An offshore buoy as a small artificial island and a fish-aggregating device (FAD) in the Mediterranean

The ODAS Italia 1 oceanographic buoy is moored in the Ligurian Sea, 37 nm from Genoa, along the Genoa-Cape Corse transect (43° 48.90 N–09° 06.80 E), over a 1270 m deep sea bottom. The underwater portion of the buoy is 37 m long and 0.60 m in diameter, acting as a small island for colonization of fouling organisms and as a fish-aggregating device (FAD). The role of the buoy in attracting and maintaining fish assemblages was investigated by visual censuses in different seasons at depths of 0–40 m. Fish from seven families, comprising 12 species, of which three are benthic, were recorded with maximum abundance in summer. Fouling was studied from samples collected on the buoy and on immersed panels. The fouling community of the buoy consisted of 34 algae and 100 animal species, including three fish. The settlement processes of the fouling community on the panels, in particular on those exposed for over 70 months at 12 m and 33 m depth, are described based on counts of settled organisms, the covering index of each taxa and biomass assessments. On the panels, 63 species were identified. The fouling biomass, on the panel submerged for 70 months, assessed as wet weight, reached 2.8 kg/m² at 12 m depth and 4.8 kg/m² at 33 m depth.Observations of benthic organisms settled directly on the buoy were made between 1988 and 1989 and when the buoy was retrieved and brought back to shore on April 15, 1991 after 52 months at sea. At this time, the fouling community along the full 37 m length of the buoy was sampled, and 91 taxa, including 83 species, were identified. Several of the species present on the buoy are shallow, coastal species, some with a very short larval period. Possible ways of colonization by such species are discussed. Despite seasonal changes, the pelagic fish community was more stable over the period of 11 years of study than the benthic community settled on the buoy (that is still developing).     

A quantitative scuba-diving survey of the sublittoral macrobenthos at subantarctic Marion Island

Summary A SCUBA-diving survey of the macrobenthos of hard substrata in the sublittoral zone at subantarctic Marion Island was conducted during March and April 1988. Dense beds (12 kg m^–2) of the kelp Macrocystis laevis occur in depths > 5 m. Durvillaea antarctica is found along the infralittoral fringe and Desmarestia rossi and Durvillaea sp. occur in a narrow zone from 3 m–6 m. Under-storey algae (chiefly rhodophytes) tend to decrease in biomass with depth, with mean values of 1.57 kg m^–2 at 5m, 0.75 kg m^–2 at 10m and 0.49 kg m^–2 at 15 m. Encrusting coralline algae are particularly abundant in shallow areas (¯x = 0.92 kg m^–2) but are insignificant in deeper areas. Total biomass of macrozoobenthos increased with depth with mean values of 0.12 kg m^–2 at 5 m, 0.34 kg m^–2 at 10 m and 0.46 kg m^–2 at 15 m. Polychaetes, crustaceans, echinoderms, molluscs, sponges and bryozoans dominated the macrozoobenthos in terms of biomass. Approximately 200 species of macrobenthic animals were recorded and numerically, polychaetes, crustaceans, molluscs, nematodes and echinoderms dominated. The sublittoral benthos at Marion Island is compared with that occurring at other subantarctic and Antarctic islands, in particular, the Kerguelen Island group. Zoogeographic trends and the possible effects of nutrient input from seabird guano are briefly discussed.     

Role of planktonic bacteria in productivity and cycling of organic matter in the Eastern Pacific Ocean

Total number, biomass, production, and respiration of bacterioplankton were measured in oligotrophic, mesotrophic and eutrophic waters of the Eastern Pacific. Total number of bacteria in the upper mixed layer and in the upper thermocline boundary layers varied from 30–60.10³ ml^-1 in oligotrophic waters to 100–400.10³ ml^-1 in mesotrophic waters of fronts and divergences, and to 1–2,5.10⁶ ml^-1 in eutrophic waters of coastal upwellings. Wet biomass varied from 5–10 mg l^-1 in oligotrophic waters, to 50–200 mg l^-1 in mesotrophic waters, and to 1–2 g m^-3 in eutrophic waters. Below the layer of maximum temperature gradient i.e. below 35–50 m, bacterioplankton density decreased 5–10 times. P/B coefficients per day were highest in the oligotrophic surface water ( 1), and lowest in the eutrophic ones (0.2–0.4). In mesotrophic waters they were intermediate (0.4–1.0). the stock of labile organic matter (LOM) accessible to microbial action varied from 0.3 to 1.6 mg Cl^-1. Its highest value occurred in the upwelling area. The stock of LOM does not noticeably decrease from the euphotic zone to a depth of 2 000 m. Its turnover time varied from 5 to 45 days in surface waters, and 30–50 years in deep oceanic waters. The role of bacterioplankton in productivity and in cycling of organic matter in surface — and deep oceanic waters is discussed.     

Controlled artificial upwelling in a fjord using a submerged fresh water discharge: computer and laboratory simulations

To enhance the conditions for producing shellfish in coastal waters, the possibility of employing artificial upwelling of nutrients is explored. The effectiveness of a submerged discharge of fresh water is studied by means of a numerical buoyant plume model, BJET, and laboratory simulations. An optimisation of the entrainment of deeper, nutrient-rich water to the proper intrusion depth is demonstrated. The studies show that a downward directed jet of fresh water below the euphotic zone can lift significant amounts of nutrients to the primary production near the surface. The outlet must be large enough to lift the deeper water through the pycnocline to the desired depth of primary production. The results are applied to a possible discharge arrangement in the Samnangerfjord, to the east of Bergen, Norway, using field data from 1999. With a discharge of up to 8 m³/s of fresh water at 35 m depth, the entrainment of deeper water into the buoyant plume, up to the 15 m depth, is 12 to 13 times as much. The chosen arrangement could give an expected vertical transport to the euphotic zone of 467 kg d^–1 N, 46 kg d^–1 P and 555 kg d^–1 Si during the summer growth period. This includes periods of coastal downwelling with greatly reduced values of nutrients, but not periods of strong, deep stratification with deeper intrusions. Further optimisation is possible using active controls of the discharge system.     

The diversity,biomass and production of zooplankton in Lake Inarijärvi

About 650 zooplankton samples were collected from Lake Inarijärvi in 1977–1979 from the littoral and pelagial zones of the lake. One hundred and twenty-three zooplankton taxa were found and most of them can be considered euplanktonic.The most important species were Holopedium gibberum, Daphnia cristata, Cyclops spp. and Eudiaptomus spp. Mean pelagial zooplankton biomass was 0.29 g m^–3 in the 0–5 m depth zone, 0.17 g m^–3 in 5–10 m and 0.11 g m^–3 in 10–20 m.The zooplankton biomass at a sandy shore was about 0.09 g m^–3, at a stony shore 0.05 g m^–3 and at a vegetated shore 0.76 g m^–3. About 70% of the whole zooplankton production consisted of crustaceans.The sum of herbivore and carnivore zooplankton production in the pelagial area during the summer was 210–330 kg ha^–1 × 3 months.     

Vertical Distribution of the Far East Trepang Apostichopus japonicusin Vostok Bay,Sea of Japan

The abundance, weight, and age structure of aggregations of the Far East trepang Apostichopus japonicuswere studied in Vostok Bay, Sea of Japan, during the first part of September 2000. The highest density of aggregations (8.3 ± 0.5 ind/m²) and biomass (131.88 ± 10.50 g/m²) of animals was characteristic for coastal sites at the depth 0.5–1.5 m. With anincrease in depth the biomass and density of A. japonicusreduced 1.5 and 3 (depth 5–6 m), 220 and 830 times (depth 8–15 m) respectively. The ratio of large animals concurrently increased in the aggregation structure. The specifics of the distribution and spatial variability of the aggregation structure are discussed in relation to uncontrolled fishing of holothurians of large sizes.     

The influence of coastal upwelling on the distribution of Calanus chilensis in the Mejillones Peninsula (northern Chile): implications for its population dynamics

A field experiment was carried out in October 1998 during active upwelling in a coastal area off the Mejillones Peninsula (23° S). Zooplankton was sampled at day and night, during two subsequent days at 4 stations inside and outside of the upwelling plume. Three depth strata were sampled: 0–20 m, 20–80 m and 80–200 m. Oceanographic data were obtained in a grid of 23 stations using a CTDO, a fluorometer and a Doppler current meter. Calanus chilensis was mostly represented by late stages, i.e. copepodid C5 and adult males and females. There were no day/night effects on vertical distribution, and abundance was significantly higher inside the upwelling plume in the upper 20-m layer at nearly 14 ind. m^–3, compared to ca. 5 ind. m^–3 outside the upwelling plume. Temperature at 10 m depth and biomass, estimated from stage numbers and their mean dry weights, were used to estimate growth and daily production of Calanus at temperature-dependent rates. The potential loss of biomass from the upwelling center because of advection in the upwelling plume was estimated from current data in the Ekman layer and biomass density. The mean cross-shelf component of the current was estimated at 10.4 km d^–1 within the upwelling plume. This yielded a loss of biomass of 9.7 mg dry weight m^–2 . Production, estimated by a temperature-dependent approach, ranged between 44 and 35 mg dry weight m^–2 d^–1, at mean temperatures of 14.6 °C and 15.8 °C inside and outside of the upwelling plume respectively. Within the plume, as much as 22% of daily production may be advected offshore. However, a higher concentration of biomass in the upwelling plume allowed a greater production compared to surrounding areas. A mass balance approach suggests that advective losses may not have a major impact on the C. chilensis population, because of very high daily production at temperature-dependent rates.     

Phytoplankton biomass and species composition in a Mediterranean coastal lagoon

Phytoplankton populations were investigated weekly at a central station in the Fusaro lagoon (Mediterranean Sea) from 27 November 1989 to 18 June 1990 to assess species composition, temporal succession and standing stock of the different species. Chlorophyll concentrations varied from 1.2 to 73.2 µg 1^–1 in surface waters, and from 1.3 to 53.5 µg 1^–1 at the 4.5 m depth. Phytoplankton communities were dominated by Prorocentrum micans Ehrenberg in December and January, and by small-sized diatoms in the rest of the sampling period. In surface waters, a maximum biomass of 9.5 mg C 1^–1 was measured in January, in correspondence with high concentrations (8.1 × 10⁶ cells 1^–1) of P. micans, whereas an abundance peak of 159.9 × 10⁶ cells 1^–1 was registered on the last sampling date due to a massive bloom of a very small diatom, Minutocellus polymorphus (Hargraves & Guillard) Hasle, von Stosch & Syvertsen. On the whole, phytoplankton populations of the Fusaro lagoon showed distinct characters as compared to those of southern Tyrrhenian coastal waters and of other lagoons.     

Growth of water hyacinth (Eichhornia crassipes (Mart.) Solms) in the middle Paraná River (Argentina)

The productivity of water hyacinth in the middle Paraná River was measured at three initial biomass levels by means of periodic harvesting of plants contained in 2 m² floating baskets. The measuring period extended between August 1981 and July 1982. Initial biomass densities were 2, 5 and 10 kg (fw) m^–2.Compared with inner island ponds, flowing waters connected to the main river showed better conditions for water hyacinth growth. The duration of the growth period was some 30% longer than in island ponds, probably due to the relatively warmer river waters. Productivity ranged between 108 and 164 g (fw) m ^–2 d^–1 (annual average) for the lower and higher initial biomass values, respectively.     

Summer bacterio- and zooplankton in coastal waters of Svalbard

The distribution of bacterio- and zooplankton was studied in West Spitsbergen (Svalbard) coastal waters in August 2009. The bacterial abundance and biomass varied from 110 to 3360 × 10³ cells/ml and from 0.66 to 44.93 mg C/m³, respectively. These values were the highest in the surface water layers and decreased with depth. The greatest abundance of bacteria (5099 individuals/m3) was found in the freshened and warm water layer in Isfjorden. The highest biomass of zooplankton (182.41 mg C/m³) was registered in the zone of interaction of coastal and transformed Atlantic waters in the archipelago inner shelf. The spatial variations of abundance and biomass of the planktonic communities were related to the thermohaline characteristics of the water masses and, probably, to microalgal blooms.     

A quantitative diving survey of benthic vegetation and fauna in Lake Kariba, a tropical man-made lake

SUMMARY. 1. The biomass distribution of submerged vascular vegetation and benthic fauna were investigated by diving in Lake Kariba. The vegetation was well correlated with transparency of the water. Maximum biomass (1400 g dryweight m^?2) and a depth penetration of 6 m were found in areas little influenced by river inflow, while these were only 110 g m^?2 and 2 m, respectively, in the basin receiving water from the Zambezi river. 2. The lake is mesotrophic-oligotrophic. The total biomass for the lake was 101,000 tons dryweight of rooted vegetation composed of Lagarosiphon ilicifolius Oberm. (52%), Najas pectinata (Parl) Magnus (33%), Vatlisneria aethiopica Frenzl (11%), Ceratophyllum demersum L. (3%) and Potamogeton octandrus L. (0.5%). Average plant biomass for the potentially colonizable depth zone of 0-12 m and for the total lake amounted to 79.9 g m^?2 and 18.8 g m^?2, respectively. 3. The distribution of the benthic fauna generally followed that of the vegetation. The total animal biomass of 118,840 tons dryweight. including shells, consisted of mussels (95.8%), snails (4.1%) and insect larvae (0.1%). Four species of mussels were found: Caelatura mossambicensis (von Martens) (81% of mussel biomass), Corbicula africana (Krauss) (9%). Mutela dubia (Gmelin) (5%) and Aspatharia wahlbergii (Krauss) (5%). Among the snails Melnoides tuberculuta (Müller), Cleopatra spp. and Bellamya capillata (Frauenfeld) dominated. 4. The average animal biomass was high compared to most other lakes perhaps due to lack of predators. For the colonizable 0-12 m depth interval and the total lake it was 96.2 g m^?2 including shells (15.0 g m^?2 shell-free dryweight) and 22.6 g m^?2 including shells (3.4 g m^?2 shell-free dryweight). respectively. Biomass of plants and animals was even higher prior to the recent lowering of the water level by 7 m, which was estimated to have stranded 84,000 tons of mussels on the shore.     

Denitrification in the water column of an intermittently anoxic fjord

Denitrification was studied in the water column in the Bunnefjord, inner part of the Oslofjord in southern Norway, using a ¹⁵N-technique (the isotope pairing method). The fjord is 150 m deep and during our surveys in September–December 1998 hydrogen sulphide was present in the deep water below 80 m. No significant denitrification was found in water samples from the surface layer (4 m depth), but high rates were observed within a deep density gradient between 62 and 78 m depth. Oxygen concentration within this layer was low (<21 mmol m^–3), and the concentration of NO₃ decreased from ca. 15 mmolm^–3 at 62 m depth to not detectable below 78 m. Pronounced peaks of NO₂ up to 4.4 mmol m^–3 were observed at 70–78 m depth. The maximum denitrification rate of 1.5 mmol N m^–3 d^–1 was observed at 70 m depth. Integrated for the whole layer, the denitrification rate was 13 mmol N m^–2 d^–1. A significant linear correlation was found between the denitrification rate and the ambient nitrate concentration which indicated that the rate was primarily controlled by the availability of nitrate in the O₂-poor water. Compared to rates reported for coastal water, denitrification in the water column in the Bunnefjord was high and the process appears to be a major sink of bioavailable nitrogen in the fjord.     

Variability of phytoplankton distribution and primary production around Elephant Island,Antarctica, during 1990–1993

The distribution and abundance of phytoplankton within a sampling grid of 50×10³ km² around Elephant Island were determined from early January to mid-March of 4 successive years, 1990–1993. The number of stations where physical-optical-biological data were obtained from the surface to a maximum of 750 m ranged from 74 in 1990 to 206 in 1993. Contour maps of chlorophyll-a (chl-a) concentrations showed marked mesoscale patchiness that varied from month to month and also interannually. The distribution patterns for chl-a were similar when plotting surface concentrations or integrated values to 100 m. Three major zones could be distinguished that differed in both physical and biological characteristics. Stations in the northwest portion of the grid (Drake Passage waters) and in the southeast portion of the grid (Bransfield Strait waters) showed the most pronounced interannual variations, with phytoplankton biomass and rates of primary production being considerably higher in 1990–91 than in 1992–93. The central portion of the sampling grid, which included the major frontal system north of Elephant Island, showed the smallest interannual variations in both biological and physical parameters and the highest rates of primary production. Phytoplankton biomass and rates of primary production were correlated with depth of the upper mixed layer (UML), which in turn was correlated with the measured wind stress. The mean depth of the UML was 50 m, while the mean depth of the euphotic zone was 90 m. Using the measured mean surface solar irradiance (550 Einsteins m^–2 s^–1), the mean irradiance experienced by cells in the UML of 50 m would be around 105 E m^–2 s^–1, which is similar to the measured I_k (light saturation) value for photosynthesis (101 Em^–2 s^–1). The mean value from all cruises for chl-a in surface waters was 0.7 mg m^–3, while the mean rate of primary production was 374 mg Cm^–2 day^–1.     

Role of plankton in the turnover of organic matter on the Great Barrier Reef,Australia

Number, biomass and production of phytoplankton, bacteria, micro- and mesozooplankton and turnover of labile and stable organic matter were measured in waters over some Capricornia round reefs, and over the reefs of Lizard Island. Primary production was 10 to 40 mg C m^–3 d^–1 but was lower over the living reefs. Microbial wet biomass in reef waters varied from 100 to 500 mg m^–3, and production from 4 to 68 mg C m^–3 d^–1, which was commensurable with primary production. The biomass of microzooplankton (ciliates, zooflagellates and larvae) in waters of Lizard Island reefs reached 100–300 mg m^–3. Mesozooplankton biomass at night in reef waters of Heron Island varied from 200 to 800 mg m^–3. Its composition depended upon the tide phase. PB coefficients in bacterioplankton were 0.3 to 1.2 per day. The food demand of bacterioplankton in waters over the reefs was 5 to 20 times higher than the primary phytoplankton production. Labile organic matter (LOM) doubled in waters after it stayed over living reef for several hours. The turnover time of LOM in reef waters was as short as 1–2 weeks.     

An experimental study of old-field succession in relation to different environmental factors

From 1984 to 1986, old-field succession on sterilized sand and loam was studied under different water- and nutrient regimes. Within one month, moss and phanerogam species appeared on all experimental plots but further succession was rather varied. Salix species established quickly on loam and formed within 3 years a shrub layer up to 3 m in height. On sand, woody plant species were observed only at a high ground-water level. On loam, the well-known old-field succession from short-living therophytes to long-living phanerophytes of clearings and woodlands proceeded very quickly. In contrast, on sand, therophytes, hemicryptophytes and herbaceous chamaephytes of ruderal- and grassland communities were still dominant after three years. A high ground-water level as well as mineral fertilization had sometimes positive, sometimes negative effects on this succession. Periodic estimates of cover, made during the succession were supplemented at the end of the experiment by the measurements of phytomass and bioelement storage. The highest amount of biomass was measured on the three loamy soils where shrub layers were well developed. In comparison with data published elsewhere, the above-ground biomass of 2.2–2.8 kg dry matter m^-2 and the below-ground biomass up to 7.2 kg dry matter m^-2 were both extraordinarily high. Over the three years, the vegetation on sandy soils accumulated between 1.2 and 5.1 g N m^-2 yr^-1 and on loamy soils between 17.1 and 24.7 g N m^-2 yr^-1.     

Vertical distribution and seasonal variation of eelgrass beds in Iwachi Bay,Izu Peninsula,Japan

The vertical distribution and population structure of eelgrass beds were surveyed in Iwachi Bay, along the Pacific coast of central Japan. Samplings were conducted from May through November 1977 by SCUBA. Eelgrass was distributed between 3 and 11 m in depth. The relative light intensity at 12 m depth was 11% at the lower range. The highest population density was 290 shoots/m² in September and the fresh weight of biomass was 888 g/m² in July at 7 m depth. The maximum mean leaf area index was about 3 at 10 m depth in July. The ratio of reproductive shoots to the total shoots was about 36% at 7 m depth in June. Eelgrass showed good growth at 7–10 m depth, which is comparatively deeper than other eelgrass habitations. The high values of water transparency and sunshine duration, as well as solar radiation compared with other localities was believed to contribute to the growth of eelgrass in deeper waters in Iwachi Bay.     

Effect of nutrient supply on growth of blue mussels (Mytilus edulis) in a landlocked bay

The growth of blue mussels (Mytilus edulis) was studied in the landlocked bay Hopavågen in central Norway for 3 years, of which in 2 years (1998 and 1999) nutrients were added to increase the primary production. Nutrients (N:Si:P) were added daily from May to October in 1998 (molar ratio 15:5:4.1) and 1999 (molar ratio 16:8:1). The doses of nutrients correspond to 0.4 and 0.8 g P l^–1 day^–1 in 1998 and 1999 respectively. The growth of blue mussels (Mytilus edulis) in 1997 was followed at four depths (1, 2, 4 and 7 m). In 1998 and 1999 growth was followed at 2 and 10 m depths at four locations in Hopavågen and at a control station outside on the coast. The nutrient supply in 1998 only slightly increased the algal biomass (chlorophyll a), whereas mean daily primary production during the summer remained at the same level as the previous year. The increased nutrient supply in 1999 caused a nearly 50 and 100% increase in mean summer biomass and daily primary production, respectively. The growth of blue mussels in Hopavågen in 1997 and 1998 was within the same size range during the summer. In 1999 the shell length of blue mussels kept at 2 m depth was significantly higher than in the previous year at end of the growth season. The recorded growth was also significant higher than for mussels at 2 m depth at the control station. No difference in shell length was observed on mussels grown at 10 m depth in Hopavågen and in the control stations in 1998 and 1999. A higher tissue content was found in blue mussels grown at 2 m depth in Hopavågen, both in 1998 and 1999 when compared to the control groups. At 10 m depth no differences were recorded.     

Micronekton and macrozooplankton in the open waters near Antarctic ice edge zones (AMERIEZ 1983 and 1986)

Summary Micronekton and macrozooplankton assemblages (0–1000 m) were sampled from the open ocean in the vicinity of marginal ice zones in the southern Scotia and western Weddell Seas using midwater trawls. Small regional differences in species composition were found in the differing hydrographic settings with the Scotia Sea being slightly more diverse. Most species exhibited broad vertical ranges with no distinct pattern of vertical movement. Exceptions were mesopelagic fish and Salpa thompsoni which undertook diel vertical migrations. Biomass was high (2.4–3.1 g DW/m²), comparable to Pacific subarctic waters. Euphausia superba and Salpa tompsoni were the numerical and biomass dominants, representing over 50% of the total numbers and standing stocks. In terms of biomass, euphausiids were the most important group at shallow depths (0–200 m) but were surpassed by salps in the Scotia Sea and mesopelagic fish in the Weddell Sea when all depths down to 1000 m were considered. Pelagic fish biomass (3.3–4.4 g WW/m²) greatly exceeded published estimates for birds (0.025–0.070 g WW/m²), seals (0.068–0.089 g WW/m²) and whales (0.167 to 0.399 g WW/m²), making mesopelagic fish the most prevalent krill predators in the Antarctic oceanic system.     

Variation of fatty acid profile with solar cycle in outdoor chemostat culture ofIsochrysis galbana ALII-4

Outdoor chemostat cultures of the marine microalgaIsochrysis galbana at constant dilution rate (0.034 h^–1 ) have been carried out under different weather conditions. Steady-state biomass concentrations were 1.61±0.03 kg m^–3 in May and 0.95±0.04 kg m^–3 in July, resulting in biomass output rates of 0.54 kg m^–3 d^–1 and 0.32 kg m^–3 d^–1 in May and July, respectively. Two patterns of daily variation with the solar cycle were observed in the fatty acid content. Saturated and mono-unsaturated fatty acids (16:0 and 16:1n7) show significant variation with the solar cycle, associated with short-term changes in environmental factors. Palmitic and palmitoleic acids are generated during daylight and consumed during the dark period. However, polyunsaturated fatty acids do not show a significant response to the solar cycle and their changes are associated with long-term variation in environmental factors. The maximum EPA productivity was obtained in May, 14.1 g m^–3 d^–1, which is close to that found in the literature for indoor continuous cultures. Nonetheless, the outdoor EPA content (up to 2.61 % d.wt) was lower than the indoor EPA content from a previous study (5% d.wt).     

Variations saisonnières de la production primaire dans les eaux de surface de la rivière du Saguenay

Experiments of primary production were carried out at weekly intervals in the surface waters at one station (maximum depth of 20 m) in the Saguenay River, near Chicoutimi, during May–December 1978. The photic zone was very thin (maximum depth of 2 m). Phosphates are very low during the season sampling (maximum of 0.1 µat-g.^–1). Maximum of production rates and biomass are respectively 3.5 mg C.m^–3.h^–1 and 3.7 mg.m^–3. The river receives both industrial and urban runoff. Trace metals (Mercury, Copper, Lead, and Iron) seemed to be one of the important limiting factors for phytoplankton growth.