Physiology and chemical composition of nitrogen-fixing phytoplankton in the central North Pacific Ocean

The magnitude and physiological characteristics of biological nitrogen fixation have been studied in the oligotrophic waters of the North pacific gyre. The filamentous blue-green algae Trichodesmium spp. and Richelia intracellularis were the important nitrogen-fixing phytoplankton. Most of the nitrogen fixation occurs in the upper 40 m of the water column, with detectable fixation as deep as 90 m, which corresponds to about the 1 % light depth. There was no evidence of photoinhibition of nitrogen fixation, although CO₂ reduction was depressed slightly at the highest light levels. The rate of nitrogen fixation in the water column varied throughout the day, being highest in mid-morning and in late afternoon. Relatively high fixation rates were also found during periods of darkness. Elevated oxygen concentrations had a marked inhibitory effect on rates of nitrogen fixation, a pO₂ of 0.4 atm causing a 75% inhibition. Data from studies of nitrogen fixation and assimilation rates of ¹⁵N-labelled nitrate, ammonium, and urea indicate that nitrogen fixation furnished about 3% of the total daily fixed nitrogen requirement for phytoplankton growth. Studies with isolated colonies of Trichodesmium spp. indicated that 100% of their nitrogen requirement was met by nitrogen fixation. Chemical composition of the Trichodesmium colonies showed that the C:N ratio was 4.1 and that their phosphorus content relative to carbon or nitrogen was much lower than that of the total particulate material in the water column. Elevated ratios of carbon: adenosine triphosphate (ATP) also suggest that phosphorus deficiency may be limiting the growth of Trichodesmium. The magnitude of nitrogen fixation in the gyre is seasonally dependent, with high rates in late summer and autumn. At these times the water column is stratified, with phosphate and nitrate barely detectable in the upper 100 m. Our data suggest that during these months of stratification, biological fixation of nitrogen amounts to about 33 g-at N/m²/day.     

Nitrogen fixation in the North Pacific Ocean

Nitrogen fixation in the euphotic zone of the ocean was measured by C₂H₂ reduction and ¹⁵N₂ incorporation associated with Trichodesmium sp. and also with Richelia intracellularis occurring within the cells of Rhizosolenia styliformis var. longispina, and R. cylindrus. The vertical distribution of N₂ fixation activity, N₂-fixing species, particulate matter and dissolved nutrients was measured. The effects of light intensity, sample concentration, length of incubation, and nutrient enrichment on the rates of C₂H₂ reduction were determined. Estimates of the importance of N₂ fixation in adding previously uncycled nitrogen to the euphotic zone are given.     

Spring phytoplankton in the Subarctic North Pacific Ocean

The report includes quantitative and qualitative data on the phytoplankton from the First Canadian Transpacific Oceanographic Cruise from March to May 1969. Nanoplanktonic species (<20 m) were numerically dominant and, together with nertic diatoms, made up the larger portion of the biomass at all stations outside the Western Subarctic and Alaskan Gyres, where Denticulopsis seminae predominated. The nanoplankton consisted mainly of haptophycean, dinophycean and cryptophycean flagellates. Tropical oceanic dinoflagellates and diatoms were abundant in the net samples from Station 17 in the warm Kuroshio current but absent from Stations 24–27 in the Western Subarctic Gyre, Stations 2, 37–39 in the Alaskan Gyre, in which cold water oceanic species predominated, and Stations 31 and 33, south of the Aleutians. Subsurface maxima were common on the outward bound (westerly) leg in March.     

Larval fishes in the zooplankton community of the North Pacific Central Gyre

Patterns of species structure are described for the larval fish assemblage in the North Pacific Central Gyre. About 30,000 larvae, primarily of mesopelagic fish species, were identified for 7 cruises. Samples were collected using Isaacs-Kidd plankton trawls on 6 cruises over a 4 1/2 year period; stratified samples were taken with an opening/closing bongo net on one late-summer cruise. Data on absolute and relative abundances, size ranges and summertime depth distributions of over 150 species of larvae are presented. The depth distribution and abundance of the ichthyoplankton are compared to those of the total zooplankton community. The larval assemblage is also examined with respect to the known adult mesopelagic fish assemblage. Seasonal changes in ichthyoplankton species structure occur but, within seasons, species structure is remarkably constant from year to year. Despite seasonal changes in species abundance and rank order of abundance, a constant cumulative frequency structure was found to exist in the ichthyoplankton, both between seasons and between years. The implications of this result are discussed in light of similar findings in a terrestrial community and with respect to possible mechanisms of regulation.     

Hyperiid amphipods from the zooplankton community of the North Pacific central gyre

The hyperiid amphipods have been examined from an extensive series of depthstratified bongo net tows taken at 28°N; 155°W in the North Pacific central gyre, an open-ocean ecosystem. Seventy-nine samples from 6 depth ranges produced 14, 581 individuals belonging to 13 families, 42 genera, and 83 species. Data on depth distributions, rank orders of abundance, sex ratios, and vertical migration are presented. Most of these species are shown to be reproductively active in the gyre, and there appears to be no seasonality to their reproductive behavior. No species endemic to the gyre are yet known.     

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.     

Deep distributions of oceanic cirripede larvae in the Sargasso Sea and surrounding North Atlantic Ocean

From deep ( 1000 m), multi-depth zooplankton samples taken in the Azores frontal region from November 1980 to June 1981, high numbers of the two larval forms ofLepas pectinata, the commonest lepadid cirripede in this area, were sampled. The nauplii and cyprid larvae are large and long-lived. Nauplii were sampled where they feed, in the upper 150 m, closely associated with the fluorescence maximum. The cyprids were mainly sampled between 300 and 400 m, with a sharp cut-off in numbers just below the distribution peak. This deep distribution is intriguing, as the cyprids, which were not migrating diurnally, are non-feeding. To complete their life cycle they must settle on floating debris at the surface, where very few were sampled. They are negatively buoyant and there is no apparent physical reason for this deep distribution. There was no obvious pattern of changes in individual length or length/dry weight distribution with depth, the population appeared homogeneous. There was also no accumulation of other zooplankton at the same depth, or evidence from gut contents of large carnivores that the distribution pattern was caused by selective predation. A close association exists betweenL. pectinata andSargassum spp. weed, the cyprids preferentially settling on small fragments of weathered weed, the supply of which may be enhanced after winter. During winter, the harsh environment at the surface of the northern Sargasso Sea may discourage cyprid settlement. The deep distribution may be an ontogenetic migration conferring some survival advantage and saving energy due to the lower temperature and oxygen levels at depth. If it is a behavioural mechanism, duration of the cyprid stage will be extended until a more clement time of year, allowing them to settle over a longer period. The cyprids of other species were also sampled at depth, so the phenomenon appears to be widespread. If subsequent research confirms these observations as an ontogenetic migration, our current understanding of the fouling behaviour of oceanic lepadid cirripedes would be considerably modified.     

Symbiosis between bacteria and oceanic clonal sea star larvae in the western North Atlantic Ocean

Bacteria were present in the majority of clonal sea star bipinnariae (Luidia sp.) collected between June 1987 and August 1990 from oceanic populations in the subtropical western North Atlantic Ocean. Light and electron microscopy revealed dense aggregations of bacteria within lobes of the epidermal cuticle in the gastric region of larvae. Gram-negative rods and less-common elongate spiral bacteria were observed, always in close association with branching epidermal microvilli. Intact and partially digested bacteria occurred in vesicles and phagosomes within epidermal cells, a probable indication of phagocytotic activity by the host. The association of larvae of Luidia sp. and bacteria ranged widely on both geographical and temporal scales. During July and August 1988, larvae were abundant (1.6 to 11.1 m³) in the mixed surface layer (<10 to 100 m) of stations in the Gulf Stream, the Sargasso Sea and the North Equatorial Drift, and 96% (mean of 5 stations, n=8 to 10 larvae in each station) of the individuals surveyed harbored bacteria beneath their cuticle. In a single station on the Florida Current sampled during winter, spring, and summer months between 1989 and 1991, 79 to 90% of the larvae harbored bacteria. The incidence of symbiosis was higher in actively cloning larvae than in non-cloning larvae. This is the first documentation of a symbiosis in field populations of echinoderm larvae.     

A report of green algal pigments in the Central North Pacific Ocean

Thin-layer chromatography showed that chlorophyll b was present in acetone extracts of 6 samples of suspended matter collected in February 1973 from the Central North Pacific Ocean. Pigment patterns showed the presence of green algae and diatoms through most of the euphotic zone down to 200 m. Ratios of chlorophyll b:a ranged from 0.2 to 0.5 at 5 and 100 m depths, but were less than 0.05 at 200 m.     

Changes in size and trends of North American sea duck populations associated with North Pacific oceanic regime shifts

Broad-scale multi-species declines in populations of North American sea ducks for unknown reasons is cause for management concern. Oceanic regime shifts have been associated with rapid changes in ecosystem structure of the North Pacific and Bering Sea. However, relatively little is known about potential effects of these changes in oceanic conditions on marine bird populations at broad scales. I examined changes in North American breeding populations of sea ducks from 1957 to 2011 in relation to potential oceanic regime shifts in the North Pacific in 1977, 1989, and 1998. There was strong support for population-level effects of regime shifts in 1977 and 1989, but little support for an effect of the 1998 shift. The continental-level effects of these regime shifts differed across species groups and time. Based on patterns of sea duck population dynamics associated with regime shifts, it is unclear if the mechanism of change relates to survival or reproduction. Results of this analysis support the hypothesis that population size and trends of North American sea ducks are strongly influenced by oceanic conditions. The perceived population declines appear to have halted >20 years ago, and populations have been relatively stable or increasing since that time. Given these results, we should reasonably expect dramatic changes in sea duck population status and trends with future oceanic regime shifts.     

Fecundity of a euphasiid crustacean,Nematoscelis difficilis,in the North Pacific Ocean

The number, size, and carbon and nitrogen contents of eggs in the pouches of a euphausiid crustacean, Nematoscelis difficilis Hansen, were examined. A clear linear relationship exists between the number of eggs in the pouch and the body weight of the maternal euphausiid. The eggs are not spherical in shape, and the size of eggs is greater in larger egg masses. The carbon content of the egg masses (50.2%) is higher than that of the bodies (40.7%). The carbon: nitrogen ratio (C/N) is also higher in the egg masses than in the bodies of euphausiids. The carbon content of the eggs is equivalent to 28.4% of the body carbon, the nitrogen content to 19.2% of the body nitrogen.     

Population structure of the planktonic copepod Calanus pacificus in the North Pacific Ocean

The pelagic copepod Calanus pacificus ranges nearly continuously across temperate-boreal regions of the North Pacific Ocean and is currently divided into three subspecies—C. pacificus oceanicus, C. pacificus californicus, C. pacificus pacificus—based on subtle morphological differences and geographic location. The relation between geography and genetic differentiation was examined for 398 C. pacificus individuals sampled from six widely distributed locations across the North Pacific, including an open ocean site and coastal sites on both sides of the North Pacific basin. For each individual copepod, the DNA sequence was determined for a 421-bp region of the mitochondrial coxI gene (mtCOI). A total of sixty-three different mtCOI sequences, or haplotypes, were detected, with a sequence divergence between haplotypes of 0.2–3.1%. The number and distribution of haplotypes varied with sampling location; 12 haplotypes were distributed across multiple sampling locations, and 51 occurred at only one location. Five genetically distinct populations were detected based on F _ST values. Haplotype minimum spanning networks, nucleotide divergence and F _ST values indicated that individuals from coastal sites in the North Pacific Ocean were more closely related to each other than to individuals from the open ocean site at Station P. These results provide genetic support for the designation of two subspecies—a coastal subspecies that consists of what is currently referred to as C. p. pacificus and C. p. californicus and an open ocean subspecies C. p. oceanicus. This work also indicates that planktonic copepods with potentially high dispersal capacity can develop genetically structured populations in the absence of obvious geographic barriers between proximate locales within an ocean basin.     

Nitrogen assimilation by phytoplankton and other microorganisms in the surface waters of the central North Pacific Ocean

Assimilation rates of ¹⁵N-labelled ammonium, urea, and nitrate by plankton in the upper euphotic zone were measured in 2 summer, 2 winter, and 1 spring cruise in the central North Pacific Ocean. Average rates of ammonium plus urea assimilation could not be determined precisely, but were estimated to be 7 to 25 g-at. N m^-3 day^-1. Indirect evidence suggested that non-photosynthetic microorganisms contributed to these rates. Nitrate assimilation was negligible in the upper waters considered in this report (above the chlorophyll maximum and the nutricline). Potential, nitrate-saturated rates were in the range 1 to 8 g-at. N m^-3 day^-1. Seasonal comparison showed lowest rates of both carbon and nitrogen assimilation rates per chlorophyll a in winter.     

Nematode community composition in hydrothermal vent and adjacent non-vent fields around Myojin Knoll,a seamount on the Izu-Ogasawara Arc in the western North Pacific Ocean

In contrast to specific large benthic invertebrates in chemosynthetic ecosystems such as hydrothermal vents, meiofaunal communities in such habitats have been reported to have strong taxonomic overlap with meiofauna in the adjacent “normal” environments. However, meiofauna have only recently been included in studies of those environments and detailed information on these communities is still rare. This is especially true in the Northwest Pacific Ocean, even though there are many seamounts with active vents in the calderas of the region. Nematode community composition at the genus level in sediments from a hydrothermal vent field in the caldera of Myojin Knoll (32°06′N, 139°52′E, depth 1,300 m), a seamount on the Izu-Ogasawara Arc, Japan, was investigated for the first time and was compared with adjacent non-vent areas inside and outside the caldera. Multivariate analyses showed that the composition of nematodes in the hydrothermal field was significantly different from that in the non-hydrothermal fields around the caldera. However, the common genera, such as Oxystomina, Pareudesmoscolex, Desmoscolex, and Microlaimus were found in two, or all three vent fields while their rank contributions differed among the three fields. When the data from Myojin Knoll were compared with those from other deep-sea vent environments in different regions (e.g., North Fiji Basin, East Pacific Rise, Mid-Atlantic Ridge), the nematode composition in the vent field of the Myojin caldera was more similar to that of the non-vent fields around the caldera than the composition in vent fields of other regions. These data from the Northwest Pacific Ocean also suggest the absence of long-range transport systems and local adaptations for meiofauna in hydrothermal vent fields.     

Temperature and metabolic rate in sedentary fish from the Antarctic,North Sea and Indo-West Pacific Ocean

Resting metabolic rate was measured in demersal stages of the teleostNotothenia neglecta Nybelin from the South Orkney Islands, Antarctica, from 1985 to 1987. The relationship between and body mass (Mb) conformed to the general relationship , wherea is a proportionality constant andb is the scaling exponent. (mg O₂ h^–1) was found to scale toMb ^{(0.82±0.011)} in the summer (November to April, 1.6 to 1 850 g,n=56) and toMb ^{(0.76±0.013)} in the winter (May to October, 0.9 to 1 850 g,n=57) (values ofb are means ± SD). Although the scaling exponents were significantly different (P<0.01), was similar in the juvenile stages of summer- and winter-caught fish matched for body mass. The effects of activity on oxygen consumption was studied using a Brett respirometer. Adult stages had a factorial aerobic scope for activity of 5.7, which is similar to that reported for demersal fish from temperate latitudes. The effects of temperature on resting metabolism was investigated in fish with similar sedentary lifestyles from the North Sea (Agonus cataphractus andMyoxocephalus scorpius) and the Indo-West Pacific (Paracirrhites forsteri, P. arcatus, Neocirrhites armatus andExallias brevis). Extrapolated values of for the tropical species approached zero at 5 to 10°C. For a standard 50 g fish, for the tropical species at 25°C was in the range 3.4 to 4.4 mg O₂ h^–1, compared with 1.3 mg O₂ h^–1 forNotothenia neglecta at its acclimation temperature. Thus, the maximum metabolic rate of sedentary tropical species at 24°C is likely to be 2 to 4 times higher than inN. neglecta at 0°C. This suggests that the energy available for sustained activity is significantly lower in cold- than in warm-water fish.     

Prediction of response of zooplankton biomass to climatic and oceanic changes

This paper examines the long-term variation in zooplankton biomass in response to climatic and oceanic changes, using a neural network as a nonlinear multivariate analysis method. Zooplankton data collected from 1951 to 1990 off the shore of northeastern Japan were analyzed. We considered patterns of the Kuroshio and the Oyashio, sea surface temperature, and meteorological parameters as environmental factors that affect zooplankton biomass. Back propagation neural networks were trained to generate mapping functions between environmental variables and zooplankton biomass. The performance of the network models was tested by varying the numbers of input and hidden units. Changes in zooplankton biomass could be predicted from environmental conditions. The neural network yielded predictions with smaller errors than those of predictions determined by linear multiple regression. The sensitivity analysis of networks was used to extract predictive knowledge. The air pressure, sea surface temperature, and some indices of atmospheric circulation were the primary factors for predictions. The patterns of the Kuroshio and the Oyashio demonstrated different effects among sea areas.     

Vertical distribution and diurnal migration of some Cyclopoida (Copepoda) in the tropical region of the Pacific Ocean

The material was collected in the western equatorial part of the Pacific Ocean at a 24 h station on December 21 to 23, 1968. Collections were made by means of 2 Juday nets from 2 winches simultaneously. A series of hauls was made at 2 h intervals. The vertical distribution of 17 Cyclopoida species, belonging to 3 families (Oithonidae, Oncaeidae and Corycaeidae) was studied. From 0 to 300 m, the absolute number of Cyclopoida individuals was practically constant throughout the 24 h period. The bulk of the species performed no diurnal vertical migration, or migrated with low intensity in the usual way (i.e. moved upward at night and downward during the day). Only 2 Oncaeidae species performed significant diuranl vertical migration. No reversed migration in Cyclopoida was discovered. To characterize the vertical distribution of Cyclopoida, the distribution of the cores of populations (abundancies between 25 and 75%) was examined. During the 24 h period, the cores of populations of various Cyclopoida species were found at different depths; this steplike distribution was not disturbed in migrating species. The cores of populations differed both with habitat depth and time of ascent of the migrating species to the surface layer. This peculiarity, probably, tends to lessen the intensity of food competition between Cyclopoida species with similar nutritional habits.     

Phosphorus metabolism of oceanic dinoflagellates: phosphate uptake,chemical composition and growth of Pyrocystis noctiluca

The phosphorus metabolism of Pyrocystis noctiluca Murray (Schuett) 1886 has characteristics which may enhance its potential for success in orthophosphate impoverished waters. The steady-state phosphate uptake rates were equal in the light and dark, and were directly proportional to both the phosphorus cell quota and the cell division rate. In contrast, nutrient-saturated uptake rates were multiphasic, faster in the light than the dark, 2 to 4 orders of magnitude greater than steady-state rates, and were inversely proportional to both the phosphorus cell quota and the cell division rate. These uptake characteristics suggest that P. noctiluca may take up phosphate coincidently at their typically low ambient concentrations as well as to exploit episodic nutrient events in nature. Cell division rates were a hyperbolic function of the ambient orthophosphate concentration. The shortest doubling time was 8.7 d, the phosphate concentration at half the maximum division rate was 0.15 M and the threshold, concentration for cell division was ca 0.05 M PO ₄ ^3- . Division rates of P. noctiluca in the ocean are much faster than predicted from the measured ambient orthophosphate concentrations. Since this dinoflagellate has high naturally occurring alkaline phosphatase activities, and can utilize organic-P compounds, we suggest that organic-P can be as important as orthophosphate in supporting the observed division rates of P. noctiluca in the sea.