Plankton of the fladen ground during FLEX 76 II. Population dynamics and production of Thysanoessa inermis (Crustacea: Euphausiacea)

Samples taken in the northern North Sea with the Continuous Plankton Recorder (CPR), the Undulating Oceanographic Recorder (UOR) and the Longhurst-Hardy Plankton Recorder (LHPR) during the Fladen Ground Experiment in 1976 (FLEX 76) are used to describe the vertical distribution and population dynamics of Thysanoessa inermis (Krøyer) and to provide estimates of the production and carbon budget of the population from 19th March to 3 June 1976. Spawning occurred in late April and early May, in near synchronisation with the start of the spring bloom of phytoplankton. Eggs, nauplii and calyptopes reached maximum abundance in succession, and furciliae were numerous when sampling ceased in early June. Adults increased in length from a mean of 12.1 mm in mid-March to 17.5 mm in early June and the estimated production was 2.40 mg m^-3 over the 74 d period. Total carbon ingested by the population of T. inermis was estimated to be 10 mg C m^-2 d^-1 in the upper 100m which was only 1.5% of the daily primary production of 0.68 gC m^-2 measured over the FLEX period 26 March to 4 June 1976. The grazing by T. inermis on the phytoplankton population was assumed to have little effect on the control and depletion of the spring phytoplankton bloom during FLEX 77.JOSDAP Contribution No. 50     

Plankton of the fladen ground during FLEX 76 I. Spring development of the plankton community

Results from plankton sampling in the northern North Sea with the Continuous Plankton Recorder (CPR) and the Undulating Oceanographic Recorder (UOR) during the Fladen Ground Experiment in 1976 (FLEX 76) are summarised. The first evidence of the spring outbreak of phytoplankton was on 19 April, the day after the first signs of vertical stability of the water column were observed. This was followed by spawning of the euphausiid Thysanoessa inermis and rapid increase in the numbers of Calanus finmarchicus. C. finmarchicus was the most abundant species over the FLEX period (19 March to 3 June) and, together with T. inermis, accounted for over 80% of the dry weight of the zooplankton standing stock. By early June the standing crop of phytoplankton had been depleted and nutrients levels were reduced to very low concentrations in the upper 50 m.JONSDAP Contribution No. 49     

High reproduction of <Emphasis Type="Italic">Calanus finmarchicus</Emphasis> during a diatom-dominated spring bloom

Feeding, egg production, hatching success and early naupliar development of Calanus finmarchicus were measured in three north Norwegian fjords during a spring bloom dominated by diatoms and the haptophyte Phaeocystis pouchetii. Majority of the copepod diet consisted of diatoms, mainly Thalassiosira spp. and Chaetoceros spp., with clearance rates up to 10 ml ind⁻¹ h⁻¹ for individual algae species/groups. Egg production rates were high, ranging from ca 40 up to 90 eggs f⁻¹ d⁻¹, with a hatching success of 70–85%, and fast naupliar development through the first non-feeding stages. There was no correlation between the egg or nauplii production and diatom abundance, but the hatching success was slightly negatively correlated with diatom biomass. However, the overall high reproductive rates suggested that the main food items were not harmful for C. finmarchicus reproduction in the area, although direct chemical measurements were not conducted. The high population egg production (>1,20,000 eggs m⁻² d⁻¹) indicated that a large part of the annual reproduction took place during the investigation, which stresses the importance of diatom-dominated spring phytoplankton bloom for population recruitment of C. finmarchicus in these northern ecosystems.     

Early reproduction and development of dominant calanoid copepods in the sea ice zone of the Barents Sea—need for a change of paradigms?

Reproduction and growth of the dominant copepods Calanus finmarchicus, C. glacialis, C. hyperboreus and Pseudocalanus minutus were studied on transects across the sea ice zone of the northern Barents Sea in May and June 1997. C. glacialis and C. finmarchicus were numerically dominant and also the largest component of the biomass. C. hyperboreus was rather rare. Moderate levels of phytoplankton and eventually high concentrations of ice algae supported maximum egg production rates of 53.6 and 48.5 eggs female^–1 day^–1 of C. glacialis in May and June, respectively. Results of incubation experiments were supported by a tremendous abundance of C. glacialis eggs in the water column ranging from 7×10³ to 4.4×10⁴ m^–2 in May and from 9.8×10³ to a maximum of 9.7×10⁴ m^–2 in June. In contrast, C. finmarchicus spawned only in the vicinity of the ice edge, at a maximum rate of 30 eggs female^–1 day^–1. Egg sacs of P. minutus were often observed in the preserved samples, but contained only few eggs, which may be due to loss during sampling. The presence of considerable concentrations of young stages in May and June indicated successful recruitment of C. glacialis and P. minutus. Back calculation using published stage duration estimates indicates March/April as the begin of the reproductive and growth period for these species under the first-year ice of the Barents Sea. Hence, secondary production in the study area starts at the same time as in open water regions and polynyas in the northern North Atlantic. Although the role of ice algae in the nutrition of copepods was not clarified here, the significant relationship between phytoplankton chlorophyll and egg production of C. glacialis suggests that high reproductive activity has already been achieved at moderate food concentrations.Communicated by O. Kinne, Oldendorf/Luhe     

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.     

Grazing rates of the copepods Calanus glacialis and C. finmarchicus in arctic waters of the Barents Sea

Natural feeding rates of Copepodite Stages IV and V, and adult female Calanus glacialis (Jaschnov) and Copepodite Stage V and adult female C. finmarchicus (Gunnerus) were estimated using fluorescence analysis of gut contents. Measurements were made on copepods sampled from arctic waters east of Svalbard (Barents Sea) during the spring phytoplankton increase, in the period from 27 May to 13 June, 1983. Observations on Copepodite Stages IV and V and adult female C. glacialis suggest that the gastric evacuation rate is independent of developmental stage, whereas C. finmarchicus Copepodite Stage V showed a lower gastric evacuation rate than adult females. Gut fullness displayed a low correlation with the ambient chlorophyll concentrations. Ingestion rates calculated for C. glacialis were 0.3, 2.3, and 11.0 g C h^-1 for Copepodite Stages IV and V and adult females, respectively. Copepodite Stage V and adult female C. finmarchicus ingested 0.9 and 1.1 g C h^-1 at a temperature of ca.-1.0°C. The maximum ingestion rate in terms of percent body carbon d^-1 was higher for adult female C. glacialis and C. finmarchicus than for the respective Copepodite Stage V's. The results are discussed both in relation to the physiological state of the species and to the environmental conditions.     

Production of DMSP and DMS during a mesocosm study of an Emiliania huxleyi bloom: influence of bacteria and Calanus finmarchicus grazing

We investigated the influence of bacteria and metazooplankton on the production of dimethylsulfoniopropionate (DMSP) and dimethylsulfide (DMS) during blooms of Emiliania huxleyi (Lohmann) Hay and Mohler in seawater mesocosms. The phytoplankton succession was marked by the rapid collapse of an initial Skeletonema costatum (Greville) Cleve bloom followed by a small E. huxleyi bloom. The collapse of the diatom bloom was accompanied by an increase in concentrations of dissolved DMSP (DMSP_d) and bacterial abundance and activity (as determined by the thymidine incorporation technique). The increase in bacterial activity was followed by a rapid decrease in DMSP_d concentrations which remained low for the rest of the experiment, even during the subsequent collapse of the E. huxleyi blooms. The absence of DMSP_d and DMS peaks during the declining phase of the E. huxleyi blooms was attributed to the high bacterial activity prevailing at that time. The influence of metazooplankton grazing on DMSP and DMS production was investigated by adding moderate (24 mg dry weight m^-3) and high (520 mg dry weight m^-3) concentrations of Copepodite Stage V and adults of Calanus finmarchicus to two of four filtered (200 m mesh net) enclosures during the E. huxleyi blooms. The addition of C. finmarchicus, even in high concentrations, had no apparent effect on the dynamics of E. huxleyi, suggesting that the copepods were not grazing significantly on nanophytoplankton. The addition of copepods in high concentrations favored an accumulation of chlorophyll a and particulate DMSP. These results suggest that copepods were preying on the herbivorous microzooplankton which, in turn, was controlling the biomass of nanophytoplankton. DMS production was also enhanced in the enclosure with maximum metazooplankton biomass, suggesting that the grazing of C. finmarchicus on microzooplankton containing DMSP may contribute to DMS production. These results provide strong support to the emerging idea that bacteria and metazooplankton grazing play a dominant role in determining the timing and magnitude of DMS pulses following phytoplankton blooms.     

Seasonal variations in the densities of fecal pellets produced by Oikopleura vanhoeffeni (C. Larvacea) and Calanus finmarchicus (C. Copepoda)

Two abundant macrozooplankters, Oikopleura vanhoeffeni (Lohmann) and Calanus finmarchicus (Gunnerus) were collected from the coastal waters off Newfoundland in different seasons during 1990–1991 and incubated in natural seawater to collect freshly egested, field produced, fecal pellets. The densities of fecal pellets from O. vanhoeffeni and C. finmarchicus were measured in an isosmotic density gradient. These are the first reported seasonal measurements of fecal pellet densities from two different types of macrozooplankters, O. vanhoeffeni, a larvacean, filter feeder and C. finmarchicus, a crustacean, suspension feeder. Pellet density ranges and medians were significantly different among seasons for both species, depending primarily on the type of phytoplankton ingested and its ability to be compacted. Winter O. vanhoeffeni and fall C. finmarchicus feces filled with nanoplankters and soft bodied organisms had less open space [packing index (% open area) = 3.5 and 4% for O. vanhoeffeni and C. finmarchicus, respectively] and were more dense (1.23 and 1.19 g cm^-3) than spring feces filled with diatoms (packing index = 15 and 23%, density = 1.13 and 1.11 gcm^-3). For copepods, these results contrast with previously published density values and with the predicted copepod fecal pellet density calculated, in the present study, using the conventional mass/volume relationship. Copepod spring and summer diatom-filled feces had a calculated density of 1.12 and 1.24 gcm^-3 vs a measured median density of 1.11 gcm^-3 for both spring and summer feces; the fall feces containing nanoplankters had a calculated density of 1.08 gcm^-3 vs a measured median density of 1.19 gcm^-3. Knowledge of the seasonal variations in fecal pellet densities is important for the development of flux models.     

Phytoplankton exudate release as an energy source for the growth of pelagic bacteria

The release of dissolved organic carbon (DOC) from phytolankton during photosynthesis, and the utilization of this carbon by planktonic bacteria, was studied using ¹⁴CO₂ and selective filtration. Natural sea water samples from a coastal area of the Northern Baltic Sea were incubated in the laboratory for detailed studies, and in situ for estimation of annual dynamics. In a laboratory incubation (at +1°C) the concentration of ¹⁴C-labelled dissolved organic carbon increased for about 2 h and then reached a steady state, representing about 0. 1% of the total DOC. Labelled organic carbon in the phytoplankton and bacterial fractions continued to increase almost linearly. The continuous increase in the bacterial fraction is thought to represent almost instantaneous utilization of the DOC released from the phytoplankton during photosynthesis. As an annual average, in 4 h in situ incubations, about 65% of the labelled organic carbon was found in the phytoplankton fraction (>3 m), about 27% in the bacterial fraction (0.2 to 3 m) and the remaining 8% as DOC (<0.2 m). Large variations in these percentages were recorded. The measured annual primary production was 93 g C m^-2 (March to December), and the estimated bacterial production due to phytoplankton exudates 29 g C m^-2. This represents a release of DOC of about 45% of the corrected annual primary production of 110 g C m^-2 (assuming a bacterial growth efficiency of 0.6).     

Aspects of the ecology of a small estuarine embayment

A 1.8 ha brackish (5 to 15 S) embayment (Osborn Cove) on the Western Shore of Chesapeake Bay (USA) was studied during 1976 to examine some hydrologic and climatic influences on its phytoplankton, bacteria, intertidal benthos, a peripheral salt marsh (equivalent to 20% of the cove surface area), and the surrounding 48 ha forest watershed. Comparisons with 1975 and 1977 for temperature, salinity, rainfall and tidal extremes, show 1976 to have had normal rainfall but a cooler autumn. Sediment moves alongshore into the cove after rainfall, and erosion causes soil breakdowns from nearby cliffs. This movement, ice damage and predators appear to mediate distribution of the intertidal benthos. Phytoplankton density, chlorophyll and photosynthesis are compared with other portions of the Chesapeake Estuary sampled in parallel programs. Phytoplankton chlorophyll oscillations observed in the Bay and Potomac River were not seen in the cove. Cove gross and net photosynthesis averaged about the same as the bay, but the cove had higher rates in spring, a result of significantly higher net assimilation ratios rather than higher biomass. River-contributed nutrients may have produced this stimulation when used by tidally inoculated phytoplankton. Large numbers of small flagellates were not seen after heavy rainfall fluishing. Net phytoplankton production in the cove was estimated at 97.6 g C m^-2 yr^-1. A portion of the cove having restricted circulation apparently contributed 48% of this production in months when its phytoplankton was dominated by small flagellates. Total estimated net production by cove phytoplankton was 1.75x10³ kg C yr^-1, compared to 0.75x10³ kg C yr^-1 for a narrow peripheral zone of Spartina alterniflora growth occupying only 13% as much area. This ratio and circumstantial evidence suggests that leaf litter from the surrounding forest dominated particulate input to the cove. Bacterial plate counts showed increases in total numbers as a function of water temperature, with surface counts exceeding bottom counts. Indigenous bird and mammal waste are suggested as important bacterial inputs. Rainfall pulses resulted in rapid increases of fecal coliform and fecal streptococcus counts.     

Population dynamics and production of euphausiids II. Thysanoessa inermis and T. raschi in the North Sea and American Coastal Waters

Results from the Continuous Plankton Recorder (CPR) survey for 1966 and 1967 are used to describe seasonal changes in abundance, size and aspects of the population structure of Thysanoessa inermis (Krøyer) and T. raschi (M. Sars) at a depth of 10 m in the North Sea and in American coastal waters from the Grand Banks to the Gulf of Maine. Production and dry weight were estimated from these data. Two year-groups were usually present in the breeding population, the proportion surviving into a second year being higher in American waters than in the North Sea. Annual production for each species was within the range 0.69 to 4.66 mg m^-3 and the ratio between production and biomass (P:B) was between 1.3 and 4.2; values outside these ranges were obtained only for American coastal waters in 1967, when the frequency of sampling was low.     

Aspects of water column primary productivity in the Chukchi Sea during summer

Measurements of primary productivity, chlorophyll a, incident solar radiation, phosphate-P, silicate-Si, nitrate-N, nitrite-N, ammonium-N, temperature and salinity were made in the Marginal Ice Zone of the Chukchi Sea in summer 1974. Low to moderate levels of primary productivity (0.07 to 0.97 g C m^-2 half-day^-1) were observed; primary productivity exceeded 3 g C m^-2 half-day^-1 at two stations. Surface primary productivity was nitrogen-limited at most stations. Mean chlorophyll a concentration in the photic zone varied from 0.4 to 17.8 mg m^-3. Higher concentrations and significant subsurface accumulation of chlorophyll a, reaching 40 mg m^-3, were observed in July at stations near the ice-edge than those in open water. No chlorophyll maximum was noted in September, when values ranged from 0.4 to 2.2 mg m^-3. It is postulated that the contribution of sea-ice algae to the total chlorophyll content can be substantial, but that the stay of these cells in the water column may not be long. Non-linear regression estimates from solar radiation and chlorophyll-specific primary productivity data showed a maximal photosynthetic rate of 18 mg C mg chlorophyll a ^-1 half-day^-1, an optimal light intensity of 54 langleys half-day^-1, and markedly reduced primary productivity at moderately higher light intensities. These features indicate that phytoplankton was shade-adapted.     

Toxin accumulation and feeding behaviour of the planktonic copepod Calanus finmarchicus exposed to the red-tide dinoflagellate Alexandrium excavatum

The planktonic copepod Calanus finmarchicus is a dominant member of the zooplankton community in the lower St. Lawrence Estuary in eastern Canada. Blooms of the toxic marine dinoflagellate Alexandrium excavatum which produces high cellular levels of paralytic shellfish poisoning (PSP) toxins, occur during the period of high C. finmarchicus production in summer in this region. To study the feeding behaviour of C. finmarchicus in the presence of Alexandrium spp., experiments were conducted in which female adult copepods collected from the St. Lawrence Estuary between May and September 1991 were exposed under controlled conditions to two toxic isolates of A. excavatum (Pr18b and Pr11f) from the estuary and to a non-toxic control (PLY 173) of a closely related species, A. tamarense isolated from the Tamar Estuary, Plymouth, U.K. Clearance rates on non-toxic A. tamarense cells averaged 5.5 ml ind^-1 h^-1 but were nearzero with either toxic isolate. When presented with a mixture of A. excavatum and the non-toxic diatom Thalassiosira weissflogii in varying proportions, C. finmarchicus fed upon the diatom but avoided the toxic dinoflagellate. Although feeding rates on A. excavatum were very low, toxin analysis by high-performance liquid chromatography with fluorescence detection (HPLC-FD) revealed that the PSP toxins were accumulated in copepods exposed to toxigenic dinoflagellates.The toxin composition in copepods was similar to that of the toxic dinoflagellate, but not necessarily identical, particularly after short-term (2-h) exposure, when relatively elevated levels of N-sulfocarbamoyl toxins were detected. The evidence suggests that C. finmarchicus ingests toxic dinoflagellate cells, either mistakenly or during exploratory bouts of feeding, and accumulates PSP toxins in its gut system and perhaps in other tissues.     

Does the spawning of Calanus finmarchicus in high latitudes follow a reproducible pattern?

In the spring of 1989, an experimental study of the spawning behaviour of Calanus finmarchicus was carried out in Malangen, northern Norway. Here, a single cohort of females reproduce from mid-March to May, approximately coinciding with the wax and wane of the spring phytoplankton bloom. An evaluation of population characteristics such as the proportion of adults, sex ratio, as well as gonad maturation and daily productivity of the females clearly reveals three phases within the population's reproductive period. In between incline and decline, the highest spawning rates (on average >20 eggs female^-1 d^-1, equivalent to 5.7% body C d^-1) occur after the males have disappeared from the population and almost all females have mature gonads. During this period, the ratio of adults to copepodid Stage Vs changes from dominance of adults to that of CVs. Although first egg production was observed prior to the phytoplankton increase, it is suggested that the onset of the phytoplankton spring bloom in the first few days of April enhances the final maturation of ovaries in the females and therefore triggers the onset of the main spawning period. The clutch sizes (max. 95 eggs clutch^-1) vary with the age of the females, while the spawning frequencies depend on the available food quantities. The overlap of an estimated minimal 4 wk spawning period for the individuals leads to a main reproductive phase for the population of ca. 3 wk, during which time mean clutch sizes and spawning frequencies are maximal (highest average clutch size: 70 eggs female^-1 clutch^-1, 100 to 60% of the females spawning). This period ends before the end of the phytoplankton bloom. Calculated by stepwise interpolation and summation of the mean daily egg production in the population, an average female produced ca. 600 eggs during the spring bloom in Malangen 1989. We suggest that reproduction and population development of C. finmarchicus in spring follows a reproducible pattern for a given temperature regime and non-limiting food conditions. In the case of clearly identifiable cohorts, it seems possible to trace the state of reproduction by evaluating population parameters.     

Phytoplankton community structure and primary production in small intertidal estuarine-bay ecosystem (eastern English Channel,France)

From May 2002 to October 2003, a fortnightly sampling programme was conducted in a restricted macrotidal ecosystem in the English Channel, the Baie des Veys (France). Three sets of data were obtained: (1) physico-chemical parameters, (2) phytoplankton community structure illustrated by species composition, biovolume and diversity, and (3) primary production and photosynthetic parameters via P versus E curves. The aim of this study was to investigate the temporal variations of primary production and photosynthetic parameters in this bay and to highlight the potential links with phytoplankton community structure. The highest level of daily depth-integrated primary production Pz (0.02–1.43 g C m⁻² d⁻¹) and the highest maximum photosynthetic rate P ^B _max (0.39–8.48 mg C mg chl a ⁻¹ h⁻¹) and maximum light utilization coefficient α^B [0.002–0.119 mg C mg chl a ⁻¹ h⁻¹ (μmol photons m⁻² s⁻¹)] were measured from July to September. Species succession was determined based on biomass data obtained from cell density and biovolume measurements. The bay was dominated by 11 diatoms throughout the year. However, a Phaeocystis globosa bloom (up to 25 mg chl a m⁻³, 2.5 × 10⁶ cells l⁻¹) was observed each year during the spring diatom bloom, but timing and intensity varied interannually. Annual variation of primary production was due to nutrient limitation, light climate and water temperature. The seasonal pattern of microalgal succession, with regular changes in composition, biovolume and diversity, influenced the physico-chemical and biological characteristics of the environment (especially nutrient stocks in the bay) and thus primary production. Consequently, investigation of phytoplankton community structure is important for developing the understanding of ecosystem functioning, as it plays a major role in the dynamics of primary production.     

In situ net primary productivity and photosynthesis of Antarctic sea ice algal, phytoplankton and benthic algal communities

Primary production at Antarctic coastal sites is contributed from sea ice algae, phytoplankton and benthic algae. Oxygen microelectrodes were used to estimate sea ice and benthic primary production at several sites around Casey, a coastal area in eastern Antarctica. Maximum oxygen export from sea ice was 0.95 mmol O₂ m⁻² h⁻¹ (~11.7 mg C m⁻² h⁻¹) while from the sediment it was 6.08 mmol O₂ m⁻² h⁻¹ (~70.8 mg C m⁻² h⁻¹). When the ice was present O₂ export from the benthos was either low or negative. Sea ice algae assimilation rates were up to 3.77 mg C (mg Chl-a)⁻¹ h⁻¹ while those from the benthos were up to 1.53 mg C (mg Chl-a)⁻¹ h⁻¹. The contribution of the major components of primary productivity was assessed using fluorometric techniques. When the ice was present approximately 55–65% of total daily primary production occurred in the sea ice with the remainder unequally partitioned between the sediment and the water column. When the ice was absent, the benthos contributed nearly 90% of the primary production.     

Trophic implications of cross-shelf copepod distributions in the Southeastern Bering Sea

Spring distributions of some numerically dominant copepods reflect associations with two distinct water masses separated along the 80- to 100-m isobaths. Seaward of this middle shelf front, the oceanic Bering Sea hosts populations of Calanus cristatus, C. plumchrus, and Eucalanus bungii bungii; Metridia pacifica, Oithona similis, and Pseudocalanus spp. are also present. The large oceanic species are much less abundant in waters shallower than 80 m where the community is seasonally dominated by smaller copepods, O. similis, Acartia longiremis, and Pseudocalanus spp. Experimental and field-derived estimates of carbon ingestion indicate that the oceanic/outer shelf copepods can occasionally graze the equivalent of the daily plant production and probably routinely remove 20–30% of the primary productivity. Conversely, stocks of middle shelf copepods rarely ingest more than 5% of the plant carbon productivity. During 45 d between mid April to late May, 1979, approximately three times more organic matter was ingested m^-2 by the outer shelf/oceanic copepod community than by middle shelf species. This imbalance in cross-shelf grazing permits middle shelf phytoplankton stocks to grow rapidly to bloom proportions, and to sink ungrazed to the seabed. Over the outer shelf and particularly along the shelf break, a much closer coupling to phytoplankton supports a large biomass of oceanic grazers. Here, copepod stocks approaching 45 g dry wt m^-2 occur in late spring as a narrow band at the shelf break.Supported by National Science Foundation Grant DPP 76-23340Contribution no. 485, Institute of Marine Science, University of Alaska, Fairbanks     

Defensive mechanisms and ecology of some tropical holothurians

The ecology, defensive behavior and toxicity of three species of reef flat holothurians (Actinopyga mauritiana, Holothuria atra and Holothuria difficilis) were studied at Eniwetok Atoll, Marshall Islands. The average diurnal population density of H. difficilis ranged from 1.4 to 32 holothurians/900 cm²; resting respiratory rates (0.05 ml O₂/g wet wt/h) were comparable during day and night; nourishment in H. difficilis may be primarily from bacteria and foraminifera in which about 2% of the dry weight of sediment consumed is utilized, and the species probably passes at least 3 g dry wt of sediment/m²/day (>1 kg/m²/year). A fundamental difference in energy flow is suggested: considerably more energy is passed from benthic algae to grazing and browsing fishes to predatory fishes on coral reefs whereas, in extra-tropical latitudes, more energy is shunted from benthic algae to invertebrates to predators. The effects of holothurin leading to death in fishes are irreversible. Holothuria difficilis is best protected from predation. Its body wall is toxic and it can accurately eject Cuvierian tubules, which are also toxic. The discharge of tubules was regulated by a circadian rhythm in May. Studies on holothurians and sponges suggest that many exposed coral reef invertebrates have evolved effective defensive mechanisms in association with high intensity predation.Supported by A.E.C. Contract AT (29-2)-226 with the University of Hawaii.     

Diel changes in phytoplankton photosynthetic efficiency in Brackish waters

From 18 to 23 September 1974, investigations on the diel changes in phytoplankton were carried out in the Baltic Sea. Every 4 h, water samples were collected from 2 and 15 m, and PO₄, chlorophyll a, temperature, salinity, pH, phytoplankton composition and phytoplankton light photosynthesis relationship were determined. Continuous measurements of surface irradiance and some estimations of zooplankton were also made. P ^B (photosynthesis per unit chlorophyll a at low light levels of 2·10^-2 cal cm^-2 min^-1) revealed only random variation during the sampling period, i.e., 1.0 to 1.6 mg C (mg chlorophyll a)^-1 h^-1. P _m ^B (Light-saturated photosynthesis per unit of chlorophyll a) displayed pronounced diel fluctuations with the highest value of about 6 mg C (mg chlorophyll a)^-1 h^-1 around noon, and the lowest value of about 2.5 mg C (mg chlorophyll a)^-1 h^-1 during the night, during which latter period the value of P _m ^B was more or less constant. Reasons for the diel fluctuations are discussed, and an equation which describes these fluctuations is proposed. Using this equation, the daily phytoplankton production estimated in incubators by a previously described method can be corrected for the time of day at which samples are collected.     

Seasonal composition of meroplankton and holoplankton in the Bristol Channel

A decreasing gradation in the plankton standing stock of the Bristol Channel was observed from the seaward section to the inner, less saline, reaches. Two sub-regions of our survey, the North Outer Channel (NOC) and the Inner Channel (IC), represented the extremes of this gradient and were selected for detailed comparison. The integrated zooplankton biomass, over the 307 d sampling period (4 November 1973 to 6 September 1974), was 2 475 mg C m^-3 (266 mg C m^-2 d^-1) in the NOC and 335 mg C m^-3 (20 mg C m^-2 d^-1) in the IC. The omnivorous plankton accounted for 76% of the standing stock in the NOC and 89% in the IC, of which 58 and 23% were meroplankton and 39 and 71% were holoplankton, respectively; the remainder was unassigned. The majority of the meroplankton in both subregions was decapod larvae and adults, whereas the holoplankton biomass was dominated in the NOC by copepods (89%) and in the IC by mysids (57%), mainly Schistomyzis spiritus. Centropages hamatus was the most abundant copepod species in the NOC and accounted for 32% of the total holoplankton omnivore standing stock. In the NOC and IC, the carnivorous plankton accounted for 24 and 11% of the total plankton biomass, respectively. In the two sub-regions, 20 and 21% of the carnivores were meroplanktonic (primarily larvae of sprats and pilchards), while the holoplankton carnivores contributed 75 and 74% to the NOC and IC, respectively (Sagitta elegans, Pleurobrachia pileus). S. elegans dominated the holoplankton carnivore biomass for the majority of the year and accounted for 96% in the NOC and 60% in the IC. The integrated total particulate carbon over the 307 d period was 200 g C m^-3 (6 600 g C m^-2) in the NOC and 838 g C m^-3 (15 084 g C m^-2) in the IC. The annual primary production ranged from 164.9 g C m^-2 yr^-1 in the Outer Channel (North and South) to 6.8 g C m^-2 yr^-1 in the IC. The zooplankton biomass reached a maximum in July. The total particulate carbon (TPC) in July was 400 mg C m^-3 in the NOC of which ca. 78 mg C m^-3 were phytoplankton and ca. 21 mg C m^-3 were zooplankton; these values compare favourably with those found in the adjoining Celtic Sea. In the IC, the TPC was 2 800 mg C m^-3, of which ca. 107 mg C m^-3 were phytoplankton and 2.8 mg C m^-3 were zooplankton. From the low primary production estimates for the IC it can be concluded that the majority of the chlorophyll, like the TPC, was allochthonous in origin. Furthermore it is suggested that zooplankton plays a minor role in this estuarine ecosystem and is not the main consumer of the suspended particulate carbon; the benthic filter-feeding communities are presumed to fulfill this role in the Bristol Channel.