Primary productivity and sizes of pools of organic carbon in the mixed layer of the ocean

We estimated primary productivity and distributions of carbon in the phytoplankton, micro-zooplankton, and suspended and dissolved matter in various areas of the World Ocean to increase our information about the organic carbon cycle in the surface layer of the sea. Primary productivity ranged from about 0.1 gC m^–2 day^–1 in the Gulf of Mexico to 9 gC m^–2 day^–1 in nutrient-rich water off Peru. Phytoplankton carbon ranged from less than 10 g/l in the former to 750 g/l in the latter and in nutrient-rich water off southwest Africa. Micro-zooplankton carbon usually was less than 50 g/l in all waters, and was dominated by ciliates, copepodids, and copepod nauplii in all areas. Concentrations of particulate carbon ranged from 12 g/l off the east coast of South America to 850 g/l off southwest Africa. Concentrations of dissolved organic carbon varied between 0.5 and 1.5 mg/l in all areas except off Peru, where maximum values of 4.5 mg/l were observed. Turnover rates of carbon by small standing crops of micro-flagellates (1 to 5 longest dimension) and dinoflagellates in nutrient-poor waters were lower than those by large standing crops of diatoms and micro-flagellates in nutrient-rich waters. Concentrations of phytoplankton usually accounted for 20 to 55% and micro-zooplankton for 2 to 30% of the particulate carbon in the surface layer of the sea. Concentrations of dissolved organic carbon were not related to concentrations of particulate carbon in most waters except off Peru, where they appear to be directly related.     

Primary productivity in the Gulf of California

Primary productivity data from 6 cruises in the Gulf of California are summarized. A total of 31 stations with 133 samples was available. The Gulf of California represents a subtropical area with exceptionally high rates of primary productivity. This fact may be accounted for by temporary seasonal upwelling, a wind-mixed water column down to the shallow thermocline, especially in the northern part of the Gulf, and by tidal mixing, especially in the Ballenas Channel between Baja California and the island Angel de la Guardia. Rates of primary productivity in the Gulf of California are comparable to those in areas such as the Bay of Bengal, the upwelling areas off the west coast of Baja California, or North Africa. They are about 2 to 3 times greater than that in the open Atlantic or the open Pacific at similar latitudes.Contribution from the Scripps Institution of Oceanography, University of California, San Diego. This work was part of the Scripps Tuna Oceanography Research Program. It was supported by the Bureau of Commercial Fisheries under contract number 14-17-0007-963. Shiptime was provided from funds of the National Science Foundation.     

Seston composition of the bottom waters of Great Lameshur Bay,St. John,US Virgin Islands

Seston composition [particulate organic carbon (POC), particulate nitrogen (PN), phyto- and microzooplankton numbers and biomass] was investigated in the bottom waters of Great Lameshur Bay, St. John, Virgin Islands (USA), in October, 1970 during Tektite II Mission 17–50. Mean values of 67.4 g POC · I^-1 and 7.2 g PN · I^-1 were determined. A mean phytoplankton carbon content of 42.2 g · I^-1 and zooplankton carbon content of 5.5 g · I^-1 were calculated from counts. The phytoplankton consisted mainly of dinoflagellates 71.2% phytoplankton carbon. Copepods were the dominant zooplankters (61.8% zooplankton carbon), followed by larvaceans (30.9% zooplankton carbon). Organic carbon content of counted zooplankton faecal pellets ranged between 0.4 and 1.6 g · I^-1, and amounted probably to about 15% of the total zooplankton carbon value. Plankton and detritus components as possible food for coral-reef animals are discussed. The ratio carbon: nitrogen of suspended particles is compared to that of sedimented matter.     

Seasonal Fluctuations in the Nutritional Value of Particulate Organic Matter in A Lagoon

Water samples were collected on a fortnightly basis in the lagoon of S. Gilla (Sardinia, Mediterranean Sea) in order to study seasonal nutritional fluctuations of particulate organic matter. the lagoon is characterized by high quantities of suspended matter throughout the year. Thermohaline conditions had no effect on particulate matter quantity and composition, but the quantity as well as quality of suspended particles was drastically affected by the wind, the major effecter of sediment resuspension. As a result of sediment resuspension, seston was always richer in inorganic fraction. However, throughout the year of investigation, most particulate organic carbon was quite appealing for filter feeding communities, although the best POM quality was available during phytoplankton blooming. the phytoplankton pool of suspended matter was just a small fraction of the bulk, accounting for only 13% on average of particulate organic carbon. in terms of energy available in the seston, the highest amount was stored in organic matter heterotrophic fraction, whilst the smallest was to be found in living phytoplankton.     

Particulate Organic Matter Composition in A Semi-Enclosed Marine System

Spatial and temporal changes in the biochemical composition of particulate organic matter in a semi-enclosed marine system (Stagnone di Marsala, Mediterranean Sea) were studied, on a monthly basis, from January to December 1994, in order to assess nutritional value of suspended particles for benthic suspension feeders. According to previous findings, the study site displayed a strong oligotrophy. Chlorophyll-a accounted for a very low fraction of the total suspended matter pool (0.1%), whereas at least 75% of POC was of detrital/heterotrophic origin. POC: PON ratio values indicate that bacterioplankton biomass accounted for a significant fraction of the total POC pool, displaying values comparable to those of the phytoplankton biomass (phytoplankton to bacterial biomass ratio was about (1). Temporal and spatial changes in the biochemical composition of particulate organic matter were rather limited and related to its sources, the main of which is represented by detrital particles released by the Posidonia oceanica (L.) beds. the comparison between our results and those encountered in other coastal lagoons indicates that the low abundance of suspension-feeding organisms observed in the study area is related to the “quality depression” of particles due to the dilution of high quality compounds (i.e., biopolymeric carbon) in a largely inorganic matrix. This result leads us to conclude that, to reach the same amount of high quality particulate food, a suspension feeder mollusc in the Marsala lagoon would need to filter a sea water volume around 3 times higher than in other Mediterranean coastal lagoons.     

Biomass of suspended bacteria over coral reefs

The biomass of bacteria suspended in water flowing over coral reefs at Lizard Island and Yonge Reef (Northern Great Barrier Reef) was estimated by measurement of muramic acid. Values ranged from 20 mg C m^-3 in the open water up to about 60 mg C m^-3 over the reef flat. Direct counts of total numbers of free bacteria were made for comparison. Values of around 2.0x10⁹ cells g^-1 muramic acid showed that there was a good agreement between direct counts and muramic acid content of free bacteria in the open water. In samples containing suspended particulate matter, ratios of direct counts to muramic acid concentration were lower, because bacteria on particles could not be counted. Thus, these ratios were used to indicate the proportions of bacteria attached to particles. Changes in the biomass and numbers of bacteria were determined in water masses identified either by a drogue or fluorescein, as they moved across the reefs. In the zone on the outside of the reef, the number of free bacteria decreased compared to open sea water, but total biomass increased, showing that particulate matter containing bacteria was thrown up into suspension. About 50% of bacteria were attached to particles. Water flowing over the reef flats contained much particulate material with bacteria attached. Bacteria constituted between about 5 and 20% of particulate organic carbon.     

Particulate Organic Matter Composition in A Semi-Enclosed Marine System

Spatial and temporal changes in the biochemical composition of particulate organic matter in a semi-enclosed marine system (Stagnone di Marsala, Mediterranean Sea) were studied, on a monthly basis, from January to December 1994, in order to assess nutritional value of suspended particles for benthic suspension feeders. According to previous findings, the study site displayed a strong oligotrophy. Chlorophyll-a accounted for a very low fraction of the total suspended matter pool (0.1%), whereas at least 75% of POC was of detrital/heterotrophic origin. POC: PON ratio values indicate that bacterioplankton biomass accounted for a significant fraction of the total POC pool, displaying values comparable to those of the phytoplankton biomass (phytoplankton to bacterial biomass ratio was about (1). Temporal and spatial changes in the biochemical composition of particulate organic matter were rather limited and related to its sources, the main of which is represented by detrital particles released by the Posidonia oceanica (L.) beds. the comparison between our results and those encountered in other coastal lagoons indicates that the low abundance of suspension-feeding organisms observed in the study area is related to the “quality depression” of particles due to the dilution of high quality compounds (i.e., biopolymeric carbon) in a largely inorganic matrix. This result leads us to conclude that, to reach the same amount of high quality particulate food, a suspension feeder mollusc in the Marsala lagoon would need to filter a sea water volume around 3 times higher than in other Mediterranean coastal lagoons.     

Nutrient cycling between the water column and a marine sediment. I. Organic carbon

The carbon flow through the sediments at a station located in 18.3 m of water off the Scripps Institution of Oceanography, San Diego, California (USA) was determined. The parameters studied [and their mean rates of input (+) or output (-) to the benthos] were macro-detritus (+0.028 gC m^-2day^-1), fallout of particulate debris (+3.3 gC m^-2day^-1), benthic net photosynthesis during the day (-0.06 gC m^-2 daylight period^-1), burial (0 gC m^-2day^-1), benthic respiration at night (-0.28 gC m^-2 night period^-1), and resuspension (-3.0 gC m^-2day^-1). Resuspension of sediment at this station was found to have a controlling effect on the sediment organic carbon content. Benthic photosynthesis was able to provide 79% of the organic carbon required by the benthos for respiration during the daylight hours. A carbon-flow diagram linking together all of the above measurements is presented.     

Copepod grazing during a declining spring phytoplankton bloom in the Northern North Sea

Grazing rates of larger (Calanus finmarchicus) and smaller (Acartia clausii Pseudocalanus elongatus etc.) copepods on naturally occurring phytoplankton populations were measured during a declining spring phytoplankton bloom. During the initial period, dominated by Chaetoceros spp. diatoms, constant ingestion rates were observed in Calanus finmarchicus at suspended particulate concentrations above 300 g carbon l^-1. Average daily intake during this time amounted to 35 to 40% of body carbon and reached a maximum of 50%. The feeding response of the smaller copepods was not so well defined, although a maximum daily intake of 56% body carbon was recorded. In both groups, feeding thresholds were at particulate concentrations around 50 g C l^-1. The feeding response of C. finmarchicus was correlated with both a change in their own population and in the food cell type. Linear regressions describing the concentration-dependent feeding response were: ingestion rate (IR)=1.16 total particulate volume (TPV)-36.15 during the initial part of the period compared with IR=0.41 TPV-12.18 for the latter period. C. finmarchicus filtered out slightly larger (x 1.2 diameter) particles than the small copepods and, in both groups, some filtering adjustment was made to accomodate to modal changes in the phytoplankton population from 20–30 m to 10 m diameter cells. Particle production during feeding was frequently evident in the smallest size ranges of particles and the ratio of particle production to ingestion rate was greater at low feeding rates.     

Determining the carbon-to-chlorophyll ratio of natural phytoplankton

For field samples, regression analyses of plots of chemically determined particulate organic carbon on chlorophyll are often employed to estimate the algal carbon-to-chlorophyll ratio (F) in the presence of appreciable amounts of nonalgal particulate organic carbon. Spurious results will be obtained, however, if the temporal rate of change of the algae or the nonplant matter reverses its sign during the sampling interval and the samples cannot be ordered correctly in time or space. Previously recognized sources of bias inherent in the chemical approach are also discussed. The great uncertainty of our present knowledge of F is pointed out. It is shown that the concentration of microscopically visible, nonliving particles in the sea isnot known. Renewed studies are suggested as a means of improving on the chemical approach to determining F. The general argument holds for the ratios of nitrogen (particle volume, etc.)-to-chlorophyll, carbon (nitrogen, particle volume, etc.)-to-ATP, and similar conversion factors.Contribution No. 932 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA. The article is a revised and expanded version of a paper contributed at the Joint Oceanographic Assembly, September 1976, at Edinburgh, Scotland, UK.     

Significance of cellular nitrate content in natural populations of marine phytoplankton growing in shipboard cultures

Phytoplankton intracellular nitrate concentrations have been monitored in a 56-h experiment on a shipboard culture of surface sea water from an upwelling region. These measurements were related to parameters of biomass (particulate nitrogen) and nitrate assimilation using the ¹⁵N isotope technique and the nitrate reducase (NR) assay. The procedure for measuring cellular nitrate concentrations is described. This parameter exhibited diurnal variations, ranging from 3.1 to 20.6 ng-at nitrate per g-at particulate nitrogen, and could be correlated positively with NR activity. Nitrogen budgets show that NR activity represents only 12% of nitrate incorporation in organic phytoplankton material when nitrate is available in the sea water. However, upon depletion of the environmental nitrate (zero uptake), NR activity can fully account for the decrease of internal nitrate. From the results, it seems that internal nitrate content is a better index of nitrate consumption by marine phytoplankton than the external concentration of nitrate-nitrogen.     

水体中悬浮颗粒物对十溴二苯醚的吸附/解吸特性研究

采用室内试验法,以沉积颗粒物和浮游植物颗粒(铜绿微囊藻和普通小球藻)作为水体中悬浮颗粒物(SPM),研究了SPM对十溴二苯醚(BDE-209)的吸附/解吸特性.结果表明,沉积颗粒物和浮游植物颗粒对BDE-209的吸附分别在1.5h和0.5h内达到最大,在3h内可以充分达到平衡;沉积颗粒物和浮游植物颗粒对BDE-209的吸附能力存在差异,这与颗粒物中有机质的浓度和类型有关;在实验浓度范围内,BDE-209在SPM上的吸附可用Freundlich等温吸附方程较好地拟合;解吸过程表现出一定的迟滞性,且沉积颗粒物比浮游植物颗粒物具有更明显的迟滞性;pH中性、温度20～30℃的水体有利于SPM对BDE-209的吸附.     

Suspended particulates in the Irish Sea and feeding conditions for fish larvae

The size distributions (2 to 160 m equivalent spherical diameter) of suspended particulate material sampled on two cruises along a transect in the Irish Sea in 1988 are described in relation to hydrographic conditions, chlorophylla concentration and carbon to nitrogen ratios. Particulates were more abundant and larger size modes in the distribution were more evident, in the upper mixed layer of stratified water than in areas where the water column was fully mixed. The detrital content was estimated at 52% of total particulate matter above the thermocline in stratified regions and at around 97% at mixed water sites. In stratified regions the predominance of larger sized phytoplankton and lower levels of detritus is argued to support a more direct and efficient transfer of energy to fish larvae via larger sizes of copepods. Conversely, in mixed areas of high detrital loading the smaller size spectrum of particulates incorporates a less efficient transfer of energy through bacterial cycling and smaller copepods.     

Zooplankton abundance and grazing at Davies Reef,Great Barrier Reef,Australia

Zooplankton abundance and grazing on autotrophic and heterotrophic particulate matter were measured along a transect across Davis Reef (18°5S; 147°39E) and in the back-reef lagoon over tidal and diel cycles during austral winter (August 1984). Zooplankton entering the reef from the surrounding shelf waters decreased in abundance over the reef flat, presumably because of predation. Within the reef lagoon, maximum daytime densities of pelagic copepods occurred during high water, suggesting an external input. At night, water-column zooplankton biomass increased by a factor of 2 to 3 due to the emergence of demersal reef zooplankton. Zooplankton grazing rates on heterotrophic particulate matter (bacteria + detritus and Protozoa) compared to phytoplankton were higher on the reef flat than on the fore-reef or lagoon. Within the lagoon, zooplankton grazing rates on heterotrophic material were maximum during high water, coincident with maximum tidal concentrations of particulate organic carbon. The combined demersal and pelagic zooplankton community were often able to crop 30% of the daily primary production by >2µm phytoplankton. However, >50% of phytoplankton biomass was in cells <2µm, presumably unavailable to these zooplankton. Our particulate production and ingestion measurements, together with zooplankton carbon demand extrapolated from respiration estimates, suggest that the zooplankton community of Davies Reef derives much of its nutrition from detritus.Joint contribution from the University of Maryland, Center for Environmental and Estuarine Studies (No. 2015), and the Microbial Ecology on a Coral Reef Workshop (MECOR No. 19)     

Phosphorus dynamics associated with phytoplankton blooms in eutrophic Mikawa Bay,Japan

Changes in the phosphorus components of the particulate matter in seawater were studied in the eutrophicated waters of Mikawa Bay, Japan, during summer 1981. The contents of particulate phosphorus and hot-water extractable intracellular phosphorus displayed remarkable changes associated with phytoplankton blooms caused by wind-induced or upwelling-associated nutrient enrichment from the bottom water layers. Nanoplankton <10 m accounted for much of the particulate phosphorus (70 to 79% in June and July, and 44 to 78% in August and September); the contribution of large-sized phytoplankton >25 m varied from 9 to 49%, the peak values being attained under red-tide conditions. The capacity for phosphorus storage in cells was low in nanoplankton cells, high in large phytoplankton species. Differences in rates of phosphorus storage and growth between nanoplankton and large phytoplankton accounted for fluctuations in particulate phosphorus which were closely associated with fluctuations in phytoplankton blooms in Mikaw Bay.     

水体中悬浮颗粒物对酞酸酯的吸附和解吸特性

选择水体中的悬浮颗粒物(SPM)--沉积颗粒物和浮游植物颗粒(铜绿微囊藻和普通小球藻),研究它们对邻苯二甲酸二丁酯(DBP)、邻苯二甲酸二异辛酯(DEHP)的吸附和解吸行为.结果表明,SPM对DBP 和DEHP的吸附在很短的时间内即可达到平衡,沉积颗粒物约为2h,浮游植物颗粒约为0.5h;SPM对DEHP的吸附能力明显大于DBP;不同SPM对同一种化合物的吸附能力差异较大,这与颗粒物中有机质的类型和结构有关;在实验浓度范围内,SPM对DBP和DEHP的吸附和解吸等温线具有较好的线性关系,解吸过程存在明显的迟滞现象,且沉积颗粒物比浮游植物颗粒物具有明显的迟滞效应,DBP比DEHP具有较大程度的迟滞效应.     

Clearance rates of the salp Thalia democratica fed naturally occurring particles

Net-captured aggregate stages of the salp Thalia democratica were fed naturally occurring particles in 2-liter bottles onboard ship. The goal was to compare clearance rates of net-captured T. democratica fed natural particles to clearance rates previously determined in laboratoryborn T. democratica fed monocultures of phytoplankton. On a cruise from 19–29 April, 1979, clearance rates were estimated by monitoring the decrease in concentration of particles with a multi-channel Coulter Counter TA II over a particle size range of 2 to 32 m equivalent spherical diameter (E.S.D.). Clearance rates were slightly lower than those measured in the laboratory, decreasing 3-fold over an 18-fold increase in the initial food concentration. Ingestion rates were slightly higher than those measured in the laboratory, increasing about 6-fold over the same range of food concentration. Within the particle concentration range examined (0.08 to 1.34 mm³l^-1), there was no lower threshold of food concentration below which clearance of particles ceased. There was no effect of the concentration of T. democratica on clearance rate over a range of 2.5 to 11.5 salps l^-1. The mean hourly ration was 2.0% of body carbon.     

Microbial clumps in seawater in the euphotic zone of Saanich Inlet (British Columbia)

Viable heterotrophic microorganisms were enumerated to be 3.7±7.3 bacterial cells per microbial clump during summer in the euphotic zone of Saanich Inlet, British Columbia, Canada. Large microbial aggregates were observed, especially after the phytoplankton bloom, when the phytoplankton biomass formed about 1/2 the total suspended organic matter in the sea. The cell number per microbial clump was minimal when the phytoplankton fraction in the total suspended organic matter was almost 0 (i.e., before the phytoplankton bloom), and again when the phytoplankton bloom occurred. The size of the microbial clumps is discussed, particularly in reference to the food chain in the sea.The work was carried out at the Fisheries Research Board of Canada, Biological Station, Nanaimo, during the tenure of a National Research Council Post-doctoral fellowship.     

Estimating carbon flux to pelagic grazers in the ice-edge zone of the eastern Bering Sea

Zooplankton ingestion of phytoplankton carbon in the iceedge zone of the Eastern Bering Sea was measured using a deck incubation approach in 1982. Using further samples collected in 1983, the plant cell carbon to cell volume ratio was estimated at 0.0604 pg m^–3 from an experimentally determined particulate carbon to seston volume relationship. The application of this conversion to the results of experimental incubations of natural plant stocks with net-caught zooplankton produced ingestion rates of 68.8 and 10.26 mg C g^–1 grazer d^–1 for copepods and euphausiids, respectively. Extrapolating these rates to in situ zooplankton biomass at the edge of the seasonal ice pack yielded carbon flux rates through the zooplankton community ranging between 6.5 and 32.8 mg C m^–2 d^–1. This consumption amounted to less than 2% of the daily phytoplankton production in the ice-edge zone.     

Origin and Distribution of Suspended Organic Matter As Inferred From Carbon Isotope Composition in A Mediterranean Semi-Enclosed Marine System

The origin and distribution of suspended organic matter, the trophic features and the stable carbon isotopic composition of particulate organic carbon (POC) were studied monthly in a Western Mediterranean semi-enclosed basin. Sampling stations were selected as a function of wind-exposure and the degree of vegetation cover and then compared with an adjacent unvegetated site. the predominant vegetation was seagrass (Posidonia oceanica and Cymodocea nodosa) and Caulerpa prolifera. Water samples were analyzed for total suspended matter (inorganic and organic fractions), photosynthetic pigments (chlorophyll-a and phaeopigments), dissolved organic carbon, particulate organic carbon and their isotopic composition. Temperature and salinity were also measured at the same sampling sites within range of Mediterranean limits. the suspended organic matter concentration was 1.77 ± 1.55 mg l^-1; the chlorophyll-a concentration was low (0.35 ± 0.24 μg l^-1); the disolved organic carbon concentration was 2,140 ± 2,010 μg l^-1; the particulate organic carbon concentration was 212 ± 106 μg l^-1 and the isotopic composition was 18.77 ± 2.51%_°. There were significant temporal differences except for phaeopigments, POC and its POC isotopic composition, and there were no spatial differences other than for δ¹³C. This picture highlighted a general seasonal trend and trophical features similar to adjacent sea.

Spatial differences in δ¹³C showed that the source of suspended organic matter was different between stations as that between sources and wind-hydrodynamic constraints. In