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Fluctuations in solar radiation are one of the key factors affecting productivity and survival in habitat forming coastal macroalgae, in this regard, photoacclimation has a direct impact on the vulnerability and the capacity of seaweed to withstand, for instance, radiation excess. Here, we study ecophysiological responses through photosynthetic activity measurements under time-dependent (one year) fluctuations in solar radiation in the brown macroalga L. spicata. The responses presented seasonal patterns, with an increase in photosynthetic capacity during summer, expressed in greater maximal electron transport rate (ETRmax) and diminished thermal dissipation (NPQmax). Moreover, we studied photoprotective compounds (phenolic compounds) and total antioxidant capacity, which demonstrated an increase during periods of high solar radiation. In addition, content of photosynthetic pigment (Chla, Chlc and Carotenoids) increased under greater solar irradiance. The L. spicata can accumulate as reservoir photoprotective and antioxidant substances to withstand periods of high solar irradiance. All ecophysiological and biochemical responses in L. spicata indicate high photoacclimation and low vulnerability in the species, especially during with greater levels of solar irradiance.

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&minus, 1 of dried mass weight), while C. officinalis showed values not exceeding 0.4 mg g&minus, 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.

Radiation continuously varies in the environment, exposing phototrophic organisms to diverse compositions and ratios among photosynthetically active radiation (PAR), UV-A (Ultraviolet A radiation) and UV-B (Ultraviolet B radiation). In this sense, UV radiation can control, either positively or negatively, the physiology and metabolism of marine organisms in intertidal areas. Our study focused on the exposure of Gracilaria tenuistipitata to variable combinations of radiation, evaluating its physiology, photoprotective strategies and DNA damage under each condition. To accomplish this, algal apical fragments were acclimated to three radiation treatments (PAR, PAR + UV-A and PAR + UV-A + UV-B) for one week. Thereafter, PAR-acclimated samples were moved to PAR (control), PAR + UV-A + UV-B and PAR + UV-B. PAR + UV-A-acclimated samples were moved to PAR + UV-A (control), PAR + UV-A + UV-B and PAR + UV-B, and PAR + UVR-acclimated thalli were placed in PAR + UV-A + UV-B (control), PAR + UV-B and PAR. Samples remained in these conditions for a period of 48 h continuously. Photosynthetic parameters from chlorophyll a fluorescence were evaluated, as well as growth rates, mycosporine-like amino acids, chlorophyll a, and carotenoid contents. In addition, DNA damage was assessed by detecting cyclobutane pyrimidin dimer. Expression of photolyases, ascorbate peroxidase and gluthatione peroxidase was also analyzed at the end of experimental period. Mycosporine-like amino acids were stimulated by UV radiation. DNA damage occurred only when UV-B was provided separately. In some cases, photoacclimation procedures were activated, while in others, it was not enough. In other words, although some tolerance was observed, the species could suffer the consequences of excessive energy present in some specific treatments, e.g., PAR + UV-B. After exposure to variable radiation, it was concluded that our model organism presented acclimation depending of spectral availability, including regulation of photoprotective strategies.

In recent decades a decline in the foundation species Gelidium corneum (Hudson) J. V. Lamouroux has been detected along the Basque coast (northern Spain). This decline has been attributed to several factors, but recent studies have found a relationship between high irradiance and the biochemical and physiological stress of G. corneum . Since physiological responses to changes in light occur well before variations in morphology, the present study seeks to use a size-class demographic approach to investigate whether shallow subtidal populations of G. corneum off the Basque coast show different frond bleaching, density and biomass under different irradiance conditions. The results revealed that the bleaching incidence and cover were positively related to irradiance, whereas biomass was negatively related. The effect of the irradiance level on frond density was found to vary with size-class, i.e. fronds up to 15 cm showed greater densities under high light conditions (126.6 to 262.2 W m − 2 ) whereas the number of larger fronds (> 20 cm) per unit area was lower. In conclusion, the results of the present study suggest that irradiance might be a key factor for controlling along-shore bleaching, frond density and biomass in G. corneum . Further research should be carried out on the physiology of this canopy species in relation to its bed structure and on the interaction of irradiance and other abiotic (nutrients, temperature, wave energy) and biotic factors (grazing pressure).

The effects of increased solar photosynthetically active radiation (PAR) and ultraviolet radiation (UVR) on biomass, mycosporine-like amino acid (MAA) content, and carrageenan yield were studied for 14 days in tank-cultivated tetrasporophytes and gametophytes of Mazzaella laminarioides from the Magellan Strait. The initial values of carrageenan yield were similar between gametophytes and tetrasporophytes, whereas the MAA content was higher in gametophytes than that in tetrasporophytes. After 14 days of exposure to different combinations of solar radiation (PAR, PAR + UV-A, and PAR + UV-A + UV-B) and two irradiance levels (high and low), differences between gametophytes and tetrasporophytes of M. laminarioides in growth rate (GR), MAA, and carrageenan yield were observed. Higher GRs were observed in gametophytes than in tetrasporophytes under all radiation and irradiance treatments. A significant decrease in total MAA content was observed in both reproductive phases, with these decreases being mo...

The combined effects of ultraviolet radiation (UVR) and nutrients on the red macroalga Gracilariopsis longissima were analyzed for 14 days in a 2 × 2 factorial experiment design. Cell ultrastructure, photosynthetic activity as in vivo chlorophyll a fluorescence and the accumulation of bioactive compounds as soluble proteins, starch, lipids, polyphenols and mycosporine-like amino acids (MAAs) were evaluated. Three main morphological alterations as defense mechanisms against UVR were observed, i.e., an increase in cell wall thickness, accumulation of floridean starch grains and multiplication of the number of layers of cortical cells. Interaction of light and nutrients affected photosynthetic variables, such as electron transport rate (estimator of photosynthetic capacity) and photoinhibition irradiance (E opt ), and also the level of N compounds (i.e., proteins and MAAs) and total carbon and polyphenol content. Other variables such as chlorophyll a , carotenoids, starch, lipids, antioxidant activity and cell viability were influenced by both factors, or only by light, but without interactive effects. G . longissima showed a high acclimation to elevated UVR and nutrients by the decrease of the photosynthetic rate and the increase in the accumulation of N compounds with photoprotection and antioxidant capacities. On the other hand, starch, lipid and polyphenols increased mainly in the presence of UVR under low nutrient availability. The biotechnological implications of the metabolic changes due to N availability and UVR in this species are discussed.

Ocean acidification increases the amount of dissolved inorganic carbon (DIC) available in seawater which can benefit photosynthesis in those algae that are currently carbon limited, leading to shifts in the structure and function of seaweed communities. Recent studies have shown that ocean acidification-driven shifts in seaweed community dominance will depend on interactions with other factors such as light and nutrients. The study of interactive effects of ocean acidification and warming can help elucidate the likely effects of climate change on marine primary producers. In this study, we investigated the ecophysiological responses of Cystoseira tamariscifolia (Hudson) Papenfuss. This large brown macroalga plays an important structural role in coastal Mediterranean communities. Algae were collected from both oligotrophic and ultraoligotrophic waters in southern Spain. They were then incubated in tanks at ambient (ca. 400–500 ppm) and high CO2 (ca. 1200–1300 ppm), and at 20 °C (ambient temperature) and 24 °C (ambient temperature +4 °C). Increased CO2 levels benefited the algae from both origins. Biomass increased in elevated CO2 treatments and was similar in algae from both origins. The maximal electron transport rate (ETRmax), used to estimate photosynthetic capacity, increased in ambient temperature/high CO2 treatments. The highest polyphenol content and antioxidant activity were observed in ambient temperature/high CO2 conditions in algae from both origins; phenol content was higher in algae from ultraoligotrophic waters (1.5–3.0%) than that from oligotrophic waters (1.0–2.2%). Our study shows that ongoing ocean acidification can be expected to increase algal productivity (ETRmax), boost antioxidant activity (EC 50 ), and increase production of photoprotective phenols. Cystoseira tamariscifolia collected from oligotrophic and ultraoligotrophic waters were able to benefit from increases in DIC at ambient temperatures. Warming, not acidification, may be the key stressor for this habitat as CO2 levels continue to rise.

In order to evaluate the photosynthetic efficiency of cave photoautotrophic biofilms, in vivo chlorophyll a fluorescence was measured using two different fluorometers, in two biofilms in the touristic karstic Nerja Cave (Spain). The study was done for entire days in summer and in winter during the first year and repeated for a second year, in order to cover the widest range of environmental conditions, i.e., atmospheric CO2, temperature, seeping water and relative humidity levels. Effective quantum yield and relative electron transport rate (rETR) were determined during periods of light whereas maximal quantum yield (Fv/Fm) was determined in situ during dark periods. On summer days, in situ photosynthetic yields in cyanobacterium biofilms (Chroococcidiopsis sp.) increased 7–16 times compared to that of winter days, whereas in biofilms comprised of green and red microalgae and various cyanobacterium species, no seasonal or yearly variations were observed. In contrast, maximal rETR in the two biofilms increased in the periods with the highest values of both CO2 and relative humidity. Positive correlations between all environmental variables and rETR were found. According to Redundancy Analysis, all environmental variables, mainly CO2 and relative humidity were related to photosynthetic variables. The effective quantum yields showed different values depending on the measuring light of the PAM. The values were higher with red light (Diving PAM) compared to blue light (Junior PAM) mainly in the site dominated by cyanobacteria. Nerja Cave is shown as an excellent place to study the effects of light and CO2, among other environmental variables of biofilm photosynthetic activity. The monitoring of photosynthetic activity by in vivo chlorophyll a fluorescence could be used to follow the effects of the treatments applied by the touristic cave managers to reduce the proliferation of biofilms composed of various species, and, consequently, the biodeterioration of speleothems could be reduced.

The microalga Chlorella fusca showed high biomass productivity in thin-layer cascades (TLCs) with large light-exposed surface (high exposed surface to total volume ratio; S/V). Two trials – E1 and E2 – were conducted in Malaga (Southern Spain) in July and October, respectively using a TLC with an S/V ratio of 27 m− 1. The third trial E3 was carried out in the Czech Republic in July using another TLC with an S/V ratio of 120 m− 1. Chl a fluorescence measurements (in situ and off-line) were carried out to monitor the photosynthetic performance of these cultures. Simultaneous measurement of irradiance and effective quantum yield through the day allowed the determination of daily electron transport rate (ETR), which led to the estimation of biomass productivity. Calculated data of biomass productivity of microalgal mass cultures were well correlated with the measured values. The E3 culture achieved the highest productivity of the three trials − 27 g DW m− 2 d− 1. Extrapolating these results to southern Spain, with a longer cultivation period and higher daily dose of the solar energy, and considering a photosynthetic efficiency (PE) of 4.4% (achieved in E3), we obtained a potential productivity of 43 metric tons ha− 1 year− 1 if the unit is operated for 8 months (from March to October). Realistically, in locations with favorable climate, TLCs can be considered a suitable system for biomass production with high PE as well as easy maintenance and low operation costs. On-line monitoring of in situ Chl a fluorescence provided data that revealed essential information about the photosynthetic performance of the culture. Here, we present a first attempt of estimating biomass productivity based on ETR measurements in microalgal mass cultures that correlate well with the measured values.

Photosynthetic activity was estimated, in 2 different sets of experiments, as in vivo chlorophyll a fluorescence in 2 calcareous red macroalgae: Neogoniolithon brassica-florida, which forms vermetid reefs with Dendropoma petraeum, and the rhodolith-forming Lithophyllum margaritae. In the first set of experiments, vermetid reefs collected from the Cabo de Gata-Nijar Natural Park (Almeria, Spain) were incubated for 6 months in outdoor mesocosms at 2 different temperatures: ambient temperature (T) and ambient temperature plus 2 oC (T+). Daily variations in effective quantum yields were related to the increase in solar radiation. Electron transport rate (ETR) was higher at T+ than at T. Oxygen production, estimated from ETR, was higher in N. brassica-florida growing in vermetid reefs in the coastal area than in those incubated in mesocosms or under laboratory conditions. In the second set of experiments, L. margaritae specimens with or without attached epifauna (mainly sponges) were collected at 2 sites (Pedregoso and CFE) on the Baja California Peninsula, Mexico, and at 2 depths (1 and 2 m). ETR values for rhodoliths collected at 1 m depth and with attached epifauna were higher than those for rhodoliths collected at 2 m depth, indicating the importance of greater incident radiation on photosynthetic activity. Photosynthetic activity was higher in L. margaritae collected from Pedregoso, the site with better water quality (high transparency and low pollution) and rocky substrate availability. In the CFE samples, the photosynthetic efficiency (alphaETR) was highest in rhodoliths with attached epifauna. The highest ETR values for N. brassica-florida at T+ and the highest alphaETR or ETR values for L. margaritae with attached sponges may be explained by the increase in CO2 available for photosynthesis due to the respiration of the associated fauna. The usefulness of in vivo chlorophyll a fluorescence for estimating photosynthetic production in calcareous macroalgae under multifactorial experiments is discussed. La actividad fotosintetica fue estimada, en 2 series experimentales, mediante la fluorescencia in vivo de la clorofila a en 2 algas rojas calcareas: Neogoniolithon brassica-florida, que forma arrecifes de vermetidos con Dendropoma petraeum, y Lithophyllum margaritae (rodolitos). En la primera serie, vermetidos recolectados en el Parque Natural Cabo de Gata-Nijar (Almeria, Espana) fueron incubados en mesocosmos a diferentes temperaturas: temperatura ambiental (T) y temperatura ambiental mas 2 oC (T+). Las variaciones diurnas en el rendimiento cuantico efectivo estuvieron relacionadas con el incremento de irradiancia solar. La tasa de transporte de electrones (ETR) fue mas alta en el tratamiento T+ que en T. La produccion de oxigeno, estimada a partir de la ETR, fue mayor en N. brassica-florida del medio natural que en vermetidos incubados en mesocosmos o en el laboratorio. En la segunda serie experimental, L. margaritae con o sin epifauna (dominada por esponjas) fue recolectada en 2 sitios (Pedregoso y CFE) de la peninsula de Baja California, Mexico, a 2 profundidades (1 y 2 m). La ETR de rodolitos recolectados a 1 m de profundidad con epifauna asociada fue mas alta que la de aquellos recolectados a 2 m de profundidad, indicando la importancia del incremento de la irradiancia sobre la produccion fotosintetica. La actividad fotosintetica fue mayor en L. margaritae de Pedregoso, el lugar con mejor calidad de agua (alta trasparencia y baja contaminacion) y con disponibilidad de sustrato rocoso. En CFE, la eficiencia fotosintetica (alfaETR) fue mas alta en los rodolitos con epifauna. Los valores mas altos de ETR para N. brassica-florida a T+ y de alfaETR o ETR para L. margaritae con esponjas fijadas podrian ser explicados por el incremento de CO2 disponible para la fotosintesis procedente de la respiracion de la fauna asociada. En este trabajo se discute la utilidad de la fluorescencia in vivo de la clorofila a para estimar la fotosintesis en macroalgas calcareas en experimentos multifactoriales.

Made available in DSpace on 2018-11-26T17:55:15Z (GMT). No. of bitstreams: 0 Previous issue date: 2018-09-01 Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) Freshwater algae inhabiting shallow waters are most effectively affected by UV radiation. The aim of this study was to analyze the photosynthetic performance (parameters derived of chlorophyll fluorescence), chlorophyll a concentrations and the content of UV-absorbing compounds (mycosporine-like aminoacids-MAAs) in three tropical lotic macroalgae, Cladophora glomerata (Chlorophyta), Spirogyra sp. (Streptophyta) and Sirodotia delicatula (Rhodophyta) in response to UV radiation exposure under laboratory conditions. Experiments were performed under three treatments: (1) PAR (400-700 nm), P; (2) PAR + UVA (320-700 nm), PA; (3) PAR + UVA + UVB (280-700 nm), PAB. Cladophora glomerata and Spirogyra sp., typical sun-adapted lotic macroalgae, had distinct responses to UV radiation exposure. A more pronounced decrease of F (v)/F (m) and increase of NPQ under UVA than UVAB was observed, whereas chlorophyll a content was lower under UVAB in C. glomerata. Spirogyra sp. had a decrease of photosynthetic yields (Delta F/F (m)' and F (v)/F (m)) under PAR, indicating that UVAB radiation may have a positive effect on the photosynthetic apparatus. Surprisingly, S. delicatula, a shade-adapted alga, exhibited less sensitivity to UV exposure. These results suggest that the presence of MAAs (shinorine and palythine) in S. delicatula is a significant shield of protection against UV radiation. Fed Inst Educ Sci & Technol Sao Paulo IFSP, Campus Votuporanga, BR-15503110 Votuporanga, SP, Brazil Univ Fed Santa Catarina, Dept Bot, Campus Trindade S-N, BR-88040900 Florianopolis, SC, Brazil Malaga Univ, Fac Sci, Dept Ecol, Campus Teatinos, E-29071 Malaga, Spain Sao Paulo State Univ, Dept Bot & Zool, Rua Cristovao Colombo 2265, BR-15054000 S Jose Rio Preto, SP, Brazil Sao Paulo State Univ, Dept Bot & Zool, Rua Cristovao Colombo 2265, BR-15054000 S Jose Rio Preto, SP, Brazil

The red alga Gelidium corneum is a dominant foundation species in the south-eastern Bay of Biscay, where a decline in its populations has been documented in the few last decades. We investigated the ecophysiological responses of G. corneum to different light conditions by means of an in situ transplant experiment. We found that the stress response measured by physiological and biochemical approaches was higher in G. corneum at higher irradiance levels, for both transplanted and control specimens, than under lower light intensities. In the former case the specimens showed a decrease in maximum quantum yield (Fv/Fm), maximum electron transport rate (ETRmax), photosynthetic efficiency (αETR), photosynthetic pigment contents, nitrogen content and thallus length, whereas the C:N ratio, MAAs and bleaching cover increased. In general terms, these responses were more evident in the apical parts of the thallus than in middle ones. Our results suggest that high light stress at depths of 3 m triggered photobiological changes in G. corneum, involving ineffective photoprotection and the occurrence of chronic photoinhibition. Therefore, considering the upward trend in summer mean surface solar radiation in the study area since the 80s, high light conditions may have played a role in the declines observed in G. corneum beds from the south-eastern Bay of Biscay.

Radiation, both photosynthetic active radiation (PAR, l = 400–700 nm) and Ultraviolet (UVR, l = 280–400 nm) is one of the key factors regulating algal distribution in aquatic environments. Pyropia acanthophora and Grateloupia turuturu have been found over upper rocky shore areas in Southern Brazil, occupying the same niche space. The first species is native and the second one is exotic and considered a potential invader of South Atlantic. The aim of the present study was to evaluate the effects of radiation on physiological responses of both species and infer mechanisms that allow their niche competition in the environment. Samples were cultured in the following conditions: associated or separated, and with an addition of PAR, PAR + UVA (PA) and PAR + UVA + UVB (PAB), totalizing six factorial treatments during 5 days of exposure. Photosynthetic responses of Fv/Fm and ETR were daily evaluated. At the beginning and at the end of the experiment, samples were analyzed for pigment content (chlorophyll a and phycobiliproteins), and mycosporine-like amino acids (MAAs), while oxygen evolution was evaluated at the end of the experiment. As the main results, G. turuturu died when cultivated in PAB conditions. P. acanthophora presented higher amounts of chlorophyll a than G. turuturu during the whole experiment. Phycoerythrin and Fv/Fm remained constant in P. acanthophora but diminished for G. turuturu in UV treatments. ETR was higher for samples that were cultivated in associative treatment. The presence of G. turuturu in the same flask enhanced MAA synthesis in P. acanthophora, regardless of radiation condition. In addition, UV radiation can be a factor controlling species distribution and could counteract the spreading of invasive species, like G. turuturu, allowing P. acanthophora survival in upper rocky shore zones of the natural ecological distribution area.

The effect of different ultraviolet radiation (UVR) treatments combining PAR (P), UVA (A) and UVB (B) on the molecular physiology of Dunaliella tertiolecta was studied during 6 days to assess the response to chronic UVR exposure. UVR reduced cell growth but did not cause cell death, as shown by the absence of SYTOX Green labeling and cellular morphology. However, caspase-like enzymatic activities (CLs), (regarded as cell death proteases), were active even though the cells were not dying. Maximal quantum yield of fluorescence (Fv /Fm ) and photosynthetic electron transport rate (ETR) dropped. Decreased nonphotochemical quenching (NPQ) paralleled a drop in xanthophyll cycle de-epoxidation under UVB. Reactive oxygen species (ROS) and D1 protein accumulation were inversely correlated. PAB exhibited elevated ROS production at earlier times. Once ROS decayed, D1 protein recovered two-fold compared with P and PA at later stages. Therefore, PsbA gene was still transcribed, suggesting ROS involvement in D1 recovery by its direct effect on mRNA-translation. We add evidence of an UVB-induced positive effect on the cells when P is present, providing photoprotection and resilience, by means of D1 repair. This allowed cells to survive. The photoprotective mechanisms described here (which are counterintuitive in principle) conform to an important ecophysiological response regarding light stress acclimation.

Gracilaria species are the main source of agar worldwide. Since laboratory cultivation is an important means of sustaining such production, this study aimed to assess the influence of two fundamental strategies of physiological algae adjustment, photoacclimation and photomorphogenesis, on G. birdiae physiology. Specifically, the effects of different spectral light qualities on reproduction, growth, and pigment content in two distinct photoperiods (8 and 14 hours) were examined. Tetrasporophytes and gametophytes were submitted to different light qualities: white, green, red, and blue over the course of nine weeks. At the end of this period, chlorophyll a, allophycocyanin, phycocyanin, and phycoerythrin contents were analysed. Gametophytes showed reproductive structures only under monochromatic radiation. A stimulatory effect on tetrasporangium differentiation was mainly observed under red light, but this high fertility was negatively correlated with growth rate, i.e. algae cultivated under red light showed the lowest growth rate. However, while growth rates were higher in algae exposed to white light, they were similar to those observed in algae subjected to green light and 14 hours of daylight. PE was the predominant pigment, irrespective of light quality. Phycocyanin and phycoerythrin concentration increased when algae were cultivated under green and blue light. Therefore, considering future management prospects for G. birdiae mariculture, we suggest that red light could be utilized as a reproductive inductor to produce tetraspores. Furthermore, if the aim is to achieve high phycoerythrin content, continuous blue light could be applied during a short photoperiod. Indeed, the combination of different wavelengths could allow better economic resource exploitation.

Rock pools are dynamic and intermittently isolated habitats in the rocky intertidal. In this study, we assessed if the photosynthesis and physiological activity of the brown macroalga Cystoseira humilis and the red macroalga Digenea simplex in a rock pool at Lanzarote Island (eastern Atlantic) was affected due by their vertical distribution. Photosynthetic responses were measured at three depth levels (0.05–1 m, 0.4–2.5 and 3.5 m) through in vivo chlorophyll a fluorescence, in particular the maximal quantum yield ( F v / F m ) as an estimator of the physiological status and photoinhibition, and the electron transport rate (ETR) as an estimator of the photosynthetic capacity. Algal photoprotection and photodamage processes were related to algal zonation; shallow-water thalli had active mechanisms of dynamic photoinhibition compared to algae from the deeper level. The progressive increase of solar radiation during the day caused different responses for each macroalga, where C. humilis showed lower photoinhibition, higher ETR values than D. simplex. Algae from the shallow level had lower pigment content and higher resistance to high solar radiation through the accumulation of photoprotective compounds and higher antioxidant activity (DPPH) compared to thalli from the deeper level. In summary, this study corroborates that algae are vertically distributed inside rock pools according to their adaptive responses to light-induced stress conditions and that photoacclimation occur in a short-term period.

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 (ETRmax), saturated irradiance (EkETR) and chl a content and the lowest antioxidant activity. Under the same conditions, E. elongata had increased EkETR, antheraxanthin and β-carotene content. At 100% irradiance, C. tamariscifolia collected at 2.0 m had higher maximal quantum yield (Fv/Fm), photosynthetic efficiency (αETR), ETRmax, maximal non-photochemical quenching (NPQmax), saturation irradiance for NPQ (EkNPQ), and antheraxanthin and polyphenol content increased, whereas in E. elongata only αETR increased. In nutrient-enriched conditions, phenolic compounds, several carotenoids and N con- tent 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 varia- tions in in situ irradiance and nutrient conditions, suggesting efficient photoacclimation to envi- ronmental changes. In E. elongata, Fv/Fm and ETRmax 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.

Three macroalgal species belonging to Chlorophyta (Ulva rigida), Rhodophyta (Ellisolandia elongata) and Phaeophyceae (Heterokontophyta; Cystoseira tamariscifolia), natu- rally 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 flu- orescence 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 con- centrations 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 com- plexity 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 comprehen- sive investigation of the observed interactive effects of the different environmental parameters.

Since the mid-1990s, chlorophyll a (chl a) fluorescence measurement has become widespread to monitor photosynthetic performance of microalgal mass cultures. This paper aims to provide practical instructions for microalgal physiologists and biotechnologists on how fluores- cence monitoring can be used to explain changes in photosynthetic activity of microalgal mass cultures. Emphasis is placed on the 2 most common fluorescence techniques—pulse-amplitude- modulation and fluorescence induction kinetics—and the interpretation of important variables that reflect changes of photosynthesis and physiological status of microalgal cultures. In particu- lar, consideration is given to problems associated with the estimation of the photochemical yield of photosystem II and its relationship to linear electron transport rate and overall photosynthesis. Emphasis is also given to the applications and limitations of these techniques through several case studies. In model experiments with microalgal cultures, typical records and their interpretation under various culture conditions are illustrated. Changes of photosynthetic activity and selected variables monitored by chl a fluorescence techniques can thus be related to changes of cultivation conditions, physiological status and growth of microalgal cultures for a given microalgal strain and cultivation system. In this way, chl a fluorescence may be used as a rapid screening technique to monitor photosynthetic activity and subsequently to estimate growth rate in both indoor and outdoor studies. This text and results formed the basis of a key lecture at the 9 th International GAP workshop held at the University of Malaga in September 2012.

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 α 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 (α 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 ).

The vulnerability and acclimation to increased UVB radiation in three macroalgae of different morphofunctional groups collected in the Mediterranean coastal waters were evaluated. The algae were submitted for 7 days to increased (PABþ) and decreased (PAB� ) UVB radiation. The thickness and morphology influenced the response to increased UVB radiation, being Cystoseira tamariscifolia the less vulnerable algae followed by Ellisolandia elongata. The highest resistance to increased UVB radiation in C. tamariscifolia was related to the accumulation of polyphenols and high antioxidant activity, whereas E. elongata was due to its high reflectance. Finally, Ulva rigida suffered the highest photoinhibition under PABþ culture. The latest species presented 10 times lower polyphenol content and antioxidant activity than C. tamariscifolia. The three species showed different acclimation patterns to the changes of UVB radiation related to the morphology, photosynthetic activity, accumulation of photoprotectors and antioxidant activities. The ecological implications of the UVB variations on macroalgae are discussed. 2014 Elsevier Ltd. All rights reserved.

Short-term variations of mycosporine-like amino acids (MAAs) in Pyropia columbina (Montage) W.A. Nelson exposed to nitrate (NO3 -) enrichment under different outdoor light treatments during the spring ozone depletion of 2008 in Punta Arenas (Chile) were investigated. Segments of P. columbina thalli were cultivated under three treatments of solar radiation without or with NO3 - supply (0.38 mmol l-1): PAR (P), PAR+UVA (PA), and PAR+UVA+UVB (PAB). Samples were taken at 8:00 h (initial value), 9:30, 12:30, 15:30, and 18:00 h on November 8 and at 9:00 h on November 9 (recovery period). A complex dynamic of MAAs affected by light quality and NO3 - supply was observed. During the light period, the highest content of MAAs was reached under PAB and NO3 - enrichment, whereas MAAs increased during the recovery period in P and PAB with no NO3 - supply. Five MAAs were identified: porphyra-334, shinorine, asterina-330, palythine, and mycosporine-glycine. The hourly accumulation of each MAA varied mainly according to the time of exposure and NO3 - supply. In general, the percentage of porphrya-334 increased, whereas the other MAAs decreased during the exposure period. These results suggest that MAA content in P. columbina varied in the short-term (hours) and the changes were related to the solar irradiance received and NO3 - availability.

Global warming is driving shifts in the geographical distribution of seaweeds, such as the contraction of the Southern limit of cold-temperate Atlantic macroalgae, including the foundation intertidal fucoid Fucus serratus. However, the ecophysiological mechanisms explaining the decline of seaweeds is not fully understood. In vivo chlorophyll a fluorescence is used as an estimator of photosynthetic activity and physiological state of seaweeds. Collected at its southern distribution limit, Fucus serratus was cultured for two weeks in different temperatures (from 8 to 30 °C with increase of 2 °C). Growth rate, reproductive stage and fluorescence parameters were measured as maximal quantum yield (Fv/Fm), Electron transport rate (ETR), non-photochemical quenching (NPQ) and the yield losses (YNO and YNPQ).At high temperature, the photosynthetic responses mimic the pattern of shade adapted plants, i.e., increase of photosynthetic efficiency (αETR) and decrease of irradiance for saturation of photosynthetic rate (EkETR). At low irradiances (

In vivo chlorophyll fluorescence associated to Photosystem II is being used to evaluate photosynthetic activity of microalgae grown in different types of photobioreactors; however, controversy on methodology is usual. Several recommendations on the use of chlorophyll fluorescence to estimate electron transport rate and productivity of microalgae grown in thin-layer cascade cultivators and methacrylate cylindrical vessels are included. Different methodologies related to the measure of photosynthetic activity in microalgae are discussed: (1) measurement of light absorption, (2) determination of electron transport rates versus irradiance and (3) use of simplified devices based on pulse amplitude modulated (PAM) fluorescence as Junior PAM or Pocket PAM with optical fiber and optical head as measuring units, respectively. Data comparisons of in vivo chlorophyll fluorescence by using these devices and other PAM fluorometers as Water- PAM in the microalga Chlorella sp. (Chlorophyta) are presented. Estimations of carbon production and productivity by transforming electron transport rate to gross photosynthetic rate (as oxygen evolution) using reported oxygen produced per photons absorbed values and carbon photosynthetic yield based on reported oxygen/carbon ratio are also shown. The limitation of ETR as estimator of photosynthetic and biomass productivity is discussed. Low cost:quality PAMs can promote monitoring of chlorophyll fluorescence in algal biotechnology to estimate the photosynthetic activity and biomass productivity.

Gelidium corneum (Hudson) J.V. Lamouroux is a very important primary producer in the Cantabrian coastal ecosystem. Some local declines in their populations have been recently detected in the Basque coast. Occurrences of yellowing and an unusual branch breakdown pattern have also been reported for some G. corneum populations. In order to gain further insight into those environmental stressors operating at a local scale, here we investigate if shallow subtidal populations of G. corneum living under potentially different conditions of irradiance (PAR and UVA) and water temperature exhibit differences in some biochemical indicators of stress, namely C:N, antioxidant activity (radical cation of 2,2′-azino-bis (3-ethylbenzothiazoline-6-sulfonate); ABTS + assay) and mycosporine-like amino acids (MAAs) (Asterine 330 and Palythine). We hypothesised that G. corneum subjected to higher ambient levels of irradiance and water temperature would show higher C:N ratios, lower antioxidant activity and higher MAA concentrations. Our results partially support this hypothesis. We found that G. corneum exposed to increased levels of irradiance (PAR, UVA) exhibited greater C:N ratios and lower antioxidant activity (higher IC 50 ), whereas no relationship was found regarding MAAs. No differences in biochemical performance in relation to temperature were detected among G. corneum exposed to comparable high light. Similarly, G. corneum growing under lower UVA radiation levels showed no differences in any of the measured biochemical variables with regard to PAR and water temperature. These findings suggest that, among the environmental factors examined, UVA radiation may be an important driver in regulating the along-shore variation in G. corneum biochemical performance. Therefore, the role of irradiance, especially UV radiation, in potential future alterations in Cantabrian G. corneum populations cannot be ruled out as a potential underlying factor.

This study evaluates the production of biomass and mycosporine-like amino acids (MAAs) throughout the year in Gracilaria vermiculophylla (Rhodophyta) collected in Ria de Aveiro (Portugal). The algae were grown in outdoor tanks in seawater with the addition of fishpond effluents under two different water flows (100 and 200 L h−1) in an integrated multi-trophic aquaculture (IMTA) system (tanks 1200 L; 1.5 m2) and different algal densities (3, 5, and 7 kg m−2). MAA content in IMTA seaweeds was significantly affected by the interaction of time and stocking density, but not by the water flow. The highest MAA content was observed in April (about 3.13 mg g−1 DW) followed by May (1.79 mg g−1 DW). Seaweed biomass productivity was higher in May (372.06 g DW m−2 week−1) than in April (353.40 g DW m−2 week−1). Four MAAs were identified by HPLC and electrospray ionization mass spectrometry (ESI-MS) in G. vermiculophylla: Porphyra-334, Shinorine, Palythine and Asterina-330. The highest levels of Porphyra-334 and Shinorine were reached from November to January and the Palythine + Asterina-330 from April to August. Taking into account the average biomass and MAA production of G. vermiculophylla growing in this IMTA system (8.56 g of MAA in 18 m2 culture along 8 months; 35.5% produced in April), a total amount of 71.33 g MAA year−1 could be produced in this system by scaling up to 100 m2. MAAs could be further used as photoprotector and antioxidant compounds in cosmetic products.

The effects of solar UV radiation on mycosporine-like amino acids (MAAs), growth, photosynthetic pigments (Chl a, phycobiliproteins), soluble proteins (SP), and C and N content of Mazzaella laminarioides tetrasporophytes and gametophytes were investigated. Apical segments of tetrasporophytes and gametophytes were exposed to solar radiation under three treatments (PAR [P], PAR+UVA [PA], and PAR+UVA+UVB [PAB]) during 18 d in spring 2009, Punta Arenas, Chile. Samples were taken after 2, 6, 12, and 18 d of solar radiation exposure. Most of the parameters assessed on M. laminarioides were significantly influenced by the radiation treatment, and both gametophytes and tetrasporophytes seemed to respond differently when exposed to high UV radiation. The two main effects promoted by UV radiation were: (i) higher synthesis of MAAs in gametophytes than tetrasporophytes at 2 d, and (ii) a decrease in phycoerythrin, phycocyanin, and SPs, but an increase in MAA content in tetrasporophytes at 6 and 12 d of culture. Despite some changes that were observed in biochemical parameters in both tetrasporophytes and gametophytes of M. laminarioides when exposed to UVB radiation, these changes did not promote deleterious effects that might interfere with the growth in the long term (18 d). The tolerance and resistance of M. laminarioides to higher UV irradiance were expected, as this intertidal species is exposed to variation in solar radiation, especially during low tide.

Barufi J.B., Mata M.T., Oliveira M.C. and Figueroa F.L. 2012. Nitrate reduces the negative effect of UV radiation on photosynthesis and pigmentation in Gracilaria tenuistipitata (Rhodophyta): the photoprotection role of mycosporine-like amino acids. Phycologia 51: 636–648. DOI: 10.2216/10.77.1 Photoprotection of the agarophyte red alga Gracilaria tenuistipitata against ultraviolet radiation (UVR) was investigated in algae submitted for 1 week to photosynthetically active radiation (PAR, 260 μmol photons m−2 s−1) or PAR + UVR (UV-A, 8.13 W m−2 and UV-B, 0.42 W m−2) under different nitrogen concentrations: 0, 0.1, and 0.5 mM of NO3−. Photosynthetic pigments decreased during the time of the experiment mainly under low nitrogen supply and UVR. Incubation under high nitrogen supply (0.5 mM) sustained the photosynthetic levels over time. In contrast, mycosporine-like amino acids (MAAs) increased up to eightfold in the presence of UVR and 0.5 mM NO3−. Under PAR + UVR, maximal quantum yield was positively correla...

The combined effect of high solar ultraviolet radiation (UVR) and nutrient supply in a phytoplankton community of a high mountain lake is analyzed in a in situ experiment for 6 days with 2 × 2 factorial design. Interactive UVR × nutrient effects on structural and functional variables (algal biomass, chlorophyll a (chl a), primary production (PP), maximal electron transport rate (ETR(max)), and alkaline phosphatase activity (APA)), as well as stoichiometric ones (sestonic N per cell and N:P ratio) were found. Under non-nutrient enriched conditions, no deleterious effects of UVR on structural variables, PP, photosynthetic efficiency and ETR(max) were observed, whereas only particulate and total APA were affected by UVR. However, percentage excreted organic carbon (%EOC), dissolved APA and sestonic C and P per cell increased under UVR, leading to a decrease in algal C:P and N:P ratios. After nutrient enrichment, chl a, total algal biomass and PP were negatively affected by UVR whereas %EOC, ETR(max) and internal C, P and N content increased. We suggest that the mechanism of algal acclimation to UVR in this high UVR flux ecosystem seems to be related to the increase of internal algal P-content mediated by physiological mechanisms to save P and by a stimulatory UVR effect on dissolved extracellular APA. The mechanism involved in the unmasking effect of UVR after nutrient-enrichment may be the result of a greater sensitivity to UVR-induced cell damage, making the negative UVR effects more evident.

We have studied the effects of nitrate supply under photosynthetic active radiation (PAR) plus ultraviolet radiation (UVR) exposure on photosynthetic pigments (chlorophyll a and carotenoids), photoprotective UV screen mycosporine-like amino acids (MAAs), and photosynthetic parameters, including the maximum quantum yield (F v/F m) and electron transport rate (ETR) on the red agarophyte Gracilaria tenuistipitata. Apical tips of G. tenuistipitata were cultivated under ten different concentrations of NO 3 − for 7 days. It has been shown that G. tenuistipitata cultured under laboratory conditions has the ability to accumulate high amounts of MAAs following a nitrate concentration-dependent manner under PAR + UVR. Two MAAs were identified, shinorine and porphyra-334. The relative concentration of the first increased under high concentrations of nitrate, while the second one decreased. The presence of antheraxanthin is reported for the first time in this macroalgae, which also contains zeaxanthin, lutein, and β-carotene. The accumulation of pigments, photoprotective compounds, and photosynthetic parameters of G. tenuistipitata is directly related to N availability. All variables decreased under low N supplies and reached constant maximum values with supplements higher than 0.5 mM NO 3 − . Our results suggest a high potential to acclimation and photoprotection against stress factors (including high PAR and UVR) directly related to N availability for G. tenuistipitata.

The effects of increased photosynthetic active radiation (PAR), UV radiation (UVR), and nutrient supply on photosynthetic activity, pigment content, C:N ratio and biomass yield were studied in tank cultivated Gracilaria conferta (Rhodophyta). Electron transport rate (ETR) and biliprotein content were higher under high nutrient supply (HNS), obtained from fishpond effluents, compared to low nutrient supply (LNS), in contrast to mycosporine-like amino acids (MAAs) dynamic. The high MAA content in LNS-algae could be explained by higher UVR penetration in the thallus and by the competition for the use of nutrients with other processes. Effective quantum yield decreased after short-term exposure to high irradiance whereas full recovery in shade was produced only under slightly heat shock. UVA radiation provoked an additional decrease in photosynthesis under high water temperature. UVB radiation reversed UVA's negative effect mainly with HNS. Results support that nutrient-sufficiency help G. conferta to resist environmental changes as short-term temperature increase.

Seaweeds growing in the intertidal zone are exposed to fluctuating nitrate and ultraviolet radiation (UVR) levels. While it has been shown that elevated UVR levels and the decrease of nitrate concentration can reduce photosynthetic levels in seaweeds, less is known about the combined effect of nitrate levels and UVR on metabolism and photoprotection mechanisms of intertidal species. Consequently, the objective of this study was to evaluate the effect of nitrate concentration and UVR treatments on photosynthesis, respiration, nitrate reductase activity and phenolic compound levels of Ulva rigida (Chlorophyta). There was a two- to threefold increase in maximal gross photosynthesis (GPmax) and respiration rates, as nitrate increased from 0 to 50 μM NO 3 − . Similarly, nitrate reductase activity increased linearly from low values in algae incubated at 0 μM NO3 to high values in tissue incubated at 50 μM NO 3 − . Phenolic compounds in the tissue of U. rigida increased approximately 60% under 50 μM NO 3 − relative to those incubated at 0 μM NO 3 − . Algae exposed to UVR (8 h) showed a significant decrease in the effective quantum yield and respiration, however, no effect was observed in the phenolic compounds levels. Full recovery of effective quantum yield was observed after U. rigida was transferred for 48 h to low PAR. Nitrate reductase also decreased after an 8-h UVR exposure, but no differences were observed among the nitrate treatments. This study shows that high nitrate levels reduced the negative effect of UVR on the effective quantum yield and increased the recovery of key metabolic enzymes. It is possible that the increase of phenolic compounds in the thallus of U. rigida under high nitrate levels provide a photoprotective mechanism when exposed to high UV levels during low tides.

The photoprotective and acclimation capacity against ultraviolet radiation (UVR) was assessed for the dinoflagellate Heterocapsa sp. We examined the effect of UVR and N availability on photosynthetic activity and on the accumulation of photoprotective substances such as mycosporine-like amino acids (MAAs) and xanthophyll cycle pigments. Cells were cultivated under two different light treatments, photosynthetically active radiation (PAR) and PAR + UVR, and at two NaN0 3 concentrations, mid nitrogen (MN, 0.1 mmol L ―1 ) and high nitrogen (HN, 1 mmol L ―1 ) for 6 d. MAA and photosynthetic pigment contents as well as maximum quantum yield of fluorescence (F v : F m ) and electron transport rate were analyzed at the initial time and after 3 and 6 d of experimentation. F v : F m decreased because of UVR and N limitation. N enrichment reduced the deleterious effect of UVR on photosynthesis. The content of photosynthetic pigments and MAAs was higher at HN than at MN supply and a positive effect of UVR on MAA and pigment accumulation was observed, suggesting that under HN conditions the deleterious UVR effect is counteracted by MAAs. Under N limitation thermal energy dissipation takes place by the xanthophyll cycle, i.e., an increase of the de-epoxidation degree was observed under these culture conditions. However, UVR seems to favor diadinoxanthin accumulation; thus, no photoprotection through the xanthophyll cycle seems available in cells exposed to PAR + UVR and MN. We conclude that not only MAA accumulation but also N availability is very important to determine the photoprotective capacity against UVR of Heterocapsa sp.

It has been demonstrated that far-red light reduces growth of marine phytoplankton and that light quality controls growth and photosynthetic metabolism in algae. The green halotolerant microalga, Dunaliella bardawil, accumulates high amounts of β-carotene (up to 10% of its dry weight) under conditions of high light or nutrient limitation. The influence of increasing irradiance and of far-red light in D. bardawil was studied. Continuous irradiance was provided by white fluorescent lamps alone (WL) or supplemented with far-red Linestra lamps (WL+FR). For both types of light, cultures were acclimatized at increasing irradiances (50-300 µmol m−2 s−1), and cell density, photosynthetic activity and pigment content were determined. Cell density increased with the photon irradiance, and was higher in WL than in WL+FR under the same irradiance, but the reverse occurred in respect of cell volume. Growth rate was higher under WL+FR. Far-red light induced faster growth but reduced the maximal cell density of the cultures. Chlorophyll a concentration was higher in white light, but total carotenoid content increased dramatically in both far-red light treatments (about 50% on a per cell basis) and with the increase of irradiance. Our results show that far-red light has a significant influence on growth and photosynthesis of D. bardawil, inducing a decrease in cell density, photosynthetic activity and chlorophyll concentration, and an increase in growth rate, cell volume and carotenoid content.

The accumulation of ultraviolet (UV)-screen substances, mycosporine-like amino acids (MAAs), in the tetrasporophytic phase of the red alga Asparagopsis armata grown in fishpond effluents of gilthead sea bream, Sparus aurata, is analyzed. Both the effects of total ammonium nitrogen (TAN) fluxes and the algal densities on MAA accumulation were investigated. MAAs increased with the TAN flux, but only until values lower than 100 μM/h. Above this flux, the MAA content decreased, whereas algal yield increased. The content of individual MAAs was related to nitrogen (N) status, that is, shinorine percentage slightly decreased and palythine increased with increasing N fluxes. The decrease of MAAs at high flux of N (up to 100 μM/h) is related to the decrease of water residence time and the decrease of the biofiltration efficiency. Under different TAN fluxes and algal densities, MAA content was negatively related to algal yield indicating that MAAs were accumulated only under a high ammonium-N availability. Thus, an energy allocation between growth (primary metabolism) and MAA accumulation (secondary metabolism) is regulated by the absorption capacity of inorganic N. In conclusion, A. armata, in addition to its high biofiltration capacity of nutrients, is a good source of MAAs as potential UV screen photoprotectors.

The possible protective role against ultraviolet (UV) radiation of the sheathing leaves that wrap the young flowers of Ruppia drepanensis and Althenia orientalis was assessed. The effects of UV radiation on the viability of their pollen grains and those of R. maritima were also analysed. The absorption of the sheathing leaves of R. depranensis for visible and UV wavelengths was analysed using a spectrophotometer. Pollen grains of these species were exposed at different times to darkness, only photosynthetic active radiation (PAR), PAR+ultraviolet-A radiation (PAR+UV-A) and PAR+UVA+ ultraviolet-B radiation (PAR+UV-A+UV-B), and pollen viability was estimated by means of fluorescein diacetate test. The sheathing leaves presented a maximum absorbance at approximately 320 nm. On the other hand, the highest loss of pollen viability was produced in the presence of UV-B, especially in the case of A. orientalis. Nevertheless, viability of pollen was more affected by UV-A radiation in both Ruppia species than in A. orientalis. The effect of UV-B radiation was higher in the species with underwater pollination than in species with surface water pollination or combined surface -underwater pollination. Ecological implications are discussed.

Foliose Grateloupia species are among the most commonly reported introduced species of red algae in European waters. Unequivocal identification at species level, however, has proven exceedingly difficult owing to a relatively low number of clearcut diagnostic characters and considerable morphological plasticity. Because of their morphological similarity many of these species were placed in synonymy by subsequent generations of phycologists. In the present paper we demonstrate that Gratelopia lanceola, a native European species, is distinct from G. turuturu, a western Pacific alga introduced in European waters. These conclusions are based on comparative gene sequences (rbcL and LSU rDNA) as well as morphology and mycosporine-like amino acid (MAA) signatures. Even though support was largely lacking from rbcL and the combined analysis, G. lanceola was clearly separated from G. turuturu as well as other foliose species. These data, together with earlier accounts on genuine G. doryphora and G. schizophylla, indicate that foliose Grateloupia species display a clear geographic structuring, which has only become obscured by the recent introduction of morphologically similar species (pseudo-cryptic species) along Atlantic coasts and in the Mediterranean Sea. Grateloupia lanceola also showed a different MAA composition when compared with G. turuturu, regardless of spatial-temporal influences. The concentrations were about 3 mg g � 1 DW and about 5 mg g � 1 DW in G. turuturu and G. lanceola, respectively. The main MAA in Grateloupia lanceola was porphyra-334, which represented almost 100% of these compounds. Only some traces (

Seasonal changes in the biochemistry and photophysiology of the brown macroalga Cystoseira tamariscifolia was analyzed in southern Spain. Total carbon and nitrogen contents, phenolic compounds, antioxidant and photosynthetic activities were seasonally determined over two years. Carbon, nitrogen and photoprotective phenolic contents were higher in winter and spring than in summer and autumn. Antioxidant levels were highest in spring and we found a positive correlation between phenolic content and antioxidant activity (EC50). Photosynthetic capacity (ETRmax) and photosynthetic efficiency (αETR) were also highest in spring, and there was a positive correlation between ETRmax and the amount of phenols present. Increased irradiance in spring enhanced algal productivity, antioxidant activity and the production of photoprotective compounds but in summer nutrient depletion due to thermal stratification of coastal waters reduced photosynthetic activity and the photoprotective capacity of C. tamariscifolia. Electron microscopy showed that phenols occurred in the cytoplasm of cortical cells inside physodes. Spring would be the best period to harvest C. tamariscifolia to extract photoprotectors and antioxidants for potential commercial uses, although the environmental impacts would need to be carefully assessed.

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 CO2 seep in oligotrophic waters off Vulcano (Italy). The algae were incubated in situ at 0.2 m depth using a combination of three mean CO2 levels (500, 700–800 and 1200 μatm CO2), 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 CO2 levels benefitted both of these algae, although their responses varied depending on light and nutrient availability. In C. compressa, elevated CO2 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 (Fv/Fm) and photosynthetic efficiency (αETR) without nutrient enrichment. In P. pavonica, elevated CO2 treatments had higher carbon content, Fv/Fm, αETR, 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.

Lanthanides are biologically non-essential elements with wide applications in technology and industry. Their concentration as environmental contaminants is, therefore, increasing. Although non-essential, lanthanides have been proposed (and even used) to produce beneficial effects in plants, even though their mechanisms of action are unclear. Recently, it was suggested that they may replace essential elements. We tested the effect of low concentrations of lanthanides on the common freshwater microalga Desmodesmus quadricauda, grown under conditions of metal ion-deficiency (lower calcium or manganese concentrations). Our goal was to test if lanthanides can replace essential metals in their functions. Physiological stress was recorded by studying growth and photosynthetic activity using a pulse amplitude modulation (PAM) fluorimeter. We found that nutrient stress reduced parameters of growth and photosynthesis, such as maximal quantum yield, relative electron transport rate, photon capturing efficiency and light saturation irradiance. After adding low concentrations of five lanthanides, we confirmed that they can produce a stimulatory effect on microalgae, depending on the nutrient (metal) deprivation. In the case of a calcium deficit, the addition of lanthanides partly alleviated the adverse effects, probably by a partial substitution of the element. In contrast, with manganese deprivation (and at even lower concentrations), lanthanides enhanced the deleterious effect on cellular growth and photosynthetic competence. These results show that lanthanides can replace essential elements, but their effects on microalgae depend on stress and the nutritional state of the microalgae, raising the possibility of environmental impacts at even low concentrations.

Combined and/or interactive effects of inorganic nitrogen (as ammonium) and irradiance on the accumulation of nitrogenous compounds, like UV-absorbing mycosporine-like amino acids (MAAs), chlorophyll a and phycobiliproteins, were examined in the red algaGrateloupia lanceola (J. Agardh) J. Agardh in a high irradiance laboratory exposure and a subsequent recovery period under low light. Also, photosynthetic activity as in vivo chlorophyll fluorescence of photosystem II, i.e. optimum quantum yield (F v/F m), electron transport rate (ETR) and quantum efficiency, were examined. Photosynthetic activity, phycobiliproteins and internal nitrogen content declined during the 3-day PAR (photosynthetically active radiation; 600 mmol s � 1 m � 2 ) and PAR + UVR (ultraviolet radiation; UVB 280–315 nm 0.8 W m � 2 , UVA 315–400 nm 16 W m � 2 ) exposure.

Thalli of the intertidal Phaeophyte Fucus spiralis L. and the subtidal Chlorophyte Ulva olivascens Dangeard were exposed to artificial UV-A, UV-B and photosynthetically active radiation (PAR) by combination of PAR + UV-A + UV-B (PAB), PAR + UV-A (PA) and PAR (P) treatments. UV-A enhanced photosynthesis and stimulated carbonic anhydrase (CA) and nitrate reductase (NR) in F. spiralis whilst PAR only had an inhibitory effect in this species. U. olivascens suffered chronic photoinhibition in all the treatments as evidenced by reduced maxima photosynthesis (Pmax) and photosynthetic efficiency (α). Non stimulatory effect was observed upon CA and NR in this species. Our results showed that artificial UV radiation triggered opposite responses in both species. We suggest that differences shown by both species might be related to their location in the rocky shore and their ability to sense UV. We propose that the ratio UV:PAR acts as an environmental signal involved in the control of photosynthesis as shown by pronounced inhibition in samples exposed to only PAR. We also suggest that UV-regulated photosynthesis would be related to carbon (C) and nitrogen (N) cycles, regulating feedback processes that control C and N assimilation.

The electron transport rate (ETR) versus gross photosynthesis (GPS) relationship varies as a function of species, temperature, irradiance, and inorganic carbon levels, but less is known about the effect of nitrogen supply on this relationship. The objective of this study was to evaluate the effect of nitrate concentration on the ETR versus GPS relationship in Ulva rigida C. Agardh from the Mediterranean Sea. Chlorophyll content and tissue absorptance increased 2-fold as nitrate in the media increased from 0 to 50 μM. Whereas internal N content increases 3-fold at 50μM, internal C increased slightly. Oxygen evolution and ETR, evaluated as in vivo chl fluorescence using pulse amplitude modulated fluorometry, in general saturated at irradiances above 100 μmol photons ·m -2 · s -1 . Both maximum ETR and GPS values increased as nitrate concentration increased. In general, the ETR versus GPS relationship showed a linear response to increasing nitrate with little variance of the data. This relationship, however, became more variable at high irradiances and high nitrate concentrations. The ETR/GPS ratio was close to the theoretical value of 4 at low nitrate concentrations, and the ratio decreased exponentially when nitrate concentration in the media increased. The variations of ETR/GPS under different inorganic nitrogen supply are discussed in terms of the effect of nitrate on the photosynthesis and respiration relationship.

Effects of solar radiation on phenolic compound concentrations and photosynthetic activity, estimated as in vivo chlorophyll fluorescence, in the brown alga Cystoseira tamariscifolia (Hudson) Papenfuss were analyzed in southern Spain from June 1988 to July 2000. Annual and diurnal variations of optimum quantum yield were negatively correlated with incident irradiance. Optimum quantum yield decreased as irradiance increased at noon, and yield values recovered in the afternoon suggesting a dynamic photoinhibition. The annual and daily fluctuations of phenolic compound concentration in the tissue of C. tamariscifolia showed contrasting patterns. There was an annual cycle of phenolic compound concentration in the apical thallus, which was positively correlated with incident irradiance. The increase in phenolic compounds, however, was twofold greater in the first half of the year than the decrease during the second half of the year. In contrast to the annual cycle, there appeared to be a negative correlation between phenolic compound concentration and irradiance in the summer months while no specific relationship was observed in the fall–winter months. Loss of phenolic compounds from the tissue to the surrounding water was increased as irradiation dosage increased. This suggests that the decrease of phenolic compounds during the diurnal cycle might be regulated by the exudation of these compounds at high irradiances in the field. Collectively, our results suggest that, like dynamic photoinhibition, the rapid synthesis and turnover time of phenolic compounds in the tissue of C. tamariscifolia might serve as photoprotective mechanisms against high irradiances.

The effect of different light qualities (white, blue, green, yellow and red light) on photosynthesis, measured as chlorophyll fluorescence, and the accumulation of photosynthetic pigments, proteins and the UV-absorbing mycosporine-like amino acids (MAAs) was studied in the red alga Porphyra leucosticta. Blue light promoted the highest accumulation of nitrogen metabolism derived compounds i.e., MAAs, phycoerythrin and proteins in previously N-starved algae after seven days culture in ammonium enriched medium. Similar results were observed in the culture under white light. In contrast, the lowest photosynthetic capacity i.e., lowest electron transport rate and lowest photosynthetic efficiency as well as the growth rate were found under blue light, while higher values were found in red and white lights. Blue light favored the accumulation of the MAAs porphyra-334, palythine and asterina-330 in P. leucosticta. However, white, green, yellow and red lights favored the accumulation of shinorine. The increase of porphyra-334, palythine and asterina-330 occurred in blue light simultaneous to a decrease in shinorine. The accumulation of MAAs and other nitrogenous compounds in P. leucosticta under blue light could not be attributed to photosynthesis and the action of a non-photosynthetic blue light photoreceptor is suggested. A non-photosynthetic photoreceptor could be also involved in the MAAs interconversion pathways in P. leucosticta.

The photosynthetic performance and pigment content of Gracilaria chilensis Bird, McLachlan, and Oliveira were measured under natural solar radiation in the Quempillen river estuary (southern Chile) in order to assess the short-term acclimation of this species to the current levels of ultraviolet (UV) radiation during midsummer. The effect of the tidal fluctuation was evaluated in algae exposed for 3 days to two light climates in cages suspended at two depths (0.5 and 1.5 m from the bottom). Responses to high solar radiation at noon were also assessed to gain insights into rapid photochemical kinetics and the degree of photoinhibition. Results indicated that G. chilensis is a shade-adapted species with a high content of photosynthetic pigments (Chl a and phycobilins), high quantum efficiency (a), and low light saturating point (Ek) between 60 and 170 Amol m � 2 s � 1 reflecting the high turbidity and marked light attenuation (Kd) of both UV and photosynthetically active radiation (PAR): 1% of the UVB radiation (280–320 nm) reaches 1.8 m depth only during high tide, whereas during ebb tide, UVB can only be detected at 30 cm depth. In the case of PAR, 1% penetration depth was close to 2.7 m during low tide. Under these conditions decreases in rates of photosynthesis at noon did not exceed 23%, even in the presence of UVB. When algae were exposed to full solar radiation at noon for 3 h and then incubated under shaded conditions for recovery, evidence for photoinhibition was observed: in algae exposed to PAR+UVA+UVB, photosynthesis decreased by 16%, while recovery was close to 88% of the control. In treatments deprived of UVB, recovery was close to 100%. These results suggest that G. chilensis is not commonly exposed to high irradiances in the field and that UVB is potentially detrimental for photosynthesis when algae are maintained close to the surface, at least during summer. This was corroborated by the low content of UV-absorbing compounds (mycosporine-like amino acids, or MAAs; between 0.89 and 1.75 mg g DW). D 2004 Elsevier B.V. All rights reserved.