Effects of increased pCO(2) and temperature on the North Atlantic spring bloom. I. The phytoplankton community and biogeochemical response

The North Atlantic spring bloom is one of the largest annual biological events in the ocean, and is characterized by dominance transitions from siliceous (diatoms) to calcareous (coccolithophores) algal groups. To study the effects of future global change on these phytoplankton and the biogeochemical cycles they mediate, a shipboard continuous culture experiment (Ecostat) was conducted in June 2005 during this transition period. Four treatments were examined: (1) 12 degrees C and 390 ppm CO2 (ambient control), (2) 12 degrees C and 690 ppm CO2 (high pCO(2)) (3) 16 degrees C and 390 ppm CO2 (high temperature), and (4) 16 degrees C and 690 ppm CO2 ('greenhouse'). Nutrient availability in all treatments was designed to reproduce the low silicate conditions typical of this late stage of the bloom. Both elevated pCO(2) and temperature resulted in changes in phytoplankton community structure. Increased temperature promoted whole community photosynthesis and particulate organic carbon (POC) production rates per unit chlorophyll a. Despite much higher coccolithophore abundance in the greenhouse treatment, particulate inorganic carbon production (calcification) was significantly decreased by the combination of increased pCO(2) and temperature. Our experiments suggest that future trends during the bloom could include greatly reduced export of calcium carbonate relative to POC, thus providing a potential negative feedback to atmospheric CO2 concentration. Other trends with potential climate feedback effects include decreased community biogenic silica to POC ratios at higher temperature. These shipboard experiments suggest the need to examine whether future pCO2 and temperature increases on longer decadal timescales will similarly alter the biological and biogeochemical dynamics of the North Atlantic spring bloom.

Phytoplankton biomass size structure and its regulation in the Southern Yellow Sea (China): Seasonal variability

Phytoplankton size structure plays a significant role in controlling the carbon flux of marine pelagic ecosystems. The mesoscale distribution and seasonal variation of total and size-fractionated phytoplankton biomass in surface waters. as measured by chlorophyll a (Chl a), was studied in the Southern Yellow Sea using data from four cruises during 2006-2007. The distribution of Chl a showed a high degree of spatial and temporal variation in the study area. Chl a concentrations were relatively high in the summer and autumn, with a mean of 142 and 1.27 mg m(-3), respectively. Conversely, in the winter and spring. the average Chl a levels were only 098 and 0.99 mg m(-3) Total Chl a showed a clear decreasing gradient from coastal areas to the open sea in the summer, autumn and winter cruises. Patches of high Chl a were observed in the central part of the Southern Yellow Sea in the spring due to the onset of the phytoplankton bloom. The eutrophic coastal waters contributed at least 68% of the total phytoplankton biomass in the surface layer. Picophytoplankton showed a consistent and absolute dominance in the central region of the Southern Yellow Sea (>40%) in all of the cruises, while the proportion of microphytoplankton was the highest in coastal waters The relative proportions of pico- and nanophytoplankton decreased with total biomass, whereas the proportion of the micro-fraction increased with total biomass. Relationships between phytoplankton biomass and environmental factors were also analysed. The results showed that the onset of the spring bloom was highly dependent on water column stability. Phytoplankton growth was limited by nutrient availability in the summer due to the strong thermocline. The combined effects of P-limitation and vertical mixing in the autumn restrained the further increase of phytoplankton biomass in the Surface layer. The low phytoplankton biomass in winter was caused by vertical dispersion due to intense mixing. Compared with the availability of nutrients. temperature did not seem to cause direct effects on phytoplankton biomass and its size structure. Although interactions of many different environmental factors affected phytoplankton distributions. hydrodynamic conditions seemed to be the dominant factor. Phytoplankton size structure was determined mainly by the size-differential capacity in acquiring resource. Short time scale events, such as the spring bloom and the extension of Yangtze River plume, can have substantial influences, both on the total Chl a concentration and on the size structure of the phytoplankton. (C) 2009 Elsevier Ltd. All rights reserved.

Distribution of calcifying and silicifying phytoplankton in relation to environmental and biogeochemical parameters during the late stages of the 2005 North East Atlantic Spring Bloom

The late stage of the North East Atlantic (NEA) spring bloom was investigated during June 2005 along a transect section from 45 to 66 degrees N between 15 and 20 degrees W in order to characterize the contribution of siliceous and calcareous phytoplankton groups and describe their distribution in relation to environmental factors. We measured several biogeochemical parameters such as nutrients, surface trace metals, algal pigments, biogenic silica (BSi), particulate inorganic carbon (PIC) or calcium carbonate, particulate organic carbon, nitrogen and phosphorus (POC, PON and POP, respectively), as well as transparent exopolymer particles (TEP). Results were compared with other studies undertaken in this area since the JGOFS NABE program. Characteristics of the spring bloom generally agreed well with the accepted scenario for the development of the autotrophic community. The NEA seasonal diatom bloom was in the late stages when we sampled the area and diatoms were constrained to the northern part of our transect, over the Icelandic Basin (IB) and Icelandic Shelf (IS). Coccolithophores dominated the phytoplankton community, with a large distribution over the Rockall-Hatton Plateau (RHP) and IB. The Porcupine Abyssal Plain (PAP) region at the southern end of our transect was the region with the lowest biomass, as demonstrated by very low Chla concentrations and a community dominated by picophytoplankton. Early depletion of dissolved silicic acid (DSi) and increased stratification of the surface layer most likely triggered the end of the diatom bloom, leading to coccolithophore dominance. The chronic Si deficiency observed in the NEA could be linked to moderate Fe limitation, which increases the efficiency of the Si pump. TEP closely mirrored the distribution of both biogenic silica at depth and prymnesiophytes in the surface layer suggesting the sedimentation of the diatom bloom in the form of aggregates, but the relative contribution of diatoms and coccolithophores to carbon export in this area still needs to be resolved.

A field study on the conversion ratio of phytoplankton biomass carbon to chlorophyll-a in Jiaozhou Bay, China

A one-year field study was conducted to determine the conversion ratio of phytoplankton biomass carbon (Phyto-C) to chlorophyll-a (Chl-a) in Jiaozhou Bay, China. We measured suspended particulate organic carbon (POC) and phytoplankton Chl-a samples collected in surface water monthly from March 2005 to February 2006. The temporal and spatial variations of Chl-a and POC concentrations were observed in the bay. Based on the field measurements, a linear regression model II was used to generate the conversion ratio of Phyto-C to Chl-a. In most cases, a good linear correlation was found between the observed POC and Chl-a concentrations, and the calculated conversion ratios ranged from 26 to 250 with a mean value of 56 A mu g A mu g(-1). The conversion ratio in the fall was higher than that in the winter and spring months, and had the lowest values in the summer. The ratios also exhibited spatial variations, generally with low values in the near shore regions and relatively high values in offshore waters. Our study suggests that temperature was likely to be the main factor influencing the observed seasonal variations of conversion ratios while nutrient supply and light penetration played important roles in controlling the spatial variations.

Phytoplankton distributions and their relationship with the environment in the Changjiang Estuary, China

This study was carried out in the Changjiang Estuary from 19 to 26 May 2003. Based on the data collected from 29 stations, including two anchor stations, phytoplankton taxonomic composition, abundance, diurnal variability and spatial distribution were examined. Eighty-seven species, including 54 species of diatoms and 16 red tide causative species, were identified. Average diversity index (H') and evenness (J) values were 1.04 and 0.40, respectively. A bloom in abundance of certain phytoplankton species, especially Prorocentrum dentatum and Skeletoneina costatum, was thought to be the cause of the low diversity index and evenness values. Total phytoplankton abundance averaged 6.75 x 10(5) cells 1(-1), and was much higher than previous investigation carried out in the same month in 1986. Abundance increased seaward showing a distinct spatial difference, and the dominant species varied with salinity. Correlation between phosphorus and abundance further supported the former conclusion that phosphorus is the controlling factor in phytoplankton growth in the Changjiang Estuary where light is not limiting. Based on the relationship between DO, pH and abundance, it is likely that the bloom was caused by rapid in situ growth of phytoplankton with high nutrients and sufficient light. The data also indicated that the duration of the bloom was not long and

Studies on growth rate and grazing mortality rate by microzooplankton of size-fractionated phytoplankton in spring and summer in the Jiaozhou Bay, China

Dilution experiments were performed to examine the growth rate and grazing mortality rate of size-fractionated phytoplankton at three typical stations, inside and outside the bay, in the spring and summer of 2003 in the Jiaozhou Bay, China. in spring, the phytoplankton community structure was similar among the three stations, and was mainly composed of nanophytoplankton, such as, Skeletonema costatum and Cylindrotheca closterium. The structure became significantly different for the three stations in summer, when the dominant species at Stas A, B and C were Chaetoceros curvisetus, Pseudo-nitzschia delicatissima, C. affinis, C. debilis, Coscinodiscus oculus-iridis and Paralia sulcata respectively. Tintinnopsis beroidea and T. tsingtaoensis were the dominant species in spring, whereas the microzooplankton was apparently dominated by Strombidium sp. in summer. Pico- and nanophytoplankton had a relatively greater growth rate than microzooplankton both in spring and summer. The growth rate and grazing mortality rate were 0.18 similar to 0.44 and 0.12 similar to 1.47 d(-1) for the total phytoplankton and 0.20 similar to 0.55 and 0.21 similar to 0.37 d-1 for nanophytoplankton in spring respectively. In summer, the growth rate and grazing mortality rate were 0.38 similar to 0.71 and 0.27 similar to 0.60 d-1 for the total phytoplankton and 0.11 similar to 1.18 and 0.41 similar to 0.72 d(-1) for nano- and microphytoplankton respectively. The carbon flux consumed by microzooplankton per day was 7.68 similar to 39.81 mg/m(3) in spring and 12.03 similar to 138.22 mg/m(3) in summer respectively. Microzooplankton ingested 17.56%similar to 92.19% of the phytoplankton standing stocks and 31.77%similar to 467.88% of the potential primary productivity in spring; in contrast, they ingested 34.60%similar to 83.04% of the phytoplankton standing stocks and 71.28%similar to 98.80% of the potential primary productivity in summer. Pico- and nanophytoplankton appeared to have relatively greater rates of growth and grazing mortality than microphytoplankton during the experimental period. The grazing rate of microzooplankton in summer was a little bit greater than that in spring because of the relatively higher incubation temperature and different dominant microzooplankton species. Microzooplankton preferred ingesting nanophytoplankton to microphytoplankton in spring, while they preferred ingesting picophytoplankton to nanophytoplankton and microphytoplankton in summer. Compared with the results of dilution experiments performed in various waters worldwide, the results are in the middle range.

Evolution of nutrient structure and phytoplankton composition in the Jiaozhou Bay ecosystem

The inventories of nutrients in the surface water and large phytoplankton( > 69 pm) were analyzed from the data set of JERS ecological database about a typical coastal waters, the Jiaozhou Bay, China, from 1960s for N, P and from 1980s; for Si. By examining long-term changes of nutrient concentration, calculating stoichiometric balance, and comparing diatom composition, Si limitation of diatom production was found to be more possible. The possibility of Si limitation was from 37% in 1980s to 50% in 1990s. Jiaozhou Bay ecosystem is becoming serious eutrophication, with notable increase of NO2-N, NO3-N and NH4-N from 0.1417 mumol/L, 0.5414 mumol/L, 1.7222 mumol/L in 1960s to 0.9551 mumol/L, 3.001 mumol/L, 8.0359 mumol/L in late 1990s respectively and prominent decrease of Si from 4.2614 mumol/L in 1980s to 1.5861 mumol/L in late 1990s; the nutrient structure is controlled by nitrogen; the main limiting nutrient is probably silicon; because of the Si limitation the phytoplankton community structure has changed drastically.

The effects of spring-neap tide on the phytoplankton community development in the Jiaozhou Bay, China

The development of the phytoplankton community was studied in the Jiaozhou Bay during the spring to neap tide in August 2001, through three cruises and a 15 d continuous observation. This investigation indicates that diatom cell abundance increased sharply following the end of a spring tide, from 9 cells/cm(3) to a peak of 94 cells/cm(3). The dominant species composition and abundance show a quick species sequence from spring to neap tide, and the dominant species at the start phase is Skeletomena costatum, then changes to Chaetoceros curvisetus, finally it changes to Eucampia zodiacus. Silicate concentration increases during spring tide, as a result of nutrient replenishment from the water-sediment interface, its initial average concentration in neap tide is 1.39 mumol/dm(3) and reached the peak average concentration of 8.40 mumol/dm(3) in spring tide. But the nitrogen concentration dropped due to dilution by the low nitrogen seawater from the Huanghai Sea, its initial average concentration in neap tide is 67 mumol/dm(3) and decreased to the average concentration of 54 mumol/dm(3) in spring tide. The degree of silicon limitation was decreased and phytoplankton, especially diatoms, responds immediately after nutrient replenishment in the water column. Skeletonmea costatum, as one of the dominant species in the Jiaozhou Bay, shows a quicker response to nutrient availability than Eucampia zodiacus and Chaetoceros curvisetus. It is proposed that dominant species composition and water column stability synchronously determine the development of phytoplankton summer blooms in the Raozhou bay.

Dynamics of dimethylsulfide and dimethylsulfoniopropionate produced by phytoplankton in the Chinese seas - Distribution patterns and affecting factors

Distribution of dimethylsulfide (DMS) and/or particulate dimethylsulfoniopropionate (DMSPp) concentrations in the Jiaozhou Bay, Zhifu Bay and East China Sea were investigated during the period of 1994 - 1998. Both DMS and DMSPp levels showed remarkable temporal and spatial variations. High values occurred in the coastal or shelf waters and low values in the offshore waters. The highest levels were observed in spring or summer and lowest in autumn. DMS or DMSPp distribution patterns were associated with water mass on a large geographical scale, while biological and chemical factors were more likely influential on smaller-scale variations. Diatoms could play an important role in total DMS or DMSPp abundance in coastal waters. Nitrate was found to have a two-phase relationship with DMSPp concentrations: positive when nitrate concentration was lower than 1 mumol/L, and negative when it was above. Anthropogenic factors such as sewage input and aquaculture also showed influences on DMS or DMSPp concentration.

Grazing impact of copepods on phytoplankton in the Bohai Sea

During spring (April/May 1999) and autumn (September/October 1998) cruises in the Bohai Sea, China, copepods were the dominant components of mesozooplankton, the most abundant species being Calanus sinicus, Centropages mcmurrichi, Paracalanus parvus, Acartia bifilosa and Oithona similis. Pigment ingestion rates by three size classes of copepods (200-500, 500-1000 and > 1000 mum) were measured. In the south of the investigation area, gut pigment content (GPC), individual pigment-specific ingestion rates and grazing impacts on phytoplankton were lower in spring than in autumn. In the central area, GPC and individual pigment-specific ingestion rates were higher in spring than in autumn. The grazing impact on phytoplankton by the copepod assemblages was lower in spring than in autumn, however, because of the relatively smaller biomass in spring. In the western area where the Bohai Sea joins the Yellow Sea, GPC, individual pigment-specific ingestion rates and grazing impacts on phytoplankton were higher in spring than in autumn. Among the three size groups, the small-sized animals (200-500 mum) contributed more than 50% (range 38-98%) of the total copepod grazing during both cruises. The grazing impact on phytoplankton by copepods was equivalent to 11.9% (range 3.0-37.1%) of the chlorophyll-a standing stock and 53.3% (range 21.4-91.4%) of the primary production during the spring cruise. Grazing impact was equivalent to 6.3% (range 2.0-11.6%) of the chlorophyll-a standing stock and >100% (range 25.7-141.6%) of the primary production during the autumn cruise. The copepod community apparently consumed only a modest proportion of the standing stock of phytoplankton during spring and autumn blooms. They did, however, sometimes graze a significant proportion of daily primary production and hence were presumably able to limit the rate of further accumulation of phytoplankton, or even to prevent it. (C) 2003 Elsevier Ltd. All rights reserved.

Size-fractionated phytoplankton biomass in autumn of the Changjiang (Yangtze) River Estuary and its adjacent waters after the Three Gorges Dam construction

A cruise was undertaken from 3rd to 8th November 2004 in Changjiang (Yangtze) River Estuary and its adjacent waters to investigate the spatial biomass distribution and size composition of phytoplankton. Chlorophyll-a (Chl-a) concentration ranged 0.42-1.17 mu g L-1 and 0.41-10.43 mu g L-1 inside and outside the river mouth, with the mean value 0.73 mu g L-1 and 1.86 mu g L-1, respectively. Compared with the Chl-a concentration in summer of 2004, the mean value was much lower inside, and a little higher outside the river mouth. The maximal Chl-a was 10.43 mu g L-1 at station 18 (122.67 degrees E, 31.25 degrees N), and the region of high Chl-a concentration was observed in the central survey area between 122.5 degrees E and 123.0 degrees E. In the stations located east of 122.5 degrees E, Chl-a concentration was generally high in the upper layers above 5 m due to water stratification. In the survey area, the average Chl-a in sizes of > 20 mu m and < 20 mu m was 0.28 mu g L-1 and 1.40 mu g L-1, respectively. High Chl-a concentration of < 20 mu m size-fraction indicated that the nanophytoplankton and picophytoplankton contributed the most to the biomass of phytoplankton. Skeletonema costatum, Prorocentrum micans and Scrippsiella trochoidea were the dominant species in surface water. The spatial distribution of cell abundance of phytoplankton was patchy and did not agree well with that of Chl-a, as the cell abundance could not distinguish the differences in shape and size of phytoplankton cells. Nitrate and silicate behaved conservatively, but the former could probably be the limitation factor to algal biomass at offshore stations. The distribution of phosphate scattered considerably, and its relation to the phytoplankton biomass was complicated.

Fast microzooplankton grazing on fast-growing, low-biomass phytoplankton: a case study in spring in Chesapeake Bay, Delaware Inland Bays and Delaware Bay

Dilution experiments were performed to examine the growth and grazing mortality rates of picophytoplankton (< 2 mu m), nanophytoplankton (2-20 mu m), and microphytoplankton (> 20 mu m) at stations in the Chesapeake Bay (CB), the Delaware Inland Bays (DIB) and the Delaware Bay (DB), in early spring 2005. At station CB microphytoplankton, including chain-forming diatoms were dominant, and the microzooplankton assemblage was mainly composed of the tintinnid Tintinnopsis beroidea. At station DIB, the dominant species were microphytoplanktonic dinoflagellates, while the microzooplankton community was mainly composed of copepod nauplii and the oligotrich ciliate Strombidium sp. At station DB, nanophytoplankton were dominant components, and Strombidium and Tintinnopsis beroidea were the co-dominant microzooplankton. The growth rate and grazing mortality rate were 0.13-3.43 and 0.09-1.92 d(-1) for the different size fractionated phytoplankton. The microzooplankton ingested 73, 171, and 49% of standing stocks, and 95, 70, and 48% of potential primary productivity for total phytoplankton at station CB, DIB, and DB respectively. The carbon flux for total phytoplankton consumed by microzooplankton was 1224.11, 100.76, and 85.85 mu g C 1(-1) d(-1) at station CB, DIB, and DB, respectively. According to the grazing mortality rate, carbon consumption rate and carbon flux turn over rates, microzooplankton in study area mostly preferred to graze on picophytoplankton, which was faster growing but was lowest biomass component of the phytoplankton. The faster grazing on Fast-Growing-Low-Biomass (FGLB) phenomenon in coastal regions is explained as a resource partitioning strategy. This quite likely argues that although microzooplankton grazes strongly on phytoplankton in these regions, these microzooplankton grazers are passive.

Preliminary study on different nutrient pools supplies for the phytoplankton growth in the Jiaozhou Bay in China in the fall of 2004

The source and significance. of two mitrients, nitrogen. and phosphorous, were investigated by a modified dilution method performed on seawater samples from the Jiaozhou Bay in autumn 2004. This modified dilution method accounted for the phytoplankton growth rate, microzooplankton grazing mortality rate, the external nutrient pools, as well as nutrient supplied through remineralization by microzooplankton. The results indicated that the phytoplankton net growth rate increased in turn from inside the bay, to outside the bay, to in the Xiaogang Harbor. The phytoplankton, maximum growth rates and microzooplankton grazing mortality rates were 1.14 and 0.92 d(-1) outside the bay, 0.42 and 0.32 d(-1) inside the bay and 0.98 and 0.62 d(-1) in the harbor respectively. Outside the bay, the remineralized nitrogen (K-r = 24.49) had heavy influence on the growth of the phytoplankton. Inside the bay, the remineralized phosphorus(K-r = 3.49) strongly affected the phytoplankton growth. In the harbor, the remineralized phosphorus (K-r = 3.73) was in larger demand by phytoplankton growth. The results demonstrated that the different nutrients pools supplied for phytoplankton growth were greatly in accordance with the phytoplankton community structure, microzooplankton grazing mortality rates and environmental conditions. It is revealed that, nutrient remineralization is much more important for the phytoplankton growth in the Jiaozhou Bay than previously believed.