Erica J. H. Head

Vertical flux of respiratory carbon by oceanic diel migrant biota

Interzonal diel migrant plankton and nekton obtain organic carbon by feeding at night above the main pycnoline of subtropical and tropical oceans, and respire part of it by day in the interior of the ocean below the pycnocline. Using data from seven oceanic stations, and conservative models to compute respiration at depth, we show that this flux of respiratory carbon ranged from 20 to 430 mg C m−2 d−1 or 13–58% of computed particulate sinking flux across the pycnocline. If this flux occurs consistently between 50°N and 50°S, it will add about 5–20% (depending on method of calculation) to current estimates of global sinking flux of organic carbon across the pycnocline.

The distribution and metabolism of urea in the eastern Canadian Arctic

Urea concentrations, uptake, and excretion were measured at various locations in northern Baffin Bay and surrounding waters during the summer of 1980. Concentrations were variable ( 2.00 mg-at. N m−3) but followed patterns of decreasing concentration with depth in the euphotic zone and with distance from land. Urea accounted for > 50% of the total dissolved nitrogen in the upper mixed layer at most stations. Urea uptake rates showed generally the same distributional patterns as did concentrations and on the average accounted for 32% of the total nitrogen (NO3− + NH4+ + urea) productivity in the eupholic zone. Ammonium, and frequently NO3−, were utilized in preference to urea. Dual isotope (14C and 15N-urea) labelling experiments suggested that most urea-C was respired as CO2 while 50 to 80% of the urea-N was incorporated by the phytoplankton. Excretion measurements suggested that the four dominant macrozooplankton species (Calanus hyperboreus, C. finmarchicus, C. glacialis, and Metridia sp.) supplied only −3% of the urea-N but –40% of the NH4+-N requirements of the primary producers.

Pigment transformation and vertical flux in an area of convergence in the North Atlantic

Abstract Particulate samples from the water column, to depths of 80–120 m, were collected over a 10-day period during a bloom in the North Atlantic, near 37°N, 40°W, between 12 and 22 April 1990. These samples were analysed by HPLC for their chlorin and carotenoid content. The same analysis was carried out for samples of faecal pellets collected from copepods from depths of 0–100 m, which had fed in situ, and for material from sediment traps in the 100–500 m depth range. Water column pigment compositions were consistent with observations that diatoms and Phaeocystis were major components of the bloom. They also sometimes showed high concentrations of chlorophyllide a and a phaephorbide a-like pigment, which may be due to algal die-off and cell autolysis. Faecal pellets contained high concentrations of pyrophaeophorbide a lower amounts of unidentified phaeophorbides and some chlorophyll a, but none of the phaeophorbide a-like pigment found in the water column. Sediment traps contained high concentrations of the phaeophorbides characteristics of copepod faecal pellets and lower, approximately equal amounts of the water column phaeophorbide a-like pigment and chlorophyll a. The sampling area appeared to be in a jet of current, which was moving southwesterly along a front between Eastern Basin Water (to the north) and North Atlantic Central Water (to the south) and which seemed to be drawing in water from both these water masses. Areas of downwelling, thought to be associated with these convergent currents, may have accelerated the sedimentation of dead and dying phytoplankton cells.

Distributions of Calanus spp. and other mesozooplankton in the Labrador Sea in relation to hydrography in spring and summer (1995–2000)

Abstract We collected mesozooplankton samples in the upper 100 m in spring or early summer each year between 1995 and 2000 along a section from Hamilton Bank (Labrador) to Cape Desolation (Greenland), and along additional sections in spring 1997 and early summer 1995. The North Atlantic waters of the central basin were characterised by the presence of the copepods Calanus finmarchicus , Euchaeta norvegica and Scolecithrocella minor and euphausiids. Calanus glacialis , Calanus hyperboreus and Pseudocalanus spp. were associated with the Arctic waters over the shelves. Amongst the other enumerated groups larvaceans were concentrated over the shelves and around the margins. Amphipods, pteropods and the copepods Oithona spp. and Oncaea spp. showed no definable relationships with water masses or bathymetry, while the diel migrant ostracods and chaetognaths were confined to deep water. Metrida longa , also a strong diel migrant, and Microcalanus spp., a mainly deep water species and possible diel migrant, were both sometimes quite abundant on the shelves as well as in the central basin, consistent with their likely Arctic origins. Analysis of community structure along the section across the Labrador Sea indicated that stations could be grouped into five different zones corresponding to: the Labrador Shelf; the Labrador Slope; the western and central Labrador Sea; the eastern Labrador Sea and Greenland Slope; and, the Greenland Shelf. The boundaries between zones varied spatially between years, but community composition was relatively consistent within a given zone and a given season (spring versus early summer). The relationship between community composition and water masses was not entirely straightforward. For example, Labrador Shelf water was generally confined to the shelf, but in spring 2000 when it also dominated the adjacent slope zone, the community in the Labrador Slope zone was similar to those found in other years. Conversely, in spring 1997, when Arctic organisms were unusually abundant in the Labrador Slope zone, there was no increased contribution of shelf water. In addition, North Atlantic organisms were often found on the shelves when no slope or central basin water was present. Although other organisms were sometimes very abundant, the mesozooplankton preserved dry weight biomass was dominated everywhere by the three species of Calanus , which together always accounted for ≥70%. One species, C. finmarchicus , comprised >60% of the total mesozooplankton biomass and >80% of the abundance of large copepods in spring and summer throughout the central Labrador Sea. In western and central regions of the central basin average C. finmarchicus biomass was ca 4 g dry weight m −2 and average abundance, ca 17 000 m −2 over both seasons. Highest levels ( ca 7 g dry weight m −2 , >100 000 m −2 ) occurred in the northern Labrador Sea in spring and in eastern and southwest regions in early summer. C. hyperboreus contributed ca 20% of the total mesozooplankton biomass in the central basin in spring and C. glacialis accounted for C. hyperboreus contributed a maximum of 54% and 3.6 g dry weight m −2 , and C. glacialis , a maximum of 29% and 1 g dry weight m −2 , to the total mesozooplankton biomass.

The western North Atlantic bloom experiment

An investigation of the spring bloom was carried out in the western North Atlantic (40–50°W) as one component of the multi-nation Joint Global Ocean Flux Study (JGOFS) North Atlantic Bloom Experiment (NABE). The cruise track included an extended hydrographic section from 32 to 47°N and process studies at two week-long time-series stations at 40 and 45°N. Biological and chemical data collected along the transect indicated that the time-series stations were located in regions where the spring bloom was well developed; algal biomass was high and surface nutrient concentrations were reduced from maximum wintertime levels. Despite similarities in the vertical structure and magnitude of phytoplankton biomass and productivity, the two stations clearly differed in physical, chemical and other biological characteristics. Detailed depth profiles of the major autotrophic and heterotrophic microplankton groups (bacteria, phytoplankton, microzooplankton) revealed a strong vertical coherence in distribution at both sites, with maximum concentrations in the upper 50 m being typical of the spring bloom. Ultraplankton (< 10 μm) were an important component of the primary producers at 40°N, whereas larger netplankton (diatoms, dinoflagellates) were more important at 45°N. Silicate depletion was clearly evident in surface waters at 45°N, where diatoms were most abundant. Despite the relative importance of diatoms at 45°N, dinoflagellates dominated the biomass of the netplankton at both sites; however, much of this community may have been heterotrophic. Bacterial biomass and production were high at both stations relative to phytoplankton levels, particularly at 45°N, and may have contributed to the unexpectedly high residual ammonium concentrations observed below the chlorophyll maximum layer at both stations. Microzooplankton grazing dominated phytoplankton losses at both stations, with consumption as high as 88% of the daily primary production. Grazing losses to the mesozooplankton, on the other hand, were small (<10%), but mesozooplankton contribution to the vertical flux of organic matter (fecal pellets) was important at 45°N. F-ratios estimated by 15N tracer methods and sediment trap fluxes were similar and suggeste that ∼30% of the daily primary production was lost by direct sedimentation during the observation period. Numerous similarities in bloom characteristics were noted between the western and eastern Atlantic study sites.

Physiological and biochemical changes in Calanus hyperboreus from Jones Sound NWT during the transition from summer feeding to overwintering condition

SummarySeveral biochemical and physiological characteristics of stage V and female adult Calanus hyperboreus from two different depth ranges (0–50 m and 200–500 m) were compared at a time near the peak of the summer pelagic algal bloom in Jones Sound, and again one month later, when the near surface chlorophyll levels were low and most copepods had migrated to their overwintering dephts. For a given stage deep water animals were larger and had higher total lipid levels than did surface animals. Feeding activities, as evidenced by gut pigment contents, may be lower in animals at depth, although potential digestive activities, as expressed in the levels of three digestive enzymes, were not very different either in different stages or at different depths. Respiration rates in animals that had migrated down for the winter were not much lower than in those at the surface although energetic considerations suggest that they may decrease later. Ammonia excretion rates however, changed dramatically. Surface, feeding animals had the highest rates and deep water animals in September had undetectably (<1 ng atom ammonia nitrogen animal-1 · day-1) low rates.

Coherent assembly of phytoplankton communities in diverse temperate ocean ecosystems

The annual cycle of phytoplankton cell abundance is coherent across diverse ecosystems in the temperate North Atlantic Ocean. In Bedford Basin, on the Scotian Shelf and in the Labrador Sea, the numerical abundance of phytoplankton is low in spring and high in autumn, thus in phase with the temperature cycle. Temperature aligns abundance on a common basis, effectively adjusting apparent cell discrepancies in waters that are colder or warmer than the regional norm. As an example of holistic simplicity arising from underlying complexity, the variance in a community variable (total abundance) is explained by a single predictor (temperature) to the extent of 75% in the marginal seas. In the estuarine basin, weekly averages of phytoplankton and temperature computed from a 13 year time-series yield a predictive relationship with 91% explained variance. Temperature-directed assembly of individual phytoplankton cells to form communities is statistically robust, consistent with observed biomass changes, amenable to theoretical analysis, and a sentinel for long-term change. Since cell abundance is a community property in the same units for all marine microbes at any trophic level and at any phylogenetic position, it promises to integrate biological oceanography into general ecology and evolution.

Copepod feeding behavior and the measurement of grazing rates in vivo and in vitro

In this paper some aspects of the use of the gut fluorescence method for estimating ingestion rates have been, examined. One assumption is that gut turnover time in feeding copepods is equal to the gut clearance time in filtered seawater. When arctic Pseudocalanus were pre-fed on Thalassiosira weisflogii, and then given a trace addition of the same C14-labelled culture, or were transferred to filtered seawater, results suggested that this assumption was probably justified. In another experiment in which Pseudocalanus were fed at the same concentration of either melted ice algae, or pelagic under ice algae, there were significant differences in both gut clearance times and gut pigment levels in the two cases.Pigment: biogenic silica ratios in epontic algae were higher than those in faecal pellets produced by Pseudocalanus feeding on the algae, suggesting that pigment destruction was occurring during grazing. In a 28 hr time course experiment ingestion rates determined by rate of disappearance of particulate chlorophyll were higher than those simultaneously determined by the gut fluorescence method, which also supports the idea of pigment destruction in copepods guts.

Gut pigment accumulation and destruction by arctic copepods in vitro and in situ

The results presented here were obtained at six locations during three cruises in 1985 (off the coast of Labrador), 1986 (at the eastern end of Viscount Melbourne Sound) and 1988 (off the coast of Labrador). In situ chlorophyll maximum concentrations were >7 μgl-1 at depths of between 0 and 30 m in all sampling areas. In feeding experiments copepods attained higher gut pigment concentrations the longer they had been previously starved and higher concentrations when fed in the dark than when fed in the light. Community ingestion rates calculated from changes in particulate chlorophyll were higher than estimates derived from gut pigment data except when copepods had been starved for 24 h. Differences between estimates by the two methods suggested pigment destruction. In feeding experiments pigment: biogenic silica ratios in food and faecal pellets suggested that the length of starvation period affected the degree of pigment destruction differently at different stations and that feeding in the light greatly increased pigment destruction. A comparison of pigment: silica ratios in the water column, and in faecal pellets collected from copepods which had fed there, suggested that pigment destruction may occur in situ sometimes and that the degree to which it occurs may be affected by feeding history, light, diel feeding behaviour and species composition.

Basin-scale variability in plankton biomass and community metabolism in the sub-tropical North Atlantic Ocean

Abstract Three trans-Atlantic oceanographic surveys (Nova Scotia to Canary Islands) were carried out during fall 1992 and spring 1993 to describe the large-scale variability in hydrographic, chemical and biological properties of the upper water column of the subtropical gyre and adjacent waters. Significant spatial and temporal variability characterized a number of the biological pools and rate processes whereas others were relatively invariant. Systematic patterns were observed in the zonal distribution of some properties. Most notable were increases (eastward) in mixed-layer temperature and salinity, depths of the nitracline and chlorophyll- a maximum, regenerated production (NH 4 uptake) and bacterial production. Dissolved inorganic carbon (DIC) concentrations, phytoplankton biomass, mesozooplankton biomass and new production (NO 3 uptake) decreased (eastward). Bacterial biomass, primary production, and community respiration exhibited no discernible zonal distribution patterns. Seasonal variability was most evident in hydrography (cooler/fresher mixed-layer in spring), and chemistry (mixed-layer DIC concentration higher and nitracline shallower in spring) although primary production and bacterial production were significantly higher in spring than in fall. In general, seasonal variability was greater in the west than in the east; seasonality in most properties was absent west of Canary Islands (∼20°W). The distribution of autotrophs could be reasonably well explained by hydrography and nutrient structure, independent of location or season. Processes underlying the distribution of the microheterophs, however, were less clear. Heterotrophic biomass and metabolism was less variable than autotrophs and appeared to dominate the upper ocean carbon balance of the subtropical North Atlantic in both fall and spring. Geographical patterns in distribution are considered in the light of recent efforts to partition the ocean into distinct “biogeochemical provinces”.