Patterns in tropical seagrass photosynthesis in relation to light, depth and habitat

Seagrass meadows across north-eastern Australia, survive a range of environmental conditions in coastal bays, reefs, estuarine and deepwater habitats through adaptation of a range of structural, morphological and physiological features. The aim of this study was to investigate the influence of spatial features (habitat type, site and depth) and photon flux on the photosynthetic performance of 11 tropical seagrass species. Pulse amplitude modulated (PAM) fluorometry was used to generate rapid light curves from which measures of maximal electron transport rate (ETR_max), photosynthetic efficiency (α), saturating irradiance (E_k) and effective quantum yield (ΔF/F_m′) were derived. The amount of light absorbed by leaves (absorption factor) was also determined for each population. In intertidal habitats many seagrass species exhibited typical sun-type responses with a close coupling of both ETR_max and E_k with photon flux. Photosynthetic performance ranged from minima in Thalassodendron ciliatum to maxima in Syringodium isoetifolium. The absence of a coupling between photosynthetic performance and photon flux in subtidal populations was most likely due to highly variable light climates and possible light attenuation, and hence the photo-biology of estuarine and deepwater seagrasses exhibited photosynthetic responses indicative of light limitation. In contrast seagrass species from shallow reef and coastal habitats for the most part exhibited light saturation characteristics. Of all the variables examined ETR_max, E_k and ΔF/F_m′ were most responsive to changing light climates and provide reliable physiological indicators of real-time photosynthetic performance of tropical seagrasses under different light conditions.     

Acclimations of macroalgae as reflected in photosynthetic parameters derived from PAM fluorometry, and possible implications for abundance patterns

The presences of the common macroalgae Ulva sp. and Jania rubens vary between seasons along the Israeli Mediterranean intertidal zone. To reveal some of the potential acclimation mechanisms of these algae, we examined their photosynthetic traits during the year using rapid light curves (RLC) derived from pulse amplitude modulated (PAM) fluorometry. In addition, the relationships between those photosynthetic traits and the relative abundances of the two algae were investigated. Ulva sp. showed high maximal electron transport rates (ETR_max) (49.7–68.8 μmol electrons m⁻² s⁻¹) and onsets of light saturation values (E_k) (75.8–85.6 μmol photons m⁻² s⁻¹) in the winter months of December–February, while low values were found in the summer months of June–July (5.2–20.6 μmol electrons m⁻² s⁻¹ and 6.0–23.7 μmol photons m⁻² s⁻¹, respectively). At noon time (during the highest irradiance of the day), the maximal effective quantum yield (Y₀) did not vary significantly during the year. These results indicate that seasonal irradiance influences the number of reaction centres per thallus area. Both algae showed depressions in Y₀ at noon (23.8% for Ulva sp. and 20.3% for Jania rubens), indicating an efficient non‐photochemical quenching mechanism. A positive correlation between the relative abundance and the photosynthetic parameter Y₀ was found for Jania rubens when Y₀ was sampled in the morning or noon, indicating that growth rate and primary production for this alga can be estimated from RLCs. No such correlation was found for Ulva sp., indicating that, in addition to photosynthetic traits, the relative seasonal abundance of this alga is influenced by other factors such as grazing and/or catastrophic events.     

Mesoscale variation in the photophysiology of the reef building coral Pocillopora damicornis along an environmental gradient

Spatial variation in the photophysiology of symbiotic dinoflagellates (zooxanthellae) of the scleractinian coral Pocillopora damicornis was examined along an environmental gradient in the Whitsunday Islands (Great Barrier Reef) at two depths (3 m and 6 m). Chlorophyll a fluorescence of photosystem II (PSII) and PAR-absorptivity measurements were conducted using an Imaging-PAM (pulse-amplitude-modulation) fluorometer. Most photophysiological parameters correlated with changes in environmental conditions quantified by differences in water quality along the gradient. For example, maximum quantum yield (F_v/F_m) increased and PAR-absorptivity decreased as water quality improved along the gradient from nearshore reefs (low irradiance, elevated nutrients and sediments) to outer islands (high irradiance, low nutrients and sediments). For apparent photosynthetic rate (PS_max) and minimum saturating irradiance (E_k), the direction of change differed depending on sampling depth, suggesting that different mechanisms of photo-acclimatisation operated between shallow and deep corals. Deep corals conformed to typical patterns of light/shade acclimatisation whereas shallow corals exhibited reduced PS_max and E_k with improving water quality coinciding with greater heat dissipation (NPQ₂₄₁). Furthermore, deep corals on nearshore reefs exhibited elevated Q₂₄₁ in comparison to outer islands possibly due to effects of sedimentation and/or pollutants rather than irradiance. These results highlight the importance of mesoscale sampling to obtain useful estimates of the variability of photophysiological parameters, particularly if such measures are to be used as bioindicators of the condition of coral reefs.     

Effects of acidified seawater on calcification,photosynthetic efficiencies and the recovery processes from strong light exposure in the coral Stylophora pistillata

The aim of this study was to investigate whether coral photosynthetic efficiencies and recovery processes are affected by CO₂‐driven ocean acidification in symbiont photosynthesis and coral calcification. We investigated the effects of five CO₂ partial pressure (pCO₂) levels in adjusted seawater ranging from 300 μatm (pre‐industrial) to 800 μatm (near‐future) and strong and weak light intensity on maximum photosynthetic efficiency and calcification of a branching coral, Stylophora pistillata, as this species has often been used in rearing experiments to investigate the effects of acidified seawater on calcification and photosynthetic algae of corals. We found that, the photosynthetic efficiencies and recovery patterns under different light conditions did not differ among pCO₂ treatments. Furthermore, calcification of S. pistillata was not affected by acidified seawater under weak or strong light conditions. Our results indicate that the photosynthetic efficiency and calcification of S. pistillata are insensitive to changes in ocean acidity.     

Use of Cell-Specific PAM-Fluorometry to Characterize Host Shading in the Epiphytic Dinoflagellate Gambierdiscus toxicus

Abstract. Cell‐specific fluorescence characteristics were used to characterize the light tolerance of the toxic benthic dinoflagellate Gambierdiscus toxicus. The fluorescence parameter F_v : F_m was measured using pulse amplitude modulation (PAM) fluorometry on individual cells collected from foliose red algae growing in the sub‐tidal margin of South Water Cay, Belize. Samples were collected over several days during sunny and cloudy conditions and compared to samples incubated in situ. The data from individual cells were used to generate both F_v : F_m frequency histograms and averages. Maximum individual cell values of F_v : F_m reached 0.81 in pre‐dawn samples, a value near the theoretical maximum for PAM fluorometry. In field samples from macroalgal hosts, average F_v : F_m values declined only slightly during the day, but cells incubated in bottles under 47 % incident sunlight showed a significant mid‐day depression. In freshly collected samples, near‐maximum F_v : F_m values could be found in individual cells during the entire day; however, the frequency histograms indicated a greater range in F_v : F_m values during the afternoon than in the morning. In contrast, cultures of G. toxicus showed a tight distribution around a mean. Field samples showed a rapid recovery to near‐maximum F_v : F_m within 2 min when assayed using a standardized actinic light series. Similar results were obtained in laboratory cultures of G. toxicus grown at 73 µmol photons · m^‐2 · s^‐1, but not at 383 µmol photons · m^‐2 · s^‐1. These data provide empirical support for suggestions that G. toxicus exploits the three‐dimensional structure of the algal host thallus to minimize light exposure. This strategy permits G. toxicus, a high‐light intolerant species in culture, to thrive in shallow, well‐lit tropical seas. It may also partially explain the observed preference of G. toxicus for complex, foliose macroalgae as hosts.     

Distribution and photo-physiological condition of phytoplankton in the tropical and subtropical North Pacific

To better understand the vertical distribution of phytoplankton in the tropical and subtropical North Pacific, we used fast repetition rate fluorometry to investigate the photo-physiological condition of the phytoplankton assemblage in this region between February and March 2007. Along 155°E, between the equator and 24°N, the peak of fluorescence (F _m), an indication of the deep chlorophyll maximum (DCM), was deeper than the top of the nitracline and occurred at the 2.4 ± 1.3 % (mean ± SD) light depth (relative to 0 m). The photochemical efficiency (F _v/F _m) and effective absorption cross-section of photosystem II (σ_PSII) were low at the surface but increased rapidly at depths between the top of the nitracline (40–138 m) and the DCM (70–158 m), an indication that the photo-physiological condition of the phytoplankton improved below the top of the nitracline. The depth of the maximal F _v/F _m [Z(F _v/F _{m max})] was 18–32 m deeper than the DCM and corresponded to the 0.8 ± 0.2 % light depth. The values of F _v/F _m at the Z(F _v/F _{m max}) were 20 % higher than those at the DCM and averaged 0.48 ± 0.01. These results suggest that the phytoplankton assemblage beneath the DCM had a high potential photosynthetic performance capacity and was growing by using the very low ambient light in this region.     

Photoadaptation of carboxylating enzymes and photosynthesis during a spring bloom

Properties of the light saturation curve of photosynthesis and ribulose-1,5-bisphosphate carboxylase (RuBPC) activity are shown to change qualitatively in a natural population of marine phytoplankton during a spring bloom. Evidence is presented to show that these changes constitute photoadapative responses to increasing irradiance. As irradiance increased during the bloom, both the level of light-saturated photosynthesis (P_m) and the initial slope of the light saturation curve (α = photosynthetic efficiency) increased whether those parameters were normalized to chlorophyll a concentration (P_m^b, α^b) or to cell numbers (P_m^c, α^c). The magnitudes of these changes were such that I_k (= P_m/α, the photoadaptation parameter) did not change, but I_m, the light intensity at which photosynthesis becomes saturated, increased. RuBPC activity, both chlorophyll a (RuBPC^b) and cell number normalized (RuBPC^c), also increased during the bloom. We suggest that these adaptations were achieved by simultaneously increasing the number of photosynthetic units, proportionately decreasing the photosynthetic unit size, and increasing both the concentrations of the enzymes of the dark reactions and possibly also of photosynthetic electron transport components.We also observed diminished levels of photoinhibition in the high light adapted cells late in the bloom and have suggested that this was a consequence of the same suite of physiological changes.In situ carbon fixation per cell increased during the bloom whereas no change occurred in this parameter when normalized to chlorophyll a concentration. Although these photoadaptive responses thus permitted carbon to be fixed in situ more rapidly per cell, at a constant efficiency with respect to investment of energy in the photosynthetic apparatus, they did not result in a change in growth rate. Based on consideratios of the role of time scale in physiological adaptation, however, it is suggested that the observed alterations in photosynthesis with increasing irradiance might permit a cell to more rapidly fill an energy quota for division, possibly an advantage in a mixing environment in which energy is patchily distributed, both spatially and temporalyy.Phosphoenolpyruvate carboxylase activity when normalized to chlorophyll a (PEPC^b) did not change during the bloom while chlorophyll a normalized dark carbon fixation decreased sharply and was quantitatively small compared to PEPC^b. On this basis and considering that RuBPC^b increased during the bloom, it is suggested that, although PEPC may be involved in dark carbon fixation, its most important quantitative role is probably an indirect one in light dependent photosynthesis.We have also considered the relevance of laboratory results on photoadaptation to interpretations of field studies and have suggested that batch culture studies must be treated with caution but that turbidistat and semi-continuous methods provide reasonable simulations of natural conditions.     

Concurrent imaging of chlorophyll fluorescence,Chlorophyll a content and green fluorescent proteins‐like proteins of symbiotic cnidarians

Research on photosynthetic cnidarians has been mainly focused on the symbiosis established between the cnidarian host and its dinoflagellates endosymbionts from genus Symbiodinium. Despite the potential of imaging techniques for assessing the spatial distribution of key parameters of cnidarian photobiology, such as photochemical activity, chlorophyll a content or green fluorescent proteins (GFPs), to our best knowledge, no study has ever attempted to simultaneous map these three features. In this study, we developed a modified imaging pulse amplitude fluorometer by applying excitation light of different wavelengths and selectively detecting short spectral bands through bandpass filters. The imaging system was used to sequentially excite and quantify chlorophyll variable fluorescence (maximum quantum yield of photosystem II, F_v/F_m), Chl a content (normalized difference vegetation index) and relative content of GFPs. The spatial distribution of these photophysiological parameters was mapped both horizontally, across the surface of the soft corals Sarcophyton cf. glaucum and Sinularia flexibilis and the zoanthid Protopalythoa sp., and vertically, throughout a vertical section of S. cf. glaucum. Results showed bleached areas within each individual coral colony and registered photophysiological changes with S. cf. glaucum tissue depth. Analysis of Protopalythoa sp. polyps’ expansion revealed differential surface patterns of NDVI and GFP concentration, and a negative relation between these latter parameters within each polyp. This novel non‐invasive approach allowed a high‐resolution characterization of the spatial relationship between these key parameters through the analysis of image information on a pixel‐by‐pixel basis, which has great potential for investigating the physiological state of symbiotic associations.     

A comparative study of the photosynthetic activity among three temperate seagrass species in Northern Japan

The photosynthetic activity of Zostera marina, Zostera asiatica and Phyllospadix iwatensis shoots from populations of Hokkaido (Northern Japan) was determined using the pulse amplitude modulated (PAM) fluorometer. Several fluorescence parameters were measured as a function of irradiance and leaf age: electron transport rate (ETR), quantum yield, photochemical quenching (qP) and non‐photochemical quenching (NPQ). The leaf age determined by the leaf position in the shoot bundle strongly influenced the photosynthetic activity of Z. marina, Z. asiatica and P. iwatensis. Young leaves had the maximum electron transport rate (Zm: Leaf 1 = 15.7, Leaf 2 = 16.3; Za: Leaf 1 = 13.0, Leaf 2 = 12.2; and Pi: Leaf 1 = 12.5, Leaf 2 = 11.7) and showed higher photoprotection (NPQ) than old leaves. Among the studied seagrass species, Z. marina had the highest photosynthetic activity (ETR_max = 15.3), in accordance with the highest production in the field in comparison with the other two seagrass species. The PAM fluorometry technique showed to be effective in determining intraspecific (among‐leaves) and interspecific (among‐species) variation in seagrass photosynthetic activity.     

Contribution of photosynthetic gains during tidal emersion to production of Zostera capricorni in a North Island,New Zealand estuary

Photosynthetic characteristics of intertidal Zostera capricorni were measured under different tidal conditions in Whangapoua Harbour on the eastern Coromandel Peninsula, New Zealand, and compared with permanently submerged seagrass beds. Photosynthetic characteristics were measured using pulse amplitude modulated (PAM) fluorom‐etry and oxygen (O2) electrode techniques. Gross light saturated photosynthesis measured as oxygen exchange averaged 5.74 and 5.36 mg O₂ g^–1 dry weight (DW) h^–1 and leaf respiration rates averaged 1.22 and 1.38 mg O₂ g^–1 DW h^–1, for intertidal and subtidal plants respectively. Photosynthesis of both intertidal and shallow subtidal plants was light saturated at between 195 and 242 μmol photons m ^–2 s^–1, suggestive of acclimation to a high light environment. Despite the period of exposure at low tide clearly being an important time for photosynthetic gains for intertidal plants, when water clarity was sufficiently high, maximum rates of photosynthesis were also possible when the beds were submerged. If average water clarity was at the clearer end of a range measured for this site (K_d = 0.85 m^–1) then it was calculated that for intertidal seagrass beds growing at mean sea level in Whangapoua, c. 50% of above‐ground production could occur while plants were submerged.     

Phytoplankton production and adaptation in the vicinity of Pemba and Zanzibar islands,Tanzania

Phytoplankton production and physiology were investigated at six selected locations during a research cruise in early October 2007 in Tanzanian coastal waters. The dataset included photosynthesis– irradiance and active fluorescence parameters, phytoplankton absorption coefficients, and pigment concentrations. Primary production was estimated to vary over the range 0.79–1.89 g C m^?2 d^?1. Diagnostic pigments indicated that micro–nanophytoplankton comprised the communities at three stations and nano–picoplankton at the other three stations. At all stations, the populations maximised their photosynthesis in the upper water column under elevated irradiance and low nutrient conditions. Significant photosynthetic activity was also observed at depth under very low light where the communities increased their quantum yield of photochemistry and the proportion of accessory chlorophylls b and c and photosynthetic carotenoids.     

Photosynthesis versus irradiance relationships in the Atlantic sector of the Southern Ocean

Eleven incubation experiments were conducted in the South Atlantic sector of the Southern Ocean to investigate the relationship between new production (ρNO^–₃), regenerated production (ρNH⁺₄), and total carbon production (ρC) as a function of varying light. The results show substantial variability in the photosynthesis–irradiance (P vs E) parameters, with phytoplankton communities at stations that were considered iron (Fe)-limited showing low maximum photosynthetic capacity (P^B_max) and low quantum efficiency of photosynthesis (α^B) for ρNO₃, but high P^B_max and α^B for ρNH₄, with consequently low export efficiency. Results at stations likely relieved of Fe stress (associated with shallow bathymetry and the marginal ice zone) showed the highest rates of P^B_max and α^B for ρNO₃ and ρC. To establish the key factors influencing the variability of the photosynthetic parameters, a principal components analysis was performed on P vs E parameters, using surface temperature, chlorophyll-a concentration, ambient nutrients, and an index for community size structure. Strong covariance between ambient nitrate (NO₃) and α^B for ρNO₃ suggests that Fe and possibly light co-limitation affects the ability of phytoplankton in the region to access the surplus NO₃ reservoir. However, the observed relationships between community structure and the P vs E parameters suggest superior performance by smaller-sized cells, in terms of resource acquisition and Fe limitation, as the probable driver of smaller-celled phytoplankton communities that have reduced photosynthetic efficiency and which require higher light intensities to saturate uptake. A noticeable absence in covariances between chlorophyll-a and α^B, between P^B_max and α^B, and between temperature and α^B may have important implications for primary-production models, although the absence of some expected relationships may be a consequence of the small dataset and low range of variability. However, significant relationships were observed between ambient NO₃ and α^B for ρNO₃, and between the light-saturation parameter E_k for ρNO₃ and the phytoplankton community’s size structure, which imply that Fe and light co-limitation drives access to the surplus NO₃ reservoir and that larger-celled communities are more efficient at fixing NO₃ in low light conditions. Although the mean P^B_max results for ρC were consistent with estimates of global production from satellite chlorophyll measurements, the range of variability was large. These results highlight the need for more-advanced primary-production models that take into account a diverse range of environmental and seasonal drivers of photosynthetic responses.     

Microphytobenthos vertical migratory photoresponse as characterised by light-response curves of surface biomass

The migratory response of intertidal microphytobenthos to changes in irradiance was studied on undisturbed estuarine sediments. Two non-destructive optical techniques were used to trace variations in vivo of surface biomass: PAM fluorometry, for measuring the minimum fluorescence level (F_o); and spectral reflectance analysis, for quantifying the normalized difference vegetation index (NDVI). Following the formation of a dense biofilm at the surface, replicated sediment samples were simultaneously exposed to six different irradiance levels, ranging from 50 to 1500 μmol m⁻² s⁻¹, during a period of 120 min. The migratory photoresponse of the biofilms was characterised by constructing biomass vs. light curves (BLC), relating the accumulation of microalgal biomass after that period (estimated by F_o or NDVI) to the irradiance level incident on the surface. BLCs allow characterising the main features of the migratory photoresponse of intact biofilms. Typical BLC showed a clear biphasic pattern, with an increase in microalgal accumulation under irradiances below 100 μmol m⁻² s⁻¹, maximum values under 100–250 μmol m⁻² s⁻¹, and a gradual decrease of surface biomass under higher irradiances, indicating a strong photophobic downward migratory response. Similar BLC patterns were obtained when measuring F_o or NDVI. The construction of BLCs for biofilms from intertidal sites with distinctive sediment characteristics and diatom taxonomic composition allowed to detected significant differences in the migratory photoresponse. Biofilms from a muddy sediment exhibited considerably larger amplitude in the migratory photoresponse than the biofilms from a sandy mud site, especially under high irradiances. The photophobic migratory response to high light was found to vary among diatom species, particularly in the case of the biofilms from the muddy sediments.     

Diurnal changes of photosynthetic quantum yield in the intertidal macroalga Sargassum thunbergii under simulated tidal emersion conditions

In this study, a three-way factorial experimental design was used to investigate the diurnal changes of photosynthetic activity of the intertidal macroalga Sargassum thunbergii in response to temperature, tidal pattern and desiccation during a simulated diurnal light cycle. The maximum (F_v/F_m) and effective (Φ_PSII) quantum yields of photosystem II (PSII) were estimated by chlorophyll fluorescence using a pulse amplitude modulated fluorometer. Results showed that this species exhibited sun-adapted characteristics, as evidenced by the daily variation of F_v/F_m and Φ_PSII. Both yield values decreased with increasing irradiance towards noon and recovered rapidly in the afternoon suggesting a dynamic photoinhibition. The photosynthetic quantum yield of S. thunbergii thalli varied significantly with temperature, tidal pattern and desiccation. Thalli were more susceptible to light-induced damage at high temperature of 25 °C and showed complete recovery of photosynthetic activity only when exposed to 8 °C. In contrast with the mid-morning low tide period, although there was an initial increase in photosynthetic yield during emersion, thalli showed a greater degree of decline at the end of emersion and remained less able to recover when low tide occurred at mid-afternoon. Short-term air exposure of 2 h did not significantly influence the photosynthesis. However, when exposed to moderate conditions (4 h desiccation at 15 °C or 6 h desiccation at 8 °C), a significant inhibition of photosynthesis was followed by partial or complete recovery upon re-immersion in late afternoon. Only extreme conditions (4 h desiccation at 25 °C or 6 h desiccation at 15 °C or 25 °C) resulted in the complete inhibition, with little indication of recovery until the following morning, implying the occurrence of chronic PSII damage. Based on the magnitude of effect, desiccation was the predominant negative factor affecting the photosynthesis under the simulated daytime irradiance period. These results may explain the distribution pattern of this species in natural habitats, where it is generally restricted to tide pools in the intertidal zone of wave-swept rocky shores which could provide shelter from desiccation stress during low tide.     

Trophic Potential and Photoecology of Endolithic Algae Living within Coral Skeletons

Abstract. The biomass of the endolithic algae Ostreobium quekettii Phyllosiphoniaceae living within skeletons of the scleractinans Mycedium elephantotus and Leptoseris fragilis averages 300 μg protein. cm^-2. This represents approximately 7% of the protein of the zooxanthekie-containing tissue of M. elephantotus and approximately 38% of that of L. fragilis. Oxygen production Pmax_net of 0. querkettii in bare skeletons of M. elephantotus averaged 0.7 μg O₂.cm^-2· h^-1 measured in large skeletal fragments. This amount is approximately 6% of the productivity of the zooxanthellae Symbiodinium microadriaticum living in the same scleractinian species at the same depth Pmax_net 11 μg O₂· Cm^-2· h^-1. Light compensation of O. quekettii - within skeletons - was reached at approximately 10 and saturation at 35 40 μE·m^-2· s^-1. Algae within the M. elephantotus skeletons receive a maximum of 4–6% of the ambient irradiance, which is approximately 0.9 μE · m^-2· s^-1 approximately 0.04% surface irradiance at a depth of 88 m. In L. fragilis at a depth of 145 m, the photon flux decreases to 0.3 μE·m^-2· s^-1, which is less than 0.004% of surface intensity. With increasing depth, the ratio of Chl b to Chl a increased in endolithic algae colonizing L. fragilis, indicating improvement of light harvesting under low light conditions. In free-living O. quekettii cultured at irradiance levels from 0.5–60 μE·m^-2· s^-1, the concentrations of chlorophylls increased and that of siphonein and β-carotene decreased with decreasing photon flux.     

Depth influence on biochemical performance and thallus size of the red alga Gelidium corneum

Recent declines in the foundation species Gelidium corneum (Hudson) J.V. Lamouroux in pristine coastal stretches of the Southeastern Bay of Biscay have been documented in which individuals showed a high bleaching incidence. Among the potential factorsthat may be responsible for these changes in G. corneum abundance, it has been suggested that high levels of irradiance may be imposing stressful environmental conditions for shallow populations. Given that light is reduced exponentially as depth increases, in this investigation we examined the influence of depth on the stress of G. corneum by comparing the performance of several biochemical parameters and thallus size in plants collected at three different depths. The results revealed that plants growing in shallow waters showed lower antioxidant activity, lower concentrations of photosynthetic pigments (Chl‐a, total carotenoids, phycoerythrin), lower internal nitrogen levels (higher C:N ratio) and shorter thallus lengths than those living in deeper waters. Some of these results may be an expression of photoacclimation, but the low antioxidant activity and total carotenoid concentration detected in plants living in shallow waters suggest that the photoprotective mechanisms of G. corneum may have failed to offset photodamage. If so, our findings support previous studies that suggest that recent changes in solar radiation are partially responsible for G. corneum declines. Nevertheless, given that the metabolic performance of macroalgae is determined by the inter‐play of irradiance with other factors, including temperature and nutrient availability, further research is needed to reach a better understanding of the factors causing stress in G. corneum.     

Enhanced carbohydrate production by Southern Ocean phytoplankton in response to in situ iron fertilization

Storage carbohydrates (e.g., water-extractable β-1,3-d-glucan in diatoms) are of key importance for phytoplankton growth in a variable light climate, because they facilitate continued growth of the cells in darkness by providing energy and carbon skeletons for protein synthesis. Here, we tested the hypothesis that synthesis of storage carbohydrates by phytoplankton in the Southern Ocean is reduced by low iron and light availability. During the EisenEx/CARbon dioxide Uptake by the Southern Ocean (CARUSO) in situ iron enrichment experiment in the Atlantic sector of the Southern Ocean in November 2000, we studied the dynamics of water-extractable carbohydrates in the particulate fraction over the period of 3 weeks following the iron release. The areal amount (integral between 0- and 100-m depth) of carbohydrates increased from 1400 to 2300 mg m⁻² inside the iron-enriched patch, while remaining roughly constant in the surrounding waters. Most of the increase inside the patch was associated with the fraction of large (>10 μm) phytoplankton cells, consistent with the shift in the community structure towards larger diatoms. Deck incubations at 60% of the ambient irradiance revealed that the diurnal chlorophyll a (Chl a)-specific production rates of water-extractable polysaccharides were significantly higher for “in-patch” than for “out-patch” samples (0.5 vs. 0.3 μg C [μg Chl a]⁻¹ h⁻¹, respectively). Together with the higher photochemical efficiency of photosystem II (F_v/F_m), this indicates enhanced photosynthetic performance in response to iron fertilization. In addition, the nocturnal polysaccharide consumption rates were also enhanced by iron release, causing a striking increase in the diel dynamics of polysaccharide concentration. An iron-stimulated increase in diel dynamics was also observed in the fluorescence and size of pico- and nanophytoplankton cells (measured by flow cytometry) and is indicative of enhanced phytoplankton growth. Diurnal polysaccharide production by phytoplankton inside the patch was light-limited when they were incubated at intensities below ca. 200 μmol m⁻² s⁻¹ (daytime average). These irradiance levels correspond to those at 20- to 30-m depth in situ, whereas the upper mixed layer was frequently several-fold deeper due to storms. Therefore, these first measurements of phytoplankton carbohydrates during an in situ iron release experiment have revealed that both light and iron availability are the key factors controlling the synthesis of storage carbohydrates in phytoplankton and, hence, the development of diatom blooms in the Southern Ocean.     

Response of the soft coral Heteroxenia fuscescens to ultraviolet radiation regimes as reflected by mycosporine-like amino acid biosynthesis

The variation in mycosporine‐like amino acid (MAA) concentration in the soft coral Heteroxenia fuscescens in relation to changes in ultraviolet radiation (UVR) regimes was investigated at the Gulf of Eilat, northern Red Sea. Solar radiation (300–700 nm) was measured for different depths and seasons. The UVR irradiance was measured to a depth of 25 m on the reef. The mean attenuation coefficient for UV‐B measured in winter was twofold that of the summer value. Separation of H. fuscescens extracts by reverse‐phase isocratic high‐performance liquid chromatography revealed a single MAA compound, palythine (λ_max = 320 nm). Possible seasonal changes in MAAs in colonies of H. fuscescens along a depth gradient were examined on different dates. Palythine concentrations in the colonies were significantly higher in summer than in the other seasons particularly in shallow water. Possible changes in MAA content in colonies of H. fuscescens as a result of UVR protection, were determined by experiments conducted for periods of 1 week, 1 month and 3 months, at a depth of 5 m. In these experiments colonies were removed from the natural substrate and placed underwater, protected from UVR by a PVC filter. Significant differences between UV‐exposed and protected colonies of H. fuscescens were found only in the 3‐month experiment conducted during the summer. These findings demonstrate that UVR is an important environmental factor regulating MAA biosynthesis in the soft coral H. fuscescens.     

Growth and photosynthetic rates of Chlamydomonas plethora and Nitzschia frustula cultures isolated from Kuwait Bay, Arabian Gulf, and their potential as live algal food for tropical mariculture

With a view to utilize local algae as food organisms in mariculture, the green alga Chlamydomonas plethora and the diatom Nitzschia frustula were isolated and studied from the unique coastal waters arid zone off Kuwait. Batch cultures of C. plethora and N. frustula had maximum division rates (μ_max) of 2.5 and 3.4 day^?1, respectively. Cultures grown for 36 h yielded comparable growth rates. Photosynthesis‐irradiance relationships in cultures harvested at various phases of growth showed that cultures attained log phase after 1‐day growth and yielded the highest assimilation numbers (P: μg C [μg Chl a]^?1 h^?1): 22.8 for C. plethora and 18.1 for N. frustula. Their initial slopes (α^B: ng C [μg Chl a]^?1 h^?1 [μmol m^?2 s^?1]^?1 were also the highest observed so far: 79.5 for C. plethora and 39.6 for N. frustula. Photoinhibition was low. Compared with these, assimilation numbers in senescent cultures of 20‐day growth were <8% of the maximum for both species, and the initial slope decreased to 17 and 13% in C. plethora and N. frustula, respectively. Two polyunsaturated fatty acids (20:5n‐3 eicosapentaenoic acid and 22:6n‐3 docosahexaenoic acid) that are essential in many marine animal diets constituted up to 24% and 1.9% of the total fatty acids, respectively. Of the two algae, N. frustula contained higher levels of 16:1n‐7, 20:5n‐3 and comparatively low levels of 22:6n‐3 fatty acids. Due to its rapid growth, high photosynthetic rate and presence of the amino acids leucine, lysine, glutamic acid and arginine N. frustula has good potential as a feed organism in mariculture applications. The capacity of these algae to produce a wide range of size groups (nano and net plankton) is an additional bonus for mariculture operations.     

Surface layer variability in the Ross Sea,Antarctica as assessed by in situ fluorescence measurements

Phytoplankton fluorescence, temperature and salinity were measured from December through February using in situ instruments deployed at two locations in the southern Ross Sea, Antarctica during the austral summers of three consecutive years (2003–2004, 2004–2005, and 2005–2006) to assess the short-term, seasonal and interannual variations in phytoplankton biomass and oceanographic conditions. The seasonal climatologies of physical forcing variables were also determined from satellite measurements, and the data from the two sites compared to the 2000–2009 mean. In situ fluorometers were deployed at three depths at 77°S, 172.7°E and 77.5°S, 180°. Significant differences between the two sites were consistently observed, confirming the anticipated high level of spatial and temporal heterogeneity. Chlorophyll fluorescence was maximal in late December, and generally decreased rapidly to modest levels in January and February. However, during 1 year (2003–2004) a secondary bloom was found, with summer maxima being similar to those observed during spring. Fluorescence displayed a strong diel cycle, with strong quenching during periods of maximum irradiance. The magnitude of this reduction was large (the minimum average fluorescence was 25% of the daily mean) and decreased with depth. Fluorescence varied interannually, with the absolute levels and temporal patterns being different among years. The two sites had different temperature/salinity properties as measured at 24 m, and both variables changed with time. During 2004–2005 we were able to continuously measure the photosynthetic quantum efficiency of PSII (F_v/F_m) at 11 m, which revealed a minimum in December, and an increase in January, whereas the absolute fluorescence (F_o) decreased simultaneously. We suggest that this reflected a mixing event, whereby available irradiance increased, allowing a short period of growth in a more favorable optical environment. While substantial variations from the mean physical forcing were observed, the linkage of these physical variations with fluorescence was not always clear. Short-term (over 24-h) changes in fluorescence occurred, and were likely related to advective events. Wind events altered fluorescence in the surface layer, and these redistributed phytoplankton in the surface. The variability in chlorophyll fluorescence and physical forcing over a variety of scales in the Ross Sea provides insights into temporal–spatial coupling of phytoplankton.