Ocean acidification may benefit algae that are able to capitalize on increased carbon availability for photosynthesis, but it is expected to have adverse effects on calcified algae through dissolution. Shifts in dominance between primary producers will have knock-on effects on marine ecosystems and will likely vary regionally, depending on factors such as irradiance (light vs. shade) and nutrient levels (oligotrophic vs. eutrophic). Thus experiments are needed to evaluate interactive effects of combined stressors in the field. In this study, we investigated the physiological responses of macroalgae near a CO 2 seep in oligotrophic waters off Vulcano (Italy). The algae were incubated in situ at 0.2 m depth using a combination of three mean CO 2 levels (500, 700-800 and 1200 μatm CO 2 ), two light levels (100 and 70% of surface irradiance) and two nutrient levels of N, P, and K (enriched vs. non-enriched treatments) in the non-calcified macroalga Cystoseira compressa (Phaeophyceae, Fucales) and calcified Padina pavonica (Phaeophyceae, Dictyotales). A suite of biochemical assays and in vivo chlorophyll a fluorescence parameters showed that elevated CO 2 levels benefitted both of these algae, although their responses varied depending on light and nutrient availability. In C. compressa, elevated CO 2 treatments resulted in higher carbon content and antioxidant activity in shaded conditions both with and without nutrient enrichment-they had more Chla, phenols and fucoxanthin with nutrient enrichment and higher quantum yield (F v /F m ) and photosynthetic efficiency (α ETR ) without nutrient enrichment. In P. pavonica, elevated CO 2 treatments had higher carbon content, F v /F m , α ETR , and Chla regardless of nutrient levels-they had higher concentrations of phenolic compounds in nutrient enriched, fully-lit conditions and more antioxidants in shaded, nutrient enriched conditions. Nitrogen content increased significantly in fertilized treatments, confirming that these algae were nutrient limited in this oligotrophic part of the Mediterranean. Our findings strengthen evidence that brown algae can be expected to proliferate as the oceans acidify where physicochemical conditions, such as nutrient levels and light, permit.

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Short-term ecophysiological and biochemical responses of Cystoseira tamariscifolia and Ellisolandia elongata to changes in solar irradiance and nutrient levels were analyzed in situ in oligotrophic coastal waters by transferring macroalgae collected at 0.5 and 2.0 m depth and exposing them to 2 irradiance levels (100 and 70% of surface irradiance) and nutrient conditions (nutrient-enriched and non-enriched). Both species were affected by changes in irradiance and nutrient levels. Few interactive effects between these 2 physical stressors were found, suggesting major additive effects on both species. C. tamariscifolia collected at 0.5 m and exposed to 70% irradiance had the highest maximal electron transport rate (ETR max ), saturated irradiance (Ek ETR ) and chl a content and the lowest antioxidant activity. Under the same conditions, E. elongata had increased Ek ETR , antheraxanthin and β-carotene content. At 100% irradiance, C. tamariscifolia collected at 2.0 m had higher maximal quantum yield (F v /F m ), photosynthetic efficiency (α ETR ), ETR max , maximal non-photochemical quenching (NPQ max ), saturation irradiance for NPQ (Ek NPQ ), and antheraxanthin and polyphenol content increased, whereas in E. elongata only α ETR increased. In nutrient-enriched conditions, phenolic compounds, several carotenoids and N content increased in C. tamariscifolia at both depths. E. elongata from 2.0 m depth at 100% irradiance and nutrient-enriched conditions showed increased N content and total mycosporine-like amino acids (MAAs). Our results show rapid photophysiological responses of C. tamariscifolia to variations in in situ irradiance and nutrient conditions, suggesting efficient photoacclimation to environmental changes. In E. elongata, F v /F m and ETR max did not change in the transplant experiment; in contrast, N content, pigment and MAAs (biochemical variables) changed. The responses of these macroalgae to nutrient enrichment indicate oligotrophic conditions at the study site and environmental stress.

Summary: Cystoseira tamariscifolia thalli collected from rocky shores and rockpools in winter and summer in Southern Spain were incubated for 7 days in UV transparent cylindrical vessels under outdoor conditions. Photosynthetic activity estimated as in vivo chlorophyll a fluorescence of photosystem II, photosynthetic pigments, antioxidant activity (DPPH assay), phenolic compounds and total internal C and N contents were determined after short-term (3 d) and mid-term (7 d) periods. Maximum quantum yield of PSII (F v /F m ) was significantly higher in field-collected algae and after 7 d incubation in winter than in summer. In rocky shores and rockpools thalli, maximum electron transport rate (ETR max ) and photosynthetic efficiency (a ETR ) were much higher in summer than in winter. ETR of outdoor-grown thalli (in situ ETR) showed a daily pattern, with a decrease at noon in both winter and summer (3 rd and 7 th days). We found much higher antioxidant activity in thalli collected in summer than in winter. However, the concentration of internal UV screen substances (polyphenols) was higher in winter than in summer, whereas the release of phenolic compounds was lower. The highest capacity of acclimation in C. tamariscifolia found in summer and RS with emersion periods was explained by the highest dynamic photoinhibition, energy dissipation (non-photochemical quenching) and antioxidant activity (EC 50 ).Keywords: antioxidant activity; Cystoseira tamariscifolia; phenolic compounds; photoinhibition; photoprotection. Patrones estacionales de fotoaclimatación en el alga intermareal, Cystoseira tamariscifolia (Ochrophyta)Resumen: Talos de Cystoseira tamariscifolia recolectados en pozas y plataformas rocosas intermareales (Sur de España) en invierno y en verano se incubaron bajo radiación solar durante 7 días en recipientes cilíndricos de metacrilato transparentes a la radiación UV. Se estimó la actividad fotosintética a través de la fluorescencia de la clorofila a asociada al fotosistema II, el contenido de pigmentos fotosintéticos y compuestos fenólicos, actividad antioxidante y el contenido total en C y N internos tras 3 y 7 días de incubación. Los valores iniciales del rendimiento cuántico máximo (F v /F m ) fueron significativamente mayores en algas recolectadas en invierno que en verano mientras que la tasa de transporte electrónico máximo (ETR max ) y la eficiencia fotosintética fueron mayores en verano que en invierno en ambas zonas. Por otra parte, la tasa de transporte electrónico determinada bajo radiación solar presentó un patrón diario, con una disminución a mediodía, tanto en invierno como en los períodos de verano. La actividad antioxidante fue mayor en algas recogidas en verano; sin embargo, la concentración interna de compuestos fenólicos fue mayor en invierno que en verano, mientras que en la tasa de excreción se observó lo contrario. La alta capacidad de aclimatación en C. tamariscifolia en algas sometidas a emersión en las plataformas rocosas en verano se explica por su alta fotoinhibición dinámica, su c...

Three macroalgal species belonging to Chlorophyta (Ulva rigida), Rhodophyta (Ellisolandia elongata) and Phaeophyceae (Heterokontophyta; Cystoseira tamariscifolia), naturally growing at the same shore level and representing 3 morpho-functional groups, were exposed to short-term changes in temperature under different carbon and nitrogen regimes. Experiments were conducted in outdoor tanks at 4 combinations of carbon and nitrogen levels under reduced solar radiation. In vivo chl a fluorescence parameters and pigment contents were monitored to assess diurnal physiological responses and potential for recovery. Strong fluctuations in chl a fluorescence parameters, but not in chl a content, were observed in response to diurnal variation in solar radiation and light climate within the tanks; sensitivity varied between algal species and, in some cases, depended on the carbon and nitrogen regime. Nitrogen uptake was similarly high in U. rigida and E. elongata and lowest in C. tamariscifolia. In U. rigida and E. elongata, chl a concentrations decreased after high-carbon treatments. Effective photosystem II quantum efficiency was reduced in all species at noon, and lowest in C. tamariscifolia. The results highlight the complexity of physiological short-term acclimations which were most likely linked to biochemical changes at the cellular level. Long-term experiments are required in future for more comprehensive investigation of the observed interactive effects of the different environmental parameters

There is currently no information regarding the role that whole mitogen activated protein kinase (MAPK) pathways play in counteracting environmental stress in photosynthetic organisms. To address this gap, we exposed Ulva compressa to chronic levels of copper (10 µM) specific inhibitors of Extracellular Signal Regulated Kinases (ERK), c-Jun N-terminal Kinases (JNK), and Cytokinin Specific Binding Protein (p38) MAPKs alone or in combination. Intracellular copper accumulation and photosynthetic activity (in vivo chlorophyll a fluorescence) were measured after 6 h, 24 h, 48 h, and 6 days of exposure. By day 6, when one (except JNK) or more of the MAPK pathways were inhibited under copper stress, there was a decrease in copper accumulation compared with algae exposed to copper alone. When at least two MAPKs were blocked, there was a decrease in photosynthetic activity expressed in lower productivity (ETRmax), efficiency (αETR), and saturation of irradiance (EkETR), accompanied by higher non-photochemical quenching (NPQmax), compared to both the control and copper-only treatments. In terms of accumulation, once the MAPK pathways were partially or completely blocked under copper, there was crosstalk between these and other signaling mechanisms to enhance metal extrusion/exclusion from cells. Crosstalk occurred among MAPK pathways to maintain photosynthesis homeostasis, demonstrating the importance of the signaling pathways for physiological performance. This study is complemented by a parallel/complementary article Rodríguez-Rojas et al. on the role of MAPKs in copper-detoxification.

UV-absorbing compounds, such as mycosporine-like amino acids (MAAs), are a group of secondary metabolites present in many marine species, including red seaweeds. In these organisms, the content and proportion of the composition of MAAs vary, depending on the species and several environmental factors. Its high cosmetic interest calls for research on the content and composition of MAAs, as well as the dynamics of MAAs accumulation in seaweeds from different latitudes. Therefore, this study aimed to survey the content of UV-absorbing MAAs in three Subantarctic red seaweeds during a seasonal cycle. Using spectrophotometric and HPLC techniques, the content and composition of MAAs of intertidal Iridaea tuberculosa, Nothogenia fastigiate, and Corallina officinalis were assessed. Some samples were also analyzed using high-resolution mass spectrometry coupled with HPLC-ESI-MS in order to identify more precisely the MAA composition. I. tuberculosa exhibited the highest MAA values (above 1 mg g−1 of dried mass weight), while C. officinalis showed values not exceeding 0.4 mg g−1. Porphyra-334 was the main component in N. fastigiata, whereas I. tuberculosa and C. officinalis exhibited a high content of palythine. Both content and composition of MAAs varied seasonally, with high concentration recorded in different seasons, depending on the species, i.e., winter (I. tuberculosa), spring (N. fastigiata), and summer (C. officinalis). HPLC-ESI-MS allowed us to identify seven different MAAs. Two were recorded for the first time in seaweeds from Subantarctic areas (mycosporine-glutamic acid and palythine-serine), and we also recorded an eighth UV-absorbing compound which remains unidentified.

Short-term effects of pCO2 (700 – 380 ppm; HC-LC) and nitrate content (50-5 βM; HN-LC) on photosynthesis, estimated by different pulse amplitude modulated (PAMs) fluorometers and by oxygen evolution, were investigated in Ulva rigida (Chlorophyta) under solar radiation (ex-situ) and in the laboratory under artificial light (in-situ). After 6-days of incubation at ambient temperature (AT), algae were subjected to a 4 oC-temperature increase (AT+4oC) for 3 d. Both in-situ and ex-situ, maximal electron transport rate (ETRmax) and in situ gross photosynthesis (GP) measured by O2 evolution presented the highest values under HCHN, and the lowest under HCLN, across all measuring systems. Maximal quantum yield (Fv/Fm), and ETRmax of PSII (ETR(II)max) and of PSI (ETR(I)max), decreased under HCLN under AT+4°C. Ex situ ETR was higher than in situ ETR. At noon, Fv/Fm decreased (indicating photoinhibition), whereas ETR(II)max and maximal non-photochemical quenching (NPQmax) increased. ETR(II)max decreased under AT+4oC in contrast to Fv/Fm, photosynthetic efficiency (αETR) and saturated irradiance (EK). Thus, U. rigida exhibited a decrease in photosynthetic production under acidification, LN levels and AT+4oC. These results emphasize the importance of studying the interactive effects between environmental parameters using in-situ vs. ex-situ conditions when aiming to evaluate the impact of global change on marine macroalgae.