Microzooplankton composition,biomass and grazing rates along the WOCE SR3 line between Tasmania and Antarctica

Microzooplankton species composition and grazing rates on phytoplankton were investigated along a transect between ∼46 and 67°S, and between 140 and 145°E. Experiments were conducted in summer between November 2nd and December 14th in 2001. The structure of the microbial food web changed considerably along the transect and was associated with marked differences in the physical and chemical environment encountered in the different water masses and frontal regions. On average microzooplankton grazing experiments indicated that 91%, 102%, and 157%, (see results) of the phytoplankton production would be grazed in the <200, <20 and <2 μm size fractions, respectively, indicating microzooplankton grazing was potentially constraining phytoplankton populations (<200 μm) along most of the transect. Small ciliates in general and especially oligotrich species declined in importance from the relatively warm, Southern Subtropical Front waters (6.8 μg C/L) to the colder waters of the southern branch of the Polar Front (S-PF), (∼0.5 μg C/L) before increasing again near the Antarctic landmass. Large changes in microzooplankton dominance were observed, with heterotrophic nanoflagellates (HNF), ciliates and larger dinoflagellates having significant biomass in different water masses. HNF were the dominant grazers when chlorophyll a was low in areas such as the Inter-Polar Frontal Zone (IPFZ), while in areas of elevated biomass such as the S-PF and Southern Antarctic Circumpolar Current (SACC), a mix of copepod nauplii and large heterotrophic and mixotrophic dinoflagellates tended to dominate the grazing community. In the S-PF and SACC water masses the tight coupling observed between the microzooplankton grazers and phytoplankton populations over most of the rest of the transect was relaxed. In these regions grazing was low on the >20 μm size fraction of chlorophyll a, which dominated the biomass, while smaller diatoms and nanoplankton in the <20 μm size fraction were still heavily grazed. The lack of grazing pressure on large phytoplankton contributes to this region's potential to export carbon with larger cells known to have higher sinking rates.     

Temporal change of clustered distribution of planktonic ciliates in Toyama Bay in summers of 1989 and 1990

Temporal change of clustered distribution in vertical profiles of three nutritional groups of planktonic ciliates, e.g. heterotrophic naked ciliates, mixotrophic naked ciliates and heterotrophic loricated ciliates, was investigated by following a drifting buoy in Toyama Bay on the Japan Sea coast of central Japan in summers of 1989 and 1990. Clustered distribution, represented as the mode of population density in the vertical plane, occurred mainly in the oligotrophic upper layer (0–50 m depth) above the subsurface chlorophyll-a maximum layer. Its clustered degree was stronger when the mode of population density in the vertical plane was formed at shallower depth, while its longevity was shorter as mentioned above. Vertical distribution of ciliates during summer in Toyama Bay is characterized by ephemeral clustered distribution, or in other wards, by rapid alternations of appearance and disappearance of the clustered distribution.     

北部湾北部海域水体异养细菌的时空分布特征研究

为探讨环境因素对异养细菌丰度的影响,2016年9月至2017年8月通过月度航次调查对北部湾北部海域异养细菌丰度的时空分布特征进行了系统研究。结果表明,调查海区异养细菌丰度介于（2.75~56.86）×105 cell/mL,平均值为（11.01±6.31）×105 cell/mL。各季节细菌丰度从高至低依次为:夏季、春季、冬季、秋季。异养细菌丰度由近岸海域向西南深水区方向逐渐降低,在近岸浅水区垂直分布均匀,在水深大于20 m的海区出现季节性分层现象:表层细菌丰度较高,底层细菌丰度较低。主成分分析显示温度对异养细菌时空分布有重要影响,秋、冬季异养细菌丰度与温度呈显著负相关,在春、夏季呈显著正相关。细菌丰度与盐度呈显著负相关,说明海水盐度变化是细菌时空分布重要影响因素。异养细菌丰度与叶绿素a和溶解氧含量呈显著正相关,表明浮游植物初级生产过程影响了异养细菌的时空分布。在秋、冬和春3季异养细菌丰度与营养盐水平呈显著负相关,二者关系受浮游植物生物量间接影响。异养细菌时空分布差异取决于环境条件的变化,温度、盐度、叶绿素a和溶解氧含量是影响异养细菌丰度分布的主要因素。     

Responses of phytoplankton and heterotrophic bacteria in the northwest subarctic Pacific to in situ iron fertilization as estimated by HPLC pigment analysis and flow cytometry

To verify the hypothesis that the growth of phytoplankton in the Western Subarctic Gyre (WSG), which is located in the northwest subarctic Pacific, is suppressed by low iron (Fe) availability, an in situ Fe fertilization experiment was carried out in the summer of 2001. Changes over time in the abundance and community structure of phytoplankton were examined inside and outside an Fe patch using phytoplankton pigment markers analyzed by high-performance liquid chromatography (HPLC) and flow cytometry (FCM). In addition, the abundance of heterotrophic bacteria was also investigated by FCM. The chlorophyll a concentration was initially ca. 0.9 μg l⁻¹ in the surface mixed layer where diatoms and chlorophyll b-containing green algae (prasinophytes and chlorophytes) were predominant in the chlorophyll biomass. After the iron enrichment, the chlorophyll a concentration increased up to 9.1 μg l⁻¹ in the upper 10 m inside the Fe patch on Day 13. At the same time, the concentration of fucoxanthin (a diatom marker) increased 45-fold in the Fe patch, and diatoms accounted for a maximum 69% of the chlorophyll biomass. This result was consistent with a microscopic observation showing that the diatom Chaetoceros debilis had bloomed inside the Fe patch. However, chlorophyllide a concentrations also increased in the Fe patch with time, and reached a maximum of 2.2 μg l⁻¹ at 5 m depth on Day 13, suggesting that a marked abundance of senescent algal cells existed at the end of the experiment. The concentration of peridinin (a dinoflagellate marker) also reached a maximum 24-fold, and dinoflagellates had contributed significantly (>15%) to the chlorophyll biomass inside the Fe patch by the end of the experiment. Concentrations of 19′-hexanoyloxyfucoxanthin (a prymnesiophyte marker), 19′-butanoyloxyfucoxanthin (a pelagophyte marker), and alloxanthin (a cryptophyte marker) were only incremented a few-fold increment inside the Fe patch. On the contrary, chlorophyll b concentration reduced to almost half of the initial level in the upper 10 m water column inside the Fe patch at the end of the experiment. A decrease with time in the abundance of eukaryotic ultraphytoplankton (<ca. 5 μm in size), in which chlorophyll b-containing green algae were possibly included was also observed by FCM. Overall, our results indicate that Fe supply can dramatically alter the abundance and community structure of phytoplankton in the WSG. On the other hand, cell density of heterotrophic bacteria inside the Fe patch was maximum at only ca. 1.5-fold higher than that outside the Fe patch. This indicates that heterotrophic bacteria abundance was little respondent to the Fe enrichment.     

Full-depth profiles of prokaryotes,heterotrophic nanoflagellates,and ciliates along a transect from the equatorial to the subarctic central Pacific Ocean

Studies in epipelagic waters report higher heterotrophic microbial biomass in the productive high latitudes than in the oligotrophic low latitudes; however, biogeographical data are scarce in the deep ocean. To examine the hypothesis that the observed latitudinal differences in heterotrophic microbial biomass in the epipelagic zone also occur at depth, abundance and biomass of heterotrophic prokaryotes, nanoflagellates (HNF), and ciliates were determined at depths of 5–5000 m in the central Pacific between August and September of 2005. Heterotrophic microbial biomass increased from the tropical to the subarctic region over the full water column, with latitudinal differences in prokaryotic biomass increasing from 2.3-fold in the epipelagic zone to 4.4-fold in the bathypelagic zone. However, the latitudinal difference in HNF and ciliate biomass decreased with depth. In the mesopelagic zone, the vertical attenuation rate of prokaryotic abundance, which was calculated as the linear regression slope of log-log plot of abundance versus depth, ranged from –0.55 to –1.26 and was more pronounced (steeper slope) in the lower latitudes. In contrast, the vertical attenuation rate of HNF in the mesopelagic zone (–1.06 to –1.27) did not differ with latitude. In the subarctic, the attenuation rate of HNF was 1.7 times steeper than for prokaryotes. These results suggest the accumulation of prokaryotes in the deep subarctic Pacific, possibly due to low grazing pressure. Although the vertical attenuation rate of ciliates was steepest in the bathypelagic zone, HNF abundance did not further decrease at depths below 1000 m, except for at 2000 m where HNF was lowest across the study area. Ciliate abundance ranged 0.3–0.8 cells l^–1 at 4000 m, and were below the detection limit (<0.1 cells l ^–1) at 5000 m. To our knowledge, this study presents the first data for ciliates below 2000 m.     

Mixing and biological production at eddy margins in the eastern Gulf of Alaska

We performed a multi-day shipboard experiment in June 2001 to test whether combining water from within an anticyclonic mesoscale eddy in the eastern Gulf of Alaska with water from outside could result in enhanced phytoplankton growth and to determine how mixing might influence planktonic assemblages. Initially, the eddy had lower standing stocks of algal pigments (chlorophyll a [chl a] and accessory pigments), nutrients, phytoplankton, and particulate organic carbon/nitrogen compared to waters outside of the eddy. The eddy possessed a greater diversity and abundance of coastal diatoms while the outside waters had a greater proportion of oceanic species, including the endemic pennate diatom, Nitzschia cylindroformis. After one week of incubation, rates of primary production were significantly higher in the mixed water compared to both the eddy and outside treatments. Pigment concentrations (except chl c₃, alloxanthin, and zeaxanthin) and the proportion of large diatoms (mainly Pseudo-nitzschia spp.) and heterotrophic dinoflagellates were greater in the mixed water than would be expected from the simple combination of inside and outside waters. Nutrient limitation (most likely by trace metals) appeared to be less severe in the mixed water. Chl a was enhanced in the mixed water, particularly when compared to the eddy water. The mixing of eddy and outside water masses stimulated primary production by ∼20%, but more importantly, the mixing resulted in a distinct planktonic assemblage. The biomass enrichment was short-lived, indicating that the maintenance of elevated chl a would require further mixing events in a physical setting that also permits an accumulation of biomass. We note that submesoscale processes, including the intensification of ageostrophic circulation that elicits strong vertical mixing in the presence of strain, might explain observed patterns of high phytoplankton standing stocks at the inner edges of Haida eddies in the field.     

Abundance, variability, and potential grazing impact of planktonic ciliates in the open subaratic Pacific Ocean

The abundance and variability of planktonic ciliates in the open subarctic Pacific were determined during four month-long cruises in 1987 and 1988. The ciliate community, numerically dominated by relatively small aloricate choreotrichs, was comparable in abundance to communities in a range of oceanic and neritic environments, including waters with much higher average chlorophyll concentrations. Integrated (0–80m) ciliate biomass was typically 100–200mgC m⁻², although 3- to 4-fold higher levels were observed on two occasions in spring. Ciliate community biomass, in general, was dominated by large (>20μm width) individuals, although in August 1988 the biomass of smaller cells was as great or greater. The estimated grazing impact of the ciliate community averaged 20% of the primary production. On one instance in May 1988, however, a large biomass of ciliates led to an estimated grazing impact equivalent to 55% of phytoplankton production. While ciliates may be major phytoplankton grazers during sporadic ciliate “blooms”, dino- and other heterotrophic flagellates, which make up the bulk of microheterotroph biomass, must normally be of equal or greater importance as herbivores in this ocean region.     

Phytoplankton composition and biomass across the southern Indian Ocean

Phytoplankton composition and biomass was investigated across the southern Indian Ocean. Phytoplankton composition was determined from pigment analysis with subsequent calculations of group contributions to total chlorophyll a (Chl a) using CHEMTAX and, in addition, by examination in the microscope. The different plankton communities detected reflected the different water masses along a transect from Cape Town, South Africa, to Broome, Australia. The first station was influenced by the Agulhas Current with a very deep mixed surface layer. Based on pigment analysis this station was dominated by haptophytes, pelagophytes, cyanobacteria, and prasinophytes. Sub-Antarctic waters of the Southern Ocean were encountered at the next station, where new nutrients were intruded to the surface layer and the total Chl a concentration reached high concentrations of 1.7 ??g Chl a L⁻¹ with increased proportions of diatoms and dinoflagellates. The third station was also influenced by Southern Ocean waters, but located in a transition area on the boundary to subtropical water. Prochlorophytes appeared in the samples and Chl a was low, i.e., 0.3 ??g L⁻¹ in the surface with prevalence of haptophytes, pelagophytes, and cyanobacteria. The next two stations were located in the subtropical gyre with little mixing and general oligotrophic conditions where prochlorophytes, haptophytes and pelagophytes dominated. The last two stations were located in tropical waters influenced by down-welling of the Leeuwin Current and particularly prochlorophytes dominated at these two stations, but also pelagophytes, haptophytes and cyanobacteria were abundant. Haptophytes Type 6 (sensuZapata et al., 2004), most likely Emiliania huxleyi, and pelagophytes were the dominating eucaryotes in the southern Indian Ocean. Prochlorophytes dominated in the subtrophic and oligotrophic eastern Indian Ocean where Chl a was low, i.e., 0.043-0.086 ??g total Chl a L⁻¹ in the surface, and up to 0.4 ??g Chl a L⁻¹ at deep Chl a maximum. From the pigment analyses it was found that the dinoflagellates of unknown trophy enumerated in the microscope at the oligotrophic stations were possibly heterotrophic or mixotrophic. Presence of zeaxanthin containing heterotrophic bacteria may have increased the abundance of cyanobacteria determined by CHEMTAX.     

Seasonal pigment patterns of surface phytoplankton in the subtropical southern hemisphere

Pigment indices were used to characterise the seasonal succession of phytoplankton, and associated changes in chlorophyll a and accessory pigments, in subtropical waters of the three ocean basins in the southern hemisphere. Diagnostic indices revealed the dominance of small flagellates and elevated biomass during winter–spring in the Pacific, mixed flagellate–prokaryote communities and intermediate biomass during early summer in the Atlantic, and prokaryote dominance with low biomass in mid-summer in the Indian Ocean. Photo-pigment indices indicated only a small variation in the chlorophyll a proportion of total pigments across the ocean basins, but the accessory pigments varied considerably. Under low temperature and irradiance conditions, the photosynthetic carotenoids were prominent, but as temperatures and irradiance increased and nutrients declined, there was a significant increase in the proportion of photoprotective carotenoids. At high temperatures and irradiances, the photoprotective carotenoids were the largest component of the pigment pool, exceeding the proportion of chlorophyll a. These variations in phytoplankton composition, and their photoacclimation status, could be explained according to environmental changes and have implications for satellite estimations of biomass and primary production.     

Seasonal variations in abundance,growth and mortality of heterotrophic bacteria in Kagoshima Bay

Seasonal changes in abundance, growth and mortality of heterotrophic bacteria were investigated monthly from collections and dilution experiments in Kagoshima Bay, the southernmost of Japan. Bacteria occurred abundantly with considerable variation in the surface layers where chlorophyll a concentrations were high, whereas seasonal variations were obscure below 100 m. Especially, bacteria showed a decline of cell density toward summer when heterotrophic nano-flagellates increased their abundance. Seasonal and vertical variations in bacterial cell number during the study period were positively correlated with those of water temperature and pico-sized chlorophyll a concentration. Maximum growth and mortality rates showed positive correlations with water temperature but no positive relationships to size-fractionated chlorophyll a. Net increase rate (i.e. in situ rate if abundance changes) was negatively correlated with cell density of heterotrophic nano-flagellates. It is concluded that bacterial abundance is controlled by the resultant (i.e. net growth rate) of the balance between maximum growth and predatory mortality by heterotrophic nano-flagelllates which are both dependent on ambient temperature.     

Dynamics of heterotrophic dinoflagellates off the Pearl River Estuary,northern South China Sea

Variations in abundance, biomass, vertical profile and cell size of heterotrophic dinoflagellates (HDFs) between summer and winter and its controlling factors were studied in the northern South China Sea (SCS). It was found that HDF abundance and carbon biomass were 4–102 × 10³ cells L⁻¹ and 0.34–12.3 mg C L⁻¹ in winter (February 2004), respectively, while they were 2–142 × 10³ cells L⁻¹ and 0.22–31.4 μg C L⁻¹ in summer (July, 2004), respectively, in the northern SCS. HDF abundance and carbon biomass decreased from the estuary to inshore and then offshore. Vertical profiles of HDF abundance were heterogeneous, which accorded well with that of chlorophyll a (Chl.a). Higher abundance of HDFs was often observed at a depth of 30–70 m offshore waters, matching well with the Chl.a maximum, while it showed high abundance at the surface in some coastal and estuary stations. Small HDFs (≤20 μm) dominated the assemblage in term of abundance accounting for more than 90%. However, large HDFs (>20 μm) generally contributed equally in terms of carbon biomass, accounting for 47% on average. HDFs showed different variation patterns for the different study regions; in the estuarine and continental shelf regions, abundance and biomass values were higher in summer than those in winter, while it was the reverse pattern for the slope waters. Hydrological factors (e.g. water mass, river outflow, monsoon and eddies) associated with biological factors, especially the size-fractionated Chl.a, seemed to play an important role in regulating HDF distribution and variations in the northern South China Sea.     

Algal class abundances,estimated from chlorophyll and carotenoid pigments,in the western Equatorial Pacific under El Niño and non-El Niño conditions

Phytoplankton samples were collected from the equatorial Pacific (10°S to 10°N along 155°E) in June 1992 as part of the Australian contribution to the JGOFS program. Chlorophyll and carotenoid pigments were measured by HPLC, and a PC-based computer program (CHEMTAX) was used to estimate the contribution of 9 algal classes to the total chlorophyll a concentration in 9 separate depth bands at each location. This cruise occurred in the middle of the prolonged 1991/1993 El Niño, and the results are compared with similar data from a cruise in October 1990 which occurred before this El Niño but after the 1988/1989 La Niña.Changes in the pigment : chlorophyll a ratios appeared consistent across algal classes and, apart from some minor exceptions, consistent between cruises. Pigments involved in light-harvesting generally increased relative to chlorophyll a with increasing depth, whereas the ratio for photoprotective pigments (e.g. diadinoxanthin) usually decreased with depth. The zeaxanthin concentration per cell for cyanobacteria decreased with depth in the surface 75 m during 1992 as would be expected for a photoprotective pigment.Based on their contribution to the total chlorophyll a concentration, haptophytes, prochlorophytes, cyanobacteria (Synechococcus) and chlorophytes were the dominant algal classes in 1992. The chlorophyte contribution to chlorophyll a in 1992 (14.8%) was almost double that in 1990 (7.8%). This increase was largely at the expense of the cyanobacteria and haptophytes, which both decreased significantly. The increase in chlorophytes in 1992 was particularly noticeable in the surface waters south of the equator at about 4°S, where there was evidence of upwelling.     

High-resolution phytoplankton diel variations in the summer stratified central Yellow Sea

Vertical distributions of phytoplankton biomass and community structure were studied in the summer stratified central Yellow Sea (YS) using a submersible spectrofluorometer (Fluoroprobe, bbe Moldaenke, Germany), along with photosynthetic pigments analysis (HPLC-CHEMTAX), and microscope observation. Above all, the results of the dominating group obtained from these methods generally coincided with each other on the transect 35°N. The concentrations of brown algae, green algae and total chlorophyll a (Chl a) biomass were highly correlated between the results of Fluoroprobe observations and pigments analysis (r?=?0.79, 0.91 and 0.82, respectively, n?=?54, p?<?0.01). In the summer stratified central YS, significant differences in phytoplankton compositions on the vertical distribution was observed. On the basis of HPLC-CHEMTAX results, the dominating group of phytoplankton composition generally changed from cyanobacteria to chrysophytes and then to diatoms, from surface to bottom. Interestingly, on the basis of high-resolution observations using Fluoroprobe, a periodic fluctuation of the pycnocline presumably due to the semidiurnal internal tides was observed at an anchor station (35°N, 123°E). In addition, both nutrients and Chl a concentrations at the depth of the subsurface chlorophyll maximum (SCM) seemed to coincide with the rhythm of the pycnocline fluctuation, indicating the latter might have a potential impact on the dynamics of SCM phenomena in the summer stratified central YS.     

High-West and Low-East Distribution Patterns of Chlorophyll a, Primary Productivity and Diatoms in the Subarctic North Pacific Surface Waters, Midwinter 1996

We have determined chlorophyll a (Chla) concentration, primary productivity, cell density and species composition of diatoms, and the number of microzooplankton at the surface in the subarctic North Pacific in January 1996. The wet weight of copepods obtained by vertical tows from 150 m to the surface was also measured during the cruise. Chla concentration and primary productivity tended to be higher in the region west of 180°, the western subarctic North Pacific (WSNP), than east of 180°, the eastern subarctic North Pacific (ESNP). The same results were observed for the total diatom cell densities and for the genera Thalassiosira and Denticulopsis. Significant linear relationships were observed between the Thalassiosira cell density and Chla concentration and primary productivity, indicating that Thalassiosira contributes to the high-WSNP and low-ESNP distribution patterns of Chla concentration and primary productivity. Moreover, naked ciliate abundance tended to be lower in the WSNP than in the ESNP, whereas copepod biomass showed an inverse trend. Significantly negative Spearman rank correlations were found between the Thalassiosira cell density and the number of naked ciliates and between the number of naked ciliates and the wet weight of copepods. These results indicate that copepod grazing indirectly controls Thalassiosira cell density via predation on the naked ciliates. We conclude that the high copepod biomass in the WSNP is a factor controlling the high-WSNP and low-ESNP Thalassiosira abundance and hence Chla concentration and primary productivity patterns.     

Population dynamics of phytoplankton, heterotrophic bacteria, and viruses during the spring bloom in the western subarctic Pacific

We characterized the community composition of phytoplankton in the western subarctic Pacific from the pre-bloom to the decline phase of the spring bloom with special reference to decreases in the silicic acid concentration in surface waters as an index for diatom bloom development. Furthermore, responses of heterotrophic bacteria and viruses to the spring bloom were also concomitantly investigated. Under pre-bloom conditions when nutrients were abundant but the surface mixed layer depth was relatively deep, chlorophyll (Chl) a concentrations were consistently low and green algae (chlorophytes and prasinophytes), cryptophytes, and diatoms were predominant in the phytoplankton assemblages as estimated by algal pigment signatures. Together with the shallowing of the mixed layer depth and the decrease in silicic acid concentration, diatoms bloomed remarkably in the Oyashio region, though the magnitude of the bloom in the Kuroshio-Oyashio transition (hereafter Transition) region was relatively small. A total of 77 diatom species were identified, with the bloom-forming diatoms mainly consisting of Thalassiosira, Chaetoceros, and Fragilariopsis species. It has become evident that the carotenoid fucoxanthin can serve as a strong indicator of the diatom carbon biomass during the spring diatom bloom. Differences in the species richness of diatoms among stations generally enabled us to separate the Oyashio bloom stations from the Transition and the Oyashio pre-bloom stations. Relatively high values of the Shannon-Wiener index for the diatom species were also maintained during the Oyashio bloom, indicating that a wide variety of species then shared dominance. In the decline phase of the Oyashio bloom when surface nutrient concentrations decreased, senescent diatom cells increased, as inferred from the levels of chlorophyllide a. Although the cell density of heterotrophic bacteria changed little with the development of the diatom bloom, viral abundance increased toward the end of the bloom, suggesting an increased likelihood of mortality among organisms including diatoms resulting from viral infection. This is the first report on the microbial trophodynamics, including viruses, during the spring diatom bloom in the western subarctic Pacific.     

Photoacclimatization in the zooxanthellae of Pocillopora verrucosa and comparison with a pelagic algal community

Photoacclimatization of zooxanthellae extracted from the coral Pocillopora verrucosa was studied through the determination of pigments, light absorption and photosynthetic parameters, for samples collected in summer and winter between 1 and 40 m on a northwestern reef of Tahiti (French Polynesia). The same measurements were also performed on phytoplanktonic samples collected at a stable oceanic site north of the island. For the zooxanthellae, the variations with depth of all the parameters were generally of small amplitude. Seasonal differences were also observed. The photosynthetic to non-photosynthetic pigments ratio was higher at depth in both seasons and was higher in winter. The intracellular concentration of chlorophyll a and photosynthetic pigments was higher in winter, as was the photosynthetic pigments/chlorophyll a ratio, whereas the non-photosynthetic pigments/chlorophyll a ratio was higher in summer. Variations in the light absorption properties were also small. The photosynthetic parameters showed limited changes with depth with the largest variations (a factor of ∼2) observed for P^B_max. The trends observed for the phytoplankton assemblage were generally of much higher amplitudes than for the zooxanthellae (e.g. for photosynthetic to non-photosynthetic pigments ratio or the saturation parameter, E_k). These results suggest that, in the very clear Polynesian waters, the amount of energy that reaches the zooxanthellae of P. verrucosa is not variable enough in the 1–40 m depth range to result in a drastic modification of the photosynthetic apparatus of the algae.     

Microzooplankton diversity: relationships of tintinnid ciliates with resources,competitors and predators from the Atlantic Coast of Morocco to the Eastern Mediterranean

We examined tintinnid (loricate ciliate microzooplankton) diversity using data from 11 stations between the Moroccan upwelling system and the oligotrophic Eastern Mediterranean. Taxonomic and morphological diversity of tintinnids was compared to phytoplankton distribution and size-structure, to the abundance of competitors in the form of oligotrich ciliates, and predators as copepods. Tintinnid taxonomic diversity was estimated as numbers of species and the Shannon Index, H′; morphological diversity was quantified by substituting size classes of lorica dimensions for species. Total chlorophyll was partitioned into micro-, nano- and pico-fractions using pigment data and a size-diversity was estimated by considering the 3 size classes as 3 species. Along a west-to-east gradient, average water column concentrations of most organism groups declined approximately an order of magnitude yielding tight correlations. However, tintinnid diversity, both taxonomic and morphological, increased from the Atlantic upwelling station into the western basin of the Mediterranean, and declined slightly towards the Eastern Mediterranean, paralleling shifts in the chlorophyll size-diversity estimate. Diversity varied with absolute or relative abundance of oligotrich or copepods, but different diversity metrics were significantly correlated only with phytoplankton size-diversity. We conclude that tintinnid diversity more closely reflects resource diversity than competitive interactions or predation.     

Monsoonal influence on the distribution of phytoplankton pigments in the Arabian Sea

Variations in the distribution of chemotaxonomic pigments were monitored in the Arabian Sea and the Gulf of Oman at the end of the SW monsoon in September 1994 and during the inter-monsoon period in November/December 1994 to determine the seasonal changes in phytoplankton composition. The Gulf of Oman was characterized by sub-surface chlorophyll maxima at 20-40 m during both seasons, and low levels of divinyl chlorophyll a indicated that prochlorophytes did not contribute significantly to the total chlorophyll a. Prymnesiophytes (19′-hexanoyloxyfucoxanthin), diatoms (fucoxanthin) and chlorophyll b containing organisms accounted for most of the phytoplankton biomass in September, while prymnesiophytes dominated in November/December. In the Arabian Sea in September, high total chlorophyll a concentrations up to 1742 ng l^-1 were measured in the coastal upwelling region and a progressive decline was monitored along the 1670 km offshore transect to oligotrophic waters at 8°N. Divinyl chlorophyll a was not detected along this transect except at the two most southerly stations where prochlorophytes were estimated to contribute 25–30% to the total chlorophyll a. Inshore, the dominance of fucoxanthin and/or hexanoyloxyfucoxanthin indicated that diatoms and prymnesiophytes generally dominated the patchy phytoplankton community, with zeaxanthin-containing Synechococcus also being important, especially in surface waters. At the southern oligotrophic localities, Synechococcus and prochlorophytes dominated the upper 40 m and prymnesiophytes were the most prominent at the deep chlorophyll maximum. During the inter-monsoon season, total chlorophyll a concentrations were generally half those measured in September and highest levels were found on the shelf (1170 ng l^-1). Divinyl chlorophyll a was detected at all stations along the Arabian Sea transect, and we estimated that prochlorophytes contributed between 3 and 28% to the total chlorophyll a, while at the two oligotrophic stations this proportion increased to 51–52%. While procaryotes were more important in November/December than September, eucaryotes still accounted for >50% of the total chlorophyll a. Pigment/total chlorophyll a ratios indicated that 19′-hexanoyloxyfucoxanthin-containing prymnesiophytes were the dominant group, although procaryotes accounted for 65% at the two southerly oligotrophic stations.     

Seasonal variation in the community and size structure of nano- and microzooplankton in Gyeonggi Bay,Yellow Sea

To investigate the seasonal variation and community structure of nano- and microzooplankton in Gyeonggi Bay of the Yellow Sea, the abundance and carbon biomass of nano- and microzooplankton were evaluated at 10-day intervals from January 1997 to December 1999. Four major groups of nano- and microzooplankton communities were classified: heterotrophic ciliates, heterotrophic dinoflagellates (HDF), heterotrophic nanoflagellates (HNF), and copepod nauplii. The total carbon biomass of nano- and microzooplankton ranged from 10.2 to 168.8 μg C L⁻¹ and was highest during or after phytoplankton blooms. Nano- and microzooplankton communities were composed of heterotrophic ciliates (7.4–81.4%; average 41.7% of total biomass), HDF (0.1–70.3%; average 26.1% of total biomass), copepod nauplii (1.6–70.6%; average 20.7% of total biomass), and HNF (0.8–59.5%; average 11.5% of total biomass). The relative contribution of individual components in the nano- and microzooplankton communities appeared to differ by seasons. Ciliates accounted for the most major component of nano- and microzooplankton communities, except during summer and phytoplankton blooming seasons, whereas HDF were more dominant during the phytoplankton blooming seasons. The abundance and biomass of nano- and microzooplankton generally followed the seasonal dynamics of phytoplankton. The size and community distribution of nano- and microzooplankton was positively correlated with size-fractionated phytoplankton. The carbon requirement of microzooplankton ranged from 60 to 83% of daily primary production, and was relatively high when phytoplankton biomass was high. Therefore, our result suggests that the seasonal variation in the community and size composition of nano- and microzooplankton appears to be primarily governed by phytoplankton size and concentration as a food source, and their abundance may greatly affect trophic dynamics by controlling the seasonal abundance of phytoplankton.     

Possible mixotrophy of pigmented nanoflagellates: Microbial plankton biomass, primary production and phytoplankton growth in the NW Iberian upwelling in spring

Microbial plankton biomass, primary production (PP) and phytoplankton growth rates (μ) were estimated along the NW Iberian margin during an upwelling relaxation event. Although the interaction between wind forcing and coastline singularities caused high spatial variability in PP (0.4-8.4 g C m⁻² d⁻¹), two domains (coastal and oceanic) could be distinguished regarding microbial plankton biomass and μ. At the coastal domain, with higher influence of upwelling, diatoms showed an important contribution (27 ± 17%) to total autotrophic biomass (AB). Nonetheless, AB was dominated by autotrophic nanoflagellates (ANF) at both realms, accounting for 62 ± 16% and 89 ± 6% of the integrated AB at the coastal and oceanic domain respectively. AB and heterotrophic biomass (HB) were significantly higher at the oceanic than at the coastal domain, with both biomasses covarying according to HB:AB = 0.33. Whereas the low phytoplankton carbon to chlorophyll a ratio (C_ph:chl a = 38 ± 3) and the high μ = 0.54 ± 0.09 d⁻¹ registered at the coastal stations suggest that phytoplankton was not nutrient limited at this domain, the values (C_ph:chl a = 157 ± 8; μ = 0.17 ± 0.02 d⁻¹) recorded at the oceanic domain point to severe nutrient limitation. However, the high Fv/Fm fluorescence ratios (0.56 ± 0.09) measured at the sea surface in the oceanic domain suggest that nutrient limitation did not occur. To reconcile these two apparently opposite views, it is suggested the occurrence of mixotrophic nutrition of ANF, with heterotrophic nutrition supplying about 75% of carbon requirements.