Epipelagic planktonic bioluminescence in the marginal ice zone of the Greenland Sea

Epipelagic planktonic bioluminescence in the marginal ice zone of the Fram Strait was investigated during April and May 1989. Vertical profiles of bioluminescence potential were measured using a HIDEX (High Input Defined EXcitation) design bathyphotometer. Mesozooplankton samples were collected with oblique tows using a 153 m mesh net. The amount of bioluminescence produced by mechanical stimulation of individual organisms was measured using an integrating sphere and photon-counting system. These measures of bioluminescence potential along with estimated abundances of bioluminescent organisms allow an independent estimated of integrated epipelagic bioluminescence potential. The zooplankton community structure was relatively simple in this region and the number of bioluminescent species correspondingly small. The majority of epipelagic bioluminescence was produced by copepods (Metridia spp.), larvaceans (Oikopleura spp.), euphausiids (Thysanoeassa spp.) and ostracods (Conchoecia spp.), although the relative contribution of these organisms to the overall bioluminescence potential varied considerably with time and location over the 2 mo period. Bioluminescent dinoflagellates were rare and did not contribute significantly to epipelagic bioluminescence. Integrated bioluminescence potential in the water column was significantly correlated with zooplankton biomass, but not with any other measured environmental parameter. No enhancement of chlorophyll, zooplankton biomass or bioluminescence was observed in the marginal ice zone compared to the adjacent open waters of the Fram Strait during this spring cruise.     

Diel patterns of planktonic bioluminescence in the northern Sargasso Sea

Day-night changes in the vertical distribution, intensity, and size of bioluminescence flashes were investigated during a series of cruises to the northern Sargasso Sea in 1987 and 1988. Overall, depth integrated bioluminescence potential and flash density estimated from in situ measurements with a pumping bathyphotometer were 2 to 5 x higher at midnight than at midday. Depths from 50 to 100 m exhibited the most substantial day to night increases in bioluminescence potential and flash density. When classified by flash size (photon output per flash event), the increase from day to night was significant for all flash sizes, but was most dramatic for small flashes producing <7 x 10⁸ photons flash^-1. Bioluminescence potential and flash density increased 2 to 3 x during bathyphotometer measurements made at dusk. Bioluminescent light budgets estimated from day and night net collections in May and August 1987 also predicted 2.5 x higher nighttime than daytime mesoplankton bioluminescence. However, large bioluminescent taxa (mesoplankton) capable of significant vertical migrations only contributed on the order of 15% of the total bioluminescence in surface waters. Our results do not support the idea that most of the nightly increase in bioluminescence potential and flash density are due to vertical migration of bioluminescent organisms; rather they are consistent with an alternate view that photoinhibition of bioluminescent flashing by dinoflagellates may be primarily responsible for the diel patterns.     

Nitrogenous nutrition in the euphotic zone of the Central North Pacific Gyre

The uptake rates of the three nitrogen compounds ammonium, nitrate, and urea were measured in the oligotrophic North Central Pacific Gyre in August–September 1985. The measurements were performed by using ¹⁵N-labelled substrates and incubating for short-time periods (3 to 4 h) under simulated in situ conditions. Ambient concentrations of the nitrogenous nutrients were generally below 0.10 mol l^-1. The average total daily nitrogen uptake rate, integrated over the euphotic zone, was 12.5 mmol N m^-2 d^-1. Diel studies in the upper water mass resulted in a calculated phytoplankton growth rate of 1.3 d^-1. Ammonium was the dominating nutrient, accounting for on the average 54% of the total nitrogen uptake, while urea uptake represented 32% and nitrate 14%. Ammonium uptake rates at a coastal station off the Hawaiian Islands were very close to the rates found at the oceanic station. Organisms <3 m dominated the nitrogen assimilation, being responsible for about 75% of the ammonium uptake. The nitrogen uptake rates in this study seem to be higher than those found by earlier investigations in the area, but correlated well with other productivity measurements performed during the same cruise.     

Pattern of planktonic bioluminescence in the northern Sargasso Sea: Seasonal and vertical distribution

The vertical structure of bioluminescence potential (BPOT) and flash density (FD) were measured on five cruises to the northern Sargasso Sea in 1987 and 1988. Depth-integrated (0 to 150 m) BPOT did not vary seasonally, remaining within the range 9 to 15 × 10¹⁵ photons m^–2 in all months sampled. Conversely, depth-integrated FD was significantly higher (> 2 × 10⁵ flashes m^–2) during winter (November and March) than during summer (< 9 × 10⁴ flashes m^–2 in May and August). The vertical patterns of BPOT and FD were well correlated within a single profile, more highly so in summer than in winter. Despite intracruise variability in the vertical pattern of BPOT and FD, there were clear summer-winter differences in the vertical distribution of BPOT and FD. During winter, BPOT and FD were maximal and relatively uniform throughout the surface mixed layer; for example in November they declined sharply within the thermocline at 130 to 150 m. During summer, BPOT and FD were greatest (12 to 25 × 10¹³ photons m^–3 and 600 to 1 200 flashes m^–3, respectively) at subsurface depths. Commonly in summer, the upper depth limit of high BPOT and FD occurred at the base of the surface mixed layer (10 to 40 m) and the lower depth limit was located at the base of the subsurface fluorescence maximum (usually at 100 to 120 m).     

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.     

Total labile carbon in the euphotic zone of the Baltic Sea as measured by BOD

Total labile organic carbon was estimated in the euphotic zone of the Baltic Sea by a method based on the determination of the biological oxygen demand (BOD); the method is extremely simple, requires little sample water, and is quite precise (±4.5 μmol O₂/l or ±40.4 μgC_lab/l). The amount found was 1,150 μgC_lab/l, or about 29% of the total organic carbon. At 1 m, a relationship between biological activity, as measured by chlorophyll content, and the amount of labile carbon was found: ln C_lab (μg/l)=ln 6.87 (μg/l) +0.272 ln chlorophyll (μg/l), with a correlation coefficient of 0.86, which is significantly greater than zero at the 0.001 level.     

Bacterioplankton in the coastal euphotic zone: Distribution,activity and possible relationships with phytoplankton

Bacterioplankton were studied in the euphotic zone of the Southern California Bight, USA, with special attention to biological factors affecting bacterial distribution and activity. Measurements were made of bacterial abundance, thymidine incorporation into acid insoluble material, primary production (particulate and dissolved), chlorophyll, phaeopigments, total microbial ATP, particulate organic carbon and nitrogen, dissolved organic carbon, dissolved primary amines, and glucose and thymidine turnover rates. The data were analyzed by pairwise rank correlations with significance tested at the P<.005 level. Bacterial abundance and thymidine incorporation both declined progressively with increasing distance from shore (to 100 km); similar trends occurred for the phytoplankton, with several stations having subsurface maxima. Bacterial abundance, thymidine incorporation, and thymidine and glucose turnover rates were all significantly correlated to each other, suggesting they are comparable as relative measures of bacterial activity. Thymidine incorporation per cell, an indicator of specific growth rate, was not correlated to bacterial abundance, suggesting density independent specific growth rates. Bacterioplankton growth rate was evidently influenced more by the standing stock of phytoplankton than by the primary production of the phytoplankton. Thus, bacterial growth may possibly be stimulated by leakage of dissolved organic matter not so much from healthy photosynthesizing cells as from phytoplankton being disrupted and incompletely digested during predation by the zooplankton and nekton.     

Planktonic bioluminescence in the pack ice and the marginal ice zone of the Beaufort Sea

An icebreaker cruise into the Beaufort Sea in the fall of 1986 provided a unique opportunity for studying planktonic bioluminescence in ice fields and in the marginal ice zone. Bathyphotometer casts (bioluminescence intensity, seawater temperature, beam attenuation coefficient, and salinity) and biological collections were made to a depth of 100 m. A light budget, which describes the planktonic species responsible for the measured bioluminescence, and a dinoflagellate species budget were constructed from the mean light output from luminescent plankton and plankton counts. The vertical distribution of bioluminescence among the ice stations was similar. The maximum intensities were 2 to 8×10⁶ photons s^-1 cm^-3 in the upper 50 m of the sea-ice interface. The marginal ice zone station (MIZ) exhibited a maximum intensity of 2 to 3×10⁸ photons s^-1 cm^-3 between 5 and 30 m depth. At Ice Station 2, Metridia longa and their nauplii contributed approximately 80% of stimulable bioluminescence in the upper 10 m but, overall, Protoperidinium spp. dinoflagellates contributed most of the light to a depth of 100 m. In the MIZ, Protoperidinium spp. dinoflagellates contributed 90% of the light within the upper 10 m, decreasing to 43% of the contributed light at a depth of 40 m. Below 40 m, dinoflagellate bioluminescence decreased to a few percent of the total to a depth of 90 m. Metridia spp. copepods contributed more than 50% of the light at depths from 40 to 90 m. Ostracods, larvaceans, and euphausiid furcilia contributed <1% of all bioluminescence at all depths sampled. Correlation analyses between measured bioluminescence (photons s^-1 cm^-3), the number of bioluminescent dinoflagellates and the light budget for the MIZ indicated highly significant associations: r=0.919, p=0.001, and r=0.912, p<0.001, respectively (Student's two-tailed t-tests). Bioluminescence was negatively correlated with seawater salinity at all stations (p=0.001). Maximum bioluminescence was measured in the less saline surface waters at all stations.     

Phytoplankton pigments in the gut concents of planktonic copepods from coastal waters off southern California

Phytoplankton xanthophylls in the gut contents of the copepods Calanus pacificus, Corycaeus anglicus, and Paracalanus parvus, collected from 5 stations off San Onofre, California, in June 1982, were measured by reverse phase, high-performance liquid chromatography (HPLC). The dinoflagellate pigment, peridinin, was usually the most abundant xanthophyll in the guts of all three species of copepods. Evidently, feeding was principally on dinoflagellates (which dominated the phytoplankton biomass). The level of feeding activity, rather than the class of phytoplankton ingested, seemed to differentiate the behaviors of the copepods. Xanthophyll content per unit copepod wet weight was higher in Corycaeus anglicus and Paracalanus parvus than in Calanus pacificus. Chlorophyll a fluorescence of the copepod gut contents was measured in conjunction with the analysis of gut xanthophylls. The xanthophyll content of the gut varied directly with the concentration of chlorophyll a in the gut. Xanthophyll content was not related to the concentration of pheopigments in the gut. Apparently, the xanthophylls that were detected were due to the presence of recently ingested phytoplankton biomass.     

Seasonal study of planktonic copepods and their benthic resting eggs in northern California coastal waters

Few investigations have examined the occurrence of zooplankton resting eggs in the sea bed of waters deeper than 20 m. In this study the distribution and abundance of planktonic copepods and their benthic resting eggs in coastal waters off northern California, U.S.A., were determined and related to environmental parameters (temperature, salinity, depth, and sediment grain size). Sediment cores, net tows, and CTD profiles were obtained in April and October 1989, and February, April, and October 1990. Water depths in the study area ranged from approximately 60 to 120 m. The mean abundance of eggs was as high as 1.2×10⁵ m^-2 for Acartia clausi Giesbrecht and 1.9×10⁵ m^-2 for Tortanus discaudatus Thompson and Scott. These egg concentrations are comparable to those reported previously for shallower more protected regions. The abundance of eggs in the sediments decreased with increasing depth of the water column. For the region as a whole, eggs were least abundant in muddy sediments. The mean abundance of eggs in the sea bed also varied seasonally and annully. Benthic resting eggs of A. clausi were more abundant in April 1989 than in April 1990, and adults of the species were never found in the plankton samples. The lack of adults is not unusual since results of previous studies indicate that A. clausi is a cold-water species, and in this region water temperatures are colder in summer, than in winter, due to upwelling. Temperature and salinity data indicated that the upwelling season had commenced by the time of the April 1990, but not the April 1989 sampling. Thus, the reduced abundance of benthic eggs in April 1990 may have been due to egg-hatching in response to reduced temperatures. The results suggest that the presence of A. clausi in coastal waters off northern California is linked to recruitment from benthic resting eggs.     

Variability of biochemical composition and size distributions of seston in the euphotic zone of the Bay of Biscay: implications for microplankton trophic structure

Particle-size distributions and several biochemical components of seston were studied from October 1986 to December 1987 in surface waters of the Bay of Biscay. Variance partitioning of hydrographic and seston parameters indicates that, although seasonal variability related to the thermal cycle was important, most of the differences in seston concentration were caused by spatial components of variance. Changes in the vertical structure of the water column appeared to be the principal source of variability. Phytoplankton blooms in spring and fall and the effects of the coastal upwelling and the thermocline during summer were traced using seston concentration and biochemical composition. The different seston measurements were scaled according to their correlations with total concentration and particle size. Photosynthetic pigment biomass was related to both large and small partieles. However, concentrations of particulate proteins, lipids and carbohydrates were more related to small particles. Although not specifically analysed, low concentrations of inorganic particles and detritus can be expected in the study area, thus most of the particulate organic matter was associated with small, non-pigmented organisms, particularly in surface waters during the period of thermal stratification. The variability in concentrations of total seston and particulate organic matter was mainly due to variations in particulate proteins, lipids and carbohydrates; pigment concentrations were of secondary importance. Expressed as the ratio chlorophyll a: particulate protein-nitrogen, phytoplankton constituted an important fraction of the microplankton biomass only during spring blooms, when it averaged 75% of the particulate protein-nitrogen. In constrast, <30% of protein-nitrogen was related to chlorophyll a during the summer. These results suggest that an oligotrophic structure based on the microbial-loop prevails in microplankton assemblages of surface waters for most of the year.     

Observations on the diel migration of marine dinoflagellates off the Baja California coast

The vertical distribution of Gonyaulax polyedra, Ceratium furca, Gymnodinium sp., Ceratium dens, Gonyaulax digitale, Prorocentrum micans, Polykrikos kofoidii, and Peridinium depressum was followed during two 36 h time-series stations, off the Baja California coast. Water samples were taken at 6 h intervals, at 7 light penetration depths, for phytoplankton identification and for chemical and biological analysis. The results indicate that the dinoflagellates were able to perform vertical migration against the physical water movements, such as upwelling internal waves and wind mixing, that existed in the area during this study. Attempts to correlate the vertical dstribution of the dinoflagellates with the seawater density and nitrate distribution lead to the conclusion that even a weak density gradient acted as a barrier to the downward migration, and that nitrate assimilation in darkness was not required for their growth. Differences in the vertical distribution at 12.00 hrs of each species supports the hypothesis that migration is of a phototactic nature. A comparison of this study with previous ones on vertical migration in other areas suggests that a species should not be defined in general as a positive or negative phototactic organism, but that the direction of the migration is related to the light intensity at the sea surface.Contribution No. 77-004 of the Bigelow Laboratory for Ocean Sciences, West Boothbay Harbor, Matne 04575, USA.     

Photosynthetic characteristics of marine Synechococcus spp. with special reference to light environments near the bottom of the euphotic zone of the open ocean

The present study aimed to resolve the question why marine Synechococcus spp. abundantly occur even at the bottom of the euphotic zone in the Kuroshio are. Photosynthesis under such conditions was examined using simulated blue-green model light (BGL). Results indicated that photosynthesis of marine Synechococcus spp. under BGL is as active enough to support growth of these organisms. Examination of light-harvesting under BGL indicated that active photosynthesis is permitted by an unusually high abundance of phycoerythrin (PE), which is the main light-harvesting pigment for photosystem II (PSII), due to a phycobilisome (PBS) structure which is different from ordinary hemidiscoidals. Although the absorption maximum of PE is located at longer wavelengths than the energy maximum of BGL, PE was found to absorb BGL significantly. Thus, BGL cannot be a typical photosystem I (PSI) light. PSII is also significantly excited by BGL. Carotenoids, which largely absorb BGL, were found to be effective in light-harvesting for PSI. Based on the results obtained, possible reasons why marine Synechococcus spp. commonly occur in warm waters were discussed. Two strains of Synechococcus spp. isolated from the Gulf Stream in 1981 and from Kuroshio, Japan in 1983 were used in the present study.     

Conservation and management-oriented ecological research in the coastal zone of Baja California, Mexico

This paper presents the main results of two studies of contrasting natural and man-induced conditions along the Pacific coast of Baja California (Mexico), based on the assumption that ecological insight can be obtained from man-induced modifications insofar as relevant activities are explicitly addressed as parts of the systems under study. The study is concerned with a fragmented coastal succulent-sage scrub and showed that several patches of different size and age may harbor as many species as non-fragmented areas, and that 83% of the original species assemblage persisted in the fragments, in which the invasion by opportunistic exotic species may not progress beyond certain limits. The study of dune-backed and urban-backed beaches showed a significantly greater abundance of the Snowy Plover (Charadrius alexandrinus) at the dune-backed beach, where a much more active back-shore feeding of adult birds and the only evidences of breeding occurred. Both studies refer to landscape features of regional concern and its results may be used in conservation management. The results on fragmented coastal succulentsage scrub may encourage alternative urban designs that keep patches of the original landscape, thus meeting existing requirements of low density urban development for most of the coastal succulent-sage scrub area. The evidence presented on the negative effects of dune destruction on the abundance, feeding and reproductive performance of a threatened bird species has a bearing on the topic of biodiversity management. It may also contribute to the conservation of the coastal dunes system.     

Floristic and physiological differences between the shallow and the deep nanophytoplankton community in the euphotic zone of the open tropical Atlantic revealed by HPLC analysis of pigments

Suspended matter sampled in 1982 in the North Equatorial Current, in the open Atlantic to the west of West Africa, was analyzed by high performance liquid chromatography. The pigment fingerprint of samples taken in the surface mixed layer was dominated by zeaxanthin and chlorophyll a, in agreement with observed dominance of coccoid cyanobacteria. Near the bottom of the euphotic zone the fingerprint was more complicated, with a sharp transition at the depth of the deep chlorophyll maximum layer to dominance of chlorophyll b, 19-hexanoyloxyfucoxanthin and an unknown fucoxanthin derivative in the lower part of this layer; this fingerprint suggests dominance of eukaryotes (green algae, Prymnesiophyceae and Chrysophyceae) at depth. Up to 90% of the chl a was contained in particles smaller than 8 m, and in the surface mixed layer even more than 50% in particles smaller than 1 m. The high concentration of zeaxanthin relative to chl a near the surface suggests adaptation of the cyanobacteria to exposure to high irradiance. Evidence of this adaptation was the very high specific phytoplankton growth rate between sunrise and sunset (=0.16 h^-1), measured by recording ¹⁴C incorporation into organic carbon and into chl a carbon after isolation of the latter by HPLC. The high concentration of chl b relative to chl a at depth was possibly caused by shade-adapted green algae containing more chl b than chl a. The specific growth rate of the deep shade community was low (<0.04 h^-1), yet net primary production, calculated on the basis of chl a increase during incubation, was greatest at depth.     

The undulating oceanographic recorder — A new instrument system for sampling plankton and recording physical variables in the euphotic zone from a ship underway

The prototype Undulating Oceanographic Recorder Mark I is an instrumented towed vehicle which can be programmed to undulate between a minimum depth of 8 m and a selected maximum depth between 15 and 70 m, with an undulation length between 3 and 30 km, at any speed between 7 and 15 knots (3.6 to 7.7 m/sec). It takes a continuous series of plankton samples, each integrated over the undulation depth range, and records data on magnetic tape, at a preselected rate between 30/min and 30/h, from which salinity, temperature and depth can be derived with accuracies of ±0.1, ±0.1C° and ±0.5 m, respectively. The instrument is automatic, self-contained and generates its own power supplies. It is towed on about 200 m of unfaired steel cable and can be handled with the winching and lifting equipment found on most research vessels. Examples of processed data from three cruises are presented: temperature and salinity contours on a vertical section through a complex stratified area in the N.W. Irish Sea; results from a survey of the waters over the continental shelf and slope to the north of Scotland, showing the distribution of water masses and associated plankton; and hydrographic data taken in Loch Etive on the west coast of Scotland.     

A comparison of zooplankton patterns in the California Current and North Pacific Central Gyre

This study compares the small-scale (100's to 1000's of meters) horizontal spatial distribution of zooplankton from stations in the California Current (29°N; 118°W) and in the North Pacific Central Gyre (31°N; 155°W). Patterns were inferred from the spacing of vertical tows and the variability in abundance of 15 taxonomic categories caught in these tows. In the California Current, 4 tows were taken at random positions within 2000 m of a drogue or fixed geographic position and 4 replicate tows were taken at the drougue or the fixed point. Four series of these 8 tows were taken around noon and midnight during two days. At the Central Gyre station, two similar series (one day and one night) were taken while following a drogue. Estimates of the scale of structure based on comparisons of replicate and random tow variability suggest that aggregated patterns in the Central Gyre are larger than a few hundred meters, while the California Current structures may be less than 100 m in size. Day-night changes in patch size were not apparent in either area. Evidence for multispecies patches was found only for the Central Gyre station. Replication samples in both areas gave more consistent measures of community structure (relative proportions of species) than the random tows. Similarity of community structure decreased with increasing distance between tows being compared. The decrease was much greater for the California Current station, a further indication of smaller patch size in this area. Diurnal changes in community structure occurred in both areas.Based in part on a dissertation submitted in partial fulfillment of the requirements of the Ph.D. degree at the University of California, San Diego. Portions of this research were supported by the Marine Life Research Program of the Scripps Institution of Oceanography.     

Localization of bioluminescence in the siphonophore Nanomia cara

The base of the tentacle of the developing physonect larva (Nanomia cara) has a bioluminescent region. The ability to produce light in the larva is transitory; this ability first appears at about two days of development and is disappearing by eight days, as the larva begins to feed. Subsequently paired bilaterally symmetrical bioluminescent organs are found on the nectophores and the bracts of the adult colony. In both the larva and the adult, bioluminescence is mediated by a calcium specific photoprotein. In all cases the photocytes lack a green fluorescent protein.     

Where the wild things are: predicting hotspots of seabird aggregations in the California Current System

Marine protected areas (MPAs) provide an important tool for conservation of marine ecosystems. To be most effective, these areas should be strategically located in a manner that supports ecosystem function. To inform marine spatial planning and support strategic establishment of MPAs within the California Current System, we identified areas predicted to support multispecies aggregations of seabirds ("hotspots"). We developed habitat-association models for 16 species using information from at-sea observations collected over an 11-year period (1997-2008), bathymetric data, and remotely sensed oceanographic data for an area from north of Vancouver Island, Canada, to the USA/Mexico border and seaward 600 km from the coast. This approach enabled us to predict distribution and abundance of seabirds even in areas of few or no surveys. We developed single-species predictive models using a machine-learning algorithm: bagged decision trees. Single-species predictions were then combined to identify potential hotspots of seabird aggregation, using three criteria: (1) overall abundance among species, (2) importance of specific areas ("core areas") to individual species, and (3) predicted persistence of hotspots across years. Model predictions were applied to the entire California Current for four seasons (represented by February, May, July, and October) in each of 11 years. Overall, bathymetric variables were often important predictive variables, whereas oceanographic variables derived from remotely sensed data were generally less important. Predicted hotspots often aligned with currently protected areas (e.g., National Marine Sanctuaries), but we also identified potential hotspots in Northern California/Southern Oregon (from Cape Mendocino to Heceta Bank), Southern California (adjacent to the Channel Islands), and adjacent to Vancouver Island, British Columbia, that are not currently included in protected areas. Prioritization and identification of multispecies hotspots will depend on which group of species is of highest management priority. Modeling hotspots at a broad spatial scale can contribute to MPA site selection, particularly if complemented by fine-scale information for focal areas.