Factors controlling the timing of major spring bloom events in an UK south coast estuary

Factors controlling the timing of major (>10 mg chlorophyll a m⁻³) spring bloom events in the estuarine waters of the Solent, on the south coast of the UK, have been investigated. Winter to summer variations in chlorophyll a concentration together with relevant meteorological and hydrographical data have been analysed for 5 years (1988, 1992, 2001, 2002 and 2003). Mean water column irradiance is demonstrated to be the main factor controlling the timing of the first major spring bloom event, usually dominated by large chain-forming diatoms. When chlorophyll a concentration first exceeds 10 mg m⁻³ in spring (usually in May) the mean water column photosynthetic active radiation (PAR) averaged for one week prior to the sampling date was always >380 W h m⁻² d⁻¹. Prior to the main spring bloom event surface incident radiation and water turbidity combine to limit chlorophyll a concentration to levels <10 mg m⁻³. Chlorophyll a concentrations >10 mg m⁻³ do not occur in the Solent until almost the entire 10 m water column is within the euphotic zone (i.e. above 1% light level) and light extinction coefficient (k) is ca. ≤0.5 m⁻¹. Statistically, river flow explains the largest percentage of the variations in k and the delayed bloom in June 2002 is due to increased cloud cover and high levels of rainfall in May, which caused a reduction in surface incident irradiance and increased turbidity. Chlorophyll a peaks during these major bloom events generally occur on spring tides when increased mixing rates favour net growth of diatoms.     

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.     

Phytoplankton absorption and pigment adaptation of a red tide in the Benguela ecosystem

Phytoplankton absorption and pigment characteristics of a red tide were investigated in coastal waters of the southern Benguela. Diagnostic indices indicated that dinoflagellates were the dominant phytoplankton group, with diatoms and small flagellates being of secondary importance. Very high biomass was observed close to the coast where chlorophyll a concentrations of up to 117 mg m^–3 were measured. Both measured (a _ph) and reconstructed pigment absorption (a _pig) displayed an increasing trend with chlorophyll a, while the package effect index (Q^* _a) decreased, indicating increased packaging with an increase in biomass. Proportioning of the total pigment absorption between 400 and 700 nm revealed that chlorophyll a accounted for 39–65% of the absorption, while photosynthetic carotenoids (15–30%) and chlorophyll cs (15–30%) were also prominent in absorbing light for photosynthesis.     

Diel Patterns in Chlorophyll <Emphasis Type="Italic">a</Emphasis> Specific Absorption Coefficient and Absorption Efficiency Factor of Picoplankton

Diel patterns in the chlorophyll a specific absorption coefficient of surface picoplankton, a*_pico (γ) (m²·[mg chlorophyll a]⁻¹), were studied at 7 stations under daily cycle of in situ light condition in the western subarctic Pacific and Japan Sea. All the data were normalized by dividing the anomaly with daily averaged a*_pico (γ). Opposite diel patterns were observed for the normalized a*_pico (443) and a*_pico (675) with maximum toward dawn or dusk and minimum toward midday at 4 stations under low-irradiance (LI) conditions and vice versa at 3 stations under high-irradiance (HI) conditions. The absorption efficiency factors at red absorption peak, Q _a (675), were determined by reconstruction with intracellular chlorophyll a concentration and cell diameter. The normalized Q _a (675) also showed diel pattern with maximum toward midday and minimum toward dawn or dusk under LI. The diel pattern in a*_pico (675) and Q _a (675) were primarily caused by changes in intracellular chlorophyll a concentration due to photoadaptation under LI. The diel pattern in a*_pico (443) was influenced by pigmentation, as recognized by blue to red ratio [a*_pico (443)/a*_pico (675)] under HI. This study proposed that the opposite diel pattern in a*_pico (γ) might occur for a wide range of algal species. The results presented here have important consequences for the interpretation of diel variations in optical properties observed in the open ocean.     

Environmental influence on phytoplankton production during summer on the KwaZulu-Natal shelf of the Agulhas ecosystem

During February 2010, studies of primary production (PP) and physiology were conducted at five selected sites in the KwaZulu-Natal (KZN) Bight of the Agulhas ecosystem as part of a programme to elucidate the influence of major physical driving forces and nutrient inputs on the structure and functioning of biological communities. These sites were located in the vicinity of the Durban lee eddy, in the midshelf region of the central part of the bight, off the Thukela Mouth, and to the north and south of Richards Bay. At four of the sites, chlorophyll a ranged from 0.10 to 1.44?mg m–3 and integrated PP ranged between 0.35 and 2.58?g C m–² d–1. The highest biomass and PP, which were comparable to those observed in a wind-driven upwelling system, were associated with a diatom community observed at the midshelf site, and varied between 0.26 and 4.27?mg m–³ and 7.22 and 9.89?g C m–² d–1, respectively. Environmental conditions at each of the sites differed substantially and appeared to be influential in initiating and controlling the development and distribution of phytoplankton biomass and production. Phytoplankton adaptation to variable environmental conditions was characterised by a decreased light-limited slope (αB) and increased rate of photosynthesis (P_m ) and light saturation (Ek) with elevated temperatures. The converse (increased α^B and decreased P_m and Ek) was observed as irradiance levels declined. Generalised additive models indicated that irradiance, temperature and biomass were important variables influencing photosynthetic parameters and photosynthetic rates.     

Spatial variability of the primary production and chlorophyll <Emphasis Type="Italic">a</Emphasis> concentration in the drake passage in the austral spring

The spatial distribution of the primary production (PP) and the chlorophyll a concentration (Chl) were investigated during two research cruises in the Drake Passage area in October–November of 2007 and 2008. The algorithm evaluating the integral PP (PP_int) for the water column in this area was developed based on the data on the surface chlorophyll (Chl_s) and the incident solar irradiance obtained in 2004–2008 in the Atlantic Sector of the Southern Ocean. The results obtained both by the experimental and model approaches suggested that the Polar Front (PF) region of the Drake Passage was characterized by low values of both the PP_int (<100 mg C/m² per day) and Chl_s (0.08–0.20 mg/m³) in October–November. Low values of the Chl_s and relatively high phaeophytine a concentrations indicated the winter succession state of the phytoplankton community in the Antarctic Ocean and the southern Polar Frontal Zone (PFZ). The seasonal warming of the surface water layers and the developing pycnocline resulted in a phytoplankton bloom and a Chl_s concentration of more than 1 mg/m³ in mid-November in this area and the Subantarctic waters.     

Phytoplankton production characteristics in the southern Atlantic and the Atlantic sector of the Southern Ocean in the austral summer of 2009–2010

In December–January of 2010 the spatial distribution of the phytoplankton production characteristics was studied along transects in the vicinity of the Greenwich meridian (I) and in the Drake Passage (II). On transect I, the surface chlorophyll a concentration and primary production varied from 0.11 up to 3.57 mg/m³ and from 4.38 up to 37.47 mgC/m³ per day, respectively. The chlorophyll a in the photosynthetic layer and the integrated primary production varied from 10.7 up to 66.1 mg/m² and from 83 to 646 mgC/m² per day, respectively. On transect II in the surface layer, the chlorophyll a concentration changed within the range of 0.09–1.02 mg/m³ and the primary production ranged from 2.08 to 9.49 mgC/m³. The integrated values ranged from 6.32 to 38.29 mg/m² and from 41 to 221 mgC/m² per day, respectively. The moderate means of themaximum quantum yield (F _v/F _m) on transects I and II (0.41 and 0.35, respectively) testify to the low activity of the phytoplankton’s photosynthetic apparatus. The studied water areas in the Southern Ocean differed both in the phytoplankton biomass expressed in the chlorophyll a concentration values and in the conditions of the primary production formation.     

Photoacclimation of the invasive alga Caulerpa racemosa var. cylindracea to depth and daylight patterns and a putative new role for siphonaxanthin

The introduced green alga Caulerpa racemosa var. cylindracea grows along a broad depth range (from very shallow to 60 m depth) in the Mediterranean basin. In the present work, the photoacclimation capacity of this invasive variety was investigated in summer, the season of its maximum spread. Natural populations from the Gulf of Naples (Italy) were analyzed for photoresponse on two scales of light variability: a spatial scale (at three stations along a depth gradient, from 0.3 to 20 m depth) and a temporal scale (on the shallowest meadow, from sunrise to sunset). These responses were studied through pigment analysis (with HPLC), and in situ measurements of photosynthetic parameters (with a Diving‐PAM fluorometer). Electron transport rate (ETR)–irradiance curve parameters showed acclimation along environmental gradients dominated by variation in irradiance. In the shallowest plants, the lack of a midday depression in both the maximum relative ETRs and the photosynthetic efficiency at sub‐saturating irradiance (α) pointed to a maintenance of energy conversion levels despite the protective lowering of light‐harvesting efficiency revealed by the trend in F_v/F_m. On the other side, variation of photosynthetic efficiency occurred with depth and buffered the effect of decreasing light on maximum photosynthetic rates. A previously undescribed xanthophyll cycle centred on lutein‐siphonaxanthin interconversion appeared to operate in the shallowest populations in addition to the violaxanthin/antheraxanthin/zeaxanthin cycle commonly occurring in Chlorophyta; this would further enhance phototolerance of the alga. A further role of siphonaxanthin is in the acclimation to low light of deep environments as indicated by its stronger increase from the surface to the deepest station with respect to siphonein and chlorophyll b.     

The effects of tropical storm Dennis on coastal phytoplankton

The effects of tropical storm Dennis were documented in the coastal waters of South Carolina during August 1981. Phytoplankton photosynthesis vs. irradiance curves showed initial depression of the parameter a followed by three- to five-fold increase of both a and the asymptotic maximum rate of photosynthesis P_m^B. Productivity rates were depressed in most samples immediately after the storm. Surface samples at the inshore stations were around 50 mg C m^?3 h^?1 at saturating light intensities, while the offshore station rates were around 10 mg C m^?3 h^?1. After a 10-day lag these rates had increased to about 200 mg C m^?3 h^?1 inshore and 75 mg C m^?3 h^?1 offshore. These changes are thought to be primarily caused by changes in species composition. Some of the dominant diatom species changed and dinoflagellate species were introduced. No significant changes in nutrient concentrations were observed. Transient depressions of water temperature, salinity and light intensity may have contributed to the observed changes.     

Saldanha Bay,South Africa II: estimating bay productivity

Saldanha Bay is a narrow-mouth bay on the west coast of South Africa linked to the southern Benguela upwelling system. Bay productivity was investigated by use of the conventional light-and-dark bottle oxygen method, and, for comparison, through assimilation of the stable isotope tracer ¹³C. Gross community production GCP and net community production NCP, as determined from the oxygen method, were respectively 2.6 and 2.4 times higher than estimates determined from the stable isotope method. Chlorophyll a (Chl a) concentrations increased with the onset of spring and well-defined subsurface maxima developed in association with increasingly stratified conditions (mean water column Chl a concentrations ranged from 5.4 to 31.5?mg m^?3 [mean 15.5?mg m^?3; SD 7.6]). A sharp decline in photosynthetic rates P* (GCP normalised to Chl a concentration) with depth was attributed to light limitation, as demonstrated by the high vertical attenuation coefficients for downward irradiance K_d, which varied from 0.29 to 0.70?m^?1 (mean 0.48?m^?1; SD 0.12). Productivity maxima were consequently near-surface despite the presence of deeper subsurface biomass maxima. The community compensation depth Z_cc, where gross community production balances respiratory carbon loss for the entire community, ranged from 2.9 to 9.2?m (mean 5.8?m; SD 2.2), and was typically shallower than the 1% light depth for PAR (photosynthetically available radiation), Z1%PAR, which is traditionally assumed to be the depth of the euphotic zone and which ranged from 6.6 to 15.9?m (mean 9?m; SD 2.6). Autotrophic communities, where organic matter is produced in excess of respiratory demand, were confined on average to the upper 5.8?m of the water column, and often excluded the bulk of the phytoplankton community, where light limitation is considered to lead to heterotrophic community metabolism. Estimates of integrated water column productivity ranged from 0.84 to 8.46?g C m–2 d^?1 (mean 3.35?g C m^?2 d^?1; SD 1.9).     

The beam attenuation to chlorophyll ratio: an optical index of phytoplankton physiology in the surface ocean?

The particulate beam attenuation coefficient (c_p) is proportional to the concentration of suspended particles in a size domain overlapping that of the phytoplankton assemblage. c_p is largely insensitive to changes in intracellular chlorophyll concentration, which varies with growth irradiance (a process termed ‘photoacclimation’). Earlier studies have shown that the ratio of c_p:chlorophyll (i.e., c_p^*) exhibits depth-dependent changes that are consistent with photoacclimation. Similar relationships may likewise be expected in the horizontal and temporal dimensions, reflecting changes in mixing depth, incident irradiance, and light attenuation. A link between c_p^* and more robust photoadaptive variables has never been explicitly tested in the field. Here we use five historical field data sets to directly compare spatial and temporal variability in c_p^* with two independent indices of photoacclimation: the light-saturated, chlorophyll-normalized photosynthetic rate, P^b_opt, and the light-saturation index, E_k. For the variety of oceanographic conditions considered, a first-order correlation emerged between c_p^* and P^b_opt or E_k. These simple empirical results suggest that a relationship exists between a bio-optical variable that can potentially be retrieved remotely (c_p^*) and physiological variables crucial for estimating primary productivity in the sea.     

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.     

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.     

Relationships between picophytoplankton and optical properties in the Azores Front region in the Atlantic Ocean

The picophytoplankton evolved to become extremely effective harvesters of light and, thus, dominating productivity in the open ocean. In this study, their distribution in relation to the underwater light field was examined in the Azores Front region of the North Atlantic. In this region, attenuation coefficients of downward irradiance varied between 0.038 and 0.065 m^− 1. Maximum absorptions were at the red and green parts of the light spectrum, typical of the oligotrophic ocean. The euphotic zone ranged from 70 to 120 m. The deep chlorophyll maximum (DCM) was found at depths where subsurface light ranged between 0.1 and 1% of its surface values. Prochlorococcus was the dominant phytoplankton group while Synechococcus (cyanobacteria) and picoeukaryotes were much less abundant. The ability of chlorophyll to absorb light, i.e., the absorption coefficient of chlorophyll (a^⁎), was found to be dependent on the photoacclimation of the cells and was lower at low-light intensities. Due to a packaging effect and probable pigment changes, a^⁎ at the DCM was the lowest while the chlorophyll per Prochlorococcus cell was the highest. This is a major adaptation of the picophytoplankton to low light (less than 1% surface light), which allows them to bloom at the DCM. This study indicates that the Atlantic Ocean models for the estimation of phytoplankton concentration and/or primary production should not use a constant carbon biomass-to-chlorophyll ratio for phytoplankton based on phytoplankton size, and should take into account the variation in chlorophyll-absorption ability.     

High-resolution observations of aggregate flux during a sub-polar North Atlantic spring bloom

An aggregate flux event was observed by ship and by four underwater gliders during the 2008 sub-polar North Atlantic spring bloom experiment (NAB08). At the height of the diatom bloom, aggregates were observed as spikes in measurements of both particulate backscattering coefficient (b_bp) and chlorophyll a fluorescence. Optical sensors on the ship and gliders were cross-calibrated through a series of simultaneous profiles, and b_bp was converted to particulate organic carbon. The aggregates sank as a discrete pulse, with an average sinking rate of ∼75 m d⁻¹; 65% of aggregate backscattering and 90% of chlorophyll fluorescence content was lost between 100 m and 900 m. Mean aggregate organic carbon flux at 100 m in mid-May was estimated at 514 mg C m⁻² d⁻¹, consistent with independent flux estimates. The use of optical spikes observed from gliders provides unprecedented coupled vertical and temporal resolution measurements of an aggregate flux event.     

Recurrent Red-tides in the Southampton Water Estuary Caused by the Phototrophic CiliateMesodinium rubrum

In the Southampton Water estuary (southern England, U.K.), red-tides caused by the planktonic, phototrophic ciliateMesodinium rubrum(=Myrionecta rubra) occur during most summers and sometimes in autumn. These events were investigated in detail between 1985 and 1987 and were characterized by levels of chlorophylla(chl a) of over 100 μg l⁻¹, cell numbers ofM. rubrumof over 1×10³ ml⁻¹, oxygen saturations of around 150%, and depleted numbers of macrozooplankton. Initiation of red-water did not appear to be triggered by irradiance or nutrients, but coincided with an increase in temperature and water column stability. This enhanced stability was promoted by increased surface to bottom gradients of both temperature and salinity, and by reduced mixing during neap tides. Development of red-water was accompanied by removal of most of the dissolved NH⁺₄from the water column, whereas some NO⁻₃persisted, presumably maintained by freshwater input. NO⁻₃and NH⁺₄gradually returned to pre-bloom concentrations as the red-water declined in late summer. Maximal biomass ofM. rubrumappeared to be limited by irradiance, and self-shading probably imposed an upper limit of around 300 mg chl a m⁻²within the water column. At the observed levels of chl a, irradiance values within the population maximum between 1 and 3 m depth were only just of the order (≈15 μmol photons m⁻² s⁻¹) required to balance estimated respiratory demands. Oxygen concentration became undersaturated during the late bloom phase, with minimal values of 20–30% saturation recorded in deeper waters; however, despite this and reduced numbers of macrozooplankton, direct deleterious effects on other organisms were not observed.     

Observation of Algal Bloom in the Northwest Arabian Sea Using Multisensor Remote Sensing Satellite Data

Algal bloom observed using Indian Remote Sensing Satellite IRS-P4 Ocean Color Monitor (OCM) derived chlorophyll images during December 23, 2003–January 8, 2004, off the Oman coast. High chlorophyll concentration (～20 mg m^?3) patches were observed. MODIS-Aqua data of January 1, 2004, were analyzed to generate normalized water leaving radiance (nLw) images for seven visible channels: 412, 443, 488, 531, 551, 667 and 678 nm. The channels 667 and 678 nm showed interesting algal bloom features. The bloom features were detected in OCM image of January 2, 2004, using Subramanian's Trichodesmium detection Protocol. MODIS-Aqua retrieved Sea Surface Temperature (SST) around the bloom patches was observed to be >240°C. The OCM chlorophyll mean observed to be very high (>10.0 mg m^?3) in two bloom pockets. Quickscat scatterometer derived wind speed was found to be optimum in the range of 3–5 m/sec.     

Evidence for the grazing hypothesis: Grazing reduces phytoplankton responses of the HNLC ecosystem to iron enrichment in the western subarctic pacific (SEEDS II)

A mesoscale iron-enrichment study (SEEDS II) was carried out in the western subarctic Pacific in the summer of 2004. The iron patch was traced for 26 days, which included observations of the development and the decline of the bloom by mapping with sulfur hexafluoride. The experiment was conducted at almost the same location and the same season as SEEDS (previous iron-enrichment experiment). However, the results were very different between SEEDS and SEEDS II. A high accumulation of phytoplankton biomass (∼18 mg chl m⁻³) was characteristic of SEEDS. In contrast, in SEEDS II, the surface chlorophyll-a accumulation was lower, 0.8 to 2.48 mg m⁻³, with no prominent diatom bloom. Photosynthetic competence in terms of F _v/F _m for the total phytoplankton community in the surface waters increased after the iron enrichments and returned to the ambient level by day 20. These results suggest that the photosynthetic physiology of the phytoplankton assemblage was improved by the iron enrichments and returned to an iron-stressed condition during the declining phase of the bloom. Pico-phytoplankton (<2 μm) became dominant in the chlorophyll-a size distribution after the bloom. We observed a nitrate drawdown of 3.8 μM in the patch (day 21), but there was no difference in silicic acid concentration between inside and outside the patch. Mesozooplankton (copepod) biomass was three to five times higher during the bloom-development phase in SEEDS II than in SEEDS. The copepod biomass increased exponentially. The grazing rate estimation indicates that the copepod grazing prevented the formation of an extensive diatom bloom, which was observed in SEEDS, and led to the change to a pico-phytoplankton dominated community towards the end of the experiment.     

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.     

Influence of light intensity on diatom physiology and nutrient dynamics in the Oyashio region

The spring diatom bloom characterizes the plankton and nutrient dynamics in the Oyashio region, the westernmost part of the subarctic Pacific. Previous studies have shown that NO₃ was not depleted during the spring bloom, and an increase in the consumption ratio of Si(OH)₄ to NO₃ (ΔSi(OH)₄:ΔNO₃) was observed as the spring bloom progressed. The increase in ΔSi(OH)₄:ΔNO₃ has been suggested to be caused by growth stresses of diatoms, e.g. light limitation by self-shading. In the present study, incubation experiments of sea-surface water from the Oyashio region under saturated irradiance showed that NO₃ was depleted first and ΔSi(OH)₄:ΔNO₃ was more or less constant until the NO₃ depletion occurred. The increase in ΔSi(OH)₄:ΔNO₃ was observed after the NO₃ depletion had occurred in contrast with the field observation. This result of the increase in ΔSi(OH)₄:ΔNO₃ under saturated irradiance after NO₃ depletion suggests that the in situ increase in ΔSi(OH)₄:ΔNO₃ before the NO₃ depletion might be caused by light limitation for diatoms. Responses to a reduction in irradiance were examined using diatom species isolated from the Oyashio region. Variable responses to a reduced irradiance were observed for cell specific C, N, Si and chlorophyll a (Chl) contents. However, the examined diatom species showed similar tendencies for increases in Si:C and Si:N and decreases in C:Chl ratios with the reduction in irradiance. We conclude that light limitation changes the uptake ratio of nutrients and the elemental composition of diatoms and that light limitation is one of the factors influencing the physiology of diatoms and nutrient dynamics in the Oyashio region during the spring bloom.