Size-Fractionated Primary Production Estimated by a Two-Phytoplankton Community Model Applicable to Ocean Color Remote Sensing

In order to estimate primary production from ocean color satellite data using the Vertical Generalized Production Model (VGPM; Behrenfeld and Falkowski, 1997), we propose a two-phytoplankton community model. This model is based on the two assumptions that changes in chlorophyll concentration result from changes of large-sized phytoplankton abundance, and chlorophyll specific productivity of phytoplankton tends to be inversely proportional to phytoplankton size. Based on the analysis of primary production data, P _opt ^B , which was one parameter in the VGPM, was modeled as a function of sea surface temperature and sea surface chlorophyll concentration. The two-phytoplankton community model incorporated into the VGPM gave good estimates in a relatively high productive area. Size-fractionated primary production was estimated by the two-phytoplankton community model, and P _opt ^B of small-sized phytoplankton was 4.5 times that of large-sized phytoplankton. This result fell into the ranges observed during field studies.     

Optimal primary production model and parameterization in the eastern East China Sea

Integrated primary production (IPP) datasets from the eastern East China Sea (ECS) were used to validate the depth dependent and time integrated model (TIM), the empirical model (EM), and the vertically generalized production model (VGPM). Employing constant maximum chlorophyll-a (Chl-a) specific carbon fixation rate (P _opt ^B ), the original TIM could not explain measured IPP variance, while the low P _opt ^B estimated using P _opt ^B models originally embedded in the EM were responsible for original EM low estimations. The disparity between P _opt ^B expressions originally embedded in the VGPM and that observed in this study was responsible for the low accuracy of the original VGPM. After locally tuning on the P _opt ^B model, TIM and VGPM could predict well, whereas EM still gave low estimates. This was probably due to the fact that, unlike TIM and VGPM, EM employs a constant factor instead of a light dependent function in describing the depth of light-saturation primary production. The high accuracy of TIM was limited by the complexity of estimating many variable inputs. EM, as a simplest model, should be used with caution in the eastern ECS. Global VGPM with locally adjusted P _opt ^B seemed to be an effective model for estimating IPP in the eastern ECS.     

深圳沿岸海域净初级生产力的遥感反演及其时空变化分析

以深圳沿岸海域为研究区,以MODIS/AQUA卫星遥感产品为数据源,结合实测浮标数据修正了VGPM中叶绿素a含量的估算进而分析深圳沿岸海域净初级生产力的时空分布规律.研究表明:(1)深圳沿岸海域2014年2、5、8、10月的净初级生产力在空间分布上从近海向外逐渐降低,初级生产力整体呈现出"西高东低"的局面,且未有明显的季节性波动.(2)4个海区的叶绿素a含量均表现为夏季最高秋冬季次之,但各海区主要影响因素不同,珠江口主要受季风造成的浮游植物种类与细胞密度的季节变化影响,大亚湾主要受营养盐限制,大鹏湾的主导因素为湾内余流的季节变化,深圳湾的叶绿素a含量主要与浮游植物细胞密度的季节变化有关.(3)珠江口的初级生产力春夏季高于秋冬季;大鹏湾的初级生产力夏季最高,且季节变化趋势与叶绿素a表现一致;深圳湾的初级生产力夏季最高,且季节变化趋势与海表温度表现一致;大亚湾的初级生产力波动明显,夏冬季海洋初级生产力数值总体高于春秋季.     

Saldanha Bay,South Africa I: the use of ocean colour remote sensing to assess phytoplankton biomass

The efficacy of ocean colour remote sensing in assessing the variability of phytoplankton biomass within Saldanha Bay is examined. Satellite estimates of chlorophyll a (Chl a) were obtained using the maximum peak-height (MPH) algorithm on full-resolution (300?m) data from the Medium Resolution Imaging Spectrometer (MERIS). Subsurface Chl a maxima often occur within Saldanha Bay below the mean detection depth of the satellite (1.5?m) during periods of thermal stratification. Consequently, the MPH product was poorly correlated to in situ data from 4?m depth (r² and average relative percentage difference [RPD] of 0.094 and 53% respectively); however, the coefficient of determination was much improved if limited to in situ data collected under conditions of mixing (r² and RPD of 0.869 and 89%, respectively). Composites of monthly MPH Chl a data reveal mean concentrations consistent with in situ seasonal trends of phytoplankton biomass, confirming the capability of the MPH algorithm to qualitatively resolve surface Chl a distribution within the bay.     

Temporal variability in physicochemical properties,phytoplankton standing crop and primary production for 7 years (2002–2008) in the neritic area of Sagami Bay,Japan

Seasonal and interannual variations in physicochemical properties (i.e., temperature, salinity, dissolved oxygen and dissolved inorganic nutrients), chlorophyll a (Chl-a), particulate carbon and nitrogen (PC and PN, respectively), and primary production were investigated in the neritic area of Sagami Bay, Kanagawa, Japan, from January 2002 to December 2008. These abiotic/biotic variables, except for NH₄ ⁺–N, repeated similar seasonal variations for all 7 years. On the basis of the analysis of data obtained on 167 sampling dates, depth-integrated primary production in this water can be easily estimated from Chl-a at the surface using the regression equations obtained in the present study. Intermittently high values of dissolved inorganic nutrients, Chl-a, PC, PN and primary productivity at the surface during the summer stratified period were induced by high freshwater discharge from the rivers after rainfalls and by the expansion of nutrient-rich Tokyo Bay Water. Temperature, salinity and dissolved inorganic nutrients showed drastic variations within a scale of a few days and/or weeks, and these variations were related to sea levels that represent the intrusion of the Kuroshio Water, Intermediate Oyashio Water or deep water from the continental slope. However, there was no consistent trend in the variations in Chl-a, PC, PN and primary production due to the complex effects of these waters.     

Phytoplankton production in two east coast estuaries: Photosynthesis-light functions and patterns of carbon assimilation in Chesapeake and Delaware Bays

Chesapeake Bay is a large and productive estuary that has received close scrutiny in recent years because of indications that its water quality and biota have been damaged by man's activities. Data on primary production for the estuary as a whole, however, are surprisingly sparse. We describe here the distribution of photosynthetic carbon assimilation by phytoplankton in Chesapeake Bay, and relate productivity patterns to hydrographic characteristics of the estuary. Between March 1982 and April 1983, a series of four cruises was conducted on Chesapeake Bay, and two cruises on the urbanized Delaware Bay for comparison. The upper Chesapeake and Delaware were highly turbid with high concentrations of suspended particulate matter and dissolved inorganic nutrients. Low chlorophyll concentrations were usually found in these areas of high turbidity, despite the abundance of nutrients, suggesting light limitation. Application of Wofsy's (1983) model of phytoplanton growth confirmed this suggestion. Chlorophyll and productivity maxima usually occurred seaward of the turbidity maxima where light penetration increased and suffient nutrients were present to support active phytoplankton growth. Further seaward of the chlorophyll maxima in the Chesapeake, the photic zone depth increased, concentrations of nutrients decreased, and phytoplankton biomass decreased, suggesting that nutrient availability, rather than light, controlled phytoplankton growth in the lower portion of the estuary. In contrast to the Chesapeake, Delaware Bay was more turbid, had generally higher nutrient concentrations, and was lower in phytoplankton productivity. The chlorophyll maxima and region of rapid phytoplankton growth occurred further toward the lower estuary and shelf regions in Delaware Bay because the high turbidity extended further seaward. Nutrients were never depleted at the shelf end of the estuary sufficiently to retard phytoplankton growth. Photosynthesis-irradiance (P-I) curves from simulated in situ and constant intensity incubations showed a strong correlation of the light-limited slope (a^B) with the light-saturated rate ( ) on each cruise. Spatial variations in corresponded to patterns of phytoplankton abundance, as did integral production (PP) and carbon-based growth rates (μ_C, μ_m), and photosynthetic parameters varied significantly with temperature.     

Mesoscale features and phytoplankton biomass at the GoodHope line in the Southern Ocean during austral summer

Two sets of high-resolution subsurface hydrographic and underway surface chlorophyll a (Chl a) measurements are used, in conjunction with satellite remotely sensed data, to investigate the upper layer oceanography (mesoscale features and mixed layer depth variability) and phytoplankton biomass at the GoodHope line south of Africa, during the 2010–2011 austral summer. The link between physical parameters of the upper ocean, specifically frontal activity, to the spatially varying in situ and satellite measurements of Chl a concentrations is investigated. The observations provide evidence to show that the fronts act to both enhance phytoplankton biomass as well as to delimit regions of similar chlorophyll concentrations, although the front–chlorophyll relationships become obscure towards the end of the growing season due to bloom advection and ‘patchy’ Chl a behaviour. Satellite ocean colour measurements are compared to in situ chlorophyll measurements to assess the disparity between the two sampling techniques. The scientific value of the time-series of oceanographic observations collected at the GoodHope line between 2004 to present is being realised. Continued efforts in this programme are essential to better understand both the physical and biogeochemical dynamics of the upper ocean in the Atlantic sector of the Southern 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.     

Factors influencing the magnitude of phytoplankton primary production in a high-energy surf zone

The main factors influencing phytoplankton primary production in the surf zone of the Sundays River Beach, Algoa Bay have been characterized. These factors include cell concentration, chlorophyll concentration, irradiance, temperature and salinity. Good relationships have been obtained between cell concentration, chlorophyll concentration and primary production. The P-I curves showed dependence on temperature with a linear regression between temperature and I_k values. Light saturation was shown to occur between 300 and 510 μmol m⁻² s⁻¹ at normal field temperatures. T_max and T_min were found to be 34°C and 0°C, respectively; P_max was 25°C. Salinity had a marked effect on primary production with S_max occurring at 60 ppt and an extrapolated S_min at 0 ppt. P_max was found to occur at 30 ppt.     

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.     

Identification and classification of vertical chlorophyll patterns in the Benguela upwelling system and Angola-Benguela front using an artificial neural network

Information on the vertical chlorophyll structure in the ocean is important for estimating integrated chlorophyll a and primary production from satellite. For this study, vertical chlorophyll profiles from the Benguela upwelling system and the Angola-Benguela front were collected in winter to identify characteristic profiles. A shifted Gaussian model was fitted to each profile to estimate four parameters that defined the shape of the curve: the background chlorophyll concentration (B ₀), the height parameter of the peak (h), the width of the peak (σ) and the depth of the chlorophyll peak (z_m ). A type of artificial neural network called a self-organizing map (SOM) was then used on these four parameters to identify characteristic profiles. The analysis identified a continuum of chlorophyll patterns, from those with large surface peaks (>10 mg m^?3) to those with smaller near-surface peaks (<2 mg m^?3). The frequency of occurrence of each chlorophyll pattern identified by the SOM showed that the most frequent pattern (～12%) had a near-surface peak and the least frequent pattern (～2%) had a large surface peak. These characteristic profile shapes were then related to pertinent environmental variables such as sea surface temperature, surface chlorophyll, mixed layer depth and euphotic depth. Partitioning the SOM output map into environmental categories showed large peaks of surface chlorophyll dominating in water with cool temperature, high surface chlorophyll concentration and shallow mixed layer and euphotic depth. By contrast, smaller peaks of subsurface chlorophyll were in water with warmer temperature, lower surface chlorophyll concentration, intermediate mixed layer and deep euphotic depth. These relationships can be used semi-quantitatively to predict profile shape under different environmental conditions. The SOM analysis highlighted the large variability in shape of vertical chlorophyll profiles in the Benguela. This suggests that an ideal typical chlorophyll profile, as used in the framework of biogeochemical provinces, may not be applicable to this dynamic upwelling system.     

Photosynthesis-irradiance relationship of phytoplankton and primary production in the vicinity of Kuroshio warm core ring in spring

The light-saturated maximum value (P ^B _max) and initial slope (α) of the photosynthesis-irradiance (P-E) curve were examined in a warm streamer, a cold streamer and a warm core ring off the Sanriku area in the subarctic western North Pacific Ocean during an ADEOS/OCTS Sanriku field campaign in early May 1997. BothP ^B _max and α were within the ranges of temperate populations. A regional difference was apparent inP ^B _max: populations in the warm streamer tended to show higher value ranging between 1.92 and 4.74 mgC (mgChla)⁻¹h⁻¹ than those in the cold streamer and the warm core ring (1.35–2.87 mgC (mgChla)⁻¹h⁻¹). A depth variation was also observed in α in both the warm streamer and the warm core ring: shallow populations tended to have lower α than deep populations. The depth variations in bothP ^B _max and α resulted in a lower light intensity of the light saturation in a deeper population than that of a shallower one. These depth-related variations in the P-E parameters were likely a manifestation of “shade-adaptation” of photosynthesis. Photoinhibition was not observed over in situ surface light intensity varying below ca 1600 μmol photon m⁻²s⁻¹. Water-column primary productivity was biooptically estimated to be 233 to 949 mgC m⁻²d⁻¹ using vertical distributions of the P-E parameters, chlorophylla, phytoplankton light absorption and underwater irradiance. Applicability of surface data sets for estimation of water-column productivity is discussed.     

On the interest of using field primary production data to calibrate phytoplankton rate processes in ecosystem models

In many ecosystem models based on empirical formulations, parameters generally are calibrated in order to achieve the best fit between measured and simulated chlorophyll a standing stocks. An accurate calibration of rate processes as primary production rarely is taken into account. In this paper, we test the usefulness of calibration of phytoplankton photosynthetic processes in an ecosystem model using field primary production data. We used 18 months of photosynthetic process data from the Baie des Veys ecosystem (Normandy, France). Five empirical formulations of photosynthesis–irradiance curve models amongst the most widely used were tested. In each formulation, the variability of photosynthetic parameters (i.e. the light-saturated rate of photosynthesis (P_max^B) and the initial slope of the photosynthesis–light curve (α^B)) was considered depending on environmental factors (temperature and nutrient availability). The fit of the five equations as well as the calibration of parameters on field measurements (i.e. the light-saturated rate of photosynthesis (P_ref^B), the initial slope of the photosynthesis–light curve (α_ref^B), the half-saturation constant for nitrogen (K_N) and silicates uptake (K_Si), and the coefficient in the exponential thermal effect (K_T)) was performed using the whole available data set of P vs. E curves (n = 143, P vs. E curves). Then, the Smith formulation allowing the best simulation of the Baie des Veys primary production and corresponding parameters were introduced in an ecosystem box model. This formulation led directly to a satisfactory representation of the Baie des Veys phytoplankton dynamics without additional calibration. Results obtained were compared with a more classical approach in which ecosystem models were calibrated using published values of parameters. This comparison showed that for the two years studied, annual primary production estimated through the ecosystem model was 13% and 26% higher with our approach than with the more classical approach. This work emphasizes the importance of accurately representing rate processes in ecosystem models in order to adequately simulate production as well as standing stocks.     

Carbon production and export from Biscayne Bay,Florida. I. Temporal patterns in primary production,seston and zooplankton

Five stations along a transect from the western shore of Biscayne Bay, Florida to the Florida Current were sampled monthly for one year. The variability and amount of seston particulate organic carbon, adenosine triphosphate, chlorophyll a, primary production and zooplankton decreased along the seaward transect. The greater inshore biomass and variability of seston were the result of the allochthonous input of detritus and inorganic nutrients via terrestrial runoff. Annual primary production in this subtropical coastal lagoon ranged from 13 to 46 g C m^?2 yr^?1. Chlorophyll a in the bay ranged from 1 to 3 mg chlorophyll a m^?2. In contrast, chlorophyll a in the surface centimetre of the sediment ranged from 50 to 300 mg chlorophyll a m^?2. In this clear, shallow (2 to 3 m), oligotrophic lagoon, over 90% of total primary production is by submerged macrophytes and benthic algae. The high zooplankton biomass in the bay is most likely sustained by macrophyte detritus and the resuspension of benthic diatoms by the high winds associated with summer squalls and winter cold fronts.     

Effects of salinity downshock on dimethylsulfide production

During time-series observations in Sagami Bay, Japan, the concentration of dissolved dimethylsulfoniopropionate (DMSP_d), a precursor of dimethylsulfide (DMS), was negatively correlated with salinity. In the laboratory, low-salinity shock reduced DMS production rates of the natural bacterial community and induced rapid DMSP release from a dinophyte, Heterocapsa triquetra, suggesting that low-salinity shock reduced DMSP_d consumption but enhanced DMSP_d production, which agrees with the negative correlation between DMSP_d and salinity observed in Sagami bay. In addition, low-salinity shock did not affect DMSP lyase activity of H. triquetra. Low-salinity shock would increase the contribution from algae in DMS production, leading to an increase in potential DMS productivity in the environment.     

Distribution and seasonal variation of vitamin B12, thiamine and biotin in the sea

The distributions and seasonal variations of vitamin B₁₂, thiamine and biotin were investigated in waters of the North Pacific Ocean, the East China Sea, and the bays and inlets along the Pacific coast of Japan, by use of microbiological assay methods. A marine diatom Cyclotella nana clone 3H for vitamin B₁₂, and marine yeast Cryptococcus albidus for thiamine, and a marine bacterium Achromobacter sp. strain yH-51 for biotin were used as assay organisms. In the surface water of Sagami Bay, monthly changes in the amounts of the three dissolved vitamins followed closely that of chlorophyll a, being highest in July and lowest during late autumn and winter. The geographical and vertical distributions of thiamine and biotin in the sea generally showed similar patterns to that of chlorophyll a, whereas vitamin B₁₂ did not always follow the trend. Amounts of particulate thiamine and biotin corresponded to about 1 % of that of the dissolved form in the surface water of the North Pacific Ocean. In the coastal waters, however, they were at times found to be 144.0 % and 53.9 % respectively.     

Estimation of standing stock of chlorophylla and primary production from remote-sensed ocean color in the Oyashio region, the Western subarctic Pacific, during the spring bloom in 1997

In order to examine the applicability of remotely-sensed ocean color for the estimation of phytoplankton biomass and primary production in the Oyashio region, the western subarctic Pacific, vertical distributions of chlorophylla concentration and primary production were observed in April and May 1997. Spring bloom was observed in both April and May, and the surface concentration of chlorophylla exceeded 40 mg m⁻³. The relationship between the standing stocks of chlorophylla within the layer from the sea surface to one optical depth (0–1/k layer) and the surface chlorophylla concentration is expressed as a Michaelis-Menten equation. The mean ratio of the standing stock of chlorophylla in the euphotic layer to that in the 0–1/k layer was 4.41, this ratio did not significantly differ from 4.61 which was obtained at homogeneous distribution of chlorophylla within the euphotic layer. These facts suggest that the distribution of chlorophylla could be assumed to be homogeneous in the euphotic layer during the spring bloom. Results of primary production measurements by simulatedin situ method were compared with those by an algorithm with two variables; chlorphylla and non-spectral PAR. Daily primary production in the euphotic layer estimated by the algorithm varied in a range of 38–274% of that estimated by incubation, although the primary productions by the algorithm agreed with those by the incubation at a half of stations. Primary production within the euphotic layer calculated using simply the surface data was the same as that estimated using vertical distribution of chlorophylla. These results show that the primary production in the euphotic layer may be estimated from the remote sensed measurements during the spring bloom in the Oyashio region.     

Physical drivers of phytoplankton production in the southern Benguela upwelling system

Investigations of primary production (PP) were undertaken in the southern Benguela ecosystem during two research surveys in October 2006 and May 2007. Significant differences in environmental conditions, as well as biomass and PP, were observed between October and May. During October, integrated biomass and PP were significantly higher, ranging from 20.43 to 355.01 mg m⁻², and 0.71 to 6.98 g C m⁻² d⁻¹, respectively, than in May, where the range was 47.92–141.79 mg m⁻², and 0.70–3.35 g C m⁻² d⁻¹, respectively. Distribution patterns indicated low biomass and PP in newly upwelled water along the coast, higher biomass and PP in the mid-shelf region, while lower values were observed at and beyond the shelf edge. Latitudinal variations showed consistently higher biomass and PP in the St. Helena Bay region compared to biomass and PP south of Cape Town. During both surveys, phytoplankton communities were comprised primarily of diatoms and small flagellates, with no significant differences. Phytoplankton adaptation to environmental variability was characterised by increased P_m^B and E_k under elevated temperatures and irradiance, while no clear relationships were evident for α^B. Generalised Additive Models (GAMs) showed that photosynthetic parameters were all significant predictors of photosynthesis rates (P_z), with P_m^B being the most important, accounting for 36.97% of the deviance in P_z. However, biomass levels and environmental conditions exerted a much greater influence on P_z, with irradiance explaining the largest proportion (68.24%) of the deviance. Multiple predictor GAMs revealed that 96.26% of the deviance in P_z could be explained by a model which included nitrate, chlorophyll a, and irradiance.     

Ocean Color Satellite Imagery and Shipboard Measurements of Chlorophyll a and Suspended Particulate Matter Distribution in the East China Sea

Satellite-derived ocean color data of Coastal Zone Color Scanner (CZCS) on board the Nimbus-7 and Ocean Color and Temperature Scanner (OCTS) on board the Advanced Earth Observing Satellite (ADEOS) are jointly used with historical in situ data to examine seasonal and spatial distributions of chlorophyll a (Chl-a) and suspended particulate matter (SPM) concentrations in the East China Sea. Ocean color imagery showed that Chl-a concentrations on the continental shelf were higher than those of the Kuroshio area throughout the year. Satellite-derived Chl-a concentrations are generally in good accordance with historical in situ values during spring through autumn (although no shipboard in situ measurement was conducted at nearshore areas). In contrast, ocean color imagery in winter indicated high Chl-a concentrations (4–10 mg m^–3) on the continental shelf where bottom depth was less than 50 m when surface water was turbid (2–72 g m^–3 of SPM at surface), while historical in situ values were usually less than 1 mg m^–3. This suggests that resuspended bottom sediment due to wind-driven mixing and winter cooling is responsible for the noticeable overestimation of satellite-derived Chl-a concentrations. The algorithm for ocean color needs to be improved urgently for turbid water.     

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.