Distinguishing the molecular diversity, nutrient content, and energetic potential of exometabolomes produced by macroalgae and reef-building corals.

Metabolites exuded by primary producers comprise a significant fraction of marine dissolved organic matter, a poorly characterized, heterogenous mixture that dictates microbial metabolism and biogeochemical cycling. We present a foundational untargeted molecular analysis of exudates released by coral reef primary producers using liquid chromatography-tandem mass spectrometry to examine compounds produced by two coral species and three types of algae (macroalgae, turfing microalgae, and crustose coralline algae [CCA]) from Mo'orea, French Polynesia. Of 10,568 distinct ion features recovered from reef and mesocosm waters, 1,667 were exuded by producers; the majority (86%) were organism specific, reflecting a clear divide between coral and algal exometabolomes. These data allowed us to examine two tenets of coral reef ecology at the molecular level. First, stoichiometric analyses show a significantly reduced nominal carbon oxidation state of algal exometabolites than coral exometabolites, illustrating one ecological mechanism by which algal phase shifts engender fundamental changes in the biogeochemistry of reef biomes. Second, coral and algal exometabolomes were differentially enriched in organic macronutrients, revealing a mechanism for reef nutrient-recycling. Coral exometabolomes were enriched in diverse sources of nitrogen and phosphorus, including tyrosine derivatives, oleoyl-taurines, and acyl carnitines. Exometabolites of CCA and turf algae were significantly enriched in nitrogen with distinct signals from polyketide macrolactams and alkaloids, respectively. Macroalgal exometabolomes were dominated by nonnitrogenous compounds, including diverse prenol lipids and steroids. This study provides molecular-level insights into biogeochemical cycling on coral reefs and illustrates how changing benthic cover on reefs influences reef water chemistry with implications for microbial metabolism.

On the Health Benefits vs. Risks of Seaweeds and Their Constituents: The Curious Case of the Polymer Paradigm.

To exploit the nutraceutical and biomedical potential of selected seaweed-derived polymers in an economically viable way, it is necessary to analyze and understand their quality and yield fluctuations throughout the seasons. In this study, the seasonal polysaccharide yield and respective quality were evaluated in three selected seaweeds, namely the agarophyte Gracilaria gracilis, the carrageenophyte Calliblepharis jubata (both red seaweeds) and the alginophyte Sargassum muticum (brown seaweed). It was found that the agar synthesis of G. gracilis did not significantly differ with the seasons (27.04% seaweed dry weight (DW)). In contrast, the carrageenan content in C. jubata varied seasonally, being synthesized in higher concentrations during the summer (18.73% DW). Meanwhile, the alginate synthesis of S. muticum exhibited a higher concentration (36.88% DW) during the winter. Therefore, there is a need to assess the threshold at which seaweed-derived polymers may have positive effects or negative impacts on human nutrition. Furthermore, this study highlights the three polymers, along with their known thresholds, at which they can have positive and/or negative health impacts. Such knowledge is key to recognizing the paradigm governing their successful deployment and related beneficial applications in humans.

Socioeconomic prospects of a seaweed bioeconomy in Sweden.

Seaweed cultivation is a large industry worldwide, but production in Europe is small compared to production in Asian countries. In the EU, the motivations for seaweed farming may be seen from two perspectives; one being economic growth through biomass production and the other being the provisioning of ecosystem services such as mitigating eutrophication. In this paper, we assess the economic potential of large-scale cultivation of kelp, Saccharina latissima, along the Swedish west coast, including the value of externalities. The findings suggest that seaweed farming has the potential of becoming a profitable industry in Sweden. Furthermore, large-scale seaweed farming can sequester a significant share of annual anthropogenic nitrogen and phosphorus inflows to the basins of the Swedish west coast (8% of N and 60% of P). Concerning the valuation of externalities, positive values generated from sequestration of nitrogen and phosphorus are potentially counteracted by negative values from interference with recreational values. Despite the large N and P uptake, the socioeconomic value of this sequestration is only a minor share of the potential financial value from biomass production. This suggests that e.g. payment schemes for nutrient uptake based on the socioeconomic values generated is not likely to be a tipping point for the industry. Additionally, seaweed cultivation is not a cost-efficient measure in itself to remove nutrients. Policy should thus be oriented towards industry development, as the market potential of the biomass will be the driver that may unlock these bioremediation opportunities.

Life Cycle Assessment of Seaweed Cultivation Systems.

Life cycle assessment (LCA) is a holistic methodology that identifies the impacts of a production system on the environment. The results of an LCA are used to identify which processes can be improved to minimize impacts and optimize production.LCA is composed of four phases: (1) goal and scope definition, (2) life cycle inventory analysis, (3) life cycle impact assessment, and (4) interpretation.The goal and scope define the purpose of the analysis; describe the system and its function, establish a functional unit to collect data and present results, set the system boundaries, and explain the assumptions made and data quality requirements. Life cycle inventory analysis is the collection, processing and organization of data. Life cycle impact assessment associates the results from the inventory phase to one or multiple impacts on environment or human health. The interpretation evaluates the outcome of each phase of the analysis. In this phase the practitioner decides whether it is necessary to amend other phases, e.g., collection of more data or adjustments of goal of the analysis. In the interpretation, the practitioner draws conclusions, exposes the limitations, and provides recommendations to the readers.The quality of LCA of seaweed production and conversion is based on data availability and detail level. Performing an LCA at the initial stage of seaweed production in Europe is an advantage: the recommended design improvements can be implemented without significant economic investments. The quality of LCA will keep improving with the increase of scientific publications, data sharing, and public reports.

Nutritional and bioactive compounds of commercialized algae powders used as food supplements.

The main nutritional/bioactive compounds (protein; aminoacids, AA; fucose; minerals; vitamins B12 and C; and total phenolic content, TPC) of nine commercial algae powders, used as food supplements, were studied. Undaria pinnatifida showed the highest protein/aminoacid contents (51.6/54.4 g 100 g-1). Among brown macroalgae, Himanthalia elongata showed the highest fucose content (26.3 g kg-1) followed by Laminaria ochroleuca (22.5 g kg-1). Mineral contents of 15-24% were observed in the algae, being particularly excellent sources of iodine (69.0-472.0 mg kg-1). Porphyra spp. and Palmaria palmata showed the highest vitamin B12 contents (667-674 µg kg-1). Vitamin C ranged among 490.4-711.8 mg kg-1. H. elongata showed the highest total phenolic content (14.0 g kg-1). In conclusion, the studied algae are excellent sources of protein, AA, minerals, vitamin C and some of them presented particularly high vitamin B12 and fucose contents, which may have a potential use as food supplements.

Nutrient removal from Chinese coastal waters by large-scale seaweed aquaculture.

China is facing intense coastal eutrophication. Large-scale seaweed aquaculture in China is popular, now accounting for over 2/3's of global production. Here, we estimate the nutrient removal capability of large-scale Chinese seaweed farms to determine its significance in mitigating eutrophication. We combined estimates of yield and nutrient concentration of Chinese seaweed aquaculture to quantify that one hectare of seaweed aquaculture removes the equivalent nutrient inputs entering 17.8 ha for nitrogen and 126.7 ha for phosphorus of Chinese coastal waters, respectively. Chinese seaweed aquaculture annually removes approximately 75,000 t nitrogen and 9,500 t phosphorus. Whereas removal of the total N inputs to Chinese coastal waters requires a seaweed farming area 17 times larger than the extant area, one and a half times more of the seaweed area would be able to remove close to 100% of the P inputs. With the current growth rate of seaweed aquaculture, we project this industry will remove 100% of the current phosphorus inputs to Chinese coastal waters by 2026. Hence, seaweed aquaculture already plays a hitherto unrealized role in mitigating coastal eutrophication, a role that may be greatly expanded with future growth of seaweed aquaculture.

Relative importance of P and N in macrophyte and epilithic algae biomass in a wastewater-impacted oligotrophic river.

The role of nutrient loading on biomass growth in wastewater-impacted rivers is important in order to effectively optimize wastewater treatment to avoid excessive biomass growth in the receiving water body. This paper directly relates wastewater treatment plant (WWTP) effluent nutrients (including ammonia (NH3-N), nitrate (NO3-N) and total phosphorus (TP)) to the temporal and spatial distribution of epilithic algae and macrophyte biomass in an oligotrophic river. Annual macrophyte biomass, epilithic algae data and WWTP effluent nutrient data from 1980 to 2012 were statistically analysed. Because discharge can affect aquatic biomass growth, locally weighted scatterplot smoothing (LOWESS) was used to remove the influence of river discharge from the aquatic biomass (macrophytes and algae) data before further analysis was conducted. The results from LOWESS indicated that aquatic biomass did not increase beyond site-specific threshold discharge values in the river. The LOWESS-estimated biomass residuals showed a variable response to different nutrients. Macrophyte biomass residuals showed a decreasing trend concurrent with enhanced nutrient removal at the WWTP and decreased effluent P loading, whereas epilithic algae biomass residuals showed greater response to enhanced N removal. Correlation analysis between effluent nutrient concentrations and the biomass residuals (both epilithic algae and macrophytes) suggested that aquatic biomass is nitrogen limited, especially by NH3-N, at most sampling sites. The response of aquatic biomass residuals to effluent nutrient concentrations did not change with increasing distance to the WWTP but was different for P and N, allowing for additional conclusions about nutrient limitation in specific river reaches. The data further showed that the mixing process between the effluent and the river has an influence on the spatial distribution of biomass growth.

Ocean Acidification Accelerates the Growth of Two Bloom-Forming Macroalgae.

While there is growing interest in understanding how marine life will respond to future ocean acidification, many coastal ecosystems currently experience intense acidification in response to upwelling, eutrophication, or riverine discharge. Such acidification can be inhibitory to calcifying animals, but less is known regarding how non-calcifying macroalgae may respond to elevated CO2. Here, we report on experiments performed during summer through fall with North Atlantic populations of Gracilaria and Ulva that were grown in situ within a mesotrophic estuary (Shinnecock Bay, NY, USA) or exposed to normal and elevated, but environmentally realistic, levels of pCO2 and/or nutrients (nitrogen and phosphorus). In nearly all experiments, the growth rates of Gracilaria were significantly increased by an average of 70% beyond in situ and control conditions when exposed to elevated levels of pCO2 (p<0.05), but were unaffected by nutrient enrichment. In contrast, the growth response of Ulva was more complex as this alga experienced significantly (p<0.05) increased growth rates in response to both elevated pCO2 and elevated nutrients and, in two cases, pCO2 and nutrients interacted to provide a synergistically enhanced growth rate for Ulva. Across all experiments, elevated pCO2 significantly increased Ulva growth rates by 30% (p<0.05), while the response to nutrients was smaller (p>0.05). The δ13C content of both Gracilaria and Ulva decreased two-to-three fold when grown under elevated pCO2 (p<0.001) and mixing models demonstrated these macroalgae experienced a physiological shift from near exclusive use of HCO3- to primarily CO2 use when exposed to elevated pCO2. This shift in carbon use coupled with significantly increased growth in response to elevated pCO2 suggests that photosynthesis of these algae was limited by their inorganic carbon supply. Given that eutrophication can yield elevated levels of pCO2, this study suggests that the overgrowth of macroalgae in eutrophic estuaries can be directly promoted by acidification, a process that will intensify in the coming decades.

Characterisation of fucoidan extracted from Malaysian Sargassum binderi.

Fucoidan is a sulphated polysaccharide that consists mainly of fucose, normally found in brown seaweeds. In this study, fucoidan was extracted from Sargassum binderi (Fsar) from Malaysia and subsequently characterised. The chemical characteristics of Fsar were found to be different than those of commercial food grade fucoidan (Fysk) and those of previously studied fucoidans. NMR analysis proposed that the main structure of Fsar is →3)fuc-2-OSO3(-)(1→3)fuc(1→. The molecular weight (47.87kDa) and degree of sulphation (0.20) of Fsar were higher than those of Fysk, at 27.98kDa and 0.15, respectively. However, Fsar's polydispersity index (1.12) and fucose content (34.50%) were lower than those of Fysk, at 1.88 and 43.30%, respectively. Both Fsar and Fysk showed similar thermo-gravimetric properties with four mass losses, amorphous in nature and negative optical rotations. Results show that Fsar has fundamental characteristics of fucoidan with different structural conformation i.e. variation in glycosidic linkages and sulphate group orientation.

Combined nutrient and macroalgae loads lead to response in seagrass indicator properties.

Excess nutrients are potential factors that drive phase shifts from seagrasses to macroalgae. We carried out a manipulative field experiment to study the effects of macroalgae Ulva pertusa loading and nutrient addition to the water column on the nitrogen (N) and carbon (C) contents (i.e., fast indicators) as well as on the morphology and structure (i.e., slow indicators) of Zostera marina. Our results showed rapid impact of increased macroalgae and nutrient load on Z. marina C/N ratios. Also, macroalgae addition resulted in a trend of decreasing belowground biomass of seagrasses, and nutrient load significantly decreased above to belowground biomass ratio. Although some morphological/structural variables showed relatively fast responses, the effects of short-term disturbance by macroalgae and nutrients were less often significant than on physiological variables. Monitoring of seagrass physiological indicators may allow for early detection of eutrophication, which may initiate timely management interventions to avert seagrass loss.

Neuroprotective effects of the cultivated Chondrus crispus in a C. elegans model of Parkinson's disease.

Parkinson's disease (PD) is the second most common neurodegenerative disorder in the elderly people, currently with no cure. Its mechanisms are not well understood, thus studies targeting cause-directed therapy or prevention are needed. This study uses the transgenic Caenorhabditis elegans PD model. We demonstrated that dietary supplementation of the worms with an extract from the cultivated red seaweed Chondrus crispus decreased the accumulation of α-synulein and protected the worms from the neuronal toxin-, 6-OHDA, induced dopaminergic neurodegeneration. These effects were associated with a corrected slowness of movement. We also showed that the enhancement of oxidative stress tolerance and an up-regulation of the stress response genes, sod-3 and skn-1, may have served as the molecular mechanism for the C. crispus-extract-mediated protection against PD pathology. Altogether, apart from its potential as a functional food, the tested red seaweed, C. crispus, might find promising pharmaceutical applications for the development of potential novel anti-neurodegenerative drugs for humans.

Efficacy and safety of a dieckol-rich extract (AG-dieckol) of brown algae, Ecklonia cava, in pre-diabetic individuals: a double-blind, randomized, placebo-controlled clinical trial.

The effects of 12 weeks of supplementation with a dieckol-rich extract (AG-dieckol) from brown algae, Ecklonia cava, on glycemic parameters, serum biochemistry, and hematology were investigated in this study. Eighty pre-diabetic male and female adults were enrolled in a randomized, double-blind, placebo-controlled trial with parallel-group design. Subjects were randomly allocated into two groups designated as placebo and AG-dieckol (1500 mg per day). Compared with the placebo group, the AG-dieckol group showed a significant decrease in postprandial glucose levels after 12 weeks. The AG-dieckol group also showed a significant decrease in insulin and C-peptide levels after 12 weeks, but there was no significant difference between the AG-dieckol and placebo groups. There were no significant adverse events related to the consumption of AG-dieckol, and biochemical and hematological parameters were maintained within the normal range during the intervention period. In conclusion, these results demonstrate that AG-dieckol supplementation significantly contributes to lowering postprandial hyperglycemia and in reducing insulin resistance. Furthermore, we believe that based on these results the consumption of phlorotannin-rich foods such as marine algae may be useful for the treatment of diabetes.

Effects of N and P enrichment on competition between phytoplankton and benthic algae in shallow lakes: a mesocosm study.

Competition for resources between coexisting phytoplankton and benthic algae, but with different habitats and roles in functioning of lake ecosystems, profoundly affects dynamics of shallow lakes in the process of eutrophication. An experiment was conducted to test the hypothesis that combined enrichment with nitrogen (N) and phosphorus (P) would be a greater benefit to phytoplankton than benthic algae. The growth of phytoplankton and benthic algae was measured as chlorophyll a (Chl a) in 12 shallow aquatic mesocosms supplemented with N, P, or both. We found that enrichment with N enhanced growth of benthic algae, but not phytoplankton. P enrichment had a negative effect on benthic algal growth, and no effect on the growth of phytoplankton. N+P enrichment had a negative effect on benthic algae, but enhanced the growth of phytoplankton, thus reducing the proportion of benthic algae contributing to the combined biomass of these two groups of primary producers. Thus, combined N+P enrichment is more favorable to phytoplankton in competition with benthic algae than enrichment with either N or P alone. Our study indicates that combined enrichment with N+P promotes the dominance of phytoplankton over benthic algae, with consequences for the trophic dynamics of shallow lake ecosystems.

Temporal pattern in the bloom-forming macroalgae Chaetomorpha linum and Ulva pertusa in seagrass beds, Swan Lake lagoon, North China.

Seagrasses that are distributed over a large area of the Swan Lake, Weihai, China, support a productive ecosystem. In recent years, however, frequent macroalgal blooms have changed the ecosystem structure and threatened the seagrasses. To understand the bloom-forming macroalgae we conducted a yearly field survey of Swan Lake. Results indicated that the macroalgae Chaetomorpha linum and Ulva pertusa both exhibited a much higher productivity and attained a greater maximum biomass (of 1712±780gDWm(-)(2) and 1511 ± 555 gDW m(-2), respectively) than was the case for the seagrasses. The mean annual atomic ratios of C/N, C/P and N/P in C. linum were 14.31 ± 4.45, 402.82 ± 130.25, and 28.12±2.08, respectively. The δ(15)N values (11.09 ± 0.91‰ for C. linum; 9.27 ± 2.83‰ for U. pertusa) indicated a land-based source of N enrichment to the macroalgal blooms. High concentrations of nitrogen and phosphorus in the lagoon, particularly near the river mouth, supported the blooms.

Nitrate and phosphate regimes induced lipidomic and biochemical changes in the intertidal macroalga Ulva lactuca (Ulvophyceae, Chlorophyta).

This study was carried out in order to understand the lipid and biochemical alterations resulting from different nutritional regimes of nitrate and phosphate in Ulva lactuca. The algal thalli cultured in artificial seawater (ASW) showed higher levels of carbohydrates and non-polar lipids and increased phosphatase activities, accompanied by degradation of polar lipids, proteins and pigments. Further, higher levels of lipid hydroperoxides indicated reative oxygen species (ROS)-mediated non-enzymatic lipid peroxidation due to nutritional limitation-induced oxidative stress. Those thalli cultured in ASW supplemented with nitrate showed responses corresponding to nitrate addition, such as an increase in pigments, monogalactosyldiacylglycerols, polyunsaturated fatty acids and nitrate reductase. In addition, these thalli showed partial induction of phosphatases, low phospholipids, and high sulfolipid and 1,2-diacylglyceryl-3-O-4'-(N,N,N-trimethyl)-homoserine (DGTS) due to phosphate limitation. Similarly, algal thalli cultured in ASW supplemented with phosphate showed down-regulation of phosphatases, an increase in phospholipids due to availability of phosphate as well as a decrease in nitrate reductase, pigment, monogalactosyldiacylglycerols and polyunsaturated fatty acids due to nitrate limitation. On the other hand, algal thalli cultured in ASW supplemented with both nitrate and phosphate showed recovery of lost pigments and proteins, a high monogalactosyldiacylglycerol/digalactosyldiacylglycerol ratio, high unsaturation and high oxylipin levels (both C18 and C20). Further, the accumulation of indole-3-acetic acid in nutrient-limited thalli and of kinetin and kinetin riboside in nutrient-supplemented thalli indicated their antagonistic roles under nutrient stress. Thus, U. lactuca copes with nitrate and phosphate nutritional stress by altering the metabolic pathways involved in lipid biosynthesis including a shift in lipid classes, fatty acids, oxylipins and indole-3-acetic acid/kinetin cross-talk.

Dominance patterns in macroalgal and phytoplankton biomass under different nutrient loads in subtropical coastal lagoons of the SE Gulf of California.

Nine macroalgal blooms were studied in five coastal lagoons of the SE Gulf of California. The nutrient loads from point and diffuse sources were estimated in the proximity of the macroalgal blooms. Chlorophyll a and macroalgal biomass were measured during the dry, rainy and cold seasons. Shrimp farms were the main point source of nitrogen and phosphorus loads for the lagoons. High biomasses were found during the dry season for phytoplankton at site 6 (791.7±34.6 mg m(-2)) and during the rainy season for macroalgae at site 4 (296.0±82.4 g m(-2)). Depending on the season, the phytoplankton biomass ranged between 40.0 and 791.7 mg m(-2) and the macroalgal biomass between 1 and 296.0 g m(-2). The bulk biomass (phytoplankton+macroalgal) displayed the same tendency as the nutrient loads entering the coastal lagoons. Phytoplankton and macroalgal biomass presented a significant correlation with the atomic N:P ratio.

'Halophyte filters': the potential of constructed wetlands for application in saline aquaculture.

World consumption of seafood continues to rise, but the seas and oceans are already over-exploited. Land-based (saline) aquaculture may offer a sustainable way to meet the growing demand for fish and shellfish. A major problem of aquaculture is nutrient waste, as most of the nutrients added through feed are released into the environment in dissolved form. Wetlands are nature's water purifiers. Constructed wetlands are commonly used to treat contaminated freshwater effluent. Experience with saline systems is more limited. This paper explores the potential of constructed saline wetlands for treating the nutrient-rich discharge from land-based saline aquaculture systems. The primary function of constructed wetlands is water purification, but other ancillary benefits can also be incorporated into treatment wetland designs. Marsh vegetation enhances landscape beauty and plant diversity, and wetlands may offer habitat for fauna and recreational areas. Various approaches can be taken in utilizing plants (halophytes, macro-algae, micro-algae) in the treatment of saline aquaculture effluent. Their strengths and weaknesses are reviewed here, and a conceptual framework is presented that takes into account economic and ecological benefits as well as spatial constraints. Use of the framework is demonstrated for assessing various saline aquaculture systems in the southwestern delta region of the Netherlands.

Nutrient supply from fishes facilitates macroalgae and suppresses corals in a Caribbean coral reef ecosystem.

On coral reefs, fishes can facilitate coral growth via nutrient excretion; however, as coral abundance declines, these nutrients may help facilitate increases in macroalgae. By combining surveys of reef communities with bioenergetics modeling, we showed that fish excretion supplied 25 times more nitrogen to forereefs in the Florida Keys, USA, than all other biotic and abiotic sources combined. One apparent result was a positive relationship between fish excretion and macroalgal cover on these reefs. Herbivore biomass also showed a negative relationship with macroalgal cover, suggesting strong interactions of top-down and bottom-up forcing. Nutrient supply by fishes also showed a negative correlation with juvenile coral density, likely mediated by competition between macroalgae and corals, suggesting that fish excretion may hinder coral recovery following large-scale coral loss. Thus, the impact of nutrient supply by fishes may be context-dependent and reinforce either coral-dominant or coral-depauperate reef communities depending on initial community states.

Spread of Alsidium corallinum C. Ag. in a Tyrrhenian eutrophic lagoon dominated by opportunistic macroalgae.

In 2007, the Rhodophyceae Alsidium corallinum C. Ag., a marine taxon, bloomed in the eutrophic lagoon of Orbetello (Tuscany, Italy) for the first time, becoming the dominant species in spring and summer. In November, its biomass collapsed. The hypothesis examined in this study is that the bloom expressed a relatively low eutrophic level of the ecosystem after intense disposal of accumulated sedimentary organic matter (OM) by dystrophic processes in the two years preceding the bloom. To verify the hypothesis, we compared water physical-chemical variables, sediment redox (Eh) and OM, and standing crops of macroalgae and seagrass from the database of routine monitoring between 2005 and 2008. We also used dissolved nutrient data obtained in 2007 and 2008, as well as data on chlorophyll and total suspended matter in the water column during the microalgal bloom of 2007, and C, N and P content in thalli of the Chlorophycea Chaetomorpha linum and the Rhodophyceae Gracilariopsis longissima and A. corallinum obtained in 2007. In 2007, unusually low values of dissolved inorganic nitrogen (DIN) were recorded. Combined with stable values of soluble reactive phosphorus (SRPs), low DIN led to a reduction of about one order of magnitude in the DIN:SRP atomic ratio with respect to the past and to 2008. G. longissima accumulated C, N and P more than the other species and A. corallinum proved to be less demanding. Sediment OM was lower in the autumn of years characterized by dystrophy, confirming that summer dystrophic events coincided with maximum energy dissipation in this ecosystem. However, as soon as OM and DIN values increased (2008), the vegetation shifted towards blooms of G. longissima and C. linum, while A. corallinum almost disappeared. The results sustain the hypothesis that the bloom of A. corallinum was due to a decline in DIN that limited G. longissima, and to intense turbidity of the water caused by microphytes that developed after the dystrophic event of summer 2006. The latter probably limited the development of C. linum, which could only develop at the edges of the lagoon.

Seaweeds early development: detrimental effects of desiccation and attenuation by algal extracts.

The effects of desiccation on the early development stages of Mazzaella laminarioides, Scytosiphon lomentaria and Lessonia nigrescens, algal species with different patterns of distribution across the intertidal zone, were examined in the laboratory. In addition, the protective effect against desiccation was evaluated using algal extracts, including those from Porphyra columbina, a macroalga tolerant to desiccation that lives in the uppermost part of the intertidal zone. Our results showed that M. laminarioides displayed the highest resistance to daily desiccation, followed by S. lomentaria, whereas L. nigrescens was the most susceptible. Spores from L. nigrescens exposed to desiccation, although being able to germinate, ceased further post-germination development. In addition, our results showed that all species exposed to extracts from desiccated P. columbina successfully completed their development and strongly suggest the occurrence of compounds with protective properties that help in attenuating the stress caused by desiccation. Finally, our results indicate that the magnitude of the effects generated by desiccation on the early algal development is related to the position of the species in the intertidal zone, and that the protective effects of P. columbina extracts reveal an exceptional metabolism of this species under desiccation stress.