defense strength of the brown algae Fucus vesiculosus and Fucus serratus were assessed with monthly evaluation. The anti-macrofouling defense was assessed by means of surface-extracted Fucus metabolites tested at near-natural concentrations in a novel in situ bioassay. Additionally, the mannitol content of both Fucus species was determined to assess resource availability for defense production. The surface chemistry of both Fucus species exhibited seasonal variability in attractiveness to Amphibalanus improvisus and Mytilus edulis. Of this variability, 50-60% is explained by a sinusoidal model. Only F. vesiculosus extracts originating from the spring and summer significantly deterred settlement of A. improvisus. The strength of macroalgal antifouling defense did not correlate either with in situ macrofouling pressure or with measured mannitol content, which, however, were never depleted.

In a context of reduced global biodiversity, the potential impacts from the loss of habitat-forming species (HFS) on ecosystem structure and functioning must be established. These species are often the main community primary producers and have a major role in the establishment of organisms through facilitation processes. This study focuses on macroalgae and mussels as HFS within an intertidal zone along the St. Lawrence estuary (Quebec, Canada). Over a 16-week period, we manipulated the in situ diversity profile (richness, evenness, identity, and abundance) of the dominant HFS (Fucus distichus edentatus, F. vesiculosus, and Mytilus spp.) in order to define their role in both the establishment of associated species and community primary production. Contrary to expectation, no general change in HFS richness, evenness, abundance, or identity on associated species community establishment was observed. However, over the study period, the HFS diversity profile modified the structure within the trophic guilds, which may potentially affect further community functions. Also, our results showed that the low abundance of HFS had a negative impact on the primary productivity of the community. Our results suggest that HFS diversity profiles have a limited short-term role in our study habitat and may indicate that biological forcing in these intertidal communities is less important than environmental conditions. As such, there was an opportunistic establishment of species that ensured rapid colonization regardless of the absence, or the diversity profile, of facilitators such as HFS.

Some seaweed species have been shown to release water-borne cues after herbivore attack, for example, to attract natural enemies of the herbivore. These cues may also be sensed by neighboring seaweeds and used to adjust their defenses in anticipation of a possible herbivore attack. Several studies indicated information transfer between seaweed individuals in the past, including the brown seaweed Fucus vesiculosus. Previous work showed induction of defenses in F. vesiculosus in response to water-borne cues released by isopod-grazed conspecifics. In contrast, another study on induced responses after exposure to cues from isopod-grazed neighbors using the same seaweed species yielded contradictory results. This study reassessed the ability of F. vesiculosus individuals to sense water-borne cues released by isopod-grazed neighbors in a series of experiments that monitored F. vesiculosus palatability in response to direct grazing by Idotea baltica and water-borne cues from isopod-grazed neighbors relative to unmanipulated seaweed pieces. Two-choice feeding assays were conducted with both fresh and reconstituted seaweed pieces. Direct grazing by I. baltica induced a chemical defense in F. vesiculosus, confirming results of previous studies. In contrast, evidence for increased herbivore resistance in seaweed pieces that were located downstream of isopod-grazed F. vesiculosus could not be provided. The lack of defense induction in response to grazing of conspecific neighbors may be explained by the environmental conditions and the scattered distribution of F. vesiculosus individuals in the intertidal zone of Helgoland, which may render resource investment in the emission and/or response to water-borne cues at this site unprofitable.

Climate change has led to alterations in assemblage composition. Species of temperate macroalgae at their southern limits in the Iberian Peninsula have shown shifts in geographical range and a decline in abundance ultimately related to climate, but with the proximate factors largely unknown. We performed manipulative experiments to compare physiological responses of Fucus vesiculosus and Fucus spiralis from Portugal and Wales (UK), representing, respectively, southern and central areas of their distribution, to different intensities of solar radiation and different air temperatures. Following exposure to stressful emerged conditions, Portuguese and Welsh individuals of both fucoid species showed increased frond temperature, high desiccation levels and reduced photophysiological performance that was evident even after a 16 h recovery period, with light and temperature acting in an additive, not an interactive, manner. The level of physiological decline was influenced by geographical origin of populations and species identity, with algae from the south and those living higher on the shore coping better with stressful conditions. The negative effect of summer conditions on photophysiology may contribute to changes in fucoid abundance and distribution in southern Europe. Our results emphasise how physiological performance of geographically distinct populations can differ, which is particularly relevant when predicting responses to climate change., (© 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.)

Nutrients such as nitrogen (N) and phosphorus (P) limit primary productivity, and recent anthropogenic activities are changing the availability of these nutrients, leading to alterations in the type and magnitude of nutrient limitation. Recent work highlights the potential for N and P to interact to limit primary production in terrestrial and freshwater systems. However, mechanisms underlying co-limitation are not well described. Documentation of ambient nutrient levels and tissue nutrients of the intertidal macroalga Fucus vesiculosus for 2 years in the southern Gulf of Maine, USA, indicates that N availability may be impacting the ability of F. vesiculosus to access P, despite relatively high ambient P concentrations. To experimentally validate these observations, F. vesiculosus individuals were enriched with N or P for 6 weeks, followed by an uptake experiment to examine how the interactions between these nutrients affected macroalgal N and P uptake efficiency. Results illustrate that exposure of seaweed to different nutrient regimes influenced nutrient uptake efficiency. Notably, seaweeds enriched with N displayed the highest P uptake efficiency at low, biologically relevant, P concentrations. Our results confirm that N availability may be mediating the ability of primary producers to access P. These interactions between limiting nutrients have implications for not only the growth and functioning of primary producers who rely directly on these nutrients but also the entire communities that they support.

Seaweed-dominated communities are predicted to disappear south of 45° latitude on North-Atlantic rocky shores by 2200 because of climate change. The extent of predicted habitat loss, however, could be mitigated if the seaweeds' physiology is sufficiently plastic to rapidly acclimatize to the warmer temperatures. The main objectives of this study were to identify whether the thermal tolerance of the canopy-forming seaweed Fucus serratus is population-specific and where temperatures are likely to exceed its tolerance limits in the next 200 years. We measured the stress response of seaweed samples from four populations (Norway, Denmark, Brittany and Spain) to common-garden heat stress (20 °C-36 °C) in both photosynthetic performance and transcriptomic upregulation of heat shock protein genes. The two stress indicators did not correlate and likely measured different cellular components of the stress response, but both indicators revealed population-specific differences, suggesting ecotypic differentiation. Our results confirmed that thermal extremes will regularly reach physiologically stressful levels in Brittany (France) and further south by the end of the 22nd century. Although heat stress resilience in photosynthetic performance was higher at the species' southern distributional edge in Spain, the hsp expression pattern suggested that this edge-population experienced reduced fitness and limited responsiveness to further stressors. Thus, F. serratus may be unable to mitigate its predicted northward shift and may be at high risk to lose its center of genetic diversity and adaptability in Brittany (France). As it is an important intertidal key species, the disappearance of this seaweed will likely trigger major ecological changes in the entire associated ecosystem., (Copyright © 2013 Elsevier B.V. All rights reserved.)

Background: The fucoid brown algae (Heterokontophyta, Phaeophyceae) are increasingly the focus of ecological genetics, biodiversity, biogeography and speciation research. The molecular genetics underlying mating system variation, where repeated dioecious - hermaphrodite switches during evolution are recognized, and the molecular evolution of sex-related genes are key questions currently hampered by a lack of genomic information. We therefore undertook a comparative analysis of male and female reproductive tissue transcriptomes against a vegetative background during natural reproductive cycles in Fucus vesiculosus., Results: Over 300 k reads were assembled and annotated against public protein databases including a brown alga. Compared with the vegetative tissue, photosynthetic and carbohydrate metabolism pathways were under-expressed, particularly in male tissue, while several pathways involved in genetic information processing and replication were over-expressed. Estimates of sex-biased gene (SBG) expression were higher for male (14% of annotated orthologues) than female tissue (9%) relative to the vegetative background. Mean expression levels and variance were also greater in male- than female-biased genes. Major female-biased genes were carbohydrate-modifying enzymes with likely roles in zygote cell wall biogenesis and/or modification. Male-biased genes reflected distinct sperm development and function, and orthologues for signal perception (a phototropin), transduction (several kinases), and putatively flagella-localized proteins (including candidate gamete-recognition proteins) were uniquely expressed in males. Overall, the results suggest constraint on female-biased genes (possible pleiotropy), and less constrained male-biased genes, mostly associated with sperm-specific functions., Conclusions: Our results support the growing contention that males possess a large array of genes regulating male fitness, broadly supporting findings in evolutionarily distant heterogametic animal models. This work identifies an annotated set of F. vesiculosus gene products that potentially regulate sexual reproduction and may contribute to prezygotic isolation, one essential step towards developing tools for a functional understanding of species isolation and differentiation.

Understanding the impacts of species extinctions on the functioning of food webs is a challenging task because of the complexity of ecological interactions. We report the impacts of experimental species extinctions on the functioning of two food webs of freshwater and marine systems. We used a linear model to partition the variance among the multiple components of the diversity effect (linear group richness, nonlinear group richness, and identity). The identity of each functional group was the best explaining variable of ecosystem functioning for both systems. We assessed the contribution of each functional group in multifunctional space and found that, although the effect of functional group varied across ecosystem functions, some functional groups shared common effects on functions. This study is the first experimental demonstration that functional identity dominates the effects of extinctions on ecosystem functioning, suggesting that generalizations are possible despite the inherent complexity of interactions.

The important role of marine epibiotic biofilms in the interactions of the host with its environment has been acknowledged recently. Previous studies with the temperate brown macroalga Fucus vesiculosus have identified polar and non-polar compounds recovered from the algal surface that have the potential to control such biofilms. Furthermore, both the fouling pressure and the composition of the epibiotic bacterial communities on this macroalga varied seasonally. The extent to which this reflects a seasonal fluctuation of the fouling control mechanisms of the host is, however, unexplored in an ecological context. The present study investigated seasonal variation in the anti-settlement activity of surface extracts of F. vesiculosus against eight biofilm-forming bacteria isolated from rockweed-dominated habitats, including replication of two populations from two geographically distant sites. The anti-settlement activity at both sites was found to vary temporally, reaching a peak in summer/autumn. Anti-settlement activity also showed a consistent and strong difference between sites throughout the year. This study is the first to report temporal variation of antifouling defence originating from ecologically relevant surface-associated compounds.

Understanding the effects of environmental change on the distribution and abundance of strongly interacting organisms, such as intertidal macroalgae and their grazers, needs a thorough knowledge of their underpinning ecological relationships. Control of grazer-plant interactions is bi-directional on northwestern European coasts: grazing by limpets structures populations of macroalgae, while macroalgae provide habitat and food for limpets. Scottish shores dominated by the macroalga Fucus vesiculosus support lower densities and larger sizes of limpets Patella vulgata than shores with less Fucus. These patterns may be due to differences in inter-size-class competitive interactions of limpets among shores with different covers of Fucus. To examine this model, densities of small and large limpets were manipulated in plots with and without Fucus. Amounts of biofilm were measured in each plot. The presence of Fucus increased survival but hindered growth of small (15 mm TL) limpets, which were negatively affected by the presence of large limpets (31 mm TL). In contrast, large limpets were not affected by the presence of Fucus or of small limpets. This suggests the occurrence of asymmetric inter-size-class competition, which was influenced by the presence of macroalgae. Macroalgae and increased densities of limpets did not influence amounts of biofilm. Our findings highlight the role of interactions among organisms in generating ecological responses to environmental change.

Stress regimes defined as the synchronous or sequential action of abiotic and biotic stresses determine the performance and distribution of species. The natural patterns of stress to which species are more or less well adapted have recently started to shift and alter under the influence of global change. This was the motivation to review our knowledge on the stress ecology of a benthic key player, the macroalgal genus Fucus. We first provide a comprehensive review of the genus as an ecological model including what is currently known about the major lineages of Fucus species with respect to hybridization, ecotypic differentiation and speciation; as well as life history, population structure and geographic distribution. We then review our current understanding of both extrinsic (abiotic/biotic) and intrinsic (genetic) stress(es) on Fucus species and how they interact with each other. It is concluded that (i) interactive stress effects appear to be equally distributed over additive, antagonistic and synergistic categories at the level of single experiments, but are predominantly additive when averaged over all studies in a meta-analysis of 41 experiments; (ii) juvenile and adult responses to stress frequently differ and (iii) several species or particular populations of Fucus may be relatively unaffected by climate change as a consequence of pre-adapted ecotypes that collectively express wide physiological tolerences. Future research on Fucus should (i) include additional species, (ii) include marginal populations as models for responses to environmental stress; (iii) assess a wider range of stress combinations, including their temporal fluctuations; (iv) better differentiate between stress sensitivity of juvenile versus adult stages; (v) include a functional genomic component in order to better integrate Fucus' ecological and evolutionary responses to stress regimes and (vi) utilize a multivariate modelling approach in order to develop and understand interaction networks., (2011 Elsevier Ltd. All rights reserved.)

The brown alga Fucus vesiculosus is a foundation species in the Baltic Sea littoral, hosting a rich faunal community. We compared the species composition and diversity of invertebrate macrofauna living on F. vesiculosus between sites differing in their eutrophication status and exposure to waves at three different times during a season. We determined the size, nitrogen and phlorotannin content of the alga. The invertebrate community differed substantially between sites near fish farms and those in more pristine environment. Snails and bivalves were more abundant on the Fucus stands near fish farms than on control stands, where crustaceans were more abundant. The abundance of molluscs decreased with the increasing shore exposure, while gammaridean amphipods dominated on the exposed shores. Abundance of several taxa increased during the proceeding growing season. The density of the most important herbivore of F. vesiculosus, Idotea balthica, varied 100-fold during the season being the lowest in June and the highest in August when the generation born in the summer started to feed on Fucus. Thus, the diversity and composition of Fucus-associated invertebrate fauna varies both with environmental conditions of the stand and seasonally. Although the negative effects of eutrophication on distribution and abundance of Fucus stands are well documented, a moderate increase of nutrients was found to increase the species richness of Fucus-associated fauna in early summer., (2010 Elsevier Ltd. All rights reserved.)

The last glacial maximum (20,000-18,000 years ago) dramatically affected extant distributions of virtually all northern European biota. Locations of refugia and postglacial recolonization pathways were examined in Fucus serratus (Heterokontophyta; Fucaceae) using a highly variable intergenic spacer developed from the complete mitochondrial genome of Fucus vesiculosus. Over 1,500 samples from the entire range of F. serratus were analysed using fluorescent single strand conformation polymorphism. A total of 28 mtDNA haplotypes was identified and sequenced. Three refugia were recognized based on high haplotype diversities and the presence of endemic haplotypes: southwest Ireland, the northern Brittany-Hurd Deep area of the English Channel, and the northwest Iberian Peninsula. The Irish refugium was the source for a recolonization sweep involving a single haplotype via northern Scotland and throughout Scandinavia, whereas recolonization from the Brittany-Hurd Deep refugium was more limited, probably because of unsuitable soft-bottom habitat in the Bay of Biscay and along the Belgian and Dutch coasts. The Iberian populations reflect a remnant refugium at the present-day southern boundary of the species range. A generalized skyline plot suggested exponential population expansion beginning in the mid-Pleistocene with maximal growth during the Eems interglacial 128,000-67,000 years ago, implying that the last glacial maximum mainly shaped population distributions rather than demography.

Hybridization and polyploidy are two major sources of genetic variability that can lead to adaptation in new habitats. Most species of the brown algal genus Fucus are found along wave-swept rocky shores of the Northern Hemisphere, but some species have adapted to brackish and salt marsh habitats. Using five microsatellite loci and mtDNA RFLP, we characterize two populations of morphologically similar, muscoides-like Fucus inhabiting salt marshes in Iceland and Ireland. The Icelandic genotypes were consistent with Fucus vesiculosus x Fucus spiralis F1 hybrids with asymmetrical hybridization, whereas the Irish ones consisted primarily of polyploid F. vesiculosus.

The success of introduced species is often attributed to release from co-evolved enemies in the new range and a subsequent decreased allocation to defense (EICA), but these hypotheses have rarely been evaluated for systems with low host-specificity of enemies. Here, we compare herbivore utilization of the brown seaweed, Fucus evanescens, and its coexisting competitors both in its native and new ranges, to test certain predictions derived from these hypotheses in a system dominated by generalist herbivores. While F. evanescens was shown to be a preferred host in its native range, invading populations supported a less diverse herbivore fauna and it was less preferred in laboratory choice experiments with important herbivores, when compared to co-occurring seaweeds. These results are consistent with the enemy release hypothesis, despite the fact that the herbivore communities in both regions were mainly composed of generalist species. However, in contrast to the prediction of EICA, analysis of anti-grazing compounds indicated a higher allocation to defense in introduced compared to native F. evanescens. The results suggest that the invader is subjected to less intense enemy control in the new range, but that this is due to an increased allocation to defense rather than release from specialized herbivores. This indicates that increased resistance to herbivory might be an important strategy for invasion success in systems dominated by generalist herbivores.

Canopy-forming macroalgae are key species on temperate rocky shores. However, there is a lack of understanding of how the relative balance of physical and biological factors controls the establishment and persistence of intertidal macroalgae. Here we present an integrated study of the relative importance of wave-induced forces and grazing for the recruitment and survival of the canopy-forming intertidal macroalgae Fucus vesiculosus and F. spiralis. A set of overtopped breakwaters provided a nearly unconfounded gradient in wave exposure between seaward and landward sides. A biomechanical analysis was performed based on empirical measurements of maximum drag forces in breaking waves, a model of long-term maximum wave height, and the breaking stress of Fucus spp. The estimated maximum flow speed (7-8 m/s) on the seaward side of the breakwaters was predicted to completely dislodge or prune Fucus spp. larger than approximately 10 cm, while dislodgment was highly unlikely on the landward side for all sizes. Experimental transplantation of Fucus spp. supported the biomechanical analysis but also suggested that mechanical abrasion may further limit survival in wave-exposed locations. Experimental removal of the limpet Patella vulgata, which was the principal grazer at this site, resulted in recruitment of Fucus spp. on the seaward side. We present a model of limpet grazing that indicates that limpet densities >5-20 individuals/m2 provide a proximate mechanism preventing establishment of Fucus spp., whereas wave action >2 m/s reduces persistence through dislodgment and battering. In a conceptual model we further propose that recruitment and survival of juvenile Fucus spp. are controlled indirectly by wave exposure through higher limpet densities at exposed locations. This model predicts that climate change, and in particular an increased frequency of storm events in the northeast Atlantic, will restrict fucoids to more sheltered locations.

Canopy-forming plants and algae commonly contribute to spatial variation in habitat complexity for associated organisms and thereby create a biotic patchiness of communities. In this study, we tested for interaction effects between biotic habitat complexity and resource availability on net biomass production and species diversity of understory macroalgae by factorial field manipulations of light, nutrients, and algal canopy cover in a subtidal rocky-shore community. Presence of algal canopy cover and/or artificial shadings limited net biomass production and facilitated species diversity. Artificial shadings reduced light to levels similar to those under canopy cover, and net biomass production was significantly and positively correlated to light availability. Considering the comparable and dependent experimental effects from shadings and canopy cover, the results strongly suggest that canopy cover controlled net biomass production and species diversity by limiting light and thereby limiting resource availability for community production. Canopy cover also controlled experimental nutrient effects by preventing a significant increase in net biomass production from nutrient enrichment recorded in ambient light (no shading). Changes in species diversity were mediated by changes in species dominance patterns and species evenness, where canopy cover and shadings facilitated slow-growing crust-forming species and suppressed spatial dominance by Fucus vesiculosus, which was the main contributor to net production of algal biomass. The demonstrated impacts of biotic habitat complexity on biomass production and local diversity contribute significantly to understanding the importance of functionally important species and biodiversity for ecosystem processes. In particular, this study demonstrates how loss of a dominant species and decreased habitat complexity change the response of the remaining assembly to resource loading. This is of potential significance for marine conservation since resource loading often promotes low habitat complexity and canopy species are among the first groups lost in degraded aquatic systems.

To date, molecular markers have not settled the question of the specific status of the closely related, but phylogenetically unresolved, brown seaweeds, hermaphroditic Fucus spiralis and dioecious Fucus vesiculosus, nor their propensity for natural hybridization. To test the degree of species integrity and to assess effect of the mating system on the population genetic structure, 288 individuals coming from parapatric (discontinuous) and sympatric (contiguous) spatial configurations at two sites were genotyped with five microsatellite loci. Using a Bayesian admixture analysis, our results show that F. spiralis and F. vesiculosus comprise clearly distinct genetic entities (clusters) generally characterized by cosexual and unisexual individuals, respectively. Genetic diversity within each entity suggests that F. spiralis reproduces primarily through selfing while F. vesiculosus is characterized by an endogamous breeding regime. Nevertheless, aberrant sexual phenotypes were observed in each cluster, no diagnostic alleles were revealed and 10% of study individuals were intermediate between the two genetic entities. This pattern can be explained by recent divergence of two taxa with retention of ancestral polymorphism or asymmetrical, introgressive hybridization. However, given (i) coincident monomorphism at three loci in spiralis clusters and (ii) that significantly more intermediates were observed in sympatric stations than in parapatric stations, we argue that interspecific gene flow has occurred after divergence of the two taxa. Finally, we show that whether recently separated or recently introgressive, the divergent breeding systems probably contribute to species integrity in these two taxa.

Reproduction of attached large brown algae is known to occur only by sexual zygotes. Using microsatellites we show evolution of asexual reproduction in the bladder wrack promoting population persistence in the brackish water Baltic Sea (< 6 psu). Here a dwarf morph of Fucus vesiculosus is dominated by a single clone but clonal reproduction is also present in the common form of the species. We describe a possible mechanism for vegetative reproduction of attached algae, and conclude that clonality plays an important role in persistence and dispersal of these marginal populations, in which sexual reproduction is impaired by low salinity.

Phlorotannins are ubiquitous secondary metabolites in brown algae that are phenotypically plastic and suggested to have multiple ecological roles. Traditionally, phlorotannins have been quantified as total soluble phlorotannins. Here, we modify a quantification procedure to measure, for the first time, the amount of cell-wall-bound phlorotannins. We also optimize the quantification of soluble phlorotannins. We use these methods to study the responses of soluble and cell-wall-bound phlorotannin to nutrient enrichment in growing and nongrowing parts of the brown alga Fucus vesiculosus. We also examine the effects of nutrient shortage and herbivory on the rate of phlorotannin exudation. Concentrations of cell-wall-bound phlorotannins were much lower than concentrations of soluble phlorotannins; we also found that nutrient treatment over a period of 41 days affected only soluble phlorotannins. Concentrations of each phlorotannin type correlated positively between growing and nongrowing parts of individual seaweeds. However, within nongrowing thalli, soluble and cell-wall-bound phlorotannins were negatively correlated, whereas within growing thalli there was no correlation. Phlorotannins were exuded from the thallus in all treatments. Herbivory increased exudation, while a lack of nutrients had no effect on exudation. Because the amount of cell-wall-bound phlorotannins is much smaller than the amount of soluble phlorotannins, the major function of phlorotannins appears to be a secondary one.

Generation and expression of cell polarity in brown algal zygotes of the Fucales involve regulation of the actin cytoskeleton and localized secretion. We used degenerate PCR to isolate cDNAs that encode two small GTPases, FdRac1 and FdRab8, from zygotes of Fucus distichus (L.) Powell. Sequence analysis placed FdRac1 in the Rho family, which regulates actin, and FdRab8 in the Rab family, which regulates vesicle transport. As expected, bacterially expressed forms of both proteins bound GTP in vitro. When expressed in budding yeast, FdRac1 showed some functional overlap with CDC42, the Saccharomyces cerevisiae Rho family gene required for yeast cell polarity. Immunolocalization revealed an asymmetric distribution of FdRac1 in polarized zygotes and embryos, with FdRac1 concentrated at or near the growing tip of the algal rhizoid. Our data support the hypothesis that FdRac1 regulates algal cell polarity, possibly via the actin cytoskeleton. Because brown algae belong to the heterokont group, which diverged from other groups early in eukaryotic evolution, we argue that the Rho family function of regulating cell polarity is ancient and may extend throughout the eukaryotes.

Stationary and nonstationary spatiotemporal pattern formations emerging from the cellular electric activity are a common feature of biological cells and tissues. The nonstationary ones are well explained in the framework of the cable model. Inversely, the formation of the widespread self-organized stationary patterns of transcellular ionic currents remains elusive, despite their importance in cell polarization, apical growth, and morphogenesis. For example, the nature of the breaking symmetry in the Fucus zygote, a model organism for the experimental investigation of embryonic pattern formation, is still an open question. Using an electrodiffusive model, we report here an unexpected property of the cellular electric activity: a phase-space domain that gives rise to stationary patterns of transcellular ionic currents at finite wavelength. The cable model cannot predict this instability. In agreement with experiments, the characteristic time is an ionic diffusive one (<2 min). The critical radius is of the same order of magnitude as the cell radius (30 microm). The generic salient features are a global positive differential conductance, a negative differential conductance for one ion, and a difference between the diffusive coefficients. Although different, this mechanism is reminiscent of Turing instability.

Zygotes of the brown alga Fucus distichus (L.) Powell develop polarity prior to the first embryonic cell division and retain a pattern of asymmetric growth during early embryogenesis. In order to identify F. distichus polypeptides secreted during asymmetric cell growth, we used a functional assay in Saccharomyces cerevisiae to screen a cDNA library generated from asymmetrically growing Fucus embryos for sequences encoding polypeptides that function as signal peptides for secretion. We isolated and sequenced 222 plasmids containing Fucus cDNAs encoding signal peptide activity. The cDNA inserts from these plasmids were translated in silico into 244 potential polypeptide sequences, 169 of which are predicted to contain signal peptides. BlastP analysis of the Fucus sequences revealed similarity between many Fucus proteins and cell surface proteins that function in development in other eukaryotes, including epidermal growth factor (EGF)-like repeat-containing proteins, plant leucine-rich repeat (LRR)-receptor kinases, and algal beta-1, 3-exoglucanase. However, most of the isolated Fucus polypeptides lack similarity to known proteins. The isolation of cDNAs encoding secreted Fucus proteins provides an important step toward characterizing cell surface proteins important for asymmetric organization and growth in fucoid embryos.

The seaweed Fucus serratus is hypothesized to have evolved in the North Atlantic and present populations are thought to reflect recolonization from a southern refugium since the last glacial maximum 18 000-20 000 years bp. We examined genetic structure across several spatial scales by analysing seven microsatellite loci in populations collected from 21 localities throughout the species' range. Spatial auto-correlation analysis of seven microsatellite loci revealed no evidence for spatial clustering of alleles on a scale of 100 m despite limited gamete dispersal in F. serratus of approximately 2 m from parental individuals. Pairwise theta analysis suggested that the minimal panmictic unit for F. serratus was between 0.5 and 2 km. Isolation by distance was significant along some contiguous coastlines. Population differentiation was strong within the Skagerrak-Kattegat-Baltic Seas (SKB) (global theta= 0.17) despite a short history of approximately 7500 years. A neighbour-joining tree based on Reynold's distances computed from the microsatellite data revealed a central assemblage of populations on the Brittany Peninsula surrounded by four well-supported clusters consisting of the SKB, the North Sea (Ireland, Helgoland), and two populations from the northern Spanish coast. Samples from Iceland and Nova Scotia were most closely aligned with northwest Sweden and Brittany, respectively. When sample sizes were standardized (N = 41), allelic diversity was twofold higher for Brittany populations than for populations to the north and threefold higher than southern populations. The Brittany region may be a refugium or a recolonized area, whereas the Spanish populations most likely reflect present-day edge populations that have undergone repeated bottlenecks as a consequence of thermally induced cycles of recolonization and extinction.