Positive species interactions structure rhodolith bed communities at a global scale.

Rhodolith beds are diverse and globally distributed habitats. Nonetheless, the role of rhodoliths in structuring the associated species community through a hierarchy of positive interactions is yet to be recognised. In this review, we provide evidence that rhodoliths can function as foundation species of multi-level facilitation cascades and, hence, are fundamental for the persistence of hierarchically structured communities within coastal oceans. Rhodoliths generate facilitation cascades by buffering physical stress, reducing consumer pressure and enhancing resource availability. Due to large variations in their shape, size and density, a single rhodolith bed can support multiple taxonomically distant and architecturally distinct habitat-forming species, such as primary producers, sponges or bivalves, thus encompassing a broad range of functional traits and providing a wealth of secondary microhabitat and food resources. In addition, rhodoliths are often mobile, and thus can redistribute associated species, potentially expanding the distribution of species with short-distance dispersal abilities. Key knowledge gaps we have identified include: the experimental assessment of the role of rhodoliths as basal facilitators; the length and temporal stability of facilitation cascades; variations in species interactions within cascades across environmental gradients; and the role of rhodolith beds as climate refugia. Addressing these research priorities will allow the development of evidence-based policy decisions and elevate rhodolith beds within marine conservation strategies.

Effects of ocean warming and pollution on Sargassum forests.

The combined effects of climate change and ocean pollution have resulted in a noteworthy decline of canopy-forming species, impacting marine biodiversity and ecosystem functioning significantly. In this context, Sargassum cymosum, which is widely distributed along the southwestern Atlantic Ocean, serves as an excellent model among canopy-forming species to investigate these impacts on populations in different regions and environmental conditions. Here, we evaluate the ecophysiological responses of two populations of S. cymosum, from Florianopolis (warm-temperate province; WTP) and Fernando de Noronha (tropical province, TP), through of interaction of temperatures and nutrient concentrations, representing marine heatwaves and acute pollution levels. Our findings revealed a decrease in biomass in both populations, highlighting the significance of nutrient enrichment as an anthropogenic filter that might potentially inhibit the expansion of the populations from tropical regions and temperature for WTP ones. These stressors directly impacted the physiological performance of S. cymosum populations, including relative growth rates, photosynthesis, chlorophylls, carotenoids and phenolic compound levels. Although there was an increase in both parameters for the TP population, a significant loss of biomass was observed, with growth rates reaching -1.5% per day. Our results highlight the need for urgent actions to manage the eutrophication process due to its negative association with global warming, which can enhance the impacts and preclude the settlement and survival of Sargassum in warm-temperate areas considering the observed and predicted tropicalization process.

Levelling-up rhodolith-bed science to address global-scale conservation challenges.

Global marine conservation remains fractured by an imbalance in research efforts and policy actions, limiting progression towards sustainability. Rhodolith beds represent a prime example, as they have ecological importance on a global scale, provide a wealth of ecosystem functions and services, including biodiversity provision and potential climate change mitigation, but remain disproportionately understudied, compared to other coastal ecosystems (tropical coral reefs, kelp forests, mangroves, seagrasses). Although rhodolith beds have gained some recognition, as important and sensitive habitats at national/regional levels during the last decade, there is still a notable lack of information and, consequently, specific conservation efforts. We argue that the lack of information about these habitats, and the significant ecosystem services they provide, is hindering the development of effective conservation measures and limiting wider marine conservation success. This is becoming a pressing issue, considering the multiple severe pressures and threats these habitats are exposed to (e.g., pollution, fishing activities, climate change), which may lead to an erosion of their ecological function and ecosystem services. By synthesizing the current knowledge, we provide arguments to highlight the importance and urgency of levelling-up research efforts focused on rhodolith beds, combating rhodolith bed degradation and avoiding the loss of associated biodiversity, thus ensuring the sustainability of future conservation programs.

Beta-1,3-glucanase inhibitors in Brazilian brown seaweed.

Beta-1,3-glucanases are enzymes that hydrolyze beta-1,3-glucans, and they are essential for the metabolism of seaweed, plants and fungi. These enzymes also participate in the digestion of herbivore and fungivore animals. Because of the importance of these enzymes in insects, beta-1,3-glucanase inhibitors may be used for the development of new control strategies against agricultural pests and disease vectors. Beta-1,3-glucanase inhibitors have been described in the brown seaweed Laminaria cichorioides, but were never recorded in Brazilian seaweed species. We evaluated the presence of beta-1,3-glucanase inhibitors in samples of Padina gymnospora, Dictyota sp., Colpomenia sinuosa, and Lobophora sp., collected in Arraial d'Ajuda (Bahia). Ethanolic or buffer extracts were used in inhibition tests against the beta-1,3-glucanase of Trichoderma sp. Extracts in buffer showed no inhibition, but ethanolic extracts from all species showed different extents of inhibition. Samples from Dictyota sp. and P. gymnospora showed inhibitions above 75% (absolute ethanol) or 50% (ethanol 50%). In summary, extraction with absolute ethanol resulted in better inhibitions, and P. gymnospora showed the higher inhibitions. Brazilian seaweed may be good sources of beta-1,3-glucanase inhibitors for biochemical and physiological studies of these enzymes. Besides that, these molecules show potential for the development of new biotechnological tools for insect control.

Calcification in free-living coralline algae is strongly influenced by morphology: Implications for susceptibility to ocean acidification.

Rhodolith beds built by free-living coralline algae are important ecosystems for marine biodiversity and carbonate production. Yet, our mechanistic understanding regarding rhodolith physiology and its drivers is still limited. Using three rhodolith species with different branching morphologies, we investigated the role of morphology in species' physiology and the implications for their susceptibility to ocean acidification (OA). For this, we determined the effects of thallus topography on diffusive boundary layer (DBL) thickness, the associated microscale oxygen and pH dynamics and their relationship with species' metabolic and light and dark calcification rates, as well as species' responses to short-term OA exposure. Our results show that rhodolith branching creates low-flow microenvironments that exhibit increasing DBL thickness with increasing branch length. This, together with species' metabolic rates, determined the light-dependent pH dynamics at the algal surface, which in turn dictated species' calcification rates. While these differences did not translate in species-specific responses to short-term OA exposure, the differences in the magnitude of diurnal pH fluctuations (~ 0.1-1.2 pH units) between species suggest potential differences in phenotypic plasticity to OA that may result in different susceptibilities to long-term OA exposure, supporting the general view that species' ecomechanical characteristics must be considered for predicting OA responses.

Golden carbon of Sargassum forests revealed as an opportunity for climate change mitigation.

Marine climate change mitigation initiatives have recently attracted a great deal of interest in the role of natural carbon sinks, particularly on coastal systems. Brown seaweeds of the genus Sargassum are the largest canopy-forming algae in tropical and subtropical environments, with a wide global distribution on rocky reefs and as floating stands. Because these algae present high amounts of biomass, we suggest their contribution is relevant for global carbon stocks and consequently for mitigating climate change as CO2 remover. We modelled global distributions and quantified carbon stocks as above-ground biomass (AGB) with machine learning algorithms and climate data. Sargassum AGB totaled 13.1 Pg C at the global scale, which is a significant amount of carbon, comparable to other key marine ecosystems, such as mangrove forests, salt marshes and seagrass meadows. However, specific techniques related to bloom production and management, or the utilization of biomass for biomaterials, should be fostered.

Environmental drivers of rhodolith beds and epiphytes community along the South Western Atlantic coast.

Environmental conditions shape the occurrence and abundance of habitat-building organisms at global scales. Rhodolith beds structure important hard substrate habitats for a large number of marine benthic organisms. These organisms can benefit local biodiversity levels, but also compete with rhodoliths for essential resources. Therefore, understanding the factors shaping the distribution of rhodoliths and their associated communities along entire distributional ranges is of much relevance for conservational biology, particularly in the scope of future environmental changes. Here we predict suitable habitat areas and identify the main environmental drivers of rhodoliths' variability and of associated epiphytes along a large-scale latitudinal gradient. Occurrence and abundance data were collected throughout the South-western Atlantic coast (SWA) and modelled against high resolution environmental predictors extracted from Bio-Oracle. The main drivers for rhodolith occurrence were light availability and temperature at the bottom of the ocean, while abundance was explained by nitrate, temperature and current velocity. Tropical regions showed the highest abundance of rhodoliths. No latitudinal pattern was detected in the variability of epiphytes abundance. However, significant differences were found between sampled sites regarding the composition of predominant taxa. The predictors influencing such differences were temperature and nitrate. The Tropical region is abundant in species with warm-water affinities, decreasing toward warm temperate region. The expressive occurrence of tropical species not referred before for warm temperate beds indicate a plausible tropicalization event.

Physiological damages of Sargassum cymosum and Hypnea pseudomusciformis exposed to trace metals from mining tailing.

The damages of Mariana's mining mud in the physiology of the brown algae Sargassum cymosum and its main epiphytic, the red algae Hypnea pseudomusciformis, were evaluated by controlled essays. Seaweeds were exposed to presence or absence of mud, isolated or in biological association, for 5 and 15 days. Measured parameters were growth rates, biochemical descriptors, and the chemical investigation of concentration and metal profile of the mud dissolved in seawater. Results showed that the highest values for metals were Al > Fe > Mn > Zn in both exposure periods. The mud also affected the growth rate with lethality in both isolated and associative treatments with H. pseudomusciformis after 15 days. According to our redundancy analysis (RDA), the profile and concentration of all metallic elements can induce different physiological responses of the organisms. We were able to observe a higher physiological adaptive ability of S. cymosum against the long-term presence of metals by the synthesis of phenolic compounds, while the deviation of metabolic routes in H. pseudomusciformis can be addressed as the main responsible for its lethality. Moreover, the presence of Hypnea in associative treatments reduces Sargassum's detoxification ability. The present results reinforce the importance of biological interaction studies in a context of physiological resilience against mining mud pollution and mutual influences of species over the individual ability to avoid oxidative stress.

Physiological and biochemical responses of a coralline alga and a sea urchin to climate change: Implications for herbivory.

Direct responses to rising temperatures and ocean acidification are increasingly well known for many single species, yet recent reviews have highlighted the need for climate change research to consider a broader range of species, how stressors may interact, and how stressors may affect species interactions. The latter point is important in the context of plant-herbivore interactions, as increasing evidence shows that increasing seawater temperature and/or acidification can alter algal traits that dictate their susceptibility to herbivores, and subsequently, community and ecosystem properties. To better understand how marine rocky shore environments will be affected by a changing ocean, in the present study we investigated the direct effects of short-term, co-occurring increased temperature and ocean acidification on a coralline alga (Jania rubens) and a sea urchin herbivore (Echinometra lucunter) and assessed the indirect effects of these factors on the algal-herbivore interaction. A 21-day mesocosm experiment was conducted with both algae and sea urchins exposed to ambient (24 °C, Low CO2), high-temperature (28 °C, Low CO2), acidified (24 °C, High CO2), or high-temperature plus acidified (28 °C, High CO2) conditions. Algal photosynthesis, respiration, and phenolic content were unaffected by increased temperature and CO2, but calcium carbonate content was reduced under high CO2 treatments in both temperatures, while total sugar content of the algae was reduced under acidified, lower temperature conditions. Metabolic rates of the sea urchin were elevated in the lower temperature, high CO2 treatment, and feeding assays showed that consumption rates also increased in this treatment. Despite some changes to algal chemical composition, it appears that at least under short-term exposure to climate change conditions, direct effects on herbivore metabolism dictated herbivory rates, while indirect effects caused by changes in algal palatability seemed to be of minor importance.

Photoacclimation strategies in northeastern Atlantic seagrasses: Integrating responses across plant organizational levels.

Seagrasses live in highly variable light environments and adjust to these variations by expressing acclimatory responses at different plant organizational levels (meadow, shoot, leaf and chloroplast level). Yet, comparative studies, to identify species' strategies, and integration of the relative importance of photoacclimatory adjustments at different levels are still missing. The variation in photoacclimatory responses at the chloroplast and leaf level were studied along individual leaves of Cymodocea nodosa, Zostera marina and Z. noltei, including measurements of variable chlorophyll fluorescence, photosynthesis, photoprotective capacities, non-photochemical quenching and D1-protein repair, and assessments of variation in leaf anatomy and chloroplast distribution. Our results show that the slower-growing C. nodosa expressed rather limited physiological and biochemical adjustments in response to light availability, while both species of faster-growing Zostera showed high variability along the leaves. In contrast, the inverse pattern was found for leaf anatomical adjustments in response to light availability, which were more pronounced in C. nodosa. This integrative plant organizational level approach shows that seagrasses differ in their photoacclimatory strategies and that these are linked to the species' life history strategies, information that will be critical for predicting the responses of seagrasses to disturbances and to accordingly develop adequate management strategies.

The influence of environmental features in the content of mycosporine-like amino acids in red marine algae along the Brazilian coast.

Mycosporine-like amino acids (MAA) are ultraviolet screen substances synthesized by marine algae. The physiological function of these substances is related to cellular protection against UV radiation and as a protective mechanism against oxidative stress. These substances can be found mainly in the ocean, among red seaweeds. Its concentration in organisms has been related to ultraviolet radiation and availability of inorganic nitrogen in the environment. We start our study of MAA content in different species to understand if environmental conditions influence the concentration of MAAs in red seaweeds. The Brazilian coast presents abiotic factors that interact to create different physical-chemical features in the environment. We collected 441 samples from 39 species of red seaweed easily found in the intertidal zone, in low tide, during the summer of 2015. The sampling encompassed a latitudinal gradient (3° S to 28°5' S) at 23 points along the coast. We quantified and identified the content of MAAs in species through the method of high performance liquid chromatography. We detected for the first time the occurrence of MAAs in certain species of red algae that have not been reported to contain MAAs before. We confirmed that some environmental factors influenced the content of MAAs. Enhanced MAA contents, for example, were found in environments with a basic pH, a high ultraviolet index, and high concentrations of phosphate and nitrate. Salinity, dissolved oxygen and variations of sea surface temperature also influenced, in a secondary way, MAA content in algae in their natural environments.

The brown seaweed Sargassum cymosum: changes in metabolism and cellular organization after long-term exposure to cadmium.

Sargassum cymosum was exposed to cadmium (Cd) to determine any physiological and ultrastructural effects. To accomplish this, S. cymosum samples were cultivated under photosynthetic active radiation (PAR) and Cd (0, 0.1, 0.2, 0.4 and 0.8 mg L-1) during 7 and 14 days in laboratory-controlled conditions (0 mg L-1 Cd at both exposure times as control). Seaweeds had high retention capacity (over 90 %) for both exposure times. Growth rates showed significant increases by 14 days, especially for 0.1 and 0.4 mg L-1 Cd. Photosynthetic parameters were unaffected by Cd treatments. Chlorophyll contents were present in higher concentrations for all Cd treatments compared to respective control. Carotenoid profile showed significant differences in total composition and proportion of fucoxanthin and ß-carotene, and no lutein was detected at 14 days. Phenolic and flavonoid compounds showed major accumulation at 14 days. Transmission electron microscopy (TEM) analyses presented major alterations in Cd-treated samples, when compared with respective control, in particular disorganization of cell wall fibrils. When compared to respective control samples, multivariate analyses showed disparate and complex interactions among metabolites in Cd-exposed seaweeds, giving evidence of physiological defence response. Thus, it can be concluded that Cd is a stressor for S. cymosum, resulting in physiological and structural alterations related to defence mechanisms against oxidative stress and toxicological effects resulting from long-term metal exposure. However, in the present paper, some observed changes also appear to result from acclimation mechanisms under lower concentration of Cd relative to the tolerance of S. cymosum to experimental conditions.

The Western South Atlantic Ocean in a High-CO2 World: Current Measurement Capabilities and Perspectives.

An international multi-disciplinary group of 24 researchers met to discuss ocean acidification (OA) during the Brazilian OA Network/Surface Ocean-Lower Atmosphere Study (BrOA/SOLAS) Workshop. Fifteen members of the BrOA Network (www.broa.furg.br) authored this review. The group concluded that identifying and evaluating the regional effects of OA is impossible without understanding the natural variability of seawater carbonate systems in marine ecosystems through a series of long-term observations. Here, we show that the western South Atlantic Ocean (WSAO) lacks appropriate observations for determining regional OA effects, including the effects of OA on key sensitive Brazilian ecosystems in this area. The impacts of OA likely affect marine life in coastal and oceanic ecosystems, with further social and economic consequences for Brazil and neighboring countries. Thus, we present (i) the diversity of coastal and open ocean ecosystems in the WSAO and emphasize their roles in the marine carbon cycle and biodiversity and their vulnerabilities to OA effects; (ii) ongoing observational, experimental, and modeling efforts that investigate OA in the WSAO; and (iii) highlights of the knowledge gaps, infrastructure deficiencies, and OA-related issues in the WSAO. Finally, this review outlines long-term actions that should be taken to manage marine ecosystems in this vast and unexplored ocean region.

Effects of eutrophic seawater and temperature on the physiology and morphology of Hypnea musciformis J. V. Lamouroux (Gigartinales, Rhodophyta).

As both food and source of a kappa-carrageenan, Hypnea musciformis represents a species of great economic interest. It also synthesizes substances with antiviral, anti-helminthic and anti-inflammatory potential and shows promise for use as a bioindicator of cadmium. In this study, we investigated the combined effects of seawater from three urbanized areas (area 1: natural runoff, NRA; area 2: urbanized runoff and sewage with treatment, RTA; area 3: urbanized runoff and untreated sewage, RUS) and three different temperatures (15, 25 and 30 °C) on the growth rate, photosynthetic efficiency, photosynthetic pigments and cell morphology of H. musciformis. After 4 days (96 h) of culture, the biomass of H. musciformis showed differences that fluctuated among the areas and temperature treatments. Specifically, the specimens cultivated in 35 °C had low values of ETRmax, α(ETR), ß(ETR), and Fv/Fm photosynthetic parameters, as well as changes in cell morphology, with reduction in photosynthetic pigments and drastic reduction in growth rates. When combined with the extreme temperatures, high concentrations of ammonium ion in seawater effluent caused an inhibition of photosynthetic activity, as well as significant variation in chlorophyll a and carotenoid contents. As observed by light microscopy, the synergism between different temperatures and pollutants found in eutrophic waters caused changes in cellular morphology with increased cell wall thickening and decreased floridean starch grains. H. musciformis also showed important changes in physiological response to each factor independently, as well as changes resulting from the synergistic interaction of these factors combined. Therefore, we can conclude that extreme temperature combined with the effect of eutrophic waters, especially RUS, caused distinct morphological and physiological changes in the red alga H. musciformis.

Anatomical and ultrastructural adaptations of seagrass leaves: an evaluation of the southern Atlantic groups.

Seagrasses, which form an integral part of the worldwide coastal habitat, are considered highly relevant from an ecological point of view. Due to the scarcity of anatomical information, the present study analyzed the morphoanatomy, histochemistry, and ultrastructure of Halophila decipiens, Halodule wrightii, and Ruppia maritima leaves, discussing their adaptations to the marine environments observed throughout the southwestern tropical and subtropical Atlantic coast. The leaves of these three species feature a uniseriate epidermis with the presence of chloroplasts in large quantities and absence of stomata. The vascular system consists of a central vascular bundle with sieve tube elements of the phloem and protoxylem lacunae, as well as small vascular bundles near the leaf margins. The leaves of H. decipiens possess trichomes, but no mesophyll in the leaf margins. The mesophyll of H. wrightii and R. maritima is homogeneous with chlorenchyma cells and air lacunae scattered throughout the leaf. The histochemistry analysis revealed the absence of amyloplasts and the presence of proteins in the outer periclinal walls of ordinary epidermal cells of the three species. It was also possible to detect the presence of idioblasts containing phenolic compounds in H. decipiens and R. maritima. The ultrastructural analysis of the three species revealed many elliptical chloroplasts, with organized thylakoids, expansion of the epidermal cell wall into the cytoplasm, and a thin cuticle. Hydropoten were also observed in the three specimens. The results show that the species analyzed have important adaptations which enable their survival in the marine environment.

The effect of cadmium under different salinity conditions on the cellular architecture and metabolism in the red alga Pterocladiella capillacea (Rhodophyta, Gelidiales).

The in vitro effect of cadmium (Cd) on apical segments of Pterocladiella capillacea was examined. Over a period of 7 days, the segments were cultivated with the combination of different salinities (25, 35, and 45 practical salinity units) and Cd concentrations, ranging from 0.17 to 0.70 ppm. The effects of Cd on growth rates and content of photosynthetic pigments were analyzed. In addition, metabolic profiling was performed, and samples were processed for microscopy. Serious damage to physiological performance and ultrastructure was observed under different combinations of Cd concentrations and salinity values. Elementary infrared spectroscopy revealed toxic effects registered on growth rate, photosynthetic pigments, chloroplast, and mitochondria organization, as well as changes in lipids and carbohydrates. These alterations in physiology and ultrastructure were, however, coupled to activation of such defense mechanisms as cell wall thickness, reduction of photosynthetic harvesting complex, and flavonoid. In conclusion, P. capillacea is especially sensitive to Cd stress when intermediate concentrations of this pollutant are associated with low salinity values. Such conditions resulted in metabolic compromise, reduction of primary productivity, i.e., photosynthesis, and carbohydrate accumulation in the form of starch granules. Taken together, these findings improve our understanding of the potential impact of this metal in the natural environment.

Evaluation of impacts of climate change and local stressors on the biotechnological potential of marine macroalgae: a brief theoretical discussion of likely scenarios

Climate change can be associated with variations in the frequency and intensity of extreme temperatures and precipitation events on the local and regional scales. Along coastal areas, flooding associated with increased occupation has seriously impacted products and services generated by marine life, in particular the biotechnological potential that macroalgae hold. Therefore, this paper analyzes the available information on the taxonomy, ecology and physiology of macroalgae and discusses the impacts of climate change and local stress on the biotechnological potential of Brazilian macroalgae. Based on data compiled from a series of floristic and ecological works, we note the disappearance in some Brazilian regions of major groups of biotechnological interest. In some cases, the introduction of exotic species has been documented, as well as expansion of the distribution range of economically important species. We also verify an increase in the similarities between the Brazilian phycogeographic provinces, although they still remain different. It is possible that these changes have resulted from the warming of South Atlantic water, as observed for its surface in southeastern Brazilian, mainly during the winter. However, unplanned urbanization of coastal areas can also produce similar biodiversity losses, which requires efforts to generate long-term temporal data on the composition, community structure and physiology of macroalgae.

The impact of coastal urbanization on the structure of phytobenthic communities in southern Brazil.

The anthropogenic pressures on coastal areas represent important factors affecting local, regional, and even global patterns of distribution and abundance of benthic organisms. This report undertakes a comparative analysis of the community structure of rocky shore intertidal phytobenthos in both pristine like environments (PLE) and urbanized environments (UBE) in southern Brazil, characterizing variations on different spatial scales. Multivariate analysis of variance indicated that the PLE is characterized by a larger number of taxa and an increased occurrence of Rhodophyta species in relation to UBE. In contrast, UBE were dominated by opportunistic algae, such as Cladophora and Ulva spp. Significance tests further indicated higher species richness and Shannon-Wiener diversity on the PLE in relation to UBE. Here we provide data showing the magnitude of seaweed biodiversity loss and discuss direct and indirect consequences of unplanned urbanization on these communities.